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Nordic Journal of Botany 31: 113–117, 2013 doi: 10.1111/j.1756-1051.2012.01804.x, © 2013 he Author. Nordic Journal of Botany © 2013 Nordic Society Oikos Subject Editor: Martin Cheek. Accepted 23 August 2012 Warneckea cordiformis sp. nov. (Melastomataceae–Olisbeoideae) from a coastal dry forest in northern Mozambique Robert Douglas Stone R. D. Stone (stonerd@ukzn.ac.za), School of Life Sciences, Univ. of KwaZulu-Natal, Private Bag X01, Scottsville 3209, South Africa. Described and illustrated is Warneckea cordiformis R. D. Stone, an evidently localized endemic of coastal dry forest in Mozambique’s Cabo Delgado Province. In ‘Flora Zambesiaca’ the new species would key to W. sansibarica (Taub.) Jacq.-Fél., from which it is readily distinguished by the much smaller, ovate to cordiform leaves and white, short-pedicellate lowers. Because of its evidently very limited occurrence as well as on-going anthropogenic threats, Warneckea cordiformis is here assessed as ‘Critically Endangered’ (CR) B1a,b(iii) according to IUCN criteria. A key is provided to the Mozambican species of Warneckea. he genus Warneckea Gilg comprises about 50 species of shrubs or small trees restricted to humid forests in Africa, Madagascar and Mauritius. Further morphological and molecular studies have shown that it is distinct from Memecylon L. (Jacques-Félix 1978b, Stone 2006, Stone and Andreasen 2010). In the treatment of Memecylon s.l. for the ‘Flora Zambesiaca’ (Fernandes and Fernandes 1978), the species with strongly trinervate to multinervate leaves are properly placed in Warneckea, except for M. zambeziense A. Fern. & R. Fern. which belongs to Lijndenia Zoll. & Moritzi, perhaps as a synonym of L. barteri (Hook. f.) K. Bremer (cf. Jacques-Félix 1983, p. 156). Additional characteristics distinguishing Warneckea from Memecylon s.s. include the often well-developed calyx lobes, imperfectly 2-loculed ovary, embryo with leshy cotyledons, and foliar sclereid cells spheroid or lacking altogether (Jacques-Félix 1977, 1978a, 1978b, Jacques-Félix et al. 1978, Rao and Jacques-Félix 1978). Warneckea was recently subdivided into subgenus Warneckea (with three sections) and subgenus Carnosae (Jacq.-Fél.) R. D. Stone, the latter group characterized by sessile inlorescences with long-pedicellate lowers subtended by persistent, imbricate-decussate bracts; hypanthium with free limb and calyx lobes obsolete; and embryo with two leshy but unequal cotyledons (JacquesFélix 1985a, 1985b, Stone and Andreasen 2010). Subgenus Carnosae is found mainly in the east-African coastal forests and northwestern Madagascar, with outlying populations in Malawi, Zimbabwe and Zambia. Previous authors have considered it to include just one broadly deined species, W. sansibarica (Taub.) Jacq.-Fél., perhaps further separable into two subspecies or varieties (Wickens 1975, Fernandes and Fernandes 1978, 1980, Jacques-Félix 1985a, 1985b, Borhidi 1993). New indings suggest that W. sansibarica as previously circumscribed includes several distinct but closely related species. For example, in the Arabuko-Sokoke forest of coastal Kenya, W. sansibarica is sympatric with a smallleaved, shrubby form known as W. melindensis (A. Fern. & R. Fern.) R. D. Stone & Q. Luke. here is no evidence of hybridization between these taxa, and they are readily discernible at both the morphological and DNA sequence levels (Vollesen 1980, Luke 2005, Stone and Luke 2009, Stone and Andreasen 2010). Recent investigations have further revealed that W. sousae (A. Fern. & R. Fern.) A. E. van Wyk, which was placed by Wickens (1975) in synonymy under W. sansibarica, is only supericially similar to that species and is properly placed in W. sect. Warneckea (Vollesen 1980, Stone and Andreasen 2010). Pending a comprehensive revision of W. subgenus Carnosae, the current work describes a distinctive new species belonging to this group, discovered during recent ield studies in the coastal forests of northern Mozambique. All cited specimens have been seen unless otherwise indicated. he conservation assessment follows the criteria of IUCN (2001). A key is provided to the currently recognized species of Warneckea in Mozambique. Warneckea cordiformis R. D. Stone sp. nov. (Fig. 1) Type: Mozambique, Cabo Delgado, west of Quiterajo on track along southern margins of Namacubi forest, 11°46′05.3″S, 40°23′50.6″E, alt. ca 90 m a.s.l., heavily disturbed dry forest with Pteleopsis, 21 Nov 2009, Goyder et al. 6095 (holotype: K!, isotypes: LMA!, NU!, P!). Evergreen shrub or small tree 2–3 m high. Young branchlets reddish to purplish brown, quadrangular and 113 Figure 1. Warneckea cordiformis sp. nov. (A) lowering branch, (B)–(D) leaves, (E) detail of lower leaf surface, (F) lower (with 1 petal and 1 stamen removed). Voucher specimens: (A), (D)–(F) Goyder 6095, NU; (B)–(C) J. Burrows and S. Burrows 10773, BNRH. Scale bars: (A) 20 mm, (B)–(D) 10 mm, (E) 5 mm, (F) 2 mm. 114 narrowly 4-winged, with age becoming grayish white and rounded; internodes (0.2–)0.5–2.0(–4.5) cm long. Leaves coriaceous, 3-nerved from the base (or the larger leaves with an additional pair of weaker, submarginal nerves), dark green on the upper surface, somewhat paler beneath; petioles 1–2(–3) mm long, adaxially grooved; blades ovate to cordiform, (1.2–)1.5–3.0(–4.0) cm long, 1.2–2.2(–3.0) cm wide, the base varying from broadly cuneate to rounded to strongly cordate (the sinus between the basal lobes up to 3 mm deep), the apex shortly and obtusely acuminate; midnerve impressed on the upper surface, prominent on the lower, often extending past the leaf apex as a short mucro; lateral nerves prominent on both surfaces (in dried material), curvilinear toward the base of the blade, forming shallow arches and becoming progressively weaker towards the leaf apex; lower surface of blade inely reticulate-areolate owing to a conspicuous network of purplish veinlets. Cymules sessile, few-lowered, agglomerated at the recently defoliated nodes just below the actual leaves (rarely in the leaf axils); lowers white, pedicellate, the pedicels 1.5– 3.0 mm long, subtended by several pairs of imbricatedecussate bracts, the bracts 0.5–1.0 mm long, ฀rhombic in outline, cucullate and ฀keeled toward the apex on the abaxial side; hypantho-calyx cupulo-patellate, 1.25 mm long  2.00 mm wide, the epigynous limb well-developed and the calyx lobes obsolete; petals spatulate, 2.0 mm long  1.5 mm wide; staminal ilaments pale blue, ca 4 mm long; anthers ca 0.7 mm long, the connective strongly incurved by an ellipsoid, dorsal gland; style iliform, ca 5 mm long. Flowering in late November, fruits not seen. Distribution and habitat Warneckea cordiformis is evidently a localized endemic known only from the Namacubi (Banana) forest west of Quiterajo, Cabo Delgado province, northern Mozambique (Fig. 2). According to data provided on specimen labels, Figure 2. Geographic distribution of Warneckea cordiformis sp. nov. in Mozambique. he only known locality is indicated with a solid circle (●). Provincial boundaries are indicated by solid lines, and selected cities by open squares ( ). 115 the habitat is in dry, semi-deciduous coastal forest dominated by Guibourtia schliebenii (Harms) J. Léonard, on sandy soil at altitudes of 90–125 m a.s.l. Within its limited area of distribution the species is locally abundant (J. Burrows, Bufelskloof Herbarium, pers. comm.). Conservation status he original extent of coastal forest in the Quiterajo vicinity may have been 300–500 km2, but the remaining forest patches are now highly fragmented due to clearance for traditional agriculture over the past 200–400 years (Timberlake 2009). he present extent of dry forest in the Rio Messalo–Quiterajo area is 55 km2 with on-going anthropogenic threats, e.g. clearing for subsistence agriculture, logging and uncontrolled ires (Timberlake et al. 2011). Accordingly, W. cordiformis is here assessed as ‘Critically Endangered’ (CR) B1a, b(iii) according to IUCN (2001) criteria. Similar species and diagnostic characters Warneckea cordiformis is placed in W. subgenus Carnosae on account of its combination of leaf blades with conspicuous network of purplish veinlets on the lower surface; sessile inlorescences agglomerated at the recently defoliated nodes; loral pedicels subtended by persistent, imbricate-decussate bracts; patelliform calyx limb with lobes obsolete; and anther connectives bearing a minute, dorsal oil-gland (Stone and Andreasen 2010). he new species would key to W. sansibarica in ‘Flora Zambesiaca’ and ‘Flora de Moçambique’ (Fernandes and Fernandes 1978, 1980), but is readily distinguished from W. sansibarica and other members of W. subgenus Carnosae by its relatively small, ovate to cordiform leaves and white lowers borne on short pedicels. In contrast, the leaves of W. sansibarica are ฀elliptic and 4.0–10.5  2.0–6.0 cm with cuneate to rounded bases and apices฀  distinctly acuminate; the loral pedicels are slender and ca 6–15 mm long; and the lower color is generally pale blue to deep blue. Amongst the other members of W. subgenus Carnosae, W. cordiformis is most similar to W. melindensis, which is known from three coastal forest sites in Kenya and two in Tanzania (Fernandes and Fernandes 1955, 1960, Vollesen 1980, Luke 2005, Stone and Luke 2009). Warneckea melindensis and W. cordiformis share a฀ ฀shrubby habit, small leaves, and young branchlets quadrangular and narrowly winged (characteristics not found in W. sansibarica). he two species difer in the shape of their leaf bases (rounded to cuneate in W. melindensis vs฀ ฀cordate in W. cordiformis), as well as the length of their loral pedicels (5–7 mm vs 1.5–3.0 mm). According to data provided on specimen labels, the Kenyan populations of W. melindensis have blue lowers while at least one of the Tanzanian populations is white-lowered. Also occurring in the same Namacubi forest near Quiterajo is another Warneckea species that appears close 116 to W. sansibarica, except that the lowers are white (vs pale to deep blue) and the lowering pedicels are relatively short (3.5–4.0 mm vs 6–15 mm). In comparison to W. cordiformis it is a larger tree (5–8 m) with leaves that are larger and diferently shaped (elliptic-lanceolate, distinctly acuminate at apex, 3.8–5.2  1.7–2.5 mm). his species may warrant formal recognition, but this needs further study. It is represented by the collections J. Burrows and S. Burrows 10753, 10765 and 10833 (BNRH) and Goyder et al. 6151 (K, LMA). Yet another Warneckea species found in the Namacubi forest is W. sousae, which supericially resembles W. sansibarica in leaf size and shape, and in having sessile inlorescences with pedicellate lowers subtended by persistent, imbricatedecussate bracts. In W. sousae however the lowers are creamy white and the anthers are lacking a dorsal gland (an apparent synapomorphy for Warneckea sect. Warneckea; cf. Stone and Andreasen 2010). It is represented by the collections J. Burrows and S. Burrows 10703, 10771 and 10772 (BNRH) and Goyder et al. 6091 (K, LMA). Supericially resembling W. cordiformis is another collection, Pedro and Pedrógão 5170 (COI n.v., LMA), made in Guibourtia sand forest at Macomia, between Ingoane and Quiterajo. Previous authors had compared these sterile specimens with the Tanzanian species W. microphylla (Gilg) Borhidi or the Kenyan species W. mouririifolia (Brenan) Borhidi (Memecylon microphyllum sensu Fernandes and Fernandes 1969; Memecylon sp. C sensu Fernandes and Fernandes 1978; Memecylon sp. A sensu Fernandes and Fernandes 1980). I have re-examined the material of Pedro and Pedrógão 5170 at LMA and have determined that it belongs to Strychnos L. (Loganiaceae), not Warneckea. Both genera have trinervate leaves and are easily confused in the absence of lowers or fruits. Other east-African Warneckea species with relatively small leaves include the Kenyan W. mouririifolia, the Tanzanian W. microphylla and W. schliebenii (Markgr.) Jacq.-Fél. and the Mozambican W. sessilicarpa (A. Fern. & R. Fern.) Jacq.-Fél. All of these species belong to Warneckea sect. Warneckea (sensu Stone and Andreasen 2010), hence their resemblance to W. cordiformis is supericial. Additional specimens examined (paratypes) Mozambique, Cabo Delgado, Namacubi Forest (the Banana Forest), west of Quiterajo, 11°45′55″S, 40°23′45″E, 90 m a.s.l., dry coastal sand forest dominated by Guibourtia schliebenii, 25 Nov 2008, J. Burrows and S. Burrows 10773 (BNRH); track through middle of Namacubi (Banana) Forest, 11°45′46″S, 40°20′19″E, 125 m a.s.l., northern edge of mixed semi-deciduous forest, sandy soil, aspect level, 27 Nov 2008, J. Burrows and S. Burrows 10837 (BNRH); Quiterajo, 11.76764°S, 40.37427°E, 115 m a.s.l., deciduous coastal forest and thicket, 19 Dec 2009, Luke 13887 (EA, K n.v., LMA n.v., P n.v.). Key to the species of Warneckea in Mozambique 1. Outer bark of trunk and older branches pale reddish brown, exfoliating in patches to reveal whitish inner bark; leaves with purplish veinlets forming a ine, reticulateareolate network on the lower surface; inlorescences  sessile, borne in dense, few- to many-lowered fascicles in the leaf axils and at the thickened nodes of the upper branchlets; bracts persistent, imbricate-decussate; lowers distinctly pedicellate; loral cup patelliform with calyx lobes short or obsolete; anther connectives with a dorsal oil-gland; embryo with two leshy but unequal cotyledons (Warneckea subg. Carnosae) …………………… 2 – Outer bark of trunk and older branches grey, inely longitudinally issured; venation pattern on lower leaf surface as above or with reticulations not so close; inlorescences and bracts as above; lowers sessile or shortpedicellate; loral cup obconic to campanulate with calyx lobes generally well-developed and concealing the corolla in bud, the bases of the outer pair of lobes often auriculate-amplexate; anther connective-gland absent; embryo with just one leshy cotyledon, the second obsolete (Warneckea sect. Warneckea) …………………… 3 2. Shrub or tree to 10 m high; young branchlets terete; leaves elliptic, 4.0–10.5  2–6 cm, bases cuneate to rounded, apices  distinctly acuminate; lowers pale blue to deep blue, borne on slender pedicels ca 6–15 mm long …………………………………… W. sansibarica – Shrub or small tree 2–3 m high; young branchlets quadrangular and narrowly 4-winged; leaves ovate to cordiform, 1.5–3.0(–4.0) 1.2–2.2(–3.0) cm, bases broadly cuneate to rounded or strongly cordate, apices shortly and obtusely acuminate; lowers white, borne on pedicels 1.5–3.0 mm long ………… W. cordiformis 3. Leaves elliptic to broadly elliptic, ovate, or almost circular, 3–9  1.5–6.5 cm; petioles 2–5 mm long; reticulations of lower leaf surface not close; lowers and fruits borne on pedicels ca 4 mm long …………… W. sousae – Leaves ovate to elliptic, (1.5–)2.0–3.5  1.0–2.5 cm; petioles ca 1 mm long; reticulations of lower leaf surface very close; lowers and fruits sessile …… W. sessilicarpa Acknowledgements – he curators of the following herbaria are thanked for providing loans of or access to specimens: BNRH, K, LMA, NU. Special thanks to John and Sandie Burrows for bringing this new species to my attention. Sandie Burrows also rendered the line drawing. References Borhidi, A. 1993. Warneckea hedbergorum sp. n. (Memecylaceae) and a short review of the east-African Memecylon s.l. – Opera Bot. 121: 149–151. Fernandes, A. and Fernandes, R. 1955. Melastomataceae africanae novae vel minus cognitae–II. – Bol. Soc. Brot. Sér. 2, 29: 47–64 (19 plates). Fernandes, A. and Fernandes, R. 1960. Melastomataceae africanae novae vel minus cognitae–IV. – Bol. Soc. Brot. Sér. 2, 34: 59–89 (26 plates). Fernandes, A. and Fernandes, R. 1969. Melastomataceae africanae novae vel minus cognitae–V. – Bol. Soc. Brot. Sér. 2, 43: 285–306 (17 plates). Fernandes, R. and Fernandes, A. 1978. Melastomataceae. – In: Launert, E. (ed.), Flora Zambesiaca. Vol. 4. R. Bot. Gard. Kew, pp. 220–276. Fernandes, R. and Fernandes, A. 1980. Melastomataceae. – In: Mendes, E. J. (ed.), Flora de Moçambique, 72. Junta de Investigações Cientíicas do Ultramar, Centro de Botânica, Lisbon. IUCN 2001. he IUCN red list categories and criteria: ver. 3.1. – IUCN Species Survival Commission. Jacques-Félix, H. 1977. La graine et l’embryon chez les Memecylon (Mélastomatacées) africains. – Adansonia Sér. 2, 17: 193–200. Jacques-Félix, H. 1978a. Les subdivisions du genre Memecylon (Melastomataceae) en Afrique. – Adansonia Sér. 2, 17: 415–424. Jacques-Félix, H. 1978b. Les genres de Memecyleae (Melastomataceae) en Afrique, Madagascar et Mascareignes. – Adansonia Sér. 2, 18: 221–235. Jacques-Félix, H. 1983. Mélastomatacées. – In: Satabié, B. and Leroy, J.-F. (eds), Flore du Cameroun. Vol. 24. Délégation Générale à la Recherche Scientiique et Technique, Yaoundé. Jacques-Félix, H. 1985a. Les Memecyleae (Melastomataceae) de Madagascar, part 1. – Bull. Mus. Natl Hist. Nat. B, Adansonia 6: 383–451. Jacques-Félix, H. 1985b. Les Memecyleae (Melastomataceae) de Madagascar, part 2. – Bull. Mus. Natl Hist. Nat. B, Adansonia 7: 3–58. Jacques-Félix, H. et al. 1978. Nervation et types foliares chez les Memecylon (Melast.) africains. – Adansonia Sér. 2, 18: 67–81. Luke, Q. 2005. Annotated checklist of the plants of the Shimba Hills, Kwale District, Kenya. – J. East Afr. Nat. Hist. 94: 5–120. Rao, T. A. and Jacques-Félix, H. 1978. Les types de sclérites foliaires et la classiication des Memecylon africains. – Adansonia Sér. 2, 18: 59–66. Stone, R. D. 2006. Phylogeny of major lineages in Melastomataceae, subfamily Olisbeoideae: utility of nuclear glyceraldehyde 3-phosphate dehydrogenase (GapC ) gene sequences. – Syst. Bot. 31: 107–121. Stone, R. D. and Andreasen, K. 2010. he Afro-Madagascan genus Warneckea (Melastomataceae): molecular systematics and revised infrageneric classiication. – Taxon 59: 83–92. Stone, R. D. and Luke, Q. 2009. Warneckea melindensis (Melastomataceae), a new combination for an east African coastal forest endemic. – J. East Afr. Nat. Hist. 98: 141–145. Timberlake, J. 2009. Coastal dry forests of Cabo Delgado – second reconnaissance trip 22 Nov – 13 Dec 2008. Botanical component progress report no. 2. – R. Bot. Gard. Kew. Timberlake, J. et al. 2011. Coastal dry forests in northern Mozambique. – Plant Ecol. Evol. 144: 126–137. Vollesen, K. 1980. Annotated check-list of the vascular plants of the Selous Game Reserve, Tanzania. – Opera Bot. 59: 1–117. Wickens, G. E. 1975. Melastomataceae. – In: Polhill, R. M. (ed.), Flora of tropical east Africa. Crown Agents for Oversea Governments and Administrations. 117