Unique within the tribe Pavetteae by the pollen with psilate tectum and by the fruit containing 2 stony pyrenes, each with a laterally flattened ovoid seed with irregularly distributed surface ridges formed by elongation of the exotesta cells.

Exallosperma longiflora De Block.

Shrubs, with Terminalia-branching pattern, branching modules consisting of a long-shoot, horizontal in orientation, never bearing inflorescences and relatively smooth, and an inflorescence-bearing short-shoot with short internodes, erect in orientation, densely beset with corky stipular remnants and alternating vegetative and reproductive nodes; vegetative parts pubescent. Leaves grouped terminally on short-shoots, deciduous, petiolate with petioles long, slender and canaliculate above; blades papyraceous; hair tuft domatia present; margins not revolute; bases rounded, subcordate, cordate or unequal, more rarely truncate or obtuse. Stipules keeled, with a dense row of large colleters interspaced with hairs at the base but otherwise glabrous on the inner surface except for the tip, dimorphic: in vegetative nodes consisting of truncate or triangular sheaths forming a cone and topped by needle-like awns, in inflorescence-bearing nodes consisting of ovate sheaths with acute or shortly acuminate tips. Inflorescences seemingly terminal but actually pseudo-axillary on erect short-shoots, pedunculate, pauciflorous, cymose with trichotomous branching; all parts (axes, bracts, bracteoles, pedicels) pubescent; bracts and bracteoles well-developed, linear. Flowers hermaphroditic, pentamerous, shortly pedicellate; all parts (ovary, calyx, corolla) pubescent outside; secondary pollen presentation present. Calyx well-developed; tube short; lobes much longer than tube. Corolla white, turning yellowish with age; tube narrowly cylindrical; lobes contorted to the left in bud and spreading at anthesis. Stamens sessile, inserted in the sinuses of the corolla lobes somewhat below the level of the throat; anthers almost completely included in the corolla tube at anthesis, basimedifixed, with sagittate base and short sterile apical appendix. Disc annular, fleshy, glabrous. Ovary cup-shaped, bilocular; placentation axile, with 3–4 ovules arising on top of a small placenta attached to the base of the septum. Style and stigma only just exserted from the corolla tube at anthesis; stigmatic lobes slender, fused over their entire length except for the very tips, receptive zone on the adaxial surfaces of the free tips and along the lines of fusion of the lobes. Fruits drupaceous, ovoid, pubescent, crowned by the persistent calyx, containing 2 pyrenes; pyrene stony, hemi-ellipsoid with the abaxial side convex and the adaxial side consisting of a flat rim but otherwise open (with the openings of the two pyrenes inside a fruit separated by the membraneous septum), with a short apical longitudinal preformed germination slit on both abaxial and adaxial sides, containing 1 seed; seed laterally flattened, ± bean-shaped; hilum superficial, irregularly ovate, moderate annulus around hilum present; seed surface not smooth but with irregularly distributed ridges formed by the seed-coat; exotesta cells with continuous plate-like thickenings along the outer tangential and upper parts of the radial walls, irregular ridges on seed surface formed by strongly elongated exotesta cells; endotesta consisting of crushed cell layers with many crystals; endosperm entire. Pollen grains 3-zonocolporate, exine psilate, supratectal elements absent.

A monospecific genus, endemic to northern Madagascar, occurring on calcareous soil.

This genus is named for its peculiar seeds.

Differing from Homollea septentrionalis De Block by the size and shape of the leaves of the first order bracts (broadly ovate to orbiculate, 5.5–9.5 × 4.2–9.5 cm vs. broadly ovate to ovate, 0.8–3.5 × 0.5–2.5 cm in H. septentrionalis), the calyx tube and lobes which are glabrous inside (vs. densely sericeous), the lower number of ovules (3–4 vs. 4–6) and the different seeds (2 seeds with irregularly distributed surface ridges, ca. 8 × 5.5 mm vs. 2–6 seeds with smooth surface, ca. 4.5 × 2.5–3 mm).

Figure 2. 

Exallosperma and Helictosperma. A–C Exallosperma longiflora: A flowering branch B inflorescence C infructescence from herbarium specimen Gautier et al. 4257 D Helictosperma poissoniana, flowering branch E, F Helictosperma malacophylla: E inflorescence F detail of inflorescence. Photographs: P. De Block (A, D), S. Dessein (E, F), L. Nusbaumer (B, C, ©: Conservatoire et Jardin botaniques de la Ville de Genève).

MADAGASCAR. Antsiranana Province, Analamerana, bank of Irodo river, close to Irodo camp, 8 Jan. 2002 (fl.), De Block, Rakotonasolo & Randriamboavonjy 1132 (holotype: BR!; isotypes: BR!, G!, K!, MO!, P!, TAN!, UPS!).

Shrub, up to 5 m tall. Young shoots bisulcate, brown, densely covered with erect hairs, rapidly becoming corky with loss of pubescence; older branches brown or greyish-brown, corky and somewhat flaking. Leaves often immature at time of flowering, 7–12 × 5.5–8.5 cm, ovate or elliptic, more rarely broadly elliptic or broadly ovate (but leaves of first order bracts broadly ovate to orbiculate); blades papyraceous, drying brown to dark brown, not discolorous, densely covered with erect hairs on both surfaces; base cordate, rounded, truncate or unequal; apex acuminate, acumen 2–15 mm long; midrib and secondary nerves raised on the lower leaf surface; midrib impressed especially in the basal half on the upper leaf surface; 8–12 secondary nerves on each side of the midrib. Petioles densely covered with erect hairs, 10–25 mm long (but shorter in leaves of first order bracts). Stipules caducous, covered with erect hairs along the base and the keel outside but rapidly becoming corky and losing the pubescence; stipules of vegetative nodes with sheaths 1.5–2.5 mm long and awns 1.5–3 mm long, those of inflorescence-bearing nodes ovate with acute or shortly acuminate tips, 4–8 mm long. Inflorescences consisting of 3–12 flowers, 1–2 × 1–2 cm; anthesis asynchronous within inflorescence; all inflorescence parts (peduncle, axes, pedicels, bracts and bracteoles) densely covered with erect hairs; peduncle 1–3 cm long; first order axes 3–10 mm long; first order bracts with stipular parts triangular and leaves broadly ovate to orbiculate, 5.5–9.5 × 4.2–9.5 cm, with strongly cordate or cordate bases and petioles 3–6(–10) mm long; higher order bracts linear, up to 1.6 cm long; bracteoles opposite on the pedicel just below the ovary, linear, 0.4–1 cm long. Flowers sessile or shortly pedicellate, pedicels 0–2 mm long. Calyx green, densely covered with erect hairs outside; tube ca. 1 mm long, glabrous and without colleters inside; lobes narrowly triangular, 12–16 × 1–1.5 mm (but shorter in young buds), densely covered with appressed hairs at the base and spreading or erect hairs in the upper half inside, bases not overlapping but closely joining, tips acute. Corolla tube 2.7–3.6 cm long, ca. 1.5 mm in diameter at the base, c. 3.5 mm in diameter at the throat, densely covered with erect hairs outside, upper half densely covered with erect hairs inside with pubescence continuing in the throat and on the base of the corolla lobes; lobes elliptic, 9–11 × ca. 3.5 mm, sparsely to moderately covered with erect hairs outside, densely covered with erect hairs at the base inside, margins densely ciliate, tips acute to apiculate. Anthers sessile, inserted in the sinuses of the corolla lobes 2–2.5 mm below the level of the throat, included in the corolla tube except for the very tips, 3–3.5 mm long. Ovary ca. 1.5 mm long, green, densely covered with erect hairs. Style and stigma white, exserted from the corolla tube for 2–5 mm at anthesis, style glabrous or with a few long spreading hairs in the upper half; stigma slender, papillae present on the inner surface of the free tips, longitudinal papillate lines running down for up to 16 mm, but papillae absent just below the tips. Fruits 7–10 × 5–8 mm (persistent calyx not included), moderately to densely covered with erect hairs, drying black and glossy when ripe; seeds ca. 8 × 5.5 × 3 mm, dark brown.

Lowland dry deciduous and semi-deciduous forest on limestone; alt. 0–450 m.

Exallosperma longiflora is only known from the northernmost tip of Madagascar in the Sava and Diana Regions. Fig. 14A.

Flowering: January–February; Fruiting: April.

Exallosperma resembles the Madagascan endemic Homollea by the pedunculate, pauciflorous, pseudo-axillary inflorescences and the pentamerous flowers with relatively long corolla tubes and long, narrow calyx lobes. Exallosperma is characterised by the Terminalia-branching pattern, the large, broadly ovate to orbiculate leaves of the first order bracts, the basally attached placentas from which 3–4 collateral ovules arise, the fruit containing 2 stony pyrenes, each with a laterally flattened ovoid seed with irregularly distributed surface ridges formed by elongation of the exotesta cells and by the pollen with psilate tectum. Exallosperma longiflora may be confused with Homollea septentrionalis, which it resembles by the dense pubescence on vegetative and reproductive organs, the pauciflorous inflorescences, the long flowers with tapering corolla lobes and the long, linear calyx lobes. The two species can be distinguished by the size and shape of the leaves of the first order bracts (broadly ovate to orbiculate, 5.5–9.5 × 4.2–9.5 cm in Exallosperma longiflora vs. broadly ovate to ovate, 0.8–3.5 × 0.5–2.5 cm in H. septentrionalis), the pubescence of the calyx tube and lobes inside (glabrous vs. densely sericeous), the number of ovules (3–4 vs. 4–6) and the different seeds (2 seeds with irregularly distributed surface ridges, ca. 8 × 5.5 mm vs. 2–6 seeds with smooth surface, ca. 4.5 × 2.5–3 mm).

Endangered: EN B1ab(i, ii, iii, iv) + 2ab(i, ii, iii, iv). The extent of occurrence (EOO) of Exallosperma longiflora is estimated to be 1,791 km² and its area of occupancy (AOO) 54 km², which both comply with the criteria for the Endangered category under sub-criteria B1 and B2. The species is known from seven collections, all but two of these collected after the year 2000, reflecting the intensified collection effort in northern Madagascar during the last 20 years. Exallosperma longiflora occurs in four locations, three of which are within protected areas, notably Réserve Spéciale d’Andrafiamena (which includes Analamerana), Loky Manambato (Daraina) and Montagne de Français. The main threat to E. longiflora is decline of its habitat both inside and outside the protected areas as a result of slash-and-burn agriculture, logging for timber and charcoal and burning to favour the growth of young grass for the grazing of cattle. Furthermore, traditional mining for gold is a serious threath in the area (Rakotondravony 2009; Nusbaumer et al. 2010). Based on the above information, the species is listed as Endangered.

MADAGASCAR. Antsiranana Province: Montagne des Français, plateau supérieur de l’Anosiarivo, 28 Jan 1966 (fl.), Capuron 24425-SF (BR, P, TEF); Massif de l’Ankitakona, 25 Apr 1966 (fr.), Capuron 24663-SF (BR, P, TEF); Analamerana, bank of Irodo river, close to Irodo camp, 6 Jan 2002 (fl.), De Block, Rakotonasolo & Randriamboavonjy 1080 (BR, MO, P, TAN, UPS); Sava, sous-préfecture de Vohemar, commune rurale de Daraina, Daraina, forêt d’Ambilondomba, W of Ambilondomba, 300 m S du point côté 341, 150 m, 8 Mar 2003 (fr.), Gautier, Wohlhauser & Nusbaumer 4257 (BR, G, K); Sava, sous-préfecture de Vohemar, commune rurale de Daraina, Daraina, forêt de Solaniampilana-Maroadabo, à 700 m du point côté 608, au 85°, 437 m, 2 Feb 2006 (fl.), Nusbaumer & Ranirison 1992 (BR, G); Sava, sous-préfecture de Vohemar, commune rurale de Daraina, Daraina, forêt de Solaniampilana-Maroadabo, à 750 m du point côté 608, au 205°, 328 m, 4 Feb 2006 (fl.), Nusbaumer & Ranirison 2151 (G).

Differing from Exallosperma by the shorter calyx lobes (3–9 mm vs. 12–16 mm long), the shorter corolla tubes (0.7–1.4 cm vs. 2.7–3.6 cm long), the completely exserted anthers at anthesis (vs. included in the corolla tube except for the tips), the pollen with microreticulate to perforate tectum (vs. psilate tectum), the fruits containing a single stony pyrene that opens into four valves, and the single seed that is rolled-in on itself like a giant pill-millipede (vs. fruits containing 2 hemi-ovoid pyrenes not opening into 4 valves, each with 1 laterally flattened, bean-shaped seed).

Helictosperma malacophylla (Drake) De Block

Shrubs or small trees, with Terminalia-branching pattern, branching modules consisting of a long-shoot, horizontal in orientation, never bearing inflorescences and relatively smooth, and an inflorescence-bearing short-shoot with short internodes, erect in orientation, densely beset with corky stipular remnants and alternating vegetative and reproductive nodes; vegetative parts glabrous or pubescent. Leaves grouped terminally on short-shoots, deciduous, petiolate with petioles long, slender and canaliculate above; blades papyraceous; domatia present; margins not revolute; bases rounded, subcordate, cordate or unequal, more rarely truncate or obtuse. Stipules keeled, with a dense row of large colleters interspaced with hairs at the base but otherwise glabrous on the inner surface, dimorphic: in vegetative nodes consisting of truncate or triangular sheaths forming a cone and topped by needle-like awns, in inflorescence-bearing nodes consisting of ovate sheaths with acute or shortly acuminate tips. Inflorescences seemingly terminal but actually pseudo-axillary on erect short-shoots, pedunculate, pauci- or multiflorous, cymose with trichotomous branching; all parts (axes, bracts, bracteoles, pedicels) glabrous or pubescent; bracts and bracteoles well-developed, linear. Flowers hermaphroditic, pentamerous, pedicellate; all parts (ovary, calyx, corolla) glabrous or pubescent outside; secondary pollen presentation present. Calyx well-developed; tube short; lobes much longer than tube. Corolla white, turning yellowish with age; tube narrowly cylindrical; lobes contorted to the left in bud and spreading at anthesis, oblong, with blunt and emarginate tips. Stamens inserted in the sinuses of the corolla lobes at the level of the throat; filaments short; anthers completely exserted from the corolla tube at anthesis, basifixed, with sagittate base and short sterile apical appendix. Disc annular, fleshy, glabrous. Ovary cup-shaped, bilocular; placentation axile, with 3 ovules arising on top of a small placenta attached to the lower half of the septum. Style and stigma exserted from the corolla tube at anthesis; stigmatic lobes fused over their entire length except for the very tips, receptive zone on the adaxial surfaces of the free tips and along the lines of fusion of the lobes. Fruits drupaceous, spherical, pubescent or glabrous, crowned by the persistent calyx, containing 1 pyrene; pyrene crustaceous, spherical, formed by the outer convex parts of the two locules (the septum remaining membraneous and pushed to the side by the developing seed), opening along 4 preformed longitudinal germination slits of which 2 run down the margins of the locules and 2 are perpendicular to those, containing 1 seed; seed spherical, rolled-in on itself in the shape of a giant pill-millipede; hilum ovate, profound, moderate annulus around hilum present; exotesta cells with continuous plate-like thickenings along the outer tangential and upper parts of the radial walls, annulus formed by strongly elongated exotesta cells; endotesta consisting of crushed cell layers with many crystals; endosperm entire. Pollen grains 3-zonocolporate, exine microreticulate to perforate, supratectal elements absent.

A genus with 2 species, endemic to western and northern Madagascar, occurring on calcareous soil.

The genus is named for the shape of the seeds, which are rolled-in on themselves in the shape of giant pill-millipedes.

Shrub 2–6 m tall, more rarely tree up to 12 m tall with trunk up to 6 m tall and dbh up to 10 cm; young shoots quadrangular, often bisulcate, brown, densely covered with erect to spreading hairs; older branches brown, pale brown, greyish or fawnish, glabrous, often flaking. Leaves often immature at time of flowering, 6–15 × 4–8.5 cm, ovate, more rarely broadly ovate, elliptic or obovate; blades papyraceous, drying brown to dark brown, more rarely greenish-brown above, brown and often somewhat paler below, densely covered with erect hairs on the lower surface, moderately to densely covered with erect or spreading hairs on the upper surface, pubescence denser on the midrib and secondary nerves on both surfaces; base rounded, subcordate, cordate or unequal, more rarely truncate or obtuse; apex acuminate, acumen 3–18 mm long; hair tuft domatia present; midrib and secondary nerves raised on the lower leaf surface; midrib impressed especially in the basal half on the upper leaf surface; 10–14 secondary nerves on each side of the midrib. Petioles densely covered with erect hairs, 14–45 mm long. Stipules caducous; densely covered with erect hairs outside but rapidly becoming corky and losing the pubescence; stipules of vegetative nodes with sheaths 3–5 mm long and awns 3–6 mm long, those of inflorescence-bearing nodes ovate with acute to shortly acuminate tips, 4–6 mm long. Inflorescences consisting of 25–90 flowers, 2.5–8 × 2–7 cm; peduncle, inflorescence axes and pedicels densely covered with erect hairs; peduncle 1–5.5 cm long; first order axes up to 2(–4) cm long; first order bracts with stipular parts narrowly triangular and leaves long-petiolate and identical in shape and size to the vegetative leaves or somewhat smaller; second order bracts of the central axis often similar to the first order bracts but leaves considerably smaller and narrower with acute to attenuate base, 1–6.5 × 0.3–3.2 cm; second order bracts of lateral axes reduced or absent; higher order bracts and bracteoles linear, moderately to densely covered with erect hairs on both surfaces, no colleters present inside; bracts up to 1.2 cm long; bracteoles subopposite on the pedicel, 0.2–0.4 cm long; first order branching often shifted above the first order bracts (up to 1 cm higher); bracts sometimes adnate to axis for up to 5 mm. Flowers pedicellate, pedicels 1–5 mm long. Calyx green, moderately to densely covered with erect hairs outside; tube ca. 0.5 mm long, with a sparse ring of appressed hairs at the base but without colleters inside; lobes erect in young bud, but rapidly becoming reflexed, oblong, 3–5 × 1–1.5 mm, the upper half sparsely covered with erect hairs inside, bases not overlapping but closely joining, tips obtuse. Corolla tube 7–8 mm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the throat, moderately to densely covered with erect hairs outside, the upper 2/3 moderately to densely covered with erect hairs inside; lobes 4–5 × 3–3.5 mm, glabrous on both surfaces, margins ciliate. Anthers 3.5–5 mm long; filaments 1–1.5 mm long. Ovary 1–1.25 mm long, green, densely covered with erect hairs. Style and stigma white, exserted from the corolla tube for 7–10 mm at anthesis; style densely covered with spreading, upwardly directed hairs in the upper half; stigma with upper 4–5 mm fusiform, longitudinal papillate lines running down for a further 3–4 mm. Fruits 4–6 mm in diameter (persistent calyx not included), moderately to densely covered with erect hairs, drying brown and glossy when ripe; seeds 3–5 mm in diameter, dark brown.

Lowland dry deciduous and semi-deciduous forest on calcareous soil, usually on sand; alt. 30–800 m.

Helictosperma malacophylla is known from the Boeny, Betsiboka and Sofia Regions (Mahajanga Province), from the Ihorombe (Fianarantsoa Province) and from the Atsimo-Andrefana and Menabe Regions (Toliara Province). Fig. 14B.

Flowering: November–February(–April); Fruiting: (November–)January–May.

Ampale (dialect Masikoro; coll. ignot. 21707-SF); nofotrakoho (coll. ignot. 19382-SF); talinala (dialect Masikoro; coll. ignot. 21708-SF); voloiravy (Randriamiera 8770-RN); zamanimbato (Rakotovao 3898-RN).

Construction wood for houses and cattle enclosures (coll. ignot. 19146-SF, 19382-SF, 21707-SF, 21708-SF); fire wood (coll. ignot. 21707-SF, 21708-SF).

Helictosperma resembles Exallosperma by the Terminalia-branching pattern, the pedunculate, pseudo-axillary inflorescences and the basally attached placentas from which three collateral ovules arise. The genera differ by pollen (tectum microreticulate to perforate in Helictosperma vs. psilate in Exallosperma) and fruit/seed characters (fruit with two pyrenes, each with a single laterally flattened seed with irregularly distributed surface ridges vs. fruit with single pyrene falling apart into four valves and containing a single seed that is rolled-in on itself). Helictosperma malacophylla resembles E. longiflora by the general hairiness of the whole plant but differs from it by the larger number of flowers per inflorescence (25–90 in H. malacophylla vs. 3–12 in E. longiflora), the longer pedicels (1–5 mm vs. 0–2 mm long), the shorter bracteoles (2–4 mm vs. 4–10 mm long) and the shorter corolla tubes (7–8 mm long vs. 26–37 mm long) and calyx lobes (3–5 mm long vs. 12–16 mm long).

Near Threathened: NT. The extent of occurrence (EOO) of Helictosperma malacophylla is estimated to be 273.476 km², which falls outside any threat category, but its area of occupancy (AOO) is 261 km², which complies with the Endangered category under the sub-criterion B2. The species occurs in ten locations and is known from more than fifty collections, twelve of which were collected recently (after 1989). The distribution of these recent collections coincides with the distribution of the older specimens (from 1892 till 1975), indicating that the species remains present throughout its original distribution area. Only few specimens were collected from protected areas, notably Ankarafantsika National Park, Tsingy de Namoroka Strict Nature Reserve and Kirindy Mitea National Park. Despite its large extent of occurrence, Helictosperma malacophylla is threathened locally by reduction of its habitat through slash-and-burn agriculture, logging for timber and charcoal and burning to improve grazing. Based on the above observations, the species is assessed as Near Threathened.

MADAGASCAR. Mahajanga Province: 2 km N of Tsarahasina, 30 m, 10 May 2006 (fr.), Andriamahay & Rakotoarisoa 1359 (K); canton Bemanevika, Analafaly forest, 6 km E of Marotaolana, 384 m, 10 May 2005 (fr.), Birkinshaw, Andrianjafy & Raha-Jean 1525 (BR, MO, P, TAN); Réserve Naturelle VII, Ankarafantsika, 120–150 m, s.dat. (fl.), coll. ignot. 30-SF (P); vallée de Marivoraona, village le plus proche Ambodifiakarana, canton Betsandraka, district Tsaratanana, bord E du sentier d’Ambatobe à Ambodifiakarana, 30 Nov 1958 (fr.), coll. ignot. 19146-SF (P, TEF); forêt d’Anatialabe, village le plus proche Kamakama, canton Ankirihitra, district Ambatoboeni, 30 Nov 1958 (fr.), coll. ignot. 19382-SF (P, TEF); Soalala district, Réserve Naturelle Intégrale de Namoroka (Réserve Naturelle 8), c. 38.5 km S of Soalala, 120 m, 2 Feb 2000 (fr.), Davis, Rakotonasolo & Wilkin 2520 (BR, K, TAN); forêt de Marohogo, 22 m, 13 Feb 1999 (fr.), De Block & Rakotonasolo 799 (BR, C, G, K, MO, P, TAN, WAG); Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 24 Apr 1952 (fl.), Rakotovao 3898-RN (P, TAN); Réserve Naturelle VIII, Tsingy de Namoroka, Andranomavo, district Soalala, 29 Dec 1952 (fl.), Rakotovao 4918-RN (P, TAN); Ambatofolaka, Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 28 Mar 1954 (fr.), Rakotovao 6154-RN (BR, P, TEF); Ambatofolaka, Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 26 Jan 1954 (fr.), Rakotovao 6239-RN (BR, P, TEF); canton Andranomavo, district Soalala, 25 Feb 1957 (fr.), Randriamiera 8770-RN (BR, P, TEF); canton Andranomavo, district Soalala, 10 Nov 1958 (fl.), Randriamiera 9724-RN (BR, P, TEF); Fianarantsoa Province: de Ihosy 47–49 km ad SE per viam ad Ivohibe in nemorosis parvis residuis juxta pascua ignita, 650–700 m, 5 Nov 1967 (fl.), Bernardi 11197 (G, K, P); bassin de la Menarahaka, près du carrefour des routes d’Ihosy à Ivohibe et Iakora, 650 m, 10 Feb 1963 (fr.), Capuron 22618-SF (P, TEF); haut bassin de la Menarahaka, E d’Ihosy, 5 Nov 1967 (fr.), Capuron 27850-SF (BR, P, TEF); vallée de la Menarahaka, E d’Ihosy, 19 Dec 1968 (fl.), Capuron 28479-SF (P, TEF); 10 km NE d’Ihosy entre Ihosy et Ambararata, 22 Feb 1970 (fr.), Capuron 29068-SF (BR, P, TEF); road Antananarivo-Ihosy, a few km before reaching Ihosy, 4 Jan 1999 (fl.), De Block & Rakotonasolo 534 (BR, K, MO, TAN); haute vallée de la Menarahaka, E d’Ihosy, 700–800 m, 28 Jan–10 Apr 1955 (fr.), Humbert 29886 (BR, P); Toliara Province: Sakaraha, commune Mahaboboka, Marotsiraka, forêt d’Analaraty, 469 m, 24 Mar 2013 (fr.), Andriamihajarivo, Miandry & Rakotoarivony 1879 (BR, MO, P, TAN); c. 10 km N of Befandriana-Sud, 150 m, 28 Nov 1962 (fl.), Appert 108 (MO, Z); Morombe district, Tanandava-Tatalavalo, 70 m, 10 Mar 1963 (fr.), Appert 114 (MO, Z); Fotivolo, Ankazobe, Feb 1963 (fr.), Bosser 17287 (BR, P, TAN); environs de Berenty, 18 Feb 1970 (fr.), Bosser 19934 (BR, P); Betsipotika, E de Morondava, 18 Jan 1962 (fl.), Capuron 20872-SF (BR, P, TEF); N de Dabara, Mahabo, 1 Apr 1970 (fr.), Capuron 29141-SF (BR, P, TEF); forêt de Mavozobe, village le plus proche Mavozobe, canton Befandriana-Sud, sous-préfecture Morombe, 22 Feb 1964 (fr.), coll. ignot. 21707-SF (TEF); forêt de Mavozobe, village le plus proche Mavozobe, canton Befandriana-Sud, sous-préfecture Morombe, 22 Feb 1964 (fr.), coll. ignot. 21708-SF (P); Kirindi forest, N part - Conoco, 7–16 m, 19 Jan 2007 (fl.), De Block, Rakotonasolo, Groeninckx & Dessein 2194 (BR, G, MO, P, TAN); Morondava, 1892 (fl.), Grevé s.n. (K); Morondava, close to site of baobabs amoureux, 27 m, 22 Jan 2007 (fl.), Groeninckx, Rakotonasolo, De Block & Dessein 132 (BR, G, MO, P, TAN, WAG); bassin de la Malio, affluent de Mangoky, près d’Ambalabe, 400–450 m, Nov 1946 (fl.), Humbert 19447 (BR, P); bassin moyen du Fiherenana entre Lambomakandro et Sakaraha, 400 m, 10 Dec 1946 (fl.), Humbert 19681 (BR, P); N of Tulear, near Mangoky river, 50 m, 1 Jan 1989 (fl.), Phillipson 3068 (BR, K, MO, P, TAN, WAG); Horombe, Beroroha, Tsivoko, forêt humide de Makay dans la zone de Menapanda, 495 m, 9 Dec 2010 (fr.), Rakotovao & Andriantiana 5558 (BR, MO, P, TAN); district Ankazoabo, commune Ankazoabo, canton Morafeno, village le plus proche Ampanihimahasoa, route Sakaraha-Ankazoabo, 12 km SE d’Ankazoabo, 599 m, 11 Mar 2004 (fr.), Randrianaivo, Ratodimanana, Razafindraibe, Randrianarisoa, Edodoky & Tsimanoa 1058 (BR, G); Sakaraha, Mahaboboka, canton Marotsiraka Betsileo, S of Ambinanintelo village and S of the intersection of the two rivers Bevoalavo and Andranoheza, 417 m, 21 Feb 2011 (fr.), Randrianasolo, Andriamihajarivo, Razanatsima, Rakotoarivony, Randrianarivony, Fagnarena, Bruno & Redilike 1417 (BR, MO, P, TAN); forêt d’Anosilamy, canton Beronono, commune Beronono, 448 m, 13 Jan 2010 (fr.), Razakamalala, Rakotovao & Andriantiana 5161 (BR, MO, P, TAN); Unplaced localities: forêt de Moailake, Feb 1892 (fl.), Douilot s.n. (P); Nandrosia, May 1897 (fr.), Perrier de la Bâthie 234 (P); Boiny, not readable further, Jan 1902 (fl.), Perrier de la Bâthie 1011 (BR, P); Without locality: s.dat. (fr.), Baron 4612 (K, P); Central Madagascar, s.dat. (fr.), Baron 4673 (K); s.dat. (fr.), Baron 4679 (P); s.dat. (st.), Douilot s.n. (P); s.dat. (fr.), Homolle 1427 (P); s.dat. (fr.), Homolle 1473 (P); s.dat. (fr.), Homolle 1495 (P).

Differing from Helictosperma malacophylla by the pauciflorous inflorescences [(1–)5–15(–20) vs. 25–90 flowers], the larger calyx lobes (7–9 × 1.5–2.5 mm vs. 3–5 × 1–1.5 mm), the glabrous corolla tube, the corolla lobes without ciliate margins and the usually glabrous vegetative and reproductive parts, but, if pubescent, then hairs appressed (vs. erect or spreading in H. malacophylla).

MADAGASCAR. Antsiranana Province, Analamerana, along Ambatabe river, 41 m, 7 Jan 2002 (fl.), De Block, Rakotonasolo & Randriamboavonjy 1095 (holotype: BR!; isotypes: BR!, K!, MO!, P!, TAN!, UPS!).

Shrub 1.5–4 m tall, more rarely small tree to 4 m tall, dbh to 7 cm; young shoots somewhat quadrangular and bisulcate, dark brown, glabrous or sparsely to densely covered with appressed hairs; older branches brown, pale or greyish-brown. Leaves often immature at time of flowering, 3–10 × 2–6 cm, ovate, rarely elliptic or obovate; blades papyraceous, drying brown to dark brown, more rarely greenish, hardly discolorous, glabrous or with midrib and secondary nerves sparsely to densely covered with appressed hairs, more rarely also higher order nerves pubescent on the lower surface, glabrous or sparsely to moderately covered with appressed hairs on the upper surface; base rounded, subcordate, cordate or unequal, more rarely truncate or obtuse; apex acuminate, acumen 3–10(–15) mm long; hair tuft or ciliate pit domatia present, sometimes also in the axils of secondary nerves; midrib and secondary nerves raised on the lower leaf surface; midrib impressed in the basal half on the upper leaf surface; 5–8 secondary nerves on each side of the midrib. Petioles glabrous to densely covered with short appressed hairs, 5–35 mm long. Stipules caducous, glabrous or sparsely to densely covered with appressed hairs outside, but rapidly becoming corky and losing the pubescence; stipules of vegetative nodes with sheaths 1.5–2.5 mm long and awns 2–5 mm long, those of inflorescence-bearing nodes ovate with acute to shortly acuminate tips, 4–7 mm long. Inflorescences consisting of (1–)5–15(–20) flowers, up to 3 × 2 cm; peduncle, inflorescence axes and pedicels glabrous or moderately to densely covered with appressed hairs; peduncle 0.5–3.5 cm long; first order axes up to 1.2 cm long; first order bracts with stipular parts narrowly triangular and leaves long-petiolate and identical in shape and size to the vegetative leaves or somewhat smaller; second order bracts of the central axis often similar to the first order bracts but leaves considerably smaller and narrower with acute to attenuate base, more rarely identical in shape to vegetative leaves with cordate or rounded base, up to 3.5 × 2.5 cm; second order bracts of lateral axes, higher order bracts and bracteoles linear, glabrous, ciliate or sparsely to moderately covered with appressed or spreading hairs on both surfaces, no colleters present inside; bracts up to 2.2 cm long; bracteoles subopposite on the pedicel, 0.2–1.2 cm long; first order branching often shifted above the first order bracts (up to 1 cm higher); bracts often adnate to axis for up to 5 mm. Flowers pedicellate, pedicels 1–6 mm long. Calyx green; tube 0.75–1 mm long, glabrous or more rarely moderately to densely covered with appressed hairs outside, glabrous and without colleters inside; lobes erect, leaf-like, 7–9 × 1.5–2.5 mm, glabrous inside and outside but with margins ciliate or more rarely sparsely covered with appressed hairs outside (mostly in basal half or along veins), bases not overlapping but closely joining, tips acute to obtuse. Corolla tube 5–14 mm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the throat, glabrous outside, densely covered with erect hairs except at the base and at the throat inside; lobes 4–5 × 3–3.5 mm, glabrous on both surfaces, margins not ciliate. Anthers 3–4 mm long; filaments 1–1.5 mm long. Ovary 1–1.5 mm long, faintly ribbed longitudinally when dry, green, glabrous or more rarely moderately to densely covered with appressed hairs. Style and stigma white, exserted from the corolla tube for 4–7 mm at anthesis; style densely covered with spreading, upwardly directed hairs over the whole length except for a further 2–3 mm. Fruits 5–7 mm in diameter (persistent calyx not included), with faint longitudinal ribs, glabrous or more rarely moderately to densely covered with appressed hairs, drying blackish and glossy when ripe; seeds ca. 5 mm in diameter, dark brown.

Lowland dry deciduous and semi-deciduous forest on limestone; alt. 0–450 m.

Helictosperma poissoniana is known from the Diana Region (Antsiranana Province) and from the Boeny and Melaky Regions (Mahajanga Province). Fig. 14C.

Flowering: October–January, May; Fruiting: January–December.

Hazontaka (Rakotovao 4081-RN); maroampotatra (Rakotovao 3914-RN); pitsopitsoka (Randriamiera 6722-RN); refeko (Leandri 573); tsarepepana (dialect Antakarana; Humbert 19013); voanievitra (Rakotovao 6240-RN).

The three flowering specimens from the Tsingy de Bemaraha (Leandri 573 & 578; Jongkind 3415) have longer flowers (corolla tube 13–14 mm long) than all other specimens of this species (corolla tube 5–9 mm long). – Some specimens in the P herbarium were annotated as Tarenna poissoniana Homolle (e.g. Poisson 21). Capuron (1973) discussed this species in his unpublished treatment of the Madagascan Rubiaceae under the same name.

Near Threathened: NT. The extent of occurrence (EOO) of Helictosperma poissoniana is estimated to be 70,048 km², which exceeds the upper limits for any threat category but its area of occupancy (AOO) is 198 km², which falls within the limits for the Endangered category under the sub-criterion B2. The species occurs in seven locations and in three protected areas: Namoroka Strict Nature Reserve, Bemaraha National Park and Ankarana Special Reserve. Helictosperma poissoniana is widespread but threathened locally as a result of the reduction of its habitat through slash-and-burn agriculture, illegal logging and fires to improve grazing. Furthermore, artisanal sapphire mining in Ankarana Special Reserve is a serious problem. Based on the above observations, the species is assessed as Near Threathened.

MADAGASCAR. Antsiranana Province: Massif de l’Ankarana, 5 Nov 1990 (fl.), Bardot-Vaucoulon 238 (P); Massif de l’Ankarana, 17 Nov 1990 (fl., fr.), Bardot-Vaucoulon 303 (K, P); plateau de l’Ankarana, W de Mahamasina (Antanatsimanaja), 23 Apr 1963 (fr.), Capuron 22670-SF (BR, P, TEF); près de Marotaolana, Anivorano Nord, 4 Nov 1966 (fr.), Capuron 24543-SF (BR, P, TEF); district Ambilobe, village Ambilomagodro, km 114, montagne d’Ambohibe, grès de l’Isalo, 300 m, 8 Feb 1960 (fr.), Cours & Humbert 5705 (P); Ankarana, close to Apondrabe river, 82 m, 26 May 1999 (fr.), De Block, Rapanarivo & Randriamboavonjy 1042 (BR, G, K, MO, P, TAN, WAG); Ankarana, following the dry river Apondrabe, close to Mahamasina, 82 m, 27 May 1999 (fr.), De Block, Rapanarivo & Randriamboavonjy 1057 (BR, K, MO, P, TAN); Analamerana, along Ambatabe River, 41 m, 7 Jan 2002 (fl.), De Block, Rakotonasolo & Randriamboavonjy 1092 (BR, MO, TAN, UPS); Ankarana, near Mahamasina, perte d’eau, 82 m, 15 Jan 2002 (fr.), De Block, Rakotonasolo & Randriamboavonjy 1242 (BR, G, K, MO, TAN, WAG); Ankarana Special Reserve, c. 5 km NW of park village near Besaboba river, 90 m, 25 Apr 1993 (fr.), Harder, Merello, Razafimandimbison & Razafindrabaeza 1704 (MO, P, TAN); Diego-Suarez, Jan 1945 (fl.), Homolle 305 (P); Ambodimagodro, plateau de l’Ankarana, Dec 1938–Jan 1939, 250 m (fl.), Humbert 19013 (P); plateau de l’Analamera, 50–400 m, Jan 1938 (fl.), Humbert 19184 (P); collines et plateaux calcaires de l’Ankarana du Nord, 30–350 m, 24 Jan–29 Feb 1960 (fr.), Humbert 32468 (BR, P); collines et plateaux calcaires de l’Ankarana du Nord, colline S du jardin botanique 8, 30–350 m, 24 Jan–29 Feb 1960 (fr.), Humbert 32626 (BR, P); collines et plateaux calcaires de l’Ankarana du Nord, 30–350 m, 24 Jan–29 Feb 1960 (fr.), Humbert 32832 (P); Ankarana du Nord, Mar 1962 (fr.), Keraudren 1687 (P); Ankarana Réserve Spéciale, close to camp des Anglais, 180 m, 18 Feb 1994 (fr.), Lewis, McDonagh, Andrianarisata, Randriamabolona, Andiratsiferama & Bled 1125 (BR, K, MO, P, WAG); Réserve Spéciale d’Ankarana, Ambondromifehy, 11 Jan 2008 (fr.), Rakotonasolo 1164 (K); Mahajanga Province: Beanka, partie sud, Sarodrano, relevé linéaire B30, 429 m, 5 Mar 2012 (fr.), Bolliger, Hanitrarivo & Rakotozafy 278 (BR, G); forêt de Marohogo, près du village de Marohogo, 7 Apr 1965 (fr.), Capuron 24091-SF (BR, P, TEF); Soalala District, Réserve Naturelle Intégrale VIII, Tsingy de Namoroka, c. 40 km S of Soalala, 130 m, 3 Feb 2000 (fr.), Davis, Rakotonasolo & Wilkin 2533 (BR, K, TAN); district Antsalova, Tsingy de Bemaraha, Réserve Naturelle IX, near Ambodiria, 150 m, 17 Mar 2004 (fr.), Davis & Rakotonasolo 3122 (BR, K); forêt de Marohogo, 22 m, 13 Feb 1999 (fr.), De Block & Rakotonasolo 797 (BR, C, G, K, MO, P, TAN, TEF, WAG); forêt de Marohogo, 22 m, 13 Feb 1999 (fr.), De Block & Rakotonasolo 798 (BR, C, G, K, MO, P, TAN, WAG); environs de Majunga, 2–15 m, 28–30 Dec 1924 (fl.), Humbert 4046 (BR, P); Tsingy de Bemaraha, N of Manambolo river, 50 m, 28 Nov 1996 (fl.), Jongkind, Andriantiana & Razanatsoa 3258 (BR, K, WAG); Tsingy de Bemaraha, N of Manambolo river, 50 m, 6 Dec 1996 (fl.), Jongkind, Andriantiana & Razanatsoa 3415 (BR, K, WAG); Réserve Naturelle IX, Bemaraha, Antsingy Nord, 22 Nov 1932 (fl.), Leandri 573 (P); calcaires de l’Antsingy, vers Ambodiriana, E d’Antsalova, 100–150 m, 9 Feb 1960 (fr.), Leandri & Saboureau 3072 (BR, P); Antsingy d’Antsalova, Tsingy de Bemaraha, Réserve Naturelle IX, Jan 1975 (fr.), Morat 4837 (P, TAN); Beanka, partie nord, bord de la rivière Bokarano, 187 m, 18 Dec 2011 (fr.), Nusbaumer, Bolliger, Hanitrarivo & Rakotozafy 3202 (BR, G); environs de Majunga, May 1908 (fl.), Perrier de la Bâthie 3266 (P); Namoroka, Andranomavo, Ambongo, Oct 1905 (fl.), Perrier de la Bâthie 3634 (BR, P); environs de Majunga, May 1908 (fr.), Perrier de la Bâthie 3766 (P); Kamakama, sur le plateau de l’Ankarana, Oct 1901 (fl.), Perrier de la Bâthie 3777 (P); Majunga, 22 Dec 1904 (fr.), Poisson 21 (P); Antsalova, Réserve Naturelle Intégrale IX, Tsingy de Bemaraha, Ambodiriana, 14 Mar 2004 (fr.), Rakotonasolo, Davis & Maurin 767 (BR, K, TAN); Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 30 Apr 1952 (fr.), Rakotovao 3914-RN (P); Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 10 Jun 1952 (fr.), Rakotovao 4081-RN (P); Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 20 Nov 1953 (fr.), Rakotovao 5672-RN (BR, P); Ambatafolaka, Réserve Naturelle VIII, Tsingy de Namoroka, canton Andranomavo, district Soalala, 4 Feb 1954 (fr.), Rakotovao 6240-RN (BR, P, TEF); Beanka, partie centrale, Andoloposa, 358 m, 26 Mar 2012 (fr.), Rakotozafy, Bolliger & Hanitrarivo 97 (BR, G); Boeny, canton Andranomavo, district Soalala, 13 Oct 1954 (fl.), Randriamiera 6722-RN (P, TEF); Boeny, canton Andranomavo, district Soalala, 18 Jan 1955 (fr.), Randriamiera 7070-RN (BR, P, TEF); Boeny, canton Andranomavo, district Soalala, 25 Feb 1957 (fr.), Randriamiera 8771-RN (BR, P, TEF); Boeny, canton Andranomavo, district Soalala, 15 Apr 1957 (fr.), Randriamiera 8795-RN (BR, P, TEF).

Differing from species within the Coptosperma assemblage by the combination of the following characters: 3 ovules pendulous from a small placenta attached to the upper half of the septum and keeled triangular stipules with well-developed awn (vs. stipules not keeled and without awn, of the “bec du canard” type).

Pseudocoptosperma menabense Capuron ex De Block

Shrubs; vegetative parts except for young shoots glabrous. Leaves persistent, petiolate with petioles short and canaliculate above; blades coriaceous; domatia absent; margins revolute. Stipules triangular with well-developed awns, keeled, with 2 or 3 rows of colleters at the base but otherwise glabrous on the inner surface. Inflorescences terminal, sessile, multiflorous, cymose with trichotomous branching; partial inflorescences compact; all parts (axes, bracts, bracteoles, pedicels) densely pubescent; bracts and bracteoles small, triangular. Flowers hermaphroditic, pentamerous, sessile to shortly pedicellate; all parts (ovary, calyx, corolla) glabrous outside; secondary pollen presentation present. Calyx with short tube and small lobes. Corolla white, turning yellowish with age; tube narrowly cylindrical, short; lobes contorted to the left in bud and spreading at anthesis. Stamens inserted in the sinuses of the corolla lobes at the level of the throat; filaments short; anthers completely exserted from corolla tube at anthesis, basifixed, with sagittate base and short sterile apical appendix. Disc annular, fleshy, glabrous. Ovary cup-shaped, bilocular; placentation axile, with 3 ovules pendulous from the base and the lateral sides of a small placenta attached to the upper half of the septum. Style and stigma exserted from the corolla tube at anthesis; stigmatic lobes fused over their entire length, receptive zone along the lines of fusion of the lobes. Fruits drupaceous, spherical, glabrous, crowned by the persistent calyx, containing 1 pyrene; pyrene crustaceous, spherical, formed by the outer convex parts of one developed and one aborted locule (the septum remaining membraneous and pushed to the side by the developing seed), with a small central apical protuberance on the adaxial side, opening along the line of fusion of the locules, containing 1 seed; seed subspherical; hilum superficial, ovate, annulus around hilum absent; exotesta cells parenchymatic and filled with tannins; endotesta consisting of crushed cell layers without crystals; endosperm ruminate. Pollen grains 3-zonocolporate, exine microreticulate to perforate, supratectal elements absent.

A genus with a single species, endemic to western Madagascar.

The genus is named for its resemblance to Coptosperma.

Differing from Coptosperma mitochondrioides Mouly & De Block by the triangular, keeled stipules with a robust awn (vs. stipules of the “bec du canard” type with rounded tip) and the smooth fruits (vs. fruits with ca. 10 longitudinal ribs).

MADAGASCAR. Mahajanga Province, forêt Tsimembo, dans la concession Barthe, 19 Dec 1953 (fl.), Martin 8252-SF (holotype: P!; isotypes: BR!, TEF!).

Shrub or small tree to 8 m tall, dbh to 10 cm; young shoots bisulcate, dark brown, densely covered with short erect hairs; older branches pale brown or fawn, glabrescent, in dried condition strongly contrasting with the blackish-brown stipules and dark brown petioles. Leaves 5–12 × 1–2.5 cm, narrowly elliptic or narrowly obovate; blades coriaceous, drying glossy and brown or more rarely greenish above, somewhat paler and dull below, glabrous on both surfaces; base cuneate to attenuate; apex acuminate, acumen 5–12 mm long; midrib raised and secondary and tertiary nerves somewhat raised on the lower leaf surface; midrib impressed on the upper leaf surface; 10–16 secondary nerves on each side of the midrib. Petioles 2–6 mm long, glabrous. Stipules drying blackish-brown, rapidly becoming corky, caducous, triangular with the robust awn as long as or longer than the basal sheath, glabrous outside, glabrous but with 2–3 basal rows of colleters inside; sheaths 1–2.5 mm long; awns 2–4 mm long. Inflorescences consisting of numerous flowers, 1–3.5 × 2–7 cm, sessile; inflorescence axes, pedicels, bracts and bracteoles densely covered with short erect hairs, green but drying dark brown; bracts with stipular parts reduced and foliar parts triangular and vaulted, 1–2 mm long, densely covered with appressed hairs and with a basal row of colleters inside, margins ciliate; central first order bracts often with stipular parts reduced and foliar parts leaf-like, 0.5–4 × (0.2–)0.4–0.9 cm, elliptic or narrowly elliptic, base attenuate or cuneate, petiole 1–2 mm long; bracteoles at the base of the ovary, broadly triangular, 0.4–0.7 mm long, tips rounded to obtuse, with appressed hairs mostly in the upper half and a single colleter at each side of the base inside; first order axes 0.5–2.5 cm long. Flowers sessile or shortly pedicellate, pedicels 0–1 mm long with central flowers mostly sessile. Calyx green, glabrous outside; tube ca. 0.25 mm long, glabrous and without colleters inside; lobes ovate, 0.2–0.3 mm long, bases not overlapping but closely joining, tips rounded to obtuse, rarely acute. Corolla tube 1.5–2.5 mm long, ca. 0.4 mm in diameter at the base, ca. 1 mm in diameter at the throat, glabrous outside, throat and upper third to half moderately to densely covered with erect hairs inside; lobes oblong, 2–2.5 × 0.75–1 mm, glabrous on both surfaces, tip blunt and emarginate. Stamens completely exserted at anthesis; filaments < 0.5 mm long; anthers 1.3–1.5 mm long. Ovary 0.5–1 mm long, green, glabrous. Style and stigma white, exserted from the corolla tube for 2–5 mm at anthesis; style densely covered with spreading, upwardly directed hairs in upper half; stigma with upper 1.5–2 mm fusiform, longitudinal papillate lines running down for a further 1–1.5 mm. Fruits spherical, 3–3.5 mm in diameter (persistent calyx not included), glabrous, drying dark brown, somewhat glossy and wrinkled when ripe; seeds ca. 2.5 mm in diameter, dark brown.

Dry deciduous forest, on sand (white sand and laterite); alt. 0–800 m.

Occurring in western Madagascar from 23° to 15° 30'S; recorded in the Atsimo-Andrefana, Menabe, Melaky and Sofia Regions. Fig. 14D.

Flowering: December–January; Fruiting: January–March.

Kerehetika (Martin 8252-SF); masonjohany (dialect Sakalava; Rabarivola 19861-SF); taolakena (dialect Sakalava; Ravelosaona 6592-SF); vahona (Harmelin 10202-RN bis).

Wood used by Sakalava against headaches (Razafimandimbison & Bremer 487).

Pseudocoptosperma menabense strongly resembles a Coptosperma species. Like Coptosperma, it has coriaceous, glabrous leaves and terminal, sessile, compact inflorescences with pentamerous white flowers with small-sized corolla tubes, bracteoles, ovaries, calyx tubes and calyx lobes. Furthermore, the fruits have a single ruminate seed. However, P. menabense is unique within the group of species currently brought together under the name Coptosperma by the combination of the keeled triangular stipules with well-developed awn and the placentation (3 ovules pendulous from a small placenta attached to the upper half of the septum). Some Coptosperma species also have three pendulous ovules but their stipules are of a different type, notably, the “bec du canard” type (Capuron 1973). In this case the stipular sheaths are flat with a rounded or obtuse apex, i.e. they are pressed against each other in such a way that their margins meet without overlapping (De Block et al. 2001: fig. 1), whereas the stipules in P. menabense are folded around each other (visible only in the youngest stipule pair). Species without the “bec du canard” stipule type usually have ovules (1 to 3) impressed in a large placenta. – Some specimens in the herbarium TEF bear the name Enterospermum menabense Capuron, but the species was hitherto not formally described.

Vulnerable: VU B1ab(i,ii,iii,iv) + 2ab(i,ii,iii,iv). The extent of occurrence (EOO) of Pseudocoptosperma menabense, estimated to be 86,558 km², exceeds the limits for the Vulnerable status under sub-criterion B1 but its area of occupancy (AOO), estimated to be 117 km², falls within the limits for the Endangered category under sub-criterion B2. The species occurs in five locations, two of which are in protected areas: Zombitse-Vohibasia National Park and Kirindy Mitea National Park. The species is known from 16 collections, half of which were collected after the year 2000. The major threat for this species is habitat loss by logging for charcoal and timber, burning for grazing and slash-and-burn agriculture both inside and outside the protected areas (Nicoll and Langrand 1989). Hence, based on the above information, the species is listed as Vulnerable.

MADAGASCAR. Mahajanga Province: Ménabé, forêt de Tsimembo, E d’Ambereny, Antsalova, 29–31 Mar 1966 (fr.), Capuron 24598-SF (BR, P, TEF); Antsalova, Ambereny, 11 Jan 1959 (fr.), Harmelin 10202-RN bis (BR, P, TEF); region of Port Bergé, along RN6, 242 m, 18 Mar 2010 (fr.), De Block, Groeninckx & Rakotonasolo 2354 (BR, G, K, MO, P, S, TAN); forêt Tsimembo, dans la concession Barthe, district Antsalova, 17 Mar 1961 (fr.), Rabarivola 19861-SF (P, TEF); Toliara Province: forêt de Jarindrano, rive gauche du haut Fiherenana, E de Maromiandry, Sakaraha, 29 Dec 1961 (fl.), Capuron 20569-SF (BR, P, TEF); forêt d’Andranomena, entre Andranomena et Marofandilia, Morondava, 19 Jan 1962 (fl.), Capuron 20895-SF (BR, P, TEF); Morondava District, forêt de Kirindi, CFPF Morondava (forêt d’Andalandahalo), jardin botanique 2, c. 45 km NE of Morondava, 10 m, 20 Feb 2000 (fr.), Davis, Rakotonasolo & Wilkin 2564 (BR, K, TAN); Kirindi forest, N part - Conoco 7, 16 m, 19 Jan 2007 (fr.), De Block, Rakotonasolo, Groeninckx & Dessein 2187 (BR, MO, P, TAN); Morondava, Kirindi Forest, close to ecotourist camp, 73 m, 20 Jan 2007 (fr.), De Block, Rakotonasolo, Groeninckx & Dessein 2208 (BR, K, MO, P, TAN); Zombitse-Vohibasia National Park, Zombitse, 31 Jan 2007 (fr.), De Block, Rakotonasolo, Groeninckx & Dessein 2257 (BR, K, MO, P, TAN); Lamboukily, 14 km of base camp in Kirindi, 42 m, 20 Jan 2007 (fr.), Groeninckx, Rakotonasolo, Dessein & De Block 102 (BR, MO, P, TAN); Lamboukily, 14 km of base camp in Kirindi, 42 m, 20 Jan 2007 (fr.), Groeninckx, Rakotonasolo, Dessein & De Block 108 (BR, MO, P, TAN); Menabe, 55 km NE of Morondava, route 8 at CPPF, Kirindy forest, 0.25 to 0.5 km NE of principal concession road, 4.5 km E of route 8, block CN4 and CN5, 35 m, 19–20 Mar 1992 (fr.), Noyes, Harder, Rakotobe, Razafindrabeaza & Abraham 1039 (BR, K, MO, P); forêt d’Andranofotsy situé 5 km N du village du même nom, Belo, Tsirihihina, 4 Jan 1953 (fr.), Ravelosaona 6592-SF (BR, TEF); Atsimo-Andrefana, Zombitse-Vohibasia National Park, along Ritik’ala trail, 700 m from the start at the carpark, 750–800 m, 3 Dec. 2003 (fl.), Razafimandimbison & Bremer 487 (UPS).

Differing from Paracephaelis sericea by the presence of shoot dimorphism, the smaller leaves grouped terminally on lateral short-shoots (blades ≤ 3.5 × 1.5 cm vs. 7–21 × 4.5–12 cm in P. sericea), the uni- or pauciflorous inflorescences (1–5 vs. 15 to numerous flowers), the trilobate bracts and bracteoles (vs. triangular), the variability in the number of calyx lobes [(4–)5–7 vs. 5], the pollen without supratectal elements (vs. supratectal elements present) and the fruit with 2 ruminate seeds (vs. 4–10 seeds with entire endosperm).

Tulearia splendida De Block.

Shrubs; shoot dimorphism present: vegetative long-shoots with well-developed internodes, reproductive short-shoots with compressed internodes and densely beset with corky stipular remnants; vegetative parts densely pubescent. Leaves grouped terminally on short-shoots, persistent, petiolate with petioles short and canaliculate above; blades < 4 × 1.5 cm, coriaceous; domatia absent; margins strongly revolute. Stipules triangular with short acuminate tip, inside densely covered with appressed hairs all over (hairs visible along the margins from the outside) and with large colleters in the lower half. Inflorescences terminal, sessile, uni- or pauciflorous, cymose with trichotomous branching; all parts (axes, bracts, bracteoles, pedicels) densely pubescent; bracts and bracteoles trilobate. Flowers hermaphroditic, pentamerous, shortly pedicellate; all parts (ovary, calyx, corolla) densely pubescent outside; secondary pollen presentation present. Calyx well-developed, either with short tube and long lobes or with lobes as long as or shorter than tube; lobes (4–)5–7(–8). Corolla white, sericeous outside; tube narrowly cylindrical; lobes contorted to the left in bud and spreading at anthesis. Stamens inserted in the sinuses of the corolla lobes at or somewhat below the level of the throat; filaments short; anthers usually partly included in the corolla tube at anthesis, basifixed, with sagittate base and short sterile apical appendix. Disc annular, fleshy, glabrous. Ovary cup-shaped, bilocular or rarely trilocular; placentation axile, with 3–7 ovules arranged along the periphery of a small placenta attached to the upper half of the septum. Style and stigma white, exserted from the corolla tube at anthesis; stigmatic lobes fused over their entire length except for the very tips, receptive zone on the adaxial surfaces of the free tips and along the lines of fusion of the lobes. Fruits drupaceous, subspherical, pubescent, crowned by the persistent calyx, containing 2 pyrenes; pyrenes crustaceous or stony, hemi-ovoid, formed by the convex outer and flat inner parts of each locule, with central apical protuberance or ridge on the adaxial side, opening along a central longitudinal preformed germination slit present over the entire length on the abaxial and adaxial sides (running through apical ridge or apical protuberance), containing 1 or very rarely 2 seeds; seed hemi-ovoid (or angular in case of 2 seeds per pyrene); hilum ovate, superficial, annulus around hilum absent; exotesta cells parenchymatic and filled with tannins; endotesta consisting of crushed cell layers without crystals; endosperm ruminate. Pollen grains 3-zonocolporate, exine microreticulate to perforate, supratectal elements absent.

A genus of two species, restricted to the dry forest and scrub of southern and southwestern Madagascar, on calcareous soil.

The genus is named for its occurrence in the region of Toliara (Tuléar).

Differing from Paracephaelis sericea by the habit (shrub vs. tree 5–16 m tall), the small leaves (1–3.5 × 0.6–1.5 cm vs. 7–21 × 4.5–12 cm), the pauciflorous inflorescences (1–5 vs. 15 to numerous flowers), the trilobate bracts and bracteoles (versus triangular and vaulted), and the fruit with 2 hemi-ovoid ruminate seeds (vs. 4–10 laterally flattened seeds with entire endosperm).

Figure 3. 

Tulearia. A–D Tulearia splendida: A habit B flowers C young inflorescence showing calyces and flower buds D young fruits E–K T. capsaintemariensis: E, F habit G branching pattern H top view of flower I lateral view of flower J ovary and calyx K fruit. Photographs: P. De Block (A, B, E, F, J), S. Dessein (C, D), M. Strack Van Schijndel (G–I), I. Van der Beeten (K).

MADAGASCAR. Toliara Province, La Table, ca. 15 km from Tuléar on RN 7, 4 Jan 1999 (fl.), De Block, Leyman, Dessein, Rakotonasolo & Randriamboavonjy 542 (holotype: BR!; isotypes: BR!, K!, MO!, P!, TAN!).

Shrub to 4 m tall, but usually smaller, densely branched; young shoots brown, moderately to densely covered with erect or spreading hairs, rapidly becoming corky with loss of pubescence; older branches brown, pale brown, fawnish or greyish, corky and somewhat flaking. Leaves elliptic, narrowly elliptic, more rarely obovate, narrowly obovate, ovate or narrowly ovate, 10–35 × 6–15 mm; blades coriaceous, drying brownish-green to brown above, somewhat paler below, densely covered with short erect and more sparsely with long appressed hairs above, lanate but often with hairs more appressed on midrib and secondary nerves below; base cuneate to attenuate; apex rounded and mucronate; midrib and secondary nerves raised on the lower leaf surface; midrib impressed on the upper leaf surface; 4–6(–7) secondary nerves on each side of the midrib. Petioles 1.5–4 mm long, densely covered with appressed or spreading hairs. Stipules caducous, densely covered with appressed hairs outside but rapidly becoming corky and losing the pubescence; sheaths triangular, 2–4 mm long, tips 1–2 mm long. Inflorescences with 1–5 flowers but usually 3-flowered, compact; inflorescence axes, pedicels, bracts and bracteoles densely covered with appressed or spreading to erect hairs; bracts and bracteoles with short petiole-like stalk, trilobate; first order bracts either with central lobe a petiolate leaf similar in size and shape to vegetative leaves and lateral lobes linear, 2–4 mm long, or central lobe narrowly triangular or linear up to 1 cm long and lateral lobes linear, usually much shorter than the central lobe; higher order bracts similar in shape and size to bracteoles; bracteoles subopposite on the pedicel just below the ovary, 8–12 mm long, consisting of a ca. 1 mm long petiole-like stalk, an ca. 1 mm high sheath, a narrowly oblong central lobe, 5–11 × 1–1.25 mm, and 2 linear lateral lobes, 3–5 mm long; bracts and bracteoles densely covered with appressed to erect hairs all over and with scattered colleters in the sheath inside. Flowers sweetly scented, shortly pedicellate, pedicels 1.5–2.5 mm long. Calyx pale green or green, considerably wider than the ovary; tube ca. 1 mm long, densely covered with appressed to spreading hairs outside, densely covered with appressed hairs and with a prominent ring of colleters at the base inside; lobes (4–)5–7(–8), leaf-like or narrowly oblong, somewhat variable in size within a flower, 7.5–10 × 1–2 mm, sometimes linear interstitial lobes up to 5 mm long present or lobes shallowly or profoundly split lengthwise (with two tips), moderately to densely covered with appressed to spreading hairs outside and inside, bases not overlapping but closely joining, tips acute. Corolla sericeous outside; tube 4–30 mm long, ca. 1.5 mm in diameter at the base, 3.5–4.5 mm in diameter at the throat, basal half densely covered with erect hairs inside; lobes oblong or rarely square, 4–8 × 3–6 mm, inner surface glabrous and drying orange or blackish-brown and contrasting with the white pubescence of the corolla tube, margins densely ciliate, tip blunt and emarginate, somewhat assymetrical. Stamens inserted in the sinuses of the corolla lobes at the level of the throat, their bases often included in the corolla tube at anthesis; filaments < 1 mm long; anthers 4–4.5 mm long. Ovary ca. 1.5 mm long, pale green or green, densely covered with erect hairs, often longitudinally ribbed when dry. Placenta attached somewhat above the midde of the septum, with (3–)4–5 ovules arranged along its periphery. Style and stigma white, exserted from the corolla tube for 4–6 mm at anthesis; style sparsely covered with upwardly directed hairs in the upper half (somewhat below the receptive zone of the stigma); stigmatic tips free and spreading for ca. 1 mm, receptive zone ca. 8 mm long, the upper 3–4 mm fusiform, the lower 4–5 mm not widened. Fruits bilobed, 4.5–5 × 5.5–6.5 mm (persistent calyx not included), densely covered with short erect hairs, drying brown or blackish, glossy and somewhat wrinkled when ripe; 2 pyrenes per fruit, crustaceous, with a central apical cuspidate protuberance on the adaxial side; seeds 1(–2) per pyrene, ca. 3.5 × 3–3.5 mm, brown.

Open-canopy dry forest, spiny forest, xerophytic thicket, dry scrub, on calcareous soil, both rocky and sandy (e.g. dunes); alt. 0–350 m.

Only known from the Atsimo-Andrefana Region in southwestern Madagascar. Fig. 14E.

Flowering: December–April; Fruiting: from March onwards.

Toalanambata (coll. ignot. 31297-SF).

Medicine for the eyes (“fanafody des yeux”; Dequaire 27496).

Tulearia splendida strongly resembles Paracephaelis by the general hairiness of the whole plant, the robust sericeous flowers, the well-developed calyx and the ovules arranged on the periphery of the placenta. Tulearia differs from Paracephaelis by its seeds (2 hemi-ovoid, ruminate seeds in Tulearia vs. 4–10 laterally flattened seeds with entire endosperm in Paracephaelis) and by its pollen (tectum without supratectal elements vs. tectum with supratectal elements).

Vulnerable: VU B1ab(i, ii, iii, iv) + 2ab(i, ii, iii, iv). The extent of occurrence (EOO) of Tulearia splendida is estimated to be 21,644 km², which exceeds the upper limits for the Vulnerable status but its area of occupancy (AOO), estimated to be 342 km², falls within the limits of the Endangered category. The species is well-collected with both recent and historic specimens. Tulearia splendida occurs in eight locations, only one of which lies in a protected area, the Tsimanampetsotsa National Park. The species is threathened by habitat loss as a result of grazing, subsistence farming, logging for timber and charcoal and burning to improve grazing. Based on the above information, the species is assessed as Vulnerable.

Some specimens in the herbarium P were annotated as Randia tulearensis Homolle (Perrier de la Bâthie 12816 & 19025). In his unpublished work on the Madagascan Rubiaceae, Capuron (1973) used the name Enterospermum tulearense (Homolle) Capuron. But no formal description of the species was hitherto attempted.

MADAGASCAR. Toliara Province: Tuléar, 16 Dec 1912 (fl.), Afzelius s.n. (P); Tuléar, 16 Dec 1912 (fl.), Afzelius 265 (S); Tuléar, Saint Augustin, 20 Dec 1912 (fl.), Afzelius 268 (S); Tuléar, 16 Dec 1912 (fl.), Afzelius 269 (S); 3 km S of Morombe towards Ampasilava, 14 m, 16 Sep 2006 (fr.), Andriamahay & Rakotoarisoa 1514 (K); Betioky, 350 m, 22 Apr 2004 (fr.), Andriamahay & Rakotoarisoa 750 (K); W d’Ejeda, 15 May 1951 (fr.), Bosser 252 (P); plateau Mahafaly, Ankalirano, W d’Ejeda, Mar 1960 (fl., fr.), Bosser 14302 (P); Tuléar, falaises du Fiherenana, Feb 1962 (fl.), Bosser 15711 (P, TAN); environs du Lac Ihotry, N de Tongobory, 8 Jan 1962 (fl.), Capuron 20713-SF (BR, P, TEF); vers le PK 28 de la route Tuléar-Sakaraha, Dec 1961 (fl.), Capuron & Chauvet 20777-SF (P, TEF); environs de Tuléar, bas de La Table, 4 Mar 1961 (fl.), Chauvet 51 (BR, P, TEF); environs de Tuléar, route de Sarodrano, 11 Mar 1961 (fl.), Chauvet 74 (P); km 22 sur route d’Antananarivo, Tuléar, 9 Nov 1961 (fl.), Chauvet 180 (BR, P, TEF); ancienne route de Sarodrano, Tuléar, 13 Nov 1961 (fl.), Chauvet 198 (BR, P, TEF); route de Sarodrano, 14 Nov 1962 (fl.), Chauvet 362 (P, TEF); La Table, Tuléar, 10 Mar 1963 (fr.), Chauvet 412 (P); Tsimanampetsotsa, Réserve de Manampetsotsa, Lac Folifoetsy, commune Behelony, district Ejeda, 13 Mar 1987 (fr.), coll. ignot. 31297-SF (TEF); vicinity of Tongobory along banks of Onilahy River, 60 m, 14 Feb 1975 (fl.), Croat 31182 (K, P, MO, TAN); dunes on road to Ifaty, 17 m, 2 Feb 2007 (fl.), De Block, Dessein, Groeninckx & Rakotonasolo 2287 (BR, MO, P, TAN, WAG); La Table, 94 m, 2 Feb 2007 (fl.), De Block, Dessein, Groeninckx & Rakotonasolo 2301 (BR, MO, P, TAN, WAG); Ampasimariry, close to Lac Tsimanampetsotsa, Ambola, commune Beheloke, district Tulear II, 14 m, 4 Feb 2007 (fl.), De Block, Dessein, Groeninckx & Rakotonasolo 2311 (BR, MO, P, TAN); Morombe, s.dat. (fr.), Decary 18720 (BR, P); La Table, Tuléar, s.dat. (fl.), Dequaire 27496 (P); environs de Tulear, Ankilibe, 5 Feb 1957 (fl.), Descoings 2326 (TAN); province de Tuléar, s.dat. (fl.), Géay 32 (P); road towards Betioky, 171 m, 3 Feb. 2007 (fr.), Groeninckx, Rakotonasolo, Dessein & De Block 209 (BR, MO, P, TAN, WAG); Lac Tsimanampetsotsa, 5 Feb 2007 (fr.), Groeninckx, Rakotonasolo, Dessein & De Block 216 (BR, G, MO, P, TAN); piste ‘Ajax’, 30 km N de Tuléar, 10 Dec 1968 (fl.), Guillaumet 2288 (BR, P, TAN); Itampolo, Lac d’Itampolo, s.dat. (fr.), Homolle s.n. (P); Tuléar, s.dat. (fr.), Homolle 1566 (P); Lac d’Itampolo, s.dat. (fr.), Homolle O5 (P); gorges de Fiherenana entre Beantsy et Anjamala, 30–300 m, 16–19 Jan 1947 (fl.), Humbert 19889 (BR, P); Manambo, près de la mèr, 20 m, 29–30 Jan 1947 (fl.), Humbert 20089 (BR, P); embouchure de la Menarandra, Bevoalava-Ankazondranto, 1–150 m, 12 Mar 1955 (fr.), Humbert & Capuron 29386 (BR, P); plateau Mahafaly, W de Betioky, 100–300 m, 17–20 Mar 1955 (fr.), Humbert & Capuron 29485 (BR, P); environs de Tuléar, sur la Table, SW flanc, Mar 1960 (fr.), Keraudren 583 (BR, P); environs de Tuléar, bord de mèr, sur les dunes près du village d’Ankilibe, Mar 1960 (fr.), Keraudren 613 (P); plateau calcaire Mahafaly, près du village d’Ankaliano, W d’Ejeda, Mar 1960 (fr.), Keraudren 854-bis (P); Mahafaly, près du village d’Ankaliano, SW de Betioky, Mar 1960 (fr.), Keraudren 856 (P); environs de Tuléar, gorges de Fiherenana, Feb 1962 (fl.), Keraudren 1347 (P); route d’Ampanihy à Androka, 37 km SW d’Ampanihy, colline E de la piste, 230–260 m, 6 Feb 1990 (fr.), Labat, Du Puy & Phillipson 2081 (K, P); Tuléar, La Table, E of town on the road to Antananarivo, 60 m, 28 Feb 1993 (fl.), Luckow 4170 (BR, MO, TAN); 20–30 km N of Tulear on road to Morombe, 5–10 m, 27 Dec 1988 (fl.), Miller & Miller 3791 (K, MO, P, TAN); 14 km SE of Tulear on RN 7, 100 m, 24 Mar 1991 (fr.), Miller & Randrianasolo 6131 (K, MO, P, TAN); route Sahodona-Ampanihy, Mar 1964 (fl.), Morat 637 (P); plateau Mahafaly, Jan 1910 (fl.), Perrier de la Bâthie 9516 (P); environs de Tuléar, Aug 1919 (fl.), Perrier de la Bâthie 12816 (BR, P); environs de Tuléar, Apr. 1933 (fr.), Perrier de la Bâthie 19025 (P); Manampetsa, Apr 1933 (fl.), Perrier de la Bâthie 19124 (P); E of Tulear, around la Table, 100 m, 5 Jan 1989 (fl.), Phillipson 3082 (BR, K, MO, P, TAN, WAG); Réserve de Tsimanampetsotsa, NW corner, 50 m, 11 Jan 1989 (fl., fr.), Phillipson & Rabesihanaka 3156 (K, MO, P, TAN, WAG); 12 km N of Betsiky on road to Tongobory along E facing calcareous escarpment, 300 m, 13 Feb 1990 (fr.), Phillipson 3498 (K, MO, P, TAN); Atsimo-Andrefana, N of Toliara, between Fiherenana and Manombo rivers, Ranobe forest, Belalanda commune, c. 1 km from RN 9 along PK 32 track, 50 m, 15 Mar 2006 (fl.), Phillipson, Ranaivojaona, Andrianjafy & Lubke 5909 (BR, MO); dunes de Befanany, 15 Feb 1921 (fr.), Poisson 149 (P); Réserve naturelle X, Lac Tsimanampetsotsa, canton Soalany, district Betioky, 21 Mar 1953 (fl.), Raodonanahany 5015-RN (BR, P, TAN); Manombo, near PK 32, 11 Dec 2004 (fl.), Rakotonasolo, Smith, Hoffmann, Ralimanana & Sharon 878 (BR, K, MO, P); 1 km before Tongobory on PK 58 from Andranovory, 12 Dec 2004 (fl.), Rakotonasolo, Smith, Hoffmann, Ralimanana & Sharon 885 (BR, K); Tuléar II, Belalanada, Ranobe, forest E of allée des baobabs, 159 m, 26 Jan 2007 (fr.), Ranaivojaona, Manjakahery & Andrianjafy 1699 (K, MO); Andatabo, 18 km S de Tuléar, bord de la RN 7, 50–100 m, 5 Feb 1999 (fr.), Randrianaivo, Ralimanana, Randrianasolo, Andriantiana, Rakotondrajaona & Rahelivololona 325 (BR, K, MO, P); on road to Saint Augustin, 25 km from Tuléar, 17 Feb 1998 (fr.), Razafimandimbison 281 (MO); Betioky, 10 km S of Tongobory, along road to Andranovory, 11 Dec 2003 (fl.), Razafimandimbison 526 (UPS); Betioky, 10 km S of Tongobory, along road to Andranovory, 11 Dec 2003 (fl.), Razafimandimbison 530 (S, UPS); Toliara II, Ranobe, 32 m, 19 Mar 2007 (fr.), Razanatsoa & Manjakahery 369 (BR, MO, P, TAN).

Differing from T. splendida by the smaller leaves (5–20 × 3.5–5.5 mm vs. 10–35 × 6–15 mm in T. splendida), the secondary nerves which are invisible on both leaf surfaces (vs. visible in T. splendida), the uniflorous inflorescences (vs. 1–5 flowers), the shorter bracteoles (up to 3 mm vs. 8–12 mm long), the shorter calyx lobes (1–2 mm vs. 7.5–10 mm long) and the longer calyx tube (1.5–3 mm vs. ca. 1 mm long).

MADAGASCAR. Toliara Province, Fort-Dauphin, road between Faux-Cap and Marovato, 124 m, 3 Apr 2010 (fl., fr.), Groeninckx, De Block & Rakotonasolo 309 (holotype: BR!; isotypes: BR!, K!, MO!, P!, TAN!).

Shrub, 0.5–1.5 m high; young shoots brown, densely covered with spreading hairs, rapidly becoming corky with loss of pubescence; older branches pale brown, fawnish or greyish, corky. Leaves elliptic, narrowly elliptic or rarely broadly elliptic, 5–20 × 3–5.5 mm; blades thickly coriaceous, drying brown to blackish brown and somewhat glossy above, somewhat paler below, densely covered with short erect hairs above, lanate but often with hairs more appressed on midrib below; base obtuse to rounded; apex rounded and mucronate; midrib raised in the basal half on the lower leaf surface, somewhat impressed on the upper leaf surface; secondary nerves invisible on both surfaces. Petioles 1–2 mm long, densely covered with appressed or spreading hairs. Stipules caducous, moderately to densely covered with appressed hairs outside but rapidly becoming corky and losing the pubescence; sheaths triangular, 1.5–2 mm long; tips 0.5–1.25 mm long. Inflorescences uniflorous; bracteoles opposite at the base of the ovary, trilobate or, rarely, reduced to a single lobe; if trilobate, then consisting of a ca. 0.5 mm high basal sheath, 2 linear or narrowly triangular lateral lobes, 0.5–1.5 mm long, and a central lobe, either linear and 1.5–3 mm long or more rarely leaflike (petiole to 2 mm long, blade to 6 × 2 mm, shape identical to that of vegetative leaves), bracteoles moderately covered with appressed or spreading hairs outside, lateral lobes and base of central lobe densely covered with appressed hairs and a few large colleters inside, central lobe higher up round in cross-section and pubescence on adaxial surface identical to that on abaxial surface. Flowers sessile. Calyx green, densely covered with erect or spreading hairs outside, densely covered with appressed hairs inside; tube 1.5–3 mm long, with a ring of colleters at the base inside, the colleters more densely present in the region of the sinuses of the calyx lobes; lobes (4–)5–7, ovate, 1–2 × ca. 1 mm, somewhat keeled when dry, bases not overlapping but closely joining, tips acute to rounded. Corolla sericeous outside; tube 8–10(–18*) mm long, 1.5–2 mm in diameter at the base, 3–4 mm in diameter at the throat, basal half densely covered with erect hairs inside; lobes oblong, 6–7(–12*) × 3.5–4(–5.5*) mm, glabrous inside, margins densely ciliate, tips rounded. Stamens inserted in the sinuses of the corolla lobes ca. 1.5 mm below the level of the throat, only upper half exserted from corolla tube at anthesis; filaments < 1 mm long; anthers ca. 4 mm long. Ovary 1.5–2 mm long, green, densely covered with erect or spreading hairs, faintly ribbed longitudinally when dry. Placenta attached to the upper half of the septum with 5–7 ovules arranged along its periphery. Style and stigma white, 12–14(–22*) mm long, exserted from the corolla tube for 4–5 mm; style densely covered with upwardly-directed spreading hairs in the lower half; stigma fusiform, stigmatic tips free and spreading for ca. 1 mm, receptive zone 6–7(–9*) mm long, widened over the entire length. Fruits bilobed or rarely trilobed, 5–5.5(–7*) × 4.5–5(–7*) mm (persistent calyx not included), densely covered with short erect hairs; when mature, fruit and persistent calyx tube black, calyx lobes remaining green; 2(–3) pyrenes per fruit, stony, with a central vertical ridge apically on the adaxial side; 1 seed per pyrene, ca. 3.5–4 × 3 mm.

Open-canopy dry scrub, on calcareous soil, alt. 0–150 m.

Only occurring along the coast in the Androy Region in southern Madagascar. Fig. 14F.

Flowering & fruiting: April.

Measurements indicated with * in the description are from a specimen grown in greenhouse conditions.

Critically Endangered: CR B1ab(i, ii, iii, iv) + 2ab(i, ii, iii, iv). The extent of occurrence (EOO) of Tulearia capsaintemariensis cannot be calculated because only two specimens have ever been collected, but it can be estimated that the EOO is below 100 km². Its area of occupancy (AOO) is 18 km² using a cell width of 3 km but 8 km² using a cell width of 2 km. The species was only discovered in 2010 and occurs in a single location which is not included in a protected area. The main threat to the species is habitat loss as a result of grazing, subsistence farming or land clearing for sisal plantations. Based on the above information, the species is assessed as Critically Endangered.

MADAGASCAR. Toliara Province: près de Cap Sainte Marie National Park, Valala, 21 m, 4 Apr 2010 (fr.), De Block, Groeninckx & Rakotonasolo 2421 (BR, G, K, MO, P, TAN).