New species, additions and a key to the Brazilian species of the Geminata clade of Solanum L. (Solanaceae) in Brazil

Abstract Two additions and four new species are described from Brazil for the large Geminata clade (Solanum: Solanaceae) bringing the total diversity in the group to 149 species, with 44 of these occurring in Brazil. New species are described from Brazil: Solanum amorimii S.Knapp & Giacomin, sp. nov. from Bahia and adjacent Minas Gerais states, Solanum filirhachis Giacomin & Stehmann, sp. nov. from Espirito Santo, Solanum psilophyllum Stehmann & Giacomin, sp. nov. from Minas Gerais and Solanum verticillatum S.Knapp & Stehmann, sp. nov. from São Paulo, Rio de Janeiro and Minas Gerais. Modern character-rich descriptions and lectotypifications are provided for Solanum apiahyense Witasek and Solanum lacteum Vell. All are illustrated, mapped and assessed for conservation status. We also provide a brief analysis of the diversity and endemism of the Geminata clade in Brazil and a key to all 44 Brazilian species.


Introduction
Solanum L. is one of the largest of fl owering plant genera, and includes ca. 1400 species occurring worldwide on all continents except Antarctica. Th e genus was traditionally divided into the "spiny" and "non-spiny" solanums (e.g., Dunal 1852), based on the presence or absence of leaf and stem prickles. Molecular phylogenetic analysis showed that the "spiny" solanums form a monophyletic group (Bohs 2005;Weese and Bohs 2007;Särkinen et al. 2013), but the "non-spiny" solanums consist of a grade comprising several distinct monophyletic groups. Th e largest of these monophyletic groups are the potato clade (ca. 178 species of potatoes and their relatives), the "M" clade (of Särkinen et al. 2013; ca. 110 species of true nightshades and dulcamaroids; see Knapp 2013) and the Geminata clade, whose Brazilian members are treated here.
Th e Geminata clade as broadly defi ned contains 149 species, all but one of which occur in the New World tropics (Knapp 2002a(Knapp , 2008. Members of the group are shrubs and small trees mostly occurring in forest understory habitats; they are often inconspicuous, rare and rarely collected, with only a few widespread and weedy species. Th e group's name comes from the morphology of sympodial units in many of the component species where leaves appear to be twinned (geminate) at a node due to concaulescence of shoot generations (Danert 1958). Th e two leaves are often of markedly diff erent sizes and occasionally even shapes (see Knapp 2002a); plants are then markedly anisophyllous. Knapp (2002a) treated the group as section Geminata (G.Don) Walp. and divided the group into several informal species groups based on seed and sympodial morphology. Species later found to belong to the monophyletic group containing members of section Geminata (e.g., S. argentinum Bitter & Lillo and S. havanense Jacq. and its relatives; Weese and Bohs 2007) were added to the group and a list of component species with a key to all taxa was provided (Knapp 2008).
Solanum trachytrichium Bitter was included in Geminata by Knapp (2002aKnapp ( , 2008) but recent molecular work in the related Brevantherum clade (Giacomin 2015) revealed that it, plus the rare Brazilian species S. apiahyense Witasek ) are sister to the Geminata clade as treated by Knapp (2008), but with low support. Solanum apiahyense together with S. trachytrichium form a strongly supported lineage that is either sister to the Brevantherum or Geminata clade depending on the marker used. We here include these two taxa in the broad circumscription of the Geminata clade for practical reasons of identifi cation and morphological similarity, while recognising that future phylogenetic studies may show S. apiahyense and S. trachytrichium to be a distinct group (see discussion under S. apiahyense). Th ey share trichome types with the Geminata clade, and their possible relationships and similarities are discussed below.
An analysis of species richness and endemism patterns in the Neotropics using a group of species including many members of the Geminata clade (Knapp 2002b) showed peaks of both diversity and endemism in the Andes and south-eastern Brazil, as had been predicted by Gentry (1982) for understory plants in general. Subsequent analysis on a country level (Knapp 2008) showed species richness of the Geminata clade was highest in Colombia, Peru and Brazil with 41 (9 endemic from Colombia, 22%; 11 endemic from Peru, 27%) and 35 (incorrectly recorded as 34; 17 endemic, 50%) species respectively. Concentrated work in Brazil focused on the Lista de Especies de Flora do Brasil ) has clarifi ed the status of several names of dubious application and brought to light new species of the Geminata clade that are described here. We here record 44 species (43 native) of the group for Brazil, instead of 35 recorded by Knapp (2008). Of these nine additional records for the country, two are range extensions (S. arboreum Dunal, S. diphyllum L.) and are documented with character-rich descriptions in the literature cited above, while two have not been described over their entire range (S. apiahyense and S. lacteum) or have been of uncertain application (S. lacteum) and we provide descriptions to a modern standard here. Five new species have been discovered since Knapp (2008). We describe four of these new taxa (one is in review elsewhere, see below) and clarify diversity and distribution of the entire clade for Brazil.

Materials and methods
Descriptions are based on fi eld observations and examination of herbarium specimens from 27 collections in Brazil and abroad (B, BM, BHCB, BR, CEPEC, CORD, ESA, F, FUEL, FURB, G, HUEFS, IAC, JPB, K, LE, MBM, MBML, NY, PMSP, RB, SP, SPSF, UEC, UT, VIC, WU). Herbarium acronyms are from Index Herbariorum (http://sciweb.nybg.org/science2/IndexHerbariorum.asp) and all specimens are cited in the text. Full data are provided in the supplemental fi le and on the Solanaceae Source website (http://www.solanaceaesource.org). Extent of Occurrence (EOO) and Area of Occupancy (AOO) were calculated using GeoCat (http://geocat.kew.org) using the standard 2 km 2 cell width for AOO calculation. Conservation status of each species was assessed using the IUCN (2014) criteria based on the GeoCat analyses (Bachman et al. 2012) combined with fi eld knowledge.

Results and discussion
Th e broadly defi ned Geminata clade has 43 species native to Brazil (Table 1); only S. diphyllum (see Knapp 2002a) is known only from cultivation and may be naturalising. Th e state distribution of each species is given in Table 1, along with endemic status and extra-Brazilian distribution of non-endemic species. Endemism of native Brazilian Geminata species now stands at 65% (28/43 native species, excluding S. diphyllum). Th e south-eastern region (following Brazilian political divisions) is the most species-rich area of the country with 24 species, followed by the southern region (19 species); the northern regions have fewer species, refl ecting the circum-Amazonian species richness of Solanum in general (Table 2). Th e distribution in federal units (states) by species is presented in the second column of Table 1 and by state in Table 3. Th e states of Paraná (19 species), Minas Gerais (17 species) and Santa Catarina (16 species) are the most species-rich, followed by São Paulo (15 species) and Rio de Janeiro (12 species). Six species are endemic to a single state; S. cordioides and S. santosii in Bahia, S. fi lirhachis in Espirito Santo, S. psilophyllum in Minas Gerais, S. gertii in Paraná, and S. sp. 1 (a new species based on collections including Giacomin et al. 1789 [BHCB, UFP] being described by M.F. Agra and currently in review) in Pernambuco.
Only seven of the native species occur exclusively outside the Mata Atlântica biome (Atlantic rainforest; as defi ned by IBGE 2012); all of these are Amazonian (see Table 1). All of the endemic species (28) occur in Mata Atlântica, with 24 of those occurring only in that biome; only S. caavurana (Caatinga + Cerrado), S. intermedium (Cerrado) and S. psilophyllum (Cerrado; in the forested capões associated with Campos Rupestres) occur in other vegetation types. Solanum caavurana is widespread in secondary habitats and S. intermedium and S. psilophyllum occur in regions where the Cerrado and Mata Atlântica meet (e.g., Serra do Cipó in Minas Gerais). Few Geminata species are widespread in Brazil; only S. caavurana, S. campaniforme, S. leucocarpon, S. pseudocapsicum, S. pseudoquina and S. stipulatum occur in more than four states.
Because of their biology and occurrence in small populations of scattered individuals, most of the species described here (with the exception of S. verticillatum) can be classifi ed as rare and of some conservation concern. Rabinowitz (1981) suggested that species become rare (and by extension subject to extinction risk) by a variety of pathways and if this were so, the ecological and evolutionary consequences of rarity would be diverse. She analysed plant rarity using a scheme that took into account range size, habitat specifi city and local abundance (population size); in her classifi cation rare species ranged from 'common' to 'endemics'. Th e ecological consequences of rarity are likely to diff er in rare taxa of the diff erent categories.
Although the south-eastern part of Brazil is the most intensively collected part of the country (Sousa-Baena et al. 2013) all of the new species and additions to the Geminata for the Brazilian fl ora come from this region. As collecting is intensifi ed in other regions (such as the western edges of the Amazon basin) we expect more of these forest understory solanums for Brazil.  ) -see also Table 3.
Ecology. Solanum amorimii is found in the understory of wet Atlantic forests (Floresta Ombrófi la Densa, Mata Atlântica; IBGE 2012) from 50-1000 m, most commonly found at around 500-900 m elevation in very preserved sites.
Phenology. Flowering specimens have been collected from July to October but appears to peak in August; fruiting specimens have been collected from September to April.
Etymology. Th e species epithet honours André M. Amorim, curator of the herbarium at CEPEC in Ilhéus, Bahia, and collector of the type specimen, whose knowledge of the fl ora of Bahia has helped many botanists in the region and beyond.
Preliminary conservation status (IUCN 2014). Near-threatened (NT) B1, 2a, b (ii, iii); EOO 20,663 km 2 (NT); AOO 40 km 2 (EN). Although the large extent of occurrence (> 20,000 km 2 ) places S. amorimii out of the vulnerable category, the small number of locations (5-10) and the fragmentation of its forest habitat mean it is of some conservation concern. Populations occur within several private protected areas (in Minas Gerais the only population is within a private reserve) so the species is afforded some protection. On the other hand, the known collections suggest the species is restricted to pristine sites, which are becoming increasingly rare. As with all Geminata species, it is possible that more populations remain to be collected; these plants are inconspicuous in the deep forest understory and usually occur in small, sparsely distributed populations.
Notes. Solanum amorimii is morphologically very similar to the sympatric S. restingae, but can be distinguished by its much smaller fl owers with long-triangular calyx lobes and by its unwinged stem. Both species grow in the understory of mostly undisturbed forests and can be small shrubs or treelets. Solanum restingae has markedly cucullate corolla lobes, and the calyx lobes are so small as to be almost non-existent, especially in fruit. Bud shape also diff ers between the two species, with those of S. amorimii being globose to somewhat ellipsoid and those of S. restingae more elongate  with a distinct "nipple" from the cucullate corolla tips. In fruit the two species can be diffi cult to distinguish, but the winged stems of S. restingae and the presence of calyx lobes in S. amorimii should enable identifi cation.
Leaves of S. amorimii are usually somewhat auriculate at the base, with the base not surrounding the stem but enlarged to a very short petiole. Plants grow in forest understory, sometimes in open places such as treefall gaps. From overall morphology this species would belong to the S. arboreum species group of Knapp (2002a), but its relationships have not been tested using molecular sequences.
Specimens examined. BRAZIL. BAHIA: Mun. Arataca, RPPN Caminho das Pedras, Serra do Peito-de-Moça, entrada a 9.5 km no Assent. Santo Antonio, mais 8.9 km ate a sede da RPPN, trilha de acesso ao topo da serra, após a Mormaço, 15°10'27"S, Description. Small erect shrubs, to 50 cm tall, often rhizomatous with a horizontal woody branch bearing several adventitious roots; young stems moderate to densely pubescent, with 4-8-celled hyaline trichomes to 2 mm long; new growth drying dark, densely pubescent; bark of older stems pale gray, glabrescent, not exfoliating. Sympodial units 3-plurifoliate, normally not geminate, if geminate, with leaves diff ering only in size. Leaves simple, 3.4-11 × 0.8-4 cm, elliptic to narrowly elliptic, membranous, slightly discolorous, shiny green adaxially when fresh, drying pale green beneath, dark above, not shiny, both surfaces moderate to densely pubescent with hyaline simple uniseriate trichomes 1-2 mm long with up to 5 cells, sometimes with a multicellular base (but see comments); primary veins 5-7 pairs, the midrib and primary veins darker abaxially, raised; base attenuate to acute, slightly decurrent onto the petiole, mostly symmetric; margins entire, not revolute, ciliate with antrorse hyaline trichomes; apex attenuate to acuminate; petioles 2.5-15 mm long, densely pubescent, with trichomes like those of the stems and leaves. Infl orescences 1.7 to 3.3 cm long, mostly lateral or less often strictly opposite the leaves, unbranched, with 3-5 fl owers, moderate to densely pubescent, with hyaline trichomes like those of the stems and leaves; peduncle 4-15 mm long; pedicels 5 to 11 mm long, articulated at base; pedicel scars closely spaced ca. 1 mm apart. Buds globose to slightly elongate, the corolla mostly included in the calyx tube, exserted only just before anthesis. Flowers all perfect, 5-merous. Calyx tube up to 1 mm long, conical, getting refl exed, the lobes up to 0.9 mm long in fl ower, to 1.7 mm long in fruit, approximately 1.6 mm wide, acuminate and discretely keeled, adaxially, glabrous or papillose, covered with tiny 1-2-celled glandular trichomes, abaxially densely pubescent, with trichomes as those of the stem, or sometimes even longer, with 2.5 mm, and normally 5-6 cells. Corolla 1.5-1.7 cm in diameter, white, stellate, membranous, lobed from 2/3 to 3/4 of the way to the base, the lobes 7.5-9 mm long, 3-3.5 mm wide, refl exed at anthesis, deltate to lanceolate, glabrescent adaxially, abaxially sparsely pubescent, with 3-4-celled delicate simple trichomes of ca. 0.5 mm along  the midvein, with tufts of few celled tiny trichomes less than 0.1 mm long on the tips and margins. Stamens 3.2-3.6 mm long; fi lament tube ca. 0.5 mm long, the free portion of the fi laments up to 0.6 mm long equal in length or slightly unequal, and when so, one fi lament slightly longer (barely visible in dried material), glabrous; anthers 2.6-2.8 mm long, 1.6-1.8 mm wide, ellipsoid, slightly connivent, yellow, slightly sagittate at the base, the pores directed introrsely, opening into longitudinal slits at maturity. Ovary glabrous; style 4.2-5 mm long, white, straight, glabrous; stigma capitate, light green. Fruit a globose berry 0.7-1.4 cm in diameter (immature?), dull green, drying dark, the pericarp glabrous and not markedly shiny; fruiting pedicels 1.2-2 cm long, ca. 0.7 mm in diam. at the base, to 1.1 mm at the apex, with a slight constriction at the receptacle; calyx lobes in fruit somewhat enlarged. Seeds approximately 70 per fruit, known only from very young fruits, possibly fl attened and with a marginal wing when fully developed.
Distribution. In the Serra do Mar mountain range in the Brazilian states of Paraná, Santa Catarina and São Paulo ( Figure 5).
Ecology. Solanum apiahyense is a rare and inconspicuous shrub of the understory and edges of well preserved and secondary fragments of the montane Brazilian Atlantic rainforest (Floresta Ombrófi la Densa of IBGE 2012; Mata Atlântica), from 600 to 900 m. Although most collections are from well preserved sites, S. apiahyense is not exclusively associated with shaded environments. Th e species is also found along unpaved roadsides close to the type locality.
Phenology. Fertile specimens are known from September to February. Mature fruits were observed only in October.
Etymology. Th e epithet refers to the type locality, the city of Apiaí in southern São Paulo state.
Preliminary conservation status (IUCN 2014). Endangered (EN) B1; B2 ab (ii, iii, iv). EOO 3,208 km 2 (EN); AOO 16 km 2 (EN). Although the species occurs in a wide latitudinal range, it is locally rare, and is known from only six localities. None of the known populations are from within protected areas.
Notes. Solanum apiahyense, described more than a century ago (Witasek 1910), has not been assigned to any infraspecifi c group of Solanum so far. Recent phylogenetic analysis using molecular data (Giacomin 2015) has shown it to be closely related to S. trachytrichium, which was previously assigned to the Geminata clade (Knapp 2002a(Knapp , 2008 and to its own subsection when originally described (subsect. Silicosolanum Bitter; Bitter 1919). Bitter (1919) based this on the unusual trichome morphology of hooked cells arising from a fl attened multicellular base that give the leaves a feeling of sandpaper in herbarium specimens. Although molecular data support a close relationship between S. apiahyense and S. trachytrichium, the affi nities of this clade are not clear-cut. Data from combined markers place it as sister to all other Geminata clade species, but with low support. In analyses of individual markers, it emerges as sister to either the Brevantherum or Geminata clades depending upon the marker used (Giacomin 2015).
Morphologically both taxa are easy to distinguish from most other Geminata species, and have the following assemblage of characters: both are small shrubs with leaves mostly not geminate, they have leaf trichomes with an expanded multicellular base and relatively large fl owers (>1.5 cm in diameter). Among them, Solanum apiahyense and S. trachytrichium are easy to distinguish: S. trachytrichium has a unique scabrous indumentum on the leaf surfaces and stems, composed of short unicellular hooked trichomes on a mound-like multicellular base, while in S. apiahyense the surface is not rough to the touch, and although some trichomes with multicellular bases can be seen on leaves, these are translucent, very long (ca. 2 mm) and mostly 5-7-celled. Th ese long trichomes of S. apiahyense are easily seen on the new growth, while S. trachytrichium trichomes are not visible to the naked eye. In addition, the fl owers of S. apiahyense are slightly smaller, 1.5-1.7 cm in diameter versus 1.6-2.2 cm in S. trachytrichium.
In the past, the epithet S. apiahyense has been applied to more than one species of the S. inornatum group (part of the Brevantherum clade; Giacomin and Stehmann 2014) by various Solanum taxonomists, although they are now known to not be closely related. Although members of the S. inornatum group (e.g., S. inornatum Witasek, S. bradei Giacomin & Stehmann and relatives) and S. apiahyense are similar in habit and in having pubescence of long, translucent trichomes, they can be readily distinguished by close examination of the trichomes; those of S. apiahyense are multicellular with 5-7(8) cells while those of members of the S. inornatum group are mostly 3-celled (probably representing modifi ed stellate hairs, Giacomin and Stehmann 2014). Fruiting specimens of S. apiahyense have peduncles longer than 1 cm and the pedicels are strongly apically expanded and constricted just beneath the calyx lobes (see Figure 1C), while in the species of the S. inornatum group species, the peduncles do not exceed 1 cm and the pedicels are never apically expanded with a distal constriction. Examination of trichomes with a 10× hand lens will allow easy identifi cation of both fl owering and fruiting material.
Th e type material found at WU (Puiggari 3711) consists of a single sheet, and does not match the photograph of a dried specimen in the original publication (Witasek 1910: tab. 30, fi g. 2). It should therefore be treated as an isotype (Mentz and Oliveira 2004). As no further material could be found in other possible herbaria where J.I. Puiggari deposited his collections, the specimen at WU is here designated as a lectotype.  Diagnosis. Diff ers from the sympatric S. campaniforme Roem. & Schultes in its deep forest habitat, leaves with ruffl ed margins, fl owers less than 1 cm in diameter, pedicels with a constriction at the distal end that are swollen in fruit, and few seeds.
Type Description. Erect shrubs to small trees, up to 3 m tall, normally branching close to the apex, the upper stems ascendant; young stems terete, glabrous; new growth brownish, glabrous. Bark of older stems turning pale greyish brown, glabrous, not exfoliating. Sympodial units difoliate, mostly geminate, with leaves not diff ering in shape or size. Leaves simple, 4.6-15.9 cm long, 1.3-4.9 cm wide, narrowly elliptic, membranous to chartaceous, slightly discolorous when dry, the adaxial surface glabrous, dark green and somewhat shiny in live plants, the abaxial surface sparsely pubescent with simple uniseriate 7-12-celled trichomes to 1 mm long in tufts in the primary vein axils, occasionally extending to the midrib; primary veins 5-9 pairs, yellowish green, discretely raised above, raised beneath; base attenuate to acute, slightly decurrent onto the petiole, sometimes asymmetric; margins entire, slightly undulate (ruffl ed) and revolute, apex long-attenuate to acuminate; petioles 1-9 mm long, glabrous. Infl orescences 3.5 to 26 cm long, opposite the leaves or internodal, unbranched, slender and very delicate, with 18-60 fl owers, but bearing normally with 4-10 fl owers at a time, glabrous; peduncle 1.8-3.8 cm long; pedicels 7-18 mm long, ca. 0.4 mm in diam. at the base, ca. 0.9 mm in diameter at the apex, with a constriction at the receptacle, articulated at base, unevenly spaced 1.7 to 10 mm apart. Buds globose, the corolla completely exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube to 1 mm long, conical, the lobes ca. 0.2 mm long, ca. 1.5 mm wide, acuminate and somwewhat keeled, papillose adaxially, glabrous abaxially. Corolla 6-8 mm in diameter, normally whitish purple adaxially, light purple abaxially, stellate, membranous, lobed more than ¾ the way to the base, the lobes 4-5 mm long, 1-1.7 mm wide, spreading at anthesis and becoming refl exed in older fl owers, deltate to lanceolate, glabrous on both surfaces, minutely papillose at tips and margins. Stamens 2.5-3 mm long; fi lament tube ca. 0.3 mm long, the free portion of the fi laments up to 0.2 mm long, equal in length or slightly unequal, and when so, two fi laments slightly longer (barely visible in dried material), glabrous; anthers 2-2.5 mm long, 1.2-1.5 mm wide, ellipsoid, slightly connivent, yellow, poricidal at the tips the pores directed introrsely, elongating to longitudinal slits with age. Ovary glabrous; style 4-6 mm long, white, straight, glabrous, the stigma light grayish green, capitate. Fruit a globose berry 1-1.5 cm in diameter, dull green at maturity, with irregular black spots ( Figure  Figure 6. Holotype specimen (sheet two) of S. fi lirhachis (Giacomin et al. 1854, BHCB019057). Reproduced with permission of the Universidade Federal de Minas de Gerais. 1F) drying grayish brown, the pericarp glabrous, not shiny; fruiting pedicels 2.0-2.4 cm long, clearly obconical, ca. 0.5 mm in diam. at the base, widening markedly towards the apex to ca. 2.5 mm in diam.; calyx lobes in fruit ca. 1.5 mm long, commonly broken off in dried fruiting material. Seeds 20-25 per berry, 2.5-4.5 mm long, 2-3.3 mm wide, ovoid-reniform to somewhat fl attened towards the margins, light to dark brown, the surface irregularly pitted, the testal cells undulate.
Distribution. Restricted to the state of Espírito Santo (Figure 7), in south-eastern Brazil. Collections are known from the central and northern parts of the state, from both sides of the Rio Doce.
Ecology. Rare in the understory of well-preserved fragments of the sub-montane and montane Brazilian Atlantic coastal rainforest (Floresta Ombrófi la Densa; IBGE 2012), normally in formations where granitic outcrops are present or close by, in elevations ranging from 200 to 750 m.
Phenology. Fertile specimens of Solanum fi lirhachis are known mostly from the rainy season (from November to March), but the type collection from June indicates that the species might be fertile for a longer period. Mature fruits were observed in specimens from November and June.
Etymology. Th e epithet refers to the long and slender infl orescence rachis, which is not observed in any of the Brazilian sympatric species, although a common feature in some species of the S. confi ne group from Colombia, Ecuador and Venezuela (Knapp 2002a).
Preliminary conservation status (IUCN 2013). Endangered (EN) B1, B2 ab (ii, iii, iv); EOO 1,136 km 2 (EN); AOO 20 km 2 (EN). Solanum fi lirhachis is currently known from only fi ve localities, and all collections are from within private properties, where agriculture (both large and small scale) is known to occur. Despite the fact that it inhabits higher elevations that are usually harder to access and not always suitable for agriculture, we strongly recommend that further eff orts to map new populations of the species should be undertaken, mainly within protected areas with similar forest types. Although the type locality of Santa Teresa in central Espirito Santo has several well preserved fragments of forest, the landscape has been rapidly transformed in the last few decades to Eucalyptus and coff ee plantations, and summer vacation homes (cottages).
Notes. Solanum fi lirhachis is remarkably similar to a suite of species of the Geminata clade with ruffl ed leaf margins (see Figure 1E) and long fi liform infl orescences (S. leptorhachis Bitter and S. nematorhachis S.Knapp from the W Andean slopes in Colombia and Ecuador and S. tenuifl agellatum S.Knapp of Venezuela). Knapp (2002aKnapp ( , 2008 treated these as members of her S. confi ne species group, all of whose members have a thin infl orescence rhachis, small fl owers and leaves with ruffl ed (undulate) margins, although this latter character is impossible to see in herbarium specimens. Solanum fi lirhachis diff ers from those species in its distribution and in the tufts of trichomes in the abaxial leaf vein axils (domatia); other members of this morphologically similar set of species are glabrous or have fi ne, golden pubescence. Th e only Brazilian species Knapp (2002) placed in this group was S. stipulatum which can be easily distinguished from S. fi lirhachis by its shorter infl orescences, fl owers with refl exed corolla lobes and winged stems with anisophyllous difoliate, geminate sympodial units. Solanum stipulatum is usually a shrub of watercourses, and often grows amongst rocks and is submerged in fl oods, while S. fi lirhachis is a slender treelet of forest understory. Th e relationships of the S. confi ne group have not yet been tested using molecular markers.
Another Brazilian species with which S. fi lirhachis could be confused is S. campaniforme that has similar (but somewhat stouter) elongate infl orescences and tufts of uniseriate trichomes in the abaxial leaf vein axils. Solanum fi lirhachis has leaves with ruffl ed margins tht normally dry pale green and smaller fl owers (0.6-0.8 cm in diameter) that (at least in the type specimen) are tinged purple; S. campaniforme has leaves with entire, non-ruffl ed margins that normally dry black or brownish black and larger fl owers (1.2-1.8 cm in diameter) with strongly cucullate corolla lobes.
We have designated a two sheet holotype for S. fi lirhachis in order to represent both fl ower and fruit in the type sheets.  Description. Shrub or small treelet 1-3 m (occasionally as small as 25-30 cm or as tall as 5 m); young stems terete, glabrous; new growth glabrous or minutely papillate; bark of older stems pale brown, with prominent paler lenticels. Sympodial units difoliate, geminate or more usually not geminate; leaves of a pair usually diff ering in size but not in shape. Leaves simple, 9.5-25 cm long, 3.5-9 cm wide, narrowly obovate, widest in the distal half, membranous, glabrous on both surfaces, the abaxial surface paler in dry specimens; primary veins 6-10 pairs, drying dark abaxially; base attenuate; margins entire; apex bluntly acute to attenuate; petiole 1-3 cm long, glabrous; minor leaves, if present, diff ering only in size from the majors. Infl orescences 0.1-0.5 cm long, terminal, more or less leaf-opposed or internodal and appearing pseudoaxillary, unbranched or occasionally furcate, with 5-10 fl owers, glabrous; peduncle 0.1-0.5 cm long, the fl owers in an apical clump; pedicels 0.9-1.1 cm long, < 0.5 mm in diameter at the base and apex, fi liform, spreading at anthesis, glabrous, articulated at the base, with a constriction at the apex just below the calyx lobes, this becoming more pronounced in fruit; pedicel scars congested and overlapping at the tip of the very short infl orescence. Buds ovoid, the corolla strongly exserted form the calyx tube before anthesis. Flowers 5-merous, perfect. Calyx tube ca. 0.5 mm long, conical, the lobes 0.5-0.75 mm long, ca. 0.5 mm wide, deltate, with scarious margins and rounded tips, glabrous. Corolla 0.9-1 cm in diameter, white, stellate, lobed ca. 2/3 of the way to the base, the lobes 3-4.5 mm long, 1.5-3 mm wide, spreading or somewhat refl exed at anthesis, the tips and margins minutely papillose. Stamens 2.5-3 mm long; fi lament tube ca. 0.5 mm long, the free portion of the fi laments < 0.5 mm long, glabrous; anthers 1.5-2 mm long, ca. 1 mm wide, ellipsoid to almost globose, yellow, poricidal at the tips, the pores lengthening to longitudinal slits with age. Ovary glabrous; style ca. 4 mm long, glabrous; stigma minutely capitate, the surface papillose. Fruit a globose to somewhat ellipsoidal berry, 0.5-1 cm in diameter, greenish white, occasionally pointed at the apex, the pericarp thin, shiny, brittle when dry; calyx lobes in fruit not markedly enlarging; fruiting pedicels 1-1.3 cm long, 0.5-1 mm in diameter at the base, enlarging gradually to 1.5-2 mm in diameter at the apex, with a slight constriction just below the calyx lobes, not markedly woody, pendant; calyx lobes in fruit not markendly enlarged. Seeds 10-20 per berry, 3-4 mm long, 2-3 mm wide, somewhat fl attened-reniform (perhaps immature?), dark to blackish brown, the surfaces minutely pitted, the margins paler and thickened; testal cells pentagonal in outline.
Distribution. South-eastern Brazil in the states of Espirito Santo, Minas Gerais and Rio de Janeiro (Figure 11).
Ecology. Solanum lacteum grows in wet Atlantic forests (Mata Atlântica, Floresta Ombrófi la Densa) in forest understory of well preserved sites, from 600 to 1500 m elevation.
Phenology. No apparent pattern in fl owering or fruiting; specimens are often collected with only infl orescences, each plant is very few-fl owered.
Etymology. Th e species epithet was coined by Vellozo (1829) to refer to the whitish colour of the plant -"Color albescens totius plantae nomen triviale dedit" (the white color of the entire plant gives it its trivial name [epithet]). We have not observed entire plants that are white in colour, but suspect Vellozo (1829) was referring to the congested infl orescence that is completely white.

Preliminary conservation status (IUCN 2014).
Near Th reatened (NT) B1, 2 a, b(ii, iii); EOO 32,466 km 2 (NT); AOO 28 km 2 (EN). In spite of its large extent of occurrence, S. lacteum is only known from six locations and we consider it to be at risk due to the fragmentation and loss of its primary forest habitat. Populations in all three states of occurrence, however, are from within protected areas. It is possible that it is more common than it appears, considering that the fl owers are so small and inconspicuous that it is easily overlooked.
Notes. Solanum lacteum is characterized by its tiny infl orescences with tightly packed fl owers and the difoliate sympodia that are usually not conspicuously geminate. Th e leaves are narrowly obovate and widest in the distal third. Th ey dry a characteristic blackish brown above and paler brown beneath. Th e infl orescences often occur internodally and are completely white, including the peduncle and pedicels. Th e colour of the leaves on herbarium specimens is similar to that of S. caavurana and S. campaniforme, but those species always have leaf pubescence on the lower leaf surfaces and more elongate infl orescences. Th e highly congested infl orescences of S. lacteum are distinctive and the species is not easily confused with any other growing sympatrically. It is somewhat similar to S. psilophyllum, which is similarly glabrous; diff erences between these two species are noted in the discussion of S. psilophyllum.
Solanum lacteum grows in the understory of undisturbed forest and can vary from being a tiny subshrub (see Figure 8B) to a small treelet ca. 5 m tall. Th is variation in height is common in members of the Geminata clade and may have to do with plant age and maturity.
Vellozo's (1831) illustration ( Figure 9) is not particularly clear, but the congested infl orescences and swollen fruiting pedicels with a slight distal constriction are clearly depicted. In addition, S. lacteum usually has prominent lenticels on the stems; these are also depicted in Vellozo's plate. We have selected an epitype from Rio de Janeiro State to support this suboptimal plate with material that is fertile and shows the key characters (Agra et al. 7298).
We have recognised S. cormanthum here as a synonym of S. lacteum; after detailed study we consider the plate of S. cormanthum (t. 113) to represent fl owering material of the same taxon as that shown in fruit in Vellozo's plate of S. lacteum (t. 93). Solanum cormanthum was used by both Sendtner (1846) and more recently in the Lista de Especies de Flora do Brasil ) to refer to a diff erent taxon we here recognise as a narrow endemic from Minas Gerais (see S. psilophyllum below). Both these authors expressed reservations about the correct application of this name. As is the case with the plate of S. lacteum, the depiction of the plant is not particularly clear, but the small fl owers, small anthers and infl orescences that appear axillary (although they are not) are characteristic of S. lacteum. Th e locality cited for S. cormanthum ("silvis maritimis Regii Praedii S. Crucis"; Vellozo 1829: 86) is well within the geographic range and habitat of S. lacteum, although today it is part of the city of greater Rio de Janeiro.
Sendtner's (1846) plate of S. glomulifl orum (f. 11-15) clearly shows the scariousmargined calyx with rounded lobes and very plump anthers characteristic of S. lacteum. In his protologue Sendtner (1846) cited two collections of S. glomulifl orum; a fl ower- ing specimen of Schott from "Serra d'Estrella" (Serra de Estrela, in Rio de Janeiro State) and a fruiting specimen of Sellow's from an unspecifi ed locality in Brazil (F neg. 2823; presumably from Berlin]. We select here the Schott specimen at F (accession number 874710; barcode F0073278F) as the lectotype of S. glomulifl orum, as it bears a label with the locality and collector in J.F. MacBride's handwriting and presumably comes from Berlin where the original is now destroyed. Th e collection number 5412 noted on this sheet was not mentioned by Sendtner (1846) Diagnosis. Like Solanum evonymoides Sendtn. but diff ering in smaller fl owers, infl orescences that are unbranched or branch only once near the base, pedicels with a constriction at the apex just below the calyx lobes and ovoid-reniform seeds. Description. Treelet to 4 m, rhizomatous with underground stems; young stems terete, glabrous; new growth completely glabrous, occasionally minutely papillate; bark of older stems greenish brown, slightly winged from the leaf bases. Sympodial units difoliate, geminate; leaves of a pair diff ering in size but not usually in shape. Leaves simple, the major leaves 10-15(-25) cm long, 4-13 cm wide, elliptic to narrowly elliptic, occasionally wider in the distal third and narrowly obovate, membranous, glabrous on both surfaces, the abaxial surface often drying paler than the adaxial surface; primary veins 8-11 pairs, drying somewhat lighter than the lamina; base attenuate, somewhat oblique; margins entire; apex acute, the tip somewhat blunt; petiole 1.5-2 cm long, glabrous; minor leaves 6-8 cm long, 2-3 cm wide, diff ering from the majors only in size and sometimes not present in dried specimens. Infl orescences 0.2-2 cm long, opposite the leaves or appearing to arise from the leaf axils, unbranched, but apparently sometimes with 2 infl orescences from one axil and appearing branched (Giacomin et al. 186), with 5-8 fl owers, glabrous; peduncle 0.1-2 cm; pedicles 1.2-1.5 cm long, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the swollen apex with a marked constriction just below the calyx lobes, slender and expanding distally, spreading or pendant at anthesis, glabrous, articulated at the base; pedicel scars 0.5 -1 mm apart, more congested in the distal part of the infl orescence. Buds obovoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers 5-merous, perfect. Calyx with the tube 0.5-1 mm long, broadly conical, the lobes 1-1.5 mm long, deltate to triangular, refl exed at anthesis, glabrous. Corolla 1.2-1.4 cm in diameter, white, stellate, lobed 1/2 to 2/3 of the way to the base, the lobes ca. 5 mm long, 2.5 mm wide, spread at anthesis, glabrous with the tips minutely papillate. Stamens 3.5-4 mm long; fi lament tube ca. 0.5 mm long, the free portion of the fi laments ca. 0.5 mm long, glabrous; anthers 2.5-3 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 5-6 mm long, glabrous; stigma not expanded, blunt, the surface minutely papillate. Fruit a globose berry, 1-1.3 cm in diameter, green, the pericarp not markedly shiny, thick; fruiting pedicels 1.5-1.7 cm long, ca. 1 mm in diameter at the base, 2.5-3 mm and expanded at the apex, woody and pendant; calyx lobes in fruit not markedly expanding, but distinctly diff erentiated from the enlarged pedicel apex. Seeds not known.
Distribution. In the south-eastern part of the state of Minas Gerais, in islands of forest (capões) associated with iron or quartzite formations in the Iron Quadrangle and Serra do Cipó regions, in the southern limit of Espinhaço mountain range (Figure 13).
Ecology. Solanum psilophyllum grows in the forest understory on thin soils associated with iron-rich or quartzite formations, at elevations from 800-900 m.
Phenology. Flowering specimens have been collected throughout the year; fruits have only been seen on the type specimen, collected in July. It is probable that this species fl owers and fruits sporadically throughout the year.  Etymology. Named for its completely glabrous leaves (from the Greek psilos smooth or bare, phyllos leaf).
Preliminary conservation status (IUCN 2014). Critically Endangered (CR) B1, 2 a, b(ii, iii, iv); EOO 26 km 2 (CR); AOO 16 km 2 (EN). Solanum psilophyllum is known from only two localities and its habitat is under severe pressure from mining and frequent forest fi res (see Notes). Th e population from which the type specimen was collected, close to a private railroad, has already been destroyed. Although the area of occupancy would suggest a status of Endangered we consider the extreme threats to these populations coupled with the habitat specifi ty of members of the Geminata clade (see above) warrant a status of Critically Endangered.One of the known collections might be from a protected area (PARNA Serra do Cipó), although not stated on the specimen label (Campos & Belisário CFSC-13505) but appears to be from a roadside, subject to occasional fi re.
Notes. Solanum psilophyllum is the species previously called Solanum cormanthum Vell. in Lista de Especies de Flora do Brasil ). Th at name, however, has been of uncertain application since Sendtner (1846) listed a collection from Minas Gerais ("Caxoeira do Campo") as belonging to S. cormanthum, but with reservations.
Th ree sheets of labelled as "Solanum cormanthum Vell." in Martius's hand in Brussels belong to this species as do presumed duplicates of this collection in F (F-680206) and G (G00016950) cited by Knapp (2008) as belonging to S. evonymoides Sendtn., a species now considered to only occur from coastal Bahia to northeastern Minas Gerais (see discussion of S. verticillatum below). Sendtner (1846) cites a collection in Martius's herbarium from "Caxoeira do Campo, prov. Minarum, Martio fl oret: Martius"; this was probably collected by Claussen. One of the three of the sheets in BR (BR00000825373) is from Martius's herbarium and is labelled "Mart. 1839." Another sheet is defi nitely attributed to Claussen and collected in 1835, while the third is attributed ("comm. Schüch fi l. 1850") to Guilherme Schüch, the Baron of Capanema (Minas Gerais, currently the active iron mine of Capanema), who sent plants to Martius.
Th e Vellozo illustration of S. cormanthum (tab. 113, Vellozo 1831) has distinctly axillary infl orescences and is said to come from what is now the city of Rio de Janeiro ("Praedii S. Crucis"), an area of very diff erent vegetation and soils than the iron or quartzite rich formations of Minas Gerais. We recognise S. cormanthum here as a synonym of S. lacteum, both on morphological and distributional grounds. Members of the Geminata clade are very similar morphologically and Vellozo's plates are often distinctly suboptimal for secure identifi cation. In view of the restricted distribution and habitat of these plants (see below) we prefer to describe this as new here rather than use S. cormanthum for these distinct and endangered populations.
Solanum psilophyllum has a very narrow distribution restricted to the Iron quadrangle, within areas that are today active mines, and to the Serra do Cipó region, were it was collected more than ten years ago, in forest fragments close to roadsides. Th e fact that no collections are known from northern areas of the Espinhaço range likely indicates that the distribution is extremely restricted to the region acround Serra do Cipó and the Iron Quadrangle. Eff orts to locate new populations of this species are urgent, especially considering that most areas where it might occur are currently owned by mining companies and are subject to an intensive land use.
Solanum psilophyllum is morphologically similar to S. verticillatum (described here below), another completely glabrous species of the Geminata clade occurring in the states of São Paulo and Rio de Janeiro. It can be distinguished from that species by its longer calyx lobes and by the swollen distal portions of the pedicels that are markedly constricted just below the calyx lobes. In addition, the leaf texture of S. psilophyllum is somewhat fl eshy, while leaves of S. verticillatum are brittle and chartaceous.
Solanum psilophyllum is also morphologically similar to S. lacteum from Atlantic forests in Rio de Janeiro, Espirito Santo and Minas Gerais states. It diff ers from that species in its larger fl owers (>1 cm in diameter), longer infl orescences, elliptic rather than obelliptic leaves that do not dry a blackish brown colour and in the non-lenticellate stem. Like S. lacteum, S. psilophyllum is completely glabrous. Solanum psilophyllum has an underground stem ( Figure 8C), like S. arboreum Dunal of northern South America and Central America (see Knapp 2002a); this characteristic may be more common in the Geminata clade than currently thought, as it is rare that the underground parts of these small shrubs are collected or even observed.    Description. Tree to 8 m, the branching appearing somewhat verticillate with branches in congested groups; young stems terete, completely glabrous, usually shiny; new growth completely glabrous and shiny, in live plants sometimes purplish green; bark of older stems pale yellow when dry, in live plants greyish brown. Sympodial units plurifoliate, the leaves clustered along the stems. Leaves simple, 4.5-16 cm long, 2-5 cm wide, elliptic to obelliptic, usually narrowly so, chartaceous and somewhat brittle, both surfaces glabrous and shiny, drying a golden brown; primary veins 6-10 pairs, drying yellowish brown, not looping in a submarginal vein; base acute to acuminate; margins entire, sometimes revolute; apex abruptly acute to attenuate; petiole (0.5-)1-2 cm long, glabrous, drying pale yellowish brown. Infl orescences 2-5 cm long, terminal, appearing axillary but this due to short internodes and congested leaves, branching 1-2 times, with 30-40 fl owers, completely glabrous; peduncle 0.5-2.5 cm long; pedicels 1.5-1.7 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, fi liform, spreading at anthesis, glabrous, articulated at the base; pedicel scars unevenly spaced 1-2 mm apart, usually clustered at the tips of the infl orescence branches. Buds ellipsoid, the corolla completely enclosed in the calyx when young, exserted 2/3 to 3/4 of the way just before anthesis. Flowers 5-merous, all perfect, intensely sweet-smelling (Custodio Filho 305). Calyx tube 1-1.5 mm long, conical, the lobes 0.9-1 mm long, ca. 1 mm wide, broadly deltate, with scarious margins and a central thickened keel ending in a rounded point, glabrous or the tips with a few papillae. Corolla (1.4-)1.6-1.8 cm in diameter, white, stellate, lobed nearly to the base, the lobes 6-8 mm long, 2.5-3.5(-4) mm wide, spreading at anthesis, densely papillate on the cucullate tips, otherwise completely glabrous. Stamens 4.5-6 mm long; fi lament tube 1 mm long or less, the free portion of the fi laments minute, <0.5 mm long, glabrous; anthers (3-)4-4.5 mm long, 1-1.2 mm wide, obellipsoid with the base narrower than the distal portion, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 5-7 mm long, glabrous; stigma minutely capitate, the surface papillose. Fruit a globose berry, 1-1.2 cm in diameter, pale green and white speckled (immature) becoming yellow or orange when ripe, the pericarp shiny and leathery, shattering when pressed and dried; fruiting pedicels 2-2.5 cm long, ca. 1 mm in diameter at the base, expanding gradually to ca. 2 mm in diameter at the apex, more or less woody, hanging; calyx lobes in fruit not markedly lengthening. Seeds 10-20 per berry, 5-5.5 mm long, 3-4 mm wide, reniform and somewhat fl attened, dark brown with paler margins, the surfaces minutely pitted and usually quite thin the embryo easily visible, the testal cells with sinuate margins.
Distribution and ecology. Endemic to south-eastern Brazil, in the states of Minas Gerais, Rio de Janeiro and São Paulo; in the Serra do Mar and Mantiequeira mountain chains ( Figure 15).
Ecology. Solanum verticillatum grows on the montane coastal forests (Mata Atlântica) as a small tree in forests and secondary growth from 700 to almost 2000 m elevation. Plants can be as large as 10 cm in diameter, and form part of the low canopy of these forests.
Phenology. Most fl owering specimens collected in the months of June and July; fruiting in November-January. Sporadic fl owering and fruiting apparently occurs throughout the year, but a fl owering peak occurs in the austral winter (May-August), which is also the drier season.
Etymology. Named for the pseudo-verticillate nature of the stems, where many branches appear to arise from a set of closely spaced nodes ( Figure 8F inset).
Preliminary conservation status (IUCN 2014). Least Concern (LC); EOO 75, 516 km 2 (LC); AOO 60 km 2 (EN). Although only described here, S. verticillatum is known from many localities along the Serra do Mar, many of which are from within protected areas (e.g., Reserva Biológica do Alto da Serra de Paranapiacaba in São Paulo state and Reserva Ecológica de Macaé de Cima, in Nova Friburgo, Rio de Janeiro state). Where it occurs, S. verticillatum is relatively common.
Notes. Solanum verticillatum was considered a montane form of S. evonymoides by Knapp (2008); fi eld collections in 2013 confi rmed the distinctness of this species. Solanum evonymoides is known from coastal forests in Bahia and adjacent Espirito Santo, and eastern Minas Gerais and although morphologically similar to S. verticillatum is distinct in both habitat and in several morphological features. Solanum verticillatum diff ers from S. evonymoides in its tree habit, branches that appear verticillate due to short internodes ( Figure 8F inset), smaller sweet-smelling fl owers (< 2 cm in diameter), shiny chartaceous leaves, and orange berries.
Solanum verticillatum also resembles S. psilophyllum (another set of specimens previously recognised as S. evonymoides by Knapp 2008) in its glabrous shiny leaves. It diff ers from that species in its more broadly deltate calyx lobes, its distinctly pedunculate infl orescences (versus infl orescences that branch only very near the base in S. psilophyllum), its berry that is orange or yellow-orange when ripe, and in its fl attened rather than ovoid seeds. Th ese two species can be very diffi cult to distinguish, but the marked constriction just below the calyx lobes at the distal end of the swollen pedicel occurs only in S. psilophyllum.
Th is species was commonly collected until approximately the 1980s and populations from the Paranapiacaba reserve are well represented in SP. It is strange that more recent collections do not seem to have been made; this may be due to the tree habit of S. verticillatum and to its similarity to the more common species S. campaniforme and S. pseudoquina A.St.Hil. It can be distinguished from S. campaniforme by its shiny, completely glabrous leaves (the leaves of S. campaniforme have tufts of trichomes in the vein axils abaxially) and from S. pseudoquina by its equal anthers (those of S. pseudoquina are markedly unequal). It diff ers from both species in its yellow or orange berries and pseudoverticillate branching.