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Acta Botanica Brasilica

Print version ISSN 0102-3306On-line version ISSN 1677-941X

Acta Bot. Bras., ahead of print  Epub Mar 19, 2018

http://dx.doi.org/10.1590/0102-33062017abb0417 

Article

New combinations and taxonomic notes for Tarenaya (Cleomaceae)

Raimundo Luciano Soares Neto1  * 
http://orcid.org/0000-0002-5643-9464

William Wayt Thomas2 
http://orcid.org/0000-0002-4996-536X

Maria Regina de Vasconcellos Barbosa3 
http://orcid.org/0000-0001-6166-3922

Eric H. Roalson4 
http://orcid.org/0000-0003-1655-3681

1 Programa de Pós-Graduação em Biologia Vegetal, Universidade Federal de Pernambuco, 50670-901, Recife, PE, Brazil

2 Institute of Systematic Botany, The New York Botanical Garden, 10458-5126, Bronx, New York, USA

3 Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, 58051-970, João Pessoa, PB, Brazil

4 School of Biological Sciences, Washington State University, 99164-4236, Pullman, Washington, USA

ABSTRACT

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.

Keywords: Cleome; Hemiscola; nomenclature; Tarenaya; type

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 In 1952, when Iltis [unpublished] revised the New World Cleome species, the genus was treated as a section, Cleome sect.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), and Costa-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 recognition of Cleome as a paraphyletic genus (Hall et al. 2002; Sanchez-Acebo 2005; Hall 2008; Inda et al. 2008; Feodorova et al. 2010, Patchell et al. 2014) has led to the deconstruction of Cleome s.l., and consequently multiple genera have been segregated and reorganized (Iltis & Cochrane 2007; 2014; 2015; Cochrane & Iltis 2014; Roalson et al. 2015; Thulin & Roalson 2017; Barrett et al. 2017; Roalson & Hall 2017; Soares Neto et al. 2017), including Tarenaya (Iltis & Cochrane 2007). This taxonomic reorganization was based on morphology, chromosome number, and phylogenetic relationships, and specific nomenclatural recombinations were made for Flora of North America (Iltis & Cochrane 2007) and Flora Mesoamericana (Iltis & Cochrane 2014; 2015).

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.

Results and discussion

Tarenaya Raf., Sylva Tellur. 111. 1838. Type: Tarenaya spinosa (Jacq.) Raf., Sylva Tellur. 111. 1838. Cleome spinosa Jacq., Enum. Syst. Pl. 26. 1760. Cleome sect. Tarenaya (Raf.) Iltis, Novon 15(1): 146. 2005.

=Hemiscola Raf., Sylva Tellur. 111. 1838. Type: Hemiscola aculeata (L.) Raf., Sylva Tellur. 111. 1838. Cleome aculeata L., Syst. Nat., ed. 12. 3: 232. 1768.

=Scolosperma Raf., Sylva Tellur. 111. 1838. Type: Scolosperma dendroides (Schult.f.) Raf., Sylva Tellur. 111. 1838. Cleome dendroides Schult.f., Syst. Veg., ed. 15 bis [Roemer & Schultes] 7(1): 28. 1829.

=Lianodes Kuntze, Lex. Gener. Phaner. 129. 1903. nom. nud., nom. inval.

=Neocleome Small, p.p., Man. S.E. Fl. 577. 1933. Type, designated here: Neocleome spinosa (Jacq.) Small, Man. S.E. Fl. 577. 1933. Cleome spinosa Jacq., Enum. Syst. Pl. 26. 1760.

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.

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 horseshoe-shaped, longitudinally striate and transversely ridged, cleft covered by a membrane attaching both tips (cotyledonar and radicular “claws”).

1. Tarenaya aculeata (L.) Soares Neto & Roalson, comb. nov. Cleome aculeata L., Syst. Nat., ed. 12. 3: 232. 1768. Type: Habitat in America, D. Zoega s.n. (lectotype, designated by Al-Shehbaz (1988): BM-LINN 850.17 [image!]). ≡ Hemiscola aculeata (L.) Raf., Sylva Tellur. 111. 1838.

2. Tarenaya afrospina (Iltis) Soares Neto & Roalson, comb. nov. Cleome afrospina Iltis, Amer. J. Bot. 54(8): 955. 1967. Type: REPUBLIC CONGO. Environs Coquilhatville, Ile du Fleuve [du R. Congo], 28 September 1928 [1925], W. Robyns 767 (holotype: BR 88677575 [image!]; isotypes: BR 8867063, [image!], BR 88677070 [image!], BR 8867520 [image!], GH GH00042345 [image!], K K000230602 [image!], US US00100469 [image!]).

3. Tarenaya atropurpurea (Schott) Soares Neto & Roalson, comb. nov. Cleome atropurpurea Schott, in Schreib. Nachr. Oestr. Naturf. 1: 129. 1820. Type: BRAZIL. Sine loco, H.W. Schott s.n. (lectotype, designated here: W 0060314 [image!]). Additional syntype: BRAZIL. Rio de Janeiro, H.W. Schott 4442 (W 0060315 [image!]).

= Cleome dendroides Schult.f., Syst. Veg., ed. 15 bis 7(1): 28. 1829. Cleome arborea Weinm., Syll. Pl. Nov. 1: 227. 1824, nom. illeg., non C. arborea Schrad. 1821. Neotype, designated here: t. 3296 in Curtis’s Botanical Magazine, v. 61, ser. 2: v. 8. 1834. ≡ Scolosperma dendroides (Schult.f.) Raf., Sylva Tellur. 111. 1838.

Notes-The species commonly known as Cleome dendroides requires the application of an older name. Here we make the combination Tarenaya atropurpurea, as Cleome atropurpurea is the oldest name for this taxon. Iltis considered this name “nom. subnud.” as annotated on the syntype Schott 4442; however, Schott clearly presents a 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.

4. Tarenaya bicolor (Gardner) Soares Neto & Roalson, comb. nov. Cleome bicolor Gardner, London J. Bot. 2: 330. 1843. Type: BRAZIL. [Rio de Janeiro] Organ Mountains, hab. in open rocky and cultivated places, April 1837, G. Gardner 309 (lectotype, designated here: BM BM000573983 [image!]; isolectotypes: E E00326206 [image!], E E00326207 [image!], F, G G00226190 [image!], G G00226191 [image!], GH 00042326 [image!], K K000220447 [image!], K K000220448 [image!], P P00076106 [image!], NY 00215151!, US 00100471 [image!], WIS 0258903!).

Notes- Gardner 309 has a large number of known duplicates designated only as “typus” without any herbarium holding a clear holotype. Many of these specimens are good representatives of the species, bearing flowers and mature fruits, characters useful in recognizing this species. The specimen BM000573983, however, is from the Gardner herbarium and is labeled with Gardner’s handwriting and, therefore, is being designated as the lectotype.

5. Tarenaya boliviensis (Iltis) Soares Neto & Roalson, comb. nov. Cleome boliviensis Iltis, Novon 15(1): 147. 2005. Type: BOLIVIA. La Paz: “Plantae Andium Boliviensium, Prov. Larecaja, vicinus Sorata, Challapampa, ad rivum, in schistosis, Reg.[ion] temp[lada], 2600 m, September 1857‒ January 1958”, G. Mandon 938 (holotype: WIS 0258904!, isotypes: BM, F, G G00226276 [image!], G G00226277 [image!], GH 00042328 [image!], K K000220474 [image!]; LE 00001884 [image!], LE 00001885 [image!], LPB, MPU 013699 [image!], MPU 013700 [image!], NY!, P P00076103 [image!], P P00741914 [image!], P P00741915 [image!], PR, S S-R-7312[image!], W, WIS 0258905!).

6. Tarenaya chapalensis (Iltis) Soares Neto & Roalson, comb. nov. Cleome chapalensis Iltis, Bol. Inst. Bot. Univ. Guadalajara 5: 428. 1998. Type: [MEXICO. Michoacán] Wet roadsides and shallow water courses, on road to La Barca, 1/3 km N of the R.R., station at Zamora, 19°58’N, 102°16’W, 1520 m, 29 July 1960, H.H. Iltis et al. 490 (holotype: WIS WIS0258906!, isotypes: BM BM000629040 [image!], ENCB, F [image!], G, GH 00042307 [image!], GUADA, IBUG IBUG0001788 [image!], IEB 000157790 [image!], K K000220427 [image!], LE, LIL, MEXU 00140923 [image!], MICH 112011 [image!], MO 107291!, MSC, NY 00387673!, SMU, TEX, UC 1332256 [image!], US, WIS WIS0258910!, WIS0258907!, WIS0259808!, WIS0258909!, WIS0258912!, WIS 0258911!, WIS0258913!, XAL XAL0106658 [image!], ZEA).

7. Tarenaya crenopetala (A.DC.) Soares Neto & Roalson, comb. nov. Cleome crenopetala A.DC., Mém. Soc. Phys. Genève 6(1): 220. t. 2. 1833. Lectotype, designated here: t. 2. in DC. & A.DC., Cinq. notice sur. les pl. rares du Jardin de Genève, Mém. Soc. Phys. Genève 6 (1). 1833).

Notes-According to the protologue, the description of this species was based on an individual growing in the gardens of Geneva from seeds from Uruguay. No specimen was found in any herbaria matching the type, and, therefore, the illustration presented together with the original description is being here designated as the lectotype.

We are also correcting here the authorship of Cleome crenopetala that has been erroneously attributed to Augustin de Candolle, but was in fact described by Alphonse de Candolle, as can be seen at the end of the original description.

8. Tarenaya diffusa (Banks ex DC.) Soares Neto & Roalson, comb. nov. Cleome diffusa Banks ex DC., Prodr. 1: 241. 1824. Type: BRAZIL. Rio de Janeiro: v. s. in h. Banks (holotype: BM BM000573979 [image!]). ≡ Cleome aculeata var. diffusa (Banks ex DC.) Kuntze, Revis. Gen. Pl. 3(3): 7. 1898. ≡ Hemiscola diffusa (Banks ex DC.) Iltis, Novon 17(4): 448. 2007.

9. Tarenaya domingensis (Iltis) Soares Neto & Roalson, comb. nov. Cleome domingensis Iltis, Brittonia 10: 56. 1958. Replaced name: Cleome erosa Urb., Symb. Antill. (Urban) 7(2): 224. 1912, nom. illeg., non Cleome erosa (Nutt.) Eaton, 1836. Type: DOMINICAN REPUBLIC. [Santo Domingo] Hab. in Sto. Domingo ad Rio Jimenoa, 1190 m alt., in margine sylvarum, May 1910, H. von Türckheim 3303 (holotype: B B100242710 [image!]; isotype: NY 00074435!).

10. Tarenaya eosina (J.F.Macbr.) Soares Neto & Roalson, comb. nov. Cleome eosina J.F.Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 170. 1929. Replaced name: Cleome microcarpa Hassl., Repert. Spec. Nov. Regni Veg. 12: 254. 1913, nom. illeg., non Cleome microcarpa Ule 1908. Type: PARAGUAY. Bella Vista, E. Hassler 11013 (holotype: G; isotype: MPU MPU013707 [image!]).

11. Tarenaya guaranitica (Chodat & Hassl.) Soares Neto & Roalson, stat. & comb. nov. Cleome rosea Vahl ex DC. var. guaranitica Chodat & Hassl., Bull. Herb. Boissier, sér. 2, 3: 797. 1903. Type: PARAGUAY. Chololo: ad marginem silvae, in valle fluminis Y-aca, December 1900, E. Hassler 6707 (holotype: G G001064449 [image!]; isotypes: B B100242716 [image!], BM BM000537896 [image!], G G00106446 [image!], G G00106447 [image!], G G00106448 [image!], K K000220457 [image!], NY 00215161!, P P00076111 [image!], S S-R-7343 [image!], WIS WIS0259073!; WIS0259074 [frag!]). ≡ Cleome guaranitica Briq., Annuaire Conserv. Jard. Bot. Genève 17: 371. 1914.

12. Tarenaya horrida (Mart. ex Schult.f.) Roalson & Soares Neto, comb. nov. Cleome horrida Mart. ex Schult. & Schult.f., Syst. Veg., ed. 15 bis 7(1): 32. 1829. Type: BRAZIL. Espírito Santo: Itapemirim, Prinz Max. von Neuwied s.n. (lectotype, designated here: BR 698551 [image!], isolectotypes: BR 698580 [image!]; WIS 0258937!). ≡ Cleome spinosa L. f. horrida (Mart. ex Schult. & Schult. f.) Eichler, Fl. Bras. 13(1): 253. 1865. ≡ Cleome spinosa Jacq. var. horrida (Mart. ex Schult. & Schult. f.) Fawc. & Rendle, Fl. Jamaica 3: 226. 1914.

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.

14. Tarenaya inermis (Malme) Soares Neto & Roalson, comb. nov. Cleome inermis Malme, Ark. Bot. 22A, no. 7: 5. 1928. Type: BRAZIL. Rio Grande do Sul: Cachoeira, in silvula clara, 10 January 1902, G. Malme II 1041 (lectotype, designated here: S S-R-7321 [image!]; isolectotypes: B B 10 0242693 [image!], S S10-16936 [image!], WIS 0258939!, WIS 0258940!).

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.

15. Tarenaya latifolia (Vahl ex DC.) Soares Neto & Roalson, comb. nov. Cleome latifolia Vahl ex DC., Prodr. 1: 239. 1824. Type: [GUIANA] In Guiana, v. s. in h. Vahl. (neotype, designated here: French Guiana, 1858, P.A. Sagot 1170, P P00076114 [image!]; isoneotypes: GOET [photo at WIS], P P00745091 [image!], P P00076129 [image!]).

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.

16. Tarenaya microcarpa (Ule) Soares Neto & Roalson, comb. nov. Cleome microcarpa Ule, Engl., Bot. Jarhb. Syst. 42: 201. 1908. Type: BRAZIL. Piauhy [Piauí]: Catinga der Serra Branca, January 1907, E. Ule 7428 (lectotype, designated here: HBG HBG-522391 [image!]; isolectotypes: B B 10 0242701 [image!], K 000220462 [image!], F neg 5766 [image!]).

Notes-HBG-522391 is being designated as lectotype, because of the two collections seen by Ule, the collection at HBG is the one labeled as “Cleome microcarpa s. nv.”, while the specimen at B was labeled first as “Cleome viridiflora s. nv.” (crossed off) and later as C. chlorantha s. nv. (ined.).

17. Tarenaya psoraleifolia (DC.) Soares Neto & Roalson, comb. nov. Cleome psoraleifolia DC., Prodr. 1: 239. 1824. Type: BRAZIL. v. s. in h. Mus. Par. [A. Ferreira s.n.] (holotype: P P00141286 [image!]).

18. Tarenaya regnellii (Eichler) Soares Neto & Roalson, comb. nov. Cleome regnellii Eichler, Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 190: 1870. Type: BRAZIL. Ad Caldas in prov. Minas Geraës [Minas Gerais], A.F. Regnell coll. III n. 147 (lectotype, designated here: BR 698554 [image!]; isolectotypes: B B 10 0242684 [image!], BR 698555 [image!], F neg. [image!], K K000220449 [image!], LD 1314574 [image!], LE 00001896 [image!], S (2 sheets) [image!], WIS 0259069!, WIS 0259070!).

Notes- We here designate specimen BR 698554 as the lectotype for several reasons. The B specimen label bears three names, each in a different script, entered in the sequence Cleome rosea, C. polygama (scratched), and then C. regnellii, the latter in a fancy penmanship that does not appear to be that of Eichler. On that label in this same calligraphy also appears a full citation of the place of publication of Eichler’s species, making it unlikely that this annotation pre-dated the publication of his 1870 paper. These facts impede the selection of the B specimen as lectoype. However, two sheets in BR, BR 698554 and BR 698555, are annotated as C. regnellii in Eichler's handwriting. Eichler, who with Urban completed the Flora Brasiliensis, certainly saw Martius’s herbarium, which included specimens collected by many others and which is now at BR. For these reasons, it seems plausible that the BR collections, having been seen by Eichler, are the proper type material. BR 698554 is a better, more complete specimen than BR 698555 and is therefore chosen as the lectotype.

19. Tarenaya rosea (Vahl ex DC.) Soares Neto & Roalson, comb. nov. Cleome rosea Vahl ex DC., Prodr. 1: 239.1824. Type: [BRAZIL. Rio de Janeiro] Ad Rio-Janeiro Brasiliae. v. s. in. h. Juss., P. Commerson 255 (holotype: P P00671649!).

20. Tarenaya siliculifera (Eichler) Soares Neto & Roalson comb. nov. Cleome siliculifera Eichler, Fl. Bras. 13(1): 260. 1865. Type: [BRAZIL. Minas Gerais] Prov. Minarum: Habitat prope Cocaes et in Serra dos Pinheiros, [1839], J. Pohl s.n. (lectotype, designated here: BR 698586 [image!] ; isotypes: F (frag. + neg.) [image!], G G00226314 [image!], M M-0010385 [image!], LE, W [image!], WIS 0259077!).

Notes-As with the type of C. regnellii above, it is most likely that the BR specimen represents material used by Eichler, and we therefore lectotypify C. siliculifera with BR 698586.

21. Tarenaya titubans (Speg.) Soares Neto & Roalson, comb. nov. Cleome titubans Speg., Anales Soc. Ci. Argent. 15: 97. 1883. Type: ARGENTINA. [Buenos Aires] Província Bonaërensi: Inter dumeta in paludosis secus, “el Rio de la Plata”, loco dicto “las Conchas”, May 1881, C.L. Spegazzini s.n. (neotype, designated here: ARGENTINA. Buenos Aires, Quilmes, loc. Bernal, March 1973, F.M. Rodríguez 191, SI 066837 [image!]; isoneotype: SI 066836 [image!]).

Notes-The Spegazzini type collection at LP does not appear to include the type of Cleome titubans, as expected. Costa-e-Silva (2000) also referenced her inability to find the type of this species for her review of Brazilian Cleome. Because no original material of C. titubans has been found among Spegazzini’s collections, a neotype is designated here.

22. Tarenaya torticarpa (Iltis & T. Ruiz Zapata) Soares Neto & Roalson, comb. nov. Cleome torticarpa Iltis & T. Ruiz Zapata, Novon 7(4): 367. 1997. Type: VENEZUELA. Estado Falcón: Distrito Federación, Parque Nacional Cueva de la Quebrada el Toro, 10°50’N, 69°07’W, en selva de galería, bajo sombra, 200 m abajo de la “toma de agua” en la estácion del Parque, 600 m, 29 October 1983, Thirza Ruiz Zapata & Teo Ruiz 4138 (holotype: MY; isotypes: COL, F, K, MER, MO!, NY NY00387680!, NY NY00345549!, US, VEN, WIS WIS0259083!, WIS0259084!, WIS0259085!).

23. Tarenaya trachycarpa (Klotzsch ex Eichler) Soares Neto & Roalson, comb. nov. Cleome trachycarpa Klotzsch ex Eichler, Fl. Bras. 13(1): 252. 1865. Type: [BRAZIL] Habitat in Brasilia austro-orientali, Sello [F. Sellow] 2059 (holotype: B 10 0242688 [image!]). ≡ Cleome psoraleifolia DC. var. trachycarpa (Klotzsch ex Eichler) Kuntze, Revis. Gen. Pl. 3(3): 7. 1898.

24. Tarenaya virens (J.F.Macbr.) Soares Neto & Roalson, comb. nov. Cleome virens J.F.Macbr., Candollea 5: 360. 1934. Type: PERU. Loreto: La Victoria on the Amazon River, August-September 1929, L. Williams 2596 (holotype: F [image!]; isotype: G G00226308 [image!]).

25. Tarenaya werdermannii (Alf.Ernst) Soares Neto & Roalson, comb. nov. Cleome werdermannii Alf.Ernst., Notizbl. Bot. Gart. Berlin-Dahlem 13: 378. 1936. Type: BOLIVIA. Depto. Sta. Cruz: Missiones Guarayos-Sta. Cruz de la Sierra, ca. 250‒300 m ü. M., October 1926, E. Werdermann 2597 (lectotype, designated here: B B10 0242682 [image!]; isolectotypes: B B10 0242698 [image!], LPB [image!], MO 1000057!, S [image!], WIS WIS0259089 [frag.]!).

Notes-Both specimens of Werdermann 2597 at B are good representatives for this species, but B 100242682 is chosen as the lectotype, because it is labeled “typus” in Ernst’s hand.

Acknowledgements

We acknowledge the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES, for the grant awarded for his doctoral scholarship, the Sandwich Doctoral scholarship (Process 88881.132147/2016-01), and the Missouri Botanical Garden for the Shirley Graham Fellowship. We thank to Olof Ryding at C for helping us with the Vahl collections and unpublished manuscripts, and Laura Iharlegui at LP for searching for Spegazzini collections for us. Many thanks to Theodore Cochrane at WIS for a thorough critique of the manuscript.

References

Al-Shehbaz IA. 1988. Capparaceae. In: Howard RA. (ed.) Flora of the Lesser Antilles. Vol. 4. Cambridge, Arnold Arboretum/ Harvard University. p. 293-310. [ Links ]

Arana MD, Oggero AJ. 2016. New combinations in Tarenaya (Cleomaceae) for the Argentinian flora. Phytotaxa 267: 162-164. [ Links ]

Barrett RL, Roalson EH, Ottewell K, et al. 2017. Resolving generic boundaries in Indian-Australasian Cleomaceae: circumscription of Areocleome, Arivela, and Corynandra as distinct genera. Systematic Botany 42: 694-708. [ Links ]

Cochrane TS, Iltis HH. 2014. Studies in the Cleomaceae VII: five new combinations in Corynandra, an earlier name for Arivela. Novon 23: 21-26. [ Links ]

Costa-e-Silva MB. 2000. O gênero Cleome L. (Capparaceae Juss.) para o Brasil. PhD Thesis, Universidade Federal Rural de Pernambuco, Recife. [ Links ]

Candolle AP. 1824. Prodromus systematis naturalis regni vegetabilis. Vol. 1. Paris, Sumptibus Sociorum Treuttel et Würtz. p. 241. [ Links ]

Eichler AW. 1865. Capparideae. In: Martius CFP. (ed.) Flora Brasiliensis. Vol. 13(1). München, Leipzig. p. 253-254. [ Links ]

Feodorova TA, Voznesenskaya EV, Edwards GE, Roalson EH. 2010. Biogeographic patterns of diversification and the origins of C4 in Cleome (Cleomaceae). Systematic Botany 35: 811-826. [ Links ]

Grisebach AHR. 1864. Flora of the British West Indian islands. London, Lovell Reeve & Co. p. 16. [ Links ]

Hall JC. 2008. Systematics of Capparaceae and Cleomaceae: an evaluation of the generic delimitations of Capparis and Cleome using plastid DNA sequence data. Canadian Journal of Botany 86: 682-696. [ Links ]

Hall JC, Sytsma KJ, Iltis HH. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. American Journal of Botany 89: 1826-1842. [ Links ]

Hooker WJ. 1834. Curtis’s Botanical Magazine. v. 61. ser. 2: v. 8. London. [ Links ]

Iltis HH. 1952. A revision of the New World species of Cleome. PhD Thesis, Washington University, St. Louis. [ Links ]

Iltis HH. 1967. Studies in the Capparidaceae XI: Cleome afrospina, a tropical African endemic with neotropical affinities. American Journal of Botany 54: 953-962. [ Links ]

Iltis HH. 2005. Studies in the Cleomaceae II: Cleome boliviensis, a new, spiny, large-flowered Andean species. Novon 15: 146-155. [ Links ]

Iltis HH, Cochrane TS. 2007. Studies in Cleomaceae V: a new genus and ten new combinations for the Flora of North America. Novon 17: 447-451. [ Links ]

Iltis HH, Cochrane TS. 2014. Studies in Cleomaceae VI: a new genus and sixteen new combinations for the Flora Mesoamericana. Novon 23: 51-58. [ Links ]

Iltis HH, Cochrane TS. 2015. Cleomaceae. In: Davidse G, Sánchez MS, Knapp S, Cabrera FC. (eds.) Flora Mesoamericana. Vol. 2(3). St. Louis, Missouri Botanical Garden. p. 256-267. [ Links ]

Iltis HH, Zapata TR. 1997. Studies in the Capparidaceae XIX: Cleome torticarpa n. sp., a Venezuelan endemic. Novon 7: 367-372. [ Links ]

Inda LA, Torrecilla P, Catalán P, Ruiz-Zapata T. 2008. Phylogeny of Cleome L. and its close relatives Podandrogyne Ducke and Polanisia Raf. (Cleomoideae, Cleomaceae) based on analysis of nuclear ITS sequences and morphology. Plant Systematics and Evolution 274: 111-126. [ Links ]

Jacobs M. 1960. Capparidaceae. In: Steenis CGGJ. (ed.) Flora Malesiana. Vol. 6(1), ser. 1. Jakarta, Noordhoff-Kolff N.V. p. 61-105. [ Links ]

Patchell MJ, Roalson EH, Hall JC. 2014. Resolved phylogeny of Cleomaceae based on all three genomes. Taxon 63: 315-328. [ Links ]

Rafinesque CS. 1838. Sylva Telluriana: trees and shrubs of North America, and other parts. Philadelphia, Author. [ Links ]

Roalson EH, Hall JC. 2017. New generic concepts for African Cleomaceae. Systematic Botany 42: 925-942. [ Links ]

Roalson EH, Hall JC, Riser JP, Cardinal-McTeague WM, Cochrane TS, Sytsma KJ. 2015. A revision of generic boundaries and nomenclature in the North American cleomoid clade (Cleomaceae). Phytotaxa 205: 129-144. [ Links ]

Sánchez-Acebo L. 2005. A phylogenetic study of the New World Cleome L. (Brassicaceae-Cleomoideae). Annals of the Missouri Botanical Garden 92: 179-201. [ Links ]

Small JK. 1933. Manual of southeastern flora. Chapel Hill, The University of North Carolina Press. [ Links ]

Soares Neto RL, Barbosa MRV, Roalson EH. 2017. Cleoserrata (Cleomaceae): taxonomic considerations and a new species. Phytotaxa 324: 179-186. [ Links ]

Soares Neto RL, Barbosa MRV, Roalson EH. 2018. Two new species of Tarenaya (Cleomaceae) for Brazil. Phytotaxa 334: 28-34. [ Links ]

Thulin M, Roalson EH. 2017. Resurrection of the genus Rorida (Cleomaceae), a distinctive Old World segregate of Cleome. Systematic Botany 42: 569-577. [ Links ]

Received: December 12, 2017; Accepted: February 19, 2018

* Corresponding author: lucianosoares.rdon@gmail.com

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