Croton adenophorus Baill. in Adansonia 1: 153. 1861

Kainulainen, Kent, Ee, Benjamin van, Razafindraibe, Hanta & Berry, Paul E., 2017, A revision of the Adenophorus Group and other glandular-leaved species of Croton (Euphorbiaceae) from northern Madagascar and Mayotte, including three new species, Candollea 72 (2), pp. 371-402 : 382-386

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Croton adenophorus Baill. in Adansonia 1: 153. 1861


Croton adenophorus Baill. in Adansonia 1: 153. 1861 View in CoL [as adenophorum] ( Fig. 1C View Fig , 2B View Fig , 4 View Fig E-F, 6).

Croton adenophorus var. genuinus Müll. Arg. in A. DC., Prodr. 15(2): 589. 1866 View in CoL [nom. inval.].

Ξ Oxydectes adenophora (Baill.) Kuntze, Revis. Gen. Pl. 2: 610. 1891 View in CoL .

Lectotypus (designated here): MADAGASCAR. Prov. Antsiranana: Diana Reg.: Nossi-bé , 1837, Richard 214 ( P [ P00123689 ]!; isolecto-: P [ P00123690 ]!). Syntypi: MADAGASCAR. Prov. Antsiranana: Nossi-bé, s.d., Richard 574 ( P [ P00123691 ]!); ibid. loc., 1849, Boivin 2187 ( G [ G00446358 , G00446359 ]!, P [ P00301515 ]!).

= Croton payerianus Baill. in Adansonia 1: 154. 1861 View in CoL [as payerianum]. Ξ Oxydectes payeriana (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891 View in CoL . Lectotypus (designated here): MADAGASCAR. Prov. Antsiranana: “ Nossibé ”, 1849, Boivin 2187 ( P [ P00389508 ]!; isolecto-: G-DC [ G00311984 ]!, G [ G00446358 , G00446359 ]!, P [ P00389509 , P00404480 , P00404481 , P00123684 ]!).

= Croton subaemulans Baill. in Bull. Mens. Soc. Linn. Paris 2: 850. 1890 View in CoL . Typus: M ADAGASCAR: “Central Madagascar”, Baron 5795 (holo-: K [ K000422590 ]!; iso-: P [ P00133593 ]!), syn. nov.

= Croton tenuicuspis Baill. in Bull. Mens. Soc. Linn. Paris 2: 927. 1891 View in CoL . Typus: M ADAGASCAR: “Madag. centr.”, Baron 5846 (holo-: P [ P00133364 ]!, iso-: K [ K000422590 ]!), syn. nov.

= Croton tulasnei Baill. in Adansonia 1: 156. 1861 View in CoL . Ξ Oxydectes tulasnei (Baill.) Kuntze, Revis. Gen. Pl. 2: 613. 1891 View in CoL . Lectotypus (designated here): FRANCE. Dept. Mayotte: Bouzi , 1850, Boivin s.n. ( P [ P00133305 ]!; isolecto-: P [ P00133306 , P00466148 ]), syn. nov.

Shrubs or trees, 1-9 m tall, dichotomously branching, internodes sometimes so short as to give the appearance of whorled branches. Branches flattened on new growth but becoming terete with age, bright green and sparsely covered with whitish stellate indument, soon turning reddish brown, glabrous, ± glossy. Bark smooth and conspicuously lenticellate. Stipules 6-17 × 2-4 mm, lanceolate or auriculate, tapering to a thin apex, sometimes with marginal glands (and glandular filaments), caducous. Leaves opposite. Petioles 0.6-8.5 cm, adaxially canaliculate, stellate-pubescent, usually with a pair of subsessile, concave, discoid, yellow, glabrous glands (0.3-0.8 mm in diam.) by the base of the lamina. Leaf blades membranaceaous to chartaceous, glandular crenulate, denticulate, or serrate, ovate, 4.2-17.0 × 2.3-11.5 cm, apex acute to acuminate, base rounded to cordate; abaxial and adaxial surfaces sparsely stellate-pubescent on young leaves, ± glabrous on older leaves, pale green when fresh (turning orange in old leaves) and drying matte pale green to brown; venation evident on the adaxial side, prominent below, with 5-10 pairs of brochidodromus, ± penninerved secondary veins (the lowermost pairs congested and appearing palmate), and crossvenulate tertiary venation; glands are usually present in some of the axils of the secondary veins on the abaxial side ( Fig 1C View Fig ). Inflorescences terminal, raceme-like thyrses, 1.5-22 cm long, terminal, often appearing unisexual, or bisexual with pistillate flowers at the base and staminate flowers at the distal end, axes stellate-pubescent, flattened; bracts narrowly triangular, 1.0- 3.5 mm long. Staminate flowers with stellate-pubescent, subglobose buds 1.8-2.3 mm diam., pedicels elongating from bud to anthesis, 1.0- 3.5 mm long; sepals 5, pale green, shortly connate at base, lobes broadly triangular to ovate, often unequal in size, 1.5-3.5 × 1.3-2.0 mm, apex acute, inflexed at anthesis, abaxially stellate-pubescent, adaxially sparsely ciliate, margins densely ciliate; petals 5, yellowish, ellipticspatulate, 1.9-2.4 × 0.7-1.0 mm, recurved at anthesis, abaxially stellate-pubescent and papillose, adaxially ciliate towards apex, margins densely ciliate; disc glands 5, opposite the sepals, sessile, deltoid, truncate or with an apical depression, c. 0.3 × 0.5 mm, yellow; stamens 10-16, white, filaments 1.6- 3.1 mm long, ciliate, anthers broadly elliptic, c. 0.7 × 0.6 mm; receptacle pilose. Pistillate flowers with stellate-pubescent buds, 2-3 mm diam., pedicels 2-4(-9) mm long; sepals 5, elliptic, spreading at anthesis, often unequal in size, 2.5-11.5 × 1.1-2.5 mm, apex acute, shortly connate at base, glandular margin, abaxially and adaxially subglabrous, sometimes bearing glands similar to those on the leaves, bright green, persistent in fruit; petals usually absent/reduced, but when present, c. 1.5 × 0.3 mm, abaxially stellate-pubescent, adaxially glabrous, margin ciliate; disc glands 5, opposite the sepals, sessile, ellipsoidal, c. 0.3 × 0.6 mm, pale yellow; ovary densely covered by ferruginous to dark brown, stellate trichomes, pale green, globoid-ellipsoid, 2-3 mm in diameter, styles 3, 2- 3 mm long, each branch flattened and 1-3 times bifurcate, the first bifurcation often congested and fused, spreading, recurved at the apices, abaxially stellate-pubescent, adaxially glabrous, (pale) yellow, turning brown, persistent. Capsules, 5.0-6.0 × 5.5-8.0 mm, smooth, pale brown, covered with contrasting dark brown stellate trichomes, exocarp not separating, endocarp woody, 0.3-0.4 mm thick ( Fig. 4E View Fig ); columella 3-4 mm long, cornute, capitate. Seeds ± compressed-ellipsoid, 3.1-4.1 × 2.3-3.1 × 1.8-2.5 mm ( Fig. 4F View Fig ); testa glossy, verrucose, brown; caruncle reniform, c. 0.5 × 1 mm.

Phenology. – We have seen flowering specimens from August to April, and in bud in May (we have not seen any specimens from June or July), so it is likely that this species flowers more or less continuously throughout the year.

Distribution, habitat and ecology. – Croton adenophorus occurs both in evergreen forest and in secondary or degraded vegetation, and it is often collected in riparian or moist habitats. It occurs from sea level to c. 450 m elevation. It is locally frequent in the Sambirano Domain ( HUMBERT, 1955), and there is also a population from the littoral forest of Analabe in northernmost Vohémar, Antsiranana Province on the northeast coast ( Fig. 2B View Fig ). Cours 3207, collected in the Mainampango forest (Andapa, SAVA Reg.), was indicated as a shade plant in a vanilla plantation. It is possible that the disjunct outliers from southern Mahajanga, i.e., Decary 2359 from Morafenobe, Service Forestier 16 from Tsingy de Namoroka, and van Ee et al. 1165 from Ambodiriana, were also escaped from cultivation, since this species is not only used as a shade plant but also in hedgerows (see Randrianarivelo et al. 145). The only known specimen from Mayotte is Boivin s.n. (the type of C. tulasnei ), collected in 1850, and it is therefore uncertain whether C. adenophorus still occurs there, and if the gathering was made from a native or introduced plant.

Conservation assessment. – Since C. adenophorus is widespread and known from numerous localities, including the protected areas of Ampasindava, Galoko, and Manongarivo, we assign the species as “Least Concern” [LC] according the IUCN Red List Categories and Criteria ( IUCN, 2012).

Vernacular names. – “Bemasaho” (Rabenantoandro et al. 1070), “Hazomafaiky” (Madiomanana & Ammann 181), “Hazomafaitra” (Nusbaumer & Tahinarivony 2977), “Matifioditra” (Cours 3207), “Mavokely” (Decary 999), “Sakaiala” (Randrianarivelo 145), “Sakaitomendry” (Randrianarivelo 145).

Notes. – The species epithet adenophorus means “bearer of glands.”

This species is distinct in the glabrous and glossy reddishbrown bark; acuminate leaves that usually dry matte green; contrasting white stellate hairs on the young leaves (and shoots), whereas the mature leaves are ± glabrous ( Fig. 1C View Fig ); the usually large, foliaceous stipules that may be auriculate at the base and then taper to a fine point ( Fig. 6 View Fig D-E); the flattened, patent stigmas; and the often unequal and foliaceous, bright green sepals that sometimes have glands similar to those on the leaves. The lectotype of C. adenophorus (P [P00123689]), which was collected by Achille Richard, bears a label with the name “ Croton acuminatum” in Richard’s hand, which BAILLON (1861) cited in his protologue. BAILLON (1861) also cited Boivin 2187 at P [P00301515] as a syntype of C. adenophorus , which he annotated as such, despite citing and annotating other sheets of the same collection number as syntypes of C. payerianus on the next page. Since C. adenophorus and C. payerianus were published in the same article, either one could be the accepted name, but we follow the precedent of LEANDRI (1939), who synonymized C. payerianus .

It is important to note that our concept of C. adenophorus differs from that of LEANDRI (1939). By including C. loucoubensis as a synonym of C. adenophorus, LEANDRI confounded this group, and his description of C. adenophorus corresponds to what we recognize here as C. loucoubensis . He then went on to recognize C. subaemulans , which is really the same as the true C. adenophorus . BAILLON (1861) separated C. tulasnei from his C. payerianus (to us, a synonym of C. adenophorus ) by its poorly developed or absent laminar glands, somewhat different leaf shape (oval-ovate and acute vs cordate and acuminate), linear stipules, and different branching pattern of the stigma. However, similar variation in the shape of the leaves and stipules, and in the number of laminar glands, is also seen within populations of C. adenophorus in Madagascar, and the stigma branches may vary from 1-3 times bifurcate ( Fig. 6 View Fig G-H). We therefore do not consider C. tulasnei as a distinct species from C. adenophorus .

Additional specimens examined. – MADAGASCAR. Prov. Antsiranana: Diana Reg., Nosy Be , env. d’Ambatozavany , 50-250 m, 14.XII.1967, Bernardi 11863 ( G, K, P); Nosy Be, 1847-1852, Boivin s.n. ( G, P); SAVA Reg., forêt de la montagne de Mainampango Est , village Ambalavoanio , c. 450 m, 1.III.1949, Cours 3207 ( P); Ampasindava , 10 m, 14.VIII.1957, Cours 5232 ( P); Analanantsoa, Ambodisakoana , 14°04’S 48°15’E, 40 m, 1.XI.1994, Derleth 145 ( G, K, MO, P, TAN); GoogleMaps ibid. loc., 14°04’S 48°15’E, 50 m, 3.IV.1996, Gautier 2966 ( G, K, MO, P); GoogleMaps Manongarivo RS, Beraty , 14°01’51”S 48°14’53”E, 65 m, 22.X.2012, Gillespie et al. 10651 ( CAN, MICH, MO, TAN); GoogleMaps Ambanja, Antsatsaka-Ambato , 13°27’23”S 48°32’39”E, 26 m, 30.VIII.2002, Hong-Wa 71 ( MO, P, TAN, TEF); GoogleMaps Vallée de la Lokoho (NE), près d’Ambalavohino , 75-300 m, 9-10.I.1949, Humbert & Cours 22834 ( K, P); Ampasindava, 7.IX.1967, Jacquemin H501J ( P); Ampasindava , 13°45’39”S 48°06’38”E, 370 m, 29.XI.2008, Madiomanana & Ammann 181 ( G, K, MO, P, TEF, WAG); GoogleMaps Vohemar Distr., Nosibe , Anjiabe , Anaborano , Analabe forest , 13°04’08”S 49°54’20”E, 5 m, 9.XII.2004, Manjakahery & Sola 80 ( MO); GoogleMaps Ampasindava , 13°48’17”S 48°10’48”E, 218 m, 29.XI.2007, Nusbaumer 2615 ( G, K, MO, P, TEF); GoogleMaps ibid. loc., 13°43’59”S 48°04’15”E, 141 m, 23.XI. 2008, Nusbaumer & Tahinarivony LN 2966 ( G, MO, P, TEF); GoogleMaps ibid. loc., 13°46’09”S 48°06’12”E, 312 m, 24.XI.2008, Nusbaumer & Tahinarivony LN 2977 ( G, K, MO, P, TEF); GoogleMaps Anaborano, 13°04’42”S 49°54’13”E, 25 m, 2.XI.2002, Rabenantoandro et al. 1070 ( MICH, MO, P); GoogleMaps Analabe forest, 13°04’14”S 49°53’40”E, 22.II.2003, Rabevohitra et al. 4481 ( MO, P, TEF); GoogleMaps ibid loc., 13°04’09”S 49°54’08”E, 18 m, 11.X.2004, Randrianarivelo et al. 145 ( MICH, MO); GoogleMaps Ambilobe Distr., Beramanja , Anketrabe Bilinta , Ambahatra , Galoko forest , 13°36’11”S 48°42’37”E, 430 m, 7.X.2013, Randriatsivery et al. 479 ( G, MO, P, TAN); GoogleMaps Analabe forest , 13°04’43”S 49°54’04”E, 13.V.2004, Razakamalala et al. 1270 ( MO, P, TEF); GoogleMaps Nosy Be , 25.IX.1951. Reserves Naturelles 2992 ( P); ibid loc., s.d., Richard 180 ( P); ibid loc., 12.III.1971, Schmitt 252 ( P); RN [Route Nationale] 6, 41 km N of Ambanja, 13°28’41”S 48°44’03”E, 25 m, 2.XI.2009, van Ee et al. 1133 ( MICH); GoogleMaps ibid loc., van Ee et al. 1134 ( MICH); ibid loc., van Ee et al. 1135 ( MICH); RN 6 , 41 km N of Ambanja, 13°31’23”S 48°41’28”E, 20 m, 2.XI.2009, van Ee et al. 1141 ( MICH); GoogleMaps RN 6, close to turn off for Ampampamena airstrip , 13°29’33”S 48°38’12”E, 8 m, 27.II.2016, Van Ee et al. 2287 ( MICH, TAN); GoogleMaps ibid loc., van Ee et al. 2288 ( MICH, TAN); RN 6 , N of the turnoff to Belinta , 13°30’22”S 48°42’05”E, 17 m, 27.II.2016, van Ee et al. 2291 ( MICH, TAN); GoogleMaps ibid loc., 13°30’09”S 48°42’20”E, 22 m, 27.II.2016, van Ee et al. 2292 ( MICH, TAN); GoogleMaps ibid loc., van Ee et al. 2293 ( MICH, TAN); ibid loc., 13°28’56”S 48°43’36”E, 17 m, 27.II.2016, van Ee et al. 2294 ( MICH, TAN); GoogleMaps RN 6 E of Ampampamena , 13°30’46”S 48°40’30”E, 34 m, 27.II.2016, Van Ee et al. 2295 ( MICH, TAN); GoogleMaps RN 6 E of Ampampamena and south of Ambatomainty , 13°28’26”S 48°44’20”E, 21 m, 27.II.2016, van Ee et al. 2299 ( MICH, TAN); GoogleMaps RN 6 , PK [ Pointe Kilometrique roadmarker ] 525, 57 km N of Ambanja, 13°26’12”S 48°47’34”E, 17 m, 8.III.2016, van Ee et al. 2382 ( MICH, TAN); GoogleMaps Ambanja, Nosy Be , 16.XII.1921, Waterlot 289 ( P); ibid loc., Waterlot 291 ( P); Manongarivo, Ambahatra , 13°58’S 48°27’E, 400 m, 15.III.1999, Wohlhauser 60050 ( G, K, MO, P); GoogleMaps Manongarivo, Antafia Be , Amatolary , Moyen Ambahatra , 13°55’S 48°27’E, 230 m, 7.V.1999, Wohlhauser SW60113 ( G, K, MO, P). GoogleMaps Prov. Mahajanga: Sofia Reg., Analalava Distr ., Maromandia ( Befotaka ), 16.IX.1922, Decary 999 ( P); Haut Bemarivo, VIII.1907, Perrier de la Bâthie 9556 ( P); Melaky Reg., env. de Morafenobe , 3.X.1923, Decary 2359 ( P); Boeny Reg., Tsingy de Namoroka , 4.IV.1933, Service Forestier 16 ( P); RN 6 , PK 449 , 19 km S of Ambanja, 13°46’56”S 48°20’59”E, 25 m, 3.XI.2009, van Ee et al. 1142 ( MICH); GoogleMaps RN 6 , PK 413 , 55 km S of Ambanja, 14°00’04”S 48°10’06”E, 100-140 m, 3.XI.2009, van Ee et al. 1147 ( MICH); GoogleMaps Betsiboka Reg., RN 4 , PK 227 , 88 km S of Moevaharana, 17°31’03”S 46°58’41”E, 200 m, 6.XI.2009, van Ee et al. 1165 ( MICH); GoogleMaps RN 6 south of Ambanja , 13°55’44”S 48°12’58”E, 201 m, 8.III.2016, van Ee et al. 2384 ( MICH, TAN); GoogleMaps Along Route Nationale 6 , at PK 399 , 14°05’22”S 48°06’42”E, 214 m, 8.III.2016, van Ee et al. 2386 ( MICH, TAN); GoogleMaps ibid loc., van Ee et al. 2388 ( MICH, TAN).










Conservatoire et Jardin botaniques de la Ville de Genève


"Alexandru Ioan Cuza" University


Harvard University - Arnold Arboretum


Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet


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Croton adenophorus Baill. in Adansonia 1: 153. 1861

Kainulainen, Kent, Ee, Benjamin van, Razafindraibe, Hanta & Berry, Paul E. 2017

Oxydectes adenophora (Baill.)

Baillon 1891: 610

Croton tenuicuspis Baill . in Bull. Mens. Soc. Linn. Paris 2: 927. 1891

Baillon 1891: 927

Croton subaemulans Baill . in Bull. Mens. Soc. Linn. Paris 2: 850. 1890

Baillon 1890: 850

Croton adenophorus var. genuinus Müll. Arg. in A. DC., Prodr. 15(2): 589. 1866

Müll. 1866: 589

Croton payerianus

Baillon 1861: 154
Baillon 1861: 612

Croton tulasnei

Baillon 1891: 613
Baillon 1861: 156
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