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