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Compositae, Senecioneae) A peer-reviewed open-access journal PhytoKeys 158: 1–106 (2020) Taxonomic revision of Xenophyllum 1 doi: 10.3897/phytokeys.158.50848 MONOGRAPH http://phytokeys.pensoft.net Launched to accelerate biodiversity research Taxonomic revision of the Andean genus Xenophyllum (Compositae, Senecioneae) Joel Calvo1, Andrés Moreira-Muñoz1 1 Instituto de Geografía, Facultad de Ciencias del Mar y Geografía, Pontificia Universidad Católica de Valpa- raíso, Avenida Brasil 2241, 2362807 Valparaíso, Chile Corresponding author: Joel Calvo ([email protected]) Academic editor: A.R. Magee | Received 5 February 2020 | Accepted 1 July 2020 | Published 3 September 2020 Citation: Calvo J, Moreira-Muñoz A (2020) Taxonomic revision of the Andean genus Xenophyllum (Compositae, Senecioneae). PhytoKeys 158: 1–106. https://doi.org/10.3897/phytokeys.158.50848 Abstract The Andean genusXenophyllum (Compositae, Senecioneae) is distributed along the high-Andes from northeastern Colombia to northern Chile and northwestern Argentina, mainly thriving in the paramo and puna ecoregions. It comprises suffruticose plants forming dense mats, hummocks, or clumps of erect stems. They are characterized by displaying involucral bracts fused at the base, supplementary bracts ab- sent, and mostly radiate capitula with white ray corollas, seldom yellow or pink (disciform in one species). Traditionally, Xenophyllum species were treated as members of the genus Werneria, a morphologically close genus that includes rosettiform or scapiform perennial herbs. As currently circumscribed, Xenophyl- lum mostly differs from Werneria in having elongate stems. Herein, the first modern and comprehensive revision of the genus recognizing twenty-two species and two subspecies is presented. Werneria decumbens is synonymized with X. weddellii, as well as X. fontii with X. humile and X. oscartovarii with X. dactylophyl- lum. Likewise, four varietal names and two sectional names are proposed as new synonyms. Seven names are lectotypified, the nameX. sotarense is epitypified,W. decumbens neotypified, and the supraspecific name W. sect. Integrifoliae Rockh. is typified. The combination X. crassum subsp. orientale comb. nov. is made. Descriptions and distribution maps are provided for all accepted species, in addition to an identifi- cation key. Ten species are illustrated, three of them for the first time. Keywords Andes, Asteraceae, Neotropics, nomenclature, taxonomy, typification, Werneria Copyright Joel Calvo, Andrés Moreira-Muñoz. 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. 2 Joel Calvo & Andrés Moreira-Muñoz / PhytoKeys 158: 1–106 (2020) Table of contents List of figures ............................................................................................................ 2 Introduction ............................................................................................................. 3 Material and methods............................................................................................... 4 Morphology ............................................................................................................. 5 Taxonomic treatment ............................................................................................... 9 Key to the species of Xenophyllum ........................................................................... 11 Key to the subspecies of Xenophyllum crassum ......................................................... 40 Unverified names.................................................................................................... 96 Acknowledgements ................................................................................................. 96 References .............................................................................................................. 96 Appendix I ............................................................................................................. 98 Appendix II .......................................................................................................... 100 List of figures 1. Number of Xenophyllum species per country ......................................................... 4 2. Main habit types in Xenophyllum .......................................................................... 6 3. Diagram of measurements used in descriptions .................................................... 8 4. Distribution map of Xenophyllum ....................................................................... 10 5. Illustration of Xenophyllum acerosum ................................................................... 14 6. Distribution map of Xenophyllum acerosum and X. humile .................................. 15 7. Illustration of Xenophyllum humile ...................................................................... 17 8. Illustration of Xenophyllum roseum ...................................................................... 31 9. Distribution map of Xenophyllum sotarense, X. roseum, and X. funkianum ........... 33 10. Illustration of Xenophyllum funkianum ............................................................. 35 11. Illustration of Xenophyllum rigidum .................................................................. 37 12. Distribution map of Xenophyllum rigidum, X. crassum subsp. crassum, and X. crassum subsp. orientale....................................................................................... 38 13. Illustration of Xenophyllum crassum subsp. crassum ............................................ 41 14. Illustration of Xenophyllum marcidum ............................................................... 48 15. Distribution map of Xenophyllum marcidum and X. ciliolatum .......................... 51 16. Illustration of Xenophyllum juniperinum ........................................................... 54 17. Distribution map of Xenophyllum juniperinum, X. weddellii, and X. incisum ..... 56 18. Illustration of Xenophyllum poposa .................................................................... 65 19. Leaves and capitulum of Xenophyllum lorochaqui, X. poposa, and X. incisum ...... 66 20. Distribution map of Xenophyllum poposa, X. lorochaqui, and X. rosenii .............. 67 21. Illustration of Xenophyllum lorochaqui .............................................................. 72 22. Illustration of Xenophyllum pseudodigitatum ...................................................... 79 23. Distribution map of Xenophyllum digitatum and X. pseudodigitatum ................. 80 Taxonomic revision of Xenophyllum 3 24. Distribution map of Xenophyllum decorum, X. amblydactylum, and X. dactylophyllum ................................................................................................ 82 25. Illustration of Xenophyllum dactylophyllum ........................................................ 86 26. Illustration of Xenophyllum staffordiae ............................................................... 93 27. Distribution map of Xenophyllum staffordiae and X. esquilachense ...................... 94 Introduction Xenophyllum V.A.Funk (Compositae, Senecioneae), is a genus of some 22 species dis- tributed along the highlands of the Andes from northeastern Colombia to northwestern Argentina and northern Chile. The genus extends in distribution from the Colombian department of Norte de Santander (paramo del Almorzadero, ca. 7°N) to La Rioja Prov- ince in northwestern Argentina (Sierra de Famatina, ca. 29°S). Peru and Bolivia harbor the highest species diversity (12 and 11 species, respectively), followed by Ecuador (7 spp.), Chile (6 spp.), Argentina (5 spp.), and Colombia (3 spp.). The largest number of endemic species is found in Ecuador, 4 of the 7 species (57%) in the country (Fig. 1). Species of Xenophyllum are mostly small suffruticose plants characterized by displaying involucral bracts fused at the base, supplementary bracts absent, radiate capitula (disci- form in one species), usually white ray corollas (yellow or pink in a few species), balus- terform filament collars, obtuse or auriculate anther bases, truncate style branches with a crown of sweeping trichomes or penicillate, and glabrous or white-villous achenes. Two main habits can be differentiated according to the type of growth form (1) species forming dense mats or hummocks; (2) species forming clumps of somewhat distantly spaced stems or even having a shrubby habit (Calvo and Funk 2020). Species of the genus were traditionally treated under the morphologically similar genus Werneria Kunth until the end of 20th century, when Funk (1997a) coined the genus Xenophyllum. This new genus was meant to segregate fromWerneria those species forming loose or tightly compressed hummocks or well-developed mats, and accordingly, the genus Werneria was re-circumscribed to accommodate the rosettiform species grow- ing solitary or in small clumps. However, the type of growth form is not unequivocal for segregating both genera because some species of Werneria can also form compressed mats (e.g., W. aretioides Wedd., W. weberbaueriana Rockh.). We did not find any diagnostic morphological synapomorphies to support either of the two genera, but the presence or absence of elongate stems allows placing most species in one genus or the other. On this basis, Werneria comprises rosettiform or scapiform perennial herbs while Xenophyllum
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