Research Article |
Corresponding author: Paul E. Berry ( peberry@umich.edu ) Academic editor: Sandy Knapp
© 2017 Paul E. Berry, Kent Kainulainen, Benjamin W. van Ee.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Berry PE, Kainulainen K, van Ee BW (2017) A Nomenclator of Croton (Euphorbiaceae) in Madagascar, the Comoros Archipelago, and the Mascarene Islands. PhytoKeys : 1-87. https://doi.org/10.3897/phytokeys.90.20586
|
All published names of Croton from Madagascar, the Comoros, and the Mascarenes are treated here. We indicate which names are currently accepted (123 native species and 1 introduced), which ones we consider to be heterotypic synonyms (188), which ones are doubtful (25), and which ones should be excluded (5). We newly designate lectotypes for 108 names, and epitypes for C. anisatus Baill., C. nobilis Baill., and C. submetallicus Baill. A total of 133 names are newly treated as synonyms. One new combination is made, Croton basaltorum (Leandri) P.E.Berry for C. antanosiensis var. basaltorum Leandri, and one new name is proposed, Croton toliarensis B.W.vanEe & Kainul. for C. tranomarensis var. rosmarinifolius Radcl.-Sm.
Euphorbiaceae , Croton , Madagascar, Comoros, Mascarenes, nomenclator, synonymy
This work is part of an effort to lay the taxonomic foundation for a thorough phylogenetic-based revision of Croton (Euphorbiaceae) in the Western Indian Ocean Region (WIOR). Madagascar alone harbors around one third of the roughly 450 species recognized to occur in the Old World (
Until 2016, taxa of Croton native to the Comoros, Madagascar, and the Mascarenes were published by just eight botanists in the following 27 books or journal articles:
The main herbaria consulted for the WIORCroton types were G, K, MICH, MAO, MO, P, TAN, and TEF. Scanned images of types from these and other herbaria available on JSTOR Global Plants (http://plants.jstor.org/) were also consulted. All original literature sources were also reviewed.
Specimens are considered holotypes when a single gathering in a particular herbarium is cited in the protologue, and there is only one specimen of that gathering housed there, or if there was a single gathering with no herbarium mentioned in the protologue, or else a single specimen exists in the herbarium where the author was based (ICN Art. 9.1,
Since the name Croton is derived from the Greek κροτων (tick), which is masculine, this is the proper gender for Croton. However, it was sometimes treated as neuter or as feminine by Baillon and Leandri, and this now requires that the epithets with those endings be orthographically corrected to the masculine ending. In such cases, the original spelling of an epithet is given, and users should be aware of the need to search databases using both the original spelling and subsequent, corrected spellings.
Within the taxon citations, if a lectotype is designated among the extant syntypes, the remaining syntypes are also cited. Type locality information is taken directly from the type specimen labels, even if it does not entirely agree with the location given in the protologue. If we add information not present on the specimen labels, such as the province name, or a more modern or accepted spelling of a place name, that information is placed within brackets. Barcode numbers of type specimens are cited when available, but as of late October 2017, the type specimens cited by
In the present treatment we recognize 114 accepted species native to Madagascar. All of these species are endemic to the island, with the exception of C. adenophorus Baill., which also occurs on Mayotte in the Comoros. Besides this species, there are four species endemic to the Comoros (C. bifurcatus Baill., C. emeliae Baill., C. humblotii Baill., and C. mayottae P.E.Berry & Kainul.). Another four species are endemic to Mauritius (C. fothergillifolius Baill., C. grangerioides Bojer ex Baill., C. tiliifolius Lam., and C. vaughanii Croizat), and a fifth (C. mauritianus Lam.) is endemic to Réunion. Croton bonplandianus Baill., a native of South America, is the only documented non-native, naturalized species of Croton in the WIOR, having previously been reported from the Mascarenes (Mauritius, Réunion, and Rodrigues;
For the 114 species that are native to Madagascar, only six are truly widespread throughout the island, occurring in all six former provinces, namely C. catatii Baill., C. goudotii Baill., C. macrobuxus Baill., C. mongue Baill., C. myriaster Baker, and C. stanneus Baill. Three others occur in five provinces (C. chrysodaphne Baill., C. hypochalibaeus Baill., and C. nitidulus Baker), five in four provinces, six in three provinces, and 29 in two provinces. Sixty-five species are known from just a single province. Of the six provinces, Toliara is the richest in Croton species with 53, followed by Antsiranana with 43, Toamasina with 39, Mahajanga with 35, Fianarantsoa with 27, and Antananarivo with 15. While we realize that these six administrative provinces of Madagascar have been superseded by a system of 22 administrative regions that represent subdivisions of the provinces, we did not attempt to further refine the distribution of species, although this will be a promising avenue to pursue in the future for conservation planning efforts. See Fig.
Given the small size of the islands in the Mascarenes and the Comoros Archipelago, and the small area of existing natural habitats, we believe that it is fairly unlikely that any additional Croton taxa will be discovered there. On the other hand, Madagascar harbors a large number of undescribed species, which are currently under study by the authors and awaiting formal description. We anticipate the description of at least 20 more new species from Madagascar based on the material we now have on hand. Many of these come from recently explored areas in the eastern part of the island, but also from relatively well-explored areas in the north and south but belonging to difficult species complexes, such as the coppery, lepidote-leaved species.
The sometimes extensive synonymy presented in this nomenclator (viz., C. adenophorus, C. catatii, C. chapelieri, or C. stanneus) suggests that many species have been poorly understood in the past, sometimes known only from sparse descriptions or poor type material, and numerous taxa are based on a single or just a few specimens. For instance, from
Another instructive example of a previously misunderstood species is Croton chrysodaphne Baill., a tree species that
Diversity of flowers in coppery-lepidote tree species of Croton from Madagascar that are vegetatively very similar. A Croton argyrodaphne, with leaves that are similar to those of several other species B Part of an inflorescence of Croton nobilis showing pistillate flower (below) with thick, reduplicate sepals and no petals, and staminate flower (above) with an intermediate number of stamens (ca. 18) C Staminate flower of C. chrysodaphne, with numerous (ca. 40) stamens and the unusual feature of ten (vs. normally five) petals D Pistillate flower of C. chrysodaphne, with patent, slender bifurcating styles and no petals E Staminate flowers of C. argyrodaphne, with only 11 stamens F Pistillate flower of C. argyrodaphne, with a stylar column topped by tightly bunched, short stigmas and also with recurved petals between the sepals (typically the pistillate flowers of this species are apetalous) G Base of an inflorescence of C. multicostatus showing three open pistillate flowers at the base (with well-developed, ligulate petals) and several open staminate flowers showing a low stamen number of 10 or 11. Photos by P. Berry.
Distribution of the number of native species of Croton in the Western Indian Ocean Region. There are 123 native species overall in the region, with 114 native to Madagascar; one of them (C. adenophorus) is shared with Mayotte, and the single species in the Union of the Comoros (C. humblotii) also occurs on Mayotte. The species on Mauritius and Réunion only occur there. The map of Madagascar shows how many species occur in each of the six former provinces (there are varying levels of overlap between provinces; see text for details).
The recent, hastily published work of
The overall disposition of the taxa described by
New species and subspecies of Malagasy Croton described by
|
As treated in this paper |
---|---|
C. adenophoroides Radcl.-Sm. | C. loucoubensis Baill. |
C. alaotrensis Radcl.-Sm. | C. droguetioides Kainul. & Radcl.-Sm. |
C. alceicornu Radcl.-Sm. | C. hypochalibaeus Baill. |
C. alchorneifolius Radcl.-Sm. | C. alchorneifolius Radcl.-Sm. |
C. bracteatus subsp. manongarivensis Radcl.-Sm. | C. nitidulus Baill. |
C. bracteatus subsp. populifolius Radcl.-Sm. | C. danguyanus Leandri |
C. commiphoroides Radcl.-Sm. | C. indrisilvae Kainul., B.W.vanEe & P.E.Berry |
C. daphniphylloides Radcl.-Sm. | C. chapelieri Baill. |
C. daphniphyllus Radcl.-Sm. | C. chapelieri Baill. |
C. dasygyne Radcl.-Sm. | C. radiatus P.E.Berry & Kainul. |
C. delicatulus Radcl.-Sm. | C. menabeensis Leandri |
C. domohineifolius Radcl.-Sm. | C. chapelieri Baill. |
C. droguetioides Radcl.-Sm. | C. droguetioides Kainul. & Radcl.-Sm. |
C. echinatus Radcl.-Sm. | C. dissimilis Baill. |
C. gracilior Radcl.-Sm. | C. gracilior Radcl.-Sm. |
C. hirsutissimus Radcl.-Sm. | C. nudatus Baill. |
C. lepidotoides Radcl.-Sm. | C. ferricretus Kainul., B.W.vanEe & P.E.Berry |
C. minimimarginiglandulosus Radcl.-Sm. | C. minimimarginiglandulosus Radcl.-Sm. |
C. mocquerysii subsp. meridionalis Radcl.-Sm. | C. thouarsianus Baill. |
C. neoholstiifolius Radcl.-Sm. | C. menabeensis Leandri |
C. oligostemon Radcl.-Sm. | C. hypochalibaeus Baill. |
C. parietarioides Radcl.-Sm. | C. droguetioides Kainul. & Radcl.-Sm. |
C. parvifructus Radcl.-Sm. | C. stanneus Baill. |
C. remotiflorus Radcl.-Sm. | C. trichotomus Geisel. |
C. rhododendroides Radcl.-Sm. | C. chapelieri Baill. |
C. scorpistogyne Radcl.-Sm. | C. heteranthus Aug.DC. |
C. submetallicoides Radcl.-Sm. | C. chyrsodaphne Baill. |
C. ustulatus Radcl.-Sm. | C. ustulatus Radcl.-Sm. |
C. vohemarensis Radcl.-Sm. | C. argyrodaphne Baill. |
New varieties of Malagasy Croton described by
|
As treated in this paper |
C. adabolavensis var. hippophaëoïdes | C. adabolavensis Leandri |
C. adabolavensis var. microlepidotus | C. adabolavensis Leandri |
C. adabolavensis var. ovalifolius | C. adabolavensis Leandri |
C. adabolavensis var. stellatipilus | C. adabolavensis Leandri |
C. alaotrensis var. integrifolius | C. dissimilis Baill. |
C. ambovombensis var. lepidotus | nom. inval. (Excluded Taxa) |
C. anisatus var. hirsutus | C. hovarum Leandri |
C. ankarensis var. ankarafantsikae | Incertae Sedis |
C. anosiravensis var. pilosus | C. anosiravensis Leandri |
C. antanosiensis var. ambohibyi | C. trichotomus Geisel. |
C. antanosiensis var. fianarantsoae | C. hypochalibaeus Baill. |
C. antanosiensis var. pubescens | C. greveanus Baill. |
C. bastardii var. bongolavae | Incertae Sedis |
C. bastardii var. meridionalis | C. muricatus Vahl |
C. bathianus var. ambatondrazakae | C. scoriarum Leandri |
C. bathianus var. ihosianus | C. ihosianus Leandri |
C. bathianus var. toliarae | C. crocodilorum Baill. |
C. bemaranus var. parvistipulatus | Incertae Sedis |
C. bemaranus var. pseudolepidotus | C. aleuritoides P.E.Berry |
C. betiokensis var. haplostylis | Incertae Sedis |
C. bifurcatus var. humblotii | C. bifurcatus Baill. |
C. boinensis var. parcelepidotus | C. greveanus Baill. |
C. boinensis var. tomentosus | C. catatii Baill. |
C. boivinianus var. brevifolius | C. nudatus Baill. |
C. cassinoides var. alaotrensis | C. hovarum Leandri |
C. catatii var. schizoplepis | Incertae Sedis |
C. catatii var. setosus | C. catatii Baill. |
C. catatii var. tricholepis | C. catatii Baill. |
C. chapelieri var. longepetiolata | C. submetallicus Baill. |
C. chrysodaphne var. meridionalis | C. cupreolepis P.E.Berry, B.W.vanEe, & Kainul. |
C. crocodilorum var. meridionalis | C. crocodilorum Leandri |
C. crocodilorum var. platyaster | C. stanneus Baill. |
C. daphniphylloides var. hirsutus | C. nitidulus Baker |
C. daphniphylloides var. stellatipilus | Incertae Sedis |
C. daphniphylloides var. triplinervius | Incertae Sedis |
C. daphniphyllum var. hirsutus | C. fianarantsoae Leandri |
C. decaryi var. subglaber | C. muricatus Vahl |
C. elaeagni var. antsirananae | C. elaeagni Baill. |
C. elaeagni var. argyrocarpos | Incertae Sedis |
C. elaeagni var. brevipedicellatus | Incertae Sedis |
C. elaeagni var. chrysocarpos | C. catatii Baill. |
C. fianarantsoae var. ambremontanus | C. minimimarginiglandulosus Radcl.-Sm. |
C. fianarantsoae var. coursii | C. fianarantsoae Leandri |
C. fianarantsoae var. masoalae | C. glomeratus Aug.DC. |
C. fianarantsoae var. microphyllus | C. fianarantsoae Leandri |
C. fianarantsoae var. obovalifolius | C. submetallicus Baill. |
C. fianarantsoae var. petiolaris | Incertae Sedis |
C. fianarantsoae var. ranomafanae | C. fianarantsoae Leandri |
C. fianarantsoae var. tandrokensis | C. fianarantsoae Leandri |
C. geayi var. paucisquamatus | C. geayi Leandri |
C. geayi var. pubescens | C. geayi Leandri |
C. goudotii var. tsaratananae | C. goudotii Baill. |
C. greveanus var. ambositrensis | C. hovarum Leandri |
C. greveanus var. micraster | C. catatii Baill. |
C. hovarum var. hirsutifructus | Incertae Sedis |
C. hovarum var. lepidotus | C. hovarum Leandri |
C. hovarum var. subglaber | C. hovarum Leandri |
C. incisus var. minor | C. incisus Baill. |
C. isomonensis var. microcarpus | C. trichotomus Geisel. |
C. ivohibensis var. aesculops | C. minimimarginiglandulosus Radcl.-Sm. |
C. ivohibensis var. alaotrensis | C. humbertii Leandri |
C. ivohibensis var. ankaranaënsis | C. minimimarginiglandulosus Radcl.-Sm. |
C. ivohibensis var. furfuraceus | C. heteranthus Aug.DC |
C. ivohibensis var. integrifolius | Incertae Sedis |
C. ivohibensis var. lepidotus | C. hovarum Leandri |
C. ivohibensis var. macrocalyx | Incertae Sedis |
C. ivohibensis var. polygynus | Incertae Sedis |
C. ivohibensis var. puncticulatus | C. bracteatus Lam. |
C. ivohibensis var. verticillatus | Incertae Sedis |
C. kimosorum var. pubescens | C. kimosorum Leandri |
C. leandri var. pubescens | C. toliarensis B.W.vanEe & Kainul. |
C. lichenisilvae var. oligostemon | C. lichenisilvae Leandri |
C. macrobuxus var. dolichobotrys | C. macrobuxus Baill. |
C. macrobuxus var. glandulifer | C. macrobuxus Baill. |
C. macrobuxus var. polygynus | C. chapelieri Baill. |
C. macrobuxus var. subfoliaceus | C. macrobuxus Baill. |
C. macrobuxus var. substrigosus | C. macrobuxus Baill. |
C. manampetsae var. angustifolius | C. manampetsae Leandri |
C. manampetsae var. chaetogyne | C. manampetsae Leandri |
C. manampetsae var. lepidotus | Incertae Sedis |
C. mavoravina var. concinnus | C. boiteaui Leandri |
C. mavoravina var. gracilis | C. mavoravina Leandri |
C. mavoravina var. gymnolepis | C. mavoravina Leandri |
C. mavoravina var. imanombensis | C. mavoravina Leandri |
C. mavoravina var. rotundifolius | C. mavoravina Leandri |
C. mavoravina var. thysanolepis | C. crossolepis P.E.Berry & Kainul. |
C. menabeensis var. furfuraceus | C. nudatus Baill. |
C. menarandrae var. pubescens | C. menarandrae Leandri |
C. meridionalis var. latifolius | Incertae Sedis |
C. meridionalis var. pseudolepidotus | C. meridionalis Leandri |
C. meridionalis var. stipularis | Incertae Sedis |
C. miarensis var. monadenius | C. miarensis Leandri |
C. mongue var. borealis | C. mongue Baill. |
C. nitidulus var. acuminatus | C. submetallicus Baill. |
C. nitidulus var. angustiglans | C. nitidulus Baker |
C. nitidulus var. bekolosiensis | C. nitidulus Baker |
C. nitidulus var. cinereus | C. submetallicus Baill. |
C. nitidulus var. eglandulosus | Incertae Sedis |
C. nitidulus var. fuscicrameus | C. nitidulus Baker |
C. nitidulus var. hypopoliotes | C. submetallicus Baill. |
C. nitidulus var. macrophyllus | C. submetallicus Baill. |
C. nitidulus var. microphyllus | C. macrobuxus Baill. |
C. nitidulus var. pubescens | C. macrobuxus Baill. |
C. oreades var. borealis | C. mongue Baill. |
C. oreades var. craspedadenius | C. mongue Baill. |
C. oreades var. periphoradenius | C. mongue Baill. |
C. peltieri var. hazofotsiensis | C. miarensis Leandri |
C. regeneratrix var. mayottensis | C. mayottae P.E.Berry & Kainul. |
C. regeneratrix var. ranomafanae | C. myriaster Baker |
C. rubricapitirupis var. macrophyllus | Incertae Sedis |
C. stanneus var. hirsutus | C. stanneus Baill. |
C. subaemulans var. tsingyensis | C. bemaranus Leandri |
C. submetallicus var. tomentosus | C. submetallicus Baill. |
C. thuarsianus var. angustifolius | Incertae Sedis |
C. thuarsianus var. longifolius | Incertae Sedis |
C. thuarsianus var. macrocalyx | C. minimimarginiglandulosus Radcl.-Sm. |
C. thuarsianus var. robustior | C. thouarsianus Baill. |
C. tranomarensis var. rosmariniifolius | C. toliarensis B.W.vanEe & Kainul. |
C. tsiampiensis var. ankaranensis | C. tsiampiensis Leandri |
C. tsiampiensis var. macrophyllus | C. tsiampiensis Leandri |
C. tsiampiensis var. microphyllus | C. tsiampiensis Leandri |
Croton adabolavensis var. hippophaëoïdes Radcl.-Sm., Gen. Croton Madag. Comoro 92. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Cap Ste. Marie Reserve, SW of Tsiombe, 25°35'S, 45°09'E, 27 Dec 1988, P.B. Phillipson 3008 (holotype: K!; isotypes: MO!, P [P00433074]!).
Croton adabolavensis var. microlepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 92. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Réserve Naturelle Intégrale d’Andohahela, along border road, 25°00'S, 46°40'E, 19 Oct 1990, A. Randrianasolo, D. Faber-Langendoen, N. Dumetz & R. Rabevohitra 174 (holotype: K!; isotypes: MO!, P [P00433039]!).
Croton adabolavensis var. ovalifolius Radcl.-Sm., Gen. Croton Madag. Comoro 92. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Beza Mahafaly Reserve, near Betioky, ridge E of Sakamena River, valley of Analafahy River, 23°58'S, 44°39'E, 28 Nov 1987, P.B. Phillipson 2621 (holotype: K!; isotype: MO!, P [P00133395]!).
Croton adabolavensis var. stellatipilus Radcl.-Sm., Gen. Croton Madag. Comoro 93. 2016, syn. nov. Type. Madagascar. Prov. Toliara: 27 km from Tulear (Toliara), 23°21'S, 43°51'E, 30 Dec 1987, P.B. Phillipson 2759 (holotype: K!; isotype: MO!, P [P00133397]!).
Madagascar. Prov. Toliara: Anosy Region, vallée moyenne du Mandrare, près d’Anadabolava, forêt sèche, 200–250 m, Dec 1933, H. Humbert 12498 (lectotype, designated here: P [P00248927]!; isolectotype: P [P00312384]!).
Shrubs; southeastern Madagascar (Toliara).
Croton payerianus Baill., Adansonia 1: 154. 1861, as ‘payerianum’. Type. Madagascar. Prov. Antsiranana: Diana Region, Nossibé, 1849, L.H. Boivin 2187 (lectotype, designated here: P [P00389508]!; isolectotypes: G-DC [G00311984]!, G [G00446358]!, G [G00446359]!, P [P00389509]!, P [P00404480]!, P [P00404481]!, P [P00123684]!).
Croton
tulasnei
Baill., Adansonia 1: 156. 1861. Type. Mayotte [French Overseas Department]: Bouzi [Ilot M’Bouzi], 1850, L.H. Boivin s.n. (lectotype, designated by
Croton subaemulans Baill., Bull. Mens. Soc. Linn. Paris 2: 850. 1890. Type. Madagascar. sin. loc., R. Baron 5795 (holotype: K [K000422590]!; isotype: P [P00133593]!).
Oxydectes adenophora (Baill.) Kuntze, Revis. Gen. Pl. 2: 610. 1891. Type. Based on Croton adenophorus Baill.
Oxydectes payeriana (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton payerianus Baill.
Oxydectes tulasnei (Baill.) Kuntze, Revis. Gen. Pl. 2: 613. 1891. Type. Based on Croton tulasnei Baill.
Croton tenuicuspis Baill., Bull. Mens. Soc. Linn. Paris 2: 927. 1891. Type. Madagascar. sin. loc., R. Baron 5846 (holotype: P [P00133364]!; isotype: K [K000422590]!).
Madagascar. Prov. Antsiranana: Diana Region, Nossi-bé [Nosy Be], 1837, J.M.C. Richard 214 (lectotype, designated by
Shrubs or small trees; northern and northwestern Madagascar (Antsiranana, Mahajanga) and Mayotte in the Comoros Archipelago.
Croton adenophorus was substantially recircumscribed from the concept of
The sheet P00466148 is listed in Sonnerat under Croton tulasnei, but without an image. We never saw this specimen at P either before or after the herbarium renovation in 2015, and the whole folder of C. tulasnei was missing during visits to P in 2016 and 2017.
Madagascar. Prov. Antsiranana: SAVA Region, summit of Marojejy on path from Mandena, 14°26'56"S, 49°43'58"E, 30 Sep 1994, B. Lewis, F. Rasoavimbahoaka & J. Rastefanonirina 1206 (holotype: K!; isotypes: MO!, P [P00131484]!).
Small trees; restricted to the Marojejy Massif in northern Madagascar (Antsiranana).
This is a distinctive high-elevation species with well-developed petals in the pistillate flowers and large capsules, belonging to the Mongue Group of
Croton bemaranus var. pseudolepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 209. 2016 [23 Dec 2016], as ‘bemarana’, syn. nov. Type. Madagascar. Prov. Antsiranana: Diana Region, Montagne des Français, à l’Est de Diego Suarez, 26 Nov 1958, Service Forestier 20088 (lectotype, designated here: P [P00312410]!; isolectotypes: B, G [G00341674]!, K [K001040349]!, MO!, P [P00206489]!, S!, TEF!, WAG).
Madagascar. Prov. Antsiranana: Diana Region, Montagne des Français, à l’Est de Diégo-Suarez, 26 Nov 1958, Service Forestier 20088 (holotype: P [P00312410]!; isotypes: B, G [G00341674]!, K [K001040349]!, MO!, P [P00206489]!, S!, TEF!, WAG).
Trees; known only from Montagne des Français in northernmost Madagascar (Antsiranana).
Oxydectes ambanivoulensis (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton ambanivoulensis Baill.
Madagascar. Prov. Toamasina: [region of the] Ambanivoules, [west of] Tamatave, dans les forêts, Dec. 1836, J.P. Goudot s.n. (lectotype, designated here: G [G00446336]!; isolectotype: P [P00301514]!).
Shrubs; eastern lowland Madagascar (Toamasina).
There are two sheets of Goudot s.n. at G within the same jacket, meaning they are considered as being part of the same collection (see
Ambanivoules were an ethnic group of eastern Madagascar located approximately 80–100 km west of Tamatave, their name derived from the Malagasy “Antanbanivolo,” or “people living at the base of the mountains covered with bamboos” (Schatz 2013).
Croton divaricatus Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 59. 1939, as ‘divaricata’, nom. illeg. non Croton divaricatus Sw., 1788.
Madagascar. Prov. Toliara: Androy Region, Ambovombe, 19 Dec 1924, R. Decary 3174 (lectotype, designated by
Shrubs; southern lowland Madagascar (Toliara).
Croton geayi var. androiensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 35. 1939. Type. Madagascar. Prov. Toliara: Androy/Anosy Regions, Bevilany, limite entre l’Anosy et l’Androy, 14 Nov 1932, R. Decary 10940 (lectotype, designated here): P [P00123697]!; isolectotypes: G, K [K001044841]!, TAN [TAN000533]!). Madagascar. Prov. Toliara: Androy Region, Ambovombe, 6 Jan 1931, R. Decary 8361 (syntypes: P [P00123696]!), TAN [TAN000532]!). Madagascar. Prov. Toliara: Kotoala, sables et dunes littorales, 21 Jan 1931, R. Decary 8408 (syntypes: K [K001040395]!, MO [sheet #04861163]!, P [P00404477]!, P [P00123698]!).
Based on Croton geayi var. androiensis Leandri
Shrubs; southern Madagascar (Toliara).
Oxydectes anisata (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton anisatus Baill.
[Cult. ex Madagascar]: Cultivated in the garden of M. Hubert, Saint Benoît, Réunion, s.d., M. Lepervanche s.n. (holotype: P [P00301513]!). Madagascar. Prov. Toamasina: Atsinanana Region, Vohibola, N to NNW of village of Andranokoditra, N of Lac Ampitabe, 18°33'34"S, 49°15'01"E, 5 m, 12 Feb 2003, P. Lowry, R. Rabevohitra, J. Rabenantoandro, R. Razakamalala & S.W.J. Lowry 6072 (epitype, designated here): P [P00548219]!; additional duplicates: DAV!, K!, MICH [MICH1210799]!, MO [sheet # 5902002]!).
Shrubs; eastern coast of Madagascar (Toamasina).
Since the holotype of Croton anisatus has only young inflorescences in bud and is known only from cultivation on a quite different island, we designate an epitype with open flowers. This littoral species can be characterized by its very congested inflorescences and the pseudoverticillate, anisophyllous, and sparsely lepidote-pubescent leaves with an entire margin. The plant is apparently quite aromatic as indicated by the descriptions of both the holotype and epitype, as well as by its specific epithet.
Madagascar. Prov. Mahajanga: Causse d’Ankara, bois rocailleux et secs sur calcaire jurassique, Dec 1900, H. Perrier de la Bâthie 9830 (lectotype, designated here: P [P00123702]!; isolectotype: P [P00123703]!). Madagascar. Prov. Mahajanga: Tsingy du Bemaraha (9e Réserve), 4 Oct 1932, J. Leandri 103bis (syntypes: K [K001040394]!, P [P00123700]!, P [P00123701]!).
Shrubs; western Madagascar (Mahajanga).
The Leandri 103bis syntype corresponds to Croton tsiampiensis.
Madagascar. Prov. Toamasina: Atsinanana Region, District Brickaville, Commune Maroseranana, Fokontany Ambodilendemy, Andrangato River, 18°26'37"S, 48°46'31"E, 446 m, 13 Mar 2011, P. Antilahimena 7554 (holotype: MICH [MICH1513200]!; isotypes: MO!, P!, TAN!).
Shrubs to small trees; eastern Madagascar (Toamasina).
Croton anosiravensis var. pilosus Radcl.-Sm., Gen. Croton Madag. Comoro 207. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Analamera, 50-400 m, Jan 1938, H. Humbert 19141 (lectotype, designated here: P [P00133307]!; isolectotype: P [P00133308]!).
Madagascar. Prov. Antsiranana: Base des escarpements de l’Anosiravo, poteau kilométrique 6 de la route de Diego Suarez à Orangéa, 12 Dec 1963, Service Forestier 22930-SF (holotype: P [P00404432]!; isotypes: K [K001040369]!, TEF [TEF000193]!).
Shrubs; northern Madagascar (Antsiranana).
In its stellate-pubescent and cordate leaves, Croton anosiravensis is superficially similar to some of the species in the Adenophorus Group. However, it does not have opposite leaves or laminar glands, and is probably not closely related. It appears to be a rare species, because besides the type from the northern slopes of Montagne des Français, it is otherwise only known from Analamera (Humbert 19141 [P]) and Befarafara in Daraina (Rakotonandrasana et al. 1048 [CNARP, MICH, MO, P, TAN]).
Madagascar. Prov. Toliara: environs de Fort-Dauphin, près de Bévilany, 200-300 m, 14 Sep 1928, H. Humbert & C.F. Swingle 5695 (lectotype, designated here: P [P00123716]!; isolectotype: P [P00123717]!). Madagascar. Prov. Toliara: Massif de Beampingaratra (Sud-Est), du col de Bevava au sommet de Bekoho, forêt sur latérite de gneiss, 1100-1500 m, 6-7 Nov 1928, H. Humbert 6416 (syntypes: P [P00123718]!, P [P00154301]!, TAN [TAN000521]!). Madagascar. Prov. Toliara: Fort Dauphin, J. Cloisel 60 (syntype: P [P00123710]!). Madagascar. Prov. Toliara: Behara, 9 Jul 1926, R. Decary 4321 (syntypes: P [P00123711]!, S [S07-14102]). Madagascar. Prov. Toliara: Ranofotsy, 29 Jul 1932, R. Decary 10175 (syntypes: G [G00446368]!, P [P00123712]!).
Shrubs to small trees; southern Madagascar (Toliara).
We believe that the lectotype chosen here best conforms to the protologue among the syntypes cited by
Oxydectes argyrodaphne (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton argyrodaphne Baill.
Croton argyrodaphne var. occidentalis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 44. 1939, syn. nov. Type. Madagascar. Prov. Mahajanga: région du Cap Saint-André, dunes, 8 Jun 1930, R. Decary 7893 (lectotype, designated here: P [P00127457]!). Madagascar. Prov. Toliara: bassin de la Tsiribihina, Jul 1911, H. Perrier de la Bâthie 9656 (syntypes: P [P00127460]!, P [P00127461]!, P [P00127462]!).] !). Madagascar. Prov. Toliara: Morondava, received 26 Dec 1878, H. Grevé 11 (syntype: P [P00127459]!).
Croton argyrodaphne var. boinensis Leandri, Adansonia, sér. 2, 12: 404. 1972, syn. nov. Type. Madagascar. Prov. Mahajanga: Ampalony, 9 Aug 1971, L.P. Schmitt 515 (holotype: P [P00389629]!).
Croton argyrodaphne var. orientalis Leandri, Adansonia, sér. 2, 12: 404. 1972, syn. nov. Type. Madagascar. Prov. Toamasina: Intendro, près de Fénérive, 9 Jul 1958, Service Forestier 19160-SF (holotype: P [P00418607]!).
Croton vohemarensis Radcl.-Sm., Gen. Croton Madag. Comoro 78. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Mantamena, Bekaroaka Range, 7 km N of Daraina (Vohemar), 13°08'S, 49°42'E, 112-330 m, 26 Nov 1990, D. Meyers 206 (holotype: MO!).
Madagascar. Prov. Antsiranana: Diana Region, Nossibé, 1837, J.M.C. Richard 218 (lectotype, first step designated by
Trees to large shrubs; mainly in northern Madagascar, but extending as far south as northern Toliara Province on the west coast and northern Toamasina on the east coast (Antsiranana, Mahajanga, Toamasina, Toliara).
The type of Croton vohemarensis consists of two small twigs with leaves that are unusually small and wide for C. argyrodaphne. However, the low stamen number (11) and the characteristic stylar column are very typical of C. argyrodaphne (see Fig.
What Leandri recognized as Croton argyrodaphne var. occidentalis is a rather distinctive element of this species, with a western, subcoastal distribution. The plants are small trees, and the leaves have yellowish pigmentation along the midvein on the adaxial leaf surfaces, but other than that they conform well to the general aspect of C. argyrodaphne.
Madagascar. Prov. Toliara: Sud-Ouest, s.d., M.G. Cours 4641 (holotype: P [P00312369]!; isotypes: K [K001044848], MO [sheet # 5737746]!), P [P00380443]!, P [P00380444]!).
Shrubs; southwestern Madagascar (Toliara).
Madagascar. Prov. Toliara: Antanimora, 16 Jun 1926, R. Decary 4346 (holotype: P [P00301486]!; isotype: K [K001044842]!).
Shrubs; southwestern Madagascar (Toliara).
Croton antanosiensis var. basaltorum Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 46. 1939. Type. Madagascar. Prov. Mahajanga: Pl. [Plateau] d’Antanimena, entre la Mahavavy et le Betsiboka, Jun 1906, H. Perrier de la Bâthie 9593 (lectotype, designated here: P [P00123726]!; isolectotype: P [P00123725]!).
Based on Croton antanosiensis var. basaltorum Leandri
Shrubs; western Madagascar (Mahajanga).
Geographically and morphologically, this species is sufficiently distinct from C. antanosiensis to merit recognition at the species level. Morphologically, it appears more similar to C. cupreolepis.
Croton appertii Leandri, Adansonia, sér. 2, 15: 331. 1976, syn. nov. Type. Madagascar. Prov. Toliara: Beharana, près de Manja, 300 m, 17 Nov 1961, O. Appert 37 (holotype: P [P00312368]!; isotypes: K [K001044847]!, MO [sheet # 2287361]!, Z [Z-000015970]!, Z [Z-000015971]!).
Madagascar. Prov. Toliara: forêt de Besomaty, entre le Fiherena et l’Isahaina (Mangoky), 750-800 m, Dec 1933, H. Humbert 11236 (lectotype, designated here: P [P00404492]!; isolectotypes: K [K001044843]!, P [P00127500]!).
Shrubs; southwestern Madagascar (Mahajanga, Toliara).
Madagascar. Prov. Mahajanga: Haut Bemarivo, Oct 1907, H. Perrier de la Bâthie 9545 (lectotype, designated by
Shrubs; northwestern Madagascar (Antsiranana, Mahajanga).
Croton subaemulans var. tsingyensis Radcl.-Sm., Gen. Croton Madag. Comoro 120. 2016, syn. nov. Type. Madagascar. Prov. Mahajanga: Tsingy du Bemaraha (9e Réserve), reçu Feb -Apr 1933, J. Leandri 116 (holotype: K!; isotypes: P [P00133368!, P00133369!]).
Madagascar. Prov. Mahajanga: causse d’Ankara, Oct 1900, H. Perrier de la Bâthie 1153 (lectotype, designated here: P [P00206491]!; isolectotypes: K [K001040396]!, P [P00389502]!). Madagascar. Prov. Mahajanga: Bemara, partie Nord de l’Antsingy, entre Ambatomiloloha et Anjohinomby, 29 Nov 1932, J. Leandri 656 (syntypes: P [P00127512]!, P [P00389503]!).
Shrubs; western and northern Madagascar (Antsiranana, Mahajanga).
Madagascar. Prov. Antsiranana: entre Andrangana et la rivière Anjambazamba, route de Sambava à Antsirabe-Nord, 2-7 Oct 1966, Service Forestier 27197-SF (holotype: P [P00312371!; isotypes: K [K001044846]!, MO [sheet #04861161]!, P [P00404483]!, TEF [TEF000192]!).
Shrubs; northeastern Madagascar (Antsiranana, Toamasina).
Madagascar. Prov. Toamasina: Menagisy, Brickaville, 11 Oct 1956, Service de Eaux et Forêts de Madagascar 12358-SF (lectotype, designated here: P [P00312375]!; isolectotypes: P [P00127519]!, TEF!).
Shrubs or small trees; eastern lowland Madagascar (Antsiranana, Toamasina).
Although the sheet designated here as lectotype has a sticker stating “TYPE” and the other sheet at P has one stating “ISOTYPE,” it is not clear who applied those labels and if it was done after Leandri's publication. In any case, there is no annotation in Leandri's hand on either sheet to indicate which of the two he intended to be the holotype.
Oxydectes bernieri (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton bernieri Baill.
Croton bernieri var. namorokensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 54. 1939, syn. nov. Type. Madagascar. Prov. Mahajanga: Tsingy de Namoroka (8e Réserve), 1933, Service Forestier 11 (holotype: P [P00127543]!).
Madagascar. Prov. Antsiranana: Diégo-Suarez, 1835, A.C.J. Bernier 306 (lectotype, designated here: P [P00127520]!; isolectotypes: G [G00434420]!, P [P00127521]!, P [P00301481]!). Madagascar. Prov. Antsiranana: Baie de Diégo-Suarez, Dec 1848, L.H. Boivin 2657 (syntypes: G [G00311744]!, G-DC [G00311744]!, P [P00127522]!, P [P00127523]!, P [P00127524]!, P [P00301480]!).
Shrubs to small trees; northern and northwestern Madagascar (Antsiranana, Mahajanga).
Madagascar. Prov. Toliara: Plateau Mahafaly à l’ouest de Betioky, 100-300 m, 17-20 Mar 1955, H. Humbert & R. Capuron 29489 (holotype: P [P00312386]!; isotypes: P [P00127545], P [P00127546]!).
Shrubs; southern Madagascar (Toliara).
Croton bifurcatus var. genuinus Müll.Arg. in A.P.de Candolle, Prodr. 16(2): 584. 1866, nom. inval.
Oxydectes bifurcata (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton bifurcatus Baill.
Croton bifurcatus var. humblotii Leandri ex Radcl.-Sm., Gen. Croton Madag. Comoro 25. 2016, syn. nov. Type. Mayotte [French Overseas Department]: forêt de Mazé, M. Bini [Majimbini], 24 May 1884, L. Humblot 1162 (holotype: K; isotype: P [P00196067]!).
Mayotte [French Overseas Department]: Cascade du Msapéré, 1849, L.H. Boivin 3380 (holotype: P [P00196066]!; isotypes: G-DC [G00311982]!), G [G00446373]!, P [P00196065]!, P [P00466147]!).
Shrubs; known only from the French island of Mayotte in the Comoro Islands.
The type of C. bifurcatus var. humblotii differs in only minor ways from typical C. bifurcatus.
Oxydectes bocquillonii (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton bocquillonii Baill.
Croton brevispicatus var. bocquillonii (Baill.) Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 27. 1939. Type. Based on Croton bocquillonii Baill.
Madagascar. Prov. Mahajanga: Ambongo, 16 Feb 1841, A. Pervillé 648 (lectotype, designated here: P [P00127552]!; isolectotypes: P [P00127553]!, P [P00127554]!, P [P00131420]!).
Shrubs; western Madagascar (Mahajanga).
The sheet chosen here as lectotype bears two labels, one on the left that lists Pervillé 648 as the collector, and one on the right in Baillon's hand that seemingly attributes the collection to Boivin, stating “ex Ambongo, cum cl. Pervillé et Bernier comm. (1846).” Baillon lists this in his protologue as a separate collection, but it looks identical to the other syntypes, so we believe it is actually part of the same collection by Pervillé.
The types lack pistillate flowers, but they are distinctive from C. brevispicatus in bearing a pair of sessile acropetiolar glands and in having relatively long petioles for the size of the leaf, and blades with a rounded-cuneate base and an acuminate apex.
Madagascar. Prov. Mahajanga: Bongolava, Boïna, Nov 1906, H. Perrier de la Bâthie 9567 (lectotype, designated here: P [P00127563]!). Madagascar. Prov. Mahajanga: Ankarafantsika, 7e reserve, chemin de Ste. Marie, Ankorika, 150-200 m, Service Forestier Madagascar 154 (syntypes: K [K001040380]!, P [P00127566]!). Madagascar. Prov. Mahajanga: environs de Madirovalo, Boïny, Nov 1902, H. Perrier de la Bâthie 9803 (syntypes: P [P00127564!], P [P00127565]!. Madagascar. sin. loc., Service Forestier Madagascar 64 (syntype: P [P00127567]!)).
Shrubs; western Madagascar (Mahajanga).
Croton mavoravina var. concinnus Radcl.-Sm., Gen. Croton Madag. Comoro 31. 2016, syn. nov. Type. Madagascar. Prov. Toliara: environs d’Ampandrandava, entre Bekily et Tsivory, 1000 m, Nov 1942, Herbier du Jardin Botanique de Tananarive 5730 (holotype: P [P00154405]!).
Madagascar. Prov. Toliara: environs de Bekily, 12 Oct 1966, P. Boiteau 384 (lectotype, designated here: P [P00312385]!; isolectotypes: P [P00131001]!, P [P00131002]!).
Shrubs; southern Madagascar (Toliara).
Furcaria boiviniana Baill., Étude Euphorb.: 356. 1858. Type. Madagascar. Prov. Antsiranana: Diana Region, Ile Nossibé, 1841, A. Pervillé 267 (lectotype, designated here: P [P00312458]!; isolectotypes: G [G00446376]!, P [P00131010]!). Madagascar. Prov. Antsiranana: Diana Region, Nossibé, 1847-1852, L.H. Boivin 2183 (syntype: P [P00312459]!).
Oxydectes boiviniana (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Furcaria boiviniana Baill.
Based on Furcaria boiviniana Baill.
Shrubs; northern Madagascar (Antsiranana).
Croton bakerianus Baill., Bull. Mens. Soc. Linn. Paris 2: 849. 1890, as ‘bakeriana’. Type. Madagascar. sin. loc., s.d., R. Baron 2091 (holotype: K [K001040351]!; isotype: P [P0013152]!).
Oxydectes bojeriana (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton bojerianus Baill.
Madagascar. sin. loc., s.d., W. Bojer s.n. (holotype: P [P00131353]!; isotypes: K [K000347489]!, M [M0110365]!).
Shrubs; central highland Madagascar (Antananarivo, Fianarantsoa, possibly Mahajanga).
Argentina. Prov. Corrientes: 1833, A. Bonpland s.n. (lectotype, designated here: P [P00623061]!; isolectotype: P [P00623060]!). Paraguay: Apr-May 1845, H.A. Weddell 3207 (syntypes: P [P00623063]!, P [P00623062]!).
Herbs to subshrubs; native to southern South America, but naturalized in the Mascarene islands of Mauritius, Réunion, and Rodrigues (
Croton bonplandianus is currently the only non-native, naturalized species of Croton in the Western Indian Ocean Region. Oddly, it is known from Mayotte and the Mascarenes, but it has not yet been observed or collected in Madagascar.
Andrichnia bracteata (Lam.) Baill., Étude. Euphorb. 362. 1858. Type. Based on Croton bracteatus Lam.
Oxydectes bracteata (Lam.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton bracteatus Lam.
Croton ivohibensis var. puncticulatus Radcl.-Sm., Gen. Croton Madag. Comoro 150. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Farafangana, Réserve spéciale de Manombo, 23°03'42"S, 47°44'26"E, 30 m, 25 Aug 1995, P. Rakotomalaza et al. 452 (holotype: K!; isotype: MO!).
Madagascar. sin. loc., s.d., P. Commerson s.n. (holotype: P-LA [P00382050]!; isotypes: G-DC [G00311196]!, G [G00446380]!, MPU [MPU014765]!, MPU [MPU014766]!, P [P00312457]!; probable isotype: P-JU (P00131377]!).
Shrubs or small trees; eastern coastal Madagascar (southern Fianarantsoa).
The collection locality of the type of Croton bracteatus is uncertain. Although most of Commerson's collections from Madagascar came from the Fort Dauphin area in Toliara Province, the type of C. bracteatus matches well specimens from the Manombo Reserve south of Farafangana in Fianarantsoa Province.
Croton brachybotryus Müll.Arg., Prodr. 15(2): 571. 1866, nom. superfl. Type. Based on Croton brevispicatus Baill.
Oxydectes brevispicata (Baill.) Kuntze, Revis. Gen. Pl. 2: 609. 1891. Type. Based on Croton brevispicatus Baill.
Madagascar. Prov. Antsiranana: [Diana Region] côte orientale, Baie de Rigny, Dec 1848, L.H. Boivin 2658 (holotype: P [P00131378]!; isotypes: G [G00018191]!, G-DC [G00311751]!), P [P00131406]!).
Shrubs; northern and western Madagascar (Antsiranana, Mahajanga).
Boivin's collection number 2658 was applied to several different collection events.
Madagascar. “Madagascar central,” P. Campenon s.n. (lectotype, designated here: P [P00389501]!; isolectotype: P [P00131482]!).
Trees; central and northern upland Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Mahajanga).
Oxydectes cassinoides (Lam.) Kuntze, Revis. Gen. Pl. 2: 610. 1891. Type. Based on Croton cassinoides Lam.
Croton delphinianus Baill., Bull. Mens. Soc. Linn. Paris 2: 928. 1891. Type. Madagascar. Prov. Toliara: Fort Dauphin, s.d., G.F. Scott-Elliot 1557 (lectotype, designated here: P [P00133052]!). Madagascar. Prov. Toliara: Fort Dauphin, s.d., G.F. Scott-Elliot s.n. (possible original material: P [P00131487]!).
Madagascar. sin. loc., s.d., P. Commerson s.n. (lectotype, designated here: P [P00131494]!; isolectotypes: G-DC [G00311976]!, MPU [MPU014767]!), P-JU [Catal. 16372]!).
Shrubs; southeastern coastal Madagascar (Toliara).
Unlike most other Malagasy Croton species described by Lamarck from Commerson specimens, there is no specimen of C. cassinoides found in the Lamarck Herbarium at P, so we designate the sheet in the general herbarium at P as lectotype. Most of Commerson's collections from Madagascar came from the Fort Dauphin area in Toliara Province, and all other specimens of C. cassinoides that we have determined are from near Fort Dauphin, so this is likely where the type came from. In the description of C. delphinianus (
Croton hilaris Baill., Bull. Mens. Soc. Linn. Paris 2: 927. 1891. Type. Madagascar. Prov. Toliara: Fort-Dauphin, G.F. Scott-Elliot 2650 (holotype: P [P00131522]!; isotype: K [K001040342]!).
Croton ranohirae Leandri, Adansonia, sér. 2, 9: 502. 1970, syn. nov. Type. Madagascar. Prov. Toliara: Plateaux et vallées de l’Isalo à l’Ouest de Ranohira, 900 m, 2 Nov-4 Dec 1946, H. Humbert 19591 (lectotype, designated here: P [P00312378]!; isolectotypes: K [K000422598]!, K [K000422599]!, P [P00389515]!, P [P00389516]!).
Croton boinensis var. tomentosus Radcl.-Sm., Gen. Croton Madag. Comoro 78. 2016, syn. nov. Type. Madagascar. “NW Madagascar”, sin. loc. cert., recd. Sep 1887, R. Baron 5477 (holotype: K!).
Croton catatii var. setosus Radcl.-Sm., Gen. Croton Madag. Comoro 45. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Forêt de Sahafary, S/P de Diego Suarez, 2 Dec 1970, M. Debray 1554 (holotype: P [P00154364]!).
Croton catatii var. tricholepis Radcl.-Sm., Gen. Croton Madag. Comoro 45. 2016, syn. nov. Type. Madagascar. Prov. Toliara: bassin de réception de la Mananara, affluent du Mandrare, entre l’Andohahela et l’Elakelaka, près de Mahamavo, Jan-Feb 1934, H. Humbert 13876 (holotype: K!; isotypes: P [P00127556!, P00127557!, P00127558]!).
Croton elaeagni var. chrysocarpos Radcl.-Sm., Gen. Croton Madag. Comoro 91. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Monte Isalo, in nemore Zombitsy, 800 m, 7 Nov 1967, L. Bernardi 11281 (holotype: K!; isotypes: G!, P [P00154383]!).
Croton greveanus var. micraster Radcl.-Sm., Gen. Croton Madag. Comoro 41. 2016, syn. nov. Type. Madagascar. Prov. Antananarivo: Ambatotsipihina, Tsinjoarivo, 21 Nov 1949, Service Forestier 1041-SF (holotype: P [P00133806]!).
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, Didy, 14 Aug 1889, L.D.M. Catat 1819 (holotype: P [P00131523]!).
Shrubs to normally trees; widespread in forested areas of Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Mahajanga, Toamasina, Toliara).
In this circumscription, Croton catatii is a variable but distinctive species. It is usually a tree and typically occurs in montane habitats, but in the drier areas of Isalo and Zombitse area in Fianarantsoa and Toliara Provinces, it can be shrubby, and plants there have more verrucose fruits than elsewhere. Although plants of C. catatii are typically finely lepidote, plants such as what was desribed as C. catatii var. setosus from Sahafary in Antsiranana Province can be softly pubescent, with trichomes having long-protruding central rays.
Oxydectes chapelieri (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton chapelieri Baill.
Croton louvelii Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 40. 1939, as ‘louveli’. Type. Madagascar. Prov. Toamasina: forêt côtière, Tampina, Jan. 1924, M. Louvel 217 (holotype: P [P00312340]!).
Croton aymoniniorum Leandri, Adansonia, sér. 2, 13: 175. 1973, as ‘aymoninorum’. Type. Madagascar. Prov. Toliara: Forêt de Mandena, Fort Dauphin, 19 Oct 1970, M. Keraudren-Aymonin & G. Aymonin 24940 (holotype: P [P00312374]!).
Croton daphniphylloides Radcl.-Sm., Gen. Croton Madag. Comoro 164. 2016. Type. Madagascar. Prov. Toamasina: Ambila-Lemaitso, 7 Nov 1951, Service Forestier 4228 (holotype: P [P00133469]!).
Croton daphniphyllus Radcl.-Sm., Gen. Croton Madag. Comoro 161. 2016, as ‘daphniphyllum’. Type. Madagascar. Prov. Toliara: Fort Dauphin in Mandena, 2 km E of Botanic Garden, 24°58'S, 47°00'E, 9 Oct 1990, D. Faber-Langendoen, N. Dumetz & A. Randrianasolo 2226 (holotype: P [P00133462]!; isotype: MO).
Croton daphniphyllus var. hirsutus Radcl.-Sm., Gen. Croton Madag. Comoro 163. 2016. Type. Madagascar. Prov. Toliara: Préfecture de Tôlañaro (Fort Dauphin), Canton de Mananbaro, Petriky Forest, c. 1.5 km W of large dune near N shore of Lake Andranany, c. 12 km WSW of Tôlañaro (Fort Dauphin), 25°03'S, 46°53'E, 14 Apr 1989, R. Gereau, R. Rabevohitra & N. Dumetz 3374 (holotype: K!; isotypes: MO [sheet # 3683138], P [P00133120]!).
Croton domohineifolius Radcl.-Sm., Gen. Croton Madag. Comoro 157. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Fanajazana-Mananjary, Jan 1955, Service Forestier 15451-SF (holotype: P [P00133471]!).
Croton macrobuxus var. polygynus Radcl.-Sm., Gen. Croton Madag. Comoro 177. 2016, syn. nov. Type. Madagascar. Prov. Toliara: near Fort Dauphin, s.d., J. Cloisel 178 (holotype: P [P00133612]!).
Croton rhododendroides Radcl.-Sm., Gen. Croton Madag. Comoro 163. 2016. Type. Madagascar. Prov. Toliara: Préfecture Tôlañaro (Fort Dauphin), forêt à 5 km de Ste. Luce, au nord de Maliafolaky, 24°47'S, 47°10'E, 21 Oct 1989, N. Dumetz, G. McPherson & R. Rabevohitra 775 (holotype: P [P00133460]!; isotype: MO!).
Madagascar. sin. loc., s.d., L.A. Chapelier s.n. (holotype: P [P00389523]!). Madagascar. Prov. Toliara: Sainte Luce, at entrance to preserve, 24°46'46"S, 47°10'17"E, 10 m, 17 Feb 2009, B. van Ee, P.E. Berry, B.L. Dorsey & H. Razanatsoa 925 (epitype, designated by
Shrubs; southeastern and eastern coastal Madagascar (Antsiranana, Fianarantosoa, Toamasina, Toliara).
Croton chapelieri was accepted by
Madagascar. Prov. Toliara: P.K. 30, route de Tulear à Tana, 16 Feb 1962, F. Chauvet 273 (lectotype, designated here: P [P00312370]!; isolectotypes: G [G00074264]!, K [K001044839]!, MO [sheet #04861159]!, P [P00380439]!, P [P00380440]!, P [P00380441]!, TEF [TEF000188]!).
Shrubs; southern Madagascar (Toliara).
Croton
arenicola
Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 57. 1939, nom. illeg. non Croton arenicola Small 1905, syn. nov. Type. Madagascar. Prov. Toliara: Ambovombe, 10 Aug 1924, R. Decary 2949 (lectotype, designated by
Croton leandrii Croizat, Trop. Woods 77: 15. 1944, as ‘leandri’, syn. nov. Type. Based on (replacement name for) Croton arenicola Leandri
Madagascar. Prov. Toliara: vallée moyenne du Mandrare, près d’Anadabolava, mont Vohitrotsy, 850 m, Dec 1933, H. Humbert 12659 (lectotype, designated here: P [P00131549]!; isolectotypes: G [G00018190]!, K [K001040365]!, P [P00131550]!; TAN [TAN000527]!).
Shrubs or small trees; southern Madagascar (Toliara).
Oxydectes chrysodaphne (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton chrysodaphne Baill.
Croton lepidotus Aug.DC., Bull. Herb. Boissier, sér. 2, 1: 565. 1901. Type. Madagascar. Prov. Toamasina: Maroantsetra, forêts à l’intérieur de la baie d’Antongil, 1897, A. Mocquerys 274 (holotype: G-DC [G00018155]!; isotypes: Z [Z-000015984], Z [Z-000015985]).
Croton meeusei Leandri, Notul. Syst. (Paris) 13: 184. 1948, syn. nov. Type. Madagascar. Prov. Toamasina: Soanierana-Antasibe, 350 m, 13 Oct 1938, H.J. Lam & B. Meeuse 5957 (holotype: P [P00312381]!; isotypes: L [L0234889]!, WAG!).
Croton submetallicoides Radcl.-Sm., Gen. Croton Madag. Comoro 58. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Betampona Réserve Naturelle Intégrale, 40 km NW of Toamasina, 275-650 m, 17°31'S, 49°07'E, 28 Sep 1993, B. Lewis & S. Razafimandimbison 656 (holotype: K!; isotypes: MO!, P [P00433129]!).
Madagascar. Prov. Toamasina: Foulpointe or near Tamatave, 21 Jul 1794 [date on attached letter], L.A. Chapelier s.n. (lectotype, designated by
Large shrubs or trees; eastern lowland Madagascar (Antsiranana, Fianarantsoa, Mahajanga, Toamasina, eastern Toliara).
Both Croton meeusei and C. submetallicoides fit well within the variation of leaf size and shape of C. chrysodaphne, but they are reported to have the normal five petals in staminate flowers, whereas specimens from the Foulpointe area (the presumed lectotype locality as well as the epitype locality) have the unusual number of ten petals. We judge this to be somewhat of an anomaly and not a feature that has been fixed in the species overall. The type of C. meeusei lacks pistillate flowers altogether, whereas the type of C. submetallicoides has the short, curved inflorescence and pistillate flowers that are typical of C. chrysodaphne.
Madagascar. Prov. Antsiranana/Mahajanga border: Massif du Tsaratanana, crête séparant les bassins du Sambirano et de la Mahavavy, entre la cote 2362 et la base du piton coté 2831 m, 11-13 Nov 1966, Service Forestier 27057-SF (holotype: P [P00312380]!; isotype: TEF [TEF000189]!).
Shrubs or small trees; high elevations of north-central Madagascar (border area of Antsiranana and Mahajanga).
Oxydectes cotoneaster (Müll.Arg.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton cotoneaster Müll.Arg.
Croton mahafaliensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 75. 1939. Type. Madagascar. Prov. Toliara: plateau Mahafaly, Menarandra, Feb 1910, H. Perrier de la Bâthie 9776 (lectotype, designated here: P [P00312455]!; isolectotype: P [P00133488]!) Madagascar. Prov. Toliara: Beheloka, 80 m, Jun 1910, H. Perrier de la Bâthie 9767 (syntypes: P [P00312456], P [P00133487]). Madagascar. Prov. Toliara: environs de Tuléar, Aug 1919, H. Perrier de la Bâthie 12804 (syntype: P [P00312454]). Madagascar. Prov. Toliara: Lac Manampetsa [Tsimanampetsotsa], Apr 1933, H. Perrier de la Bâthie 19047 (syntype: P [P00312453]).
Madagascar. Prov. Toliara: St. Augustin, 1837, L. Bouton s.n. (holotype: K [K001040358]!).
Shrubs; southern Madagascar (Toliara).
Croton bathianus var. toliarae Radcl.-Sm., Gen. Croton Madag. Comoro 115. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Beza Mahafaly Reserve near Betioky, E of Sakamena River, near Ambinda, 23°40'S, 44°38'E, 3 Nov 1987, P.B. Phillipson 2511 (holotype: K!; isotypes: MO, P [P00127504]!).
Croton crocodilorum var. meridionalis Radcl.-Sm., Gen. Croton Madag. Comoro 112. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Ihosy, in monte Lalanandro, 6 Nov 1967, L. Bernardi 11259bis (holotype: K!; isotypes: G!, P [P00154382]!).
Madagascar. Prov. Mahajanga: vallée de la Betsiboka à Marovoay, colline de Marovoay, 50 m, 4–5 Sep 1924, H. Humbert & H. Perrier de la Bâthie 2347 (lectotype, designated here: G [G00018189]!; isolectotypes: P [P00133026]!, P [P00312451]!, TAN [TAN000528]!). Madagascar. Prov. Mahajanga: Boiny, Nov., H. Perrier de la Bâthie 9831 (syntypes: P [P00133027]!, P [P00312452]!).
Shrubs; mainly southern and western Madagascar (Mahajanga, Toliara), but also in Fianarantsoa Province.
The Geneva sheet was chosen as the lectotype of Croton crocodilorum, since it is the most complete sheet among the syntypes and contains many seeds in the packet. It was annotated by Leandri as ‘Croton crocodilorum Leandri n. sp.,’ before it was acquired by Geneva as part of the Herbier Delessert in 1949.
The type of Croton bathianus var. toliarae supposedly differs from the type of C. crocodilorum in its rugulose (vs. smooth) seeds (
Croton mavoravina var. thysanolepis Radcl.-Sm., Gen. Croton Madag. Comoro 32. 2016. Type. Madagascar. Prov. Toliara: environs de Tuléar, 10-12 Jan 1947, H. Humbert 19816 (holotype: K!; isotype: P [P00433491]!).
Madagascar. Prov. Toliara: Atsimo-Andrefana Region, along Route Nationale 7 on S side or road, ca. 30 km as the bird flies E of Toliara, 23°19'54"S, 43°55'15"E, 190 m, 8 Feb 2009, B. van Ee, P.E. Berry, B.L. Dorsey & H. Razanatsoa 823 (holotype: MICH [MICH1210791]!; isotypes: G!, MO!, P!, TAN).
Shrubs; dry forest and scrub in southwestern Madagascar (Toliara).
Croton
nobilis
var.
delphinensis
Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 48. 1939. Type. Madagascar. Prov. Toliara: Anosy Region, environs de Fort-Dauphin, forêt de Manantantely, 60–300 m, 15 Sep 1928, H. Humbert & C.F. Swingle 5742 (lectotype, designated by
Croton chrysodaphne var. meridionalis Radcl.-Sm., Gen. Croton Madag. Comoro 62. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Préfecture de Fort-Dauphin, Forêt de Manantantely, 50-200 m, 9 Nov 1990, R. Rabevohitra 2428 (holotype: K!; isotypes: MO!, P [P00133653]!).
Madagascar. Prov. Toliara: Anosy Region, Domaine de la Cascade private preserve (Manantantely-Soanerana), several km north of Route Nationale 13 at Point Kilométrique 9 west of Fort Dauphin, 24°59'05"S, 46°55'39"E, 168–283 m, 2 Aug 2015, B. van Ee & P.E. Berry 2153 (holotype: MICH!; isotypes: MO, P, TAN!).
Trees, eastern montane forests of Madagascar (Antsiranana, Fianarantsoa, Toamasina, Toliara).
Croton bracteatus subsp. populifolius Radcl.-Sm., Gen. Croton Madag. Comoro 138. 2016, syn. nov. Type. Madagascar. Prov. Mahajanga: Route Tananarive-Majunga, environs de la Betsiboka, Jan 1949, Service des Eaux et Forêts 148SF (holotype: P [P00154434]!).
Madagascar. Prov. Mahajanga: environs de Madirovalo (Boïny), Nov 1907, H. Perrier de la Bâthie 9832 (lectotype, designated here: P [P00389520]!; isolectotype: P [P00327989]!). Madagascar. Prov. Mahajanga: Bongolava, Boïny, Nov 1906, H. Perrier de la Bâthie 9565 (syntypes: P [P00389519]!, P [P00133040])!). Madagascar. [probably Prov. Mahajanga, Réserve de Namoroka], Service Forestier 76 (syntype: P [P00389517]!). Madagascar. sin. loc., received 4 Apr 1933, Service Forestier 104 (syntype: P [P00389518]!).
Shrubs; western Madagascar (Mahajanga).
Croton bevilaniensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 79. 1939, syn. nov. Type. Madagascar. Prov. Toliara: confines de l’Anosy et de l’Androy, Bevilany, 8 Aug 1932, R. Decary 10258 (lectotype, designated here: P [P00127548]!; isolectotypes: K [K000253646]!, TAN [TAN000525]!).
Madagascar. Prov. Toliara: Massif de l’Angavo, à l’Est d’Antanimora, 19 Jul 1926, R. Decary 4446 (lectotype, designated here: P [P00133044]!; isolectotypes: K [K000253647]!, P [P00133043]!, TAN [TAN000531]!).
Shrubs; southern Madagascar (Toliara).
Croton echinatus Radcl.-Sm., Gen. Croton Madag. Comoro 13. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: environs d’Antalaha, 23 Nov. 1948, H. Humbert & R. Capuron 21918 (holotype: P [P00131511]!).
Croton alaotrensis var. integrifolius Radcl.-Sm., Gen. Croton Madag. Comoro 11. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Mananara Avaratra, Antanambe, above Mahavohobe River, 16°27'S, 49°47'E, 26 Oct 1994, G. Prance & J. Andriantiana 30783 (holotype: K!).
Madagascar. Prov. Toamasina: Fito (Ambanivoules), 1833, J.P. Goudot 7 (holotype: P [P00154404]!).
Shrubs; eastern lowland Madagascar (Antsiranana, Toamasina).
Croton droguetioides Radcl.-Sm., Gen. Croton Madag. Comoro 16. 2016 [23 Dec 2016]. Type. Madagascar. Prov. Toamasina: marais et bordures de Torotorofotsy, 18°52'S, 48°20'E, 24 Feb 1997, P. Rakotomalaza et al. 1170 (holotype: K!; isotype: MO!), nom. illeg.
Croton alaotrensis Radcl.-Sm., Gen. Croton Madag. Comoro 10. 2016, syn. nov. Type. Madagascar. “Central Madagascar”, received Dec 1883, R. Baron 3006 (holotype: K!, isotype: P [P00131488]!).
Croton parietarioides Radcl.-Sm., Gen. Croton Madag. Comoro 16. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatoharanana, près Antsevabe, 1000 m, 6 Mar 1951, G. Cours 4111 (holotype: K!; isotype: P [P00418635]!).
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, Moramanga District, Andasibe, Berano, Ambatovy mine concession, 18°47'59"S, 48°20'31"E, 1009 m, 22 Mar 2016, B. van Ee, P. Antilahimena, K. Kainulainen & P.E. Berry 2447 (holotype: MICH [MICH1513195]!, isotypes: MO!, P!, TAN!).
Shrubs; eastern montane Madagascar (Toamasina).
Croton elaeagni var. antsingyensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 34. 1939, syn. nov. Type. Madagascar. Prov. Mahajanga: Andranoboka, 21 Nov 1932, J. Leandri 551 (lectotype, designated here: P [P00133092]!; isolectotypes: P [P00133093]!, S [S07-14588]!). Madagascar. Prov. Mahajanga: Amborokontsy, 6 Oct 1932, J. Leandri 124 (syntypes: P [P00133090]!, P [P00133091]!).
Croton elaeagni var. antsirananae Radcl.-Sm., Gen. Croton Madag. Comoro 90. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Réserve Spéciale d’Ankarana, à environ 106 km au Sud d’Antsiranana par route, et 12 km à l’Ouest de Mahamasina au Lac Vert, 12°55'13"S, 49°05'10"E, 12 Dec 1995, O. Andrianantoanina & R. Bezara 908 (holotype: K!, isotypes: MO, P [P00433269]!).
Madagascar. Prov. Toliara: Andakabé, près Morondava, s.d., H. Grevé 82 (lectotype, designated here: P [P00133081]!; isolectotypes: G [G00018188]!, K [K001040385]!, P [P00133082]!, P [P00133083]!, P [P00133084]!, P [P00133085]!, P [P05481486]!).
Shrubs; northern, western and southern Madagascar (Antsiranana, Mahajanga, Toliara).
Madagascar. Prov. Toliara: Fort-Dauphin, s.d., G.F. Scott-Elliot 2970 (lectotype, designated here: P [P00133118]!, isolectotype: K [K000347488]!).
Shrubs; southern Madagascar (Toliara).
Croton bifurcatus var. emeliae (Baill.) Müll.Arg. in A.P.de Candolle, Prodr. 15(2): 584. 1866. Type. Based on Croton emeliae Baill.
Mayotte [French Overseas Department]: Jul 1849, H. Boivin 3381 (lectotype, designated here: P [P00404495]!; isolectotypes: G-DC [G00311983]!, P [P00133119]!, P [P00213565]!).
Shrubs; known only from the French island of Mayotte in the Comoro Islands.
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, along dirt road north of Route Nationale 2, past village of Savahoana, 981 m, 18°55'06"S, 48°20'38"E, 14 Aug 2015, B. van Ee, P.E. Berry & H. Razafindraibe 2212 (holotype: MICH [MICH1513196]!; isotypes: MICH [MICH1513197]!, MO, P, TAN).
Shrubs; montane forests of eastern Madagascar (Toamasina).
Croton horridulus Baill., Bull. Mens. Soc. Linn. Paris 2: 977. 1891, nom. illeg. non Croton horridulus (Baill.) Müll.Arg., 1866.
Croton lapiazicola Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 42. 1939, syn. nov. Type. Madagascar. Prov. Mahajanga: Tsingy du Bemaraha (9e Réserve), Feb. 1933, J. Leandri 945 (lectotype, designated here: P [P00312338]!; isolectotype: P [P00133400]!). Madagascar. Prov. Mahajanga: Tsingy du Bemaraha (9e Réserve), 5 Oct 1932, J. Leandri 139 (syntype: P [P00312339]!), ibid. loc., Oct 1932, J. Leandri 162 (syntype: P [P00389500]!).
Madagascar. R. Baron 5579 (holotype: P [P00133163]!); isotype: K [K000422591]!.
Shrubs; northern and western Madagascar (Antsiranana, Mahajanga).
Oxydectes farinosa (Lam.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton farinosus Lam.
Croton scottii Baill., Bull. Mens. Soc. Linn. Paris 2: 967. 1891, as ‘scotti’, syn. nov. Type. Madagascar. Prov. Toliara: Fort-Dauphin, s.d., G.F. Scott-Elliot 2987 (holotype: P [P00154436]!; isotype: K [K000253644]!).
Croton moraharivensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 68. 1939, syn. nov. Type. Madagascar. Prov. Toliara: vallée de la Manambolo, rive droite (bassin du Mandrare) aux environs d’Isomono (confluent de la Sakamalio), Mont Morahariva, 1000-1400 m, Dec 1933, H. Humbert 13176 (lectotype, designated here: P [P00132989]!; isolectotypes: G [G00018151]!, P [P00132990]!). Madagascar. Prov. Toliara: bassin de réception de la Mananara, affluent du Mandrare, pentes occidentales des montagnes entre l’Andohahela et l’Elakelaka, 600-800m, Feb 1934, H. Humbert 14041 (syntypes: K [K000815882]!, P [P00132991]!).
Croton vohibariensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 78. 1939, syn. nov. Type. Madagascar. Prov. Toliara: vallée moyenne du Mandrare, près d’Anadabolava, Mont Vohibaria (sommet), 800-810 m, Dec 1933, H. Humbert 12627 (lectotype, designated here: P [P00133322]!; isolectotypes: G [G00446404]!, K [K000253645]!, P [P00133323]!, P [P00133324]!). Madagascar. Prov. Toliara: vallée moyenne du Mandrare, près d’Anadabolava, Mont Vohitrotsy, sommet, vers 850 m, Dec 1933, H. Humbert 12673 (syntype: P [P00133325]!).
Madagascar: sin. loc., s.d., P. Commerson s.n. (holotype: P-LA [P00382067]!; isotypes: G [G00446383]!, G [G00446384]!, LINN [LINN-HS1492-5], P [P00404490]!).
Shrubs; southeastern Madagascar (Toliara).
Although there is a wide elevational variation in this species as here circumscribed, the three names listed in synonymy share the characteristic glaucous-farinose lower leaf surface, and all appear to have an affinity for igneous outcroppings.
Croton lepidotoides Radcl.-Sm., Gen. Croton Madag. Comoro 71. 2016 [23 Dec 2016], syn. nov. Type. Madagascar. Prov. Toamasina: ca. 15 air-km NE of Moramanga, ca. 11 km E of Antanambao, Ambatovy, 18°51'08"S, 48°18'40"E, 30 Jan 1997, P. Rakotomalaza et al. 1024 (holotype: K!; isotype: MO!).
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, Moramanga district, Andasibe, Berano, Ambatovy mine concession, on “cuirasse” between the workers houses and the Ambatovy supply road, 18°51'02"S, 48°18'29"E, 1142 m, 21 Mar 2016, B. van Ee, P. Antilahimena, K. Kainulainen & P.E. Berry 2436 (holotype MICH [MICH1513194]!; isotypes: G!, K!, MAPR!, MO!, P!, TAN!).
Shrubs; eastern Madagascar (Moramanga District of Toamasina Province).
Croton nitidulus var. grandifolius Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 39. 1939, as ‘grandifolia’. Type. Madagascar. Prov. Fianarantsoa: Vondrozo, 17 Sep 1926, R. Decary 5333 (holotype: P [P00133643]!; isotypes: K [K001040382]!, P [P00425637 - wood], S [S07-17118]!, TAN [TAN000539]!).
Croton nitidulus var. tandrokensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 39. 1939, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Massif d’Andringitra, col du Tandroka, versant Est, 1200 m, Sep 1910, H. Perrier de la Bâthie 9749 (holotype: P [P00133638]!).
Croton fianarantsoae var. grandifolius Leandri, Adansonia, sér. 2, 13: 297. 1973, as ‘grandifolia’, syn. nov. Type. Madagascar. Prov. Toamasina: Sahasanato, Antajonomby, Canton Befody, Distr. Nosy-Varika [N de Manajary], 21 May 1955, Service Forestier 14556 (lectotype, designated here: P [P00154399]!; isolectotypes: P [P00154400]!, P [P00154401]!, TEF [TEF000191]!).
Croton daphniphylloides var. hirsutus Radcl.-Sm., Gen. Croton Madag. Comoro 165. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Parc National de Ranomafana, Talakately, 21°15'S, 47°27'E, 27 Jul–6 Aug 1993, A. Kotozafy 131 (holotype: K!; isotypes: MO, P [P00433137]!).
Croton fianarantsoae var. coursii Radcl.-Sm., Gen. Croton Madag. Comoro 154. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Rahobevava, 1300 m, 14 Mar 1951, G. Cours 4370 (holotype: P [P00133614]!).
Croton fianarantsoae var. microphyllus Radcl.-Sm., Gen. Croton Madag. Comoro 155. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Parc National de Ranomafana, SE of Savondronona, Maharira, 21°18'S, 47°23'E, 1200-1400 m, 21-23 Apr 1993, S. Malcomber, C. Hemingway & A. Randriamanantena 2441 (holotype: K!; isotypes: MO, P [P00422457]!).
Croton fianarantsoae var. ranomafanae Radcl.-Sm., Gen. Croton Madag. Comoro 156. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Parc National de Ranomafana, trail S from Cabine de Recherche to Vato camp, 21°15'S, 47°27'E, 1060 m, 11-15 Nov 1991, S. Malcomber, A. Leeuwenberg, C. Rakotomazana, H. Ranarljadna & G. Rahajasoa 1042 (holotype: K; isotypes: MO, P [P00422461]!).
Croton fianarantsoae var. tandrokensis (Leandri) Radcl.-Sm., Gen. Croton Madag. Comoro 156. 2016, syn. nov. Type. Based on Croton nitidulus var. tandrokensis Leandri
Madagascar. Prov. Fianarantsoa: Andrambovato, Parcelle B5 [Fort Carnot], 23 Mar 1954, Service Forestier 10169-SF (holotype: P [P00338570]!; isotype: P [P00154402]!, TEF [TEF000190]!).
Shrubs to small trees; eastern Madagascar (Antsiranana, Fianarantsoa, Toamasina).
Croton fianarantsoae is part of a difficult species complex, and further work is needed to differentiate it from similar species such as C. nitidulus.
Oxydectes fothergillifolia (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton fothergillifolius Baill.
Mauritius. sin. loc., L.M.A. Du Petit-Thouars s.n. (lectotype, designated here: P [P00404184]!; isolectotype: P [P00404183]!).
Shrubs; Mauritius.
Croton geayi var. paucisquamatus Radcl.-Sm., Gen. Croton Madag. Comoro 94. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Plateau Mahafaly à l’Ouest de Betioky, 17-20 Mar 1955, H. Humbert & R. Capuron 29490 (holotype: P [P00123679]!).
Croton geayi var. pubescens Radcl.-Sm., Gen. Croton Madag. Comoro 95. 2016, syn. nov. Type. Madagascar. Prov. Toliara: vallée inférieure de l’Onilahy, 12 Aug 1928, H. Humbert & C.F. Swingle 5235 (holotype: P [P00133137]!; isotype: G!).
Madagascar. Prov. Toliara: sin. loc., 1906, M.F. Geay 28 (holotype: P [P00133132]!).
Shrubs; southern Madagascar (Toliara).
The two varieties of Croton geayi described by
Croton lamianus Leandri, Notul. Syst. (Paris) 13: 184. 1948, as ‘Lamiana’, syn. nov. Type. Madagascar. Prov. Toamasina: Soanierana-Ambahoabe, 50 m, 5 Dec 1938, H.J. Lam & B. Meeuse 5756 (lectotype, designated here: P [P00312383]!; isolectotypes: K [K001040375]!, L [93970419], P [P00312382]!). Madagascar. Prov. Toamasina: Soanierana-Ambahoabe, 100 m, 3 Dec 1938, H.J. Lam & B. Meeuse 5604 (syntypes: P [P00133398]!, L [939171491]).
Croton fianarantsoae var. masoalae Radcl.-Sm., Gen. Croton Madag. Comoro 154. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Manarivola, Sahamalaza, Vinanivao, Antalaha, Parc Masoala, 15°48'25"S, 50°17'15"E, 12–22 Feb 1996, R. Bernard et al. 213 (holotype: K!; isotypes: MO, P [P00433259]!).
Madagascar. Prov. Toamasina: Maroa [Maroantsetra], 1898, A. Mocquerys 298 (lectotype, designated here: G [G00446385]!). Madagascar. Prov. Toamasina: Maroa [Maroantsetra], s.d., A. Mocquerys 317 (syntype: G [G00446386]!).
Shrubs; northeastern Madagascar (Antsiranana, Toamasina).
Croton platanifolius Bojer ex Baker, J. Bot. 20: 268. 1882. Type. Madagascar. edges of woods in East Betsileo, received Jul 1880, R. Baron 262 (holotype: K [K000422589]!; isotype(fragment): P [P00133700, upper packet]!).
Croton emirnensis Baker, J. Linn. Soc., Bot. 20: 252. 1883. Type. Madagascar. sin. loc. s.d., R. Baron 1854 (lectotype, designated here: K [K000422588]!). Madagascar. sin. loc., s.d., R. Baron 1841 (syntypes: K [K001040356]!, P [P00133701]!).
Oxydectes goudotii (Baill.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton goudotii Baill.
Croton mollivelus Baill. Bull. Mens. Soc. Linn. Paris 2: 926. 1891, as ‘mollivelum’. Type. Madagascar. sin. loc., s.d., C.M. Le Myre de Vilers s.n. (holotype: P [P00310180]!).
Croton tsaratananae Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 74. 1939, syn. nov. Type. Madagascar. Prov. Antsiranana/Mahajanga border: Mont Tsaratanana, 2000 m, Dec 1912, H. Perrier de la Bâthie 9720 (lectotype, designated here: P [P00389496]!; isolectotypes: P [P00154411]!, P [P00154412]!).
Croton goudotii var. tsaratananae (Leandri) Radcl.-Sm., Gen. Croton Madag. Comoro 128. 2016, syn. nov. Type. Based on Croton tsaratananae Leandri
Madagascar. Prov. Antananarivo: environs de Tananarivo, rec’d. 1840, J.P. Goudot s.n. (holotype: G [G00018187]!; isotype (fragment from G): P [P00133700, lower packet]!
Trees; upland forests across Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Mahajanga, Toamasina, Toliara).
Madagascar. Prov. Mahajanga: Tsingy de Bemaraha, N of the Malambo River, 19°09'S, 44°49'E, 27 Nov 1996, C.C.H. Jongkind, J. Andriantiana & H. Razanatsoa 3235 (holotype: K!; isotypes: MO!, WAG).
Shrubs; western Madagascar (Mahajanga – Melaky Region).
Croton boutonianus Müll.Arg., Linnaea 34: 80. 1865. Type. Mauritius. in sylvis montosis, 1833, W. Bojer s.n. (lectotype, designated here: G-DC [G00311198]!; isolectotypes: G [G00446409]!, M [M0110371]!). Mauritius. 1857, L. Bouton s.n. (syntype: G-DC [G00311197]!).
Oxydectes boutoniana (Müll.Arg.) Kuntze, Revis. Gen. Pl. 2: 611. 1891. Type. Based on Croton boutonianus Müll.Arg.
Oxydectes grangerioides (Bojer ex Baill.) Kuntze, Revis. Gen. Pl. 2: 610. 1891. Type. Based on Croton grangerioides Bojer ex Baill.
Mauritius. s.d., J. Néraud s.n. (lectotype, designated here: G [G00446387]!; possible isolectotype: P [P00404205]!). Mauritius. à l’embrasure, au-dessous du Pouce, s.d. , L.M.A. Du Petit-Thouars s.n. (syntypes: P [P00404204]!, P [P0040179]!). Mauritius. crêtes de la montagne de Port-Louis, vers l’extrémité au-dessus de l’anse Courtois, Oct 1849, L.H. Boivin s.n. (syntype: P [P00404203]!). Mauritius. 1850, J.N.E. Vesco s.n. (syntype: P [P00404180]!). Mauritius. W. Bojer s.n. (original material: MAU [1402; specimen not seen], possible type: G [G00446413]!, M [M0110457]!).
Shrubs; Mauritius.
Croton greveanus var. borealis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 45. 1939. Type. Madagascar. Prov. Mahajanga: région littorale, entre Masoarivo et Ambato, Nov 1932, J. Leandri 536 (lectotype, designated here: P [P00133803]!; isolectotypes: G [G00446389]!, P [P00133804]!, P [P00133805]!). Madagascar. Prov. Mahajanga: Tsingy du Bemara, 9e Réserve Naturelle, Tsiombivositra, 10 Nov 1932, J. Leandri 517 (syntype: P [P00133802]!). Madagascar. Prov. Mahajanga: Trano Passage, J. Leandri 264 (P [P00133800], P [P00133802]!). Madagascar. Prov. Mahajanga: Savalika, 11 Dec 1932, J. Leandri 341 (P [P00133785]!, P [syntype: P00133786]!). Madagascar. Prov. Mahajanga: Boina, Firingalava, Dec 1898, H. Perrier de la Bâthie 777 (syntypes: P [P00133788]!, P [P00133789]!). Madagascar. Prov. Mahajanga: causse d’Ankara, bois rocailleux calcaire de Kamakama, Dec 1901, H. Perrier de la Bâthie 9798 (syntype: P [P00154464]!).
Croton antanosiensis var. pubescens Radcl.-Sm., Gen. Croton Madag. Comoro 43. 2016, syn. nov. Type. Madagascar. Prov. Toliara: 44 km Tulear-Ihosy, 12 Nov 1967, L. Bernardi 11411 (lectotype, designated here: G [“Hb. G 0044814”]!; isolectotypes: G [“Hb. G 0044804”]!, P [P00154375]!).
Croton boinensis var. parcelepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 78. 2016, syn. nov. Type. Madagascar. Prov. Mahajanga: Ampasimandroro, Maintirano, 12 May 1956, Service Forestier 16322-SF (lectotype, designated here: P [P00347747]!; isolectotype: P [P00154435]!).
Madagascar. Prov. Toliara: Bé-Kapaké [Bekopaka], ad. riv. Morondava, s.d., H. Grevé 239 (lectotype, designated here: P [P00404488]!; isolectotypes: P [P00133768]!, P [P00133769]!, P [P00133770]!, P [P00133771]!).
Large shrubs or small trees; western, central, and southern Madagascar (Fianarantsoa, Mahajanga, Toliara).
Madagascar. Prov. Antananarivo: forêt basse a feuilles persistantes, restes de forêt du Mt. Ambohiby (SE de Tsiroanomandidy), 1600 m, 11-16 Nov 1952, J. Leandri et al. 1790 (lectotype, designated here: P [P00312376]!; isolectotypes: K [K001044845]!, P [P00404478]!, P [P00404479]!).
Shrubs; central Madagascar (Antananarivo, Toamasina).
Croton ivohibensis var. furfuraceus Radcl.-Sm., Gen. Croton Madag. Comoro 149. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Sous-Préfecture d’Andapa, Bealampona, Befingotra, Réserve Anjanaribe-Sud on Rn Andapa-Bealanana, Antsahanifelana, near Ampiferantany, 14°47'45"S, 49°27'54"E, 22 May 1995, D. Ravelonarivo & R. Rabesonina 816 (holotype: K!; isotype: MO!).
Croton scorpistogyne Radcl.-Sm., Gen. Croton Madag. Comoro 166. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Ambohihalanana Ct., Antalaha Distr., 15 Dec 1956, Réserves Naturelles 8059-Rn (holotype: P [P00133301]!).
Madagascar. Prov. Toamasina: Maroa [Maroantsetra], forêts à l’intérieur de la baie d’Antongil, 1897, A. Mocquerys 228 (holotype: G [G00018162]!; isotype: Z [Z-000015980]).
Large shrubs or small trees; northeastern Madagascar (Antsiranana, Toamasina).
Croton heterochrous Baill., Bull. Mens. Soc. Linn. Paris 2: 862. 1890, nom. illeg. non Croton heterochrous Müll.Arg. 1865.
Croton belintae Leandri, Cat. Pl. Madag., Euphorb.: 30. 1935, syn. nov. Type. Madagascar. Prov. Antsiranana: Vavatobé, Belinta, Feb 1880, J.M. Hildebrandt 3326 (lectotype, designated here: P [P00127506]!; isolectotypes: G [G00018184]!, G [G00018185]!, JE [JE00015890], JE [JE00015891]!, K [K001040370]!, M [M-0110366], P [P00127507]!, P [P00133155]!, P [P00133156]!).
Madagascar. Prov. Antsiranana: Pasandava-bai [Bay], Kisimani, Jun 1879, J.M. Hildebrandt 3013 (lectotype, designated here: G [G00075617]!; isolectotypes: G [G00075618]!, G [G00018186]!, JE [JE00000063]!, K [K000347497]!, M [M0110367]!), P [P00133157]!, P [P00133158]!, P [P00133159]!).
Shrubs; northern and northwestern Madagascar (Antsiranana, Mahajanga).
The type of Croton belintae differs from the typically lepidote plants of C. hildebrandtii only in the presence of prominent porrect rays emerging from the center of the lepidote scales, which gives the plant a more fuzzy-pubescent appearance. In all other characters, however, such as leaf shape, petiolar glands, and the small flowers with somewhat recurved pedicels, they are identical. We therefore treat C. belintae as a synonym of the earlier name C. hildebrandtii.
Croton rubricapitirupis Leandri, Adansonia, sér. 2, 13: 173. 1973, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Ambatomenaloha, à l’W d’Itremo, 20 Jan 1955, R. Capuron 11580 (lectotype, designated here:P [P00380442]!; isolectotypes: P [P00154295]!, P [P00154296]!, P [P00154297]!, TEF [TEF000185]).
Croton anisatus var. hirsutus Radcl.-Sm., Gen. Croton Madag. Comoro 14. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatondrazaka Distr., Ambatosoratra Canton, 26 Aug 1959, Réserves Naturelles 10857 (holotype: P [P00133189]!).
Croton cassinoides var. alaotrensis Radcl.-Sm., Gen. Croton Madag. Comoro 10. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatondrazaka Distr., Ambatosoratra Canton, near the shore of Lac Alaotra, 16 Oct 1958, Réserves Naturelles 9633-Rn (holotype: P [P00131518]!).
Croton greveanus var. ambositrensis Leandri ex Radcl.-Sm., Gen. Croton Madag. Comoro 39. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Andina, environs d’ Ambositra, Dec 1921, H. Perrier de la Bâthie 18606 (holotype: P [P00133790]!).
Croton hovarum var. lepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 179. 2016, syn. nov. Type. Madagascar. sin. loc., Oct 1881, R. Baron 678 (holotype: K [not seen]; isotype: P [P00133165]!).
Croton hovarum var. subglaber Radcl.-Sm., Gen. Croton Madag. Comoro 179. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Rn 5, Ambalavao Distr., Sendrisoa Canton, 12 Nov 1953, Réserves Naturelles 5871 (lectotype, designated here: P [P00133188]!; isolectotype: P [P00133187]!).
Croton ivohibensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 37. 1939, syn. nov. Type. Madagascar. Prov. Fianarantsoa: chaine du Vohibory, à l’ouest d’Ivohibe, 1000-1300 m, 1 Nov 1924, H. Humbert 3051 (lectotype, designated here: P [P00133264]!; isolectotypes: G [G00018157]!, K [K001040387]!, P [P00133263]!, P [P00133265]!, TAN [TAN000535]!).
Croton ivohibensis var. lepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 149. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Réserve Naturelle V, Antaniloha Canton, Ivohibe District, 29 Nov 1951, Réserves Naturelles 35-29 Rn (lectotype, designated here: P [P00133632]!; isolectotype: P00133631]!).
Madagascar. Prov. Antananarivo: Imerina, Ifanangoavana, Jan 1881, J.M. Hildebrandt 3811 (lectotype, designated here: P [P00133177]!; isolectotypes: G [G00446390]!, G [G00446405]!, JE [JE00015925]!, K [K001040345]!, M [M0110359]!), P [P00133176]!). Madagascar. sin. loc., s.d., R. Baron 675 (syntype: P [P00133164]!), Baron 678 (syntypes: K [K001040346]!, P [P00133165]), R. Baron 3151 (syntypes: K [K001040347]!, P [P00133166], P [P00133167]!). Madagascar. Prov. Antananarivo: 20 km E de Tananarive, sur la route de Tamatave, 30 Oct 1928, R. Decary 6813 (syntypes: P [P00133171]!, G [G00018161]!, S [S07-14579]!), ibid. loc.. 30 Sep 1928, R. Decary 6834 (syntypes: P [P00133172]!; MO [5598313]!). Madagascar. Prov. Antananarivo: au Nord d’Ankazobe, 9 Mar 1930, Decary 7292 (syntypes: P [P00133173], GB [GB-0047692]!), ibid. loc., 11 Mar 1930, R. Decary 7380 (syntypes: P [P00133174], MO [1602793]!). Madagascar. Prov. Antananarivo: Manankazo, au NE d’Ankazobe, Nov 1913, H. Perrier de la Bâthie 9867 (syntypes: P [P0013383]!, P [P0013384]!, P [P0013385]!), ibid. loc., H. Perrier de la Bâthie 9877 (P [P00133186]!). Madagascar. Prov. Toamasina: Andovoranto, Moramanga, bord de la Sahamarirana entre Ampasimpotsy et Bevalanirano, 24 Oct 1912, R. Viguier & H. Humbert 989 (syntypes: P [P00422391]!, P [P00422392]!, K [K001040344]!). Madagascar. Prov. Fianarantsoa: Vakinankaratra, Ambatolampy, bois entre Tsinjoarivo et Ambohimasina, 2 Oct 1912, R. Viguier & H. Humbert 1924 (syntypes: P [P00133190]!, P [P00133191]!).
Shrubs; central upland Madagascar (Antananarivo, Fianarantsoa, Toamasina).
Croton ivohibensis var. alaotrensis Radcl.-Sm., Gen. Croton Madag. Comoro 148. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Befody – forêt de l’Est, Ambatondrazaka, 29 Aug 1952, Service de Eaux et de Forêts de Madagascar 4052-SF (holotype: P [P00133207]!).
Madagascar. Prov. Toamasina: forêt d’Analamazaotra, Dec 1932, J. Leandri 709 (lectotype, designated here: P [P00133198]!; isolectotypes: P [P00133197]!, P [P00133199]). Madagascar. Prov. Toamasina: forêt d’Analamazaotra, 1000m, 19 Oct 1912, R. Viguier & H. Humbert 805 (syntypes: P [P00133209]!, P [P00133210]!, B [B100153963]!), ibid. loc., 21 Oct 1912, R. Viguier & H. Humbert 830 (G [G00018160]!, P [P00133210]!, P [P00133211]!, P [P00133212]), ibid. loc., Feb 1912, H. Perrier de la Bâthie 9739 (syntypes: P [P00133200]!, P [P00133201]!, P [P00133202]!).
Shrubs; eastern montane Madagascar (Toamasina).
Croton humblotii var. anjuanensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 44. 1939, syn. nov. Type. Union of the Comoros: Anjouan, Apr 1905, Lavanchie s.n. (holotype: P [P00154408]!).
Mayotte [French Overseas Department]: forêt de Combani, 10 Oct 1884, L. Humblot 1298 (lectotype, designated here: P [P00196068]!; isolectotypes: K [K000347496]!, K [K001040343]!, LD [LD1210694], LD [LD1210214], LG [LG000009002795]), P [P00196069], P [P00196070].
Shrubs or trees; Comoro Islands, occurring on Mayotte and the three islands of the Union of the Comoros (Anjouan, Grande Comore, and Mohéli).
Croton squamiger var. acutifolius Müll.Arg. in A.P.de Candolle, Prodr. 15(2): 523. 1866, syn. nov. Type. Madagascar. in sylvis ins. Madag., s.d., W. Bojer s.n. (holotype: P [P00133274]!, isotype: M [M0110356]!).
Croton alceicornu Radcl.-Sm., Gen. Croton Madag. Comoro 70. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatovy, 18°51'34"S, 48°18'25"E, 3 Mar 1997, P. Rakotomalaza et al. 1220 (holotype: K!; isotype: MO!).
Croton antanosiensis var. fianarantsoae Radcl.-Sm., Gen. Croton Madag. Comoro 42. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Ranomafana National Park, 7 km S of the National Road 25 W of Ranomafana, 21°15'30"S, 47°25'00"E, 31 Mar 1993, D. Turk et al. 378 (holotype: K!; isotype: G [G00414720]!, MO!, P [P00418629]!).
Croton oligostemon Radcl.-Sm., Gen. Croton Madag. Comoro 45. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Analamazava, Binara Range, SW of Daraina (Vohemar), 13°15'S, 49°38'E, 26 Apr 1990, D. Meyers 90 (holotype: K!; isotypes G00414721!, MO!, P [P00433104]!).
Madagascar. sin. loc., s.d., R. Baron 5635 (lectotype, designated by
Shrubs; montane forests of Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Toamasina, Toliara).
Croton barorum var. mangokyensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 68. 1939, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Vallée d’Ihosy, bassin du Mangoky, 800-1000 m, 29-30 Oct 1924, H. Humbert 2992 (lectotype, designated here: P [P00301485]!; isolectotypes: P [P00404482]!, P [P00127484]!).
Croton bathianus var. ihosianus Radcl.-Sm., Gen. Croton Madag. Comoro 114. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: environs d’Ihosy, Mar 1934, H. Humbert 14445 (holotype: P [P00127476]!).
Madagascar. Prov. Fianarantsoa: environs d’Ihosy, Dec 1963, J. Bosser 18722 (lectotype, designated here: P [P00328080]!; isolectotypes: P [P00133216]!, TAN!).
Shrubs; south-central Madagascar (Fianarantsoa).
Oxydectes incisa (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton incisus Baill.
Croton incisus var. minor Leandri, Notul. Syst. (Paris) 13: 183. 1948, syn. nov. Type. Madagascar. Prov. Toamasina: Reserve Naturelle de Betampona, près de Tamatave; 450 m, 19 Feb 1938, H.J. Lam & B. Meeuse 6014 (holotype: P [P00133221]!; isotypes: G [G00446382]!, K!, L, S!).
Madagascar. sin. loc., s.d., L.M.A. Du Petit-Thouars s.n. (holotype: P [P00389622]!).
Shrubs; eastern lowland Madagascar (Toamasina).
Some indices, such as
Croton commiphoroides Radcl.-Sm., Gen. Croton Madag. Comoro 21. 2016. [23 Dec 2016], syn. nov. Type. Madagascar. Prov. Toamasina: Périnet-Analamazaotra, 8-9 Aug 1961, Service Forestier 20317 (holotype: P [P00133230]!).
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, Moramanga district, Analamazaotra National Park, on trail E of visitor center, 18°56'45"S, 48°25'33"E, 975 m, 11 Aug 2015, B. van Ee, P.E. Berry & H. Razafindraibe 2175 (holotype: MICH [MICH1513201]!; isotypes: G, MO, TAN).
Small shrubs; eastern montane Madagascar (Toamasina - Andasibe area).
Madagascar. Prov. Toliara: pays arides des Antandroi, Fort Dauphin, Jun-Jul, received Sep 1890, G.F. Scott-Elliot 2986 (lectotype, designated here: P [P00133237]!; isolectotype: K [K000422592]!).
Small shrubs; southern Madagascar (Toliara).
Croton
brevispicatus
var.
isalensis
Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 28. 1939. Type. Madagascar. Prov. Fianarantsoa: Isalo, 1910, H. Perrier de la Bâthie 9788 (lectotype, designated by
Based on Croton brevispicatus var. isalensis Leandri
Shrubs; southern Madagascar (Fianarantsoa, Toliara).
In his publication elevating Croton brevispicatus var. isalensis to C. isalensis,
Madagascar. Prov. Toliara: vallée de la Manambolo, rive droite (basin du Mandrare) aux environs d’Isomono (confluent de la Sakamalio), Mont Morahariva (Mahamena), 1000–1400 m, Dec 1933, H. Humbert 13247 (lectotype, designated here: P [P00133243]!; isolectotypes: G [G00018158]!, K [K001040341]!, P [P00133244]!, TAN [TAN000534]!). Madagascar. Prov. Toliara: vallée de la Sakamalio, affluent de la Manambolo (basin du Mandrare), 900-1100 m, Dec 1933, H. Humbert 13367 (syntypes: P [P00133245]!, P [P00133246]!). Madagascar. Prov. Toliara: bassin de reception de la Mananara, affluent du Mandrare, pentes occidentales des montagnes entre l’Andohahela et l’Elakelaka au Vatazo (S d’Imonty), 900-950 m, Feb 1934, H. Humbert 14076 (syntypes: P [P00133249]!, P [P00133250]!). Madagascar. Prov. Toliara: bassin de reception de la Mananara, affluent du Mandrare, pentes occidentales des montagnes entre l’Andohahela et l’Elakelaka entre Ampahiso et Mahamavo, 800 m, Jan-Feb.1934, H. Humbert 13768 (syntype: G [G00018159]!).
Large shrubs to small trees; southeastern Madagascar (Toliara).
Croton squamiger Baill., Adansonia 1: 168. 1861, as ‘squamigerum’. Type. Madagascar. Prov. Antsiranana: sin. loc., s.d., J.M.C. Richard 576 (lectotype, designated here: P [P00133578]!; isolectotype: P [P00133579]!). Madagascar. Prov. Antsiranana: Diégo-Suarès et baie de Rigny, 1837, J.M.C. Richard 176 (syntype: P [P00133577]!). Madagascar. Prov. Antsiranana: Cap d’Ambre, 1847-1852, L.H. Boivin 2658 (syntype: P [P00133273]!).
Croton squamiger var. obtusifolius Müll.Arg. in A.P.de Candolle, Prodr. 15(2): 523. 1866, nom. inval. Type. Based on Croton squamiger Baill.
Oxydectes jennyana (Gris ex Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton jennyanus Gris ex Baill.
Oxydectes squamigera (Baill.) Kuntze, Revis. Gen. Pl. 2: 613. 1891. Type. Based on Croton squamiger Baill.
Madagascar. Prov. Antsiranana [Diana Region]: vers les bas-fonds humides, lorsqu’on s’élève par les torrents au dessus des montagnes qui se trouvent au nord du port Lugwata [Diego Suarez], 1833, J.P. Goudot s.n. (holotype: G [G00434419]!; isotypes: P [P00133278]!, P [P00154519]!).
Shrubs; northern and western Madagascar (Antsiranana, Mahajanga).
We follow here the precedent of
Concerning the type of Croton squamiger, three other sheets of Boivin 2658 correspond to C. brevispicatus, so this is clearly a mixed collection. In the description of C. squamiger,
We have corrected the termination of the specific epithet to “squamiger” [masc. gender, Adj. group A nom., see
Croton kimosorum var. pubescens Radcl.-Sm., Gen. Croton Madag. Comoro 73. 2016, syn. nov. Type. Madagascar. Prov. Toliara: NW of Tôlañaro, Andohahela Réserve Intégrale, 24°57'S, 46°39'E, 23 Dec 1993, S. Malcomber 2642 (holotype: K!; isotype: MO!).
Madagascar. Prov. Toliara: vallé de la Sakamalio, affluent de la Manambolo (basin du Mandrare), 500-800 m, Dec 1933, H. Humbert 13320 (holotype: P [P00133294]!; isotypes: G [G00018156]!, K [K001040384]!, TAN [TAN000536]!).
Shrubs; southern Madagascar (Toliara).
Madagascar: “Central Madagascar”, Oct 1882, R. Baron 1951 (lectotype, designated by
Shrubs; eastern montane forests (Antananarivo, Toamasina).
See the note above under Croton cassinioides and its synonym C. delphinianus regarding the Scott-Elliot 1557 specimen (P00133052, upper right), which was also cited by
Croton lichenisilvae var. oligostemon Radcl.-Sm., Gen. Croton Madag. Comoro 198. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Anony, Sihanaka, 3 Sep 1937, Herb. Jard. Bot. Tan. 2953 (holotype: P [P00131512]!).
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, environs d’Analamazaotra, s.d., 1000 m, H. Perrier de la Bâthie 9637 (lectotype, designated here: P [P00389499]!; isolectotype: P [P00133451]!).
Shrubs; eastern montane Madagascar (Toamasina).
Croton adenophorus var. loucoubensis (Baill.) Müll.Arg. in A.P.de Candolle, Prodr. 15(2): 589. 1866. Type. Based on Croton loucoubensis Baill.
Croton adenophoroides Radcl.-Sm., Gen. Croton Madag. Comoro 117. 2016. Type. Madagascar. Prov. Antsiranana: Besinkara, Ambalafary, Andvakena: premier cours d’eau sur le chemin de Bekolosy, 14°04'S, 48°17'E, 500 m, 12 Nov 1994, L. Gautier & P. Derleth 2529 (holotype: K!; isotypes: G!, MO!, P [P00433174]!).
Madagascar. Prov. Antsiranana: Diana Region, Nossibé, forêt de Loucoubé, Mar 1851, L.H. Boivin s.n. (lectotype, designated by
Large shrubs or small trees; northern Madagascar (Antsiranana).
Croton sambiranensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 41. 1939. Type. Madagascar. Prov. Antsiranana: Haut Sambirano, 500 m, Dec 1912, H. Perrier de la Bâthie 9699 (lectotype, designated here: P [P00133571]!; isolectotypes: P [P00133572]!, P [P00404491]!).
Croton macrobuxus var. dolichobotrys Radcl.-Sm., Gen. Croton Madag. Comoro 176. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatovy, NE of Moramanga, 18°51'07"S, 48°18'26"E, 28 Feb 1998, G. McPherson 17500 (holotype: K; isotype: MO!).
Croton macrobuxus var. glandulifer Radcl.-Sm., Gen. Croton Madag. Comoro 176. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatovy, 18°51'25"S, 48°17'50"E, 28 Feb 1997, P. Rakotomalaza et al. 1194 (holotype: K; isotype: MO, P [P00433501]!).
Croton macrobuxus var. subfoliaceus Radcl.-Sm., Gen. Croton Madag. Comoro 177. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Torotorofotsy R., Berano, 15 air-km NE of Moramanga, 11 km E of Antanambao, between Ambatovy-South & Analamay-East, 18°50'32"S, 48°19'55"E, 20 Feb 1997, P. Rakotomalaza, G. Razafimanantsoa & F. Andriatsiferana 1149 (holotype: MO!).
Croton macrobuxus var. substrigosus Radcl.-Sm., Gen. Croton Madag. Comoro 177. 2016, syn. nov. Type. Madagascar. Prov. Antananarivo: Sambaina-Manjakandriana, Ankatsohazo, 29 Oct 1968, R. Razafindrambao H534R (holotype: P [P00154348]!).
Croton nitidulus var. microphyllus Radcl.-Sm., Gen. Croton Madag. Comoro 173. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Reserve Naturelle Marojejy, along the trail to the summit of Marojejy Est, 14°26'S, 49°15'E, 10 Oct 1988, J. Miller et al. 3524 (holotype: K!; isotype: MO!).
Croton nitidulus var. pubescens Radcl.-Sm., Gen. Croton Madag. Comoro 173. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Marojejy, N d’Andapa, 14°29'S, 49°38'E, coll. 21-22 Jan 1994, F. Rasoavimbahoaka et al. 30 (holotype: K!; isotype: MO!).
Madagascar. Madag. Centr., s.d., R. Baron 3063 (lectotype, designated here: K [K001040373]!; isolectotype: P [P00133472]!).
Shrubs; upland forests of Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Mahajanga, Toamasina, Toliara).
Both Croton nitidulus var. microphyllus and C. nitidulus var. pubescens from high elevations in the Marojejy massif are smaller-leaved plants than most other specimens of C. macrobuxus. They should be studied in further detail to determine if they should be recognized as a distinct, related species.
Madagascar. Prov. Antsiranana: Massif de l’Ambohimirahavavy, rebord Sud du plateau de Marofamano, 6 Feb 1951, Service des Eaux et Forêts de Madagascar 984-SF (lectotype, designated here: P [P00389625]!; isolectotypes: K [K000422594]!, MO [sheet #04861162], P [P00389623]!, P [P00389624]!, TAN!, TEF [TEF000186]!).
Trees; northern and northwestern Madagascar (Antsiranana, Mahajanga).
Croton manampetsae var. angustifolius Radcl.-Sm., Gen. Croton Madag. Comoro 27. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Beza Mahafaly Reserve, near Betioky, 23°41'S, 44°35'E, 28 Oct 1987, P.B. Phillipson 2473 (holotype: K; isotypes: MO, P [P00133494]!).
Croton manampetsae var. chaetogyne Radcl.-Sm., Gen. Croton Madag. Comoro 27. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Fort Dauphin, Ranopiso, Manatalinjo, Réserve Naturelle Intégrale d’Andohahela, 24°49'S, 46°37'E, 26-30 Oct 1994, S. Eboroke 875 (holotype: K; isotypes: MO, P [P00433495]!).
Madagascar. Prov. Toliara: Lac Manampetsa, Apr 1933, H. Perrier de la Bâthie 19087 (lectotype, designated here: P [P00133491!); isolectotypes: K [K001040386]!, P [P00133492]!).
Shrubs; southern Madagascar (Toliara).
Except for Croton manampetsae var. lepidotus (see Incertae Sedis section), the varieties recognized by
Halecus mauritianus (Lam.) Raf., Sylva Tellur.: 62. 1838. Type: Based on Croton mauritianus Lam.
Klotzschiphytum mauritianum (Lam.) Baill., Étude Euphorb.: 383. 1858. Type: Based on Croton mauritianus Lam.
Oxydectes mauritiana (Lam.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type: Based on Croton mauritianus Lam.
Réunion [French Overseas Department]: Île de Bourbon, s.d., P. Commerson s.n. (holotype: P-LA [P00382069]!; isotypes: G-DC [G00311200]!, G [G00446393]!, G [G00446394]!, MPU [MPU014846]!, MPU [MPU014847]!, MPU [MPU014848]!, P [P00121732]!, P [P00404317]!, P-JU Catal. 16377!, P [P00404321]!, P [P00404316]!, P [P00404318]!; possible isotypes, K [K001040358]!, P [P00404319]!, P [P00404320]!).
Large shrubs or small trees; restricted to the island of Réunion.
Lamarck clearly attributed this species to the Île de Bourbon, the former name of Réunion, so it is unclear why he named it “mauritianus.” At the time of Lamarck's publication, both Réunion (Île de Bourbon) and Mauritius (Île de France) were occupied by the French, and Réunion was administered out of Port Louis, Mauritius. So perhaps Lamarck used the name in a general sense for the islands administered out of Mauritius.
Croton mauritianus Lam. is the type of Croton sect. Klotzschiphytum Baill. (Baillon, 1861), which is one of the few described sections of Croton with an Old World species as its type.
Croton capuronii Leandri, Bull. Soc. Bot. France 103: 604. 1957 [‘1956’]. Type. Madagascar. Prov. Toliara: Anosy Region: environs de Bevilany, route Ambovombe-Fort Dauphin, 23 Sep 1953, R. Capuron 8493bis (lectotype, designated here: P [P00312141]!; isolectotype: P [P00347487]!). Madagascar. Prov. Toliara: environs de Fort Dauphin, près de Bevilany, 14 Sep 1928, H. Humbert & C.F. Swingle 5673 (syntypes: G, K, P [P00133508]!).
Croton mavoravina var. gracilis Radcl.-Sm., Gen. Croton Madag. Comoro 31. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Beza Mahafaly Reserve near Betioky, 23°40'S, 44°35'E, 5 Jan 1988, P.B. Phillipson 2785 (holotype: K; isotypes: MO, P [P00133515]!).
Croton mavoravina var. gymnolepis Radcl.-Sm., Gen. Croton Madag. Comoro 31. 2016, syn. nov. Type. Madagascar. Prov. Toliara: District d’Amboasary, Canton de Tranomaro, Ambatomika, 25 May 1957, G. Cours 5221 (holotype: P [P00133507]!).
Croton mavoravina var. imanombensis Radcl.-Sm., Gen. Croton Madag. Comoro 32. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Imanombo, Nov 1952, J.M. Bosser 3837 (holotype: P [P00133495]!).
Croton mavoravina var. rotundifolius Radcl.-Sm., Gen. Croton Madag. Comoro 32. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Beza Mahafaly Reserve near Betioky, hills E of the Sakamena River, valley of the Ambinda stream, 23°40'S, 44°39'E, 26 Oct 1987, P.B. Phillipson 2450 (holotype: K; isotypes: MO, P [P00133513]!).
Madagascar. Prov. Toliara: de Tsivory à Anadabolava, Mandrare moyen, 300-400 m, Dec 1933, H. Humbert 12319 (lectotype, designated here: P [P00312342]!; isolectotype: P [P00133510]!).
Shrubs; southern Madagascar (Toliara).
A fourth variety of Croton mavoravina that was described by
Croton regeneratrix var. mayottensis Radcl.-Sm., Gen. Croton Madag. Comoro 202. 2016. Type. Mayotte [French Overseas Department]: Rassi Maoussi, 30 m, 24 Apr 997, O. Pascal 915 (holotype: K!; isotypes: BR, G!, K, MAO!, MO!, P [P00144592]!, WAG).
Mayotte [French Overseas Department]: Grande-Terre, Chiconi, village, 16 Jan 2001, F. Barthelat, M’Changama & A.B. Sifary 225 (holotype: P [P00229211]!; isotypes: G!, K!, MAO!, MO!).
Shrubs; endemic to the island of Mayotte in the Comoro Islands.
Croton subaemulans var. minor Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 82. 1939, syn. nov. Type. Madagascar. Prov. Mahajanga: causse d’Ankara, Kamakama, Dec 1900, H. Perrier de la Bâthie 9814 (lectotype, designated here: P [P00133377]!; isolectotype: P [P00133376]!). Madagascar. Prov. Mahajanga: embouchure de la Soahanina, Tsiampihy, 15 Oct 1932, J. Leandri 298 (syntypes: K [K001040361]!, P [P00133370]!).
Croton delicatulus Radcl.-Sm., Gen. Croton Madag. Comoro 20. 2016, syn. nov. Type. Madagascar. Prov. Mahajanga: station forestière d’Ampijoroa, ca. 3 km N d’Andranofasika, 16°20'S, 46°51'E, 8 Apr 1984, L. Dorr & L. Koenders 2962 (holotype: P [P00131505]!; isotypes: K!, MO!).
Croton neoholstiifolius Radcl.-Sm., Gen. Croton Madag. Comoro 120. 2016, syn. nov. Type. Madagascar. Prov. Mahajanga: Tsingy de Bemaraha, Antsalova-Tsiandro Berano, 18°39'S, 44°44'E, 24 Nov 1992, J. Labat, T. Deroin, R. Edmond, H. Rabarion & O. Laivao 2153 (holotype: K!; isotype: P [P05606030]!).
Madagascar. Prov. Mahajanga: Antsingy, vers Andobo (E d’Antsalova), vers Tsiandro, 300 m, 5-8 Feb 1960, J. Leandri & P. Saboureau 3021bis (lectotype, designated here: P [P00389507]!; isolectotype: P [P00132918]!).
Shrubs; western Madagascar (Mahajanga, Toliara).
Croton menarandrae var. pubescens Radcl.-Sm., Gen. Croton Madag. Comoro 87. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Environs d’Ampandrandava, entre Bekily et Tsivory, Oct 1942, A. Seyrig 139bis (holotype: P [P00154452]!).
Madagascar. Prov. Toliara: Berge Menarandra, Ampanihy, 25 Sep 1953, Service Forestier 8273 (lectotype, designated here: P [P00347525]!; isolectotypes: P [P00132921]!, TEF [TEF000187]!).
Shrubs; southern Madagascar (Toliara).
Croton sublinearis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 60. 1939. Type. Madagascar. Prov. Toliara: Lahimanara, env. d’Ambovombe, 8 Jun 1931, R. Decary 8991 (holotype: P [P00133597]!; isotypes: G [G00018136]!, K [K001040363]!, S [S07-16870], TAN).
Croton tranomarensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 60. 1939, syn. nov. Type. Madagascar. Prov. Toliara: Anosy Region, Tranomaro, au NE d’Ambovombe, 19 Jun 1931, R. Decary 9022 (lectotype, designated here: P [P00133389]!; isolectotypes: G [G00018135]!, K [K001040392]!, S [S07-16871], TAN [TAN000544]).
Croton meridionalis var. pseudolepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 194. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: 47-49 km SE of Ihosy on the road to Ivohibe, 5 Nov 1967, L. Bernardi 11179 (holotype: K!; isotypes: G!, P [P00154376]!).
Madagascar. Prov. Toliara: bassin supérieur du Mandrare (Sud-Est), vallée de la Manombolo, 300-400 m, 23-24 Nov 1928, H. Humbert 6798 (lectotype, designated here: P [P00132930]!; isolectotypes: G [G00018153]!), P [P00132929]!). Madagascar. Prov. Toliara: plateaux et vallées de l’Isalo, environs de Fanjahira 300-600 m, 9-12 Oct 1924, H. Humbert 2737 (syntype: P [P00132928]!). Madagascar. Prov. Toliara: Vallée de l’Onilahy, aux environs de Tongobory, 100-200 m, 1-8 Oct 1924, H. Humbert 2737 (syntypes: G [G00018154]!, P [P00132927]!).
Shrubs; southern Madagascar (Fianarantsoa, Toliara).
Most individuals of this species have pairs of falcate, foliaceous, clasping stipules and stellate pubescence. Plants with similar stipules, but usually broader and more silvery-lepidote lower leaf surfaces were described under Croton meridionalis var. latifolius and C. meridionalis var. stipularis, which may represent a different species (see Incertae Sedis).
Croton peltieri Leandri, Adansonia, sér. 2, 10: 184. 1970, syn. nov. Type. Madagascar. Prov. Toliara: Lac Manampetsa (“Tsimanampetsotsa”), 23 Oct 1940, R. Decary 16060 (lectotype, designated here: P [P00338571]!; isolectotypes: P [P00154280]!, P [P00154281]!).
Croton peltieri var. hazofotsiensis Radcl.-Sm., Gen. Croton Madag. Comoro 74. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Hazofotsy Reserve 11, 28 Apr 1971, A. Richard 123 (holotype: K!).
Croton miarensis var. monadenius Radcl.-Sm., Gen. Croton Madag. Comoro 76. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Cap Sainte-Marie et environs Nord du Cap, 17 Dec 1968, Service Forestier 28552-SF (lectotype, designated here: P [P00154277]!; isolectotype: P [P00154276]!).
Madagascar. Prov. Toliara: Miari, Nov 1956, J. Bosser 10492 (holotype: P [P00312379]!, isotype TAN!).
Shrubs; southern Madagascar (Toliara).
This is a very distinctive species from southern Madagascar with silvery, ovate, long-petiolate leaves that usually have a single acropetiolar gland. It is also very unusual in having 1- or 2-locular capsules, and the branching is strongly dichotomous. All of the taxa placed here in synonymy share these features.
Madagascar. Prov. Mahajanga: Milanja, près de Soalala, 18 Nov 1954, Conservation de Réserves Naturelles 6863RN (holotype: P [P00389506]!).
Shrubs; western Madagascar (Mahajanga).
Croton fianarantsoae var. ambremontanus Radcl.-Sm., Gen. Croton Madag. Comoro 153. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: National Park of Montagne d’Ambre, path from station to Bianamalao, 12°32'S, 49°10'E, 1120 m, 4 Jun 1989, B. Du Puy, D. Du Puy, & G. Guy 218 (holotype: K!; isotype: P [P00060699]!).
Croton ivohibensis var. aesculops Radcl.-Sm., Gen. Croton Madag. Comoro 148. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Reserve Spéciale d’Ankarana, 12°54'43"S, 49°06'39"E, 19 Feb 1994, M. Andrianarisata et al. 43 (holotype: K!; isotypes: MO!, P [P00433266]!).
Croton ivohibensis var. ankaranaensis Radcl.-Sm., Gen. Croton Madag. Comoro 148. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Reserve Naturelle Intégrale d’Ankarana, 12°51'S, 49°05'E, 18 May 1987, M. Nicoll & J.P. Abraham 684 (holotype: K!; isotypes: MICH [MICH1514788]!, MO!, P [P00133271]!).
Croton thouarsianus var. macrocalyx Radcl.-Sm., Gen. Croton Madag. Comoro 183. 2016, syn. nov. Type. Madagascar. Antsiranana: Ankarana Reserve, Tsingy area, 12°54'42"S, 49°06'42"E, 22 May 1993, C.C.H. Jongkind & S. Rapanarivo 960 (holotype: K!; isotypes: MICH [MICH1514787]!, MO!, P [P00433151]!).
Madagascar. Prov. Antsiranana: au sud d’Antsiranana, près de Joffreville dans le Parc National de Montagne d’Ambre, 12°27'S, 49°13'E, 3-10 Aug 1993, O. Andrianantoanina & Rocsceohclher 283 (holotype: K!; isotype: MO!).
Shrubs; northern Madagascar (Antsiranana).
This species is restricted to Montagne d’Ambre and tsingy habitats in the Ankarana Special Reserve. It has somewhat anisophyllous terminal leaves like C. thouarsianus, but the calyx of the pistillate flowers is much larger and more accrescent in fruit, with a long pedicel. Plants vary considerably in pubescence, with some hirsute stems in the type of C. thouarsianus var. macrocalyx. Quoting from
Madagascar. Prov. Toamasina: Maroa [Maroantsetra], forêts à l’intérieur de la baie d’Antongil, 1897, A. Mocquerys 256 (holotype: G [G00018152]!; isotype: Z [Z-000015988]!).
Shrubs or small trees; eastern lowland Madagascar (Antsiranana, Toamasina).
Oxydectes mongue (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton mongue Baill.
Croton oreades Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 72. 1939, syn. nov. Type. Madagascar. Prov. Toamasina: Analamazaotra, 800 m, Dec, H. Perrier de la Bâthie 5287 (lectotype, designated here: P [P00389513]!; isolectotypes: P [P00389512]!, P [P00389514]!). Madagascar. Prov. Fianarantsoa: haute vallée de la Rienana, province de Farafangana, 3 Oct 1926, R. Decary 5563 (syntypes: K [K001040348]!, P [P00133687]!). Madagascar. Prov. Fianarantsoa: Sud-Est de Fianarantsoa, 1000-1200 m, 27 Oct 1926, R. Decary 5842 (syntypes: P [P00133688]!, S [S07-17117]!).
Croton oreades var. occidentalis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 73. 1939, syn. nov. Type. Madagascar. Prov. Mahajanga: plateau de Miangaka [Marangaka, Ankaizina], 1000 m, Dec 1922, H. Perrier de la Bâthie 15130 (lectotype, designated here: P [P00133691]!; isolectotype: P [P00133690]!).
Croton mongue var. vatambensis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 72. 1939. Type. Madagascar. Prov. Toliara: Vatambe, Fort-Dauphin, 7 Sep 1932, R. Decary 10601 (holotype: P [P00132950]!, isotypes: K [K001040389]!, TAN [TAN000537]!).
Croton mongue var. borealis Radcl.-Sm., Gen. Croton Madag. Comoro 131. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Réserve Naturelle Intégrale no. 12 de Marojejy, le long d’un affluent de la rivière Manantenina, 11 km NW de village Manantenina, 14°26'S, 49°44'E, 1220 m, 27 Oct 1996, P.J. Rakotomalaza, N. Messmer & D. Ravelonarivo 793 (holotype: K!; isotypes: MO!, P [P00433430]!).
Croton oreades var. borealis Radcl.-Sm., Gen. Croton Madag. Comoro 132. 2016, syn. nov. Type. Madagascar. Prov. Mahajanga: Befandriana-Nord, Matsoandakana, Réserve Spéciale Anjanaharibe-Sud, village d’Anjiamazava, suivant la route Nationale d’Andapa-Bealanana, piste vers le Nord approchant le sommet de Bevitsika, 14°42'S, 49°27'E, 1100 m, 14 Dec 1994, D. Ravelonarivo & R. Rabesonina 551 (holotype: K!; isotype: MO!).
Croton oreades var. craspedadenius Radcl.-Sm., Gen. Croton Madag. Comoro 132. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Andapa, Ambalamanasy II, Andasibe Kobahina, suivant la piste entre Andasibe Kobahina et Andranovola, dans la Réserve Naturelle Intégrale de Marojejy, 14°31'S, 49°38'E, 603 m, 1-3 Feb 1994, F. Rasoavimbahoaka et al. 86 (holotype: K!; isotypes: MO!, P [P00433034]!).
Croton oreades var. periphoradenius Radcl.-Sm., Gen. Croton Madag. Comoro 133. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Ampamaherana, 29 Sep 1949, Service Forestier 2033-SF (holotype: P [P00132967]!).
Madagascar. sin. loc., s.d., L.A. Chapelier s.n. (lectotype, designated here: P [P00132944]!; isolectotype: P [P00132945]!).
Trees; montane forests of Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Mahajanga, Toamasina, Toliara).
It is surprising to us that neither
Oxydectes multicostata (Müll.Arg.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type: Based on Croton multicostatus Müll.Arg.
Croton vernicosus Baker in J. Linn. Soc., Bot. 22: 519. 1887. Type. Madagascar. sin. loc., s.d., R. Baron 4935 (lectotype, designated here: K [K000347500]!, isolectotype: K [K000347498]!).
Croton sclerodorus Baill., Bull. Mens. Soc. Linn. Paris 2: 968. 1891, as ‘sclerodorum’. Type. Madagascar. sin. loc., s.d., R. Baron 4735 (holotype: P [P00133318]!).
Madagascar. Prov. Toliara: Fort Dauphin, no collector indicated, (holotype: P-JU [Catalogue # 16338]!; isotype: Madagascar j. maut. No. 39, P-LA [P00382066]!).
Large shrubs or small trees; southeastern Madagascar (Toliara, possibly also in Fianaranstsoa).
See
Oxydectes muricata (Vahl) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton muricatus Vahl in E.F.Geiseler
Croton denisii Leandri, Bull. Mus. Natl. Hist. Nat., sér. 2, 3: 367. 1931, as ‘denisi’, syn. nov. Type. Madagascar. Prov. Toliara: Behara, 9 Jul 1926, R. Decary 4320 (holotype: P [P00389632]!).
Croton bastardii var. meridionalis Radcl.-Sm., Gen. Croton Madag. Comoro 204. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Antanimora, Dec 1959, J. Bosser 13913 (holotype: P [P00133042)!; isotype: [P00133041]!).
Croton decaryi var. subglaber Radcl.-Sm., Gen. Croton Madag. Comoro 124. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Antanimora, Dec 1959, J. Bosser 13913 (holotype: P [P00133041)!; isotype: P [P00133042]!).
Madagascar. sin. loc., s.d., P. Commerson s.n. (holotype: P-JU [Catal. # 16373]!), isotype: P [P00312343]!).
Shrubs; southern Madagascar (Toliara).
Different sheets of the same collection were designated as types of Croton bastardii var. meridionalis and C. decaryi var. subglaber by
Croton calomeris Baill., Bull. Mens. Soc. Linn. Paris 2: 860. 1890. Type. Madagascar. Madag. centr., s.d., R. Baron 5929 (lectotype, designated here: P [P00132997]!; isolectotypes: K [K001040364]!, P [P00132998]!).
Croton myriaster var. austromadecassus Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 58. 1939, as ‘austromadecassa’. Type. Madagascar. Prov. Fianarantsoa: Pic d’Ivohibe (Bara), 1500-2000 m, 5-11 Nov 1924, Humbert 3198 (lectotype, designated here: P [P00224708]!; isolectotypes: P [P00338294]!, TAN [TAN000538]!). Madagascar. Prov. Fianarantsoa: Befotaka (Prov. de Farafangana), 20 Aug 1926, R. Decary 5147 (syntype: P [P00133001]). Madagascar. Prov. Fianarantsoa: Massif d’Andringitra, 1700 m, Sep 1911, H. Perrier de la Bâthie 9762 (syntype: P [P00133016]!). Madagascar. Prov. Fianarantsoa: haute vallée de la Rienana (Bassin du Matitanana), 1000-1400 m, 18-22 Nov 1924, H. Humbert 3624 (syntype: P [P00133005]!). Madagascar. Prov. Toliara: Massif du Beampingaratra, vallée de la Maloto, 800-1500 m, 31 Oct-1 Nov 1938, H. Humbert 6328 (syntypes: P [P00133008]!, US [US01050275]!). Madagascar. Prov. Toliara: Massif de l’Andohahela, 1800-1979 m, 21-22 Oct 1928, H. Humbert 6213 (syntype: P [P00133007]!). Madagascar. Prov. Fianarantsoa: Massif de l’Ikongo (Prov. de Farafangana), 17 Oct 1926, R. Decary 5692 (syntype: P [P00133002]!). Madagascar. Prov. Fianarantsoa: Massif de l’Ikongo (Prov. de Farafangana), 17 Oct 1926, R. Decary 5779 (syntype: P [P00133003]!).
Croton regeneratrix var. ranomafanae Radcl.-Sm., Gen. Croton Madag. Comoro 202. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: W of Ranomafana, Parc National Ranomafana, 21°16'S, 47°28'E, 900-1100 m, 8-10 Sep 1992, S. Malcomber & R. Rakoto 1551 (holotype: K!; isotype: MO!).
Madagascar. sin. loc., s.d., R. Baron 223 (lectotype, designated here: K [K000422585]!; isolectotypes: [K000422584]!; P [P00132995]!).
Trees; montane forests of Madagascar (Antananarivo, Antsiranana, Fianarantsoa, Mahajanga, Toamasina, Toliara).
Croton microprunus Baill., Bull. Mens. Soc. Linn. Paris 2: 861. 1890. Type. Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, Didy, 14 Aug 1889, L.D.M. Catat 1812 (holotype: P [P00132942]!).
Croton macrochlamys Baill., Bull. Mens. Soc. Linn. Paris 2: 863. 1890. Type. Madagascar. sin. loc., s.d., R. Baron 4074 (holotype: K [K001040393]!).
Croton fuscirameus Baill., Bull. Mens. Soc. Linn. Paris 2: 927. 1891. Type. Madagascar. “Central Madagascar, received Dec 1883, R. Baron 2988 (lectotype, designated here: K [K001040379]!; isolectotype: P [P00154398]!).
Croton nitidulus var. meridionalis Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 39. 1939. Type. Madagascar. Prov. Fianarantsoa: Chaîne du Vohibory, à l’Ouest d’Ivohibe, 1000-1300 m, 1 Nov 1924, H. Humbert 3051bis (lectotype, designated here: P [P00133622]!; isolectotypes: G [G00446406]!), K [K001040388, K001040390]!. Madagascar. Prov. Fianarantsoa: haute vallée de l’Iantara, bassin du Manampatra, 500-800 m, 16-17 Nov 1924, H. Humbert 3433 (syntype: P [P00133623]!).
Croton nitidulus var. parvifolius Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 39. 1939, as ‘parvifolia’. Type. Madagascar. Prov. Antsiranana: Analamahitso (haut Bemarivo), 1000 m, Aug 1907, H. Perrier de la Bâthie 9534 (holotype: P [P00133648]!).
Croton nitidulus var. spatulatus Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 40. 1939, as ‘spatulata’. Type. Madagascar. Prov. Toamasina: Bassin de l’Onive, Mangoro, forêt d’Andasibe, 1000 m, Nov 1911, H. Perrier de la Bâthie 9659 (lectotype, designated here: P [P00133626]!; isolectotypes: P [P00133627]!, P [P00133628]!).
Croton nitidulus var. fuscirameus (Baill.) Radcl.-Sm., Gen. Croton Madag. Comoro 171. 2016. Type. Based on Croton fuscirameus Baill.
Croton bracteatus subsp. manongarivensis Radcl.-Sm., Gen. Croton Madag. Comoro 137. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Manongarivo Massif, above Ambodisakoana, E of Ankaramy, 14°05'S, 48°20'E, 17 Oct 1994, G. McPherson & H. van der Werff 16377 (holotype: K!; isotype: MO!).
Croton nitidulus subsp. angustiglans Radcl.-Sm., Gen. Croton Madag. Comoro 170. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Route d’Anjiro-Moramanga (M28), Nov 1938, G. Cours 813 (holotype: P [P00148095]!).
Croton nitidulus subsp. bekolosiensis Radcl.-Sm., Gen. Croton Madag. Comoro 170. 2016, syn. nov. Type. Madagascar. Prov. Antsiranana: Réserve Spéciale de Manongarivo, R. Bekolosy, 1180 m, 14°02'S, 48°18'E, 16 May 1995, L. Gautier & C. Chatelain LG2681 (holotype: K!; isotype: G!).
Madagascar. Madag. Centr., s.d., R. Baron 1302 (lectotype, designated here: K [K000347491]!; isolectotype: P [P00133607]!). Madagascar. Madag. Centr., s.d., R. Baron 1349 (syntypes: K [K000347490]!, P [P00133610]!, P [P00133611]!).
Shrubs; montane forests of eastern and central Madagascar (Antsiranana, Antananarivo, Fianarantsoa, Toamasina, Toliara).
Oxydectes nobilis (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton nobilis Baill.
Madagascar. sin. loc.,s.d., L.M.A. Dupetit-Thouars s.n. (lectotype, designated here: P [P00133651]!; isolectotypes: P [P00133652]!, P [P00312353]!). Madagascar. Prov. Toliara: Anosy Region, southern slopes of Col de Maningotry, road to Ranomafana, 150 m, 16 Feb 2009, B. van Ee, P.E. Berry, B.L. Dorsey & H. Razanatsoa 938 (epitype, designated here: MICH [MICH1514784]!; additional duplicates: MO, P, TAN).
Trees; submontane moist forests of southeastern Madagascar (Toliara).
The existing syntypes of Croton nobilis at P are all large-leaved (most around 18 × 7 cm), sterile branches. They are consistent with oversized sucker shoots that can sometimes be found on basal growth or regrowth of cut trees. The leaves on these specimens also resemble sucker leaves of C. chrysodaphne, which comes from eastern coastal Madagascar. However,
Oxydectes noronhae (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton noronhae Baill.
Madagascar. Prov. Toamasina: Côte est, s.d., L.A. Chapelier s.n. (lectotype, designated here: P [P00133669]!; isolectotypes: MPU [MPU014849]!, P [P00133668]!). Madagascar. Prov. Toamasina: Foulpointe, F. Noronha s.n. (syntypes: G [G00018150]!, G-DC [G00311743]; ibid. loc., W. Bojer s.n. (syntypes: K [K001040398]!, P [P00133664]!). Madagascar. sin. loc., L.M.A. Du Petit-Thouars s.n. (syntypes: P [P00133680]!, P [P00133681]!).
Shrubs; eastern coastal Madagascar (Fianarantsoa, Toamasina).
A Bojer s.n. specimen at G-DC (G00311742) has a label that states “Insula Mauritius, M. Bojer 1833,” and is indicated as a type. The only Bojer specimen cited in the protologue of Croton noronhae (
Oxydectes nudata (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton nudatus Baill.
Croton boivinanus var. brevifolius Radcl.-Sm., Gen. Croton Madag. Comoro 12. 2016. Type. Madagascar. Prov. Antsiranana: Fivonandra Antsiranana II, 79 km au Sud d’Antsiranana par route RN6, et 15 km à l’Est de l’ancien chantier du Colas à Marotaolana, Campement à l’Est du village d’Ampantsona, 12°51'20"S, 49°18'10"E, 394–551 m, 3–6 Jun 1997, O. Andrianantoanina & R. Bezara 1068 (holotype: K!; isotypes: G!, MO!, P [P00433267]!).
Croton hirsutissimus Radcl.-Sm., Gen. Croton Madag. Comoro 99. 2016. Type. Madagascar. Prov. Antsiranana: versant Est du Massif de l’Ankerana (partie S du Massif de Mafokovo, au N de Vohémar), 50-400 m, 17 Dec 1966, Service Forestier 27363-SF (holotype: P [P00154439]!).
Croton menabeensis var. furfuraceus Radcl.-Sm., Gen. Croton Madag. Comoro 101. 2016. Type. Madagascar. Prov. Mahajanga: Sofia Region, Antsohihy District, Antsatrana, Bora, 17 Jul 1970, Service Forestier 30011 (holotype: P [P00154438]!).
Madagascar. Prov. Antsiranana: Diana Region, Baie de Diego-Suarès, Dec 1848, L.H. Boivin 2659 (holotype: P [P00389498]!). Madagascar. Prov. Antsiranana: Diana Region, Antsiranana II, Orangea, road going uphill from military checkpoint at entrance to Orangea, 12°14'08"S, 49°21'40"E, 50 m, 25 Oct 2009, B. van Ee, P.E. Berry, K.J. Wurdack, E.A. Haber & H. Razafindraibe 1081 (epitype, designated by
Shrubs; northern and western Madagascar (Antsiranana, Mahajanga).
Madagascar. Prov. Antsiranana: Diana Region, Ramena, Ankorikakely, Baie des Sakalava, 12°16'40"S, 49°23'01"E, 25 m, 9 Dec 2004, J. Razafitsalama et al. 692 (holotype: MICH [MICH1517188]!; isotypes: CNARP, MO!, P [P05484901]!, TAN).
Shrubs; northern Madagascar (Antsiranana).
Madagascar. Prov. Toamasina. Alaotra-Mangoro Region: Moramanga District, in primary moist montane forest along road heading south from highway Route Nationale 2 towards Lakato, ca 9.8 km south by line-of-sight from Rn 2, 19°03'05"S, 48°21'32"E, 1030-1060 m, 13 Aug 2015, B. van Ee, P.E. Berry & H. Razafindraibe 2198 (holotype: MICH [MICH1513198]!; isotypes: MAPR!, MO!, MICH [MICH1513199]!, P,! TAN!).
Large shrubs or small trees; montane forests of eastern Madagascar (Toamasina).
Madagascar. Prov. Toliara: Cap Sainte Marie, 1-150 m, 5-7 Mar 1955, H. Humbert & R. Capuron 29220 (lectotype, designated here: P [P00312372]!; isolectotypes: K [K000422596]!, P [P00389510], P [P00389511]!).
Shrubs; southern Madagascar (Toliara).
Croton dasygyne Radcl.-Sm., Gen. Croton Madag. Comoro 19. 2016 [23 Dec 2016], syn. nov. Type. Madagascar. Prov. Toamasina: Chutes du Maningory, Lac Alaotra, s.d., Herb. Jard. Bot. Tan. 3769 (S-39) (holotype: P [P00131514]!).
Madagascar. Prov. Toamasina: Alaotra-Mangoro Region, Moramanga District, Commune Ambohibary, Fokontany Ampitambe, forêt Sahaevo, 18°49'59"S, 48°17'47"E, 1118 m, 11 Dec 2006, J. Razanatsoa & T. Marcellin 279 (holotype: MICH [MICH1458525]!; isotypes: MO!, TAN!).
Shrubs; eastern montane forests of Madagascar (Toamasina).
Madagascar. Prov. Antsiranana: Ampitambarimena, Antalaha, 14 Mar 1955, Réserves Naturelles de Madagascar (Rakotoniaina) 7057 (lectotype, designated here: P [P00312373]!; isolectotypes: G [G00190674]!, K [K000422597]!, MO [sheet #04861160]), P [P00404493]!, P [P00404494]!).
Shrubs; northern Madagascar (Antsiranana).
The lectotype was annotated in Leandri's hand as “Croton rakotoniainii”, which is in accord with the collector's actual surname, so the spelling is corrected here from the variant it was published as.
Croton regeneratrix var. perrierianus Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 64. 1939, as ‘perrieriensis’. Type. Madagascar. Prov. Antananarivo: planté près d’Ambatolampy, provenant de l’Ankaratra, May 1921, H. Perrier de la Bâthie 13769 (lectotype, designated here: P [P00133537]!; isolectotype: P [P00133538]!).
Madagascar. Prov. Fianarantsoa: versant nord du Pic d’Ivohibe, 1200-1400 m, 28 Sep 1926, R. Decary 5708 (lectotype, designated here: P [P00133532]!; isolectotypes: K [K000422586]!, P [P00394468]).
Trees; montane forests of central Madagascar (Antananarivo, Fianarantsoa).
Madagascar. Prov. Antsiranana: Diana Region, Sahafary forest in the Saharaina river basin, road off of Rn 6 to the E towards the “red tsingy”, 12°36'19"S, 49°26'23"E, 250 m, 26 Oct 2009, B. van Ee, P.E. Berry, K.J. Wurdack, E.A. Haber, H. Razafindraibe & L.J. Razafitsalama 1089 (holotype: MICH [MICH1517187]!; isotypes: P!, TAN).
Shrubs; northern Madagascar (Antsiranana).
Croton sakamaliensis var. microphyllus Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 55. 1939, as ‘microphylla’. Type. Madagascar. Prov. Toliara: vallée de la Manambolo, rive gauche (Bassin du Mandrare) aux environs d’Isomono (confluent de la Sakamalio), Monts Kotriha et Isomonobe, 400-600 m, Dec.1933-Jan 1934, H. Humbert 12826 (lectotype, designated here: P [P00133552]!; isolectotypes: K [K001040361]!, P [P00133551]!, P [P00133553]!).
Madagascar. Prov. Toliara: Vallée de Sakamalio, affluent de la Manambolo (Bassin du Mandrare), 500-800 m, Dec 1933, H. Humbert 13322 (lectotype, designated here: P [P00389497]!; isolectotype: P [P00133549]!).
Shrubs; southern Madagascar (Toliara).
Croton salviformis var. rufopunctatus Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 57. 1939, as ‘rufopunctata’. Type. Madagascar. Prov. Toliara: Atsimo-Andrefana Region, du lac Manampetsa au delta de la Linta (côte Sud-Ouest), au N d’Itampolo, 1-10 m, 24-28 Aug 1928, H. Humbert & C.W. Swingle 5380 (lectotype, designated here: P [P00133569]!; isolectotype: TAN [TAN000541]!). Madagascar. Prov. Toliara: Atsimo-Andrefana Region, du lac Manampetsa au delta de la Linta, 1-10 m, 17-24 Aug 1928, H. Humbert & C.W. Swingle 5297 (syntypes: G [G00446401]!, K [K001040367]!, P [P00133567]!, P [P00133568]!).
Madagascar. Prov. Toliara: Anosy Region, Fort-Dauphin, s.d., G.F. Scott-Elliot 2990 (holotype: P [P00133566]!, isotype: K [K000422583]!).
Shrubs; southern Madagascar (Toliara).
Croton bathianus var. ambatondrazakae Radcl.-Sm., Gen. Croton Madag. Comoro 114. 2016. Type. Madagascar. Prov. Toamasina: Menaloha (G-3), Distr. d’Ambatondrazaka, Nov 1937, G. Cours 587 (holotype: P [P00127483]!).
Madagascar. Prov. Antsiranana-Mahajanga border: Centre, au lieu dit Analankeboka, à l’Ouest de Bealanana, 20 Nov 1966, Service Forestier 27107-SF (lectotype, designated by
Shrubs or small trees; northern and central Madagascar (Antsiranana, Mahajanga, Toamasina).
Croton
perrieri
Leandri, Bull. Mus. Natl. Hist. Nat., sér. 2, 3: 369. 1931. Type. Madagascar. Prov. Mahajanga: Le Berizoka [Beritsoka], Oct 1897, H. Perrier de la Bâthie 353 (lectotype, designated by
Croton
baldauffii
Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 55. 1939, as ‘baldauffi’. Type. Madagascar. Prov. Toliara: forêt de Besomaty, entre le Fiherenana et l’Isahena (Mangoky), 750-800 m, Oct 1933, H. Humbert 11249 (lectotype, designated by
Croton
ikopae
Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 83. 1939. Type. Madagascar. Prov. Antananarivo: Analamanga Region, vallée de l’Ikopa, au NW d’Ankazobe, 15 Mar 1930, R. Decary 7554 (lectotype, designated by
Croton crocodilorum var. platyaster Radcl.-Sm., Gen. Croton Madag. Comoro 112. 2016. Type. Madagascar. Prov. Toliara: forêt du Zombitsy, près de Sakaraha, Mar 1960, M. Keraudren 510 (holotype: P [P00154485]!).
Croton parvifructus Radcl.-Sm., Gen. Croton Madag. Comoro 122. 2016. Type. Madagascar. Prov. Toliara: Forêt de Zombitsy, au NE de Sakaraha (150 km NE Tuléar), 600-800 m, 2 Nov 1960, J. Leandri & J. Ratoto 3605 (holotype: P [P00132992]!).
Croton stanneus var. hirsutus Radcl.-Sm., Gen. Croton Madag. Comoro 64. 2016. Type. Madagascar. Prov. Fianarantsoa. Andranovola, near Lomanosiny, Antambohobe Canton, Ivohibe District, 13 Aug 1967, Service Forestier 26381 (holotype: P [P00133592]!).
Madagascar. “Madag. centr.”, received Nov 1885, R. Baron 3382 (holotype: K [K001040368]!, isotype: P [P00133580]!).
Shrubs to small trees; across most of Madagascar (Antananarivo, Antsiranana, Fianarantosa, Mahajanga, Toamasina, Toliara).
See
Croton chapelieri var. longepetiolata Radcl.-Sm., Gen. Croton Madag. Comoro 152. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: W of Vavatenina, Réserve Naturelle Intégrale Zahamena, Amboditamenaka Forest, 17°44'S, 49°00'E, 15–20 Sep 1993, S. Malcomber B. Rasolondraibe, L.M. Randrianjanaka & J.P. Abraham 2511 (holotype: K!; isotypes: MO [MO-2990285]!, P [P00433109]!).
Croton fianarantsoae var. obovalifolius Radcl.-Sm., Gen. Croton Madag. Comoro 155. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: W of Ranomafana, 21°16'S, 47°28'E, 8–10 Sep 1992, S. Malcomber & R. Rakoto 1550 (holotype: K; isotypes: MO, P [P00422460]!).
Croton nitidulus var. acuminatus Radcl.-Sm., Gen. Croton Madag. Comoro 169. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Itinéraire de Didy à Brickaville (forêt orientale), reçu Apr 1954, G. Cours 4871 (lectotype, designated here: P [P00133641]!; isolectotypes: P [P00133639]!; P [P00133640]!; P [P00133642]!).
Croton nitidulus var. cinereus Radcl.-Sm., Gen. Croton Madag. Comoro 170. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Ambatovy, 18°51'34"S, 48°18'25"E, 3 Mar 1997, P. Rakotomalaza et al. 1217 (holotype: K!; isotype: MO!).
Croton nitidulus var. hypopoliotes Radcl.-Sm., Gen. Croton Madag. Comoro 172. 2016, syn. nov. Type. Madagascar. “Central Madagascar”, received Dec 1883, R. Baron s.n. (holotype: K!).
Croton nitidulus var. macrophyllus Radcl.-Sm., Gen. Croton Madag. Comoro 172. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: Itinéraire de Didy à Brickaville (forêt orientale), s.d., G. Cours 4660 (lectoype, designated here: P [P00133616]!; isolectotypes: P [P00133615]!, P [P00133617]!).
Croton submetallicus var. tomentosus Radcl.-Sm., Gen. Croton Madag. Comoro 58. 2016, syn. nov. Type. Madagascar. Prov. Toamasina: forêt de l’Analamazaotra, reçu 3 Dec 1934, E. Ursch 21 (holotype: P [P00133354]!).
Madagascar. sin. loc., s.d., R. Baron 5286 (lectotype, designated here: K [K000422582]!; isolectotypes: K [K000422581]!, P [P00133327]!). Madagascar. Prov. Toamasina: Alaotra-Mangoro Region. Forest W of Eulophiella Hotel, south of Rn 2 and Andasibe, 18°59'08"S, 48°25'55"E, 900 m, 15 Aug 2015, B. van Ee, P.E. Berry & H. Razafindraibe 2222 (epitype, designated here: MICH [MICH1514786]!; additional duplicates: G, K, MO, P, TAN, US).
Shrubs or trees; eastern montane forests of Madagascar (Fianarantsoa, Toamasina).
The type of Croton submetallicus is deficient for characterizing this species properly; it lacks pistillate flowers, which are very diagnostic. Therefore we are designating an epitype that is well distributed, has photographs in Tropicos®, and is also sampled molecularly. The only other described species name that could be applied here is C. macrochlamys, but it has a very poor type specimen, and we are treating it here as a synonym of C. nitidulus because of its smaller leaf size.
For Croton nitidulus var. acuminatus,
Madagascar. Prov. Fianarantsoa: Ifandana (Prov. de Farafangana), 8 Sep 1926, R. Decary 5070 (holotype: P [P00133355]!; isotypes: K [K001044840]!, TAN [TAN000543]!).
Shrubs; eastern montane forests of Madagascar (Fianarantsoa).
Madagascar. Prov. Mahajanga: Ménabé, Tsingy du Bemaraha (9e Réserve), Feb 1933, J. Leandri 941 (lectotype, designated here: P [P00133358]!; isolectotypes: K [K001040355]!, P [P00133359]!, P [P00133360]!). Madagascar. Prov. Mahajanga: Boïna, bords du Jabohazo près du Mont Tsitondroina, Dec 1902, H. Perrier de la Bâthie 9799 (syntypes: P [P00133361]!, P [P00133362]!, P [P00133363]!).
Shrubs; western Madagascar (Mahajanga).
Oxydectes thouarsiana (Baill.) Kuntze, Revis. Gen. Pl. 2: 613. 1891. Type. Based on Croton thouarsianus Baill.
Croton mocquerysii var. meridionalis Radcl.-Sm. Radcl.-Sm., Gen. Croton Madag. Comoro 160. 2016, syn. nov. Type. Madagascar. Toliara Province: Réserve Naturelle Intégrale # 11, Andohahela, à vicinité d’Eminiminy, 24°40'S, 46°48'E, 13–25 Jan 1993, B. Randriamampionona 77 (holotype: K!; isotypes: MO!, P [P00433050]!).
Croton thouarsianus var. robustior Radcl.-Sm., Gen. Croton Madag. Comoro 183. 2016, syn. nov. Type. Madagascar. Toliara Province: Préfecture de Fort Dauphin, Forêt d’Analalava, Manantenina, côte Est, 1 Nov 1990, N. Dumetz 1378 (holotype: K!; isotype: MO!).
Madagascar: sin. loc., s.d., L.M.A. Du Petit-Thouars s.n. (lectotype, designated here: P [P00389504]!; isolectotype: P [P00389505]!).
Shrubs; southeastern Madagascar (Toliara).
Oxydectes tiliifolia (Lam.) Kuntze, Revis. Gen. Pl. 2: 613. 1891. Type. Based on Croton tiliifolius Lam.
Mauritius. collector unknown [likely P. Commerson s.n.] (holotype: P-LA [P00382046]!). Mauritius. s.d., P. Commerson s.n. (likely original material or isotype: P [P00121778]!).
Trees; Mauritius.
Croton tranomarensis var. rosmarinifolius Radcl.-Sm., Gen. Croton Madag. Comoro 196. 2016. Type. Madagascar. Prov. Toliara: 38 km SW of Ampanihy, on road to Androka, 24°50'S, 44°25'E, 5 Feb 1990, P.B. Phillipson J.-N. Labat, D. Du Puy & B. Du Puy 3434 (holotype: K! isotypes: DAV!, G!, MO!, P [P00433298]!).
Based on Croton tranomarensis var. rosmarinifolius Radcl.-Sm.
Shrubs; southern Madagascar (Toliara).
The specific epithet “rosmarinifolius” is previously occupied in Croton (Croton rosmarinifolius Salisb. (1796), which is itself an illegitimate replacement name for C. cascarilla (L.) L.), hence the new name coined here. In both this species as well as the similar C. chlaenacicomes, the leaf margins often become revolute when the plant is undergoing drought stress; that is most likely what caused the strongly inrolled margins on the type of C. tranomarensis var. rosmarinifolius.
Croton pulchellus Baill., Adansonia 1: 161. 1861, as ‘pulchellum’. Type. Madagascar. sin. loc., s.d., J. Martin s.n. (holotype: G [G00446399]!; isotype: P [P00133530]!).
Oxydectes pulchella (Baill.) Kuntze, Revis. Gen. Pl. 2: 612. 1891. Type. Based on Croton pulchellus Baill.
Oxydectes trichotoma (Geiseler) Kuntze, Revis. Gen. Pl. 2: 613. 1891. Type. Based on Croton trichotomus Geiseler
Croton trichotomus var. pulchellus (Baill.) Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 50. 1939. Type. Based on Croton pulchellus Baill.
Croton remotiflorus Radcl.-Sm., Gen. Croton Madag. Comoro 48. 2016, syn. nov. Type. Madagascar. Prov. Toliara: Réserve Intégrale # 11, Andohahela, vicinity of Eminiminy, 24°40'S, 46°48'E, 4-24 May 1993, B. Randriamampionona 385 (holotype: K!; isotypes: DAV!, G [G00414974]!, MICH!, MO!).
Croton antanosiensis var. ambohibyi Leandri ex Radcl.-Sm., Gen. Croton Madag. Comoro 42. 2016, syn. nov. Type. Madagascar. Prov. Antananarivo: Mont Ambohiby, SE de Tsiroanomandidy, 1600 m, 11-16 Nov 1952, J. Leandri, R. Capuron & A. Razafindrakoto 1775 (lectotype, designated here: P [P00154305]!; isolectotypes: P [P00154302]!, P [P00154303]!, P [P00154304]!).
Croton isomonensis var. microcarpus Radcl.-Sm., Gen. Croton Madag. Comoro 50. 2016, syn. nov. Type. Madagascar. Prov. Fianarantsoa: Antambohobe Canton, Ivohibe Distr., 6 Mar 1962, Réserves Naturelles 12150-Rn (holotype: P [P00154428]!).
Madagascar. “Madagascar f. maut., No 38” (lectotype, designated here: P-LA [P00382065]!).
Shrubs; mainly an eastern coastal species of Madagascar in Toamasina and Toliara Provinces, as far south as the Fort Dauphin area, but also recorded from Antananarivo and Fianarantsoa Provinces in upland forests (as C. antanosiensis var. ambohibyi and C. isomonensis var. microcarpus).
In the protologue of Croton trichotomus,
In our view, Croton trichotomus is primarily a littoral species of the eastern coast, but it also occurs in a number of more inland and higher elevation situations, which is an exception among Malagasy Croton species.
For Croton antanosiensis var. ambohibyi,
Croton tsiampiensis var. ankaranensis Radcl.-Sm., Gen. Croton Madag. Comoro 110. 2016. Type. Madagascar. Prov. Antsiranana: Diana Region, Massif de l’Ankarana, 4 Nov 1990, M. Bardot-Vaucoulon 224 (holotype: P00123707!).
Croton tsiampiensis var. macrophyllus Radcl.-Sm., Gen. Croton Madag. Comoro 110. 2016. Type. Madagascar. Prov. Mahajanga: Reserve Naturelle Bemaraha, Ambodiriana, ca. 9 km E of Antsalova, 18°39'S, 44°43'E, 100–125 m, 13–15 Dec 1990, L. Gillespie 4139 (holotype: K!; isotype: MO).
Croton tsiampiensis var. microphyllus Radcl.-Sm., Gen. Croton Madag. Comoro 111. 2016. Type. Madagascar. Prov. Antsiranana: Diana Region, P.K. 10 de la route Diego Suarez-Orangea, 13 Dec 1963, Service Forestier 22956 (holotype: P00123706!).
Madagascar. Prov. Mahajanga: Tsiampihy, près de l’embouchure de la Soahanina, 15 Oct 1932, J. Leandri 311 (lectotype, designated by
Shrubs; northern and western Madagascar (Antsiranana, Mahajanga).
Madagascar. Prov. Toliara: 8-16 km E of Tuléar (Toliara) on road to Tananarive (Antananarivo), 50 m, 7 Feb 1975, T. Croat 30998 (holotype: K!; isotypes: MO!, P [P00433216]!).
Shrubs; southwestern Madagascar (Toliara).
Mauritius. Perrier, near the Mare aux Vacoas, 500 m, 12 May 1938, R.E. Vaughan MAU accession number 863 (holotype: A [A00047560]; isotype: WIS [WIS00000605MAD]). Mauritius. [in flower] 14 Feb 1934, R.E. Vaughan MAU accession number 863 (paratype: K [K000422600]!). Mauritius. [in fruit] 1 Apr 1934, R.E. Vaughan MAU accession number 863 (paratype: K [K000422600]!), MAU, MAU).
Shrubs; Mauritius.
Croton vaughanii is listed by the IUCN as Critically Endangered (
Madagascar. Prov. Toamasina: environs de Vatomandry, près de lagunes, Nov 1921, H. Perrier de la Bâthie 14084 (lectotype, designated by
Shrubs or small trees; eastern littoral forests of Madagascar (Fianarantsoa, Toamasina).
This section includes 22 taxa described by
Madagascar. Prov. Mahajanga: Ampijoroa, Ankarafantsika, 26 Oct 1970, A. Richard 476 (holotype: K!).
It is uncertain what this variety is, but probably does not belong to Croton ankarensis.
Madagascar. Prov. Mahajanga: Bongolava Hills, Port Bergé, 15°40'S, 47°30'E, 23 Nov 1987, E. Bisset 38 (holotype: K!).
It is uncertain what this is, but it is probably not a member of Croton bastardii.
Madagascar. Prov. Toliara: 45 km N of Tulear on Morombe road E of junction to Manombo, 23°01'S, 43°36'E, 14 Dec 1988, P.B. Phillipson 2884 (holotype: K!; isotype: MO!).
Croton bemaranus is restricted to tsingy habitats in Antsiranana and Mahajanga Provinces and has entire leaves and elongate stipules. This variety does not fit the species at all and requires further study to determine its affinities.
Madagascar. Prov. Fianarantosoa: Ihosy-Ivohibe, 21 Dec 1965, J. Peltier & M. Peltier 5534 (holotype: P [P00154466]!).
This is a very scrappy specimen, making it difficult to determine what it may be. It does not appear to belong to Croton betiokensis, which is confined to southern Toliara Province.
Madagascar. Prov. Toliara: 45 km NE of Morondava on Beroboka road, 20°03'S, 44°37'E, 7/8 Dec 1990, L. Gillespie 4117 (holotype: K!; isotype: MO!).
This variety has much smaller leaves than typical Croton catatii and is only a small shrub, so it is not clear yet whether it fits within another species or of it may represent an undescribed species.
Madagascar. Prov. Antsiranana: Marojejy, N of Mandena, 14°29'S, 49°49'E, 22 Apr 1993, A. Randrianasolo 322 (holotype: K!; isotype MO!).
It is unlikely this belongs to C. chapelieri (in which C. daphniphylloides is treated here as a synonym); it occurs in the Marojejy Reserve at 500 m elevation and is a 8 m tall tree, whereas C. chapelieri is confined to the eastern littoral zone near sea level. It could also be a northern form of C. submetallicus, with more acute leaves and lacking the submetallic leaf undersides of that species.
Madagascar. Prov. Antsiranana: Mt. Anjenabe, vallée inférieure de l’Androranga, affluent de la Bemarivo aux environs d’ Antongondriha, 3-7 Nov 1950, H. Humbert & R. Capuron 24021 (holotype: K!; isotypes: MO!, P [P00433396]!).
Madagascar. [Prov. Antananarivo?]: environs d’Ambohikely, s.d. [1952-1955], J. Dequaire 27466 (holotype: P [P00154267]!).
This does not appear to belong to C. elaeagni, and its placement is uncertain.
Madagascar. [Prov. Antananarivo?]: environs d’Ambohikely, s.d. [1952-1955], J. Dequaire 27460 (holotype: P [P00154282]!).
This may be the same as the preceding variety.
Madagascar. Prov. Antsiranana: Andapa, Andrakata, Analamboahangy Marojejy RNI, aux environs de Manenobasy, 14°35'16"S, 49°41'20"E, 1171 m, 18-24 Jan 1995, F. Rasoavimbahoaka 488 (holotype: MO; isotypes: P [P00433385]!, TAN).
This may correspond to Croton submetallicus, but the leaves are acuminate and not submetallic on the undersides, and it occurs well north of the known distribution of that species.
Madagascar. Prov. Toamasina: Valoala Triangulation Point, 17°25'S, 48°15'E, 6 Jan 1945, A.M. Homolle 2193 (holotype: P [P00133178]!).
This is a fruiting specimen with small pistillate sepals and rusty stellate pubescence that is probably not C. hovarum, but it cannot be placed yet in another species.
Madagascar. Prov. Toliara: bassin de réception de la Mananara, affluent du Mandrare, pentes occidentales des montagnes entre l’Andohahela et l’Elakelaka, Mt Apiky au dessus de Mahamava, 800-900 m, Jan-Feb 1934, H. Humbert 13821 (lectotype, designated here: P [P00133268]!; isolectotypes: P [P00133269]!, P [P00133270]!).
This is likely an undescribed species from the lower end of evergreen forests in southeastern Toliara Province. The pistillate flowers have a very long pedicel, and the calyx is very well developed. It is likely the same as C. nitidulus var. eglandulosus and C. thouarsianus var. angustifolius (see below).
Madagascar. sin. loc., received Jan 1892, R. Baron 6134 (holotype: K!; isotype: P [P00154509]!).
This could be a form of Croton hovarum, but the leaves seem too coarsely serrate for that species, and not knowing where the specimen came from, hinders our ability to place it.
Madagascar. Prov. Toamasina: Lac Alaotra, Herb. Jard. Bot. Tan. 3777 (holotype: P [P00133298]!).
This is an interesting specimen that bears some resemblance to Croton tanalorum. Additional material is needed to assess this further. Another specimen of it is Cours 631 (P [P00133297]).
Madagascar. Prov. Fianarantsoa: Reserve Speciale de Pic d’ Ivohibe, Marovitsika Forest, 22°28'49"S, 46°56'49"E, 850 m, 17 Oct 2000, P. Hoffmann et al. 223 (holotype: K; isotype: MO!).
This does not appear to belong to Croton hovarum (where we include C. ivohibensis), and it needs further study to determine its affinities.
Madagascar. Prov. Mahajanga: bassin supérieur du Bemarivo (Boina), 800 m, Sep 1907, H. Perrier de la Bâthie 9550 (lectotype, designated here: P [P00133404]!; isolectotypes: P [P00133403]!, P [P00133405]!).
The type is somewhat similar to Croton ericius, but the leaves are much narrower and it is less hirsute than that species.
Madagascar. Prov. Toliara: environs de Tuléar (Toliara), 30 km Tulear-Tananarive, Mar 1960, M. Keraudren 541 (holotype: P [P00133295]!).
This variety differ from Croton manampetsae in that the leaves are mostly alternate (vs. opposite) and in the lepidote indumentum on the abaxial side of the leaves (vs. stellate trichomes). It will likely be described as a distinct species.
Madagascar. Prov. Toliara: haute vallée du Mandrare près d’Andetra (Andotsy), 26 Nov 1928, H. Humbert 6849 (holotype: G!; isotypes: K, P [P00418562]!).
This plant is similar to C. meridionalis, except that it has considerably wider leaves that are silvery-lepidote on the lower surface and turn dark blackish green on the upper surface when dried. Further study is needed to determine if it should be recognized as a distinct species.
Madagascar. Prov. Toliara: 45 km N of Tulear (Toliara) on road to Morombe E of junction to Manombo, 23°01'S, 43°36'E, 14 Dec 1988, P.B. Phillipson 2885 (holotype: K!; isotypes: DAV!, MO!, P [P00123724]!).
This is likely the same as the preceeding variety.
Madagascar. Prov. Toliara: bassin de réception de la Mananara, affluent du Mandrare, entre Andohahela et Elakelaka, Mt. Apiky au-dessus de Mahamavo, Jan-Feb 1934, H. Humbert 13848 (lectotype, designated here: P [P00133646]!; isolectotypes: G!, P [P00133647]!).
This is probably the same undescribed species as Croton ivohibensis var. integrifolius and C. thouarsianus var. angustifolius.
Madagascar. Prov. Antsiranana: Andapa, pentes occidentales du Massif de Marojejy (Nord- Est), Bassin de la Lokoho, a l’Est d’ Ambalamanasy II, 500-800 m, 28 Nov-6 Dec 1948, H. Humbert & R. Capuron 22126 (holotype: K; isotypes: P [P00433398]!, P [P00433399]!).
The type of this variety is a large-leaved shrub with a very large, foliaceous pistillate calyx on a long pedicel. It may represent an undescribed species.
Madagascar. Toliara Province: NW of Tolanaro, Reserve Naturelle Intégrale # 11 (Andohahela), NW of Eminiminy, beside River Itrotroky, 24°38'S, 46°46'E, 6–13 Feb 1993, S. Malcomber et al. 2159 (holotype: K!; isotypes: G!, MO).
This is probably the same undescribed species as Croton ivohibensis var. integrifolius and C. nitidulus var. eglandulosus.
Madagascar. Prov. Fianarantsoa: Réserve Speciale de Manombo, 23°01'43"S, 47°43'51"E, 26 Oct 2000, P. Hoffmann et al. 270A (holotype: K!; isotypes: BR, G, MO).
This is probably an undescribed species from the inland area of the Manombo Reserve in Fianarantsoa Province close to the coast, but not related to Croton thouarsianus.
Madagascar. Prov. Toliara: Anosy Region, vallée de la Manombolo, rive droite (basin du Mandrare) aux environs d’Isomono (confluent de la Sakamalio), mont Morahariva, 1000-1200 m, Oct 1933, H. Humbert 13105 (lectotype, designated here: P [P00133393]!; isolectotypes: G [G00446403]!, P [P00133391]!, P [P00133392]!; K [K001040390]!, TAN [TAN000546]).
The leaves of these specimens are too small to conform to Croton meridionalis, and
Madagascar. Prov. Toliara: Anosy Region, Ambovombe, Behara, 9 Jan 1931, R. Decary 8370 (holotype: P [P00133390]!; isotypes: K [K001040391]!, TAN [TAN000545]).
The leaves of the type of this variety are too small to conform to Croton meridionalis (for which C. tranomarensis is a synonym).
Croton ambovombensis var. lepidotus Radcl.-Sm., Gen. Croton Madag. Comoro 192. 2016, nom. inval. (Art. 40.7). This designation is not validly published because
Croton arenicola f. pubescens Leandri, Ann. Mus. Colon. Marseille, sér. 5, 7(1): 58. 1939, nom. inval. Protologue lacking Latin description or diagnosis (
Croton leandrii var. pubescens Radcl.-Sm., Gen. Croton Madag. Comoro 188. 2016, nom. inval. (Art. 40.6).
Democratic Republic of the Congo: Yaosuka [Yangambi], 26 Sep 1947, J. Léonard 1448 (holotype: BR [BR00000889422]; isotypes: K [K000347428], YBI [YBI159606671]).
Croton haumanianus is known from Madagascar from a single collection, Bosser 17019 (P [P00133154]!, TAN), from the Station d’Essais “Caroline,” Ilaka-Est [Toamasina Province, south of Vatomandry]. According to
Oxydectes macrostachya (Hochst. ex Delile) Kuntze, Revis. Gen. Pl. 2: 610. 1891. Type. Based on Croton macrostachyus Hochst. ex Delile
[Ethiopia] Abyssinicum: prope Djeladjeranne, 10 Aug 1840, W.P. Schimper 1665 (syntypes: BR [BR0000008252005], K [K000347438], K [K000347439], M [M0110345], M [M0110346], MO [1905790], MPU [MPU007279], P [P00540347], P [P00540348], P [P00540349]; possible isosyntype: MPU [MPU007276]). [Ethiopia]: in regione inferiori septentrionali montis Scholoda, 20 Jun 1837, W.P. Schimper 196 (syntypes: BR [BR0000008252012], BR [BR0000008367969], HBG [HBG516429], HBG [HBG516430], K [K000347440], MPU [MPU007275], NY [NY00688543], S [S07-16861]).
Croton furcellatus Baill., Bull. Mens. Soc. Linn. Paris 2: 967. 1891. Type. Mexico. Estado Sonora: [Ravines and mesas about] Guaymas, 1887, E.W. Palmer 180 (holotype: P [P00404489]!; isotypes: GH [GH00303156]!, K [K000476756]!, US [00851399]!, US [00851400]!).
Mexico. Estado Sonora: Sierra de Nayos [Nariz], Jul 1855, A.C.V. Schott III. 17 (holotype: NY [NY00246490]!; isotype: F [F0093623F]!).
Due to mislabeled herbarium specimens,
Kurkas tiglium (L.) Raf., Sylva Tellur.: 62. 1838. Type. Based on Croton tiglium L.
Tiglium officinale Klotzsch, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19 (Suppl. 1): 418. 1843. Type. Based on Croton tiglium L.
Croton officinalis (Klotzsch) Alston in H. Trimen, Handb. Fl. Ceylon 64 (Suppl.): 264. 1931, nom. superfl. Type. Based on Croton tiglium L.
Oxydectes tiglium (L.) Kuntze, Revis. Gen. Pl. 2: 614. 1891. Type. Based on Croton tiglium L.
Sri Lanka. (lectotype, first-step designated by
There is an early collection of this species from Madagascar, namely Boivin s.n. (G), from the côte orientale de Madagascar, 1846-1852, but it has not been recorded since.
Croton luteobrunneus Baker, J. Linn. Soc., Bot. 20: 254. 1883, as ‘luteo-brunneum.’ Type. Madagascar. sin. loc., s.d., R. Baron 1770 (holotype: K [K000431062]!).
Based on Croton luteobrunneum Baker
We are grateful for financial support provided by the National Geographic Society (grant 8568-08) and the U.S. National Science Foundation (awards DEB-1353162 and 1353070). We also wish to thank the curators of herbaria at G, K, MICH, MO, P, TAN, and TEF for their assistance during our visits there. We are grateful to Nicolas Fumeaux (G), Gill Challen (K), Pete Phillipson (P), and Véronique Durand-Andro (P) for their assistance in researching collections at their respective herbaria. We also thank John Boggan (US) and Walter Kittredge (GH) for help in finding isotypes of Croton furcellatus and sending us images. We especially thank Thomas Haevermans and the Museum Nationale d’Histoire Naturelle in Paris for their support of a final visit to P to research their collections. Lastly, we are very appreciative to Roy Gereau, Sandra Knapp, Gordon McPherson, and Pete Phillipson for their careful reviews of our original manuscript and helpful suggestions for improvement.