Taxonomic Studies on Malagasy Dalbergia (Fabaceae). III. Two New Species from Southeastern Madagascar and an Emended Description of the Rosewood Species Dalbergia maritima Crameri, Simon Phillipson, Peter B. Rakotonirina, Nivohenintsoa Wilding, Nicholas Andriamiarisoa, Roger Lala Lowry Ii, Porter P. Widmer, Alex Systematic Botany 2022 Basel, Switzerland 2022-06-14 47 2 397 416 BSZ3W Crameri, Phillipson & N.Wilding Crameri, Phillipson & N. Wilding 2022 [848,1487,1823,1846] Magnoliopsida Fabaceae Dalbergia Plantae Fabales 15 411 Tracheophyta species razakamalalae sp. nov.   Deciduous treeto ca. 20 mtall, or shrub-like when resprouting after felling, bole to ca. 15 mhigh, DBH to at least 40 cm; bark gray-brown, smooth at first, becoming fissured with age. Branchesglabrous, pale brown to purple-brown in vivo (dark brown to dark purple in sicco) when young, becoming gray-brown, lenticels present. Leavesalternate, 7–13(–16) cm long, with 11–19(–23) alternate leaflets, petiole and rachis purplish-green in vivo, dark brown to dark purple in sicco, glabrous; petiole (9–)12–20(–25) mm long; stipules narrowly ovate, ca. 3.0 3 1.0 mm, glabrous, early caducous; leaflets (8–)13–25(–35) 3 (4–)5–10(–14) mm, often noticeably smaller toward base or/and apex; petiolule 1.0–2.0 mm long, yellow-green in vivo, dark brown to dark purple in sicco, glabrous; lamina narrowly ovate to narrowly elliptic, rarely ovate to elliptic, thinly coriaceous, base cuneate, margins not revolute in sicco, apex emarginate, rarely obtuse, venation brochidodromous, with 5–7 principal lateral veins per side; upper surface matt, mid-green in vivo, dark purple-brown in sicco, glabrous, venation inconspicuous (slightly raised in sicco), midrib inconspicuous or forming a groove; lower surface paler than upper in vivo and in sicco, glabrous, venation forming a loose network with higher-order veins (often paler than matrix in sicco), midrib prominent. Inflorescencesracemose, composed of (1–)4–12 alternate flowers each, often with imparipinnate reduced leaves subtending individual flowers especially near base (thus becoming single-flowered), sometimes pseudo-paniculate with smaller racemes (bearing abortive flower buds) branching off from close to base, 2–5 cmlong; axes green to purple-green especially at apex in vivo, dark brown to dark purple in sicco, glabrous; anthesis before or concurrent with leaf emergence; peduncle to 6 mmlong. Flowersoften subtended by glabrous, imparipinnate reduced leaves, 22–57 mmlong, with 7–13 alternate, narrowly ovate to narrowly elliptic leaflets, leaving visible scar, bracts not seen (early caducous and leaving visible scar); pedicel 2–4(–6) mm long, slender, glabrous; bracteoles lanceolate, ca. 2.5–3.0 3 0.6–0.8 mm, glabrous, early caducous; calyx base to apex of longest petal 10–14 mmlong in sicco; calyx bright green to purple and brightly dotted especially at base in vivo, purple-brown, darker at base in sicco, with a 3.1–4.0 mm long tube, 7.0– 8.2 mmlong from base to apex of lower lobe, glabrous, persistent, 2 upper sepals long-connate, their lobes 2.3–3.9 3 2.5–2.9 mm, apex obtuse to rounded, 2 lateral sepals triangular, 3.2–4.2 3 1.5–2.2 mm, lowest sepal triangular, margins weakly incurved, apex slightly hooked, 3.2–4.2 3 1.4–2.2 mm; petals glabrous, white with often pink or bluish tinged veins at anthesis, dark yellow to dark cream in sicco; standard petal broadly obovate to orbicular, claw and lamina forming an obtuse angle, margins slightly incurved backwards when in full flower in vivo, base attenuate, apex notched, 9.6–11.5 3 5.8–9.2 mm, including 2.4–3.9 mmlong claw; wing petals 7.3–10.8 3 2.2–3.2 mm, including 2.0– 2.8 mmlong claw, base distinctly auriculate; keel petals 7.3–9.3 3 2.4–2.9 mm, including 2.0– 2.9 mmlong claw, base distinctly auriculate; androecium glabrous, diadelphous, 6.4–10.3 mmlong; stamens 9 1 1, free for upper 1.7–5.0 mm; gynoecium 7.0– 7.7 mmlong, glabrous; stipe ca. 3.5 mmlong; ovary 4.3–5.5 mmlong, with 3–5 ovules; style slender, slightly incurved, 1.4–1.8 mmlong. Fruits(immature) purple-red to carmine in vivo, purple-brown in sicco, with 1–3 seeds, body oblong or narrowly elliptic, 3.5–5.5 3 0.8–1.5 cmwhen single-seeded, up to 7.5 3 1.7 cmwhen 3-seeded, base cuneate, apex rounded or acute, surface indistinctly net-veined, glabrous; stipe ca. 7–10 mmlong; style rarely persistent. Seeds(immature) sub-reniform, flattened, brown, ca. 6 3 3 mm. Figures 1E, 2D, 5.    Etymology— Dalbergia razakamalalaeis named in honor of the botanist Richardson Razakamalala, who has made more than 9000 high-quality collections over the last two decades, contributing significantly to the knowledge of the flora of Madagascar, and which have included collections of this and many other  Dalbergiaspeciesmade while working together with local guides and other members of the Missouri Botanical Garden’ s research team in Madagascar.  Vernacular Names and Uses—Sambalahimanga ( Andriamihajarivo et al. 2455), Tombobitsy lahy ( Razakamalala & S. N. Andrianarivelo 8035), Tongobitsy or Tambobitsy ( Reserves Naturelles 1689). The heartwood of  Dalbergia razakamalalaeis beautifully veined and burgundy-colored (  Bernardet al. 2645, Karatra & Rakotovao 242, Ramanitrinizaka & Sandratriniaina 1). It is considered to be a high-quality rosewood (  Humbert20607) and is used in cabinet making (  Humbert20355bis).   Habitat, Distribution, and Phenology— Dalbergia razakamalalaeoccurs in inland low-elevation evergreen humid forests on lateritic soils, at 20–510 melevation in southeastern Madagascar(Anosy and Atsimo-Atsinanana Regions). It occurs in and around the Tsitongambarika protected area, the Ankarabolava protected area, in the northern parcel of the Manombo protected area, and in the Ampotaky forest farther north and more inland of Tsitongambarika ( Fig. 3D).  Dalbergia razakamalalaehas been collected in full flower from November to February. Immature to mature fruits have been recorded from December to February.    Conservation Status— Dalbergia razakamalalaeis known from 42 positively identified collection records that represent 6 extant occurrences and 5 occurrences that appear to have been extirpated. Its former extent of occurrence (EOO) was at least 4293 km 2and its former area of occupancy (AOO) was at least 80 km 2(based on a 4 km 2grid), whereas its current documented geographic range has the form of an EOO of 2096 km 2and an AOO of 60 km 2, and comprises four subpopulations. The species occurs in forest ecosystems (  MadagascarCatalogue 2021). Forest cover decline between 1953 and 2017 was estimated from the forest cover time series of Vieilledent et al. (2018a, 2018b) to be 73% in the altitudinal range of 20–510 mand within the minimum convex polygon encompassing all known collections of this species. Therefore,  D. razakamalalaeis inferred to have undergone and to be undergoing continuing decline in EOO, AOO, quality of habitat, number of subpopulations, and number of mature individuals. This species occurs at five locations with respect to the most serious plausible threat, which is selective logging for trade in its high-quality heartwood, as inferred from recent field observations of exploited trees at several sites. The occurrences within the protected areas of Ankarabolava, Manombo, and Tsitongambarika represent three separate locations. Occurrences outside of the Tsitongambarika protected area represent the fourth location. The subpopulation from the Ampotaky forest at Beampingaratry, which is situated at higher elevation and appears to be less accessible, represents the fifth location. For these reasons,  D. razakamalalaeis assigned a preliminary IUCN conservation status of Endangered: EN B1ab(i,ii,iii,iv,v)12ab(i,ii,iii,iv,v).   Notes—Material of  Dalbergia razakamalalaehas previously been included in or associated with  D. maritimasensu Bosser and Rabevohitra (2002), mainly owing to their overlapping morphological variation with respect to leaflet size, shape, and number, and to inflorescence structure. However,  D. razakamalalaediffers in its leaflet texture and margins, and in its larger flowers. Its flowers are similar in size to those of  D. louvelii s.l., and it inhabits inland low-elevation evergreen humid forests like  D. maritimasubsp. pubescens, but unlike these taxa, its leaves are consistently glabrous, as summarized in Figs. 1E, 2D, 3A–Band Table 3. A specimen with both flowers ( Reserves Naturelles 1689) and immature fruits ( Reserves Naturelles 1689bis, collected 19 d later) was examined by Bosser and Rabevohitra in 1995, who associated it with both  D. maritimaand  D. louveliion account of its small and glabrous leaflets (as in  D. maritimasubsp. maritima) and large flowers (as in  D. louvelii). They suggested that this collection might be a hybrid between these two taxa, without any evidence for the presence of  D. louveliiin the region, and evidently without realizing that its morphology is consistent with other collections they saw from the same region and habitat type, viz. Reserves Naturelles 1124and Service Forestier 22334, both included in their broad definition of  D. maritima, and potentially also  Humbert20355bisand 20607, two sterile collections present in the Paris herbarium at the time. The collections made in the 1940s (  Humbert20355bis, Reserves Naturelles 1689and 1689bis) from forests around Manantantely, and the collection from the 1960s from the Ivola forest ( Reserves Naturelles 1124) increase the documented distribution range of  D. razakamalalaesouthwards, including to the southern part of the Tsitongambarika protected area, but no extant occurrences are known from these areas, despite extensive recent collection efforts, so these populations are presumed to have been extirpated. Likewise, a sterile collection from Ambila in the Manakara district ( Service Forestier 38-R-118) increases the documented range of  D. razakamalalaeby ca. 110 kmto the north, but it probably dates from the 1950s and originates from a site that is not included in a protected area, so this possible subpopulation likewise probably no longer exists. A recently made sterile collection from the degraded forests of Fotobohitra in the Ifanadiana district ( Ravaomanalina 71) may also represent  D. razakamalalaeand would further increase its range northwards, but the physical specimen could not yet be examined.   Additional Specimens Examined—   Madagascar.— ANOSY[ Toliara]: Ampotaky forest( Beampingaratry),  3 Dec 2019,  Karatra& Rakotovao241( DBEV, MO, P, TAN, ZT); same locality,  3 Dec 2019,  Karatra& Rakotovao242( DBEV, MO, P, TAN, ZT);  Ivola forest( Tolagnaro district), s. d. (y.fr),  Reserves Naturelles1124(P);  Manampanihy valley( Ampasimena),  18 Mar 1947,  Humbert20607(P);  Manatantely forest( Tolagnaro district),  1 Mar 1947,  Humbert20355bis( MO, P, TAN); same locality,  20 Nov 1948(fl),  Reserves Naturelles1689(P); same locality,  9 Dec 1948(y.fr),  Reserves Naturelles1689bis(P);  Tsitongambarikaprotected area and surroundings ( Iabakohocommune),  6 Feb 2019(fr),  Ramanitrinizaka& Sandratriniaina1( DBEV, MO, P, TAN, ZT); same locality,  9 Feb 2019,  Ramanitrinizaka& Sandratriniaina12( DBEV, P); same locality,  9 Feb 2019(fr),  Ramanitrinizaka& Sandratriniaina13( DBEV, MO, P, TAN); same locality, same date,  Ramanitrinizaka& Sandratriniaina18( DBEV, MO, P); same locality,  12 Feb 2019,  Ramanitrinizaka& Sandratriniaina25( DBEV, MO, P); same locality, 7 Feb  2019,  Ramanitrinizaka& Sandratriniaina57( DBEV, MO, P); same locality,  16 Feb 2019,  Sandratriniaina& Ramanitrinizaka23( DBEV, MO, P); same locality, same date,  Sandratriniaina& Ramanitrinizaka26( DBEV, MO, P); same locality, same date,  Sandratriniaina& Ramanitrinizaka27( DBEV, MO, P); same locality, same date,  Sandratriniaina& Ramanitrinizaka29( DBEV, MO, P); same locality,  1 Apr 2014,  Razakamalala7736( MO, P, TAN); same locality, same date,  Razakamalala7761( MO, P, TAN); same locality, same date,  Razakamalala7762( MO, P, TAN); same locality, same date,  Razakamalala7764( MO, P, TAN); same locality,  12 Feb 2016,  Razakamalala& S. N. Andrianarivelo8036( MO, P, TAN, TEF, ZT); same locality,  14 Feb 2016(fr),  Razakamalala& S. N. Andrianarivelo8040( MO, P, TAN, TEF, ZT); same locality,  4 Nov 2019,  S. A. Andrianarivelo& Razakamalala53( DBEV, MO, P, TAN, ZT); same locality,  Feb 1963(fl),  Service Forestier( Capuron) 22334(P, TEF);  Tsitongambarikaprotected area and surroundings ( Mananteninacommune/ Ivohibe-Bemangidy/ Antsotso),  11 Feb 2016,  Razakamalala& S. N. Andrianarivelo8032( MO, P, TAN, TEF, ZT); same locality, same date,  Razakamalala& S. N. Andrianarivelo8035( MO, P, TAN, TEF, ZT); same locality,  1 Nov 2019(fl),  Razakamalala& S. A. Andrianarivelo8558( DBEV, MO, P, TAN, ZT); same locality,  2 Nov 2019,  Razakamalala& S. A. Andrianarivelo8560( DBEV, MO, P, TAN, ZT); same locality,  6 Feb 2019(fr),  Razakamalalaet al. 8266( DBEV, MO, P, TAN, ZT); same locality, same date (fr),  Bernardet al. 2641( DBEV, MO, P, TAN, ZT); same locality,  9 Feb 2019,  Bernardet al. 2645( DBEV, MO, P, TAN, ZT); same locality,  11 Feb 2016,  S. N. Andrianarivelo& Razakamalala255( MO, P, TAN, TEF, ZT);  ATSIMO-ATSINANANA[ Fianarantsoa]: Amparihy( Ambitananona - Amparihy - Vangaindrano),  23 Nov 1953(fl),  Service Forestier7110(P, TEF);  Ankarabolavaprotected area,  20 Feb 2021,  Andriamiarisoaet al. 2620( MO, P, TAN);  Manombo Special Reserve,  5 Nov 2019,  Rakotovao& Andriamiarisoa7522( DBEV, MO, P, TAN, ZT); same locality, same date,  Rakotovao& Andriamiarisoa7523( DBEV, MO, P, TAN, ZT); same locality, same date,  Rakotovao& Andriamiarisoa7528( DBEV, MO, P, TAN, ZT); same locality,  28 Jan 2014,  Emeline23( MO, P, ZT); same locality,  4 Nov 2019,  Andriamiarisoa& Rakotovao2424( DBEV, MO, P, TAN, ZT); FITOVI- NANY[ Fianarantsoa]: Canton Ambila ( Manakara district), s. d.,  Service Forestier38-R-118(P). 2019-12-03 DBEV, MO, TAN, ZT Karatra & Rakotovao Madagascar Beampingaratry Ampotaky forest 17 413 1 Toliara 2019-12-03 DBEV, MO, TAN, ZT Karatra & Rakotovao Madagascar Beampingaratry Ampotaky forest 17 413 1 Toliara Reserves Naturelles Madagascar Tolagnaro district Ivola forest 17 413 1 1947-03-18 Madagascar Ampasimena Manampanihy valley 17 413 1 1947-03-01 MO, TAN Madagascar Tolagnaro district Manatantely forest 17 413 1 1948-11-20 MO, TAN Reserves Naturelles Madagascar Tolagnaro district Manatantely forest 17 413 1 1948-12-09 MO, TAN Reserves Naturelles Madagascar Tolagnaro district Manatantely forest 17 413 1 2019-02-06 DBEV, MO, TAN, ZT Ramanitrinizaka & Sandratriniaina Madagascar Iabakoho Tsitongambarika 17 413 1 2019-02-09 DBEV Ramanitrinizaka & Sandratriniaina Madagascar Iabakoho Tsitongambarika 17 413 1 2019-02-09 DBEV, MO, TAN Ramanitrinizaka & Sandratriniaina Madagascar Iabakoho Tsitongambarika 17 413 1 2019-02-09 DBEV, MO Ramanitrinizaka & Sandratriniaina Madagascar Iabakoho Tsitongambarika 17 413 1 2019-02-12 DBEV, MO Ramanitrinizaka & Sandratriniaina Madagascar Iabakoho Tsitongambarika 17 413 1 [625,800,1975,1992] 2019-02-12 DBEV, MO Feb Madagascar Iabakoho Tsitongambarika 17 413 1 [848,1360,159,177] DBEV, MO Ramanitrinizaka & Sandratriniaina 17 413 1 2019-02-16 DBEV, MO Sandratriniaina & Ramanitrinizaka 17 413 1 2019-02-16 DBEV, MO Sandratriniaina & Ramanitrinizaka 17 413 1 2019-02-16 DBEV, MO Sandratriniaina & Ramanitrinizaka 17 413 1 2019-02-16 DBEV, MO Sandratriniaina & Ramanitrinizaka 17 413 1 2014-04-01 MO, TAN Razakamalala 17 413 1 [910,1435,303,321] 2014-04-01 MO, TAN Razakamalala 17 413 1 2014-04-01 MO, TAN Razakamalala 17 413 1 2014-04-01 MO, TAN Razakamalala 17 413 1 2016-02-12 MO, TAN, TEF, ZT Razakamalala & S. N. Andrianarivelo 17 413 1 2016-02-14 MO, TAN, TEF, ZT Razakamalala & S. N. Andrianarivelo 17 413 1 2019-11-04 DBEV, MO, TAN, ZT S. A. Andrianarivelo & Razakamalala 17 413 1 1963-02 TEF Service Forestier & Capuron 17 413 1 2016-02-11 MO, TAN, TEF, ZT Razakamalala & S. N. Andrianarivelo Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 2016-02-11 MO, TAN, TEF, ZT Razakamalala & S. N. Andrianarivelo Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 2019-11-01 DBEV, MO, TAN, ZT Razakamalala & S. A. Andrianarivelo Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 2019-11-02 DBEV, MO, TAN, ZT Razakamalala & S. A. Andrianarivelo Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 2019-02-06 DBEV, MO, TAN, ZT Razakamalala Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 2019-02-06 DBEV, MO, TAN, ZT Razakamalala Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 [848,1487,687,706] 2019-02-09 DBEV, MO, TAN, ZT Razakamalala Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 2016-02-11 MO, TAN, TEF, ZT S. N. Andrianarivelo & Razakamalala Tsitongambarika Ivohibe-Bemangidy Manantenina 17 413 1 1953-11-23 TEF Service Forestier Ambitananona - Amparihy - Vangaindrano Amparihy 17 413 1 Fianarantsoa 2021-02-20 MO, TAN Andriamiarisoa Ankarabolava 17 413 1 Fianarantsoa 2019-11-05 DBEV, MO, TAN, ZT Rakotovao & Andriamiarisoa Manombo Special Reserve 17 413 1 Fianarantsoa 2019-11-05 DBEV, MO, TAN, ZT Rakotovao & Andriamiarisoa Manombo Special Reserve 17 413 1 Fianarantsoa 2019-11-05 DBEV, MO, TAN, ZT Rakotovao & Andriamiarisoa Manombo Special Reserve 17 413 1 Fianarantsoa [848,1297,903,922] 2014-01-28 MO, ZT Emeline Manombo Special Reserve 17 413 1 Fianarantsoa 2019-11-04 DBEV, MO, TAN, ZT Andriamiarisoa & Rakotovao Manombo Special Reserve 17 413 1 Fianarantsoa Service Forestier Manakara district 17 413 1 Fianarantsoa