Xyloolaena richardii Asteropeia multiflora Thouars (Asteropeiaceae) X. richardii X. sambiranensis X. speciosa Scleroolaena richardii Endemic families of Madagascar. VIII. A synoptic revision of Xyloolaena Baill. (Sarcolaenaceae) Lowry Ii, Porter P. Schatz, George E. Wolf, Anne-Elizabeth Adansonia 2002 3 24 1 7 19 (Baill.) Baill. [180,558,1397,1424] Magnoliopsida Sarcolaenaceae Xyloolaena Plantae Huerteales 5 12 Tracheophyta species richardii   Bull. Mens. Soc. Linn. Paris 1: 412 (1884). —  Scleroolaena richardiiBaill., Hist. Pl.4: 224 (1872), “richardi”. — Type:  Richards.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 mtall, ranging from sea level to over 1,500 melevation 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 Toamasinato 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 multifloraThouars (Asteropeiaceae)(see SCHATZ et al. 1999b). Previous authors (e.g., CAVACO 1952a,b; CAPURON 1970) have defined  X. richardiibroadly to include material placed here in  X. sambiranensisand  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 cmtall and 2.6-3.8 cmin diameter. The type material of  Scleroolaena richardiiconsists 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 richardiiis regarded as a species of Least Concern (LC).  MATERIAL EXAMINED. — Baron 5799, 6593, without precise locality; Bernard 201, Masoala PN;  Humbert18052, 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;  Schatzet 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 sambiranensisLowry & G.E. Schatz,  sp. nov.  Frutex vel arbor 3-12 malta. 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 acutaacuminatave 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 mmaltis unaquaque staminodia c. 25 gerente constante. Fructus involucro 2.1- 3.2(-3.5) cm alto, 2.2-2.5 cmdiam. subtentus.  TYPUS. —  Schatz3222, Madagascar, Prov. Antsiranana, Manongarivo RS, 30 kmSE 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 mtall. 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 mmlong, 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 mmlong, 2-3 mmdiam., 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 mmlong, 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 mmhigh, each bearing c. 25 filiform staminodes c. 3-12 mmin 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 mmlong, glabrous, anthers ellipsoid to ovoid, 1-1.5 mmlong, basifixed, reflexed; ovary cylindrical, densely whitish to light brownish woolly tomentose, style c. 15 mmlong, 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  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,  Schatz3222; B-C, H, Gautier 2984; F, Perrier de la Bâthie 15052). with vertical striations on inner surface, globose to ellipsoid or ovoid, 2.1-3.2(-3.5) cm tall, 2.2-2.5 cmdiam., the wall c. 3-4 mmthick, with an apical opening c. 7-10 mmdiam., the collar lined inside with the persistent bract-like processes; fruit globose, entirely contained within the involucre, c. 10-12 mmdiam., 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 mmlong, c. 2-3 mmdiam. at the base, densely brown to orange hirsute, tapering to the glabrous upper portion. — Fig. 2.   Xyloolaena sambiranensisis a shrub to tree 3- 12 mtall. 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 sambiranensishas an area of occupancy < 2,000 km 2and 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:  Derleth160, Manongarivo RS, Besinkara, chemin d’Ambodisakoanaà Ambalafary, sur le plateau, 14°04’S, 48°17’E,  300 m,  3 Nov. 1994, fr. (G, MO, P, TAN, TEF);   Gautier2984, Manongarivo RS, Besinkara, chemin d’Ambodisakoanaà 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);   Malcomberet 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 Bâthie15052, bassin de l’Ambazona[=Ambazoana], [ 13°35’S, 48°38’E], bud, fr. ( P);   Perrier de la Bâthie15424, same locality, grès liasiques,  Feb. 1923, bud, fl. [K, P(2 sheets)];   Saboureau60, chaîne de Bekaka, Ambanja, [ 13°44’S, 48°30’E],  150 m, fl. ( P);   Service Forestier2956, Antentezanambo, [ 13°49’S, 48°06’E],  16-30 Mar. 1951, fl., y.fr. ( P, TAN, TEF);   Service Forestier3161, Bekaka, Ambanja, [ 13°44’S, 48°30’E],  20 Mar. 1950, ster. ( P);   Service Forestier7734, Bandrakorony, Ambanja, [ 13°49’S, 47°52’E],  6 Oct. 1953, fr. ( P, TEF);   Service Forestier9286, Benavony, Ambanja, [ 13°41’30”S, 48°29’E],  22 Mar. 1954, bud, fr. ( P, TEF);   Service Forestier10411, route de Maromandiaà Ambanja,  10 Aug. 1954, fr. ( P);   Service Forestier10451, rivière Antsahakolana, dist. d’Ambanja, Manongarivo RS, [ 14°04’30”S, 48°16’E],  360 m,  19 July 1954, fr. ( P, TEF);   Totozafy Be555, Manongarivo RS, Besinkara, bord de chemin, 14°04’S, 48°17’E,  4 June 1996, fr. (G, MO, P).    Prov. Mahajunga:  Decary2130, Maromandia, terrain gréseux, [ 14°12’S, 48°05’E],  15 May 1923, fl. [ P(2 sheets)];   Perrier de la Bâthie3025, collines gréseuses des environs de Maromandia, [ 14°12’S, 48°05’E],  Mar. 1909, fl. [ P(2 sheets)];   Service Forestier19396, Amparihibe, canton Maromandia, colline sur sol gréseux, [ 14°18’30”S, 48°07’E],  9 Jan. 1959, bud ( P, TEF).  5.  Xyloolaena speciosaLowry & G.E. Schatz,  sp. nov.  Frutex vel arbor usque ad 20 malta. Folia lamina ellipticausque 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 mmalto staminodia 200 ad 250 gerente constante. Fructus involucro 2.0- 2.7 cmalto, 2.0- 2.5 cmdiam. 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 mtall. 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 mmlong, 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 mmlong, 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 mmlong, 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 mmlong, 2-3 mmdiam., 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 mmlong, 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 mmhigh bearing c. 200-250 spreading to recurved, filiform staminodes c. 8-12 mmin length; stamens c. 100, filaments slender, erect at anthesis, free, 20-22 mmlong, glabrous, anthers ellipsoid, basifixed, reflexed; ovary narrowly ovoid, densely rusty brown tomentose, style c. 15 mmlong, 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 cmtall, 2-2.5 cmdiam., the wall c. 4-5 mmthick, with an apical opening c. 6-8 mmdiam., the collar lined inside with the persistent bract-like processes; fruit globose to depressed globose, entirely contained within the involucre, c. 12-14 mmdiam., 3- lobed, densely brownish orange wooly tomentose, also with scattered erect, white strigose trichomes, apex forming a distinct conical rostrum. — Fig. 3.   Xyloolaena speciosais 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. sambiranensisthey are almost always at least 2.5 times as long as wide).  Xyloolaena speciosacan 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 25590was 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 speciosais assigned a preliminary status of Critically Endangered (CR).   PARATYPES. — MADAGASCAR,  Prov. Antsiranana:  Antilahimenaet al. 321, Ambato FC, 13°26’42”S, 48°33’18”E,  21 Nov. 1996, fr. ( MO, P, TEF);   Bosser20243, route Ambilobe-Ambanja, PK20, [ 13°18’S, 48°54’E], forêt sempervirente,  18 Apr. 1970, fl., y.fr. ( P);   Humbert& Capuron25590, collines gréseuses au SW d’Ambilobedans les vallonnements et près de ruisseaux temporaires, [ 13°16’S, 49°01’E],  10 m,  12 Mar. 1951, fl., fr. ( MO, P);   Humbert& Capuron25873, vallée de l’Ifasyen aval d’Anaborano, vers Irary( Dist. d’Ambilobe), grès du Karrooet alluvions, [ 13°34’S, 48°49’E],  50-200 m,  31 Mar. 1951, fl. ( P);   Perrier de la Bâthie3020, schistes liasiques de la presqu’île d’Ambato, [ 13°26’S, 48°33’E],  Jan. 1909, ster. ( P); Revision of  Xyloolaena(Sarcolaenaceae)  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).  Randrianaivo et al. 236, Bevoay, Ambato FC, 40 kmN 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). P Madagascar -15.266666 Angonsti 1285 50.483334 Richard 5 12 1 Antsiranana 1991-03-23 1991-03-25 1991-03-23 MO, P, TEF Manongarivo RS Madagascar 200 -14.133333 Ankaramy 1289 48.35 6 13 1 Antsiranana 1994-11-03 MO, P, TAN, TEF Manongarivo RS & Besinkara Madagascar Prov. 300 -14.066667 d'Ambodisakoana 1289 48.283333 Derleth 8 15 1 Antsiranana paratype 1996-04-04 MO, P, TAN, TEF Manongarivo RS Madagascar Gautier 200 -14.066667 d'Ambodisakoana 1289 48.283333 Besinkara 8 15 1 paratype 1992-11-15 MO, P Madagascar 100 -13.966666 Malcomber 1289 48.166668 8 15 1 paratype P Perrier de la Bathie & de l'Ambazona Madagascar -13.583333 1290 48.633335 8 15 1 paratype 1923-02 P Perrier de la Bathie Madagascar -13.583333 1290 48.633335 8 15 1 paratype P de Bekaka Madagascar 150 -13.733334 Ambanja 1290 48.5 Saboureau 8 15 1 paratype 1951-03-16 1951-03-30 1951-03-16 P, TAN, TEF Madagascar -13.816667 Antentezanambo 1289 48.1 Service Forestier 8 15 1 paratype 1950-03-20 P Madagascar Service Forestier -13.733334 Ambanja 1290 48.5 Bekaka 8 15 1 paratype 1953-10-06 P, TEF Madagascar Service Forestier -13.816667 Ambanja 1289 47.866665 Bandrakorony 8 15 1 paratype 1954-03-22 P, TEF Madagascar Service Forestier -13.691667 Ambanja 899 48.483334 Benavony 8 15 1 paratype 1954-08-10 P de Maromandia Madagascar Ambanja Service Forestier 8 15 1 paratype 1954-07-19 P, TEF Manongarivo RS Madagascar Service Forestier 360 -14.075 d'Ambanja 897 48.266666 Antsahakolana 8 15 1 paratype 1996-06-04 MO, P Manongarivo RS Madagascar -14.066667 Besinkara 1289 48.283333 Totozafy Be 8 15 1 paratype 1923-05-15 P Madagascar Prov. -14.2 Maromandia 1289 48.083332 Decary 8 15 1 Mahajunga paratype 1909-03 P de Maromandia Madagascar -14.2 Perrier de la Bathie 1289 48.083332 8 15 1 paratype 1959-01-09 P, TEF Madagascar Service Forestier -14.308333 Maromandia 896 48.116665 Amparihibe 8 15 1 paratype 1951-03-12 P, MO, TEF du Levika & de Matsoborilava Madagascar -13.266666 Antsiranana 1291 49.016666 8 15 1 Antsiranana 1996-11-21 MO, P, TEF Antilahimena & Ambato FC Madagascar -13.445 Antsiranana 21 48.555 9 16 1 Antsiranana paratype 1970-04-18 P Bosser & Ambilobe-Ambanja, PK Madagascar -13.3 1291 48.9 9 16 1 Prov. paratype 1951-03-12 MO, P Capuron & SW d'Ambilobe Madagascar 10 -13.266666 1291 49.016666 9 16 1 Prov. paratype 1951-03-31 P Capuron & de l'Ifasy & d'Anaborano & Irary & Dist. d'Ambilobe & du Karroo Madagascar 125 -13.566667 1290 48.816666 9 16 1 Prov. paratype 1909-01 P Perrier de la Bathie & d'Ambato Madagascar -13.433333 1291 48.55 9 16 1 Prov. paratype 1927-01 1998-05-16 1927-01 MO, TEF, P Ambato FC 55 -13.2 d'Ambanja 1291 49.083332 11 18 1