Revised delimitation of Croton campestris (Euphorbiaceae), including description of two new species, molecular phylogenetic, anatomical and micromorphological data

Croton section Adenophylli is the largest section in the genus, with around 223 species distributed in the neotropical region. One of the main diagnostic characteristics of this section is the fruit columella, which has three prominent ascending terminal appendages. During a taxonomic review of this section in Brazil, we observed that C. campestris has a problematic taxonomic circumscription, thereby being confused with some other congeners. We investigate the circumscription of C. campestris through the examination of several herbarium specimens across its entire geographic range, and in its different habitats. We conducted macro-and micro-morphological studies as well as molecular analyses based on DNA sequences. Our results allow us to propose two new species ( C. hatschbachii and C. stellatorotatus ), update the nomenclature of C. campestris and C. subvillosus , and define the systematic positions, phylogenetic relationships, morphology and leaf anatomy of the above-mentioned taxa. The species are described, and their geographic distributions, morphological relationships, flowering and fruiting times, and conservation statuses are commented on. The two new species are also illustrated. Seventeen new synonyms, thirteen lectotypes and two neotypes are proposed. We also provide a key for the identification of 11 taxa belonging to the section Adenophylli , previously confused with C. campestris .


Introduction
Croton is the second largest genus of Euphorbiaceae, with from 1200 to 1300 species widely distributed in the tropics and subtropics (Govaerts et al. 2000;Berry et al. 2005).
These species inhabit dry or moist vegetation regions and exhibit habits ranging from small herbs, shrubs, and lianas to large trees, and display a wide diversity of trichomes, floral and extrafloral nectary glands, and interesting floral patterns (van Ee et al. 2011;Thaowetsuwan et al. 2020).Croton is represented by approximately 700 species in the C. heliotropiifolius, C. lanatus, C. subacutus, C. subvillosus and two undescribed species (see Tab. 1).Detailed examinations of all of these species led us to: a) update the circumscriptions of C. campestris as well as C. subvillosus -a species that is often confused with the former in taxonomic and floristic studies of the genus in the midwestern region of Brazil (Sodré et al. 2014;2017;Secco et al. 2018); b) describe and illustrate C. hatschbachii and C. stellatorotatus as new species; c) propose 17 new synonyms and 14 lectotypes for names related to C. campestris; and d) update the geographic distributions, conservation statuses, and phenologies of all recognized species, with illustrations, photographs and maps.We also compared C. campestris, C. hatschbachii, C. stellatorotatus and C. subvillosus in terms of the anatomies of their leaves and the micromorphologies of their foliar trichomes, as studies of that nature have frequently been found to be valuable for separating taxa in the genus (e.g., Caruzo et al. 2016;Sodré et al. 2019;Sodré & Silva 2020).We positioned the new species, as well as C. campestris and C. subvillosus, in a molecular phylogeny derived from trnL-F (cpDNA) and ITS (nrDNA) markers to confirm their phylogenetic positions and better understand the relationships between them.We also present a key to differentiate C. campestris from the other 11 species of Croton sect.Adenophylli with which it has previously been confused.

Phylogenetic Studies
Our sampling included accessions of 69 species of all four subgenera of Croton L., among them 34 species of the section Adenophylli Griseb., ten species of section Cyclostigma Griseb., C. chimboracensis P.E.Berry  DNA was extracted from silica-dried specimens following the cetyltrimethyl ammonium bromide (CTAB) protocol of Doyle & Doyle (1987) with modifications.ITS and trnL-F regions of DNA were amplified with PCR and sequenced with primers from White et al. (1990), Taberlet et al. (1991) and Urbatsch et al. (2000) followed the same laboratory procedures as in previous phylogenetic studies in the genus (van Ee et al. Revised delimitation of Croton campestris (Euphorbiaceae), including description of two new species, molecular phylogenetic, anatomical and micromorphological data 2011; Riina et al. 2009).The sequences were aligned using the CLUSTALW in MEGA version 6 (Tamura et al. 2013).The data matrix obtained was analyzed by Bayesian Inference, using MrBayes version 3.1.2(Ronquist & Huelsenbeck 2003).The evolutionary model was previously selected by JModelTest 2.1.5(Darriba et al. 2012) based on Akaike Information Criterion (AIC) (Akaike 1973).The model used to analyze the ITS datasets was SYM + G and to analyze the trnL-trnF was TVM + G).The analysis consisted of two independent runs of four Markov chain Monte Carlo, each run with 10,000,000 generations, with trees sampled every 1,000 generations.After 25 % of the initial samples collected (burn in) being discarded, the remainder were used to determine the values of posterior probability (PP).The statistical support of each node of the cladograms obtained was estimated through the bootstrap statistics (Felsenstein 1985) from 10,000 heuristic pseudo-replicates using a random addition of taxa and TBR branch swapping performed in PAUP* version 4.0b10.Wellsupported clades are defined as those with Bayesian posterior probabilities (PPs) and bootstrap support (BS) greater than or equal to 0.95 and 85 %, respectively.

Scanning Electron Microscopy
For the scanning electron microscopy (SEM) analyses, the leaves of the two new species, C. subvillosus and C. campestris were removed from field and herbarium specimens, fixed on stubs, and coated with gold using a spray applicator (Leica EM SCD050, Heidelberg, Germany).The samples were examined using a scanning electron microscope (Zeiss EVOMA100, Jena, Germany) at the High-Resolution Microscopy Multiuser Laboratory (LabMic), at the Institute of Physics, Federal University of Goiás (UFG).The trichomes were classified according to the terminology of Webster et al. (1996).
The conservation statuses of the species were assessed according to IUCN (2017) criteria, with the Extent of Occurrence (EOO) calculated using the Geospatial Conservation Assessment Tool GeoCAT (Bachman et al. 2011).Our descriptions follow the terminology of the specialized literature on Croton.Geographic distributions were based on georeferenced data from the herbarium specimens analyzed, and were plotted using QGIS v. 2.14 (QGIS 2020 development team).
The typifications of the species were based on literature review and consultations of high-resolution photographs of type collections available on the aforementioned websites.The lectotype proposals follow article 9.3 of the Shenzhen Code (Turland et al. 2018).When the protologue of a specific taxon indicated an herbarium with only a single exsiccatae type collection, it was considered the holotype, while any other exsiccatae in other herbaria were considered isotypes.However, when the taxon was described without any mention of a host herbarium, and if only a single exsiccatae was found, we chose to treat it as a syntype (considering that other exsiccatum from that same collection might be found in other herbaria during further studies).
The synonymizations proposed here are based on analyses of protologue descriptions, type collections, and variations found in populations in their respective areas of occurrence.None of the varieties or forms previously described for the species are recognized here, as we consider the characteristics that differentiate them as being continuous between populations of their respective species.Future molecular and morphological studies will need to increase the sampling of the section Adenophylli, including species previously admitted in the section Velamea and species related to C. chimboracensis.Although we don't admit C. chimboracensis as a member of section Adenophylli, due to low or absent support in the ITS and trnL-F analysis, it will have fundamental importance in future studies with the section.This species appears as a sister to the C. sect.Adenophylli in ITS analysis, and still presents differentiated columella with an entire apex, rare in species of the section, which can help in the investigation of how this unique pattern of fruit columella emerged in the genus.Regarding the sampling of the C. sect.Velamea, it may come to confirm whether the group could emerge as a subsection within the section Adenophylli.-I, 4J, K) varying in length, number, size, and arrangement of their rays.The trichomes have slightly thickened cell walls, and some of them show hexagonal cell structures with transverse openings externally -interpreted here as secretory structures of epidermal origin (Fig. 4P).Regarding their density, the trichomes are slightly dense in C. campestris (Fig. 3A), sparsely distributed in C. hatschbachii (Fig. 3D) and C. subvillosus (Fig. 3J), and strongly dense in C. stellatorotatus (Fig. 3G).The stellateporrect trichome type is characterized by having lateral rays (7)(8)(9)(10)(11)(12)(13) arising in a single plane (Fig. 3B, C, J, K) (with a more developed central ray (0.2-1 mm), called the porrect ray (Fig. 3B, C, J, K)) on both leaf surfaces of C. campestris (Fig. 4A) and C. subvillosus (Fig. 4N, O).The multiradiate trichomes of C. hatschbachii show 16 to 30 lateral rays arising in 2 or 3 whorls (Fig. 3E, F).The stellate-rotate trichomes show 13-20 lateral rays joined along 1 / 10 of their lengths, and (usually) a vestigial central ray (20-50 µm long); they are exclusive to C. stellatorotatus among the species studied here (Fig. 3H, I).The species also differ in terms of the lengths of the lateral rays of the trichomes, being (usually) short in C. hatschbachii (250-350 µm), of medium length in C. stellatorotatus (300-600 µm) and C. campestris (350-1000 µm), and long in C. subvillosus (500-2000 µm).

Leaf anatomy
The leaves of C. campestris are hypostomatic, but amphistomatic in C. hatschbachii, C. stellatorotatus and C. subvillosus, with stomata distributed at the same level as the common epidermal cells, with discrete substomatal chambers (Fig. 4C, G, L, Q).The epidermis in all species is unistratified, has rectangular and quadrangular cells with straight internal periclinal and anticlinal walls, and slightly sinuous external periclinal walls covered by a thin cuticle (e.g., Fig. 4I, L).Croton hatschbachii also shows a discontinuous hypodermis composed of longitudinally rectangular cells with thin walls on the adaxial surface (Fig. 4S).All of the species also show sessile or shortly stipitate ovoid, ellipsoidal, or subglobose colleters along the leaf margins, or on the base of the leaf blade, as seen in C. campestris (Fig. 4T).
The mesophyll is dorsiventral, with a palisade parenchyma composed of a layer of juxtaposed cells in C. campestris, C. hatschbachii, and C. stellatorotatus (Fig. 4C, G, L), and one or two layers in C. subvillosus (Fig. 4Q), as well as spongy parenchyma composed of cells of different sizes and shapes, with conspicuous intercellular spaces (Fig. 4C, G, L, Q).Druses occur in the palisade and spongy parenchyma of all species (e.g., C. hatschbachii (Fig. 4H), and C. subvillosus, (Fig. 4Q)), as well as laticifers, as seen in C. hatschbachii (Fig. 4S).The leaf margins of all species have obtuse outlines, with parenchymatous cells in subepidermal positions (Fig. 4D, I, M, R).
The vascular bundles are collateral and of large calibers, with annular collenchyma cells adjacent to the phloem on the abaxial surface (Fig. 4U); the smaller caliber vascular bundles are surrounded by a conspicuous parenchymatous sheath (Fig. 4V). Figure 4u shows cells of the epidermis with external periclinal walls with conspicuous pectocellulosic thickenings.
Midrib -the indument in that region is composed of the same types of trichomes as the leaf blade itself.The epidermis is unistratified, and composed of rectangular, rounded, or quadrangular cells with thin walls, covered by a thin cuticle, and without stomata (Fig. 5A-H).
The cortex consists predominantly of annular and angular collenchyma, and the ground parenchyma has rounded cells of different sizes (Fig. 5 A-H).A palisade parenchyma continuity is also noted between the two faces of the leaf blade (Fig. 5A, C, E, G).Druses were observed throughout the cortex tissues on both surfaces (e.g., C. campestris (Figs.5B, I) and C. hatschbachii (Fig. 5D)), and laticifers Rodolfo Carneiro Sodré, Alexandre Antônio Alonso and Marcos José da Silva Revised delimitation of Croton campestris (Euphorbiaceae), including description of two new species, molecular phylogenetic, anatomical and micromorphological data Rodolfo Carneiro Sodré, Alexandre Antônio Alonso and Marcos José da Silva Revised delimitation of Croton campestris (Euphorbiaceae), including description of two new species, molecular phylogenetic, anatomical and micromorphological data  (showing different thicknesses of their primary walls) are dispersed among the tissues adjacent to the vascular bundle (Fig. 5B, D, F, H), and the walls of the species studied here show distinct outlines (Fig. 5K-N).
The vascular bundles are collateral and U-shaped, with phloem surrounded by a pericycle of parenchyma cells (Fig. 5A-H).Druses were observed only in the phloematic parenchyma of C. campestris and C. hatschbachii (Fig. 5B, D).
Petiole -The petiolar indumenta are similar to those described for the leaf blades and midribs, with stellateporrect trichomes having secretory structures of epidermal origins (Fig. 6H, L).The petiolar outline in the cross section is planar-convex (Fig. 6A, C, E, G), with the epidermis composed of common round cells with a thin cuticle, but without stomata.
The cortex is composed of annular collenchyma and ground parenchyma, both with cells of varying sizes (e.g., C. campestris (Fig. 6B) and C. stellatorotatus (Fig. 6F)).Druses were found in the petioles of all species, as exemplified by C. hatschbachii (Fig. 6D).
Additional notes -The anatomical characteristics observed in the studied species, their trichome types, the presence of a discontinuous hypoderm, the distribution patterns of the stomata, the presence of accessory vascular bundles in the petiole, the presence libriform fibers in the pericycle, and the numbers of vascular bundles in the petiole all proved useful in species differentiation (Tab.2), as was reported by Barros & Soares (2013) and Vitarelli et al. (2015) for other Croton species.Among these features, the types of trichomes can be highlighted, as they have been used to differentiate species in the genus even since Müller Argoviensis (1873), and were emphasized by Webster et al. (1996) and other authors such as Inamdar & Gangadhara (1977), Nepumoceno & Oliveira (1979) and Lucena & Sales (2006).Despite the taxonomic value of the trichomes in some cases at both sectional and species levels, we emphasize that different type of trichomes appear to have evolved independently within the genus, as they occur in taxa from distinct sections.The stellate-rotate trichomes reported here for C. stellatorotatus, for example, were cited by Webster et al. (1996), Gordillo & Matías (2005) and Gomes et al. (2018), for members of the sections Lasiogyne, Barhamia, and Astraeopsis.respectively; the multiradiate trichomes of C. hatschbachii were likewise observed in taxa belonging to the sections Barhamia, Cyclostigma, and Geiseleria by Gordillo & Matías (2005), Feio et al. (2018), and Sodré et al. (2019) respectively.Similarly, the stellateporrect trichomes observed here in C. campestris and C. subvillosus, seem to be the most frequent in the genus (e.g., Lucena & Sales 2006;Carneiro-Torres et al. 2011;Feio et al. 2018).We also highlight the structures interpreted here as trichomes truly represent those structures -unlike structures with similar appearances reported for other species of the genus by Vitarelli et al. (2016) and Sodré et al. (2019), which are simply trichome-like emergences, not found in the species here studied.Those observations evidence the micromorphological richness found in Croton, and likewise point to the need to determine the correct type of indument in the genus.
Rodolfo Carneiro Sodré, Alexandre Antônio Alonso and Marcos José da Silva Revised delimitation of Croton campestris (Euphorbiaceae), including description of two new species, molecular phylogenetic, anatomical and micromorphological data Leaf anatomical characters such as a unistratified epidermis, the presence of druse crystals, laticifers, angular collenchyma, and parenchyma cells in the primary veins, as well as collateral vascular bundles are common in Croton species as well as in Euphorbiaceae in general (e.g., Metcalfe & Chalk 1950;Mendonça et al. 2008;Barros & Soares 2013;Vitarelli et al. 2016).Although the species studied here preferentially occupy environments with certain water restrictions (such as Cerrado, and the Caatinga domain), they show an epidermis covered by only a thin cuticle, with the pericycle usually composed of parenchymal cells -different from reports by Morretes & Ferri (1959), Morretes (1969), and Bieras & Sajo (2009), who described a thickened cuticle on the epidermis and sclerified cells surrounding vascular tissues, for example, in Cerrado species from different plant families.The lack of a thickened cuticle on the leaf epidermis of the studied Croton species is possibly compensated with the presence of trichomes which sometimes form a dense indument, as already observed in other species of Croton by Sodré et al. (2019).Our observations show that living in the aforementioned dry environments does not necessarily lead to the development of obvious xeromorphic anatomical adaptations.
Remarks -Croton campestris was described and illustrated by Saint-Hilaire (1827) based on his own collection (A.F.C.P. Saint-Hilaire 2155bis) from the Congonhas da Serra region, in rocky fields of the south-central portion of Minas Gerais State.Baillon (1864) described the Croton sect.Velamea and allocated ten species to it, including C. campestris.That author complemented its description and recognized four varieties for the species: vars.subacutum, nigricans, atratum, and dupraei, differentiated, mainly by the type and color of the branches, leaves and capsules, the size and shape of the leaves, and length of the racemes.Müller Argoviensis (1866) recognized two other new varieties for C. campestris: C. campestris var.angustifolius and C. campestris var.laetifolius, and elevated Croton campestris var.atratum to the level of species, naming it Croton versicolor, which stresses its variable colors and long inflorescences with congested flowers.In the Flora Brasiliensis, Müller Argoviensis ( 1873) maintained the varieties of Croton campestris recognized until then, except for C. campestris var.subacutus, which he treated as a distinct species, Croton subacutus, by having branches with smooth and yellowwhitish trichomes, stamens with glabrous filaments, and small fruits.However, while analyzing the type collections of Croton campestris, its varieties, C. versicolor, and several specimens from the same geographic area, we observed great variations and overlaps in the characters identified as important for differentiating them, even in the same Revised delimitation of Croton campestris (Euphorbiaceae), including description of two new species, molecular phylogenetic, anatomical and micromorphological data population, or being continuous between populations, and we therefore treat them as synonyms.
Müller Argoviensis (1873) 1873) by few and overlapping characteristics related to the dimension of the petiole, indument density on the adaxial surfaces of the leaf blades, presence of trichomes on the petals and stamens, and capsule shape.With the exception of C. incertus and C. ferruginellus, the former described based on a collection from São Paulo State, and the latter from an unknown location in Brazil, all of the other species were described based on collections from the grasslands and rocky fields of Minas Gerais, as well as C. campestris and its varieties.Additionally, while analyzing the populations of Croton campestris in the field, as well as approximately 100 other exsiccatum (collected in rocky fields in the municipality of Ouro Preto to the municipality of Diamantina in Minas Gerais), we were able to observe wide variation, sometimes within the same population, in the characters used until then to differentiate them.The indumentum can be whitish, yellowish, or ferruginous, usually dense, tomentose or hirsute to the touch, but sometimes sparse, especially on the adaxial surface of the leaf blade; the leaves are usually elliptic, from wide to narrow, sometimes oblong, lanceolate, oval or oboval, and the thyrsus can be short to elongated (2.3-25 cm long) with varying numbers and densities of pistillate flowers/capsules (2-)5-15 per inflorescence.Additionally, we collected most of the representative morphotypes of those species, and found no reproductive or vegetative characteristics that could actually differentiate them from C. campestris, and therefore consider them all as conspecific.
Still considering the works of Baillon (1864) and Müller Argoviensis (1866,1873), two varieties of Croton agrarius described by the first author (C.agrarius var.cremostachyus and C. agrarius var.augustinianus) and later circumscribed in the concept of Croton grandivelus by Müller Argoviensis (1866), demonstrated morphologies most similar to C. campestris by the smooth or slightly striate branches and axes of the inflorescences, stipules reduced to 2-4 glands, leaf blades only 4-8(-10) times longer than the petioles, and tiny bracts (1.2-2.3 mm long).In contrast, C. grandivelus, the stipules are narrowly lanceolate and 2-8 mm long with numerous glands at the base, leaf blades 15-25 times larger than the petioles, and bracts 3.4-6.3mm long; in C. fulvus (species that currently includes C. agrarius in its concept) the branches and axes of the inflorescences are strongly striate, the leaf blades are usually 10-12 times longer than petioles, and the bracts 3.5-5 mm long.
Historically, C. campestris has been confused by different authors with several other species (see Tab.In order to more easily recognize those species, we provide here a key to their identification, and photographs (Fig. 14), but did not include C. hadrianii (C.sect.Geiseleria) and C. glandulosobracteatus (C.sect.Barhamia), the only taxa that does not belong to the sect.Adenophylli and are therefore easily differentiated by the absence of three prominent and ascending appendices at the apex of the columella.
Phenology -Flowering and fruiting from September to June.
The name Croton heliotropiifolius has been accepted by several authors to designate the most widely distributed species of section Adenophylli in the east and northeast of Brazil (e.g., Govaerts et al. 2000, Carneiro-Torres 2009, Caruzo et al. 2021).We analyzed specimens of C. heliotropiifolius from the type locality in the Andes and an extensive list of Brazilian materials, especially from the northeast region, and we agree that the specimens are very similar.Therefore, until population level phylogenetic studies are conducted on this species covering its entire range, it will not be possible to know whether they are cryptic species.
Proposed conservation status -Croton hatschbachii has an Extent of Occurrence (EOO) of 526.4 km 2 , is known from approximately ten localities in three municipalities in Minas Gerais (EOO) and only one known population occurs in a protected area (the Grão Mogol State Park).Although the species inhabit rocky areas at high altitude and of difficult access that are not always suitable for agriculture, stellate-rotate with 14-19 lateral rays in a single whorl, each ray 0.5-1 mm long, united up to 1 / 10 of their length, stipe 0.2-1 mm long, central ray vestigial, up to 0.05 mm long, or sometimes developed, 0.1-0.6(-1.3)mm long.
Phenology -Found flowering and fruiting throughout the year.
Remarks -Croton stellatorotatus has usually been identified in herbaria as C. campestris, and it was described and illustrated under that name by Carneiro-Torres ( 2009), as it is a non-viscous subshrub or shrub with branches and leaves with dense yellowish or whitish indument of stellate trichomes, leaves without nectary glands, petioles up to 1.7 cm long, shortly pedicellate pistillate flowers (pedicels up to 0.5 mm long), with 2-partite styles.Croton stellatorotatus differs, however, by having stellate-rotate trichomes with 13-20 lateral rays united for 1 / 10 of their length, and the central ray usually vestigial (ca.15 µm long) on its branches, leaves, axes of the thyrses, sepals of the pistillate flowers and ovary; leaf blades usually orbicular, ovate or largely elliptic, thyrses up to 6 cm long with 3-7 pistillate flowers, staminate flowers with small petals (1.6-2.2 mm long) and stamens (2.1-2.8 mm long), and styles 2.
Phenology -Flowering from August to December, and fruiting from September to February.
Remarks - Sodré et al. (2017), when considering the species of the Crotoneae tribe for the Chapada dos Veadeiros region, Goiás State, separately described and illustrated C. subvillosus and C. campestris, and differentiated them by trichome densities on the adaxial surfaces of their leaf blades, the presence of trichomes on the pedicels of the flowers, and style orientation.After studying larger numbers of collections of those two putative species, as well as field observations, we found that the differences pointed out by Sodré et al. (2017) were not consistent in terms of their differentiation, as they show variability among individuals in the same population.
As such, the specimens identified as C. campestris by those authors, by Sodré et al. (2014), and by Secco et al. (2018), are C. subvillosus, which is different from C. campestris in terms of its herbaceous or subshrub habit up to 50 cm in height, with little branching, with leaf trichomes with lateral rays 1-2 mm long, short petiolate leaves ((petioles (6-) 12-50 times smaller than the leaf blades)), leaf blades with an acute or short attenuate base, 7-15 pairs of secondary veins, axes of the thyrses and external surfaces of the floral calyxes of both sexes with a hirsute indument, as well as the pedicels of staminate flowers 2.3-4.7 mm long.Croton campestris, however, is characterized by a subshrub or shrub habit up to 2 meters tall, densely branched, lateral rays of leaf trichomes 0.35-1 mm long, petiole only 3.8-10 (-13) times smaller than the leaf blade, the leaf blades with obtuse bases, slightly cordate, or less commonly acute, 5-10 pairs of secondary veins, the axes of the thyrsus and external surfaces of the calyxes of the flowers of both sexes tomentose, and pedicels of the staminate flower up to 2 mm long.
& Riina (species sister to C. sect.Adenophylli according to Riina & Berry 2010) and one species of genus Brasiliocroton P.E.Berry & Cordeiro as an outgroup.Two datasets were built: one with 70 samples for the nuclear ribosomal ITS region and another with 65 samples for the plastid spacer trnL-trnF region.Sequences were retrieved from GenBank depending on DNA availability, except for five of them (C.campestris A.St.-Hil., C. echioides Baill., C. hatschbachii Sodré & M.J. Silva sp.nov., C. stellatorotatus Sodré & M.J. Silva sp.nov.and C. subvillosus Müll.Arg.), which were generated in this study.Information about all sequences used in this study with their vouchers and GenBank accession numbers are presented in Tab.S1.

Figure 1 .Figure 2 .
Figure 1.Tree obtained from the Bayesian analysis of ITS sequences.Values above and below the branches are Bayesian posterior probabilities and bootstrap percentages respectively.The scale bar indicates the mean number of nucleotide substitutions per site.In bold the species treated in this study.Asterisk indicates that the sample identification has been updated.

Figure 7 .
Figure 7. Croton campestris A. St.-Hil.: A. Subshrub with pendent branches in campo sujo vegetation in the municipality of Belo Horizonte, MG, Brazil, R.C. Sodré 3439.B. Subshrub with erect branches in campo sujo vegetation in the municipality of Santana do Riacho, MG, R.C. Sodré 3443.C. Flowering branch showing whitish indument, large and elliptic leaf blades, and elongate inflorescences in cerrado denso vegetation in the municipality of Diamantina, MG, R.C. Sodré et al. 3456.D. Flowering branch showing ferruginous indument and narrowly elliptic leaf blades in cerrado rupestre vegetation in the municipality of Datas, MG, R.C. Sodré 3449.E. Flowering branch showing ferruginous indument, short leaves, and inflorescences in cerrado rupestre vegetation in the municipality of Diamantina, MG, R.C. Sodré et al. 3454.F. Flowering branch showing whitish dense indument of the leaves and inflorescences in campo limpo vegetation in the municipality of Gouveia, MG, R.C. Sodré 3450.G. Flowering branch showing the yellowish indument in campo limpo vegetation in the municipality of Belo Horizonte, MG, R.C. Sodré 3438.H. Inflorescences with one pistillate flower near the base, and numerous staminate flowers along the axes, R.C. Sodré 3437.I. Young inflorescences with ca. 15 pistillate flowers towards the base, and staminate buds towards the distal end, R.C. Sodré 3437.J-K.Staminate flowers with greenish-yellow calyxes and whitish petals.L. Salmoncolored staminate flowers, note whitish indument on the calyx.M. Staminate flowers with ferruginous calyxes.N. Inflorescence base with only two sparsely distributed pistillate flowers, note brownish and hirsute indument of the ovary.O. Inflorescence base with ca. 10 whitish and densely distributed pistillate flowers.P. Fecundated pistillate flowers, note dark styles and whitish indument on the sepals and ovary.Q. Young pistillate flowers with brownish indument.R. Greenish-yellow capsules.S. Whitish capsules.T. Clear green capsules.U. Brownish capsules.V. Dehisced capsule, showing the columella with three apical lobes (J, N, R R.C. Sodré 3438; K, O, S R.C. Sodré 3450; L, P, T R.C. Sodré 3456; M, Q, U R.C. Sodré 3449) Photographs by R.C. Sodré.

Figure 10 .
Figure 10.Croton hatschbachii Sodré & M.J. Silva, sp.nov.: A. Subshrub habit in "cerrado rupestre" vegetation.B. Fertile branch ramifying in trifurcations.C. Fertile branch showing spiral alternate leaves, with lime-green adaxial surfaces.D. Short inflorescence showing sparse pistillate flowers, whitish and long-pedicellate staminate flower, and greenish-yellow buds, note the inconspicuous bracts.E. Inflorescence showing two capsules at base, one of them still young, and staminate flowers and buds along the axes.F. Inflorescence showing one developing fruit at base, and sparse staminate cymules along the axes, note the long pedicels of the staminate flowers.G. Staminate cymules, with flowers and buds, note whitish petals and stamens, and greenish calyx.H. Inflorescence base with two pistillate flowers, note floccose indument of the axes, greenish sepals, and brownish ovary and styles.I. Oblong cleargreen capsules.Photographs by R.C. Sodré from R.C. Sodré et al. 3464 and 3465.

Table 1 .
Authors who misinterpreted different materials such as C. campestris, with their respective vouchers, locations and valid species to which they correspond.

Rodolfo Carneiro Sodré, Alexandre Antônio Alonso and Marcos José da Silva triangular
, apex acute, globose glands at the base, sometimes with two short lobes, externally with hirsute indument of stellate-porrect trichomes with 6-7 lateral rays 0.7-1.5 mm long, porrect ray 1.2-1.5 mm long, sessile or subsessile, stipe up to 0.1 mm long, or covered by simple trichomes the same length as the rays, internally glabrous.