Notes on new and misunderstood taxa of Cyrtanthus (Amaryllidaceae: Cyrtantheae) from the Western Cape, Eastern Cape and KwaZulu- Natal, South Africa

Cyrtanthus aureolinus Snijman is a new, rare species of fire lily, which is localized in a vlei on the northern slopes of the Groot Swartberg, Western Cape. The upright or slightly spreading, yellow to cream-coloured flowers and the perigone tube which gradually widens to the throat suggest that it is closely related to the Western Cape endemic, C. ochroleucus (Herb.) Burch, ex Steud., and C. mackenii Hook.f., a variable species from southern KwaZulu-Natal and Eastern Cape. The species differs mainly by the shape, size and position of the tepals and the length of the filaments. Cyrtanthus mackenii var. cooperi (Baker) R.A.Dyer is raised to subspecies rank as C. mackenii subsp. cooperi (Baker) Snijman. Its hysteranthous leaf habit and grassland habitat differ from the riverine habitat of the evergreen C. mackenii subsp. mackenii. Described in detail are C. aureolinus, C. ochroleucus, and C. mackenii.


INTRODUCTION
Found in southern and East Africa, Cyrtanthus Aiton, comprising about 55 species, is one of the largest gen era in the family Amaryllidaceae. Throughout its distri bution, Cyrtanthus has species adapted to cope with fire and because of this specialization they are commonly known as fire lilies.
In the Cape Floristic Region, some of the first plants to flower in response to summer fires and sufficient moisture are species of Cyrtanthus which characteris tically produce their inflorescences in advance of the leaves. In the heat of summer, however, their flowering is often fleeting, hence many populations of Cyrtanthus in the Cape Fold Mountains are infrequently seen or prob ably yet to be recorded. This became evident after a wild fire burnt a large tract of arid fynbos on the dry north ern slopes of the Groot Swartberg, east of Oudtshoom, Western Cape, in December 2004. Two to three weeks later, a large population of Cyrtanthus appeared in a blackened seepage zone on the lower slopes and the stand of yellow flowers promptly attracted attention from local nature conservationists and botanists. Field and morphological studies of the newly discovered popu lation in relation to other species of Cyrtanthus with tubular, yellow flowers have confirmed that the plants are distinct from all the currently known species of the genus and they are described here as a new species, C. aureolinus.
Due to the discovery of C. aureolinus, its putative close allies, C. ochroleucus (Herb.) Burch, ex Steud. and C. mackenii Hook.f, were examined in detail. The mor phological variability within and among populations of the two varieties of C. mackenii was re-assessed in rela tion to phenology and habitat. Consequently, the rank of C. mackenii var. cooperi (Baker) R.A.Dyer is raised to that of subspecies as C mackenii subsp. cooperi (Baker) Snijman. To accommodate the taxonomic changes and new data, C. mackenii and C. ochroleucus are described and mapped in full. Semi-evergreen, bulbous geophyte, 220-300 mm tall when flowering. Bulb solitary, hypogeal, ovoid, up to 30-45 X 25-30 mm, narrowed to a slender neck 40-60 X 8-15 mm; outer tunics brown and papery; inner tunics cream-coloured and fleshy. Leaves developing sequen tially, either absent or 2 or 3 newly emerging at flower ing, finally up to 4, strap-shaped, up to 300 x 5-9 mm, suberect to recurved, ± channelled, glabrous; abaxial surface with 3 or 4 prominent median veins, not keeled; margin smooth. Inflorescence 4-10-flowered, solitary or occasionally 2 per bulb; scape erect, up to 250 mm long, 5-8 mm diam. at base, tapering distally up to 4-6 mm at apex, round to slightly compressed in t/s, green with a reddish blush towards base, hollow; spathe valves 2, outer sheathing inner at base, lanceolate, exceeding the pedicels, up to 45-65 x 7-9 mm (at base), membranous, suberect and flushed ox-blood red when fresh, later turn ing brown and reflexing; bracteoles several, linear, up to 15 mm long; pedicels suberect to slightly spreading, up to 40 x 1.5 mm at anthesis, lengthening up to 50 mm and becoming more rigid when fruiting, green to pale brown. Flowers erect to spreading slightly above the horizontal, opening sequentially, sulphur-yellow, sometimes with 6 diffuse green median bands from perigone base to tepal tips, firm, unscented, apparently without nectar; peri gone tube dilated gradually to throat, straight or slightly curved distally, 2 9^0 mm long, ± 3 mm diam. at base, widening gradually to 6-8 mm diam. at throat; tepals narrowly ovate, slightly spreading, neither reflexed nor rolled back, 12-15 x 5-7 mm, subacute at apex; outer tepals sometimes slightly narrower than inner, shortly mucronate. Stamens biseriate, regular, attached slightly below and in perigone throat, both whorls shortly exserted; outer filaments ± 5 mm long, attached ± 1 mm lower than inner; inner filaments ± 7 mm long; anthers dorsifixed, ± 4.5 mm long before dehiscing, yellow; pol len yellow. Ovary ellipsoidal, somewhat 3-angled, ± 5 x 2 mm, green; ovules axile, ± 25 per locule; style usually pressed against upper tepals, as long as or slightly longer than perigone, reaching up to 40 mm; stigmatic branches 3, spreading, slender, ± 2.0 x 0.5 mm, truncate, shortly papillate towards apex. Capsule ovoidal, ±15x8 mm. Seeds unknown. Plate 1.
Phenology and pollination: the first flowers appear two to three weeks after summer fires. Individual flowers fade after a few days, but a many-flowered inflorescence may remain attractive for approximately one week. The response of individual plants is also staggered, so the entire flowering period lasts approximately three weeks. Although most individuals produce at least one leaf while flowering, the ftill complement of leaves is pres ent only later in the season. Thereafter the plants remain evergreen until burnt by another fire.
Cyrtanthus aureolinus shows the typical fire lily response of not flowering in the seasons between fires. Observations in the early morning indicate that the flow ers are visited by honeybees which alight on the tepals but do not enter the perigone tube. In the absence o f nec tar, pollen appears to be their only floral reward.
Diagnostic features: in its floral characters, particu larly the form, position and colour of the flowers, Cyrtanthus aureolinus resembles C. ochroleucus from the Langeberg, Western Cape, and to a lesser degree the Eastern Cape and KwaZulu-Natal species, C. mackenii. All three species have somewhat tubular flowers with ± equally long perigone tubes that widen gradually from the base to the throat and have small tepal lobes.
Although the flowers o f C. mackenii and C. ochroleucus vary in colour from yellow to cream-coloured or rarely white, they resemble the clear yellow flowers of C. aure olinus in being upright to somewhat spreading, and are seldom horizontally spreading.
Despite their many similarities, these species can be distinguished florally mainly by the shape, size and position of the tepals, and by the length o f the fila ments (Table 1). The narrowly ovate to oblong tepals in Cyrtanthus aureolinus and C. ochroleucus are more than twice as long as broad and slightly spreading, unlike those of C. mackenii which are ovate, less than twice as long as broad and patent or rolled back at anthesis. The filaments, however, are most diagnostic. In C. aureolinus both the outer and inner whorls of filaments (5 and 7 mm long respectively) extend beyond the perigone throat. Those of C. ochroleucus are slightly shorter (± 2 and ± 4 mm respectively) and only the inner filaments extend beyond the throat. In contrast, both the outer and inner filaments in C. mackenii are very short (± 1 and ± 1 mm respectively) and the stamens remain included in the perigone throat or within the upper part of the tube.
If the features of the tepals and stamens that charac terize the species are also significant for determining affinities, then Cyrtanthus aureolinus is closest to C. ochroleucus. The perigone throat of C. aureolinus, how ever, is wider than that of C. ochroleucus and its leaves are broad and smooth, unlike the narrow leaves of C. ochroleucus which are minutely papillate on the margin and midvein. Both species are found in the fynbos of the Western Cape where they occupy different habitats and display dissimilar flowering habits.
Distribution and habitat: Cyrtanthus aureolinus is confined to damp habitats on the northern foothills of the Groot Swartberg, west of Meiringspoort ( Figure 1). The only known population, covering an area of ± 15 x 40 m in size, is found in a permanent seepage zone in arid fyn bos, where sandstone-derived soils meet a band o f loamrich soils at ± 800 m. In winter the area is waterlogged but it remains sufficiently wet in summer to support a population o f the clicking stream frog, Strongylopus grayii (Smith). In periods between fires (of ± 18-^0 years) the seepage zone is dominated by large clumps of restioids which form dense, fibrous tussocks at their base. Most individuals o f C. aureolinus grow between these clumps, but in the wetter parts o f the vlei, they are found on the raised tussocks. They are absent fi-om the wettest parts of the site. Re-assessment o f Cyrtanthus mackenii and C. ochroleucus The discovery of Cyrtanthus aureolinus, which is putatively most closely related to C. ochroleucus, makes it necessary to focus attention on the question able identity of a group of hysteranthous-leaved plants from the grasslands of the Eastern Cape, northeast of King William's Town-a problem raised by Reid & Dyer (1984) and to some extent by Turrill (1960). Reid «fe Dyer (1984) pointed out that these populations are found in the same area as C. mackenii var. cooperi, but remarked that the plants might possibly represent out liers o f C. ochroleucus, a species originally collected by William Burchell whose travels in the Cape never extended northeast of the Great Fish River (McKay 1943). The inconsistent naming of these particular plants is reflected in Batten & Bokelmann (1966: 31, t. 25, fig. 4) who referred to the plants from near King William's Town as Cyrtanthus sp., and those from near East London as C. ochroleucus (t. 24, fig. 1). In contrast. Manning (2001: 74, fig. 1 & 81, fig. 4) referred to both as C mackenii.
In an attempt to resolve the relationships of the Eastern Cape plants to Cyrtanthus mackenii and C. ochroleucus, all the South African herbarium mate rial (BOL, GRA, NBG, NH, PRE, SAM) belonging to this complex was studied together with the types of C. ochroleucus, C. mackenii var. mackenii and C. mackenii var. cooperi.
The types of Cyrtanthus mackenii var. cooperi-Cooper 1806 (K, NH, PRE) collected in British Kaffraria, the eastern part of today's Eastern Cape Province-all lack leaves which strongly suggests that the leaves are hysteranthous. Furthermore, the pressed inflorescences (on the sheets at K and PRE) closely resemble those of the hysteranthous-leaved plants from the Eastern Cape referred to by Reid & Dyer (1984) with regard to flower number, perigone tube length, size o f the perigone throat (4 mm wide), and shape and size of the tepals (ovate and 4 X 3.5 mm). This close similarity is in contrast to the more distant likeness found between C. ochroleucus and the hysteranthous-leaved Eastern Cape plants which are in question. Despite their shared hysteranthous-leaved state, they differ in leaf width (± 3 vs 7-11 mm); tepal shape (narrowly ovate to oblong vs ovate); tepal size (7-11 X 3-4 vs 3.5-6.0 x 2.5-^.0 mm); and flowering time (Oct.-Dec. vs Aug.-Oct.). Moreover, no intermediates between the two elements are known and their distribu tion ranges are disjunct, separated by a gap o f ± 750 km.
Based on the above comparison, the resemblance of the hysteranthous-leaved plants ft'om the Eastern Cape to the type of Cyrtanthus mackenii var. cooperi is con sidered to be sufficiently close to warrant their inclusion in C. mackenii var. cooperi. This is in agreement with Baker's (1896) circumscription of the taxon which he originally named C. lutescens var. cooperi Baker.
Following the inclusion here of the Eastern Cape's populations o f hysteranthous-leaved plants into Cyrtan thus mackenii var. cooperi, is the need to re-assess the classification of C. mackenii into two varieties which Dyer (1939) and Reid & Dyer (1984) distinguished pri marily by the possession o f yellow to cream-coloured rather than white flowers. To do so, this comparison has incorporated field data on flowering and leafing phenolo gies and habitat which are often missing from herbarium records.
The descriptions of Cyrtanthus mackenii (Dyer 1939;Turrill 1960;Reid & Dyer 1984) mention that the spe cies reproduces vegetatively from bulblets. Field data, however, indicate that this habit is confined to the typi cal variety and leads to the formation of large clumps of bulbs with several shiny, evergreen leaves. Furthermore, the flowers o f C. mackenii var. mackenii are sweetly scented, the perigone has a fairly broad throat (6-8 mm wide), and the tepals (5-10 x 3.5-5.0 mm) are ovate and vary from patent to rolled back at anthesis. Distributed from near Port Shepstone in southern KwaZulu-Natal southwards along the Eastern Cape coast, the plants grow most often on the shaded banks of permanent streams, in patches of remnant forest which seldom or never bum. They flower mainly in winter, fi'om June to September, or rarely as late as November.
In contrast, Cyrtanthus mackenii var. coopeh is deciduous and has solitary bulbs that produce two to five glaucous leaves each season. The flowers have a strong spicy scent with a 4.0-5.5 mm wide perigone throat, and short, ovate tepals (3.5-6.0 x 2 .5^.0 mm) that are slightly recurved to rolled back at anthesis. Populations belonging to this taxon are plentiful in seasonally damp places in open, rolling grassland but occasional plants can be found in drier situations. They have been recorded from the Midlands of southern KwaZulu-Natal, near Kokstad, through to King William's Town, Eastern Cape, and close to East London where they approach the coast. Although the plants do not depend on fire to flower, they are known to flower most prolifically in the spring (August to October) following winter fires. With few exceptions, the bulbs produce new leaves slightly later in the season.
Despite the subtle nature of the floral features that separate these taxa, they may play a role in attracting a slightly different range of pollinators. This, together with the slightly asynchronous flowering times and their discrete habitats, may impose some level of reproductive isolation upon the taxa. It is apparent, nevertheless, that these morphological, phenological and habitat differ ences reflect a level of discontinuity that warrants more than varietal delimitation, especially as the populations of the two taxa occupy different, albeit partially sympatric, geographical ranges, namely the coast versus the Midlands of southern KwaZulu-Natal and the Eastern Cape. The only places where populations of the two taxa co-occur are where patches of forest extend inland into the temperate grasslands along major river courses. This interpretation, consequently, excludes the coastal, ever green plants with yellow flowers that Dyer (1939) and Reid & Dyer (1984) previously considered to belong to C. mackenii var. cooperi.
Phenology'-, flowering starts in June and usually lasts until September, but can extend into early November. The leaves are evergreen.
Diagnostic features: the plants, commonly known as Ifafa lilies, are clump-forming and produce sev eral leaves which persist throughout the year. The usu ally sweet-smelling flowers have ovate tepals that are patent to rolled back at anthesis and the stamens are characteristically included in the fairly broad perigone throat (Table 1)

Specimens examined
Deciduous, bulbous geophyte, 180-530 mm tall when flowering. Bulb solitary. Leaves developing sequentially, absent at flowering, rarely lor 2 emerging at end of flow ering period, finally up to 5, green or glaucous; margin smooth or minutely rough. Inflorescence solitary or occa sionally 2 per bulb; scape glaucous with a pinkish blush. , suberect to spreading horizontally, creamcoloured to dull yellow, often suffused with apricot-pink on tube; scent spicy, usually bittersweet; perigone tube 25-47 X ± 2 mm diam. at base, widening gradually up to 4.0-5.5 mm diam. at throat; tepals ovate, slightly recurved to rolled back, 3.5-6.0 x 2 .5^.0 mm; outer tepals slightly narrower than inner, with somewhat subacute and shortly mucronate tips; inner tepals with round tips. Stamens included in upper part of perigone tube and throat, outer attached ± 5 mm below throat, inner attached ± 2 mm below throat. Ovary green or pinkish grey.
Phenology: the flowering period which often follows winter fires starts at the end of July at low altitudes (30 m) and peaks in September, but a few individuals con tinue flowering until November at high altitudes (1 800 m). The plants flower in the seasons between fires so are not true fire lilies. The leaves are absent during the main flowering period in September and begin to emerge from scattered individuals in October. Leafing contin ues throughout summer until the onset of the dry season when the bulbs become dormant over winter.
Diagnostic features: the plants are solitary and decidu ous, and the leaves mostly reach maturity after flowering. The perigone tube has a relatively narrow throat with small, ovate tepals (Table 1). The flowers are most often dull yellow but in populations from around Maclear, Umtata, Stutterheim, and King William's Town, the peri gone tube is pinkish cream to apricot. Often the flowers have a spicy scent. Deciduous, bulbous geophyte, 130-360 mm tall when flowering. Bulb hypogeal, ovoidal, 30-40 x 30-35 mm, narrowed to a short neck up to 10 x 7-10 mm; outer tunics brown and papery; inner tunics cream-coloured and fleshy. Leaves developing sequentially, either absent or 1 newly emerging at flowering, 2 or more, linear, at least 100 X up to 3 mm, suberect; abaxial surface with a prominent median vein, keeled; margin and midvein minutely papillate. Inflorescence 2-6-flowered, soli tary per bulb; scape erect, up to 320 x 5 mm (at base), tapering distally up to ± 3 mm at apex, round to slightly compressed in t/s, dull green, hollow; spathe valves 2, outer sheathing inner at base, lanceolate, exceeding pedicels, up to 25 x ± 3.5 mm (at base), membranous, suberect and pinkish when fresh, later reflexing and turning brown; bracteoles several, linear, up to 15 mm long; pedicels suberect to slightly spreading, dull green, up to 12 X ± 1.5 mm at anthesis, lengthening up to 15 mm and becoming more rigid when fruiting. Flowers erect to slightly spreading, opening ± simultaneously, cream-coloured to dull yellow, suffused with pink or green on perigone tube sometimes extending to tepals, with a bittersweet scent; nectar unknown; perigone tube dilated gradually to throat, slightly curved, 33-50 mm long, ± 2 mm diam. at base, widening gradually up to 5-6 mm diam. at throat; tepals narrowly ovate to oblong, slightly spreading, neither reflexed nor rolled back, 7-11 X 3 -^ mm, outer tepals ± as wide as inner tepals, shortly mucronate at apex; inner tepals obtuse. Stamens biseriate, regular; outer stamens included; inner stamens shortly exserted; outer filaments ± 2 mm long, attached ± 3 mm below perigone throat; inner filaments ± 4 mm long, attached in perigone throat; anthers dorsifixed, 2 mm long before dehiscing, dark yellow; pollen yellow. Ovary ellipsoidal, ±5x3 mm, dull green; ovules axile, ± 25 per locule; style usually pressed against upper tepal, slightly shorter than perigone, reaching up to 47 mm; stigmatic branches 3, short, spreading, slender, ± 1 mm long, truncate, shortly papillate towards apex. Capsule unknown.
Phenology, the plants flower from October to December, mostly in advance of the leaves, but in a few individuals one newly emerging leaf may be present. After flowering, the leafing habit remains poorly known.
Diagnostic features: Cyrtanthus ochroleucus is dis tinguished by its hysteranthous leaves which are nar row (± 3 mm wide) and minutely papillate on the mar gin and midvein, by the suberect tubular flowers which are cream-coloured to dull yellow, and by the short (± 2 mm) outer filaments and longer (± 4 mm) inner filaments so that only the inner stamens are exserted from the peri gone throat. The similarities and differences between C ochroleucus and its close allies, C. aureolinus and C mackenii, are summarized in Table 1.
Distribution and habitat: this species is only known from fynbos on the lower northern slopes o f the Langeberg, and on the Albertinia coastal plain. Western Cape (Figure 2). Populations have been found in stony or sandy habitats and, although the bulbs flower after fire, they also flower in the intervals between fires. The plants grow singly and are scattered in open patches.
Nomenclatural note: Herbert (1821) based his origi nal description of Monella ochroleuca Burch, (the basionym of C. ochroleucus) on the herbarium collections of Burchell, but he failed to cite any particular specimen. The original diagnosis o f M. ochroleuca states that it is allied to M odora (Ker Gawl.) Burch, (based on C. odorus Ker Gawl.) but that it is distinguished by having yellow-white flowers. Cyrtanthus odorus itself is a dis tinctive species which is characterized by hysteranthous leaves, suberect, tubular flowers with small tepals and, in contrast to C. ochroleucus, has dark red flowers.
Herbert (1837) later adopted the name C. lutescens for M. ochroleuca, which he explicitly referred to as a syno nym, and he cited Burchell 7144 as the only preserved specimen. There can be little doubt that the diagnosis of M ochroleuca clearly distinguishes it from other taxa and that the name is validly published. Thus C. lutes cens is considered to be nomenclaturally superfluous and is illegitimate. The lectotype of C. ochroleucus that has been designated here is Burchell 7144 (K) which con sists of leafless, flowering bulbs with narrowly tubular flowers that appear to be held upright.