KEW BULLETIN (2016) 71:55
DOI 10.1007/S12225-016-9673-2
ISSN: 0075-5974 (print)
ISSN: 1874-933X (electronic)
Inversodicraea Engl. resurrected and I. pepehabai sp. nov.
(Podostemaceae), a submontane forest species from the Republic
of Guinea
Martin Cheek1 & Pepe Haba2,3
Summary. Inversodicraea, shown by previous workers to merit resurrection, is restored here, to include all those
species previously included in Ledermanniella subg. Phyllosma. The final seven species remaining in subg. Phyllosma
are formally transferred to Inversodicraea as I. mortonii, I. gabonensis, I. boumiensis, I. harrisii, I. paulsitae, I. torrei and
I. thollonii. A new species from the submontane forest of Mt Ziama, in Guinea-Conakry is formally described as
I. pepehabai. Similar to I. adamesii, it is assessed as Endangered according to the IUCN 2012 standard.
Key Words. Conservation, forest, Ledermanniella, rheophyte, Ziama.
Introduction
Podostemaceae are a pantropical family of annual or
perennial herbs. All species of the family are restricted
to rocks in rapids and waterfalls of clear-water rivers,
and are therefore rheophytes. However this very
habitat is being increasingly exploited for hydropower
at some risk to the survival of the Podostemaceae they
contain (Schenk et al. 2015; Cheek et al. 2015; Cheek &
Ameka 2016). Most of the African species of
Podostemaceae are narrow endemics, many being
species known from only a single waterfall. New
discoveries of species are still being made frequently
(Rial 2002; Cheek 2003; Schenk & Thomas 2004;
Beentje 2005; Kita et al. 2008; Schenk et al. 2015;
Cheek et al. 2015; Cheek & Ameka 2016). Important
characters in defining genera in Podostemaceae are
the position of the flower in the unruptured
spathellum, and the shape, and sculpture of the ovary.
At species level, important characters are the shape
and relative proportions of spathellae, stigmas, anthers, filaments, gynophores, pedicels, and leaves.
The current generic classification of African
Podostemaceae is based on the framework established
by Cusset (1973, 1974, 1978, 1983, 1987). This work
has been compiled and updated by Rutishauser
(2004). Recently, combined morphological and molecular phylogenetic studies of African Podostemaceae
have shown that Ledermanniella (as delimited by
Cusset) is paraphyletic, including all other sampled
genera of Podostemaceae recognised in Africa (Thiv
et al. 2009, employing plastid markers matK, trnD-trnT,
rpoB-trnC in sampling 9 genera and 17 species of
African Podostemaceae, and Schenk et al. (2015),
employing plastid markers matK, trnL, rpoB-trnC, ndhF,
rbcL and matR in sampling 10 genera and 27 species of
African Podostemaceae).
Materials & Methods
Nomenclatural changes were made according to the
Code (McNeil et al. 2012). Herbarium material was
examined with a Leica Wild M8 dissecting binocular
microscope fitted with an eyepiece graticule measuring in units of 0.025 mm at maximum magnification.
The drawing was made with the same equipment
with a Leica 308700 camera lucida attachment.
The format of the description follows Cusset
(1983), e.g in referring to scale-like leaves as “leaflets”
and long, subfiliform, to narrowly ribbon-like
dichotomously branched leaves as “leaves”. All specimens marked “!” have been seen. The conservation
assessment follows the IUCN (2012) standard. The
main online search address used for retrieving specimen data from P was https://science.mnhn.fr/institution/mnhn/collection/p/item/p00179355?listIndex=
128&listCount=610. Herbarium codes follow Index
Herbariorum (Thiers, continuously updated).
Accepted for publication 9 November 2016.
1
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK. e-mail: m.cheek@kew.org
2
Herbier National de Guineé, Université de Gamal Abdel Nasser, BP 680, Conakry, République de Guinée.
3
Guinée-Biodiversité., Conakry, République de Guinée.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2016
55
Page 2 of 8
Completing the resurrection of Inversodicraea
Engl.
Molecular phylogenetic sampling of African
Podostemaceae has been only c. 30% complete at
species level (Schenk et al. 2015). Thiv et al. (2009)
and Schenk et al. (2015), have convincingly shown
that Ledermanniella subg. Phyllosma C. Cusset forms
a well-supported clade of species that are sister to
the rest of all other African Podostemaceae sampled, mainly comprising Ledermanniella subg.
Ledermanniella and, embedded within it, seven
smaller genera: Macropodiella, Winklerella, Djinga,
Dicraeanthus, Leiothylax, Letestuella and Stonesia. Thiv
et al. (2009) and Schenk et al. (2015) both advocate
resurrecting the generic name Inversodicraea Engl.,
as the oldest generic name for the species of
Ledermanniella subg. Phyllosma and both transfer
species formerly included under the last taxon to
Inversodicraea. The last seven remaining species
names of subg. Phyllosma are placed into
Inversodicraea below.
Species of the resurrected Inversodicraea
(Ledermanniella subg. Phyllosma as erected, delimited
and revised by Cusset (1983)) are easily recognised
due to the unique synapomorphy (derived feature)
in having their stems more or less densely covered
in spirally inserted, dorsiventrally flattened,
often lobed, rarely needle-like, scale-leaves or
“leaflets”, often in addition to larger, but caducous,
branched, filiform leaves which are also commonly
seen in other African Podostemaceae. Such
scale-leaves are otherwise absent in African
Podostemaceae.
1. Inversodicraea mortonii (C. Cusset) Cheek comb.
nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60472989-2
Basionym: Ledermanniella mortonii C. Cusset, Bull. Mus.
Natl. Hist. Nat. 4e sér., 5, B, Adansonia 4: 368 (1983).
Type: Sierra Leone, Morton SL 2899 (holotype WAG;
isotype K!).
2. Inversodicraea gabonensis (C. Cusset) Cheek comb.
nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60472990-2
Basionym: Ledermanniella gabonensis C. Cusset, Bull.
Mus. Natl. Hist. Nat. 4e sér., 5, B, Adansonia 4:
368 (1983). Type: Gabon, Le Testu 6017
(holotype P).
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2016
KEW BULLETIN (2016) 71:55
3. Inversodicraea boumiensis (C. Cusset) Cheek comb.
nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60472991-2
Basionym: Ledermanniella boumiensis C. Cusset, Bull.
Mus. Natl. Hist. Nat. 4e sér., 5, B, Adansonia 4: 370
(1983). Type: Gabon, Le Testu 6536 (holotype P).
4. Inversodicraea harrisii (C. Cusset) Cheek comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60472992-2
Basionym: Ledermanniella harrisii C. Cusset, Bull. Mus.
Natl. Hist. Nat. 4e sér., 5, B, Adansonia 4: 374 (1983).
Type: Sierra Leone, Harris 41 (holotype P).
5. Inversodicraea paulsitae (C. Cusset) Cheek comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60472993-2
Basionym: Ledermanniella paulsitae C. Cusset, Bull. Mus.
Natl. Hist. Nat. 4e sér., 5, B, Adansonia 4: 378 (1983).
Type: Gabon. Le Testu 6510 (holotype P).
6. Inversodicraea torrei (C. Cusset) Cheek comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60473134-2
Basionym: Ledermanniella torrei C. Cusset, Bull. Mus.
Natl. Hist. Nat. 4e sér., 5, B, Adansonia 4: 386 (1983).
Type: Mozambique, Torre 5155 (holotype LISC).
7. Inversodicraea thollonii (Baill.) Cheek comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60473135-2
Basionym: Podostemum thollonii Baill., Bull. Mens. Soc.
Linn. Paris 2: 877 (Baillon 1890). Type: Gabon,
Thollon 825 (holotype P).
Inversodicraea pepehabai
Pepe Haba 292, here described as Inversodicraea pepehabai,
is one of the first records of Podostemaceae from the
Guineé Forestière Region in the Loma-Man Highlands
of the Republic of Guinea (Guinea-Conakry) possibly as
a result of insufficient previous collecting effort. In
contrast, the waterfalls of the Fouta Djalon Highlands
of that country are much better known for multiple
species of Podostemaceae (see for example Taylor 1953).
KEW BULLETIN (2016) 71:55
Page 3 of 8 55
Since the flowers of Pepe Haba 292 are inverted in
the spathellum, and the ellipsoid ovaries and fruits are
circular in section, with eight longitudinal ribs, this
specimen would clearly have been placed in
Ledermanniella Engl. in the prevailing classification of
African Podostemaceae by Cusset (1973, 1974, 1978,
1983, 1987). However, since the stems possess numerous scale-like leaflets, it is placed here in the genus
Inversodicraea (formerly Ledermanniella subgenus
Phyllosma C. Cusset, revised by Cusset (1983)). In
Cusset’s 1983 treatment it keys out as L. adamesii (G.
Taylor) C. Cusset (now Inversodicraea adamesii G.
Taylor) since it has irregularly and shallowly 2 – 3lobed leaflets which are scattered, covering < 50% of
the stems. However, it differs from that species in the
characters detailed in Table 1 below.
Inversodicraea pepehabai Cheek sp. nov. Type: GuineaConakry, Guineé-Forestère, Seredu, Mt Ziama,
8°21'25.8"N, 9°18''30.7"W, fl. fr. 8 Feb. 2010, Pepe M.
Haba 292 (holotype HNG!, isotypes K!, P!).
http://www.ipni.org/urn:lsid:ipni.org:names:60473136-2
Annual (probably), erect, densely branched herb, 3 –
7.5 cm tall (excluding the leaves), attachment to
substrate unknown. Stems in wefts, each main axis with
5 – 15 lateral branches, one or two of which are as
equally developed as the main axis, so that 8 – 20 stem
apices are developed distally, equidistant from the
base (Fig. 1A and B). Stems erect, terete, 0.5 – 0.9 mm
diam., ± evenly (densest at stem apices) 30 – 40%
covered, in leaflets. Leaflets patent, ovate to obovate in
outline, (0.3 –) 0.4 – 0.6 mm long, entire, or with 1 or
2 shallow, short (about 1/10 the length of the leaflet,
i.e. c. 0.05 mm long) triangular lateral lobes, apex
acute to obtuse, basal attachment as broad, or almost
as broad as the leaflet. Leaves falling from stems at
anthesis (fruiting stems leafless), arising at stem apex,
terminal, flattened, 0.4 – 1.5 cm × 0.1 mm dichoto-
mously branching 2 – 4-times in the distal halves,
apices acute. Spathellae single, terminal on stems,
probably obovoid in bud, c. 2 × 0.75 – 1.2 mm. Flower
inverted in bud; pedicel 2 – 2.5 (– 3) mm long at
anthesis, not accrescent; tepals two, filiform, 0.6 –
0.75 mm long, inserted on the gynophore. Androecium
of a single stamen, erect, not extending above the
ovary, (1.2 –) 2 – 4 mm long, anther cells 4, equal, 0.25
(– 0.5) mm long. Pollen in monads. Gynoecium:
gynophore 0.2 – 0.8 (– 1) mm long; ovary unilocular,
narrowly ellipsoid, 0.9 – 1.3 × 0.5 mm, orbicular, not
flattened in transverse section, with 8 subequal,
equidistant, parallel longitudinal ribs, those of the
sutures as prominent as the remaining 6; stigmas two,
botuliform 0.7 – 1.0 × 0.15 mm, held at c. 180° apart.
Fruit of similar dimensions to ovary, ribs cylindrical in
section c. 0.05 mm diam., styles caducous. Seed planoconvex, papillate, 0.2 – 0.25 × 0.1 × 0.8 mm. Figs 1 – 3.
RECOGNITION. Similar to Inversodicraea adamesii in
the irregularly-lobed (lobes 1, 2 or absent) leaflets
which cover < 50% of the stem. Distinct in the
leaflets being patent (not ± appressed), the flowers
with 1 stamen (not 2), the fruits 8-ribbed (not 6).
DISTRIBUTION. Republic of Guinea (Guinea-Conakry).
SPECIMEN EXAMINED. GUINEA-CONAKRY . GuineéF o r e s tè r e , S e r e d u , Mt Z i a m a , 8 ° 21 ' 2 5 . 8 " N ,
9°18'30.7"W, fl. fr. 8 Feb. 2010 Pepe Haba 292
(holotype HNG!, isotypes K!, P!).
ECOLOGY. Rapids in submontane evergreen forest; c.
600 m asl.
CONSERVATION. Inversodicraea pepehabai is here assessed
as Endangered under Criterion D of IUCN (2001),
since less than 250 individuals are estimated to
have been observed at the single location known
(EN D1). The area of occupancy (AOO) is
assessed as 1km2 using the 1km2 grid preferred
by IUCN for riverine organisms. Currently there
are no obvious threats, but should tea and/or
quinine plantations, or timber extraction be resumed inside the Ziama Foret Classé, there is a
Table 1. The significant diagnostic characters separating Inversodicraea adamesii and I. pepehabai.
Habit
Leaflet shape
No. stamens/flower
No. ribs on fruit
Leaf: Flower length
Pedicel length
Androecium:Gynoecium length
Filament length
Stigma shape
Inversodicraea adamesii
Inversodicraea pepehabai
branches spreading, tending to horizontal
lanceolate
2
6
1:3
c. 10 mm
2:1
5 – 8 mm
subulate
stems and branches always erect
ovate to obovate
1
8
2:1
2 – 2.5 ( – 3) mm
0.8:1
(1.2 –) 2 – 4 mm
botuliform
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2016
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KEW BULLETIN (2016) 71:55
Fig. 1. Inversodicraea pepehabai. A habit of larger plant, with fruit, leaves have been shed; B habit of smaller, flowering plant, leaves still
attached; C detail of B, showing 2 – 4 times dichotomously branched leaves; D stem with flower at anthesis, and fruits before dehiscence;
E flower emerging inverted from spathellum; F flower approaching anthesis: ovary not yet erect; G leaflet variation: entire, 2 – 3 lobed; H
fruit, transverse section showing 8 ± equally developed ribs; J seed, surface papillate. From Pepe Haba 292. DRAWN BY ANDREW BROWN.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2016
KEW BULLETIN (2016) 71:55
risk that surface run-off would raise levels of
nutrients and silt in entering the water course in
which I. pepehabai is found, increasing algal-growth,
that would compete with, and (due to the
increased silt levels), reduce establishment of new
plants of the species.
It is to be hoped that additional records of this
species will be found at further sites, lowering its
threat assessment.
Inversodicraea pepehabai is one of several plant
species unique to the Ziama Massif inside the
Ziama Man & Biosphere reserve. It is likely to be
proposed in future as an “Important Plant Area”
for Guinea (see Darbyshire, continuously updated).
ETYMOLOGY. Named for the collector of the only
known specimen, Pepe Haba of N’zerekore, one of the
most expert field botanists in Guinea.
Discussion
The species described here as Inversodicraea
pepehabai is unusual among African Podostemaceae
being found in the rapids of a submontane forest
area. Other species with this ecology are Saxicolella
Page 5 of 8 55
amicorum J. B. Hall (Hall 1971; pers. obs. Cheek),
Ledermanniella letouzeyi Cusset (Cusset 1987; Cheek
et al. 2004), L. onanae Cheek (Cheek 2003; Cheek
et al. 2004) and L. pollardiana Cheek & Ameka
(Cheek & Ameka 2008). Most other species of the
genus occur in rapids of lowland forest or woodland habits.
The affinities of Inversodicraea pepehabai are
probably with other West African species of the
genus which have slender, usually lobed, leaflets. At
first sight Inversodicraea pepehabai appears very
similar to the relatively common and widespread
I. ledermannii Engl., but the leaflets in that species
are consistently, deeply and equally 3-lobed, and
cover most of the stem, being ± appressed, not
patent. The leaflets of I. pygmaea G. Taylor
(L. taylori C. Cusset) are also similar, but are also
consistently 3-lobed resembling I. ledermannii excepting that the central lobe is always longer than
the laterals, and again, the stems are short (1 –
2 cm long) and completely covered in leaflets. In
I. mortonii the leaflets are needle-like, and
completely lacking lobes. It is with I. adamesii that
the closest similarity of leaflet shape occurs, but the
Fig. 2. Inversodicraea pepehabai. Plants in flower at the type locality at Mt Ziama, Feb. 2010.
PHOTO BY PEPE HABA.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2016
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KEW BULLETIN (2016) 71:55
Fig. 3. Inversodicraea pepehabai. Plants in situ on vertical rock face at the type locality at Mt Ziama, Feb. 2010.
differences in habit, flower and fruit leave no
doubt that they are distinct species (see Table 1).
Inversodicraea pepehabai is among a host of new
species that have been brought to light and
published from the Republic of Guinea in the last
seven years, following decades of relative inactivity
in plant species discovery in that country and its
neighbours. Examples of such newly discovered
species in Guinea are: Brachystephanus oreacanthus
Champl. (Champluvier & Darbyshire 2009),
Xysmalobium samoritourei Goyder (2009), Gymnosiphon
samoritoureanus Cheek (Cheek & van der Burgt
(2010)), Striga magnibracteata Eb. Fisch. & I.
Darbysh. (Fischer et al. 2011), Eriosema triformum
Burgt (van der Burgt et al. 2012), Eriocaulon
cryptocephalum S. M. Phillips & Mesterházy (2015),
Napoleonea alata Jongkind (Prance & Jongkind
2015) and Psychotria samoritourei Cheek (Cheek &
Williams 2016). Just over the border in Mali,
Calophyllum africanum Cheek & Luke (2016) was
recently found, and in Ivory Coast Macropodiella
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2016
PHOTO BY PEPE HABA.
cussetiana Cheek (Cheek & Ameka 2016). Even a
new rheophytic genus, Karima Cheek & Riina has
come to light in Guinea (Cheek et al. 2016).
Acknowledgements
Professor Basile Camara and Dr Seydou
Magassouba, respectively Director General and
Director-Adjoint, Université de Gamal Abdel
Nasser-Herbier National de Guinée, are thanked
for arranging permits and for their long term
support and collaboration. Janis Shillito is thanked
for typing the manuscript. George Gosline assisted
with access to electronic specimen data. This paper
was completed under the project Important Plant
Areas in the Republic of Guinea supported by the
Darwin Initiative of the Department of the Environment Food and Rural Affairs (DEFRA), UK
government (project Ref. 23 – 002). Two anonymous reviewers are thanked for constructive comments on an earlier draft of the paper.
KEW BULLETIN (2016) 71:55
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