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<I>Alectra Sessiliflora</I> Systematic Botany (2011), 36(1): pp. 141–152 © Copyright 2011 by the American Society of Plant Taxonomists DOI 10.1600/036364411X553234 Taxonomic Revision of the Alectra sessiliflora Complex (Orobanchaceae) Jeffery J. Morawetz 1 , 2 and Andrea D. Wolfe Department of Evolution, Ecology and Organismal Biology, The Ohio State University, 300 Aronoff Laboratory, 318 W.12 th Avenue, Columbus, Ohio 43210 U. S. A. 1 Current address: Rancho Santa Ana Botanic Garden, 1500 North College Avenue, Claremont, California 91711 U. S. A. 2 Author for correspondence ( [email protected] ) Communicating Editor: Daniel Potter Abstract— Alectra sessiliflora (Orobanchaceae) is the most widespread species within the genus, occurring throughout sub-Saharan Africa, and into India, China and the Philippines. Three varieties are currently recognized (A. sessiliflora var. monticola, A. sessiliflora var. senegalensis , and A. sessiliflora var. sessiliflora ) and are distinguished by geographic range, calyx pubescence and stamen filament pubescence. Due to the overlapping nature of the characters used to distinguish among these varieties, accurate assignment of a specimen to a single variety is nearly impossible. We undertook a phenetic study of morphological characters to assess the validity of these varieties. Principal coordinate analysis and the unweighted pair-group method using arithmetic averages were used to explore whether specimens would cluster into the currently recognized varieties. Our analyses revealed no clustering based exclusively on geographic distribution. A small cluster of seven specimens was seen in the principal coordinate analysis using Gower’s coefficient of similarity as input values, but this cluster was not diagnosable by unique characters. Based on these results we recommend that Alectra sessiliflora be recognized without infraspecific taxa. Finally, additional names are included as synonyms under Alectra sessiliflora based on extensive study of field-collected and herbarium specimens. Keywords— hemiparasite , multivariate analysis , Orobanchaceae , principal coordinate analysis , species complex , UPGMA. Alectra Thunb. is a genus of 12 primarily hemiparasitic spe- Melch. Additionally, A. sessiliflora var. sessiliflora has been cies of herbaceous plants distributed mainly in sub-Saharan divided into two forms: A. sessiliflora var. sessiliflora forma ses- Africa (Morawetz, unpubl. data). Species of Alectra are char- siliflora and A. sessiliflora var. sessiliflora forma barbata (Hiern) acterized by sessile flowers with a yellow, marcescent corolla, Hilliard & B. L. Burtt. These varieties all overlap in their distri- a recurved or “horseshoe-shaped” style, and clavate stigma butions within Africa, and it is only A. sessiliflora var. monticola ( A. fruticosa Eb. Fisch. is the exception to habit, inflorescence that extends outside of Africa. The three varieties historically and corolla color typical of the genus; see Morawetz and have been distinguished in keys based upon calyx pubes- Wolfe 2009 ). Species of Alectra occur in damp to inundated cence (margins, nerves, or both), and stamen filament pubes- grasslands where they generally parasitize grasses and sedges cence (all glabrous or longer pair pubescent), although these ( A. fruticosa is again an exception, occurring in high elevation characters are not diagnostic. Relative leaf size and shape also montane rainforests in Madagascar; A. schoenfelderi Dinter & often have been considered diagnostic, particularly for A. ses- Melch. is also exceptional, occurring in arid areas of southeast- siliflora var. monticola ; plants of this variety possess the small- ern Angola, northeastern Namibia, and adjacent southwestern est, most coarsely toothed leaves (see Melchior 1941 ; Hepper Zambia). The characters most often used to separate species 1960 ; Philcox 1990 ). The calyx pubescence and stamen fila- are presence or absence of pubescence on the stamen filaments ment pubescence characters that are most commonly used (which are didynamous, typical for the family), the shape of to distinguish among the varieties are summarized concisely Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Sat, 02 Oct 2021 05:50:54 the anther theca base, and the shape of the leaves (see Hepper by Philcox (1990) : plants of A. sessiliflora var. sessiliflora have 1963 ; Philcox 1990 ; Fischer 2006 ; Ghazanfar et al. 2008 ). calyces that are ciliate on the “upper margin” and rarely on Alectra sessiliflora (Vahl) Kuntze is the most widespread spe- nerves, filaments that are glabrous or the longer pair bearded; cies with a range extending from the entirety of sub-Saharan plants of A. sessiliflora var. senegalensis have calyces that are Africa into India, China, and the Philippines; the plant is a ciliate on nerves and margins, the longer pair of filaments weed of grasslands throughout its range ( Morawetz 2007 ). always bearded; and plants of A. sessiliflora var. monticola Given this wide distribution and habit, it is unsurprising have calyces ciliate on nerves and margins, the longer pair of that many species have been described from throughout the filaments always bearded (the latter two varieties are iden- range, with several having already been placed into synon- tical in description). Hilliard and Burtt (1986) further distin- ymy ( Melchior 1941 ; Hepper 1960 ; Wood 1997 ). Through the guished among the character states within A . sessiliflora var. examination of taxonomic literature and loaned specimens for sessiliflora by defining forma sessiliflora as having all stamen the purpose of producing a monograph of Alectra (Morawetz, filaments glabrous, and defining forma barbata has having the unpubl. data), several additional species have been iden- two longer filaments bearded (summarized in Table 1 ). These tified as conspecific with A. sessiliflora : A. aberdarica Chiov., authors stated that they “do not feel that the trivial variants A. asperrima Benth ., A. congolensis Troupin, A. hundtii Melch., commonly ranked as formae have any place in a nomencla- A. ibityensis Eb. Fisch., A. ledermannii Engl., A. moeroensis Engl., tive classification,” and also noted that they observed mixed A. schliebenii Melch., and A. trinervis Hemsl. Considering the populations where both states occur. The purported distribu- lack of a current comprehensive understanding of A. sessili- tion of traits among putative varieties makes it impossible to flora , a taxonomic revision of this species is necessary. identify any of these infraspecific taxa with confidence. Due Recognition of the number of infraspecific taxa has also var- to the lack of reliable resolution provided by these characters, ied over time and specimens often are not annotated below combined with personal observations of continuous variation species level. Currently three varieties of A. sessiliflora are rec- within individual plants (for leaf size and shape, calyx pubes- ognized: A. sessiliflora var. sessiliflora , A. sessiliflora var. senega- cence), within populations and among the alleged varieties, a lensis (Benth.) Hepper, and A. sessiliflora var. monticola (Engl.) study of this species complex is necessary. 141 142 SYSTEMATIC BOTANY [Volume 36 Table 1. Summary of calyx and filament pubescence character states ters based on the characters scored. Given the unclear boundaries of these for each variety and form of Alectra sessiliflora . X = presence, - = absence, infraspecific taxa, it is rarely possible to unambiguously assign specimens and +/− = sometimes present/sometimes absent. to a single taxon. The characters used were chosen based on differences among varieties noted in previous treatments ( Melchior 1941 ; Hepper Calyx ciliate Filaments 1960 ; Hilliard and Burtt 1986 ; Philcox 1990 ) and personal observations by JJM. Several characters were scored in pairs, referring to the condition at Margins Nerves Glabrous Two longer bearded the distal and proximal extremes of the plant. Particularly for leaf size, var. sessiliflora forma sessiliflora X +/− X - shape, and petiole length, differences between apex and base have been forma barbata X +/− - X proposed to be important for distinguishing among varieties (e.g., A . ses- var. senegalensis XX - X siliflora var. sessiliflora is listed in Philcox (1990) as never having the upper var. monticola X X - X leaves petiolate). The distal leaves measured were always from the first set of leaves below the inflorescence on the main axis, if branched; the lowest leaves measured were from the lowest pair on the stem (specimens were generally not included in the study if the lowest pair of leaves were Here we address the species limits of Alectra sessiliflora , and not intact and thus unavailable for measurement). Measurements were whether the currently recognized infraspecific taxa are taxo- recorded from one leaf of the pair, favoring the least damaged or folded nomically meaningful. leaf; in cases where both leaves in the pair were in the same condition, one leaf was chosen at random for measurement. Length was measured from the base of the blade to its apex, and width was measured at the widest Materials and Methods point of the blade. Where margin was coded “entire,” the number of teeth per leaf (or teeth per bract) was coded 0. In the case of length/width ratios More than 1,300 specimens of this species complex were obtained on and for shape, the quotient was calculated (e.g. length divided by width) loan for study by JJM, and field collections
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