Diospyros malandy H.N. Rakouth, Randrianaivo, G.E. Schatz & Lowry, 2021

Schatz, George E., Lowry Ii, Porter P., Rakouth, Hasina N. & Randrianaivo, Richard, 2021, Taxonomic studies of Diospyros (Ebenaceae) from the Malagasy region. VI. New species of large trees from Madagascar, Candollea 76 (2), pp. 201-236 : 222-234

Diospyros malandy H.N. Rakouth, Randrianaivo, G.E. Schatz & Lowry , sp. nov. ( Fig. 12 , 13A ).

Holotypus: MADAGASCAR. Reg. DIANA [Prov. Antsiranana]: Ankarana, 6.X.2018, fr., Randrianaivo et al. 3240 (MO-6956007!; iso-: DBEV!, P [ P00722673 ]!, TAN!) .

Diospyros malandy H.N. Rakouth, Randrianaivo, G.E. Schatz & Lowry resembles other members of the Sclerophylla group in having spherical fruit with accrescent, strongly reflexed calyx lobes, but is distinguished by its leaves with a cuneate to rounded base, margin that is slightly revolute and undulate, and rounded to retuse apex, and densely reticulate, evident tertiary venation.

Shrub to tree, 2–25 m tall, 3–29.5 cm DBH. Young stems densely covered with short, erect, yellowish to pale brown trichomes 0.1 mm long. Leaves distichous, alternate, borne 15 – 25 mm from one another, lamina 3.0 – 8.5 × 2 – 5 cm, obovate, chartaceous to subcoriaceous, discolorous in sicco, light grayish to pale olive green above, nearly glaucous below, matte on both surfaces, initially densely covered above with curly, semi-appressed, yellowish to pale brown trichomes c. 0.1 mm long, glabrescent, base cuneate to rounded, symmetric, margin slightly revolute and undulate, apex rounded to retuse, midvein flat above, raised below, to 2 mm wide at the base, narrowing toward the apex, with dense, semiappressed, whitish to light yellowish trichomes c. 0.1 mm long, glabrescent, venation semicraspedodromous, with 5–9 secondary veins per side, 2–3 originating from the base, raised above, more so below, tertiary venation reticulate, distinctly visible; petiole short, 1–2 mm long, c. 2 mm in diam., darker brown than the lamina, with dense, semi-appressed, yellowish to pale brown trichomes c. 0.1 mm long, glabrescent. Male and female flowers not seen. Fruits axillary, solitary, pedicel in fruit 5 – 11 mm long, 2 mm in diam. (to 3 mm in diam. distally), yellowish, covered with dense, appressed, pale yellow to whitish trichomes 0.1 mm long; fruiting calyx strongly accrescent, the base forming a short collar c. 2–4 × 3–4 mm, slightly quadrangular, 3–4-lobed, the lobes 9–15 × 6–8 mm, oblong, spreading to reflexed, adaxially convex, the surfaces discolorous in sicco, sparsely covered on both surfaces with appressed, whitish to light yellowish trichomes c. 0.2 mm long, glabrescent, margin weakly revolute, apex obtuse to rounded with distinct venation; fruit spherical to obloid or ovoid, 11–15 × 10–13 mm, minutely verrucose, green in vivo, with sparse, appressed, whitish to light yellow trichomes c. 0.2–0.4 mm long, glabrescent, apex apiculate. Seeds 4, 1 per locule, narrowly flattened ellipsoid, 2.5–3 × 0.9–1 mm, weakly verrucose, dark brown to black.

Etymology. – The specific epithet refers to the fact that the color of the bark is grayish white, which is atypical for the genus and is expressed in its local name, ‘Mapingo malandy’ (= white Diospyros ).

Vernacular names. – “Mapingo malandy” (Randrianaivo

3240).

Distribution and ecology. – Diospyros malandy is restricted to northern Madagascar, where it is known from two protected areas, Andrafiamena Andavokoera and Ankarana. It occurs in dry, semi-deciduous forest on tsingy, dry forest on basaltic soil, and dense, dry, degraded forest on reddish sand, as well as river banks, from c. 90 to 320 m elevation.

Phenology. – Flowering material of Diospyros malandy has been collected in February, and immature fruits have been collected in December.

Conservation status. – Diospyros malandy has a geographic range in the form of an Extent of Occurrence of 15 km ² and a minimum Area of Occupancy of 15 km ². It is present in two protected areas, Andrafiamena Andavokoera and Ankarana. It is also present along the roadside in the buffer zone of the Ankarana protected area, where it is threatened by forest clearing for agriculture, charcoal production, fire, and exploitation for firewood and house construction material, all of which will result in continuing decline. With respect to the most serious plausible threat of exploitation for firewood and house construction material, it exists at four locations. Therefore, D. malandy can be assessed for its risk of extinction as “Endangered” [EN B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].

8 mm

10 mm

[A: Bardot-Vaucoulon et al. 1316, P; B–D: Bardot-Vaucoulon et al. 1186] [Drawing: Alain Jouy]

branch with immature fruit; B. Diospyros mimusops G.E. Schatz & Lowry , branch with fruit; C–E. Diospyros rakotovaoi G.E. Schatz & Lowry : C. Branch with fruit; D–E. Male flowers.

[A: Randrianaivo 3240; B: Razakamalala 8563; C: Razakamalala et al. 3190; D–E: Razafitsalama 1141] [Photos: A: R. Randrianaivo;

B: S. Andrianarivelo; C: C. Rakotovao; D–E: J. Razafitsalama]

Notes. – Diospyros malandy belongs to the Sclerophylla group (see notes under D. bardotiae for a list of its diagnostic features), within which it can be distinguished by its small fruits with a strongly accrescent, reflexed calyx, and leaves in dried material that have evident, densely reticulate tertiary venation and are usually light grayish to pale olive green above (often appearing matte to nearly glaucous), with a cuneate to rounded base, margin that is slightly revolute and undulate, and rounded to retuse apex.

Additional specimens examined. – MADAGASCAR. Reg. Diana [Prov. Antsiranana]: Ankarana , 14. I .2003, fr., Bardot-Vaucoulon 1186 ( K, MO, P, TAN); ibid. loco, 25. I .2003, fr., Bardot-Vaucoulon 1316 ( K MO, P, TAN); Ampondrabe, 12.II.2006, ♂ fl., Rakotondrajaona 387 ( CNARP, MO, P, TAN) ;

Ankarana, 146 m, 17. I.2014, ster., Randrianaivo 2406 ( BR, DBEV, G, MO, P) ; Sajoavato, Sahafary, 25. I.2014, ster., Randrianaivo 2460 ( BR, DBEV, G, MO, P) ; Sahafary, 25. I.2014, ster., Randrianaivo 2462 ( BR, DBEV, G, MO, P) ; ibid. loco, 8.X.2018, ster., Randrianaivo et al. 3263 ( DBEV, MO, P, TAN) ; ibid. loco, ster., Randrianaivo et al. 3264 ( DBEV, G. MO, P) ; ibid. loco, ster., Randrianaivo et al. 3265 ( DBEV, G, P) ; ibid. loco, 9.X.2018, y.fr., Randrianaivo et al. 3275 ( DBEV, G, MO, P, TAN) ; Mahamasina, Ankarana, 9.X.2018, y.fr., Randrianaivo et al. 3276 ( DBEV, G, MO, P, TAN) ; Ankarana, 22.III.2020, ster., Ravaoherinavalona et al. 118 ( DBEV, MO, P, TAN) ; ibid. loco, ster., Ravaoherinavalona et al. 135 ( DBEV, MO, P, TAN) ; ibid. loco, ster., Ravaoherinavalona et al. 137 ( DBEV, MO, P, TAN) ; ibid. loco, ster., Ravaoherinavalona et al. 138 ( DBEV, MO, P, TAN) ; ibid. loco, ster., Ravaoherinavalona et al. 139 ( DBEV, MO, P, TAN) ; ibid. loco, 26.III.2020, ster., Ravaoherinavalona et al. 144 ( DBEV, MO, P, TAN) .

Diospyros melanocarpa G.E. Schatz & Lowry , sp. nov. ( Fig. 14 ).

Holotypus: MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Farankaraina, 19.IX.1957, fr., Service Forestier 18343 (MO-6956008!; iso-: G [ G00341738 ]!, K!, P [ P00722678 , P03829465 ]!, TEF [ TEF000891 ]!) .

Diospyros melanocarpa G.E. Schatz & Lowry resembles other members of the Decaryana group in having male flowers borne in cymose inflorescences and 1–3 axillary, short pedicellate fruits with an urceolate calyx and broadly triangular calyx lobes with revolute margins and a reflexed apex, but is distinguished by having shiny, glabrous, black fruits and small leaves [1.5–6.5(–8.7) × 0.8–4 cm].

Tree 15–20 tall, 30 cm DBH. Young stems terete, sparsely covered with very short, appressed and erect, whitish to golden trichomes c. 0.3 mm long, and an irregular, white, crystal-like, waxy substance (perhaps an artifact of drying), also sometimes present on leaves, pedicels, and fruits. Leaves pseudodistichous, lamina 1.5–6.5(–8.7) × 0.8–4 cm, elliptic to rarely slightly obovate, chartaceous, glabrous above, initially rather densely covered with very short, appressed, white trichomes c. 0.3–0.4 mm long below, more densely so along the midvein, glabrescent, base cuneate, margin slightly revolute, apex acute to rounded, midvein flat to slightly impressed above, slightly raised at the base to flat at the apex below, venation weakly brochidodromous, with c. 7 secondary veins per side, mostly indistinct above, flat and indistinct below, tertiary venation finely reticulate; petiole 4–6 mm long, 1 mm in diam., canaliculate, sparsely covered with erect, white trichomes c. 0.3 mm long. Male flowers in axillary cymose inflorescences, 2– 15 flowers per inflorescence, to 6 mm long, axes 0.5 mm in diam., rather densely covered with appressed, light golden trichomes c. 0.1–0.2 mm long, pedicel c. 1 mm long, 0.1 mm in diam., rather densely covered with appressed, light golden trichomes c. 0.2–0.3 mm long; calyx obconical, square in cross-section, 2– 2.5 × 2 mm, 4-lobed, triangular (1 × 1 mm), with moderately appressed, light golden trichomes 0.3 –0.5 mm long outside, glabrous inside; corolla narrowly obconic, square in cross-section, 4.5 × 2.5 mm, 4-lobed, the lobes broadly ovate to triangular, 1 × 1 mm, densely covered with appressed, light golden trichomes to 1 mm long on the outside of the lobes and extending downward along the 4 corners, the rest of corolla glabrous outside, glabrous inside except for a tuft of trichomes at the apex of the lobes; stamens 12, inserted on the corolla at two levels 1/3 and 2/3 of the way up the corolla; filaments 0.5 mm long; anthers 0.8 mm long, ovoid; pistillode present, hemispherical, 0.5 mm long, glabrous, black in sicco. Female flowers not seen. Fruits 1–3 in axils of leaves or fallen leaves, sessile or pedicel in fruit c. 1–2 mm long, subtended by several bracteoles; fruiting calyx urceolate, 4 mm long, 4 mm in diam. at base, to 7–9 mm in diam. at the apex, 4–5-lobed, the lobes 4–5 × 5–6 mm, broadly triangular, densely covered with appressed, whitish gray trichomes c. 0.2–0.5 mm long, margins strongly revolute, apex strongly reflexed; fruit depressed ellipsoid-oblongoid to depressed spherical, 12–15 × 9–14 mm, drying black, initially rather densely covered with semiappressed, whitish golden trichomes c. 0.5– 1.3 mm long, glabrescent, crowned by the style/stigma remnant, 1–2 mm long. Seeds 4– 5, spherical wedge-shaped, 13–14 × 4–5 mm, glabrous, black, shiny.

Etymology. – This specific epithet refers to the black color of the fruits.

Distribution and ecology. – Diospyros melanocarpa occurs primarily in northeastern Madagascar, from the area around the Baie d’Antongil north to Anjambalava-Ambavala situated to the north of Sambava, with a single collection from Tampolo ( MADAGASCAR CATALOGUE, 2021). It is found in low- to mid-elevation humid forest.

Phenology. – Flowering material has been collected in October, and collections with fruits have been collected in April, May, September, and October.

Conservation status. – Diospyros melanocarpa has a geographic range in the form of an Extent of Occurrence of 17,047 km ² and a minimum Area of Occupancy of 28 km ². It is present in three protected areas, Makirovana, Masoala, and Tampolo. Outside of the protected areas and also at Tampolo, it is threatened by forest clearing for agriculture, fire, and exploitation for firewood and house construction material, all of which will result in continuing decline. With respect to the most serious plausible threat of exploitation for firewood and house construction material, D. melanocarpa exists at seven locations, and was recently assessed for its risk of extinction as “Vulnerable” [VU B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)] ( IUCN, 2021).

Notes. – Diospyros melanocarpa is a member of the Decaryana group, characterized by male flowers borne in axillary, cymose inflorescences, short pedicellate female flowers borne in 1–3-flowered axillary fascicles, and depressed spherical fruit with an urceolate calyx and broadly ovate to triangular calyx lobes, the margins revolute and the apex reflexed. Other members of the group include D. bezofensis H. Perrier , D. decaryana H. Perrier , D. decaryoides G.E. Schatz & Lowry , and D. implexicalyx H. Perrier , as well as two new species that remain to be described. Within the Decaryana group, D. melanocarpa is the only species with shiny, glabrous, black fruits, and can also be distinguished by its small leaves (1.5–6.5(–8.7) × 0.8–4 cm), a feature shared only with D. implexicalyx .

Additional specimens examined. – MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Tampolo , 11.X.2001, y.fr., Rabevohitra et al. 3869 ( MO, P, TAN, WAG) . Reg. SAVA [Prov. Antsiranana]: Ranovery , 26.X.2010, ♂ fl., Bernard & Ndrano 1695 ( MO, P, TAN) ; Mt. Anjenabe , 3–7.XI.1950, fr., Humbert & Capuron 18343 ( P) ; Nosiarina , 7. V.2000, y.fr., Rabenantoandro et al. 208 ( MO, P, TAN) ; Tsihomanaomby, 13. I.2021, fr., C. Rakotonirina & Martial 663 ( DBEV, MO, P, TAN) ; ibid loc., 12.XII.2020, y.fr., Razakamalala et al. 8821 ( MO, P, TAN) ; massif de l’Anjenabe, 7.IX.1950, fr., Service Forestier 770 ( G, K, MO, P [ P03829464 ], TEF, W) ; Andrapengy , 11.IV.1967, y.fr., Service Forestier 27730 ( G, MO, P, TEF) ; Cap-Est , 19–21.IV.1967, fr., Service Forestier 27763 ( G, MO, P, TEF) .

Diospyros mimusops G.E. Schatz & Lowry , sp. nov. ( Fig. 13B , 15 ).

Holotypus: MADAGASCAR. Reg. Anosy [Prov. Toliara]: Antsotso, 24.V.2006, fr., Antilahimena et al. 4887 (MO- 6336318!; iso-: P [ P06490534 ]!, TAN [ TAN002011 ]!, TEF!) .

Diospyros mimusops G.E. Schatz & Lowry resembles other members of the Tetraclis group in having leaves with a mucronate apex, male flowers borne in cymose inflorescences, and often valvate corolla aestivation, but is distinguished by having glabrous and sometimes glaucous young stems and leaves.

Tree 5–20 m tall, 8–17 cm DBH. Young stems terete to flattened, glabrous, sometimes glaucous in sicco. Leaves distichous, lamina 6–11.5 × 2.2–4.2 cm, elliptic to obovate, coriaceous, glabrous, sometimes shiny or glaucous above in sicco, base attenuate to acute, margin revolute, apex acute to shortly acuminate, occasionally rounded to retuse, acumen 2–3 mm long, usually with a very short, black mucron, midvein slightly impressed above, raised below, venation weakly brochidodromous, loops of secondaries ¼–1/ 3 in from margin, with 10–15 secondary veins per side, slightly raised on both surfaces, tertiary venation densely reticulate, raised and evident above and below; petiole 12–22 mm long, 1–1.5 mm in diam., glabrous. Male flowers borne in axillary, 3–5-flowered cymes, 1 inflorescence per axil, peduncle 10–20 mm long, 0.5–1 mm in diam., sparsely covered with appressed, white trichomes c. 0.2 mm long, pedicel 5–13 mm long, 0.5–0.8 mm in diam., rather sparsely covered with appressed, white trichomes c. 0.2 mm long; calyx obconic, 4–5 × 5–6 mm, 4-lobed, the lobes broadly triangular, 2.5 × 3.5 mm, apex shortly acuminate, rather sparsely covered outside with appressed, white trichomes c. 0.2 mm long, glabrous inside; corolla tubular, 10.5 × 6 mm, 4-lobed, the lobes valvate, 3.5 × 3 mm, triangular, rather sparsely covered with appressed, white trichomes c. 0.2 mm long on both surfaces; stamens c. 30, sparsely covered throughout with appressed, white trichomes c. 0.2 mm long, of variable lengths, inserted on the corolla at the midpoint, lower series with slender, terete, sigmoid filaments c. 1.5 mm long, upper series with flattened filaments 3.5 mm long, anthers 2.5 mm long, lanceolate, apex acuminate, dehiscing by apical pores; pistillode absent. Female flowers not seen. Fruits axillary, solitary, pedicel in fruit 4–5 mm long, 4–5 mm in diam., sparsely covered with appressed, white trichomes c. 0.2 mm long; fruiting calyx broadly cupuliform, 7 × 18– 22 mm, with a distinct basal collar c. 2 mm long, 6 – 7 mm in diam., (3–)4-lobed, sometimes very shallowly so with nearly indistinct lobes, the lobes broadly triangular, 2–9 × 10–18 mm, margin flat or revolute, sparsely covered with appressed, white trichomes c. 0.2 mm long, glabrescent; fruit depressed spherical, 16–28 × 17–32 mm, crowned with a very short style/stigma remnant, sparsely covered with appressed, white trichomes c. 0.2 mm long. Seeds 4, spherical wedgeshaped, 23 × 17 mm, glabrous.

Etymology. – The specific epithet refers to the fact that the vegetative features of this species superficially resemble those of some species of the genus Mimusops L. ( Sapotaceae ).

Distribution and ecology. – Diospyros mimusops is known from the Bemangidy forest near Iaboakoho on the eastern slope of the Vohimena mountains and from near the Col de Maningotry in Parcel 1 of Andohahela National Park ( MADAGASCAR CATALOGUE, 2021). It occurs in low- and midelevation humid forest.

Phenology. – Flowering material has been collected in December, and collections with fruit have been made in April, May, and November.

Conservation status. – Diospyros mimusops has a very restricted geographic range in the form of an Extent of Occurrence of 12 km ² and an Area of Occupancy of 12 km ². Its distribution is wholly contained within the protected areas of Andohahela and Tsitongambarika. Nevertheless, it occurs near the edge of the forest at Iaboakoho and is thus threatened by ongoing exploitation of trees for firewood and house construction material, which will result in continuing decline of quality of habitat and mature individuals. With respect to the most serious plausible threat of exploitation of trees for firewood and house construction material, D. mimusops exists at two locations, and was recently assessed for its risk of extinction as “Endangered” [EN B1ab(iii,v)+2ab(iii,v)] ( IUCN, 2021).

Notes. – Diospyros mimusops is a member of the Tetraclis group (see notes under D. ambanjensis for a list of its diagnostic features), and along with D. beberonnii (see above), is the only member found in far southern Madagascar. Within the group, D. mimusops can be distinguished by its glabrous and sometimes glaucous young stems and leaves.

Additional specimens examined. – MADAGASCAR. Reg. Anosy [Prov. Toliara]: road towards Ranomafana Sud , 5.XII.1989, ♂ fl., McPherson 14621 ( G, MO, P, TAN); Antsotso, 2.IV.2008, fr., Rabenantoandro et al. 1911

[A, C, E: Antilahimena et al. 4887, P; B, D: McPherson 14621, P] [Drawing: Alain Jouy]

(MO, P, TAN); ibid. loco, 22. V.2006, fr., Randriatafika et al. 676 ( MO, P, TAN) ; 8.XII.2007, ♂ fl., Razakamalala et al. 3771 ( MO, P, TAN) ; ibid. loco, 1.IV.2014, ster., Razakamalala 7737 ( MO, P, TAN) ; ibid. loco, 11.II. 2019, ster., Razakamalala et al. 8285, 8286, 8289 ( DBEV, G, MO, P, TAN) ; Antanitsara, 5.XI.2019, y.fr., Razakamalala & Andrianarivelo 8563 ( DBEV, MO, P, TAN) .

Diospyros quadrangularis G.E. Schatz & Lowry , sp. nov. ( Fig. 16 ).

Holotypus: MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Ambinanitelo, 9.IX.2004, fr., Antilahimena 2876 (MO-6336086!; iso-: P [ P03975182 ]!, TAN!) .

Diospyros quadrangularis G.E. Schatz & Lowry resembles other members of the Subsessilifolia group in having flattened young stems and leaves with a short petiole and a rounded to cordate base, but is distinguished by having young stems that are 4-angled and square in cross section, and relatively large fruits (40 –60 × 40 –53 mm).

Tree 4–12 m tall, 10–20 cm DBH. Young stems distinctly 4-angled, square in cross-section, glabrous to rarely very sparsely covered with minute (<0.1 mm long), appressed trichomes, initially reddish brown becoming white with age. Leaves distichous, lamina 3–13.7 × 1.4–7.8 cm, narrowly obovate to broadly elliptic, chartaceous to subcoriaceous, glabrous on both surfaces, base rounded to subcordate to cordate, rarely obtuse, margin flat, apex acute to rounded, midvein slightly impressed above, prominently raised below, distinctly narrowing from base to apex, venation brochidodromous, with 7–10 secondary veins per side, flat to slightly raised above and below, tertiary venation indistinct; petiole 1–5 mm long, 1–2 mm in diam., glabrous. Male and female flowers not seen. Fruits axillary, solitary, pedicel in fruit 1–7 mm long, 5–9 mm in diam., densely covered with semi-appressed, golden brown trichomes 0.5 mm long, pedicel scar 6–10 mm in diam.; calyx in fruit cupuliform to broadly cupuliform, 10–12 × 24–32 mm, 4-lobed, the lobes broadly triangular, 4–10 × 16–20 mm, densely covered with semi-appressed, golden brown trichomes 0.5 mm long on both surfaces, margins flat; fruit spherical to slightly ellipsoid, 40–60 × 40–53 mm, glabrous except very sparsely covered with appressed golden trichomes c. 0.5 mm long at the base where the fruit is covered by the calyx. Seeds 6–8, spherical wedgeshaped, 30–33 × 14–15 mm, glabrous, reddish brown, slightly shiny.

Etymology. – This specific epithet refers to the fact that the young stems are 4-angled.

Vernacular names. – “Fandramanan’ala” (Razanatsima 127),

“Hazomainty” (Service Forestier 15101).

Distribution and ecology. – Diospyros quadrangularis occurs in northeastern Madagascar, from Tampolo north to Ambondrobe, with outlier subpopulations farther south at Ambalabe in the Corridor Ankeniheny-Zahamena protected area, and farther north at Andrafiamena ( MADAGASCAR CATALOGUE, 2021). It is found in low- and mid-elevation humid forest.

Phenology. – Fruiting material has been collected in March, and from May to November.

Conservation status. – Diospyros quadrangularis has a geographic range in the form of an Extent of Occurrence of 54,568 km ² and a minimum Area of Occupancy of 60 km ². It is present in seven protected areas, i.e., Andrafiamena Andavokoera, Anjanaharibe-Sud, Corridor AnkenihenyZahamena (Ambalabe), COMATSA Sud, Makira, Makirovana-Tsihomanaomby, and Masoala. Outside of protected areas, it is threatened by forest clearing for agriculture, fire, and exploitation for firewood and house construction material, all of which will result in continuing decline. With respect to the most serious plausible threat of forest clearing for agriculture, it exists at 12 locations. Therefore, D. quadrangularis can be assessed for its risk of extinction as “Near Threatened” [NT], as it is close to qualifying for VU status under criterion B2.

Notes. – Diospyros quadrangularis is a member of the Subsessilifolia group, characterized by flattened young stems and leaves with a short petiole and a rounded to cordate base. Other members of the group include D. mangabensis Aug. DC. , D. mangorensis H. Perrier , D. sphaerosepala , and D. subsessilifolia H. Perrier , as well as more than a dozen new species awaiting description. Within the Subsessilifolia group, D. quadrangularis can be distinguished by its 4-angled young stems that are square in cross section, and relatively large fruits (40–60 × 40–53 mm).

Additional specimens examined. – MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Anjahana , 19.VII.2002 , y.fr., Antilahimena et al. 1214 ( MO, P, TAN); Tampolo , 18.VIII.1955 , fr., Service Forestier 15101 ( MO, P, TEF); ibid. loco, 23. I.2014, ster., Razakamalala & Bernard 7710 ( BR, G, MO, P, TAN); Île Ste. Marie , 18.XII.2017 , y.fr., Razakamalala 8186 ( MO, P, TAN); ibid. loco et datum, Razakamalala 8188 ( MO, P, TAN); Ambohitsitondroina de Mahalevona , 29.XI.1953 , fr., Service Forestier 8690 ( MO, P, TEF); Tampolo , 28–29.VIII.1957 , fr., Service Forestier 18154 ( MO, P, TEF); Île Sainte-Marie , 16. V.1969, fr., Service Forestier 28806 ( MO, P, TEF); ibid. loco, 16. V.1969, y.fr., Service Forestier 28819 ( G, MO, P, TEF). Reg. Atsinanana [Prov. Toamasina]: Ambalabe , 12. VI.2006, fr., Razanatsima 127 ( G, MO, P, TAN); ibid. loco, 5. V.2010, y.fr., Razanatsima 885 ( MO, P, TAN); ibid. loco, 12.VII.2017 , y.fr., Razanatsima et al. 1753 ( MO, P, TAN). Reg. SAVA [Prov. Antsiranana]: Anjanaharibe-Sud Reserve , 7.IX.1997 , fr., Birkinshaw et al. 486 ( MO, P, TAN); Antsirabe-Nord, Andavanambo, Tsihomanaomby , access from Antanambaonisokitra , 23.IV.2014 , y.fr., Birkinshaw et al. 2013 ( MO, P, TAN); Anjahankely , 18.XI.2010 , fr., Burivalova 33 ( G, MO, P, TAN); massif d’Antsahabe , 14.X.2004 , fr., Callmander et al. 220 ( G, MO, P, TAN); Amdisatrana , 2.VIII.1997 , fr., McPherson 17178 ( MO, P, TAN); Antranohofa , 9. V.1998, y.fr., Ravelonarivo 1094 ( MO, P, TAN); Manakana , 13.III.2004 ,

Candollea 76, 2021 New species of Diospyros (Ebenaceae) from Madagascar – 231

[A: Birkinshaw et al. 486, P; B–D: Antilahimena 2876, P] [Drawing: Alain Jouy]

fr., Razakamalala et al. 1016 (G, MO, P, TAN); ibid. loco, 20.V.2004, fr., Razakamalala 1346 (G, MO, P, TAN).

Diospyros rakotovaoi G.E. Schatz & Lowr y, sp. nov. ( Fig. 13C–E , 17 ).

H o l o t y p u s: M A D A G A S C A R. R e g. S AVA [Pr o v. Antsiranana]: Antsahaberaoka, 21.II.2007, fr., Rakotovao et al. 3692 (MO-6214314!; iso-: G!, P [ P04539975 ]!, TAN [ TAN002010 ]!) .

Diospyros rakotovaoi G.E. Schatz & Lowry resembles other members of the Tetraclis group in having fauve to rusty indumentum, leaves generally with a mucronate apex, male flowers borne in cymose inflorescences, and often valvate corolla aestivation, but is distinguished by its obovate leaves that are glabrous and shiny above and densely covered with semi-erect light brown trichomes along the midvein and secondary veins below, rendering it somewhat scabrous, and its relatively large spherical fruit 29–37 mm in diam., densely covered with erect, rusty brown trichomes 0.3 mm long.

Tree 6–25 m tall, 6–15 cm DBH. Young stems terete, densely covered with slightly curly, erect, light brown trichomes 0.5 mm long, distinctly lenticellate. Leaves distichous, lamina 6.5–11.5 × 2.3–3.7 cm, obovate, chartaceous to subcoriaceous, glabrous, shiny above, densely covered with semi-erect, whitish trichomes c. 1 mm long along the midvein and secondary veins, and sparsely covered with semi-erect light brown trichomes c. 0.5 mm long elsewhere below, somewhat scabrous, glabrescent, base attenuate to cuneate, margin flat, apex rounded, rarely somewhat retuse, with a distinct mucron c. 1 mm long, midvein impressed above, prominently raised below, venation brochidodromous, with 7–9 secondary veins per side, flat above, raised below, tertiary venation reticulate, flat above, slightly raised below; petiole 5–9 mm long, 1 mm in diam., densely covered with slightly curly, erect, light brown trichomes 0.5 mm long, glabrescent. Male flowers in cymose, axillary inflorescences, each with 7 flowers; inflorescence 15–20 mm long, the axes densely covered with erect, rusty brown trichomes 0.3 mm long, pedicel 3–13 mm long, 1.5 mm in diam.; calyx cupuliform, 5 × 8 mm, 1 mm thick, densely covered outside with erect, rusty brown trichomes 0.3 mm long, glabrous inside except for a ring of very short trichomes at the base, 4-lobed, 4 × 5 mm, broadly triangular; corolla cylindrical, 8 × 7 mm, 1 mm thick, densely covered with appressed, light brown trichomes 0.2 mm long on both surfaces, 4-lobed, the lobes valvate, 4.5 × 4.5 mm, triangular; stamens 8, inserted in a single whorl on the corolla c. 1/3 from the base, filaments 3 mm long, with a small hook-like basal appendage, densely covered with appressed, light brown trichomes 0.5–0.7 mm long, anthers 3 mm long, lanceolate, apex apiculate, dehiscing by apical pores; torus with a ring 3 mm in diam. of dense, erect trichomes 1 mm long; pistillode absent. Female flowers not seen. Fruits axillary, solitary, pedicel in fruit 4 mm long, 4 mm in diam., densely covered with erect, rusty brown trichomes 0.3 mm long; fruiting calyx broadly cupuliform, 10–12 × 25–27 mm, 4-lobed, the lobes broadly triangular, 10–12 × 17–19 mm, margin flat to slightly revolute, densely covered with erect, rusty brown trichomes 0.3 mm long; fruit spherical, 29–37 mm in diam., densely covered with erect, rusty brown trichomes 0.3 mm long. Seeds 1–4, ovoid, 17 mm long, 10 mm in diam., black, matte.

Etymology. – This species honors our friend and colleague Charles Rakotovao, one of Madagascar’s most prolific field botanists, who has made more than 8500 collections starting in 1994, including 200 numbers of Diospyros , contributing immensely to our knowledge of the Malagasy flora. Since joining the staff of the Missouri Botanical Garden in 2005, Charles has conducted field work throughout the island and has taken more than 4100 excellent photos ( MADAGASCAR CATALOGUE, 2021), documenting both his own collections and those of many colleagues.

Distribution and ecology. – Diospyros rakotovaoi occurs in the mountains of northern Madagascar, from the Anjanaharibe-Sud reserve north to Antsahabe in the Daraina region, and extending into the Sambirano region in the northwestern part of the island at the Galoko-Kalobinono and Tsaratanana protected areas ( MADAGASCAR CATALOGUE, 2021), where it is found in mid- to high-elevation humid to subhumid forest.

Phenology. – Flowering material has been collected in November, and specimens with fruit have been made in February, May, August, and November.

Conservation status. – Diospyros rakotovaoi has a restricted geographic range in the form of an Extent of Occurrence of 12,793 km ² and a minimum Area of Occupancy of 28 km ². Its distribution is wholly contained within six protected areas, i.e., Anjanaharibe-Sud, COMATSA Nord, GalokoKalobinono, Loky Manambato, Makirovana, and Tsaratanana. Nevertheless, at several localities it has been collected at the forest edge where there are active threats consisting of forest clearing for agriculture, fire, and exploitation of trees for firewood and house construction material, which will result in continuing decline of quality of habitat and mature individuals. With respect to the most serious plausible threat of exploitation of trees for firewood and house construction material, D. rakotovaoi exists at seven locations, and was recently assessed for its risk of extinction as “Vulnerable” [VU B1ab(iii,v)+2ab(iii,v)] ( IUCN, 2021).

Notes. – Diospyros rakotovaoi is another member of the

Tetraclis group (see notes under D. ambanjensis for a list of its

[A, B, D: Raharimampionona et al. 303, P; C, E, F: Razafitsalama 1141, P] [Drawing: Alain Jouy]

diagnostic characters), within which it is characterized by its leaves with an obovate lamina with a glabrous, shiny adaxial surface and the abaxial surface densely covered with semierect, light brown trichomes along the midvein and secondary veins, rendering it somewhat scabrous, as well as its relatively large spherical fruits (29– 37 mm in diam.) that are densely covered with erect, rusty brown trichomes 0.3 mm long.

Additional specimens examined. – MADAGASCAR. Reg. Diana [Prov. Antsiranana]: Réserve Intégrale de Tsaratanana , 31.VIII.2000, fr., Antilahimena 596 ( MO, P, TAN) ; Kalebenono , 25.XI.2006, ♂ fl., Razafitsalama & Torze 1141 ( MO, P, TAN) . Reg. SAVA [Prov. Antsiranana]: forêt d’Antsahabe , 26.XI.2004, ♂ fl., Gautier 4755 ( G, MO, P, TAN) ; Belalona , 25. V.2009, fr., Raharimampionona et al. 303 ( MO, P, TAN, TEF) ; Réserve Spéciale d’Anjanaharibe-Sud , 9.XI.1999, fr., Rakotomalaza & Ravelomanantsoa 2145 ( G) ; ibid. loco, 3.XI.1994, fr., Ravelonarivo & Rabesonina 508 ( L, MO, P, TAN, WAG) ; Tsaratanana , 6.II.2006, fr., Razakamalala et al. 3190 ( G, MO, P, TAN) .

Figures

Fig.12.– Diospyros malandy H.N. Rakouth, R. Randrianaivo, G.E. Schatz & Lowry. A. Branch in fruit; B. Detail of leaf (adaxial surface); C. Fruit; D. Fruiting calyx (seen from below).

Fig. 13. – Photographs of Diospyros L. species. A. Diospyros malandy H.N. Rakouth, R. Randrianaivo, G.E. Schatz & Lowry,

Fig. 14. – Diospyros melanocarpa G.E. Schatz & Lowry. A. Branch in fruit; B. Fruit; C. Fruit (seen from above); D. Detail of leaf (adaxial surface). [A–D: Service Forestier 18343, P] [Drawing: Alain Jouy]

Fig. 15. – Diospyros mimusops G.E. Schatz & Lowry. A. Branch with fruit; B. Branch with male inflorescences; C. Detail of leaf (abaxial surface); D. Schematic section of male flower; E. Fruit.

Fig. 16. – Diospyros quadrangularis G.E. Schatz & Lowry. A. Branch; B. Fruiting branch with stem cross-sections; C. Detail of leaf (adaxial surface); D. Fruits.

Fig. 17.– Diospyros rakotovaoi G.E. Schatz & Lowry. A. Branch with fruits; B. Detail of leaf (abaxial surface); C. Branch with male inflorescences; D. Fruiting calyx (seen from below); E. Male flower; F. Schematic section of male flower.