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Systematic Anatomy of Euphorbiaceae Subfamily Oldfieldioideae I

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Systematic Anatomy of Euphorbiaceae Subfamily Oldfieldioideae I University of Richmond UR Scholarship Repository Biology Faculty Publications Biology 1994 Systematic Anatomy of Euphorbiaceae Subfamily Oldfieldioideae I. Overview W. John Hayden University of Richmond, [email protected] Follow this and additional works at: http://scholarship.richmond.edu/biology-faculty-publications Part of the Botany Commons, Other Plant Sciences Commons, and the Plant Biology Commons Recommended Citation Hayden, W. John. "Systematic Anatomy of Euphorbiaceae Subfamily Oldfieldioideae I. Overview." Annals of the Missouri Botanical Garden 81, no. 2 (1994): 180-202. This Article is brought to you for free and open access by the Biology at UR Scholarship Repository. It has been accepted for inclusion in Biology Faculty Publications by an authorized administrator of UR Scholarship Repository. For more information, please contact [email protected]. SYSTEMATIC ANATOMY OF W.John Hayden2 EUPHORBIACEAE SUBFAMILY OLDFIELDIOIDEAE. I. OVERVIEW' ABSTRACT The biovulate subfamily Oldfieldioideaeof Euphorbiaceae, characterized by spiny pollen, is an otherwise apparently diverse assemblage of mostly Southern Hemisphere trees and shrubs that traditionally have been allied with genera of Phyllanthoideae and Porantheroideae sensu Pax and Hoffmann. Although fairly diverse anatomically, the following structures characterize the subfamily with only a few exceptions: pinnate brochidodromousvenation with generally randomly organized tertiary and higher order venation; foliar and petiolar glands absent; unicellular or unbranched uniseriate trichomes; latex absent; mucilaginous epidermis or hypodermis; brachyparacytic stomata; vessel elements with simple perforation plates and alternate, often very small, intervascular pits; thick-walled nonseptate imperforate tracheary elements; numerous narrowheterocellular rays; and abundantaxial xylem parenchyma in diffuse to somewhat banded patterns and often bearing prismatic crystals. Anatomically, the shrubby Australian ericoid genera form a well-definedgroup with obvious affinities to the more arborescent Australasian genera, which show clear relationships to each other; the African and neotropical genera bearing compound leaves form another distinct group; the remaining genera are somewhat more isolated and seem to represent, in various cases, elements that are primitive within the subfamily or elements derived from the group bearing compound leaves. Presence of theoid teeth and palmately compound leaves in Oldfieldioideaeare features consistent with Dilleniid origin for Euphorbiaceae. As a taxonomic entity the euphorbiaceous sub- dendron was shown to be a member of Oldfieldioi- family Oldfieldioideae K6hler & Webster dates deae (Hayden, 1977; see also Hayden et al., 1984, conceptually from the palynological studies of Punt and Hakki, 1985), an anatomical survey of the (1962) and K6hler (1965) who noted the spiny entire subfamily was initiated (Hayden, 1980) to pollen that characterizes the group; the assemblage assess relationships from data independent of re- recognized by pollen structure was subsequently productive (including pollen) morphology. Based in formalized nomenclaturally as a new subfamily part on these anatomical studies, classification of (Webster, 1967). In essence, pollen characters Oldfieldioideaehas been modified from Webster's circumscribed a group of genera that previously (1975) original proposal; two broadly similar clas- had been assigned to the biovulate subfamilies Phyl- sifications, one by G. L. Webster (the preceding lanthoideae and Porantheroideae in the system of issue) and another by G. A. Levin and M. G. Pax & Hoffmann (1931). Webster's first (1975) Simpson (this issue), are proposed elsewhere. classification of oldfieldioid genera contained sev- eral novel taxonomic associations at variance with HISTORY earlier classifications, notably those of Pax & Hoff- mann (1931) and Hutchinson (1969). Classifications of Euphorbiaceae from the first Interest in the anatomy of Oldfieldioideae half of the 19th century (e.g., Jussieu, 1824; End- stemmed from the need for comparative data to licher, 1836-1840) offer little insight into rela- aid placement of the problematic genus Picroden- tionships among oldfieldioid genera. At this for- dron Planchon (Hayden, 1977). Despite a number mative time in the definition of Euphorbiaceae of publications dealing with the anatomy of old- (Webster, 1987) the few oldfieldioidgenera known fieldioid genera (Appendix 2), detailed information were often scattered widely in the family; more- for most of the subfamily is lacking. Once Picro- over, some were referred to other families. Genera ' This study is based largely on a Ph.D. dissertation submitted to the University of Maryland, College Park. I thank W. L. Stern for assistance and guidance; H. K. Airy Shaw, L. J. Hickey, G. A. Levin, A. M. W. Mennega, and G. L. Webster were particularly helpful in providing specimens and/or comments; numerous others also provided specimens. 2 Department of Biology, University of Richmond, Richmond, Virginia 23173, U.S.A. ANN. MISSOURIBOT. GARD. 81: 180-202. 1994. Volume 81, Number 2 Hayden 181 1994 EuphorbiaceaeSubfamily Oldfieldioideae of Pseudanthinae (or Caletieae in the traditional/ included, using current nomenclature, Dissiliaria, narrow sense) were the first to be classified to- Longetia, Austrobuxus, Hyaenanche, Mischo- gether, undoubtedly because of their shared ericoid don, Oldfieldia, Piranhea Baillon, and only one xeromorphic habit and common provenance in nonoldfieldioidgenus, Bischofia Blume, within gen- Australia. Baillon (1858) grouped Micrantheum era 46-54. Although these plants constituted three Desf. with Pseudanthus Sieber ex Sprengel, to discrete groups in his conspectus, Bentham (1878) which Agardh (1858) added Stachystemon Plan- expressed some doubt about their relatedness and chon, establishing the composition of Caletieae that he chose to associate Neoroepera and Petalostig- was followed in all subsequent studies until the ma with other phyllanthoid genera. Pax (1890) addition, first, of Neoroepera Muell. Arg. by Koh- treated oldfieldioidgenera much as had Bentham; ler (1965) and, now, the inclusion of all Austral- Tetracoccus Engelm. ex C. Parry, however, was asian oldfieldioidsas proposed by Levin & Simpson included, and Pax & Hoffmann (1931) added Peta- (1994, this issue) and Webster (1994). During the lostigma, Androstachys Prain, and Aristogeiton- intervening years, however, the use of cotyledon ia Prain to various subtribes consisting largely of width as the primary criterion for subdivision of oldfieldioid genera. Perhaps the first clear indica- the family (Mueller, 1866; Pax, 1890; Pax & tion of the relatedness of oldfieldioid genera can Hoffmann, 1931) relegated the genera of Pseu- be found in Pax's (1925) essay on Euphorbiaceae, danthinae (as Caletieae) to the Stenolobeae, a small which includes the phylogenetic tree reproduced group of Australian xerophytes with narrow cot- in Figure 1. Although much of the detail in this yledons. Their somewhat isolated position in Sten- phylogenetic tree may be challenged seriously in olobeae minimizedassociation of Pseudanthinaewith the light of present knowledge, it does show old- other oldfieldioid genera. fieldioid genera (as usual, minus Pseudanthinae) Other oldfieldioidgenera similarly suffered early comprising a clade distinct from other biovulate taxonomic assignments that delayed consideration Euphorbiaceae. of relationshipswith the rest of the subfamily. Bail- In the disposition of genera, Hurusawa's (1954) lon (1858) submerged Podocalyx Klotzsch within classification of Euphorbiaceae essentially follows the phyllanthoidgenus Richeria Vahl, a disposition that of Pax & Hoffmann (1931) and thus contains followed well into this century. Oldfieldia Benth. no new insights on classification of oldfieldioidgen- & Hook. f. was temporarily considered sapinda- era. In light of Pax's (1925) phylogeny and because ceous (Mueller, 1866; Baillon, 1878). Worse, the of its disregard for the palynological data available relationships of Picrodendron were long obscured at the time (Punt, 1962; K6hler, 1965), Hutch- by a series of misassignments at both the generic inson's (1969) classification was a retrograde de- and familial levels (Hayden et al., 1984). velopment; oldfieldioidgenera were widely distrib- Aside from Caletieae sensu stricto, the genera uted in five of 12 biovulate tribes, with three tribes of Oldfieldioideae were scattered widely in Muel- mixing oldfieldioid and phyllanthoid elements. ler's (1866) treatment of the family for the Prod- Hutchinson was the first, however, to associate romus; Mischodon Thwaites was included with Paradrypetes Kuhlm. with other oldfieldioid gen- uniovulate genera, following an earlier error of era. Baillon (1858), Oldfieldia was still banished to Airy Shaw (1965) entertained serious doubts Sapindaceae, and the remaining five oldfieldioid about inclusion of several oldfieldioid genera in genera in his treatment were each assigned to dif- Euphorbiaceae. Accordingly, he proposed Andro- ferent subtribes of Phyllantheae. Greater cohe- stachydaceae, of uncertain relationships,to accom- siveness is apparent in a later publication of Baillon modate Androstachys; he recognized Picroden- (1878). For example: Choriceras Baillon was in- draceae as distinct from but allied to Euphorbiaceae; cluded within Caletieae; Austrobuxus Miq. (as Bur- and he viewed Aristogeitonia, Calaenodendron aeavia Baill.), Longetia Baillon ex Muell. Arg.,
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