Menispermaceae) 9 Doi: 10.3897/Phytokeys.100.21828 MONOGRAPH Launched to Accelerate Biodiversity Research
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A peer-reviewed open-access journal PhytoKeys 100: 9–89A taxonomic(2018) revision of Curarea Barneby & Krukoff (Menispermaceae) 9 doi: 10.3897/phytokeys.100.21828 MONOGRAPH http://phytokeys.pensoft.net Launched to accelerate biodiversity research A taxonomic revision of Curarea Barneby & Krukoff (Menispermaceae) Rosa del C. Ortiz1 1 Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, USA Corresponding author: Rosa del C. Ortiz ([email protected]) Academic editor: C. Morden | Received 23 October 2017 | Accepted 13 April 2018 | Published 21 June 2018 Citation: Ortiz RdC (2018) A taxonomic revision of Curarea Barneby & Krukoff (Menispermaceae). PhytoKeys 100: 9–89. https://doi.org/10.3897/phytokeys.100.21828 Abstract A monograph of Curarea, a neotropical genus in the plant family Menispermaceae, is presented. Curarea is distinguished from related genera by the combination of staminate flowers with sepals in two whorls and pistillate flowers with three petals, three carpels and usually elongated carpophores bearing three ses- sile drupelets. Nine species are recognised, amongst them two new to science, C. gentryana from Ecuador and C. barnebyana, from Ecuador and Peru. Additionally, two new combinations, C. iquitana and C. tomentocarpa, are proposed for distinct taxa recovered in a multivariate analysis of quantitative characters of the broadly distributed and morphologically variable C. toxicofera. The anatomy and morphology of species in the genus is documented, identification key, species descriptions, distribution maps and a pre- liminary conservation assessment for all accepted species are also provided. Of the nine species recognised here, C. barnebyana is assigned a preliminary status of Vulnerable, C. crassa (known only from the coastal Atlantic Forest in Brazil) and C. gentryana (endemic to western Ecuador) are both assigned a preliminary status of Endangered. Keywords Arrow poison, curare, dioecious liana, neotropical clade, Ranunculales Copyright Rosa del C. Ortiz. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 10 Rosa del C. Ortiz / PhytoKeys 100: 9–89 (2018) Table of contents Introduction ........................................................................................................... 10 Materials and methods ........................................................................................... 13 General morphology ...................................................................................... 19 Habit and stem .............................................................................................. 19 Wood anatomy .............................................................................................. 19 Leaves ............................................................................................................ 20 Indumentum .................................................................................................. 21 Venation pattern ............................................................................................ 21 Leaf anatomy: adaxial cuticle, transverse sections of petiole, lamina and mid-vein ...23 Inflorescences ................................................................................................. 26 Flowers........................................................................................................... 28 Infructescences and drupelets ......................................................................... 29 Pollen morphology ......................................................................................... 30 Phytochemistry .............................................................................................. 30 Ethnobotany .................................................................................................. 30 Geographical distribution and habitat ............................................................ 31 Phylogenetic affinities .................................................................................... 32 Multivariate analyses of Curarea toxicofera s.l. ................................................. 32 Taxonomic treatment ............................................................................................. 35 Curarea Barneby & Krukoff ............................................................................... 35 Key to the species of Curarea .......................................................................... 37 1. Curarea barnebyana R.Ortiz, sp. nov........................................................... 38 2. Curarea candicans (Rich. ex DC.) Barneby & Krukoff ................................ 43 3. Curarea crassa Barneby ............................................................................... 49 4. Curarea cuatrecasasii Barneby & Krukoff .................................................... 53 5. Curarea gentryana R.Ortiz, sp. nov. ............................................................ 58 6. Curarea iquitana (Diels) R.Ortiz, comb. nov. ............................................. 61 7. Curarea tecunarum Barneby & Krukoff ...................................................... 66 8. Curarea tomentocarpa (Rusby) R.Ortiz, comb. nov. .................................... 72 9. Curarea toxicofera (Wedd.) Barneby & Krukoff .......................................... 78 Tribute ................................................................................................................... 84 Acknowledgments .................................................................................................. 85 References .............................................................................................................. 85 Supplementary material 1 ....................................................................................... 89 Introduction Curarea Barneby & Krukoff of the plant family Menispermaceae, tribe Tiliacoreae, is a genus of understory or canopy dioecious lianas that are widely distributed in the humid forests of tropical America, mostly at low elevation from eastern Brazil to Costa Rica in Central America. Within Tiliacoreae, Curarea shares with remain- A taxonomic revision of Curarea Barneby & Krukoff (Menispermaceae) 11 ing members of the tribe endocarps ornamented by longitudinal ribs, seeds hippo- crepiform and without endosperm and fleshy cotyledons (Ortiz et al. 2016). Curarea was segregated from Chondrodendron Ruiz & Pav. to include species that differed in several floral and carpological features (Barneby and Krukoff 1971). InCurarea , the calyx of staminate flowers is composed of two whorls of three, scarcely fleshy and greenish sepals, these being variously described as sepaloid scales (Barneby and Kru- koff 1971: 7) or as sepals (Barneby 1996). InChondrodendron , the calyx is formed by four or more whorls of three, membranous sepals that are cream coloured and, hence, described as petaloid sepals (Barneby and Krukoff 1971: 7). Staminate flow- ers of both genera have 6 petals that are membranous and much shorter than the innermost sepals (Table 1). Pistillate flowers ofCurarea have only 3 petals and 3 carpels, whilst those of Chondrodendron have 6 petals and 6 carpels. The drupelets of both genera have narrow supportive structures, but in Curarea, these are elongate carpophores that bear atop sessile fruits, while in Chondrodendron the fruits are con- spicuously stipitate (Table 1). The distinct nature of Curarea is also supported in a family-wide phylogenetic analysis of chloroplast markers (Ortiz et al. 2016), where it is recovered as monophyletic and sister to Sciadotenia Miers, in a neotropical clade that also includes Chondrodendron (Fig. 1A; Ortiz et al. 2016). Within this neotropi- cal clade, Curarea shares with Sciadotenia sessile drupelets borne on narrow elongated carpophores bearing six carpels in Sciadotenia and only three in Curarea. The two whorls of sepals in the staminate flowers distinguish Curarea from Chondrodendron and Sciadotenia, both of which have 3–7 whorls of sepals. When first described, Curarea included four species: C. candicans, C. cuatrecasasii, C. tecunarum and C. toxicofera, with C. toxicofera designated as the type species of the genus (Barneby and Krukoff 1971). While C. cuatrecasasii and C. tecunarum were newly described, both C. candicans and C. toxicofera were transferred from Chondro- dendron. Another species was later added to the original four with the recognition of C. crassa, an endemic new species from the Brazilian Atlantic Forest (Barneby 1996). Although the etymology of the generic epithet was never provided, the name Curarea doubtlessly refers to curare, the South American arrow-poison for which species of Curarea provide one of the ingredients. In this first treatment of Curarea (Barneby and Krukoff 1971), which was also the most comprehensive one at that time, C. toxicofera was broadly interpreted and included entities formerly described as various taxa, rendering it a large complex with unclear patterns of morphological variation. However, subsequent works generally fol- lowed that of Barneby and Krukoff (1971) and used the five species they recognised in treatment of local, regional and national floras, e.g. Flora of Surinam (Jansen-Jacobs 1976); Flórula de Las Reservas Biológicas de Iquitos, Perú (Ortiz 1997); Flora of Ecua- dor (Ott