646. CYLINDROCLINE LORENCEI
Compositae
Plant in Peril, 31
Nicholas Hind, Michèle Sánchez and Carlos Magdalena
Summary. Cylindrocline lorencei A. J. Scott, a ‘Plant in Peril’, is described and
illustrated, and its cultivation requirements provided. The species’ conservation
status is commented on: once declared extinct in the wild, attempts have now
been made to re-introduce the species into Mauritius. The relationships of
Cylindrocline are discussed.
The subject of this plate, the Mauritian Cylindrocline lorencei A. J. Scott
(Compositae: Inuleae) is, like the other species in the genus (Cylindrocline
commersonii Cass.), a botanical curiosity. It attains horticultural merit
with age as an architectural plant that has a good bark structure and
interesting leaves, as well as being relatively pest free (q.v. Hind, 1996).
It does have one point of considerable interest, hence the subtitle of
this paper as a ‘Plant in Peril’, in that it is extinct in the wild. Its
existence in cultivation has provided a success story, both in terms of
its continued propagation and its eventual repatriation to Mauritius,
for the Royal Botanic Gardens, Kew.
Cylindrocline Cass. was placed in the tribe Inuleae Cass. (including the
everlastings, edelweiss and cudweeds) by Cassini (1817). Bentham’s
broad treatment of the Inuleae (Bentham, 1873) included nine subtribes which he considered a well-defined group. Anderberg aimed
to ‘refute this taxonomic misconception’ (Anderberg, 1989, 1991c)
along with Bremer (1987) and, in a short series of papers (Anderberg,
1991a; 1991b; 1991c), sought to divide the Inuleae sensu lato into three
tribes, the Inuleae Cass. sensu stricto, Gnaphalieae (Cass.) Lecoq & Juillet
and the Plucheeae (Benth.) Anderb. As the result of cladistic analyses,
of predominantly morphological characters, Cylindrocline was found to
be ‘sister’ to the rest of the tribe Plucheeae as recognised by Anderberg
(1991c), i.e. basal to the rest of the tribe. The separate tribes were
presented by Bremer (1994). However, matters are slowly turning full
circle, and molecular systematics has proposed that the genera of the
Plucheeae are nested within the Inuleae (cf. Funk et al., 2005; Anderberg
& Eldenäs, 2007), albeit based on a rather poor sampling (c. 14%
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Plate 646 Cylindrocline lorencei
renato moraes
of genera). The more recent analysis of the Inuleae sensu lato (i.e.
including the Plucheeae) has placed Cylindrocline, along with the bulk of
the so-called core genera of the Plucheeae (e.g. Coleocoma F. Muell. – 1
sp., Doellia Sch.Bip. ex Walp. – 2 spp., Epaltes Cass. – c. 14 spp.,
Karelinia Less. – 1 sp., Laggera Sch.Bip. ex Koch, p.p. – 17 spp,
Pluchea Cass. – c. 80 spp., Porphyrostemma Benth. ex Oliv. – 4 spp.,
Pseudoconyza Cuatrec. – 1 sp., Sphaeranthus L. – c. 40 spp., Streptoglossa
Steetz – 1 sp., and Tessaria Ruiz & Pav. – 1 sp.), in a comblike polytomy (Anderberg et al., 2005). None have any particular
horticultural merit. One of the authors (N.H.) still prefers to recognise
a broader concept of the Inuleae (Hind, 2007) within which Cylindrocline
is still placed, recognising the three subtribes Inulinae, Gnaphaliinae Less.
and Plucheinae Benth.
Obvious generic relationships of Cylindrocline have become far
less clear with each new set of analyses. Originally, Cassini (1822)
placed Cylindrocline in his ‘Inulées-Prototype’ in a group of four genera
with paleaceous receptacles – Rhanterium Desf., Molpadia Cass. (=
Buphthalmum L.) and Neurolaena R. Br. (now in the Heliantheae Cass.).
Bentham (1873), in his broad concept of the Inuleae, placed our genus
in the subtribe Filaginae Benth., along with Blepharispermum Wight ex
DC. and Athroisma DC. based upon achene characteristics. Both of
the latter are sometimes placed in the tribe Athroismeae Panero. The
more recent analyses of Anderberg (1991c) and Anderberg et al.,
(2005) suggest no close allies, indicating that the genus should be
treated as somewhat isolated.
In the earlier special issue of Curtis’s Botanical Magazine (Volume 13,
Part 4, 1996) dealing with the Mascarene Islands, Wendy Strahm
provided an assessment of the need for conservation on the islands
(Strahm, 1996). The incredible richness of the flora, in terms of
endemics (73% of flowering plants, 22% of pteridophytes) was emphasised, as well as the number of plants now considered extinct – 80
species on Mauritius alone. In tabulating the ‘top thirty’ threatened
plants of Mauritius’, Cylindrocline lorencei was listed noting, regrettably,
that there was only ‘One shrub which has since died’. It was last
known alive in the Black River Gorge National Park, Mauritius.
The subject of the present plate is one large stock plant that has
been growing at Kew since 2001 in the Temperate Unit of the
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Tropical Nursery; there are several other plants at Kew, from the
original clone and subsequent propagations from our stock plant.
Cultivation. Achenes from the one remaining plant in Plaine
Champagne, Mauritius, were collected by Jean Yves Lesouëf in 1980.
Germination tests in the Conservatoire Botanique National de Brest in
1994 and 1995 were failures. However, newly developed techniques,
to grow the embryos in vitro, resulted in three clones which, when
grown on, produced seeds in 2003. The fact that any live material was
found in the achenes resulted from a series of staining experiments in
Brest. Small clusters of live cells were found in very few seeds when
extracted from the achenes, leading to the production of three clones.
The Royal Botanic Gardens, Kew accession is a donation from 2001
by Conservatoire Botanique National de Brest from one of the clones.
Cylindrocline lorencei is a tender shrub, requiring, for cultivation in
Great Britain, a temperate glasshouse, with a minimum temperature
of 8◦ C. No supplemental lights or humidity are necessary. At The
Royal Botanic Gardens, Kew it is grown in Zone 1 of the Tropical
Nursery. Relative humidity is ambient, and ventilation is improved
by the use of top fans hung over the benches, as well as roof vents.
Temperatures can reach 28–30◦ C in summer. Irrigation water is
treated by a reverse osmosis process, which neutralises pH and
removes dissolved salts.
Good plant growth and health were achieved using well-drained
multipurpose compost based on a loam/coir/Perlite medium with
added slow release fertiliser. Soil should be allowed to almost dry
between watering. Plants do not respond well to water-logging.
Feeding with a well balanced fertiliser (NPK) is carried out every
fortnight. They flower well in summer.
Pests. The plants are relatively pest free, with occasional minimal
occurrence of mealy bug. Mechanical pest control is recommended,
as plants are very sensitive to chemical foliar sprays and are scorched
when chemical sprays are applied.
Propagation. Propagation by seed has been unsuccessful at Kew,
possibly due to a self-incompatibility problem. The seed produced by
selfed plants appears not to be viable. Propagation can be achieved
by tissue culture or cuttings. Cuttings are difficult and slow to root,
sometimes taking up to four months. Successful apical cuttings can
be taken in the spring when they are potted into equal parts of
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Fig. 1. Cylindrocline lorencei. Habit sketch of mature plant cultivated at RBG, Kew. Drawn
by Renato Moraes.
peat/Perlite (70:30) and placed in a cabinet on a heated bench with a
minimum temperature of 20◦ C. As soon as rooted, they are potted on
into a loam/coir/Perlite mix and transferred to a cooler zone. Tissue
culture has proved to be the most efficient method to propagate this
species. Plants were weaned into a coir/Perlite (70:30) mix when
1–2 cm or more in size inside a closed propagator on a heated bench
for 1–2 weeks.
Reintroduction and repatriation. In 2006, micropropagated
plants in jars were repatriated, featuring in a headline article on
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‘Conservation’ in Kew Scientist (Issue 29, April 2006). Unfortunately
the plants failed to establish during the weaning process on Mauritius.
In another attempt, in March 2007, 12 mature plants (about 60 cm
tall) were repatriated by one of the authors (C.M.). This repatriation
was successful and a year later some were still growing at the Native
Plant Propagation Centre at Currepipe, Mauritius. However, the
climatic conditions at this nursery are not ideal for this species, as
Cylindrocline lorencei grew at a higher altitude than the lowland, and
more tropical, Currepipe facility.
Cylindrocline lorencei A. J. Scott in Kew Bull. 42(2): 476 (1987). Type:
‘Mauritius. Plaine Champagne, low thicket heath community, Lorence
15532’.[Plaine Champagne (along road to Bel Ombre Nature Reserve c.
1/ mile from main road). Shrub 5 feet tall in low thicket heath community.
4
Flowers violet mauve, leaves olive green. 15. 1. 73. Lorence 15532.] Holotype:
MAU; isotypes: MAU (see note below), MO.
Illustration: Flore des Mascareignes, 109: 48, pl. 14 (1993).
Description. Treelet or small tree to 2 m. Trunk to c. 7.5 cm diameter
at base, sparsely to moderately well-branched above and densely leafy in
terminal rosettes (these far denser and conspicuous in herbarium material).
Stems at first densely lanate, later becoming glabrescent and conspicuously
marked with scars of fallen leaves, older leaves retained when dead (at least
in cultivated material where they are easily removed by hand but clearly
do not fall). Leaves spiralled, distinctly rosettiform (in the wild) or forming
a very loose rosette towards branch apices or simply leafy upper stems (in
cultivated material), sessile to short-petiolate, petiole 10–15 mm long, lamina
oblanceolate, 120–180 (–200) mm long by (37–) 45–55 ( −60) mm wide, olive
green (to ‘golden tomentose’ – Lorence 2196) and slightly discolorous and lighter
beneath, indumentum above uniform and densely velutinous beneath, areolae
densely pale brown to off-white velutinous but over venation dirtier ambercoloured lanate indumentum with much longer, contorted hairs, all trichomes
eglandular, uniseriate and often conspicuously septate, base cuneate, primary
venation yellowish-green above and beneath, secondary venation insculpate
above, prominent beneath, margins entire, apex acute. Inflorescence terminal but
rapidly overtopped by axillary vegetative growth (in cultivated material), one
to few per shoot per season, peduncle to 14 cm long (usually much shorter
in herbarium material), basal section below first leaf-like bract c. 5.5 cm, all
densely lanate, eventually peduncles pendulous by fibres of retted peduncle.
Capitula heterogamous, disciform, sessile and often forming small glomerules at
apices of inflorescence branches; involucre cylindrical, 12–14 mm tall by c. 5 mm
diameter; phyllaries 5-seriate, 5 mm long by 2 mm wide, midribs often somewhat
keeled, margins conspicuously long-ciliate, apices acute, with tuft of long, often
crisped hairs usually over a lilac (in fresh material) or purplish (in dried material)
apical blotch; receptacle conical, paleaceous, paleae linear-lanceolate, pale brown
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G
M
J
D
H
E
L
F K
A
B
C
Fig. 2. Cylindrocline lorencei. A, capitulum; B, outer phyllary; C, inner phyllary; D, l.s.
capitulum; E, palea; F, marginal female floret; G, disc floret with accompanying palea; H, corolla,
anther cylinder and style of disc floret; J, corolla of disc floret opened out showing attachment of
anther filaments and complete style; K, apices of achene setulae, much enlarged; L, apex of pappus
seta of female floret, much enlarged; M, apex of pappus seta of disc floret, much enlarged. Scale
bars: A, D = 5 mm; B, C, E, F, G, H, J = 1 mm. Drawn by Renato Moraes from living material in
RBG, Kew.
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or straw-coloured and often purplish towards apices, margins long-laciniate.
Marginal florets female, corollas filiform, 3 mm long, short-dentate, sparsely short
stipitate-glandular and glandular-punctate about apex; style base lacking basal
node, narrowing from base upwards, glabrous, exserted, style arms c. 0.8 mm
long, lilac, short-pubescent, hair apices acute. Disc florets hermaphrodite, at first
male later becoming female, corollas purplish, tubular, 3–4 mm long, fused to
apex of achene, 5-lobed, lobes c. 1 mm long, conspicuously glandular outside,
apices obtuse and somewhat thickened; anther cylinder exserted, apical anther
appendages triangular, apices acute, basal anther appendages short-tailed; style
base lacking basal node, but with annular nectary, glabrous, style arms c. 0.8 mm
long, lilac, short-pubescent, hair apices acute. Achenes triquetrous and with a
broader flatter face outside(marginal florets) to cylindrical and c. 8-ribbed, (disc
florets), 3–3.5 mm long, brownish, body sparsely setuliferous on angles towards
apex, setulae of twin-hairs, apices asymmetrical and scarcely divided, one apical
cell conspicuously bigger than other, fine-acute; carpopodium an inconspicuous
narrow basal flange; pappus setae 8–12 (often fewer in marginal florets), usually
appearing uniseriate but clearly biseriate on some disc achenes, 3–4 mm,
setae free, flattened with laciniate margins (originally described as plumose!)
(Figs 1 and 2).
Distribution. Mascarenes: endemic to Mauritius, before it became
extinct, on the Plaine Champagne.
Habitat and Ecology. Growing amongst low ericoid scrub with Philippia
and Helichrysum, on lateritic soil; 700 m.
Flowering time. December to January, and June in Mauritius; June to
July in cultivation, but is somewhat variable.
Conservation status. When Lorence collected the material used by
Scott as the holotype (1973), he noted that there was only one plant in the Plaine
Champagne. Lesuoëf’s collection of fruits in 1980 confirmed the continued
existence of this plant. At this point the species could have been classed as
Endangered. However, Strahm’s note (Strahm, 1996) recorded that the plant
had died: the species at that point would be recorded as Extinct. The plant was
considered extinct in the wild in 1990 yet the ‘1997 Red List’ (Walter & Gillett,
1998) still recorded the species as Endangered. Although a number of plants
have been repatriated to Mauritius it still remains to be seen if these will survive
in the Native Plant Propagation Centre at Currepipe and, if replanted in their
original habitat, will continue to grow in the Plaine Champagne.
Although aspects of the capitulum and its florets have been recorded, nothing
has been recorded of the biology of the developing capitulum. The first author
was alerted to the stock plant flowering by the artist Renato Moraes, who has
painted the subject of this plate. Upon investigation the capitula were found to
go through the following cycle. The outer florets are filiform and female, the
styles a pinkish lilac matched by the apices of the innermost phyllaries; the disc
portion of the capitulum is still green at this point and no other structures are
visible. After a few days the first of the inner, scattered outer disc florets begin
to open in a spiral. These have purplish corollas and are at first male with the
pollen mass being forced out of the top of the anther cylinder as the style grows.
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In some, as the style begins to grow up through the anther cylinder, the anther
cylinder is forced over sideways. The disc florets then gradually become female
as the style grows through the anther cylinder and the style arms separate.
Whilst most of the ‘recent’ collections of Cylindrocline lorencei have been made
since the mid-1970s, the earlier collections date from Blackburn’s in the 1860s,
Bojer’s from the 1830s, and apparently the earliest, that of Du Petit-Thouars
from the 1790s. It is likely that both the two latter collections were mixtures of
C. commersonii and C. lorencei, vide Du Petit-Thouars s.n., s.loc. (P).
It is worth drawing attention to the protologue statement by Scott (1987),
in the material examined, that the holotype was ‘MAU, 2 sheets’. Many years
ago the first author examined these sheets when they were photographed for
record. It is quite clear that only one sheet is actually labelled as the holotype
and that the other is a similar sized specimen merely determined by A. J. Scott;
its type status was not indicated. This second specimen is merely a duplicate
and not indicated as part of the holotype; it is to be counted as an isotype. This
second specimen possesses a habit sketch drawing, by Lorence, pinned to the
sheet, together with a small portion of bark from the trunk.
Key to the species of CYLINDROCLINE (translated from Scott, 1993)
Leaves narrowly oblanceolate, to 14 cm long, ± discolorous and slightly paler
beneath, apices acute; peduncles 1 to few, to 14 cm long . . . . . . . . . C. lorencei
Leaves spathulate, oblanceolate to obovate, to 7 cm, conspicuously discolorous
and white woolly beneath, apices obtuse or broadly acute; peduncles few to
numerous, to 6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. commersonii
Acknowledgements. The authors would like to thank Renato Moraes, the
18th Artist Scholar of the Margaret Mee Fellowship Programme, for producing
the colour plate, text habit figure and the black and white dissections, whilst
under the tutelage of Christabel King during his scholarship in 2006.
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