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Key to the Species of Dioon

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Key to the Species of Dioon It is important to remember that leafl et partially shields next distal leafl et Key to the Species considerable variation exists within spe- (incubous) or next proximal leafl et (suc- cies in this genus. While we have tried cubous) when viewed from above. of Dioon to account for the major contingencies, keeled. Vee-shaped; resembling boat keel (Jones, 2002). exceptions are regularly encountered. Tim Gregory, Jody Haynes lanceolate. Lance-shaped, much longer than For example, prickles may or may not be broad; wide base, tapered apex, widest & Jeff Chemnick present along the leafl et margins in many below center. instances, and some leaves in the same linear. Long and narrow; sides parallel or INTRODUCTION species may be fl at or slightly keeled in nearly so. The genus Dioon is particularly diffi cult cross section. To make matters worse, nitid. Smooth and shiny (Daydon Jackson, to work with taxonomically because the differences in leaf/leafl et morphology can 1965). Cf. glaucous. differences between species are subtle. sometimes be found even within a single petiole. Section of leaf axis below lowermost leafl ets, pinnacanths, or spines. The range of form within the world’s other cohort of leaves on an individual plant, phylloproximal. Leafl et margin that, in un- major cycad genera—Cycas, Ceratozamia, which, in turn, may differ slightly from twisted leafl et, is directed toward proximal Encephalartos, Macrozamia, and Zamia—is leaves in previous cohorts. Thus, there is part of leaf (Grobbelaar, 2003). See also more extensive, thus making identifi cation no substitute for a large sample size. basiscopic; Cf. acroscopic, phyllodistal. a slightly easier task. The following key It is our hope that, in spite of these pinnacanth. Sharply-pointed structure inter- is designed to enable workers to identify daunting subtleties and seeming ambigui- mediate between leafl et and spine, usually all known species within the genus Dioon, ties, this key will aid in the identifi ca- green. as well as several distinct ‘types’ that are tion of species within this most ‘diffi cult’ pp-angle. Pinna-pinna angle (pinna = leafl et) measures angle, on adaxial side of leaf, be- currently under investigation. cycad genus. tween planes in which long axes of leafl ets When working with keys, it is impor- occur (Grobbelaar, 2002). tant to remember that they are contriv- BACKGROUND pr-angle. Pinna-rachis angle (pinna = leafl et) ances to facilitate identifi cation, not The following key was originally pre- measures angle between imaginary line phylogenetic or taxonomic schemes. As sented as an illustrated poster at the 7th drawn from middle of leafl et attachment such, the individual pairings as well as the International Conference on Cycad Biology through distal end of leafl et and rachis in overall arrangement of species may not held in January 2005 in Xalapa, Veracruz, distal direction (Grobbelaar, 2002). necessarily imply relationship. Perhaps Mexico. The poster has been reproduced prickle. Small, sharp protuberance of epider- mal origin, usually green and somewhat not coincidentally, the species within (with permission) by the Cycad Society irregularly distributed. the “Spinulosum” Clade differ signifi - (TCS) and is available for sale on the TCS proximal. Nearest to point of attachment; cantly from the species in the “Edule” website (www.cycad.org). basal. Cf. distal. Clade, and these clades, by defi nition, do pubescent. Densely covered with fi ne, short represent purported phylogenetic group- GLOSSARY hairs. Cf. glabrous. ings (see Moretti et al., 1993; De Luca The following technical terms are used pungent. Terminating in a stiff, sharp point. et al., 1995; Bogler & Francisco-Ortega, in the key below. Except where noted, the rachis. Section of leaf axis where leafl ets are 2004). Similarly, the traits that separate defi nitions were taken from Osborne and attached. recurved. Bent or turning abaxially. the three species within the “Spinulosum” Walters (2004). ‘Cf’ indicates a related or revolute. Margin rolled abaxially. Clade create a hierarchical structure for opposite term. serrated. Margin of saw-toothed, sharply- the respective species that coincides with tipped protrusions pointing apically. Cf. published phylogenetic relationships (sum- abaxial. Side of organ facing away from central entire. marized in Haynes & Bonta, 2007). axis, e.g. lower side of leaf or leafl et. Cf. tomentose. Densely woolly in a fi nely matted The glossary from the proceedings adaxial. fashion. Cf. glabrous. of the Cycad Classifi cation Concepts acuminate. Tapering to protracted point, with workshop—held in Miami, FL, USA, in April sides somewhat concave. REFERENCES 2002—was used as the standard for defi ni- adaxial. Side of organ facing towards central Bogler, D.J. & J. Francisco-Ortega. 2004. Molec- tions of most cycad morphological terms axis, e.g. upper side of leaf or leafl et. Cf. ular systematic studies in cycads: evidence from trnL intron and ITS2 rDNA sequences. (Osborne & Walters, 2004). Grobbelaar’s abaxial. basiscopic. Leafl et margin closest to the base Botanical Review 70:260-273. (2003) instructive angles of leafl et inser- of the leaf. See also phylloproximal; Cf. Daydon Jackson, B. 1965. A Glossary of Botanic tion, not included in the aforementioned acroscopic, phyllodistal. Terms, with their Derivation and Accent, 4th glossary, are also referred to herein. declinate. Gently curving abaxially (Daydon Ed. Hafner Publishing Co., New York. Workers may need to reference both Jackson, 1965). De Luca, P., A. Moretti, G. Siniscalco, P. Capu- decurrent. Leafl ets in which attachment ex- works to fully appreciate the key. to, S. Cozzolino, L. Gaudio, D.W. Stevenson, tends downward along rachis. It was our intention to provide a key E.T. Wurtel & R. Osborne. 1995. Molecular defl exed. Bent abruptly abaxially (Norstog & that could be used in the absence of systematics of cycads. Pp. 131-137, In: Nicholls, 1997). Cf. refl exed. reproductive structures; not only are the P. Vorster (ed.), Proceedings of the Third distal. Furthest from point of attachment; api- International Conference on Cycad Biology. strobili of Dioon species much less dis- cal. Cf. proximal. Cycad Society of South Africa, Stellenbosch. tinct compared to the other large cycad elliptic. Two-dimensional structure widest near genera, but they are also often lacking in middle but narrowed toward each rounded Moretti, A., P. Caputo, S. Cozzolino, L. Gaudio, habitat. Because identifi cation of Dioon end. G. Siniscalco-Gigliano & D.W. Stevenson. seedlings is also quite diffi cult, eophylls entire. Continuous margin; not toothed or 1993. A phylogenetic analysis of Dioon are mentioned only when diagnostic. lobed. Cf. serrated. (Zamiaceae). American Journal of Botany 80:204-214. Thus, the vegetative key below can be eophyll. First leaf produced by seedling. applied almost entirely using adult leaf falcate. Sickle shaped. Grobbelaar, N. 2002. Cycads, with Special glabrous. Smooth surface, without hair of any Reference to the Southern African Spe- material. On occasion, reference is made kind. Cf. pubescent, tomentose. cies. Published by the author, Pretoria, to newly emerging leaves, but each ap- glaucous. Surface covered by bluish-gray, waxy plicable couplet also contains character or powdery substance. Cf. nitidus. South Africa. separations based on ‘hardened’ leaves. imbricate. Leafl et arrangement in which one IA. Adult leaflets wide (usually 15 mm); eophyll leaflets wide, with evenly serrated margins ................................. “Spinulosum” Clade 2A. Median leaflets of adult leaves elliptic-acuminate, pungent; eophylls resembling adult leaves, with proximal leaflets reducing to pinnacanths and with petioles short (< 20% of leaf length) or absent; emerging leaves densely, persistently tomentose......................................................................................................................... D. mejiae 2B. Median leaflets of adult leaves lanceolate, not pungent; eophylls with proximal leaflets nearly equal in size to median and distal leaflets and with long, unarmed petioles ( 50% of leaf length); emerging leaves glabrous to pubescent 3A. Adult leaves to 2 m long, usually lacking petioles; proximal leaflets lanceolate, reducing to pinnacanths; median leaflets symmetrically lanceolate, to 21 cm long and 2 cm wide; distal and median leaflet margins usually serrated; leaflet attachments only slightly decurrent; emerging leaves glabrous ........................................................ D. spinulosum 3B. Adult leaves to 1.4 m long, with unarmed petioles 10-15 cm long; proximal leaflets linear-lanceolate to linear, reducing but not to pinnacanths; median leaflets asymmetrically lanceolate, to 19 cm long and 2.5 cm wide; distal and median leaflet margins usually entire; leaflet attachments strongly decurrent; emerging leaves pubescent ................................................................................................................................... D. rzedowskii IB. Adult leaflets narrow (< 15 mm); eophyll leaflets narrow, only distally serrated ....................................................... “Edule” Clade 4A. Leaflets flat, not declinate or deflexed, margins sometimes strongly revolute 5A. Leaves flat or slightly keeled (pp-angle > 165q) 6A. Leaflets usually entire (> 99% of individuals) 7A. Leaflets < 6 mm wide, flat ..........................................................................................D. angustifolium 7B. Leaflets > 7 mm wide, flat to strongly revolute ..........................................................................
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