Cryptantha cycloptera (Greene) Greene forma truncata M.E. Mabry & Rebman, 2016

Mabry, Makenzie E., Dowdy, Regina A., Simpson, Lee M., Rebman, Jon P. & Simpson, Michael G., 2016, Taxonomy of the winged popcorn flower: Cryptantha pterocarya (Boraginaceae), Phytotaxa 253 (2), pp. 97-130 : 117-118

Cryptantha cycloptera (Greene) Greene forma truncata M.E. Mabry & Rebman , forma nov.

Type:— USA: California, San Bernardino County, Whipple Mountains , west slope just below saddle at north end of Savahia Peak, steep slopes of dark volcanic boulders and outcrops, annual, flowering, 762 m, 34.276633˚ N 114.5328˚ W, 4 May 2003, Sarah J. De Groot 2834, with J. Mark Porter, Naomi Fraga (holotype RSA 721252!, isotypes CAS 1123044!, ARIZ 393611)

Diagnosis: — Cryptantha cycloptera forma truncata is similar to the typical form of Cryptantha cycloptera ( forma cycloptera ) in having only appressed trichomes along the upper inflorescence stem and in having all four nutlets winged, with the wing extending along the base of the nutlet body. It differs in having heteromorphic nutlets, with one nutlet having the wing reduced on the sides and base.

Etymology: —The epithet “ truncata ” refers to the reduced, somewhat truncate basal wing of the odd nutlet of this form, in contrast to the rounded, well-developed basal wing of the typical form of Cryptantha cycloptera .

Paratypes: —See Appendix 2.

As with all strictly taxonomic studies, a future detailed molecular phylogenetic analysis will be extremely important in elucidating evolutionary relationships within the complex. If forma pseudocycloptera and forma truncata each are found to constitute a monophyletic clade, then their recognition at a higher taxonomic rank may be warranted. A similar study of C. pterocarya var. purpusii and var. stenoloba may test their monophyly and possible introgression, and may help elucidate their interrelationships to var. pterocarya , the type for the species. Other interesting questions are the molecular mechanism for development of heteromorphism and the other traits distinguishing the taxa and forms of this complex. Finally, the interrelationships of C. oxygona and C. utahensis to C. pterocarya and the evolutionary direction and possible adaptive significance of nutlet number and wing development will be an interesting avenue of future research.

The nomenclature we accept from our study of this complex of taxa is summarized in Table 3. A revised key is presented here of Johnston’s (1925) series Pterocaryae (modified in part from Kelley et al. 2012), clarifying the identity of the entities that we propose in this paper:

1. Nutlets 1(2), margin a ± flat narrow rim to sharp-angled, tubercles often spinulose especially toward apex ............... C. utahensis

- Nutlets generally 4(3), margin occasionally sharp-angled, generally narrow- to wide-winged along entirety, nutlet tubercles not or slightly spinulose .............................................................................................................................................................................. 2

2. Flower limb typically (3) 4–8 mm in diameter ................................................................................................................ C. oxygona

- Flower limb less than 4 mm in diameter .......................................................................................................................................... 3

3. At least some nutlets with marginal wing encircling entire nutlet body, including base; nutlets homomorphic with all 4 winged ( forma cycloptera ), or heteromorphic with odd nutlet having reduced wing at sides and base ( forma truncata ) ....... C. cycloptera

- All nutlets lacking a marginal wing at base of nutlet body, nutlets homomorphic or heteromorphic .............................................. 4

4. At least some nutlets with narrow, lobed or toothed wings <0.6 mm wide, or with narrow, linear, marginal rim; nutlet body 1.2–2.2 mm long .................................................................................................................................................. C. pterocarya var. purpusii

- At least some nutlets with prominent, apically lobed or toothed wings> 0.6 mm wide; nutlet body 1.8–3.2 mm long .................. 5

5. Calyx body in fruit ovoid, 3–4(6) mm long, sepals lance-ovate to lanceolate (length:width ratio 2–4); nutlets heteromorphic with one nutlet unwinged ( forma pterocarya ) or homomorphic with all four nutlets winged ( forma pseudocycloptera ), nutlet body generally 1.8–2.8 mm long, wing width of consimilar nutlets 0.6 mm and greater ................................. C. pterocarya var. pterocarya

- Calyx body in fruit lance-ovoid, 5–8 mm long, sepals lanceolate (length:width ratio 3–5); nutlets heteromorphic, the odd nutlet unwinged, nutlet body generally 2.8–3.2 mm long .............................................................................. C. pterocarya var. stenoloba

Cryptantha cycloptera (Greene) Greene (1887: 120) .

Krynitzkia cycloptera Greene (1885: 207) (Basionym). Cryptantha pterocarya (Torrey) Greene var. cycloptera (Greene) J. F. Macbride (1916: 44) . Type:— USA. Arizona: Pima County, hills near Tucson, 15 April 1884, Cyrus Guernsey Pringle s.n. (holotype CAS 202, barcode 0006884).

Cryptantha cycloptera (Greene) Greene forma truncata M.E. Mabry & Rebman , forma nov.

Type:— USA. California : San Bernardino County, Whipple Mountains, west slope just below saddle at north end of Savahia Peak, 4 May 2003, Sarah J. De Groot 2834 (holotype RSA 721252, isotypes: CAS 1123044, ARIZ 393611).

Cryptantha pterocarya (Torrey) Greene (1887: 120) .

Eritrichium pterocaryum Torrey (1859: 142) (Basionym). Krynitzkia pterocarya (Torrey) A. Gray (1885: 276). Type:— USA: Washington, Wallah Wallah, s.d., 1859, Wilkes Expedition 1047 (holotype NY 335248).

= Eritrichium pterocaryum Torrey var. pectinatum A. Gray (1874: 61). Krynitzkia pterocarya (Torrey) A. Gray var. pectinata (A. Gray) A. Gray (1885: 276). Cryptantha pterocarya (Torrey) Greene var. pectinata (A. Gray) Brand (1931: 55). Type:— USA. Utah, 1874, C. C. Parry 168-49 (isotypes ISC-v-0000202, NY00335256).

Cryptantha pterocarya (Torrey) Greene var. pterocarya forma pseudocycloptera M.E. Mabry & M.G. Simpson , forma nov.

Type:— USA. California : Imperial County, In-Ko-Pah Mountains, Jacumba Natural Area, along road ascending to Valley of the Moon, 13 April 2001, Rebman 7221 (holotype SD 155821, isotypes RSA 700474, UC 1787754, UCR 155804).

Cryptantha pterocarya (Torrey) Greene var. purpusii Jepson (1925: 849) .

Type:— USA. California : Inyo County, Argus Mountains, June 1897, C. A. Purpus 5433 (holotype JEPS 2548).

Cryptantha pterocarya (Torrey) Greene var. stenoloba I. M. Johnston (1939: 391) .

Type:— USA. Nevada: Clark County, desert 15 mi E. of Glendale, 19 May 1933, B. Maguire & H. L. Blood 4466 (holotype GH 00096072).