Xyloolaena richardii (Baill.) Baill.

Lowry Ii, Porter P., Schatz, George E. & Wolf, Anne-Elizabeth, 2002, Endemic families of Madagascar. VIII. A synoptic revision of Xyloolaena Baill. (Sarcolaenaceae), Adansonia (3) 24 (1), pp. 7-19 : 12-18

3. Xyloolaena richardii (Baill.) Baill.

Bull. Mens. Soc. Linn. Paris 1: 412 (1884). — Scleroolaena richardii Baill., Hist. Pl. 4: 224 (1872), “richardi”. — Type: Richard s.n., Madagascar, Prov. Antsiranana, Angonsti , [15°16’S, 50°29’E], fr. (holo-, P!) .

Xyloolaena richardii , as circumscribed here, is a tree c. 4-20 m tall, ranging from sea level to over 1,500 m elevation in areas with a humid bioclimate in NE Madagascar, with one population recorded in the far SE (Fig. 1). This species occurs in littoral forest on sand from Lac Nosive S of Toamasina to Ampahana N of Antalaha, but is also found at many inland sites, where it likely occurs on quartzite sands, a pattern also seen in Asteropeia multiflora Thouars (Asteropeiaceae) (see SCHATZ et al. 1999b). Previous authors (e.g., CAVACO 1952a,b; CAPURON 1970) have defined X. richardii broadly to include material placed here in X. sambiranensis and X. speciosa . It can, however, be distinguished by its elliptic to ovate leaves that are usually less than two times as long as broad and whose apex lacks a mucro, and by its yellow flowers and large woody involucre in fruit that measures 3-4 cm tall and 2.6-3.8 cm in diameter.

The type material of Scleroolaena richardii consists only of detached fruits, but their large size makes it abundantly clear that they are conspecific with other collections from humid and subhumid areas in eastern Madagascar.

VERNACULAR NAMES. — Fakody, Fombaotoafo, Pesoala, Takodibe, Tamboro, Tsikody zahana, Vahitoambody, Voakoropetaka, Voantiambody, Voantsatroka.

CONSERVATION STATUS. — With six subpopulations recorded from the current network of protected areas and an area of occupancy> 2,000 km 2, Xyloolaena richardii is regarded as a species of Least Concern (LC).

MATERIAL EXAMINED. — Baron 5799, 6593, without precise locality; Bernard 201, Masoala PN; Humbert 18052, Analavory; Humbert & Capuron 24206 (= Service Forestier 804), vallée de l’Andalangy; Humblot 199, Lac Nossi-Ve [= Nosive]; Lowry 5194, Tanambao; Morat 4482, Antsiasiaka; Perrier de la Bâthie 2162, Maroantsetra; Rabe 146, Masoala PN; Rahajasoa 245, Cap Est; Rahajasoa 628, Masoala PN; A. Rakotozafy 815, Maroantsetra; A. Rakotozafy 2055, Ft. Dauphin; Rasoavimbahoaka 190, Marojejy PN; Réserves Naturelles 2129, Marotandrano RS; Réserves Naturelles 8806, Masoala RNI; Richard s.n., Angonsti; Schatz et al. 3812, Cap Est; Schatz & Villiers 1817, Maroantsetra; Service Forestier 804 (= Humbert & Capuron 24206), vallée de l’Andalangy; Service Forestier 1372, Andatsakala; Service Forestier 3445, Ampanavoana; Service Forestier 7492, Anjanazana; Service Forestier 9084, Beanana to Sahajinja; Service Forestier 10606, Angijombarika, Mandritsara; Service Forestier 10612, Bemanarana; Service Forestier 10720, Ampahana; Service Forestier 15066, Besinkara to Maromandia; Service Forestier 18195, Manompana; Service Forestier 25845, Ambalafarihy.

4. Xyloolaena sambiranensis Lowry & G.E. Schatz , sp. nov.

Frutex vel arbor 3-12 m alta. Folia lamina anguste ovata usque oblonga , (5-)7-13.5(-15.5) × 2.7-5 (-5.5) cm, longitudine latitudinem fere semper plus quam 2.5-plo excedente, apice acuta acuminatave plerumque manifeste mucronata, nervis secundariis alternis suboppostisve (11- ad) 13- ad 16-jugatis. Inflorescentia plerumque triflora ; bracteis longe ante anthesin caducis; involucro florem solitarium subtendente. Flos petalis albis; disco extrastaminali ex appendicibus 5 ligulatis rotundatis usque elongatis 5-6 mm altis unaquaque staminodia c. 25 gerente constante. Fructus involucro 2.1- 3.2(-3.5) cm alto, 2.2-2.5 cm diam. subtentus.

TYPUS. — Schatz 3222, Madagascar, Prov. Antsiranana, Manongarivo RS, 30 km SE of Ankaramy , W slopes of Antsatrotro, 14°08’S, 48°21’E, 200 m, 23-25 Mar. 1991, fl. (holo-, MO!; iso-, G!, K!, P!, TEF!) .

Large shrubs to trees 3-12 m tall. Young twigs glabrous to densely short stellate, consistently glabrous with age. Leaves narrowly ovate to oblong, khaki green to tan or chocolate brown and often shiny above (in dry material), usually greener and dull below, chartaceous to subcoriaceous, (5-)7-13.5(-15.5) × 2.7-5(-5.5) cm, almost always greater than 2.5 times as long as wide, glabrous, apex acute to acuminate, usually with a distinct mucro, margin entire, minutely thickened and occasionally slightly revolute, base acute, sometimes nearly obtuse, venation brochidodromous, with (11-)13-16 pairs of alternate to subopposite secondary veins, midrib flat to weakly channeled above, raised below; petiole 8-13 mm long, glabrous; stipules 2, lateral, free, hyaline, lanceolate, 13-15 × 3-4 mm, glabrous, apex acuminate, margin entire, flat to involute throughout or only toward the apex. Inflorescences terminal on principal and lateral branches, primary axis more or less condensed, with c. 5-8 pairs of bract scars and sometimes 1-2 scars of reduced caducous leaves, axes choco- late brown to nearly black, glabrous to densely white strigulose, especially just below the bract scars, with a single terminal peduncle, and usually 2 alternate lateral axes each with a smaller and later-maturing flower, bracts caducous well before anthesis, broadly ovate or rarely bilobed, c. 7-12 × 8-10 mm, concave adaxially, apex broadly acute, sometimes with 2-3 small triangular lobes, light chocolate brown, weakly coriaceous, outer surface obscured by densely whitish wooly tomentose indument, terminal peduncle (ultimate axis below the involucre) stout, 1.5- 3 mm long, 2-3 mm diam., densely white to brownish strigose, peduncle of the lateral axes somewhat more slender, involucre in flower broadly cupulate, tawny to light brown or rusty orange short wooly tomentose, containing a single sessile flower, topped by a prominent ring or flange of spirally arranged rounded-triangular to lanceolate bract-like processes, the smallest ones borne in the lower cycle, gradually larger above, to c. 1.5 mm long, very densely light tawny to rusty brown strigose; sepals 5, imbricate, connate at the base, the outer 2 smaller, narrowly triangular or elliptic to nearly rhombic, c. (11-)15-20 × 5-11 mm, somewhat asymmetrical, the inner 3 very widely ovate to oblate, distinctly asymmetrical, with one lobe broadly expanded laterally (but usually hidden by the adjacent sepal), c. 12- 22 × 15-26 mm, glabrous and dark chocolate brown on adaxial surface (in dry material), densely light tawny strigose on abaxial surface, apex acute; petals 5, imbricate, white, membranaceous when dry, ovate to widely ovate, glabrous, apex broadly acute to rounded; extrastaminal disc of 5 basally fused rounded to elongate, ligulate appendages c. 5-6 mm high, each bearing c. 25 filiform staminodes c. 3-12 mm in length, c. 20 borne along the apical margin and 5-7 about midway on the inner surface; stamens c. 100, filaments slender, free, 10-25 mm long, glabrous, anthers ellipsoid to ovoid, 1-1.5 mm long, basifixed, reflexed; ovary cylindrical, densely whitish to light brownish woolly tomentose, style c. 15 mm long, densely whitish to light brownish woolly tomentose at the base, glabrous above, stigma terminal, hemispherical, glabrous. Involucre in fruit woody, dark brown, smooth or slightly wrinkled on outer surface (in dry material), smooth

mm 15

with vertical striations on inner surface, globose to ellipsoid or ovoid, 2.1-3.2(-3.5) cm tall, 2.2-2.5 cm diam., the wall c. 3-4 mm thick, with an apical opening c. 7-10 mm diam., the collar lined inside with the persistent bract-like processes; fruit globose, entirely contained within the involucre, c. 10-12 mm diam., with 3 weakly defined lobes, densely brown to orangish hirsute, with a persistent, thickened lower portion of the style (sterile upper portion of the fruit?) c. 10 mm long, c. 2-3 mm diam. at the base, densely brown to orange hirsute, tapering to the glabrous upper portion. — Fig. 2 .

Xyloolaena sambiranensis is a shrub to tree 3- 12 m tall. It can be recognized by its white flowers and narrowly ovate leaves with (11-)13- 16 pairs of secondary veins. This species appears to be most closely related to X. speciosa , but can be distinguished by its more numerous secondary veins and caducous inflorescence bracts, its extrastaminal disc comprising 5 well-defined lobes each with c. 25 staminodes, and its more southerly distribution in the Sambirano region (subhumid bioclimate) from S of Maromandia N to the Ambanja area (Fig. 1).

V ERNACULAR NAMES. — Sofiakomba, Sofiankomba, Sofikomba, Sofinakomba, Sofinkomba.

CONSERVATION STATUS. — Xyloolaena sambiranensis has an area of occupancy <2,000 km 2 and only two of the five recorded sub-populations occur within the network of protected areas, and is therefore assigned a preliminary status of Vulnerable ( VU).

PARATYPES. — MADAGASCAR, Prov. Antsiranana: Derleth 160, Manongarivo RS, Besinkara, chemin d’Ambodisakoana à Ambalafary , sur le plateau, 14°04’S, 48°17’E, 300 m, 3 Nov. 1994, fr. (G, MO, P, TAN, TEF) ; Gautier 2984, Manongarivo RS, Besinkara, chemin d’Ambodisakoana à Ambalafary , à mi-montée de la corniche, 14°04’S, 48°17’E, 200 m, 4 Apr. 1996, fl., fr. (G, MO, P, TAN, TEF) ; Malcomber et al. 1708, S of Ankaramy, RN 6, beside road, 13°58’S, 48°10’E, 100 m, 15 Nov. 1992, fr. ( MO, P) ; Perrier de la Bâthie 15052, bassin de l’Ambazona [=Ambazoana], [13°35’S, 48°38’E], bud, fr. ( P) ; Perrier de la Bâthie 15424, same locality, grès liasiques, Feb. 1923, bud, fl. [K, P (2 sheets)] ; Saboureau 60, chaîne de Bekaka, Ambanja , [13°44’S, 48°30’E], 150 m, fl. ( P) ; Service Forestier 2956, Antentezanambo , [13°49’S, 48°06’E], 16-30 Mar. 1951, fl., y.fr. ( P, TAN, TEF) ; Service Forestier 3161, Bekaka, Ambanja , [13°44’S, 48°30’E], 20 Mar. 1950, ster. ( P) ; Service Forestier 7734, Bandrakorony, Ambanja , [13°49’S, 47°52’E], 6 Oct. 1953, fr. ( P, TEF) ; Service Forestier 9286, Benavony, Ambanja , [13°41’30”S, 48°29’E], 22 Mar. 1954, bud, fr. ( P, TEF) ; Service Forestier 10411, route de Maromandia à Ambanja , 10 Aug. 1954, fr. ( P) ; Service Forestier 10451, rivière Antsahakolana, dist. d’Ambanja , Manongarivo RS, [14°04’30”S, 48°16’E], 360 m, 19 July 1954, fr. ( P, TEF) ; Totozafy Be 555, Manongarivo RS, Besinkara , bord de chemin, 14°04’S, 48°17’E, 4 June 1996, fr. (G, MO, P) . Prov. Mahajunga: Decary 2130, Maromandia , terrain gréseux, [14°12’S, 48°05’E], 15 May 1923, fl. [ P (2 sheets)] ; Perrier de la Bâthie 3025, collines gréseuses des environs de Maromandia, [14°12’S, 48°05’E], Mar. 1909, fl. [ P (2 sheets)] ; Service Forestier 19396, Amparihibe, canton Maromandia , colline sur sol gréseux, [14°18’30”S, 48°07’E], 9 Jan. 1959, bud ( P, TEF) .

5. Xyloolaena speciosa Lowry & G.E. Schatz , sp. nov.

Frutex vel arbor usque ad 20 m alta. Folia lamina elliptica usque ovata, (6-)8-14.5 × (4-) 4.5-7.5 cm, longitudine latitudinem plerumque minus quam 2.3- plo excedente, apice rotundata acutave plerumque manifeste mucronata, nervis secundariis alternis suboppositisve 9- vel 10- (ad 12-)jugatis. Inflorescentia (2- vel) 3- ad 5-flora; bracteis omnibus plerumque ad anthesin nonnullis saepe sub fructum persistentibus; involucro florem solitarium subtendente. Flos petalis salmoneis; disco extrastaminali ex annulo humili 3-5 mm alto staminodia 200 ad 250 gerente constante. Fructus involucro 2.0- 2.7 cm alto, 2.0- 2.5 cm diam. subtentus.

TYPUS. — Service Forestier (Capuron) 3051, Madagascar, Prov. Antsiranana, base des collines gréseuses du Levika [=Leviky], aux environs de Matsoborilava, [13°16’S, 49°01’E], 12 Mar. 1951, fl., fr. (holo-, P!; iso-, K!, MO!, P!, TEF) .

Large shrubs or trees to 20 m tall. Twigs glabrous. Leaves elliptic to ovate, chocolate brown or occasionally brownish green and often shiny above (in dry material), brownish to khaki green and dull below, subcoriaceous, (6-)8-14.5 × (4-) 4.5-7.5 cm, usually less than 2.3 times as long as wide, glabrous, apex rounded to acute, usually with a distinct mucro, margin entire, minutely thickened and usually slightly revolute, base rounded to obtuse, occasionally almost truncate, venation brochidodromous, with 9-10(-12) pairs of alternate to subopposite secondary veins, midrib flat to weakly channeled above, raised below; petiole (6-) 10-15 mm long, glabrous; stipules 2, lateral, free, crustaceous to subcoriaceous, ovate, c. 6-8 × 3.5-4.5 mm, glabrous, apex acute to acuminate, margin entire. Inflorescences terminal on main branches and short lateral shoots, a compact cyme with (2-)3-5 flowers, primary axis c. 2-20 mm long, with 2-10 pairs of bract scars, brown (orangish tan on bract scars), glabrous at base, upper internodes densely light tawny wooly tomentose, secondary axes 2-12 mm long, whitish to light tawny wooly tomentose throughout, bracts prominent, usually persistent at least until anthesis and often at least some in fruit, broadly ovate, c. 10-18 × 8-16 mm, strongly concave adaxially, apex broadly acute or with 2-3 irregular lobes, light chocolate brown (in dry material), crustaceous to coriaceous, outer surface often obscured by dense whitish wooly tomentose indument, terminal peduncle (ultimate axis below the involucre) stout, 1-2 mm long, 2-3 mm diam., densely brown to rusty orange wooly tomentose, those of the lateral, later-flowering axes somewhat more slender, involucre in flower densely rusty orange wooly, whitish on the flared rim, containing a single flower, topped by a more or less prominent ring of spirally arranged narrowly triangular bract-like processes, the smallest ones borne in the lower cycle, gradually larger above, to c. 1 mm long, very densely rusty brown wooly, tawny on the tips of the processes; sepals 5, imbricate, connate at the base, the outer 2 smaller, narrowly triangular, c. 16-22 × 9- 13 mm, nearly symmetrical, the inner 3 widely ovate, weakly asymmetrical, with one lobe slightly expanded laterally (but often hidden by the adjacent sepal), c. 16-22 × 22-25 mm, glabrous and dark chocolate brown on the adaxial surface (in dry material), densely whitish to light tawny short tomentose on the abaxial surface, apex rounded to broadly acute; petals 5, imbricate, salmon pink, membranaceous when dry, broadly ovate to nearly circular, glabrous, apex rounded; extrastaminal disc a low, weakly 5-lobed rim c. 3-5 mm high bearing c. 200-250 spreading to recurved, filiform staminodes c. 8-12 mm in length; stamens c. 100, filaments slender, erect at anthesis, free, 20-22 mm long, glabrous, anthers ellipsoid, basifixed, reflexed; ovary narrowly ovoid, densely rusty brown tomentose, style c. 15 mm long, densely rusty brown tomentose at the base, glabrous above, stigma terminal, flattopped, glabrous. Involucre in fruit woody, dark brown, smooth on outer surface (in dry material), smooth and light brown on inner surface, globose to ellipsoid or ovoid, 2-2.7 cm tall, 2-2.5 cm diam., the wall c. 4-5 mm thick, with an apical opening c. 6-8 mm diam., the collar lined inside with the persistent bract-like processes; fruit globose to depressed globose, entirely contained within the involucre, c. 12-14 mm diam., 3- lobed, densely brownish orange wooly tomentose, also with scattered erect, white strigose trichomes, apex forming a distinct conical rostrum. — Fig. 3 .

Xyloolaena speciosa is a large shrub or tree whose flowers are reported to be salmon pink. Along with X. sambiranensis , which appears to be its closest relative, this species has narrowly ovate to oblong leaves, although their length rarely exceeds 2.3 times the width (in X. sambiranensis they are almost always at least 2.5 times as long as wide). Xyloolaena speciosa can be further distinguished by having only 9-10(-12) pairs of secondary leaf veins, persistent inflorescence bracts, and an extrastaminal disc that forms a low rim with c. 200-250 staminodes. It is narrowly distributed in an area with a dry to subhumid bioclimate extending from the Ambato Peninsula and the lower Ifasy river valley N to near Ambilobe (Fig. 1).

Humbert & Capuron 25590 was collected at the same locality and on the same day as the type, and is virtually identical in all aspects, suggesting that the specimens of both numbers were taken from the same individual.

VERNACULAR NAME. — Marabanty.

CONSERVATION STATUS. — With no sub-populations recorded from within the current protected areas network, and thus a predicted population decline of> 80% in the next three generations, Xyloolaena speciosa is assigned a preliminary status of Critically Endangered (CR).

PARATYPES. — MADAGASCAR, Prov. Antsiranana: Antilahimena et al. 321, Ambato FC, 13°26’42”S, 48°33’18”E, 21 Nov. 1996, fr. ( MO, P, TEF) ; Bosser 20243, route Ambilobe-Ambanja, PK 20, [13°18’S, 48°54’E], forêt sempervirente, 18 Apr. 1970, fl., y.fr. ( P) ; Humbert & Capuron 25590, collines gréseuses au SW d’Ambilobe dans les vallonnements et près de ruisseaux temporaires, [13°16’S, 49°01’E], 10 m, 12 Mar. 1951, fl., fr. ( MO, P) ; Humbert & Capuron 25873, vallée de l’Ifasy en aval d’Anaborano, vers Irary (Dist. d’Ambilobe), grès du Karroo et alluvions, [13°34’S, 48°49’E], 50-200 m, 31 Mar. 1951, fl. ( P) ; Perrier de la Bâthie 3020, schistes liasiques de la presqu’île d’Ambato, [13°26’S, 48°33’E], Jan. 1909, ster. ( P) ;

Revision of Xyloolaena (Sarcolaenaceae)

Randrianaivo et al. 236, Bevoay, Ambato FC, 40 km N d’Ambanja , 13°27’43”S, 48°32’33’E, 50-60 m, 16 May 1998, fr. (K, G, MO, TEF, P); Service Forestier 23426, presqu’île d’Ambato, [13°26’S, 48°33’E], 16 Mar. 1964, fl. [ MO, P (2 sheets)]; Ursch 279, forêt à l’est d’Ambilobe, [13°12’S, 49°05’E], 30- 200 m, Jan. 1927, fr. ( P) .

Figures

Fig. 2. — Xyloolaena sambiranensis: A, flowering branch; B, leaf (adaxial surface); C, leaf (abaxial surface); D, flower (cross sectional view); E, portion of extrastaminal disc; F, flower from below; G, inner sepal; H, fruiting involucre (mature and immature), I, cross section of fruiting involucre. (A, D, E, G, Schatz 3222; B-C, H, Gautier 2984; F, Perrier de la Bâthie 15052).

Fig. 3. — Xyloolaena speciosa: A, flowering branch; B, leaf (adaxial surface); C, leaf (abaxial surface); D, flower near anthesis; E, flower (cross sectional view); F, flower from below; G, inner sepal; H, fruiting involucre. (A-C, F-G, Service Forestier 3051; D-E, Service Forestier 23426; H, Antilahimena et al. 321).