A revision of the Adenophorus Group and other glandular-leaved species of Croton (Euphorbiaceae) from northern Madagascar and Mayotte, including three new species Kainulainen, Kent Ee, Benjamin van Razafindraibe, Hanta Berry, Paul E. Candollea 2017 2017-09-27 72 2 371 402 8WT3Y Kainul. & P.E. Berry Kainul. & P. E. Berry 2017 [801,1221,876,901] Magnoliopsida Euphorbiaceae Croton Plantae Malpighiales 23 394 Tracheophyta species orangeae sp. nov.     Typus: MADAGASCAR. Prov.Antsiranana: Diana Reg., Ramena, Ankorikakely, Baie des Sakalava, 12°16’40”S 49°23’01”E,  25 m,  9.XII.2004,  Razafitsalama et al. 692(holo-: MICH[ MICH1517188]!; iso-: CNARP, MO!, P[ P05484901]!, TAN).    Croton orangeae Kainul. & P.E. Berryis similar to C. adenophorus Baill.in its denticulate, cordate leaves and glabrous stems, but differs in its stipules that are awn-shaped (vs foliaceous), its mature leaves that are scabrously stellate-pubescent (vs subglabrous), and the bark of the branches matte and tan to gray (vs ± glossy and reddish brown) and soon turning flaky (vs smooth).  Shrubs 1-4 mtall, dichotomously branching, internodes sometimes so short as to give the appearance of whorled branches. Branches flattened on new growth but becoming terete with age, pale green and sparsely covered with whitish stellate indument, soon turning tan to gray, glabrous, matte. Bark flaky, gray to brown ( Fig. 10B). Stipules 5-10 mmlong, awn-shaped, early caducous. Leavesopposite, deciduous. Petioles 0.5-10 cmlong, adaxially canaliculate, stellate-pubescent, usually with a pair of subsessile, concave, discoid glands 0.4-0.6 mmdiam. at the junction with the lamina. Leaf blades chartaceous, (subentire) dentate to denticulate, ovate, 2.2-14.0 × 2.5-14 cm, apex (acute) acuminate, base (truncate) cordate; both sides with a persistent, scabrous stellate indument, green when fresh, drying matte pale green to brown; venation evident, with 6-10 pairs of brochidodromus, ± penninerved secondary veins (the lowermost pairs congested and appearing palmate), and cross-venulate tertiary venation; venation prominent, sometimes with glands in some of the axils of the secondary veins on the lower surface ( Fig. 1F), but absent in most leaves. Inflorescencesterminal, racemelike thryses 1-6.5 cmlong, bisexual with pistillate flowers towards the base and staminate flowers towards the distal end, axes stellate-pubescent, flattened; bracts narrowly triangular, c. 2.5 mmlong, caducous. Staminate flowerswith stellate-pubescent, subglobose buds, c. 1.8 mmdiam., pedicels elongating from bud to anthesis, 1-5 mmlong; sepals 5, pale green, shortly connate at base, lobes broadly triangular-ovate, 2-2.5 × 1.5-1.8 mm, apex acute, inflexed at anthesis, abaxially stellate-pubescent, adaxially glabrous, margins ciliate; petals 5, elliptic to spatulate, 1.5-2.5 × 0.7-0.9 mm, recurved at anthesis, abaxially glabrous and papillose, adaxially ciliate towards apex, margins densely ciliate; disc glands 5, opposite the sepals, sessile, ellipsoidal, c. 0.4 × 0.6 mm, yellow; stamens 13-21, white, filaments 1.5-2.5 mmlong, ciliate, anthers broadly elliptic, c. 0.8 × 0.7 mm; receptacle pilose. Pistillate flowerswith stellate-pubescent buds 2-2.5 mmdiam., pedicels 1.5- 5.5 mmlong; sepals 5, elliptic, not spreading at anthesis, 1.8-2.8 × 1-1.7 mm, apex acute, shortly connate at base, abaxially and adaxially stellate-pubescent, persistent in fruit; petals sometimes absent or reduced, when present 1.4-1.8 × 0.7-0.9 mm, spatulate, glabrous except for the ciliate margin, abaxially papillose; disc glands 5, opposite the sepals, sessile, ellipsoidal, 0.3 × 0.7 mm, yellow; ovary densely covered by ferrugineous to dark brown, stellate trichomes, globoid-ellipsoid, 2-3 mmdiam., styles 3, c. 2 mmlong, each branch flattened and twice bifurcate, spreading, recurved at the apices, abaxially stellate-pubescent, adaxially glabrous, persistent. Capsulesbroadly globoid, 5 × 6.5 mm, smooth, pale brown, covered with contrasting dark brown stellate trichomes ( Fig. 4K), exocarp not separating, endocarp woody, c. 0.5 mmthick; columella 4-5 mmlong, cornute, capitate. Seeds± compressed-ellipsoid, c. 3.5 × 3 × 2 mm; testa matte, verrucose, pale brown; caruncle narrowly reniform c. 0.4 × 1 mm( Fig. 4L).   Etymology. –The epithet refers to the Orangea (Oronjia) Peninsula in Antsiranana Province, where the typewas collected and the species is apparently restricted.   Phenology. –Only a few specimens are known, collected in bud in March, in flower in August-December, and in fruit in December.   Distribution, habitat and ecology.– This species is so far only known from the Orangea (Oronjia) Peninsula in northern Antsiranana Provincewhere it grows in dry scrub vegetation and littoral forest on limestone and sand, from sea level to c. 50 min elevation ( Fig. 2D).   Conservation assessment. – Croton orangeaeis known only from the Orangea Peninsula. The deciduous forests of Orangea are currently threatened by deforestation despite their recent additon to Madagascar’s protected area network. Deforestation is mostly related to agricultural practices. Given this and the extent of occurrence is < 5000 km 2, the new species is assigned as “Endangered” [EN B1 ab(iii)], according the IUCN Red List Categories and Criteria ( IUCN, 2012).   Notes. – Croton orangeaeis distinct for its glabrous, grayish branches with flaky bark, and dentate to denticulate leaves with a persistent, scabrous indument on both sides. In the shape of the leaves and in the pale brown capsules with white and contrasting brown stellate trichomes, it may appear similar to  C. adenophorus, but that species has a glossy reddish bark and subglabrous leaves, and also differ in its usually longer inflorescences ( 1.5-22 cmvs 1-6.5 cmin  C. orangeae), and distinctly foliaceous stipules and calyces. No vernacular names have been noted for this species.   Additional specimens examined.–   MADAGASCAR. Prov. Antsiranana: Diana Reg., Antsiranana Distr., Orangea,  25.XI.1970,  Debray1525-D( K, P);  ibid. loc., 12°14’20”S 49°21’49”E,  13 m,  29.VIII.2001,  Rabenantoandro& Razanatsoa579( MO, P);  ibid. loc., 12°14’08.2”S 49°21’40.3”E,  50 m,  25.X.2009,  van Ee et al. 1080( MICH);  ibid. loc., 12°13’54”S 49°21’28”E,  19 m,  5.III.2016,  van Ee et al. 2343( MICH, TAN);  ibid. loc., 12°14’08”S 49°21’41”E,  30 m,  5.III.2016,  van Ee et al. 2351( MICH, TAN);  ibid. loc.,  5.III.2016,  van Ee et al. 2352( MICH, TAN);  ibid. loc.,  5.III.2016,  van Ee et al. 2353( MICH, TAN). 3463906314 2004-12-09 MICH Razafitsalama et al Madagascar Baie des Sakalava 25 -12.277778 Diana Reg. 21 49.383614 Ankorikakely 23 394 MICH1517188 1 holotype 3463906402 2004-12-09 CNARP, MO, P, TAN Madagascar Baie des Sakalava 25 -12.277778 Typus 21 49.383614 Ankorikakely 23 394 P05484901 1 isotype 3463906329 1970-11-25 K, P Debray Madagascar Diana Reg. Orangea Antsiranana Distr. 24 395 1 Prov. Antsiranana 3463906450 2001-08-29 MO, P Rabenantoandro & Razanatsoa Madagascar Diana Reg. 13 -12.238889 Orangea 21 49.36361 Antsiranana Distr. 24 395 1 Prov. 3463906385 2009-10-25 MICH van Ee et al Madagascar Diana Reg. 50 -12.235611 Orangea 1 49.361195 Antsiranana Distr. 24 395 1 Prov. 3463906324 2016-03-05 MICH, TAN van Ee et al Madagascar Diana Reg. 19 -12.231667 Orangea 21 49.357777 Antsiranana Distr. 24 395 1 Prov. 3463906301 2016-03-05 MICH, TAN van Ee et al Madagascar Diana Reg. 30 -12.235556 Orangea 21 49.36139 Antsiranana Distr. 24 395 1 Prov. 3463906505 2016-03-05 MICH, TAN van Ee et al Madagascar Diana Reg. 30 -12.235556 Orangea 21 49.36139 Antsiranana Distr. 24 395 1 Prov. 3463906432 2016-03-05 MICH, TAN van Ee et al Madagascar Diana Reg. 30 -12.235556 Orangea 21 49.36139 Antsiranana Distr. 24 395 1 Prov.