Blumea 64, 2019: 165 –176 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE https://doi.org/10.3767/blumea.2019.64.02.07 A revision of the Crepidorhopalon whytei complex (Linderniaceae) in eastern Africa I. Darbyshire1, B. Wursten 2, Q. Luke 3, E. Fischer 4 Key words Chimanimani hepperi IUCN Red List assessment Kwale Namuli Nguru taxonomy Abstract The Crepidorhopalon whytei (Linderniaceae) species complex is revised using morphological analyses. Based primarily on variations in indumentum, floral morphology, corolla colour and seed morphology, four species are formally recognised within this group in eastern Africa. Crepidorhopalon whytei s.str. is widespread in the highlands of eastern Africa, extending from South Sudan and Ethiopia in the north through to western Tanzania in the south. A new combination in Crepidorhopalon is made for Lindernia flava (= C. flavus), which is confined to the Manica Highlands of the Mozambique-Zimbabwe border. Two new species are described, C. namuliensis which is known only from Mt Namuli in northern Mozambique and C. kwaleensis which is known only from the coastal lowlands of southeast Kenya. A fifth, imperfectly known species is documented from the Nguru Mountains of Tanzania where it is so far known from a single collection. Three names are lectotypified. The habitat requirements and distribution are documented and the extinction risk is assessed for each species. Crepidorhopalon flavus is assessed as globally Vulnerable and C. kwaleensis as globally Endangered, while C. namuliensis and C. whytei are currently considered to be of Least Concern, although the latter is declining markedly in parts of its range. The botanical importance of the key sites for the newly recognised taxa is discussed. Published on 30 July 2019 INTRODUCTION The genus Crepidorhopalon Eb.Fisch. (Fischer 1989) in the family Linderniaceae comprises species with a combination of aulacospermous seeds, generally non-rosulate habit, and the presence of anatomically complex multicellular clavate hairs on the lower lip of the corolla (Fischer 1989, 1992, Fischer et al. 2014). In the few species that have a rosulate habit, notably Crepidorhopalon welwitschii (Engl.) Eb.Fisch., C. malaissei Eb.Fisch. and C. robynsii Eb.Fisch., this is the result of a condensed main stem, with single flowers in the axils of leaflike bracts rather than with lateral bracteate inflorescences in the leaf axils as in Craterostigma Hochst. or Linderniella Eb.Fisch., Schäferh. & Kai Müll. In the ‘Flora of Tropical East Africa’ account of the Scrophulariaceae (Ghazanfar et al. 2008), Crepidorhopalon was not upheld, and species that had previously been transferred to that genus were treated in the genera Lindernia All. and Torenia L. However, molecular studies (Rahmanzadeh et al. 2005, Fischer et al. 2013) support the morphological evidence for recognition of Crepidorhopalon as a distinct genus, and it is now widely accepted (e.g., African Plant Database, continuously updated, Stevens 2001 onwards). for example C. chironioides (S.Moore) Eb.Fisch. and C. symo­ ensii Eb.Fisch., respectively restricted to the Kundelungu Mountains and the Upemba region of D.R. Congo (Fischer 1989, 1992) and C. mutinondoensis Eb.Fisch. & I.Darbysh., known only from the Mutinondo Wilderness Area in eastern Zambia (Fischer et al. 2014). Many species occur in seasonally wet flushes over rock or in more permanently marshy areas. One of the more frequently encountered and well-known species of the genus is Crepidorhopalon whytei (Skan) Eb.Fisch. which is a widespread trailing or scrambling herb of upland marshes and seepage areas in eastern Africa (Fischer 1992, 1999, Philcox & Ghazanfar 2008 as Lindernia whytei ). The main range of this species is from Ethiopia and South Sudan Crepidorhopalon is currently recorded from tropical Africa, where 30 species are known, and from a single species on Madagascar (African Plants Database, continuously updated). The centre of diversity for the genus is in the Haut Katanga area of the Democratic Republic of the Congo where 19 species are found (Fischer 1999). As in other genera of Linderniaceae, many species of Crepidorhopalon are highly range-restricted, 1 2 3 4 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom; corresponding author e-mail: i.darbyshire@kew.org. Herbarium, Nieuwelaan 38, Meise 1860, Belgium. East African Herbarium, National Museums of Kenya, P.O. Box 45166, Nairobi 00100, Kenya. Institut für Integrierte Naturwissenschaften – Biologie, Universität KoblenzLandau, Universitätsstraße 1, 56070 Koblenz, Germany. Map 1 Distribution of species in the Crepidorhopalon whytei complex: C. whytei (●); C. namuliensis (■); C. flavus (✶); C. sp. aff. whytei (▼); C. kwaleensis (▲); ‘Lindernia gossweileri ’ (●). © 2019 Naturalis Biodiversity Center You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 166 through Uganda, eastern D.R. Congo and Kenya to Rwanda, Burundi and western Tanzania (Map 1), where it has blue or purple flowers (Fig. 1a – c). However, Philcox (1990) included both Lindernia flava S.Moore, described from Mozambique and Zimbabwe, and L. gossweileri S.Moore, described from Angola, within a broader circumscription of C. whytei although noted that these taxa have yellow flowers. He said: “unfortunately, Blumea – Volume 64 / 2, 2019 I have seen only two collections from the Flora Zambesiaca area and one, Walters 2788, lacks any descriptive notes while the other, Goldsmith 72/68, cites the flowers as being yellow”. This was in addition to the two syntypes of L. flava, both of which are cited as having yellow flowers. Philcox went on to say that, despite this flower colour difference, “I do not hesitate to widen the range of colour in my description to include both extremes” Fig. 1 Crepidorhopalon whytei and allies. a – c. Crepidorhopalon whytei, Butare, Rwanda; d – e. C. namuliensis, Mt Namuli, Mozambique; f– h. C. flavus, foothills of the Chimanimani Mountains, Mozambique. — Photos: a – c. E. Fischer; d – g. B. Wursten; h. I. Darbyshire. I. Darbyshire et al.: Crepidorhopalon whytei complex in eastern Africa 5 – 230 m 1940 – 2010 m 330 –1130(–1340) m Ellipsoid or subquadrangular, with longitudinal ridges and furrows and with faint horizontal ribs 1100 – 2250 m Seed characteristics Altitude range 1730 –1870 m Sub-square in face view, circular in lateral view, with a row of subcircular depressions (and with one longitudinal furrow on the posterior side) Unknown Ellipsoid, with longitudinal ridges very shallow and inconspicuous 0.5 –1.2 mm Anterior stamen spur length Ellipsoid, with longitudinal ridges and furrows 1.8 – 2.4 mm ± 0.8 mm 0.5 – 0.8 mm Either foliaceous or ± muchreduced, then ovate-acuminate to linear-lanceolate Bracts 1.2 –1.75 mm [Except proximal-most pairs] lanceolate or linear-lanceolate Foliaceous Foliaceous or ± much-reduced, then ovate-acuminate to lanceolate-acuminate Largely glabrous or with short eglandular hairs; with or without scattered (sub)sessile glands Indumentum of pedicels and calyces and fertile portion of stems (excluding calyx lobe margins) Foliaceous Glabrous or with few sessile glands on young stems Glabrous or with few sessile glands on young stems Glabrous or with few sessile glands Blue or purple with white patches in the mouth Corolla colour: lower lip Glandular-pubescent Unknown C. whytei Character This study is based primarily upon the investigation by the first author (I.D.) of specimens from a range of herbaria, most notably the collections at K and BM but with consultation of relevant specimens (either physically or digitally) from BR, L, LMA, P, SRGH and WAG. Further, it builds upon the monographic work on African Linderniaceae conducted by the final author here (E.F.) who studied specimens of this species complex from a range of other herbaria (B, BRLU, BRVU, M, MJG – see Fischer 1992). The LISC herbarium (IICT/University of Lisbon), important for historical specimens from Mozambique, is currently being rehoused at the Museum of the University of Lisbon (MUHNAC) and so some of the Mozambican specimens of Crepidorhopalon were not available for study but we were kindly sent images of some relevant specimens by Dr Maria Romeiras at MUHNAC. Photographs of the species in the field were also consulted wherever available. Abbreviations for herbaria follow the standard form listed on Index Herbariorum (Thiers, continuously updated), except for the fledgling herbarium of the Micaia Foundation (http://micaia.org, last accessed May 2019) at Ndzou Camp in Moribane Forest Reserve which is listed as ‘Ndzou’. Only specimen duplicates that have been seen by one or more of the authors are cited; duplicates that have not been seen are not listed except where they have been sought for and not located (marked as ‘[not found]’), and except for isotype specimens that have not been seen (marked as ‘[not seen]’). Those for which only online images have been seen are marked with an asterisk (*); these were accessed via JSTOR Global Plants (https://plants.jstor.org/, last accessed May 2019), or via online herbarium catalogues. Only a maximum of three specimens per country are cited for Crepidorho­ palon whytei; all material seen is cited for the other taxa. All measurements were made on dried material except for those of the flowers, which were soaked in Aerosol OT 5 % solution prior to dissection and measurement. Table 1 The main morphological characters separating species in the Crepidorhopalon whytei complex. MATERIALS & METHODS C. namuliensis C. flavus As herbarium material from across the range of C. whytei s.l. was reviewed, three further interesting taxa were uncovered. The first is a species discovered during recent surveys on Mt Namuli in northern Mozambique which, while close to C. whytei and found in a similar ecological niche, has several diagnostic characters (Downes & Darbyshire 2017). The second is a species from the coastal lowlands of Kenya in Kwale District that has been named as C. whytei in the past but has several distinct characters as well as being ecologically separated from true C. whytei. The third is a small-flowered taxon known only from a single collection from the Nguru Mountains of Tanzania – the only record of the C. whytei complex in the ancient Eastern Arc mountain chain of East Africa. A synoptic revision of the C. whytei complex has therefore been conducted in order to delimit the taxa in this challenging group. Yellow C. sp. aff. whytei (= Manktelow et al. 91081) C. kwaleensis Recent field observations around the foothills of the Chimanimani Mountains (Darbyshire and Wursten, pers. obs.), have demonstrated that the populations in this area consistently have corollas with a bright yellow lower lip (the upper lip is typically yellow-brown or purplish tinged) – see Fig. 1f– h. Other collections from the region, not seen by Philcox for Flora Zambesiaca (Biegel 3583, Pope & Müller 1295) also support this observation. These populations are found in low- to mid-altitude marshy sites, usually less than 1 100 m a.s.l. Plants from the core range of C. whytei in East Africa never have yellow flowers and are found at higher altitude sites, over 1 100 m a.s.l. These observations led the current authors to a renewed interest in the C. whytei group. White or tinged pale blue with a yellow patch in the mouth (p. 63). Philcox’s approach has been followed in subsequent publications (e.g., Fischer 1992). Purple with white patches in the mouth 167 168 The categories and criteria of the IUCN Red List of Threatened Species (IUCN 2012) are used to assess the conservation status (extinction risk) of each species. Extent of occurrence (EOO) and area of occupancy (AOO) were calculated using the GeoCat tool (Bachman et al. 2011), last accessed May 2019. The species distribution map was produced using SimpleMappr (Shorthouse 2010). RESULTS Based on detailed morphological analyses, we formally recognise four species within the Crepidorhopalon whytei complex in eastern Africa and informally recognise a fifth, imperfectly known species (Table 1). Crepidorhopalon whytei s.str. is widespread in the highlands of eastern Africa, extending south as far as western Tanzania; it is somewhat variable in corolla size and bract morphology but these differences appear to be clinal. The other three species are highly range-restricted: C. flavus (S.Moore) I.Darbysh. & Eb.Fisch. is confined to the Manica Highlands of the Mozambique-Zimbabwe border, centred on the Chimanimani massif, C. namuliensis I.Darbysh. & Eb.Fisch. is known only from Mt Namuli in northern Mozambique, C. kwaleensis I.Darbysh. & Eb.Fisch. is known only from the coastal lowlands of Kenya in Kwale County and the imperfectly known species is so far recorded only from a single collection from the Nguru Mountains of Tanzania (Map 1). Crepidorhopalon whytei, C. flavus and C. namuliensis are clearly closely related and, while easily separable morphologically (Table 1 and key below), the rank at which to recognise these taxa is open to debate. However, in view of their wide geographic separation, and the expected ensuing genetic isolation, coupled with the consistent morphological differences, we choose to recognise them as distinct species rather than subspecies. This decision is in keeping with the established convention within African Linderniaceae of recognising all distinct taxa at species rank (see, e.g., Fischer 1992). Based on differences in seed morphology, C. kwaleensis is easily separated from C. whytei, C. flavus and C. namuliensis; it has previously also been confused with C. hepperi Eb.Fisch. from western Tanzania, which is therefore included in the key to species in the Taxonomic Account presented below. TAXONOMIC ACCOUNT Key to the Crepidorhopalon whytei complex 1. Corolla lacking short glands on external surface of tube; imperfectly known species from the Nguru Mountains of Tanzania . . . . . . . . . . . . . . . . . . . . . . . . . C. sp. aff. whytei 1. Corolla with short glands conspicuous on distal portion of tube externally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Lower lip of corolla yellow; seeds with only shallow, almost imperceptible longitudinal furrows; restricted to the Manica Highlands of Mozambique and Zimbabwe . . . . . C. flavus 2. Lower lip of corolla blue, purple or white; seeds either with conspicuous longitudinal furrows or subcircular depressions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Lower lip of corolla white or at most tinged pale blue; calyces, pedicels and distal internodes with numerous glandular hairs; restricted to Mt Namuli, Mozambique . . . . C. namuliensis 3. Lower lip of corolla deep blue or purple; calyces, pedicels and distal internodes largely glabrous or with eglandular hairs, sometimes with scattered (sub)sessile glands but these inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Seeds not longer than wide, with subcircular depressions along the face (Fig. 2i – k, 4i); spurs of anterior pair of sta- Blumea – Volume 64 / 2, 2019 mens markedly long and slender, 1.8–2.4 mm long; restricted to Kwale region, Kenya . . . . . . . . . . . . . . . . C. kwaleensis 4. Seeds ± longer than wide, with longitudinal furrows along the face (Fig. 2a – c, g – h); spurs of anterior pair of stamens shorter and less slender, 0.5–1.2 mm long . . . . . . . . . . . 5 5. Stems procumbent, scrambling, trailing or at most weakly decumbent; calyces with lobes longer than tube; plants of highland marshes and flushes, widespread in eastern Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. whytei 5. Stems ± erect; calyces with tube longer than or equal to lobes; plants of seasonally wet grassland and miombo woodland in western Tanzania . . . . . . . . . . . . . . . . . C. hepperi Crepidorhopalon whytei (Skan) Eb.Fisch. — Fig. 1a–c, 2a–c Fischer (1989) 444; (1992) 165, t. 85, 86; (1999) 110, pl. 44; (2006) 270, f. 160.25; Darbyshire et al. (2015) 329. — Lindernia whytei Skan in Hemsley & Skan (1906) 340; Malaise (1985) 402; Philcox & Ghazanfar (2008) 70, pro maj. parte; Agnew (2013) 300. — Type: Whyte s.n. (lecto K [K000379658], chosen here), Kenya, 1st and 2nd days’ march from Mumias, fl. 7 fr. 6 –7 Dec. 1898. Additional syntypes: Hannington s.n. (K), Tanzania, Msalala, fl. without date [1894]; Scott­Elliot 7225 (K [K000379657]), Kenya, Nandi Country, fl. Dec. 1893 – see note. Torenia mildbraedii Pilg. (in Mildbraed & Pilger 1911: 285). — Type: Mildbraed 701 (B destroyed; lecto K [K000379666], chosen here), Luhondo Valley, fl. 13 Aug. 1907 – see note. Annual or perennial herb, with procumbent, scrambling, trailing or at most weakly decumbent stems up to 60 cm long or more; stems brittle, markedly 4-angular, winged along the angles, glabrous to sparsely or rarely more densely pale eglandularpubescent, distal internodes sometimes with scattered (sub)sessile glands; internodes often long, the leaves widely spaced. Leaves sessile, broadly ovate to suborbicular, 7–31 by 5.5–29 mm, base rounded or shallowly cordate, margin serrulate with 2 – 8(–10) teeth per margin, these sometimes minute, apex acute or obtuse; primary venation palmate, with 5 or 7 main veins from base; surfaces glabrous except sometimes for short hairs along margin towards base and/or on main veins beneath, rarely pubescent throughout, surfaces also with sunken glands drying brown, most visible on lower surface. Flowers either axillary and solitary or together forming a lax terminal raceme (this depending on size of bracts); bracts either foliaceous throughout or rapidly reducing in size distally, then ovate-acuminate to linear-lanceolate, 1.5–7.5 by 0.6–6(–9.5) mm, margin toothed or entire; bracteoles absent; pedicels 1.2 –7.5 mm long, glabrous or with (sub)sessile glands, rarely eglandular-pubescent. Calyx 5–10.5(–14) mm long, lobes linear-lanceolate, somewhat unequal in length, longest lobes 3–7.5(–10.5) mm long, apices divergent to recurved at maturity, each lobe 1-veined and with hyaline margin, external surface glabrous except for short pale eglandular hairs along lobe margins and sometimes extending onto tube, rarely eglandular-pubescent throughout, with or without scattered (sub)sessile glands. Corolla 10.5 – 24 mm long, lower lip blue or purple with white patches in the mouth, or very rarely individuals within a population pure white (Napier 5349), external surface glandular-puberulous mainly on tube, upper lip usually with pale eglandular hairs; tube (5.7–)7–11 mm long, cylindrical but slightly contracted in midpoint and widened at mouth, internal surface with 4 lines of subsessile glands in proximal half; upper lip ovate, 3 – 6.5 mm long, apex emarginate or rounded, internal surface with numerous minute glands; lower lip 5–12 mm long, 3-lobed, median lobe rounded-obovate, 4–7.5 by 5–7 mm, margins of lobes somewhat irregular, bosses on palate of lower lip with blunt-tipped multicellular hairs. Stamens 4, ventral stamens spurred, spurs clavate, 0.5 –1.2 mm long, papillate, filaments above spurred portion 3.8 –7 mm long; posterior stamens with filaments 1.1– 3 mm long; anthers of the two pairs of stamens adhering, thecae 0.7–1 mm long. I. Darbyshire et al.: Crepidorhopalon whytei complex in eastern Africa 169 threats to its habitats in parts of its range, and the species is rapidly decreasing at least in Rwanda and Burundi. While the species was widespread in swamps around Butare (Central Rwanda, Southern Province, Huye district) between 1984 and 1998, the intensive agricultural land use has led to a major decline of the populations, and the species has now almost disappeared (E. Fischer, pers. obs.). If the same situation is true across the species’ range then it could qualify as threatened under IUCN criterion A. Notes — Crepidorhopalon whytei is rather variable in flower size and inflorescence form. Plants from the Lake Victoria basin (Uganda, West Kenya, North Tanzania, e.g., E. & C. Godman 87, BM) have the smallest corollas and generally have more clearly developed racemes as the bracts are ± much-reduced in size in comparison to the proximal leaves. Those from the central Kenyan Highlands (e.g., Davis 60, BM) have the largest corollas and have foliaceous bracts, hence the racemes are leafy and the flowers appear to be axillary. Similar plants to these Kenyan populations are found elsewhere in this species’ range, for example in Ethiopia and along the Albertine Rift. However, there are also populations that are intermediate between the two forms, particularly in Uganda and neighbouring regions, and it has not been possible to separate any clear infraspecific taxa on the basis of flower and bract size. Fig. 2 Seeds of Crepidorhopalon whytei and allies. a – c. Crepidorhopalon whytei (E. Brown 343, Uganda); d – e. C. namuliensis (Patel 7319, Mozambique); f. C. flavus (Goldsmith 72/68, Zimbabwe); g – h. C. hepperi (Bidgood et al. 6937, Tanzania); i – k. C. kwaleensis (Luke 2899, Kenya). — Scale bar = 0.5 mm. Ovary ± 1.4 mm long; style 5.5 – 9 mm long; stigma fan-shaped and minutely fringed. Capsule narrowly ellipsoid to lanceoloid, brown, ± equal in length to calyx or slightly longer, 7–13 mm long, apex somewhat tapered, surface glabrous; seeds ellipsoid or subquadrangular ± 0.6 – 0.85 by 0.35 – 0.5 mm, with longitudinal ridges and furrows and with faint horizontal ribs. Distribution & Ecology — Crepidorhopalon whytei is widespread in East Africa, extending from southwest Ethiopia and southern-most South Sudan, through Uganda, Kenya, eastern D.R. Congo, Rwanda and Burundi to northern and western Tanzania (Map 1). It occurs in upland marshes, seepage areas, stream and pool margins at 1100 – 2250 m elevation. Selected specimens seen. Burundi, route Karuzi-Muhinga, étang Nzibariba, fl. & fr. 9 Jan. 1958, Van der Ben 1780 (BR, K); Mwaro Prov., Mwaro, fl. & fr. 10 Jan. 1968, Lewalle 2685 (BR, K); Bubanza Prov., route RweguraNdora, fl. 19 Oct. 1976, Reekmans 5438 (BR, K). – d.r. Congo, Aru-Mahagi, fl. & fr. Aug. 1931, Lebrun 3728 (BR, K); Munagana, fl. & fr. 1934, De Witte 1872 (BR, K); terr. Mahagi, Nioka-La Korda, fl. 2 Dec. 1957, Bamps 78 (BR, K). – Ethiopia, Kaffa, near Sombo, fl. & fr. 13 Nov. 1960, Mooney 8612 (K); about 7 km along the road from Jimma to Bonga, fl. & fr. 20 June 1969, J.J.F.E. de Wilde 5267 (BR, K, M, P, WAG*); Kochi, about 5 km E of Jimma along road to Addis Ababa, fl. & fr. 2 Jan. 1973, Friis et al. 2060 (K). – KEnya, South Mt Kenya, between Rupinganzi and Thiba R., fl. 16 Sept. 1951, Davis 60 (K); bridge over Gathiba R. below Kamweti Forest Station, fl. 9 Aug. 1971, Robertson 1565 (K); Western Province, Mumias, fl. & imm. fr. 11 May 1979, Bridson 80 (K). – rwanda, Rugezi, terr. Biumba, fl. 7 fr. 6 Oct. 1958, Van der Ben 2349 (BR, K); Butare Préfecture, Rubona, centre I.S.A.R., fl. 22 Oct. 1974, Troupin 15499 (BR, K); Butare, domaine de l’I.N.R.S., fl. 9 June 1978, Raynal 20379 (K, P). – South Sudan, Imatong Mts, between Gilo and Mt Konoro, fl. 18 Nov. 1980, Friis & Vollesen 316 (K). – tanzania, Bukoba, fl. June 1931, Haarer 2035 (K); Bukoba, fl. & fr. 18 Jan. 1959, Lind 2373 (K); Kahama Dist., fl. 16 June 1975, Kahuranga et al. 2781 (K). Conservation — Crepidorhopalon whytei is a widespread species with an EOO of 854 750 km 2 and is known from numerous localities. It would therefore qualify as of Least Concern under criterion B of IUCN (2012). That said, there are serious Richards & Arasululu 26150 (K) from the Uruwira-Mpanda road in southwest Tanzania is unusual in having markedly long calyces, 12.5–14 mm long with lobes 8.5 –10.5 mm long, compared to typical C. whytei with calyces 5 –10.5 mm and lobes 3 –7.5 mm long. This plant is otherwise a good match for other material of C. whytei. It would be interesting to see more material from western Tanzania to confirm if this difference is consistent; if so, it might be considered a regional subspecies. Plants of C. whytei are usually subglabrous or sparsely pubescent but occasional more densely pubescent plants are recorded across its range (e.g., Bagshawe 1476a, BM). The corolla usually has conspicuous eglandular hairs on the external surface of the upper lip but these can be absent in some populations, most notably in the isolated Ethiopian populations around Jimma; these plants are otherwise a close match for typical C. whytei. Skan (in Hemsley & Skan 1906) cited three specimens in the protologue of Lindernia whytei; Whyte s.n. is here selected from among these as the lectotype as it is an informative specimen and it was Whyte’s collection of this species that led Skan to choose the epithet. The holotype of Torenia mildbraedii is believed to have been destroyed in the bombing of the Berlin Herbarium; the extant duplicate at K comprises a small section of a leafy branch with a flower bud, a terminal branch section with immature leaves and an accompanying illustration of a dissected flower. This specimen is here selected as the lectotype of T. mildbraedii. Crepidorhopalon namuliensis I.Darbysh. & Eb.Fisch., sp. nov. — Fig. 1d – e, 2d – e, 3 Closely resembling Crepidorhopalon whytei but differing in the inflorescence and distal internodes having a conspicuous glandular-pubescent indumentum (vs inflorescence and stems essentially glabrous or eglandular-pubescent only, sometimes with scattered (sub)sessile glands); in the lower lip of the corolla being white or at most tinged pale blue (vs lower lip of corolla blue, purple or mauve); and in having the combined characters of axillary inflorescences and foliaceous bracts with corollas 13.5 –16.5 mm long (in C. whytei, if the bracts are foliaceous and so the flowers appear axillary, then the corolla is 16 – 24 mm long). — Type: Patel 7319 (holo K; iso LMA), Mozambique, Zambezia Prov., Namuli Mt, Muretha Plateau, fl. & fr. 25 May 2007. Etymology. Crepidorhopalon namuliensis is named after Mt Namuli, a key locality for plant diversity and endemism in Mozambique and the only known site for this species. 170 Blumea – Volume 64 / 2, 2019 i j h a c f g b e l n d k m Fig. 3 Crepidorhopalon namuliensis I.Darbysh. & Eb.Fisch. a. Habit; b. stem indumentum; c. detail of eglandular and glandular stem hairs; d. mature leaf, abaxial surface; e. bract, abaxial surface; f. pedicel and calyx; g. side view of calyx lobe showing marginal glandular hairs and hairs along midrib; h. dissected corolla with androecium; i. detail of hairs on exterior of corolla tube; j. detail of hairs on exterior of lower corolla lip; k. detail of palate of lower corolla lip; l. anterior stamen; m. pistil; n. mature fruit within calyx (a: Mphamba et al. 3, K; b – n: Patel 7319, K). — Scale bars: a = 3 cm; b, f, h, n = 2 mm; c, i – j = 250 μm; d – e = 1 cm; g, k– m = 1 mm. — Drawn by Andrew Brown. I. Darbyshire et al.: Crepidorhopalon whytei complex in eastern Africa Perennial herb, with numerous stems radiating from a central rootstock, at first erect but becoming scrambling or trailing, up to 50 cm long or more; stems brittle, markedly 4-angular, somewhat winged along the angles, with short spreading or ascending multicellular eglandular hairs and short spreading glandular hairs, the gland tip orange, becoming sparse on mature stems; internodes of fertile leafy stems 10 – 22 mm long when mature, those of sterile stems sometimes much longer. Leaves sessile, broadly ovate, 9 – 20 by 8 –18 mm, base rounded or shallowly cordate, margin serrate or shallowly so with 2 – 5 teeth per margin, apex acute or obtuse; primary venation palmate, with 5 or 7 main veins from base; sparsely puberulous with glandular and eglandular hairs mainly on margin and main veins beneath, surfaces also with sunken glands drying brown, most visible on lower surface. Flowers axillary, solitary; bracts foliaceous, only slightly reducing in size distally; bracteoles absent; pedicels 2.7– 5.5(– 6.5) mm long, shortly glandular-pubescent. Calyx 5–7.8 mm long, lobes lanceolate, somewhat unequal in length, longest lobes 3.3 – 4.2 mm long, apices can become divergent at maturity, 5-veined, somewhat hyaline between the veins, external surface shortly glandular-pubescent, margins of lobes eglandular-pubescent. Corolla 13.5 –16.5 mm long, tube and upper lip whitish to purple, lower lip white or at most tinged pale blue, with (?or without) a yellow patch in the mouth, external surface glandular-puberulous, upper lip also with pale longer eglandular hairs; tube 7.5 – 8.5 mm long, cylindrical but slightly contracted in midpoint where 1.3 – 2 mm wide and widened at mouth where 2 – 4 mm wide, internal surface with 4 lines of subsessile glands in proximal half; upper lip ovate, 3.5 – 5 by 4 – 5 mm, apex emarginate, internal surface with numerous minute glands; lower lip 6.5 – 8.5 mm long, 3-lobed, median lobe rounded-obovate, 3.7– 4 by 4.5 – 5.3 mm, lateral lobes rounded-elliptic, 4 – 4.5 by 4.2 – 5 mm, margins of lobes somewhat irregular, bosses on palate of lower lip with blunt-tipped multicellular hairs. Stamens 4, ventral stamens spurred, spurs clavate, 1.2 –1.75 mm long, papillate, filaments above spurred portion 4–4.3 mm long; posterior stamens with filaments 1.3–2 mm long; anthers of the two pairs of stamens adhering, thecae 0.7– 0.9 mm long. Ovary 1.3 –1.7 mm long, with sessile glands on surface; style 6.7– 8.3 mm long; stigma fan-shaped and minutely fringed. Capsule ellipsoid, brown, ± equal in length to calyx or slightly shorter, to 7.5 mm long, apex acute; seeds ± 1–1.15 by 0.75 mm, with longitudinal ridges and furrows. Distribution & Ecology — Crepidorhopalon namuliensis is apparently endemic to Mt Namuli in Zambezia Province of northern Mozambique. It occurs in montane grassland at 1730–1870 m elevation, favouring damp ground over rock where it can be associated with typical seepage assemblages including Cyperus, Xyris, Pimpinella and Senecio species together with Loudetia simplex (Nees) C.E.Hubb. (fide Mphamba 3). Additional specimens seen. MozaMBiquE, Zambezia Prov., Namuli Mt, Muretha Plateau, fl. & imm. fr. 24 May 2007, T. Harris 158 (K, LMA [not found]); Namuli Mt, Muretha Plateau, Mukocha Forest, fl. 28 May 2007, Timberlake 5063 (K, LMA [not found]); Namuli Mt, Muretha Plateau, fl. 17 Nov. 2007, Mphamba 3 (K, LMA); below base of Namuli Peak, Serra de Gurue, fl. 25 Apr. 2008, Würsten 186 (BR). Conservation — This species has been assessed as of Least Concern (Matimele et al. 2017). Although it is known only from Mt Namuli there are no known threats to this species or its habitat at present as the montane grasslands of Namuli are not subject to settled agriculture and the human impact is low. However, this species and the other Namuli grassland endemics should be carefully monitored as any disturbance or change to this habitat could quickly result in them becoming threatened with extinction. The impact from deliberate dry season burning should be carefully monitored in particular. 171 Notes — This species is separated from other members of the Crepidorhopalon whytei complex primarily by having a conspicuous glandular indumentum and pale corollas. In addition, the internodes of the majority of the stems are rather short such that the plants appear more densely leafy than is seen in other taxa within this complex. The corollas are typically smaller than those of forms of C. whytei that have foliaceous bracts, but they fall well within the overall range of corolla size in C. whytei (see Note to that species). Mt Namuli is emerging as a site of high importance for plant diversity and endemism, with 14 strict endemic species and two strict endemic subspecies now known. Most of these taxa, such as Coleus caudatus E.Downes & I.Darbysh. and Crotalaria namuliensis Polhill & T.Harris, are restricted to the montane grassland and wet flushes where Crepidorhopalon namuliensis is found (Harris et al. 2011, Downes & Darbyshire 2017). These habitats are particularly rich in flowering herbs and geophytes during the main rainy season (J. Timberlake, pers. comm., based on observations made on Mt Namuli in Feb. 2017), a time when there has been almost no botanical collecting on Namuli to date, hence the chances of discovering further new endemic species at this site are high (Downes & Darbyshire 2017). This locality is certain to qualify as an Important Plant Area (IPA) under the ongoing programme of IPA identification in Mozambique (see https://www.kew.org/science/projects/ tropical-important-plant-areas-tipas-mozambique; Darbyshire et al. 2017). Crepidorhopalon flavus (S.Moore) I.Darbysh. & Eb.Fisch., comb. nov. — Fig. 1f– h, 2f Lindernia flava S.Moore in J. Linn. Soc., Bot. 40: 153 (1911). — Type: Swynnerton 1966 (lecto BM [BM000930712], chosen here; isolecto K [K000379639]), Zimbabwe, near Chirinda, fl. 14 June 1906. Lindernia whytei sensu auct.: Philcox (1990) 62, p.p.; Philcox & Ghazanfar (2008) 70, p.p. Perennial herb, trailing or scrambling, laxly branched; stems brittle, markedly 4-angular, winged along the angles, glabrous or with few short eglandular hairs above each node, distal internodes sometimes with few sessile glands; internodes of leafy stems (9–)28–70 mm long when mature. Leaves sessile, broadly ovate, 9.5– 22 by 6 – 20 mm, base rounded or shallowly cordate, margin serrate or shallowly so with 1– 4 teeth per margin, apex acute or obtuse; primary venation palmate, usually with 5 main veins from base; glabrous except for few eglandular hairs on margin towards leaf base, surfaces also with sunken glands drying brown, most visible on lower surface. Flowers axillary, solitary, one flower per node, usually together forming a lax raceme; bracts either foliaceous or reducing in size and relative width distally, distal pairs of bracts often lanceolate-acuminate or ovate-acuminate, 2–6 by 0.7–4.5 mm, the pairs often markedly uneven in size, the bract subtending the flower smaller and narrower; bracteoles absent; pedicels 2 –7.5(–10.5) mm long, glabrous, with sparse sessile glands. Calyx 5 – 9.5 mm long, lobes lanceolate, somewhat unequal in length, longest lobes 3.2 – 6.5 mm long, becoming divergent at maturity and with apices somewhat recurved, 5-veined, tubular portion hyaline between the veins, margins of lobes eglandular-pubescent towards base, elsewhere glabrous, with few sessile glands. Corolla 10 –14 mm long, tube and upper lip yellow-brown or upper lip slightly purple-tinged, lower lip bright yellow, external surface glandular-puberulous, upper lip with pale longer eglandular hairs; tube 5.7– 8.5 mm long, cylindrical towards base where 0.9 –1.2 mm wide, widened towards mouth where 2 – 2.5 mm wide, internal surface with 4 lines of subsessile glands in proximal half; upper lip ovate, 2.3 – 3.5 by 2.3 – 3 mm, apex emarginate, internal surface with numerous 172 glands; lower lip 4.3–6 mm long, 3-lobed, median lobe roundedobovate, 2.5 – 3.5 mm long and wide, lateral lobes somewhat smaller, bosses on palate of lower lip with blunt-tipped multicellular hairs. Stamens 4, ventral stamens with clavate spurs, 0.5 – 0.8 mm long, papillate, filaments above spurred portion 2.7–3.5 mm long; posterior stamens with filaments 1.2 –1.4 mm long; anthers of the two pairs of stamens adhering, thecae 0.6 – 0.9 mm long. Ovary 1.1–1.4 mm long, with sessile glands on surface; style 4–6 mm long; stigma fan-shaped and minutely fringed. Capsule narrowly ellipsoid or more gradually tapered, brown, ± equal in length to calyx, 5.5 – 9.5 mm long; seeds ellipsoid, ± 0.5 – 0.6 by 0.45 – 0.5 mm, longitudinal ridges very shallow and inconspicuous. Distribution & Ecology — This species is recorded in swamps, riverbanks and seasonally wet open grassland with sandrich peaty soils, including swampy clearances in forest, at 330 –1130(–1340) m elevation. Additional specimens seen. MozaMBiquE, Manica Prov., Mt Maruma, fl. & fr. 13 Sept. 1906 [syntypes], Swynnerton 1922 (BM, K); Dombe, entre Chissanto e Zinesse, fl. & fr. 24 Nov. 1965, Pereira & Marques 876 (BR, WAG*); E of Makurupini R., fl. 11 June 1971, Biegel 3583 (WAG*); Sussendenga Dist., eastern foothills of Chimanimani Mts, c. 28 km W of Dombe, fl. & fr. 25 Apr. 1974, Pope & Müller 1295 (K); Chimanimani Mts foothills, near Zomba Community, Magorogodo hill, fl. 27 Oct. 2013, Würsten & Dondeyne BE875 (BR); Chimanimani foothills, Maronga, Wandowani, W of Comeni’s compound, fl. 14 Nov. 2015, Darbyshire 895 (K, LMA, Ndzou); Maronga, between Comeni’s compound and Murere River, fl. 15 Nov. 2015, Darbyshire sight record. – ziMBaBwE, Manicaland Prov., Melsetter [Chimanimani], fl. 7 fr. 1 Oct. 1919, Walters 2728 (K); ibid., Walters 2758 (K); Umtali [Mutare], fr. Oct. 1937, Brain s.n. in SRGH 10828 (SRGH*) - see note; Tarka Forest Reserve, banks of Chisengu R., fl. & fr. May 1968, Goldsmith 72/68 (K, P*). Conservation — Crepidorhopalon flavus is a highly rangerestricted species, known only from the southern Manica Highlands of the Mozambique-Zimbabwe border region, where it is centred on the foothills of the Chimanimani Mountains. The EOO is 4 440 km 2, or only 646 km 2 if the Walters and Brain specimens are excluded in view of the fact that it is unclear as to whether the former was collected near Chimanimani town or in the Chimanimani Mountains and there is some uncertainty over the latter collection (see note). Six to eight locations are defined based on threats. Those sub-populations that lie within the formally protected areas of the Chimanimani Mts – the Chimanimani National Park in Zimbabwe and the Chimanimani National Reserve in Mozambique – are likely to be secure. The only activity recorded recently in the seasonal wetlands within the National Reserve was cutting of grass for thatching (I. Darbyshire, pers. obs.) and this is unlikely to impact the Crepidorhopalon significantly. However, the majority of locations lie outside of these protected areas and there has been much disturbance and destruction of natural habitats within its range due to high human population pressure and continued expansion of subsistence agriculture. This includes areas of seasonally wet soils which, away from areas of quartzite, are fertile and so favoured for cultivation (B. Wursten, pers. obs.). Some sub-populations may also have been impacted by artisanal gold mining activities along some of the rivers and streams in the southern foothills of the Chimanimani Mountains. With a continuing decline in extent and quality of habitat inferred, and with fewer than 10 locations known within an EOO of considerably less than 20 000 km 2, this species is assessed as Vulnerable under criterion B1 – VU B1ab(iii). Notes — In view of the consistently different flower colour, coupled with the marked range (and hence genetic) disjunction and differing habitat requirements of the Zimbabwe/Mozambique plants, it is considered most appropriate to reinstate Lindernia flava as a good species and the new combination in Crepidorhopalon is made here. This species is otherwise very similar to C. whytei although the seeds differ slightly in having Blumea – Volume 64 / 2, 2019 almost imperceptible longitudinal furrows, these being clearly visible in C. whytei (see Fig. 2). In the specimen citations under Lindernia whytei, Philcox (1990) wrongly attributes Swynnerton 1922 as having been collected from ‘serra do Gúruè, E of Picos Namuli, near source of R. Malema, c. 1800 m’; it was actually collected from Mt Maruma in Manica Province. The specimen Brain s.n. at SRGH does not have any flowers and there are no notes on flower colour; it is recorded from further north than the other collections and at a higher altitude (the maximum elevation recorded above). It is strange that this species has not been re-recorded from the well-botanised area around Mutare but there is no reason to doubt the provenance of this specimen. Further material from the Mutare area would be useful to confirm the identity of this specimen. When describing Lindernia flava, Moore (1911) noted the close similarity to his L. gossweileri S.Moore (1907: 87) from near Capopa in Malange Province, Angola (Map 1), but said that they differ in L. gossweileri having calyces divided to the base and having a larger corolla with a longer tube and with the lips almost equal in size rather than markedly unequal. To our knowledge, L. gossweileri has never been recollected in Angola and the type specimen (Gossweiler 1086, BM [BM000930701], K [K000379630]) is rather scanty, such that it is difficult to draw firm conclusions on its relationship to C. flavus. However, the differences noted by Moore seem to be rather minor and that of differing flower size and proportions may not be of great significance in view of the differences in flower size recorded in other species of Crepidorhopalon including C. whytei. Therefore, it is possible that these two yellow-flowered taxa represent a single species, although the extreme range disjunction seems improbable. More material from Angola is needed for confirmation, but if they do prove to be the same species then L. gossweileri would have nomenclatural priority and the epithet is unoccupied in Crepidorhopalon. The wet grasslands in the foothills of the Chimanimani Mountains, the stronghold for this species, are of wider botanical interest as they hold several interesting species besides C. fla­ vus. Of particular note is Mesanthemum africanum Moldenke which is endemic to the Chimanimani Mountains, occurring in damp sites on quartzitic sands both at higher altitudes and (at much lower abundance) in the foothills, probably a result of small populations being established from seed washed down along rivers from the high massif. During surveys of the Maronga area of Mozambique in 2015, C. flavus was found growing alongside two species of Xyris (Xyridaceae). The first of these proved to be X. angularis N.E.Br. (Darbyshire 918), a widespread West African species that had not previously been recorded from Mozambique but is known from the Zimbabwe side of Chimanimani (Lock 2010); the Chimanimani population is a significant outlier for this species. The second Xyris species collected (Darbyshire 938) is an unmatched, potentially new species. The first Mozambican records of Edrastima angolen­ sis (K.Schum.) Neupane & N.Wilkstr. (Darbyshire 920) in the Rubiaceae and Fimbristylis aphylla Steud. in the Cyperaceae (Darbyshire 926) were also made at the same wetland site (Timberlake et al. 2016). Crepidorhopalon sp. aff. whytei (= Manktelow et al. 91081) Straggling herb, stems markedly 4-angular, glabrous except for inconspicuous sessile orange glands. Leaves sessile, broadly ovate, 11.5 –17 by 8 –13.5 mm, base rounded or subcordate, margin with 2 – 3 teeth along each margin, apex acute or obtuse; primary venation palmate, with 5 main veins from base; surfaces glabrous except for sunken glands drying brown, most conspicuous on lower surface. Flowers axillary, solitary; bracts I. Darbyshire et al.: Crepidorhopalon whytei complex in eastern Africa foliaceous, only slightly reducing in size distally; bracteoles absent; pedicels 6 –11 mm long, glabrous. Calyx 6.5 – 8.5 mm long, lobes lanceolate, somewhat unequal in length, longest lobes 4.5 – 5.5 mm long, becoming recurved at maturity, surfaces glabrous. Corolla 8.5 –10 mm long, colour unknown, external surface glabrous; tube 4.5 – 6 mm long, cylindrical, ± 1.4 mm wide centrally; upper lip rounded-ovate, 1.8–2.6 mm long, margin irregular, internal surface with minute glands; lower lip ± 3.8 mm long, 3-lobed, median lobe rounded-obovate, ± 2.3 mm long, margins of lobes somewhat irregular, bosses on palate of lower lip with blunt-tipped multicellular hairs. Stamens 4, ventral stamens spurred, spurs shortly clavate, ± 0.8 mm long, papillate, filaments above spurred portion ± 4 mm long; posterior stamens with filaments ± 0.7 mm long; anthers of the two pairs of stamens adhering, thecae ± 0.5 mm long. Ovary not seen; style ± 5 – 5.5 mm long; stigma fan-shaped and minutely fringed. Capsule not seen. Distribution & Ecology — Known only from a single collection from the Nguru Mts where it was recorded from the ericaceous shrub zone growing with Xerophyta sp. at 1940–2010 m elevation. Specimens seen. tanzania, Morogoro, Nguru Mts, on top of Kwasenjuga, 2.5 km S Maskati Mission, fl. 1 Feb. 1991, Manktelow et al. 91081 (K). Note — The specimen cited is close to forms of C. whytei with leafy bracts, but differs in having very small corollas to only 10 mm long with the tube only 4.5 – 6 mm long and lacking conspicuous short glands on the external surface of the distal portion of the tube. Unfortunately, the flower colour is not recorded on the specimen label, but the flowers have dried rather pale compared to specimens of typical C. whytei which often retain some of their colour. It is possible that the flowers were yellow as in C. flavus, or white as in C. namuliensis but it differs from both those species in having smaller flowers which lack conspicuous short glands on the external surface of the distal portion of the tube, and also differs from the latter in lacking glandular hairs on the stems, pedicels and calyces. More material is needed to investigate this population further. However, the isolated location is notable, and the Nguru Mountains – a part of the ancient Eastern Arc mountain chain – contain many endemic and near-endemic species, for example Impatiens messumbaensis G.M.Schulze in Balsaminaceae (Grey-Wilson 1982), Pilea nguruensis Friis & I.Darbysh. in Urticaceae (Friis et al. 2015) and Streptocarpus bambuseti B.L.Burtt, S. burttianus Pócs and S. stomandrus B.L.Burtt. in Gesneriaceae (Darbyshire 2006). Crepidorhopalon kwaleensis I.Darbysh. & Eb.Fisch., sp. nov. — Fig. 2i–k, 4, 5 Most similar to Crepidorhopalon whytei but differing in the anterior pair of stamens having markedly longer, slender spurs 1.8 – 2.4 mm long (vs spurs 0.5–1.2 mm long), in the seeds being no longer than wide and with subcircular depressions on the surface (vs seeds longer than wide, with longitudinal furrows on the surface), and in the plants being erect or decumbent (vs procumbent, scrambling, trailing or at most weakly decumbent in C. whytei ). Also potentially confused with C. hepperi Eb.Fisch. but again differing in the longer and more slender staminal spurs (these 0.5 – 0.7 mm long in C. hep­ peri), the differing seeds (those of C. hepperi similar to C. whytei but more angular and with more conspicuous minute transverse furrows perpendicular to the longitudinal furrows – Fig. 2), and in having usually more broadly ovate leaves with length : width ratio 0.9 –1.4 : 1 (vs leaves ovate to lanceolate, length : width ratio 1.35 – 3.4(– 4.9) : 1 in C. hepperi). — Type: Luke 3796 (holo K; iso EA [not located], MO [not seen], US [not seen]), Kenya, Kwale County, Majoreni area, 5 km NE, fl. & fr. 18 Aug. 1993. Etymology. Crepidorhopalon kwaleensis is named after Kwale County of Kenya, where this species is endemic. Lindernia whytei sensu auct.: Luke (2005) 90; Philcox & Ghazanfar (2008) 70, p.p. quoad Drummond & Hemsley 4007. 173 Lindernia hepperi sensu Philcox & Ghazanfar (2008) 72, p.p. quoad spec. ex Kenya. Annual (possibly sometimes short-lived perennial) herb, with one to few stems from a small rootstock, erect or becoming scrambling or trailing, up to 20 – 40 cm long, branching in proximal half; stems markedly 4-angular, winged along the angles, proximal portions of stems pilose with ± numerous pale multicellular eglandular hairs 0.3 –1 mm long, sometimes with few interspersed glandular hairs, hairs becoming more sparse distally, fertile portions of stems (raceme rachis) ± glabrous except for inconspicuous sessile orange glands; internodes of fertile leafy stems 16 – 44(– 90) mm long when mature. Leaves sessile, broadly ovate, 12.5 –18 by 9 –16 mm, base rounded or subcordate, margin with 3 – 5 minute teeth along each margin, apex acute, attenuate or obtuse; primary venation palmate, with 5 or 7 main veins from base; ± sparsely eglandular pilose on margin and main veins beneath, surfaces also with sunken glands drying brown, visible on lower surface. Flowers held in a lax terminal raceme, each inflorescence node singleflowered; proximal-most pair of bracts ovate to suborbicular and acuminate, 3.8 –7.5 by 2.2 – 6 mm, rapidly reducing upwards where lanceolate or linear-lanceolate, 1.5 – 2.7 by 0.3 – 0.8 mm; bracteoles absent; pedicels 1.5 – 2.8 mm long, glabrous. Calyx 3.7– 5.5 mm long, lobes lanceolate, somewhat unequal in length, longest lobes 2 – 4 mm long, apices can become divergent at maturity, 5-veined, hyaline between the veins, surfaces glabrous. Corolla (8 –)10.5 –12 mm long, purple, lower lip with white patches in the mouth, external surface glandular-puberulous; tube 5.2 – 6 mm long, cylindrical, ± 1.5 mm wide centrally, internal surface with 4 lines of subsessile glands in proximal half; upper lip ovate, (2 –)2.5 – 4.3 mm long, apex slightly emarginate, internal surface with minute glands; lower lip (3.5 –)5 – 6 mm long, 3-lobed, median lobe roundedobovate, 3–3.5 mm long, lateral lobes elliptic-obovate, 2.3–3.5 mm long, margins of lobes somewhat irregular, bosses on palate of lower lip with blunt-tipped multicellular hairs. Stamens 4, ventral stamens spurred, spurs linear-clavate, 1.8 – 2.4 mm long, apex papillate, filaments above spurred portion ± 4 mm long; posterior stamens with filaments ± 2 mm long; anthers of the two pairs of stamens adhering, thecae 0.45 – 0.7 mm long. Ovary not seen; style ± 4.5 mm long; stigma fan-shaped and minutely fringed. Capsule narrowly ellipsoid to ovoid with a gradually tapered apex, green-brown, longer than calyx, 5 – 9 mm long; seeds sub-square in face view and circular in lateral view, ± 0.4–0.5 mm diam, with a row of subcircular depressions and with a marked longitudinal furrow on the posterior side. Distribution & Ecology — Crepidorhopalon kwaleensis is known only from the coastal lowlands of southeast Kenya in Kwale County. It occurs in seepage areas and pools in grassland, palm swamps and grassland along forest margins, at 5 – 230 m elevation. At one site, it was found growing along the margin of a pond used by mammals as a watering hole (Drummond & Hemsley 4007). Additional specimens seen. KEnya, Mwasangombe Forest, 15 miles SW of Kwale, fl. & fr. 27 Aug. 1953, Drummond & Hemsley 4007 (K); Kwale County, Gazi village N, fl. & fr. 12 Oct. 1991, Luke 2899 (EA [not located], K); near Lunguma, fl. & imm. fr. 20 Aug. 1994, Luke & Gray 4061 (EA [not located], K); Majoreni area, 5 km NE, fl. & fr. 18 Aug. 1993, Luke 3796 (EA [not located], K); Ramisi-Lungalunga Rd before Majoreni turn-off, fl. 20 Dec. 2018, Luke et al. 18735 (EA). Conservation — Based on the occurrence records cited above and two additional sight records (Q.L.), this species has an EOO of 691 km2 and an AOO of 28 km 2. The area in which most of the localities are found (near Majoreni) is undergoing rapid habitat conversion to sugar plantations and maize fields (see Fig. 5d), with some sub-populations almost certainly having been lost. The forest patch that adjoined the most northern 174 Blumea – Volume 64 / 2, 2019 i e a d b c f g h Fig. 4 Crepidorhopalon kwaleensis I.Darbysh. & Eb.Fisch. a. Habit; b. stem indumentum; c. mature leaf, abaxial surface; d. flower, face view; e. flower, posterior view with upper lip folded back to show stamens and stigma; f. stamens, anterior pair to the right; g. partial inflorescence with bracts, calyx and immature fruit; h. mature fruit in calyx; i. seeds in (l to r) anterior, apical and posterior views (a, e – f, h: Drummond & Hemsley 4007, K; b – c, g, i: Luke 3796, K, d from photo by Q. Luke). — Scale bars: a = 3 cm; b = 1 mm; c, d = 1 cm; e, g – h = 5 mm; f = 2 mm; i = 500 μm. — Drawn by Andrew Brown. I. Darbyshire et al.: Crepidorhopalon whytei complex in eastern Africa 175 Fig. 5 Crepidorhopalon kwaleensis. a – b. Habit; c. detail of flowers showing long abaxial staminal appendages; d. recent destruction of habitat for C. kwa­ leensis near Majoreni, Kwale County. — Photos: W.R.Q. Luke. record (Kaya Lunguma) has been destroyed. There is only one locality inside a protected area (Mwasagombe Forest, now believed to be inside the Shimba Hills National Reserve) but it has not been recollected there since 1953. With four threat-defined locations identified, this species is assessed as Endangered under criterion B – EN B1ab(i,ii,iii,iv,v) + B2ab(i,ii,iii,iv,v). does superficially look somewhat intermediate between these two species, but clearly differs from both in the stamen and seed characters mentioned above. It additionally differs from C. hepperi in having broader leaves; although the leaf shape in C. hepperi is somewhat variable, it typically has lanceolate or more narrowly ovate leaves. Notes — This species is most easily recognised by the combination of the long slender spurs on the anterior pair of stamens, longer than in the other species of the C. whytei complex, and in the striking seeds that are no longer than wide and with subcircular depressions, rather than ± longer than wide and with longitudinal furrows on the surface in other members of this group (all species have a more marked longitudinal furrow on the posterior side). It is otherwise similar to forms of C. whytei that have a racemose inflorescence due to the bracts being markedly reduced in relation to the cauline leaves, but the growth habit is more erect or decumbent than in C. whytei where the stems are trailing. The coastal lowlands of southeast Kenya are known for their botanical interest, with a variety of endemic and range-restricted species. These include species restricted to similar habitats to C. kwaleensis. For example, the pteridophyte Marsilea fade­ niana Launert is known only from seasonal waterholes and dry riverbeds around Kwale (Launert 2003), while in the Cy­ peraceae, Cyperus boreobellus Lye is restricted to damp soils over rock and small pools in the Kwale region and Bulbostylis afroorientalis (Lye) R.W.Haines is known with certainty only from the seasonally wet grasslands of this region (Hoenselaar et al. 2010). In the ‘Flora of Tropical East Africa’ account of Lindernia (Philcox & Ghazanfar 2008), material here assigned to this new taxon was treated under two different species – L. whytei and L. hepperi. Crepidorhopalon hepperi is easily separated from C. whytei due to its more erect, short, much-branched habit, narrower leaves, smaller flowers and calyces with the lobes usually shorter than the tube. However, C. kwaleensis Acknowledgements We thank Andrew Brown for producing the illustrations of the new species. Ranee Prakesh (BM), Peris Kamau (EA), Maria Romeiras (LISC), Inês Chelene (LMA) and Anthony Mapaura (SRGH) are thanked for checking and (where appropriate) providing access to the relevant herbarium specimens at their institutions. We are grateful to Stephen Manktelow (Swedish Species Information Centre, Agricultural University of Sweden, Uppsala) for checking his notes on his collection of Crepidorhopalon sp. aff. whytei from the Nguru Mountains in Tanzania. I.D. would also like to thank Federico 176 Fabriani (K) for his assistance with the use of the microscope camera set-up when taking images of the seeds of Crepidorhopalon. Fieldwork on Mt Namuli in 2007 that resulted in the discovery of Crepidorhopalon namuliensis was sponsored by the Darwin Initiative award 15/036: ‘Monitoring and Managing Biodiversity Loss in South East Africa’s Montane Ecosystems’ completed in 2009. 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