PhytoKeys 5: 5–19 (2011)
A peer-reviewed open-access journal
The identity of Albuca caudata Jacq. (Hyacinthaceae)
doi: 10.3897/phytokeys.5.1166
5
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The identity of Albuca caudata Jacq. (Hyacinthaceae)
and a description of a new related species: A. bakeri
Mario Martínez-Azorín1,2,3, Manuel B. Crespo1, Anthony P. Dold2 , Nigel P. Barker3
1 CIBIO (Instituto Universitario de la Biodiversidad), Universidad de Alicante, P.O. Box 99, E-03080 Alicante, Spain 2 Selmar Schonland Herbarium, Department of Botany, Rhodes University, Grahamstown 6140,
South Africa 3 Department of Botany, Rhodes University, Grahamstown, 6140, South Africa
Corresponding author: Mario Martínez-Azorín (mmartinez@ua.es)
Academic editor: W. John Kress | Received 24 February 2010 | Accepted 13 June 2010 | Published 27 July 2011
Citation: Martínez-Azorín M, Crespo MB, Dold AP, Barker NP (2011) The identity of Albuca caudata Jacq.
(Hyacinthaceae) and a description of a new related species: A. bakeri. PhytoKeys 5: 5–19. doi: 10.3897/phytokeys.5.1166
Abstract
The name Albuca caudata Jacq. has been widely misunderstood or even ignored since its description in
1791. After studying herbarium specimens and living populations in South Africa, plants fitting Jacquin´s
concept of that species are found to be widely distributed in the Eastern Cape, mainly in the Albany
centre of Endemism. Furthermore, some divergent specimens matching Baker´s concept of A. caudata are
described as a new related species: A. bakeri. Data on typification, morphology, ecology, and distribution
are reported for both taxa. Affinities and divergences with other close allies are also discussed.
Keywords
Albuca bakeri sp. nov., Albuca caudata, distribution, taxonomy, typification
Introduction
The genus Albuca L. is accepted to include about 60 species in recent treatments (cf.
Speta 1998; Manning et al. 2002), although up to 131 when considered in a wider
sense (Phillips 1951), with over 180 names available in its traditional sense (cf. IPNI
2011). Recent molecular works have however considered the circumscription of Albuca in quite different ways. On the one hand, Manning et al. (2009) enlarged the tradi-
Copyright Mario Martínez-Azorín et al. This is an open access article distributed under the terms of the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Mario Martínez-Azorín et al. / PhytoKeys 5: 5–19 (2011)
tional concept of Albuca to include other genera such as Stellarioides Medik., Coilonox
Raf., Trimelopter Raf. and Battandiera Maire, reaching a total of 110–180 species, and
hence being very heterogeneous in morphology. On the other, Martínez-Azorín et al.
(2011) recovered the traditional concept of Albuca on the basis of phylogenetic, morphologic and biogeographic evidences.
Taxa of Albuca in its traditional sense (cf. Martínez-Azorín et al. 2011) are distributed mainly in southern and eastern Africa, with only a few species extending to Ethiopia, Saudi Arabia and north of Chad and Nigeria. The only comprehensive revision
of Albuca is that of Baker (1897, 1898), who focused on South African and tropical
African taxa. Recent accounts (cf. Müller-Doblies 1994, 1995, 2006; Manning and
Goldblatt 2009) have greatly increased the knowledge of groups with nodding-flowers,
namely A. subg. Albuca and A. subg. Falconera (Salisb.) Baker. However, information
on groups with erect-flowers, e.g. A. subg. Mitrotepalum U. Müll.-Doblies (= A. sect.
Branciona (Salisb.) J.C. Manning & Goldblatt), has remained almost unchanged from
the late 19th century, and is the focus of our current research (Martínez-Azorín et al.
in press a, b). In addition, no identification keys are available for all currently accepted
taxa of Albuca, most of which are still poorly known or in need of further studies (cf.
Phillips 1926; Dyer 1947). For instance, Albuca caudata Jacq., a plant described from
the Cape, has long been misinterpreted or even ignored. The study of herbarium specimens as well as natural populations of wild plants has revealed the existence of different biological entities to which the name A. caudata has been applied. In the present
contribution, the name A. caudata is typified to warrant further usage in the sense it
was originally published, and data are also presented to describe a new species to which
that name was applied erroneously.
Materials and methods
Herbarium specimens from the following herbaria were studied: BOL, BNRH, GRA,
J, K, KEI, KMG, NBG, NH, NU, PEU, PUC, UFH, WIND (acronyms according to
Thiers 2011). Moreover, a detailed morphological study of both taxa was undertaken
based on plants from natural populations. Authors of the cited taxa follow IPNI (2011).
results and discussion
Albuca caudata was described by Jacquin (1791) and later illustrated (Jacquin 1795) (Fig.
1) after plants collected in “Promontorio bonae Spei”, which “Apud nos in caldariis floret
Decembri & Januario”. The characters mentioned by Jacquin (1791) are: bulb roundish to
ovate; leaves scarce, glabrous, linear-lanceolate, canaliculate, acute, with terete apex; stem
weak and inclined; raceme lax, with patent pedicels; tepals white with green bands, the
inner tepals with yellowish apices; all six stamens bearing anthers; and style clavate, thick
and trigonous. Plants fitting this description are widespread in the Eastern Cape. They are
The identity of Albuca caudata Jacq. (Hyacinthaceae)
Figure 1. Albuca caudata Jacq. from Jacquin, Ic. Pl. Rar. 2(16): 20, t. 442. 1795.
7
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Mario Martínez-Azorín et al. / PhytoKeys 5: 5–19 (2011)
usually easy to recognise by their solitary partially hypogeal bulbs, covered by short, papery, brown to grey scales that reach different heights, and their inclined unilateral racemes
with all pedicels erect. However, a wide range in variation of vegetative and reproductive
features can be observed, such as morphology of the scales of the bulb neck, leaf length
and width, and flower disposition with regard to the inflorescence axis. A number of herbarium specimens labelled A. caudata differ significantly from Jacquin’s description, but
match Baker’s concept of that taxon (cf. Baker 1869). These plants are characterized by
their hypogeal roundish bulbs with fleshy outer tunics, all of them reaching the top of the
bulb and ending into a long epigeal neck surrounded by transversally banded cataphylls.
The particular characteristics of those collections, which were illustrated by Baker (1869)
(Fig. 2), do not fit any of the known species of A. subg. Mitrotepalum. As differences with
the typical A. caudata are remarkable, segregation at the species rank is here favoured and
a new species is described for them. Information on both taxa is provided below.
Albuca caudata Jacq., Collectanea 4: 203 (1791).
http://species-id.net/wiki/Albuca_caudata
Neotype (here designated). Jacquin, Ic. Pl. Rar. 2(16): 20, t. 442 (1795), ex Promontorio bonae Spei. Apud nos in caldariis floret Decembri & Januario (Fig. 1).
Epitype (here designated). SOUTH AFRICA. Eastern Cape: Alexandria, Addo
National Park, 400 feet, 29.X.1954, S.M. Johnson 1077 (GRA).
Description. Evergreen bulbous plants. Bulb mostly solitary and hypogeal, ovoid to
oblong, up to 10 × 6 cm, usually with its wide basal plate elongated into a domed axis
where the fleshy scales are attached; tunics fleshy, short and usually not reaching the top
of the bulb, imbricate, persistent, papery grey or brownish in the upper part, sometimes
with transversal prominent dark ridges, giving a brownish multiscaled appearance to the
bulb in outline. Roots fleshy, thick and usually tuberose, white, numerous, up to 200 ×
4 mm. Leaves 4-10, disposed in an apical rosette, linear-lanceolate, 15-120 × 0.5-2 cm,
straight up and curving down when old, infolded, canaliculate, persistent, pale bright
green to glaucous, glabrous, usually minutely papillate on nerves and margins, with a
terete apex evident in young leaves. Inflorescence inclined, unilateral raceme, 11–40 cm
long; peduncle 12–55 cm long; pedicels 3.5–9 cm long at base becoming smaller, up
to 0.2–1 cm long near top, patent and being usually all erect; bracts ovate-lanceolate to
triangular, long acuminate, 11–25 × 5–9 mm, papery white with brownish distant nerves
that converge at the tips, much shorter than pedicels in the lower part of the inflorescence.
Flowers erect; tepals white with a green median stripe 2–4 mm wide, sometimes with the
tips yellowish; outer tepals oblong, 18–28 × 4–7 mm, apex slightly cucullate; inner tepals
ovate, 15–24 × 4–10 mm, with apex strongly cucullate. Stamens all six bearing fertile
anthers; outer anthers 1.5–2.5 mm long; inner anthers 3–4 mm long; outer filaments
10–16 × 1.5–2 mm, linear lanceolate to narrowly oblong, not pinched down; inner filaments 10–17 × 1.5–3 mm, linear oblong, wider and pinched in the lower half. Ovary oblong to obovate, up to 6–8 × 2–3.5 mm, stipitate, with prominent paraseptal crests that
The identity of Albuca caudata Jacq. (Hyacinthaceae)
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Figure 2. Albuca bakeri Mart.-Azorín & M.B. Crespo from Baker, Refug. Bot. (Saunders) vol. 1, t. 45.
1869 (as A. caudata).
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Mario Martínez-Azorín et al. / PhytoKeys 5: 5–19 (2011)
are divergent in the lower part and form three prominent ridges; style subobpyramidal or
clavate, trigonous, up to 7–10 × 2 mm, stigma yellowish green. Capsule ovate, 14–20 ×
10–14 mm, trigonous to subsphaerical in section, pale-brown when mature; valves splitting in the upper quarter. Seeds flat, c. 5–6 × 4–5 mm, dark brown to black, flattened and
semidiscoidal, biseriate and horizontally stacked in each locule. (Fig. 3)
Flowering time. September to November; capsules dehiscing at the end of November and December.
Habitat. Plants of A. caudata are often associated with bush-clumps, where the
inclined inflorescence is supported by woody plants.
Distribution. Currently known from Addo in the west to Grahamstown in the
east, below 600 m, with an outlying population as far inland as Somerset East, reaching 900 m (Fig. 4).
Diagnostic characters. Albuca caudata can be easily identified by its bulb mostly
solitary covered by brownish papery scales usually disposed at different heights and bearing long thick tuberose roots, its long and narrow canaliculate or infolded leaves, its
inclined raceme, with usually all pedicels patent and erect, giving a unilateral appearance
to the inflorescence, and its white erect flowers with a median green stripe (Fig. 3).
Etymology. The specific epithet ‘caudata’ presumably refers to the rather pointed,
tail-like leaves, although Jacquin did not specifically mention it (E.E.A. Gledhill, unpubl. ms. in NBG).
Relationships. The recently described Albuca batteniana Hilliard & B.L. Burtt
(Hilliard and Burtt 1985) shares some morphological characters with A. caudata, such
as the inclined scape bearing a horizontally arcuate inflorescence with erect pedicels,
and the flower morphology. This species, however, differs in the coriaceous recurved
much broader and flattened leaves, the longer tepals (30–42 mm long), and the structure of the bulb, being proliferous, epigeal, and composed by scales truncate at the top,
disposed into a long domed axis and ending at different heights, without membranous
neck (Table 1).
Observations. A. caudata shows some variability in the colour of the membranous bulb scales, being pale coloured with orange transversal ridges in some inland
populations whilst those from the coastal areas are usually brown coloured with darker
transversal ridges.
Selected specimens studied. SOUTH AFRICA. Alexandria, 4 miles east of Sandflats, 1000 feet, 17/12/1953, E.E.A. Archibald 5431 (GRA); Eastern Cape, along road
in Springs Reserve, north of Uitenhage, 39 m, 21.X.2009, A.B. Low 16732 (GRA);
Addo Elephant National Park, in main Botanical Reserve, 20.X.1996, K. Johnson 241
(GRA); Alexandria, Nanaga, opposite Glen Rosa turn-off, 1100 feet, 23.X.1953, E.E.A.
Archibald 5315 (GRA); Eastern Cape, Albany, Queen´s Road, 10 miles north of Grahamstown, 2000 feet, 05.X.1953, S. Johnson 774 (GRA); Eastern Cape, Albany, a few
yards from Archibald 5636, Pluto´s Vale, 2000 feet, 22.IX.1954, E.E.A. Archibald 5636
(GRA); Eastern Cape, Grahamstown, c. 5 miles on Cradock road, 626 m, 11.XII.2009,
M. Martínez-Azorín & A.P. Dold 85 (GRA); Eastern Cape, Redhouse, thicket west of
village, 6 m, 27.XI.2009, M. Martínez-Azorín, A.P. Dold & A. Martínez-Soler 45 (GRA).
The identity of Albuca caudata Jacq. (Hyacinthaceae)
11
Figure 3. Albuca caudata Jacq. Eastern Cape, Redhouse (M. Martínez-Azorín, A.P. Dold & A. MartínezSoler 45 GRA) a Inflorescence b Outer tepals c Inner tepals d Outer stamen e Inner stamen f Ovary,
lateral views g Mature capsule h Dehiscing capsule i Bulb and leaves j Bulb in longitudinal section k Bulb
with tuberose roots. Scales 1 cm.
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Mario Martínez-Azorín et al. / PhytoKeys 5: 5–19 (2011)
Figure 4. Known distribution of A. caudata Jacq.
Albuca bakeri Mart.-Azorín & M.B. Crespo, sp. nov.
urn:lsid:ipni.org:names:77112770-1
http://species-id.net/wiki/Albuca_bakeri
Holotype. SOUTH AFRICA. Eastern Cape: North of Grahamstown, on Cradock
Road turn off to Kwandwe, 592 m, 05.IX.2010, 33°12'39"S, 26°24'07"E, M. Martínez-Azorín & A. Martínez-Soler 218 (GRA Holo.; ABH, K, NBG, PRE Iso.).
Diagnosis: Species insignis ex Albuca subg. Mitrotepalum characteribus floralibus
ad Albucam caudatam accedit, sed valde differt et facile distinguitur bulbo hypogaeo
solitario carnoso tunicis omnibus apicem attingentes in collum angustum supra solum
desinentes, e basi cataphyllis albido-membranosis manifeste transversaliter fusco-striatis obtectum qui habitum pulchre zebrinum exhibent, insuper racemo subdeltoideo
non secundo floribus spiraliter dispositis.
Illustrations: Baker (1869) in Refugium Botanicum, vol. 1, tab. 45 (Fig. 2); Fig. 5.
Description. Evergreen or deciduous bulbous plants. Bulb mostly solitary, occasionally growing in small clumps, hypogeal, ovoid to spherical, 3.2–7 × 2.5–6.5 cm,
with soft outer tunics that are pale and fleshy, ending in a long epigeal neck, up to
10 × 2 cm, covered with whitish open and sheathing membranous cataphylls bearing
transversal sinuous ridges with their lower side pale to dark brown coloured, giving
a zebra banding horizontal pattern; tunics fleshy, whitish, all reaching the top of the
bulb, concentrically arranged. Roots fleshy, narrow, white, up to 90 × 2 mm. Leaves
The identity of Albuca caudata Jacq. (Hyacinthaceae)
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Table 1. Main diagnostic characters among Albuca caudata, A. bakeri and A. batteniana.
A. batteniana
Proliferous
Narrowly oblong
Outer tunics somewhat
coriaceous,
green to brown
Mostly hypogeal
Hypogeal
Mostly epigeal
Imbricate scales mostly
All scales reaching the top of Imbricate scales ending at
ending at different heights
the bulb
different heights
Neck absent or short and
Neck long and thin, covered Neck usually absent
thick, covered by brown to
by transversally banded
grey membranous scales
sheathing cataphylls
Roots
Usually numerous, thick and Thin and scarce
Thin or slightly thickened
tuberose
Leaves
Narrow, infolded and
Narrow, infolded and
Wide, flattened and
canaliculate
canaliculate
usually recurved
Inflorescence Inclined and secund
Erect and helicoidal
Inclined and secund
Outer tepals 18–28 mm
19–23 mm
30–42 mm
Seeds
5–6 × 4–5 mm
4–5 × 3–4 mm
5–7 × 4–5 mm
Bulb
A. caudata
Mostly solitary
Ovoid to oblong
Outer tunics membranous,
brown to grey
A. bakeri
Mostly solitary
Ovoid to spherical
Outer tunics fleshy,
white to yellow
2-6, disposed in an apical rosette, linear-lanceolate to oblong, 9-40 × 0.4-1.3 cm, erect
when young and later curving downwards, infolded, canaliculate, persistent or usually
deciduous, pale bright green to glaucous, glabrous, usually minutely papillate on nerves
and margins, exceptionally with long papillate margins. Inflorescence an erect raceme
or subcorymb, 3–15 cm long; peduncle 9–22 cm long; pedicels helicoidally disposed,
3–7.5 cm long, longer at the base, up to 0.2–0.7 cm long near top, erect-patent; bracts
ovate-lanceolate to triangular, long acuminate, 9–27 × 4–10 mm, papery white with
brownish separated nerves that converge at the tips, much shorter than pedicels at least
in the lower part of the inflorescence. Flowers erect; tepals white with a green median
stripe 2–3 mm wide, sometimes with the tips yellowish; outer tepals lanceolate-oblong,
19–23 × 5–7 mm, with apex slightly cucullate; inner tepals ovate, 13–17 × 6–7 mm,
with apex strongly cucullate. Stamens all six bearing fertile anthers; outer anthers 1.5–3
mm long, inner anthers 4–6 mm long; outer filaments 10–13.5 × 1.5–2 mm, linear
lanceolate to narrowly oblong, not pinched down; inner filaments 10.5–14.5 × 2–3.5
mm, linear oblong, wider and pinched in the lower half. Ovary oblong to obovate, up
to 6–7 × 2–3.5 mm, stipitate, with prominent paraseptal crests that are divergent in the
lower part and form three prominent ridges; style subobpyramidal or clavate, trigonous,
up to 7–11 × 3.5–4.5 mm, stigma yellowish green. Capsule ovate, 14–16 × 11–12 mm,
trigonous to subsphaerical in section, pale-brown when mature; valves splitting in the
upper quarter. Seeds flat, c. 4–5 × 3–4 mm, dark brown to black, flattened and semidiscoidal, biseriate and horizontally stacked in each locule. (Fig. 5)
Flowering time. July to September; capsules dehiscing at the end of September
and November.
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Figure 5. Albuca bakeri Mart.-Azorín & M.B. Crespo. North of Grahamstown, turn off to Kwandwe
(holotype: M. Martínez-Azorín & A. Martínez-Soler 218 GRA) a Plant b Flower c Outer tepals d Inner
tepals e Outer stamen f Inner stamen g–h Ovary, lateral views i Mature capsule j Capsule, longitudinal
section k Capsule, transversal section l Bulb neck with membranous banded cataphylls. Scales 1 cm.
The identity of Albuca caudata Jacq. (Hyacinthaceae)
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Figure 6. Known distribution of A. bakeri Mart.-Azorín & M.B. Crespo.
Habitat. Albuca bakeri is found growing singly in dry, stony, open ground at low
altitude reaching c. 650 m.
Distribution. from Jansenville to Alice and the Keiskamma river in the Eastern Cape,
with two outlying populations near Calitzdorp in the Western Cape karroo (Fig. 6).
Diagnostic characters. Albuca bakeri can be easily identified by its solitary hypogeal fleshy bulb ending in an epigeal neck, covered by whitish transversally banded
membranous cataphylls, giving a conspicuous zebra banding pattern (Fig. 5). Moreover, its erect and helicoidal raceme with white and green erect flowers, and the smaller
seeds (c. 4–5 × 3–4 mm), separate it from A. caudata.
Etymology. Name honouring John Gilbert Baker (1834–1920), a leading expert
on monocotyledons, who worked at the Royal Botanic Gardens, Kew, and was the
keeper of the herbarium K.
Relationships. No other Albuca with erect flowers have been described with the
characteristic long, thin, zebra banded bulb neck of A. bakeri. The closest species
appears to be A. caudata, though the structure of the bulb and inflorescence clearly
distinguish them (Table 1).
Observations. The peculiar zebra banded cataphylls of A. bakeri are similar to
those found in some other groups of Hyacinthaceae. As pointed out by Müller-Doblies
and Müller-Doblies (1981), zebrine cataphylls are present in evolutive distant taxa
such as Rhadamanthus fasciatus B. Nord., Tenicroa exuviata (Jacq.) Speta (Urgineoideae), several species of Ledebouria Roth (Scilloideae), Coilonox zebrinum (Baker) Speta
and some species of Nicipe Raf. [= Ornithogalum sect. Vaginaspasia U. Müll.-Doblies
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Mario Martínez-Azorín et al. / PhytoKeys 5: 5–19 (2011)
& D. Müll.-Doblies] (Ornithogaloideae). Moreover, Stellarioides arida (Oberm.) Speta
and Battandiera stapffii (Schinz) Mart.-Azorín, M.B.Crespo & Juan show the neck of
the bulb covered with membranose transversally banded cathaphylls, indicating that
the zebrine cataphylls could have evolved independently in at least five lineages of the
Ornithogaloideae (e.g. Albuca, Battandiera, Coilonox, Nicipe and Stellarioides), possibly
as a result of convergent evolution in dry climates of southern Africa.
Some morphological variation has been found within A. bakeri. Some individuals
from Janseville and Port Elizabeth have a slightly setose bulb neck with the characteristic transversally banded membranous cataphylls of this species. Other specimens
from Grahamstown and Port Elizabeth showed somewhat proliferous bulbs, resulting
in a small clump of plants growing together, and with shorter scales not so markedly
banded.
When Baker (1897) described and illustrated his concept of A. caudata (Fig. 2), he
mentioned: “Bulb two to three inches thick, round or oblong, crowned as in the preceding [A. fastigiata Dryand.] with brown fibres. Leaves about a foot long, four lines
broad, more rigid than in the preceding, clasping the stem at the base and more or less
concave on the face upwards, and keeled on the back”. This description is vague and
inaccurate, since the illustration he presented did not show “fibres” at all, and no specific comments on the transversal banding of the upper scales of the bulb were made.
However, when Baker reconsidered A. caudata in later works, his previous concept was
changed to “Bulbus globosus 2–3 poll. crassus viridis apice squamosus” (Baker 1872),
or “bulb globose, 2–3 in. diam.; tunics not splitting into fibres at the top” (Baker
1897), or “Bulb globose, 2–3 in. diam.” (Baker 1898).
Materials studied. SOUTH AFRICA. Eastern Cape, Alexandria, 1½ miles east
of Paterson, 1000 feet, 24.VIII.1953, E.E.A. Archibald 5972 (GRA); Eastern Cape,
Alexandria, south-west end of Zuurkop, Addo National Park, 1000 feet, 23.IX.1953,
S.M. Johnson 751 (GRA); Eastern Cape, Albany, 5 miles north of Alicedale, on Riebeck East road, 1500 feet, 21.IX.1954, E.E.A. Archibald 5638 (GRA); Victoria East:
Alice, dry stony places on Sandilis Kop on north east side, 08.IX.1934, M.H. Giffen
614 (GRA); Victoria East: Alice, Sandilis Kop western side among grass, 13.IX.1935,
M.H. Giffen 618 (GRA); Hillside, Gowie´s Kloof, Grahamstown, IX.1947, Hill
s.n. (GRA); Grahamstown, West Hill, Pine plantation, VIII.1956, V. van Niekerk
s.n. (GRA); Cradock road, Grahamstown, 01.IX.1945, E. Barrat 28 (GRA); In graminosis prope Grahamstown, M. Daly & M. Sole 316 (BOL); In graminosis prope
Grahamstown, 2000 feet, VIII.1893, Schonland s.n. (NBG); Grahamstown (3326
BC): Ecca Reserve, south near old Queens Road/Quarry, 20.VIII.1992, T. Dold
153 (GRA); Leander Beacon, VIII.1943, L. Miles s.n. (GRA); Port Elizabeth, Summerstrand, grassy roadside, IX-X.1990, H.J. Vanderplank s.n. (GRA); Port Elizabeth
(3325CD): 3 km south of Uitenhage towards van Stadens, 01.IV.1978, P.L. Perry
601 (NBG); Port Elizabeth (3325CB): Kirkwood District, farm Brakleegte, 300 m,
28.VIII.1985, M.T. Hoffman 1064, 1065 (NBG); ibidem, 14.IX.1985, M.T. Hoffman
1002 (NBG); Graaff-Reinet (3224DC): District Janseville, just south of Janseville
(+/- 1 km) in municipal-owned land, 11.VIII.1985, M.T. Hoffman 1063 (NBG);
The identity of Albuca caudata Jacq. (Hyacinthaceae)
17
Ladismith (3321BC): Calitzdorp dam, 22.II.1981, P.L. Perry 1521 (NBG); Ladismith (3321CC): Sopieshoogte, north entrance to Garcia´s Pass, Riversdale, 1600
feet, 15.IX.1981, A. Fellingham 149 (NBG); Eastern Cape, Grahamstown, hills above
Botanic Garden, 591 m, 14.XI.2009, 33°19’04”S, 26°31’15”E, M. Martínez-Azorín
& A. Martínez-Soler 12 (GRA); Eastern Cape, Grahamstown, Burnkraal, 649 m,
24.XI.2009, 33°16'40"S, 26°29'41"E, M. Martínez-Azorín & A.P. Dold 34 (GRA);
Eastern Cape, Redhouse, thicket west of village, 6 m, 27.XI.2009, 33°50'01"S,
25°33'56"E, M. Martínez-Azorín, A.P. Dold & A. Martínez-Soler 44 (GRA); Eastern
Cape, north of Grahamstown, Table Hill farm, 587 m, 11.XII.2009, 33°15'21"S,
26°27'17"E, M. Martínez-Azorín & A.P. Dold 83 (GRA); Eastern Cape, north of Grahamstown, on Cradock road turn off to Kwandwe, 594 m, 31.I.2010, 33°12'38"S,
26°24'07"E, M. Martínez-Azorín, M.B. Crespo & A. Martínez-Soler 118 (GRA);
Eastern Cape, Quamnyana, between Breakfast Vlei and Commitees Drift, 411 m,
14.VIII.2010, 33°06'56"S, 22°55'57"E, C. Peter (GRA); ibidem, 27.VIII.2010, M.
Martínez-Azorín & A.P. Dold 207 (GRA); Eastern Cape, Port Elizabeth, Settler´s
Park, 28 m, 03.IX.2010, 33°58'21"S, 25°36'09"E, M. Martínez-Azorín & A.P. Dold
210 (GRA); Eastern Cape, Grahamstown, Sunny side, Hillsview street, 570 m,
07.IX.2010, 33°19'08"S, 26°32'00"E, M. Martínez-Azorín & A. Martínez-Soler 221
(GRA); Eastern Cape, Alicedale, railway cross to Burchell Game Reserve, 288 m,
14.IX.2010, 33°18'51"S, 26°06'01"E, M. Martínez-Azorín & A. Martínez-Soler 226
(GRA); Eastern Cape, Alice, Fort Hare University, Sandili's Kop, 582 m, 17.IX.2010,
32°47'02"S, 26°51'38"E, M. Martínez-Azorín & A. Martínez-Soler 235 (GRA); Eastern Cape, Keiskamma River, Linedrift, 141 m, 12.XI.2010, 33°04'29"S, 27°13'02"E,
M. Martínez-Azorín, A.P. Dold & A. Martínez-Soler 525 (GRA).
Acknowledgements
This work was supported by the Fundación Ramón Areces (Spain). We thank the
curators of the herbaria who provided access to specimens examined and the Cacadu
Department of Economic Development and Environment Affairs for permission to
collect herbarium specimens. Thanks to Cameron McMaster for providing additional
images of A. batteniana.
references
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