New combinations and taxonomic notes for Tarenaya ( Cleomaceae )

Tarenaya clade includes 37 species based on phylogenetic relationships and on the stipular spine synapomorphy, however only 10 species thought to belong to the genus have had names established in Tarenaya. Besides the two new species are being described, we present 25 new combinations for the species and refine the typification of 13 species. Ten lectotypes and three neotypes are designated here. One generic synonym is also typified.


Introduction
Tarenaya was segregated from Cleome by Rafinesque (1838) with one species, T. spinosa (based on Cleome spinosa), because he did not agree with the broad concept of Cleome.He based his proposal on the habit, type of leaf, number of leaflets, number of stamens, presence and morphology of nectaries, length of gynophore, and geographic distribution.Tarenaya was generally not adopted in taxonomic treatments and was instead considered to be a synonym of Cleome for the next ~170 years.In 1952, when Iltis [unpublished] revised the New World Cleome species, the genus was treated as a section, Cleome sect.Tarenaya, comprising 24 species and characterized by spines and prickles of epidermal origin (characters not shared by three species and one subspecies, currently recognized under Cleoserrata Iltis and unarmed forms in various species), seeds with an aril and a large cleft.Later, Jacobs (1960), Iltis (1967), Iltis &Zapata (1997), andCosta-e-Silva (2000) used Tarenaya informally in their classifications, and the name was finally formally established as a section by Iltis (2005) when he described a new species of Cleome from the Andes.By that time Iltis (2005) considered section Tarenaya to comprise a group of 40 species from the New World and one from East Africa.
The Tarenaya clade is defined by an easily recognized synapomorphy: a pair of spines at the base of the petioles (stipular spines) and is well supported in all phylogenetic studies of the family (Hall et al. 2002;Sanchez-Acebo 2005;Hall 2008;Inda et al. 2008;Feodorova et al. 2010, Patchell et al. 2014;Barrett et al. 2017).Not all of the presumed species of Tarenaya have been sampled for molecular phylogenetic studies, but based on those sampled to date, and on the stipular spine synapomorphy, Tarenaya comprises 37 species.Hemiscola, another segregate genus recognized by Iltis & Cochrane (2007), shares the stipular spines and is embedded within Tarenaya.For these reasons, the unification of these genera in a broader concept of Tarenaya has been proposed (Feodorova et al. 2010, Patchell et al. 2014).Its species range from Mexico to Argentina, except for one species found exclusively in East Africa (Iltis 2005;Iltis & Cochrane 2015).
Despite broad agreement that Tarenaya should be recognized at the generic level, only 10 of the 37 species thought to belong to the genus have had names established in Tarenaya (Iltis & Cochrane 2007;2014;Arana & Oggero 2016;Soares Neto et al. 2018).In addition to these species, two new species are being described (RL Soares Neto et al. unpubl. res.).In this study, we present 25 new combinations for those species hypothesized to be part of the Tarenaya lineage, but that do not have nomenclatural combinations within Tarenaya.We also include combinations necessary to include Hemiscola with Tarenaya.Ten lectotypes, three neotypes, and one typification of a generic name are provided to stabilize the nomenclature discussed here.

Materials and methods
Type specimens that have been examined in person by the authors are annotated with exclamation marks.Images of type specimens examined online are annotated as "[image!]", and a barcode number, if available, is given.Original protologues were studied for all names treated here.Notes- Small (1933), when delimiting the genus Neocleome, made two new combinations, N. spinosa and N. serrata, but did not designate a type species for the new genus.One of these species is now placed in Tarenaya (N.spinosa), but the other (N.serrata) is assigned to the genus Cleoserrata.We believe Small intended to include in Neocleome a broader diversity of Tarenaya (he states: "… about 70 species, natives, mainly, of tropical regions.")and we therefore typify Neocleome based on Tarenaya spinosa.

Results and discussion
Herbs to subshrubs or shrubs, annual or perennial, branched from the base; pubescent to puberulent-glandular indument at branches or totally glabrous; stipular spines at the base of petioles; leaves palmately-compound with 3-7(-12) leaflets; racemes corymbiform, flowers bracteate, the lower with leaf-like bracts with 3-5 leaflets, becoming 1-foliolate in the inflorescence axis; flowers tetramerous, zygomorphic petals unguiculate, white or white becoming pink or purplish at apex, pink to purplish, or a pair of each color; nectary annular; stamens 6, elongated by a short androgynophore enrolled by the nectary; mature capsules cylindrical, ellipsoid, fusiform, oblanceoloid, sessile or short to long elongated stalks; seeds horseshoeshaped, longitudinally striate and transversely ridged, cleft covered by a membrane attaching both tips (cotyledonar and radicular "claws").
diagnosis of this species with his publication of Cleome atropurpurea.For this reason, we apply that name here.There are two original collections of Schott at W, and we designate W 0060314 as the lectotype as it is the more complete specimen with leaves and flower buds.The other syntype material is of significant importance, though, as it includes mature fruits, which are lacking from the lectotype.
Cleome arborea Weinm.was described based in an individual cultivated in the Horto Imperiali Paulowskiensi, near St. Petersburg, from seeds from Brazil.Schultes filius (1829) renamed Cleome arborea as C. dendroides, taking into account the earlier homonym, C. arborea Kunth.Schultes filius cited one collection of Weinmann's, probably the type, which was not found in LE nor in any other herbarium.Hooker (1834) in Curtis's Botanical Magazine presented notes about this Cleome, improving its description and also presenting a beautiful illustration highlighting the stipular spines, purple flowers, fruits, and seeds.This illustration is here chosen as the neotype for this name.
Notes-Two specimens at BR are annotated as the holotype.Both are from the same collection by Prinz Max.von Neuwied and so are presumed syntypes.We here designate BR 698551 as the lectotype because it has both flowers and fruits on it, both of which are necessary for distinguishing T. horrida from closely related species.
13. Tarenaya houstonii (R.Br.)Soares Neto & Roalson, comb. nov. Cleome houstonii R.Br. Hort. Kew., ed. 2 [W.T. Aiton] 4: 131. 1812. Lectotype, designated here: t. 45. in Martyn, Hist. Plant. Rar. v. 5, 1736.Notes-Cleome houstonii was described based on Sinapistrum indicum, which was described based on plants grown from seeds from Cuba and cultivated in England by Houston in the year 1730.Although Candolle (1824) cited dried material from Jamaica, and Grisebach (1864) also cited a material collected by Houston from Jamaica, no specimen has been found for this collection.The illustration presented with the description of Sinapistrum indicum is a good plate highlighting a branch with bud, flowers, and fruits and provides details of the petals, stamens, and ovary.This plate provides all of the characters necessary to recognize the species and is therefore here designated as the lectotype.Notes-There are two type collections of Malme 1041 in S, and therefore one needs to be designated the lectotype.We designate S-R-7321 as the lectotype as it is more mature, having expanded inflorescences and mature fruits.
Notes-De Candolle described Cleome latifolia based on a specimen seen in the Vahl herbarium; however, we have found no records of such specimen in C (Olof Ryding, pers.comm.), nor in G-DC.Given such, we are designating as the neotype Sagot 1170, one of the specimens studied by Eichler (1865) for Flora Brasiliensis and Iltis (1952) in his revision of New World Cleome.