Dalbergia razakamalalae Crameri, Phillipson & N.Wilding, 2022

Dalbergia razakamalalae Crameri, Phillipson & N.Wilding View in CoL , sp. nov. TYPE: MADAGASCAR. Anosy [Toliara]: For^et d’ Analamahavondjaky (commune de Iaboakoho), 7 Dec 2019 (fl), T. Andriamihajarivo, N. H. Rakotoarivelo & F. Rakotoarivony 2455 (holotype: P [P00853053]!, isotypes: MO, TAN image!, ZT [ZT-00169820]).

Dalbergia razakamalalae is similar to D. maritima R.Vig. in possessing leaves with rather small leaflets and racemose inflorescences, but differs by its consistently glabrous leaves (vs. glabrous or pubescent), larger flowers (10–14 mm vs. 8–10 mm long), and narrowly ovate to narrowly elliptic leaflets (vs. ovate to elliptic) that are thinly coriaceous (vs. coriaceous) and have plane (vs. revolute) margins and frequently an emarginate (vs. obtuse to rounded) apex.

Deciduous tree to ca. 20 m tall, or shrub-like when resprouting after felling, bole to ca. 15 m high, DBH to at least 40 cm; bark gray-brown, smooth at first, becoming fissured with age. Branches glabrous, pale brown to purple-brown in vivo (dark brown to dark purple in sicco) when young, becoming gray-brown, lenticels present. Leaves alternate, 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. Inflorescences racemose, 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 cm long; 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 mm long. Flowers often subtended by glabrous, imparipinnate reduced leaves, 22–57 mm long, 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 mm long 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 mm long 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 mm long claw; wing petals 7.3–10.8 3 2.2–3.2 mm, including 2.0– 2.8 mm long claw, base distinctly auriculate; keel petals 7.3–9.3 3 2.4–2.9 mm, including 2.0– 2.9 mm long claw, base distinctly auriculate; androecium glabrous, diadelphous, 6.4–10.3 mm long; stamens 9 1 1, free for upper 1.7–5.0 mm; gynoecium 7.0– 7.7 mm long, glabrous; stipe ca. 3.5 mm long; ovary 4.3–5.5 mm long, with 3–5 ovules; style slender, slightly incurved, 1.4–1.8 mm long. 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 cm when single-seeded, up to 7.5 3 1.7 cm when 3-seeded, base cuneate, apex rounded or acute, surface indistinctly net-veined, glabrous; stipe ca. 7–10 mm long; style rarely persistent. Seeds (immature) sub-reniform, flattened, brown, ca. 6 3 3 mm. Figures 1E View FIG , 2D View FIG , 5 View FIG .

Etymology — Dalbergia razakamalalae is 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 Dalbergia species made 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 razakamalalae is beautifully veined and burgundy-colored ( Bernard et al. 2645, Karatra & Rakotovao 242, Ramanitrinizaka & Sandratriniaina 1). It is considered to be a high-quality rosewood ( Humbert 20607) and is used in cabinet making ( Humbert 20355bis).

Habitat, Distribution, and Phenology — Dalbergia razakamalalae occurs in inland low-elevation evergreen humid forests on lateritic soils, at 20–510 m elevation 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 View FIG ). Dalbergia razakamalalae has been collected in full flower from November to February. Immature to mature fruits have been recorded from December to February.

Conservation Status — Dalbergia razakamalalae is 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 2 and its former area of occupancy (AOO) was at least 80 km 2 (based on a 4 km 2 grid), whereas its current documented geographic range has the form of an EOO of 2096 km 2 and an AOO of 60 km 2, and comprises four subpopulations. The species occurs in forest ecosystems ( Madagascar Catalogue 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 m and within the minimum convex polygon encompassing all known collections of this species. Therefore, D. razakamalalae is 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. razakamalalae is 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 razakamalalae has previously been included in or associated with D. maritima sensu 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. razakamalalae differs 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. maritima subsp. pubescens , but unlike these taxa, its leaves are consistently glabrous, as summarized in Figs. 1E View FIG , 2D View FIG , 3A–B View FIG and 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. maritima and D. louvelii on account of its small and glabrous leaflets (as in D. maritima subsp. 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. louvelii in 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 1124 and Service Forestier 22334, both included in their broad definition of D. maritima , and potentially also Humbert 20355bis and 20607, two sterile collections present in the Paris herbarium at the time. The collections made in the 1940s ( Humbert 20355bis, Reserves Naturelles 1689 and 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. razakamalalae southwards, 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. razakamalalae by ca. 110 km to 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. razakamalalae and 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 & Rakotovao 241 ( DBEV, MO, P, TAN, ZT) ; same locality, 3 Dec 2019, Karatra & Rakotovao 242 ( DBEV, MO, P, TAN, ZT) ; Ivola forest ( Tolagnaro district ), s. d. (y.fr), Reserves Naturelles 1124 (P) ; Manampanihy valley ( Ampasimena ), 18 Mar 1947, Humbert 20607 (P) ; Manatantely forest ( Tolagnaro district ), 1 Mar 1947, Humbert 20355bis ( MO, P, TAN) ; same locality, 20 Nov 1948 (fl), Reserves Naturelles 1689 (P) ; same locality, 9 Dec 1948 (y.fr), Reserves Naturelles 1689bis (P) ; Tsitongambarika protected area and surroundings ( Iabakoho commune), 6 Feb 2019 (fr), Ramanitrinizaka & Sandratriniaina 1 ( DBEV, MO, P, TAN, ZT) ; same locality, 9 Feb 2019, Ramanitrinizaka & Sandratriniaina 12 ( DBEV, P) ; same locality, 9 Feb 2019 (fr), Ramanitrinizaka & Sandratriniaina 13 ( DBEV, MO, P, TAN) ; same locality, same date, Ramanitrinizaka & Sandratriniaina 18 ( DBEV, MO, P) ; same locality, 12 Feb 2019, Ramanitrinizaka & Sandratriniaina 25 ( DBEV, MO, P) ; same locality, 7 Feb

2019, Ramanitrinizaka & Sandratriniaina 57 ( DBEV, MO, P) ; same locality, 16 Feb 2019, Sandratriniaina & Ramanitrinizaka 23 ( DBEV, MO, P) ; same locality, same date, Sandratriniaina & Ramanitrinizaka 26 ( DBEV, MO, P) ; same locality, same date, Sandratriniaina & Ramanitrinizaka 27 ( DBEV, MO, P) ; same locality, same date, Sandratriniaina & Ramanitrinizaka 29 ( DBEV, MO, P) ; same locality, 1 Apr 2014, Razakamalala 7736 ( MO, P, TAN) ; same locality, same date, Razakamalala 7761 ( MO, P, TAN) ; same locality, same date, Razakamalala 7762 ( MO, P, TAN) ; same locality, same date, Razakamalala 7764 ( MO, P, TAN) ; same locality, 12 Feb 2016, Razakamalala & S. N. Andrianarivelo 8036 ( MO, P, TAN, TEF, ZT) ; same locality, 14 Feb 2016 (fr), Razakamalala & S. N. Andrianarivelo 8040 ( MO, P, TAN, TEF, ZT) ; same locality, 4 Nov 2019, S. A. Andrianarivelo & Razakamalala 53 ( DBEV, MO, P, TAN, ZT) ; same locality, Feb 1963 (fl), Service Forestier (Capuron) 22334 (P, TEF) ; Tsitongambarika protected area and surroundings (Manantenina commune/ Ivohibe-Bemangidy / Antsotso ), 11 Feb 2016, Razakamalala & S. N. Andrianarivelo 8032 ( MO, P, TAN, TEF, ZT) ; same locality, same date, Razakamalala & S. N. Andrianarivelo 8035 ( MO, P, TAN, TEF, ZT) ; same locality, 1 Nov 2019 (fl), Razakamalala & S. A. Andrianarivelo 8558 ( DBEV, MO, P, TAN, ZT) ; same locality, 2 Nov 2019, Razakamalala & S. A. Andrianarivelo 8560 ( DBEV, MO, P, TAN, ZT) ; same locality, 6 Feb 2019 (fr), Razakamalala et al. 8266 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (fr), Bernard et al. 2641 ( DBEV, MO, P, TAN, ZT) ; same locality, 9 Feb 2019, Bernard et al. 2645 ( DBEV, MO, P, TAN, ZT) ; same locality, 11 Feb 2016, S. N. Andrianarivelo & Razakamalala 255 ( MO, P, TAN, TEF, ZT) ; ATSIMO-ATSINANANA [Fianarantsoa]: Amparihy ( Ambitananona - Amparihy - Vangaindrano ), 23 Nov 1953 (fl), Service Forestier 7110 (P, TEF) ; Ankarabolava protected area, 20 Feb 2021, Andriamiarisoa et al. 2620 ( MO, P, TAN) ; Manombo Special Reserve , 5 Nov 2019, Rakotovao & Andriamiarisoa 7522 ( DBEV, MO, P, TAN, ZT) ; same locality, same date, Rakotovao & Andriamiarisoa 7523 ( DBEV, MO, P, TAN, ZT) ; same locality, same date, Rakotovao & Andriamiarisoa 7528 ( DBEV, MO, P, TAN, ZT) ; same locality, 28 Jan 2014, Emeline 23 ( MO, P, ZT) ; same locality, 4 Nov 2019, Andriamiarisoa & Rakotovao 2424 ( DBEV, MO, P, TAN, ZT) ; FITOVI- NANY [Fianarantsoa]: Canton Ambila ( Manakara district ), s. d., Service Forestier 38-R-118 (P) .