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Acanthogilioideae

 Subfamily

Acanthogilioideae J. M. Porter & L. A. Johnson

Aliso 19: 60, 2000.

    Shrubs with persistent, alternate, spiny, pinnately lobed primary leaves and deciduous, linear secondary leaves that are axillary and fascicled on short shoots.  Calyx and corolla radially symmetric, corolla salverform.  Fruits dehiscing only about half the length, either loculicidal or both loculicidal and septicidal.  Seeds winged. (Phylogenetic definition.—The monophyletic group descended from the first ancestor of Acanthogilia gloriosa to evolve persistent, spinescent primary leaves, formed by the lignification of the vascular bundles of the leaf, and simultaneously zonocolporate pollen bearing verruca.)  Type: Acanthogilia A. G. Day & Moran.

 Membership.Acanthogilia.

 

 Tribe

Acanthogilieae V. E. Grant

Amer. J. Bot. 85: 744, 1998.

    Shrubs with persistent, alternate, spiny, pinnately lobed primary leaves and deciduous, linear secondary leaves that are axillary and fascicled on short shoots.  Calyx and corolla radially symmetric, corolla salverform.  Seeds winged. (Phylogenetic delineation.See Acanthogilioideae, with which this taxon is redundant; recognition of this tribal rank is maintained for uniformity of the classification.)  Type: Acanthogilia A. G. Day & Moran.

Membership.Acanthogilia.


 Genus

Acanthogilia A.G. Day & Moran, Proc. Calif. Acad. Sci. 44: 111.  1986.

     Stiff, spiny shrub with dimorphic leaves, the primary alternate, woody-persistent, pinnate, with terete spinose divisions, the secondary appearing fascicled on axillary short shoots, with persistent bases and deciduous, flat, linear, herbaceous blades.  Calyx tubular, with equal spinose lobes and narrower scarious intervals that rupture in fruit, glandular.  Corolla radial to slightly zygomorphic, salverform.  Stamens subequally attached near middle of tube, superficially adnate below, subequal in length, exserted.  Pollen yellow, the grains zonocolporate, perreticulate, supraverrucate.  Seeds elongate, flat, usually winged, mucilaginous when wet.  Chromosomes: x = 9.

Type species—Gilia gloriosa Brandegee, Proc. Calif. Acad. Sci. Ser. 2, 2: 184, pl. 9. 1889.

Holotype specimen (UC101896). High resolution image available at UC-JEPS herbarium  (http://ucjeps.berkeley.edu/cgi-bin/display_smasch_img.pl?smasch_accno=UC101896).

Discussion

 

Acanthogilia gloriosa (Brandegee) A.G. Day and Moran

Gilia gloriosa Brandegee, Proc. Calif. Acad. Sci. Ser. 2, 2: 184, pl. 9. 1889.  Loeseliagloriosa (Brandegee) I. M. Johnston, Proc. Calif. Acad. Sci. Ser. 4, 12: 962. 1924.  Leptodactylon gloriosum (Brandegee) Wherry, Am. Midl. Naturalist 34: 383. 1945.  lpomopsis gloriosa (Brandegee) A.D. Grant in V.E.  Grant, Aliso 3: 357. 1956.  Acanthogilia gloriosa (Brandegee) A.G. Day and Moran, Proc. Calif. Acad. Sci. 44: 115.  1986.  TypeT. S. Brandegee s.n., Mexico, Baja California, Ubi, 8 May 1889 (holotype, UC; isotypes, CAS [DS], GH, NY, US).  According to Day and Moran (1986), this location is at the tinaja (waterhole) of Yubay, near 29°11' N, 113°59' W, ca. 650 m elevation, ca. 9 km NE of the abandoned mine of Desengaño and ca. 52 km from the Pacific coast.  Acanthogilia gloriosa has not been relocated at this site.

    Shrub 1–3 m, erect to more often decumbent and sprawling, 1–5 m broad, much branched at base, wood brittle, spiny, the young parts glandular-viscid, trichomes two-many celled, mostly less than 0.5 mm long, tipped with yellowish, globose, multicellular gland. Stems to 6 cm diameter, the bark light to dark gray, separating into small plates or strips; lower branches arching, sometimes rooting at nodes. Branching sympodial, the stems flowering terminally the first year and so not elongating further, 1–125 cm long, 1–4 mm diameter the first year, green becoming tan or gray, subterete, persistently spiny with remnant leaves; internodes mostly 5–8 mm, exceeded by leaves. Primary leaves subopposite to mostly alternate, 1–3 cm long, vascular bundles reinforced with fibers, green becoming tan and finally dark gray, and woody-spinose, rigidly divaricate and straight except terminal segment usually declined, persisting two to three (sometimes to six) years but weak after first or second year, the lowermost often simple but most pinnate with narrowly linear rachis and one to two (sometimes to three) pairs of spreading spinose lobes to 9 mm long, the base 1.5–3.0 mm wide. Secondary leaves appearing fascicled in axils, few and short to 25 in axils of leaves of flowering or older branches, secondary leaf bases whitish to tan, persistent, the blades herbaceous, mostly simple, rarely with one to two short lobes, linear-oblanceolate, spinose-tipped, flattened, 5–20 mm long, to 1 mm wide, rather sparsely glandular, deciduous throughout plant all at about one time; first leaves of new shoot with enlarged semiglobular bases to 1.5 mm wide and blades sometimes less than 1 mm long. Short shoots producing leaves for (1–)3–4(–6) years, to 8 mm long, often becoming long shoots. Inflorescence densely glandular-puberulent and glutinous, a thyrse to 2 dm long on a new, long shoot, with terminal flower opening first and with up to 30 short one-few-flowered branches below; or inflorescence reduced to short one-few-flowered shoot, though sometimes several such shoots borne on one older branch to form two-generational inflorescence. Pedicels erect to spreading, 1–6 mm long, 0.5–1.0 mm thick. Calyx 10–16 mm long, 3.0–4.5 mm wide, cylindric, tapering to rounded at base, densely glandular and glutinous abaxially, sparsely so adaxially (internal tube), tubular portion 7–9 mm long, composed of green, herbaceous strips alternating with white, hyaline regions; lobes equal, erect or slightly outcurved, 3–8 mm long, triangular-lanceolate, pungent-acuminate, cartilaginous, with many crowded veins within, scarious-margined except near apex; sinuses V-shaped, the scarious intervals much narrower than green ribs, distended at anthesis, folding inward as segments later converge, mostly rupturing in fruit. Corolla salverform, 3.0–4.5 cm long, glandular without, in bud pale yellow becoming orangish; tube stout, slightly upcurved, 2.0–3.2 cm long, 2.5–3.5 mm wide below, gradually flaring to orifice 5–7 mm wide, dull orange-red to orange-brown, becoming paler and more purplish; throat yellow; segments 2–4 cm wide, white and muriculopapillose adaxially, rose-veined abaxially, the lobes in bud convolute, in anthesis widely spreading or somewhat reflexed, in age strongly reflexed, 8–20 mm long, 4–12 mm wide, obliquely oval to strap-shaped, obtuse to slightly emarginate, sometimes apiculate, with 30–50 close-spaced parallel veins per lobe. Filaments glabrous, 15–27 mm long, subequally attached at middle of tube or slightly above; appearing superficially adnate below, the filament portion of the corolla tube flanged, the flange abutting the ovary, producing five nectar-channels between the filaments; the free portion lacking this flange, terete, subequally exserted 4–13 mm from throat; anthers oblong, sagittate, 4–5 mm long before anthesis, dehiscing as corolla begins to open. Pollen grains suboblate to spheroidal (polar diameter 55–64 mm, equatorial diameter 61–71 mm), zonocolporate, insulate. Nectary disk green, ca. 2 mm wide, shallowly cupped or fluted, the margin regularly undulate to form erect, retuse lobes opposite filament vasculature and spreading, truncate lobes opposite central corolla lobe vasculature. Ovary tricarpellate, trilocular, 4–5 mm long, ca. 1.5 mm diameter; style 20–40 mm long, slightly shorter to slightly longer than anther height (more or less uniform per individual); stigma lobes acute, 1.5–2.0 mm long, reflexed by third day. Ovules 14–24 per locule, ovoid, ca. 0.6 mm long. Capsules 7–15 mm long, 4–5 mm in diameter, cylindroidal, beaked, yellowish brown, loculicidally dehiscent, the valves often recurving and occasionally each valve splitting septicidally also. Seeds overlapping, 1–6 per locule, narrowly oblong, flat to angled, 6–7 mm long, ca. 1.5 mm wide, the golden-brown body, often with a marginal, elongated membranous wing, the body and wing swelling and mucilaginous when wet; embryo green. Chromosomes: n= 9 (Day and Moran 1986).