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Home , Cichorioideae, Distephanus, Ethulia, Pteronia

Chapter 26

Eremothamneae Harold Robinson and Vicki A. Funk

HISTORICAL OVERVIEW AND MORPHOLOGY and Thyret (1971) suggested that Eremothamnus should form a monogeneric subtribe in Arctotideae and in one The Eremothamneae was fi rst described by Robinson case, on the basis of achene anatomy, Reese (1989) seemed and Brettell (1973) on the basis of Eremothamnus marlo- to agree that Eremothamnus was in the subfamily Cichori- thianus O. Hoff m. of Namibia. The combination of yel- oideae, but not in Arctotideae. low rays and capillary pappus caused Hoff mann (1890– Robinson and Brettell (1973) discovered two particu- 1894) to place the in the tribe Senecioneae and larly distinctive features in Eremothamnus. One was the in the subtribe Liabinae (currently tribe ), at that structure of the endothecial cells, with annular thicken- time appended to Senecioneae (Figs. 26.1–26.2). Liabinae ings on the lower half but none on the upper half. A with its opposite , milky sap, and mostly neotropical second and more striking character was the rather long distribution was an unusual place for the alternate-leaved sweeping hairs divided longitudinally into two or three Eremothamnus. At the time Liabinae included the then cells, each cell with its own point (Fig. 26.3). These mostly African, alternate-leaved, Newtonia O. Hoff m. sweeping or collecting hairs are borne on slender style ( = Cass.), but all these entities have now been branches and are distributed on the style as in excluded from Senecioneae. In Robinson and Brettell and Liabeae with no sharp basal delimitation. They dif- (1973) the basic diff erences in stigmatic surfaces, lengths fer from the shorter, single-celled sweeping papillae on of disk corolla lobes, and elongated or calcarate bases broader style branches and upper shafts in Arctotideae. of the anther thecae were noted, characters that distin- The latter’s scabrae end below in an abrupt basal collar. guish all the members of the subtribe Liabinae from most Karis (1992) discovered that a second genus, Hoplo- Senecioneae. The diff erences now seem to be important phyllum DC., from western South Africa, was related to distinctions of the subfamily (in the present Eremothamnus. The position was confi rmed by Robinson sense, Vernonieae, Liabeae, Arctotideae, plus (1992, 1994) in studies of the sweeping hairs and the pol- smaller tribes) from Asteroideae to which Senecioneae, len (Fig. 26.4). The type of Hoplophyllum had a , Heliantheae, etc., belong (see Chapter 1). There much longer history than Eremothamnus, being described have been suggestions that Eremothamnus be placed in the as spinosa L. f. in 1781. Placement in Pteronia African genus Pteronia L. of the tribe Astereae (Muschler (Astereae), in the wrong subfamily, was the only taxo- 1911; Hutchinson and Phillips 1917; Dinter 1927), or pos- nomic placement that Eremothamnus and Hoplophyllum sibly in Inuleae (Moore 1929), along with the suggestion shared before the study by Karis (1992). Pteronia spinosa that it be returned to the overly broad tribe Senecioneae was transferred by De Candolle (1836) to his new genus (Merxmüller 1954). Nevertheless, all more recent com- Hoplophyllum which was placed in Vernonieae. In spite parisons of Eremothamnus have been with members of of its yellow fl orets (Fig. 26.2), it was left in Vernonieae Cichorioideae, especially Arctotideae (Leins 1970). Leins until the study by Karis (1992). It should be noted that 412 Robinson and Funk

Fig. 26.1. Eremothamnus marlothianus O. Hoff m. A habit, note prostrate shrubby nature and dry rocky, sandy habitat; B fl owering heads during a year (2006) of unusually high rainfall, note the large number of fl orets; C fl owering heads during a year (2007) of low rainfall, note the small number of fl orets. [Photographs: A, B, C. Mannheimer; C, V.A Funk, Funk 12684; all from the same population.] Chapter 26: Eremothamneae 413

Fig. 26.2. Hoplophyllum spinosum (L. f.) DC. A habit; B fl owering heads, note absence of ray fl orets; C involucral , note hyaline margins. Taxon can be locally common in somewhat disturbed areas. [Photographs, V.A. Funk; Funk 12650.] 414 Robinson and Funk

Fig. 26.3. Sweeping hairs. A, B Hoplophyllum spinosum (L. f.) DC. (B detail from A). Note the slender style branches bearing comparatively long sweeping hairs that are divided longitudinally into two or three cells. [Photographs, H. Robinson; Barker 9757, US.]

Fig. 26.4. Pollen. A–C Eremothamnus marlothianus O. Hoff m.; D–F Hoplophyllum spinosum (L. f.) DC. The pollen grain structure is unique in the but obviously shared between the two genera. Scale bars for A, B, D, E = 10 μm; scale bars for C, F = 1 μm. [Photographs: A–C, J.J. Skvarla, Geiss and Vauuren 710, PRE; D–F, J.J. Skvarla, Koekemoer 2045, PRE.] Chapter 26: Eremothamneae 415 the two genera had such diff erent histories primarily be- well supported (Funk et al. 2004), and the two subtribes cause Eremothamnus had ray fl orets and Hoplophyllum did sometimes fall into sister group relationships with other not (Figs. 26.1–26.2). As pointed out by Karis (1992), the tribes. In summary, one can say that the subfamily analysis genera also share spinose or spiniform leaves, as seen in (Chapter 23) indicates that Eremothamneae is close to, but many Arctotideae and Cardueae, and they share caveate probably not in, the tribe Arctotideae (or its subtribes), and pollen (Fig. 26.4), a feature most common in the subfam- for now it is best to leave it in its own tribe. ily Asteroideae, but found in a few Cichorioideae such as some Liabeae (Robinson and Marticorena 1986) and Vernonieae ( L.; Bolick 1978).

Tribe Eremothamneae H. Rob. & Brettell in Phytologia PHYLOGENY 26: 164. 1973 – Type: Eremothamnus O. Hoff m. Erect branching , stems very sparsely puberulous Karis et al. (1992) performed a cladistic analysis of Cichori- in Hoplophyllum to densely tomentose in Eremothamnus, oideae s.l. based on morphological data and concluded hairs of tomentum long and simple, highly contorted, uni- that Eremothamnus was related to Arctotis (the only sample seriate, multicellular, crosswalls oblique; pith solid. Leaves from Arctotideae) but the sampling was very limited. In alternate, spiniform in Hoplophyllum, or obovate with few a separate paper in 1992 (p. 197) Karis concluded that short spines on distal margins and somewhat succulent in “Hoplophyllum … is more closely related to Eremothamnus Eremothamnus. Heads solitary and terminal or in upper than to any taxon in the Vernonieae. The unique endoth- axils, or in terminal clusters, sessile or subsessile; involu- ecial confi guration shared with Eremothamnus constitutes cral bracts multiseriate, gradate, distally papyraceous and a synapomorphy for these two taxa.” Another putative usually with apical spine; epaleaceous. Corollas synapomorphy is the accessory apical vein in the lobes yellow; without ray fl orets and with ca. 5 disk fl orets in of the disc fl orets, occurring in both genera. He also Hoplophyllum, with 12–20 rays and 25–30 disk fl orets in mentioned that Robinson (pers. comm.) had pointed out Eremothamnus, rays when present without anthers, with the unique hairs (Fig. 26.3). In 1995 Bergqvist et al. 3 short apical teeth; disc fl orets regular, deeply 5-lobed, used chloroplast DNA restriction site data and showed lobes linear, outside with glands and few hairs; anther th- that Eremothamnus was the sister group to , how- ecae calcarate at base and shortly tailed; endothecial cells ever, again, the sampling was limited to only one mem- oblong with annular thickenings crowded in basal half, ber of the tribe Arctotideae. In 2001 (Karis et al.) the lacking from distal half; apical appendage oblong with chloroplast gene ndhF was used to investigate the place- apiculate tip in Hoplophyllum, soft and broadly elongate ment of Eremothamnus and Hoplophyllum. The analysis in Eremothamnus; style base partly immersed in nectary, included fi ve species of Arctotideae and it resulted in a without enlarged basal node; branches narrow with stig- poorly supported trichotomy among the two subtribes of matic papillae covering entire inner surfaces; upper scape Arctotideae and the Eremothamnus-Hoplophyllum clade. of style and outer surfaces of branches not broadened, Recent molecular studies involving a large selection covered with elongate sweeping hairs formed by two of taxa and several additional gene regions support the or three cells separated by longitudinal walls. Achenes hypothesis that Eremothamnus and Hoplophyllum are sister prismatic, densely and sericeously setuliferous, with long taxa (Funk et al. 2004). However, the taxa are found on a biseriate setulae not or scarcely divided at tips, with thick long branch that can be found in more than one place on cell walls, achene surface smooth, shape slightly trigonous the phylogeny depending on the outgroups selected and to pentagonal, walls without phytomelanin, with linear the gene regions used in the anlaysis. In fact, at this point, raphids; pappus of many capillary bristles in 2–3 series, the molecular data give more information about where somewhat gradate in length, tapered at tips. the clade does not belong than where it does belong. Eremothamneae are presently seen to include the two Certainly it is in the subfamily Cichorioideae and not in genera Eremothamnus and Hoplophyllum, from Namibia any of the tribes of Asteroideae. Within the subfamily it and adjacent South Africa. Eremothamnus is monotypic is never found within Vernonieae, Liabeae or Cichorieae. with one species, Eremothamnus marlothianus O. Hoff m., Furthermore, it does not fi t in either of the well defi ned restricted to one area in Namibia, and Hoplophyllum has subtribes of Arctotideae. Sometimes it can be found as the two species, Hoplophyllum spinosum (L. f.) DC. and H. ferox weakly supported sister group of one or the other of these Sond.; the former is the more common of the two being two subtribes, most often Gorteriinae, but at other times found occasionally in the Western and Northern Cape it is the sister group of the unplaced genus or in Provinces along roadsides, while the latter is restricted various polytomies with all of the tribes except Cichorieae. in its distribution to the Western Cape. The general To add to the problem, the of Arctotideae is not taxonomic position determined by Robinson and Brettell 416 Robinson and Funk

(1973) and expanded by Karis (1992) has proven correct. BIOGEOGRAPHY The structure of the anthers with calcarate bases, the long lobes of the disk corollas, the stigmatic papillae cover- Because the Eremothamneae clade is small and its place- ing the entire inner surface of the style branches and the ment is not secure, it is not possible at this time to dis- pollen place the tribe defi nitely outside of what is now cuss its biogeography, but its presence in Cichorioideae recognized as the subfamily Asteroideae. strengthens the overall African presence of this subfamily The Southern African distribution is reminiscent of (see Fig. 44.4). groups such as Arctotidinae in Cichorioideae with which Leins (1970) and others have compared Eremothamnus. However, the structure alone shows a form of narrow style CHROMOSOME NUMBERS, CHEMISTRY, BIOLOGY, and unique form of sweeping hairs (Fig. 26.3) that would ECOLOGY, ETHNOBOTANY preclude inclusion of Eremothamneae in Arctotideae, Platycarpheae, or in a relationship with Heterolepis Cass. Nothing is known.

POLLEN Acknowledgements. The authors thank Marjorie Knowles (US), for proof-reading the manuscript and Tim Lowrey (UNM) and The grains are spherical, tricolporate, spinulose with Sterling Keeley (HAW) for their helpful reviews. We thank John larger spines in intercolpi and smaller crowded spines J. Skvarla (OKL) for the pollen SEM photos and C. Mannheimer around poles, with microperforations in exine and an (WIND) for photos of Eremothamnus. Funds for the fi eld trip interior of exine caveate, with thin foot-layer, individual (by Funk) to South Africa and Namibia were provided by the spines fi stulose at the base (Fig. 26.4). It is unique in the Smithsonian Institution’s Unrestricted Endowment Fund. family and shared by both genera.

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