New species and species reports of Croton L. (Euphorbiaceae) from the eastern forest corridor of Madagascar Kainulainen, Kent Ee, Benjamin van Antilahimena, Patrice Razafindraibe, Hanta Berry, Paul E. Candollea 2016 2016-11-23 71 2 327 356 8WT3G Kainul., B.W. van Ee & P.E. Berry Kainul., B. W. van Ee & P. E. Berry 2016 [139,723,282,306] Magnoliopsida Euphorbiaceae Croton Plantae Malpighiales 10 337 Tracheophyta species ferricretus sp. nov.    Typus:   MADAGASCAR. Prov.Toamasina: Alaotra- Mangoro Region, Moramanga Distr., Andasibe, Berano, Ambatovy mine concession, on “cuirasse” between the workers houses and the Ambatovy supply road, within sight of the Ambatovy office building, 18°51’02”S 48°18’29”E,  1142 m,  21.III.2016,  van Ee, Antilahimena, Kainulainen& Berry2436( holo- MICH[ MICH1513194]!; iso-: G!, K!, MAPR!, MO!, P!, TAN!).    Croton ferricretus Kainul., B.W. van Ee & P.E. Berryis similar to C. antanosiensis Leandriin its medium-sized, lepidote leaves and elongate inflorescences, but differs from it in the more ferrugineous indumentum (vs. silvery), shorter petioles, and obloid (vs. globoid) capsules.   Shrubs 0.4-4 mtall, dichotomously branching, internodes sometimes contracted giving the appearance of whorled branches. Branches± flattened on new growth and densely ferrugineous-lepidote, brown or gray, becoming terete and glabrous with age. Stipuleslinear-triangular, 0.5-1 mm. Leavesalternate along stem, subopposite or whorled at apex. Petioles 3-17(-28) mm, adaxially canaliculate, without any apparent glands. Leaf blades subcoriaceous, entire, elliptic, 18-85 × 9-28 mm, apex acute to shortly acuminate (rarely obtuse), base cuneate; adaxial surface glabrous, glossy, dark green when fresh (turning orange in old leaves) and drying gray-green; venation impressed, obscure, with 10-20 pairs of brochidodromus, ± penninerved secondary veins; abaxial surface densely silverylepidote and ferrugineous-punctate, the ferrugineous scales scattered among silvery ones, but more prevalent on the leaf veins; midrib prominent, ferrugineous-lepidote. Inflorescencesracemose, 20-85 mmlong, axillary or terminal, with mostly staminate flowers, usually with 1-2(-5) pistillate flowers at the base; axes densely ferrugineous-lepidote, flattened; bracts linear-lanceolate, 1-3 mmlong. Staminate flowerswith ferrugineous-lepidote, subglobose buds 1.5-2.5 mmin diam., pedicels elongating from bud to anthesis, 2-5 mmlong; sepals 5, firm, connate about halfway from the base, lobes broadly triangular to ovate, 1.0-1.4 × 0.8-1.7 mm, apex acute, inflexed at anthesis, abaxially ferrugineous-lepidote, adaxially glabrous, pale green; petals 5, white, narrowly obovate-spatulate, 1.8-2.5 × 0.5-1.1 mm, recurved at anthesis, abaxially lepidote, adaxially ciliate, margins densely ciliate; disc glands/nectaries 5, opposite the sepals, sessile, ellipsoid with an apical depression, 0.6-0.8 × 0.5-0.7 mm, yellow; stamens 14-19, white to pale yellow, filaments 1-2 mmlong, ciliate, anthers elliptic, 0.5- 0.7 mmlong; receptacle pilose. Pistillate flowerswith ferrugineous-lepidote, elliptic buds 2.8-3.0 × 1.7-2.5 mm, pedicels 1-8 mmlong; sepals 5, firm, narrowly triangular, ascending at anthesis, 1.8-5.3 × 0.5-1.3 mm, apex acute, shortly connate at base, abaxially ferrugineous-lepidote, adaxially glabrous, pale green, persistent in fruit; petals absent/reduced; disc glands/ nectaries 5, opposite the sepals, sessile, pentagonal, 0.8-1.0 × 0.6-0.8 mm, yellow; ovary obloid, 2.0- 2.5 mmdiam., lepidote, styles 3, c. 3 mmlong, each bifurcating twice (or ultimately thrice), spreading, recurved at the apices, abaxially ferrugineous-lepidote, adaxially glabrous but with yellowish pilose trichomes at the base, white, turning brown, persistent. Capsules4.5-9.0 × 3.5-6.0 mm, smooth, gray-green, lepidote, exocarp not separating, endocarp woody, c. 0.2 mmthick; columella 4-6 mmlong, cornute, capitate. Seeds± compressed-ellipsoid, c. 5 × 3 × 2 mm; testa glossy, rugulose, punctate, dark brown; caruncle reniform c. 1.3 × 0.5 mm.   Etymology. –The specific epithet refers to the ultramafic ferricrete soils to which this species is restricted.   Vernacular names. –“Fotsiavadika”, “Lazalaza”.   Phenology. –Flowering and fruiting specimens have been collected from May through March, indicating that  Croton ferricretusis quite aseasonal in its phenology.   Distribution, habitat and ecology. –  Croton ferricretusis so far only known from the Ambatovy-Analamay forest in Madagascar’s Moramanga Distr., occurring at 1000-1150 maltitude on ultramafic ferricrete soils ( Fig. 1A, 8A). It is a common and sometimes dominant component of the scrublands found on the most extreme rock-like crusts, and it also occurs in adjacent evergreen forests with better-developed soils. When growing on bare ferricrete (“cuirasse”),  C. ferricretusdoes not occur with other species of  Croton,but in forested areas it can be found growing alongside  C. lichenisilvaeLeandri,  C. macrobuxusBaill.,  C. nitidulusBaker,  C. submetallicusBaill., or  C. droguetioides.   Notes. –  Croton ferricretuswas recognized in the inventory of the Ambatovy-Analamay forest by PHILLIPSON et al. (2010)under the working name “ Croton lepidotoides” (ined. by the late Radcliffe-Smith), and considered a species of concern (SOC2, i.e., found only in the area of the mining footprint and the surrounding conservation zone). This is a very distinctive species among Malagasy Crotonsthat have small- to medium-small leaves with coppery-lepidote indumentum ( Fig. 8B-C). Other species that have these characteristics tend to have glomerulate inflorescences in the leaf axils, such as  C. jennyanusGris ex Baill.and  C. hypochalibaeusBaill., rather than racemose inflorescences ( Fig. 8D-F). It resembles  C. antanosiensisLeandrifrom southeastern Madagascar, but differs in its much more coppery-lepidote indumentum, shorter petioles, and more obloid capsules ( Fig. 7A). Since this is a locally common species that is capable of growing on bare, ferricrete substrates, we believe that it may be an excellent candidate for revegetation efforts once the mining operations at Ambatovy-Analamay have ceased. Its shrubby habit would enable it to reproduce more quickly than tree species, and it could help stabilize slopes in previously mined areas.    Paratypi. –   MADAGASCAR. Prov. Toamasina: Alaotra-Mangoro Region, Moramanga distr., Ambatovy-Analamay forest, 18°50’47”S 48°18’28”E,  1137 m,  9.II.2005,  Antilahimena et al.3323( G, K, MO, P, TAN);  ibid.loc., 18°51’27”S 48°19’07”E,  1113 m,  30.IX.2005,  Antilahimena& Edmond3862( MICH, MO, P, TAN);  ibid.loc., Antilahimena& Edmond3863( MICH, MO, P, TAN);  ibid.loc., 18°48’28”S 48°20’11”E,  1049 m,  2.XII.2005,  Antilahimena et al.4324( G, MO, P, TAN);  ibid.loc., 18°48’29”S 48°20’21”E,  1104 m,  21.XII.2005,  Antilahimena4513( G, K, MO, P, TAN);  ibid.loc., 18°51’01”S 48°18’30”E,  1119 m,  6.III.2008,  Antilahimena et al.6140( MICH, MO, P, TAN),  18°51’01”S 48°18’30”E,  1119 m,  6.V.2008,  Antilahimena et al.6144( MICH, MO, P, TAN);  ibid.loc., 18°51’11”S 48°19’05”E,  1095 m,  27.X.2008,  Antilahimena et al.6765( MO, TAN);  ibid.loc., 18°51’00”S 48°18’29”E,  1114 m,  12.XI.2009,  Antilahimena& Edmond7175( MICH, MO, P, TAN);  ibid. loc., 18°48’34”S 48°19’08”E,  1080 m,  5.X.2007,  Bernard& Phillipson596( MICH, MO, P, TAN);  ibid.loc., 18°48’32”S 48°20’42”E,  11.X.2007,  Bernard& Phillipson619( MICH, MO, P, TAN);  ibid.loc., 18°51’14”S 48°18’56”E,  1114 m,  18.I.2010,  Bernard et al.1532( MO, P, TAN);  ibid.loc., 18°48’12”S 48°21’56”E,  995 m,  3.VI.2007,  Miandrimanana et al.201( MO, P, TAN);   22.II.1965,  Peltier& Peltier5169( P);  ibid.loc., Peltier& Peltier5994( P);  18°48’31”S 48°20’39”E,  1115 m,  6.X.2007,  Phillipson et al.6034( MICH, MO, TAN);  ibid.loc., 18°51’08”S 48°18’40”E,  1121 m,  30.I.1997,  Rakotomalaza et al.1024( MO, P, TAN);  ibid.loc., 18°48’28”S 48°20’04”E,  1083 m,  4.IV.2005,  Rakotovao& Edmond1801( MO, TAN);  18°48’22”S 48°20’13”E,  1075 m,  3.VIII.2005,  Rakotovao et al.1961( MICH, MO, P, TAN);  18°48’29”S 48°19’17”E,  1122 m,  16.I.2005,  Ranaivojaona et al.1101( MICH, MO, P, TAN);  ibid.loc., 18°50’04”S 48°17’46”E,  1107 m,  15.II.2005,  Razanatsoa et al.113( MICH, MO, P, TAN);  ibid.loc., 18°51’06”S 48°18’40”E,  1114 m,  18.II.2005,  Razanatsoa et al.187( G, MO, P, TAN);  ibid.loc., 18°51’24”S 48°19’02”E,  1114 m,  26.IX.2005,  Razanatsoa et al.401( G, MO, P, TAN);  18°49’20”S 48°19’40”E,  1132 m,  20.X.2005,  Razanatsoa& Razafindasy548( MO, P, TAN);  ibid.loc.,  4.III.1987,  Service Forestier31265( TEF);  ibid.loc., 18°48’55”S 48°19’28”E,  1114 m,  21.III.2016,  van Ee et al.2437C( MICH, MO, P, TAN);  ibid.loc., van Ee et al.2438( MICH, MO, P, TAN);  ibid.loc., 18°48’28”S 48°20’11”E,  1050 m,  22.III.2016, v an Ee et al.2454 ( MICH, MO, P, TAN);  ibid.loc., van Ee et al.2455( MICH, MO, P, TAN). 3420017433 2016-03-21 MICH van Ee & Antilahimena & Kainulainen & Berry Madagascar Moramanga Distr. 1142 -18.850555 Berano 21 48.308056 Andasibe 10 337 MICH1513194 1 Prov. Toamasina holotype 3420017358 2016-03-21 G, K, MAPR, MO, P, TAN van Ee & Antilahimena & Kainulainen & Berry Madagascar Moramanga Distr. 1142 -18.850555 Berano 21 48.308056 Andasibe 10 337 1 Prov. Toamasina isotype 3420017411 2005-02-09 G, K, MO, P, TAN Antilahimena et al. Madagascar Moramanga distr. 1137 -18.84639 Ambatovy-Analamay forest 21 48.307777 11 338 1 Prov. Toamasina paratype 3420017450 2005-09-30 MICH, MO, P, TAN Antilahimena & Edmond Madagascar Moramanga distr. 1113 -18.8575 Ambatovy-Analamay forest 21 48.31861 11 338 1 Prov. Toamasina paratype 3420017347 2005-09-30 MICH, MO, P, TAN Antilahimena & Edmond Madagascar Moramanga distr. 1113 -18.8575 Ambatovy-Analamay forest 21 48.31861 11 338 1 Prov. Toamasina paratype 3420017368 2005-12-02 G, MO, P, TAN Antilahimena et al. Madagascar Moramanga distr. 1049 -18.807777 Ambatovy-Analamay forest 21 48.336388 11 338 1 Prov. Toamasina paratype 3420017379 2005-12-21 G, K, MO, P, TAN Antilahimena Madagascar Moramanga distr. 1104 -18.808054 Ambatovy-Analamay forest 21 48.339165 11 338 1 Prov. Toamasina paratype 3420017385 2008-03-06 MICH, MO, P, TAN Antilahimena et al. Madagascar Moramanga distr. 1119 -18.850279 Ambatovy-Analamay forest 21 48.308334 11 338 1 Prov. Toamasina paratype 3420017414 2008-05-06 MICH, MO, P, TAN Antilahimena et al. Madagascar Moramanga distr. 1119 -18.850279 Ambatovy-Analamay forest 21 48.308334 11 338 1 Prov. Toamasina paratype 3420017417 2008-10-27 MO, TAN Antilahimena et al. Madagascar Moramanga distr. 1095 -18.853056 Ambatovy-Analamay forest 21 48.318054 11 338 1 Prov. Toamasina paratype 3420017447 2009-11-12 MICH, MO, P, TAN Antilahimena & Edmond Madagascar Moramanga distr. 1114 -18.85 Ambatovy-Analamay forest 21 48.308056 11 338 1 Prov. Toamasina paratype 3420017454 2007-10-05 MICH, MO, P, TAN Bernard & Phillipson Madagascar Moramanga distr. 1080 -18.809444 Ambatovy-Analamay forest 21 48.31889 11 338 1 Prov. Toamasina paratype 3420017400 2007-10-11 MICH, MO, P, TAN Bernard & Phillipson Madagascar Moramanga distr. 1080 -18.808887 Ambatovy-Analamay forest 21 48.344997 11 338 1 Prov. Toamasina paratype 3420017418 2010-01-18 MO, P, TAN Bernard et al. Madagascar Moramanga distr. 1114 -18.85389 Ambatovy-Analamay forest 21 48.315556 11 338 1 Prov. Toamasina paratype 3420017396 2007-06-03 MO, P, TAN Miandrimanana et al. Madagascar Moramanga distr. 995 -18.803333 Ambatovy-Analamay forest 21 48.365555 11 338 1 Prov. Toamasina paratype 3420017472 [116,450,1016,1036] 1965-02-22 P Peltier & Peltier Madagascar Moramanga distr. 995 -18.803333 Ambatovy-Analamay forest 21 48.365555 11 338 1 Prov. Toamasina paratype 3420017349 [459,764,1016,1036] 1965-02-22 P Peltier Madagascar Moramanga distr. 995 -18.803333 Ambatovy-Analamay forest 21 48.365555 11 338 1 Prov. Toamasina paratype 3420017428 2007-10-06 MICH, MO, TAN Phillipson et al. Madagascar Moramanga distr. 1115 -18.80861 Ambatovy-Analamay forest 21 48.344166 11 338 1 Prov. Toamasina paratype 3420017444 1997-01-30 MO, P, TAN Rakotomalaza et al. Madagascar Moramanga distr. 1121 -18.852222 Ambatovy-Analamay forest 21 48.31111 11 338 1 Prov. Toamasina paratype 3420017333 2005-04-04 MO, TAN Rakotovao & Edmond Madagascar Moramanga distr. 1083 -18.807777 Ambatovy-Analamay forest 21 48.334442 11 338 1 Prov. Toamasina paratype 3420017372 2005-08-03 MICH, MO, P, TAN Rakotovao et al. Madagascar Moramanga distr. 1075 -18.80611 Ambatovy-Analamay forest 21 48.336945 11 338 1 Prov. Toamasina paratype 3420017309 2005-01-16 MICH, MO, P, TAN Ranaivojaona et al. Madagascar Moramanga distr. 1122 -18.808054 Ambatovy-Analamay forest 21 48.32139 11 338 1 Prov. Toamasina paratype 3420017346 2005-02-15 MICH, MO, P, TAN Razanatsoa et al. Madagascar Moramanga distr. 1107 -18.834446 Ambatovy-Analamay forest 21 48.296112 11 338 1 Prov. Toamasina paratype 3420017431 2005-02-18 G, MO, P, TAN Razanatsoa et al. Madagascar Moramanga distr. 1114 -18.851667 Ambatovy-Analamay forest 21 48.31111 11 338 1 Prov. Toamasina paratype 3420017391 2005-09-26 G, MO, P, TAN Razanatsoa et al. Madagascar Moramanga distr. 1114 -18.856667 Ambatovy-Analamay forest 21 48.317223 11 338 1 Prov. Toamasina paratype 3420017470 2005-10-20 MO, P, TAN Razanatsoa & Razafindasy Madagascar Moramanga distr. 1132 -18.822224 Ambatovy-Analamay forest 21 48.327778 11 338 1 Prov. Toamasina paratype 3420017336 [257,693,1336,1356] 1987-03-04 TEF Service Forestier Madagascar Moramanga distr. 1132 -18.822224 Ambatovy-Analamay forest 21 48.327778 11 338 1 Prov. Toamasina paratype 3420017406 2016-03-21 MICH, MO, P, TAN van Ee et al. Madagascar Moramanga distr. 1114 -18.815277 Ambatovy-Analamay forest 21 48.324444 11 338 1 Prov. Toamasina paratype 3420017445 [250,692,1390,1409] 2016-03-21 MICH, MO, P, TAN van Ee et al. Madagascar Moramanga distr. 1114 -18.815277 Ambatovy-Analamay forest 21 48.324444 11 338 1 Prov. Toamasina paratype 3420017420 2016-03-22 MICH, MO, P, TAN v an Ee et al. Madagascar Moramanga distr. 1050 -18.807777 Ambatovy-Analamay forest 21 48.336388 11 338 1 Prov. Toamasina paratype 3420017426 [200,633,1443,1462] 2016-03-22 MICH, MO, P, TAN van Ee et al. Madagascar Moramanga distr. 1050 -18.807777 Ambatovy-Analamay forest 21 48.336388 11 338 1 Prov. Toamasina paratype