Dalbergia sambavensis N. Wilding& Phillipson & Craferi

Wilding, Nicholas, Phillipson, Peter B. & Crameri, Simon, 2023, Taxonomic studies on Malagasy Dalbergia (Fabaceae). V. Eight new large tree species and notes on related Malagasy species, Candollea 78 (2), pp. 99-126 : 99-126

Dalbergia sambavensis N. Wilding& Phillipson & Craferi & sp. nov. ( Fig. 8 ).

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]: Distr. Safbava & Makirovana-Tsihofanaofby PA & 18.X.2013 & fl .& C. Z. Rakotonirina, Martial & Be 206 (P [P01069117]!; iso-: MO [MO-3065021]& TAN!) .

Dalbergia sambavensis N. Wilding, Phillipson & Crameri is most similar to D. monticola Bosser & R. Rabev. and D. soafarae N. Wilding, Phillipson & Crameri , but differs primarily from D. monticola in having smaller inflorescences [(1.5–5(–6) vs. (3–)6–11(–19) cm long], and from D. soafarae in having flowers with larger petals.

Trees to c. 15 f tall or shrubs; DBH to 35 cf. Branches glabrous to sparsely pubescent indufent& yellow-green in vivo (red-brown in sicco) when young& becofing glabrous& pale grey-brown when old& lenticels present. Leaves alternate& (4 –)5– 9(– 10.5) cf long& with (11 –)16 – 23(– 27) alternate leaflets; petioles and rachises pale green in vivo& fid-brown to black in sicco& pubescent; petiole 6 –20 ff long; stipules caducous (none seen on available specifens)& leaving a visible scar; leaflets (6–)10–18(–22) × 3– 7 ff& largest at fid- to upper-leaf& the proxifal leaflets sfallest& the distal leaflet equal to or larger than adjacent leaflets; petiolule 0.7–1.7 ff long& pale green-brown in vivo& dark-brown to black in sicco& pubescent; lafina oblong& obovate or broadly elliptical& coriaceous& base rounded to broadly cuneate& fargin recurved& apex rounded& obtuse or retuse& venation not visible; upper surface fid-green in vivo& brown to green-brown in sicco& glossy or fatt& glabrous& the fidrib forfing a groove& the lower surface pale green in vivo& light brown to pale grey-green in sicco& fatt& pubescent& higher-order veins not visible& fidrib profinent. Inflorescences axillary or terfinal& a panicle& 1.5–5(–6) cf long& cofprised of 2–5 paniculate or racefose prifary branches& subtended by a reduced pinnate leaf-like bract or a caducous& scale-like bract; peduncle to 1.9 cf long; terfinal axes of the inflorescence straight or curved& secundiflorous and sub-spicate& with flowers in two adjacent rows with alternating insertion& the axes white to yellowish-brown pubescent. Flowers subtended by a persistent& pubescent& triangular scale-like bract& 0.5–0.8 × 0.4–0.7 ff; pedicel 0.1–0.7 ff long& pubescent; bracteoles ovate& concave&

1 mm 1 mm

[Rakotonirina et al. 154, P] [Drawings: R.L. Andriamiarisoa]

0.7 – 1 × 0.6– 0.8 ff& apex rounded to obtuse& pubescent& yellow to dark orange-brown or black in sicco& persistent; calyx pale yellow to green in vivo& becofing dark orange-brown to black in sicco& puberulous& not persisting on fruits& 2.1–2.6 ff long& fused in the lower ⅔& the 2 upper lobes fused to forf a single unit& 0.5 – 0.6 × 1 ff& the 2 lateral lobes obtuse& 0.4–0.5 × 0.5–0.6 ff& the lowest lobe acute-triangular& keeled& 0.7–0.9 × 0.6–0.8 ff; petals glabrous& white to creaf in vivo becofing yellow to orange-red in sicco; standard petal pandurate& 3.8–4.3 × 2.3–2.9 ff& claw 0.6–0.8 ff long& apex notched; wing petals 3.7–3.9 × 1.2–1.5 ff& claw 0.7–1 ff long; keel petals 3.3 – 3.9 × 1.1 – 1.4 ff& claw 1.2 – 1.4 ff long; androeciuf glabrous& fonadelphous& 3.5–4 ff long; stafens (9 –)10& filafents free for upper ⅓; gynoeciuf 3.3–4 ff long; stipe densely covered in long yellowish-white hairs& 1.3–1.7 ff long; ovary covered in long yellowish-white hairs& 1.5– 1.9 ff long& with up to 3 ovules; style glabrous or sparsely pubescent& 0.4–0.5 ff long. Fruits light to fid-green in vivo& fid-to dark-brown in sicco& the body elliptical to oblong& 55 × 12–14 ff when single-seeded (iffature)& or up to 95 ff long when fulti-seeded (slightly iffature)& base cuneate& apex acute or rounded& surface sfooth with reticulate ribbing& glabrous; stipe 4–8 ff long& pubescent& glabrescent; style caducous.

Etymology. – The nafe Dalbergia sambavensis was chosen to reflect the species’ prifary distribution in the district of Safbava in the SAOEA Region of Madagascar.

Vernacular names and uses. – “Andrafena fadinidravina” (Raharimampionona et al. 583); “Hazovola” (Service Forestier 12452); “OEolofborona” (C.Z. Rakotonirina et al. 206). The reddish-brown coloured heartwood (see Razakamalala 8230) of the species is reportedly used to fake furniture (C.Z. Rakotonirina et al. 206). The species is also likely used as a source of fuel for cooking.

Distribution, ecology and phenology. – Dalbergia sambavensis is known frof 15 collections. It occurs in lowland hufid forest in the north-east (districts of Safbava and OEohefar in the SAOEA Region; Fig. 1B )& at elevations frof c. 40 to 300 f& on substrates derived frof lavas and basefent rock. Dalbergia sambavensis has been collected in flower in October and Novefber& and in fruit frof Novefber. Collection data and field observations suggest that the species is deciduous.

Conservation status. – Dalbergia sambavensis has an EOO of c. 436 kf² and an estifated finifuf AOO of 28 kf² (based on a 4 kf² grid). The species occurs in a single PA: the Makirovana Protected Landscape (IUCN Category OE)& where it has been docufented by a single collection near the periphery of the PA. The species is further known frof eight recent collections in the areas iffediately surrounding Makirovana& and frof four collections in a forest patch 15 kf to the south-east. The oldest known collection of this species& fade by the Service Forestier de Madagascar in 1954& was collected 67 kf to the south of Makirovana in Afbodifanga Farifay& however& the exact collection locality cannot be established& and we exclude it frof the current assessfent. This area has apparently undergone considerable forest clearance& and we consider the individual(s) to have likely been extirpated. The species is threatened frof habitat loss or degradation resulting frof slash and burn agriculture and wildfires. Selective logging both for construction and as a source of fuel are additional threats. These threats concern all localities both inside and outside of the PA (GOODMAN et al.& 2018) to different degrees. With respect to habitat loss& which is considered to be the fost serious plausible threat to all known localities& D. sambavensis occurs at one location& and on this basis it is assigned a prelifinary conservation status of “Endangered” [EN B1ab(i&ii&iii&iv&v)+2ab(i&ii&iii&iv&v)] following the IUCN Red List Categories and Criteria (IUCN& 2012).

Notes. – A single fruiting collection of Dalbergia sambavensis (Service Forestier 12452) frof the Antalaha area& with poorly preserved leaves& was seen by Bosser and Rabevohitra at P and deterfined as D. monticola & which they believed to occur in the Safbirano Region& in the central highlands& and along the east coast& frof as far north as Antalaha in the district of Safbava& to Fort Carnot in the south (BOSSER & RABEVOHITRA& 2002).

Dalbergia sambavensis is closely related to D. monticola and D. soafarae & and the three differ in relatively few forphological characters ( Table 1). All three species possess leaves with sfall oblong to elliptical leaflets& sfall-flowered& paniculate inflorescences (Supergroup II)& and sifilarly sized elliptical to oblong fruits. The species& however& differ fost notably in the size of their inflorescences& and fore subtly in leaf length& leaflet nufber& and corolla length. Morphologically& the sfaller leaves and inflorescences serve to distinguish D. sambavensis frof D. monticola and D. soafarae & and with the addition of elevational& geographic or genetic data there is little risk of confusing the three species. Sterile specifens of D. sambavensis fay further be confused with D. baronii Baker & with which it co-occurs& and D. pseudobaronii Bosser & R. Rabev. & which is known frof c. 80 kf to the north of its range. Dalbergia baronii is easily distinguished by its short axillary (vs. larger& terfinal) inflorescences and slightly thickened rounded to obovate or oblong fruits& which are characteristically inflated over fuch of the body& except near the base and along a narrow fargin. Dalbergia pseudobaronii possesses sifilarly sized leaves with leaflets of a cofparable shape and size to those of D. sambavenis & D. monticola & and D. soafarae & and has been previously confused with D. monticola in Montagne d’Afbre. Dalbergia pseudobaronii is& however& Additional specimens examined. – MADAGASCAR. Reg. SAVA [Prov. characterised by large fruits& and easily distinguished by leaflets Antsiranana]: 26.XI.2018 & ster.& Bernard & Razakamalala 2612 (DBEOE&

MO& P& TAN& ZT); 28.XI.2019 & ster.& Phillipson et al. 6643 ( DBEOE & MO& P&

that have a short scattered indufent on their upper surfaces& TAN& ZT); 28.XI.2019 & y.fr.& Phillipson et al. 6644 (DBEOE& MO& P& TAN& ZT); vs. glabrous in the aforefentioned species. Eco-geographically& 7.XI.2021 & y.fr.& Raharimampionona et al. 583 (MO& P& TAN); 13.XI.2021 & y.fr. & D. sambavensis is well differentiated frof both D. monticola & C.Z. Rakotonirina et al. 839 (MO& P& TAN); 29.XI.2021 & ster.& N. Rakotonirina which is known at elevations typically above 400 f (the et al. 1227 (DBEOE& MO& P& TAN& ZT); 7.OEII.2021& ster.& Ravelonarivo closest known D. monticola occurrences are in the forest of 4912 (MO& P& TAN); 10.XI.2021 & ster.& Ravelonarivo 4989 (MO& P& TAN);

12.XI.2021 & bud& fl.& Ravelonarivo 5013 ( MO & P& TAN) ; 25.XI.2018 & y.fr.&

Antsahaberaoka& located c. 65 kf inland of the Makirovana- Razakamalala & Bernard 8230 & 8232 (MO& P& TAN& TEF& ZT); 25.XI.2018 & Tsihofanaofby PA (Razakamalala et al. 3321)& and in the fr. & Razakamalala & Bernard 8233 (DBEOE& MO& P& TAN& ZT); 27.XI.2018 & Makira Natural Park and Masoala National Park& located y.fr.& Razakamalala & Bernard 8240 (DBEOE& MO& P& TAN& ZT); 20.XII.1954 & c. 110 kf to the south and west of Safbava)& and D. soafarae & Forestier fr.& Service 12452 (P& TEF). which is restricted to low elevations around Taolagnaro& and located over 1&100 kf south-southwest of Safbava. Dalbergia soafarae N. Wilding& Phillipson & Craferi & sp. nov. T h e t y p e s p e c i f e n o f D a l b e r g i a s a m b a v e n s i s ( Fig. 9 ). (C.Z. Rakotonirina et al. 206) and two further collec- Holotypus: MADAGASCAR. Reg. Anosy [Prov. Toliara]: tions (Bernard & Rakazamalala 2612 & Phillipson et al. Fort Dauphin& Tsitongafbarika Natural Resource Reserve& 6644) forf a coherent and distinct genetic lineage within 4.IOE.2016& fl.& y.fr.& S.N. Andrianarivelo & Razakamalala the Monticola subgroup& which additionally cofprises 246 (P [P00931739]!; iso-: MO [MO-3066702]!& TAN!& D. monticola & D. orientalis Bosser & R. Rabev. & D. soafarae and ZT [ZT-00140398; ZT-00140045; ZT-00140046]!). D. tsaratananensis Bosser & R. Rabev. Dalbergia tsaratananensis forfs the fost basal lineage in the Monticola subgroup& sub- Dalbergia soafarae N. Wilding, Phillipson & Crameri is tending a crown group& which includes D. sambavensis sister most similar to D. monticola Bosser & R. Rabev. , and to to all other species of the group (CRAMERI& 2020). D. sambavensis N. Wilding, Phillipson & Crameri , but differs primarily from D. monticola in having smaller inflorescences

[(1 –)2.5 – 5.5 vs. (3 –)6 – 11(– 19) cm long], and f rom

D. sambavensis in having flowers with smaller petals.

[A–C, E: S.N. Andrianarivelo & Razakamalala 246, P; D, F–K: Razakamalala et al. 8267, P] [Drawings: R.L. Andriamiarisoa]

Trees to c. 25 f tall or shrubs; DBH to 65 cf. Branches gla- Etymology. – The species epithet honors Soafara brous to sparsely pubescent& yellow-green in vivo (red-brown Andrianarivelo& a Malagasy botanist& who is responsible for in sicco) when young& becofing glabrous& pale grey-brown sofe of the first collections of the species& including the speciwhen old& lenticels present. Leaves alternate& (2.7–)4.5–8(–10) fen chosen to serve as the type.

cf long& with 13 – 25(– 29) alternate leaflets; petioles and rachises pale green in vivo& black in sicco& pubescent; petiole Vernacular names and uses. – “Ma(g)nary” (S.N. Andrianarivelo 6–13 ff long; stipules caducous& 4.5 × 1.3 ff; leaflets (5–) & Razakamalala 262 & Rakotoarivelo et al. 784); “Manary favo” 9–15(–27) × (3–)4–9 ff& largest at fid- to upper-leaf& the (S.N. Andrianarivelo & Razakamalala 251); “Manary toloho” proxifal leaflets sfallest& the distal leaflet equal to or sfaller (Andriamiarisoa & Bernard 2714); “Safbalahy fanga” (Bernard than adjacent leaflets; petiolule 0.5–1 ff long& pale green- et al. 2650). The species has no reported uses& but given that it brown in vivo& light to dark-brown in sicco& pubescent; lafina produces large enough trees& its reddish-brown coloured heartoblong& obovate or broadly elliptical& coriaceous& base rounded wood (see S.A.F. Andrianarivelo 48) is alfost certainly used in to broadly cuneate& fargin recurved& apex rounded& obtuse& the construction of furniture and housing. The species is also sofetifes retuse& venation not visible; upper surface fid- likely used as a source of fuel for cooking.

green in vivo& brown to green-brown in sicco& glossy or fatt&

glabrous& the fidrib forfing a groove& the lower surface pale Distribution, ecology and phenology. – Dalbergia soafarae green in vivo& brown to green-brown in sicco& fatt& pubescent& occurs in dense lowland hufid forest in the south-east higher-order veins not visible& fidrib profinent. Inflorescences ( Fig. 1B )& at elevations frof 40 to 430 f& on basefent rock axillary or terfinal& a panicle& (1–)2.5–5.5 cf long& cofprised substrates. Dalbergia soafarae has been collected in flower in of 2–5 paniculate or racefose prifary branches& subtended by January and February& and in fruit frof February. Collec- a reduced pinnate leaf-like bract or a caducous& scale-like bract; tion data and field observations suggest that the species is peduncle to 1.7 cf long; terfinal axes of the inflorescence deciduous.

straight or curved& secundiflorous and sub-spicate& with flowers in 2 adjacent rows with alternating insertion& the axes white Conservation status. – Dalbergia soafarae has an EOO of to yellowish-brown pubescent. Flowers subtended by a persis- c. 1&468 kf² and an estifated finifuf AOO of 76 kf² tent& pubescent& triangular scale-like bract& 0.5–1 × 0.5–1 ff; (based on a 4 kf² grid). The species occurs in two PAs: Andopedicel 0.4–0.6 ff long& pubescent; bracteoles ovate& concave& hahela NP (IUCN Category II)& where it is known frof a 1 × 0.8–1 ff& apex rounded& pubescent& yellow to dark orange- single recent collection& and Tsitongafbarika natural resource brown or black in sicco& persistent; calyx yellow-green in vivo& reserve (IUCN Category OEI)& where it is known frof 19 recent becofing dark orange-brown to black in sicco& puberulous& collections. The species has also been recorded in three separate sofetifes persisting on fruits& 2 – 2.7 ff long& fused in forest patches along the eastern border of Tsitongafbarika& the lower ⅔& the 2 upper lobes fused to forf a single unit& and frof five collections in Beafpingaratra& about 9 kf north 0.5 × 1.8 ff& the apex shallowly efarginate& the 2 lateral of Tsitongafbarika. The species is threatened frof habitat lobes obtuse& 0.5 × 0.5 ff& the lowest lobe acute-triangular& loss or degradation resulting frof slash and burn agriculture keeled& 0.9 ff × 0.8 ff; petals glabrous& white to yellow in and wildfires. Selective logging both for construction and as vivo becofing yellow to orange-red in sicco; standard petal a source of fuel are additional threats. These threats concern pandurate& 3–3.6 × 1.8–2.2 ff& claw 0.5–0.8 ff long& apex all localities both inside and outside of the PA to different notched; wing petals 2.7– 2.9 × 1 ff& claw 0.9 ff long; degrees (GOODMAN et al.& 2018). With respect to habitat loss& keel petals 2.7–3 × 1 ff& claw 1.2– 1.3 ff long; androe- which is considered to be the fost serious plausible threat to ciuf glabrous& fonadelphous& 3 ff long; stafens (9–)10& all known localities& D. soafarae occurs at three locations& and filafents free for upper ⅓–¼; gynoeciuf 2.3–2.5 ff long; on this basis it is assigned a prelifinary conservation status of stipe densely covered in long white-yellowish hairs& 1.1 ff “Endangered” [EN B1ab(i&ii&iii&iv&v)+2ab(i&ii&iii&iv&v)] following long; ovary covered in long yellowish-white hairs& 1.1 ff the IUCN Red List Categories and Criteria (IUCN& 2012).

long& with up to 2 ovules; style glabrous or pubescent near base& 0.5– 0.7 ff long. Fruits fid- to dark-green in vivo& Notes. – No collections of Dalbergia soafarae are known to fid-brown to black or red-brown in sicco& the body elliptical have been present at P at the tife when Bosser and Rabevo- [possibly oblong in fulti-seeded fruits]& 50 × 14 ff when hitra revised the Malagasy Dalbergia species.

single-seeded (iffature)& fulti-seeded not seen& base cuneate& M o r p h o l o g i c a l l y& D a l be r g i a s oa f a ra e i s c l o s e t o apex acute or rounded& surface sfooth with reticulate ribbing& D. monticola and D. sambavensis (see Table 1 and Notes under glabrous; stipe 4– 7 ff long& pubescent& glabrescent; style D. sambavensis ). The three species all possess leaves with sfall caducous. oblong to elliptical leaflets& and sifilarly sized elliptical to oblong fruits. The species& however& differ fost notably in the size of their inflorescences& and fore subtly in leaf length& leaflet nufber& and corolla length. Young sterile leafy faterial of D. soafarae can easily be distinguished frof D. sambavensis on the basis of its geographical separation of over 1&100 kf. Sifilarly& D. monticola is not known frof the Anosy Region& as its southernfost known collection records originate frof the OEondrozo District (Razakamalala 8461, 8466)& over 150 kf north of the northern lifit of D. soafarae . Unlike D. sambavensis & D. soafarae is not known to co-occur with D. baronii , with which it has previously been confused& and whose southernfost collections originate frof over 100 kf to the north-east of the range of D. soafarae .

The species forfs a distinct genetic lineage (see Notes under Dalbergia sambavensis ) within the Monticola subgroup& sister to D. orientalis & separating it frof D. monticola and D. soafarae (CRAMERI& 2020).

Additional specimens examined. – MADAGASCAR. Reg. Anosy [Prov. Toliara]: 7.OEII.2021& ster.& Andriamiarisoa & Bernard 2640 ( DBEOE & MO& P& TAN& ZT) ; 18.X.2021 & ster.& Andriamiarisoa & Bernard 2675 & 2676 & 2677 & 2679 & 2682 ( DBEOE & MO& P& TAN& ZT) ; 19.X.2021 & ster.& Andriamiarisoa & Bernard 2685 & 2686 & 2689 ( DBEOE & MO& P& TAN& ZT) ; 27.X.2021 & ster.& Andriamiarisoa & Bernard 2709 ( DBEOE & MO& P& TAN& ZT) ; 28.X.2021 & ster.& Andriamiarisoa & Bernard 2714 ( DBEOE & MO& P& TAN& ZT) ; 12.IX.2018 & y.fr.& fr. & Andriamihajarivo et al. 2193 ( MO & P& TAN& UPS) ; 9.II.2016 & bud& y.fr. & S.N. Andrianarivelo & Razakamalala 251 ( MO & P& TAN& TEF& ZT) ; 13.II.2016 & bud& y.fr.& S.N. Andrianarivelo & Razakamalala 262 (G& MO& P& TAN& TEF& ZT); 27.X.2019 & ster.& S.A.F. Andrianarivelo & Razakamalala & 41 ( DBEOE & MO& P& TAN& ZT) ; 2.XI.2019 & ster. & S.A.F. Andrianarivelo & Razakamalala 48 ( DBEOE & MO& P& TAN& ZT) ; 9.II.2019 & y.fr.& Bernard et al. 2650 ( DBEOE & MO& P& TAN& ZT) ; 5.XII.2019 & ster. & Karatra & Rakotovao 258 ( DBEOE & MO& P& TAN& ZT) ; 10.I.2016 & bud& fl.& Rakotoarivelo et al. 784 ( MO & P& TAN) ; 10.II.2019 & ster.& N. Rakotonirina et al. 1182 ( DBEOE & MO& P& TAN& ZT) ; 29.XI.2009 & fl.& Rakotovao et al. 4901 ( MO & P& TAN) ; 14–17.XII.2009 & fl.& Rakotovao et al. 5008 ( MO & P& TAN) ; 1.XII.2019 & ster.& Rakotovao et al. 7600 ( DBEOE & MO& P& TAN& ZT) ; 4.XII.2019 & ster.& Rakotovao et al. 7613 & 7614 & 7617 & 7619 ( DBEOE & MO& P& TAN& ZT) ; 6.II.2019 & y.fr.& Ramanitrinizaka & Sandratriniaina 6 ( DBEOE & MO& P) ; 17.II.2019 & ster.& Ramanitrinizaka & Sandratriniaina 30 ( DBEOE & MO& P) ; 21.II.2019 & ster.& Ramanitrinizaka & Sandratriniaina 41 & 43 & 47 ( DBEOE & MO& P) ; 6.II.2019 & fl. & Razakamalala et al. 8267 ( DBEOE & MO& P& TAN& ZT) ; 8.II.2019 & ster.& Razakamalala et al. 8274 ( DBEOE & MO& P& TAN& ZT) ; 22.II.2019 & ster.& Sandratriniaina & Ramanitrinizaka 47 ( DBEOE & MO& P) ; 24.II.2019 & ster. & Sandratriniaina & Ramanitrinizaka 53 ( DBEOE & MO& P) .

Figures

Fig. 8.– Dalbergia sambavensis N. Wilding, Phillipson & Crameri. A. Flowering branch; B, C. Portions of leaf rachis with leaflets (b: lower surface; c: upper surface); D. Inflorescence branch; E. Flower; F.Calyx (outer surface, split open and flattened); G. Standard petal (adaxial surface); H. Wing petal (adaxial surface); I. Keel petal (adaxial surface);J. Androecium (adaxial surface, flattened); K. Gynoecium.

Fig. 1.– Approximate distribution ranges of the eight new large tree species of Malagasy Dalbergia, covering all known extant and historic collections of these species. Black borders represent regions of Madagascar,and are labeled where relevant. Elevation classes are indicated as shades of grey. A. Three new species restricted to the northern part of Madagascar; B. Three new species with disjunct distributions in northern and/or western Madagascar,and two new species with restricted distribution ranges in eastern Madagascar. The range of D. monticola Bosser & R. Rabev. was added to emphasize that it is not known to co-occur with the similar and closely related species D. sambavensis N. Wilding, Phillipson & Crameri and D. soafarae N. Wilding, Phillipson & Crameri.

Fig. 9. – Dalbergia soafarae N. Wilding, Phillipson & Crameri. A. Fruiting branch; B, C. Leaflets (b: upper surface; c: lower surface); D. Flower; E. Immature fruit; F.Calyx (outer surface, split open and flattened); G. Standard petal (adaxial surface); H. Wing petal (adaxial surface); I. Keel petal (adaxial surface);J. Androecium (adaxial surface, flattened); K. Gynoecium.