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A Taxonomic Revision of Commicarpus (Nyctaginaceae) in Southern Africa

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A Taxonomic Revision of Commicarpus (Nyctaginaceae) in Southern Africa South African Journal of Botany 84 (2013) 44–64 Contents lists available at SciVerse ScienceDirect South African Journal of Botany journal homepage: www.elsevier.com/locate/sajb A taxonomic revision of Commicarpus (Nyctaginaceae) in southern Africa M. Struwig ⁎, S.J. Siebert A.P. Goossens Herbarium, Unit for Environmental Sciences and Management, North-West University, Private Bag X6001, Potchefstroom 2520, South Africa article info abstract Article history: A taxonomic revision of the genus Commicarpus in southern African is presented and includes a key to the Received 19 July 2012 species, complete nomenclature and a description of all infrageneric taxa. The geographical distribution, Received in revised form 30 August 2012 notes on the ecology and traditional uses of the species are given. Eight species of Commicarpus with five in- Accepted 4 September 2012 fraspecific taxa are recognized in southern Africa and a new variety, C. squarrosus (Heimerl) Standl. var. Available online 8 November 2012 fruticosus (Pohn.) Struwig is proposed. Commicarpus species can be distinguished from one another by vari- fl Edited by JS Boatwright ation in the shape and indumentum of the lower coriaceous part of the ower and the anthocarp. Soil anal- yses confirmed the members of the genus to be calciophiles, with some species showing a specific preference Keywords: for soils rich in heavy metals. Anthocarp © 2012 SAAB. Published by Elsevier B.V. All rights reserved. Commicarpus Heavy metals Morphology Nyctaginaceae Soil chemistry Southern Africa Taxonomy 1. Introduction as a separate genus (Standley, 1931). Heimerl (1934), however, recog- nized Commicarpus as a separate genus. Fosberg (1978) reduced Commicarpus Standl., a genus of about 30–35 species, is distributed Commicarpus to a subgenus of Boerhavia, but this was not validly throughout the tropical and subtropical regions of the world, especially published (Harriman, 1999). Recent molecular studies (Douglas and in Africa and western Asia (Douglas and Spellenberg, 2010). With 12 Manos, 2007) have provided further evidence that Boerhavia and species, northeastern Africa and southern Arabia are the center of diver- Commicarpus are indeed two separate, monophyletic genera. sity for the genus (Thulin, 1990). Members of Commicarpus have a In southern Africa (Botswana, Lesotho, Namibia, South Africa and preference for arid environments, exemplified by their clustering in Swaziland), eight species of Commicarpus are recognized. These are Somalia, Ethiopia and adjacent parts of southern Arabia. In southern widely distributed in the region with Namibia being the main center Africa a secondary center of diversity, eight species, are confined to of diversity (Germishuizen and Meyer, 2003). The most recent revision the arid parts of Botswana, Namibia and South Africa (Meikle, 1978). of Commicarpus in southern Africa is that of Stannard (1988) for Flora Members of Commicarpus are morphologically similar to those of Zambesiaca, although he only treated members of five species, namely Boerhavia and the genus was originally treated as a section of Boerhavia C. chinensis (L.) Heimerl subsp. natalensis Meikle, C. helenae (Roem. (Boerhavia sect. Adenophorae Heimerl; Heimerl, 1889). This classifica- and Schult.) Meikle var. helenae, C. pentandrus (Burch.) Heimerl, tion was followed until 1909 when Standley separated the genus C. pilosus (Heimerl) Meikle and C. plumbagineus (Cav.) Standl. var. Commicarpus from Boerhavia due to differences in habit (scrambling plumbagineus. Commicarpus helenae var. helenae, C. pentandrus and or climbing in Commicarpus;diffuseinBoerhavia), structure of the C. plumbagineus var. plumbagineus were also treated in the Flora of fruit (10-ribbed with large viscid and mucilaginous glands in Tropical East Africa (Whitehouse, 1996)andtheFlora of Somalia Commicarpus;3–5-winged or 5-ribbed with multicellular trichomes (Thulin, 1993). Three currently-recognized species have not featured presentorabsentinBoerhavia), and the shape of the perianth in any earlier treatments, and this has resulted in misidentifications in (funnel-shaped in Commicarpus; bell-shaped in Boerhavia). This distinc- the Flora of Southern Africa region (Struwig et al., 2011). tion was maintained until 1931 when Standley included Commicarpus This contribution is a first attempt at a comprehensive taxonomic in Boerhavia “to aid in reducing … the increasing number of genera”, revision for the genus in southern Africa, and includes a key to the though he stated that he still believed in the validity of Commicarpus species, complete nomenclature, and a formal description of all the infrageneric taxa. Taxon accounts are supplemented with geographi- ⁎ Corresponding author. Tel.: +27 18 2992505; fax: +27 18 2992370. cal distribution records, notes on the ecology, and known traditional E-mail address: [email protected] (M. Struwig). uses. Red list assessments are also included. 0254-6299/$ – see front matter © 2012 SAAB. Published by Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.sajb.2012.09.009 M. Struwig, S.J. Siebert / South African Journal of Botany 84 (2013) 44–64 45 2. Materials and methods 3. Results and discussion Plant material was collected during field work in Namibia and South 3.1. Vegetative morphology Africa. Specimens from the following southern African herbaria were studied:BLFU,BOL,GRA,J,KMG,KSAN,NBG,NH,NMB,NU,PRE,PRU, Commicarpus species are all perennial, but two distinct growth PUC, SAM, UCBG, UNIN, WIND and ZULU (acronyms according to forms are discernable. Commicarpus fallacissimus, C. pilosus and Holmgren et al., 1990). C. squarrosus are semi-woody, sub-shrubs of 0.6–1.0 m high (Fig. 1a), Micrographs of the leaves were taken with a Canon 450D fitted while C. chinensis subsp. natalensis, C. decipiens, C. helenae var. helenae, with a 100 mm Sigma macro lens. The length and breadth of the C. pentandrus and C. plumbagineus var. plumbagineus are herbaceous leaves as well as the length of the petiole were measured. Fresh floral forbs with procumbent, decumbent or scrambling stems, or sometimes material was collected in situ in 4% paraformaldehyde and flowers upright (Fig. 1b). Younger plants of C. fallacissimus and C. pilosus are, taken from herbarium specimens were rehydrated for 10 min in however, often herbaceous forbs, but tend to develop a semi-woody, boiling water. Three flowers per species (one each from Namibia, suffrutescent growth form with age. Botswana and South Africa, where available) were dissected and the The shape of the leaves is variable (Fig. 2)andthreegroupsare length of the peduncle, pedicel, upper and lower portions of the flow- evident. The leaves of C. chinensis subsp. natalensis, C. helenae var. er, stamens and ovary were measured. Mean length and width of helenae, C. pentandrus and C. plumbagineus var. plumbagineus are trian- anthocarps were determined by measuring ten herbarium specimens gular (cordate, ovate and deltoid), those of C. pilosus and C. squarrosus per species and three anthocarps per specimen (n=30). Micrographs are roundish (orbicular and ovate) and those of C. fallacissimus and of the flowers and anthocarps were taken with a Nikon Digital C. decipiens are elongated with a roundish base (lanceolate-ovate). Camera DXM 1200F fitted on a Nikon SMZ 1500 stereomicroscope. Meikle (1978) stated that all the Commicarpus species are superfi- Morphological terminology follows Hickey and King (2000). cially similar in growth form and foliage, and the southern Africa spe- Distribution records and habitat information were obtained from cies considered here are also not easily distinguishable in the field herbarium specimens as well as observations made during field trips. despite the slight variations in habit and leaf shape discussed above. Localities and species distribution patterns were mapped using Vegetative characters therefore do not provide evidence on which ArcView 9.2 (ESRI, 2006). Information on the uses was gathered to base a classification system. from herbarium label information, and a broad literature survey. Where fresh plant material was sampled, soil from the top 100 mm of the root zone of a plant was also collected. This was 3.2. Reproductive morphology repeated for five plants per population in 100 m2 to make-up a composite soil sample for analysis. All soil samples were air dried The inflorescence in Commicarpus is an umbel, sometimes with up to and sent to Eco-Analytica for analysis (North-West University, four whorls together in the same inflorescence. The peduncles are 20– Potchefstroom). Soils were analyzed for cation exchange capacity 150 mm long (Table 1). The flowers of Commicarpus are infundibiliform (CEC), electrical conductivity (EC), pH (H2O and KCl), phosphorus (Fig. 3). The flowers can be divided into two parts, an upper petaloid − 2− (using Bray method), nitrogen (NO3 ), sulfur (SO4 ), and extract- part and a distinct lower, coriaceous part (Fig. 4). The upper petaloid able K+,Mg2+,Ca2+,andNa+. A total extract of 29 metals was part has white, pink or purple flavonoid pigmentation and is done per composite sample. Additionally, particle size distribution 2–17 mm long (Table 1). The lower, coriaceous part of the flower is was conducted to determine the soil texture. 2–7 mm long and can be cylindrical (C. chinensis subsp. natalensis, C. Red list assessments of the species indigenous to South Africa fallacissimus and C. plumbagineus var. plumbagineus), clavate (C. were taken from Raimondo et al. (2009). However, the Namibian en- decipiens and C. pentandrus)
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