10 11646@phytotaxa 417 1 1

Phytotaxa 417 (1): 001–105 ISSN 1179-3155 (print edition)
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Monograph PHYTOTAXA
Copyright © 2019 Magnolia Press ISSN 1179-3163 (online edition)
https://doi.org/10.11646/phytotaxa.417.1.1
PHYTOTAXA
417
Taxonomic synopsis of Croton section Geiseleria

(Euphorbiaceae) in Brazil, including description of a new
species
RODOLFO CARNEIRO SODRÉ1*, MARGARETH FERREIRA DE SALES2, PAUL EDWARD
BERRY3 & MARCOS JOSÉ DA SILVA4
1
Programa de Pós-graduação em Ciências Biológicas - Botânica, Departamento de Botânica, Instituto de Biociências, Universidade
Estadual Paulista “Júlio de Mesquita Filho”, CP 510, 18618-970, Botucatu, SP, Brazil. email: sodrerodolfo@gmail.com
2
Universidade Federal Rural de Pernambuco, Av. D. Manuel de Medeiros, s.n., Dois Irmãos, 52.171-930, Recife, PE, Brazil.
3
University of Michigan Herbarium and Department of Ecology and Evolutionary Biology, 3600 Varsity Drive, 48108-2287, Ann Arbor,
MI, USA.
4
Laboratório de Morfologia e Taxonomia Vegetal, Instituto de Ciências Biológicas, Departamento de Botânica, Universidade Federal
de Goiás, CP 131, 74001-970, Goiânia, GO, Brazil.
Magnolia Press
Auckland, New Zealand
Accepted by Hajo Esser: 5 Sept. 2019; published: 19 Sept. 2019 1

RODOLFO CARNEIRO SODRÉ, MARGARETH FERREIRA DE SALES, PAUL EDWARD BERRY &
MARCOS JOSÉ DA SILVA
Taxonomic synopsis of Croton section Geiseleria (Euphorbiaceae) in Brazil, including description of a
new species
(Phytotaxa 417)
105 pp.; 30 cm.
19 September 2019
ISBN 978-1-77670-769-0 (paperback)
ISBN 978-1-77670-770-6 (Online edition)
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ISSN 1179-3155 (print edition)

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2 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Abstract
Croton is the second largest genus of Euphorbiaceae with about 1200 tropical species. Of these, over 715 species are native
to the Americas and have been assigned to 31 sections. Croton sect. Geiseleria includes about 80 species distributed from
the United States to Argentina; they are especially diverse in Brazil, where about two thirds of its species occur. Many speci-
mens from this section have been erroneously identified and require proper typification. We present a taxonomic synopsis of
C. sect. Geiseleria for Brazil, based on the analysis of over 2000 specimens from 60 herbaria and our own collections from
dozens of excursions to multiple regions of the country. Fifty species in the section are recognized here for Brazil, 37 of them
endemic to the country, among which C. seccoi is described as new to science. Three species (C. carinatus, C. catariae, and
C. mollis) are newly transferred to the section based on morphological analysis. Twenty-seven new synonyms, 70 lectotypes
and one neotype are proposed. All of the species are included in an identification key, and then comments on their geographic
distributions, habitats, phenologies, conservation status, and diagnostic characteristics are provided. Finally, distribution
maps, original illustrations, and photographs are provided for most of the species.
Key words: South America, Crotoneae, endemism, taxonomy, typification
Introduction
Euphorbiaceae are one of the largest and most diverse families of flowering plants, with about 6300 species and 330
genera worldwide (Wurdack & Farfan-Rios 2017). Among the genera, Euphorbia Linnaeus (1753: 450) and Croton
Linnaeus (1753: 1004) are the largest, with around 2000 and 1200 species, respectively. Croton is especially diverse in
the Americas and particularly in Brazil, where it is represented by 316 species (van Ee et al. 2011, BFG 2015).
Croton includes herbs, shrubs, trees, and sometimes lianas, with several types of trichomes (stellate, fasciculate,
multiradiate, rosulate, dendritic, lepidote, papillate or glandular), hyaline or colored sap, usually alternate leaves with
glands at either the apex of the petiole, the base of the blade, or along the leaf margins; its flowers are usually arranged
in thyrses with cymules, with the pistillate ones basal and the staminate ones distal. The pistillate flowers generally
have petals reduced to glands, but sometimes they are fully developed, and the stigmas are 2-fid to multifid. The
staminate flowers have filaments that are curved inwards in bud (Webster 1993, Berry et al. 2005, van Ee et al. 2011).
The species occur in diverse habitats, from near desert to moist rainforests and savannas.
In the current infrageneric classification of the New World species of Croton, van Ee et al. (2011) recognized four
subgenera and 31 sections, the latter differentiated mainly by habit, type of trichomes, presence and location of leaf
glands, type of prefloration, pistillate flower calyx morphology, number of stamens, number of stigma branches, and
features of the columella tip.
Among the American sections of Croton, C. section Geiseleria (Gray 1856: 391) Baillon (1858: 359) is one of
the three largest ones, with about 80 species of shrubs or subshrubs occurring mainly in open and dry areas (van Ee
et al. 2011), with Brazil being its main diversity center. Over 50 species are found there, especially in the Cerrado
biome (Sodré & Silva 2017a). Species of this section can be recognized by the shrubby or subshrubby habit, alternate
leaves with sessile or stipitate glands at the apex of the petiole or at the base of the blade and/or in the sinuses of the
leaf margins; inflorescences with unisexual cymules, and pistillate flowers with 5–8(–10) free sepals, glandular or
eglandular and often unequal, rudimentary petals, bifid to multifid stigmas, and fruits with an irregularly fimbriate
columella apex (Fig. 1).
Croton section Geiseleria was described as a separate genus, Geiseleria, by Klotzsch (1841b), in honor of the
botanist and German physician Eduard Ferdinand Geiseler. However, that name was illegitimate, according to van Ee
& Berry (2010), because the type species, Croton glandulosus Linnaeus (1759: 1275), had already been used as the
type for an earlier published genus name, Decarinium Rafinesque (1825: 1). The name was later validly published
by Gray (1856) as a subgenus of Croton. Baillon (1858) was the first to treat Geiseleria at the sectional level, which
was followed by van Ee et al. (2011), who broadened its circumscription to include the members of Croton sections
Decarinium (Rafinesque 1825: 1) Müller Argoviensis (1865: 78), Ocalia (Klotzsch 1841a: 195) Baillon (1858: 366),
Podostachys (Klotzsch 1841a: 193) Baillon (1858: 365), and Octolobium Chodat & Hassler (1905: 496) sensu Webster
(1993).
CROTON SECTION GEISELERIA IN BRAZIL Phytotaxa 417 (1) © 2019 Magnolia Press • 3
FIGURE 1. Diagnostic characters for Croton sect. Geiseleria. a subshrubby habit (C. pycnadenius), b shrubby habit (C. parodianus),
c leaves showing acropetiolar stipitate glands to the left and marginal stipitate glands to the right (C. sincorensis), d leaves showing
acropetiolar sessile glands (C. inaequilobus), e thyrses without a sterile zone between pistillate and staminate flowers (C. hirtus), f and g
thyrses with a sterile zone between pistillate and staminate flowers (f C. lundianus, g C. sclerocalyx), h staminate cymules (C. sincorensis),
i pistillate flowers with 8-fid stigmas (C. sincorensis), j pistillate flowers with 2-fid stigmas (C. tetradenius), k dehisced capsule showing
columella with irregularly fimbriate apex (C. agoensis); l–p: morphological variation of pistillate flowers. l C. antisyphiliticus, m C.
krukoffianus, n C. sincorensis, o C. hostmannii and p C. catariae. Most photographs here and in subsequent figures taken by R.C. Sodré,
except for Fig 18e–g by André Luiz de Rezende Cardoso and Fig 29e–g by Marcelo Fragomeni Simon.

Most Brazilian species of Croton sect. Geiseleria were described by Baillon (1864) or Müller Argoviensis (1865,
1873). The treatment in Flora Brasiliensis (Müller Argoviensis 1873) is still perhaps the most comprehensive work on
Croton in Brazil. After this classic work, about 15 new species of the section were described as new to different regions
of Brazil (e.g., Ule 1908, Croizat 1944, Steyermark 1958, Carneiro-Torres et al. 2011, Secco 2009, Secco et al. 2012,
Silva et al. 2014, 2015a, Sodré & Silva 2017a, b, 2018a). A few others were included in floristic works in different
regions of the country (Smith et al. 1988, Lima & Pirani 2003, Caruzo & Cordeiro 2007, Secco 2008, Carneiro-Torres
2009, Silva et al. 2010, Alves 2012, Sodré et al. 2017).
Given the richness of Croton sect. Geiseleria in Brazil, there are currently issues concerning typification of
many species names, widespread faulty identifications of herbarium specimens, and poor documentation about their
geographical distribution and conservation status. In response, we present here a new taxonomic synopsis of the
section for Brazil, as a first step towards its overall taxonomic revision, including the description of a new species
from the North and Northeast regions of Brazil. This study includes the typification of the species treated and their
synonymy, geographical distributions, conservation status, and diagnostic characteristics. An identification key is also
provided for the species, along with maps with their distribution in Brazil, illustrations of their diagnostic characters,
and photographs of living specimens for most of the species.
Material and methods
Brazil is the fifth largest country in area with about 8.5 million km2, but it is the most biodiverse country on earth
(Mittermeier et al. 1997). To assist with citations of distributions within the country, as well as its major vegetation
types, we provide here a map of the six different Phytogeographical Domains: Cerrado, Pampas, Amazon, Atlantic
Forest, and Caatinga (Fig. 2a), as well as a map of the 27 federative units (states and Federal District) and their five
aggregate regions (North, Northeast, South, Southeast and Midwest) (Fig. 2b).
Forty-two field trips were made to different regions of Brazil (Table 1), mostly in the Midwest, to observe of
populations of Croton sect. Geiseleria in the field and to obtain photographic records and herbarium vouchers. The
material collected is deposited at UFG and was analyzed together with about 6000 specimens from 60 other herbaria:
ALCB, ASE, B, BHCB, BR, CEN, CEPEC, CESJ, CGMS, COR, EAC, ESA, F, FLOR, G, HBR, HBRA, HCF,
HEPH, HERBAM, HJ, HMG, HRB, HUCS, HUEFS, HUEF, HUFU, HUTO, IAN, IBGE, ICN, INPA, K, LD, M,
MBM, MBML, MFS, MG, MO, MOSS, NX, P, PACA, PAMG, PEUFR, RB, RON, SP, TEPB, TUB, UB, UCS,
UEC, UFG, UFMT, UFRN, US, VIES, and W (acronyms according to Thiers, continuously updated). Information
on the phenology of the species, their preferred habitats and popular names was taken from the labels of herbarium
specimens or through field observations. The classification of vegetation types follows Ribeiro & Walter (1998) for the
Cerrado and IBGE (2012) for the other phytogeographic domains. The terminology adopted to designate vegetative
and reproductive structures follows Hickey (1973), Webster et al. (1996), Silva et al. (2009), van Ee et al. (2011),
Caruzo & Cordeiro (2013), and Sodré et al. (2017). For species with more than ten collections analyzed, only one or
two were cited for each political unit in which it occurs. However, all material examined for this paper is cited in the
exsiccatae list (Appendix 1).
The typification of the species was based on the International Code of Nomenclature (ICN) (Turland et al. 2018),
on a literature review and consultation of high resolution photographs of the type collections available at the JSTOR
Global Plants (2018) or Reflora (2018) websites or made available directly in the virtual herbarium collections of B,
BR, F, G, K, NY, P, US and W. Synonymizations proposed here are based on the analysis of the protologues, type
collections, and consideration of the variation found in populations in their respective areas of occurrence. None of
the varieties or forms previously described for the species are recognized here, since we consider the characteristics
that differentiate them to be continuous among populations of their respective species. Leaf characters such as petiole
length, shape, margin and venation of blade, presence, number and shape of glands and floral characters such as
number and density of flowers along the axis of the inflorescence, length of bracts, number and shape of sepals of
pistillate flowers, number of division and presence of trichomes on the stigmas, number of stamens and presence of
trichomes on their filaments were the main characters analyzed to distinguish the species.
The species were illustrated for their main diagnostic characters (leaf blades, details of glands, and both pistillate
and staminate flowers). Distribution maps were made using Quantum GIS Development Team software (QGIS) version
2.8.1 from geographic coordinates obtained during the collections or cited on the specimen labels. The conservation
status of the species follows the recomendations of IUCN red list categories (IUCN 2017), supplemented by the
occurrence extension calculation (EOO) that was calculated through the Geospatial Conservation Assessment Tool
(GeoCAT - http://geocat.kew.org) (Bachman et al. 2011), which led us to recognize the conservation status of the
species as either Critically Endangered (CR), Data Deficient (DD), Least Concern (LC), Near Threatened (NT), or
Vulnerable (VU), considering also the number and density of their populations, their overall distribution area, and their
occurrence in designated protected areas.
TABLE 1. Field expeditions conducted throughout this study.

State Locality Date
BA Chapada Diamantina December, 2017
BA Municipality of Caetité December, 2017
DF Surroundings of Brasília December, 2016
September, October, 2015; January, February, June, October,
GO Chapada dos Veadeiros
2016; January, February, June, September, 2017; April, 2018
GO Silvânia National Forest and adjacent areas May, October, 2016
Linda Serra dos Topázios Private Reserve of Natural
GO April, 2014
Heritage
GO Pirineus State Park and Corumbá de Goiás March, 2016; October, 2016
GO Municipality of Minaçu May, 2016
GO Serra do Caiapó and Emas National Park April, 2016; August, 2016
GO Northeast of Goiás December, 2016
GO Serra Dourada State Park February, 2017
MG Municipality of Sacramento January, 2017
MG Serra do Cipó National Park February, 2017
MG Grão-Mogol State Park July, 2016; February, 2017
MG Municipalities of Diamantina, Datas and Gouveia July, 2016; January, 2017
Triangulo Mineiro Region, Araraquara and Botanical
MG; SP July, 2016; March, 2017
Garden of São Paulo
Southeast of Mato Grosso and Pantanal of Mato Grosso
MS; MT August, 2016; April, 2017; September, 2016
do Sul
Barra do Garças region, Serra do Roncador and Chapada
MT March, 2015; March, 2016
dos Guimarães State Park
MT Northeast and central east of Mato Grosso October, 2016
Santa Bárbara State Park, Serra das Araras Ecological
MT November, December, 2016
Station and adjacent areas
MT North of Mato Grosso April, 2017
Serra Geral do Tocantins Ecological Station and
TO April, 2016; May, 2017
surroundings of Jalapão region

FIGURE 2. a Political-administrative and regional map of Brazil; b Map of the Phytogeographical Domains present in Brazil. AC=Acre,
AL=Alagoas, AM=Amazonas, AP=Amapá, BA=Bahia, CE=Ceará, DF= Distrito Federal, ES=Espírito Santo, GO=Goiás, MA=Maranhão,
MG= Minas Gerais, MS=Mato Grosso do Sul, MT=Mato Grosso, PA=Pará, PB=Paraíba, PE=Pernambuco, PI=Piauí, PR=Paraná, RJ=Rio
de Janeiro, RN=Rio Grande do Norte, RO=Rondônia, RR=Roraima, RS=Rio Grande do Sul, SC=Santa Catarina, SE=Sergipe, SP=São
Paulo, TO=Tocantins.
Results and Discussion
Fifty species of Croton sect. Geiseleria are recognized here for Brazil, corresponding to two thirds of the total species
cited for the section by van Ee et al. (2011). Three of these species present a broad Neotropical distribution, namely,
Croton glandulosus, C. hirtus L’Héritier de Brutelle (1785: 17), and C. trinitatis Millspaugh (1900: 57); both C. hirtus
and C. glandulosus are also found in the Old World as introduced species, mainly in Africa and Australia. Ten species
are found in at least two South American countries (C. aberrans Müller Argoviensis (1873: 232), C. antisyphiliticus
Mart. in Spix & Martius (1823: 282), C. catariae Baillon (1864: 349), C. hostmannii Schltdl. ex Miquel (1848: 477), C.
krukoffianus Croizat (1944: 453), C. lundianus (Didrichsen 1857: 137) Müller Argoviensis (1866: 662), C. sclerocalyx
(Didrichsen 1857: 134) Müller Argoviensis (1865: 134), C. sipaliwinensis Lanjouw (1939: 698), C. mollis Bentham
(1854: 375), and C. subincanus Müller Argoviensis (1865: 139), whereas the remaining 37 species are all restricted to
Brazil.
Within Brazil, the Midwest region was the richest with 32 species, followed by the Southeast and North regions,
with 25 each, and the Northeast and South, with 24 and seven, respectively. Looking at Phytogeographic Domains,
the Cerrado presented the greatest diversity (32 species), followed by the Caatinga (15), the Amazon Forest (15), the
Atlantic Forest (12), the Pantanal (2), and the Pampa (1).
Five new species were discovered during this extended study: Croton ramosissimus Sodré & Silva (2017a: 128),
C. rhodotrichus Sodré & Silva (2017b: 338), C. uliginosus Sodré & Silva (2018a: 101), C. macrosepalus Sodré &
M.J. Silva in Sodré et al. (2019) and C. seccoi, the last one published in this paper. Twenty-three species had their
distribution extended to other states, regions, or countries (e.g., C. aberrans, C. abaitensis Baillon (1864: 337), C.
campinarensis Secco et al. (2012: 2), C. tetradenius Baillon (1864: 343)), while others had their distribution further
restricted by the lack of verifiable vouchers for their previously published areas of occurrence (e.g., C. adenodontus
(Müller Argoviensis 1866: 684) Müller Argoviensis (1873: 267), C. antisyphiliticus, C. goyazensis Müller Argoviensis
(1865: 120)).
Taxonomically, 27 new synonyms, 70 lectotypes and one neotype are proposed. Croton carinatus Müller
Argoviensis (1873: 126), C. catariae, and C. mollis, taxa previously assigned by van Ee et al. (2011) to other sections
such as C. sect. Barhamia (Klotzsch 1853: 104) Baillon (1858: 367) and C. sect. Adenophylli Grisebach (1859: 40),
are transferred here to sect. Geiseleria based on re-examination of their morphological characteristics.
Regarding their conservation status, most of the species (39) are considered as Least Concern; Croton planaltoanus
Silva et al. (2014: 89) is classified as Near Threatened; C. bidentatus Müller Argoviensis (1865: 140), C. inaequilobus
Steyermark (1958: 7), C. macrosepalus, C. sublepidotus Müller Argoviensis (1873: 260), and C. uliginosus as
Vulnerable; C. arenosus Carneiro-Torres et al. (2011: 123), C. carinatus, and C. rhodotrichus are recognized as Critically
Endangered; and C. adenodontus and C. crustulifer Croizat (1944: 451) are considered to be Data Deficient.
Morphological aspects of species of Croton sect. Geiseleria in Brazil
Habit: Most of the species studied (44) are subshrubs, which can be either erect (Fig. 3) or subdecumbent (Fig. 4a,
b). Six others, namely C. asperrimus Bentham (1854: 376) (Fig. 4c), C. parodianus Croizat (1944: 452) (Fig. 4d), C.
nepetifolius Baillon (1864: 344) (Fig. 1b), C. inaequilobus (Fig. 4e), C. hadrianii Baillon (1864: 313) (Fig. 4f), and C.
ramosissimus (Fig. 4g) are shrubs from 1 to 5 m tall. Both shrubs and subshrubs have stems that are generally branched
from the base or from the lower half of the stem. Some specimens of C. antisyphiliticus (Fig. 3f) and C. junceus Baillon
(1864: 339) (Fig. 3e) have mostly unbranched stems. The pattern of branching of the species is sympodial (Fig. 3c),
with dichotomies or trichotomies in the upper half of the main stem. Some species present main stems with similarly
sized, shortened secondary branches that give the plant a monopodial growth aspect, as in C. ramosissimus (Fig. 4g).
In most cases, the branches are cylindrical, cinereous or brownish, smooth or discretely fissured in the adult parts, or
angulose greenish, yellowish or rusty in the young portions, and they vary from glabrous or pubescent.
Latex: All species studied have latex that is usually scarce and hyaline, although it may become slightly orange after
oxidation, as in C. desertorum Müller Argoviensis (1873: 265) and C. nepetifolius.

FIGURE 3. Growth patterns of some subshrubby species: stems erect and densely ramified in a Croton hirtus, b C. desertorum and c C.
glandulosus; stems erect and sparsely ramified in d C. goyazensis, e C. junceus and f C. antisyphiliticus.
FIGURE 4. Growth patterns of some subshrubby and shrubby species: subshrubby habit with decumbent stems in a Croton pycnadenius
and b C. odontadenius; shrubby habit densely ramified in c C. asperrimus, d C. parodianus, e C. inaequilobus, f C. hadrianii, and g C.
ramosissimus.

Trichome morphology: This is one of the most important characters in the taxonomy of Croton, and trichomes are found
in vegetative and reproductive parts of all species studied. Baillon (1864) and Müller Argoviensis (1873) recognized
two main groups of eglandular trichomes in Croton: the stellate ones, with free rays, and lepidote ones, with the rays
united. Froembling (1896) and Webster et al. (1996) studied these in greater detail and proposed subclasses and new
types, such as dendritic, fasciculate, multiratiate, and rosulate trichomes. In this paper, we followed Webster et al.
(1996) and recognized five types of trichomes for the species studied as follows:
i. Stellate-porrect (Fig. 5a-d): This type occurs in all species studied and is characterized by 8–12 lateral rays that
are free or shortly united (up to 15% of the length), arranged in a single plane, with a prominent central ray. The
central ray usually is as long as the lateral rays (Fig. 5d) or up to 20 times longer than the laterals (Fig. 5b, c). In a
few species, mainly on the abaxial leaf surface, the central porrect ray is greatly shortened (Fig. 5a).
ii. Simple (Fig. 5e, f): These trichomes are present mainly on the branches and leaf blades of C. aberrans, C.
krukoffianus, C. odontadenius Müller Argoviensis (1873: 227), and C. sclerocalyx. Simple trichomes are also
found on the filaments of the stamens and on the margin of the petals and receptacles of the staminate flowers.
iii. Multiradiate (6a-d): This type of trichome is present in 19 of the 50 species studied and is characterized by the
presence of 15–30 free lateral rays, arranged in two or more planes (Fig. 6b). For us, these trichomes correspond
to Webster et al.’s (1996) rosulate and multiradiate trichomes and to the geminate trichomes of Theobald (1979).
These trichomes may be sessile or stipitate, and they often have a porrect ray that is usually slightly differentiated
from the other rays. They are differentiated from dendritic trichomes because they have cycles of basal rays that
do not extend from an elongated central axis, as in Croton suberosus Kunth (1817: 86) (Webster et al. 1996).
iv. Fasciculate (Fig. 6e, f): This kind of trichome occurrs in C. antisyphiliticus, C. gracilescens, and C. pycnadenius
Müller Argoviensis (1873: 216) and is characterized by the presence of up to ten ascending lateral rays. They differ
from stellate-porrect and stellate-rotate trichomes, both recognized by Webster et al. (1996), which have the lateral
rays parallel to the leaf blade.
v. Stellate-rotate (Fig. 6g, h): This type was found only in C. gracilescens and is recognized by the presence of 10 to
15 lateral rays united up to 30% of their length.
Foliar glands: These are cited for 72 genera and 377 species of Euphorbiaceae (Weber et al. 2015), highlighting the
family as the third largest in the world in number of species with these structures. In Croton, extrafloral nectaries are
generically called glands in taxonomic treatments (e.g., Caruzo & Cordeiro 2013, Sodré et al. 2017), and they are
usually found on the petioles and leaf blades and have been used in the differentiation of sections and species since the
early works of Baillon (1858, 1864) and Müller Argoviensis (1873). In this study, they were found in all 50 species, at
the apex of the petiole (= acropetiolar; Fig. 7a-c), at the base of the leaf blade (= basilaminar; Fig. 7d), and along the
leaf margins (Fig 7e, f) usually in the indentations (Fig. 8f). In 41 of the species studied, a pair or more of acropetiolar
or basilaminar glands were observed per leaf, however, these may be absent in C. planaltoanus and C. ramosissimus,
or vary from four to eight in C. agoensis Baillon (1864: 348) (Fig. 13v), C. catariae (Fig. 17u), C. gracilirameus
Silva et al. (2015a: 162) (Fig. 24b), C. mollis (Fig. 39b), C. sublepidotus (Fig. 39k), C. tetradenius (Fig. 39p), and
C. macrosepalus (Fig. 42o). Along the leaf margin, glands are distributed more or less randomly in C. agoensis, C.
catariae, and C. gracilirameus, whereas in other species they are restricted to the sinuses, usually solitary and facing
the abaxial surface, or sometimes paired, or on both surfaces as in C. parodianus. Regarding the shape of the glands
found on the petioles and leaf blades, they are usually patelliform (disc-shaped), and may be sessile (Fig. 7g-i), shortly
stipitate (when the length of the stipe is up to twice the diameter of the apical secretory region, Fig. 7j-1), or long-
stipitate with a longer stipe (Fig. 8a, b). The cylindrical and obconic glands vary in length/diameter ratio from 2:1 (Fig.
8d, f, g, h, i) to 10: 1 (Fig. 8e, j), with a slightly concave surface or with an apical orifice as in C. goyazensis (Fig. 8d).
Capitate glands are usually stipitate and marginal (Fig. 8k).
Stipules and leaves: The stipules in Croton sect. Geiseleria are generally persistent, the smaller ones 0.3-1 mm long
and fairly inconspicuous, sometimes covered by the indument of the branches (e.g., C. goyazensis, C. hadrianii, C.
planaltoanus), and larger ones up to 11 mm long and brownish (e.g., C. antisyphiliticus, C. krukoffianus, C. sclerocalyx)
or greenish (e.g., C. aberrans [Fig. 12f] and C. odontadenius). They are generally linear or linear-lanceolate, sometimes
oval, triangular or lanceolate, with an acute or rarely obtuse apex and usually chartaceous. The stipules have two to
eight subglobose or ovoid glands at base in most species studied (e.g., C. abaitensis, C. adenodontus, C. asperrimus,
C. desertorum, C. hadrianii, C. junceus, C. odontadenius, C. parodianus, and C. macrosepalus), but the glands are
stipitate-capitate in C. catariae and C. gracilirameus.
FIGURE 5. Diversity of trichomes in Croton sect. Geiseleria: a stellate-porrect with short lateral and porrect rays (Croton abaitensis),
b trichomes with long porrect rays (Croton macrosepalus), c stellate-porrect trichomes on pin-cushion, see trichome base in detail (C.
tetradenius), d stellate-porrect trichomes with porrect ray shorter than lateral rays (C.parodianus), e simple trichomes, see detail showing
the scars of lateral rays at the base (C. aberrans), f simple trichomes on the leaf blade and stellate-porrect trichomes along the midvein
(C. lundianus), g 2–7-radiate trichomes, with or without short lateral rays arising at the base (C. sclerocalyx), h 2–6-radiate trichomes (C.
odontadenius).

FIGURE 6. Diversity of trichomes in Croton sect. Geiseleria: a–d multiradiate (a and b Croton goyazensis, c C. inaequilobus and d C.
carinatus), e and f fasciculate (C. antisyphiliticus) and g and h stellate-rotate (C. gracilescens).
FIGURE 7. Diversity of foliar glands in Croton sect. Geiseleria. a–f photographs in situ, a Croton abaitensis, acropetiolar and stipitate, b
C. hadrianii, acropetiolar and sessile, c C. inaequilobus, acropetiolar and subsessile, d C. uliginosus, basilaminar and subsessile, e stipitate
glands of the leaf margin of C. agoensis, indicated by the white arrows, f sessile glands of the leaf margin of C. parodianus, indicated by
the black arrows. g–h SEM photographs; g–i sessile patelliform glands, g and h C. antisyphiliticus, i C. desertorum; j–l patelliform and
shortly stipitate glands, j C. gracilirameus, k C. inaequilobus, l C. lundianus.

FIGURE 8. Diversity of foliar glands in Croton sect. Geiseleria. a and b patelliform and long-stipitate, a Croton abaitensis, b C. lundianus;
c–e cylindrical, c C. carinatus, d C. goyazensis, e C. aberrans; f–j obconic, f C. parodianus, g C. rhodotrichus, h C. krukoffianus, i C.
asperrimus, j Croton macrosepalus; k stipitate-capitate, C. gracilirameus.
The leaves in Croton sect. Geiseleria are simple and spirally alternate, or else subopposite or subverticillate in the
dichotomy or trichotomy of the branches or immediately below the inflorescences, as in C. glandulosus (Fig. 9a) and C
. goyazensis (Fig. 22g). The leaves are always petiolate, with the petiole varying from 0.1 to 3.5 cm long, or sometimes
in C. hirtus and C. tetradenius they can reach 7 cm long. The petioles are cylindrical or angulose, discretely canaliculate
above, and always pubescent. Shapewise, the leaf blades are usually oval or lanceolate (e.g., C. adenodontus, C.
glandulosus, C. goyazensis, C. gracilirameus, C. hirtus, C. inaequilobus, C. parodianus [Fig. 9b], C. sclerocalyx, C.
spica, C. tetradenius [Fig. 9g], and C. trinitatis), but they can also be lanceolate (e.g., C. asperrimus, C. spica [Fig.
9c], C. uliginosus), linear or linear-lanceolate (e.g., C. carinatus [Fig. 9d], C. gracilescens, C. junceus), elliptic (e.g.,
C. krukoffianus, C. planaltoanus [Fig. 9e], C. macrosepalus), or oblong (e.g., C. agoensis [Fig. 9f], C. ramosissimus).
In Croton abaitensis, C. aberrans, C. agoensis, C. antisyphiliticus, C. lundianus, and C. odontadenius, however, the
leaf blades can present a combination of the shapes mentioned above. As for the dimensions, the leaf blades vary from
1.5–20.5 cm and 0.2–8.4 cm wide, with the smallest size recorded for C. gracilirameus, C. lundianus and C. uliginosus
(1.5–6 × 0.7–4.6 cm) and the largest for C. antisyphiliticus, C. hadrianii, C. hirtus, C. nepetifolius, C. tetradenius,
C. sincorensis Martius ex. Müller Argoviensis (1873: 86), and C. spica (8–20 × 6–8.4 cm). As for the margins, the
studied species have serrate blades (e.g., C. gracilirameus, C. hirtus, C. inaequilobus, C. planaltoanus [Fig. 9e] and
C. uliginosus), serrulate (e.g., C. adenodontus [Fig. 13p], C. asperrimus [Fig. 17f], C. hadrianii [Fig. 24k], doubly-
serrate (e.g., C. aberrans, C. antisyphiliticus, C. gracilescens, C. krukoffianus, C. lundianus (Fig. 28t), C. odontadenius
and C. sipaliwinensis [Fig. 36o]), crenate (e.g., C. goyazensis, C. pycnadenius and C. parodianus [Fig. 9h, 31j]), or
entire to subentire (e.g., C. carinatus [Fig. 9d], C. junceus, C. tetradenius [Fig. 9g], C. rhodotrichus, and Croton
macrosepalus). The apex of the leaf blades is usually acute or shortly acuminate, except in C. tetradenius, where it is
slightly caudate (Fig. 9g) and in C. agoensis, C. pycnadenius, and C. spica, where it is aristate or caudate (Fig. 9f). The
leaf bases are usually obtuse, although there are also slightly cordate ones (e.g., C. abaitensis, C. agoensis [Fig. 9f], C.
goyazensis, C. ramosissimus, C. sclerocalyx, C. spica and C. trinitatis), conspicuously cordate (e.g., C. gracilirameus,
C. parodianus [Fig. 9h]), rounded (e.g., C. adenodontus, C. asperrimus, C. hadrianii [Fig. 24k], or sometimes acute
(e.g., C. odontadenius, C. planaltoanus [Fig. 9e]), truncate (e.g., C. trinitatis [Fig. 42a]), or asymmetric bases (e.g., C.
odontadenius [Fig. 31f]. Most of the species have membranaceous leaves, except for C. antisyphiliticus, C. agoensis,
C. carinatus, C. goyazensis, C. nepetifolius, C. pycnadenius, and C. parodianus, which have chartaceous blades. The
foliar venation pattern in the species studied is brochidodromous in most of species, e.g., C. gracilescens [Fig. 21t], C.
glandulosus [Fig. 21h], C. rhodotrichus [Fig. 36a], C. tetradenius (Fig. 39o), actinodromous-brochidodromous, with 5
or 7 veins leaving from the base of the blade (e.g., C. abaitensis [Fig. 13a], C. adamantinus Müller Argoviensis (1873:
115) [Fig. 13k], C. catariae [Fig. 17t] e C. nepetifolius [Fig. 31a]), semicraspedodromous (i.e., C. hirtus [Fig. 24u],
C. sipaliwinensis [Fig. 36o]), craspedodromous (i.g., C. lundianus [Fig. 28t], C. trinitatis [Fig. 38a]), or apparently
absent in C. junceus and C. carinatus [Fig. 9d], due to secondary veins immersed in the leaf blade and therefore
poorly visible. Commonly on the blades of all species the primary and secondary veins are impressed on the adaxial
surface or prominent on the abaxial surface. The leaves are pubescent on both surfaces, varying only in the density of
the trichomes, which is usually greater on the abaxial surface. They are often discolorous, although in some species
the color difference between the surfaces is very tenuous, as in C. aberrans, C. planaltoanus (Fig. 34 a,b) and C.
uliginosus.
Inflorescences: The inflorescences in the species studied are contracted bisexual thyrses, often with a racemose
appearance, with basal pistillate flowers and distal staminate ones. Generally the pistillate flowers open prior to the
staminate ones. In most species, the staminate flowers are continuous with the pistillate ones along the inflorescence
axis (Fig. 10 a), but in C. krukoffianus, C. lundianus, and C. sclerocalyx, the pistillate flowers are separated from the
staminate ones by a sterile zone (Fig. 10b). The inflorescences are usually terminal, although they may also occur
in the dichotomy of branches. Lengthwise, the inflorescences vary from less than 0.7 cm long in C. trinitatis (Fig.
10c) to 16.5 cm long in C. carinatus and C. tetradenius. The number of pistillate cymules per thyrse varies from one
to twenty, with the lowest number (one or two) found in C. uliginosus and the largest (8–20) in C. abaitensis and C.
hirtus, always with one flower per cymule. Staminate cymules are generally numerous and can be grouped laxly (Fig.
10d) or congested (Fig. 10e) in the thyrses, with 1–6 flowers per cymule.

FIGURE 9. Leaf patterns in species of C. sect. Geiseleria. a leaves alternate along the branch and whorled below the dichotomy of
the branches in Croton glandulosus, b leaf blades ovate in C. parodianus, c oblong in C. spica, d linear in C. carinatus, e elliptic in C.
planaltoanus, f leaf base slightly cordate and apex acuminate in C. mucronifolius, g leaf base rounded and apex attenuate in C. tetradenius,
h base cordate in C. parodianus.
FIGURE 10. Patterns of inflorescences and staminate flowers in Croton sect. Geiseleria. a inflorescences without a sterile zone between
staminate and pistillate flowers, the latter distributed in more than half of the inflorescence axis in C. hirtus, b thyrses with a sterile zone
between staminate and pistillate flowers in C. lundianus, c short inflorescences with few flowers in C. trinitatis, d lax thyrses in C. junceus,
e congested thyrses in C. spica, f staminate flowers with short pedicel and calyx united for 1/2 the length in C. odontadenius, g staminate
flowers with long pedicels and calyx shortly connate at the base in C. goyazensis.

Bracts: The bracts of members of Croton sect. Geiseleria resemble the stipules and are located at the base of the
pedicels of both staminate and pistillate flowers. They are usually linear, linear-lanceolate, or lanceolate, but sometimes
oval, oval-lanceolate, elliptic, oblong, or triangular, and usually persistent, sometimes even after the fall of the flowers.
Their dimensions vary between 0.4–7 mm in length and 0.1–2.6 mm in width. In most species, the bracts have globose,
subglobose, ellipsoid or ovoid glands at the insertion with the axis of the inflorescence. Pyriform glands were observed
in the bracts of C. adenodontus, C. glandulosus, C. rhodotrichus, and C. macrosepalus; in C. hirtus they are stipitate-
pyriform, in C. asperrimus they are patelliform and stipitate; and in C. gracilirameus they are stipitate-capitate.
Staminate flowers: The staminate flowers are pedicellate, actinomorphic, with developed sepals and petals, staminal
filaments inflexed in bud, and segmented, glabrous or pubescent discs. In general, in the genus they are usually treated
as morphologically homogeneous and not very informative in differentiating species by several authors (e.g., Croizat
1941, Gomes 2006, Silva 2009). However, in the species studied, the staminate flowers showed variation in pedicel
size, calyx connation, number of sepals and petals, number of stamens, and also the presence or absence of indument,
which are useful in the delimitation of the species. The pedicels can be very short (up to 0.4 mm long, Fig. 10f), as
in C. odontadenius, C. trinitatis and C. sclerocalyx, or long (up to 5 mm long) in C. goyazensis (Fig. 10g) and C.
gracilirameus. The perianth is normally pentamerous, except in C. aberrans, where it is 6-merous and in C. sclerocalyx
where it varies from 3–5-merous. The calyx is generally shortly joined in the basal 1/4 to 1/6 of its length (Fig. 11g), or
up to half its length (Fig. 10f) in C. aberrans, C. odontadenius, C sclerocalyx, and C. sipaliwinensis. These lobes are
generally oval, with an acute or obtuse apex, greenish (Fig. 10a), yellow-whitish (Fig. 10d, g), brownish (Fig. 10c, f),
or salmon (Fig. 35h), glabrous on the inner face, with sparse or dense trichomes on the outer face. The petals are free,
whitish, generally elliptic, and obovate or oblanceolate, although they can also be oblong, linear-lanceolate, lanceolate,
or oval-lanceolate, with an obtuse or rounded apex. They are always ciliate along the margins and may have long basal
trichomes on the outer face; however, they can also be pubescent on both surfaces, but especially above on the inner
side, which is usually villous, strigose, or glabrescent. The androecium in the species studied usually consists of 11
free stamens, one of them central to the rest, with two concentric series of 5, practically undifferentiated in size, or
else it may vary from 11 to 24 stamens in C. odontadenius, 14 to 16 in C. tetradenius, 8 to 11 in C. lundianus and C.
trinitatis, 7 or 8 in C. uliginosus, or just 5 stamens, as in C. sipaliwinensis. The filaments can be glabrous or villous,
either in part or overall. The disc is 5-segmented, with yellowish, usually suborbicular or subovoid segments, and the
receptacle is either villous or glabrescent.
Pistillate flowers: These are usually monochlamydeous or sometimes dichlamydeous; in the latter case the flowers are
apparently monochlamydeous with petals usually reduced to glands. In the species studied, the calyx is valvate, with
sepals equal or more often unequal in size. The pedicels are cylindrical, fairly short (0.1–2 mm long) in most species
(Fig. 11b), or they can be longer (3–7.5 (–11) mm long) in C. goyazensis and C. gracilirameus (Fig. 11a). The calyx is
free, pentamerous in most species, but with 6 to 8 sepals in C. aberrans, C. hadrianii, C. krukoffianus, C. lundianus,
C. odontadenius, C. sclerocalyx, C. sincorensis, and C. uliginosus. The sepals are strongly unequal in C. abaitensis,
C. adenodontus, C. hirtus, C. inaequilobus, C. rhodotrichus (Fig. 11c), C. spica, C. parodianus, C. macrosepalus,
and in some individuals of C. glandulosus, being equal or moderately unequal in the other species (Fig. 11a, b, d, e).
The sepals are quite variable in shape (e.g., obovate, oblong, oblanceolate, elliptic, lanceolate, linear, triangular, or
spatulate), and their apices are acute (Fig. 11c, d) or sometimes acuminate. They are greenish (Fig. 11a, b) or white
(Fig. 11d, e), with trichomes on the outer surface or on both surfaces, with hyaline, whitish or sometimes brownish or
reddish colors. In some species it is possible to observe petals, usually small and in the form of glands (ovoid, subulate,
subglobose or filiform. The ovary is subglobose or oblong, usually densely pubescent, but sometimes glabrate as in C.
aberrans and C. sclerocalyx. The stigmas are 2-fid (with 6 terminal tips; Fig. 11a, c, d) in most species (40), 4-fid (with
12 terminal tips; Fig. 11b) in C. antisyphiliticus, C. hadrianii, C. lundianus, C. odontadenius, and C. sclerocalyx, and
8–16(–32)-fid (multifid, with 24–48(–96) terminal tips) in C. krukoffianus. They are glabrous in C. asperrimus and C.
trinitatis, but pubescent in most species. The disc in most species is 5-segmented, but the number of segments equals
that of the sepals, and it is usually completely adhered to the base of the sepals. The receptacle is glabrous, glabrescent,
or pubescent.
FIGURE 11. Pistillate flowers (a–e) and fruits (f–k) of some members of Croton sect. Geiseleria. a flower with long pedicel and glandular
sepals in C. gracilirameus, b flower with short pedicel and 4-fid stigmas in C. hadrianii, c 5-merous flower with unequal sepals in C.
rhodotrichus, d 6-merous flower with equal sepals in C. lundianus, e 6-merous flowers with 2-fid stigmas in C. uliginosus, f greenish
capsules with whitish indument in C. adenodontus, g green-lustrous capsule with hyaline indument and short pedicel in C. aberrans, h
green capsules with white lines on the sutures of the septa and locules in C. glandulosus, i light green capsule in C. gracilirameus (note
also the glandular-stipitate sepals), j brown capsules with long pedicels in C. goyazensis, k light green capsule with brownish trichomes
in C. rhodotrichus.

Fruits: The fruits in Croton sect. Geiseleria are globose, subglobose, or oblongoid capsules. They vary from 3.3 to 7 mm
in length, with the shortest ones found in C. adenodontus (3.3–3.5 × 3.3–3.5 mm) (Fig. 11f) and C. trinitatis (3.7–3.9
× 3.5–3.8 mm) and the longest ones in C. aberrans (6.5–7 × 7–7.3 mm) and C. triangularis Müller Argoviensis (1865:
128) (8–9 mm, Fig. 11g). Shortly pedicellate capsules (pedicels 0.3–1 mm long) are found in C. antisyphiliticus, C.
desertorum, C. glandulosus, C. gracilescens, and C. sclerocalyx, whereas filiform pedicels up to 9 mm long are found
in C. goyazensis (Fig. 11i) and in C. gracilirameus. The capsules are predominantly opaque greenish with whitish or
hyaline indument (Fig. 11f, h, i), but can be lustrous in C. aberrans (Fig. 11g), C. lundianus, C. odontadenius, and
C. sclerocalyx, or else they can have dark red trichomes in C. rhodotrichus (Fig. 11k). The capsules generally have a
persistent calyx, sometimes exceeding the capsule length in C. sclerocalyx and Croton macrosepalus, or when smaller
reaching at least 1/4 of its length.
Seeds: Seeds in the section are carunculate, maculate, and oblong to ellipsoid. In the species studied, elliptic, oblong,
ovoid, elliptic or suborbicular seeds were found, with a well developed caruncle (most species), rudimentary one
(e.g., C. asperrimus), or ecarunculate (e.g., C. adenodontus). The caruncles are oval, triangular, hat-shaped, elliptic,
reniform, or amorphous. They are usually sessile or shortly stipitate in C. rhodotrichus, and can be visible only on
the dorsal side (e.g., C. antisyphiliticus, C. gracilescens) or visible on both sides of the seed (e.g., C. goyazensis, C.
gracilirameus, C. rhodotrichus). As for the texture, the seeds have a crustaceous testa and are closely foveolate, with
brownish or grayish colors.
Taxonomic treatment
Croton sect. Geiseleria (Gray 1856: 391) Baillon (1858: 359) = Croton subgenus Geiseleria Gray (1856: 391). Type:
Croton glandulosus L.
Herbs or subshrubs, erect or decumbent, or shrubs up to 5 m tall, annual or perennial, sometimes xylopodiferous
(with a thickened, underground stem), monoecious; latex scarce or abundant; trichomes stellate-porrect, multiradiate,
fasciculate, stellate-rotate, or simple. Leaves simple, always petiolate, alternate, sometimes subopposite or subverticillate
near the dichotomy or trichotomy of the branches or near the inflorescences, almost always with 2 (–8) sessile or
stipitate glands at the base of the leaf blade or at the apex of the petiole; margins serrate to crenate, often with sessile
or stipitate glands in the sinuses. Inflorescences thyrses, bisexual, lax or congested, terminal or in the dichotomies or
trichotomies of the branches, pistillate cymules 1–20 in the basal portion of the rachis with one flower per cymule,
staminate cymules with 1–6 flowers, continuous with the pistillate ones or else separated from them by a bare, sterile
zone; bracts frequently with sessile or stipitate, persistent glands. Staminate flowers actinomorphic, dichlamydeous,
calyx (3–)5(–6)-lobed, lobes united at the base or up to half their length, petals (3–)5(–6), free, stamens (5–)11(–24),
glabrous or villous. Pistillate flowers actinomorphic, zygomorphic or asymmetric, sepals 5–8, free, unequal or equal
in size, valvate aestivation; petals, when present, vestigial and glanduliform or laminar, stigmas 2–4(–16)-fid, with 6–
12(–48) terminal tips, glabrous or pubescent. Capsules with persistent styles, stigmas and sepals, the latter frequently
accrescent, columella irregularly fimbriate at the apex; seeds usually carunculate, foveolate, uniformly colored or
maculate.+-
Key to the species of Croton sect. Geiseleria occurring in Brazil

1. Leaf margins entire or subentire ....................................................................................................................................................... 2
- Leaf margins serrate, doubly serrate, serrulate, crenate or erose ................................................................................................... 13
2. Leaf blades linear, up to 1 cm wide .................................................................................................................................................. 3
- Leaf blades ovate, elliptic or lanceolate, 1.5–8 cm wide .................................................................................................................. 5
3. Inflorescences congested; staminal filaments villous; pistillate sepals unequal ........................................................... 30. C. mollis
- Inflorescences lax; staminal filaments glabrous; pistillate sepals equal .......................................................................................... 4
4. Petioles without glands, or rarely with cylindric glands; staminate bracts 1.8–3.3 mm long; pistillate sepals linear-lanceolate .......
................................................................................................................................................................................... 11. C. carinatus
- Petioles with patelliform glands; staminate bracts 0.8–1.3 mm long; pistillate sepals ovate ..................................... 26. C. junceus
5. Inflorescence with a distinct sterile zone between pistillate and staminate flowers; pistillate sepals 6–8; stigmas multifid, with
24–48 terminal tips .............................................................................................................................................. 27. C. krukoffianus
- Inflorescence without a distinct sterile zone between pistillate and staminate flowers; pistillate sepals 5, sometimes 6 or 7 in C.
hadrianii; stigmas (2–)4-fid, with (6–)12 terminal tips .................................................................................................................... 6
6. Petioles 0.1–0.2 cm long; leaf blades with an aristate or caudate apex; pistillate sepals linear-lanceolate; stigmas glabrous ...........
.............................................................................................................................................................................. 45. C. sublepidotus
- Petioles 0.6–7.1 cm long; leaf blades with an acute or acuminate apex; pistillate sepals obovate, oblong, elliptic or ovate; stigmas
stellate ............................................................................................................................................................................................... 7
7. Petiolar glands patelliform and sessile ............................................................................................................................................. 8
- Petiolar glands cylindric, obconic, or stipitate-patelliform .............................................................................................................. 9
8. Leaf blades ovate, oval-lanceolate, or less often elliptic or oblong; inflorescences with 2–4(–6) pistillate flowers; pistillate sepals
5–7; stigmas (2–)4-fid; fruiting pedicel 0.5–1 mm long .......................................................................................... 21. C. hadrianii
- Leaf blades elliptic; inflorescences with 8–12 pistillate flowers; pistillate sepals 5; stigmas 2-fid; fruiting pedicel 1.5–2 mm long
................................................................................................................................................................................ 44. C. subincanus
9. Aromatic plants (when dry); petioles 1.1–7.1 cm long; leaves usually whorled below the inflorescences or the dichotomies of
the branches, with acuminate apex and actinodromous-brochidodromous venation; pistillate sepals equal or subequal; stamens
(11–)14–16 ............................................................................................................................................................. 46. C. tetradenius
- Non-aromatic plants; petioles 0.6–1(–1.5) cm long; leaves predominantly alternate or rarely opposite immediately below the
inflorescences, with acute apex and brochidodromous venation; pistillate sepals unequal; stamens 11 ....................................... 10
10. Stipules 4.8–10 mm long; pistillate sepals ovate or oval-lanceolate, covering the fruit ................................... 29. C. macrosepalus
- Stipules 0.8–4 mm long; pistillate sepals spatulate-obovate, oblanceolate, oblong or linear-lanceolate, not covering the fruit ... 11
11. Leaves with 2–8 acropetiolar glands; bracts 2.5–4 mm long; stamens with villous filaments and anthers with acute apex; stigmas
2–4-fid ........................................................................................................................................................................... 30. C. mollis
- Leaves with 2(4) acropetiolar glands; bracts 1–2 mm long; stamens with glabrous filaments and anthers with truncate or obtuse
apex; stigmas 2-fid ......................................................................................................................................................................... 12
12. Leaf margins with patelliform glands; inflorescence with 8–12 pistillate flowers; bracts without glands; seeds not maculate, the
caruncle sessile ...................................................................................................................................................... 24. C. hostmannii
- Leaf margins with ovoid or globose glands; inflorescence with 2–4 pistillate flowers; bracts with glands; seeds maculate with a
stipitate caruncle ................................................................................................................................................. 38. C. rhodotrichus
13. Pistillate flowers with 6–8(–10) sepals ........................................................................................................................................... 14
- Pistillate flowers with 5 sepals ....................................................................................................................................................... 22
14. Acropetiolar glands narrowly cylindric; inflorescences less than 1.5 cm long ........................................................... 2. C. aberrans
- Glands acropetiolar or basilaminar, patelliform, obconic or largely cylindric; inflorescences 4–20 cm long ............................... 15
15. Thyrses without a distinct sterile zone between pistillate and staminate flowers .......................................................................... 16
- Thyrses with a distinct sterile zone between pistillate and staminate flowers ............................................................................... 19
16. Petioles 0.1–0.3 cm long; leaf blades linear-lanceolate, without glands in the sinuses; stigmas usually 2-fid ....... 49. C. uliginosus
- Petiole 0.5–3.5 cm long, leaf blades ovate, oblong or elliptic, with glands in the sinuses; stigmas 4- or multifid, rarely 2-fid ... 17
17. Decumbent subshrubs 0.1–0.6 m tall; leaf bases usually acute ......................................................................... 33. C. odontadenius
- Shrubs 0.5–4 m tall; leaf bases obtuse, rounded or slightly cordate .............................................................................................. 18
18. Shrubs with erect stems; petiolar glands sessile; leaf margins serrulate or crenulate, with patelliform sessile glands in the sinuses;
stigmas (2–)4-fid ...................................................................................................................................................... 21. C. hadrianii
- Shrubs with pendent stems; petiolar glands stipitate; leaf margins serrate or crenate, with cylindric or stipitate-patelliform glands
in the sinuses; stigmas multifid ............................................................................................................................. 41. C. sincorensis
19. Pistillate flowers with rigid sepals, lobed or deeply serrate at the apex, covering the gynoecium and fruit ........ 39. C. sclerocalyx
- Pistillate flowers with membranaceous sepals, entire or sometimes serrulate, not covering the gynoecium and fruit ................. 20
20. Stamens 5; ovary glabrous or glabrescent ........................................................................................................ 42. C. sipaliwinensis
- Stamens 8–11; ovary stellate .......................................................................................................................................................... 21
21. Petioles 3 to 5 times shorter than the leaf blades; pistillate flowers with oblanceolate or obovate, whitish sepals; stigmas 2(–4)-
fid ............................................................................................................................................................................. 28. C. lundianus
- Petiole 8 to 20 times shorter than the leaf blades; pistillate flowers with oblong, elliptic, lanceolate or linear, greenish sepals;
stigmas 8–16(–32)-fid ......................................................................................................................................... 27. C. krukoffianus
22. Prostrate subshrubs, petiole length equal to that of the leaf blade, or up to half as long; leaf blades reniform or largely ovate ........
.......................................................................................................................................................................... 16. C. glechomifolius
- Erect shrubs or subshrubs, petioles 3 to 30 times shorter than the leaf blades; leaf blades ovate, elliptic, oblong, lanceolate or
linear ............................................................................................................................................................................................... 23
23. Acropetiolar or basilaminar glands sessile, patelliform, often absent in C. ramosissimus ............................................................ 24
- Acropetiolar or basilaminar glands stipitate-patelliform, obconic or cylindric, often absent in C. planaltoanus and C.
sublepidotus .................................................................................................................................................................................... 33
24. Pistillate flowers with sepals externally stellate and internally glabrous; stigmas glabrous or sometimes glabrescent; staminate
flowers with petals internally villous and externally glabrous ....................................................................................................... 25
- Pistillate flowers with sepals stellate on both surfaces; stigmas stellate; staminate flowers with petals villous on both surfaces 27
25. Few-branched subshrubs less than 0.5 m tall; stems brownish; petioles 0.5–2 cm long; rough-textured leaves; stigmas 4-fid;
capsules hirsute .................................................................................................................................................. 6. C. antisyphiliticus
- Usually much-branched subshrubs 0.6–1.2 m tall; light green, with yellowish or ferrugineous stems; petioles 0.1–0.3(–1) cm long;
leaves not rough-scabrous; stigmas 2-fid; capsules tomentose or pubescent ................................................................................. 26
26. Leaf blades oval-lanceolate, elliptic or sometimes oblong, margins serrate and usually without glands, discolorous, yellowish or
light green, densely pubescent on the abaxial surface with whitish trichomes; thyrses congested; ovary hirsute .............................
................................................................................................................................................................................ 14. C. desertorum
- Leaf blades linear or narrowly elliptic, margins serrate or serrulate with glands in the sinuses, concolorous, dark green, sparsely
pubescent on the abaxial face with hyaline trichomes; thyrses lax; ovary pubescent ......................................... 18. C. gracilescens
27. Plants densely branched and appearing monopodial; petioles and base of the leaf blades usually without glands; inflorescences lax;

stamens villous; capsules and seeds oblongoid ................................................................................................. 37. C. ramosissimus
- Plant not densely branched and appearing sympodial; acropetiolar or basilaminar glands always present; inflorescences congested;
stamens glabrous; capsules globose, seeds ovoid, ellipsoid or orbicular ....................................................................................... 28
28. Flowers in obvious thyrses; stigmas 4-fid ...................................................................................................................................... 29
- Flowers in raceme-like thryses; stigmas 2-fid ................................................................................................................................ 30
29. Stipules 0.2–0.6 mm long; petioles 6 to 8 times shorter than the leaf blades; leaf blades with obtuse or rounded base and margins
doubly crenulate or serrulate; bracts 0.7–1.1 mm long ............................................................................................ 21. C. hadrianii
- Stipules 4–8 mm long; petioles 3 to 4 times shorter than the leaf blades; leaf blades with cordate base and margins doubly serrate
or erose; bracts 2–5 mm long ............................................................................................................................... 32. C. nepetifolius
30. Leaf blades elliptic, the base base usually cuneate, the margins serrulate, without glands .................................. 44. C. subincanus
- Leaf blades ovate, oblong, lanceolate, or rarely elliptic with obtuse, rounded or lightly cordate base, the margins serrate, usually
with glands in the sinuses ............................................................................................................................................................... 31
31. Subshrubs less than 1 m tall; leaf blades oblong or lanceolate with obtuse or slightly cordate base and acuminate, aristate or caudate
apex; bracts 3–6 mm long; stigmas 3.5–6 mm long ........................................................................................................ 43. C. spica
- Shrubs 0.8–2.3 m tall; leaf blades ovate or oval-lanceolate with obtuse, rounded or truncate base and usually acute apex; bracts 2–3
mm long; stigmas 1.5–2.3 mm long ............................................................................................................................................... 32
32. Leaf glands acropetiolar, usually reddish; inflorescences 1.2–2.9 cm long, with 2 or 3 pistillate flowers; disk 4-segmented; seeds
maculate .............................................................................................................................................................. 25. C. inaequilobus
- Leaf glands basilaminar, yellowish; inflorescences 2.1–8 cm long, with 6–12 pistillate flowers, disk 5-segmented; seed not
maculate ............................................................................................................................................................ 40. C. seccoi sp. nov.
33. Stigmas 4–6-fid .............................................................................................................................................................................. 34
- Stigmas 2-fid .................................................................................................................................................................................. 35
34. Stipules 8–12 mm long, leaf blades elliptic, oblong or oval-elliptic; thyrses 1.5–3.5 cm long; stamens 14 or 15; pistillate sepals
with even margins ............................................................................................................................................. 10. C. campinarensis
- Stipules 3–5 mm long, leaf blades triangular or ovate; thyrses 4.5–25 cm long; stamens 10 or 11; pistillate sepals with revolute
margins ................................................................................................................................................................. 47. C. triangularis
35. Petioles 10 to 30 times shorter than the leaf blades ........................................................................................................................ 36
- Petioles 4 to 8 times shorter than the leaf blades ............................................................................................................................ 40
36. Leaves with patelliform or obconic glands, obtuse or rounded base and acute apex; bracts with glands; pistillate sepals elliptic;
fruiting calyx as long as the capsule ........................................................................................................................ 8. C. asperrimus
- Leaves with cylindric and/or stipitate-capitate glands, base slightly cordate and apex acuminate, aristate or caudate; bracts without
glands; pistillate sepals lanceolate or oval-lanceolate; fruiting calyx up to one third of the capsule length .................................. 37
37. Acropetiolar glands usually lacking; leaf blades concolorous, with sparse trichomes and impressed and inconspicuous secondary
veins on both surfaces; stigmas glabrous ............................................................................................................ 45. C. sublepidotus
- Acropetiolar/basilaminar glands evident; leaf blades discolorous, with dense trichomes and conspicuous secondary veins on
abaxial surface; stigmas pubescent ................................................................................................................................................. 38
38. Leaf blades membranaceous, not bullate on the adaxial surface, with irregularly serrate margins and tertiary veins impressed on
the abaxial surface; stipules and pistillate sepals usually with glands; seeds maculate .............................................. 5. C. agoensis
- Leaf blades chartaceous, bullate on the adaxial surface with crenulate ou crenate-serrate margins and tertiary veins prominent on
the abaxial surface; stipules and pistillate sepals without glands; seeds not maculate .................................................................. 39
39. Decumbent plants; leaf blades largely ovate or orbicular with cordate base and eglandular margins; staminate cymules with 2
flowers; capsules greenish ................................................................................................................................... 36. C. pycnadenius
- Erect plants; leaf blades oval-lanceolate, oblong or elliptic with obtuse or slightly cordate base and glandular margins; staminate
cymules with 3–5 flowers; capsules yellowish or withish ................................................................................ 31. C. mucronifolius
40. Stipules, leaf margin, bracts, and pistillate sepals with stipitate-capitate glands ........................................................................... 41
- Stipules, leaf margin, bracts, and pistillate sepals with or without glands of other types .............................................................. 42
41. Stems delicate and shortly tomentose; leaf blades 2.4–4.6 × 1.4–4.2 cm with slightly cordate base, margins with sparse glands,
secondary veins impressed or discretely prominent on abaxial surface, and tertiary veins inconspicuous; stipules, bracts and
pistillate sepals with few, discreet glands .......................................................................................................... 19. C. gracilirameus
- Stems robust and velutinous; leaf blades 4.7–7.8 × 3.7–5.7 cm with cordate base, margins with dense and regularly distributed
glands, secondary and tertiary veins conspicuous and prominent on abaxial surface; stipules, bracts and pistillate sepals with
numerous, conspicuous glands ................................................................................................................................... 12. C. catariae
42. Pistillate flowers with equal sepals ................................................................................................................................................. 43
- Pistillate flowers with unequal sepals ............................................................................................................................................. 50
43. Leaf glands not visible to the eye; pistillate sepals with revolute margins .................................................................................... 44
- Leaf glands visible to the eye; pistillate sepals with even margins ................................................................................................ 45
44. Petioles 1.3–2.5 cm long; leaf blades 5.5–7 × 2.8–4 cm with acuminate apex, serrate margins, and actinodromous-brochidodromous
venation .................................................................................................................................................................. 13. C. crustulifer
- Petioles 0.4–0.8 cm long; leaf blades 2–4.8 × 0.9–2 cm with acute apex, serrulate margins, and brochidodromous venation ..........
....................................................................................................................................................................................... 22. C. harleyi
45. Pistillate flowers with gynoecium equal to or up to twice as long as the pedicel .......................................................................... 46
- Pistillate flowers with gynoecium 3 to 8 times longer than the pedicel ......................................................................................... 48
46. Leaf venation craspedodromous or semi-craspedodromous; thyrses 0.3–1.5 cm long; staminate flowers 1.5–2.6 mm long, the
pedicels and calyx glabrous or glabrescent; stigmas 0.8–1 mm long ........................................................................ 48. C. trinitatis
- Leaf venation brochidodromous; thyrses 2.5–8 cm long; staminate flowers 4–10 mm long, with the pedicel and calyx densely
pubescent; stigmas 2–3 mm long ................................................................................................................................................... 47
47. Non-aromatic subshrubs 0.3–1 m tall; indument on young branches and inflorescences canescent; leaves with cylindric or conic
basilaminar glands with a pronounced orifice at the apex; thyrses with 2–6 pistillate flowers; capsules oblongoid .........................
................................................................................................................................................................................. 17. C. goyazensis
- Aromatic shrubs (even when dry) 1.2–2 m tall; indument brownish or ferrugineous on young branches and inflorescences; leaves
with patelliform basilaminar glands without an orifice; thyrses with 8–20 pistillate flowers; capsules globose .. 20. C. grewioides
48. Petioles with 2–6 cylindric or obconic glands; leaf blades with caudate apex and eglandular margins; stamens (11–)14–16; stigmas
densely pubescent .................................................................................................................................................. 46. C. tetradenius
- Petioles with 2 stipitate-patelliform glands; leaf blades with acute or acuminate apex and glandular margins; stamens 9–11; stigmas
glabrous or glabrescent ................................................................................................................................................................... 49
49. Shrubs 1.2–2 m tall; leaf base cordate; bracts eglandular; pistillate sepals ovate or elliptic; capsules oblong, uniformly greenish ..
............................................................................................................................................................................... 3. C. adamantinus
- Herbs or subshrubs 0.1–1 m tall; leaf base obtuse; bracts glandular; pistillate sepals obovate, oblanceolate or spatulate; capsules
globose, green, but whitish on the sutures of the septa and locules ..................................................................... 15. C. glandulosus
50. Stems fistulose in cross-section; stems and leaves hirsute; bracts with stipitate glands ................................................ 23. C. hirtus
- Stems solid in cross-section; stems and leaves pubescent or tomentose; bracts with or without sessile glands ............................ 51
51. Petioles with 2–8 glands; leaf blades narrowly elliptic or linear-lanceolate, 0.5–1.8 cm wide, with 8–10 pair of secondary veins;
stigmas 2–4-fid .............................................................................................................................................................. 30. C. mollis
- Petioles with or without 2(4) glands; leaf blades ovate, lanceolate, elliptic or orbicular, 2–8.6 cm wide, with 2–6 pair of secondary
veins; stigmas 2-fid ......................................................................................................................................................................... 52
52. Leaf margins without glands .......................................................................................................................................................... 53
- Leaf margins with glands ............................................................................................................................................................... 55
53. Shrubs 0.7–1.5 m tall; petioles 0.2–0.5 cm long; leaf blades chartaceous; inflorescences with 7–10 pistillate flowers; bracts without
glands; stigmas stellate; stamens with villous filaments ............................................................................................. 7. C. arenosus
- Herbs or subshrubs 0.1–0.5 m tall; petioles 0.7–3 cm long; leaf blades membranaceous; inflorescences with 2–5 pistillate flowers;
bracts with glands; stigmas glabrous or glabrescent; stamens with glabrous filaments ................................................................. 54
54. Leaf blades with acuminate apex; bracts 2–3 mm long; capsules uniformly greenish, pedicels 2–4 mm long; endemic to Espírito
Santo and Rio de Janeiro states ................................................................................................................................ 9. C. bidentatus
- Leaf blades with acute apex; bracts 1.2–1.8 mm long; capsules green, but whitish on the sutures of the septa and locules, pedicels
0.3–0.8 mm long; widespread in the Neotropics .................................................................................................. 15. C. glandulosus
55. Few-branched subshrubs less than 0.5 m tall; leaf blades concolorous, usually without acropetiolar or basilaminar glands and with
cuneate or acute base .......................................................................................................................................... 34. C. planaltoanus
- Much-branched shrubs or subshrubs 0.7–4 m tall; leaf blades discolorous, with acropetiolar or basilaminar glands and with obtuse,
cordate or rounded base .................................................................................................................................................................. 56
56. Inflorescences congested, 0.5–1.7 cm long, with 8–12 staminate flowers; staminate bracts 0.4–1.2 mm long .................................
................................................................................................................................................................................ 4. C. adenodontus
- Inflorescences lax, 2–25 cm long, with 20-many staminate flowers; staminate bracts 1.5–6 mm long ........................................ 57
57. Leaf blades with obtuse or rounded base; flowers in raceme-like thryses; bracts with pyriform glands ...................... 40. C. seccoi
- Leaf blades with slightly or strongly cordate base; flowers in obvious thyrses; bracts with ellipsoid or ovoid glands ................ 58
58. Pistillate flowers with ovate, oblong, or elliptic sepals and glabrous or glabrescent stigmas; stamens with glabrous filaments;
species of the Caatinga Biome .............................................................................................................................. 50. C. virgultosus
- Pistillate flowers with usually obovate sepals; stigmas densely pubescent; stamens with villous filaments; species of the Cerrado
or Amazonian Biome ...................................................................................................................................................................... 59
59. Subshrubs 0.6–1.2 m tall; petioles 0.4–1.5(–2) cm long; leaf blades with slightly cordate or sometimes obtuse base, 3 or 4 pairs of
secondary veins, margins with only one gland per sinus only on abaxial surface; species of the Cerrado Biome .............................
.................................................................................................................................................................................... 1. C. abaitensis
- Shrubs 1–2.5 m tall; petioles 2–5(–7) cm long; leaf blades with strongly cordate base, 4–6 pairs of secondary veins; margins with
glands on both surfaces, two glands per sinus; species of the Amazonian Biome ................................................ 34. C. parodianus
1. Croton abaitensis Baillon (1864: 337).—Oxydectes abaitensis (Baill.) Kuntze (1891: 610). Type:—BRAZIL. Minas
Gerais: Abaité, 1844, H.A. Weddell 1818 (holotype: P 00623674!; isotypes: A 00047215!, F V0056098F!). (Figs.
12a–e, 13a–e).
Croton longinervius Müller Argoviensis (1873: 215).—Croton longinervius var. major Müller Argoviensis (1873: 215); nom. inval.—
Oxydectes longinervia (Müll.Arg.) Kuntze (1891: 613). Type:—BRAZIL. Goiás: “Serra de Christaës”, s.d., J.B.E. Pohl 826
(lectotype: W 0051292!, designated by Sodré et al. 2017; isolectotype: G 00434553!).
Croton longinervius var. minor Müller Argoviensis (1873: 216). Type:—BRAZIL. Goiás: ad Rio S. Marcos prope S. Luzia, no date, J.B.E.
Pohl 1637 (lectotype: W 0051293!, designated by Sodré et al. 2017; isolectotypes: A 00257952!, G 00434554!, K 000186104!, K
000252628!, W 0051294!).
Croton megaponticus Müller Argoviensis (1873: 210).—Oxydectes megapontica (Müll.Arg.) Kuntze (1891: 613). Type:—BRAZIL. Goiás:
“prope Megaponte”, s.d., J.B.E. Pohl 1110 (lectotype: W 0004076!, designated by Sodré et al. 2017; isolectotypes: G 00434616!, F
24501!).
Croton decarianus Pilger ex Glaziou (1913: 620), nom. nud.
Distribution and habitat:—Croton abaitensis was reported for the Distrito Federal and states of Goiás, Mato Grosso
and Minas Gerais by Sodré et al. (2017) and it is also reported here for the states of Mato Grosso do Sul and Tocantins
(Fig. 14a). It grows in campo sujo, campo limpo and in cerrado s.str. between 645 and 1,280 m elevation.

FIGURE 12. Croton abaitensis a habit, detail of acropetiolar glands in upper right corner, b thyrse showing pistillate flowers, buds and
staminate flowers, c thyrse showing pistillate flowers, d thyrse showing staminate flowers, e capsules. Croton aberrans f habit, detail of
stipule in lower right corner, g staminate flowers, h pistillate flowers and staminate buds, i capsules.
FIGURE 13. a–e Croton abaitensis a leaf, b basilaminar glands, c marginal gland, d staminate flower, e pistillate flower; f–j C. aberrans
f leaf, g acropetiolar glands, h marginal gland, i staminate flower, j pistillate flower; k–o C. adamantinus k leaf, l basilaminar glands, m
marginal gland, n staminate flower, o pistillate flower; p–t C. adenodontus p leaf, q basilaminar glands, r marginal gland, s staminate
flower, t pistillate flower; u–y C. agoensis u leaf, v basilaminar glands, w marginal gland, x staminate flower, y pistillate flower. from a–e
drawn from M. J. Silva 3286, f– from G. Hatschbach 47331, j from J. C. Lindeman s.n. (ICN 8799), k–o from V. J. Santos 267, p–t from
R. C. Sodré 3145, u–y from W. R. Anderson 9039.

FIGURE 14. Geographical distribution of some species of Croton sect. Geiseleria in Brazil. a Croton abaitensis, C. aberrans, C.
adamantinus, C. adenodontus and C. agoensis; b C. antisyphiliticus, C. arenosus, C. asperrimus, C. bidentatus and C. campinarensis.
Phenology:—Collected in flower from September to February and in fruit from January to March.
Preliminary conservation status:—Croton abaitensis is widely distributed (Extent of Occurrence estimated at
roughly 850,000 km2), its populations are abundant, and many are found in protected areas, so we categorize it as Least
Concern.
Relationships:—Croton abaitensis resembles C. agoensis, C. mucronifolius (Müller Argoviensis (1873: 216),
and C. pycnadenius by its actinodromous-brochidodromous pattern venation, with a pair of basal veins exceeding
half the length of the blade; short petioles (0.3–0.8 cm long) with shortly stipitate glands at the apex, and linear bracts
2–6.3 mm long. However, C. abaitensis differs from the other three by having leaf margins with patelliform glands (vs.
stipitate-capitate in C. agoensis and C. mucronifolius, and glands absent in C. pycnadenius), villous staminal filaments
(vs. glabrous in all others), staminate petals villous on both surfaces (vs. glabrous in all others); obovate, ovate, or
orbicular, unequal and eglandular sepals in pistillate flowers (vs. linear-lanceolate or lanceolate, equal and usually
glandular sepals in all others); and stellate stigmas (vs. glabrous in all others).
Representative specimens examined:—BRAZIL. Distrito Federal: Brasília, Reserva Ecológica do IBGE, área do
córrego Taquara, 15°55’55”S, 47°53’51”W, 1015 m, 28 October 1999, M.L. Fonseca & D. Alvarenga 2184 (IBGE, SP).
Goiás: Campo Alegre de Goiás, BR-050, próximo ao rio Imburuçu, 29 November 1992, G. Hatschbach & E. Barbosa
58277 (MBM, SP). Mato Grosso: Cuiabá, Parque Nacional da Chapada dos Guimarães, próximo ao estacionamento
da cidade de Pedra, 26 March 2016, R.C. Sodré et al. 2242 (UFG). Mato Grosso do Sul: Corumbá, trilha do Pelego,
morro atrás do Porto do Amolar, 18°1’17.9”S, 57°31’53.8”W, 10 February 2002, G.A. Damasceno Júnior et al. 2586
(COR). Minas Gerais: Uberlândia, saída pela BR 452, sentido Belo Horizonte, 18°55’22.9”S, 48°12’10.8”W, 940 m,
06 July 2016, R.C. Sodré et al. 2520 (UFG). Tocantins: Dianópolis, Serra Geral do Tocantins, região das nascentes do
rio Palmeiras, ca. 5 km após a ponte sobre o rio Limoeiro, 11°36’24.1”S 46°28’23.2”W, 708 m, 07 May 2017, R.C.
Sodré & U.S. Amaral 3200 (UFG).
2. Croton aberrans Müller Argoviensis (1873: 232).—Oxydectes aberrans (Müll.Arg.) Kuntze (1891: 613). Type:—
BRAZIL. Mato Grosso do Sul: “in campis arenosis secus Rio Pardo”, September 1826, L. Riedel 546 (lectotype: LE
00003045!, designated here; isolectotype: G 00434413) (Figs. 12f–i, 13f–j).
Croton guaraniticus Chodat & Hassler (1905: 496). Type:—PARAGUAY. Cordillera: “Cordillerae centralis; in region collium: Cerros
de Tobaty”, September 1900, E. Hassler 6314 (lectotype: G 00307062!, designated here; isolectotypes: A 00047309!, B† [photo
at F 0BN005109!], F 0056134F, G, K 000574182!, MPU 014831!, NY 00246548!, P 00623669!, P 00623670!, S 07-12802, W);
remaining syntype:—PARAGUAY. “pr. Tacuaral” September 1885, E. Hassler 1083 (not located).
Croton guaraniticus f. latifolius Chodat & Hassler (1905: 497), syn. nov. Type:—PARAGUAY. Canindeyú: “in campo Nandurucay Sierra
de Maracayu”, E. Hassler 4529 (lectotype: P 00623671!, designated here; isolectotype: NY 00246549!).
Croton guaraniticus var. virgatus Chodat & Hassler (1905: 497), syn. nov. Type:—PARAGUAY. Canindeyú: “Iter ad Yerbales montium
Sierra de Maracayu, in regione vicine Igatimi”, December 1898–1899, E. Hassler 5604 (lectotype: G 00306908!, designated here;
isolectotypes: G 00306910!, G 00306930!, G 00306931!, GH 00047310!, MPU 014829!, P 00634499!); remaining syntype:—
PARAGUAY. Concepción: “in campo in regione cursus superioris fluminis Apa”, December 1902, E. Hassler 7993a (G 00306906!,
G 00306907!, NY 00246550!).
Croton guaraniticus f. intermedia Chodat & Hassler (1905: 497), syn. nov. Type:—PARAGUAY. Concepción: “in campis siccis pr.
Bellavista in regione cursus superioris fluminis Apa”, November 1901–1902, E. Hassler 7933 (lectotype: G 00306934!, designated
here; isolectotypes: G 00306912!, G 00306913!, G 00306914!, G 00306935!, MPU 014830!, P 00623672!, P00623673).
Croton guaraniticus f. microphylla Chodat & Hassler (1905: 497), syn. nov. Type:—PARAGUAY. Concepción: “In regione cursus superioris
fluminis Apa”, November 1901–1902, E. Hassler 7933b (lectotype: G 00306911!, designated here; isolectotypes: G00306932!, G
00306933!).
Distribution and habitat:—Croton aberrans occurs in Argentina (Misiones), Bolivia (Santa Cruz), Brazil (Mato
Grosso do Sul, Paraná, Rio Grande do Sul, Santa Catarina, and São Paulo [Fig. 14a]), and Paraguay (Amambay,
Canindeyú). It is usually found in campos and cerrado s.str. between 345 and 1,388 m elevation.
Phenology:—Collected with flowers from September to February and with fruits from October to April.
Preliminary conservation status:—Since this species is widespread and occurs in an area greater than 500,000
km2, it was categorized as Least Concern.
Relationships:—Croton aberrans is most often confused with C. hirtus and was treated as a synonym of that
species by Caruzo & Cordeiro (2007), probably due to its branches and leaves with hirsute trichomes and petioles with
cylindric glands. However, C. aberrans has densely congested inflorescences 0.9–1.4 cm long, bracts with sessile
glands, pistillate flowers with 6–8(9) subequal, oblong or lanceolate sepals, and large capsules and seeds with 6.5–7
mm and 4.9–5.7 × 3.3–3.7 mm, respectively. In C. hirtus the inflorescences are more elongate inflorescences 2–6
cm long, the bracts have stipitate glands, the pistillate flowers are 5-merous with unequal sepals, the largest obovate,
oblanceolate or spatulate, and the capsules and seeds are 4.5–5 mm long and 3.6–3.7 × 2.5–2.7 mm, respectively.
Taxonomic notes:—We lectotypified Croton aberrans based on a collection at LE, as it was the most complete
collection and was also annotated with Müller’s handwriting. Although Caruzo & Cordeiro (2007) treated the specimen
at G as the holotype, Müller Argoviensis (1873) did not indicate any herbarium for the type, which led us to designate
a lectotype. We also updated the type locality of Croton aberrans, interpreted for the province of São Paulo by Müller
Argoviensis (1873), but which actually corresponds to the state of Mato Grosso do Sul, according to Riedel’s itinerary
in Flora Brasiliensis (Urban 1906).
Regarding the lectotype of Croton guaraniticus, we chose E. Hassler 6314 at G (barcode 00307062), because it
contains illustrations made by Chodat, which were likely used to draft the original description for this name. For the
three forms of C. guaraniticus var. virgatus, we also chose collections from G, because these were the most complete
and best represented description; while for C. guaraniticus var. guaraniticus f. latifolius, was chosen a collection at P
(none were found so far at G). Croton guaraniticus was synonimized under C. aberrans by Peña-Chocarro et al. (2010),
and we agree with that because they exhibit continuous and overlapping characters. Govaerts (2011) recognized the
forms of C. guaraniticus as synonyms of C. hirtus, but that species is easily differentiated from C. aberrans. Thus we
propose here the synonymization of all other infraspecific categories of C. guaraniticus under C. aberrans.
Representative specimens examined:—ARGENTINA. Misiones: Posadas, prope “La Granja” Prodium, 10
November 1907, E.L. Ekman 452 (LD). Bolivia, Santa Cruz: Vallegrande, Entre Las Vueltas y Corosito, 18°52’10”S
63°57’27”W, 1388 m, 28–29 November 2011, G.A. Parada et al. 3856 (MO). BRAZIL. Mato Grosso do Sul: Bodoquena,
ca. 10 km Serra de Bodoquena, 4 km a direita da MS-178, sentido Bonito, em estrada acessada próximo a mineradora
Mont Serrat, 20°37’47.1”S 56°42’26.4”W, 345 m, 22 September 2016, R. C. Sodré & R.A. Pereira-Silva 2636 (UFG);
ibid., 17 April 2017, R.C. Sodré & L.L.C. Antunes 3176 (UFG). Paraná: Guarapuava, Fazenda Três Capões, 10 January
1984, G. Hatschbach 47331 (C, MBM). Rio Grande do Sul: Passo Fundo, 20 km E de Passo Fundo, 30 October 1971,
J.C. Lindeman et al. s.n. (ICN 8799). Santa Catarina: Abelardo Luz, 11 km North of Abelardo Luz, 25 December 1956,
L.B. Smith & R. Reitz 9237 (HBR). São Paulo: Botucatu, 18 km N of Botucatu (14 km east of São Manuel) along the
São Manuel–Piracicaba highway, near ex-RR station 13 de Maio, 30 November 1972, I.S. Gottsberger 2101 (UB).
PARAGUAY. Amambay: Pedro Juan Caballero, 10 km divisa com Brazil, na altura do km 16, 22°33’S, 55°46’W, 18
November 1963, J.C. Gomes Jr. 1449 (SP, UB).
3. Croton adamantinus Müller Argoviensis (1873: 115).—Oxydectes adamantina (Müll.Arg.) Kuntze (1891: 613).
Type:—BRAZIL. Minas Gerais: “Prope Jiquitinhonha in distr. Adamantium”, December 1824, L. Riedel 1252
(lectotype: LE 00003049!, designated here; isolectotypes: G 00434411!, K 000185988!, LE 00003046!, LE 00003047!,
LE 00003048!, LE 00003051!, LE 00003052!) (Figs. 13k–o, 15a–e).
Vernacular name:—canela-de-urubu, velame-bravo.

Distribution and habitat:—This species is endemic to Brazil, in Bahia, Ceará, Minas Gerais, Paraíba, Pernambuco,
Piauí, Rio Grande do Norte, and Sergipe (Fig. 14a). It grows in Caatinga vegetation on sandy or clayey soils, between
150 and 1,000 m elevation (Carneiro-Torres 2009, Alves 2012).
Phenology:—Collected with flowers and fruits throughout the year.
Preliminary conservation status:—This species has an Extent of Occurrence estimated at 750,000 km2 and,
therefore, considered as Least Concern.
Relationships:—Croton adamantinus can be confused with C. virgultosus Müller Argoviensis (1873: 104)
because of its shrubby habit 1.5–4 m tall, stipitate acropetiolar or marginal glands, ovate or oval-lanceolate leaf blades
with actinodromous-brochidodromous venation, androecium with 10 or 11 stamens, and 2-fid stigmas. However, C.
adamantinus differs from C. virgultosus by its thyrses 1–4 cm long with 3–6 pistillate flowers (vs. 6–11 cm long, with
8–12 pistillate flowers in C. virgultosus), pistillate flowers with equal, ovate sepals (vs. unequal, oblong to lanceolate),
glanduliform petals (vs. filiform) and gynoecium 4 to 6 times longer than the pedicels (equal to or up to twice as long
as the pedicel).
Taxonomic notes:—Croton adamantinus was described based on L. Riedel 1252 from the state of Minas Gerais,
Diamantina region, and six of the eight mounted sheets of this collection are deposited in LE. Our choice for the sheet
LE 00003049 was due to the fact that a short description of the species was written on it with Müller’s handwriting.
FIGURE 15. Croton adamantinus a habit, b Flowering branch, detail of the acropeciolar glands in the lower right corner, c thyrse showing
staminate flowers visited by a bee, d pistillate flowers, e capsules. C. adenodontus f habit, detail of the acropetiolar gland in insert, g
flowering branch, with detail of the staminate flowers in the insert, h pistillate flowers, i thyrse showing capsules and a staminate flower.

Representative specimens examined:—BRAZIL. Bahia: Caetité, 8 km NW de Caetite na BR-430 rumo a Bom
Jesus da Lapa, 14°0’S, 42°35’W, 910 m, 09 November 1984, A.C. Allem et al. 2906 (CEN). Ceará, Aracati, BR-304,
próximo ao km 61, córrego de Ubarana, 11 March 1979, E. Nunes et al. s.n. (UEC 0005707, HUEFS 000083381);
Barbalha, estrada de Seu Adão, 7°18’40”S, 39°18’15”W, 914 m, 02 February 2012, C.G. Crepaldi 33 (PEUFR). Minas
Gerais: Januária, s.loc., 04 November 1978, L. Krieger s.n. (CESJ 16085, HUEFS 000084263). Pernambuco: Sertânia,
BR-110, estrada Moderna/Ibimirim, 8°30’23”S, 37°35’2”W, 495 m, 29 June 2007, D.S. Carneiro Torres et al. 964
(HUEFS). Piauí: São João do Piauí, saída para Nova Santa Rita, 8°17’0”S, 42°10’32”W, 329 m, 7 February 2007,
D.S. Carneiro-Torres et al. 805 (HUEFS). Rio Grande do Norte: BR 101, litoral Norte indo p/ Oeste, 19 km depois da
entrada para Pitangui, 5°18’13”S, 35°11’29”W, 05 August 2008, M. Andrade Neto s.n. (EAC 0044186).
4. Croton adenodontus (Müll.Arg.) Müller Argoviensis (1873: 267).—Croton glandulosus var. adenodontus Müller
Argoviensis (1866: 684).—Oxydectes adenodonta (Müll.Arg.) Kuntze (1891: 613). Type:—BRAZIL. Rio de Janeiro:
no further locality, 5 April 1831, L. Riedel 65 (lectotype: LE 00003057!, designated here; isolectotypes: A 00257929!,
GH 00257928!, K 000254391!, LE 00003055!, LE 00003056!, LE 00018229!, LE 00018230!, NY 0008435!, P
00623667!); remaining syntypes:—BRAZIL. Rio de Janeiro: no further locality, 1834, P.W. Lund 130 (G 00312857!);
BRAZIL. Rio de Janeiro: no further locality, s.d., J. Lhotsky 55 (G 00312856!) (Figs. 13p–t, 15f–i).
Croton fluminensis (Kuntze) Schumann (1900: 348).—Oxydectes fluminensis Kuntze (1898: 288), syn. nov. Type:—BRAZIL. Rio de
Janeiro: no further locality, December 1892, O. Kuntze s.n. (lectotype: NY 00503940!, designated here; isolectotypes: B† [photo F
0BN005096!], K 000574180!).
Distribution and habitat:—Croton adenodontus was described based on collections from the state of Rio de Janeiro
and is reported here for the first time for Espírito Santo and Mato Grosso do Sul (Fig. 14a). The species grows in
restingas in Espírito Santo and Rio de Janeiro and in ferruginous rock outcrops, locally known as “cangas” in Mato
Grosso do Sul, between 30 and 207 m elevation. Although C. adenodontus was cited for Bahia, Ceará, and Tocantins
by BFG (2015), we verified that the specimens of these states identified as C. adenodontus correspond to other species
of section Geiseleria, such as C. agoensis e C. virgultosus.
Phenology:—Collected with flowers and fruits in April, June and August, and only flowers in September and
December.
Preliminary conservation status:—We consider the conservation of this species as Deficient Data, because it is
known from only three locations, one of which is more than 1200 km away from the other two and, therefore, more
field investigations should be conducted to determine their distribution and abundance.
Relationships:—Croton adenodontus is morphologically most similar to C. glandulosus, but differs from it by
the serrulate leaf margin with stipitate glands in the sinuses (vs. serrate, crenate or serrulate and usually without glands
in the sinuses in C. glandulosus); pistillate flowers with unequal sepals, the two largest elliptic or obovate (vs. equal
or unequal, obovate, oblanceolate or spatulate); pubescent stigmas (vs. glabrescent); small capsules (3.3–3.5 mm long)
without distinct color on the sutures of the septa and locules (vs. 4–5 mm long, with lines whitish on the sutures of the
septa and locules); and seeds usually ecarunculate (vs. carunculate).
Taxonomic notes:—Croton glandulosus var. adenodontus was described based on L. Riedel 65, P.W. Lund 130 and
J. Lhotsky 55, all from the Rio de Janeiro state, and when it was elevated to species level by Müller Argoviensis (1873),
he added G. Gardner 113, also from Rio de Janeiro. The latter, however, is not original material for C. glandulosus var.
adenodontus. Among the three collections, L. Riedel 65 is the most representative, especially in LE, where 5 mounted
sheets were found, one of which contains Mueller’s own annotation of C. adenodontus, and this one was chosen as a
lectotype.
Representative specimens examined:—BRAZIL. Espírito Santo: Guarapari, Parque Natural Municipal Morro
da Pescaria, 20°39’29.2”S 40°28’27.5”W, 30 m, 8 September 2013, A.C.S. Dal col & J. Rodrigues Filho 131 (VIES).
Mato Grosso do Sul: Aquidauana, Piraputanga, estrada de terra CG.–Aquidauana, 16 December 1990, U.M. Resende
et al. 308 (CGMS, SP); a direita da estrada Parque de Piraputanga (MS-450), ca. 7 km do distrito de Piraputanga em
direção ao distrito de Palmeiras, 20°27’17.8”S, 55°29’1.8”W, 207 m, 15 April 2017, R.C. Sodré & L.L.C. Antunes 3145
(UFG), 3146 (UFG), 3147 (UFG), 3148 (UFG), 3149 (UFG), 3150 (UFG), 3151 (UFG), 3152 (UFG), 3153 (UFG),
3154 (UFG), 3155 (UFG). Rio de Janeiro: s.loc., August 1836, G. Gardner 113 (K, NY, US); Pedra das Cabritas, 9 June
1887, C.A.W. Schwacke 5607 (RB).
5. Croton agoensis Baillon (1864: 348).—Oxydectes agoensis (Baill.) Kuntze (1891: 610). Type:—BRAZIL. Minas
Gerais: “Brazilia Meridionalis, prope Olho d’Agua”, 1816–1821, A.F.C.P. Saint-Hilaire s.n. (lectotype: P 00623665!,
designated here; isolectotypes: A 00047218!, P 00623664!) (Figs. 13u–y, 16a–d).
Vernacular name:—alcachofrinho, algodãozinho, malvinha, pustemeira, velame.

Distribution and habitat:—Known only from Brazil, occurring in Bahia, Goiás, Maranhão, Minas Gerais, Piauí,
and Tocantins states (Fig. 14a). It inhabits cerrado tipico, cerrado denso or cerradão, usually in sandy soils, between
224 and 1340 m elevation.
Phenology:—Flowering and fruiting throughout the year.
Preliminary conservation status:—Croton agoensis is widely distributed (Extent of Occurrence estimated at
roughly 500,000 km2) and, therefore, it is designated here as Least Concern.
Relationships:—This species is morphologically similar to C. mucronifolius and C. pycnadenius, especially in its
flowers and fruits. However, it can be distinguished by its membranaceous leaves (vs. chartaceous in C. mucronifolius
and C. pycnadenius), with serrate or serrulate margin with stipitate-capitate glands (vs. crenate or serrate and usually
without glands), as well as by the flowers of both sexes with the calyx sparsely pubescent externally (vs. densely
tomentose).
Taxonomic notes:—Baillon (1864) designated A.F.C.P. Saint-Hilaire s.n. from Minas Gerais as the type of C.
agoensis; this collection has two duplicates with different barcodes at P and one in A, the latter donated by P. We
designate P 00623665 as lectotype of C. agoensis because it is the most complete and most representative of the
original description of the species. This collection was treated as the holotype of the species by Sodré et al. (2017).
Representative specimens examined:—BRAZIL. Bahia: Correntina, Rio Piau, ca. 225 km SW of Barreiras
on road to Posse, 850 m, 12 April 1966, H.S. Irwin et al. 14596 (C). Goiás: Posse, área do Aeroporto, 14°6’58”S,
46°20’48”W, 840 m, 16 October 2001, M.L. Fonseca et al. 2900 (IBGE); Maranhão: Pastos Bons, 31.6 km west along
main road from turnoff to Pastos Bons, 27 January 1970, G. Eiten & L.T. Eiten 10381 (SP). Minas Gerais: Januária, 7.5
km by road W of Rio Pandeiros on road from Januária to Serra das Araras, 600 m alt., 18 April 1994, W.R. Anderson et
al. 9039 (F, INPA). Piauí: Ribeiro Gonçalves, Estação Ecológica Uruçuí-Una, 24 July 1983, A. Fernandes s.n. (EAC
12149). Tocantins: Lizarda, Bacia do Tocantins, 9°16’48”S, 47°0’36”W, 351 m, 21 November 2008, C.W. Fagg &
F.C.A. Oliveira 2062 (IBGE, SP).
6. Croton antisyphiliticus Martius (in Spix & Martius 1823: 282).—Croton antisyphiliticus var. genuinus Müll.Arg.,
Prodr. 15: 593. 1866, nom. inval.—Oxydectes antisyphilitica (Mart.) Kuntze (1891: 611). Type:—BRAZIL. São Paulo:
“in campis editis e.g. ad Ypanema, ubi Erva Mular”, 18 January 1818, C.F.P. Martius s.n. (holotype: M 0086128, photo
in F 19474!) (Figs. 16e–i, 17a–e).
Croton perdicipes Saint-Hilaire (1827: tab. 59).—Croton antisyphiliticus var. perdicipes (A. St.-Hil.) Müller Argoviensis (1865: 110).—
Croton perdicipes var. genuinus Baillon (1864: 337), nom. inval. Type:—BRAZIL. Minas Gerais: “campos au près de Tanque, comarca
de Rio das Mortes”, 1816–1821, A.F.C.P. Saint-Hilaire cat. C1, n. 227 (lectotype: P 00623643!, designated here; isolectotypes: MPU
014855!, P 00623644).
Croton antisyphiliticus var. cordiifolius Müller Argoviensis (1865: 110).—Ocalia cordiifolia Klotzsch (1841a: 195), nom. nud.—Ocalia
grandifolia Klotzsch (1841a: 195), nom. nud. Type:—BRAZIL. s.loc., s.d., F. Sellow s.n. (lectotype: BR 0000008763891!, designated
here; isolectotype: K 000254362!, P05566748!).
Croton antisyphiliticus var. latifolius (Baill.) Müller Argoviensis (1866: 593).—Croton perdicipes var. latifolius Baillon (1864: 337).
Type:—BRAZIL. Minas Gerais: s.loc., s.d., A.F.C.P. Saint-Hilaire cat. C2, n. 6118 (lectotype: P 05566736!, designated here);
remaining syntypes:—BRAZIL. s.loc., s.d., A.F.C.P. Saint-Hilaire cat. C2, n. 1393 bis (P 05566735!); BRAZIL. Sao Paulo: s.loc.,
s.d., A.F.C.P. Saint-Hilaire cat. C1, n. 1058 (P 05566737!)
Croton antisyphiliticus var. minor (Baill.) Müller Argoviensis (1866: 593).—Croton perdicipes var. minor Baillon (1864: 337).—Ocalia
sellowiana Klotzsch (1841a: 195), nom. nud. Type:—BRAZIL. Minas Gerais: s.loc., s.d., A.F. Regnell I 400 (lectotype: P 05566672!,
designated here); remaining syntypes:—BRAZIL. s.loc., s.d., C. Gaudichaud-Beaupré 953 (P 05566676!); s.loc., s.d., F. Sellow s.n.
(K 000641027!, P 05566746!); BRAZIL. Minas Gerais: s.loc., 1838, P. Claussen 769 (P 05566675!); BRAZIL. s.loc., 1858, H.A.
Weddell 1086 (G 00311951!, P 05566742!).
Croton antisyphiliticus var. mollis Müller Argoviensis (1866: 593). Type:—BRAZIL. Minas Gerais: “in editis campis S. de Grao Magor
et versus Contagem de S. Maria in confin. Goyaz”, s.d., C.F.P. Martius s.n. (holotype: B†, lectotype: M 0086130!, designated here;
isolectotype: M 0086129!).
Croton antisyphiliticus var. echiifolius Müller Argoviensis (1865: 110).—Ocalia echiifolia Klotzsch (1841a: 195), nom. nud. Syntypes:—
BRAZIL. s.loc., 1834, P.W. Lund s.n. (G 00311949!); s.loc., s.d., F. Sellow s.n. (not found); Minas Gerais: s.loc., s.d., A.F. Regnell
s.n. (not found).

Croton antisyphiliticus var. heterotrichus Müller Argoviensis (1873: 208). Type:—BRAZIL. Minas Gerais: “Villa de Uberaba”, October
1848, A.F. Regnell III 1086 (BR 0000008763235!, designated here); remaining syntypes:—BRAZIL. Minas Gerais: “ad Barbacena”,
September 1862, L.S.M. Netto s.n. (syntype: BR 0000008764553!); “prope Barbacena”, s.d., L. Riedel (not found); BRAZIL. s.loc.,
s.d., Sellow s.n. (not found).
Croton antisyphiliticus var. hypoleucus Müller Argoviensis (1873: 207). Type:—BRAZIL. Minas Gerais: “ad S. Luzia in glareosis, et
prope Barbacena”, October 1824, L. Riedel s.n. (lectotype: LE 00018215!, designated here; isolectotype: LE 00018214!).
Croton antisyphiliticus var. intermedius Müller Argoviensis (1873: 208). Type:—BRAZIL. Minas Gerais: “prope Barbacena”, s.d., J.B.E.
Pohl 1632 (catalogue number 3917) (lectotype: W 0060075!, designated here; isolectotype: K 000254361!); remaining syntypes:—
BRAZIL. Minas Gerais: “ad Lagoa Santa”, 17 July 1863, J.E.B. Warming 1639 (F V0093333F!, LE!); “prope Barbacena”, s.d., L.
Riedel (not found).
Croton antisyphiliticus var. minutulus Müller Argoviensis (1873: 209). Type:—BRAZIL. Minas Gerais: “inter Tejuco seu Cidade
Diamantina et Rio Paranahyba”, s.d., W. Burchell 5891 (holotype: BR 0000008763563!).
Croton antisyphiliticus var. nitidulus Müller Argoviensis (1873: 209). Type:—BRAZIL. Minas Gerais: “ad Congonhas do Campo”, 1844,
M.D. Stephan s.n. (lectotype: BR 0000008762573!, designated here); remaining syntypes:—BRAZIL. “Brazilia occidentali”, s.d.,
Tamberlik s.n. (not found); BRAZIL. Minas Gerais: “ad Lagoa Santa”, s.d., J.E.B. Warming s.n. (not found); BRAZIL. Minas Gerais:
“ad Barbacena”, s.d., L. Riedel 2808 (LE 00018216!).
Vernacular name:—alcanforeira, alcânfora, canela-de-perdiz, cocaleira, curraleira, erva-mular, minuano, pé-de-

perdiz, pé-de-perdiz-macho, pé-de-perdiz-peluda.
Distribution and habitat:—This is a relatively widely distributed species known from Bolivia (Møller-Jørgensen
et al. 2014), Brazil, and Paraguay (Caruzo & Cordeiro 2007). In Brazil, we confirmed its occurrence in the Distrito
Federal and in the states of Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Santa Catarina, and São
Paulo (Fig. 14b). Although Caruzo & Cordeiro (2007) and Carneiro-Torres (2009) reported C. antisyphiliticus from
other Brazilian states, we verified that the specimens indentified by these authors as this species correspond instead
to C. gracilescens, C. agoensis, or C. parodianus. Carneiro-Torres (2009) also reported C. antisyphiliticus for Central
American countries (e.g., Mexico, Panama, Nicaragua, Guatemala, and Honduras), but after examining material from
this region in numerous herbaria, we could not verify its presence there. It grows in cerrado s.str. and campos sujos in
clay or stony soils between 410 and 1250 m elevation.
Phenology:—Recorded flowering from August to December and fruiting from September to January.
Preliminary conservation status:—Croton antisyphiliticus can be considered Least Concern because of its wide
distribution and frequency.
Relationships:—Croton antisyphiliticus is morphologically similar to C. gracilescens, a species which has been
considered one of its synonyms by Govaerts et al. (2000) and van Ee (2011). However, Croton gracilescens can
be distinguished by its stellate-rotate trichomes on the branches and leaves (vs. stellate-porrect or fasciculate in C.
antisyphiliticus), petioles 0.1–0.4 cm long (vs. 0.5–2 cm long) leaf blades linear or narrowly elliptic (vs. usually oblong
or elliptic), ovary with pubescent indument of stellate trichomes (vs. hirsute indument of stellate-porrect trichomes),
and stigmas always 2-fid (vs. 4-fid). Additionally, C. gracilescens is generally of larger stature, up to 1.3 m tall, and
its indument is sparse and inconspicuous in vegetative and reproductive organs, whereas C. antisyphiliticus is a small
subshrub up to 0.5 m tall, usually densely pubescent.
Taxonomic notes:—Govaerts et. al. (2000) synonymized all varieties of C. antisyphiliticus under this species, a
position corroborated by van Ee (2011). However, we do not agree with the synonymization of C. antisyphiliticus var.
angustifolius and C. antisyphiliticus var. weddellianus under C. antisyphiliticus, varieties that are here synonimized
under C. gracilescens. We assigned lectotypes for all varieties that were described based on two or more collections,
or on a single collection, but with duplicates in the same herbarium under different barcodes, or in different herbaria,
always prioritizing the specimens that best reflected the original description or that contained some annotation of the
author of the species, as was the case of the lectotype chosen for C. antisyphiliticus var. hypoleucus.
Representative specimens examined:—BOLIVIA. Depto. Santa Cruz: Chiquitos, Santiago de Chiquitos, al este
de la zona del arco de piedra, subiendo hacia la cumbre de la meseta, 18°20’30”S, 59°33’4”W, 880 m, 24 October 2010,
J.R.I. Wood & D. Villarroel 26937 (USZ, UB). BRAZIL, Distrito Federal: Brasília, Jardim Botânico de Brasília, ca.
20 km a SE de Brasília, 29 August 1985, Equipe do Jardim Botânico de Brasília 560 (HEPH); Reserva Ecológica do
IBGE, 15°56’S, 47°53’W, 1100 m, 13 September 2000, M. Aparecida da Silva 4497 (IBGE); Campus da Universidade
de Brasília, Centro Olímpico, 15°46’1”S, 47°51’16”W, 1012 m, 14 September 2006, M.Q. Matos et al. 27 (UB, ESA).
Goiás: Anápolis, Campus UEG, 28 September 2004, J.E.Q. Faria Júnior 42 (HUEG); Cocalzinho, Parque Estadual
da Serra dos Pireneus, após a segunda porteira, saindo da fazenda Capitão do Mato, sentido Morro do Cabeludo,
FIGURE 16. Croton agoensis a habit, with detail of the basilaminar glands in the lower right corner and of the marginal glands in the
upper left corner, b thyrses showing pistillate flowers and staminate buds and flowers, with detail of two staminate flowers in the insert, c
pistillate flower, d pistillate flower and capsule. C. antisyphiliticus e habit, f thyrses with pistillate flowers and staminate buds, g staminate
flowers, h pistillate flowers, i a pair of capsules.

FIGURE 17. a–e Croton antisyphiliticus a leaf, b acropetiolar glands, c marginal gland, d staminate flower, e pistillate flowers; f–j C.
asperrimus f leaf, g acropetiolar glands, h marginal gland, i staminate flower, j pistillate flower; k–n C. bidentatus k leaf, l acropetiolar
glands, m staminate flower, n pistillate flower; o–s C. campinarensis o leaf, p basilaminar glands, q marginal gland, r staminate flower, s
pistillate flower; t–x C. catariae t leaf, u basilaminar glands, v marginal glands, w staminate flower, x pistillate flower. Drawn from a–e
R. C. Sodré 1449, f–j R. C. Sodré 3110, k–n L. Kollmann 1432, o–s D. C. Daly 982, t–x R. C. Sodré 1205.
15°48’0”S, 48°48’0”W, 1100 m, 24 October 2004, P.G. Delprete et al. 8957 (UFG); Goiás, Distrito de Jeroaquara,
Serra de Santa Rita, 23 October 1971, J.A. Rizzo & A. Barbosa 7162-A (UFG). Mato Grosso: Cuiabá, Mirante Centro
Geodésico da América do Sul, ca. 3 km antes de chegar no município de Chapada dos Guimarães, 15°28’48”S,
55°41’20”W, 25 March 2016, R.C. Sodré et al. 2222 (UFG); Nova Brazilândia, estrada Nova Brazilândia–Planalto
da Serra, ca. 20 km NE (em linha reta) de Nova Brazilândia, 14°52’S, 54°49’W, 10 October 1997, V.C. Souza et al.
20704 (ESA, UFMT). Mato Grosso do Sul: Costa Rica, Fazenda Fortuna, 18°24’4.4”S, 52°59’32.7”W, 29 October
2004, A.S. Penha et al. 197 (CGMS); Ponta Porã, próximo ao Posto Fiscal, entre a rodovia MS e a ferrovia, 5 km W de
Ponta Porã, 22°21’50.1”S, 55°43’57.5”W, 680 m, 26 October 2004, A. Pott & V.J. Pott 12093 (CGMS). Minas Gerais:
Joaquim Felício, Serra do Cabral, 20 September 2005, G. Hatschbach & E. Barbosa 79358 (HUCS); Perdizes, EPDA-
Galheiros, Céu do cavalo, 23 November 2002, E.H. Amorim et al. 321 (HUFU, SP). Paraná: Palmeira, Rio Capivara,
20 October 1989, V. Nicolack & J. Cordeiro 70 (FLOR); Ponta Grossa, Passo do Pupo, 5 October 1995, J.M. Silva &
J. Cordeiro 1564 (BHCB, ESA); Buraco do Padre, 02 November 2007, M.G. Caxambu et al. 1739 (HCF, CGMS).
Santa Catarina: Mafra, campo West of Mafra on the road to Barracas, 7 December 1956, L.B. Smith & R. Klein 8438
(NY). São Paulo: Bom Sucesso de Itararé, fazenda Santa Maria do Espinho e Saco Grande, 24°16’14”S, 49°15’35”W,
16 December 1997, S.I. Elias et al. 222 (ESA); Itapeva, estrada de terra de Taquarituba–Itapeva, 29 September 1994,
J.Y. Tamashiro et al. 741 (ESA); Itararé, estrada para 400 alqueires, 24 September 1989, C.A.M. Scaramuzza & V.C.
Souza 525 (ESA); Pedregulho, Estreito, perto do rio Grande, 06 November 1997, W.M. Ferreira et al. 1573 (ESA).
PARAGUAY. Pedro Juan Cabalero, 5 km O, 16 October 1984, G. Hatschbach & R. Kummrow 48505 (MBM, F).
7. Croton arenosus Carn.-Torres & Cordeiro (in Carneiro-Torres et al. 2011: 123). Type:—BRAZIL. Bahia: Reserva
Biológica Esquentada, 9º56’58”S, 38º58’48”W, 440 m, 18 February 2004, E.B. Miranda, R.M. Harley, F.H. Silva,
B.M. da Silva & Carlos 613 (holotype: HUEFS 000002805!).
Vernacular name:—caatinga-de-cheiro.
Distribution and habitat:—This species is only known from the Canudos Biological Station in the northeast
portion of Bahia state (Fig. 14b), where it grows in sandy caatingas, between 400 and 450 m elevation (Carneiro-Torres
et al. 2011).
Phenology:—Collected with flowers in January, February, March, June and September and with fruits in March
and June.
Preliminary conservation status:—Carneiro-Torres et al. (2011) categorized the conservation status of Croton
arenosus as Critically Endangered, because it has been collected in just one location and its geographic range is smaller
than 100 km2.
Relationships:—Authough the authors of C. arenosus compared the species with C. triangularis, we compare it
more closely to C. harleyi Carneiro-Torres et al. (2011: 128) by the shape and size of its leaves and the morphology of
the pistillate flowers. Even so, C. arenosus differs from C. harleyi by the chartaceous leaf blades with shortly stipitate
patelliform glands at the base; inflorescences 2–4.5 cm long with 5–10 pistillate flowers; bracts 0.5–1 mm long; and
pistillate flowers with oblong or lanceolate sepals with plane margins. In C. harleyi the leaves are membranaceous with
cylindric glands at the base; inflorescences up to 2 cm long with 2 to 4 pistillate flowers, the bracts 3–4 mm long, and
the pistillate flowers with ovate sepals with revolute margins.
Representative specimens examined:—BRAZIL. Bahia: Canudos, Reserva Biológica de Canudos, 9°56’43”S,
39°0’56”W, 434 m, 30 June 2007, D.S. Carneiro-Torres et al. 993 (HUEFS, SP).
8. Croton asperrimus Bentham (1854: 376).—Oxydectes asperrima (Benth.) Kuntze (1891: 611). Type:—BRAZIL.
Pará: “in vicinibus Obidos”, November–December 1849, R. Spruce 478* (lectotype: K 001181152!, designated here;
isolectotypes: K 001181155!, P 00623631!) (Figs 17f–j, 18a–d).
Distribution and habitat:—Croton asperrimus is endemic to the Amazon Forest biome, described from Pará, then
cited for Maranhão by Secco (2008). Here it is also reported for Amazonas, Mato Grosso, and Rondônia (Fig. 14b). It
grows near water courses in periodically flooded areas on sandy soils between 85 and 245 m of elevation.
Phenology:—Collected with flowers and fruits from September to May.
Preliminary conservation status:—The species has an estimated Extent of Occurrence of about 1,000,000 km2
and, therefore, is classified as Least Concern.
Relationships:—Croton asperrimus can be recognized by its lanceolate leaf blades with serrulate margins and
obconic acropetiolar and marginal glands; thyrses up to 15 cm long with 5–13 pistillate flowers; bracts with 1 or 2
stipitate-patelliform glands; pistillate flowers glossy green with elliptic and equal sepals; and seeds with very small
caruncles (ca. 0.5 mm long). Even so, this species could be confused with C. sipaliwinensis, from which it differs
mainly by the petiole 10 to 15 times shorter than the leaf blades (vs. petioles 4 to 8 times shorter than the leaf blades in
C. sipaliwinensis), thyrses without a sterile zone between pistillate and staminate flowers (vs. with long sterile zone);
staminate flowers with pedicels of 1.2–2.9 mm long (vs. 0.2–0.4 mm long), 9–11 stamens (vs. 5), and stigmas 2-fid
(usually 4-fid).
Taxonomic notes:—Bentham (1854) described C. asperrimus based on R. Spruce 478*, with an asterisk only
to differentiate from R. Spruce 478, a collection of the genus Psychotria (Rubiaceae). The first collection has two
duplicates at K and one at P, and we chose the K specimen with barcode 001181152 because it was part of the former
Bentham Herbarium, author of the species, and, therefore authentic material.
Representative specimens examined:—BRAZIL. Amazonas: Itapiranga, 11 September 1903, M. Silva 1903
(MG). Mato Grosso: Alta Floresta, próximo a balsa do rio Teles Pires, ca. 37 km ao norte de Alta Floresta, 9°38’24.7”S,
56°0’54.7”W, 245 m, 04 April 2017, R.C. Sodré & A.O. Souza 3110 (UFG). Maranhão: Alzilândia, Rio Pindaré, 3°45’S,
46°5’W, 100 m, 11 December 1978, J. Jangoux & R.P. Bahia 374 (NY, RB). Pará: Prainha, 6 December 1873, J.W.H.
Traill 730 (P). Rondônia: Jaci-Paranã, margem direita do rio Madeira (Cachoeira de Jirau), 9°19’37”S, 64°43’35”W,
85 m, 20 March 2010, G. Pereira-Silva 15058 (CEN, INPA).
9. Croton bidentatus Müller Argoviensis (1865: 140).—Oxydectes bidentata (Müll.Arg.) Kuntze (1891: 611). Type:—
BRAZIL, Rio de Janeiro: s.loc., September 1822, H.K. Beyrich s.n. (holotype: B†; lectotype: P 00623619!, designated
here). (Fig. 17k–n)
Distribution and habitat:—Croton bidentatus is known from just three collections in Espírito Santo and Rio de
Janeiro (Fig. 14b). It apparently grows on rocky outcrops, from sea level to 400 m elevation.
Phenology:—Collected with flowers and fruits in the months of September and December.
Preliminary conservation status:—We categorize its conservation status as Vulnerable due to its estimated
Extent Occurrence of 10,000 km2, and because it is known from fewer than ten localities in legally unprotected areas.
Relationships:—Croton bidentatus shares with C. adenodontus the branches and leaves with ferrugineous
indument, leaf margins with stipitate glands, bracts with glands at the base, and pistillate flowers with strongly unequal
sepals, besides it being found in the same geographical area. Despite this, C. bidentatus is readily distinguished by its
stipules 4–6 mm long, petioles with two stipitate-patelliform or cylindric glands, leaf blades with a cordate base and
acuminate apex, linear or spatulate bracts 2–3 mm long, and glabrous stigmas; while in C. adenodontus the stipules
are 0.4–1.6 mm long, the petioles have obconic or shortly stipitate patelliform glands, the leaf blades have a rounded
or obtuse base and acute apex, the bracts are 0.4–1.2 mm long, and the stigmas are pubescent.
Taxonomic notes:—Müller Argoviensis (1865) described C. bidentatus based on a specimen, without collector or
number, coming from Rio de Janeiro and deposited at B, as indicated in the protologue. We could not find any specimen
corresponding to the protologue of C. bidentatus in B, which led us to assume that this material has been destroyed, like
so many others during the Second World War (Hiepko 1987). At P there is a specimen (barcode 00623619) attributed
to C. bidentatus whose label states that it came from B and indicates that Beyrich was the collector. We interpret this
as either the original or the duplicate of the holotype and so designated it here as the lectotype. However, we caution
that this specimen of C. bidentatus is more similar to C. adenodontus than M.A. zu Wied-Neuwied s.n., which was used
to complement the description and to illustrate C. bidentatus by Müller Argoviensis (1873) in Flora Brasiliensis. Thus,
we believe that is necessary to conduct future studies to consider these binomials and their types.
Representative specimens examined:—BRAZIL. Espírito Santo: Santa Teresa, Vale do Canaã, 19°54’22”S,
40°36’07”W, 400 m, 30 December 1998, L. Kollmann & E. Bausen 1432 (MBML, SP); Barra do Jacú, s.d., M.A. zu
Wied-Neuwied s.n. (BR, G, LE).
10. Croton campinarensis Secco et al. (2012: 2). Type:—BRAZIL. Pará: Acará, campina do Guajará, Alça Viária, 20
November 2009, R.S. Secco & A.S. Rosário PEA 928 (holotype: MG 205681!; isotypes: K, MICH, NY, RB, SP) (Fig.
17o–s, 18e–g).
Vernacular name:—cróton-da-campina.
Distribution and habitat:—Croton campinarensis was reported from Pará in the municipalities of Acará, Belém,
Breu Branco and Tucuruí by Secco et al. (2012) and it is newly reported for Venezuela here (Fig. 14b). It grows in
‘campinas’ or ‘campinaranas’ on sandy soils between 20 and 100 m elevation.
FIGURE 18. Croton asperrimus a habit, b thyrse showing pistillate flowers and staminate buds, c staminate flowers and buds, d capsules.
C. campinarensis e habit, f and g flowering branch. C. carinatus h habit, detail of the acropetiolar glands in insert, i staminate flower, j
pistillate flowers, k capsules.

Phenology:—Collected with flowers and fruits from September to February, and only with flowers in May.
Preliminary conservation status:—It can be categorized as Least Concern because its geografic range estimated
at roughly 200,000 km2.
Relationships:—Croton campinarensis could be confused with C. krukoffianus, since both have usually elliptic
leaf blades with stipitate glands at the base and margins, inflorescences with 1–5 pistillate flowers, and stigmas with
12 or more terminal tips. However, C. campinarensis can be easily distinguished by the petioles 1–2.7 cm long (vs.
0.1–0.5 cm long in C. krukoffianus), leaf base cordate or slightly cordate (vs. obtuse), thyrses 1.5–3.5 cm long without
a distinct sterile zone between pistillate and staminate flowers (vs. thyrses (1.6–)4–12 cm long, usually with a distinct
sterile zone between pistillate and staminate), pistillate flowers 5(6)-merous (vs. 6–8(–9)-merous), stamens 14 or 15
(vs. 11(–13)), and stigmas 4–6-fid (vs.8–16-fid).
Representative specimens examined:—BRAZIL. Pará: approx. 18 km east of Tucuruí and rio Tocantins, by BR-
263, 3°30’S, 49°32’W, 28 October 1981, D.C. Daly et al. 982 (INPA, HRB). Tucuruí, s.loc., 2 September 1983, F.E.L.
Miranda et al. 414 (INPA); margem direita do rio Tocantins, 29 May 1980, M.G. Silva & C. Rosário 5244 (INPA);
Campinas de Santa Rosa, 4 October 1983, J. Revilla et al. 8522 (INPA). Acará, Campina do Guajará, 8 December
2010, N. Cardoso 157 (IAN); Vila Cruzeirinho, Ramal do areial, km 28, 2°0’58”S, 48°19’9”W, 19 February 2014,
L.M.M. Carreira & F.C.A. Lucas 3306 (MFS), 3307 (MFS). VENEZUELA. Bolívar: Caicara, 100 m, 10 June 1940, L.
Williams 13250 (F, US).
11. Croton carinatus Müller Argoviensis (1865: 126).—Oxydectes carinata (Müll.Arg.) Kuntze (1891: 611). Type:—
BRAZIL. Tocantins: Dianópolis, Serra do Duro, September 1839, G. Gardner 3438 (lectotype: BR 8764232!, designated
here; isolectotypes: B† [photo at F 0BN005065!], E 00346718!, F 0056112F!, G-DC 00312238!, K 000186098!, OXF
00091584!, K 000186099!, P 00623084!, P 00623085!) (Fig. 18h–k).
Distribution and habitat:—Croton carinatus is known only from the municipality of Dianópolis, in Tocantins (Fig.
19a). Although BFG (2015) cites it exclusively for the state of Goiás, probably because of the locality “Prov. Goyaz,
Serra do Duro”, appearing on the labels of its type collection, we verified that the locality “Serra do Duro” actually
belongs to the municipality of Dianópolis, in Tocantins State. It grows in cerrado s.str. on sandy soils between 590 and
750 m elevation.
Phenology:—Collected with flowers from October to March and with fruits from December to May.
Preliminary conservation status:—This species is proposed here as Critically Endangered because of its
distribution in an unprotected area in an area of less than 100 km2.
Relationships:—Croton carinatus is easily recognized by its branches with whitish or yellowish indument;
petioles 1–3 mm long; chartaceous, linear leaf blades, with entire margins without glands; thyrses 1.7–19.5 cm long,
lax; staminate cymules with 3–5 flowers; and narrowly lanceolate, equal and non-accrescent pistillate sepals. This
species could be confused with other narrow-leaved species like C. junceus and C. gracilescens, but both species can
be separated by the features shown in Table 2.
TABLE 2. Main morphological characters distinguishing Croton carinatus, C. junceus and C. gracilescens.
Character C. carinatus C. gracilescens C. junceus
Acropetiolar glands Cylindric, usually absent Patelliform, always present Patelliform, always present
Leaf margin Entire Serrate or serrulate Entire
Marginal glands Stipitate-capitate, usually absent Patelliform, always present Absent
Pistillate sepals Linear-lanceolate Ovate Ovate or lanceolate
Taxonomic notes:— Croton carinatus was described by Müller Argoviensis (1865) based on G. Gardner 3438
from Tocantins state, as explained in the comment on geographical distribution. Of the eight duplicates of this collection
we found in six herbaria, the specimen at BR was the only one with an annotation by Müller, and it was chosen as the
lectotype.
In the recent infrageneric classification proposed by van Ee et al. (2011), Croton carinatus was not assigned to any
of the sections, but we place it here in sect. Geiseleria due to its subshrubby habit, stellate and multiradiate trichomes,
leaves with acropetiolar glands, and floral similarity with other species belonging to the section, such as C. agoensis
and C. sublepidotus.
FIGURE 19. Geographical distribution of some species of Croton sect. Geiseleria in Brazil. a Croton carinatus, C. catariae, C. crustulifer,
C. desertorum and C. glandulosus; b Croton glechomifolius, C. gracilescens, C. gracilirameus, C. goyazensis and C. grewioides.

Representative specimens examined:—BRAZIL. Tocantins: Dianópolis, Ponto 404(2), 11°33’35”S 46°28’48”W,
746 m, 24 September 2003, A.O. Scariot et al. 678 (CEN); Serra Geral do Tocantins, ca. 17 km ao longo da rodovia
que leva à Garganta, acessada pela TO-040 (Novo Jardim–Dianópolis), na altura do IFTO-Dianópolis, 11°35’46.6”S
46°40’16.8”W, 591 m, 06 May 2017, R.C. Sodré & U.S. Amaral 3189 (UFG); ibid., região das nascentes do rio
Palmeiras, ca. 30 km a leste de Dianópolis (em linha reta), ca. 2 km após a ponte sobre o rio Limoeiro, 11°36’11.5”S
46°30’29.6”W, 651 m, 07 May 2017, R.C. Sodré & U.S. Amaral 3192 (UFG), 3195 (UFG); ibid., ca. 5 km após a ponte
sobre o rio Limoeiro, 708 m, 11°36’24.1”S 46°28’23.2”W, 07 May 2017, R.C. Sodré & U.S. Amaral 3201 (UFG), 3202
(UFG), 3203 (UFG).
12. Croton catariae Baillon (1864: 349). Type:—BRAZIL. Mato Grosso: s.loc., 1833, C. Gaudichaud-Beaupré 247
(holotype: P 00623092!; isotypes: A 00047263!, RB photo 00663842) (Figs. 17t–x, 20a–d).
Croton chaetocalyx Müller Argoviensis (1865: 133).—Oxydectes chaetocalyx (Müll.Arg.) Kuntze (1891: 611). Type:—BRAZIL.
Megaponte, Villa Boa, 1837 or 1839, J.B.E. Pohl 1626 (holotype: B†; lectotype: BR 0000008764560!, designated here; isotypes: A
00063009!, F, G 00434439!, K 000186109!, W 0051197!, W 0051198!).
Vernacular name:—velame-bravo.
Distribution and habitat:—Croton catariae has been collected in Bolivia, Brazil (Silva et al. 2015a), and
Suriname (Secco 2008). In Brazil, it was reported for the Distrito Federal and for Goiás, Mato Grosso (Silva et al.
2015a), Pará (Secco 2008), and Tocantins (BFG 2015); here it is newly reported for Bahia, Maranhão, Mato Grosso do
Sul, and Minas Gerais (Fig. 19a). It grows in Cerrado s.str. and along forest edges, in sandy or clayey soils (Silva et al.
2015a), between 200 and 1,500 m elevation.
Phenology:—Collected in flower from September to April and in fruit from November to June.
Preliminary conservation status:—Croton catariae can be considered Least Concern because of its broad range,
numerous localities, and its Extent of Occurrence estimated at 1,500,000 km2.
Relationships:—Croton catariae is similar in many respects to C. gracilirameus, most notably in the conspicuous
stalked glands on the pistillate calyx. However, C. catariae can be distinguished by its robust branches with velutinous
indument; large leaf blades (4.7–7.8 × 3.7–5.7 cm), with secondary and tertiary veins prominent on the abaxial surface;
stipules, leaf margins, bracts and pistillate sepals with multiple, clustered, showy glands; and thyrses with 4–12 pistillate
flowers. In C. gracilirameus the branches are delicate and tomentose, the leaf blades are smaller (2.4–4.6 × 1.4–4.2
cm), with secondary veins impressed or discretely prominent on the abaxial surface and tertiary veins inconspicuous;
the glands of the stipules, leaf margins, bracts and sepals are few and discreet; and the thyrses have only one or two
pistillate flowers.
Taxonomic notes:—Although Croton catariae has traditionally been recognized as a member of C. sect. Barhamia,
the species shares many of the characteristics cited for the section Geiseleria, such as leaves with stipitate glands at the
base and in the sinuses of the leaf margins, inflorescences with unisexual cymules, and pistillate flowers with 5 free
and glandular sepals and bifid stigmas. Moreover, we note that C. gracilirameus, the morphologically closest species
to C. catariae, was recently described and assigned to Croton sect. Geiseleria (Silva et al. 2015), which supports our
position to transfer C. catariae to the same section.
Representative specimens examined:—BOLIVIA, Santa Cruz: José Miguel de Velazco, a 16 Km de SE de San
Ignacio, 16°30’44”S 60°53’4”W, 481 m, 25 January 2004, V. Solis Neffa et al. 1237 (US). Brazil, Bahia: Ibicoara,
região da Serra de Sincorá, 19 February 1943, R.L. Fróes 20149 (NY). Distrito Federal: Brasília, 3 Km on paved road
N of Fercal (intersection with dirt road to Sobradinho) ca. 25 Km NNE of the Brasília TV tower, 27 February 1988, S.
Ginzbarg 723 (UB). Goiás: Caldas Novas, at headwaters of the creek, Rio Quente, at hotel, “Pousada do Rio Quente”,
13 km WSW of City of Caldas Novas, 17°48’S, 48°45’W, 22 December 1974, E.P. Heringer and G. Eiten 14163 (US).
Monte Alegre de Goiás, 8 km by road W of Monte Alegre, 600–700 m, 11 March 1973, W.R. Anderson et al. 6837
(UB, US). Maranhão: Loreto, “Ilha de Balsas” region between the Balsas & Parnaíba Rivers, 35 km south of Loreto,
1 km east of main house of Fazenda Morros, 7°23’S, 45°4’W, 300 m, 13 February 1970, G. Eiten & L.T. Eiten 10590
(SP, UB, US). Mato Grosso: Nova Xavantina, approx. 10 km S of Xavantina, 14°44’S, 52°20’W, 11 January 1977,
J.A. Ratter et al. 4080 (UEC). Mato Grosso do Sul, Rio Verde de Mato Grosso, Sete Quedas, 8 February 1975, G.
Hatschbach et al. 34816 (MBM). Minas Gerais: Paracatu, Fazenda Ponta, Sítio do Sr. Dozinho, 31 March 1983, B.A.S.
Pereira 417 (IBGE). Pará: Conceição do Araguaia, Redenção, Area of cerrado vegetation about 4 km west of town
center along highway PA-150, 8°2’S, 50°4’W, 200 m, 21 February 1980, T. Plowman et al. 8995 (INPA). Tocantins:
Presidente Kennedy, road from highway BR-153 to Itaporã, 12 km west of village of Presidente Kennedy, Fazenda
Primavera along Ribeirão Feinho, 3°25’S, 48°37’W, 400 m, 31 January 1980, T. Plowman et al. 8199 (NY). Paranã,
cerca de 2 km a norte da vila Rosário, no sentido de São Salvador, 12°57’0”S, 48°8’19”W, 310 m, 26 March 2007, G.
Pereira-Silva & G.A. Moreira 11604 (CEN).
FIGURE 20. Croton catariae a habit, b staminate flowers and buds, c pistillate flowers, d young capsules. C. desertorum e habit, f thyrses
showing fruits and staminate flowers and buds, g staminate flowers and buds, h pistillate flowers, i and j fertile branches, k capsules.

13. Croton crustulifer Croizat, Darwiniana 6: 451. 1944.—TYPE: Brazil, Minas Gerais or Rio de Janeiro: Serra da
Mantiqueira, 1882, A.F.M. Glaziou 14254 (holotype: A 00047277!; isotype: K 000574178!, P 04831116, P 04831126)
(Fig. 21a, b).
Distribution and habitat:—Croton crustulifer is represented only by its type collection, coming from the Serra da
Mantiqueira, between the states of Minas Gerais and Rio de Janeiro (Fig. 19a). Its environmental conditions are
not known, but this mountain range has a mosaic of high-mountain vegetation composed of upper montane forests,
Araucária forests, high-altitude grasslands and rocky outcrops (Meireles et al. 2014).
Phenology:—There are no records about the time of flowering and fruiting of the species.
Preliminary conservation status:—As the species is known for only one location, we chose to consider it Data
Deficient.
Relationships:—Croton crustulifer resembles C. virgultosus in its shrubby habit, conspicuous stipules (more
than 2 mm long), petiole 1.3–2.5 cm long, and leaf blades usually ovate with coarsely serrate or dentate margin and
actinodromous-brochidodromous venation. Despite this, C. crustulifer differs by its rounded or obtuse leaf base (vs.
cordate in C. virgultosus), leaf glands not visible to the eye (vs. prominent glands), inflorescences up to 3 cm long (vs.
6–11 cm long), equal pistillate sepals with revolute margins (vs. unequal sepals with plane margin), and styles ca. 2
mm long (vs. 0.5–1 mm long).
Taxonomic notes:— Croizat (1944) described C. crustulifer based on Glaziou 14254, which has four duplicates,
one at A, one at K, and two at P (which Croizat apparently did not see). The collection at A has Croizat’s annotation
indicating that it is the holotype of species, although the author does not cite this in the protologue. Therefore, we treat
the collection at A as the holotype.
14. Croton desertorum Müller Argoviensis (1873: 265).—Oxydectes desertorum (Müll.Arg.) Kuntze (1891: 613).
Type:—BRAZIL. Minas Gerais: in deserto, s.d., C.F.P. von Martius s.n. (lectotype: M 0086104! [photo at F V006600!],
designated by Sodré et al. 2017; isolectotype: G 00434490!). (Figs. 20e–k, 21c–g)
Distribution and habitat:—Croton desertorum occurs in Goiás (Sodré et al. 2017), Bahia, and Minas Gerais (BFG
2015) (Fig. 19a). It inhabits campos sujos and cerrado s.str. on rocky soils between 900 and 1,540 m elevation.
Phenology:—Collected with flowers and fruits throughout the year.
Preliminary conservation status:—It can be categorized as Least Concern because its geographic range is
greater than 50,000 km2, besides having several populations growing in protected areas such as Chapada dos Veadeiros
National Park and Serra dos Pireneus State Park.
Relationships:—Croton desertorum is morphologically close to C. spica, but the latter is easily distinguishable
by the leaf blades with slightly cordate base and acuminate apex; staminate bracts 2.5–5 mm long; and pistillate
flowers with unequal sepals, pubescent on both surfaces and densely stellate stigmas. In C. desertorum the leaf base is
obtuse and the apex is acute; staminate bracts 1–1.5 mm long; and pistillate flowers with equal sepals, pubescent only
on the dorsal surface, and the stigmas are glabrous or glabrescent.
Representative specimens examined:—BRAZIL. Goiás: Água Fria de Goiás, a direita do km 61 da GO 118,
sentido São Gabriel a São João d’Aliança, 14°59’23.4”S, 47°36’16.4”W, 1119 m, 30 October 2014, R.C. Sodré et al.
1470 (UFG); Alto Paraíso de Goiás, cerca 30 km a esquerda da GO 118 de São João d’Aliança a Alto Paraíso de Goiás,
na estrada para catarata do rio dos Couros, 14°17’23.6”S, 47°42’33.2”W, 1049 m, 31 October 2014, R.C. Sodré et al.
1494 (UFG), 1496 (UFG), 1497 (UFG); São João d’Aliança, 10 km de São João d’Aliança/Alto Paraíso, margem da
rodovia GO-118, 25 November 1994, M. Aparecida da Silva et al. 2434 (UFG).
15. Croton glandulosus Linnaeus (1759: 1275).—Oxydectes glandulosa (L.) Kuntze (1891: 614). Type:—JAMAICA,
s.loc., s.d., P. Browne (lectotype: LINN 1140-7!, designated by Fawcett & Rendle (1920)). (Figs. 21h–k, 22a–e).
Croton divaricatus Swartz (1788: 100).—Croton glandulosus var. divaricatus (Sw.) Müller Argoviensis (1866: 684). Type:—“India
occidentalis”, s.d., s.coll., s.n. (holotype: S S-R-1305!)
Croton glandulosus var. gardneri Müller Argoviensis (1866: 684). Type:—BRAZIL. “Prov. Piauhy et Goyaz”, 1844, G. Gardner 3435
(lectotype: BR 0000008761965!, designated here; isolectotypes: F 0056126F!, G-DC 00312839!, NY 00246544!, NY 00246545!, P
00623602!, P 00623603!, P 00623604!).
Croton glandulosus var. intermedius Müller Argoviensis (1866: 684). Type:—BRAZIL. Bahia: Muritiba, 1842, J.S. Blanchet 3656
(lectotype: G-DC 00312838!, designated here; isolectotypes: G 0060034!, LE 00018217!); remaining syntypes:—BRAZIL. Bahia:
“in apricis”, 1830, P. Salzmann 495 (G 00312837!); MÉXICO, Vera Cruz, s.d., C.J.W. Schiede 62 (B).
Croton glandulosus var. occidentalis Müller Argoviensis (1873: 270). Type:—BRAZIL. “Brazilia occidentali”, s.loc., s.d., Tamberlik s.n.
(lectotype: W 0060012!, designated here; isolectotype: W 0060013!).
Croton scordioides Lamarck (1786: 215).—Croton glandulosus var. scordioides (Lam.) Müller Argoviensis (1866: 685). Type:—
BRAZIL. Rio de Janeiro: “Des environs de Rio Janeiro”, June 1767, P. Commerson s.n. (holotype: P-LAM!; isotypes: A 00277217!,
P 00623601!; P-JUSS 00674059!).
Croton glandulosus var. subincanus Müller Argoviensis (1866: 685), syn. nov. Type:—BRAZIL. Pará: Santarém, March 1850, R. Spruce
s.n. (lectotype: G 00312828!, designated here; isolectotype: TCD 0007623!).
Croton glandulosus var. warmingii Müller Argoviensis (1873: 269). Type:—BRAZIL. Minas Gerais: “in ruderatis prope Lagoa Santa”,
s.d., J.E.B. Warming 1646 (holotype: C n.v.; photo at F).
Vernacular name:—gervão, gervão-branco, malva-vermelha, velame.

Distribution and habitat:—Croton glandulosus is the most widely distributed species of section Geiseleria
in the Americas, occurring from Argentina to the central United States. It is introduced in Africa and Australia (van
Ee et al. 2011). In Brazil, it is found in all states (BFG 2015) (Fig. 19a), growing on forest edges, road margins, and
anthropic environments on rocky, clayey or sandy soil, from sea level to 1,250 m elevation.
Preliminary conservation status:—The wide distribution of this species and its considerable ecological range
and weedy nature place it in the category of Least Concern.
Relationships:—Croton glandulosus is often confused in herbaria with C. hirtus, C. lundianus, or C. trinitatis,
due to their herbaceous habits, similar aspect of growth and leaf forms, however, they differ in the characters listed in
Table 3.
TABLE 3. Morphological comparison between C. glandulosus and related species.

Characters C. glandulosus C. hirtus C. lundianus C. trinitatis
Thyrses Continuous Continuous Discontinuous Continuous
Length of bracts (mm) 1.2–2 1.8–2.2 1.4–3 0.4–0.7
Number of pistillate flowers 3–6 5–16 2 or 3 2–5
per thyrse
Glands of bracts Sessile Stipitate Absent Absent
Calyx of pistillate flowers 5-merous 5-merous 6(–8)-merous 5-merous
Length of stigmas (mm) 2–2.5 2–2.5 2.5–3 0.8–1

Capsule color Clear green, white lines Dark green without Dark green without Greenish or orange,
on the sutures lines on the sutures lines on the sutures without lines on the
sutures
Length of fruiting pedicel 0.3–0.8 1–1.5 1–2 2–4
(mm)
Representative specimens examined:—ARGENTINA: Corrientes: Ituzaingó, Ruta 34, ca. 3 km da ruta

14°16’41”S, 55°18’4”W, 140 m, 18 January 2007, J. Paula-Souza et al. 7506 (ESA). BRAZIL, Amapá: Macapá, na
estrada Pôrto Santana–Pôrto Platon, no km 100, 16 March 1962, J. Mattos & N. Mattos 9977 (SP). Amazonas: Manaus,
Rosa de Maio, km 8, December 1979, D.F. Coelho s.n. (INPA 92453). Bahia: Ilhéus, Fazenda Theobroma, próximo à
margem do rio Santana, 14°52’S, 39°4’W, 18 March 1988, S. Ginzbarg 777 (SP); Iramaia, Distrito de Novo Acre, 2
km ao NE de Jiquí, na estrada de Barra da Estiva para Iramaia, 13°30’S, 41°5’W, 24 March 1988, S. Ginzbarg et al.
862 (SP). Ceará: São Gonçalo do Amarante, Dunas do Pecém, próximo ao Porto, 13 June 2004, A.V. Vieira s.n. (EAC
34235). Espírito Santo: Santana, a 10 km oeste de Santana, via Nanuque-Santana, 15 December 1962, J. Mattos & H.
Bicalho 10776 (SP). Maranhão: Balsas, northern edge of city of Balsas, 7°32’S, 46°21’W, 13 March 1962, G. Eiten
& L.T. Eiten 3601 (SP); São Luís, Itaquihacanga, Campus Universitário UFMA, 27 April 1998, R.S. Secco & N.A.
Rosa 877 (MG, SP). Minas Gerais: Carangola, Perímetro Urbano, 20°43’S, 42°1’W, 8 November 1988, L.S. Leoni 508
(SP); Uberaba, BR Uberaba–Uberlândia, EPAMIG, 27 January 1984, L.C. Frutuoso 61 (UFMT). Pará: Belém, estrada
do Cafezal, na entrada da Reserva Mocambo, 3 February 1988, S. Ginzbarg et al. 670 (SP). Paraíba: Mamanguape,
estrada para Barra de Mamanguape, cerca de 5 km da BR 101, 6°54’56”S, 35°4’25”W, 50 m, 27 July 2001, V.C. Souza
& J.P. Souza 26599 (ESA, SP). Paraná: Paranaguá, Ilha das Cobras, 02 November 1986, W.S. Souza & S.M. Silva
25096 (UEC). Pernambuco: Petrolina, 8.1 km NNE of city of Petrolina, 9°20’S, 40°28’W, 8 March 1970, G. Eiten &
L. T. Eiten 10869 (SP). Piauí: Gilbués, Boqueirão, 15 February 1995, S.M. Rodrigues 230 (SP). Rio de Janeiro: Rio
de Janeiro, Parque Nacional da Tijuca, Vista Chinesa, no bordo da estrada, 21 November 1987, S. Ginzbarg & M.
Emmerich 655 (SP). Rio Grande do Sul: Alegrete, Cerro do Tigre, February 1990, M. Sobral & D.B. Falkenberg 6477
(SP). Rio Grande do Norte, Carnaúba dos Dantas, Fazenda Fundão, 28 May 2004, L.P. Felix et al. 10392 (HUEFS).
Rondônia: Nova Mamoré, Beira da estrada entre Guajará Mirim e Nova Mamoré, 10°13’8”S, 65°16’9.2”W, 112 m, 15
July 2011, N.C. Bigio et al. 318 (RON). Roraima: Boa Vista, Campus do Museu Integrado de Roraima (MIRR), Parque
Anauá, 2°50’23”S, 60°41’2”W, 3 March 2004, R.I. Barbosa 112 (INPA). Santa Catarina: Campos Novos, Rodovia
BR-470, 6 km L de Campos Novos, 11 February 1996, O.S. Ribas et al. 1265 (MBM, ESA). São Paulo: Angatuba,
Estação Ecológica de Angatuba, estrada de terra a oeste do Centro Administrativo da UC, 23°14’32”S, 48°12’58”W,
707 m, 18 December 2012, O.L.M. Silva et al. 42 (SP). Sergipe: Nossa Senhora do Socorro, Floresta Nacional do Ibura,
10°50’23”S, 37°8’37”W, 24 May 2013, T.R. Silva & J.P. Santana 144 (ASE). Tocantins: São Salvador do Tocantins,
Eixo A da barragem São Salvador, 12°48’12”S, 48°14’52”W, 250 m, 21 February 2002, G. Pereira da Silva et al. 5927
(CEN, SP). PARAGUAY: San Pedro: Alto Paraguay, Primavera, 24 November 1960, A.L. Woolston 1220 (SP)
16. Croton glechomifolius Müller Argoviensis (1865: 126).—Oxydectes glechomifolia (Müll.Arg.) Kuntze (1891:
611). Type: —BRAZIL. Paraná: Central East Mesoregion, 15 November 1828, F. Sellow s.n. (holotype: B†, photo
at F 249290!); neotype:—BRAZIL. São Paulo: “in campis graminosis ad furnas”, March 1826, L. Riedel 263 (LE
00018218!, designated here) (Fig. 21l–o)
Croton tragifolius Baillon (1865: 16). Type:—BRAZIL. São Paulo: “bords du chemin, près le rio Verde”, 1816–1821, A.F.C.P. Saint-
Hilaire cat. C2 1387 (lectotype: P 00493389!, designated here; isolectotype: P 00493390!).
Vernacular name:—velame.
Distribution and habitat:—This species is distributed in Argentina, Paraguay, and Brazil, in the latter occurring
in Paraná, Rio Grande do Sul, Santa Catarina, and São Paulo states (Caruzo & Cordeiro 2007) (Fig. 19b) in campo
limpo or campo sujo and on the edge of capão, on rocky or shallow soils, between 500 and 1,300 m elevation.
Phenology:—Collected with flowers and/or fruits from September to April.
Preliminary conservation status:—Croton glechomifolius has an estimated Extent of Occurrence of about
140,000 km2 and, therefore, it is classified here as Least Concern.
Characterization:—Croton glechomifolius is unique in Croton sect. Geiseleria in Brazil by its subshrubby habit
with generally prostrate stems; hirsute, stellate-porrect trichomes, petiole half to equal to the length of the blade, and
reniform or broadly ovate leaf blades with coarsely serrate or dentate margins without glands in the sinuses.
Taxonomic notes:—The neotypification of C. glechomifolius is necessary since its holotype at B was destroyed
during World War II. Riedel 263 at LE was mentioned for the species by Müller Argoviensis (1873) in Flora Brasiliensis
and reflects the original description. It was also annotated by Muller, so it was chosen here as neotype of the species.
Croton tragifolius was described by Baillon (1865) in the same year as C. glechomifolius based on Saint-Hilaire
cat. C2 1387, which has two duplicates at P. Eight years later Müller Argoviensis (1873) considered these species as
synonymous, which we concur with.
Representative specimens examined:—ARGENTINA. Misiones, Dep. Gral. Manuel Belgrano, Ruta Provincial
17, Campinas de América, Cementerio Campina o campo natural, 21 October 2006, H.A. Keller 3722 (CTES, SP).
BRAZIL. Paraná: Curitiba, Bairro Santo Inácio, Parque Bariguí, campo sujo na beira da Alameda Ecológica Burle
Marx (em frente ao Expo Renault Bariguí), 14 January 2015, M.B.R. Caruzo et al. 184 (SP); Lapa, localidade de
Serrinha, 25°43’49.2”S, 49°42’22.8”W, 920 m, 18 October 2012, I. Cordeiro et al. 3371 (SP); Porto Amazonas, ca.
50 km a O. de Curitiba, ca. 3 km ao S do entroncamento entre BR-227 e BR-376, beira de capão a leste da estrada
de terra, 11 February 1988, S. Ginzbarg et al. 687 (SP); Piraí do Sul, Serra do Piraí, km 15 em direção à Ventania, 03
April 2010, S.E. Martins & S.L. Pompéia 1287 (SP). Santa Catarina: Água Doce, Usina Eólica Horizonte, 26°34’53”S,
51°41’54”W, 1294 m, 23 September 2013, J.T. Motta et al. 4091 (ICN, MBM, SP); Campo Erê, Fazenda Campo São
Vicente, 24 km west of Campo Erê, 900–1000 m, 26–28 December 1956, L.B. Smith et al. 9483 (US, NY, HBR, R).
São Paulo: Itararé, Fazenda Espinho, 24°16’12”S, 49°16’8”W, 1200 m, 06–10 September 1993, V.C. Souza et al.
4175 (ESA, SP). Rio Grande do Sul: Cruz Alta, 16 December 1937, B. Rambo s.n. (PACA-AGP 3047). PARAGUAY.
Amambay: “in campis siccis Ponta Porã”, 1907–1908, E. Hassler 9915 (G)
FIGURE 21. a, b Croton crustulifer a leaf, b pistillate flower; c–g C. desertorum c leaf, d basilaminar glands, e marginal gland, f
staminate flower, g pistillate flowers; h–k C. glandulosus h leaf, i acropetiolar glands, j staminate flower, k pistillate flowers; l–o C.
glechomifolius l leaf, m acropetiolar glands, n staminate flower, o pistillate flower; p–s C. goyazensis p leaf, q basilaminar glands, r
staminate flower, s pistillate flower; t–x C. gracilescens t leaf, u basilaminar glands, v marginal gland; w staminate flower, x pistillate
flower. Drawn from a, b A. F. M. Glaziou 14254, c–g R. C. Sodré 1628, h–k G. Hatschbach 74374, l–o J. T. Motta 4091, p–s R. C. Sodré
2908, t–x R. C. Sodré 1413.

17. Croton goyazensis Müller Argoviensis (1865: 120).—Oxydectes goyazensis (Müll.Arg.) Kuntze (1891: 611).
Type:—BRAZIL. Goiás: “in campis petrosis Catalão”, August 1834, L. Riedel 2473 (lectotype: LE 00003114!,
designated by Sodré et al. 2017; isolectotypes: B†, photo at F F0BN005103!, G 00434536!, G-DC 00312494!, GH
00047305!, K 000186100!, LE 00003111!, LE 00003112!, LE 00003113!); remaining syntypes:—BRAZIL. Goiás:
“in campis Catalão”, September 1834, P.W. Lund s.n (F V0093325F!); s.loc., s.d., J.B.E. Pohl 1618 (BR 8762290!, K
000913002!, K 000913003!) (Figs 21p–s, 22f–k).
Croton luzianus Müller Argoviensis (1873: 218).—Oxydectes luziana (Müll.Arg.) Kuntze (1891: 613). Type:—BRAZIL, Goiás: “ad
Ponto Alto prope S. Luzia”, s.d., J.B.E. Pohl 1618 (catalogue number 1034) (lectotype: W 0051288!, designated by Sodré et al. 2017;
isolectotypes: F V0056157!, G 00434543!, W 0051287!); remaining syntypes:—BRAZIL. Goiás: “ad Ponto Alto”, s.d., J.B.E. Pohl
s.n. (catalogue number 1037) (W 0051286!).
Croton goyazensis var. rotundifolius Glaziou (1913: 619), nom nud.
Vernacular name:—alcanforzinho.
Distribution and habitat:—Croton goyazensis occurs in the Distrito Federal and in the states of Goiás and
Minas Gerais (Fig. 19b). Although it was reported by Carneiro-Torres (2009), Alves (2012), and BFG (2015) for
other Brazilian states as well as for Paraguay and Bolivia, we do not agree with these reports since we verified that
the specimens from these localities identified as C. goyazensis were in fact other species belonging to Croton sect.
Geiseleria, such as C. parodianus and C. seccoi. Croton goyazensis grows in campo sujo, cerrado ralo or cerrado
typical in clay or stony soils, between 750 and 1350 m elevation.
Phenology:—Flowering and fruiting the whole year, but especially from July to December.
Preliminary conservation status:—Croton goyazensis can be categorized as Least Concern because its Extent of
Occurrence was estimated at roughly 60,000 km2, besides having several populations in protected areas (e.g., Chapada
dos Veadeiros National Park, National Forest of Silvânia, Ecological Reserve of the IBGE and Brasília National
Park).
Relationships:—Croton seccoi was identified as C. goyazensis by Carneiro-Torres (2009) and Alves (2012)
in herbarium collections. However, C. goyazensis differs by its subshrubby habit up to 1 m tall, cylindric or conic
basilaminar glands with a pronounced orifice at the apex, eglandular bracts, pedicels of staminate flowers 2.8–5 mm
long, pistillate flowers with equal, lanceolate or linear sepals with 0.6–1 mm large, and oblong capsules with curved
pedicels 3.3–9 mm long. Croton seccoi is characterized by a shrubby habit 0.8–2.3 m tall, shortly stipitate patelliform
glands without an orifice, glandular bracts, pedicels of staminate flowers 1.7–1.9 mm long, pistillate flowers with
unequal, oblong, oval-lanceolate or oval-elliptic sepals 1–1.7 mm large, and globose capsules with erect pedicels 1.3–2
mm long.
Taxonomic notes:—Chodat and Hassler (1905) described two additional varieties for C. goyazensis, var.
angustifolius based on E. Hassler 7865 and var. major based on E. Hassler 5041, 5871 and 7292, all from Paraguay.
After analyzing these collections, we found that they do not correspond to Croton goyazensis, but rather to a member
of C. sect. Adenophylli, as evidenced by the columella with three prominent lobes.
Representative specimens examined:—BRAZIL. Distrito Federal: Brasília, Reserva Ecológica do IBGE, limite
leste da reserva, 12 December 2016, R.C. Sodré 2908 (UFG), 2913 (UFG), 2915 (UFG), 2916 (UFG), 2918 (UFG).
Goiás: Goiânia, a esquerda da rodovia Goiânia para Trindade, no km 12, 13 April 1968, A. Barbosa & J.A. Rizzo 415
(UFG). Minas Gerais: Uberlândia, saída pela BR 452, sentido Belo Horizonte, 18°55’22.9”S, 48°12’10.8”W, 940 m,
06 July 2016, R.C. Sodré et al. 2518 (UFG), 2519 (UFG).
18. Croton gracilescens Müller Argoviensis (1865: 110).—Oxydectes gracilescens (Müll.Arg.) Kuntze (1891: 611).
Type:—BRAZIL. Goiás: “prope Cavalcante”, s.d., J.B.E. Pohl 297 (holotype: B 100242753!, photo at F F0BN005104;
isotypes: A 00257930!, BR 0000008762627!, G-DC 00434535!, K 000254364!; K 000254366!) (Figs. 21t–x, 23a–f).
Croton antisyphiliticus var. angustifolius Müller Argoviensis (1865: 110), syn. nov.—Ocalia angustifolia Klotzsch (1841a: 195), nom.
nud. Type:—BRAZIL. “Brazilia Meridionali”, s.d., F. Sellow s.n. (B†).
Croton antisyphiliticus var. weddellianus (Baill.) Müller Argoviensis (1866: 593), syn. nov..—Croton perdicipes var. weddellianus Baillon
(1864: 337). Type:—BRAZIL. Goiás: Mara Rosa, Sertão d’Amaroleite, Sep–October 1844, H.A. Weddell 2753 (lectotype: P
05566744!, designated here; isolectotype: P 05566743!).
Vernacular name:—pé-de-perdiz, pé-de-perdiz-fêmea, pé-de-perdiz-lisa.

Distribution and habitat:—Croton gracilescens occurs in Goiás (Sodré et al. 2014), and is also reported here for
Mato Grosso and Tocantins (Fig. 19b). It grows in campo sujo or cerrado ralo, usually on rocky soil between 290 and
1040 m elevation.
FIGURE 22. Croton glandulosus a habit, b short inflorescences with pistillate and staminate flowers, c inflorescences showing staminate
flowers and buds, see detail of the glands at the base of the bracts in the insert, d pistillate flowers, e capsules with accrescent sepals
and white lines on the capsules. C. goyazensis f habit, g flowering branch, h staminate flowers, i pistillate flowers, j young capsules, k
capsules.

Phenology:—Collected with flowers and fruits from September to February.
Preliminary conservation status:—Croton gracilescens can be categorized as Least Concern because it has
an estimated Extent of Occurrence of about 85,000 km2 and because it is common in protected areas such as Serra
Dourada State Park and Chapada dos Veadeiros National Park.
Relationships:—Croton gracilescens is sometimes mistakenly identified as C. antisyphiliticus, under which it
was synonimized by Govaerts et al. (2000) and van Ee (2011). Nevertheless, it can be distinguished by the characters
listed in the discussion of C. antisyphiliticus. Others species similar to Croton gracilescens are C. carinatus and C.
junceus, especially due to their subsessile, very narrow leaves, but they can be distinguished by the features shown in
Table 2.
Taxonomic notes:—After analysis of the original diagnoses of Croton perdicipes var. weddellianus and C.
antisyphiliticus var. angustifolius, we verified that the characters listed by Baillon (1864) and Müller Argoviensis
(1865), such as thin and elongated branches and narrowly lanceolate leaves, are typical of C. gracilescens and not C.
antisyphiliticus, so we propose the synonymization of these two varieties under C. gracilescens. We chose as lectotype
of Croton antisyphiliticus var. weddellianus the sheet at P with barcode 05566744, because it contains more branches
and leaves than the other sheets at P, and it has fruits and flowers of both sexes.
Representative specimens examined:—BRAZIL. Goiás: Alto Paraíso de Goiás, estrada para o Vale da Lua,
14°10’25.4”S, 47°47’5.1”W, 1,040 m, 15 October 2010, M.J. Silva 3057 (UFG); ibid., 22 October 2011, M.J. Silva
et al. 3856 (UFG), 3878 (UFG); Minaçu, estrada Minaçu–balsa “Porto dos Paulistas”, cerca de 8.2 km de Minaçu
(km marcados no rio Bonito), 13°31’0”S, 48°9’21”W, 380 m, 7 November 2000, G. Pereira-Silva et al. 4346 (CEN).
Mato Grosso: Alto Araguaia, 22 km from Alto Araguaia towards Rondonópolis, 700 m, 22 September 1988, R. Kral
& M.G.L. Wanderley 75015 (NY). Tocantins: Dianópolis, próximo à área de empréstimo de cascalho para a nova
rodovia, 11°36’9”S, 46°31’2”W, 29 September 2003, T.B. Cavalcanti et al. 3323 (CEN). Paranã, margem esquerda do
rio Tocantins, 12°48’45”S, 48°14’35”W, 290 m, 25 November 2007, G. Pereira-Silva et al. 12347 (CEN).
19. Croton gracilirameus M.J.Silva, Sodré & P.E.Berry (in Silva et al. 2015a: 162). Type:—BRAZIL. Goiás: Alto
Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, 3 km a oeste do Morro do Buracão, 14º6′33.6″S,
47º43′32.7″W, 1181 m, 10 February 2013, M.J. Silva et al. 4716 (holotype: UFG 60363!; isotype: MICH!) (Figs.
23g–j, 24a–e).
Distribution and habitat:—Croton gracilirameus occurs from the southwest to northeast of the state of Goiás (Fig.
19b), with records for the municipalities of Alto Paraíso de Goiás, Cavalcante, Goiás, Mossâmedes and Piranhas. It
grows in cerrado s.str. on sandy soil and rocky outcrops, between 949 and 1405 m elevation.
Phenology:—Collected with flowers from October to May and with fruit from November to April.
Preliminary conservation status:—Silva et al. (2015a) classified C. gracilirameus as Least Concern because
it is locally common in its area of occurrence and because it inhabits protected areas, such as Chapada dos Veadeiros
National Park and Serra Dourada State Park.
Relationships:—Croton gracilirameus and C. catariae are morphologycally similar, but the two species differ
from each other in several features discussed under C. catariae.
Representative specimens examined:—BRAZIL. Goiás: Alto Paraíso de Goiás, Serra do Pouso Alto,
aproximadamente 3 km a direita da GO 118, sentido Alto Paraíso de Goiás a Teresina de Goiás, entrando no cruzeiro,
13°50’7.3”S, 47°15’22.5”W, 953 m, 06 April 2013, R.C. Sodré et al. 680 (UFG); Cavalcante, Parque Nacional da
Chapada dos Veadeiros, base da Serra de Pouso Alto, 09 March 2012, R.C. Sodré 47 (UFG). Mossâmedes, na cabeceira
do rio Índio Grande, 1º transecto, 13.I.1994, fl., J.A. Rizzo 10907 (UFG).
20. Croton grewioides Baillon (1864: 365).—Croton glycosmeus Müller Argoviensis (1865: 140), nom illeg. Type:—
BRAZIL. Bahia: Marais de Japira, 1840, J.S. Blanchet 3100 (lectotype: P 00623585!, designated here; isolectotypes: A
00257931, A 00257932!, BR 0000008768223!, C 10011172!, F 0056131F!, F0056132F!, G 00312861!, G 00434532!,
G-DC 00434533!, K 000254381!, K 000254382!, K000254383!, LE 00003110, MO 260056!, NY 00246546!, OXF
00091947!, P 00623583!, P 00623584!, TUB 009091!) (Fig. 24f–j).
Croton zehntneri Pax & Hoffmann (1923: 176). Type:—BRAZIL. Bahia: Lapa, Caatinga da Mulata, April 1912, L. Zehntner 4084
(holotype: M 0089093!, photo at F 209696[6621]).
Vernacular name:—alecrim-de-cabocla, angico, angélica, caatinga-de-cheiro, canelinha, canelinha-de-cheiro,

cravinho, criola, marmeleiro-de-folha-miúda.
FIGURE 23. Croton gracilescens a habit, b fertile branch, c staminate flowers, d and e pistillate flowers, f capsules. C. gracilirameus g
habit, h staminate flowers, i pistillate flower, j capsules (note the stalked glands on the calyx).

FIGURE 24. a–e Croton gracilirameus a leaf, b basilaminar glands, c marginal gland; d staminate flower, e pistillate flower; f–j C.
grewioides f leaf, g basilaminar glands, h marginal gland, i staminate flower, j pistillate flower; k–o C. hadrianii k leaf, l acropetiolar
glands, m marginal gland, n staminate flowers, o pistillate flowers; p–t C. harleyi p leaf, q basilaminar glands, r marginal gland, s
staminate flower, t pistillate flowers; u–x C. hirtus u leaf, v acropetiolar glands, w staminate flower, x pistillate flower. Drawn from a–e
R. C. Sodré 680, f–i V. J. Santos 254, j V. J. Santos 255, k–o R. C. Sodré 2872, p–s O. F. Oliveira 135, t R. C. Sodré 3337, u–y R. C. Sodré
2830.
Distribution and habitat:—Croton grewioides is known from Bolivia (Møller-Jørgensen et al. 2014) and Brazil,
and it is also reported here for Peru. In Brazil, it occurs in Alagoas, Bahia, Ceará, Minas Gerais, Paraíba, Pernambuco,
Piauí, Rio Grande do Norte, and Sergipe (Silva et al. 2010) (Fig. 19b). It inhabits in shrubby caatinga or arboreal
caatinga, usually near rocky outcrops, on sandy or clayey soil, between 200 and 1200 m elevation (Carneiro-Torres
2009).
Preliminary conservation status:—Due to its wide distribution and considering that many of its populations
occur in protected areas, we treat C. grewioides as Least Concern.
Relationships:—Croton grewioides resembles C. seccoi by its shrubby habit up to 2.5 m tall, branches with
ferrugineous or brownish indument, leaves with shortly stipitate patelliform glands, obtuse or lightly cordate leaf base,
thyrses with 6 or more pistillate flowers and flowers of both sexes with evident pedicels (more than 0.8 mm long).
Despite this, C. grewioides is an aromatic plant growing with a monopodial aspect and its pistillate flowers have equal,
lanceolate or linear-lanceolate sepals 0.4–1 mm wide and the gynoecium equal to or up to twice as long as the pedicel;
whereas C. seccoi is a non-aromatic plant growing with a sympodial aspect with pistillate flowers with oblong, oval-
lanceolate or oval-elliptic, unequal sepals, 1–1.7 mm wide, and the gynoecium 3 to 6 times longer than the pedicels.
Taxonomic notes:—In the protologue of C. grewioides, Baillon (1864) cited Blanchet 3100 at “herb. Mus. et
Less.”, signifying P, where he worked, and at the Delessert Herbarium, which was later incorporated into G. Therefore,
the lectotype has to be either a sheet at P or else one at G that would have been part of the original Delessert Herbarium.
To be most consistent and since Baillon worked at P, we chose as lectotype the sheet at P with barcode 00623585,
which is also one of the most complete with fully developed leaves and flowers of both sexes.
Representative specimens examined:—BOLIVIA. Chuquisaca: Belisario Boeto, Subiendo del Puente Rosita
em el Rio Grande em dirección a Nuevo Mundo, 18°52.153’S, 64°17.531’W, 1389 m, 24 November 2004, J.R.I. Wood
& H Huaylla 21063 (HSB, MICH). BRAZIL. Alagoas: Olho d’Água do Casado, Fazenda Capelinha, 9°30’50.5”S,
37°49’6.3”W, 11 July 200, R.A. Silva & D. Moura 1569 (SP). Bahia: Senhor do Bonfim, Serra de Santana, base da
serra, 7 February 2003, V.J. Santos et al. 254 (HRB, HUEFS), 255 (HRB, HUEFS). Ceará: Aiuaba, Estação Ecológica
de Aiuaba, Serra do Êrmo, 26 February 1997, L.W. Lima-Verde et al. 518 (EAC, SP). Minas Gerais: Grão-Mogol, Vale
do Rio Itacambiruçu, 6°53’S, 41°48’W, 670 m, 05 September 1990, T.R.S. Silva et al. CFCR 13287 (SP). Paraíba:
Borborema, November 1936, H. Zenaide 24a (SP). Pernambuco: Gravatá, 14 April 1995, L.P. Félix s.n. (SP 332132);
Buíque, Parque Nacional Serra do Catimbau, Serra das Torres, ápice da Serra, 8°33’53”S, 37°14’34”W, 833 m, 28 June
2007, D.S. Carneiro-Torres et al. 956 (HUEFS, SP). Piauí: Teresina, 36 km northwest of Picos on BR 316 to Teresina,
6°53’S, 41°48’W, 24 January 1993, W. Thomas et al. 9608a (NY, SP). Rio Grande do Norte: Mossoró, Serra do Carmo,
5°14’S, 37°6’W, 08 May 1979, O.F. Oliveira 687 (MOSS). Sergipe: Feira Nova, 22 July 2014, L.A.S. Santos 1104
(ASE). Peru, Amazonas: Luya District, Camporredondo, em la subida a Guadalupe, 1280 m, 1991, J. Campos 2130
(MO, MICH).
21. Croton hadrianii Baillon (1864: 313). Type:—BRAZIL, s.loc, 1783–1792, A.R. Ferreira s.n. (comm. Étienne
Geoffroy Saint-Hilaire) (lectotype: P 00623573!, designated by Sodré & Silva (2018b); isolectotype: P 00623574!,
photo at F 1039147). (Figs. 24k–o, 25a–f)
Distribution and habitat:—Croton hadrianii was recently rediscovered in Pará, Mato Grosso, and Rondônia (Sodré
& Silva 2018b) (Fig. 26a), in the Amazon and Cerrado Phytogeographic Domains or in transition areas between them.
It inhabits hillsides on shallow or stony soils, in open vegetation or on the edge of forests, between 85 and 810 m
elevation.
Phenology:—Collected in flower and fruit from December to July.
Preliminary conservation status:—Sodré & Silva (2018b) categorized C. hadrianii as Least Concern due to it
wide distribution (Extent of Occurrence estimated at roughly 500,000 km2).
Relationships:—Croton hadrianii is morphologically most similar to C. krukoffianus, but it differs by its
inconspicuous stipules (0.2–0.6 × 0.3–0.5 mm), petioles 0.7–1.2 cm long with sessile, patelliform acropetiolar glands
1.3–1.7 mm diam., oval-lanceolate leaf blades with inconspicuous sessile patelliform glands in the sinuses of the
margins, and stigmas with 6–12 terminal tips. In C. krukoffianus the stipules are 1.2–5 × 0.5–1.7 mm, the petioles
0.3–0.5 cm long with cylindric or stipitate patelliform glands up to 1 mm diam., the leaf blades are usually elliptic with
conspicuous stipitate glands in the sinuses, and the stigmas have 24–48 terminal tips. Croton hadrianii can also be
confused with C. subincanus by the petioles with sessile patelliform glands and leaf blades with subentire margins and
whitish indument on the abaxial surface, but C. hadrianii is easily differentiated by its usually ovate or oval-lanceolate
FIGURE 25. Croton hadrianii a habit, b and c thyrse showing staminate flowers and buds, d thyrse showing pistillate flowers and
staminate buds, e pistillate flower, f capsules. C. harleyi g habit, detail of a flowering branch in the lower right corner and a thyrse in the
upper left corner, h staminate flower, i and j pistillate flowers.
leaf blades; thyrses with 2–4(–6) pistillate flowers with 5–7 sepals and usually 4-fid stigmas; and fruiting pedicels
0.5–1 mm long; while in C. subincanus the leaf blades are elliptic; the inflorescences have 8–12 pistillate flowers with
5 sepals and 2-fid stigmas, and the fruiting pedicels are 1.5–2 mm long.
Representative specimens examined:—BRAZIL. Mato Grosso: Alta Floresta, Complexo Hidrelétrico Apiacás,
PCH Cabeça-de-boi, Acesso Península, 10°21’18”S, 56°58’40”W, 292 m, 18 December 2014, C.R.A. Soares-Lopes et
al. 8198 (HERBAM); Aripuanã, Região da Serra Morena e Vale do Ouro, a direita da estrada de terra que liga as sedes
dos municípios de Aripuanã e Castanheira (MT-420), 10°33’18.8”S, 58°53’3.2”W, 473 m, 08 April 2017, R.C. Sodré
& A.O. Souza 3133 (UFG); Porto Estrela, Estação Ecológica Serra das Araras, ca. 1 km a direita da porteira de entrada
da reserva, acessada pela Fazenda Coqueiro, 15°46’36.3”S, 57°15’9.6”W, 810 m, 05 December 2016, R.C. Sodré &
N.A.G. Escobar 2872 (UFG). Pará: Jacareacanga, UHE São Manoel-Sete Quedas, 9°19’5”S, 56°46’4”W, 11 March
2016, L.P. Zanzini et al. 76 (HERBAM); Marabá, Serra dos Carajás setor n 4, 25 March 1977, fl., fr., M.G. Silva & R.
Bahia 2917 (INPA); Parauapebas, Serra dos Carajás, 25–30 km NW of Serra Norte mining camp., 5°55’S, 50°26’W,
5 December 1981, D.C. Daly et al.1712 (HRB, INPA). Rondônia: Itapuã do Oeste, Floresta Nacional do Jamari,
12 km from km 602 of BR-364 in Itapuã do Oeste along Estrada Principal, Trilha da Pedra Grande, 9°11’53.59”S,
63°4’53.51”W, 15 March 2012, W. Castro et al. 154 (RON); Porto Velho, estrada Porto Velho-Cuiabá, BR 364, km
159, 9 February 1983, L.O.A. Teixeira et al. 1414 (INPA); Jaci-Paraná, margem direita do rio Madeira, em frente à
corredeira de Jirau, 9°19’17”S, 64°43’35”W, 85 m, 23 March 2010, fl., G. Pereira-Silva et al. 15140 (CEN, HUEFS,
RON).
22. Croton harleyi Carn.-Torres & Cordeiro (in Carneiro-Torres et al. 2011: 128). Type:—BRAZIL. Bahia: Rio de
Contas, Serra do Rio de Contas, 13º49′16″S, 41º35′50″W, 5 December 2004, R.M. Harley et al. 55288 (holotype:
HUEFS 000002836!; isotypes: SP, HUESB) (Figs. 24p–t, 25g–j).
Distribution and habitat:—Croton harleyi occurs in Bahia, Pernambuco, Piauí, and Rio Grande do Norte (Carneiro-
Torres et al. 2011) (Fig. 26a), in arboreal caatinga on sandstone outcrops between 415 and 1013 m elevation.
Phenology:—Collected in flowerand fruits from December to April.
Preliminary conservation status:—According to Carneiro-Torres et al. (2011) C. harleyi can considered as
Least Concern, due to its wide distribution, including in protected areas, such as the Serra das Confusões National Park
in Piauí.
Characterization and relationships:—Croton harleyi is distinguished from the others species in the section by
its small, elliptic leaf blades (2–4.8 × 0.9–2 cm) with serrulate, eglandular margins; short inflorescences (1–2 cm long);
linear bracts 3–4 mm long, and pistillate flowers with large, equal, ovate to elliptic sepals (3.5–6 × 2.5–3 mm) that are
accrescent with revolute margins. The morphological relationships of C. harleyi with C. arenosus are discussed in the
comments of that species.
Representative specimens examined:—BRAZIL. Bahia: Casa Nova, margem do lago de Sobradinho, 9°33’43”S,
41°46’6”, 415 m, 29 February 2008, C. Correia et al. 428 (HUEFS); Palmeiras, em direção à Serra Negra, 12°31’51.6”S,
41°35’17.4”, 736 m, 19 December 2017, R.C. Sodré et al. 3337 (UFG), 3338 (UFG), 3339 (UFG), 3340 (UFG); Sento
Sé, Alta do Tabuleiro, topo da serra, próximo à torre, PNBO, 10°36’24.3”S, 41°31’32.9”, 1013 m, 27 January 2010,
A.P. Fontana et al. 6401 (HUEFS). Rio Grande do Norte: São Vicente, José Antônio, a 2 km do Saco de Santa Cruz,
6°9’30”S, 36°40’00”, 500 m, 01 March 1980, O.F. Oliveira, J.D.O. Freitas & J.E. Araújo 135 (MOSS). Piauí: Caracol,
Serra das Confusões, 08 December 1980, A. Fernandes & Del’Arco s.n. (EAC 9099, HUEFS 000003043)
23. Croton hirtus L’Héritier de Brutelle (1785: 17).—Croton glandulosus var. hirtus (L’Hér.) Müller Argoviensis (1866:
684).—Podostachys hirta (L’Hér.) Klotzsch (1841a: 194).—Brachystachys hirta Klotzsch (1843: 47).—Oxydectes
glandulosa var. hirta (L’Hér) Kuntze (1891: 614).—Croton glandulosus subsp. hirtus (L’Hér.) Croizat (1948: 401).
Type:—FRENCH GUIANA. s.loc., s.d., L.C. Richard s.n. (holotype: P 00623551!; isotype: P 00623550!) (Fig. 24
u–x, 27a–d).
Croton glandulosus var. tenellus Müller Argoviensis (1866: 685), syn. nov. Type:—VENEZUELA. s.loc., s.d., A. Fendler 1236 β (holotype:
G 00312827!).
Croton glandulosus var. sagotii Müller Argoviensis (1873: 269), syn. nov. Type:—FRENCH GUIANA. Karouany, 1857, P.A. Sagot
503 (lectotype: BR 0000008762085!, designated here; isolectotypes: G 00312852!, S S-R-10503!); remaining syntype:—BRAZIL.
Tocantins: “inter Funil et S. João”, s.d., W. Burchell 8913 (BR 0000008763136!).
Croton glandulosus var. martii Müller Argoviensis (1873: 268). Type:—BRAZIL. Maranhão: “in campis et pascuis ad flumen Itapicurú
prope Caxias”, s.d., C.F.P. Martius 2557a (lectotype: M 0089077!, designated here; isolectotypes: K 000254389!, L 0234460!, M

0089078!, M 0089079!).
Croton glandulosus var. schomburgkianus Müller Argoviensis (1873: 268). Type:—GUYANA. s.loc., 1836, Rob. Schomburgk 101
(lectotype: BM 001125625!, designated here; isolectotypes: BM 001124406!; F 0056127F!, G 00312849!).
Croton glandulosus var. sieberi Müller Argoviensis (1873: 268), syn. nov. Type:—TRINIDAD AND TOBAGO. “in insula Trinitatis”,
s.d., Sieber Fl. Trinit. 106 (lectotype: L 0234397!, designated here; isolectotypes: L 0234398!, G 00312850!); remaining syntype:—
VENEZUELA. Vargas: near La Guayra, 1860, A. Fendler 1236 (G 00312851!).
Croton glandulosus var. cordovensis Müller Argoviensis (1874: 203), syn. nov. Type:—ARGENTINA, “in fruticetis et silvaticis umbrosis
propre Cordoba”, 1870, P.G. Lorentz 294 (lectotype: G 00378337!, designated here; isolectotypes: B†, CORD 00003076!)
Vernacular name:—croto, malva-de-pêlos, malva-grande.

Distribution and habitat:—Croton hirtus is widely distributed in the Americas, from Mexico to Argentina
(Govaerts et al. 2000), but can also be found as an adventive in some Old World countries (van Ee et al. 2011). In
Brazil, it occurs in most states (AL, AM, AP, BA, CE, DF, GO, MA, MG, PA, PB, PE, PI, PR, RN, RS, SE, SC, SP and
TO; Caruzo & Cordeiro 2007; Carneiro-Torres 2009; BFG 2015) and is being cited here for the states of Mato Grosso
and Mato Grosso do Sul (Fig. 26a). It grows in different types of vegetation, usually associated with disturbed areas,
in clayey, sandy or stony soils, from sea level to 1,100 m elevation
Preliminary conservation status:—Croton hirtus can be considered as Least Concern for its wide distribution,
efficient ability to colonize environments, and frequency of occurrence.
Relationships:—Croton hirtus can be recognized by the following combination of characters: subshrubby habit
0.2–0.6(–1) m tall; vegetative and reproductive organs with simple and stellate-porrect, stiffly hirsute trichomes; petioles
with narrowly cylindric glands at the apex; dark green, ovate, membranaceous leaf blades with rounded or truncate
base, serrate or crenate margins usually without glands, semicraspedodromous venation with 3–5 veins arising at the
base; inflorescences 0.9–6 cm long, without a distinct sterile zone between pistillate and staminate flowers; bracts with
4–6 stipitate-pyriform glands; pistillate flowers with unequal sepals, the largest obovate, oblanceolate or spatulate; and
dark green capsules with hyaline trichomes. The morphological relationship of C. hirtus with C. aberrans is discussed
in the notes of that species, and Table 3 shows the main differences between C. hirtus, C. glandulosus, C. lundianus,
and C. trinitatis.
Taxonomic notes:—We interpret P 00623551 as the holotype of C. hirtus because it is part of the original Richard
Herbarium, whereas P 00623550 was part the de Franqueville Herbarium, which was later incorporated into the Drake
Herbarium, and finally this was donated to P after the death of Drake in 1904. Therefore it is unlikely that L’Héritier
de Brutelle (1785) could have seen this latter specimen.
After analyzing all the types of taxa described under Croton hirtus and C. glandulosus, we propose the
synonymization of C. glandulosus var. tenellus Müll.Arg., C. glandulosus var. sagotii Müll.Arg., C. glandulosus var.
sieberii, and Croton glandulosus var. cordovensis Müll.Arg. under C. hirtus because they have branches and leaves
with hirsute trichomes and bracts with long-stipitate pyriform glands, characteristics unique to C. hirtus. Croton
glandulosus var. martii Müll.Arg. and C. glandulosus var. schomburgkianus were treated as synonyms of C. hirtus by
van Ee (2011) and Zuloaga et al. (2008), respectively, which is followed here.
Representative specimens examined:—ARGENTINA. Córdoba: Cólon, Colanchanga, 31°8’43”S, 64°21’27”W,
945 m, 09 March 2013, P. Demaio 312 (WU). BRAZIL. Alagoas: Porto Calvo, Fazenda Macacos, 05 November
1981, C.R. Campêlo & M.S. Silva 1626 (UEC). Bahia: Iaçu, rodovia Iaçu/Milagres (BA-046), km 13, início do ramal
para as fazendas Suíbra e Morros, 12°49’S, 40°10’W, 240 m, 22 March 1988, S. Ginzbarg et al. 798 (MBML).
Ceará: Iguatu, fazenda Trussu/RPPN Trussu, 6°19’54”S, 39°22’63”W, 29 May 2008, B.G. Lima 101 (MOSS). Distrito
Federal: Brasília, borda da Mata do Córrego Riacho Fundo, no lado direito, atrás da casa, 15°52’00”S 48°00’00”W,
1100 m, 09 February 2001, E.S.G. Guarino et al. 641 (CEN). Goiás: Cavalcante, porção central do povoado Kalunga
Engenho II, 16 February 2013, R.C. Sodré & M.T. Faria 535 (UFG). Maranhão: São Luís, Praia do Coalho, 02
February 1989, Marques & Ferreira 67 (SP). Mato Grosso: Cuiabá, Campus da Universidade Federal do Mato Grosso,
estacionamento do Instituto de Biociências, 15°36’34.7”S 56°03’46.1”W, 02 December 2016, R.C. Sodré & N.A.G.
Escobar 2830 (UFG). Mato Grosso do Sul: Aquidauana, Aldeia Terena Limão Verde, 20°19’44”S, 55°41’29”W, 300
m, 17 August 2003, A. Pott & V.J. Pott 11301 (HMS). Minas Gerais: Belo Horizonte, campus da UFMG, February
1978, J.A. Oliveira 2321 (BHCB). Pará: Belém, estrada do Cafezal, na entrada da Reserva Mocambo, 3 February
1988, S. Ginzbarg et al. 671 (SP). Paraíba, s.loc., 3 January 1935, H. Zenaide 8 (SP). Paraná: Campo Mourão, 13
October 1965, G. Hatschbach et al. 12961 (MBM). Pernambuco: Ilha Fernando de Noronha, próximo ao Hotel de
Transito do Governo, 32°25’S, 3°50’W, 31 May 1993, A.M. Miranda et al. 786 (SP). Rio Grande do Norte: Almino
FIGURE 26. Geographical distribution of some species of Croton sect. Geiseleria in Brazil. a Croton hadrianii, C. harleyi, C. hirtus, C.
hostmannii and C. inaequilobus; b C. junceus, C. krukoffianus, C. lundianus, C. macrosepalus and C. mollis.

Afonso, Sítio Serra de São Miguel, 6°10’S, 37°45’W, 450 m, 24 March 1994, F.E. Sobrinho 145 (MOSS). São Paulo:
Mogi Guaçú, Martinho Prado, Reserva Biológica da Fazenda Campininha, 22 December 1980, W. Mantovani 1450
(SP). Sergipe: Poço Redondo, 10 km da cidade, 9°49’S, 37°40’W, 23 August 2005, D.S. Carneiro-Torres et al. 478
(HUEFS). Tocantins: Ananás, acampamento de ENGEVIX, próximo ao aeroporto, 6°7’59”S, 48°20’19”W, 140 m, 15
April 2004, G. Pereira-Silva et al. 8647 (CEN). Panamá, Canal Zone: in west Ft. Kobbe, towards Vera Cruz, 31 July
1963 J.D. Duke & H.W. Mussell 6562 (LD). Peru, Prope Tarapoto: Peruviae orientalis, June 1855, R. Spruce 4237 (LD).
Sri Lanka, Sabaragamuwa Province: Balangoda road, Imbulpe village, close to Belihul Oya rest house, 154 mile post;
Balangoda–Belihul Oya road, 6°42’11.6”N, 80°45’0”E, 500 m, 16 January 2003, H.S. Kathriarachchi et al. 37 (WU).
24. Croton hostmannii Schltdl. ex Miquel (1848: 477). Type:—SURINAME, s.loc., s.d., F.W.R. Hostmann 1106
(holotype: U 0001885!; isotypes: A 00063011!, G 00434525!, G 00312276!, K 000574186!, K 000574187!, S-R-
10529!) (Fig. 28a–e).
Vernacular name:—carrapicho-do-cerrado.
Distribution and habitat:—Croton hostmannii is known from Brazil and Suriname (Secco 2008) and also
reported here for Guyana. In Brazil, it was cited for Maranhão by Secco (2008) and is being reported here for Roraima
(Fig. 26a). It grows in areas of campina on sandy soils between 50 and 450 m elevation.
Phenology:—Collected with flowers in February, and with flowers and fruits in May and August.
Preliminary conservation status:—Croton hostmannii has an Extent of Occurrence estimated at 200,000 km2
and, therefore, considered as Least Concern.
Relationships:—Croton hostmannii is similar to C. subincanus by its shrubby habit 1–3 m tall, elliptic leaf
blades with entire or subentire margins, and thyrses 5–14 cm long with 8–12 pistillate flowers and without a distinct
sterile zone between pistillate and staminate flowers. However, C. hostmannii can be distinguished by its shortly
stipitate patelliform acropetiolar and marginal glands (vs. petiole with sessile glands and eglandular leaf margin in C.
subincanus), leaf blades glabrous or sparsely pubescent on the adaxial surface (vs. densely pubescent) and pistillate
calyx with lobes united up to halfway from base (vs. free sepals).
Representative specimens examined:—BRAZIL. Roraima: Boa Vista, estrada para a Colônia do Apiaui, 2°50’N,
60°40’W, 07 August 1986, J.A. Silva et al. 663 (NY). Maranhão: Santa Quitéria, Fazenda Bom Jesus, 15 February
2006, Carreira 2702 (MG). GUYANA. Potaro-Siparuni: Kaieteur Plateau, 7 May 1944, B. Maguire & D.B. Fanshawe
23291 (IAN, RB).
25. Croton inaequilobus Steyermark (1958: 7). Type:—BRAZIL. Goiás: Chapada dos Veadeiros, 14 km South of
Veadeiros, in sandstone rocky area on west bank of stream and above, 47°30’S, 14°20’W, 25 April 1956, E.Y. Dawson
14685 (holotype: R 000104977!; isotypes: F V0056145F!, RSA 0002666!) (Figs. 27e–i, 28f–j).
Distribution and habitat:—Croton inaequilobus is endemic to the Chapada dos Veadeiros, in the northeast portion of
Goiás (Sodré et al. 2017) (Fig. 26a). It grows in cerrado rupestre in rock crevices usually close to rivers, in dark sandy
soil between 876 and 1,500 m elevation.
Phenology:—Collected with flowers and fruits in all months of the year, although its flowering and fruiting is
abundant from December to May (Sodré et al. 2017).
Preliminary conservation status:—Although Croton inaequilobus occurs within the Chapada dos Veadeiros
National Park (CVNP), its Extent of Occurrence is estimated at only about 80 km2, and therefore we consider its
conservation status as Vulnerable.
Relationships:—Silva et al. (2014) compared Croton inaequilobus to C. planaltoanus, but C. inaequilobus is
unique by the following set of characteristics: shrubby habit 0.6–2.3 m tall; black stems with lenticels; vegetative
and reproductive organs with whitish or pinkish, tomentose indument of stipitate, multirradiate or stellate trichomes;
petioles with two reddish, orange or yellowish, patelliform, sessile glands; ovate or oval-lanceolate leaf blades with
serrulate or crenulate margin; congested inflorescences s 1–2.9 cm long, with only 2 or 3 pistillate flowers; bracts
1.6–4.3 mm long, without glands; pistillate flowers with unequal sepals, the largest obovate or oblanceolate; clear
green capsules; and black, ovoid or oblongoid seeds with clear spots.
Representative specimens examined:—BRAZIL. Goiás: Alto Paraíso de Goiás, Parque Nacional da Chapada
dos Veadeiros, ca. 2 km a norte do alojamento principal do Parque, 14°8’37”S, 47°47’37”W, 978 m, 05 June 2015,
M.J. Silva et al. 6748 (UFG). Cavalcante, Parque Nacional da Chapada dos Veadeiros, após às Corredeiras do rio Preto,
14°9’17.4”S, 47°50’8.5”W, 876 m, 8 March 2013, R.C. Sodré et al. 593 (UFG), 594 (UFG).
FIGURE 27. Croton hirtus a habit, b fertile branch, c inflorescences showing pistillate and staminate flowers, d capsules, note the stipitate
glands of the bracts. C. inaequilobus e habit, f vegetative branch, with detail of the acropetiolar glands in the insert, g inflorescence with
pistillate flowers and staminate flowers and buds, h staminate flowers, i capsules, note the persistent bracts even after the staminate flowers
fall.

FIGURE 28. a–e Croton hostmannii a leaf, b acropetiolar glands, c marginal gland, d staminate flower, e pistillate flower; f–j C.
inaequilobus f leaf, g acropetiolar glands, h marginal gland, i staminate flower, j pistillate flower; k–n C. junceus k leaf, l acropetiolar
glands, m staminate flower, n pistillate flower; o–s C. krukoffianus o leaf, p acropetiolar glands, q marginal gland, r staminate flower, s
pistillate flower; t–w C. lundianus t leaf, u acropetiolar glands, v staminate flower, w pistillate flower. Drawn from a–e B. Maguire 23291,
f–j /M. J. Silva 6748, k–n R. C. Sodré 2598, o–s G. T. Prance 28847 e t–w G. Hatschbach 5362.
26. Croton junceus Baillon (1864: 339). Type:—BRAZIL. Mato Grosso: “entre Goyas et Cujaba”, November–
December 1844, M. Weddell 2940 (lectotype: P 00623522!, designated here); remaining syntypes:—BRAZIL. Mato
Grosso: “sources du Paraguay, envir. de Diamantino”, November–December 1844, M. Weddell s.n. (P 00623523!, P
00623524!) (Figs. 28 k–n, 29a–d).
Distribution and habitat:—Croton junceus is known from Goiás, Mato Grosso, and Mato Grosso do Sul (Sodré &
Silva 2018a) (Fig. 26b). It grows in campo sujo or cerrado ralo on sandy soil between 500 and 840 m elevation.
Phenology:—Collected with flowers from September to January and fruits from October to January.
Preliminary conservation status:—Croton junceus has an Extent of Occurrence of about 140,000 km2, and
therefore we categorized its conservation status as Least Concern.
Relationships:—Croton junceus is a linear-leaved species that is morphologically similar to C. carinatus and C.
gracilescens, from which it differs by the characters shown in Table 2; it also shares similarities with C. uliginosus, as
discussed by Sodré & Silva (2018a). In general, C. junceus can be recognized by its dark green stems and leaves with
sparse hyaline trichomes; petioles with sessile patelliform glands; concolorous, linear leaf blades 0.2–0.7 mm wide,
margins entire without glands and brochidodromous venation with secondary veins inconspicuous and impressed on
both surfaces; lax thyrses and pistillate flowers with equal, ovate to lanceolate sepals.
Representative specimens examined:—BRAZIL. Goiás: Caiapônia, Bacia do rio Caiapó, 17°4’20”S, 51°34’2”W,
752 m, 17 November 2007, S.S. Silva et al. 492 (IBGE). Chapadão do Céu, Parque Nacional das Emas, às margens
do rio Formosa, ponte próximo à sede do Parque, 17 January 2012, M.B.R. Caruzo et al. 154 (SP). Mato Grosso:
Água Boa, 46 km S of Água Boa by BR 158, 8 October 1988, R. Kral et al. 75229 (NY), Araguainha, campo limpo
com gramíneas ca. 1,5 m alt., parcialmente alagado, acima da sede da fazenda Ribeirão das Pedras (prop. Licionora
Gonzaga), 18 January 2011, R.D. Sartin et al. 160 (UFG). Barra do Garças, ca. 50 km N. of Barra do Garças on road
to Xavantina, 15°53’S, 52°15’W, 16 October 1964, H.S. Irwin & T.R. Soderstrom 6941 (NY). Nova Xavantina, ca.
6 km S of Xavantina, 14°38’S, 52°14’W, 9 September 1967, G. Argent et al. 6324 (NY, UB). Paranatinga, estrada
Chapada–Paranatinga, Fazenda Roncador, 04 October 1980, M. Macedo & S. Assumpção 1737 (UFMT, UEC). Mato
Grosso do Sul: Costa Rica, Adjacências do Parque Nacional das Emas, Fazenda Pontal, próximo a ferrovia Ferronorte,
ca. 4 km ao sul da estrada que leva de Chapadão do Céu à trijunção GO/MT/MS, 18°22’38.4”S, 53°0’46.2”W, 838 m,
21 September 2016, R.C. Sodré 2598 (UFG).
27. Croton krukoffianus Croizat (1944: 453). Type:—BRAZIL. Rondônia: near Tabajara, upper Machado River
region, 1 December 1931, B.A. Krukoff 1480 (lectotype: NY 00233106!, designated here; isolectotypes: G 00434571!,
P 00623521!, S 07-12786!) (Fig. 28o–s, 29e–g).
Croton spiraeifolius Jablonszki (1965: 164), syn. nov. Type:—VENEZUELA. Amazonas: Cerro Yapacana, savanna 3, 1 January 1951, B.
Maguire, R.S. Cowan & J.J. Wurdack 30571 (holotype: NY 00688115!; isotypes: F V0093314F!, S 07-12772!, US 00810683!).
Croton subserratus Jablonszki (1965: 164), syn. nov. Type:—VENEZUELA. Amazonas: Cerro Guanay, summit, 1800 m, 2 February
1951, B. Maguire, K.D. Phelps, C.B. Hitchcock & G. Budowski 31731 (holotype: NY 00688114!; isotypes: US 00810679!, VEN
749581).
Distribution and habitat:—Croton krukoffianus is widely distributed in campinaranas in Brazil and Amazonian
Venezuela between 17 and 1800 m elevation. In Brazil, it is known from the states of Amazonas, Mato Grosso,
Roraima and Rondônia (Secco 2008) and is newly cited here for Pará (Fig. 26b).
Phenology:—Collected with flowers the year round, and with fruits from November to February.
Preliminary conservation status:—Croton krukoffianus can be categorized as Least Concern because its Extent
of Occurrence was estimated at roughly 500,000 km2.
Relationships:—Croton krukoffianus is similar to C. campinarensis e C. hadrianii, but is distinguished by the
characters listed in the comments of those species.
Taxonomic notes:—Croton krukoffianus was described by Croizat (1944) based on Krukoff 1480 from the
Province of Mato Grosso, in a region currently belonging to Rondônia state. Since Croizat (1944) did not mention
any herbarium for the type in the protologue of the species, we selected the NY sheet since it contains a label with
Krukoff’s annotations, and this was also his home institution. Jablonski (1965) later described C. spiraeifolius and C.
subserratus, however, he did not discuss possible similarities or differences between them, or with C. krukoffianus.
We conclude that the three names constitute the same species after the analysis of approximately 30 specimens of the
three species from the northern region of Brazil, where we could observe a continuous variation in the form of the
petiole glands and in the degree of serration of the margins and in the density of trichomes of the leaf blades. These
FIGURE 29. Croton junceus a habit, b staminate flowers, c pistillate flowers, d capsules and a leaf, with detail of the acropetiolar glands
in the insert. C. krukoffianus e flowering branch, f thyrses showing young capsules and staminate flowers and buds, g thyrse showing
staminate staminate flowers and buds. C. lundianus h habit, i detail of the leaf base showing the stipules and the acropetiolar glands, j
thyrse showing a pistillate flower separated from the staminate buds by a sterile zone, k staminate flowers, l capsule.
species, on the other hand, always have short petioles (less than 8 mm long), elliptic leaf blades with stipitate glands in
the margin sinuses, thyrses with 1–4 pistillate flowers separated from the staminate ones by a sterile zone, as well as
pistillate flowers with 5–8 oblong or lanceolate sepals and stigmas with 24 to 48 tips, characters that unify them and
that allowed us to treat them as conspecific.
Representative specimens examined:—BRAZIL. Mato Grosso: Apiacás, rio Juruena, 8°53’27.6”S, 58°33’7.2”W,
November 2007, M. Sobral et al. 11070 (BHCB). Amazonas: Apuí, Igarapé Coca-Cola, 7°26’3”S, 61°56’22”W, 86 m,
19 August 2009, P.A.C.L. Assunção & F.A. Carvalho 1141 (INPA, UFG); Presidente Figueiredo, km 125 da estrada
Presidente Figueiredo–Caracaraí, 14 October 1997, M.G.G. Vieira et al. 1356 (INPA, UFG). Pará: Mundurukú, Alto
Tapajós, region of Missão Velha, ca. 2 km N of the Rio Cururú, 7°45’S, 57°20’W, 200 m, 13 February 1974, W.R.
Anderson 10900 (NY); Rondônia: Porto Velho, ao longo da BR-364, 9 km E do povoado Jirau, ramal ao sul, 9°21’39”S,
64°39’31”W, 114 m, 23 June 2013, M.F. Simon et al. 1889 (CEN, INPA, IAN). Roraima: Caracaraí, Parque Nacional
Viruá, 1°14’46”N, 61°7’24”W, 62 m, 22 January 2010, M.C. Dávila et al. 5904 (INPA).
28. Croton lundianus (Didr.) Müller Argoviensis (1866: 662).—Podostachys lundianus Didrichsen (1857: 137).—
Croton lundianus var. genuinus Müller Argoviensis (1866: 663), nom. inval. Type:—BRAZIL. São Paulo: Taubaté,
November 1833, P.W. Lund s.n. (holotype: C 10011178!) (Figs. 28t–w, 29h–l).
Croton teucridium Baillon (1864: 349), syn. nov. Type:—BRAZIL. Espírito Santo: s.loc., 1816–1821, A.F.C.P. Saint-Hilaire cat. C2, 2e
part., 262-b (lectotype: P 00634743!, designated here; isolectotypes: A 00047452!, F 0056208F!, P 00634742!).
Croton yacaensis Croizat (1944: 459), syn. nov. Type:—PARAGUAY. Alto Valle del Rio Y-acá, December 1900, E. Hassler 6804
(lectotype: G 00307115!, designated here; isolectotypes: A 00047464!, F V0192148F!, G 00307116!, G 00307114!, P 00634779!, P
00634780!).
Croton similis Müller Argoviensis (1865: 134).—Croton lundianus var. similis Müller Argoviensis (1866: 663).—Podostachys incana
Klotzsch (1841a: 194), nom. nud.—Podostachys sellowiana Klotzsch (1841a: 194), nom. nud.—Croton klotzschii var. incana Baillon
(1864: 347), nom. nud. Type:—BRAZIL. “Brazilia meridionali”, s.loc., s.d., F. Sellow s.n. (lectotype: K 000254394!, designated
here; isolectotypes: BR 0000013317614!, K 000254395!, P 00633410!).
Croton klotzschii var. bahiensis Baillon (1864: 347).—Croton lundianus var. bahiensis Müller Argoviensis (1866: 663). Type:—BRAZIL.
Bahia: s.loc., April 1834, J.S. Blanchet 1665 (lectotype: P 00633411!, designated here; isolectotypes: A 00257944!, A 00257945!, A
00257946!, G 00434544!, G 00312475!, P 00633412!).
Croton lundianus var. glabrescens Müller Argoviensis (1866: 664). Type:—BRAZIL. Bahia: s.loc., 1834, J.S. Blanchet 1824 (lectotype: G
00434547!, designated here); remaining syntype:—BRAZIL. Bahia: “In collibus aridis”, 1830, Salzmann 494 (G 00312468!).
Croton lundianus var. gracilentus Müller Argoviensis (1866: 664). Type:—BRAZIL. Ceará: s.loc., Kalkmann 141 (holotype: MEL;
isotype: G 00312466!).
Croton klotzschii var. hilarii Baillon (1864: 347).—Croton lundianus var. hilarii (Baill.) Müller Argoviensis (1866: 663). Type:—
BRAZIL. São Paulo: s.loc., 1816–1821, A.F.C.P. Saint-Hilaire cat C1 1116 (lectotype: P 05565739, designated here; isolectotype: A
00257947!); remaining syntype:—BRAZIL. São Paulo: s.loc., s.d., F. Sellow s.n. (P).
Croton klotzschii var. latifolius Baillon (1864: 347).—Croton lundianus var. latifolius (Baill.) Müller Argoviensis (1866: 663). Type:—
BRAZIL. Rio de Janeiro: s.loc., 1831–1833, C. Gaudichaud-Beaupré 1137 (lectotype: P 00633407!, designated here; isolectotype:
G 00312476!); remaining syntype:—BRAZIL. Rio de Janeiro: Serra d’estrella, 1844, H.A. Weddell 821 (P 00633408!).
Croton lundianus var. leptophyllus Müller Argoviensis (1866: 664). Type:—BRAZIL. Pernambuco: s.loc., November 1837, G. Gardner
1129 (holotype: B†; lectotype: K 000254405!, designated here; isolectotypes: G 00434549!, K 000254404!)
Croton lundianus var. major Müller Argoviensis (1866: 663). Type:—BRAZIL. s.loc., s.d., J.B.E. Pohl s.n. (holotype: B).
Croton klotzschii var. microphylla Baillon (1864: 347).—Croton lundianus var. microphyllus (Baill.) Müller Argoviensis (1866: 664).
Type:—BRAZIL. São Paulo: s.loc., s.d., A.F.C.P. Saint-Hilaire 1115 (holotype: P 05565736)
Croton lundianus var. mollis Müller Argoviensis (1866: 664). Type:—BRAZIL. Espirito Santo: s.loc., s.d., M.A. zu Wied-Neuwied s.n.
(holotype: B†; lectotype: M 0086076!, designated here; isolectotype: G 00434545!)
Croton lundianus var. pubescens Müller Argoviensis (1866: 664). Type:—BRAZIL. Minas Gerais: s.loc., s.d., C.F.P. Martius (holotype:
B†).
Croton lundianus var. serratus Müller Argoviensis (1866: 664).—Podostachys serrata Klotzsch (1841a: 194), nom. nud.—Croton
dentifolius Baillon (1864: 347), nom. nud. Type:—BRAZIL. Bahia or Espírito Santo: s.loc., s.d., F. Sellow s.n. (lectotype: BR
0000008710857!, designated here; isolectotypes: P 00633409!, K 000254396!); remaining syntypes:—BRAZIL. Bahia: s.loc., s.d.,
J.S. Blanchet 1824 (A 00257943!, G 00434546!, G 00312469!)
Croton lundianus var. hirtus Müller Argoviensis (1873: 230). Type:—BRAZIL. “Brazilia meridionali”, s.d., F. Sellow s.n. (lectotype: BR
0000008710833!, designated here; isolectotypes: G 00434551!, LE 00018219!); remaining syntypes:—BRAZIL. s.loc, s.d., F.C.
Raben 763 & 783 (BR 0000008710826!).

Croton lundianus var. intercedens Müller Argoviensis (1873: 229). Type:—BRAZIL. São Paulo: s.loc, s.d., W. Burchell 3910 (lectotype:
K 000254400!, designated here; isolectotype: G 00434548!).
Croton lundianus var. psilophyllus Müller Argoviensis (1873: 229). Type:—BRAZIL. Minas Gerais: “Padre de Correia, Sumidouro et
Ribeirâo”, s.d., J.B.E. Pohl 1611 (catalogue number 14) (holotype: W 0060098!).
Croton lundianus var. rufidulus Müller Argoviensis (1873: 227). Type:—BRAZIL. “forte in prov. Goyaz”, s.loc, s.d., J.B.E. Pohl s.n. (not
found).
Podostachys subfloccosa Didrichsen (1857: 135).—Croton lundianus var. subfloccosus (Didr.), Müller Argoviensis (1866: 663). Type:—
BRAZIL: São Paulo: Taubaté, s.d., P.W. Lund s.n. (lectotype: C 10011179!, designated here; isolectotype: C 10011180!).
Vernacular name:—gervão, gervão-miúdo, gervão-branco.

Distribution and habitat:—Croton lundianus occurs in Argentina, Brazil, and Paraguay (Silva et al. 2010).
In Brazil, the species occurs in all regions (AL, BA, CE, ES, GO, MG, PE, PI, PR, RJ, SC, SE, SP and TO) (Fig.
26b). Although C. lundianus was cited for the Amazon Forest Domain, in Amazonas, Maranhão, Mato Grosso, Pará,
and Rondônia, as well as in Venezuela and Guyana by Carneiro-Torres (2009) and BFG (2015), we verified that
the specimens that would justify such a distribution correspond to other species, such as C. campinarensis or C.
sipaliwinensis, and for this reason we do not include these distributions here. Croton lundianus inhabits forest edges,
cerrado s.str., shrubby caatinga, campos rupestres and in disturbed environments, on sandy or clayey soils, between
200 and 1,400 m elevation (Cordeiro 1992; Lima & Pirani 2003; Caruzo & Cordeiro 2007; Carneiro-Torres 2009;
Lucena 2009; Silva et al. 2010).
Preliminary conservation status:—Due to its wide distribution and capacity to inhabit different types of
vegetation, we categorized its conservation status as Least Concern.
Relationships:—Croton lundianus shows wide morphological variation, with decumbent or erect plants, branches
and leaves glabrescent to densely pubescent, and leaf blades from widely ovate to narrowly lanceolate with serrulate,
serrate, deeply or doubly serrate margins; however, its thyrses with discontinuous flowers and pistillate flowers with 6
or 7 white, oblanceolate or obovate sepals with serrulate margins are characters that clearly identify the species. Croton
lundianus is morphologically closest to C. sipaliwinensis mainly due to the ovate or oval-lanceolate leaf blades, thyrses
with 2–4 pistillate flowers, the latter usually 6-merous and separated from the staminate flowers by a sterile zone, and
the glabrous or glabrescent calyx of the staminate flowers. However, C. sipaliwinensis has obconic leaf glands (vs.
stipitate patelliform in C. lundianus), 4-merous staminate flowers with the calyx joined for 1/3 to 1/2 of its length
(vs. 5-merous with calyx joined for 1/4 or 1/5 of its length), stamens 5 (vs. 8–10); linear, oblong-linear or lanceolate
pistillate sepals (vs. oblanceolate or obovate), and ovary always glabrous (vs. usually pubescent).
Representative specimens examined:—BRAZIL. Bahia: Igrapiúna, Faz. Michelin, 27 February 2001, S. Sampaio
17 (HRB); Mucugê, Unidade de Manejo Sustentável, próximo à confluência do rio Mucugê com o rio Combuca,
12°59’41”S, 41°21’14”W, 970 m, 03 January 1997, H.P. Bautista et al. 40 (HRB). Espírito Santo: Guarapari, Parque
Estadual Paulo César Vinha, 20°36’1”S, 40°28’0”W, 23 May 1997, J.M.L. Gomes 2814 (VIES, UFG), 2826 (VIES,
UFG). Goiás: Posse, aproximadamente 15 km ao Norte da cidade de Posse em sentido ao Posto do Rosário, 27 January
2017, A.O. Souza 1912 (UFG). Maranhão: Anajatuba, 25 January 1976, B.G.S. Ribeiro & G.S. Pinheiro s.n. (HRB
12817). Minas Gerais: Belo Horizonte, margem da Represa da Pampulha, 30 November 1982, T.S.M. Grandi & R.C.F.
Carvalho 119 (BHCB); Santana do Riacho, APA da Serra do Cipó, estrada para Conceição do Mato Dentro, próximo a
ponte do antigo km 123, 1200–1400 m, 9 January 2001, M. Alves et al. 2176 (SPF, BHCB, IAN). Paraná: Cerro Azul,
Rio Turvo p/ Cerro Azul, In silva caedua, 20 November 1957, G. Hatschbach 4313 (PACA); Ponta Grossa, Parque
Estadual de Vila Velha, 19 March 2012, J.M. Silva et al. 8211 (HUCS). Rio de Janeiro: Parque Nacional do Itatiaia,
30 January 1975, T.M. Pedersen 10917 (C). Teresópolis, Parque Nacional da Serra dos Órgãos, 900 m, March 1953,
J. Vidal 6606 (RB, INPA). Santa Catarina: Florianópolis, Fazenda Experimental da Ressacada, Universidade Federal
de Santa Catarina, 27°41’40”S, 48°32’45”W, 06 November 2012, B. Munhoz 2 (FLOR). São Paulo: São José dos
Campos, parte inferior da encosta leste do vale do córrego da Ressaca, 6 1/2 km ao SW (ao longo da Rodovia Pres.
Dutra) da saída para São José dos Campos, e dentro de 500 m ao SE da rodovia, 560–590 m, 14 November 1961, I.
Mimura 82 (WU); São Paulo, Reserva do Instituto de Botânica de São Paulo, 15 November 1980, N.A. Rosa & J.M.
Pires 3743 (INPA). Sergipe: Itabaiana, Serra de Itabaiana, subida e topo da Serra, 10°41’S, 37°26’W, 31 March 1988,
M.C. Ferreira 126 (HRB).
29. Croton macrosepalus Sodré & M.J. Silva (in Sodré et al. 2019: 237). Type:—BRAZIL. Tocantins: Dianópolis,
Serra Geral do Tocantins, in the region where the source of the river Palmeiras is located, ca. 30 km east to the urban
area of Dianópolis (in a straight line), ca. 5 km after the bridge over the river Limoeiro, 11°36’24.1”S, 46°28’23.2”W,
708 m, 7 May 2017, R.C. Sodré & U.S. Amaral 3198 (holotype: UFG!; isotypes: K!, P!, RB!, SP! (Figs. 30a–b,
31a–d).
Vernacular name:—velame-roxo.
Distribution and habitat:—An endemic species of Brazil, with records from Bahia, Goiás, and Tocantins (Fig.
26b). It grows in cerrado s.str. on sandy soils between 450 and 890 m elevation.
Phenology:—Collected with flowers and fruits from December to May.
Preliminary conservation status:—Sodré et al. (2019) recommended the conservation of this species as
Vulnerable due to its estimated Extent Occurrence of 6,000 km2, and because it is known from fewer than ten localities
with populations smaller than 20 individuals each, located in areas unprotected and usually subject to agricultural
activities.
Relationships:—Croton macrosepalus was compared to C. spica by Sodré et al. (2019) by the particular
morphology of their pistillate flowers. However, we believe that C. macrosepalus can also be confused with some
species of C. sect. Geiseleria also have entire or subentire leaf margins, such as C. mollis, C. rhodotrichus, and C.
subincanus. In addition to the similar leaf margins, these species also share with C. macrosepalus the usually elliptic
and discolorous leaf blades with whitish, tomentose indument on the adaxial surface and brochidodromous venation
with 4–8(–10) pairs of secondary veins impressed on both surfaces and bracts with sessile glands, although they can
be differentiated by other characters shown in Table 4.
TABLE 4. Comparative morphology of Croton macrosepalus and related species.

Character C. macrosepalus C. mollis C. rhodotrichus C. subincanus
Habit Subshrubs Shrubs Subshrubs Shrubs
Height (m) 0.2–0.6 1–2 0.5–1 1–3
Marginal glands Globose or ovoid cylindrical Ovoid or pyriform (lacking)
Inflorescence (cm) 1.3–2.8 4.5–16 2–5 5–7.5
No pistillate flowers per inflorescence 2 or 3 5–14 2–4 6–12
Length of bracts (mm) 3.2–4.8 3–5 1.3–2 0.9–1
Staminal filaments Glabrous Villous Glabrous Glabrous
Shape of the largest pistillate sepals Ovate or largely Lanceolate or Spatulate-obovate Oblong or obovate
ovate oblong
Largest pistillate sepals (mm) 5.5–8.3 × 3.3–5.3 3–5 × 1.2–2.3 3.2–5 × 1.2–2.1 3.5–5 × 1.4–2
Representative specimens examined:—BRAZIL. Bahia: Correntina, Fazenda Jatobá, 13°–14°S, 45°45’–

46°15’W, 3 March 1991, L.G. Viollati et al. 226 (UB, IBGE). Goiás: Posse, ca. 2 km da área urbana de Posse,
em direção à BR 020, 14°06’27.4”S 46°19’28.3”W, 823 m, 15 December 2016, R.C. Sodré & C.C. Oliveira 2979
(UFG). Tocantins: Dianópolis, Serra Geral do Tocantins, região das nascentes do rio Palmeiras, ca. 30 km a leste da
área urbana de Dianópolis (em linha reta), ca. 2 km após a ponte sobre o rio Limoeiro, 11°36’11.5”S 46°30ʹ29.6”W,
651 m, 07 May 2017, R.C. Sodré & U.S. Amaral 3192 (UFG).

FIGURE 30. Croton macrosepalus a habit, b fertile branch, detail of a congested thyrse with young capsules and staminate flowers in
the upper right corner and a capsule, with the large, unequal sepals in the lower right corner. C. mucronifolius c habit, with detail of the
acropetiolar glands in the insert, d pistillate flowers, e capsules.
FIGURE 31. a–d Croton macrosepalus a leaf, b acropetiolar glands, c staminate flower, d pistillate flower; e–i C. mollis e leaf, f
acropetiolar glands, g marginal gland, h staminate flower, i pistillate flower; j–n C. nepetifolius j leaf, k basilaminar glands, l marginal
gland, m staminate flower, n pistillate flower; o–s C. odontadenius o leaf, p acropetiolar glands, q marginal gland, r staminate flower,
s pistillate flower; t–x C. parodianus t leaf, u basilaminar glands, v marginal gland, w staminate flower, x pistillate flower. Drawn from
a–d R. C. Sodré 3192, e–i W. Mantovani 12713, j–m E. M. F. Chaves 552, n M. R. Del’Arco s.n. (TEPB 1359), o–s H. D. Ferreira 243,
t–x R. C. Sodré 3099.

30. Croton mollis Bentham (1854: 375). Type:—BRAZIL. Amazonas: Manaus, October 1851, R. Spruce 1806
(lectotype: K 000254384!, designated here; isolectotypes: A 00047350!, B n.v., photo at F 0BN005135!, BR
0000008710871!, G00434609!, G 00434610!, G00312858!, GOET 003356!, K 000254385!, LD 1629579!, LE
00003129!, NY 00262929!, OXF 00091967!, P 00634504!, P00634505!, TCD 0007627!). (Fig. 31e–i).
Croton gardnerianus Baillon (1864: 301), syn. nov. Type:—BRAZIL. Ceará: s.loc., August–November 1838, G. Gardner 944 (holotype:
P 00623194!, isotype: A 00257926!).
Croton wittianus Ule (1908: 75), syn. nov. Type: BRAZIL. Amazonas: Manaus, “am Strande bei Ouvidor”, January 1901, E.H.G. Ule 5330
(lectotype: HBG 516405!, designated here; photo at F 0BN005197!; isolectotypes: A 00047351, B 100127735!).
Vernacular name:—velame-da-praia.
Distribution and habitat:—Croton mollis occurs in Brazil (Amazonas, Roraima and Ceará [Fig. 26b]) and
Venezuela. It grows in seasonally flooded areas along riverbanks, near waterfalls, igapó forests and campinaranas, in
white sandy beaches between 30 and 60 m elevation.
Preliminary conservation status:—Croton mollis has an Extent of Occurrence of about 800,000 km2, and so we
categorized its conservation status as Least Concern.
Relationships:—The morphological relationships of C. mollis with C. macrosepalus, C. rhodotrichus, and C.
subincanus are discussed under under C. macrosepalus; and the main characteristics to differentiate them are shown in
Table 4.
Taxonomic notes:—We analyzed around 40 specimens from the northern and northeastern region of Brazil
identified as C. wittianus, C. mollis, and C. gardnerianus, as well as all types and protologues of these species, and we
verified that the specimens were quite variable in the shape of leaf blades (linear, narrowly elliptic, elliptic, lanceolate,
narrowly lanceolate, and oval-elliptic) and indumentum (hirsute, pannose, tomentose or puberulent, but always
composed of stellate trichomes). In contrast to this variation in leaf blade shape and indument, we observed a set of
characteristics with variations that were continuous in the populations and that, therefore, enabled us to treat them as
conspecific, namely: shrubby habit 1–3 m tall; 2–8 cylindrical acropetiolar glands; leaf blades elliptic, lanceolate and
sometimes linear, with entire or serrulate margins, and brochidodromous venation with 8–10 pairs of secondary veins;
congested thyrses 5–16 cm long, without a sterile zone between pistillate and staminate flowers; bracts 2.5–4 mm
long, with or without pyriform glands; stamens with villous filaments and anthers with acute apex; 5-merous pistillate
flowers with lanceolate, elliptic, obovate or oblong, unequal sepals, 2–4-fid stigmas; and oblong capsules with whitish
indument and long pedicels (2.5–5 mm).
Although Croton mollis was described on the basis of a single collection, R. Spruce 1806, we have seen 15
duplicates of this collection from 12 different herbaria. We designated as lectotype K 000254384, since it belonged to
the original Bentham Herbarium and has flowers, buds, and young fruits. For C. wittianus, we chose as its lectotype
the specimen at HBG, where the main collections of Ule are deposited.
Representative specimens examined:—BRAZIL. Amazonas: Manaus, Praia Grande, Rio Negro, 16 July 1981,
W. Mantovani & D.M.S. Rocha 12713 (UEC); 3°2’S, 60°30’W, 9 August 1991, S. Mori & C. Gracie 21980 (INPA,
NY); rio Araras, ilha do Cumaru, 9 May 1973, A. Loureiro et al. 38047 (INPA); Rio Negro, próximo à Praia do Caju,
26 August 1978, L. Coelho & O.P. Monteiro 827 (INPA); between Manaus and Igarape Tarumã, 14 October 1966,
G.T. Prance et al. 1966 (NY); Barcelos, Rio Aracá, imediações da comunidade de Bacuquara, 0°9’16”S, 63°10’41”W,
35–50 m, 16 April 2014, A. Amorim et al. 8606 (RB); junction of rio Aracá & Rio Cuieiras (Jauari), 0°31’30”S,
63°29’30”W, 8 July 1985, G.T. Prance et al. 29416 (INPA, MG, NY, SP); rio Negro, boca do lago Tupé, 28 March
1979, F.H. Schmidt & G. Pabst 9680 (INPA); baixo rio Negro, praia próximo do lago Tupé, 17 December 1958,
W. Rodrigues 721 (INPA); Ponta Negra, 3 January 1961, W. Rodrigues & L. Coelho 2043 (INPA). São Gabriel da
Cachoeira, Rio Negro above Camanaus, 2 November 1971, G.T. Prance et al. 16034 (INPA, NY). Roraima: Caracaraí,
Parque Nacional Serra da Mocidade, rio Capivara, 1°3’33”N, 61°44’48”W, 55 m, 25 March 2012, R.C. Forzza et al.
6974 (RB); Parque Nacional do Viruá, margem do Rio Anauá, 01 October 2012, F.N. Cabral et al. 450 (UEC)
31. Croton mucronifolius Müller Argoviensis (1873: 216). Type:—BRAZIL, Piauí: s.loc., August 1839, G. Gardner
2722 (lectotype: G 00434608!, designated here; isolectotypes: K 000186101!, K 000186102!, OXF 00091966!) (Fig.
30c–e).
Vernacular name:—alcachofrinho, malvinha, pustemeira, velame.

Distribution and habitat:—Croton mucronifolius is restricted to Brazil and occurs in Bahia, Goiás, Maranhão,
Minas Gerais, and Piauí, and it is newly reported here for Mato Grosso and Tocantins (Fig. 32a). It grows in ‘cerrado
típico’ or ‘cerradão’, on sandy soil between 440 and 1170 m elevation.
FIGURE 32. Geographical distribution of some species of Croton sect. Geiseleria in Brazil. a Croton mucronifolius, C. nepetifolius, C.
odontadenius, C. parodianus and C. planaltoanus; b C. pycnadenius, C. ramosissimus C. rhodotrichus, C. sclerocalyx and C. seccoi.

Phenology:—Flowering and fruiting throughout the year, but with abundant flowers and fruits from October to
June.
Preliminary conservation status:—Croton mucronifolius has an estimated Extent of Occurrence of about
800,000 km2 and, therefore, is classified as Least Concern.
Characterization and Relationships:—This species has a subshrubby habit up to 1.3 m tall; very short petioles
1–3 mm long; leaf blades with a slightly cordate base and 2–6 cylindrical glands, serrate margins with or without
irregularly distributed stipitate-capitate glands and actinodromous-brochidodromous venation, with a pair of basal
veins exceeding half the length of the blade, staminate cymules with 3–5 flowers, and pistillate flowers with equal,
lanceolate sepals. The morphological relationships between C. mucronifolius and C. agoensis, which it resembles most
closely, are discussed under C. agoensis.
Representative specimens examined:—BRAZIL. Bahia: BR 349, em direção a Correntina, próximo à fazenda
Entre Rios, lado esquerdo da estrada, 13°29’25.4”S, 44°58’28”W, 776 m, 12 July 2014, A.O. Souza et al. 1221 (UFG);
Mucugê, estrada para Guiné na BA-142 sentido Barra da Estiva, 7 km da estrada de chão, 12°57’54”S, 41°28’26”W,
1024 m, 19 November 2005, D.S. Carneiro-Torres et al. 613 (HUEFS, SP). Goiás: Cavalcante, Serra do Tombador,
GO 241, 74,5 km da cidade de Cavalcante, 13°34’3.2”S, 47°31’26.6”W, 1173 m, 06 August 2016, J.A. Oliveira et al.
144 (UFG). Maranhão: BR-230, próximo a São Domingos de Jaboatão, Benedito Leite, 26 April 1979, P. Martins & E.
Nunes s.n. (EAC 0005943). Mato Grosso: Campo Novo do Parecis, km 805 da BR 364, entre Campo Novo e Tangará
da Serra, ca. 3 km antes da entrada para Diamantino, 14°17’53.1”S, 57°58’10”W, 09 April 2017, R.C. Sodré & A.O.
Souza 3138 (UFG). Minas Gerais: Rio Pardo de Minas, Areião II, 15°26’32”S, 42°26’29”W, 986 m, 06 November
2006, A.C. Sevilha et al. 4768 (CEN). Piauí: estrada Gilbués–Corrente, a 10 km de Gilbués, 12 November 1979, A.
Fernandes & Matos s.n. (EAC 0007223). Tocantins: Ponte Alta do Tocantins, TO-130, entre os municípios Pindorama
do Tocantins e Ponte alta, 10°51’46.6”S, 47°29’44.9”W, 439 m, 09 April 2016, R.C. Sodré et al. 2294 (UFG).
32. Croton nepetifolius Baillon (1864: 344). Type:—BRAZIL. Minas Gerais: s.loc., 1816–1821, A.F.C.P. Saint-Hilaire
cat B2 2115 (lectotype: P 00634517!, designated here; isolectotype: P 00634518!); remaining syntype:—BRAZIL.
Minas Gerais: “mont. près du Penhino”, 1816–1821, A.F.C.P. Saint-Hilaire cat B1 2063 (P 00634519!, P 00634520!,
P 00634521!, P 00634522!) (Figs. 31j–n, 33a–d).
Vernacular name:—canelinha-de-areia, malva-peluda, malva-preta, marmeleiro-preto, marmeleiro-roçarento,

velame.
Distribution and habitat:—Croton nepetifolius is widely distributed in the Caatinga Domain, in Alagoas, Bahia,
Ceará, Minas Gerais, Paraíba, Pernambuco, Piauí, Rio Grande do Norte, and Sergipe (Carneiro-Torres 2009; Silva et
al. 2010) (Fig. 32a). It grows in shrubby or arboreal caatinga usually in rocky outcrops, in sandy soil between 30 and
1,130 m elevation. Armbruster et al. (2007) and Møller-Jørgensen et al. (2014) cited C. nepetifolius for Venezuela
and Bolivia, respectively. However, we analyzed the collections cited by these authors and we conclude that the
collections identified as C. nepetifolius in Venezuela correspond in fact to C. campinarensis (see comments below
on the morphological relationships of theses species). Regarding the collections from Bolivia, they correspond to
C. roborensis, a species synonymized by Møller-Jørgensen et al. (2014) under C. nepetifolius, which although very
similar to Brazilian specimens of C. nepetifolius, we believe that a meticulous study considering these species is
needed to verify if they are conspecific.
Phenology:—Flowering from October to February and fruiting from January to May, but likely to be found with
flowers or fruits throughout the year.
Preliminary conservation status:—It can be categorized as Least Concern because its geographic range is
estimated at roughly 800,000 km2.
Relationships:—Croton nepetifolius mophologically resembles C. campinarensis in its shrubby habit 2–4 m
tall, ovate, oval-oblong or oval-elliptic leaf blades with slightly cordate base and doubly serrate or erose margins, and
4-fid stigmas. However, C. nepetifolius can be easily recognized by having petioles 3 to 4 times shorter than the leaf
blades, sessile patelliform basilaminar glands, thyrses more than 5 cm long, as well as equal pistillate sepals, whereas
C. campinarensis has petioles 4 to 6 times shorter than the leaf blades, stipitate patelliform basilaminar glands, thyrses
1.5–3.5 cm long, and unequal pistillate sepals.
FIGURE 33. Croton nepetifolius a habit, b flowering branch, c thyrse showing staminate flowers and buds, d lower part of a thyrse
showing pistillate flowers. C. odontadenius e habit, f thyrse with pistillate and staminate flowers, g part of thyrse showing staminate
flowers and buds, h pistillate flowers, i capsule.

Representative specimens examined:—BRAZIL. Bahia: Abaíra, estrada Abaíra–Catolés, ca. 300 metros a leste
do Riacho Machado, 13°17’22.5”S, 41°44’42.7”W, 878 m, 17 December 2017, R.C. Sodré et al. 3312 (UFG). Ceará:
Pecém, 16 January 2008, A.M. Miranda et al. 5632 (HST, SP). Minas Gerais: Piauí: Parnaíba, Embrapa Meio Norte,
3°4’31”S, 41°44’29”W, 46 m, 01 February 2016, K.N.C. Castro & J.B.A. Souza 482 (CEN); Caracol, Serra das confusões,
06 December 1980, M.R. Del’Arco & A.G. Fernades s.n. (TEPB 1359); Cocal, Jacaré do Tabuleiro, 16 January 2004,
E.M.F. Chaves & E.M. Sérvio Júnior 552 (TEPB); Brazileira, Fazenda Angelim, 4°7’9.4”S, 41°50’29.5”W, 135 m, 20
January 2001, J.M. Costa 179 (SP, TEPB). Rio Grande do Norte, Coronel João Pessoa, Mata Redonda, Sa. São José,
600 m, 19 July 1991, M.A. Figueiredo et al. 257 (EAC).
33. Croton odontadenius Müller Argoviensis (1873: 227). Type:—BRAZIL. Minas Gerais: “in campis aridis petrosis
prope Lagoa Santa”, 5 October 1863, J.E.B. Warming 1642 (lectotype: C 10011184!, designated by Sodré et al. 2017;
isolectotype: G 00434597!); remaining syntypes:—BRAZIL. Minas Gerais: “in campis aridis petrosis prope Lagoa
Santa”, 5 October 1863, J.E.B. Warming 1641 (C 10011185!) (Figs. 31o–s, 33e–i).
Croton caperoniifolius Müller Argoviensis (1873: 206), syn. nov. Type:—BRAZIL. Minas Gerais: Lagoa Santa, Rio das Velhas, 4 November
1863, J.E.B. Warming 1572 (lectotype: C 10011160!, designated by Caruzo & Cordeiro 2007; isolectotype: G 00434446!); remaining
syntypes:—BRAZIL. Mato Grosso do Sul: “ad Rio Pardo”, s.d., L. Riedel s.n. (LE 00003073!).
Croton sclerocalyx var. lanceolatus Müller Argoviensis (1865: 135). Type:—BRAZIL. Minas Gerais: 1845, J.F. Widgren s.n. (S R-
10551!).
Distribution and habitat:—Croton odontadenius is known from Minas Gerais and Goiás (Sodré et al. 2017) and it is
newly reported here for the Distrito Federal and for Mato Grosso do Sul (Fig. 32a). It occurs in ‘campo limpo’, ‘campo
sujo’, ‘cerrado rupestre’ or sometimes in ‘cerrado típico’, in sandy to clayey soils between 849 to 1,542 m elevation.
Phenology:—Flowering from September to May and fruiting usually from December to June.
Preliminary conservation status:—Croton odontadenius can be categorized as Least Concern because its Extent
of Occurrence was estimated at roughly 450,000 km2, besides having several populations in protected areas such as
Chapada dos Veadeiros National Park and Ecological Reserve of the IBGE.
Relationships:—Croton odontadenius is sometimes mistaken for C. aberrans because of its usually decumbent,
subshrubby habit, commonly hirsute stems and leaves; oblong, elliptic or oval-lanceolate, sometimes linear, concolorous,
dark green leaf blades, congested inflorescences without a distinct sterile zone between pistillate and staminate flowers,
staminate calyx united up to half its length, and pistillate flowers with 6–8 sepals. However, in C. odontadenius the
acropetiolar and marginal glands are largely cylindrical or shortly stipitate-patelliform, with a length-to-diameter ratio
of 0.8–2 (vs. narrowly cylindrical, length-to-diameter ratios of 8–10 in C. aberrans), the inflorescences 2–4 cm long
(vs. less than 1.5 cm long), the stigmas are multifid (vs. 2-fid), and the capsules measure 5–5.8 × 5.3–6 mm (vs. 6.5–7
× 7–7.3 mm).
Taxonomic notes:—When he described Croton caperoniifolius, Müller Argoviensis (1873) related it to C.
antisyphiliticus citing that the latter was differentiated by inflorescences with more flowers and muricate ovaries.
Recently, Caruzo & Cordeiro (2007) proposed a lectotype for C. caperoniifolius and synonymized it under C.
antisyphiliticus without any further explanation. However, we do not agree with this synonymization, since we believe
that C. antisyphiliticus can be distinguished from C. caperoniifolius by its rough-textured leaves with stellate-porrect
trichomes, lax thyrses, 5-merous pistillate flowers with ovate and not accrescent sepals and staminate flowers with
calyx shortly joined at the base; whereas in C. caperoniifolius the hirsute leaves are not rough-scabrous, the thyrses
are congested, the pistillate flowers are 6–8-merous with oblong, lanceolate, linear or oblanceolate sepals, and the
staminate flowers have their calyx joined up to half their length. However, when we compare C. caperoniifolis
with C. odontadenius, we see that their diagnostic characters are identical, so we consider them synonymous. As C.
odontadenius and C. caperoniifolius have equal priority, we chose to keep the first one because it is more commonly
used in Brazilian herbaria.
Representative specimens examined:—BRAZIL. Distrito Federal: Brasília, Reserva Ecológica do IBGE, em
frente à nascente do córrego Escondido, 18 December 1986, M. Aparecida da Silva & D. Alvarenga 236 (IBGE); APA
Gama–Cabeça de Veado, Região Administrativa do Núcleo Bandeirante, lagoa do Córrego do Cedro, 15°53’46”S,
47°56’36”W, 990 m, 09 October 2002, M.L. Fonseca & D. Alvarenga 3637 (IBGE). Goiás: Anápolis, 15 October
1956, L.B. Smith & Macedo 4707 (SP, US); Mineiros, Parque Nacional das Emas, 14 March 1990, H. D. Ferreira
2274 (UFG). Serranópolis, 25 March 1984, H.D. Ferreira 243 (UFG). Mato Grosso do Sul: Corumbá, Morro Urucum,
19°10’47”S, 57°35’41”W, 849 m, 3 December 2012, J.P. Souza et al. 11718 (UFG). Minas Gerais: Ritápolis, trevo
para Ritápolis, 22 October 2013, M. Sobral & D.F. Lima 15558 (HUFSJ, SP)
34. Croton parodianus Croizat (1944: 452). Type:—BRAZIL. Maranhão: Island of São Luiz, February–March 1939,
R.L. Fróes 11775 (holotype: A 00047371; isotypes: G 00434656!, K 000254441!, NY, S S-R-10555!, U 0008010!)
(Figs. 31t–x, 34a–e).
Croton strobiliformis Secco (2009: 249), syn. nov. Type:—BRAZIL. Pará: Tucuruí, margem direita do Rio Tocantins, BR-263, km 16,
campina de solo arenoso, 30 January 1980, P.L.B Lisboa et al. 1286 (holotype: MG 071358; isotypes: NY, UB!).
Vernacular name:—velame.
Distribution and habitat:—Croton parodianus was reported from Pará, Maranhão, and Roraima by Secco
(2008), and it is newly reported here for Amazonas, Mato Grosso, and Rondônia (Fig. 32a). It grows in campinarana
or less commonly in “terra firme” forest, usually associated with rocky outcrops on hillsides, in sandy soils, between
Phenology:—Collected with flowers in November and with flowers and fruits from January to April.
Preliminary conservation status:—Species with Extent of Occurrence estimated at 2,000,000 km2 and, therefore,
considered as Least Concern.
Relationships:—Croton parodianus is morphologically related to C. sincorensis by its shrubby habit more than
2 m tall, petiole 1–3(–8) cm long and leaf blades 5.2–15 × 3.1–8.5 cm, stipitate glands in the sinuses of leaf margins,
and long thyrses (4.8–10.5 cm). However, C. parodianus differs by its cordate leaf base with stipitate patelliform
glands, inflorescences with 6–12 pistillate flowers with 5 sepals and 2-fid stigmas. Croton sincorensis has cylindrical
acropetiolar glands, leaf blades with a rounded, obtuse or truncate base, thyrses with 1–4 pistillate flowers, these with
6–10 sepals and multifid stigmas.
Taxonomic notes:—Croton strobiliformis was described on the basis of a single collection (P.L.B. Lisboa et
al. 1286) and although it was related to C. hirtus by Secco (2009), its diagnostic characteristics (e.g., palmatinerved
leaves with stipitate glands at the base, slightly rough on the abaxial surface, congested inflorescences, internally
villous staminate flower petals, and capsules with dense indumentum of stellate trichomes) fall within the range of
morphological variation of C. parodianus.
Representative specimens examined:—BRAZIL. Amazonas: Humaitá, estrada para Velho-Humaitá (BR-319,
km 675), 7°31’S, 63°10’W, 5 April 1976, M.R. Estrela & W. Bellusci 94–5476 (SP). Mato Grosso: Aripuanã, Região da
Serra Morena, na estrada de terra que leva ao Vale dos Sonhos, ca. 5 km da MT-208 (Juruena-Aripuanã), 10°22’57.3”S,
58°52’27.7”W, 289 m, 08 April 2017, R.C. Sodré & A.O. Souza 3125 (UFG); Guarantã do Norte, Serra do Cachimbo, a
direita do km 1116 da BR 163, sentido Novo Progresso-PA, ca. 8 km antes da divisa MT/PA, 9°35’5.1”S, 54°53’17.4”W,
458 m, 04 April 2017, R.C. Sodré & A.O. Souza 3099 (UFG). Pará: Itaituba, estrada Santarém-Cuiabá, BR-163, km
171, próximo a divisão Pará-MT, 9°35’S, 54°35’W, 22 April 1983, I.L. Amaral et al. 872 (INPA); Serra do Cachimbo,
BR 163, Cuiabá-Santarém Highway, km 768,5, 10 km S of Mato Grosso, 490 m, 12 November 1977, G.T. Prance et
al. 25277 (MG, NY, SP, UEC); Acará, Vila Cruzeirinho, ramal do areial, km 28, 2°0’58”S, 48°19’9”W, 19 February
2014, L.M.M. Carreira & F.C.A. Lucas 3305 (MFS); Belém, rio Guajará, 14 May 1954, G.A. Black 54-16182 (IAN,
UB). Rondônia: Porto Velho, Serra dos Parecis, estrada Guajará-Mirim–Abunã, km 11, 03 February 1983, L. Carreira
et al. 503 (IAN).
35. Croton planaltoanus M.J.Silva & Sodré (in Silva et al. 2014: 216). Type:—BRAZIL. Goiás: Alto Paraíso de Goiás,
cerca de 11 km da cidade em direção a Teresina de Goiás após o Morro do Japonês, 14°02’00”S, 47°31’40.3”W, 1540
m, 24 November 2012, M.J. Silva & J.P. Santos 4577 (holotype: UFG 60551!, isotype: UB!) (Figs. 34f–k, 35a–d).
Distribution and habitat:—Croton planaltoanus is apparently endemic to the state of Goiás, known from Serra
Dourada (Mossâmedes and Goiás municipalities) and from the Chapada dos Veadeiros (Água Fria de Goiás, Alto
Paraíso de Goiás and Cavalcante municipalities; Silva et al. 2014; Sodré et al. 2017) (Fig. 32a). It occurs in campo
limpo, campo sujo, and cerrado típico, in gravel or sandy soil between 900 and 1540 m elevation.
Phenology:—Flowering and fruiting throughout the rainy season from September to April.
Preliminary conservation status:—Silva et al. (2014) categorized the conservation status of Croton planaltoanus
as Near Threatened due to its few collections from only two localities, Serra Dourada and Chapada dos Veadeiros, and
its Extent of Occurence around 20,000 km2.

FIGURE 34. Croton parodianus a habit, b thyrse showing pistillate flowers, young fruits and staminate flowers and buds, c staminate
flowers, d pistillate flowers, e capsules. C. planaltoanus f habit, g vegetative branch, h distal portion of an inflorescence with staminate
flowers and buds, i staminate flowers and buds, j pistillate flowers and staminate buds, k capsules.
FIGURE 35. a–d Croton planaltoanus a leaf, b marginal gland, c staminate flower, d pistillate flower; e–i C. ramosissimus e leaf, f
basilaminar glands, g marginal gland, h staminate flower, i pistillate flower; j–m rhodotrichus j leaf, k acropetiolar glands, l staminate
flower, m pistillate flower; n–r C. sclerocalyx n leaf, o basilaminar glands, p marginal gland, q staminate flower, r pistillate flower; s–w
C. sincorensis s leaf, t acropetiolar glands, u marginal gland, v staminate flower, w pistillate flower. Drawn from a–d R. C. Sodré 1485,
e–i R. C. Sodré 2579, j–m R. C. Sodré 2232, n–r R. C. Sodré 2984, s–w H. Q. Boudet Fernandes 1689.

Relationships:—Although C. planaltoanus was compared to C. inaequilobus when it was described (Silva et al.
2014), we believe that it is more related to C. abaitensis because of its subshrubby habit, short petiole (0.4–1.3 cm
long), long inflorescences (1.5–9.8 cm) with 4–10 pistillate flowers distributed along the basal 1/3 of the axis, linear-
lanceolate staminate bracts, and pistillate flowers with equal or unequal sepals. However, C. planaltoanus is easily
differentiated because it is a few-branched subshrub up to 60 cm tall with usually eglandular petioles and usually
elliptic, concolorous leaf blades, with an acute or obtuse base and brochidodromous venation, with the lower secondary
veins never exceeding half the length of the blades. Croton abaitensis, on the other hand, is a much-branched subshrub
up to 1.2 m tall, with usually with glandular petioles and ovate, discolorous leaf blades with an obtuse or slightly
cordate base, actinodromous-brochidodromous venation, and the pair of pair of veins exceeding half the length of the
blades.
Representative specimens examined:—BRAZIL. Goiás: Água Fria de Goiás, a direita do km 61 da GO-118,
sentido São Gabriel a São João d’Aliança, 14°59’23.4”S, 47°36’16.4”W, 1119 m, 30 October 2014, R.C. Sodré et al.
1466 (UFG), 1469 (UFG). Alto Paraíso de Goiás, ca. 12 km N.W. de Veadeiros, estrada para Cavalcante, 14°S, 47°W,
1200 m, 21 October 1965, H.S. Irwin et al. 9439 (UB); estrada para Colinas do Sul a 14 km de Alto Paraíso de Goiás,
24 November 1994, F.C.A. Oliveira 224 (IBGE); a noroeste de Alto Paraíso, ca. 1 km a direita da entrada da cidade,
14°7’59”S, 47°31’47.9”W, 1278 m, 31 October 2014, R.C. Sodré et al. 1485 (UFG). Cavalcante, estrada para povoado
Kalunga, Engenho II, 13°42’1.4”S, 47°27’50.3”W, 898 m, 05. April 2013, R.C. Sodré et al. 653 (UFG). Goiás, Serra
Dourada, 08 September 1976, P. Gibbs et al. 2799 (UEC). Mossâmedes, Serra Dourada, próximo ao portão da Reserva
Ecológica da Serra Dourada, 16°4’45”S, 50°11’21”W, 991 m, 19 September 2003, R. Mello-Silva et al. 2224 (SP).
36. Croton pycnadenius Müller Argoviensis (1873: 216). Type:—BRAZIL. Tocantins: “inter Funil et S. João”, s.d.,
W.J. Burchell 8988 (lectotype: BR 8761354!, designated here; isolectotypes: A 00257984!, G 00434631!, K 00186103!)
(Fig. 36a–f).
Croton pardinus Müller Argoviensis (1873: 217), syn. nov. Type:—BRAZIL. Mato Grosso do Sul: “in campis siccis secus Rio Pardo”,
s.d., L. Riedel 467 (lectotype: LE 00018220!, designated here; isolectotypes: G 00434657! [photo F 24513!], LE 00018221!).
Distribution and habitat:—Croton pycnadenius occurs in Bahia, Goiás, Maranhão, Mato Grosso, Mato Grosso do
Sul, Minas Gerais, Pará, and Tocantins (Fig. 32b). It grows in ‘cerrado típico’ and ‘campo sujo’ in sandy soil between
Phenology:—Flowering from October to May and fruiting from December to August.
Preliminary conservation status:—The species can be categorized as Least Concern, because of its Extent of
Occurrence estimated at 1,100,000 km2.
Relationships:—Croton pycnadenius is morphologically most similar to C. mucronifolius, but differs from it
by being a decumbent plant with largely ovate or orbicular leaf blades with a cordate base and eglandular margins,
staminate cymules with 2 flowers, and greenish capsules. In contrast, C. mucronifolius is an erect plant with oval-
lanceolate, oblong or elliptic leaf blades with an obtuse or slightly cordate base and glandular margins; it also has
staminate cymules with 3–5 flowers and yellowish or withish capsules. Croton pycandenius is also similar to C.
abaitensis and C. agoensis, but is distinguished by the characters listed in the comments of those species.
Taxonomic notes:—Croton pycnadenius was described for the former Goyaz Province between Funil and São
João, both of which now corresponding to Tocantins state. We designated as lectotype BR 8761354, because it contains
notes in Müller’s hand. Croton pardinus was published in the same work based on L. Riedel 467, which was erroneously
interpreted by Müller Argoviensis (1873) as coming from the Província of São Paulo. It has since been clarified by
Urban (1906) that Rio Pardo is located in the state of Mato Grosso do Sul. This collection is represented by a specimen
at G and two at LE; we chose the LE sheet annotated by Müller Argoviensis as the lectotype since it is the ones best
reflecting the original description.
Representative specimens examined:—BRAZIL. Goiás: Guarani de Goiás, km 303 da BR-020, 15 km N do
entroncamento de Posse, 950 m, 16 August 1990, T.B. Cavalcanti et al. 747 (CEN). Maranhão: Carolina, BR 010,
transamazônica, margem esquerda da rodovia, lugar Pedra Caída, 13 April 1983, M.F.F. Silva et al. 1056 (INPA). Pará:
Serra do Cachimbo, along BR 163, Cuiabá-Santarém road, km 838–839, 490 m, 18 February 1877, J.H. Kirkbride Jr.
& E. Lleras 2905 (INPA); ibid., km 823, 7 November 1977, G.T. Prance et al. P25004 (NY, MG). Tocantins: Mateiros,
entre Ponte Alta e Mateiros, 10°38’19.3”S, 47°17’52.9”W, 396 m, 18 December 2014, L.L.C. Antunes et al. 1299
(UFG), 1303 (UFG).
FIGURE 36. Croton pycnadenius a habit, b vegetative branch, c flowering branch, d thyrse showing pistillate flowers and staminate
flowers and buds, e pistillate flowers, f capsule and leaf base. C. ramosissimus g habit, h staminate flowers, i pistillate flowers, j fruiting
branch, k capsules.

37. Croton ramosissimus Sodré & M.J.Silva (in Sodré & Silva 2017a: 128). Type:—BRAZIL. Goiás: Corumbá
de Goiás, hillside opposite to the belvedere of Salto de Corumbá, to the left of BR-414 road, towards Cocalzinho,
15°50’53.2”S 48°46’07”W, 1050 m, 10 October 2016, R.C. Sodré, R.D.S. Nabarro & F. Roa 2668 (holotype: UFG!;
isotypes: CEN!, K!, MICH!, NY!) (Figs. 35e–i, 36g–k).
Vernacular name:—alcacuz.
Distribution and habitat:—This species is quite broadly distributed in the Distrito Federal and in the state of
Goiás (Fig. 32b), where it is found in cerrado típico, cerrado ralo or campo sujo, in gravelly, sandy or clayey soils
between 560 and 1220 m elevation.
Phenology:—Collected with flowers and/or fruits from September to May, but its flowers are more frequent from
September to December and the fruits abundant from November to February.
Preliminary conservation status:—Sodré & Silva (2017a) categorized the conservation status of Croton
ramosissimus as Least Concern because its geographic range is greater than 20,000 km2 (here estimated at ca. 80,000
km2), besides having populations found in protected areas.
Relationships:—Croton spica is the species most morphologically similar to C. ramosissimus by virtue of
the oblong, elliptic or lanceolate leaf blades with brochidodromous venation and tomentose indument of stipitate,
multiradiate trichomes on the abaxial surface, racemose inflorescences, usually linear-lanceolate bracts, and accrescent
calyces in fruit. Nevertheless, C. ramosissimus is distinguished by being a much-branched shrub or subshrub 1–2 m
tall, growing with a monopodial aspect, with prominent leaf venation on the abaxial surface, lax inflorescences, bracts
1.1–2.7 mm long, villous staminal filaments, pistillate flowers with equal or subequal, ovate or oval-lanceolate sepals,
and oblongoid capsules and seeds, the latter without spots. On the other hand, C. spica is characterized by its few-
branched subshrubby habit up to 1.2 m tall, sympodial growth habit, leaf blades with impressed veins on the abaxial
surface, congested inflorescences, bracts 2.3–6 mm long, glabrous staminal filaments, pistillate flowers with unequal
sepals, the largest of which are obovate, elliptic or orbicular, and globose capsules and maculate, ovoid seeds.
Representative specimens examined:—BRAZIL. Distrito Federal: Brasília, Cachoeira do Pipiripau, 12 km de
Planaltina Leste, 31 December 1974, E.P. Heringer 14274 (IBGE, NY, K, US, UB, CEN); Lago Sul, Parque Ecológico
do Rasgado, próximo ao Setor de Mansões Don Bosco, 15°50’12”S, 47°29’04”W, 1062 m, 28 October 2003, M.
Aparecida da Silva & D. Alvarenga 5584 (IBGE, SP); Recanto das Emas, estrada vicinal a direita do km 13 da BR
060, sentido Brasília, após o posto da Polícia Rodoviária Federal e antes do trevo para Santo Antônio do Descoberto,
15°55’53” S, 48°9’51” W, 1,066 m, 12 December 2016, R.C. Sodré 2892 (UFG), 2893 (UFG), 2894 (UFG). Goiás:
Caldas Novas, estrada de acesso a UHE-Corumbá, 2,7 km antes do portão de acesso à obra, 18°00’S, 48°34’W, 12
February 1993, T.A.B. Dias et al. 413 (CEN). Cristalina, RPPN Linda Serra dos Topazios, 16°45’S, 47°40’W, 27
February 1998, C. Proença, D. Araújo, M.A. Rodrigues 1940 (UB); BR-040, km 89, sentido Cristalina-Luziânia,
logo atrás de alguns eucaliptos e uma casa abandonada, 16°44’9.7”S, 47°37’9”W, 1219 m, 08 September 2014, L.S.
Inocencio & A.O. Souza 134 (UFG), 136 (UFG), 140 (UFG). Mossâmedes, Serra Dourada, divisa dos municípios de
Mossâmedes ao Sul e Goiás ao Norte, área da UFG, 08 November 1969, J.A. Rizzo 4556 (UFG, RB).
38. Croton rhodotrichus Sodré & Silva (2017b: 338). Type:—BRAZIL: Mato Grosso: Cuiabá, north side of the
National Park Chapada dos Guimarães, on the road of Quilombo, ca. 10 km from the road MT-020, 15°16’51”S,
55°52’13”W, 680 m, 06 March 2015, M.J. Silva, R.C. Sodré & G.H. Silva 6518 (holotype: UFG 60353!; isotype:
UFMT 43524!) (Figs 35j–m, 37a–f).
Distribution and habitat:—Croton rhodotrichus is known only from Mato Grosso state, in the municipalities of
Chapada dos Guimarães, Cuiabá and Ribeirão Cascalheira (Sodré & Silva 2017b) (Fig. 32b). It occurs in cerrado
típico, cerrado denso or cerrado ralo, in sandy soil between 300 and 680 m elevation.
Phenology:—Collected with flowers from December to March and with fruits only in March.
Preliminary conservation status:—The species is known from only two localities, in the Chapada dos
Guimarães National Park (CGNP) in estimated area at about 40 km2 and by a single collection in the municipality of
Ribeirão Cascalheira approximately 500 km away from the CGNP; we consider its conservation status as Critically
Endangered.
Relationships:—Croton rhodotrichus was related with C. glandulosus and C. hirtus when described by Sodré
& Silva (2017b). However, we compare C. rhodotrichus with C. macrosepalus, C. mollis and C. subincanus in Table
4, because we believe that they can be confused by their usually elliptic and discolorous leaf blades with entire or
subentire margin, whitish, tomentose indument on the adaxial surface and brochidodromous venation with 4–8(–10)
pairs of secondary veins impressed on both surfaces and bracts with sessile glands.
FIGURE 37. Croton rhodotrichus a habit, b flowering branch, c staminate flowers, d pistillate flowers, e inflorescence showing pistillate
flowers and staminate bracts persistent even after flowers fall off, f nearly mature capsule on the left and dehisced capsule with columella
on the right. C. sclerocalyx g habit, h fertile branch, i thyrse with pistillate flowers and staminate flowers and buds, j staminate flowers and
buds, k dehisced capsule, with conspicuous persistent sepals.

Representative specimens examined:—BRAZIL. Mato Grosso: Chapada dos Guimarães, porção norte do Parque
Nacional da Chapada dos Guimarães, na estrada do Quilombo, ca. 8 km da MT 020, 15°17’24.4”S, 55°51’18.4”W,
666 m, 26 March 2016, R.C. Sodré et al. 2232 (UFG), 2233 (UFG), 2234 (UFG); porção norte do Parque Nacional
da Chapada dos Guimarães, ca. 1 km a direita do km 8 da estrada do Quilombo a partir da MT 020, 15°16’57.4”S,
55°51’35.1”W, 26 March 2016, R.C. Sodré et al. 2235 (UFG), 2236 (UFG). Cuiabá, porção norte do Parque Nacional
da Chapada dos Guimarães, na estrada do Quilombo, ca. 10 km da MT 020, 15°16’51”S, 55°52’13”W, 680 m, 06
March 2015, R.C. Sodré et al. 1712 (UFG), 1721 (UFG); ibid., 15°16’47.7”S, 55°52’8.4”W, 611 m, 06 March 2015,
M.J. Silva et al. 6515 (UFG), 6516 (UFG), 6517 (UFG), 6520 (UFG); porção norte do Parque Nacional da Chapada
dos Guimarães, na estrada do Quilombo, ca. 7 km da MT 020, 15°17’46.4”S, 55°50’51.2”W, 677 m, 26 March 2016,
R.C. Sodré et al. 2230 (UFG), 2231 (UFG); porção norte do Parque Nacional da Chapada dos Guimarães, na estrada do
Quilombo, ca. 8,5 km da MT 020, 15°17’22.2”S, 55°51’22.2”W, 669 m, 26 March 2016, fl., fr., R.C. Sodré et al. 2239
(UFG); Parque Nacional da Chapada dos Guimarães, ca. 15 metros a direita do início da trilha que leva a cacheira Véu
de Noiva, 15°24’17”S, 55°49’48”W, 680 m, 07 March 2015, R.C. Sodré et al. 1739 (UFG); ibid., 26 March 2016, R.C.
Sodré et al. 2248 (UFG); 2249 (UFG); 2250 (UFG); 2251 (UFG); 2252 (UFG); 2253 (UFG); 2254 (UFG). Ribeirão
Cascalheira, between km. 256, Xavantina – Cachimbo road, and Base camp., 12°49’S, 51°46’W, 23 December 1967,
D. Philcox et al. 3707 (NY, RB, UB).
39. Croton sclerocalyx (Didr.) Müller Argoviensis (1865: 134).—Podostachys sclerocalyx (Didrichsen 1857: 134).—
Croton sclerocalyx var. genuinus Müller Argoviensis (1865: 134), nom. inval. Type:—BRAZIL. São Paulo: “campis
St. Carlos”, 1834, P.W. Lund s.n. (lectotype: C 10011194!, designated here; isolectotypes: C 10011195!, G 00312480!)
(Figs. 35n–r, 37g–k).
Croton mentiens (S.Moore) Pax (1897: 211).—Heterocroton mentiens Moore (1895: 461). Type:—BRAZIL. Mato Grosso: Santa Cruz,
1891–1892, S.L. Moore s.n. (holotype: BM 000939427!).
Croton sclerocalyx var. hirsutissimus Müller Argoviensis (1865: 134). Type:—BRAZIL. São Paulo: s.loc., s.d., L. Riedel 1626 (lectotype:
LE 00018222!, designated here; isolectotypes: GH 00277216!, LE 00018223!, LE 00018224!); remaining syntypes:—BRAZIL.
Minas Gerais: 1840, P. Claussen 316 (BR 0000008761132!, BR 0000008761163!); s.loc., s.d., C.F.P. Martius s.n. (not found).
Croton sclerocalyx var. rufidulus (Baill.) Müller Argoviensis (1866: 662).—Croton rufidulus Baillon (1864: 347). Type:—BRAZIL.
Minas Gerais: 1838 or 1839, P. Claussen 776 (lectotype: P 00493385!, designated here; isolectotypes: P 00493386!, G 00434695!);
remaining syntypes:—BRAZIL. Minas Gerais: s.loc., 1816–1821, A.F.C.P. Saint-Hilaire cat C1 277 (syntypes: A 00257990!, P
00493387!, P 00493388!).
Croton sclerocalyx f. luxurians Müller Argoviensis (1873: 226). Type:—BRAZIL. Minas Gerais: Lagoa Santa, s.d., J.E.B. Warming 1640
(holotype: G 00434697!)
Croton sclerocalyx var. intermedius Müller Argoviensis (1873: 226). Type:—BRAZIL. Minas Gerais: Caldas, 1869, A.F. Regnell III 1088
(lectotype: BR 0000008761170!, designated here; isolectotypes: US 00109681!, US 00997753!, US 01014155!, A 00257991!);
remaining syntypes:—BRAZIL. Minas Gerais: Caldas, 29 December 1854, J.F. Widgren 348 (BR 0000008761149!, S 07-12774!, S
07-12775!, S 07-12776!)
Croton sclerocalyx var. hispidus Müller Argoviensis (1873: 226). Type:—BRAZIL. São Paulo: Jundiahy, G.H. Langsdorff s.n. (lectotype:
LE 00018225!, designated here; isolectotype: G 00434694!).
Distribution and habitat:—This species is found in Brazil and Paraguay (Caruzo & Cordeiro 2007). In Brazil it was
reported from the Federal District and the states of Goiás, Mato Grosso, Minas Gerais, and São Paulo by Caruzo &
Cordeiro (2007), and to these we are adding here reports for Mato Grosso do Sul and Tocantins (Fig. 32b). It inhabits
in cerrado s.str., Cerradão, or campo sujo, on sandy or clayer soils, between 278 and 1150 m of elevation.
Phenology:—Flowering and fruiting throughout the year, although flowers and fruits are more common in the
rainy season, from October to April.
Preliminary conservation status:—Species with conservation interpreted here as Least Concern, due to it wide
occurrence and its somewhat weedy nature and efficient colonization of different types of vegetation.
Relationships:—Croton sclerocalyx is very similar to C. sipaliwinensis by their ovate or oval-lanceolate leaf
blades with double-serrated margins and semi-craspedodromous venation that is 3–5-nerved at the base and prominent
on the abaxial surface, thryses with discontinuous flowers, 6- or 7-merous pistillate flowers, usually 4-fid stigmas,
staminate flowers with pedicels 0.3–0.7 mm long and calyx usually united in 1/2 to 1/4 of the length, and capsules with
accrescent sepals. However, C. sclerocalyx differs by the branches, leaves, inflorescence axis, and pistillate flowers
covered by a tomentose or hirsute indument of simple, 2–7(–14)-radiate, or stellate-porrect trichomes (vs. glabrescent
or pubescent indument of simple or stellate trichomes in C. sipaliwinensis), shortly stipitate patelliform basilaminar
and marginal glands (vs. obconic acropetiolar and marginal glands), pistillate flowers with fleshy, obovate sepals with
deeply serrated margins (vs. membranaceous, lanceolate or linear sepals with entire margin), glabrous or pubescent
ovary (vs. always glabrous) and androecium with 6–11 stamens (vs. always 5).
Taxonomic notes:—Caruzo & Cordeiro (2007) cited Lund s.n. at C as the holotype of C. sclerocalyx. However,
there are two specimens of this collection at C and one in G with the same locality “campis St. Carlos”., We selected
C 10011194 as the lectotype, since it contains illustrations and comments made by Didrichsen (1857) that probably
supported the initial description of the species, under the genus Podostachys.
Similar to Govaerts et al. (2000), we also consider all the varieties described for C. sclerocalyx as synonyms of
the species, except for C. sclerocalyx var. pubescens, which has been synonymized here under C. sipaliwinensis. We
propose lectotypes for C. sclerocalyx var. hirsutissimus and C. sclerocalyx var. hispidus based on specimens at LE
with annotations in the hand of Müller Argoviensis; while for C. sclerocalyx var. rufidulus and C. sclerocalyx var.
intermedius we chose Claussen s.n. 776 (P 00493385) and Regnell III 1088 (BR 8761170), respectively, because they
are in good condition and reflect the original description of the varieties.
Representative specimens examined:—BRAZIL. Distrito Federal: Brasília, Bacia do Rio São Bartolomeu,
adjacências do córrego Papuda, 1 April 1981, E.P. Heringer et al. 6695 (IBGE). Goiás: Bela Vista de Goiás, margem
direita da GO 020, entre os municípios de Bela Vista de Goiás e Cristianópolis, à aproximadamente 15 km desta última,
17°1’12.7”S, 48°57’37”W, 800 m, 23 January 2014, R.C. Sodré et al. 1089 (UFG); 1090 (UFG). Formosa, Serra do
Pinduca, margem direita do km 57 da BR 020, sentido Formosa para Alvorada do Norte, ca. 5 km antes do povoado
JK, 15°16’10.8”S, 47°9’26.9”W, 805 m, 15 December 2016, R.C. Sodré & C.C. Oliveira 2984 (UFG). Mato Grosso:
Barra do Garça, margem direita da BR-070, ca. 20 km após o entroncamento com a BR-158, sentido General Carneiro,
15°48’51.5”S 52°28’37.5”W, 358 m, 25 March 2016, R.C. Sodré et al. 2208 (UFG). Diamantino, imediações do Posto
Gil, ca. 2 km a direita da BR 364, sentido Nobres a Diamantino, em direção à fazenda Pedra Branca, 14°34’2.3”S
56°13’43”W, 429 m, 03 December 2016, R.C. Sodré & N.A.G. Escobar 2846 (UFG). Nossa Senhora do Livramento,
Fazenda Coqueiro, ca. 100 metros do rio Jauquara, 15°49’08.5”S 57°11’55.6”W, 05 December 2016, R.C. Sodré &
N.A.G. Escobar 2871 (UFG). São Felix do Araguaia, estrada entre vila de Pontinópolis e a Serra dos Magalhães,
11°33’39.4”S, 51°13’0.7”W, 21 March 1997, V.C. Souza et al. 14733 (UFMT). Mato Grosso do Sul: Campo Grande,
RPPN da Universidade Federal do Mato Grosso do Sul, 20°30’26.1”S 54°37’2.5”W, 02 September 2016, R.C. Sodré
et al. 2578 (UFG). Minas Gerais: Araxá, BR 262, ca. 2 km de Araxá, saída para Belo Horizonte, 19°32’54.6”S,
46°56’25.6”W, 968 m, 07 July 2016, R.C. Sodré et al. 2528 (UFG). Belo Horizonte, Serra do Taquaril, 15 January
1934, M. Barreto 2689 (BHCB). Francisco Sá, estrada de Brejo das Almas, 9 November 1938, F. Markgraf et al. 3187
(BHCB). Parque Nacional Grande Sertão Veredas, 15°24’19”S, 45°55’5”W, R. Rodrigues da Silva et al. 215 (IBGE).
São Paulo: Araraquara, 20 November 1888, A. Loefgren 1027 (SP). Tocantins: Palmas, Fazenda São João, 10 January
2006, M. Sobral & J. Larocca 10394 (BHCB).
40. Croton seccoi Sodré & M.J.Silva, sp. nov. (Fig. 38)
Similar to Croton parodianus Croizat in its ovate leaf blades and flowers in inflorescences s up to 9 cm long, but C. seccoi differs in the
yellow-ferrugineous indument of branches, leaves and inflorescences (vs. whitish in C. parodianus), rounded, obtuse, truncate or
assymetrical leaf base (vs. cordate), serrate or serrulate leaf margin (vs. crenate-serrate), staminate bracts 1.4–1.6 mm long (vs. 2–2.3
mm), oval-lanceolate, oblong or lanceolate pistillate sepals (vs. usually obovate), and maculate seeds (vs. not maculate).
Type:—BRAZIL. Piauí: Piracuruca, Parque Nacional das Sete Cidades, (5a cidade), 4°6’12”S, 41°41’12”W, 174 m, 9 February 2007, fl.,
fr., D.S. Carneiro-Torres, M.M. Silva-Castro & D. Cardoso 836 (holotype: HUEFS 116972!; isotype: HUESB 4191!).
Subshrub or shrub 0.5–2.3 m tall, erect, monoecious, densely branched; yellowish and ferrugineous stems and
branches, with stellate-porrect, shortly stipitate trichomes with 11–16 lateral rays; latex not seen. Entire leaves,
spirally alternate along the stems and subopposite or subverticillate in the dichotomy or trichotomy of the branches
or immediately below the inflorescences; petioles 0.6–2.5 cm long; blades 2.6–7.2 × 1.2–3.9 cm, ovate or sometimes
oval-lanceolate, with obtuse, rounded, truncate or asymmetrical base with 2(–4) shortly stipitate or sessile, patelliform
glands 0.6–1.1 mm diam on the abaxial surface, acute or acuminate apex, serrate or serrulate margin, with subsessile or
stipitate patelliform glands 0.2–0.3 mm diam in the sinuses, membranaceous, discolorous, green-ferrugineous, adaxial
surface rough with subsessile, stellate-porrect trichomes with 9–11 lateral rays and a shortly adpressed porrect ray and
yellowish to whitish, tomentose indument on abaxial surface of stipitate, stellate-porrect trichomes with 12–15 lateral
rays and discrete porrect ray; brochidodromous venation, with 3–6 pairs of secondary veins, primary and secondary
veins usually prominent on the abaxial surface; stipules 0.6–3.3 × 0.3–0.9 mm, linear, lanceolate, narrowly triangular

FIGURE 38. Croton seccoi: a fertile branch. b detail of the marginal gland. c. stipule. d leaf blade. e detail of the stipitate basilaminar
glands. f. detail of the sessile basilaminar glands. g staminate bract. h detail of the staminate bract base showing glands. i staminate flower.
j ventral surface of the staminate petals. k segment disk of the staminate flower. l. stamen. m pistillate flower. n. stellate-porrect trichome
on the pistillate sepal. o. ventral surface of the pistillate sepal. p detail of the pistillate petal. q gynoecium. r capsule with accrescent sepals.
s ventral side of the seed. t dorsal side of the seed. Drawn from the holotype.
or ovate with 4–8 ovoid and sessile glands at the base, covered with stellate-porrect trichomes, densely pubescent
on the abaxial surface and with few trichomes on the adaxial surface. Inflorescences 1–9.2 cm long, terminal or in
dichotomies of branches, bisexual, with (2–)5–9 pistillate flowers at the base of the inflorescence axis and numerous
staminate flowers continuous along the inflorescence axis; staminate bracts 1.4–1.6 × 0.7–0.8 mm, oval-elliptic with
4 ovoid or pyriform sessile glands at the base, concave, glabrous or glabrescent on the adaxial surface, abaxial surface
convex with stellate-porrect trichomes; pistillate bracts 1.3–1.5 × 0.6–0.7 mm, oval-lanceolate. Staminate flowers
4.8–5.2 mm long, yellowish, actinomorphic; pedicels 1.7–1.9 mm long; calyx 5-lobed, lobes 1.8–2.1 × 1.1–1.3 mm,
equal, oval-oblong or ovate, with obtuse apex, slightly united at the base, ventrally glabrous, dorsally stellate-porrect;
petals 2–2.5 × 0.8–1 mm, elliptic, with acute or obtuse apex, villous on both surfaces, ciliate on the margins, the
longest cilia at the base; stamens 11, 3 mm long, with glabrous filaments; disk with 5 oblong yellowish segments;
receptacle villous. Pistillate flowers 4.5–6 mm long; pedicels 0.7–2.2 mm long; sepals 5(–7), unequal or subequal,
decreasing continuously from the size of the external sepals to the internal sepals, 2–4 × 0.9–1.7 mm, oval-lanceolate,
oblong or lanceolate, sometimes elliptic or ovate, with acute apex or less commonly obtuse or acuminate, margins
entire, plane, eglandular, stellate-porrect on both surfaces with greater density of trichomes on the dorsal surface, free,
valvate prefloration; petals 5, 0.3–0.8 × 0.1–0.2 mm, ovoid, subulate or filiform, glabrous; ovary 1.1–1.3 × 1.2–1.5
mm, globose, yellowish-ferrugineous with tomentose indument of stellate-porrect trichomes; stigmas 3, deeply bifid,
2–2.5 mm long, covered with stellate-porrect trichomes on the upper surface, glabrous on lower surface; disk with 5
segments; receptacle glabrous. Capsules 4.3–5 × 4.5–5.2 mm, globose, greenish with ferruginous or whistish indument,
accrescent sepals up to half the length of the fruit or similar to the length of the fruit, pedicels 1.3–2 mm long; seeds
3.7–4 × 2.6–2.9 mm, largely ellipsoid, grayish, with discretely dark spots, slightly foveolate; caruncle hat-shaped,
sessile, yellow-whitish, visible on ventral surface of the seed.
Vernacular name:—melosinha, velame.
Distribution and habitat:—Occuring in Ceará, Maranhão, Pará, and Piauí (Fig. 32b), in cerrado típico or restinga
vegetation on sandy soils between 15 and 225 m elevation.
Phenology:—Collected with flowers from November to July and fruits from April to July.
Preliminary conservation status:—We recommend a conservation status of Least Concern for this species,
because it has an Extent of Occurrence estimated at 60,000 km2, and some of its populations are inside protected areas,
such as the Sete Cidades National Park in Piauí.
Etymology:—The specific epithet honors Ricardo Secco, an expert on Amazonian Euphorbiaceae, for his valuable
contributions to the taxonomic knowledge of Croton.
Additional specimens examined (paratypes):—BRAZIL. Ceará: Barroquinha, Distrito de Bitupitá, 2°55’23”S,
41°12’14”W, 15 m, 10 June 2016, E.B. Souza et al. 4236 (HUVA, HUEFS, EAC), 4246 (HUVA, HUEFS, EAC).
Maranhão: Barreirinhas, Tabocas, 30 June 2012, K. Martins TB 64 (IAN). Piauí: Brasileira, Baixa do Barreiro (6ª
cidade)–PARNA de Sete cidades, 4°7’23”S, 41°42’13”W, 190 m, 30 June 2000, M.E. Alencar & R.S. Albino 1098
(TEPB, UEC, SP); Picada dos 100 pts (Baixa do Barreiro–6ª cidade)–PARNA de Sete cidades, 4°7’18”S, 41°42’21.6”W,
225 m, 12 July 2001, M.E. Alencar 1384 (TEPB, SP); s.loc., 24 April 2002, M.E. Alencar 1504 (SP). Piracuruca,
Parque Nacional de Sete Cidades, Boqueirão, 13 September 1877, G.M. Barroso 84 (RB, US); 28 February 1980,
M.R. Del’Arco s.n. (EAC 8157, HUEFS 82288); ibid., M.R. Del’Arco s.n. (EAC 8150); ibid., M.R. Del’Arco s.n. (EAC
8152, HUEFS 106348); ibid., M.R. Del’Arco s.n. (EAC 8164); ibid., M.R. Del’Arco & M.G.M. Arrais s.n. (HUEFS
92863) ibid., A. Fernandes s.n. (EAC 8131); 09 August 1985, A. Fernandes & Matos s.n. (EAC 13309, HUEFS
106367), 5ª cidade, próximo à furna do índio, 16 November 2001, A. Carvalho et al. 77 (TEPB, SP); em direção ao
Centro dos Visitantes na bifurcação (5ª cidade)–PARNA de Sete Cidades, 4°6’31”S, 41°41’21”W, 210 m, 23 Fev 2000,
M.E. Alencar et al. 856 (TEPB, UEC, HUEFS); Parque Nacional das Sete Cidades, entrada do Parque (1ª cidade),
4°6’12”S, 41°41’12”W, 174 m, 09 February 2007, D.S. Carneiro-Torres et al. 835 (HUEFS, HUESB). Pará, Maracanã,
Martins Pinheiro, Campina do Mangaba, 27 February 1975, L. Coradin 103 (IPEAN, RB); ca. 73 airline km NE of
Castanhal; savana adjoining the village of Martins Pinheiro, 0°52’S, 47°35’W, 50 m, 06 April 1980, G. Davidse et al.
17969 (INPA, MO, US)
Discussion and relationships:—Croton seccoi is placed here in Croton sect. Geiseleria because of its shrubby or
subshrubby habit, stellate indument, leaves with basilaminar and marginal glands, and unequal pistillate sepals. The
new species is morphologically most similar to C. parodianus, a species also found in northern Brazil, due to its ovate
leaf blades with frequently stipitate glands, petioles usually more than 1 cm long, flowers in inflorescences s up to 9
cm long, bracts with ovoid sessile glands at the base, staminate petals villous on both surfaces and pubescent, 2-fid
stigmas. However, C. seccoi differs from this species by the yellow-ferruginous indument on branches, leaves and
inflorescences, leaves with rounded, obtuse, truncate or assymetric bases and serrate or serrulate margins, staminate
bracts 1.4–1.6 mm long, pistillate flowers with oval-lanceolate, oblong, lanceolate, or sometimes ovate or elliptic
sepals, and maculate seeds. In C. parodianus the indument is whitish, the leaf blades have a cordate base and crenate-
serrate margins, the staminate bracts are 2–2.3 mm long, the pistillate sepals are usually obovate, and the seeds are not
maculate. Additionaly, C. seccoi has smaller petioles 0.6–2.5 cm long and smaller leaf blades 2.6–7.2 × 1.2–3.9 cm
(vs. petioles 2–8 cm long and leaf blades 5.2–15 × 3.1–8.5 cm in C. parodianus), sessile or shortly stipitate patelliform
basilaminar glands (vs. long-stipitate), leaf margins with glands only on the abaxial surface, one per sinus (vs. glands
on both surfaces, usually two glands per sinus) and staminal filaments glabrous (vs. villous at the base).
Many specimens of the C. seccoi have been identified erroneously in herbaria and in works of Carneiro-Torres
(2009) and Alves (2012) as C. goyazensis, a species endemic to Goiás, Minas Gerais, and the Distrito Federal. However,
these species are separated easily by morphological characters listed in the comments under C. goyazensis.
41. Croton sincorensis Martius ex. Müller Argoviensis (1873: 86). Type:—BRAZIL. Bahia: “in silvis Catingas ad
Sincorá”, s.d., C.F.P. von Martius 1952 (lectotype: M 0089119!, designated here; isolectotypes: G 00434687!, M
0089117!, M0089118!) (Figs. 35s–w, 39a–e).
Vernacular name:—cipó-lixa.
Distribution and habitat:—Croton sincorensis is endemic to Brazil and occurs in the states of Bahia, Espirito
Santo, Minas Gerais, and Pernambuco (Carneiro-Torres 2009; Silva et al. 2010) (Fig. 40a). It grows in capoeira,
bordering semi-deciduous forests, ombrophilous forests or caatinga shrublands on sandy or clayey soils between 35
and 1000 m of elevation.
Phenology:—Flowering from September to June and fruiting from January to June.
Preliminary conservation status:—Croton sincorensis has an Extent of Occurrence estimated at 350,000 km2,
and, therefore, we classify it as Least Concern.
Relationships:—Croton sincorensis is a shrub with pendent branches that reaches up to 5 m tall. Its leaves are
chartaceous, oval-lanceolate or oblong, with obtuse or truncate base, and a double-serrate or crenate margin with
stipitate glands in the sinuses. Its thyrses have 1 or 2 pistillate flowers with 6–10 lanceolate sepals, multifid stigmas
and usually with a sterile zone between the staminate and pistillate flowers, and the staminate flowers in numerous
3–5-flowered cymules. The morphological relation of this species with C. parodianus is discussed in the commentary
of that species.
Representative specimens examined:—BRAZIL. Bahia: 16 km S of Cumuruxatiba on the road to Prado,
39°15’S, 17°13’W, 50 m, 18 January 1877, R.M. Harley et al. 18083 (US); Lençóis, 3 km N pela ligação com a rodovia
BR-242, 9 April 1992, G. Hatschbach, M. Hatschbach and E. Barbosa 56940 (MBM, INPA). Espírito Santo: Santa
Teresa, fazenda Tabajara, Penha, 25 November 1985, H.Q. Boudet Fernandes 1689 (MBML).
42. Croton sipaliwinensis Lanjouw (1939: 698). Type:—SURINAME. Sipaliwini: sup. Kamp. XI, 10 December 1935,
H.E. Rombouts 357 (holotype: U 0001899; isotype: K 000254424!) (Fig. 41a–e)
Croton sclerocalyx var. pubescens Müller Argoviensis (1865: 134), syn. nov. Type:—GUYANA. s.loc., 1842–1843, Rich. Schomburgk 851
(holotype: B†; lectotype: P 00634708!, designated here; isolectotype: G n.v., photo at F 24529, K 000186092!).
Distribution and habitat:—Croton sipaliwinensis was previously reported for Brazil and Suriname by Secco (2008),
and it is also reported here for French Guiana and Guyana. In Brazil, it occurs in Amazonas, Pará, and Rondônia (Secco
2008) and is newly reported in Mato Grosso and Maranhão as well (Fig. 40a). It grows in riparian vegetation (e.g.,
igapó forest), and in campinas or capoeiras, between 50 and 200 m of elevation.
Phenology:—Collected with flowers and fruits from October to July.
Preliminary conservation status:—For presenting an Extent of Occurrence of 2.3 million km2, C. sipaliwinensis
has its conservation status here classified as Least Concern.
Relationships:—This species is morphologically close to C. sclerocalyx and C. lundianus, although they may be
differentiated by the characteristics discussed under C. lundianus.
Representative specimens examined:—BRAZIL. Amazonas: Barcelos, Tapereira (cemetary and old fort) and
São Tomé and vicinity; campina with rock (gneiss) 0°25’S, 64°25’W, 29 June 1979, J.M. Poole 1725 (RB); São Gabriel
da Cachoeira, margem da estrada Perimetral Norte, 17 July 1979, L.A. Maia 652 (INPA). Maranhão: Imperatriz,
behind rodobrás station on South side of city of Imperatriz, 5°33’S, 47°28’W, 6 January 1970, G. Eiten & L.T. Eiten
10214 (US); São Luís, Reserva da CAEMA, 28 April 1998, R.S. Secco & N.A. Rosa 879 (INPA). Mato Grosso:
Apiacás, rio Juruena, 8°53’28”S, 58°33’7”W, November 2007, M. Sobral et al. 11138 (RB); Cotriguaçu, Fazenda
FIGURE 39. Croton sincorensis a fertile branch, detail of the acropetiolar (above) and marginal glands (below), b portion of a thyrse
with staminate flowers, c thyrse with basal pistillate flowers and staminate buds, d pistillate flower, e young capsule. C. spica f habit, g
and h inflorescences showing pistillate flowers and staminate buds, i infructescence showing capsules and persistent staminate bracts, j
staminate flowers, k pistillate flowers.

FIGURE 40. Geographical distribution of some species of Croton sect. Geiseleria in Brazil. a Croton sincorensis, C. sipaliwinensis, C.
spica, C. subincanus and C. sublepidotus; b C. tetradenius, C. triangularis, C. trinitatis, C. uliginosus and C. virgultosus.
FIGURE 41. a–e Croton sipaliwinensis a leaf, b acropetiolar glands, c marginal gland, d staminate flower, e pistillate flower; f–j spica
f leaf, g basilaminar glands, h marginal gland, i staminate flower, j pistillate flower; k–n C. subincanus k leaf, l acropetiolar glands, m
staminate flower, n pistillate flower; o–r C. sublepidotus o leaf, p acropetiolar glands, q marginal gland, r staminate flower, s pistillate
flower; t–w C. tetradenius t leaf, u acropetiolar glands, v staminate flower, w pistillate flower. Drawn from: a–e M. Sobral 11138, f–j R.
C. Sodré 2768, k–n E. L. Sette Silva 529, o–r M. L. Fonseca 6622, t–w E. P. Queiroz 258.

São Nicolau, margem esquerda do rio Juruena, 16 October 2009, A. Petini-Benelli 153 (UFG), 154 (UFMT), 155
(UFMT); Paranaíta, Ilha do Manteiga, 9°13’57”S, 57°0’23”W, 132 m, 12 June 2012, C.R.A. Soares 6641 (HERBAM).
Pará: Oriximirá, Cachoeira Porteira, Weedy field towards the boat landing, south of the settlement, 18 June 1980, C.
Davidson & G. Martinelli 10349 (INPA, RB); Tucuruí, BR-230, km 48, capoeira na parte alta da ribanceira de um
riacho, 5 November 1979, M.F.F. Silva et al. 28 (INPA). Rondônia: Santa Bárbara, rodovia BR-364, km 120, 12 km da
sede da mineração, 9°10’S, 63°7’W, 29 May 1982, L.O.A. Teixeira et al. 850 (INPA). FRENCH GUIANA. Camopi:
Haute Camopi, Mont Belvédère lisière inferieure, 200 m, 21 November 1984, J.J. Granville 6944 (INPA).
43. Croton spica Baillon (1864: 362). Type:—BRAZIL. Mato Grosso: s.loc., 1831–1833, C. Gaudichaud-Beaupré
249 (holotype: P 00634647!, photo and fragment at F 0056198F!) (Figs. 39f–k, 41f–j)
Croton tamberlikii Müller Argoviensis (1873: 265), syn. nov. Type:—BRAZIL. “Brazilia occidentali”, s.d., Tamberlik s.n. (lectotype: W
0051319!, designated here; isolectotypes: F 0056204!, G 00169991!, W 0051320!).
Croton chapadensis Müller Argoviensis (1873: 262).—Oxydectes chapadensis (Müll.Arg.) Kuntze (1891: 613). Type:—BRAZIL. Mato
Grosso: Aldêa da Chapada, September 1827, L. Riedel 1136 (lectotype: LE 00003082!, designated here; isolectotypes: A 00257904,
LE 00003083!, P 05505085).
Distribution and habitat:—Croton spica was reported by Sodré et al. (2017) for the Distrito Federal, Goiás, and
Mato Grosso and is newly reported here for Mato Grosso do Sul and Tocantins (Fig. 40a). It grows in cerrado s.str. in
clay or rocky soil between 200 and 1,200 m elevation. Croton spica was also cited for Bolivia by Møller-Jørgensen et
al. (2014) as C. tamberlikii, based on the collection A. Rodriguez and J. Surubí 642 (F) identified by Grady Webster.
However, this collection actually corresponds to C. urticifolius Lamarck (1786: 219), so we do not acknowledge the
distribution of C. spica for Bolivia.
Phenology:—Collected with flowers and fruits from November to May.
Preliminary conservation status:—Silva et al. (2015b) classified the conservation of this species as of Least
Concern, a decision supported here, given its estimated Extent of Occurrence of 500,000 km2 and its abundant
populations, some of them located in protected areas, such as National Parks of Chapada dos Veadeiros and Chapada
dos Guimarães.
Relationships:—Croton spica can be confused with C. desertorum and C. ramosissimus, but can be distinguished
from them by the characters already noted in the comments of those species.
Taxonomic notes:—We designate Tamberlik s.n. from W [barcode 0051319] as lectotype of C. tamberlikii, since
it contains notes written by Müller, including one indicating that this specimen was used to illustrate the species in
Tabula 38 of Flora Brasiliensis (Müller Argoviensis 1873). Croton tamberlikii is synonymized under C. spica after
comparing its type collection, from the state of Mato Grosso, with collections of C. spica from the same state, and
finding no significant differences among them.
Representative specimens examined:—BRAZIL, Distrito Federal: Brasília, Recanto das Emas, estrada vicinal
a direita do km 13 da BR 060, sentido Brasília, após o posto da Polícia Rodoviária Federal e antes do trevo para
Santo Antônio do Descoberto, 15°55’53”S, 48°9’51”W, 1066 m, 12 December 2016, R.C. Sodré 2907 (UFG). Goiás:
Mineiros, Parque Nacional das Emas, 18°8’22.73”S, 52°53’10.22”W, 30 January 2009, C. Aoki 371 (CGMS, SP).
Niquelândia, beira da GO 237, 45 km após a Polícia Militar Rodoviária, esquerdo, sentido Niquelândia, 14°28’11”S,
48°46’8”W, 30 November 1999, S.M. Verboonen et al. 120 (CEN); Pirenópolis, Serra dos Pirineus, subida para o
Parque, 15°49’1”S, 48°53’38”W, 1085 m, 18 January 2005, J.P. Souza et al. 4134 (ESA). Mato Grosso: Barra do
Garças, Serra do Roncador, ca. 60 km a norte de Barra do Garças pela BR-158, ca. 5 km a esquerda desta rodovia
(sentido N. Xavantina) por uma vicinal que se inicia após o povoado Vale dos Sonhos, 15°20’54”S, 52°13’54”W,
642 m, 24 March 2016, R.C. Sodré et al. 2169 (UFG); Porto Esperidião, ca. 9 km ao longo da estrada que leva ao
Parque Estadual Serra de Santa Bárbara, acessada pela MT 265 / Vila Picada, 16°10’15.7”S, 59°11’2.2”W, 200 m, 03
November 2016, R.C. Sodré et al. 2768 (UFG). Vila Bela da Santíssima Trindade, ca. 55 km a noroeste de Pontes e
Lacerda, pela BR 174 e estrada vicinal que leva à Mineradora São Francisco, 14°52’08.6”S 59°35’52.7”W, 558 m, 04
November 2016, R.C. Sodré 2777 (UFG), 2778 (UFG). Mato Grosso do Sul: Corumbá, áreas de trabalho 2 e 3 (Mina 5)
MCR Rio Tinto, 7 December 2004, I.H. Ishii et al. 686 (COR); Morraria Urucum, 16 February 2006, E. Tameirão Neto
4314 (BHCB, COR). Tocantins: Monte do Carmo, margem direita da TO 255, sentido Ponte Alta do Tocantins a Monte
do Carmo, ca. 5 km desta última, 10°47’13”S, 48°4’11”W, 535 m, 10 April 2016, R.C. Sodré et al. 2327 (UFG), 2328
(UFG), 2329 (UFG). Palmas, estrada para Aparecida do Rio Negro, Parque Estadual do Lajeado, Fazenda Agronorte,
750 m, 13 February 2001, M. Alves et al. 2224 (SP).
44. Croton subincanus Müller Argoviensis (1865: 139). Type:—GUYANA. Roraima, 1842–1843, Rob. Schomburgk &
Rich. Schomburgk 665/1029 (lectotype: G-DC 00434675!, designated here; isolectotypes: B†, photo at F 0BN005173!,
BM 000560063!, G 00312864!, G 00434676, K 000254414!, P00634733, P00634734, P 00634735!) (Fig. 41k–n).
Distribution and habitat:—Croton subincanus is known from Guyana, Suriname, Venezuela, and Roraima state in
Brazil (Fig. 40a), growing near rivers in open vegetation, at altitudes of 700–1,200 m (Webster et al. 1999; Secco
2008).
Phenology:—Collected with flowers and fruits in November and June.
Preliminary conservation status:—Croton subincanus can be considered Least Concern because of its broad
range.
Relationships:—Croton subincanus can be recognized by its branches and leaves with a whitish indument of
stellate, tomentose trichomes; elliptic leaf blades with a cuneate base and sessile patelliform glands and entire or
serrulate margins without glands; congested thyrses with 6–12 pistillate flowers; bracts ca. 1 mm long; pistillate flowers
with elliptic, unequal sepals; and 2-fid stigmas. The species can be confused with C. hadrianii and C. hostmannii, but
it differs by the features shown in the comments of C. hostmannii. In Table 4 we also present the main differences
between C. subincanus and related species with entire or subentire leaf margins.
Representative specimens examined:—BRAZIL. Roraima: Serra Tepequem, 1036 m, 29 November 1954, B.
Maguire & C. Maguire 40097 (NY, UB); Boa Vista, Platô da Serra Tepequém, leste da corrutela, 3°45’N, 61°50’W, 6
June 1986, E.L. Sette Silva 529 (INPA).
45. Croton sublepidotus Müller Argoviensis (1873: 260). Type:—BRAZIL. Piauí or Tocantins: s. loc., 1844, G.
Gardner 3437 (holotype: BR 0000008760715!, isotype: G 00434674!, OXF 00091994!) (Fig. 41o–s).
Distribution and habitat:—Croton sublepidotus, a species previously known only from the type collection with an
unspecified locality, is confirmed here for Tocantins state (Fig. 40a). It occurs in cerrado s.str. in the municipalities of
Dianópolis and Mateiros, Jalapão region, in sandy soils between 440 and 700 m elevation.
Phenology:—Flowering and fruiting from October to May.
Preliminary conservation status:—Croton sublepidotus presents an estimated Extent of Occurrence at about
5,000 km2 and although some of its populations are found in protected areas such as Jalapão State Park and Serra
Geral do Tocantins Ecological Station, we treat it as Vulnerable, because it has only seven known locations, and the
populations consist of a small number of individuals.
Relationships:—Even though the trichomes of Croton sublepidotus are stellate, they have numerous, short rays
that give it a more lepidote appearance than most other species of the section. This species is morphologically similar to
C. mucronifolius, C. agoensis and C. pycandenius by its petioles 0.2–0.5 cm long, ovate or oval-lanceolate sometimes
oblong or elliptic leaf blades with lightly cordate, truncate or rarely obtuse base, entire, serrulate or serrate margin with
or without stipitate-capitate glands; long, lax thyrses (7.8–14.5 cm), with staminate cymules of 1–4 flowers; bracts 2.5–
3 cm long; and pistillate flowers with glabrous, equal, lanceolate sepals and 2-fid stigmas. However, C. sublepidotus
differs by its glabrescent and dark green branches, acropetiolar glands usually lacking; leaf blades concolorous, with
sparse trichomes and impressed and inconspicuous secondary veins on both surfaces and glabrous stigmas. On the
other hand, in C. agoensis, C. mucronifolius and C. pycandenius the branches are yellowish, whitish or grayish and
densely pubescent; the acropetiolar or basilaminar glands are always present; the stigmas are pubescent; and the leaf
blades are discolorous, with dense trichomes and conspicuous secondary veins on abaxial surface.
Representative specimens examined:—BRAZIL. Tocantins: Dianópolis, cerrado antes da cachoeira do rio
Palmeira, 11°33’53”S, 46°19’27”W, 700 m, 26 September 2003, A.O. Scariot et al. 824 (CEN); 825 (CEN). Mateiros,
Jalapão, estrada para São Felix do Jalapão, ca. 10 km de Mateiros, 10°41’31”S, 46°27’27”W, 8 October 2007, J.
Paula-Souza et al. 9187 (SP); estrada Rio Nova/Mateiros, 469 m, 10°36’S, 46°36’W, 9 May 2001, L.H. Soares e Silva
et al. 973 (UB, UFG); ESEC Serra Geral do Tocantins, região da Cachoeira da Velha, 10°22’1”S, 47°2’20”W, 440
m, 29 November 2012, M.L. Fonseca et al. 6622 (IBGE, HUEFS, HUTO); Serra Geral do Tocantins, 10°33’44.3”S,
46°26’33.4”W, 528 m, 18 December 2014, L.L.C. Antunes et al. 1333 (UFG), 1335 (UFG); estrada de São Félix do
Tocantins para Mateiros, 10°23’42”S, 46°29’33”W, 484 m, 06 November 2015, B.T.P.M. Goes et al. 142 (IBGE); ca.
23 km Mateiros-BA, 10°35’48”S, 46°17’46”W, 603 m, 11 November 2009, E. Melo et al. 7220 (HUEFS).

46. Croton tetradenius Baillon (1864: 343).—Type: BRAZIL. Minas Gerais: s.loc., 1816–1821, A.F.C.P. Saint-Hilaire
cat B1 1500 (lectotype: P 00634739!, designated here, photo and fragment at F 0056207F; isolectotypes: P 00634740!,
P 00634741!) (Figs. 41t–w, 42a–f).
Croton pulegiodorus Baillon (1864: 361), syn. nov.—Croton pulegioides Müller Argoviensis (1873: 259), nom illeg.—Croton tristis
Müller Argoviensis (1865: 140), nom illeg. Type:—BRAZIL. Bahia: Jacobina, St Thomé, June 1844, J.S. Blanchet 3786 (lectotype: P
00634607!, designated here; isolectotypes: BR 0000008761477!, G 00312860, G 00434633!, P 00634608!, P 00634609!); remaining
syntypes:—BRAZIL. Minas Gerais: s.loc., 1839, P. Claussen 764 (A 00257980!, G 00434634!, G 00434635!, P 00634610!, P
00634611!).
Croton leptobotryus Müller Argoviensis (1873: 155), syn. nov. Type:—BRAZIL. Minas Gerais: “ad Lagoa Santa super rupes calcareas et in
virgultis ad Sumidouro”, 2 December 1864, J.E.B. Warming 1628 (holotype: C 10011177!; isotypes: A 00257981!, G 00434560!).
Croton regelianus Müller Argoviensis (1873: 133), syn. nov. Type:—BRAZIL. Minas Gerais: “in umbrosis ad rivulum Cipó”, November
1824, L. Riedel 869 (lectotype: LE 00018228!, designated here; isolectotypes: G 00434626!, GH 00257987!, K 000186148!, LE
00018226!, LE 00018227!).
Croton regelianus var. matosii Radcliffe-Smith (1993: 150), syn. nov. Type:—BRAZIL. Ceará: Chapada da Ibiapaba, Cocalzinho–Viçosa,
14 January 1982, A. Fernandes & F. Matos 11077 (holotype: K 000186147!; isotype: EAC).
Vernacular name:—alecrim-de-são-joão, barba-de-bode, caatinga-de-bode, velame, velaminho, velaminho-da-serra,

velandinho, zabelê.
Distribution and habitat:—Croton tetradenius is widely distributed in Brazil, especially in the Northeast, where
it is present in all states except Maranhão (Carneiro-Torres 2009; Lucena 2009). There are also reports from Goiás,
São Paulo, and Minas Gerais (Caruzo & Cordeiro 2007), and it is newly reported here for the Distrito Federal, Mato
Grosso, Mato Grosso do Sul, and Tocantins (Fig. 40b). It grows in cerradão, semideciduous seasonal forests, caatinga,
capoeira, brejos de altitude, usually in calcareous outcrops, sandy or clayey soils, between 10 and 1,160 m elevation.
Preliminary conservation status:—This species can be categorized as Least Concern because its Extent of
Occurrence is estimated at roughly 2,000,000 km2.
Taxonomic notes and characterization:—Croton tetradenius is a polymorphic species, remarkable in the 2–6
obconic to cylindric acropetiolar glands and membranaceous leaf blades with acuminate or caudate apex and entire
or serrulate margins. Croton pulegiodorus, C. pulegioides, C. tristis, C. leptobotryus, C. regelianus and C. regelianus
var. matosii are treated here as synonyms, as already suggested by Lucena (2009) and Alves (2012) in their respective
doctoral thesis and master’s dissertation. Croton tetradenius, in the present circumscription, is also recognized for
being an aromatic subshrub 0.4–1.3 m tall, with thin, dichotomously branched stems, usually whorled leaves below
the inflorescences or at the dichotomy of the branches, petioles 1.1–7.1 cm long, actinodromous-brochidodromous
leaf venation, lax thyrses 1.2–15.9 cm long without a sterile zone between pistillate and staminate flowers, short bracts
(0.8–1.4 cm long), and staminate cymules of 1–3-flowers; pistillate flowers with equal oblong or oval-lanceolate
sepals and 2-fid stigmas; and (11–)14–16 stamens.
Representative specimens examined:—BRAZIL. Bahia: Oliveira dos Brejinhos, ca. 6 km ao sul da cidade,
12°22’2.7”S, 42°54’14.3”W, 884 m, 20 December 2017, R.C. Sodré et al. 3343 (UFG). Ceará: Viçosa do Ceará, 28
October 1975, A. Fernandes s.n. (EAC 0002633). Distrito Federal: Brasília, perto do Rio Salinas, 15°31’S, 47°57’W,
840 m, 26 February 1981, J.H. Kirkbride 3939 (UFG); APA da Cafuringa, próximo ao rio da Palma, 2 December 2011,
M.R.V. Zanatta & J.E.Q. Faria 1052 (UB, UFG). Goiás: Colinas do Sul, cerca de 2 km da ponte do rio Bagagem,
14°09’S, 48°04’W, 450 m, 23 November 1992, R.F. Vieira et al. 1380 (CEN); Posse, Fazenda Sabonete, 14°03’53”S,
46°29’15”W, 03 November 2000, M. Aparecida da Silva et al. 4639 (IBGE). Mato Grosso: Alto Paraguay, Fazenda
Currupira, Serra das Araras, 15°10’S, 56°51’W, 26 January 1995, B. Dubs 1784 (ESA). Mato Grosso do Sul: Bonito,
Fazenda Esperança, 12 km N de Bonito, 17 June 2001, A. Pott et al. 9100 (HMS, CGMS, UFG); Fazenda Santo
Antônio, 2,5 km NE da rodovia MS-345, entrada a 10 km N de Bonito na rodovia MS-345, 21°1’33”S, 56°26’10”W,
300 m, 4 December 2007, A. Pott & V.J. Pott 14823 (CGMS, UFG). Minas Gerais: Missões, pedreira a 7 km de Ita,
14°54’S, 44°06’W, 620 m, 14 November 1991, R.F. Vieira et al. 1051 (CEN, HEPH); Monte Azul, 2 km ao norte de
Monte Azul na BR-122, trecho Monte Azul–Espinhosa, 14°55’S, 42°50’W, 685 m, 08 November 1984, A.C. Allem et
al. 2894 (CEN). Pernambuco: Inajá, Reserva Biológica de Serra Negra, 09 March 1995, M.F. Sales et al. 561 (PEUFR);
Brejo da Madre de Deus, fazenda Buriti, 8°9’00”S, 36°22’15”W, 29 March 1996, L.F. Silva et al. 202 (PEUFR). Piauí:
São João do Piauí, Embrapa Meio Norte, 8°20’39”S, 42°19’35”W, 24 March 2015, K.N.C. Castro & J.B.A. Souza 474
(CEN). Rio Grande do Norte: Macaíba, Reserva do Assentamento José Coelho, 16 June 2004, R.T. Queiroz 87 (UFRN,
MOSS). Tocantins: Aurora do Tocantins, estrada para o Balneário, 2,5 km do balneário, 12°46’55”S, 46°21’45”W, 25
January 2005, J. Paula-Souza 4621 (ESA, SP); Natividade, Mineradora a 3 km da cidade, 11°37’10”S, 47°41’42”W,
350–550 m, 7 March 2015, A.M. Amorim et al. 9330 (RB).
FIGURE 42. Croton tetradenius a habit, detail of the acropetiolar glands in the insert, b staminate flowers, c and d pistillate flowers, e
portion of a thyrse with young capsules and staminate flowers and buds, f capsules, note the strongly accrescent sepals. C. triangularis
g habit, h leaves, with detail of the acropetiolar glands in the insert, i distal portion of a thyrse showing staminate flowers and buds, j
staminate cymule with a flower and buds, k cymule with a pistillate flower and staminate buds, l young capsule, note the strongly revolute
sepals.

47. Croton triangularis Müller Argoviensis (1865: 128). Type:—BRAZIL. Bahia: “inter Campos et Vittoria”, December
1836, F. Sellow 1242 (holotype: B 100127737!, photo at F 0BN005188!; isotype: A 00047455!) (Figs. 42g–l, 43a–e).
Distribution and habitat:—Croton triangularis was reported by Carneiro-Torres (2009) for Bahia and Minas Gerais
states, and it is also cited here for Piauí (Fig. 40b). It grows in shrubby caatinga and in transition areas between
Caatinga and Cerrado, in rocky, clayey or sandy soils between 400 and 1,169 m elevation.
Phenology:—Collected with flowers from September to March and with fruits from February to April.
Preliminary conservation status:—Although Carneiro-Torres (2009) estimated the Extent of Occurrence of
the species at only 2,000 km2 and categorized it as Vulnerable, but the analysis of a larger number of specimens has
increased its Extent of Occurrence to 180,000 km2, for which we now treat its conservation status as Least Concern.
Relationships:—Croton triangularis can be confused with C. adamantinus and C. virgultosus by its stipitate
acropetiolar glands and leaf blades with a cordate base and actinodromous-brochidodromous venation that is 3–5-
nerved at the base. However, C. triangularis is easily recognized by its chartaceous, ovate or triangular, strongly
discolorous leaf blades with whitish indument with ferrugineous veins on the abaxial surface; long, robust, ferrugineous
thyrses (up to 30 cm long), with staminate cymules of 3–5 flowers; pistillate flowers with equal sepals with revolute
margins and 4-fid stigmas, and long capsules (8–9 mm long). On the other hand, C. adamantinus and C. virgultosus
have leaf blades that are ovate, lanceolate or oval-oblong, lightly discolorous and greenish on the abaxial surface, the
thyrses are up to 9 cm long with staminate cymules of 2 flowers, the pistillate flowers have unequal pistillate sepals
with plane margins and 2-fid stigmas, and the capsules are 6–7 mm long.
Representative specimens examined:—BRAZIL. Bahia: Caetité ca. 5 km a noroeste de Caetité, sentido Bom
Jesus da Lapa, 14°2’38.1”S, 42°31’11.17”W, 981 m, 16 December 2017, R.C. Sodré et al. 3295 (UFG); Cândido
Sales, povoado de Santa Cruz, 15°29’30”S, 41°18’29”W, 17 December 2011, M.M. Silva-Castro et al. 1580 (HUESB);
Correntina, mata de dentro à 12 km da cidade, 13°22’S, 44°38’, 620 m, 18 November 1991, R.F. Vieira et al. 1150 (CEN,
SP); Igaporã, cerca de 5 km a partir do entroncamento da BR-430, na estrada que vai para Tanque novo, 17 January
2004, M. Machado & M.V.M. Oliveira 175 (HUEFS); Licínio de Almeida, 2 km antes da entrada da cidade, 14°32’4”S,
42°31’51”W, 10 January 2006, T.S. Nunes et al. 1621 (HUEFS); São Félix do Coribe, 13°33’18”S, 44°7’26”W, 705
m, 11 January 2008, R.F. Souza-Silva et al. 308 (HUEFS). Minas Gerais: Grão Mogol, próximo à Fazenda Varjão do
Morro, 16°24’52.86”S, 42°57’23.28”W, 840 m, 11 April 2016, O.L.M. Silva et al. 293 (SP). Francisco Sá, km 16 da
rodovia BR-251, trecho Montes-Claros–Francisco Sá, 16°30’S, 43°30’W, 620 m, 07 November 1984, A.C. Allem et
al. 2883 (CEN, SP); Janaúba, km 5 da BR-122 Francisco Sá–Janaúba, 16°30’S, 43°35’W, 690 m, 07 November 1984,
A.C. Allem et al. 2883 (CEN, RB, SP); Matias Cardoso, sentido Manga, Parque Lagoa do Cajueiro, 14°56’15”S,
43°53’55”W, 496 m, 22 April 2006, D.S. Carneiro-Torres et al. 746 (HUEFS, NY); Riacho dos Machados, Serra da
Bocaina, km 400 da BR-251, próximo à torre da Telemig, 10 March 1999, A. Rapini & M.L. Kawasaki 725 (HUEFS);
Salinas, estrada Salinas/Aeroporto, a 2 km do entroncamento, 16°11’46”S, 42°20’18”W, 704 m, 7 April 2002, T.
Ribeiro et al. 297 (HUEFS); Turmalina, Posses, estrada entre Posses e Botumirim, 2 km sul da passagem de balsa do
rio Jequitinhonha, 16°59’51”S, 42°47’51”W, 445 m, 24 January 2004, R. Riina et al. 1353 (HUEFS). Piauí: Caracol,
Parque Nacional da Serra das Confusões, entre Lajeado do Canto Verde e a Toca do Urubú, 8°54’9”S, 43°26’40”W,
580 m, 18 February 2013, G. Martinelli et al. 17917 (RB).
48. Croton trinitatis Millspaugh (1900: 57). Type:—GUYANA. s.loc., 1838, Rob. Schomburgk 241 (lectotype: BM
001124405!, designated by van Ee & Berry 2010; isolectotype: G 00312806!) (Fig. 43f–i, 44a–b).
Croton miquelensis Ferguson (1901: 49), nom. illeg. Type:—U.S.A. Florida: Escambia Co., wharf at Pensacola, 20 September 1886, A.H.
Curtiss 15 (lectotype: GH 277262!, designated by van Ee & Berry 2010); remaining syntypes:—GUYANA, 1837, Rob. Schomburgk
241 (BM); TRINIDAD, s.d., H. Crueger s.n. (TRIN).
Croton tragioides Blake (1922: 11). Type:—GUATEMALA. Izabal: shores of Lake Izabal, 2 June 1919, S.F. Blake 7854 (holotype: US
00028448!).
Croton buchtienii Rusby (1927: 284). Type:—BOLIVIA. Beni: distr., General José Ballivián Segurola, Reyes, 1000 ft, 26 October 1921,
H.H. Rusby 1332 (lectotype: NY 246520!, designated by van Ee & Berry 2010; isolectotypes: GH 47237!, MICH 1104791!, NY
579361!, PH 00008862!, US 01232343/00109512!).
Vernacular name:—guanxuma, vassourinha.

Distribution and habitat:—A widespread American species distributed from Central America to Bolivia and
Paraguay (Govaerts et al. 2000; van Ee & Berry 2010). In Brazil, it occurs in all the states of the north, and in the
Midwest (MT, MS), Northeast (BA, MA, PB, PI, PE) and Southeast (MG, RJ) regions (Fig. 40b). It inhabits campo
de várzea, campo limpo, campinarana, edge of riparian forest and in anthropic environments (BFG 2015) on sandy or
clayey soils, from sea level to 750 m of elevation.
Preliminary conservation status:—Due to wide distribution, C. trinitatis has conservation status considered as
Least Concern.
Relationships:—Croton trinitatis it is often confused with C. glandulosus, C. lundianus and C. hirtus in
herbarium collections, however, these four species can be differentiated by the features shown in Table 3 (under C.
glandulosus).
Representative specimens examined:—BRAZIL. Acre: Cruzeiro do Sul, Along road from Cruzeiro do Sul to
Barão do Rio Branco, Nortwest of Cruzeiro do Sul; Vicinity of Rio Moa, 5 km East of Rio Moa, 7°37’S, 72°26’W,
25 August 1986, T.B. Croat & A. Rosas Jr. 62652 (INPA). Amapá: Santana, Coração, 07 October 1979, D.F. Austin et
al. 6975 (INPA). Amazonas: Barcelos, Rio Negro, lugar Acanga, pouco abaixo de Barcelos, 27 June 1979, L.A. Maia
et al. 88 (INPA). Maranhão: Balsas, Gerais de Balsas, Vila de Carli, 46°5’S, 70°35’W, 300 m, 9 March 1998, R.C.
Oliveira & G.P. da Silva 1016 (IPA, HEPH). Minas Gerais: Pirapora, setor industrial, 9 January 1990, G. Hatschbach
et al. 54129 (MBM, C). Pará: Almeirin, Monte Dourado, Rio Jarí, canteiro de Germinação de Gemelina, 1 October
1968, N.T. Silva 1073 (IAN). Rondônia: Candeias do Jamari, estrada Porto Velho–Cuiabá, BR 364, km 48 a margem
da estrada, 5 February 1983, M.R.C. Bilby et al. 12 (INPA). Tocantins: Lagoa da Confusão, Ilha do Bananal, Parque
Nacional do Araguaia, Lago do Quitandeiro, 10°23’43”S, 50°26’43”W, 170 m, 25 March 1999, M. Aparecida da Silva
et al. 4147 (IBGE, UFMT). COLOMBIA. Córdoba: Ayapel, Palotal, 16 May 1949, R. Romero-Castañeda 1678 (IAN).
GUYANA. Essequibo ISL-W Demerara: Parika backdam, 6°50’S, 58°28’W, 0–10 m, 18 August 1993, T.W. Henkel et
al. 2671 (INPA). FRENCH GUIANA: Village Trou Poisson, Route Cayenne–St Laurent, PK 117, avant Trou Poisson,
27 December 1986, G. Cremers 9537 (INPA). PANAMA. Colón: 1 km off Pipeline Road, 2 km from end, 17 km
NW of Gamboa, 60 m, 7 April 1974, M. Nee 11123 (LD). PERU. Loreto: Iquitos, 22 October 1940, E. Asplund (LD).
VENEZUELA. Lower Orinoco, 1856, H.H. Rusby & R.W. Squires 29 (WU).
49. Croton uliginosus Sodré & M.J. Silva (in Sodré & Silva 2018a: 101). Type:—BRAZIL. Goiás: Mineiros, Emas
National Park, valley of the Avoador stream (tributary of the Jacúba River), ca. 16 km from the headquarters of the
Mineiros Park, 1.5 km from the road, at the margin of the stream, 18°0’34.4”S, 52°56’14.6”W, 767 m, 15 April 2016,
R.C. Sodré et al. 2385 (holotype:UFG!; isotypes HJ!, CGMS!, K!) (Figs. 43j–m, 44c–g).
Distribution and habitat:—Croton uliginosus is known from the states of Goiás and Mato Grosso (Fig. 40b), in
campo limpo or campo sujo, on hydromorphic soils between 760 and 930 m of elevation (Sodré & Silva 2018a).
Phenology:—Collected with flowers and fruits from November to May.
Preliminary conservation status:—Although Sodré & Silva (2018a) recognized the conservation of C. uliginosus
as Least Concern, in this study we propose to change it to Vulnerable, since the Extent of Occurrence this species is
estimated at 10,000 km2. In addition, C. uliginosus is known from only five localities, of which only the Emas National
Park is a protected area.
Relationships:—Croton uliginosus is a linear-leaved species that was compared to C. junceus by Sodré & Silva
(2018a). However, it can be differentiated from the latter by its ascending, lanceolate or linear-lanceolate leaves with
short petioles (up to 3 mm long), with serrate margins lacking glands in the sinuses; and by its thyrses with only two
pistillate flowers at the base; staminate flowers with 7 or 8 stamens, and 6-merous pistillate flowers.
Representative specimens examined:—BRAZIL. Goiás: Caiapônia, estrada Caiapônia–Jataí, margem esquerda
junto a fonte de água sulfurosa, 14 October 1968, Sidney & Onishi 1537 (UB); Cachoeira do Sereno, próximo à BR 158
e a entrada do brejo Vereda, 16°56’39”S, 51°49’58”W, 21 November 2011, R.F. Monteiro et al. 406 (RB). Mineiros,
74 km from Alto Araguaia to Goiânia, 930 m, 30 September 1963, J.M. Pires 56979 (UB). Mato Grosso: Alto Taquari,
entre a ferrovia Ferronorte e a área urbana de Alto Taquari, 17°51’21”S, 53°16’41”W, 28 February 2002, V.J. Pott et
al. 5092 (HMS); ibid., 20 September 2016, R.C. Sodré & R.A. Pereira-Silva 2584 (UFG); ca. 500 m abaixo do lago da
cidade, paralelo à ferrovia, 17°51’1.8”S, 53°17’0.7”W, 861 m, 6 November 2016, R.C. Sodré 2814 (UFG).

FIGURE 43. a–e Croton triangularis a leaf, b acropetiolar glands, c marginal gland, d staminate flower, e pistillate flower; f–i trinitatis
f leaf, g basilaminar glands, h staminate flower, i pistillate flower; j–m C. uliginosus j leaf, k basilaminar glands, l staminate flower, m
pistillate flower; n–r C. virgultosus n leaf, o basilaminar glands, p marginal gland, q staminate flower, r pistillate flower. Drawn from a–e
R. C. Sodré 3295, f–i R. C. Sodré 2256, j–m R. C. Sodré 2381, n–r M. L. Silva 203.
FIGURE 44. Croton trinitatis a fertile branch, detail of a short inflorescences showing pistillate flowers and staminate flowers and buds
in the upper left corner, b fertile branch, detail of the capsules in the upper right corner. C. uliginosus c flowering branch, with detail of
the basilaminar glands in the insert, d staminate flowers and buds, e and f pistillate flowers, g capsule. C. virgultosus h flowering branch,
i thyrse showing basal pistillate flowers and distal staminate flowers and buds, with a gap in between the two sections.

50. Croton virgultosus Müller Argoviensis (1873: 104). Type:—BRAZIL. Bahia: ad margines silvarum siccarum et
in pascuis prope Caxoeira, February, C.F.P. Martius 2202 (lectotype: M 0089126!, designated here; isolectotypes: G
00434719!, M 0089124!, M 0089125!, M 0089127!, M 0089128!) (Figs. 43n–r, 44h, i).
Distribution and habitat:—This species is restricted to Brazil with records from the states of Bahia, Ceará, Minas
Gerais, Pernambuco, Piauí, and Rio Grande do Norte (BFG 2015) (Fig. 40b). It grows in arboreal caatingas, on sandy
or stony soils, between 80 and 880 m elevation.
Phenology:—Flowering from November to April and fruiting from January to May.
Preliminary conservation status:—This species presents an Extent of Occurrence estimated at 500,000 km2,
and, therefore, is here classified in the category Least Concern.
Relationships:—Croton adamantinus, C. crustulifer and C. triangularis are the morphologically closest species
to C. virgultosus, as discussed in the comments of these other species. In general, C. virgultosus can be distinguished
by having oval to oblong leaves with a cordate base, long stipitate glands at the apex of the petioles and in the sinuses
of the leaf margins, thyrses 5–10 cm long with 5–8 pistillate flowers with unequal, lanceolate or elliptic sepals; and
villous staminate petals on both surfaces.
Taxonomic notes:—It was necessary to make a lectotypification of Croton virgultosus, because there are five
sheets of C.F.P. Martius 2202 at M and one at G. We chose M 0089126 as the lectotype, since it was the only one
annotated by Müller and the one with the most complete branches that best reflects the original description.
Representative specimens examined:—BRAZIL. Bahia: Morro do Chapéu, rio Ferro Doido, um pouco embaixo
à Cachoeira, 11°37’15”S, 40°59’46”W, 876 m, 03 March 1997, P. Gasson et al. 5970 (ALCB, UFBA). Minas Gerais,
Januária, 04 November 1978, L. Krieger 16085 (SP). Rio Grande do Norte: Mossoró, Distrito de Alagoinha, Fazenda
Rafael Fernandes, 5°3’14”S, 37°24’26”W, 87 m, 20 April 2006, M.L. Silva, R.C. Oliveira & R.B. Silva 31 (MOSS).
Final considerations
After this revision, including synonyms and discoveries of new species in Croton section Geiseleria, we now estimate
a total of 80 species in the section, 50 of which occur in Brazil, and 37 of them endemic there. However, further
collections, especially in the North and Midwest regions of Brazil, will likely reveal new species for this group.
Although we have estimated the conservation status for all of the species, these should be considered preliminary
assessments, because we still know little about their areas of occurrence and the risk factors of their populations.
Acknowledgements
We are grateful to Cordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the scholarship grant
to the first author; the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the productivity
grant (process no. 302648/2016-0) supplied to Marcos José da Silva; Renato Galhardo Neto for his skillful illustrations,
to curators; and to the staff of the herbaria cited for allowing access their collections; André Luiz de Rezende Cardoso
and Dr. Marcelo Fragomeni Simon for providing some of the photographs of the plants in the field; and Ricarda Riina
for providing photographs of type collections of some of the species examined. This work was also supported by NSF
grants DEB-0212481 to P.E. Berry and DEB-0508725 to P.E. Berry and B. van Ee.
References
Alves, A.S.A. (2012) Taxonomia de Croton sect. Ocalia (Klotzsch) Baill. no Brazil e Croton sect. Geiseleria (A. Gray) Baill. s.l.
(Euphorbiaceae) no Nordeste do Brazil. Master’s dissertation. Universidade Federal Rural de Pernambuco, Recife, 91 pp.
Armbruster, W.S., Berry, P.E., Esser, H.-J., Gillespie, L.J., Hayden, W.J., Heald, S.V., Levin, G.A., Punt, W., Secco, R.S. & Webster, G.L.
(2007) Euphorbiaceae. In: Funk, V., Hollowell, T., Berry, P., Kelloff, C. & Alexander, S.N. (Eds.) Checklist of the Plants of the
Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro, Guyana, Surinam, French Guiana). Contributions of United States
National Herbarium 55: 290–338.
Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial
conservation assessment tool. ZooKeys 150: 117–126.
https://doi.org/10.3897/zookeys.150.2109
Baillon, H.E. (1858) Étude générale du groupe des Euphorbiacées. Librairie de Victor Masson, Paris, 684 pp.
Baillon, H.E. (1864) Species Euphorbiacearum. Euphorbiacées Américaines. Première partie: Amérique austro-orientale. Adansonia 4:
257–377.
Baillon, H.E. (1865) A L’énumération des espèces d’Euphorbiacées du Brésil. Adansonia 6: 15–16.
Bentham, G. (1854) On the North Brazilian Euphorbiaceae in the collections of Mr. Spruce. Hooker’s Journal of Botany and Kew Garden
Miscellany 6: 321–377.
Berry, P.E., Hipp, A.L., Wurdack, K.J., Van Ee, B. & Riina, R. (2005) Molecular phylogenetics of the giant genus Croton and tribe Crotoneae
(Euphorbiaceae sensu stricto) using ITS and trnL-trnF DNA sequence data. American Journal of Botany 92: 1520–1534.
https://doi.org/10.3732/ajb.92.9.1520
BFG—The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085–1113.
https://doi.org/10.1590/2175-7860201566411
Carneiro-Torres, D.S. (2009) Diversidade de Croton L. (Euphorbiaceae) no bioma Caatinga. Doctoral thesis. Universidade Estadual de
Feira de Santana, Feira de Santana, 296 pp.
Carneiro-Torres, D.S., Cordeiro, I., Giulietti, A.M., Berry, P.E. & Riina, R. (2011) Three new species of Croton (Euphorbiaceae s.s.) from
the Brazilian Caatinga. Brittonia 63: 122–132.
https://doi.org/10.1007/s12228-010-9145-1
Caruzo, M.B.R. & Cordeiro, I. (2007) Sinopse da tribo Crotoneae Dumort. (Euphorbiaceae s.s.) no Estado de São Paulo, Brazil. Hoehnea
34: 571–585.
https://doi.org/10.1590/S2236-89062007000400011
Caruzo, M.B.R. & Cordeiro, I. (2013) Taxonomic revision of Croton section Cleodora (Euphorbiaceae). Phytotaxa 121: 1–41.
Chodat, R.H. & Hassler, E. (1905) Plantae hasslerianae soit énumération des plantes récoltés au Paraguay par le Dr. Emile Hassler de
1885–1902. Bulletin de l’Herbier Boissier, ser. 2, 5: 481–506.
Cordeiro, I. (1992) Flora da Serra do Cipó, Minas Gerais: Euphorbiaceae. Boletim de Botânica da Universidade de São Paulo 13: 169–
217.
Croizat, L. (1941) Preliminaries for the study of Agentine and Uruguayan species of Croton. Darwiniana 5: 417–462.
Croizat, L. (1944) Additions to the genus Croton L. in South America. Darwiniana 6: 442–468.
Croizat, L. (1948). Euphorbiaceae. In: Maguire, B. (Ed.) Plant explorations in Guiana in 1944, chiefly to the Tafelberg and the Kaieteur
Plateau—IV. Bulletin of the Torrey Botanical Club 75: 400–408.
Didrichsen, F. (1857) Plantas nonnullas musei Universitatis Hauniensis. Videnskabelige Meddelelser fra den naturhistoriske Forening i
Kjöbenhavn 8–10: 123–151.
Fawcett, W. & Rendle, A.B. (1920) Flora of Jamaica IV. British Museum, London, 369 pp.
Ferguson, A.M. (1901) Crotons of the United States. Annual Report Missouri Botanical Garden 12: 33–73.
Froembling, W. (1896) Anatomisch-systematische Untersuchung von Blatt und Axe der Crotoneen und Euphyllantheen. Botanisches
Centralblatt 65: 129–422.
Glaziou, A.F.M. (1913) Plantae Brasiliae centralis a Glaziou lectae. Bulletin de La Société Botanique de France 59 (3g): 611–648.
Gomes, A.P.S. (2006) Revisão das espécies sul-americanas de Croton L. subgen. Croton sect. Argyroglossum Baill. (Crotonoideae-
Euphorbiaceae). Doctoral thesis. Universidade Federal Rural de Pernambuco, Recife, 140 pp.
Govaerts, R., Frodin, D.G. & Radcliffe-Smith, A. (2000) World checklist and bibliography of Euphorbiaceae (and Pandaceae). Royal
Botanic Gardens, Kew, 1622 pp.
Govaerts, R.H.A. (2011) World checklist of selected plant families published update. Facilitated by the Trustees of the Royal Botanic
Gardens, Kew. Available from: http://apps.kew.org/wcsp/prepareChecklist.do (accessed 4 September 2018)
Gray, A. (1856) A manual of the botany of the Northern United States. 2nd ed. George P. Putnam & Co., New York, 739 pp.
Grisebach, A.H.R. (1859) Flora of the British West Indian Islands 1. Reeve & Co., London, pp. 1–96.
Hiepko, P. (1987) The collections of the Botanical Museum Berlin-Dahlem (B) and their history. Englera 7: 219–252.
Hickey, L.J. (1973) Classification of the Architecture of Dicotyledonous Leaves. American Journal of Botany 60: 17–33.
IBGE—Instituto Brasileiro de Geografia e Estatística (2012) Manual Técnico da Vegetação Brasileira. 2nd ed. Instituto Brasileiro de
Geografia e Estatística, Rio de Janeiro, 272 pp.
IUCN (2017) Guidelines for using the IUCN red list categories and criteria. Version 13. Prepared by the Standards and Petitions
Subcommittee of the IUCN Species Survival Commission. IUCN, Gland and Cambridge. Available from: http://cmsdocs.
s3.amazonaws.com/RedListGuidelines.pdf (accessed 12 September 2018)

Jablonski, E. (1965) Euphorbiaceae. In: Maguire, B. (Ed.) Botany of the Guayana Highlands—Part VI. Memoirs of The New York Botanical
Garden 12: 150–178.
JSTOR Global Plants (2018) JSTOR Global Plants. Available at: http://plants.jstor.org/ (accessed 18 September 2017)
Klotzsch, J.F. (1841a) Neue und weniger gekannte südamerikanische Euphorbiaceen-Gattungen. Archiv für Naturgeschichte 7: 175–204.
Klotzsch, J.F. (1841b) Die Crotoneae der Flora von Nord-Amerika. Archiv für Naturgeschichte 7: 250–255.
Klotzsch, J.F. (1843) Euphorbiaceae. In: Bentham, G. (Ed.) XIV. Contribution towards a Flora of South America—Enumeration of Plants
Collected by Mr. Schomburgk in British Guiana. London Journal of Botany 2: 42–52.
Klotzsch, J.F. (1853) Euphorbiaceae. In: Seemann, B.C. (Ed.) The botany of the voyage of H.M.S. Herald. Reeve & Co., London, pp.
99–106.
Kunth, K.S. (1817) Euphorbiaceae. In: Humboldt, F.W.H.A, Bonpland, A.J.A. & Kunth, K.S. (Eds.) Nova Genera et Species Plantarum
2. Librairie Graeco-Latino-Germanicae, Paris, 401 pp.
Kuntze, C.E.O. (1891) Revisio Generum Plantarum 2. Arthur Felix, Leipzig, pp. 375–1011.
Kuntze, C.E.O. (1898) Revisio Generum Plantarum 3 (3). Arthur Felix, Leipzig, 576 pp.
Lamarck, J.B. (1786) Encyclopédie méthodique. Botanique 2. Panckoucke, Paris, 774 pp.
Lanjouw, J. (1939) New or noteworthy Euphorbiaceae from Suriname. Recueil des Travaux Botaniques Néerlandais 36 (2): 697–704.
L’Héritier de Brutelle, C.L. (1785) Stirpes novae aut minus cognitae. Philippi-Dionysii Pierres, Paris, 184 pp.
Lima, L.R. & Pirani, J.R. (2003) O gênero Croton L. na Cadeia do Espinhaço, Minas Gerais, Brasil. Boletim de Botânica da Universidade
de São Paulo 21: 299–344.
https://doi.org/10.11606/issn.2316-9052.v21i2p299-344
Linnaeus, C. (1753) Species plantarum. Laurentius Salvius, Stockholm. 1200 pp.
Linnaeus, C. (1759) Systema Naturae ed. 10, 2. Laurentius Salvius, Stockholm, pp. 825–1384.
Lucena, M.F.A. (2009) Diversidade de Euphorbiaceae s.l. no Nordeste do Brazil. Doctoral thesis. Universidade Federal de Pernambuco,
Recife, 197 pp.
Meireles, L.D., Kinoshita, L.S. & Shepherd, G.J. (2014) Composição florística da vegetação altimontana do distrito de Monte Verde
(Camanducaia, MG), Serra da Mantiqueira Meridional, Sudeste do Brasil. Rodriguésia 65: 831–859.
https://doi.org/10.1590/2175-7860201465403
Millspaugh, C.F. (1900) Plantae Utowanae. Publications of the Field Columbian Museum, Botanical Series 2: 1–110.
Miquel, F.A.W. (1848) Symbolae ad floram Surinamensem. Euphorbiaceae Juss. Linnaea 21: 473–479.
Mittermeier, R.A., da Fonseca, G.A.B., Rylands, A.B. & Mittermeier, C.G. (1997) Brazil. In: Mittermeier, R.A., Robles Gil, P. &
Mittermeier, C.G. (Eds.) Megadiversity. earth’s biologically wealthiest nations. CEMEX, Mexico City, pp. 38–73.
Møller-Jørgensen, P., Nee, M. & Beck, S.G. (2014) Catálogo de las plantas vasculares de Bolivia. Missouri Botanical Garden Press, St.
Louis, 1741 pp.
Moore, S.L.M. (1895) The phanerogamic botany of the Matto Grosso Expedition, 1891–1892. Transactions of the Linnean Society of
London 4: 265–516.
Müller Argoviensis, J. (1865) Euphorbiaceae. Vorläufige Mittheilungen aus dem für De Candolle’s Prodromus bestimmten Manuscript
über diese Familie. Linnaea 34: 1–224.
Müller Argoviensis, J. (1866) Euphorbiaceae (excl. Euphorbieae). In: de Candolle, A.P. (Ed.) Prodromus systematis naturalis regni
vegetabilis 15(2). Treuttel et Würtz, Paris, pp. 189–1261.
Müller Argoviensis, J. (1873) Euphorbiaceae. In: Martius, C.F.P. von & Eichler, A.W. (Eds.) Flora Brasiliensis 11(2). Fleischer, München,
752 pp.
Müller Argoviensis, J. (1874) Euphorbiaceae novae. Journal of Botany, British and Foreign 12: 200–233.
Paña-Chocarro, M.C., Espada-Mateos, C., Vera, M., Céspedes, G. & Knapp, S. (2010) Updated checklist of vascular plants of the
Mbaracayú Forest Nature Reserve (Reserva Natural del Bosque Mbaracayú), Paraguay. Phytotaxa 12: 1–224.
Pax, F. (1897) Euphorbiaceae. In: Engler, H.G.A. & Prantl, K.A.E. (Eds.) Die natürlichen Pflanzenfamilien, Nachtrag und Register zu Teil
II–IV. Engelmann, Leipzig, 380 pp.
Pax, F. & Hoffmann, K. (1923) Euphorbiaceae americanae novae. I. Repertorium specierum novarum regni vegetabilis 19: 174–177.
Radcliffe-Smith, A. (1993) A new Croton variety of pharmacognostic significance from eastern Brazil. Kew Bulletin 49: 149–150.
Rafinesque, C.S. (1825) Neogenyton, or indication of sixty-six new genera of plants of North America. Lexington, Ky., 4 pp. Available
from: https://hdl.handle.net/2027/coo.31924073202768?urlappend=%3Bseq=7 (accessed 17 November 2018)
Reflora (2018) Herbário Virtual Reflora. Jardim Botânico do Rio de Janeiro. Available from: http://www.herbariovirtualreflora.jbrj.gov.
br (accessed 7 January 2018)
Ribeiro, J.F. & Walter, B.M.T. (1998) Fitofisionomias do Bioma Cerrado. In: Sano, S.M. & Almeida, S.P. (Eds.) Cerrado: ambiente e
flora. Empresa Brazileira de Pesquisa Agropecuária, Centro de Pesquisa Agropecuária dos Cerrados, Planaltina, pp. 89–166.
Rusby, H.H. (1927) Descriptions of new genera and species of plants collected on the Mulford Biological Exploration of the Amazon
Valley, 1912–1922. Memoirs of the New York Botanical Garden 7: 205–387.
Saint-Hilaire, A.F.C.P. (1827) Plantes usuelles des Brésiliens. Grimbert Libraire, Paris, 340 pp.
Schumann, K. (1900) Neue Arten der Siphonogamen 1898. Just’s Botanischer Jahresbericht 26 Abt. 1: 323–397.
Secco, R.S. (2008) Sinopse das espécies de Croton L. (Euphorbiaceae) na Amazônia brasileira: um ensaio taxonômico. Museu Paraense
Emílio Goeldi, Belém, 169 pp.
Secco, R.S. (2009) Uma nova espécie de Croton sect. Geiseleria (Euphorbiaceae) da Amazônia Oriental Brasileira. Revista Brasileira de
Botânica 32: 249–252.
https://doi.org/10.1590/S0100-84042009000200005
Secco, R.S., Rosário, A.S. & Berry, P.E. (2012) Croton campinarensis (Euphorbiaceae), a new species from eastern Amazonian Brazil.
Phytotaxa 49: 1–5.
Silva, J.S. (2009) O gênero Croton L. (Euphorbiaceae) em Pernambuco, com ênfase nas espécies da microrregião do Vale do Ipanema.
Master’s dissertation. Universidade Federal Rural de Pernambuco, Recife, 163 pp.
Silva, J.S., Sales, M.F. & Carneiro-Torres, D.S. (2009) O gênero Croton L. (Euphorbiaceae) na microrregião do Vale do Ipanema,
Pernambuco, Brasil. Rodriguésia 60: 879–901.
https://doi.org/10.1590/2175-7860200960409
Silva, J.S., Sales, M.F., Gomes, A.P.S. & Carneiro-Torres, D.S. (2010) Sinopse das espécies de Croton L. (Euphorbiaceae) no estado de
Pernambuco, Brasil. Acta Botanica Brasilica 24: 441–453.
https://doi.org/10.1590/S0102-33062010000200015
Silva, M.J., Sodré, R.C. & Berry, P.E. (2015a) Novelties in Croton (Euphorbiaceae) from Goiás, Brazil. Systematic Botany 40: 162‒167.
https://doi.org/10.1600/036364415X686468
Silva, M.J., Sodré, R.C. & Berry, P.E. (2015b) An enigmatic species of Croton (Euphorbiaceae) is rediscovered after more than 100 years
in midwestern Brazil. Phytotaxa 221: 83‒88.
Silva, M.J., Sodré, R.C. & Sales, M.F. (2014) A new species of Croton L. (Euphorbiaceae s.s.) from the Brazilian Cerrado. Systematic
Botany 39: 216–221.
https://doi.org/10.1600/036364414X678125
Smith, L.B, Downs, R.J. & Klein, R.M (1988) Euforbiáceas. In: Reitz, R. (Ed.) Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues,
Itajaí, pp. 1‒408.
Sodré, R.C., Berry, P.E. & Silva, M.J. (2017) The tribe Crotoneae (Euphorbiaceae, Crotonoideae) in the Chapada dos Veadeiros, Goiás,
Brazil. Phytotaxa 321: 1–59.
Sodré, R.C. & Silva, M.J. (2017a) An overlooked new species of Croton section Geiseleria (Euphorbiaceae) from the Brazilian Cerrado,
with a key to the species of this section occurring in the state of Goiás and in the Distrito Federal. Phytotaxa 329: 127–138.
Sodré, R.C. & Silva, M.J. (2017b) Croton rhodotrichus: A new species of Croton section Geiseleria (A. Gray) Baill. (Euphorbiaceae) from
the state of Mato Grosso, Brazil. Systematic Botany 42: 338–345.
https://doi.org/10.1600/036364417X695600
Sodré, R.C. & Silva, M.J. (2018a) A new species of Croton L. (Euphorbiaceae) from the wet fields of Midwestern Brazil. Journal of the
Torrey Botanical Society 145: 100–107.
https://doi.org/10.3159/TORREY-D-17-0007.1
Sodré, R.C. & Silva, M.J. (2018b) Rediscovery of Croton hadrianii (Euphorbiaceae), a well distributed but long-forgotten species in
Brazilian Amazon. Phytotaxa 349: 141–151.
Sodré, R.C., Souza, A.O., Alonso, A.A. & Silva, M.J. (2019) Molecular, morphological, and anatomical data support a new species of
Croton sect. Geiseleria (Crotonoideae, Euphorbiaceae) Plant Systematic and Evolution 305: 233–246.
https://doi.org/10.1007/s00606-019-1566-z
Spix, J.B. von & Martius, C.F.P. von (1823) Reise in Brasilien auf Befehl Sr. Majestät Maximilian Joseph I. Königs von Baiern in den
Jahren 1817 bis 1820. Erster Theil. M. Lindauer, München, 412 pp.
Steyermark, J.A. (1958) The Machris Brazilian expedition. Botany: Phanerogamae, Euphorbiaceae, Lentibulariaceae, Rubiaceae. Los
Angeles County Museum Contributions in Science 21: 3–21.
Swartz, O. (1788) Nova genera & species plantarum seu prodromus descriptionem vegetabilium maximam partem incognitorum quæ sub
itinere in Indiam Occidentalem annis 1783-87 digessit Olof Swartz. M. Swederi, Stockholm, Uppsala & Aboa, 152 pp.

Theobald, W.L., Krahulik, J.L. & Rollins, R.C. (1979) Trichome description and classification. In: Metcalfe, C.R. & Chalk, L. (Eds.)
Anatomy of the Dicotyledones, ed. 2, v. 1. Oxford, Claredon Press, pp. 40–53.
Thiers, B. (2018 [Continuously updated]) Index Herbariorum: a global directory of public herbaria and associated staff. New York
Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/ih/ (accessed 7 January 2018)
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Husber, W.-H., Li, D.-Z., Marhold,
K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for algae,
fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum
Vegetabile 159. [Glashütten: Koeltz Botanical Books]
https://doi.org/10.12705/Code.2018
Ule, E. (1908) III. Beiträge zur Flora der Hylaea nach den Sammlungen von Ules Amazonas-Expedition. Verhandlungen des Botanischen
Vereins der Provinz Brandenburg 50: 69–123.
Urban, I. (1906) Vitae itineraque collectorum botanicorum, notae collaboratorum biographicae. In: Martius, C.F.P. von, Eichler, A.W. &
Urban, I. (Eds.), Flora Brasiliensis 1, 1. R. Oldenbourg, Munich, pp. 1–154.
van Ee, B.W. (2011) The contribution of Johann Friedrich Klotzsch to the taxonomy of Croton (Euphorbiaceae) and associated genera.
Willdenowia 41: 15–33.
https://doi.org/10.3372/wi.41.41102
van Ee, B.W. & Berry, P.E. (2010) Typification notes for Croton (Euphorbiaceae). Harvard Papers in Botany 15: 73–84.
van Ee, B.W., Riina, R. & Berry, P.E. (2011) A revised infrageneric classification and molecular phylogeny of New World Croton
(Euphorbiaceae). Taxon 60: 791–823.
Weber, M.G., Porturas, L.D. & Keeler, K.H. (2015) World list of plants with extrafloral nectaries. Available from: http://www.
extrafloralnectaries.org/ (accessed 17 March 2018)
Webster, G.L. (1993) A provisional synopsis of the sections of the genus Croton (Euphorbiaceae). Taxon 42:93–823.
https://doi.org/10.2307/1223265
Webster, G.L., Berry, P.E., Levin, G.A., Esser, H.-J., Hayden, J., Armbruster, W.S., Secco, R.S. & Heald, S. (1999) Euphorbiaceae.
In: Steyermark, J.A., Berry, P.E., Yatskievych, K. & Holst, B.K. (Eds.) Flora of the Venezuelan Guayana. Euriocaulaceae–
Lentibulariaceae. Vol. 5. Missouri Botanical Garden Press, St. Louis, pp. 72–228.
Webster, G.L., Del-Arco-Aguilar, M.J. & Smith, B.A. (1996) Systematic distribution of foliar trichome types in Croton (Euphorbiaceae).
Botanical Journal of the Linnean Society 121: 41–57.
https://doi.org/10.1111/j.1095-8339.1996.tb00744.x
Wurdack, K.J. & Farfan-Rios, W. (2017) Incadendron: a new genus of Euphorbiaceae tribe Hippomaneae from the sub-Andean cordilleras
of Ecuador and Peru. Phytokeys 85: 69–86.
https://doi.org/10.3897/phytokeys.85.14757
Zuloaga, F., Morrone, O. & Belgrano, M. (2008) Catálogo De Las Plantas Vasculares Del Cono Sur. Monographs in Systematic Botany
107. [Missouri Botanical Garden Press, St. Louis]
APPENDIX 1. List of species of Croton sect. Geiseleria (Euphorbiaceae) occurring in Brazil and list of all the
herbarium specimens examined.
Numerical list of species:

1. Croton abaitensis Baill.
2. Croton aberrans Müll.Arg.
3. Croton adamantinus Müll.Arg.
4. Croton adenodontus Müll.Arg.
5. Croton agoensis Baill.
6. Croton antisyphiliticus Mart.
7. Croton arenosus Carn.-Torres & Cordeiro
8. Croton asperrimus Benth.
9. Croton bidentatus Müll.Arg.
10. Croton campinarensis Secco, A.Rosário & P.E.Berry
11. Croton carinatus Müll.Arg.
12. Croton catariae Baill.
13. Croton crustulifer Croizat
14. Croton desertorum Müll.Arg.
15. Croton glandulosus L.
16. Croton glechomifolius Müll.Arg.
17. Croton goyazensis Müll.Arg.
18. Croton gracilescens Müll.Arg.
19. Croton gracilirameus M.J.Silva, Sodré & P.E.Berry
20. Croton grewioides Baill.
21. Croton hadrianii Baill.
22. Croton harleyi Carn.-Torres & Cordeiro
23. Croton hirtus L.’Hér.
24. Croton hostmannii Schltdl. ex Miq.
25. Croton inaequilobus Steyerm.
26. Croton junceus Baill.
27. Croton krukoffianus Croizat
28. Croton lundianus (Didr.) Müll.Arg.
29. Croton macrosepalus Sodré & M.J.Silva
30. Croton mollis Benth.
31. Croton mucronifolius Müll.Arg.
32. Croton nepetifolius Baill.
33. Croton odontadenius Müll.Arg.
34. Croton parodianus Croizat
35. Croton planaltoanus M.J.Silva & Sodré
36. Croton pycnadenius Müll.Arg.
37. Croton ramosissimus Sodré & M.J.Silva
38. Croton rhodotrichus Sodré & M.J.Silva
39. Croton sclerocalyx (Didr.) Müll.Arg.
40. Croton seccoi Sodré & M.J.Silva
41. Croton sincorensis Mart. ex Müll.Arg.
42. Croton sipaliwinensis Lanj.
43. Croton spica Baill.
44. Croton subincanus Müll.Arg.
45. Croton sublepidotus Müll.Arg.
46. Croton tetradenius Baill.
47. Croton triangularis Müll.Arg.
48. Croton trinitatis Millsp.
49. Croton uliginosus Sodré & M.J.Silva
50. Croton virgultosus Müll.Arg.
List of all specimens examined:
Specimens examined are arranged in alphabetical order by first collector’s last name and initials, followed by collection
number. The corresponding species number from the list above is given in parentheses after each collection number.
Herbarium acronyms and barcode numbers are provided when collector number is missing. s.coll. = unknown collector,
s.n. = without collection number.
Albuquerque J.M. 45 (48); Alencar, M.E. 856 (40), 1098 (40), 1384 (40), 1504 (40); Allem, A. 2645 (12), 2883 (47),
2894 (46), 2906 (3), 3027 (31), 3068 (18), s.n. ICN 26971 (2); Alvarenga, D. 725 (12); Alves, M. 2176 (28), 2224 (43);
Amaral, A.G. 1635 (17); Amaral, I.L. 872 (34); Amorim, E.H. 321 (6); Amorim Neto, L. 848 (15); Amorim, A. 8606
(30); Amorim, A.M.A. 7409 (28), 9330 (46); Anderson, W.R. 6837 (12), 7304 (12), 9039 (31), 10716 (48), 10900 (27),
36539 (29), 36665a (29); Andrade Neto, M. s.n. EAC 0044186 (3); Antar, G.M. 271 (31), 375 (31); Antunes, L.L.C.
1299 (36), 1303 (36), 1327 (31), 1331 (31), 1333 (45), 1335 (45); Aoki, C. 312 (1), 371 (43); Araújo, A.O. 90 (6);
Arbo, M.M. 5621 (31); Árbocz, G. 3399 (12), 3405 (43), 6466 (48); Archer, W.A. 8184 (48); Argent, G. 6324 (26);
Armando, M. 1107 (25); Assunção, P.A.C.L. 1141 (27); Austin, D.F. 6975 (48), 4061 (48), 7014 (48);
Baldwin, J.T. 3056 (26); Barbosa, A. 415 (17), 657 (17), 668 (17), 916 (17), 965 (17); Barbosa, E. 240 (48); Barbosa,
R.I. 112 (15); Barreto, K.D. 3208 (6), 63305 (28); Barreto, M. 10444 (39); Batalha, M.A. 2242 (1), 2260 (1), 2370 (1),
2389 (1), 2397 (15), 3610 (1), 3739 (1); Batista, L.R. 26907 (2); Bauduíno-Rambo, S.J. s.n. PACA 9383 (2), s.n. PACA
9669 (2), s.n. PACA 9881 (2), s.n. PACA 53006 (2); Bautista, H.P. 40 (28); Beyrich, H.K. s.n. P 00623619 (9); Bigio,
N.C. 318 (15); Bilby, M.R.C. 12 (48); Black, G.A. 47-2094 (48), 50-9620 (48), 50-10287 (48), 54-16182 (34); Blake,
S.F. 7854 (48); Blanchet, J.S. 1665 (28), 1824 (28), 3100 (20), 3656 (15), 3786 (46); Boone, W. 821 (28); Bortolotto,
I.M. 1471 (48); Bosquetti, L.B. 448 (6); Bouças, P. 464 (48); Boudet Fernandes, H.Q. 1689 (41); Braga, M.M.N.
417 (6); Brandão, M. 19558 (28), 23909 (28), 26036 (28), 27638 (28); Braz, T. 351 (6); Brazão, C.M.S.C. 125 (21);
Browne, P. s.n. LINN 1140-7 (15); Burchell, W. 3910 (28), 5891 (6), 8913 (23), 8988 (36);
Cabral, F.N. 450 (30); Camargos, J.C. 80 (6); Campêlo, C.R. 1626 (23); Carneiro-Torres, D.S. 478 (23), 613 (36),
699 (36), 746 (47), 805 (3), 836 (40), 956 (20), 964 (3), 993 (7); Carreira, L.M.M. 503 (34), 2702 (24), 3306 (10),
3307 (10); Caruzo, M.B.R. 154 (26), 184 (16); Carvalho, A. 77 (40); Carvalho, F.A. 2145 (27); Carvalho, F.J. 150 (6);
Carvalho, S.M.F. 109 (17); Carvalho-Sobrinho, J.G. 1284 (48); Castro, D.B. s.n. UFMT-ICLMA 1443 (6); Castro,
K.N.C. 474 (46), 482 (32); Castro, W. 154 (21); Cavalcanti, T.B. 747 (36), 767 (36), 1911 (18), 3323 (18); Caxambu,
M.G. 1739 (6); Chacon, R.G. 1114 (1); Chagas, J. s.n. INPA 6326 (48); Chaves, E.M.F. 552 (32); Cid, C.A. 1691
(48); Claussen, P. 316 (39), 764 (46), 769 (6), 776 (39); Coêlho, D.F. s.n. INPA 92453 (15); Coelho, L. 827 (30);
Commerson, P. s.n. P 00623601 (15); Conceição, C.A. 1713 (48); Coradin, L. 103 (40); Cordeiro, I. 224 (27), 384
(1), 385 (1), 3371 (16), 3418 (1), 3463 (5) s.n. BHCB 013552 (6); Cordeiro, J. 2399 (28); Correia, C. 428 (22); Costa,
C.C.A. 66 (23); Costa, D.S. 860 (21); Costa, J.M. 179 (32); Costa, J.P. 10 (2); Costa, M.A.S. 1018 (48); Coutinho,
B.A. 59 (2), 66 (2); Cremers, G. 9537 (48); Crepaldi, C.G. 33 (3); Croat, T.B. 62652 (48); Crueger, H. s.n. TRIN (48);
Curtiss, A.H. 15 (48);
Dal Col, A.C.S. 131 (4); Daly, D.C. 982 (10), 1712 (21); Damasceno Júnior, G.A. 2586 (1); Dantas, M. 1072 (48);
Dantas, M.M. 131 (39); Davidson, C. 1035 (48), 10349 (42); Dávila, M.C. 5904 (27); Dawson, E.Y. 14685 (25);
Dél’Arco, M.R. s.n. TEPB 1359; Delfini, C. 642 (35); Delprete, P.G. 8927 (6), 8957 (6), 8975 (6), 9292 (1), 9294 (1),
9953 (1), 9962 (1), 10361 (1), 10425 (6), 10463 (6); Demaio, P. 312 (23); De Paula, J.R. 15 (37); Dias, T.A.B. 413 (37);
Dubs, B. 1784 (46); Duke, J.D. 6562 (23);
Echternacht, L. 456 (28); Eiten, G. 3601 (15), 3649 (31), 3654 (31), 3666 (5), 10336 (31), 10214 (42), 10381 (31)10869
(15); Ekman, E.L. 452 (2); Elias, S.I. 222 (6); Esgario, C. 32 (28); Estevan, D.A. 876 (6); Esteves, G.L. 2518 (31);
Estrela, M.R. 94-5476 (34);
Fagg, C.W. 2041 (36), 2062 (31); Faria-Júnior, J.E.Q. 42 (6) 1131 (18); Farias, R. 392 (36), 653 (28); Felix, L.P. 10392
(15), s.n. SP 332132 (20); Fendler, A. 1236 (23), 1236 β (23); Fernandes, A. 11077 (46); s.n. EAC 9099 (22), s.n. EAC
12149 (31); s.n. EAC 2633 (46); Ferreira, A.R. s.n. P 00623574 (21); Ferreira, F.M. 276 (6); Ferreira, H.D. 2274 (33),
243 (33), 2485 (6), 4067 (6), 4291 (37); Ferreira, M.B. 1405 (17); Ferreira, M.C. 26 (48), 126 (28); Ferreira, W.M.
1573 (6); Filgueiras, T.S. 1689 (39), 1708 (1); Figueiredo, S. 123 (6); Figueiredo, M.A. 257 (32); Fonseca, M.L. 59
(1), 570 (18), 2184 (1), 2190 (1), 2652 (36), 2653 (36), 2900 (5), 2909(5), 3637 (33), 3646 (17), 3913 (1), 5036 (15),
5616 (31), 6566 (36), 6622 (45), 6626 (36); Fontana, A.P. 6401 (22); Forzza, R.C. 4659 (31), 6974 (30); Fraga, C.N.
2258 (28); França, F. 3702 (36), 3716 (31); Francener, A. 507 (6); Franco s.n. HUEG 3650 (1); Freitas, J.C. 199 (48);
Freitas, R. 748 (6), 804 (6), 860 (6); Fróes, R.L. 11775 (34), 20149 (12), 34116 (48); Frutuoso, L.C. 61 (15), s.n. UFMT
12445 (15), s.n. UFMT 3357 (48);
Galeão, R. 10 (48); Gardner G. 113 (4), 944 (30), 1129 (28), 2722 (31), 3435 (15), 3437 (45), 3438 (11); Gasson, P.
5970 (50); Gates, H. 97 (33); Gaudichaud-Beaupré, C. 247 (12), 249 (43), 953 (6), 1137 (28); Gibbs, P. 2799 (35);
Gimenes, L.L.S. 5 (6); Ginzbarg, S. 655 (15), 670 (15), 671 (23), 687 (16), 723 (12), 777 (15), 798 (23), 862 (15), 852
(23); Ginzberger, A. 4348 (28); Glaziou, A.F.M. 4980 (28), 14254 (13); Goes, B.T.P.M. 142 (45); Gomes Jr. J.C. 1449
(2), 1468 (2) 1755 (15); Gomes, J.M.L. 531 (28), 2814 (28); Gomes-Klein, V.L. 3303 (6); Gouvêa, A.P.M.L. 82 (1);
Grandi, T.S.M. 119 (28), 1167 (6), 5748 (23); Guarino, E.S.G. 641 (23); Guedes, M.L. 2882 (31);
Hagelund, K. 4767 (2); Hassler, E. 1083 (2), 4529 (2), 6314 (2), 5604 (2), 6804 (28), 7933 (2), 7993a(2), 7933b (2),
9915 (16); Harley, R.M. 18083 (41); Hatschbach, G. 4313 (28), 12961 (23), 21819 (2), 34816 (12), 36040 (12), 43067
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6695 (39), 14163 (12), 14232 (15), 14274 (37), 16669 (23), 16710 (39); Hostmann, F.W.R. 1106 (24); Huber, J. 1096
(48); Hurbath, F. 251 (31);
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Jangoux, J. 374 (8); Jardim, J.G. 3647 (36); Júnior, J.E.C. 71 (6), 75 (6), 79 (6);
Kalkmann 141 (28); Kathriarachchi, H.S. 37 (23); Keller, A. 3722 (16); Kirkbride Jr., J.H. 3093 (43), 3939 (46); Klein,
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s.n. HUEFS 000084263 (3); Krukoff, B.A. 1480 (27); Kuhlmann, E. s.n. SP 69946 (2); Kuntze, O. s.n. NY 00503940
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8831 (2), s.n. ICN 8997 (2); Lisboa, P.L.B. 1286 (34); Loiola, M.I.B. 898 (23); Loefgren, A. 1027 (39); Lombardi,
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23291 (24), 30571 (27), 31731 (27), 40097 (44); Maia, L.A. 88 (48), 652 (42); Mantovani, W. 1450 (23), 12713
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639 (6), 705 (6), 744 (6); Medeiros, H. 1625 (21); Mello-Silva, R. 2224 (35), 2249 (18), 2300 (31), 2253 (14), 2344
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283 (6), 2533 (6); Monteiro, R.F. 406 (49); Moore, S.L. 826 (48), s.n. BM 000939427 (39); Moreira, J.C. 343 (23);
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(6); Murça Pires, J. 675 (48);
Nee, M. 11123 (48); Nelson, B.W. 1123 (48); Netto, L.S.M. s.n. BR 0000008764553 (6); Nicolack, V. 70 (6); Nóbrega,
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Paiva, V.F. 786 (6); Parralego, Y. 18 (6); Passos, F.B. 01 (17); Pastore, J.F.B. 350 (1), 1551 (31); Paulino-Filho H.F. 54
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1608 (36), 2976 (36); Petini-Benelli, A. 153 (42), 154 (42), 155 (42); Philcox, D. 3106 (12), 3707 (38); Pinto, C.G.P.
203 (36); Pinto, T.M.E.S. 11 (6); Pires, J.M. 56979 (49); Pires, R. 15 (6); Pivetta 553 (2); Plowman, T. 12078 (48);
Pohl, J.B.E. 297 (18), 826 (1), 1110 (1), 1611 (28), 1618 (17), 1626 (12), 1632 (6), 1637 (1), s.n. (W 0051286) (17);
Poole, J.M. 1725 (42); Porto, M.L. 1808 (2); Pott, A. 8334 (2), 9100 (46), 11301 (23), 12093 (6), 12154 (6), 14823
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Queiroz, L.P. 2024 (31), 2081 (36), 2119 (31), 4110 (31), 3916 (31); Queiroz, R.T. 87 (46);
Raben, F.C. 763 & 783 (28); Rambo, B. s.n. PACA-AGP 3047 (16); Ramos, A.E. 313 (1); Ramos-Júnior, E. 67 (15);
Rapini, A. 725 (47); Ratter, J.A. 4080 (12); Rebellato, L. 207 (15) 207*(48); Regnell, A.F. I-400 (6), III-1086 (6), III-
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Resende, U.M. 308 (4); Rezende, A.V. 88 (31), 122 (31); Ribas, O.S. 1265 (15); Ribeiro, B.G.S. 1758 (48), s.n. HRB
12817 (42); Ribeiro, D.G. 30 (49); Ribeiro, T. 297 (47); Richard, L.C. s.n. P 00623551 (23); Riedel, L. 65 (4), 263
(16), 467 (36), 546 (2), 869 (46), 1136 (43), 1252 (3), 1626 (39), 2473 (17), 2808 (6), s.n. (LE 00003073) (33), s.n.
(LE 00018215) (6); Riina, R. 1353 (47); Rios, M.N.S. 151 (17); Rizzo, J.A. 712 (17), 1909 (17), 2848 (6), 2851 (39),
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(39), 12500 (6), 12641 (6), 12691 (6), 12700 (39), 12871 (39); Rocha, M. 41 (1), 69 (1); Rodrigues, A.S. 158 (1);
Rodrigues, E. 48 (48); Rodrigues, S.M. 230 (15); Rodrigues, W. 721 (30), 2043 (30), s.n. INPA 942 (48); Rombouts,
H.E. 357 (42); Romero-Castañeda 1678 (48); Rosa, N.A. 3743 (28); Rosas, A. 46 (48); Roveratti, J. 558 (1); Rusby,
H.H. 29 (48), 1332 (48);
Saavedra, M.M. 753 (28); Sagot, P.A. 503 (23); Saint-Hilaire, A.F.C.P. 1115 (28), cat B1 1500 (46), cat B1 2063 (32),
cat B2 2115 (32), cat C1 n. 227 (6), cat C1 277 (39), cat. C1 n. 1058 (6), cat C1 n. 1116 (28), cat. C2 262-b (28), cat.
C2 n. 6118 (6), cat. C2 1387 (16), cat. C2 n. 1393 bis (6), s.n. P 00623665 (5); Sakuragui, C.M. 453 (6); Sales, M.F.
561 (46); Salino, A. 10422 (28), 10603 (28); Sampaio, S. 17 (28); Sanchez, M. 2011 (6), 2020 (6), 2217 (6); Santos,
E.R. 1791 (31), 2312 (31); Santos, J.P. 609 (36), 650 (31), 659 (36), 662 (31), 675 (5), 739 (39), 751 (6), 772 (39), 828
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(21), 1931 (21), 2128 (21); Scaramuzza, C.A.M. 525 (6); Scariot, A.O. 678 (11), 824 (45), 825 (45); Schessl, M.
260992-1-7 (48), 131092-4-1 (48), 281092-177/2 (15), 3203b (48), 4309 (48), 5273 (48); Schiede, C.J.W. 62 (15);
Schmidt, F.H. 9680 (30); Schomburgk, Rob. 101 (23), 241 (48); 665 (44); Schomburgk, Rich. 851 (42) Schwacke
C.A.W. 5607 (4); Secco, R.S. 124 (21), 877 (15), 879 (42), 928 (10); Sellow, F. 1242 (47), s.n. BR 0000008710833
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Sieber Fl. Trinit. 106 (23), Silva, A.S.L. 1761 (21); Silva, C.M. 70 (23); Silva, G.P. 1967 (1), 4346 (18), 4580 (12),
5501 (37), 5754 (23), 5896 (12), 5927 (15), 6866 (37), 7173 (12), 8647 (23), 11143 (12), 12347 (18), 13973 (21),
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(1), 6866 (1), 6895 (15), 7484 (1); Silva, M.C. 762 (17); Silva, M.F.F. 28 (42), 1056 (36); Silva, M.L. 31 (50); Silva,
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(33), 4084 (33), 4087 (33), 4132 (33), 4133 (33), 4161 (1), 4162 (1), 4163 (1), 4186 (33), 4201 (33), 4488 (35), 4489
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(33), 4822 (33), 4830 (19), 4832 (19), 4888 (25), 5079 (1), 5010 (25), 5013 (25), 5084-B (12), 5210 (25), 5221 (25),
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(1), 5852 (12), 5853 (12), 5854 (1), 6515 (38), 6516 (38), 6517 (38), 6518 (38), 6520 (38), 6748 (25); Silva, N.T. 1516
(48), 2638 (48), 4200 (48); Silva, O.L.M. 42 (15), 293 (47); Silva, R.A. 1569 (20); Silva, R.R. 215 (39), 734 (6); Silva,
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1889 (27), 2330 (12); Skorupa, L.A. 77 (1), 112 (18), 555 (29), 654 (6); Smith, L.B. 8438 (6), 9237 (2), 9338 (2), 9572
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(48), 8401 (21), 8441 (48), 8484 (21); Sobral, M. 6477 (15), 10394 (39), 11070 (27), 11138 (42); Sobrinho, F.E. 145
(23); Sodré, R.C. 04 (14), 08 (14), 09 (14), 19 (14), 26 (33), 29 (33), 34 (12), 37 (12), 38 (12), 42 (14), 47 (19), 48 (19),
50 (33), 54 (33), 55 (33), 71 (14), 74 (14), 75 (14), 77 (14), 78 (1), 79 (1), 80 (1), 81 (1), 83 (14), 84 (14), 85 (14), 86
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(14), 174 (14), 176 (14), 179 (14), 186 (14), 187 (14), 188 (14), 236 (35), 237 (35), 238 (35), 239 (35), 240 (35), 241
(35), 257 (19), 268 (19), 270 (19), 275 (19), 322 (37), 326 (37), 327 (37), 329 (37), 366 (33), 367 (33), 268 (33), 373
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753 (33), 758 (33), 763 (37), 764 (37), 773 (12), 786 (25), 787 (25), 788 (25), 812 (25), 815 (25), 819 (33), 838 (12),
853 (17), 854 (17), 855 (17), 856 (17), 857 (17), 858 (17), 859 (17), 860 (17), 862 (17), 954 (12), 985 (1), 990 (1), 991
(1), 997 (1), 998 (1), 999 (1), 1000 (1), 1015 (37), 1016 (37), 1017 (37), 1018 (37), 1032 (12), 1034 (12), 1035 (12),
1036 (12), 1037 (12), 1038 (12), 1043 (18), 1052 (12), 1069 (43), 1070 (43), 1071 (43), 1072 (43), 1074 (43), 1077
(43), 1089 (39), 1090 (39), 1096 (39), 1099 (39), 1164 (12), 1166 (1), 1167 (1), 1168 (12), 1174 (12), 1178 (23), 1184
(12), 1194 (12), 1211 (12), 1216 (14), 1220 (1), 1242 (18), 1250 (12), 1288 (39), 1305 (14), 1313 (14), 1314 (14), 1361
(33), 1371 (12), 1386 (14), 1413 (18), 1414 (18), 1415 (14), 1416 (14), 1417 (18), 1418 (14), 1419 (14), 1420 (18),
1421 (14), 1422 (17), 1423 (17), 1425 (14), 1439 (17), 1446 (35), 1466 (35), 1469 (35), 1470 (14), 1477 (18), 1478
(18), 1480 (35), 1481 (35), 1482 (35), 1485 (35), 1493 (17), 1494 (14), 1496 (14), 1497 (14), 1525 (14), 1575 (1), 1576
(33), 1585 (33), 1588 (33), 1577 (1), 1581 (1), 1584 (1), 1602 (33), 1624 (1), 1628 (14), 1712 (38), 1721 (38), 1739
(38), 2150 (48), 2151 (48), 2152 (48), 2153 (48), 2154 (48), 2155 (48), 2156 (48), 2157 (48), 2158 (48), 2159 (48),
2160 (48), 2161 (48), 2162 (48), 2163 (48), 2164 (48), 2165 (48), 2169 (43), 2170 (43), 2171 (43), 2172 (43), 2175
(43), 2176 (43), 2177 (43), 2178 (43), 2196 (6), 2197 (48), 2198 (48), 2199 (48), 2200 (48), 2201 (48), 2208 (39), 2218
(43), 2222 (6), 2223 (6), 2225 (43), 2227 (6), 2229 (43), 2230 (38), 2231 (38), 2232 (38), 2233 (38), 2234 (38), 2235
(38), 2236 (38), 2239 (38), 2242 (1), 2244 (43), 2248 (38), 2249 (38), 2250 (38), 2251 (38), 2252 (38), 2253 (38), 2254
(38), 2256 (48), 2257 (48), 2258 (48), 2259 (48), 2260 (48), 2261 (48), 2263 (43), 2266 (48), 2269 (43), 2270 (43),
2272 (6), 2327 (43), 2328 (43), 2329 (43), 2376 (39), 2377 (39), 2378 (39), 2379 (39), 2380 (39), 2381 (49), 2382 (49),
2383 (49), 2384 (49), 2385 (49); 2386 (49), 2389 (49), 2390 (49), 2391 (49), 2392 (49), 2393 (49), 2394 (49), 2395
(49), 2396 (49), 2397 (49), 2398 (49), 2518 (17), 2519 (17), 2520 (1), 2526 (1), 2527 (1), 2528 (39), 2529 (1), 2530
(1), 2531 (1), 2532 (1), 2578 (39), 2579 (37), 2580 (37), 2581 (37), 2582 (37), 2583 (37), 2584 (49), 2585 (49), 2586
(49), 2587 (49), 2588 (49), 2598 (26), 2599 (26), 2600 (26), 2601 (26), 2602 (26), 2604 (26), 2605 (26), 2606 (26),
2607 (26), 2608 (26), 2609 (26), 2610 (26), 2611 (26), 2612 (26), 2636 (2), 2668 (37), 2669 (37), 2670 (37), 2671 (37),
2672 (37), 2673 (37), 2674 (37), 2675 (37), 2676 (37), 2681 (39), 2726 (43), 2727 (43), 2728 (43), 2729 (43), 2730
(43), 2731 (43), 2768 (43), 2770 (43), 2772 (43), 2777 (43), 2778 (43), 2779 (43), 2780 (43), 2781 (43), 2782 (43),
2783 (43), 2784 (43), 2785 (43), 2786 (43), 2790 (43), 2797 (43), 2803 (39), 2814 (49), 2815 (49), 2816 (49), 2817
(49), 2818 (49), 2819 (49), 2821 (26), 2822 (26), 2823 (26), 2824 (26), 2825 (26), 2826 (37), 2827 (37), 2828 (37),
2829 (37), 2830 (23), 2838 (43), 2839 (43), 2846 (39), 2850 (36), 2851 (36), 2852 (36), 2853 (36), 2854 (36), 2855
(36), 2856 (36), 2857 (36), 2858 (36), 2871 (39), 2872 (21), 2873 (21), 2874 (21), 2875 (21), 2876 (21), 2877 (21),
2878 (21), 2879 (21), 2880 (21), 2881 (21), 2892 (37), 2893 (37), 2894 (37), 2896 (37), 2899 (43), 2906 (43), 2907
(43), 2908 (17), 2913 (17), 2915 (17), 2916 (17), 2918 (17), 2919 (39), 2920 (39), 2942 (5), 2943 (5), 2944 (5), 2945
(5), 2984 (39), 2979 (29), 3100 (34), 3101 (34), 3102 (34), 3103 (34), 3104 (34), 3105 (34), 3106 (34), 3107 (34), 3108
(34), 3109 (34), 3110 (8), 3125 (34), 3128 (34), 3129 (34), 3130 (34), 3131 (34), 3132 (34), 3133 (21), 3134 (21), 3135
(21), 3136 (21), 3138 (31), 3145 (4), 3146 (4), 3147 (4), 3148 (4), 3149 (4), 3150 (4), 3151 (4), 3152 (4), 3153 (4),
3154 (4), 3155 (4), 3176 (2), 3177a (29), 3177b (29), 3185 (29), 3189 (11), 3192 (29), 3196 (29), 3197 (29), 3198 (29),
3199 (29), 3200 (1), 3201 (11), 3202 (11), 3203 (11), 3295 (47), 3312 (32), 3337 (22), 3338 (22), 3339 (22), 3340 (22)
3343 (46); Sousa, S. 419 (6); Sousa, G. 266 (28); Souza, A.O. 1205 (5), 1221 (31), 1519 (31), 1617 (39), 1912 (28);
Souza, E.B. 1960 (27), 4236 (40), 4246 (40); Souza, J.P. 210 (6), 295 (6), 4134 (43), 4621 (46), 4773 (29), 7506 (15),
9187 (45), 11571 (43), 11718 (33); Souza, L.F. 70 (6), 443 (37), 1284 (37), 1329 (37), 1358 (37), 1404 (37), 1489 (37),
1668 (37), 4630 (49), 5296 (6), 5305 (6), 5928 (49); Souza, M.C. 224 (48); Souza, V.C. 2089 (6), 2369 (6), 4175 (16),
4253 (6), 5358 (17), 5365 (17), 5371 (17), 7106 (6), 7335 (6), 14072 (15), 14096 (15), 14638 (48), 14733 (39), 15543
(48), 15610 (21), 16904 (36), 20081 (6), 20125 (6), 20187 (6), 20267 (6), 20517 (6), 20555 (6), 20704 (6), 20816 (6),
21314 (6), 21558 (18), 23907 (18), 26309 (31), 26599 (15); Souza, W.S. 25096 (15); Souza-Silva, R.F. 308 (47);
Spruce, R. 478* (8), 1806 (30), 4237 (23), s.n. G 00312828 (15); Stannard, B. 5358 (36); Stefanello, T.H. 364 (48);
Stephan, M.D. s.n. BR 0000008762573 (6);
Tamashiro, J.Y. 741 (6); Tamberlik s.n. W 0051319 (43), W 0060012 (15); Tameirão Neto, E. 413 (28), 612 (6), 4314
(43); Teixeira, L.O.A. 850 (42), 1414 (21); Teles, S.B. s.n. UFG 19200 (39); Thomas, W. 9608a (20); Torres, R.B.
18446 (21); Traill, J.W.H. 730 (8); Tuma, I.L. 45 (6), 63 (17);
Ule, E.H.G. 5330 (30);
Vale, G.D. 37 (37), 38 (6); Vasconcelos, S.H. 70 (23), 71 (23), 72 (23); Verboonem, S.M. 120 (43), 134 (18); Vidal, J.
6606 (28); Vidal, J.R.B. 164 (17); Vieira, A.V. s.n. EAC 34235 (15); Vieira, R.F. 596 (1), 1051 (46), 1150 (47), 1380
(46); Vieira, M.G.G. 1356 (27); Vilhalva, D.A.A. 77 (15); Viollati, L.G. 70 (31), 154 (31), 226 (29);
Walter, B.M.T. 592 (19), 1070 (39), 4529 (12), 4541 (6); Wanderley, M.G.L. 1246 (26); Warming, E. 1572 (33),
1616 (28), 1628 (46), 1639 (6), 1640 (39), 1641 (33), 1642 (33), 1646 (15); Weddel, H.A. 821 (28), 1086 (6), 1818
(1), 2753 (18), 2940 (26), s.n. P 00623523 (26); Widgren, J.F. 348 (39), s.n. LD 18844359 (6), s.n. S R-10551 (33);
Wied-Neuwied, M.A. s.n. BR 0000008762245 (9), s.n. M 0086076 (28); Wood, J.R.I. 26937 (6); Woolston, A.L. 1220
(15);
Zanatta, M.R.V. 907 (37), 976 (37), 1052 (46), 1171 (1); Zanzini, L.P. 76 (21); Zehntner, L. 4084 (20); Zenaide, H. 8
(23), 24a (20);
s.coll. s.n. S-R-1305 (15).

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Phytotaxa 417 (1): 001–105 ISSN 1179-3155 (print edition)

Taxonomic synopsis of Croton section Geiseleria

Accepted by Hajo Esser: 5 Sept. 2019; published: 19 Sept. 2019 1

FIRST PUBLISHED IN 2019 BY

© 2019 Magnolia Press

ISSN 1179-3155 (print edition)

2 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Key words: South America, Crotoneae, endemism, taxonomy, typification

4 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Material and methods

TABLE 1. Field expeditions conducted throughout this study.

6 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Morphological aspects of species of Croton sect. Geiseleria in Brazil

8 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

10 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

12 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

14 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

16 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

18 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

20 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Key to the species of Croton sect. Geiseleria occurring in Brazil

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24 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

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Vernacular name:—canela-de-urubu, velame-bravo.

30 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—alcachofrinho, algodãozinho, malvinha, pustemeira, velame.

32 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—alcanforeira, alcânfora, canela-de-perdiz, cocaleira, curraleira, erva-mular, minuano, pé-de-

34 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

38 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

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Vernacular name:—gervão, gervão-branco, malva-vermelha, velame.

TABLE 3. Morphological comparison between C. glandulosus and related species.

Length of stigmas (mm) 2–2.5 2–2.5 2.5–3 0.8–1

Representative specimens examined:—ARGENTINA: Corrientes: Ituzaingó, Ruta 34, ca. 3 km da ruta

46 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—pé-de-perdiz, pé-de-perdiz-fêmea, pé-de-perdiz-lisa.

48 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—alecrim-de-cabocla, angico, angélica, caatinga-de-cheiro, canelinha, canelinha-de-cheiro,

50 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

54 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—croto, malva-de-pêlos, malva-grande.

56 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

58 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

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Vernacular name:—gervão, gervão-miúdo, gervão-branco.

TABLE 4. Comparative morphology of Croton macrosepalus and related species.

Representative specimens examined:—BRAZIL. Bahia: Correntina, Fazenda Jatobá, 13°–14°S, 45°45’–

64 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

66 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—alcachofrinho, malvinha, pustemeira, velame.

68 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

Vernacular name:—canelinha-de-areia, malva-peluda, malva-preta, marmeleiro-preto, marmeleiro-roçarento,

70 • Phytotaxa 417 (1) © 2019 Magnolia Press SODRÉ ET AL.

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