Diospyros beberonnii 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 : 214-220

Diospyros beberonnii G.E. Schatz & Lowry , sp. nov. ( Fig. 7 ).

Holotypus: MADAGASCAR. Reg. Anosy [Prov. Toliara]: Parcelle 1 de la RN d’Andohahela, 10 – 27.IX.1993, fr., Randriamampionona 640 (MO-6956127!; iso-: BR!, CAS!, K!, F!, G [ G00341740 ]!, MO-6956128!; NY!, OXF!, P [ P03829510 ]!, PRE!, S!, TAN!, US!, W!, WAG!) .

Diospyros beberonnii G.E. Schatz & Lowry resembles other members of the Tetraclis group in having fauve to rusty indumentum and leaves generally with a mucronate apex, but is distinguished by having white trichomes on its young stems and leaves, a small leaf lamina [(3–)4–8 × 1–2.8 cm], a small fruiting calyx (5 mm long with lobes 6–7 × 7– 8 mm), and small fruits (13–17 mm long).

Tree 14–20 m tall, 20 –60 cm DBH. Young stems terete, densely covered with semi-appressed, whitish trichomes 0.5 mm long. Leaves distichous, lamina (3–)4–8 × 1–2.8 cm, narrowly ovate to elliptic to rarely obovate, chartaceous, initially sparsely covered with appressed, whitish trichomes c. 0.8 mm long above, more densely so along the midvein and margin, glabrescent, sparsely covered with appressed, whitish trichomes c. 0.5– 0.8 mm long below, densely so along the midvein, base cuneate to attenuate, margin slightly thickened below, apex acute to acuminate, with a distinct mucron 1–1.5 mm long, midvein impressed above, prominently raised below, venation brochidodromous, with 10–12 secondary veins per side, tertiary venation reticulate, raised above and below, very evident above; petiole 4–8 mm long, 1 mm in diam., densely covered with semi-appressed, whitish trichomes 0.5 mm long, glabrescent. Male flowers not seen. Females flowers not seen. Fruits axillary, solitary, pedicel in fruit 5–7 mm long, 4–5 mm in diam., very densely covered with erect, orangebrown trichomes 0.5–0.7 mm long completely obscuring the surface; fruiting calyx 5 × 12–13 mm, 4-lobed, the lobes ovatetriangular, 6–7 × 7–8 mm, very densely covered with erect, orange-brown trichomes 0.5– 0.7 mm long, spreading, not appressed to the fruit wall, margin flat, pedicel scar 4 mm in diam.; fruit spherical to broadly ellipsoid, 13–17 × 15–17 mm, initially densely covered with appressed, orange-brown trichomes to 1.1 mm long, glabrescent except at the apex and at the base where protected by the calyx, crowned by a very short style/stigma remnant <1 mm long, surface verrucose in sicco. Seeds 1–2?, 11 × 6 mm, flattened ovoid, black, somewhat shiny.

Etymology. – This species honors Beberonn Randriamampionona, who began conducting botanical field work in 1989 and subsequently worked as a field botanist for Missouri Botanical Garden between 1992 and 1996, making nearly 1350 excellent collections, primarily in Andohahela National Park.

Distribution and ecology. – Diospyros beberonnii is known only from Parcel 1 of Andohahela National Park in southeastern Madagascar, with a single collection made just outside the southwestern limit of the parcel ( MADAGASCAR CATALOGUE, 2021). It occurs in low-elevation humid forest, occasionally in transition areas toward drier forest.

Phenology. – Fruiting material has been collected in March,

July, and September.

Conservation status. – Diospyros beberonnii has a very restricted geographic range in the form of an Extent of Occurrence of 264 km ² and an Area of Occupancy of 20 km ². Its distribution is almost entirely contained within the protected area of Andohahela. Nevertheless, it is present near the road that traverses Andohahela Parcel 1 and at a site just outside the southwestern limit of the parcel, 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 number of mature individuals. With respect to the most serious plausible threat of exploitation of trees for firewood and house construction material, D. beberonnii exists at five locations, and was recently assessed for its risk of extinction as “Endangered” [EN B1ab(iii,v)+2ab(iii,v)] ( IUCN, 2021).

Notes. – Diospyros beberonnii is one of two species in the Tetraclis group (see notes under D. ambanjensis for a list of its diagnostic features) restricted to southeastern Madagascar, along with D. mimusops (described below). Within the group, it can be distinguished by the white trichomes on its young stems and leaves, and the overall small dimensions of its lamina [(3–)4–8 × 1–2.8 cm], fruiting calyx (5 mm long with lobes 6–7 × 7–8 mm), and fruit (13–17 mm long).

Additional specimens examined. – MADAGASCAR. Reg. Anosy [Prov. Toliara]: Andohahela, Parcelle 1, 16–24.III.1993, fr., Randriamampionona 234 ( MO, P, TAN); ibid. loco, 8.VII.1994, fr., Randriamampionona 829 ( MO, P, TAN); ibid. loco, 9.IX.2019, ster., Razakamalala & S . Andrianarivelo 8578 ( DBEV, MO, P, TAN); ibid. loco, ster., Razakamalala & S . Andrianarivelo 8582 ( DBEV, MO, P, TAN) .

Diospyros crassipedicellata G.E. Schatz & Lowry , sp. nov. ( Fig. 8 , 9A–C ).

Holotypus: MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Farankaraina, 16.IX.1957, fr., Service Forestier 18318 (MO-6956005!; iso-: G [ G00341737 ]!, P [ P00722707 , P03829422 ]!, TEF) .

Diospyros crassipedicellata 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 having a distinctively thickened pedicel in fruit (5–9 mm in diam.) and narrow leaves (3.5–11.5 × 1.1–5 cm).

Tree 6 – 20 m tall, 11– 30 cm DBH. Young stems terete, densely covered with straight, appressed or wavy semiappressed, whitish trichomes 0.5 mm long. Leaves distichous, lamina 3.5–11.5 × 1.1– 5 cm, elliptic to narrowly obovate to obovate, subcoriaceous to coriaceous, initially sparsely to densely covered with appressed trichomes 0.5 mm long above, more densely so along the midvein and margin, glabrescent, sparsely to densely covered with appressed trichomes to 0.8 mm long below, densely so along the midvein, base cuneate to attenuate, margin slightly thickened below, apex obtuse to acute to acuminate, occasionally rounded, usually with a distinct mucron 1–1.5 mm long, midvein slightly impressed above, prominently raised below, venation brochidodromous, with 9–12 secondary veins per side, tertiary venation reticulate, flat above and below; petiole 3–8 mm long, 1 mm in diam., densely covered with semi-appressed, whitish trichomes 0.5 mm long, glabrescent. Male flowers in cymose, axillary inflorescences, 1–3 inflorescences per axil, 2–7 flowers per inflorescence, or occasionally flowers solitary; inflorescence 5–17 mm long, the axes densely covered with erect, fauve trichomes c. 0.8–1 mm long, pedicel 3– 9 mm long, 0.8– 2 mm in diam., densely covered with erect, fauve trichomes c. 0.8–1 mm long; calyx cupuliform, 3.5–4 × 6–7 mm, 4-lobed, the lobes triangular, 2– 4 × 3 – 4 mm, densely covered outside with erect, fauve trichomes c. 0.8–1 mm long, glabrous inside; corolla cupuliform, 8 mm long, 5 mm in diam., 4-lobed, the lobes valvate, 4 × 4 mm, ovate-triangular, rather densely covered outside with appressed trichomes to 0.5–0.8 mm long, rather densely covered inside with appressed trichomes c. 0.8–1 mm long

2 mm

[A, E: Service Forestier 18318, P; B–C: Service Forestier 12941, P] [Drawing: Alain Jouy]

from the point of insertion of the stamens to the apex, glabrous below the point of insertion of stamens, stamens 20, inserted on the corolla 1/3 from base, filaments 0.1 mm long, bearing a tuft of curved trichomes 1 mm long, anthers 1.8 mm long, apex apiculate, dehiscing by apical pores; pistillode absent, torus densely covered with erect trichomes. Females flowers not seen. Fruits axillary, solitary, pedicel in fruit 8–13 mm long, 5– 9 mm in diam., very densely covered with erect, orangebrown trichomes 0.5–0.7 mm long completely obscuring the surface; fruiting calyx broadly cupuliform, 8– 12 × 20 mm, 3 –4-lobed, the lobes triangular to ovate to broadly ovate, 7–13 × 12–15 mm, densely covered with semi-erect, wavy, light brown trichomes c. 0.4– 0.8 mm long, obscuring the calyx surface, erect and adhering to the fruit, margin flat, pedicel scar 5–6 mm in diam.; fruit ellipsoid, 25– 38 × 20– 25 mm, densely covered with semi-erect, wavy, light brown trichomes c. 0.4–0.8 mm long, obscuring the surface of the fruit. Seeds 4, 13 × 5 mm, flattened ovoid, black, shiny.

Etymology. – The specific epithet refers to the distinctively thickened pedicel in fruit.

Vernacular name. – “Hazomainty” (Service Forestier 10738).

Distribution and ecology. – Diospyros crassipedicellata occurs in eastern Madagascar, from Manombo reserve north to the area around the Baie d’Antongil, mostly in low-elevation humid forest near the coast, but also farther inland in midelevation humid forest at Ambalabe and Ankerana, both of which are situated within the Corridor Ankeniheny-Zahamena protected area ( MADAGASCAR CATALOGUE, 2021).

Phenology. – Flowering collections have been made from January to March, and fruiting material has been collected in May, June, and from August to November.

Conservation status. – Diospyros crassipedicellata has a geographic range in the form of an Extent of Occurrence of 32,769 km ² and a minimum Area of Occupancy of 36 km ². It is present in four protected areas, i.e., Analalava, Corridor Ankeniheny-Zahamena (Ambalabe and Ankerana), Masoala, and Manombo. Outside of the protected areas, it is threatened by forest clearing for agriculture, fire, grazing, 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, D. crassipedicellata exists at seven locations, and was recently assessed for its risk of extinction as “Vulnerable” [VU B2ab(i,ii,iii,iv,v)] ( IUCN, 2021).

Notes. – Diospyros crassipedicellata belongs to the Tetraclis group (see notes under D. ambanjensis for a list of its diagnostic features), within which it can be distinguished by its distinctively thickened pedicel in fruit, which is 5–9 mm in diameter.

Additional specimens examined. – MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Ambanizana , 20.IX.2002, fr., Antilahimena 1421 ( MO, P, TAN) ; ibid. loco, 9.XI.1994, fr., Rabe 195 ( MO, P, TAN) ; ibid. loco, 27. VI.1994, fr., Randriamarosoa et al. 172 ( MO, P, TAN) ; Farankaraina , 27.VIII.1954, fr., Service Forestier 10738 ( MO, P, TEF) ; ibid. loco, 16.IX.1957, fr., Service Forestier 18318 ( G, MO, P, TEF) ; Île Sainte-Marie , 18. V.1969, fr., Service Forestier 28842 ( MO, P, TEF) . Reg. Atsinanana [Prov. Toamasina]: Mahavelona, Foulpointe , 7.X.2011, fr., Andriamiarinoro & Amosa 247 ( MO, P, TAN) ; Maroseranana , 24.III.2011, ♂ fl., Antilahimena 7783 ( MO, P, TAN) ; Analalava Reserve , 13 Sep 2017, fr., Grevais 120 ( MO, P, TAN) ; ibid. loco, 15. I.2017, bud, Lowry et al. 7520 ( MO, P, TAN) ; ibid. loco, 18. VI.2012, y.fr., Miandrimanana et al. 556 ( MO, P, TAN) ; Ambalabe, Sahanionaka , 7.II.2011, ♂ fl., A. Randrianasolo et al. 1385 ( MO, P, TAN) ; Ankerana , 900 m, 14.III.2011, fl., Ravelonarivo & Edmond 3666 ( MO, P, TAN) . Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: Manombo , 27.VII.1955, ster., Réserves Naturelles 13956 ( P) ; ibid. loco, 10. I.1955, ♂ fl., Service Forestier 12940 ( G, MO, P, TEF) .

Diospyros grandiflora G.E. Schatz & Lowry , sp. nov. ( Fig. 9D–F , 10 ).

Holotypus: MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: Betampona Reserve, 11.I.2017, fr., Lowry et al. 7482 (MO-6956129!; iso-: G [ G00341741 ]!, K!, P [ P00722717 ]!, TAN!, W!) .

Diospyros grandiflora G.E. Schatz & Lowry most closely resembles D. toxicaria Hiern in its glabrous, coriaceous leaves with reticulate tertiary venation raised both above and below, but differs by its smaller leaf blades (6 –12.5 × 3–5.4 cm vs. 8–19 × 4 – 6.5 cm), often shorter petioles (10 –13 mm vs. 6 –18 mm), and larger female flowers with a calyx 10 –15 × 16 –18 mm (vs. 7–9 × 4–5 mm) and corolla c. 15 mm long (vs. 4–5 mm).

Tree 20 – 24 m tall, 20 cm DBH. Young stems terete, sparsely covered with appressed white to light brown t r i c h om e s 0.5 m m l on g. Lea ves d i s t i c h o u s, l a m i n a 6–12.5 × 3–5.4 cm, elliptic to oblong to slightly ovate, coriaceous, glabrous above and below, shiny above, base obtuse to rounded, margin flat, apex rounded to acute, recurved, midvein impressed above, prominently raised below, venation weakly brochidodromous, with 6 –10 secondary veins per side, slightly raised above and below, tertiary venation reticulate, slightly raised above and below; petiole 10–13 mm long, 1.5–2 mm in diam., canaliculate, glabrous. Male flowers solitary, axillary, sessile, surrounded by c. 4 distichous, concave bracts, broadly ovate to circular to oblate, 2–7 × 5–10 mm, increasing in size from the basal bract to the apical bract, the apex deeply retuse, densely covered outside with appressed, fauve trichomes c. 0.3 mm long, glabrous inside; calyx in bud densely covered outside with appressed, fauve trichomes c. 0.5 mm long, glabrous inside; corolla in bud 4–5-lobed, densely covered outside with appressed, fauve trichomes c. 1 mm long from the lobes downward nearly to the base, glabrous elsewhere outside, glabrous inside; stamens 20, inserted on the torus, filaments 1 mm long, anthers 2 mm long, lanceolate, the apex acuminate; pistillode present, densely covered with short, erect trichomes. Female flowers solitary, axillary, surrounded by c. 8 oblate bracts, 13– 15 × 17– 21 mm, the apex deeply retuse, densely covered outside with appressed, fauve trichomes c. 0.5 mm long, glabrous inside, pedicel 2.5–3 mm long, 5 mm in diam.; calyx cupuliform to urceolate, 10– 15 × 16– 18 mm, 4-lobed, the lobes ovate-triangular, 6–7 × 8–10 mm, densely covered outside with erect, rusty brown trichomes 1 mm long; corolla c. 15 mm long, densely covered outside with appressed, rusty brown trichomes 3 mm long on the back of the lobes downward, mauve in vivo. Fruits axillary, solitary, pedicel in fruit 4–5 mm long, c. 8 mm in diam., with persistent, short, erect, dark ferruginous brown trichomes, with evident bract scars; fruiting calyx broadly hemispherical, c. 15 × 30 mm, 5-lobed, the lobes broadly triangular, c. 8 × 10 mm, apex reflexed, densely covered with persistent, erect, dark ferruginous brown trichomes c. 1 mm long on both surfaces; fruit sub-spherical, slightly square in transverse section, c. 25–30 × 25–28 mm, densely covered with persistent, erect, dark ferruginous brown trichomes, the apex depressed, with a persistent stylar remnant c. 3 mm long.

Etymology. – The specific epithet refers to the large size of the flowers.

Vernacular names. – “Hazomafana” (Réserves Naturelles 2642),

“Hazomainty” (Réserves Naturelles 9731).

Distribution and ecology. – Diospyros grandiflora occurs in the Betampona reserve and a small remnant forest patch c. 11 km to the west-southwest ( MADAGASCAR CATALOGUE, 2021), where it grows in mid-elevation humid forest.

Phenology. – Flowering material has been collected in July and August, and fruits are known from January.

Conservation status. – Diospyros grandiflora has an extremely restricted geographic range in the form of an Extent of Occurrence of <5 km ² and an Area of Occupancy of <5 km ². It is present in the protected area of Betampona. Outside of the protected area, it is threatened by forest clearing for agriculture and exploitation of trees for firewood and house construction material, which will result in continuing decline. With respect to the most serious plausible threat of forest clearing for agriculture, D. grandiflora exists at two locations, and was recently assessed for its risk of extinction as “Endangered” [EN B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)] ( IUCN, 2021).

Notes. – Diospyros grandiflora resembles D. toxicaria , a widespread species occurring in low- to mid-elevation humid forest in eastern Madagascar ( MADAGASCAR CATALOGUE, 2021), but has smaller leaves, with the lamina 6– 12.5 × 3–5.4 cm (vs. 8–19 × 4–6.5 cm in D. toxicaria ) and petioles 10–13 mm long (vs. 6–18 mm long), as well as considerably larger female flowers whose calyx is 10–15 × 16–18 mm (vs. 7–9 × 4–5 mm) and corolla c. 15 mm long (vs. 4 –5 mm long). The female flowers of D. grandiflora are the largest of any known member of the genus on Madagascar.

Additional specimens examined. – MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: RN 1 [= Betampona], 9.VIII.1950, ♂ fl., Réserves Naturelles 2642 ( TAN, P); ibid. loco, 10.VII.1958, ♀ fl., Réserves Naturelles 9731 ( P, TEF); ibid. loco, 8.VII.1958, ♀ fl., Lewis 621 ( MO, P); Betampona, 11. I .2017, ster., Lowry et al. 7481 A ( TAN, MO); Sahambala , 24.VII.2017, bud, G . Rakotonirina et al. 195 ( K, MO, P, TAN) .

Figures

Fig. 7. – Diospyros beberonnii G.E. Schatz & Lowry. A. Branch with fruits; B. Fruit; C. Fruiting calyx (seen from below). [A: Randriamampionona 234, P; B–C: Randriamampionona 640, P] [Drawing:Alain Jouy]

Fig. 8. – Diospyros crassipedicellata G.E. Schatz & Lowry. A. Branch with fruit; B. Branch with male inflorescences; C. Schematic section of male flower; D. Male inflorescence; E. Fruit.

Fig. 9. – Photographs of Diospyros L. species. A–C. Diospyros crassipedicellata G.E. Schatz & Lowry: A. Branch with male flowers; B. Male inflorescence; C. Male flowers; D–F. Diospyros grandiflora G.E. Schatz & Lowry: D. Branch with fruit; E. Fruit (side view); F. Fruit (top view). [A: Lowry et al. 7520; B: Antilahimena 7783; C: Randrianasolo et al. 1385; D–F: Lowry et al. 7482] [Photos: A, D–F: P. Lowry; B: P. Antilahimena; C: F. Rakotoarivony]

Fig. 10. – Diospyros grandiflora G.E. Schatz & Lowry. A. Branch in fruit; B. Branch with female flowers; C. Schematic section of female flower; D. Corolla of female flower; E. Fruit.