Dalbergia pseudomaritima Crameri, Phillipson & N.Wilding, 2022

Dalbergia pseudomaritima Crameri, Phillipson & N.Wilding View in CoL , sp. nov. TYPE: MADAGASCAR. Anosy [Toliara]: Sainte Luce, 13 Feb 2019 (fr), N. Rakotonirina, R. Razakamalala & R. Bernard 1190 (holotype: P [P01069698]!, isotypes: DBEV, MO, TAN image!, ZT [ZT-00169818]).

Dalbergia pseudomaritima is similar to D. chapelieri Baill. in possessing paniculate inflorescences that appear before or at the same time as the emerging, glabrous leaves, but differs by its shorter leaves [(4–)5–8(–10) cm vs. (8–)10–18(–26) cm long] with distinctly smaller leaflets [distal leaflets 7–15(–22) 3 5–8(–12) mm vs. 22–48 3 11–25 mm and sometimes reaching 90 3 40 mm on coppice shoots] that are broadly elliptic to orbicular (vs. elliptic to oblong-elliptic or obovate), resembling those of Dalbergia maritima R.Vig. in number and size.

Deciduous tree to ca. 12 m tall, or shrub-like when resprouting after felling, bole to ca. 7 m high, DBH to at least 25 cm; bark pale gray to brown, smooth at first, becoming fissured with age. Branches glabrous, orange-brown in vivo (dark brown to dark purple in sicco) when young, becoming gray, lenticels present. Leaves alternate, (4–)5–8(–10) cm long, with (8–)10–17(–21) alternate leaflets, petiole and rachis bright green in vivo, purple-brown in sicco, glabrous; petiole (6–)8–10(–12) mm long; stipules obovate, 4.0–6.5 3 1.0–2.0 mm, glabrous, early caducous; leaflets (5–)7–14(–22) 3 (4–)5– 8(–12) mm, sometimes noticeably smaller toward base, but often rather uniform; petiolule 0.5–2.0 mm long, yellow-green in vivo, dark brown to black in sicco, glabrous; lamina broadly elliptic to orbicular, rarely obovate, thinly coriaceous, base broadly cuneate, margins thickened but not revolute in sicco, apex shallowly retuse, sometimes mucronulate or rounded, venation brochidodromous, with 5–9 principal lateral veins per side; upper surface matt, yellow-green in vivo, olive-green to red-brown in sicco, glabrous, venation inconspicuous (slightly raised in sicco), midrib inconspicuous or forming a groove; lower surface matt, paler than upper in vivo and in sicco, glabrous, venation forming a dense network of higher-order veins, contrasting and often darker than matrix in sicco, highest-order veins often open-ended, midrib prominent. Inflorescences paniculate, composed of 6–20 flowers each, compact, 2–5 cm long, with (2–)4–6 partial inflorescences composed of (1–)2–6 flowers each; axes green in vivo, dark brown in sicco, glabrous or sparsely and minutely ciliate at junctions; anthesis before or concurrent with leaf emergence; peduncle to 8 mm long. Flowers subtended by glabrous or minutely ciliate, oblong to obovate bracts, 3.5–6.0 3 1.0– 1.5 mm, early caducous; pedicel 0.5–2.5 mm long, glabrous; bracteoles obovate, (1.5–)3.5–4.5 3 (0.5–)1.0– 1.5 mm, glabrous or minutely ciliate, early caducous; calyx base to apex of longest petal 8–12 mm long in sicco; calyx green, reddish at base in vivo, yellow-brown to purple-brown in sicco, with a 2.5–3.4 mm long tube, 5–8 mm long from base to apex of lower lobe, glabrous or sparsely and minutely ciliate, persistent, 2 upper sepals long-connate, their lobes 1.8–2.5 3 1.9–2.5 mm, apex obtuse to subacute, 2 lateral sepals cymbiform, 3.1–4.2 3 1.3–1.9 mm, lowest sepal triangular, margins incurved, apex often distinctly hooked, 3.4–4.6 3 1.0– 2.5 mm; petals glabrous, white or pinkish-white at anthesis, becoming cream post anthesis, yellow to brown in sicco; standard petal ovate to elliptic to obovate, claw and lamina almost perpendicular, margins incurved forwards in vivo, base truncate to subcordate, apex notched, 8.8–10.0 3 3.7–4.7 mm, including 2.5–3.5 mm long claw; wing petals 7.3–10.3 3 2.0– 2.8 mm, including 1.5–2.9 mm long claw, base distinctly auriculate; keel petals 7.4–9.4 3 2.5–3.1 mm, including 1.8–2.7 mm long claw, base distinctly auriculate; androecium glabrous, monadelphous or diadelphous, 9.4–10.6 mm long; stamens 9–10 or 9 1 1, free for upper 2.7–4.0 mm; gynoecium 6.4–8.2 mm long, glabrous; stipe 3–4 mm long; ovary 3.5–4.5 mm long, with 3–5 ovules; style slender, slightly incurved, 1.9–2.5 mm long. Fruits (immature) yellow-green becoming red-brown in vivo, yellow-brown to red-brown in sicco, with 1–3(–4) seeds, body elliptic to oblong, 4.5–6.5 3 1.6–2.3 cm when single-seeded, up to 8.5 3 2.5 cm when 3-seeded, base attenuate, apex rounded or obtuse, surface with reticulate veins, glabrous; stipe 5–10 mm long; style persistent. Seeds (immature) sub-reniform, flattened, brown, 8.0–9.0 3 5.0–6.0 mm. Figures 1F View FIG , 2F View FIG , 4 View FIG .

Etymology —The epithet reflects the superficial similarity to and confusion with Dalbergia maritima .

Vernacular Names and Uses —Manary (Ramamonjiarisoa 4), Manary toloho (Ramamonjiarisoa 10), Sambalahy (Ramison & Ramisy 108), Tombobitsy ( Raza fi mandimby et al. 237).

The heartwood of Dalbergia pseudomaritima is orange-brown in color (S. A. Andrianarivelo & Razakamalala 58, Razakamalala & S. A. Andrianarivelo 8566). Its wood is used as firewood and for charcoal production (R. Randrianaivo pers. comm.).

Habitat, Distribution, and Phenology — Dalbergia pseudomaritima occurs in littoral forests on sand and adjacent swamp forests (Razakamalala et al. 6675), with one collection from inland low-elevation evergreen humid forests on sandy lateritic soils near a stream ( Bernard et al. 2654), at 0–30 m elevation. It is restricted to southeastern Madagascar (Anosy Region), occurring mainly in the protected areas of Mandena and Sainte Luce ( Fig. 3D View FIG ). Dalbergia pseudomaritima has been collected in full flower from October to January. Immature fruits have been recorded from October, and mature fruits have been observed only once in late March (Gereau et al. 3326).

Conservation Status — Dalbergia pseudomaritima is known from 42 collection records that represent 5 extant occurrences and 1 occurrence that appears to have been extirpated. Its former extent of occurrence (EOO) was at least 275 km 2 and its former area of occupancy (AOO) was at least 56 km 2 (based on a 4 km 2 grid), whereas its current documented geographic range has the form of an EOO of 252 km 2 and an AOO of 44 km 2, and comprises three subpopulations. The species mainly occurs in forest ecosystems and rarely extends to marshes ( 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 35% in the altitudinal range of 0–30 m and within the minimum convex polygon encompassing all known collections of this species. Therefore, D. pseudomaritima 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 four locations with respect to the most serious plausible threat, which is habitat degradation or loss due to land clearing and fire for subsistence agriculture. The occurrences within the protected areas of Mandena and Sainte Luce represent two separate locations. Occurrences outside of the Sainte Luce protected area, including sites north of the Ebakika river, represent the third location. The Ampasy forest subpopulation, which is comparatively less accessible, represents the fourth location. For these reasons, D. pseudomaritima 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 pseudomaritima 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 and number, and due to their occurrence in littoral forest. However, D. pseudomaritima differs by numerous characters of its leaves, inflorescences, flowers and fruits, as summarized in Fig. 3A–B View FIG and Table 3. By contrast, the inflorescence and flower characters of D. pseudomaritima are similar to those of the closely related D. chapelieri s.l., with which it shares an often conspicuous reticulate venation with open-ended highest-order veins on the lower leaflet laminae. The currently known geographic ranges of D. chapelieri s.l. and D. pseudomaritima do not appear to overlap locally, but the

widely distributed D. chapelieri s.l. occurs both to the east and north of D. pseudomaritima . A single collection of D. pseudomaritima is known from a site located outside of the species’ typical (remaining or former) littoral forest habitat ( Bernard et al. 2654), on sandy lateritic soils near a stream. In the same general area, D. pseudomaritima might come into contact with neighboring populations from low-elevation evergreen humid forests attributed to the most closely related lineage within D. chapelieri s.l. (e.g. Razakamalala 7739, Razakamalala 7765, and S. A. Andrianarivelo & Razakamalala 51, Fig. 1H View FIG ). However, D. pseudomaritima clearly differs from these individuals by its shorter leaves [(4–)5–8(–10) cm vs. (8–) 10–13 cm long] with distinctly smaller leaflets [distal leaflets 7–15(–22) 3 5–8(–12) mm vs. 24–40(–51) 3 14–20(–25) mm] that are broadly elliptic to orbicular (vs. ovate to elliptic) and thinly coriaceous (vs. coriaceous) with plane (vs. strongly revolute) margins ( Figs. 1F–H View FIG , 3A View FIG ; Table 3), and no individuals with an intermediate leaf morphology ( Fig. 3A View FIG ) or genotype (Crameri 2020) have yet been found.

Additional Specimens Examined — Madagascar. — ANOSY [Toliara]: Ambanihazo village ( Iabakoho commune), 31 Aug 2012 (st), Ludovic 1570 ( TAN) ; same locality, 25 Nov 2011 (fl), Razakamalala et al. 6675 ( MO, P, TAN) ; Ampasy forest ( Iabakoho commune), 10 Feb 2019 (st), Bernard et al. 2654 ( DBEV, MO, P, TAN, ZT) ; Mandena protected area and surroundings, 21 Nov 1977 (st), Ramamonjiarisoa 2 (P) ; same locality, same date (fl), Ramamonjiarisoa 4 (P) ; same locality, same date (st), Ramamonjiarisoa 5 (P) ; same locality, same date (st), Ramamonjiarisoa 10 (P) ; same locality, 12 Jun 1991 (st), Zarucchi et al. 7593 (K, MO, P) ; same locality, 7 Dec 1989 (fl), Dumetz & McPherson 1139 (K, MO, P) ; same locality, 7 Apr 2014 (st), Razakamalala 7783 ( MO, P, TAN) ; same locality, Nov 1978 (fl), Service Forestier 30547 (P) ; same locality, 16 Oct 1989 (bud), Service Forestier 33262 5 Rabevohitra 2033 (K, MO, P, TEF, WAG) ; Mandromodromotra , 6 Dec 2006 (fl), Ramison & Ramisy 108 ( MO, P, TAN) ; same locality, same date (y.fr), Ramison & Ramisy 109 ( MO, P, TAN) ; Sainte Luce protected area and surroundings, 22 Nov 2011 (y.fr), Ratovoson 1713 ( MO, P, TAN) ; same locality, 16 Jan 1990 (y.fr), McPherson et al. 14804 ( MO) ; same locality, 16 Oct 2008 (fl, y.fr), Raza fi mandimby et al. 237 ( TEF) ; same locality, 18 Nov 2004 (fl), Raharimampionona et al. 1 ( MO, P, TEF) ; same locality, 4 Nov 2003 (fl), Rabenantoandro et al. 1556 ( MO, P, TEF) ; same locality, 15 Dec 2000 (fl), Faliniaina et al. 10 (L, MO, P, TEF, WAG) ; same locality, 18 Dec 1993 (y.fr), Luckow 4150 (BH, K, MO, TAN, WAG, Z) ; same locality, 16 Jan 1990 (y.fr), Dumetz 1195 (K, MO, P) ; same locality, 26 Apr 1989 (st), Rabevohitra 1928 ( MO, P) ; same locality, 15–16 Jan 1990 (y.fr), Rabevohitra 2145 (K, MO, P, TEF) ; same locality, 17–18 Jan 1990 (fl), Rabevohitra 2178 (K, MO, P, TEF) ; same locality, 6 Nov 2019 (st), Razakamalala & S. A. Andrianarivelo 8566 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (fl), Razakamalala & S. A. Andrianarivelo 8567 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (st), Razakamalala & S. A. Andrianarivelo 8568 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (y.fr), Razakamalala & S. A. Andrianarivelo 8569 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (st), Razakamalala & S. A. Andrianarivelo 8570 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (y.fr), Razakamalala & S. A. Andrianarivelo 8571 ( DBEV, MO, P, TAN, ZT) ; same locality, 5 Apr 2014 (st), Razakamalala et al. 7767 ( MO, P, TAN) ; same locality, 20 Oct 2012 (fl), Razakamalala et al. 7228 ( MO, P, TAN) ; same locality, 17 Oct 2012 (fl, y.fr), Ramananjanahary et al. 780 ( MO, P, TAN) ; same locality, 20 Oct 2012 (bud, fl, y.fr), Ramananjanahary et al. 830 ( MO, P, TAN) ; same locality, 29 Mar 1989 (fr), Gereau et al. 3326 (K, MO, P, WAG) ; same locality, 7 Nov 2019 (st), S. A. Andrianarivelo & Razakamalala 58 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (fl), S. A. Andrianarivelo & Razakamalala 60 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (fl, y.fr), S. A. Andrianarivelo & Razakamalala 63 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (st), S. A. Andrianarivelo & Razakamalala 64 ( DBEV, MO, P, TAN, ZT) ; same locality, same date (fl, y.fr), S. A. Andrianarivelo & Razakamalala 65 ( DBEV, MO, P, TAN, ZT) .