Two new species of Cineraria (Senecioneae, Asteraceae) from KwaZulu-Natal, South Africa

Two new species of Cineraria L. (Senecioneae) are described: C. glandulosa Cron, endemic to KwaZulu-Natal and C. dryogeton Cron from the Umtamvuna Nature Reserve on the border of KwaZulu-Natal and the Eastern Cape. Both species are narrow endemics, restricted to specific habitats and geological formations, and are therefore vulnerable to human-induced disturbance and the threat of extinction. However both species occur in nature reserves, which reduces the threat of extinction. Introduction Cineraria L. (Senecioneae, Asteraceae) comprises mainly perennial herbs and subshrublets with heterogamous, radiate capitula with yellow florets. As the name suggests (from ‘cinereus’ meaning ash-coloured), many of the species have grey leaves and stems due to a tomentose or cobwebby indumentum. The genus is distinguished by its obovate, compressed cypselae with narrow wings or margins, and palmately veined, commonly auriculate leaves (Hilliard 1977, Cron 1991, Bremer 1994). The capitula have a uniseriate, calyculate involucre and the cypselae have a distinct carpopodium. As a senecioid member of the Senecioneae, Cineraria has balusterform (dilated) filament collars in its anthers and discrete stigmatic areas (Nordenstam, 1978; Bremer, 1994). The style apex is obtuse, usually fringed with sweeping hairs. Cineraria is essentially an African genus, ranging from the Cape Peninsula and mountains of South Africa along the eastern highlands of Africa to Ethiopia. In the west it occurs in the mountainous areas of Namibia and southern Angola. One species, C. abyssinica Sch.Bip. ex A.Rich extends from Ethiopia into Yemen and Saudi Arabia and another, C. anampoza (Baker) Baker, is endemic to Madagascar. Cineraria has an afromontane (to afroalpine) affinity, occurring at 1600–4300 m in the mountains of tropical East Africa (Hedberg 1957), and 1200–3300 m in the Drakensberg range in South Africa (Hilliard 1977, Cron 1991), but also reaches sea-level in the Western Cape, a pattern consistent with the biogeographic relationship of altitude and latitude.


Introduction
Cineraria L. (Senecioneae, Asteraceae) comprises mainly perennial herbs and subshrublets with heterogamous, radiate capitula with yellow florets. As the name suggests (from 'cinereus' meaning ash-coloured), many of the species have grey leaves and stems due to a tomentose or cobwebby indumentum. The genus is distinguished by its obovate, compressed cypselae with narrow wings or margins, and palmately veined, commonly auriculate leaves (Hilliard 1977, Cron 1991, Bremer 1994. The capitula have a uniseriate, calyculate involucre and the cypselae have a distinct carpopodium. As a senecioid member of the Senecioneae, Cineraria has balusterform (dilated) filament collars in its anthers and discrete stigmatic areas (Nordenstam, 1978;Bremer, 1994). The style apex is obtuse, usually fringed with sweeping hairs.

Cineraria glandulosa
CONSERVATION STATUS. Rare and vulnerable, known from few localities and restricted in distribution with small populations. This species has been collected twice from Loteni Nature Reserve, where it is protected. Classified as "Near threatened" as its habitat is not very prone to destruction by humans.
NOTES. Cineraria glandulosa resembles C. erodioides in the decurrent auricles on the petiole and reniform leaves (Figure 2a, 3a, b), but differs in the dense glandular indumentum of the leaves and stem and calyculus bracts (and sometimes the involucral bracts) and the generally smaller capitula (five ray florets and 18-24 disc florets). C. erodioides is a very variable species with 5-8 (-13) ray florets and (20-) 25-40 (-80) disc florets, but the populations with smaller capitula mainly occur at low altitude in the Eastern Cape. Hilliard (1977) recognised this entity as a potentially distinct species (no. 6, p. 379-380), but included in her description Moll 3037 (K, NU) from Richmond District, matched by Cron & Scott-Shaw 9 (J, K, MO) from the same district. Although these specimens have a glandular indumentum (Figure 2b), the leaves resemble those of Cineraria atriplicifolia in their deltoid shape and dissection, especially of the uppermost leaves (Figures 3c, d;Hilliard, 1977: Figure 15Da). The auricles are not decurrent and also differ in being sharply toothed and dissected. The specimens have many more capitula on shorter peduncles and their growth form resembles C. atriplicifolia more closely than C. glandulosa. C. atriplicifolia also has small capitula in a compact synflorescence, but has caducous auricles and has glandular hairs only in the angles of the lobes of its leaves. The striate patterning of the epicarp of the cypselae (Figure 2c) also matches C. atriplicifolia (Cron 1991). We suggest that these specimens are the result of reticulate evolution between C. glandulosa and C. atriplicifolia. The manuscript name of 'C. collina' Hutch & Taylor ined. is indicated on the Medley Wood 1898 specimens (NH, K), but the name has never been published.

Specimens examined
CONSERVATION STATUS. Rare, possibly endangered due to its limited distribution, very likely restricted to this forest and similar forests in the region. EN B2ab(iii); D.
NOTES. Cineraria dryogeton shows an affinity with C. geraniifolia in its growth form and leaf shape, but differs markedly in the dense indumentum of its stems and leaves ( Figure 3). C. geraniifolia is typically a multi-stemmed, slender, perennial herb with reniform, lobed leaves and two or three fairly large capitula on long peduncles, with 12 or 13 involucral bracts and 8 -13 rays. In contrast, C. dryogeton is usually a single-stemmed trailing annual or biennial herb, bearing smaller capitula with 8 involucral bracts and 5 (rarely 7 or 8) rays. It is difficult to ascertain whether C. dryogeton is annual or a shortlived perennial as its habitat is dense, tall grassland along forest margins subject to regular burning (A. Abbott and A.E. van Wyk, pers. com.). This may also account for the long, unbranched stem of C. dryogeton (Figure 2d) that enables it to compete with the tall grass, which in turn serves to support it. The cypselae of both species are glabrous with a distinct pale narrow wing or margin, but are broadly obovate in C. geraniifolia and narrowly-obovate in C. dryogeton (Figure 2f). Occasionally, the lobing of the leaves of C. geraniifolia is deeper than usual, but then the lobes are not as large nor as sharply acute (Figure 3 h, i, j) as in C. dryogeton (Figures 3e, f, g). The auricles also differ, being much more sharply toothed and deeply dissected and conspicuous in C. dryogeton than in C. geraniifolia.
Cineraria geraniifolia is a widespread and quite variable species, extending from East London in the Eastern Cape through KwaZulu-Natal to Graskop in Mpumalanga. It occurs mainly at higher altitudes (1200-2500 m) than C. dryogeton and is found in the Amatole and Great Winterhoek Mountains in the Eastern Cape, and in the Drakensberg of KwaZulu-Natal. Some of the lower-altitude, coastal specimens from Gonubie in the Eastern Cape are unusual, having very elongate leaves and growing in marshy areas (Cron 1991). Another odd form of C. geraniifolia grows in grazed fields of the Hogsback, and has extremely dissected leaves, with very narrow lobes (e.g. Johnson 1187, Phillipson 1276, Cron & Goodman 563). Allied forms of C. geraniifolia in the mountains near Maclear in the Eastern Cape [e.g. Bester 2459 (PRU)] have smaller capitula with 8 involucral bracts and 5 rays. These forms differ from C. dryogeton in their degree of lamina dissection and shape of the leaf lobes, indumentum, and the very small or absent auricles.

Endemic regions and conservation status.
A considerable proportion (22%) of species in Cineraria are narrowly endemic. Most of them are associated with centres of plant endemism (Van Wyk and Smith 2001), as are the two species described here.

KwaZulu-Natal Midlands
The KwaZulu-Natal Midlands are the centre of diversity for Cineraria within southern Africa: 34% (12/35) of the species occur in the province with the largest number in degree grid squares 2929 and 3029. This centre of diversity is part of the larger Maputaland Pondoland Phytogeographic Region (Van Wyk and Smith 2001). The area is dissected by river gorges and the altitudinal range is considerable (600-1500 m). Grasslands predominate with pockets of forest in areas protected from fire. The Midlands merge into the foothills of the Drakensberg escarpment. All these factors contribute to a varied landscape which can accommodate a wide diversity of plants.
In KwaZulu-Natal, the Asteraceae outnumber other families of flowering plants: 551 indigenous species in 113 genera, with 30% endemic to the province (Hilliard 1978). Hilliard (1978) described two minor centres of endemism in KwaZulu-Natal, one associated with the Natal Group and Msikaba Formation sandstone (where about 4% of the Asteraceae endemic to KwaZulu-Natal occur); the other on the sandy Maputaland (= Tongaland) coastal plain. Cineraria occurs in five of the twelve associations/groups of Asteraceae in KwaZulu-Natal recognised by Hilliard (1978). C. glandulosa and C. dryogeton both appear to fall into Group 3, viz. KwaZulu-Natal endemics not confined to the Drakensberg, and some in the " Table Mountain Sandstone" outcrops and gorges, as they were previously known.
Cineraria glandulosa grows at the base of cliffs above river gorges or on the slopes of river valleys, at altitudes ranging from 1000 to 1800 m in Mphendle, Umlazi and New Hanover Districts and at lower altitude on the Oribi Flats near Murchison (Port Shepstone District). The populations resulting from putative hybridization between C. glandulosa and C. atriplicifolia occur in grassland or scrub associated with river gorges in the Richmond District. C. atriplicifolia is also rare with a scattered distribution and is classified at a "lower risk" level of threat (Scott-Shaw 1999). C. glandulosa is certainly also rare with small, scattered populations, with a relatively small range (estimated at less than 20 000 km ). Its habitat is restricted, but the threat by expansion of rural 2 communities is limitied due to its location on river banks and at the base of cliffs and its occurrence in the Loteni Nature Reserve provides some protection. It is therefore classified as "near threatened".

Pondoland
Cineraria dryogeton is thus far known only from the Umtamvuna Nature Reserve on the border between KwaZulu-Natal and the Eastern Cape, an important conservation area in the Pondoland Centre of Plant Endemism (Abbott et al. 2000). This area is rich in palaeoendemics and also neoendemics with a strong Cape and Afromontane affinity (Van Wyk and Smith 2001). The predominant rock type in the Pondoland Centre is sandstone of the Msikaba Formation with a quartz content of 70-96%, with potassium feldspar and mica next most abundant (Hobday and Mathew 1974). The sandstone islands are surrounded by sediments of the Karoo Supergroup and basement rocks (granites, gneisses, schists) of the Natal Metamorphic and Structural Province. The Msikaba Formation sandstones show more similarity to the Table Mountain Group sandstone of the Cape than to the Natal Group sandstone (Van Wyk and Smith 2001). The Umtamvuna River Gorge is one of the narrow river gorges with forest on the steep sides and a grassland plateau above that characterise the Pondoland sandstone region. A large number of plant species are endemic or largely confined to these sandstone areas, including at least 17 endemic/near-endemic herbs and shrublets occurring inside forests or on forest margins, three of which are Asteraceae (Van Wyk 1990).
Van Wyk and Smith (2001) noted that there is a clear path of migration between the Pondoland Centre and KwaZulu-Natal Drakensberg via the Ngeli range. This might have been the migration route for C. dryogeton if, as suggested here, it is closely related to C. geraniifolia, which occurs in the Drakensberg and the Eastern Cape mountains. C. albicans, a near-endemic to the region, also occurs in the adjacent mountains of the Eastern Cape and the KwaZulu-Natal Midlands (Hilliard 1977, Cron 1991.
Much of the grassland in the Pondoland Centre has been floristically depleted or destroyed by anthropogenic activities and the remaining small patches of forest are under intense human pressure (Van Wyk 1993, Abbott et al. 2000. C. dryogeton, although protected in the Umtamvuna Nature Reserve, is known only from a few small populations in the reserve, and would be under considerable threat of extirpation in similar habitats outside the reserve where it might occur. It is therefore considered endangered.