Tulearia splendida De Block, 2018

Block, Petra De, Rakotonasolo, Franck, Ntore, Salvator, Sylvain G. Razafimandimbison, & Janssens, Steven, 2018, Four new endemic genera of Rubiaceae (Pavetteae) from Madagascar represent multiple radiations into drylands, PhytoKeys 99, pp. 1-66 : 1

Tulearia splendida De Block sp. nov. Figs 3A-D , 11 , 13G-I, L, M

Diagnosis.

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).

Type.

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.

Habitat.

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

Distribution.

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

Phenology.

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

Vernacular names.

Toalanambata (coll. ignot. 31297-SF).

Vernacular uses.

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

Critical note.

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).

Preliminary IUCN assessment.

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.

Critical notes.

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.

Additional specimens examined.

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, Reserve 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 O 5 (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); Reserve 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) .

Figures

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).

Figure 11. Tulearia splendida. A, flowering branch B stipule C bracteole, ovary and calyx D bract E corolla, style, stigma and anthers F fruit G seed H cross-section through seed. A-E De Block et al. 542 F-H Groeninckx et al. 209.

Figure 13. Pollen of Pseudocoptosperma and Tulearia. A-C, J Pseudocoptosperma menabense D-F, K Tulearia capsaintemariensis G-I, L, M T. splendida. A, D, G polar view B, E, H equatorial view C, F, I mesocolpium J-L ectoaperture M pollen grain wall. A-C, J Capuron 20569 - SF; D-F, K Groeninckx et al. 309 G-I, L, M Capuron 20777 - SF.

Figure 14. Distribution maps. A Exallosperma longiflora B Helictosperma malacophylla C H. poissoniana D Pseudocoptosperma menabense E Tulearia splendida F T. capsaintemariensis.