Cryptantha maritima (Greene) Greene var. vizcainensis Rebman & M.G.Simpson, 2021

Simpson, Michael G. & Rebman, Jon P., 2021, Research in Boraginaceae: A new variety of Cryptantha maritima, Cryptantha pondii resurrected, and Johnstonella echinosepala transferred back to Cryptantha, Phytotaxa 509 (2), pp. 185-210 : 190-195

Cryptantha maritima (Greene) Greene var. vizcainensis Rebman & M.G.Simpson , var. nov. ( Figs. 3 & 4 )

Type:— MEXICO. Baja California Sur: Picachos de Santa Clara, north slope of SE peak, occasional on north slope, 600 meters elevation, 27˚07’ N, 113˚37’ W, 3 February 1973, R. Moran & J. L. Reveal 19707 (holotype SD92511 !, isotype US02908745= US 2796919!) [Note: georeferenced coordinates that were estimated from label data are indicated with “*”, otherwise verbatim from label data.]

Paratypes (alphabetical by collector and number): — MEXICO. Baja California Sur: Vizcaino peninsula south of Laguna Ojo de Liebre, at the eastern base of the “Sierra de Vizcaino ” in a portion of the range known as Sierra Campo Nuevo, at the end of a small track winding 6.1 miles west of the road to Bahia Asuncion, 3.6 miles south of the junction with the road to Bahia Tortugas, rocky upper bajada and steep rocky slopes with low scrub of Ambrosia chenopodiifolia , Pachycormus discolor , Stenocereus , Pachycereus , Errazurizia and some Larrea , common annual, flowers white, 300– 500 meters elevation, near 27°21’ N, 114°10’W, 30 April 1993, S. Boyd et al. 8086 ( MEXU 666794; RSA 576907!); rocky slopes and outwash plain with Pachycormus , Bursera , Fouquieria , Ambrosia and Jatropha , E base of Sierra de Placeros, 40 km SE of San Jose de Castro, flowers white, 366 meters elevation, 27.44347˚ N, 114.116654˚ W, 24 March 1984, D. E. Breedlove 60891 ( MEXU 484431, RSA 497744!); 35 km SE of Bahia Tortugas, pacific slope of Sierra de Placeros, rocky slope with Bursera , Jatropha , Pachycormus , Ambrosia and Simmondsia , flowers white, 460 meters elevation, 27.60463˚ N, 114.603˚ W *, 7 March 1985, D. E. Breedlove 62324 ( MEXU 484782, RSA 497109!); eastern bajada of Sierra Calvario, Systema de Sierra Vizcaino , small perennial bush, desert of dispersed succulent trees and suffrutescent shrubs, 60–243 meters elevation, 27.4125˚ N, 114.220833˚ W *, 10–15 March 1947, H. S. Gentry 797 ( ARIZ 123309!); Picachos de Santa Clara, 350 meters elevation, 27˚09’ N, 113˚40’ W, 3 February 1973, R. Moran & J. L. Reveal 19663 ( SD 92540!, US 02908744= US 2796920); 8 road miles northwest of Asunción, fairly common in bed of arroyo, 70 meters elevation, 27˚13’ N, 114˚21’ W, 4 February 1973, R. Moran & J. L. Reveal 19758 ( SD 92558!, US 02908741= US 2796922); Arroyo Malarrimo, 11 miles south of the mouth, occasional in arroyo, 75 meters elevation, 27˚29’ N, 114˚29’ W, 6 February 1973, R. Moran & J. L. Reveal 19878 ( SD 92358!); at pass at head of Arroyo Largo, occasional, 480 meters elevation, 27˚36’ N, 114˚39’ W, 8 February 1973, R. Moran & J. L. Reveal 19949 ( SD 92556!, US 02908742= US 2796924); north slope of Cerro Azul, fairly common near base, 400 meters elevation, 27˚32’ N, 114˚32’ W, 9 February 1973, R. Moran & J. L. Reveal 19973 ( SD 92328!); Arroyo Calvario 6.0 miles north of San Andrés, 130 meters elevation, 27˚20’ N, 114˚26’ W, 10 February 1973, R. Moran & J. L. Reveal 20004 ( MEXU 220305, SD 92509!, US 02908748= US 2796916).

Description:—Plants annual herbs, 12–40 cm tall, canescent. Stems stout, woody in texture, 2–5 mm in diameter, not reddish, with an erect primary shoot giving rise to several inclined to ascending or decumbent secondary shoots at base and along primary axis, secondary shoots slightly shorter than or as long as primary, stem vestiture canescent, antrorse-appressed strigose, trichomes numerous, whitish, ca. 1 mm long. Leaves spiral, absent at plant base (possibly deciduous) or present, these densely clustered, generally shriveled, cauline leaves larger, becoming reduced toward inflorescence, mostly flat, ca. 10–50 mm × 1–3 mm, sessile, base widely cuneate, margin entire to irregularly dentate, apex acute to rounded, midrib flat to very slightly raised, both surfaces hirsute and epustulate to hispid and basally pustulate, trichomes whitish, horizontal to ascending, wider at base, point of attachment swollen (termed a “pustule”), consisting of 1–3 concentric rows of white to transparent, radially elongate cells, especially prominent in shriveled, basal leaves. Inflorescence a series of ascending, racemosely arranged, circinate scorpioid cymules, these clustered in younger plants, elongating at maturity, peduncles ca. 5–20 mm long, commonly with a flower at junction with generating axis, cymules basally bracteate, bracts similar to leaves but reduced in size, ca. 20 flowers per cymule, fruits not touching at maturity, mostly 2–3 mm apart, more so near cymule base, axes canescent antrorse appressedstrigillose, trichomes ≤ ca. 0.5 mm long. Flowers typically bracteate near base of cymule, sporadically bracteate distally. Calyx ascending to inclined in fruit, lance-ovoid in shape, 1–2 mm long in flower, 2–3 mm in fruit, sepals distinct, erect, narrowly lanceolate, apically acute to rounded, apices slightly recurved, adaxial surface glabrous in lower half, sparsely appressed hirsute in upper, trichomes <0.5 mm, abaxial surface hirsute with often dense, appressed to ascending white trichomes ca. 0.5 mm long along and inside margin, midrib thickened and hispid (mostly on sepals away from cymule axis), trichomes ascending to inclined, stout, basally thickened, white, straight to sometimes curved, ca. 1.5–2 mm long. Corolla white, rotate, tube same length as calyx, limb 3–5 (1–1.5) mm in diameter [Note: measured from dried herbarium material of all types], appendages (fornices) present, color unknown. Gynobase very narrowly conical, ≥3/4 nutlet length. Style ca. 0.3 mm long, tip/stigma extending ca. 0.1 mm beyond large nutlet. Ovary lobes/ovules two. Nutlets two, erect, brown, symmetric, lance-ovate, adaxially shallowly 2-planed convex, abaxially broadly convex, heteromorphic, larger nutlet smooth, shiny, adjacent to cymule axis (axial), more firmly attached to gynobase, 1.5–1.7 mm × ca. 0.6 mm, ventral groove slightly open, groove sides not raised, minutely bifid at base with no to a very small areole, smaller nutlet, 1.2–1.3 mm × ca. 0.5 mm, minutely tuberculate both surfaces, with ca. 16–18 tubercles across dorsal face at greatest width, tubercles low, brown to light brown, papillae not observed, ventral groove closed, minutely bifid at base, no areole observed.

Diagnosis:— Cryptantha maritima var. vizcainensis is similar to most individuals of C. maritima var. cedrosensis in having relatively large (3–5 mm) corollas. It differs in having a canescent stem vestiture with appressed trichomes only, as opposed to a strigose and hirsute vestiture with both appressed and spreading trichomes in the latter, and in having two ovary lobes and ovules per ovary, these typically developing into two nutlets, as opposed to four ovary lobes and ovules and up to four nutlets per fruit in the latter.

Distribution, Habitat, and Endemism:— Cryptantha maritima var. vizcainensis is endemic to the western Vizcaíno Desert region of Baja California Sur ( Fig. 2A,B ). The habitat of the Boyd 8086, Breedlove 60891 & 62324, and Gentry 797 paratype specimens is described from specimen label information as “rocky upper bajada and steep rocky slopes with low scrub,” “rocky slopes and outwash plain,” “rocky slope,” and “bajada,” respectively.Associated plants recorded from these collections were: Ambrosia , Ambrosia chenopodiifolia (Benth.) W.W.Payne , Bursera , Errazurizia , Fouquieria , Jatropha , Larrea , Pachycereus , Pachycormus , Pachycormus discolor (Benth.) Coville ex Standl. , Simmondsia , and Stenocereus, The habitat and vegetation were not described in the label information of the vouchers collected by Reid Moran, including the holotype and isotypes. However, based on Moran’s field notes (http:// bajaflora.org/MoranNotesSearch.aspx) for the period of time of these collections (3–10 February 1973), the following dominant plants are recorded as occurring in the vegetation type where some of his specimens were collected (listed with current taxonomy): Ambrosia magdalenae (Brandegee) W.W.Payne , Apiastrum angustifolium Nutt. in Torr. & A.Gray, Bahiopsis microphylla (Vasey & Rose) E.E.Schill. & Panero , Bebbia juncea (Benth.) Greene , Condea emoryi (Torr.) Harley & J.F.B.Pastore , Cylindropuntia cholla (F.A.C.Weber) F.M.Knuth, Ebenopsis confinis (Standl.) Barneby & J.W.Grimes , Encelia stenophylla Greene , Emmenanthe penduliflora Benth. , Eriogonum fasciculatum Benth. var. emphereium Reveal , Gambelia juncea (Benth.) D.A.Sutton, Jatropha cinerea (Ortega) Müll.Arg. , Lophocereus schottii (Engelm.) Britton & Rose , Prosopis glandulosa var. torreyana (L.D.Benson) M.C.Johnst. , Salvia cedrosensis Greene , Stenocereus gummosus (Engelm.) A.C.Gibson & K.E.Horak, and Yucca valida Brandegee.

Phenology:— Based on data from available specimens, Cryptantha maritima var. vizcainensis appears to flower from February to April.

Etymology:— The varietal epithet vizcainensis means “of the Vizcaíno Desert ,” where it is endemic.

Suggested Common Name:— Vizcaíno Desert Cryptantha .

Rationale for rank:— This new taxon is unique in its combination of a larger corolla, two ovules and two 1-seeded nutlets per fruit, and canescent-strigose stem vestiture. One collection (Moran 19758, SD92558) was mixed with one individual typical of the new variety and one with small corollas (limb 1–1.5 mm wide), a potential intermediate to C. maritima var. maritima . Thus, we chose to retain this new taxon as a variety of Cryptantha maritima , as it is otherwise quite similar to the other members of that species.

Conservation status:— This new variety occurs in a limited region of the western Vizcaíno Desert of Baja California Sur and is currently known from only 11 collections (see Fig. 2B ). This region has been relatively undercollected and is dependent on late monsoonal, hurricane driven, or early winter rains for annuals to grow. It should be noted that this part of the peninsula lies between the normal northern winter and southern summer rainfall regimes; as a result, rains are highly unpredictable, and the region is very dry in most years. Given the sparsity of collections and limited range, we suggest that Cryptantha maritima var. vizcainensis be listed as rare, equivalent to a CNPS ranking of 1B.1 ( CNPS 2021).

Identification of the varieties of Cryptantha maritima :— Aside from the features mentioned earlier, we present a comparison of the fruiting calyces of the four varieties of C. maritima in Fig. 5A . Cryptantha maritima vars. maritima and pilosa in particular can be difficult to distinguish because they are similar in stem vestiture, corolla size, and nutlet morphology and overlap significantly in their distribution. Note that C. m. var. maritima can be finely hirsute along and on the surface of the abaxial sepal margins, but the trichomes tend to be fewer, shorter, and appressed; the midribs are hispid with stouter trichomes more or less spreading in orientation ( Fig. 5B ). In contrast, C. m. var. pilosa is finely hirsute along and on the surface of the abaxial sepal margins, the trichomes more numerous, longer, and upwardly inclined to erect; the midrib is hispid, but the trichomes are fewer, thinner, and more ascending to erect in orientation ( Fig. 5C ).

A revised key to the four varieties of Cryptantha maritima , including our new taxon, is presented here:

1. Stem trichomes appressed and spreading; ovules 4, nutlets 1–4; endemic to Cedros Island, Baja California .................................... ................................................................................................................................................................ C. maritima var. cedrosensis

- Stem trichomes appressed and spreading or appressed only; ovules 2, nutlets 1–2; islands and mainland of the Baja California peninsula, southwestern United States, and/or Sonora, Mexico........................................................................................................2

2. Corolla limb 3–5 mm wide; stem strigose-canescent, trichomes only or predominantly appressed; endemic to the foothills and mountains of the western Vizcaíno Desert , Baja California Sur ........................................................... C. maritima var. vizcainensis

- Corolla limb 1–2 mm wide; stems strigose and hispid, trichomes both appressed and spreading; widespread ................................3

3. Calyx of mature fruits densely white-hirsute on marginal abaxial surface, trichomes inclined to erect, longest generally ≥ 1 mm long; midribs hispid, trichomes thin, ascending to erect................................................................................. C. maritima var. pilosa

- Calyx of mature fruits sparsely to densely white-hirsute on marginal abaxial surface, trichomes appressed, longest generally <1 mm long; midribs hispid, trichomes stout, generally horizontal to reclined .............................................. C. maritima var. maritima

CRYPTANTHA PONDII

Figures

FIGURE 2. Distribution maps of varieties of Cryptantha maritima. Data from Baja Flora (2021), CCH2 (2021), GBIF (2020), and SEINet (2021), all specimens annotated by the authors (Appendix 1). A, B. Distribution in North America. A. Full distribution map. B. Close-up of Vizcaíno Peninsula and adjacent islands. Note location of type populations and restriction of C. maritima var. cedrosensis to Cedros Island and of C. maritima var. vizcainensis to the western Vizcaíno region. Black dots=C. maritima var. maritima; red dots=C. maritima var. pilosa; green triangles= C. maritima var. cedrosensis; yellow squares= C. maritima var. vizcainensis. C. Distribution of C. maritima var. pilosa in Argentina, South America, data from GBIF (2020).All maps from ©Google 2021, INEGI Data.

FIGURE 3. Cryptantha maritima var. vizcainensis. Images from Moran 19878 (SD92358).A. Close-up of base of primary shoot, showing strigose stem vestiture and pustulate trichomes of leaves. B. Corolla face, showing relatively large limb diameter and fornices. C. Mature fruit, calyx opened, showing two nutlets and style, extending just beyond larger, smooth nutlet. D. Close-up of cymule, showing appressedstrigose stem vestiture, flower bract, calyces with hirsute marginal regions of ascending trichomes, and hispid midribs with stout, inclined or spreading trichomes, the latter more prominent on the side away from the inflorescence axis.

FIGURE 4. Cryptantha maritima var. vizcainensis. Images from Moran 19878 (SD92358). A. Gynobase, nutlets, and style/stigma detached from a single fruit. B. Larger, smooth nutlet, in dorsal (left) and ventral (right) views. C. Smaller, tuberculate nutlet, in dorsal (left) and ventral (right) views. Note ventral groove, short bifid at base.

FIGURE 5. Comparisons of fruiting calyces of Cryptantha maritima varieties. A. Cryptantha maritima varieties, left-to-right: var. cedrosensis, var. maritima, var. pilosa, and var. vizcainensis. B. Two samples of var. maritima; note hirsute calyx margin region with few to many appressed trichomes and hispid calyx midrib with stout, mostly spreading trichomes. C. Two samples of var. pilosa; note densely hirsute calyx margin region with ascending to erect trichomes and hispid midribs with few, relatively thin, erect to ascending trichomes.