taxonID	type	description	language	source
6D7A87BA6533FFF8FD2D1A642771FB23.taxon	materials_examined	TYPE. — Leptolaena multiflora Thouars.	en	Schatz, George E., Lowry Ii, Porter P., Wolf, Anne-Elizabeth (2001): Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae). Adansonia (3) 23 (2): 171-189, DOI: 10.5281/zenodo.4605057
6D7A87BA6537FFF4FD2D1AFA22C7FD2C.taxon	description	J. Linn. Soc. Bot. 25: 296 (1889). — Leptolaena multiflora var. cuspidata (Baker) H. Perrier, Bull. Soc. Bot. France 78: 60 (1931). — Type: Baron 5835 [cited in protologue as “ next 5836 ”], Madagascar, chiefly from north-west Madagascar, fr. (holo-, K!; iso-, P!). Leptolaena cuspidata is a shrub to small tree that occurs in the subhumid forest of the Sambirano region in NW Madagascar, below 500 m elevation (Fig. 3). It can be recognized by its distinctive densely golden yellow granular farinose involucre and lanceolate to ovate, longcuspidate leaves. 58 km N of Ambanja; Hildebrandt 3306, Vavatobe; Humbert 18752, Ambanja; Humbert 19041, Ambodimagodro; Humbert 25580, SW of Ambilobe; Humbert 32649, Ankarana RS; Humbert 32873, Ambohipiraka; Perrier de la Bâthie 3021, Ambanja; Perrier de la Bâthie 3023, Maromandia; Perrier de la Bâthie 5340, Ambohipiraka; Perrier de la Bâthie 15677, bas Sambirano; Phillipson 2017, Beramanja; Ravololonanahary 37, Ambato; Service Forestier 3078, Irano (Irarona) to Beramanja; Service Forestier 3141, 3142, Andilamboay; Service Forestier 3171, Tsaratanana RNI; Service Forestier 7501, Bekaka to Benavony; Service Forestier 8225, Anjahakely (= Ampahakely); Service Forestier 10434, Ambilobe; Service Forestier 11486, Manongarivo RS; Service Forestier 12987, Anjakely; Stiefel 68, Manongarivo RS; Totozafy Be 563, Manongarivo RS. 3. Leptolaena delphinensis G. E. Schatz & Lowry, sp. nov. Haec species a congeneris ad Leptoaenam sensu stricto pertinentibus foliis glabris apice obtusis usque rotundatis, involucro urceolato granulari dense ferrugineo c. 3 mm alto atque sepalis involucrum c. 1.5 mm excedentibus distinguitur. TYPUS. — McPherson, Dumetz & Rabevohitra 14222, Madagascar, Prov. Toliara, North of Fort Dauphin near Ste. Luce (Manafiafy), NW of village in forest remnant on sand, 24 ° 47 ’ S, 47 ° 10 ’ E, 20 m, 22 Oct. 1989, fl. (holo-, MO!; iso-, P!, TAN!, TEF!). Fig. 3. — Distributions of Leptolaena, mapped on the bioclima- Shrubs to trees 3 - 10 m tall, trunk to 20 cm tic zones of Madagascar (after CORNET 1974; see SCHATZ 2000). L. cuspidata (Δ), L. pauciflora (•). in diam., stems glabrous. Leaves chartaceous, narrowly to broadly elliptic, 1.8 - 5.2 × 0.8 - 3.2 cm, reduced in the inflorescence, glabrous, VERNACULAR NAMES. — Zahana, Zahéna, base obtuse to rounded, margins slightly to Zana. strongly revolute, apex rounded to occasionally CONSERVATION STATUS. — With an extent of obtuse, venation obscurely brochidodromous, occurrence <20,000 km 2, an area of occupancy secondary veins (8 -) 11 - 13 per side, midrib <2,000 km 2, and only three of 14 sub-popula- flat above, raised below; petiole 3 mm long, tions encompassed within protected areas, L. cus- c. 1 mm diam., canaliculate, glabrous, stipules pidata is assigned a preliminary status of indistinct, caducous. Inflorescences terminal, Vulnerable (VU). corymbose, 10 - 50 - flowered, the axes glabrous; peduncle 0.7 - 0.8 mm long, involucre urceolate,	en	Schatz, George E., Lowry Ii, Porter P., Wolf, Anne-Elizabeth (2001): Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae). Adansonia (3) 23 (2): 171-189, DOI: 10.5281/zenodo.4605057
6D7A87BA6537FFF4FD2D1AFA22C7FD2C.taxon	materials_examined	MATERIAL EXAMINED. — Baron 5836, 6283, with- 3 mm tall, 2.5 mm diam. at apex, densely granout precise locality; Baum 333, Ambodimanga; ular ferruginous farinose, apical margin with 8 - Birkinshaw 1, Antsatsaka; Bosser 20152, Ampasindava; Cours 5674, Mt. Ambohiparaka; Decary 938, 1165, 9 subequal triangular teeth, 0.5 mm tall, 2095, Maromandia; Decary 14775, 14794, Ambilobe; 0.4 mm broad at base; sepals exserted 1 - 1.5 mm Derleth 89, Manongarivo RS; Gentry 11870, 57 - beyond involucre, densely white hirsute outside, glabrous to sparsely hirsute inside; petals narrowly elliptic, 7 × 2.5 mm, white fading to yellow, exserted 6 mm beyond involucre at anthesis; stamens 10, free to the base, filaments c. 7 - 8 mm long, anthers 0.8 mm tall; ovary conic-ovoid, densely whitish tomentose, style slender, exserted, c. 7 - 8 mm long, stigma hemispheric, with 3 weakly differentiated lobes, c. 0.8 mm across. Fruit globose to widely ovoid, somewhat asymmetric, with a longitudinal channel and concave apex when dry, c. 3 - 4 mm tall, 2 - 3 mm diam., glabrous, entirely enclosed within the persistent sepals and involucre. — Fig. 4. Leptolaena delphinensis is a shrub to small tree, and is restricted to humid littoral forest on sand near Ft. Dauphin in SE Madagascar (Fig. 2). It can be distinguished by its leaves with an obtuse to rounded apex, small, urceolate, densely granular ferruginous involucre 3 mm tall, and sepals that are exserted 1 - 1.5 mm beyond the involucre.	en	Schatz, George E., Lowry Ii, Porter P., Wolf, Anne-Elizabeth (2001): Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae). Adansonia (3) 23 (2): 171-189, DOI: 10.5281/zenodo.4605057
6D7A87BA6537FFF4FD2D1AFA22C7FD2C.taxon	description	VERNACULAR NAMES. — Fotobavy, Fotoda, Fotondahy, Fonto, Fotona. CONSERVATION STATUS. — With no sub-populations recorded from within the current protected areas network, and thus a predicted population reduction of> 80 % in the next three generations, L. delphinensis is assigned a preliminary status of Critically Endangered (CR). Along with L. raymondii, the status of L. delphinensis demonstrates the urgent need for additional protection of littoral forest along the East coast of Madagascar. PARATYPES. — MADAGASCAR, Prov. Toliara: Decary 10064, Vinanibe, [25 ° 03 ’ S, 46 ° 56 ’ E], 10 July 1932, fl. (P); Dumetz et al. 683, Mandena STF, 24 ° 57 ’ S, 47 ° 00 ’ E, 0 - 10 m, 17 Apr. 1989, bud (K, MO, P, TAN, TEF); Humbert 5955, environs de Fort Dauphin, sables entre le pic St. Louis et la mer, [25 ° 03 ’ S, 47 ° 00 ’ E], 20 Sep. - 6 Oct. 1928, fl. (MO, P); Jacquemin 1149, Mandena STF, [24 ° 57 ’ S, 47 ° 00 ’ E], 8 Dec. 1972, fr. (MO, P); Johnson WII 37, Mandena STF, [24 ° 57 ’ S, 47 ° 00 ’ E], 16 July 1963, bud (P); McPherson et al. 14142, Mandena STF, low forest on sand, 24 ° 57 ’ S, 47 ° 00 ’ E, 25 m, 17 Oct. 1989, fl. (K, MO, P, TAN, TEF); McPherson & Dumetz 14665, Mandena STF, 24 ° 57 ’ S, 47 ° 00 ’ E, 25 m, 7 Dec. 1989, fr. (MO, TAN); McPherson et al. 14835, Ste. Luce (Manafiafy), 24 ° 47 ’ S, 47 ° 10 ’ E, 20 m, 18 Jan. 1990, fr. (MO, TAN); Rabenantoandro et al. 272, Mandena STF, 24 ° 57 ’ 11 ” S, 47 ° 00 ’ 16 ” E, 5 m, 27 Sep. 2000, fl. (G, K, MO, P, TEF); Rabenantoandro et al. 308, Mandena STF, 24 ° 55 ’ 20 ” S 47 ° 02 ’ 23 ” E, 0 - 50 m, 4 Oct. 2000, fl. (G, K, MO, P, TEF); Rabevohitra 2053, Ste. Luce (Manafiafy), forêt au N. du village, 24 ° 46 ’ S, 47 ° 09 ’ E, 10 m, 20 Oct. 1989, fl. (MO, P, TAN, TEF); Rabevohitra & Rabenantoandro 3701, Mandena STF, 24 ° 57 ’ 03 ” S 47 ° 00 ’ 03 ” E, 5 m, 4 Oct. 2000, fl. (G, K, MO, P, TEF); Service Forestier 399, M a n d e n a S T F, f o r ê t l i t t o r a l e, [24 ° 57 ’ S, 47 ° 00 ’ E], 16 Feb. 1949, fr. (P, TEF); Service Forestier 8207, same locality, 24 July 1953, fl. (P, TEF); Service Forestier 14597, same locality, 1 Sep. 1955, fl. (P); Service Forestier 28656, vestiges forestiers, sur dunes anciennes, sur la route de Ste. Luce, [24 ° 47 ’ S, 47 ° 11 ’ E], 11 Dec. 1968, fr. (P, TEF); Zarucchi et al. 7504, Mandena STF, coastal low forest on sand, 24 ° 57 ’ S, 47 ° 02 ’ E, 10 m, 26 May 1991, bud (K, MO, P, TAN). 4. Leptolaena gautieri G. E. Schatz & Lowry, sp. nov. Haec species a congeneris ad Leptoaenam sensu stricto pertinentibus foliis ovatis vel late ovatis apice acutis usque acuminatis secus costam subtus indumento manifeste vestitis, involucro granulari-farinoso dense ferrugineo plerumque trichomatibus suberectis albidis c. 0.5 mm longis vestito atque petalis 10.5 - 12 mm longis distinguitur. TYPUS. — Gautier 2920, Madagascar, Prov. Antsiranana, Manongarivo RS, Bekolosy, 14 ° 02 ’ S, 48 ° 18 ’ E, 1240 m, 29 Mar. 1996, fl. (holo-, MO!; iso-, G, P!, TAN!, TEF!). Trees 3 - 15 m tall, stems initially hirsutetomentose, then glabrescent. Leaves chartaceous, ovate to broadly ovate, 1.1 - 4.8 × 0.5 - 3.7 cm, reduced in the inflorescence, blade glabrous, base obtuse to rounded or subcordate, apex acute to acuminate, the very tip rounded, occasionally somewhat falcate, venation obscurely brochidodromous, secondary veins 9 - 14 per side, midrib flat to slightly raised above, sparsely to moderately hirsute-tomentose, raised below with sparsely to moderately dense sericeous to hirsute indumentum; petiole 2.5 - 4 mm long, 1 mm in diam., canaliculate, moderately to densely hirsute, stipules indistinct, caducous. Inflorescences terminal and axillary from the uppermost leaves, corymbose, 7 - 30 - flowered, the axes densely grey hirsute-tomentose; peduncle 0.1 - 0.5 mm long, indistinct; involucre urceolate, 3 - 3.5 mm tall, 2.5 - 3 mm diam. at apex, tapering evenly to a cuneate base, densely granular ferruginous farinose, usually also with sparse to moderately dense whitish golden partially erect hairs to 0.5 mm, especially towards the base, apical margin with 8 - 10 subequal triangular teeth, 1 mm tall, 0.5 - 1 mm broad at base; sepals exserted 1.5 mm beyond involucre, densely whitish golden hirsute outside, glabrous inside; petals narrowly oblong to oblanceolate, 10.5 - 12 × 1.5 - 2 mm, white to pale yellow, with evident venation upon drying, exserted 7 - 9 mm beyond involucre teeth at anthesis, the apex rounded; stamens 10 - 12, free to the base, filaments 6 - 8 mm long, anthers 0.5 - 0.9 mm tall; ovary ovoid, densely light brown tomentose, style slender, exserted, 6 - 8 mm long, stigma obtriangular to hemispheric, 1 mm in diam. Fruit globose, somewhat asymmetric, with a longitudinal channel and concave apex when dry, c. 2.5 mm tall, 2.5 mm diam., glabrous, entirely enclosed within the persistent sepals and involucre. — Fig. 5. Leptolaena gautieri is a small to medium-sized tree that occurs in mid-elevation subhumid forest from 800 m to over 1,200 m (Fig. 6). It can be CONSERVATION STATUS. — With an area of occupancy <2,000 km 2, and only four of eight sub-populations encompassed within the protected areas network, L. gautieri is assigned a preliminary status of Vulnerable (VU). PARATYPES. — MADAGASCAR, Prov. Antsiranana: Gautier & Chatelain 2344, Manongarivo RS, Bekolosy, vallon en amont de la chute, 14 ° 02 ’ S, 48 ° 18 ’ E, 1100 m, 15 June 1994, fl. (G, TAN); Gautier 2361, Manongarivo RS, Bekolosy, 14 ° 02 ’ S, 48 ° 18 ’ E, 800 m, 15 June 1994, fl. (G, MO, P, TAN); Gautier 3510, Manongarivo RS, Ambahatra, cours supérieur, 13 ° 59 ’ S, 48 ° 26 ’ E, 1210 m, 8 Mar. 1999 (G, MO); Humbert & Capuron 24147, Mt. Anjenabe, vallée inférieure de l’Androranga, affluent de la Bemarivo aux environs d’Antongondriha, [14 ° 18 ’ S, 49 ° 45 ’ E], 1130 m, 3 - 7 Nov. 1950, fr. (P); Malcomber & Rakotomalala 2608, Manongarvio RS, Bekolosy, 14 ° 02 ’ S, 48 ° 18 ’ E, 1000 - 1200 m, 4 Dec. 1993, fr. (MO, P, TAN); McPherson & van der Werff 16359, Manongarivo RS, above the vil- lage of Ambodisakoana, 14 ° 05 ’ S, 48 ° 20 ’ E, 1100 m, 16 Oct. 1994, fr. (MO, P, TAN); McPherson & van der Werff 16396, same locality, 1100 - 1200 m, 18 Oct. 1994, bud (MO, P, TAN); Rakotomalaza 48, Manongarivo RS, Bekolosy, 14 ° 04 ’ S, 48 ° 17 ’ E, 1000 - 1480 m, Apr. 1993, fl. (MO, P, TAN); Rakotozafy 336, plateau de Beankany, Ambanja, 12 Dec. 1963, bud (P, TAN); A. Randrianasolo 307, Marojejy RNI, 14 ° 29 ’ S, 49 ° 49 ’ E, 300 - 800 m, 21 Apr. 1993, fl. (MO, P, TAN); A. Randrianasolo 325, same locality, 500 - 600 m, 22 Apr. 1993, fl. (MO, P, TAN); Réserves Naturelles 2237, Tsaratanana RNI, Ambavala, Marovato, Ambanja, 26 Sep. 1950, fr. (MO, P, TAN); Schatz 3233, Manongarivo RS, Bekolosy, 14 ° 06 ’ S, 48 ° 17 ’ E, 1000 m, 28 Mar. 1991, fl. (K, MO, P, TAN). Prov. Fig. 6. — Distributions of Leptolaena, mapped on the bioclima- Toamasina: Labat et al. 3068, Moramanga, Andasibe, tic zones of Madagascar (after CORNET 1974; see SCHATZ 2000). Forêt de Maromizaha, 18 ° 57 ’ 56 ” S, 48 ° 27 ’ 34 ” E, L. masoalensis (Δ), L. gautieri (•), L. raymondii (*). 1020 m, 1 Apr. 1999, fr. (BR, G, K, P, MO, TAN); McPherson 17467, Ambatovy, 18 ° 51 ’ 12 ” S, 48 ° 18 ’ 48 ” E, 1100 m, 26 Feb. 1998, bud (MO, P, recognized by its ovate to broadly ovate leaves TAN); Perrier de la Bâthie 14887, Analamazaotrawith an acute to acuminate apex and evident Périnet Rakotomalaza RS, [18 ° 56 1046 ’ S ,, 48 Ambatovy ° 26 ’ E], Oct,. 18 1920 ° 51, ’ fl 07. (” P S);, indumentum on the midvein below, its densely 48 ° 18 ’ 47 ” E, 1100 m, 30 Jan. 1997, bud (MO, P, ferruginous granular farinose involucre that usu- TAN); Rakotomalaza 1279, same locality, 18 ° 51 ’ 08 ” S, ally also has slightly erect whitish trichomes 48 ° 18 ’ 59 ” E, 990 m, 25 Mar. 1997, bud (MO, P, 0.5 mm long, and petals that are 10.5 - 12 mm TAN); Rakotomalaza 1286, same locality, 18 ° 51 ’ 28 ” S, 48 ° 17 ’ 43 ” E, 26 Mar. 1997, bud (MO, P, TAN); long. Rakotomalaza 1309, same locality, 18 ° 51 ’ 10 ” S, 48 ° 18 ’ 49 ” E, 1050 m, 13 May 1997, fl. (MO, P, TAN); ETYMOLOGY. — The species epithet honors Randriambololona et al. 17, Mantady PN, 18 ° 56 ’ S, Laurent GAUTIER, our friend and colleague at the 48 ° 26 ’ E, 960 m, 27 Oct. 1997, fr. (MO, P, TAN); herbarium in Geneva, who has significantly Service Forestier 10341, Mangorokely, Anosibe, Moramanga, [19 ° 26 ’ S, 48 ° 12 ’ E], 19 May 1954, fl. (P). advanced our knowledge of the flora of the Prov. Toliara: Humbert 20704, Mt. Vohimavo au Nord Manongarivo Special Reserve. d’Ampasimena, bassin de la Manampanihy, [24 ° 19 ’ S, VERNACULAR NAMES. — Anjananjana, Zahana. 47 ° 08 ’ E], 830 m, 27 - 28 Mar. 1947, fl. (MO, P). 5. Leptolaena masoalensis G. E. Schatz & Lowry, sp. nov. Haec species a congeneris ad Leptoaenam sensu stricto pertinentibus foliis glabris apice obtusis usque rotundatis, involucro ellipsoideo plerumque glabro interdum sparse albido-hirsuto c. 7 mm alto sepalos omnino includente atque staminibus tantum 6 ad 8 distinguitur. TYPUS. — Rahajasoa, Rabe & Bernard 790, Madagascar, Prov. Antsiranana, Masoala PN, Réserve Naturelle d’Ambato, 15 ° 17 ’ S, 50 ° 21 ’ E, 380 m, 8 Oct. 1994, fl. (holo-, MO!; iso-, K!, P!, TAN!). Shrubs to 2 m tall; stems glabrous to sparsely pubescent. Leaves subcoriaceous, elliptic to slightly obovate, 2.4 - 4 × 1.1 - 1.7 cm, glabrous, discolorous when dry, dark olive brown above, lighter olive green below, base cuneate to acute, margins strongly revolute, apex rounded, venation weakly brochidodromous, secondary veins c. 8 per side, midrib flat above, raised below with very sparse scattered trichomes or glabrous; peti- ole 2 - 3 mm long, 0.6 - 0.8 mm diam., shallowly canaliculate, stipules caducous. Inflorescences axillary in the terminal leaves, 2 - 8 - flowered, axes probably on similar substrates (Fig. 2). In the sparsely to densely puberulous; peduncle 1 - 2 mm past, this species has been broadly circumscribed long, distinct, involucre ellipsoid to oblong, to include all of the new taxa described here, as glabrous, 7 mm tall, 3 mm diam., apical margin well as the Sambirano endemic L. cuspidata. As with 8 narrowly triangular, subequal teeth 1 mm defined here, L. multiflora can be distinguished tall, 1 mm broad at base, glabrous or sometimes by its glabrous, elliptic, usually almost equilateral sparsely short whitish hirsute; sepals completely leaves, the largest of which usually exceed 5.5 cm included within the involucre, 4 × 4 mm, circular, in length and dry an olive green to brownish strongly inrolled, densely whitish hirsute outside, color. In addition, the sepals at anthesis exceed glabrous inside; petals narrowly oblong, 12 × the involucre teeth by (0 - 8 -) 1 - 1.8 mm, a feature 3 mm, clear yellow with evident venation upon that separates L. multiflora from L. abrahamii. drying, glabrous, exserted 5 - 6 mm beyond involucre; stamens 6 - 8, free to the base, filament 8 - V E R N AC U L A R N A M E S. — Amaninombi- 10.5 mm long, anthers unknown; ovary ellipsoid, lahimena, Amaninombilahy, Amaninombilahy à 1 mm diam., densely whitish golden hirsute- petites feuilles, Anjananjana, Birana, Dilatra, strigose, the trichomes to 2 mm long, style Dilatry Lahy, Fotona, Fotona Beravina, Fotonala, 11.5 mm long, slender, exserted 4 - 5 mm above Hazoambo, Hazomasy, Helana, Jana, involucre, stigma capitate, 1 mm in diam. Fruit Manilombilahy, Manina ombilahy, Sarifatra, unknown. — Fig. 7. Tsilaitra à petites feuilles. CONSERVATION STATUS. — Despite an extent Leptolaena masoalensis is known only from the of occurrence and area of occupancy far in excess type collection made at Ambato, a hill in the of 20,000 km 2 and 2,000 km 2 respectively, with northeastern part of the Masoala Peninsula (Fig. 6) only four of c. 19 sub-populations occurring in well known for its large population of Nepenthes protected areas, and thus indicating a possible masoalensis Schmid-Hollinger. At first glance, population reduction in the next three genera- L. masoalensis may be mistaken for L. delphinensis, tions of> 50 %, L. multiflora is assigned a prelimwith which it shares leaves with an obtuse to inary status of Endangered (EN). rounded apex. However, L. masoalensis can be easily distinguished by its much larger, ellipsoid MATERIAL EXAMINED. — Baron 2632, Bojer s. n., to oblong involucre 7 mm tall, sepals that are Forbes s. n., Louvel 6, Thouars s. n., without precise completely included within the involucre at locality; Andrianjafy 66, Rantabe; Bernard 372, anthesis, and leaves with only 8 secondary veins Masoala PN; Birkinshaw 354, Mananara RB; Boivin s. n., Foulpointe; d’Alleizette 268, Nosy-Be (?); Decary per side versus (8 -) 11 - 12 in L. delphinensis. 5198, Ifandana; Humblot 349, Nosive; Perrier de la Bâthie 14186, Mahanoro; Rabe 157, Beankoraka; CONSERVATION STATUS. — With an area of oc- Rabenantoandro 563, Mahatsara STF; Rabevohitra cupancy <100 km 2, and only a single known pop- 3808, Mahabo; Rahajasoa 424, Masoala PN; ulation that appears to lie just outside the limit of Ilaka-Est Raholivelo, 37 458, Tampolo, Tampolo STF STF; A;. Réserves Randrianasolo Naturelles 292, Masoala National Park, L. masoalensis is assigned a 2746, 3403, 5514, 9113, 9124, 9310, Masoala RNI; preliminary status of Critically Endangered (CR). Richard s. n., Foulpointe; Richard 4, 40, Ste. Marie; Schedl 135, Ambila-Lemaitso STF; Service Forestier 4 - R- 7, Antetezana STF; Service Forestier 89 - R- 303, 6. Leptolaena multiflora Thouars Manampano Ampasinambo ;; Service Service Forestier Forestier 745 - R- 174 1, - Ambila- R- 116, Lemaitso STF; Service Forestier 1371, Andatsakala; Hist. Vég. Isles Austral. Afriq.: 41, t. 11 (1805). — Service Forestier 1616, Ambila-Lemaitso STF; Service Type: Thouars s. n., Madagascar, without precise local- Forestier 2414, Soanierano Ivongo; Service Forestier ity (P!). 3680, Tohakandra; Service Forestier 3809, Misevo; Service Forestier 4679, Antetezana STF; Service As here recognized, Leptolaena multiflora Forestier 4813, Ifanirea; Service Forestier 5107, Berano; occurs primarily in coastal forest mostly on sand, Service Forestier 5677, Foulpointe; Service Forestier with a few populations recorded farther inland, 6252, Rantabe; Service Forestier 7494, Fampotakely; Service Forestier 7510, Antetezana STF; Service Forestier 8901, Rantabe; Service Forestier 9501, Ampangalana Atsimo; Service Forestier 10053, 10076, Tampolo STF; Service Forestier 12401, Amporoforo; Service Forestier 12929, Manombo STF; Service Forestier 13906, Amporoforo; Service Forestier 14504, Ifanirea; Service Forestier 15215, Tampolo STF; Service Forestier 15302, Manombo RS; Service Forestier 15305, Berano; Service Forestier 15378, Andrazaha; Service Forestier 15493, Analavory; Service Forestier 16118, Ambazato; Service Forestier 16215, Evato; Service Forestier 16472, 17820, Tampolo STF; Service Forestier 18082, Vohibola; Service Forestier 19173, Tampolo STF; Service Forestier 19535, Ampangalana Atsimo; Service Forestier 21217, Ambila-Lemaitso STF; Service Forestier 21471, Ambohimary; Service Forestier 21493, Manombo STF; Service Forestier 29717, Lakato; Service Forestier 29860, Tampolo STF; Service Forestier 29892, Ambila-Lemaitso STF; Service Forestier 32831, 34550, Mahatsara; Service Forestier 34575, Sahamalaza; Service Forestier 35140, Mahatsara; Ursch 116, Tampina.	en	Schatz, George E., Lowry Ii, Porter P., Wolf, Anne-Elizabeth (2001): Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae). Adansonia (3) 23 (2): 171-189, DOI: 10.5281/zenodo.4605057
6D7A87BA653FFFE8FF701D0927FDFB4C.taxon	description	J. Linn. Soc. Bot. 20: 96, 97 (1883). — Lectotype (here designated): Baron 1390, Madagascar, in forests of the province of Imerina (K!; iso-, MO!, P (3 sheets)!). Leptolaena turbinata Baker, J. Linn. Soc. Bot. 20: 97 (1883). — Leptolaena pauciflora var. turbinata (Baker) H. Perrier, Bull. Soc. Bot. France 78: 60 (1931). — Type: Baron 1560, Madagascar, East coast (holo-, K!; iso-, P!). Leptolaena parviflora Scott-Elliot, J. Linn. Soc. Bot. 29: 6 (1891). — Lectotype (here designated): Scott- Elliot 2554, Madagascar, woods near Fort Dauphin (K!; iso-, P (2 sheets)!). Leptolaena rubella Scott-Elliot, J. Linn. Soc. Bot. 29: 7 (1891). — Leptolaena pauciflora var. rubella (Scott- Elliot) H. Perrier, Bull. Soc. Bot. France 78: 60 (1931). — Type: Scott-Elliot 2369, Madagascar, woods near Fort Dauphin (holo-, K!; iso-, P (2 sheets)!). Leptolaena pauciflora is a small shrub to medium-sized tree occurring from humid littoral forest at sea level to subhumid and montane woodland and thicket up to 2,300 m (Fig. 3). Other than habit, it is remarkably uniform in vegetative and reproductive characters throughout its broad ecological range, and is easily recognized by its small, broadly ovate to circular or rhombic leaves usually not exceeding 2 cm in length. Of the two syntypes cited in the protologue of L. pauciflora (Baron 1390 and Meller s. n.), the former is here chosen as the lectotype. Syntypes were also cited for L. parviflora (Scott-Elliot 2554 and Scott-Elliot 2713), the first of which is here chosen as the lectotype. VERNACULAR NAMES. — Amaninambilahy, Anjananjana, Anjananjanalahy, Fotina, Foto, Fotona, Fotona Madinidravina, Fotondahy, Hazomaritra, Laro, Madiorano, Milaliambomadinika, Taolandaro. CONSERVATION STATUS. — With an extent of occurrence far exceeding 20,000 km 2, and c. 33 sub-populations, six of which are present in protected areas, L. pauciflora is considered not threatened, and assigned a preliminary status of Lower Risk (LR).	en	Schatz, George E., Lowry Ii, Porter P., Wolf, Anne-Elizabeth (2001): Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae). Adansonia (3) 23 (2): 171-189, DOI: 10.5281/zenodo.4605057
6D7A87BA653FFFE8FF701D0927FDFB4C.taxon	materials_examined	MATERIAL EXAMINED. — Baron 1390, 1560, 2624, Meller s. n., without precise locality; Andrianjafy 11, Andranofeno-Sud; Barnett 502, Mt. Ibity; Benoist 759, Ambila-Lemaitso STF; Boiteau 2568, Mahialambo; Bosser 7944, Tampoketsa d’Ankazobe; Bosser 9969, Sahatsio; Bosser 14138, pic St. Louis; Bosser 15978, Tampoketsa; Bosser 18772, col des Tapias; Cours 2981, Ambila-Lemaitso STF; Cours 4646, Didy; Cremers 1624, 2020, Ambohitantely RS; Cremers 2311, Marosiky; Croat 29218, Antsirabe; Croat 29910, Itremo; Decary 4026, 4038, Fort-Dauphin; Decary 6353, 6388, 6502, 6508, Ambila-Lemaitso STF; Decary 7302, Ankazobe; Decary 7473, Ambohitantely RS; Decary 10011, pic Saint Louis; Decary 13063, 13226, Ambatofinandrahana; Decary 14023, 14036, Faliarivo; Decary 14927, Tampoketsa d’Ankazobe; Decary 17214, Manankazo STF; Decary 17311, Ambositra; Decary 17333, Ambatofinandrahana; Decary 17403, Ambositra; Descoings 338, Fort-Dauphin; Dorr 3842, col des Tapias; Edmondson 95 - 57, Mandena STF; Geay 7507, 7618, 7707, 7926, 8708 Mananjary; Gentry 11363, Ambila-Lemaitso STF; Gentry 11836, Ambohitantely RS; Gereau 3294, Evatra; Gereau 3423, Manafiafy; Gereau 5690, Anjozorobe; Harder 1536, Anjozorobe; Humbert 4640, col des Tapias; Humbert 5864, Enivaha; Humbert 5899, pic St. Louis; Humbert 7111, col des Tapias; Humbert 11090, Ambohitantely RS; Humbert 13835, Andohahela RNI; Humbert 14080, Imonty; Humbert 14477, Faliarivo; Humbert 20623, Ampasimena; Humbert 20702, Vohimavo; Humbert 28059, Ambatofinandrahana; Humbert 29938, Itremo; Jacquemin 934, Ambavarano; Jacquemin 1158, Mandromodromotra; Johnson WII 33, Mandena STF; Jongkind 828, 932, col des Tapias; Keraudren 1062, pic Saint Louis; Keraudren 24894, 24997, Fort Dauphin; Keraudren 25712, Ivato- Ambatofinandrahana; Keraudren 26009, Itremo; Labat 3010, col des Tapias; Leeuwenberg 14443, c. 20 km Ivato-Itremo; Leeuwenberg 14589, Ambila- Lemaitso STF; Leeuwenberg 14648, Anjozorobe; Lewis 728, Ambila-Lemaitso STF; Louvel 33 bis, Analamazaotra-Périnet RS; Lowry 4375, 4397, Anjozorobe, 5215, Mandena STF; McPherson 14164, Ste. Luce; McPherson 14278, Analalava (Manantenina); Messmer 781, Itremo; Miller 8777, Anjozorobe; Morat 3315, Ivato; Morat 4411, Andohahela RNI; Morat 4512, Antsiatsiaka; Peltier 2159, Fiadanana; Peltier 2184, Anjoma; Perrier de la Bâthie 3004, Andringitra RNI; Perrier de la Bâthie 3007, Ambatofinandrahana; Perrier de la Bâthie 3012, Matatana; Perrier de la Bâthie 4488, Vatovavy; Perrier de la Bâthie 5337, Manankazo STF; Perrier de la Bâthie 5341, Ibity; Perrier de la Bâthie 5342, Tamatave; Perrier de la Bâthie 13180, Tsitondroina; Perrier de la Bâthie 13213, Beveromay; Perrier de la Bâthie 13574, Ibity; Phillipson 1850, Ambila-Lemaitso STF; Rabenantoandro 62, Itremo; Rabevohitra 3770, Mahabo; Rakotoarisoa 22, 32, Ambatofinandrahana; Rakotoarisoa 59, Faliarivo FC; Rakotozafy 2682, Anjozorobe; Randriamampionona 331, 569, Andohahela RNI; Randriamanantena 119, Andohahela RNI; Randrianaivo 75, Ambohitantely RS; A. Randrianasolo 224, Ambila-Lemaitso STF; A. Randrianasolo 232, I v a t o; A. Randrianasolo 283, M a n a n t a n t e l y; A. Randrianasolo 347, Manafiafy; Rauh 1445, Fort- Dauphin; Réserves Naturelles 13 - RN-XI, Andohahela RNI; Réserves Naturelles 168, 202, Ambohitantely RS; Réserves Naturelles 706, Andringitra RNI; Réserves Naturelles 1416, 1499, Ambila-Lemaitso STF; Réserves Naturelles 6507, Andringitra RNI; Réserves Naturelles 6706, Ambohitantely RS; Réserves Naturelles 7166, Andringitra RNI; Réserves Naturelles 11247, Zahamena RNI; Réserves Naturelles 11691, Andringitra RNI; Schatz 1320, Ambila-Lemaitso STF; Schatz 2961, Anjozorobe; Schatz 3955, Ankafobe; Schatz 3964, Itremo; Schedl 133, Ambila-Lemaitso STF; Scott-Elliot 2369, 2554, 2713, 2713 bis, 2753, Fort-Dauphin; Service Forestier 3 - R- 3, Ambohitantely RS; Service Forestier 54 - R- 10, Itremo; Service Forestier 232, Manandona; Service Forestier 1078, 1122, 1569, Ambila-Lemaitso STF; Service Forestier 2155, Fianarantsoa; Service Forestier 2708, Ampandrambato; Service Forestier 3259, Ambila-Lemaitso STF; Service Forestier 3267, Mahanoro; Service Forestier 3811, Anosivelo; Service Forestier 4157, 4707, 4711, Ambila- Lemaitso STF; Service Forestier 4829, Mangatsiatra; Service Forestier 4887, 4908, Ambila-Lemaitso STF; Service Forestier 5625, Vohipaho; Service Forestier 5631, Marohita; Service Forestier 5737, Ambila- Lemaitso STF; Service Forestier 6077, Mandena STF; Service Forestier 7035, Mananjary; Service Forestier 7258, Ambila-Lemaitso STF; Service Forestier 7292, Misevo; Service Forestier 7419, Mandena STF; Service Forestier 7598, Fierenana; Service Forestier 7771, Mandromodromotra; Service Forestier 7820, Mandena STF; Service Forestier 8416, Ambohitantely RS; Service Forestier 9506, Ampangalana Atsimo; Service Forestier 9540, Marohita; Service Forestier 9592, Ambohitantely RS; Service Forestier 9647, Ambila-Lemaitso STF; Service Forestier 9668, Manantenina; Service Forestier 11542, Faliarivo; Service Forestier 12988, Vatovary (= Vatovory); Service Forestier 13180, Ambila-Lemaitso STF; Service Forestier 13472, Ankijana; Service Forestier 13701, Ampangalana Atsimo; Service Forestier 14047, Antanambao; Service Forestier 14661, Nosy-Varika; Service Forestier 14670, Analalava; Service Forestier 14763, Ankijana; Service Forestier 15156, Anjozorobe; Service Forestier 15806, Sahavoay; Service Forestier 16113, Nosy-Varika; Service Forestier 16119, Ambazato; Service Forestier 16806, Manankazo S T F; Service Forestier 18361, A m b o h i t a n t e l y RS; Service Forestier 18771, Manohilahy; Service F o r e s t i e r 1 9 6 2 6, I a m b o a l a; S e r v i c e F o r e s t i e r 19695, Ambohimanana; Service Forestier 19874, Ambohitantely RS; Service Forestier 22456, Andohahela RNI; Service Forestier 25366, Ampasimpotsy STF; Service Forestier 32214, Ambila- Lemaitso STF; Service Forestier 34202, Ambohitantely RS; Zarucchi 7345, Anjozorobe; Zarucchi 7411, Ambila-Lemaitso STF. 8. Leptolaena raymondii G. E. Schatz & Lowry, sp. nov. Haec species a congeneris ad Leptoaenam sensu stricto pertinentibus foliis falcatis apice longe cuspidatis marginibus valde revolutis subtus indumento persistente ex trichomatibus brevibus adpressis etiam saepe trichomatibus suberectis> 1 mm longis constante distinguitur. TYPUS. — Randrianasolo & Ranaivojaona 638, Madagascar, Prov. Toamasina, Dist. Fénérive-Est, Mahatsara STF, 17 ° 38 ’ S, 49 ° 29 ’ E, 5 m, 10 Nov. 1999, fl. (holo-, MO!; iso-, G!, K!, P!, TAN!). Trees 5 - 10 m tall, to 20 cm dbh, stems short sericeous-tomentellous. Leaves chartaceous, ovate to broadly ovate, usually strongly falcate, 2 - 4.5 × 0.7 - 2.3 cm, glabrous above, appressed short sericeous below often mixed with slightly erect trichomes> 1 mm long, sometimes subglabrescent, base strongly asymmetrical with one side obtuse to rounded and the other side cuneate, margins strongly revolute, apex long cuspidate, the very tip rounded, venation obscurely brochidodromous, secondary veins 8 - 10 per side, midrib slightly raised above, Revision of Leptolaena (Sarcolaenaceae) glabrous, raised below, moderately densely sericeous; petiole 2 - 4 mm long, 0.7 - 0.8 mm in diam., canaliculate, densely sericeous, stipules indistinct, caducous. Inflorescences terminal and axillary from the uppermost leaves, corymbose, 9 - 20 - flowered, axes densely gray hirsute-tomentose; peduncle 0.5 - 1 mm long, indistinct to somewhat distinct; involucre urceolate, base acute to obtuse, 2.5 - 3.5 mm tall, 2.5 - 3 mm diam. at apex, densely granular ferruginous farinose and also with moderately dense white appressed trichomes to 0.3 mm long, apical margin with 8 - 10 unequal triangular to narrowly triangular teeth, 1 - 1.3 mm tall, 0.3 - 1 mm broad at base; sepals exserted 1 mm beyond involucre, 3 × 3 mm, very widely obovate, apex rounded, strongly inrolled, densely whitish hirsute outside, glabrous inside; petals narrowly oblong to oblanceolate, 8 - 9 × 2 - 3 mm, white to pale yellow, glabrous, exserted 4 - 5 mm beyond involucre, with evident venation upon drying, the apex rounded; stamens (10 -) 11 (- 12), free, filaments 4.5 - 5.5 mm long, anthers oblong 0.6 - Acknowledgements 0.9 × 0.3 mm; ovary obovoid, 1 × 0.8 mm, We wish to thank K. SIKES, S. ANDRIAMBOLOLONERA densely whitish hirsute, style slender, 5 - 6 mm and J. RAHARIMAMPIONONA for specimen and data long, exserted 4 mm above involucre, sparsely management; A. JOUY and Roger LALA for the fine whitish velutinous towards the base, stigma illustrations; J. RABENANTOANDRO for help with field capitate, 1.2 - 1.5 mm diam. Mature fruit work noses ;; R Ph. G. EREAU MORAT for and assistance his staff with for the hospitality Latin diagunknown. — Fig. 8. extended at the Laboratoire de Phanérogamie in Paris; and the curators of several herbaria (K, TAN and Leptolaena raymondii is a small to medium- TEF) for giving us access to their specimens. Field sized tree known only from Mahatsara Station work was conducted under collaborative agreements Forestière (Fig. 6). It is the only species of between Botanique the et Missouri Zoologique Botanical de Tsimbazaza Garden and the and Parc the Leptolaena sensu stricto with persistent indumen- Direction de la Recherche Forestière et Piscicole, tum throughout the lower surface of the leaf FOFIFA, Antananarivo, Madagascar. We gratefully blade, consisting of both short appressed tri- acknowledge courtesies extended by the Government chomes and often longer partially erect trichomes of Madagascar (Direction Générale de la Gestion des Ressources greater than 1 mm long. The leaves of this species Nationale pour Forestières la Gestion) des and Aires by the Protégées Association. Special are further characterized by their strongly revo- thanks go to the Département de Biologie et Écologie lute margins. Végétale, Université d’Antananarivo for encouragement and assistance. This research was conducted with ETYMOLOGY. — The species epithet honors support from U. S. National Science Foundation grants DEB- 9024749 and DEB- 9627072 and from our friend and colleague Raymond RABEVOHITRA, the John D. and Catherine T. MACARTHUR Foundation, Head of the FOFIFA herbarium (TEF), who has the Liz CLAIBORNE and Art ORTENBERG Foundation, contributed so much to our knowledge of LWO Inc., and the National Geographic Society. Madagascar’s woody flora. VERNACULAR NAMES. — Amaninombilahy, Anjananjana. REFERENCES CONSERVATION STATUS. — With an area of occupancy <100 km 2, and only a single known ALVERSON W. S., KAROL K. G., BAUM D. A, CHASE population that is not included within the cur- M. W., SWENSEN S. M., MCCOURT R. & SYTSMA rent protected areas network, L. raymondii is K. J. 1998. — Circumscription of the Malvales and assigned a preliminary status of Critically relationships to other Rosidae: evidence from rbc L Endangered (CR). Along with L. delphinensis, the sequence data. Amer. J. Bot. 85: 876 - 887. BAYER C., FAY M. F., DE BRUIJN A. Y., SAVOLAINEN V., status of L. raymondii demonstrates the urgent MORTON C. M., KUBITZKI K., ALVERSON W. S. & need for additional protection of littoral forest CHASE M. W. 1999. — Support for an expanded along the East coast of Madagascar. family concept of Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid atp B and rbc L DNA sequences. Bot. J. Linn. PARATYPES. — MADAGASCAR, Prov. Toamasina: Soc. 129: 267 - 303. Rabenantoandro et al. 562, Mahatsara STF, CAPURON R. 1970. — Observations sur les Sarco- 17 ° 38 ’ 13 ” S, 49 ° 29 ’ 03 ” E, 2 m, 7 July 2001, bud laenacées. Adansonia, sér. 2, 10: 247 - 265. (G, K, MO, P, TAN); Razafindramora et al. 150, CAVACO A. 1951. — Remarques sur les genres same locality and date, bud (K, MO, P, TEF); Service Leptolaena et Xerochlamys (Chlaenacées). Un nou- Forestier 32648, Marovovonana, Mahatsara STF, veau genre de Chlaenaceae. Bull. Mus. Natl. Hist. [17 ° 38 ’ S, 49 ° 29 ’ E], 15 June 1985, bud (MO, TEF); Nat., sér. 2, 23: 133 - 139. Service Forestier 34104, Mahatsara STF, [17 ° 38 ’ S, CAVACO A. 1952 a. — Recherches sur les Chlénacées, 49 ° 29 ’ E], 19 Oct. 1990, fl. (MO, TEF); Service famille endémique de Madagascar. Mém. Inst. Sci. Forestier 34407, Andranotsara, Mahatsara STF, Madagascar, sér. B, Biol. Vég. 4: 59 - 92. [17 ° 38 ’ S, 49 ° 29 ’ E], 1 Oct. 1992, bud (G, K, MO, P, CAVACO A. 1952 b. — Chlénacées. Fl. Madagascar TEF, WAG); Service Forestier 34554, point d’essai, 126: 1 - 37. Mahatsara STF, [17 ° 38 ’ S, 49 ° 29 ’ E], 1 Dec. 1993, COETZEE J. A. & MULLER J. 1984. — The phytogeobud, fr. (MO, TEF); Service Forestier 35141, same graphic significance of some extinct Gondwanan locality, 30 Aug. - 4 Sep. 1999, bud (MO, TEF). pollen types from the Tertiary of the southwestern Cape (South Africa). Ann. Missouri Bot. Gard. 71: SCHATZ G. E. 2000. — Endemism in the Malagasy 1088 - 1099. tree flora: 1 - 9, in LOURENÇO W. R. & GOODMAN CORNET A. 1974. — Essai de cartographie bioclima- S. M. (eds.), Diversity and Endemism in Madagascar. tique à Madagascar. Notic. Explic. ORSTOM No. 55. Mém. Soc. Biogéogr., Paris. DU PUY D. J. & MOAT J. 1996. — A refined classifica- SCHATZ G. E., BIRKINSHAW, C. R., LOWRY II P. P., tion of the vegetation types of Madagascar, and RANDRIANTAFIKA F. & RATOVOSON F. 2000 a. — their current distribution: 205 - 218, in LOURENÇO The endemic plant families of Madagascar project: W. R. (ed.), Biogéographie de Madagascar. Editions integrating taxonomy and conservation: 11 - 24, in de l’ORSTOM, Paris. LOURENÇO W. R. & GOODMAN S. M. (eds.), LOWRY II P. P., SCHATZ G. E. & PHILLIPSON P. B. Diversity and Endemism in Madagascar. Mém. Soc. 1997. — The classification of natural and anthro- Biogéogr., Paris. pogenic vegetation in Madagascar: 93 - 123, in SCHATZ G. E., LOWRY II P. P., LESCOT M., WOLF GOODMAN S. M. & PATTERSON B. D. (eds.), A. - E., ANDRIAMBOLOLONERA S., RAHARIMALALA V. Natural change and human impact in Madagascar. & RAHARIMAMPIONONA J. 1996. — Conspectus of Smithsonian Inst. Press, Washington & London. the vascular plants of Madagascar: a taxonomic and LOWRY II P. P., SCHATZ G. E. & PHILLIPSON P. B. conservation electronic database: 10 - 17, in VAN DER 1998. — Classification de la végétation naturelle et MAESEN L. J. G., VAN DER BURGT X. M. & VAN anthropique de Madagascar. Bull. Acad. Natl. MEDENBACH DE ROOY J. M. (eds.), The Biodiversity Madagascar, n. s., 73: 219 - 233. of African Plants. Proc. XIV AETFAT Congress. LOWRY II P. P., SCHATZ G. E., LEROY J. - F. & WOLF Klower Academic Publishers, Wageningen, The A. - E. 1999. — Endemic families of Madagascar. III. Netherlands. A synoptic revision of Schizolaena (Sarcolaenaceae). SCHATZ G. E., LOWRY II P. P. & WOLF A. - E. 1998. — Adansonia, sér. 3, 21: 183 - 212. Endemic families of Madagascar. I. A synoptic LOWRY II P. P., HAEVERMANS T., LABAT J. - N., SCHATZ revision of Melanophylla Baker (Melanophyllaceae). G. E., LEROY J. - F. & WOLF A. - E. 2000. — Adansonia, sér. 3, 20: 233 - 242. Endemic families of Madagascar. V. A synoptic SCHATZ G. E., LOWRY II P. P. & WOLF A. - E. revision of Eremolaena, Pentachlaena, and 1999 a. — Endemic families of Madagascar. II. A Perrierodendron (Sarcolaenaceae). Adansonia, sér. 3, synoptic revision of Sphaerosepalaceae. Adansonia, 22: 11 - 31. sér. 3, 21: 107 - 123. NILSSON S. & R ANDRIANASOLO A. 1999. — SCHATZ G. E., LOWRY II P. P. & WOLF A. - E. Morphology and functional aspects of the pollen in 1999 b. — Endemic families of Madagascar. IV. A the Sarcolaenaceae. Palaeoecology of Africa 26: 191 - 200. synoptic revision of Asteropeia (Asteropeiaceae). PERRIER DE LA BÂTHIE H. 1931. — Remarques sur les Adansonia, sér. 3, 21: 255 - 268. Chlaenacées. Bull. Soc. Bot. France 78: 46 - 65. SCHATZ G. E., LOWRY II P. P. & WOLF A. - E. RANDRIANASOLO A. & M ILLER J. S. 1999. — 2000 b. — Endemic families of Madagascar. VI. A Taxonomic revision of the genus Sarcolaena (Sarco- synoptic revision of Rhodolaena (Sarcolaenaceae). laenaceae). Ann. Missouri Bot. Gard. 86: 702 - 722. Adansonia, sér. 3, 22: 239 - 252. Manuscript received 18 September 2001; revised version accepted 2 October 2001.	en	Schatz, George E., Lowry Ii, Porter P., Wolf, Anne-Elizabeth (2001): Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae). Adansonia (3) 23 (2): 171-189, DOI: 10.5281/zenodo.4605057
