Two New Species, Two Rediscoveries and a Range Extension in Lachenalia (Asparagaceae: Scilloideae) from Southern and Western South Africa

Home , Elytropappus, Freesia, Graham Duncan (botanist), Lachenalia, Lachenalia contaminata

Phytotaxa 316 (3): 261–270 ISSN 1179-3155 (print edition) http://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2017 Magnolia Press Article ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.316.3.5

Two new species, two rediscoveries and a range extension in Lachenalia (Asparagaceae: Scilloideae) from southern and western South Africa

GRAHAM D. DUNCAN South African National Biodiversity Institute, Kirstenbosch National Botanical Garden, Private Bag X7, Claremont, 7735, South Africa; e-mail: [email protected]

Abstract

Two recently discovered species of Lachenalia from South Africa, Lachenalia barbarae from the Overberg district of the Western Cape and Lachenalia adamii from the Bokkeveld escarpment of the Northern Cape, are described. In addition, details of the rediscovery of two species previously thought to be possibly extinct in the wild, Lachenalia martleyi from the Overberg district in the Western Cape and Lachenalia macgregoriorum from the Bokkeveld plateau in the Northern Cape, are provided, as well as a range extension for the critically endangered Lachenalia moniliformis from the Breede River Val- ley in the Western Cape.

Key words: Hyacinthaceae, IUCN evaluation, Liliaceae, narrow African endemics, re-discovered species considered extinct, southern African flora

Introduction

The southern African genus Lachenalia Jacquin in Murray (1784: 314) is endemic to South Africa and Namibia, occurring primarily in the western and southwestern winter rainfall zones of these two countries. High levels of endemism are evident throughout its range, with the main centre of diversity (38 species) around Worcester in the southwestern part of the Western Cape (Duncan 2012). The species are deciduous, and almost all are winter-growing. Many species are adapted to arid conditions, and several are pyrophytes (Duncan & Edwards 2007). Three obvious pollination syndromes occur in the genus, viz. mellitophily, ornithophily and generalist flowers visited by various insects including monkey beetles, blister beetles, hoverflies, march flies, butterflies and day-flying moths (Duncan 2012). A phylogenetic morphological study formed the basis of classification in the 133 species then known, which were placed in two subgenera, Lachenalia and Polyxena (Duncan 2012). Perianth shape was used to subdivide the species of subgenus Lachenalia into five sections (Angustae, Lachenalia, Latae, Oblongae and Urceolatae; Duncan 2012, Duncan et al. 2005). Lachenalia arenicola Duncan & Helme (2014: 297) from the Namaqualand coastal plain was recently described, and with the addition of the two species described here, the total number of species now stands at 136. The genus Lachenalia includes numerous species under threat or facing extinction in the wild, due mainly to the destruction, degradation or alteration of wild habitats by human interference (SANBI 2012). Agricultural expansion for winter cereal crops, housing and industrial development account for the greatest loss, followed by road construction, overgrazing and trampling by livestock, vineyard expansion, alien plant infestation, mining, road widening and eutrophication. Currently, more than 50 Lachenalia species fall within the threat categories of critically endangered, endangered and vulnerable (Duncan 2012). Morphological studies were made from wild and cultivated plants of the type collections of Lachenalia barbarae and L. adamii grown in the bulb nursery at Kirstenbosch National Botanical Garden. Lachenalia martleyi and L. moniliformis were studied in habitat, and L. macgregoriorum was studied from recent habitat photographs. Pressed material in the herbaria BOL and NBG was examined (Thiers 2014). I include the grid-reference system and orthography of geographical names in accordance with Leistner & Morris (1976). Morphological measurements were made from wild and cultivated plants, and details of the seeds were examined using a light microscope.

Accepted by Mark Chase: 20 Jul. 2017; published: 8 Aug. 2017 261 Taxonomy

Lachenalia barbarae G.D.Duncan, sp. nov. (Figs. 1–3). Type:—SOUTH AFRICA. Western Cape: Haarwegskloof Renosterveld Reserve between Bredasdorp and Swellendam (3420AC), scattered solitary plants and small clumps in quartz patch overlaying shale on northwest-facing hillside, between low scrub and moss-covered stones, in full sun, elevation 245 m, 34.335°S, 20.322°W, 14 Dec. 2015, Duncan 669 (holotype: NBG).

This new species differs from L. magentea Duncan (2012: 343) in having suberect, light pink flowers with shorter, elliptical outer tepals 3–5 mm long, included or exserted stamens with spreading, much shorter filaments 3–5 mm long, longer, deep maroon peduncles, 60–165 mm long, shorter pedicels 3–5 mm long, larger, ovoid seeds 1.5–1.6 × 1.1–1.2 mm, longer, adult leaves 180–300 mm long, dying back shortly before flowering begins, prostrate or spreading primary seedling leaves with flat surfaces, ovoid, solitary bulbs and a much later flowering time in December and January. Deciduous, winter-green geophyte 85–260 mm high. Bulb ovoid, 15–22 mm in diam., solitary; tunic multilayered, outer layers dark brown, inner layer[s] translucent white; cataphyll 15 mm long, translucent white below, apex acute, green. Leaves (1) 2(3), linear, 180–300 × 3–6 mm, suberect or recurved, light green, dying back shortly before flowering, upper surface unmarked, lower surface spotted or barred with dull maroon towards base; clasping leaf base mostly subterranean, white, 10–15 mm long; primary seedling leaf linear, prostrate or spreading, brownish maroon. Inflorescence a raceme, 3–10-flowered; peduncle erect or suberect, 60–165 mm long, rigid, deep maroon, glaucous; rachis 25–95 mm long, light maroon in lower half, shading to light pink above; bracts cup-shaped at base of inflorescence, becoming lanceolate above, 1–2 × 1–3 mm, white; pedicels 3–5 mm long, shortest at top of rachis, white or light pink. Perianth zygomorphic, narrowly campanulate, suberect, tube shallowly cup-shaped, 2 mm long, light pink, outer tepals elliptical, 3–5 × 2–3 mm, light pink, apical gibbosity oblong, deep pink, inner tepals obovate, 4.0–5.0 × 3.0–3.5 mm, protruding up to 1 mm beyond outer tepals, light pink, median keel deep pink. Stamens included or exserted, filaments white, 3–5 mm long, anthers oblong, 1 mm long, pollen yellow. Ovary ovoid, light green, 1.1– 2.0 × 1.2–2.0 mm; style straight, white, 3–4 mm long; stigma capitate. Capsule obovoid, 5–6 × 4–6 mm, suberect. Seeds ovoid, 1.5–1.6 × 1.1–1.2 mm, glossy, black; strophiole 0.3–0.4 mm long, ridged. Flowering time: December to January. Etymology:—The specific epithet barbarae honours Mrs Barbara Taylor, mother of Mr Oren Taylor who has made significant financial contributions towards supporting efforts to save renosterveld vegetation from extinction in the Western Cape. Other material examined:—SOUTH AFRICA. Western Cape: Haarwegskloof Renosterveld Reserve (3420AC), quartz patch overlaying shale, 34.338°S, 20.326°W, 5th January 2011, Curtis s.n. (Haarwegskloof Private Herbarium); December 2014, Groenewald s.n. (NBG). Notes:—The first collection of plants was made at Haarwegskloof Renosterveld Reserve on 5 January 2011 by Odette Curtis, Director of the Overberg Renosterveld Conservation Trust. The two specimens collected were in late flowering stage, and she was unaware that they represented a new species. During subsequent flowering seasons, Jannie Groenewald, Manager of the Reserve, considered it to be an undescribed species, since he was unable to identify it from the literature. Following the 2014 flowering season, seeds and a single bulb collected by Groenewald were brought to the bulb collection at Kirstenbosch National Botanical Garden by Rhoda McMaster, a member of the local Custodians of Rare and Endangered Wildflowers (CREW). The bulb flowered in cultivation at Kirstenbosch in early December 2015 and was confirmed to be an undescribed species; a line drawing was prepared by Vicki Thomas. Shortly thereafter, the author visited the site at Haarwegskloof to study the plants in habitat and collect type material. Diagnostic features and affinities:—Lachenalia barbarae is recognised in flower by an erect or suberect, deep maroon, rigid peduncle bearing a raceme of small, light pink, narrowly campanulate, suberect flowers carried on white or light pink pedicels (Figs. 1, 2). It has shallowly cup-shaped perianth tubes and weakly spreading tepals. The inner tepals protrude slightly and have deeper pink median keels, and the outer tepals have prominent deeper pink apical gibbosities. The flowers have short, included or exserted, weakly spreading white stamens, and bracts that are cup- shaped at the base of the inflorescence and lanceolate above. It is further recognised by usually two, or rarely one or three linear, suberect or recurved green leaves with acute apices, unmarked upper surfaces, and lower surfaces that are lightly spotted or barred with dull maroon towards the base. The inflorescence emerges directly after the leaves have withered, and the primary seedling leaf is linear with prostrate or spreading orientation. The fruit is an obovoid capsule containing ovoid, glossy, black seeds with short, ridged strophioles.

262 • Phytotaxa 316 (3) © 2017 Magnolia Press DUNCAN FIGURE 1. Lachenalia barbarae, drawn from Groenewald s.n. A. Flowering plant. B. Single flower in lateral view. C. Single flower in front view. Scale bars: 10 mm. Illustration byVicki Thomas. species, rediscoveries and a range extension in Lachenalia Phytotaxa 316 (3) © 2017 Magnolia Press • 263 FIGURE 2. Lachenalia barbarae, general view of quartz habitat in Eastern Rûens Shale Renosterveld, Haarwegskloof Renosterveld Reserve, southern Cape (A.) and flowering plants (B, C). Photographs by Graham Duncan.

264 • Phytotaxa 316 (3) © 2017 Magnolia Press DUNCAN Lachenalia barbarae is included in subgenus Lachenalia because of its zygomorphic perianth and in section Angustae due its narrowly campanulate perianth shape (Duncan 2012). It appears closely allied to L. magentea from the southern Cape coastal belt. Lachenalia magentea has a similar narrowly campanulate perianth with a shallowly cup-shaped tube, subequal tepals, suberect pedicels and obovoid capsules, but differs in having a larger, cernuous or spreading perianth with longer, white tepals with magenta apical gibbosities and magenta median keels, and longer, shortly to well-exserted, straight filaments. It differs further in having heavily maroon-dotted, light green peduncles and subterete, synanthous adult leaves, terete, erect primary seedling leaves, offset-forming bulbs and globose seeds 1.1–1.2 × 1.3 mm. Lachenalia magentea flowers earlier (August to October) and occurs close to the coast in sandy soils in Agulhas Sand Fynbos and Agulhas and Canca Limestone Fynbos vegetation (Duncan 2012). Distribution and habitat:—Lachenalia barbarae occurs within the Fynbos Biome and is endemic to the Overberg region of the Western Cape, in the east coast Renosterveld Bioregion (Mucina & Rutherford 2006). It is highly localised to a northwest-facing hillside within the Haarwegskloof Renosterveld Reserve between Bredasdorp and Swellendam, at an elevation of 245 m (Fig. 3). The plants occur as scattered individuals or in small groups of up to six plants on quartz patches overlaying shale, in Eastern Rûens Shale Renosterveld, a vegetation type that is critically endangered due to over 80% having been transformed to cultivation (Mucina & Rutherford 2006). The shallowly seated bulbs occur on moss-covered ground between quartz pebbles, stones and rocks in association with the quartz-endemic succulent Drosanthemum asperulum (Salm-Dyck, 1842: 50) Schwantes (1927: 30) (Aizoaceae), the grass Pentameris eriostoma (Nees, 1841: 304) Steudel (1841: 298) (Poaceae), and the renosterveld shrub Elytropappus rhinocerotis (Linnaeus, 1781: 391) Lessing (1832: 344) (Asteraceae). When in flower, L. barbarae is variable in height, ranging from short specimens 85 mm high in exposed situations to relatively tall plants up to 260 mm high, emerging through vegetation. The species flowers late in the season (early December to early January) in mid-summer and is visited by the small, common hairtail butterfly Athene definita (Lycaenidae) and a small, unidentified solitary wasp.

FIGURE 3. Distribution of Lachenalia barbarae (black dot) in southern South Africa and Lachenalia adamii (black star) in western South Africa.

species, rediscoveries and a range extension in Lachenalia Phytotaxa 316 (3) © 2017 Magnolia Press • 265 Lachenalia adamii G.D.Duncan, sp. nov. (Figs. 3–5). Type:—SOUTH AFRICA. Northern Cape: Oorlogskloof Nature Reserve south of Nieuwoudtville (3119AC), occasional on steep river banks east of Kareebos on Rock Pigeon hiking trail, in shale scree, elevation 600 m, 31.480°S, 19.093°E, 11 Sept. 2011, Harrower 4977 (holotype: NBG!; isotypes: NBG!, PRE!).

This new species differs from L. latimeriae Barker (1979: 196) in having a subreclinate raceme of narrowly campanulate flowers with white outer tepals, translucent white and green inner tepals with the lower inner tepal diverging markedly from the two upper ones, weakly spreading filaments, a linear, reclinate or spreading leaf, the development of numerous bulblets around the base of the mother bulb, a broadly ellipsoid capsule, and a larger, matte black seed 1.8 × 1.6 mm with rugose secondary sculpturing and a longer strophiole 0.8 mm long. Deciduous, winter-green geophyte, 130–195 mm high. Bulb ovoid, 12–15 mm in diam., bulblet- forming around base; tunic 2-layered, outer layer dark brown, inner layer thinner, light brown; cataphylls 2, membranous, tightly surrounding leaf base, apices translucent, obtuse, lower cataphyll 5 mm long, light brown, upper cataphyll 15 mm long, white. Leaf solitary, linear, 90–150 × 6–12 mm, spreading, reclinate, conduplicate in lower third, deeply canaliculate in upper two thirds, bright green, base maroon or marked with maroon transverse bands, apex attenuate; clasping base with light brownish purple transverse bands basally, shading to green distally. Inflorescence a subreclinate raceme, 4–20-flowered; peduncle 65–110 mm long, light green; bracts cup-shaped at base of inflorescence, becoming ovate above, 1 × 1–2 mm, translucent white; pedicels suberect, 5–9 mm long, white. Perianth zygomorphic, narrowly campanulate, cernuous, lightly spicy-scented; tube cup-shaped, 2 mm long, white; outer tepals narrowly ovate, 8–9 × 4–5 mm, white, margins straight, apex slightly recurved, apical gibbosity greenish brown; inner tepals obovate, 9–11 × 4–5 mm, weakly spreading when fully open, translucent white, apices recurved, lower tepal 1 mm narrower than upper two tepals and diverging markedly, apices recurved, median keels brownish green. Stamens exserted, filaments weakly spreading, 10–11 mm long, white, anthers oblong, 0.5 mm long, dull red prior to anthesis, pollen yellow. Ovary ovoid, 3 × 2 mm, light green; style declinate, 8–9 mm long, exserted, white; stigma minutely capitate. Capsule broadly ellipsoid, 8–9 × 6–7 mm. Seed globose, 1.8 × 1.6 mm, matte black, secondary sculpturing rugose; strophiole 0.8 mm long, ridged. Flowering time: late September to mid-October. Etymology:—The specific epithet adamii honours Adam Harrower who discovered the species and made the first scientific collection of plants. Notes:—The only known collection of this species was made by Kirstenbosch horticulturist Adam Harrower in the Oorlogskloof Nature Reserve south of Nieuwoudtville in September 2011. The plants were found in a vegetative state, and habitat photographs were taken. Bulbs were cultivated in a pot at Kirstenbosch National Botanical Garden, and when they flowered later that year, it became evident that they represented an undescribed species. Photographs of the flowering plants were taken. The following year a line drawing was prepared by Vicki Thomas, and the type material was pressed. Diagnostic features and affinities:—Lachenalia adamii is recognised by a usually sparsely flowered, subreclinate raceme of cernuous, narrowly campanulate, translucent greenish white flowers with cup-shaped perianth tubes and protruding inner tepals with brownish green median keels. The inner tepals are weakly spreading when fully open, and the lower inner tepal diverges markedly from the two upper inner tepals (Figs. 4, 5). The flowers have exserted, narrowly spreading stamens, prominent, suberect pedicels, and bracts that are cup-shaped at the base of the inflorescence and become ovate above. It is further recognised by a solitary, linear, reclinate or spreading, green leaf, with the base maroon or lightly marked with maroon transverse bands, and the adult bulb forms bulblets basally (Figs. 4, 5). The seeds are globose and matte black with rugose secondary sculpturing and have ridged strophioles. Lachenalia adamii is included in subgenus Lachenalia section Angustae due to its zygomorphic, narrowly campanulate perianth (Duncan 2012). It appears closely allied to L. latimeriae, which is geographically widely separated in the southern Western Cape and southwestern Eastern Cape. Although L. latimeriae is placed in a different section (Oblongae) on account of its oblong-campanulate perianth, the overall appearance of the flowering plant most closely resembles that of L. adamii. Lachenalia latimeriae has a similar cernuous perianth with a cup-shaped tube, ovate outer tepals with greenish brown apical gibbosities, exserted stamens and prominent suberect pedicels; it differs in its 1 or 2, lanceolate, erect inflorescence, light pink to lilac, oblong-campanulate perianth with the lower inner tepal not diverging markedly from the upper inner tepals, and in its more or less straight stamens. It differs further in having obovoid capsules, smaller, glossy black seeds 0.9–1.0 × 1.3–1.4 mm without secondary sculpturing and a shorter strophiole 0.6 mm long. It flowers earlier than L. adamii, July to early September, and occurs in different vegetation types in Western and Eastern Little Karoo, Kouga Sandstone Fynbos and Gamtoos Thicket (Duncan 2012).

266 • Phytotaxa 316 (3) © 2017 Magnolia Press DUNCAN FIGURE 4. Lachenalia adamii, drawn from Harrower 4977. A. Flowering plant with bulblets. B. Single flower in lateral view. C. Dissected flower in lateral view. Scale bars: 10 mm. Illustration by Vicki Thomas.

species, rediscoveries and a range extension in Lachenalia Phytotaxa 316 (3) © 2017 Magnolia Press • 267 FIGURE 5. Lachenalia adamii, view of plants leafing out in shale scree, Oorlogskloof Nature Reserve, northwestern Cape (A.) and flowering plants in cultivation (B.). Photographs by Adam Harrower.

268 • Phytotaxa 316 (3) © 2017 Magnolia Press DUNCAN Distribution and habitat:—Lachenalia adamii occurs within the Fynbos Biome and the Karoo Renosterveld Bioregion and is only known from the type collection east of Kareebos in the Oorlogskloof Nature Reserve south of Nieuwoudtville on the Bokkeveld escarpment (Fig. 3). The plants grow among moss-covered rocks in shale scree in scattered small groups of up to 15 plants on steep, southwest-facing river banks below sandstone cliffs, at an elevation of 600 m, in Vanrhynsdorp Shale Renosterveld. This vegetation type has previously been noted for its lack of endemic geophytes (Mucina & Rutherford 2006), and thus L. adamii appears to be an exception. Associated plant species include the dwarf geophyte Xenoscapa fistulosa (Spreng. ex Klatt 1863: 781) Goldblatt & Manning (1995: 172) (Iridaceae), the dwarf succulents Adromischus hemisphaericus (Linnaeus, 1753: 429) Lemaire (1852: 60) (Crassulaceae) and Crassula tomentosa Thunberg (1778: 333) (Crassulaceae), and the shrub Stachys rugosa Aiton (1789: 303) (Lamiaceae).

Rediscovery of Lachenalia martleyi Material of Lachenalia martleyi G.D.Duncan (2012: 270) was first collected at Lourensford Farm in Somerset West east of Cape Town in October 1929, and in September 1935, a cultivated specimen of unrecorded origin flowered in cultivation in a Stellenbosch garden. These collections formed the basis for the species, which was published in the monograph The genus Lachenalia (Duncan 2012). Since no collections appeared to have been made since 1935, the species was assessed as critically endangered (possibly extinct) in the Red List of South African Plants (von Staden & Duncan 2014). Subsequently, an overlooked, unnamed collection in the Compton Herbarium at Kirstenbosch, made by M.L. Thomas (Thomas s.n. in NBG) on 3 September 1975 near the Theewaterskloof Dam between Grabouw and Villiersdorp was noted by the author in 2013 and identified as L. martleyi. In September 2014, an expedition to the site was undertaken by the author and Elgin residents Peter and Barbara Knox-Shaw, where a small population was found in flower at the exact location where it had been collected by Thomas in 1975 (Duncan 651, NBG). The plants were found in moist stony clay in semi-shade of exotic pines, in remnant Elgin Shale Fynbos (Mucina & Rutherford 2006). Due to the degraded nature of the habitat, lack of official protection and total known population of less than 1000 individuals, its conservation status should be re-assessed as critically endangered.

Rediscovery of Lachenalia macgregoriorum Lachenalia macgregoriorum Barker (1979: 199) was described from plants collected in the area known as Charlies Hoek on the Bokkeveld Plateau near Nieuwoudtville in the Northern Cape on 5 November 1962. Lachenalia macgregoriorum is distinctive in having widely campanulate, maroonish-magenta flowers that appear late in the season, early summer, directly after the glaucous, narrowly lanceolate leaves have withered. For a period of 59 years following its discovery, no further records of the species were made despite several searches. It was thought to have possibly become extinct or that specific environmental conditions were required for it to flower in the wild, and its threat status was assessed as critically endangered (possibly extinct) (Duncan & von Staden 2015). On 26 October 2016, several flowering plants were found near the original site by Eugene Marinus, Curator of the Hantam National Botanical Garden at Nieuwoudtville, while attending to perimeter fence repairs within the grounds of this botanical garden. The plants were isolated specimens on red dolerite flats among dry grasses and low scrubby cover. Since this species flowers regularly in cultivation at Kirstenbosch without any special treatment, it is assumed the short-lived, somewhat cryptic maroonish-magenta perianth colour may have accounted for it having escaped the attention of collectors for so long. As the species is currently only known from one locality and consists of well below 100 individuals, its threat status should be altered to critically endangered.

Range extension for Lachenalia moniliformis Lachenalia moniliformis Barker (1983: 438) was previously known from a single, small population in sandy depressions and stony shale ground in karroid scrub south of Worcester in the Breede River Valley, in the southwestern part of the Western Cape. Due to extensive habitat loss from expanding vineyards and erosion due to overgrazing, its threat status is assessed as critically endangered (Raimondo & Duncan 2011). In September 2013, a new population was discovered in flower by James Deacon 65 km east of the type locality on the Stockwell farm just east of Ashton (Deacon 2851, NBG). At this locality, the plants occur in small numbers on seasonally moist gravelly slopes, in close association with a large colony of L. contaminata Aiton (1789: 460). A subsequent visit to this site by the author and Deacon was made in September 2014. Due to erosion from overgrazing, L. moniliformis is unfortunately also severely threatened at this site, and its threat status should remain as critically endangered.

I thank Odette Curtis, Jannie Groenewald and Adam Harrower for kindly providing habitat information and making the original collections. Rhoda and Cameron McMaster and Barbara and Peter Knox-Shaw provided much appreciated assistance in the field. Vicki Thomas is thanked for the line drawings and Michelle Smith for preparing the map. An anonymous reviewer and the editor made helpful suggestions.

References

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