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Identifications of Karima were confirmed at the National Herbarium of Sierra Leone at the University of Njala, and further confirmed at K using photographs of the study subjects. All photographs associated with this paper were taken during this study. The electronic version of this article in Portable Document Format PDF in a work with an ISSN or ISBN will represent a published work according to the International Code of Nomenclature for algae, fungi, and plants [ 19 ], and hence the new names contained in the electronic publication of a PLOS article are effectively published under that Code from the electronic edition alone, so there is no need to provide printed copies.
In addition, the new names contained in this work have been submitted to IPNI, from where they will be made available to the Global Names Index. The pollen grains are spheroidal, inaperturate and have a crotonoid exine sculpture pattern with supratectal subunits attached to the upper edge of low, circular muri each enclosing a central lacuna. The subunit rings are 2. Subunits are ovoid-spheroidal, 1—1. The floors of the central lacunae each have 7—20 scattered granules, each c. Fig 1. All from Momoh The concatenated dataset combining trnL-F and part of rbcL contained 33 accessions and aligned positions, of which were parsimony-informative.
In the case of rbcL we were only able to amplify part of this region for the newly sampled taxa, therefore we only used the corresponding fragment of the published sequences downloaded from Genbank. The trnL-F dataset with aligned positions has more parsimony-informative characters than the rbcL data set with aligned positions and 84 parsimony-informative characters.
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The rbcL fragments for the two accessions of Karima scarciesii are identical. The two trnL-F sequences are nearly identical and differ only in one position, however, one of the accessions Jongkind has 55 characters missing at the beginning of the sequence. Preliminary Bayesian analyses of the individual datasets did not show evidence of topological incongruence between the two genetic markers results not shown.
The Bayesian consensus tree Fig 2 resulting from the analysis of the concatenated matrix shows a phylogenetic structure congruent with the phylogenetic analyses of [ 2 ] their figures 1, 2, 3. As in [ 2 ], clade C2 is highly supported but resolution at the backbone is quite low. Clades C1, C2 and inaperturate crotonoids according to Wurdack et al. The clade formed by Domohinea perrieri Leandri and Tannodia cordifolia Baill.
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The morphological differences of Karima from Croton sensu stricto are extensive Table 1. It is surprising that no other researchers have remarked upon the absence in Croton scarciesii that is, the new genus, Karima of the key diagnostic characters of Croton , such as lepidote scales or stellate hairs, the presence of nectary glands at the junction of the petiole and leaf blade, a crotonoid inflorescence thyrsoid inflorescence with female and male flowers at base, male towards apex , and inflexed staminal filaments in flower buds.
The placement of Karima near Neoholstia was reached by molecular analysis but is strongly supported by morphology Table 2. The two genera share stalked, glandular, red-apexed glands on the bracts unusual in Crotonoideae , and leaf texture and venation are also similar, as is the general indumentum type simple, erect, robust, translucent hairs. The tuberculate fruits of both genera are also similar, apart from differences in the styles. However there are major qualitative characters that divide the two taxa. These characters are usual in serving to distinguish genera, rather than species, in Euphorbiaceae as can be seen in [ 21 ].
Data for Neoholstia taken from [ 23 ] and from observations on material at K. Karima was erroneously misplaced in Croton as Croton scarciesii by Scott-Elliot [ 24 ]. In that paper most of the species are described from his specimens by the corresponding family specialists at Kew. Where no family specialist was available, Scott-Elliot himself took on responsibility for placing and describing the taxa that he considered to be new to science. Evidently, at that time, Kew had no Euphorbiaceae specialist. The placement of the taxon in Croton appears never to have been challenged.
Opportunities to detect the error were missed when the Croton accounts for the Flora of West Tropical Africa were written by the respective editors Hutchinson [ 25 ] and, in the revised edition, by Keay [ 26 ]. It is possible that they were under pressure of time to complete these accounts and so could not critically review Scott-Elliot's generic placement of this species. Perhaps they attributed the obviously anomalous morphology within Croton of the species to its rheophytic ecology.
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It is also possible that they did not dissect the flowers, which would have revealed additional characters that militated against placement in Croton Table 1. Despite the strong morphological separation of Karima from Croton Table 1 , examination of the pollen indicates that grains are spheroidal, inaperturate and with crotonoid exine sculpturing. This characterises subfamily Crotonoideae, and together with the gross morphological characters of the indumentum, leaf and flower and utilisation of the keys in Radcliffe-Smith [ 21 ] and Webster [ 27 ], suggests that Karima is best placed in Crotonoideae in the region of Codiaeae and Aleuritidieae.
The molecular phylogeny also confirms that Karima is part of Crotonoideae, and that it belongs to the inaperturate crotonoid clade C2 of Wurdack et al. This clade, emended in this paper Fig 2 , is comprised of all sampled members 27 genera of the following tribes as delimited by Radcliffe-Smith [ 21 ]: Codiaeae Pax Hutch. Webster; Ricinocarpeae Muell. However 15 genera c. The topology of clade C2 may change if any of these genera are included in a future analysis. Key to the putative genera of the Karima clade of C2 inaperturate Crotonoids sensu Wurdack et al.
L Webster Radcliffe-Smith [ 21 ]: — Genera in bold did not appear in the phylogenetic analysis of the combined dataset in Wurdack et al. The DNA sequence data indicate that Karima scarciesii is most closely related to the southeastern tropical African Neoholstia tenuifolia Fig 2.
We did not include the new taxon within the monotypic genus Neoholstia [ 21 ] because morphological differences between the two taxa are considerable see above and Table 2. Furthermore, the current phylogeny of the C2 clade of Wurdack et al. Phylogenetic analyses including additional species from each genus and using more informative molecular markers are still needed to better address taxonomic delimitation within clade C2 of the inaperturate crotonoids.
Monoecious shrub, indumentum simple, hairs sometimes gland-tipped. Stems with sparingly branched erect principal axes, and extremely short, lateral spur stems.
Leaves alternate, simple, entire, nervation pinnate with the basal nerve pair acute, glandular tissue absent; petiolate; stipules persistent, becoming indurated, margins glandular-hairy when young. Inflorescences terminal on spur short lateral shoots; male inflorescences several-flowered, glomerulate, less usually spike-like, bracts enclosing bracteoles and multiple contracted cymules, flowers emerging sequentially; female inflorescences 1-flowered. Petals 5, alternating with sepals, included in calyx, elliptic, glabrous, except for a transverse line of hairs on adaxial surface.
Disc glands 5, alternating with petals, angular-ellipsoid, erect, free, glabrous except for distal tuft of hairs. Stamens 6 —8 , erect in bud, exserted, filaments terete, glabrous apart from a band of hairs above the base, anthers dehiscing laterally, connective expanded, flattened, anther cells opposite.
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Pollen spheroidal, inaperturate and with crotonoid exine sculpturing. Female flowers resembling but about twice the size of the male flowers, calyx divided nearly to the base into 5 —6 sepals. Petals 5 —6 , twice the size of the male and visible from the exterior in the sepal sinuses; disc flattened, pentagonal; ovary shallowly 3-lobed, densely hairy; locules 3, each 1-ovulate, ovules pendulous, epitropous; styles 3, free, each bifid in distal quarter, flattened and glabrous on adaxial surface.
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Fruits dry, dehiscing septicidally into 3, single-seeded cocci, leaving an axile erect placenta; the seed retained in the cocci and visible through a ventral aperture, sepals and styles persistent in fruit. Seeds ellipsoid, carunculate, marbled, with a raphe line; testa membranous, translucent; tegmen mechanical, crustaceous, comprised of a single layer of subcolumellar dark brown sclereids.
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Embryo with cotyledons orbicular, flattened, radicle stout, almost as long as cotyledons Fig 3. E , Jordan K-N. Karima scarciesii Scott-Elliot Cheek comb. Types: Sierra Leone, Scarcies, near Mofari, fr. K ; Guinea, Scarcies, near Sasseni, fl.
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Lectotype selected here: Scott-Elliot K! Rheophytic, fastigiate, monoecious, partially deciduous shrub 0. Stems 5—8 mm diam. Primary axes sparingly branched, internodes 1. Spur shoot leaves sometimes deciduous when submerged? Stems at apex terete, 0. Leaves densely crowded on spur shoots, more widely spread on primary axes, alternate, simple, chartaceous, stipulate, petiolate.
Blades rhombic-elliptic, rhombic-ovate or obovate 1. Petiole canaliculate, 1—2 —3 mm long; indumentum as stem. Stipules persistent, subulate, rapidly becoming indurated, brown, narrowly triangular, 3—3.