New species in the section Multinerviae of Cliffortia ( Rosaceae )

Three species complexes in section Multinen iae DC. of Cliffortia L. are discussed. Two of the varieties of C. ilicifolia L. are raised to specific rank: C. reniformis and C schlechteri. as are both varieties of C. grandifolia Eckl. & Zevh.: C denticulata and C recurvata Two new species are described that were previously included within the variable species C dregeana C. Presl: C acanthophvlla and C. ceresana. Three further new species belonging to section Multineniae are also described: C oligodonta, C. prionota and C seandens


INTRODUCTION
The sectional classification of the genus Cliffortia L. is in need of revision (Whitehouse 2003).However, the cur rent circumscription (Weimarck 1934(Weimarck . 1948) ) of section Multineniae DC. can be retained for the most part.Species of section Multinen iae are generally medium to tall shrubs with tough, rigid, unifoliate leaves that are often sharply pointed and toothed and have several veins from the base.They also have male flowers with numer ous stamens (2 0 +), which are almost always white to yel low in colour, and female flowers tucked down at the base of the leaves and consequently often hidden from sight.
Within section Multinen iae there are three species com plexes, which have previously been regarded as three vari able species each with a number of varieties.The C. ilici folia L. complex has a southern Cape distribution, between Cape Town and Port Elizabeth, whereas C. dregeana C. Presl has a distribution along the western Cape mountains, from the Riviersonderend Mountains northwards to the Cederberg.The other complex.C. grandifolia Eckl.& Zeyh., is restricted to the southwestern Cape mountains and Langeberg.In this paper, four of the species complex varieties are raised to specific status and two new species belonging to the C. dregeana complex described.Three other new species belonging to section Multineniae are also described.
C ILICIFOLIA COMPLEX C. ilicifolia L. has been subdivided into five varieties based upon the shape of their leaves.Var.cordifolia (Lam.)Harv.and var.incisa Harv.are doubtfully distinct from the typical variety.In particular, var.incisa is proba bly a sh(x>t from a resprouting plant, the leaves of which are often more markedly toothed in this species than mature leaves.Var.cordifolia is possibly worthy of recog nition.as it is generally restricted to the southeastern mountains of the Cape Floristic Region, although there is intergradation with the typical variety and some indication that the form may be associated with aridity or altitude. In contrast, the two varieties described by Weimarck (1934) are very distinctive, no intermediates have been found between them and the typical variety, from which they are allopatric.They are.therefore, here elevated to the rank of species.Erect, tall shrub, up to 3 m high, resprouting after fire and spreading clonally; dominant main stem with determi nate length side branches held horizontally; only forming long branches: young stems 1.3-2.6 mm wide, completely sheathed by leaf bases.Leaves unifoliate, subcircular, 12-20 x 16-26 mm.clasping stem at base, apex sharply long-acuminate to pungent.0.2-0.9mm long, margins flat, with 6-11 broad teeth.2.8-5.0 mm long: lamina 11-17veined from base, tough and rigid.0 .2-0.5 mm thick, held at right angles to stem, glaucous, glabrous: sheath 1.4-6.2mm long, abaxially glabrous, adaxially glabrous except for fringe of hairs at apex: stipules similar in texture to lamina, 2.6-5.9 mm long, fused into a single stipule on reverse side of stem, margin smooth: petiole absent.Flow ers solitary in axil of undifferentiated leaves; bracteoles hairy on keel, margins smooth: sepals 3. glabrous.Male flow ers: bracte oles 3.6-4.9mm long; pedicel 0.9-1.4mm long; sepals broadly ovate.6.4-9.2 x 4.5-6.6 mm.acute to acuminate at apex; stamens 51-68: filaments 3.8-6.8mm long, greenish white; anthers yellow .Female flow ers: bracteoles 3.9-5.0mm long, longer than immature receptacle: sepals broadly ovate to triangular. 2 .1-2.8 x 1.0 -1.8 mm.erect: carpel 1; stigma 1.4-2.4mm long, greenish white, feathery, hidden at base of leaves; immature receptacle 1.7-2.5 x 1.5-2.3mm, glabrous, smooth.Achene 8.0-8.7 x 4.8-5.5 mm.medium to dark brown, glabrous: ribs rounded.Flowering time: ± all year, but mature fruits only found between August and November.Figure 1.
Habitat: tall, streamside vegetation and wet fynbos of lower valley slopes, on soils derived from  Distribution: restricted to the Langeberg at Garcia's Pass and the immediately surrounding valleys (Figure 2).Conservation status: highly restricted but very com mon and frequently dominant in the area.
Etymology: reniformis means kidney-shaped, refer ring to the leaves which are almost circular in outline and clasp the stem at their base.
This species is very distinctive on account of its unique growth habit: a single stem often up to 3 m tall with short branches all the way up, spreading at right angles to the main stem.The leaves are also diagnostic, being held at right angles and almost circular in outline with a leaf-like stipule formed by fusion of the two sep arate ones on the reverse side of the stem.Erect, medium, densely branched shrub, up to 1.8 m high, killed by fire; only forming long branches spread ing at right angles; young stems 0 .9-2.0 mm wide, com pletely sheathed by leaf bases.Leaves unifoliate, subcir cular to broadly ovate or oblong, 5 .5 -10.9 x 5 .7-12.0 mm, clasping stem at base, apex sharply long-acuminate to pungent, 0.3-1 mm long, margins flat or rounded, with 2-4 broad teeth, 0.6-2.6 mm long; lamina 7-10-veined from base, tough and rigid, 0.2-0.4mm thick, held at right angles to stem, glaucous, glabrous above and beneath; sheath 1.4-2.1 mm long, abaxially glabrous, adaxially glabrous except for fringe of hairs at apex; stip ules 1.6-5.8mm long, free, margin smooth; petiole absent.Flowers solitary in axil of undifferentiated leaves; bracteoles hairy on keel, margins smooth; sepals 3, glabrous.Male flowers: bracteoles mm long; pedicel 0.9-1.5 mm long; sepals broadly ovate, 5-8 x 3.2-5.7 mm, acute to acuminate at apex; stamens 2 8 ^1 ; filaments 2.8-5.0 mm long, greenish white; anthers yel low.Female flowers: bracteoles 2.3-3.2mm long, longer than immature receptacle; sepals ovate to triangular, 1.2-1.7 x 0 .9-1.2 mm, erect; carpel 1; stigma 1.1-1.4mm long, greenish white, feathery, hidden at base of leaves; immature receptacle 1.5 -1.8 x 1.0 -1.3 mm, glabrous.Achene 6.7-8.7 x 3.4-5.4mm, medium to dark brown, glabrous; ribs rounded.Flowering time: ± all year, but mature fruits only found between May and November.Figure 3.
Habitat: fynbos on very well-drained, wind-blown sands and limestone-derived soils in full sun; altitude 0-300 m.
Distribution: widespread on the Agulhas Plain from Gansbaai to Gouritz River mouth (Figure 2).
Conservation status: widespread and often common, occurring in several reserves, as well as along road verges.However, it may be susceptible to local extinc tions caused by unnatural fire regimes as it does not resprout after fire.
Etymology: named after Rudolph Schlechter, who was the first person to collect this species.

C. DREGEANA COMPLEX
The C. dregeana C.Presl complex still needs much revi sion.The species are characterized by their rigid, tough, pungently tipped leaves and large achenes.Weimarck (1934) divided the species into two varieties, var.dregeana and var.meyeriana (CPresl) Weim., the former having entire leaves, whereas the latter has 2-4 stout teeth.However, he admitted that the boundary between the two varieties was vague.Furthermore, as populations are found with both entire and toothed leaves, the varieties cannot be maintained.
Conservation status: limited in its distribution, but well protected within the Cederberg Wilderness Area, however, the Graafwater populations are all on private land.
Habitat: fynbos on rocky shale slopes and amongst sandstone rocks at high altitudes, on well-drained soils derived from Table Mountain Series in full sun; altitude 350-1 850 m.
Distribution: found from the western parts of the Hex River Mountains, north through the Witzenberg Range to the Groot Winterhoek Mountains (Figure 6 ).
Conservation status: common throughout the area in which it grows, especially at higher altitudes; it is con served within the Groot Winterhoek Wilderness Area and its resprouting habit means that it is not particularly sus ceptible to unnatural fire regimes.
Etymology: named after the town of Ceres, where it is common on the surrounding mountains, and has fre quently been collected in Michell's Pass, which leads to the town.
Habitat: in damp depressions and beside watercours es on well-drained soils from Table Mountain Series in full sun; altitude 350-1 350 m.
Distribution: Franschhoek to the southern Hottentots Holland Mountains around Nuweberg (Figure 8 ).within the Hottentots Holland Nature Reserve; however, being a reseeder it may be susceptible to local extinction caused by too frequent fires.
Etymology: denticulata means very small teeth, which surround the margins of the leaves.
Habitat: in damp depressions and beside watercours es on well-drained soils from Table Mountain Series in full sun; altitude 300-800 m.
Distribution: confined to the Kogelberg between the Steenbras and Palmiet River Valleys (Figure 8 ).
Conservation status: only known from the middle slopes of the Kogelberg, extent of occurrence, < 75 knv, and restricted to a very particular habitat where it is uncommon; being a reseeder it is vulnerable to too fre quent fire regimes; however, it is conserved entirely within the Kogelberg Biosphere Reserve.
Flowering time: September to May. Figure 10C, D.
Habitat: south-facing slopes and in the shelter of rocks at high altitudes, in well-drained rocky soil from Table Mountain Series; altitude 1 450-1 800 m.
Distribution: very narrow endemic, found only on the upper slopes of the Wemmershoek Mountains between Winterberg and Wemmershoek Peak (Figure 11), Conservation status: endemic to a very small area, extent of occurrence, < 2 0 knv, but common there at high altitudes.
Habitat: in clayish soil on shale bands of Table Moun tain Series rocks in full sun. in well-drained soils or along the edge of damp gullies; altitude 50-350 m.
Distribution: restricted to the lower mountain slopes of the Palmiet River Valley and behind Kleinmond (Figure 11).
Conservation status: endemic to a small area and uncommon there forming scattered clonal patches.Popu lations behind Kleinmond may be threatened by urban expansion but the majority of the populations are pro tected within the core region of the Kogelberg Biosphere Reserve and being a resprouter it is not particularly threatened by unnatural fire regimes.
Etymology: prionota means like a saw, referring to the leaves, which normally have fine teeth along their edge; it also alludes to the fact that the species is confined to the Palmiet River area and palmiet is the common name for the plant Prionium serratum.
Previously classified with C. intermedia, this species actually bears closer resemblance to the Langeberg endemic, C. lanceolata but lacks the hairs on the anthers that are diagnostic for that species.C. prionota has a very narrow range, being endemic to the Kogelberg, and for that area the long, narrowly lanceolate leaves on a clonally, spreading erect shrub, prevent confusion with other species.However, the species is comparatively variable within its range, from having leaves that are tough, rigid and entire to ones that are relatively soft and flexible and bearing numerous small teeth.This degree of variation may be the result of introgression with other species, such as the sympatric C. heterophylla (e.g.Levyns 7042).
Conservation status: a poorly known species, record ed from just two areas of the Riviersonderend Mountains and apparently very localized there, but the mountain range as a whole is relatively unexplored; invasive alien plants and unnatural fire regimes are threats, but the majority of the mountain range is conserved within a nature reserve.
Etymology: scandens means scrambling, referring to the way the plants form a tangled growth.This species has previously been placed under C. uncinata or C. acutifolia.However, both those species are only known from mountains north of Tulbagh.The species is easily recognized within its distribution range by its long, narrow, toothed leaves, which form brachy blasts, and the plant's scrambling habit.However, hybrid swarms between C. dregeana and C. pungens have been confirmed in the Riviersonderend Mountains (Whitehouse 2003) and may involve this species as well.Such hybrids may include the types of C. the odor i-friesii Weim.var.puberula Weim.or C. meyeriana C.Presl.Much work in this geographical area is needed to identi fy the parent and hybrid populations, especially to deter mine the limits of the species involved and the parents of the various hybrid populations.Despite this apparent confusion of species limits, preliminary molecular evi dence supports the identification of C. scandens as a dis tinct species, as it has both nuclear and chloroplast mark ers not found in any other species (Whitehouse 2003).
the rest of the lamina, whereas C. dregeana also has thickened margins but are not so pronounced.The distri butions of the two species do not overlap, and both are replaced by the more easily discernible C. ceresana in the intervening mountains.However, some specimens of C. dregeana from the Hex River Mountains do have thickened margins similar to C. acanthophylla: .ilicifoliaauct.nonL.: Weim.:99 (1934)pro parte.
Table Mountain Series in full sun; altitude 300-1 600 m. with C Weimarck (1934)ts broad, toothed leaves.However, in other characteristics it is closer to C. dregeana, with which it has generally been placed in herbaria.The broadly oblong, few toothed leaves, combined with the limited distribution range, make this species easily distinguishable.However, in the south of its range, specimens that are more intermediate in character with C. dregeana do occur:The members of the C. grandifolia Eckl.&Zeyh.complexare all tall, sparsely branched shrubs with large leaves.Weimarck (1934)divided C. grandifolia into three varieties based upon the shape of the leaves.The varieties are easily identifiable, as well as allopatric in their distributions, especially var.grandifolia, which is endemic to the Langeberg.Intermediates have not been FIGURE 7.-Lectotype of Cliffortia denticulata.C .grandifolia Eckl.& Zeyh.var.denticulata Weim., Monograph of the genus Cliffortia: 115, fig.33C (1934).C. intermedia auct.non Eckl.& Zeyh.: Harv.: 295 (1862) pro parte.