Croton tsiampiensis var. ankaranensis Radcl.-Sm. , Gen. Croton Madag. Comoro 110. 2016 Croton tsiampiensis var. macrophyllus Radcl.-Sm. , Gen. Croton Madag. Comoro 110. 2016 Croton tsiampiensis var. microphyllus Radcl.-Sm. , Gen. Croton Madag. Comoro 111. 2016 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 ZRBY Leandri Leandri Magnoliopsida Euphorbiaceae Croton Plantae Malpighiales 28 399 Tracheophyta species tsiampiensis     Lectotypus(designated here):  MADAGASCAR. Prov. Mahajanga: Tsiampihy, près de l’embouchure de la Soahanina,  15.X.1932,  Leandri311( P[ P00389521]!; isolecto-: P[ P00133302]!).   =     Croton tsiampiensisvar. ankaranensis Radcl.-Sm., Gen. Croton Madag. Comoro 110. 2016. Typus: MADAGASCAR. Prov. Antsiranana: Diana Reg., Massif de l’Ankarana,  4.XI.1990,  Bardot-Vaucoulon224(holo-: P[ P00123706]!), syn. nov.  =     Croton tsiampiensisvar. macrophyllus Radcl.-Sm., Gen. Croton Madag. Comoro 110. 2016. Typus: MADAGASCAR. Prov. Mahajanga: Reserve Naturelle Bemaraha, Ambodiriana, c.  9 kmE of Antsalova, 18º39’S 44º43’E,  100-125 m,  13-15.XII.1990,  Gillespie4139(holo-: K!; iso-: MICH!, MO), syn. nov.  =     Croton tsiampiensisvar. microphyllus Radcl.-Sm., Gen. Croton Madag. Comoro 111. 2016. Typus: MADAGASCAR. Prov. Antsiranana: Diana Reg., P.K. 10 de la route Diego Suarez-Orangea,  13.XII.1963,  Service Forestier22956(holo-: P[ P00123706]!), syn. nov .   Shrubsor trees 2-6 mtall, dichotomously branching, internodes sometimes so short as to give the appearance of whorled branches; in all parts covered in whitish to ferrugineous, stellate trichomes many of which have a bristly, porrect (to c. 4 mm) central radius. Branches flattened on new growth but becoming terete with age, green at first, soon turning a matte pale gray (to reddish brown); red latex present in freshly cut stems. Stipules 8-30 × 1-5 mm, lanceolate, entire or incised, with marginal glands, caducous. Leavesdeciduous, opposite. Petioles 1.2-12 cmlong, adaxially canaliculate, stellatepubescent, usually with a pair of acropetiolar, ± stipitate glands with a concave, discoid, yellow apex ( 0.5-1 mmdiam.). Leaf blades membranaceous to papyraceous, glandular-denticulate, broadly ovate to suborbicular, 3-18 × 3-15 cm, apex acuminate, base rounded to cordate; abaxially and adaxially conspicuously hirsute, pale green when fresh and drying matte pale green to brown; venation evident on both sides, prominent below, with 5-9 pairs of brochidodromus, ± penninerved secondary veins (the lowermost pairs congested and appearing palmate), and cross-venulate tertiary venation; with ± stipitate glands in some of the axils of the secondary veins ( Fig. 1I). Inflorescencesterminal, raceme-like thryses 5-20 cmlong, with pistillate flowers toward the base and staminate flowers towards the distal end, axes stellate, flattened; bracts triangular, 1.0- 2.5 mmlong, caducous. Staminate flowerswith stellate, subglobose buds 1.5-2.9 mmdiam., pedicels 1-3 mmlong; sepals 5, shortly connate at base, lobes ovate, c. 2 × 1.2 mm, apex acute, inflexed at anthesis, abaxially stellate, adaxially glabrous, margins ciliate; petals 5, white, elliptic-spatulate, 2.5-2.7 × 1-1.5 mm, recurved at anthesis, abaxially subglabrous and papillose, adaxially glabrous, margins ciliate; disc glands 5, opposite the sepals, sessile, ellipsoidal with an apical depression, c. 0.4 × 0.3- 0.6 mm, yellowish; stamens 12-15, white, filaments 1.5- 3.3 mmlong, glabrous or pilose at base, anthers broadly elliptic, c. 0.5-1 × 0.5-0.8 mm; receptacle pilose. Pistillate flowerswith stellate, ellipsoid buds c. 2.2 mmdiam., pedicels 1-4 mmlong; sepals 5, ovate, spreading at anthesis, 2.5-3 × 1.3- 2.3 mm, apex acute, shortly connate at base, abaxially stellate, adaxially glabrous except towards the apex, persistent in fruit; petals often reduced, 0.8-2.5 2.5 × 0.2-1 mm, linear to spathulate, recurved, glabrous but with a ciliate margin; disc glands 5, opposite the sepals, sessile, ellipsoidal, c. 0.4 × 0.8 mm; ovary densely hirsute, globoid, 1.5-3.7 mmdiam; styles 3, 3- 4.2 mmlong, each branch flattened and 2-3 times bifurcate, spreading, abaxially stellate, adaxially glabrous, persistent. Capsules4-8 × 5-8 mm, densely hirsute, exocarp not separating ( Fig. 4O), endocarp woody, 0.4-0.6 mmthick; columella 3.5-5.5 mmlong, cornute, capitate. Seeds±compressed-ellipsoid, 3.5-5.6 × 3-4.3 × 2-2.9 mm( Fig. 4P); testa glossy, verrucose, brown; caruncle narrowly reniform c. 0.7 × 2 mm.   Phenology. –This species has been collected only from October to March, presumably because it is deciduous during the remaining drier months of the year. Specimens in flower have been collected from October to February, and in fruit from October to January.   Distribution, habitat and ecology. – Croton tsiampiensishas a disjunct distribution in northern Antsirananaand southern MahajangaProvinces, respectively ( Fig. 2B). It has been collected in deciduous, semideciduous, and evergreen forests, as well as in anthropogenic savannas, at 45-500 min elevation, on sandy substrates and on tsingy limestone.   Conservation assessment.– This species appears to be widespread with fragmented populations in deciduous forests of northern and western Madagascar, including the the protected areas Ankarana, Beanka, Bemaraha, and Daraina. We therefore treat this species as “Near Threatened” [NT] according the IUCN Red List Categories and Criteria ( IUCN, 2012).   Notes. –The species epithet  tsiampiensisrefers to the typelocality, the Tsiampihy forestin Mahajanga Province.   Croton tsiampiensisis distinct for its broadly cordate leaves, large incised stipules, and very hirsute pubescence. It is superficially similar to  C. ankarensisLeandri, and Leandri’s description of that species was likely based on specimens representing different taxa, one of which is likely  C. tsiampiensis. The type of  C. ankarensis, Perrier de la Bâthie 9830from Ankara (Betsiboka Reg.), lacks laminar glands and has alternate leaves, as well as sparsely stellate-pubescent capsules. It is clearly different from  C. tsiampiensisand does not belong in the Adenophorus Group (it was placed in the Ankarensis Group by LEANDRI, 1939). The Leandri 103bis syntypefrom Bemaraha is nearly leafless, but the leaf scars are opposite. The pistillate flowers are much larger than in Perrier de la Bâthie 9830, but they conform to  C. tsiampiensis, as does the markedly hirsute and pale gray bark of the young branches. We therefore tentatively include Leandri 103bisin  C. tsiampiensis. The varieties  C. tsiampiensisvar. macrophyllusand var. microphyllusthat RADCLIFFE-SMITH (2016)described based on differences in leaf size are synonymized here. As with many of the new names in his manuscript, only single specimens are cited, and from our studies of a broader sample of specimens that are now available, it is evident that more variation in leaf size is found within, rather than between, populations. Nor is it meaningful to recognize  C. tsiampiensisvar. ankarensis, which RADCLIFFE-SMITH (2016)distinguished from the typevariety by its shorter petioles (< 1 cmvs> 1.5 cm) and inflorescences, as well as slightly larger fruits. We have collected large-leaved specimens (van Ee et al. 2371)from the Ankarana population with leaves up to 16 × 11 cmand petioles up to 11 cmlong, and Gillespie et al. 10696, also from Ankarana, is closer in inflorescence length and fruit size to the typeof  C. tsiampiensisthan to  C.var. ankarensis. There are currently no vernacular names recorded for  Croton tsiampiensis.   Additional specimens examined.–   MADAGASCAR. Prov. Antsiranana: Diana Reg., Ankarongana, Analafandro, 12°37’50”S 49°31’28”E,  46 m,  23.II.2006,  Andrianjafyet al. 1619( CNARP, MICH, MO, P, TAN);  Ankarana,  13.X.1990,  Bardot-Vaucoulon131( P);  ibid. loc.,  4.XI.1990,  Bardot-Vaucoulon224( P);  ibid. loc., 12°55’S 49°05’E,  150 m,  5.XI.1990,  Bardot-Vaucoulon252( P, TAN);  ibid. loc.,  19.I.1991,  Bardot-Vaucoulon392( P);  ibid. loc.,  200 m,  3.II.1960,  Cours& Humbert5614( P);  ibid. loc., 12°54’S 49°07’E,  150 m,  29.XI.1990,  Gillespie4085( MICH, MO);  ibid. loc., 12°55’23”S 49°05’09”E,  110 m,  2.XI.2012,  Gillespieet al. 10696( CAN, MICH, MO, TAN);  ibid. loc.,  30-350 m,  24.I-29.II.1960,  Humbert32654( P);  SAVA Reg., Vohemar Distr., Daraina, Antsahabe forest, 13°12’37”S 49°33’27”E,  520 m,  11.I.2004,  Nusbaumer902( G, MICH, MO, P);  ibid. loc., 13°12’36”S 49°33’43”E,  500 m,  21.I.2004,  Nusbaumer1076( G, MICH, MO);  ibid. loc., 13°12’35”S 49°33’38”E,  500 m,  5.XI.2006,  Randrianaivoet al.1400( CNARP, MICH, MO, P, TAN);  Ankarana National Park, Maeva trail, 12°57’23 “S 49°07’04”E,  128 m,  6.III.2016,  van Ee et al. 2370( MICH, TAN);  ibid. loc., 12°57’23 “S 49°07’00”E,  153 m,  6.III.2016,  van Ee et al. 2371( MICH, TAN);  ibid. loc., Benavony trail, 12°56’55 “S 49°07’39”E,  124 m,  7.III.2016,  van Ee et al. 2378( MICH, TAN).   Prov. Mahajanga: Melaky Reg., collines au N d’Antsalova, bord de la Soahanina,  18.X.1932,  Leandri281( K, P, TAN);  ibid loc.,  Leandri283( P);  Beanka, partie S, Ambinda-Est, 18°02’41 “S 44°30’08”E,  261 m,  24.XI.2011,  Gautier& Tahinarivony5718( G, MO);  Tsingy du Bemaraha,  4.X.1932,  Leandri103bis( K, P);  Beanka, partie S; Ambinda-Est, 18°02’59”S 44°30’17”E,  260 m,  11.XII.2011,  Nusbaumeret al. 3051( G, MICH). 3463906515 1932-10-15 P Leandri Madagascar Tsiampihy 28 399 P00389521 1 Prov. Mahajanga lectotype 3463906386 [174,465,1122,1146] 1932-10-15 P de la Soahanina Madagascar Mahajanga 28 399 P00133302 1 Mahajanga isolectotype 3463906504 1990-11-04 P Bardot-Vaucoulon MADAGASCAR Massif de l'Ankarana Diana Reg. 28 399 P00123706 1 Prov. Antsiranana 3463906317 1990-12-13 1990-12-15 1990-12-13 K Gillespie MADAGASCAR 113 -18.65 Reserve Naturelle Bemaraha 1274 44.716667 28 399 1 Prov. Mahajanga 3463906461 [212,531,1495,1519] MICH, MO 28 399 1 3463906381 1963-12-13 P Service Forestier MADAGASCAR Diana Reg 28 399 P00123706 1 Prov. Antsiranana 3463906535 2006-02-23 CNARP, MICH, MO, TAN Andrianjafy Madagascar Prov. 46 -12.630556 Ankarongana 21 49.524445 Diana Reg. 30 401 1 Prov. Antsiranana 3463906443 [824,1242,357,376] 1990-10-13 P Bardot-Vaucoulon Madagascar Ankarana 30 401 1 3463906351 1990-11-04 P Bardot-Vaucoulon Madagascar Ankarana 30 401 1 3463906392 1990-11-05 P, TAN Bardot- Vaucoulon Madagascar 150 -12.916667 Ankarana 1292 49.083332 30 401 1 3463906372 [1039,1425,410,430] 1991-01-19 P Bardot-Vaucoulon Madagascar 150 -12.916667 Ankarana 1292 49.083332 30 401 1 3463906309 1960-02-03 P Cours & Humbert Madagascar 200 -12.916667 Ankarana 1292 49.083332 30 401 1 3463906395 1990-11-29 MICH, MO Gillespie Madagascar 150 -12.9 Ankarana 1292 49.116665 30 401 1 3463906380 2012-11-02 CAN, MICH, MO, TAN Gillespie Madagascar 110 -12.923056 Ankarana 21 49.08583 30 401 1 3463906391 [899,1404,517,536] 1960-01-24 1960-02-29 1960-01-24 P Humbert Madagascar 190 -12.923056 Ankarana 21 49.08583 30 401 1 3463906349 2004-01-11 G, MICH, MO, P Nusbaumer Madagascar SAVA Reg. 520 -13.210278 Daraina 21 49.5575 Vohemar Distr. 30 401 1 3463906511 2004-01-21 G, MICH, MO Nusbaumer Madagascar Reg. 500 -13.21 Daraina 21 49.561943 Vohemar Distr. 30 401 1 3463906544 2006-11-05 CNARP, MICH, MO, P, TAN Randrianaivo Madagascar Reg. 500 -13.209722 Daraina 21 49.560555 Vohemar Distr. 30 401 1 3463906444 2016-03-06 MICH, TAN van Ee et al Madagascar 128 -12.956388 Maeva trail 21 49.117775 Ankarana National Park 30 401 1 3463906446 2016-03-06 MICH, TAN van Ee et al Madagascar 153 -12.956388 Maeva 21 49.116665 Ankarana National Park 30 401 1 3463906503 2016-03-07 MICH, TAN van Ee et al Madagascar 124 -12.948611 Benavony trail 21 49.1275 Ankarana National Park 30 401 1 3463906377 1932-10-18 K, P, TAN Leandri Madagascar Melaky Reg. Prov. 30 401 1 Prov. Mahajanga 3463906368 [824,1059,810,830] 2011-11-24 P, G, MO Leandri Madagascar Leandri 261 -18.044722 Beanka 21 44.502224 Beanka 30 401 1 Mahajanga 3463906364 2011-11-24 P, G, MO Gautier & Tahinarivony Madagascar Leandri 261 -18.044722 Beanka 21 44.502224 Beanka 30 401 1 Mahajanga 3463906407 [824,1305,864,883] 1932-10-04 K, P Leandri Tsingy du Bemaraha 30 401 1 3463906445 2011-12-11 G, MICH Nusbaumer Madagascar 260 -18.04972 Beanka 21 44.504723 30 401 1