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Gayana. Botánica

versión impresa ISSN 0016-5301versión On-line ISSN 0717-6643

Gayana Bot. vol.80 no.2 Concepción dic. 2023

http://dx.doi.org/10.4067/S0717-66432023000200103 

Original Article

The genus Carex (Cyperaceae) in Chile: a general update of its knowledge, with an identification key

El género Carex (Cyperaceae) en Chile: una actualización general de su conocimiento, con una clave de identificación

Paulo Muñoz-Schüler1  * 

Pablo García-Moro2 

José Ignacio Márquez-Corro2  3 

Diego Penneckamp4 

María Sanz-Arnal2 

Santiago Martín-Bravo2 

Pedro Jiménez-Mejías2 

1Herbario CONC, Departamento de Botánica, Facultad de Ciencias Naturales y Oceanográficas, Universidad de Concepción, Casilla 160-C, Concepción, Chile.

2Área de Botánica, Departamento de Biología Molecular e Ingeniería Bioquímica, Universidad Pablo de Olavide. Carretera de Utrera km 1 sn, 41013 Sevilla, Spain.

3Jodrell Laboratory, Royal Botanic Gardens, Kew, TW9 3AB, Richmond, United Kingdom.

4Laboratorio de Biodiversidad y Ecología del Dosel, Facultad de Ciencias Forestales y Recursos Naturales, Universidad Austral de Chile, P.O. Box 567, Valdivia, Chile.

ABSTRACT

The knowledge of the genus Carex L. (Cyperaceae) in South America is scattered, without recent comprehensive treatments for any country. Within the continent, Chile is the country that harbors the most diversity, being the second in species richness and the first in endemic taxa. However, Chilean botanists must resort to several Argentinian floras for species identification, although these works leave uncovered many Chilean species. This has led many researchers and amateurs to neglect the amazing diversity of the genus in the country. In this work, we summarize the results of several years of research on the genus Carex in Chile. We here formally report six new national records, including the reinstatement of the Nahuelbuta endemism Carex reicheana Boeckeler. Also, relevant changes (regional additions or significant distributional changes) are reported for another 33 species, and nomenclatural comments on problematic names are given. The catalogue of Carex for Chile is updated with our own findings and previous published reports from 82 to 96 species.

Keywords: Carex; Chile; identification key; taxonomy; Uncinia

RESUMEN

El conocimiento sobre el género Carex L. (Cyperaceae) en América del Sur está muy fragmentado y no existen tratamientos comprehensivos para ningún país. En el continente, Chile es el país que contiene la mayor diversidad de especies, siendo el segundo en riqueza y el primero en endemismos; paradojalmente, los botánicos de Chile deben recurrir a floras argentinas para identificar las especies, a pesar de que en ellas obviamente no figuran los endemismos de Chile. Esto se ha traducido en un desconocimiento generalizado sobre la fascinante diversidad del género en Chile. En este trabajo, resumimos los resultados de varios años de investigación sobre el género y reportamos formalmente seis nuevas especies para Chile, incluyendo la restauración del endemismo de Nahuelbuta Carex reicheana Boeckeler. Además, se reportan modificaciones distribucionales para 33 especies, junto a comentarios nomenclaturales sobre algunos nombres problemáticos. En este trabajo, se actualiza el catálogo de taxones de Carex en Chile mediante nuestros propios hallazgos y reportes publicados anteriormente, lo que lleva el número de especies conocidas para el país de 82 a 96.

Palabras clave: Carex; Chile; clave de identificación; taxonomía; Uncinia

INTRODUCTION

The genus Carex L., with around 2000 species, is one of the most species rich genera of vascular plants in the world (Roalson et al. 2021; POWO 2023). Morphologically is well characterized, with all of its species having unisexual flowers concealed by a sheathing bract-derived structure that in most cases fully encloses the female flower, forming a unique diaspora called utricle (Jiménez-Mejías et al. 2016a). The genus is greatly diversified in the temperate regions of the Northern Hemisphere, with multiple independent colonization events of the Southern Hemisphere landmasses and islands (Martín-Bravo et al. 2019). Because of both the lesser extent of continental surface in the southern hemisphere and the evolutionary history of the genus, its diversity is also reduced here. However, among the many lineages that have successfully colonized and established in temperate regions of South America and/or New Zealand, several exhibit unique cases of in situ radiations, resulting in a high number of endemic elements reported for these areas (Benítez-Benítez et al. 2021; García-Moro et al. 2022, Martín-Bravo et al. 2022). While in New Zealand and Australia the diversity of Carex has been relatively well studied from both a taxonomical and evolutionary approach (e.g., Hamlin 1959; Healy & Edgar 1980; Wilson 1993, 1994; Moar & Wilmshurst 2003; Vollan et al. 2006; Ford 2007), the diversity of South America has been neglected, with the knowledge of endemic species scattered and mostly lacking comprehensive systematic treatments.

In South America, Chile is one of the countries that harbors the largest Carex diversity, both in terms of species number and endemism (96 and 31, respectively, including results in this paper; also see Jiménez-Mejías et al. 2016b and updates), making into the third most speciose genus of the country after Senecio L. (ca. 235 spp. Compositae; Rodríguez et al. 2018; Calvo & Moreira-Muñoz 2020; Calvo & Saldivia 2022) and Adesmia DC. (134 spp. Leguminosae; Rodríguez et al. 2018). In Chile, Carex species can be dominant in certain environments (e.g., Hauenstein et al. 2002, 2009; Amigo & Castro 2015) or act as primary companions in other vegetation types (e.g., Amigo & Castro 2015). However, despite its diversity and abundance, the knowledge of the genus in Chile is quite fragmentary, as it is in the vast majority of South America. There has been only one major revision for the genus in Chile (Leveillé 1915), which recognized 44 species and provided a general key for the country. However, this work has been mostly neglected because it is written in French and is hard to find. So usually, for sedge identification, Chilean botanists and amateurs must resort to Flora Patagonica (Barros 1969) or other Argentinian floras (e.g., Genera et Species Plantarum Argentinarum (Barros 1943); Flora de la Provincia de Buenos Aires (Pedersen 1969); Flora de San Juan (Wheeler 2009)). However, these works are not directly related to Chile or leave uncovered a large area of it, thus many Chilean species may be not present in these. The problem is exacerbated considering the high endemism rate of Carex in the Pacific slopes of the Southern Cone. In addition, there is a number of recent taxonomic discoveries and taxonomic rearrangements (e.g., GCG 2015; Jiménez-Mejías et al. 2021a, 2021b; Ridley & Jiménez-Mejías 2022) that affect the treatment of the genus in Chile but remain beyond the reach of the unspecialized botanist or the amateur.

After years of taxonomic work including herbarium revision, literature search, and fieldwork, we present in this paper an update of the knowledge of the genus Carex in Chile -including the first exhaustive identification key in over 100 years- using as starting point the main floristic checklist of Chile (Catálogo de las plantas vasculares de Chile; Rodríguez et al. 2018).

MATERIALS AND METHODS

A 29-day fieldwork campaign from Coquimbo to Los Lagos (Chiloé) administrative regions was performed along the Coastal and Andes cordilleras, visiting areas where it was previously reported Carex species richness or may potentially harbor interesting records (see below). In addition, the full Carex collection of CONC, ULS, VALD and most of SI and LIL, were revised in situ, as well as type material housed in BAB and the online repository Global Plants JStor.

New or relevant chorological records were identified with respect to the information reported in the Catálogo de las plantas vasculares de Chile (Rodríguez et al. 2018) and its online updates. Distributions reported for South America and/or the rest of the World are based either in Flora del Cono Sur (Zuloaga et al. 2019), or Plants of the World Online (POWO 2023) checklist. To avoid redundancy and ease the reading, we will avoid continuously citing these works unless necessary. If additional distributional data from alternative sources are used, these are listed under the corresponding species epigraph.

Relevant specialized literature was consulted for systematics, taxonomic and chorological data. Each of these publications is cited under the corresponding species epigraph too.

For the presentation of comments in critical taxa, we follow the systematic arrangement provided by Roalson et al. (2021), with species grouped in formal sections or informal clades as suggested by these authors.

An identification key is provided for the identification of the 96 Carex species of the flora of Chile. For the preparation of this key, the last author of this paper (PJ-M) studied in situ the full collections of South American Carex of MA, K, MICH, MO, NY, and US (acronyms according to Thiers 2020). This was complemented with primary observations by the first author (PM-S) on the materials housed at CONC, SI, and LIL, as well as from DP on the collection of VALD. All the other authors of this paper later provided comments. We considered as additional reference works Léveillé (1915), Barros (1947, 1969), Wheeler (1986, 1987, 1988a, 1989, 1990, 1994, 2007a, 2009), Wheeler & Guaglianone (2003) and Jiménez-Mejías et al. (2021a, 2021b).

FIGURE 1 Carex species of Chile. A: C. negeri (Nahuelbuta range, Biobío Region); B: C. leporina (Puyehue National Park, Los Lagos Region); C: C. microglochin (Cordillera Pelada, Los Ríos Region); D: C. firmicaulis (Aguas Calientes valley, Ñuble Region); E: C. banksii (Nahuelbuta range, Araucanía Region); F: C. boelckeiana (Pino Hachado, Araucanía Region); G: C. antucensis (Aguas Calientes valley, Ñuble Region); H: C. wheeleri (Las Raíces range, Araucanía Region); I: C. pendula (Valdivia, Los Ríos Region); J: C. turbaria (Nahuelbuta range, Araucanía Region); K: C. hookeri (Chiloé, Los Lagos Region); L: C. patagonica (Las Trancas valley, Ñuble Region). Photos by: Paulo Muñoz-Schüler (A, D, E, I, J, K, L), José Ignacio Márquez-Corro (B, C, G, H), and Pedro Jiménez-Mejías (F). 

RESULTS

Rodríguez et al. (2018) reported 71 species of Carex as it did not include the names for the 21 reported species of the former genus Uncinia, which would have increased the number to 92. After a thorough revision of the names listed in Rodríguez et al. (2018) a final number of 62 accepted names were pin down from this checklist, which increased to 82 when including the 20 valid names for sect. Uncinia. In addition, six species were newly reported for Chile in this paper: Carex boelckeiana Barros, Carex dusenii Kük, Carex ecuadorica Kük, Carex leporina L., Carex pendula Huds., and the reinstatement of Carex reicheana Boeckeler. This, together with the published five reports of Carex skottsbergiana Kük (Kükenthal 1910), Carex nelmesiana Barros (Wheeler 1986), Carex patagonica Speg. (Zuloaga et al. 2019), Carex divisa Huds. (Calvo & Jiménez-Mejías 2020), and Carex phylloscirpoides Saldivia, S.Gebauer, Martín-Bravo & Jim.Mejías (Jiménez-Mejías et al. 2021b), increases the species richness in Chile to 93. Additionally, three unpublished new species are added both in the general count and in the key (Morales-Alonso et al. under review; Muñoz-Schüler et al. under review). Comments on the updates of Table 2 are presented here below whenever it is a new report, if not, it is referenced in the table and not discussed further. When the name we accept here is different from the one used in Rodríguez et al. (2018), we listed it as a synonym. A summarized list of the taxa here accepted and their regional distribution in Chile is given in Supplementary Material.

Carex subg. Psyllophorae (Degl.) Peterm.

A.1. Carex sect. Junciformes (Boeckeler) Kük.

Carex argentina Barros, Darwiniana 8: 409. 1948.

A widely distributed species of steppe environments. It has been reported from Mendoza to Santa Cruz provinces in Argentina, marking the northern and southern limits of this species distribution. However, in Chile it has been barely collected and reported only for two administrative regions (Metropolitana and Maule; García 2007, 2010), probably being overlooked because of its small size and inflorescence consisting of an acaulescent spike burrowed among leaf bases. Here we report new collections from Valle Nevado, in the Metropolitana Region and Pino Hachado, in the Araucanía Region. This last record establishes a new southern limit for this species in Chile. Its presence on the Patagonian steppe of Aysén and Magallanes administrative regions might be confirmed in future prospects. The plants of both localities were growing on dry sandy and rocky soils.

SPECIMENS: CHILE. Región Metropolitana de Santiago: Prov. Santiago, Farellones, estación de esquí Valle Nevado, 3.210 m, 16-I-2023, 1PJM-CL23 (CONC 193498). Región de la Araucanía: Prov. de Cautín, Paso Pino Hachado, laderas inmediatamente al sur del paso internacional, 1.900 m, 21-I-2023, 21PJM-CL23 (CONC 193499).

Carex boelckeianaBarros, Colecc. Ci. Inst. Nac. Tecnol. Agropecu 8(2): 70 (1969)

A very rare species so far known for very few locations in the Argentinian Andes of Neuquén and Mendoza provinces. Here we report its presence in Chile, which constitutes its first record for the country. This rarely collected species was found growing on dry stony slopes, intermingled within grass tussocks. While chorologically relevant, this finding is not surprising, since the plant was reported from the highest altitudes of the Argentinian side of Pino Hachado pass.

SPECIMEN: CHILE. Región de la Araucanía: Prov. Malleco, paso del Pino Hachado, laderas inmediatamente al sur del paso internacional, comunidades de gramíneas sobre suelo arenoso-pedregoso, 38°39’49.5”S 70°53’57.7”W, 1.900 m, 21-I-2023, P. Jiménez Mejías, P. García Moro & J.I. Márquez Corro, 18PJM-CL23 (CONC 193500).

Carex camptoglochin V.I.Krecz., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 7: 34. 1937.

A phylogenetically isolated species (Benítez-Benítez et al. 2021) sometimes subsumed as a variety or subspecies of the superficially similar C. microglochin Wahlenb. (Barros 1969; Wheeler & Guaglianone 2003), despite the fact that they actually belong to different Carex subgenera (Jiménez-Mejías et al. 2016c). The species is a South American endemism with a disjunct distribution in Patagonia and the Northern Andes, being absent from Central and Northern Chile and Argentina as well as from Bolivia. Here we report a new regional record and the northernmost limit of the Patagonian distribution of the species in Los Ríos Region. The species grows in boggy soils, usually associated with Sphagnum.

SPECIMEN: CHILE. Región de Los Ríos: Prov. Ranco, Cordillera Pelada. Parque Nacional Alerce Costero, turbera con Nothofagus antarctica en claro de bosque de Fitzroya cupressoides y Nothofagus sp., 40°09’56.9”S 73°30’18.1”W, 954 m, 1-II-2023, J.I. Márquez Corro, P. Muñoz Schüler, P. Jiménez Mejías, S. Martín Bravo, M. Sanz Arnal & P. García Moro, 71JMC23 (CONC 193502).

Carex chlorolepis Steud., Syn. Pl. Glumac. 2: 204. 1855.

Carex phalaroides auct. non Kunth, Enum. Pl. 2: 482. 1837.

The group of Carex phalaroides is a small and poorly understood taxonomic complex of species. Here we use the name C. chlorolepis as it is the prior one applying to the Chilean populations of the complex. The group of Carex phalaroides is distributed in temperate Chile (incl. Juan Fernández archipelago; Wheeler 2007), temperate Atlantic Argentina, Uruguay, and Brazil, as well as the Andes from Argentina to Venezuela. It reaches marginally Central America in Guatemala (Jiménez-Mejías et al. 2018). From Chile it has been reported discontinuously from Valparaíso to Los Lagos administrative regions. Remarkably, the group is known in Andean Argentina from La Rioja to Jujuy, just north of the Chilean populations, at the other side of the Arid Diagonal. The Chilean populations should be considered as an independent species, C. chlorolepis, as revealed by unpublished phylogenies that show them as an isolated lineage (see work in progress in Morales-Alonso et al. 2022). Carex chlorolepis is a mesophytic species that inhabits partly shadow understory in sclerophyllous forest and shrublands in the Mediterranean regions of Chile, as well as in exposed, open areas of the evergreen rainforests of southern Chile, and may even appear in disturbed (urban) habitats (pers. obs.).

We provide here new reports for O’ Higgins, Maule and Los Ríos administrative regions. Still unknown from the Metropolitana and Ñuble Regions, the presence of C. chlorolepis there should be confirmed in future prospections. Carex chlorolepis is a rather inconspicuous, slender and grass-like plant that may have been overlooked by collectors and could probably be present in these areas.

SPECIMENS: CHILE. Región de O’ Higgins: Prov. Cachapoal, Coltauco, Santuario de la Naturaleza Cerro Poqui, 34°12’32.148”S 71°3’31.752”W, 440 m, 23-IV-2019, N. García 5576 (EIF 15684); Prov. Colchagua, Nancagua, fundo Santa Elena, camino a cumbre del cerro Rucatalca, 34°36’20.772”S 71°11’44.376”W, 650 m, 21-IX-2019, N. García, D. Tapia, B. Navarrete & D. Van Sint Jan 5668 (EIF 15774). Región del Maule: Prov. Cauquenes, Cauquenes, Tiquel, cerros camino a Quirihue. En talud a orilla de camino vehicular que recorre pendiente de exposición S, 36º8’27.5’’S 72º23’17.1’’W, 426 m, 28-XI-2022, P. Muñoz Schüler & M. Mancilla Baeza, PMS-187 (CONC 193522). Región de los Ríos: Prov. Valdivia, Corral, 39º57’20.8’’S 73º22’29.8’’W, 24 m, 23-I-2023, J.I. Márquez Corro, P. Jiménez Mejías & M. Sanz Arnal, 26JMC23 (CONC 193523).

Carex nelmesiana Barros, Bol. Soc. Argent. Bot. 6(3-4): 209. 1957.

An endemic species from Patagonia, so far reported as endemic from Argentina.

In the revision of Patagonian species of sect. Junciformes, Wheeler (1986) gave an extensive discussion about the similarities between C. nelmesiana and three other species of the section and cited the material “Rio Nirehuao, 22 Nov 1908, Skottsberg (UPS!)” as coming from Argentina. However, this is a confusion, and this material comes from a locality within the limits of Chile. This material was collected by Skottsberg just a few years after the governments of Chile and Argentina signed an agreement in 1902, where they agreed to modify the limits of both countries between 40° and 52°S, along central Patagonia. This agreement directly affected the zone where the Ñirehuao river is located, passing from Argentine to Chilean sovereignty, and probably unnoticed on the official maps for some time.

SPECIMEN: CHILE. Región de Aysén: Prov. Aysén, río Ñirehuao, 22-XI-1908, Skottsberg (UPS!).

Carex patagonica Speg., Revista Fac. Agron. Univ. Nac. La Plata 32-33: 626. 1897.

A species primarily known from Argentinian Patagonia and reported marginally from Chile (Zuloaga et al. 2019; POWO 2023), not recorded in Rodríguez et al. (2018) or Zuloaga et al. (2019). Here we provide specific locations for Metropolitana, O’Higgins, Ñuble, and Biobío Regions. One specimen (N. Lavandero & L. Santilli, 958) was collected on El Roble Natural Sanctuary, just a few meters far from the Valparaiso regional limit, and its presence for this Region is highly probable but cannot be confirmed in this work. This species is an inhabitant of the understory of montane Nothofagus forests.

SPECIMENS: CHILE. Región Metropolitana de Santiago: Prov. Chacabuco, Tiltil, SN cerro El Roble, en el límite regional con Valparaíso, Ladera exposición S, 32º58’45.9’’S 71º1’5’’W, 2068 m, 28-X-2020, N. Lavandero & L. Santilli, NL-958 (CONC 193521). Región de O´Higgins: Prov. Colchagua, sierra s de Bellavista, Alto Huemul, 34º50’47.8794’’S 70º40’51.8596’’W, 1366 m, I-2006, García & al. 3289 (CONC 180616). Región del Ñuble: Prov. Diguillín, valle de Las Trancas, camino a termas de Chillán, cerca del estero Renegado, 36º55’0.1’’S 71º27’11.8’’W, 1307 m, 18-I-2023, S. Martín Bravo, P. García Moro, P. Jiménez Mejías, J.I. Márquez Corro, P. Muñoz Schüler & M. Sanz Arnal, 22SMB23 (CONC 193520). Región del Biobío: Prov. Biobío, Antuco, valle del Polcura, 37º12’56.5’’S 71º26’24.1’’W, 1000 m, XI-2016, Macaya & al. 394 (CONC 185922).

Carex pungens Boeckeler, Linnaea 39: 46. 1875.

Carex berteroana É.Desv. in C.Gay, Fl. Chil. 6: 198. 1854[1853]. nom. illeg., non C. berteroniana Steud., Flora 25: 604. 1842.

Carex setifolia Kunze var. berteroana (É.Desv.) Gunckel, Rev. Chil.Hist. Nat. 11: 145. 1954.

The name Carex berteroana É.Desv. was published in 1854 and it is to be considered a later homonym of C. berteroniana Steud. (1842) (see ICN: Arts. 53.2, 61.1 and 61.2). Both parahomonymous names honour the same person, are very similar orthographically and both apply to species of Carex (N. Turland, pers. comm.). The earliest legitimate name at species rank is C. pungens Boeckeler. Despite often being treated as a variety of C. setifolia Kunze (C. setifolia var. pungens (Boeckeler) Kük.), the available phylogenetic data seems to support its species status (Benítez-Benítez et al. 2021), and thus we consider it as such.

Carex reicheana Boeckeler, Allg. Bot. Z. Syst. 2: 173. 1896.

TYPE: Chili [CHILE], in locis altiss. Cordillerae de Nahuelbota [Nahuelbuta] invenit Karl Reiche (B, destroyed). NEOTYPE designated here: CHILE. Región de la Araucanía: Prov. Malleco, Parque Nacional Nahuelbuta, cerca del límite regional con la Región del Biobío, en suelos secos y sombríos en sotobosque de Araucaria araucana-Nothofagus pumilio, 37°18’07.31’’S 73°01’02.60’’W, 1264 m, 21-1-2023, S. Martín-Bravo, P. Muñoz-Schüler & M. Sanz-Arnal, 38SMB23, CONC 193526; isoneotypes EIF 16579; UPOS sn).

An interesting species overlooked to date as an independent taxon. Carex reicheana was synonymized by Kükenthal (1909) to C. trichodes Phil. (=C. lateriflora Boot; see Penneckamp 2022). Since the type, housed at B, was destroyed during WWII, it was subsequently listed as a synonym by Wheeler (1989), who just followed Kükenthal’s opinion (“I have not seen a duplicate of Reiche’s collection, but Boeckeler’s description of C. reicheana fits very closely the features of C. lateriflora”). Carex reicheana was described from what is today the Nahuelbuta National Park, in the Coastal Range, isolated and far from the other reported populations of the closely related C. trichodes by at least, 150 km., all in the Andes range (Wheeler 1989; Penneckamp 2022). During our fieldtrip in January-February 2023, we collected specimens similar to C. trichodes in Nahuelbuta National Park. The plants matched the description of C. reicheana protologue. Moreover, these displayed clear differences from Andean specimens of C. trichodes. In Table 1 we provide a detailed comparison of features between C. reicheana and C. trichodes according to the descriptions provided in Wheeler (1989), Penneckamp (2022) and our own observations. The differences found between the two species (Fig. 2) are notable regarding the discriminant characters employed for species distinction in sect. Junciformes (= sect. Aciculares G.A.Wheeler; Wheeler 1989) and clearly justify the reinstatement of the name C. reicheana and its treatment as an independent species.

Since the type material of C. reicheana is lost, we here proceed to neotypify the name on our own collection.

Carex setifolia Kunze ex Kunth

Uncinia setifolia Kunze ex Rodríguez et al., Cat. Pl. Vasc. Chile: 59 (2018), nom. inval.

The name Uncinia setifolia Kunze is listed in the Catálogo de las plantas vasculares de Chile (Rodríguez et al. 2018), but it seems to be a mistranscription of Carex setifolia Kunze ex Kunth. We did not find this name in any of the major plant nomenclatural databases (Zuloaga et al. 2019; POWO 2023; IPNI 2023), nor it appears in C. setifolia protologue. Accordingly, this name should be removed.

TABLE 1 Character comparison of C. trichodes and C. reicheana, two morphologically close species of sect. Junciformes. / Comparación de caracteres entre C. trichodes y C. reicheana, dos especies morfológicamente próximas de la sect. Junciformes

Carex trichodes Carex reicheana
Plant habit Culms erect at maturity, bearing the inflorescence well-above the ground. Culms erect to strongly curved at maturity, often decumbent, bearing the inflorescence at ground level.
Number of utricles per spike (3)4-6(8) (1)2-3
Utricle coloration Reddish-brown, stramineous or yellowish Blood-like red to reddish-brown
Utricle shape Flattened, plano-convex or slightly biconvex Inflated, strongly biconvex
Distribution Nothofagus pumilio forests of southern Chile’s Andean range Araucaria araucana-Nothofagus pumilio forests in the Nahuelbuta range

FIGURE 2 Carex reicheana. A. Growing habit; B-D: Detail of the spike and utricles; E: Habitat, the material studied was collected in the shady areas within the picture. Photos by: Paulo Muñoz-Schüler (A, C, E) and Santiago Martín-Bravo (B, D). / Carex reicheana. A. Hábito de crecimiento; B-D: Detalle de la espiga y los utrículos; E: Hábitat, el material estudiado se recolectó en las zonas sombreadas dentro de la foto. Fotos de: Paulo Muñoz-Schüler (A, C, E) y Santiago Martín-Bravo (B, D). 

Table 2 Additions, name changes, and distributional changes updated from the Catálogo de las plantas vasculares de Chile (Rodríguez et al. 2018). The abbreviations used for administrative regions are according to Rodríguez et al. (2018). / Adiciones, cambios de nombre, y cambios distribucionales actualizados a partir del Catálogo de las plantas vasculares de Chile (Rodríguez et al. 2018). Las abreviaciones para las regiones administrativas son las utilizadas en Rodríguez et al. (2018) 

Taxon name Update regarding Rodríguez et al. (2018) References
A. Subg. Psyllophorae
A.1. Sect. Junciformes
Carex argentina Barros Addition to ARA This paper
Carex austroamericana G.A.Wheeler Addition to RME This paper
Carex berteroana É.Desv. Illegitimate name (later homonym). The earliest legitimate name to be considered is Carex pungens Boeckeler -
Carex boelckeiana Barros Addition to Chile (ARA) This paper
Carex camptoglochin V.I.Krecz. Addition to LRI This paper
Carex lateriflora Phil. To be synonymized to C. trichodes Boot and addition to LRI Penneckamp 2022
Carex nelmesiana Barros Addition to Chile (AIS) Wheeler 1986; this paper
Carex patagonica Speg. Addition to Chile (RME, LBO, NUB, BIO) POWO 2023; Zuloaga et al. 2019; this paper
Carex phalaroides Kunth To be treated as Carex chlorolepis Steud., an endemic species from Chile. Addition to LBO, MAU and LRI This paper
Carex reicheana Boeckeler Upgrade as an independent species from C. lateriflora and added as an endemism from ARA This paper
Carex toroensis G.A.Wheeler Addition to AIS Jiménez-Mejías et al. 2016b
Uncinia setifolia Kunze Invalid name This paper
B. Subg. Vignea
B. 1. Sect. Divisae
Carex divisa Huds. Addition to Chile (VAL), and new regional report (RME) Calvo & Jiménez-Mejías 2020; this paper
B.2. Annectens Clade
Carex firmicaulis Kalela Addition to BIO Jiménez-Mejías et al. 2016b
Carex gayana É.Desv. Intraspecific treatment to be withdraw -
Carex nebularum Phil. To be considered endemic from Chile. Jiménez-Mejías et al. 2016b;
Addition to BIO this paper
Carex ecuadorica Kük. Addition to Chile (ANT) as native species. This paper
Carex pycnostachya É.Desv To be synonymized to Carex pleioneura G.A.Wheeler Jiménez-Mejías et al. 2021a
B.3. Sect. Cyperoideae
Carex leporina L. Addition to Chile (BIO, ARA, LRI, LLA) This paper
B.4. Sect. Glareosae
Carex canescens L. To be considered native in Chile and Argentina. This paper
Carex skottsbergiana Kük Addition to Chile (AIS) as an endemism. Kükenthal 1910; POWO 2023
B.5. Sect. Bracteosae
Taxon name Update regarding Rodríguez et al. (2018) References
Carex bracteosa Kunze ex Kunth Addition to MAU This paper
Carex trachycystis Griseb. To be synonymized to Carex bonariensis Desf. ex Poir. Jiménez-Mejías et al. 2021a
B.6. Disticha clade
Carex maritima auct. non Gunnerus To be synonymized to Carex melanocystis É.Desv. Addition to RME Jiménez-Mejías et al. 2021a
B.7. Vulpinae-Multiflorae alliance
Carex brongniartii Kunth Addition to LRI Ramírez et al. 1980; Teillier et al. 2013
C. Subg. Uncinia
C.1. Sect. Uncinia
Carex aspericaulis (G.A.Wheeler) J.R.Starr New combination from Uncinia aspericaulis G.A.Wheeler Global Carex Group 2015
Carex delacosta Kuntze New combination from Uncinia macloviana Gaudich. Distribution on JFE must be withdrawn Global Carex Group 2015
Carex dolichophylla J.R.Starr New combination from Uncinia macrophylla Steud. Global Carex Group 2015
Carex erinacea Cav. New combination from Uncinia erinacea (Cav.) Pers. Global Carex Group 2015
Carex fernandesiana (Nees ex Boeckler) J.R.Starr New combination from Uncinia douglasii Boott Global Carex Group 2015
Carex firmula (Kük) J.R.Starr New combination from Uncinia tenuis Poepp. ex Kunth Global Carex Group 2015
Carex firmula subsp. tenuis (Poepp. ex Kunth) Jim.Mejías & R.Ridl. New combination. Addition to Chile (JFE) as an endemism Ridley & Jiménez-Mejías 2022
Carex lechleriana (Steud.) J.R.Starr New combination from Uncinia lechleriana Steud. Global Carex Group 2015
Carex macloviformis (G.A.Wheeler) J.R.Starr New combination from Uncinia macloviformis G.A.Wheeler Global Carex Group 2015
Carex macrotrichoides J.R.Starr New combination from Uncinia chilensis G.A.Wheeler Global Carex Group 2015
Carex meridensis (Steyerm.) J.R.Starr New combination from Uncinia macrolepis Decne. Global Carex Group 2015
Carex multifaria (Nees ex Boott) J.R.Starr New combination from Uncinia multifaria Nees ex Boott. Distribution on JFE must be withdrawn Global Carex Group 2015
Carex negeri (Kük.) J.R.Starr New combination from Uncinia negeri Kük.; addition to BIO Global Carex Group 2015; this paper
Carex phleoides Cav. New combination from Uncinia phleoides (Cav.) Pers. Global Carex Group 2015
Carex plurinervata J.R.Starr New combination from Uncinia costata Kük. Global Carex Group 2015
Carex quinquin Jim.Mejías & Dorr New combination from Uncinia trichocarpa C.A.Mey. Jiménez-Mejías & Dorr 2018
Carex salticola J.R.Starr New combination from Uncinia andina G.A.Wheeler Global Carex Group 2015
Carex scabrida J.R.Starr New combination from Uncinia scabriuscula G.A.Wheeler. Distribution on JFE must be withdrawn Global Carex Group 2015; Wheeler 2007; this paper
Carex triangula J.R.Starr New combination from Uncinia triquetra Kük. Global Carex Group 2015
Carex turbaria J.R.Starr New combination from Uncinia austroamericana G.A.Wheeler. Addition to ARA, and LLA Global Carex Group 2015; this paper
Carex wheeleri J.R.Starr New combination from Uncinia araucana G.A.Wheeler Global Carex Group 2015
Taxon name Update regarding Rodríguez et al. (2018) References
D. Subg. Carex
D.1. Hirta Clade
Carex acutata Boott Misused name for Carex multispicata Kunze Jiménez-Mejías et al. 2020
Carex aematorhyncha É.Desv. Intraspecific treatment to be withdrawn This paper
Carex excelsa Poepp. ex Kunth Addition to ATA and LRI Ramírez et al. 1980; Teillier et al. 1980; this paper
Carex pumila Thunb. Addition to NUB This paper
Carex werdermannii Gross Addition to LRI and reinstate its presence in LLA This paper
D.2. Flacca clade
Carex banksii Boott Intraspecific treatment to be withdrawn This paper
Carex dusenii Kük. Addition to Chile (LLA, MAG) Barros 1969; this paper
Carex ochrostachya Phil. To be synonymized under Carex banksii Boott This paper
D.4. Sect. Spirostachyae
Carex fuscula d’Urv Intraspecific treatment to be withdrawn This paper
Carex araucana Phil. To be subsumed under Carex fuscula d’Urv Escudero et al. (2009); this paper
Carex distenta Kunze ex Kunth To be subsumed under Carex fuscula d’Urv Escudero et al. (2009); this paper
Carex inconspicua Steud. To be subsumed under Carex fuscula d’Urv Escudero et al. (2009); this paper
Carex lamprocarpa Phil. Addition to LRI and distribution in BIO and LLA must be withdrawn This paper
Carex trifida Cav. Addition to LLA This paper
D.5. Sect. Rhynchocystis
Carex pendula Huds. To be considered as a naturalized species This paper
D.6. Sect. Phacocystis
Carex chillanensis Phil. To be synonymized to Carex antucensis Kunth This paper
D.7 Sect. Racemosae
Carex phylloscirpoides Saldivia, S.Gebauer, Martín-Bravo & Jim.-Mejías Addition to Chile (ANT) as an endemism Jiménez-Mejías et al. 2021b
E. Subg. Euthyceras
E.1. Capitata Clade
Carex arctogena Harry Sm. To be used as the correct name for Southern Hemisphere populations ascribed to Carex capitata L. Villaverde et al. 2015

B. Carex subg. Vignea (P. Beauv. ex T. Lestib.) Heer

B.1 Carex sect. Divisae H.Christ ex Kük.

Carex divisa Huds., Fl. Angl. 348. 1762.

A Western Palearctic native recently added to South America (Jiménez-Mejías et al. 2018, 2021a) and Chile (Calvo & Jiménez-Mejías 2020) from the coasts of Valparaíso Region. Here we provide a new report from the Metropolitan Region of Santiago in the Central Valley for this introduced species.

SPECIMEN: CHILE. Región Metropolitana de Santiago: Prov. Chacabuco, Batuco, 33°16’S 70°47’W, 475 m, IX-1994, Bliss, 554 (CONC 145201).

B.2 Annectens Clade

Carex gayana E. Desv., Fl. Chil. 6: 205, t. 73, f. 3. 1853[1854] A species widely distributed in South America (from Perú to Tierra del Fuego), with a highly variable morphology (Jiménez-Mejías et al. 2021a). We advise not to consider infraspecific variants until biosystematic data are available. Published studies on the genetic variation of the species (Pfeiffer et al. 2018) do not seem to clearly match morphologic variation, but geographic structuring, which is in line with our proposal.

Carex nebularum Phil., Anales Univ. Chile 93: 492. 1896.

An endemism from montane forest and Andean streams from the central regions of Chile. Here we report it for Biobío Region. Records from Argentina are actually confusions with other species of the group such as Carex firmicaulis Kalela or Carex hypoleucos É.Desv. (Jiménez-Mejías et al. 2016b, 2021a). Identically, reports from Aysén need to be confirmed.

SPECIMEN: CHILE. Región del Biobío: Prov. Biobío, Reserva Nacional Ralco, Pelado Mallín, 37º53’S 71º23’W, 1830 m, I-2022, M. Mihoc et al., 4240 (CONC 158895).

Carex ecuadorica Kük., Beibl. Bot. Jahrb. Syst. 78: 7. 1904. Carex macrorrhiza auct., non Boeckeler

A Tropical Andean species, known from NW Argentina to Ecuador (Jiménez-Mejías et al. 2021a). In the southern part of its distribution it has been mistakenly reported as C. macrorrhiza Boeckeler (Jiménez-Mejías et al. 2016b, 2021a). It inhabits wet meadows and pastures at high altitudes. The material we report here is an unequivocal individual from C. ecuadorica and constitutes the first record of the species for Chile. The species was collected on Andean bofedales (peatbogs) in the Antofagasta Region.

Collections of similar plants from the more northern Region of Tarapacá are morphologically adscribed to C. macrorrhiza (Jiménez-Mejías et al. 2021). These populations (as well as the entire group of C. ecuadorica, C. macrorrhiza and the Patagonian C. subfuegiana G.A.Wheeler) need further study to clarify the biosystematics scenario.

SPECIMEN: CHILE. Región de Antofagasta: Prov. El Loa, vegas de Turi, 22°15’S 68°19’W, 3025 m, 1980, Villagrán & Armesto, 2309 (CONC 68822).

B.3 Carex sect. Cyperoideae G.Don

Carex leporina L., Sp. Pl. 2: 973. 1753.

Carex macloviana auct. p.p., non d’Urv.

A Western Palearctic species whose introduction has been reported already in North America (Mastrogiuseppe et al. 2002) and New Zealand (Dawson 2014). This species inhabits wet and mesic meadows. Its presence in South America (Chile and Argentina) was already noted by us (but not explicitly reported) in the multiaccession phylogenetic tree in Martín-Bravo et al. (2019). Here we formally report the presence of this species in Chile, in Biobío, Araucanía, Los Ríos, and Los Lagos administrative regions. The species has been mistaken for C. macloviana and considered as a particular form of it (Carex macloviana var. pseudoleporina Kük).

Remarkably, the Argentinian and Chilean populations seem to grow in undisturbed habitats, not as C. leporina does in North America and New Zealand, where it behaves as invasive. Despite this is not a clear sign of their native status, other Carex species that occur naturally in South America have their closest counterparts in Europe (W Palearctic-Neotropic disjunction, e.g., C. riparia Curtis-C. chilensis Brong., C. punctata Gaudin-C. fuscula d’Urv). Conversely, there are at least two introduced species of W Palearctic Carex in South America (C. divisa and C. divulsa Stokes.; Jiménez-Mejías et al. 2021a) Additional studies are necessary to unravel the true status of these apparently introduced populations of C. leporina in South America.

SPECIMENS: CHILE. Región del Biobío: Prov. Biobío, valle del río Laja, camino entre Polcura y salto del Abanico, margen de pantano de Cyperus eragrostis y Juncus sp., sobre suelo seco, 37°18’13.4”S 71°39’20.8”W, 154 m, 20-I-2023, P. Muñoz Schüler, P. García Moro & S. Martín Bravo, PMS-244 (CONC 193508). Región de la Araucanía: Prov. Malleco, Curacautín, ruta 181, cunetas temporalmente inundadas, 38°27’40.2”S 71°42’23.0”W, 600 m, 21-I-2023, P. Jiménez Mejías, P. García-Moro & J.I. Márquez-Corro, 17PJM-CL23 (CONC 193509). Prov. Malleco, Lonquimay, salto Alaska, estero La Cascada, 38º30’º16.8’’S 71º24’45.2’’W, 1003 m, 22-I-2023, P. García Moro, J.I. Márquez Corro & P. Jiménez Mejías, 4pgm2023 (CONC 193514). Prov. Malleco, Lonquimay, Galletué, 38°41’33.45’’S 71°20’18.38’’W, 1156 m, 06-II-2022, D.

Penneckamp, 771 (UPOS). Región de Los Ríos: Prov. Valdivia, cerro entre Santa Elvira y Cayumapu, bosque de Nothofagus obliqua, a orilla de camino, 39°44’54.1”S 73°09’22.9”W, 118 m, 23-I-2023, P. Muñoz Schüler, P. García-Moro & S. Martín-Bravo, PMS-265 (CONC 193510). Prov. Valdivia, Corral, pradera-juncal en borde de carretera, 39°56’59.5”S 73°18’50.2”W, 8 m, 23-I-2023, J.I. Márquez Corro, P. Jiménez Mejías & M. Sanz Arnal, 24bisJMC23 (CONC 193511). Prov. Valdivia, Corral, sendero Darwin, Parque Nacional Alerce Costero, Catrileufu, en la orilla del sendero, 39°59’9.62”S 73°25’16.37”O, 630 m, 19-XII-2022, D. Penneckamp, 927 (UPOS). Región de los Lagos: Prov. Osorno, Parque Nacional Puyehue, 40º43’54.5’’S 72º18’40.5’’W, 513 m, 26-I-2023, J.I. Márquez Corro, P. Jiménez Mejías & M. Sanz Arnal, 48JMC23 (CONC 193513). Prov. Osorno, Parque Nacional Puyehue, 40º46’51.5’’S 72º12’39.6’’W, 993 m, 26-I-2023, J.I. Márquez Corro, P. Jiménez Mejías & M. Sanz Arnal, 59JMC23 (CONC 193512).

B.4 Carex sect. Glareosae G.Don

Carex canescens L., Sp. Pl.: 974. 1753.

A circumpolar species widely distributed in the temperate northern hemisphere but also reported for the Pacific Southwest (New Guinea and SE Australia) and Southern South America. It has been considered as an exotic species in most South American checklists (Rodríguez et al. 2018; Zuloaga et al. 2019). However, this species usually inhabits undisturbed and often remote places such as moorlands or mountain trails, pointing to a long-stand interaction with its environments. In addition, the available molecular data is compatible with the native status of the species in South America, with samples forming exclusive clades in phylogenies and displaying exclusive unique plastidial haplotypes (Villaverde et al. 2017a). Bipolar distributions in Carex are known in a few species and depicts the dispersal and colonization ability of this genus (Márquez-Corro et al. 2017; Villaverde et al. 2017b). We strongly suggest the consideration of C. canescens in Argentina and Chile as a native species.

Carex skottsbergiana Kük., Repert. Spec. Nov. Regni Veg.: 8: 7. 1910.

Very little is known about this species, apparently endemic from southern Chile and apparently distinct from the visually similar C. canescens. So far it has only been collected from its type locality at lago O’ Higgins, in the Aysén Region, as in the original Kükenthal cited it from “Chilenisches Patagonien: Lago San Martin” (Kükenthal, 1910). It is not reported for Chile neither in Rodríguez et al. (2018) nor in Zuloaga et al. (2019).

B.5 Carex sect. Bracteosae Pax

Carex bracteosa (Kunze ex Rchb.) Kunth, Enum. Pl. 2: 379. 1837.

Despite being one of the most common species in Chile, C. bracteosa has not been reported for Maule earlier. Carex bracteosa is an endemic species, fairly common in the central valley and the Coastal Cordillera, distributed continuously from Valparaiso to Los Ríos Regions and associated with temporal or permanently wet meadows, often near the mouth of rivers or in the sclerophyllous vegetation of central Chile (Wheeler & Muñoz-Schick 2007).

SPECIMENS: CHILE. Región del Maule: Prov. Curicó, Hualañé, El Cobre, fundo La Palma, 34°55’32.89”S 71°35’34.57”W, 130 m, 17-XII-2020, D. Penneckamp 467 (UPOS). Prov. Curicó, Sagrada Familia, cerros hacia el suroeste de Villa Prat, 35°7’39.55”S 71°38’50.47”W, 214 m, 22-X-2021, D. Penneckamp & E. Picero, 739 (CONC).

B.6 Disticha clade

Carex melanocystis É.Desv., Fl. Chil.: 6: 203, t. 73, f. 5. 1853[1854].

Carex maritima auct. non Gunnerus

A species usually subsumed under the similar C. maritima Gunn. but recently segregated due to the South American plants having closer phylogenetic relationships with other related species than to C. maritima s.s. (Jiménez-Mejías et al. 2021a). The species is an Andean endemic found from Ecuador (Jiménez-Mejías et al. 2023) to Tierra del Fuego. In the Southern Cone it is disjunctly distributed in the Northern and Central regions at high altitudes, and then in Magallanes Region, where it can be found at much lower elevations. It is usually found growing at the margins of high-altitude wetlands in its northern distribution in Chile (Teillier, pers. com.). We report here as new for Metropolitana Region of Santiago.

SPECIMEN: CHILE. Región Metropolitana de Santiago: Prov. Santiago, Farellones, estación de esquí Valle Nevado, Tres Puntas, 33º11’48.3’’S 70º8’56.8’’W, 3210 m, 16-I-2023, P. Jiménez Mejías, J.I. Márquez Corro, S. Martín Bravo & P. Muñoz Schüler, 3PJM-CL23 (CONC 193515).

C. Carex subg. Uncinia (Pers.) Peterm.

C.1. Carex sect. Uncinia (Pers.) Baill.

Formerly treated as an independent genus (Uncinia Pers.), the hooked sedges were transferred into Carex according to long-discussed molecular evidence that indicates the position of the former genus Uncinia as deeply nested within Carex (GCG 2015). Later molecular works reinforce its inclusion in Carex (Jiménez-Mejías et al. 2016c; Martín-Bravo et al. 2019; Villaverde et al. 2020). Recently, a clade-focused study (García-Moro et al. 2022) clarified the position of the species of the former genus Uncinia as sect. Uncinia and within the recently named subgenus Uncinia (Villaverde et al. 2020; Roalson et al. 2021), which included other sectional groups traditionally included in Carex. A more detailed discussion will be given in an upcoming revision of subgenus Uncinia in South America (Muñoz-Schüler et al. in prep). Accordingly, the change in the generic status of the hooked sedges caused all the names in use to be transferred to Carex. The list of the accepted names and their corresponding synonyms is given in Table 2.

Carex delacosta Kuntze, Revis. Gen. Pl. 3(3): 332. 1898. Uncinia macloviana Gaudich., Voy. Uranie: 412. 1829.

The name Uncinia macloviana has been widely used as a hotchpotch to refer to similar looking species outside its range. It has been cited as occurring in the Juan Fernández archipelago both in Rodríguez et al. (2018) and Zuloaga et al. (2019), referring to other endemic species in the islands. Its presence in the archipelago was already dismissed by Wheeler (2007).

Carex multifaria (Nees ex Boott) J.R.Starr, Bot. J. Linn. Soc. 179: 34. 2015.

Uncinia multifaria Nees ex Boott in J.D.Hooker, Fl. Antarct. 2: 369. 1846.

Mistakenly reported as occurring in the Juan Fernández archipelago both in Rodríguez et al. (2018) and Zuloaga et al. (2019), as already noted by Wheeler (2007). It was probably confused with the robust specimens of C. phleoides Cav. (= Uncinia phloides (Cav.) Pers.) growing in the archipelago (see Jiménez-Mejías & Dorr 2018).

Carex negeri (Kük.) J.R.Starr, Bot. J. Linn. Soc. 179: 34. 2015. Uncinia negeri Kük., Bot. Centralbl. 76: 210. 1898.

Endemic from the temperate forests of Southern-Central Chile, with marginal occurrences in adjacent Argentina (Wheeler 1997). It is one of the few species of sect. Uncinia that inhabits Araucaria araucana (Molina) K.Koch forests but is also very typical in Nothofagus pumilio (Poepp. & Endl.) Krasser forests (Amigo & Castro 2015). We report it as new for the Biobio Region, on the Nahuelbuta range, which becomes its northernmost known location. The species must be searched in Los Ríos Region, where it is very probably present.

SPECIMEN: CHILE. Región del Biobío: Prov. Arauco, Parque Nacional Nahuelbuta, estacionamiento del mirador Piedra del Águila, en el límite regional con la Araucanía, 37º49’21.4’’S 73º2’7.6’’W, 1411 m, 21-I-2023, S. Martín Bravo, P. Muñoz Schüler & M. Sanz Arnal, 41SMB23 (CONC 193518).

Carex scabrida J.R.Starr, Bot. J. Linn. Soc. 179: 36. 2015. Uncinia scabriuscula G.A.Wheeler, Hickenia 2(45): 215. 1997. Mistakenly reported as occurring in the Juan Fernández archipelago both in Rodríguez et al. (2018) and Zuloaga et al. (2019), as already noted by Wheeler (2007). This species is an endemic element of the temperate forests of southern Chile and Argentina (Wheeler 1997), and resembles the insular endemism of the Alejandro Selkirk island Carex aspericaulis J.R. Starr (G.A.Wheeler), to which it is also phylogenetically related (Wheeler 2007; Muñoz-Schüler et al. in prep.). Further notes on its taxonomy and chorology are given in the upcoming revision of the subgenus Uncinia in South America.

Carex turbaria J.R.Starr, Bot. J. Linn. Soc. 179: 34. 2015. Uncinia austroamericana G.A.Wheeler, Darwiniana 43: 271. 2005.

A recently described species, initially considered a Subantarctic element and restricted to Magallanes Region in Chile (Wheeler 2004). Here we present novel reports for Araucanía and Los Lagos administrative regions, greatly expanding its distributional range and setting a new northernmost limit for the species. This species was collected growing on boggy soils in the highest areas of the Nahuelbuta coastal range and in subandean wet meadows at the Puyehue National Park. In its northernmost limit it was found very near to the regional limit with Biobío. Further fieldwork might confirm its presence in contiguous bogs at the Biobío part of Nahuelbuta range.

SPECIMENS: CHILE. Región de la Araucanía: Prov. Malleco: Parque Nacional de Nahuelbuta, en el límite regional con la Región del Biobío, 37°48’07.01”S 73°01’04.87”W, 1265 m, 21-1-2023, S. Martín-Bravo, P. Muñoz-Schüler & M. Sanz-Arnal, 28SMB23 (CONC 193526). Región de Los Lagos: Prov. Palena: volcán Hornopirén, 41°53’S 72°25’W, 500 m, III-1993, Godoy et al., 111 (CONC 145778).

Carex wheeleri J.R.Starr, Bot. J. Linn. Soc. 179: 37. 2015. Uncinia araucana G.A.Wheeler, Aliso 15: 3. 1996[1997].

A rare species currently known only from two locations at the cordillera de Las Raíces, in the Andean mountains of Araucanía Region. We present here additional collections from the type locality that confirm its persistence there. Further fieldwork might confirm its presence on contiguous mountain ranges in the Biobío Region.

SPECIMEN: CHILE. Región de la Araucanía: Prov. Malleco, Lonquimay, salto Alaska, estero La Cascada, 38°30’16.78”S 71°24’45.22”W, 1003 m, 22-1-2023, P. García-Moro, J.I. Márquez-Corro & P. Jiménez-Mejías, 1pgm23 (CONC 193531).

D. Carex subg. Carex

D.1. Carex Hirta Clade

Carex aematorhyncha É.Desv., Fl. Chile 6: 224, t. 73, f. 22. 1853.

One of the most singular species of subg. Carex in the Southern Cone. It is readily recognized from any other species by its densely tomentose utricles with prominent nerves and an obscurely bifid and red colored beak that contrasts with the rest of the body. Philippi (1864) described Carex corralensis Phil., which was later combined by Kükenthal (1909) as C. aematorhyncha var. corralensis (Phil.) Kük. It was reportedly distinguished by the type variety by having a shorter culm and narrower leaves and spikes (Culmus humilior gracilior. Folia spiculaeque angustiora). Barros (1969) accepted this treatment in Flora Patagonica, essentially recording Kükenthal’s synopsis for the varieties distinction and not extending further with measurements. In addition, the description made by Barros (1969) for the type variety overestimates the measure of some characters, and might have been partially taken on material referable to the similar Carex hookeri, as the culm and utricle length, as well as the leaves width are greater than the size exhibited by the materials of C. aematorhyncha that we have studied, and no reference to the distinctive red colored beak is mentioned. After a thorough revision in the studied herbaria, we strongly suggest not to consider an infraspecific treatment for Carex aematorhyncha as it seems to be an artificial segregation of the wide natural variation that this species exhibits.

Carex excelsa Poepp. ex Kunth, Enum. Pl. 2: 502. 1837.

A widely distributed species formerly treated as Carex pseudocyperus L. is here newly reported for Atacama Region and consequently expanding its northernmost distributional range. We also confirm its presence for Los Ríos Region, where it has been previously reported in floristic works (Ramírez et al. 1980; Teillier et al. 2013), but surprisingly not reported in Rodríguez et al. 2018.

SPECIMENS: CHILE. Región de Atacama: Prov. Huasco, río Chollay, crece en suelo húmedo, junto a otras malezas, 29°00’S 70°09’W, 1.900 m, I-1994, A.S.L., 94083 (ULS 4457). Región de Los Ríos: Prov. Valdivia, Panguipulli, mirador Rucatrehua, orillas del lago Panguipulli, 39°45’2.98”S 72°9’45.36”W, 147 m, 21-XI-2019, D. Penneckamp 162 (CONC). Prov. Ranco, lago Maihue, río Pillanleufú, cerca de la desembocadura, humedal en sitio sombrío de la orilla, junto a Cyperus eragrostis, 40°11’50.5”S 72°00’22.4”W, 154 m, I-2023, P. Muñoz Schüler, P. García Moro & S. Martín Bravo, PMS-272 (CONC 193505). Prov. Ranco, Río Bueno, carretera de Crucero a Entre Lagos, cunetas inundadas, 40°35’35.0”S 72°38’34.8”W, 210 m, 26-I-2023, J.I. Márquez Corro, P. Jiménez Mejías & M. Sanz Arnal, 40JMC23 (CONC 193506).

Carex hookeriKunth, Enum. Pl. 2: 490. 1837.

A poorly understood taxon of intermediate appearance between Carex chilensis Brongn. and C. aematorhyncha. It was described in the 19th Century, and to date it was known from a few collections (Kunth 1837; and as C. beecheyana in Boott (1858)). Here we present the first recent record for this species in more than 100 years, a population that constitutes a dominant stand on brackish waters in Chiloé island (Los Lagos Region). Despite the morphological similarity to the two above-cited species may indicate a hybrid origin, in the reported population only a few isolated individuals of C. aematorhyncha were found but no C. chilensis. In addition, all the collected specimens were fully fertile and had no signs of malformation. The distribution of this species according to Rodríguez et al. (2018) is far larger than expected for such a rare species, and it might be overestimated because of misidentification with the resembling C. chilensis or C. aematorhyncha. Further revision of herbarium collections of this group of Carex might correct this distribution.

SPECIMEN: CHILE. Región de Los Lagos: Prov. Chiloé, camino a la playa Mar Brava, río Pudeyi, pastos estacionalmente inundados dominados por Schoenoplectus cf. pungens y Carex hookeri, acompañadas de C. aematorhyncha, 41°55’04.6”S 73°59’45.3”W, 0 m, 29-I-2023, P. Jiménez Mejías, P. García Moro, J.I. Márquez Corro, S. Martín Bravo, P. Muñoz Schüler & M. Sanz Arnal, 42PJM-CL23 (CONC 193507).

Carex pumila Thunb. ex Murray, Syst. Veg., ed. 14.: 846. 1784. A western Pacific species widely distributed on the coasts of NE Asia as in Australia and New Zealand (POWO 2023). It has been reported as introduced in Chile, where it is widely distributed and may form dominant communities on coastal dune fields. We present a new report for Ñuble. We consider that the introduced status of the species should be re-evaluated.

SPECIMEN: CHILE. Región del Ñuble: Prov. Itata, Trehuaco, Boca Itata, 36º22’S 72º51’W, 15 m, I-2011, José L. Solís B., s.n. (VALD 696).

Carex werdermannii Gross, Notizbl. Bot. Gart. Berlin-Dahlem, 10: 763. 1929.

An endemic from southern Chile that inhabits peatlands of the Coastal and Andean ranges up to 1000 m. Previously reported only from its type locality, in Llanquihue on Los Lagos Region, it was erroneously reported by Rodríguez et al. (2018) as exclusive from Los Ríos Region. Here we provide additional locations from Los Ríos and rectify its presence in Los Lagos by providing new collections from Chiloé, thus effectively expanding its southernmost range limit.

SPECIMENS: CHILE. Región de los Ríos: Prov. Valdivia, Corral, 39°56’54.60”S 73°27’35.12”W, 517 m, 23-1-2023, J.I. Márquez-Corro, P. Jiménez-Mejías & M. Sanz-Arnal, 30JMC23 (CONC 193528). Prov. Ranco, cordillera Pelada, Parque Nacional Alerce Costero, 40°09’56.87”S 73°30’18.13”W, 954 m, 1-2-2023, J.I. Márquez-Corro, P. Muñoz-Schüler, P. Jiménez-Mejías, S. Martín-Bravo, M. Sanz-Arnal & P. García-Moro, 73JMC23 (CONC 193529). Prov. Ranco, cordillera Pelada, Parque Nacional Alerce Costero, 40°10’04.69”S 73°30’28.87”W, 965 m, 1-2-2023, J.I. Márquez-Corro, P. Muñoz-Schüler, P. Jiménez-Mejías, S. Martín-Bravo, M. Sanz-Arnal & P. García-Moro, 67JMC23 (CONC 193530). Región de los Lagos: Prov. Chiloé, cordillera de Piuchué, San Pedro, 42°19’24.20”S 73°53’42.32”W, 620 m, 30-1-2023, P. Muñoz-Schüler, P. García-Moro, P. Jiménez-Mejías, J.I. Márquez-Corro, S. Martín-Bravo & M. Sanz-Arnal, PMS-276 (CONC 193527).

D.2. Carex Flacca Clade

Carex banksii Boott, Trans. Linn. Soc. London 20: 119. 1846. Carex fonkii Phil., Linnaea 29(1): 83. 1858.

Carex banksii Boott var. fonkii (Phil.) Kük., Bot. Jahrb. Syst. 27(4): 522. 1899.

A very characteristic and widely distributed species in Chile and adjacent Patagonian Argentina. The taxonomy of C. banksii and allied names is highly problematic, with several infraspecific taxa differently subsumed or divided depending on the author (Leveillé 1915; Barros 1969). Wheeler (2007) created the sect. Pellucidae G.A.Wheeler to accommodate C. banksii together with what he considered other two closely related but independent species: C. odontolepis and Carex stuessyi G.A.Wheeler. However, while Wheeler performed a comparison between C. banksii and C. stuessyi, he did not provide any information about the differences among these species and C. odontolepis. Unpublished data by Benítez-Benítez et al. show that samples assignable to sect. Pellucidae are grouped in three clades: one from Patagonia, another from the central Andes from Maule to Valparaíso, and a last one from Juan Fernández. The identity of the plants in each clade corresponds with the three names considered by Wheeler, namely, C. banksii, C. odontolepis and C. stuessyi.

The examination of specimens regarded by Wheeler as C. odontolepis readily reveals some characters that allow the distinction of this species from the more widely distributed C. banskii. Those characters are the ones we include in our identification key (see below).

Although we have not seen the type of C. fonkii, given the geographic origin of the type in the Andes of Los Ríos Region, we provisionally consider it to be a synonym of C. banksii.

Carex dusenii Kük., Ark. Bot. 7(2): 13, pl. 4, 7. 1907.

A very poorly understood species described from Argentinian Patagonia (lago San Martín, Santa Cruz; Dusén 1907) and so far, known only from the type collection. This species was compared by Kükenthal to C. macrosolen Steud., pointing to a relationship with the acaulescent species placed in former sect. Abditispicae G.A.Wheeler. Remarkably, this affinity is supported by unpublished data by Benítez-Benítez et al., which makes C. dusenii the only species of this group with well-developed stems.

We present the second and third known record for the species, first ones for Chile and also first ones in about 100 years. These constitutes a large disjunction of more than 600 km South and 700 North from the type locality, setting its new northermost and southernmost distribution limits. Very probably, this species could be present in the vast area between the three locations, but it has been so far neglected by collectors.

SPECIMEN: CHILE. Región de Los Lagos: Prov. Palena, comuna de Hualaihué, San Ignacio de Huinay, subida al cerro Tambor, 12-XII-2013, R. Fitzek, LM7347 (CONC, MA). Región de Magallanes: Prov. Tierra del Fuego, valle de Los Castores, turbera pulvinada en el bosque, 54°5’1.33”S 68°52’8.53”W, 420, 7-III-1995, M.T.K. Arroyo & A.M. Humaña, 95-1946 (CONC).

Carex odontolepis Phil., Linnaea 29: 82. 1858.

Carex banksii Boott var. gilliesii (Phil.) Kük., Pflanzenr. IV. 20: 555. 1909.

Carex dessaueri Phil., Anales Univ. Chile 93: 496. 1896.

Carex gilliesii Phil., Anales Univ. Chile: 556. 1873.

Carex ochrostachya Phil., Anales Univ. Chile 93: 500. 1896.

After the examination of the type material of the names listed here, stored in JStor Global Plants, we found them to be a better match to C. odontolepis than to C. banksii, thus creating a reliable synonymy for the first. For further information on C. odontolepis and its relationship with C. banksii, see the text under this latter.

D.3. Carex sect. Fecundae Kük.

Carex catamarcensis C.B. Clarke ex Kük., Bot. Jahrb. Syst. 27(4): 518. 1899.

An extremely rare and poorly collected species, described from Argentina and with a single and old report from Chile based on an immature collection from Coquimbo Region (C. latibracteolata Kük., Bot. Jahrb. Syst. 27(4): 518 (1899); río Torca, Philippi, SGO digital image!). It is the southernmost occurring member of sect. Fecundae, a group of Carex with tropical affinities endemic to the Americas. Here we report a new collection of several well-developed specimens, the first recent report of the species in Chile, not far from the original collection in the country. Since the vegetation and climate in this part of the Cordillera does not seem to be similar for other species of sect. Fecundae, nor resembles the areas where C. catamarcensis has been reported in Argentina (P.J.-M. pers. obs.), we presume that the reported population must be restricted to very specific environments. Accordingly, the persistence of the reported population must be confirmed in the immediate future and measures for its preservation taken if necessary.

SPECIMEN: CHILE. Región de Coquimbo: Prov. Limarí, El Polvo, 31°08’S 70°35’W, 2.300 m, XII-1976, Jiles, 6393 (CONC 103461).

D.4. Carex sect. Spirostachyae Drej. ex L.H.Bailey

Carex fuscula d’Urv., Fl. Iles Malouin.: 28. 1825.

Carex araucana Phil., Anales Univ. Chile 93: 495. 1896.

Carex distenta Kunze ex Kunth, Enum. Pl. 2: 449. 1837. Carex inconspicua Steud., Syn. Pl. Glumac. 2: 221. 1855.

A widely distributed species in South America, spanning across the Andes from South Bolivia to Tierra del Fuego. A closely related species, C. catharinensis Boeckeler, appears in the Atlantic highlands of S Brazil, Uruguay, and the Argentinian province of Buenos Aires (Jiménez-Mejías et al. 2016b).

In the most strict sense, C. fuscula is a Patagonian plant. Specimens from temperate Chile has been variously described as different taxa (see synonyms listed here below). Our observations show that C. fuscula s.l. is extremely polymorphic, and that within a same stand, forms assignable to different taxa can be found. This raises doubts about the taxonomic validity of such syntopic morphs. Accordingly, we strongly suggest treating all these names under a broadly conceived C. fuscula until a thorough biosystematic study unravels this intricate situation.

Carex lamprocarpa Phil., Anales Univ. Chile 93: 497. 1896.

A very rare species from mainland Chile whose closest affinities seems to be with the Juan Fernandez endemic C. berteroniana Steud.

Carex lamprocarpa is only known from two old collections in southern central Chile: “Valdivia, Philippi, Feb 1888” (K!) from Los Ríos Region; and “Temuco, Maquehue, R. Morton Middleton, Dec 1905” (BM013822604!) from Araucanía Region. In the original protologue, Philippi provides additional information that places the type locality at the southern side of the Valdivia river: “In praedio meo San Juan (prov. Valdivia) loco dicto Paso de las Piedras in umbrosis humidis inverni”). We extensively searched the species in remnants of original vegetation at both locations (Temuco and caleta de San Juan) without success. According to the indications, this species must grow in shady river forests.

The affinities of the mainland C. lamprocarpa with the island C. berteroniana were discussed by Wheeler (1988b, 2007), and previously noted in herbarium labels by the determination of the Los Ríos material as the Juan Fernández species (under the name C. paleata Boott). Preliminary results show that C. berteroniana is deeply nested within a clade of New Zealand taxa (Míguez et al. 2022), supporting a long-distance dispersal event from the western to the eastern Pacific. According to morphological evidence, the presence of C. lamprocarpa in mainland Chile should be also related to this colonization event.

Wheeler (1988b, 2007) proposed the independence of C. lamprocarpa and C. berteroniana mainly on the basis of their different distributions. Implicitly, he also presumably relied on differences on utricle morphology, as showed in the utricle comparison performed in the accompanying figure: a suborbicular utricle abruptly constricted in the beak and base in C. berteroniana vs. a broadly fusiform utricle, attenuated towards both the beak and the base, in C. lamprocarpa. However, he studied a very limited amount of material, which may have biased this observation. The study of a larger number of collections of C. berteroniana by the last author of this paper (P.J.-M.) revealed that, sometimes, the utricle morphology of the insular species approaches that of the mainland one, blurring this presumed limit between the two taxa. This opens a window to the possibility of C. lamprocarpa populations being an accidental introduction of C. berteroniana in the continent. Indeed, Philippi visited the archipelago a few years before the description of the mainland plant as C. lamprocarpa, which -curiously- he found in a land of his property: “in maeo predio”, it is, “in my land”. It seems to us necessary to search and study material from the mainland C. lamprocarpa in order to address its independence, discard a possible human introduction, and urge conservation measures if necessary.

The alien C. pendula superficially resembles C. lamprocarpa.

It is an introduced species recently naturalized in Valdivia town environs. Carex pendula is much larger than C. lamprocarpa, and the utricles are smaller, duller, and lack red spots.

After the thorough revision performed by the authors of this paper, and the almost total absence of vouchers, we consider the reports of C. lamprocarpa from Los Lagos and Biobío erroneous.

Carex poeppigii C.B.Clarke ex G.A.Wheeler, Aliso 12(1): 97. 1988.

A mysterious species known from a single collection from the Antuco area (Biobío Region). Duplicates of this collection are found in BM (digital image!), K(!), LE (digital image!), and P (digital image!). This species is closely related to C. berteroniana and C. lamprocarpa (Wheeler 1988b; see above).

The label of the original material (Poeppig 247) only bears the label “Antuco”, handwritten by Poeppig. There is an additional collection from the area (cordillera de Polcura, Zoellner 2913, 1968, CONC), labeled as C. lamprocarpa and currently unlocated, that could potentially belong to this species. The labeling of material by Poeppig has been regarded by some as problematic (S. Teillier, pers. comm.). However, Poeppig seemed to be fairly constant when naming places (D. Alarcón, pers. comm.). During his visit to the valley of the Laja river in the year 1828, Poeppig distinctly indicated when he visited the volcano (“mont. Igniv. Antuc.”, see below C. antucensis) or other places such as the surrounding mountains of Antuco (e.g., “In Chil. Austr. Rupib. Cacuminis Pico de Pilque [Cerro Pilque]”, in the syntype of Azara alpina Poepp. & Endl.). Whether the indication “Antuco” in C. poeppigii original label refers to the volcano or the town in the valley is unclear to us. According to Poeppig (1960) himself, only one particular place not far from Antuco town was explicitly mentioned: Quillailebu river, a stream on the south side of Laja valley that goes upslope Sierra Velluda volcano. That may well be the location where Poeppig may have found the plant, although there seem not to be other Poeppig’s collections from this area.

During our 2023 campaign we failed in finding C. poeppigii on the slopes over Polcura town. We were unable to enter Sierra Velluda from the Laja valley or go upstream of the Polcura river. Much additional fieldwork in the area is needed to locate this taxon or discard a possible mislabeling by Poeppig.

Carex trifida Cav., Icon. 5: 41, t. 465. 1799.

A Circumantarctic large sedge species known from the southernmost tip of the South Cone, the Malvinas, as well as from New Zealand and its satellite archipelagos. It is a species or maritime affinities, known from rocky shores and cliffs, often growing associated with seabird colonies (Martín-Bravo pers. comm.; also see Johnson 1976; Bergstrom et al. 2006). In Chile it was reported from Magallanes and Aysén administrative regions. Here we report two surprising records from Chiloé (Los Lagos Region), which becomes the northernmost records of the species in its entire area.

SPECIMENS: CHILE. Región de los Lagos: Prov. Chiloé, isla Guafo. 43°36’S 74°44’W, 100 m, II-1986, Villagrán & Leiva, 7492 (CONC 75926). Prov. Chiloé, isla Quilán, 43°24’S 74°15’W, 25 m, X-2-1986, Villagrán & Leiva, 7214 (CONC 76722).

D.5. Carex sect. Rhynchocystis Dumort.

Carex pendula Huds., Fl. Angl.: 352. 1762.

A mainly European species widely cultivated, naturalized in certain areas of North America, New Zealand (Míguez et al. 2017, 2018) and China (as C. tenuirostrata X.F.Jin & D.F.Wu; Jin et al. 2012). Here we present the first reports of the species growing escaped from cultivation in Chile and South America.

SPECIMENS: CHILE. Región de los Ríos: Prov. Valdivia, 39°48’35.59”S 73°15’27.83”W, P. Muñoz Schüler, P. García-Moro & S. Martín-Bravo, PMS-268 (CONC 193532). Prov. Valdivia, Valdivia, 39°50’S 73°14’W, 15 m, 12-1961, Gunckel, 37566 (CONC 67878).

D.6. Carex sect. Phacocystis Dumort.

Carex antucensis Kunze ex Kunth, Enum. Pl. [Kunth] 2: 412.1837.

LECTOTYPE here designated: “Nº 29, Carex Nº 3 // Poeppig Coll. pl. Chil. III. 248. (29) In mont. igniv. Antuc., Chil. austr. Febr. lecta” (P-00304419 digital image!)

= Carex chillanensis Phil., Anales Univ. Chile 21: 384. 1862.

HOLOTYPE: CHILE: “Valle de las Aguas Calientes, 27 Feb 1862” R.A.Philippi 46 (SGO-37602 digital image!)

Carex goodenoughii sensuBarros (1969), non Carex goodenoughii Asch. & Graebn., Mitteleurop. Fl. ii. pt. 11. 94.

Section Phacocystis is one of the most problematic groups of Carex, whose taxonomy is much affected by the faint morphological limits between species and hybridization events (Benítez-Benítez et al. 2021). The group in South America makes no exception.

On the one hand the name Carex chillanensis Phil. was described by R.A. Philippi in 1862 (Philippi 1862) based on material collected in Chillan (valle de las Aguas Calientes, Philippi, 27 Feb 1862; SGO-37602 digital image!). Barros (1969) recorded the species as C. goodenoughii Asch. & Graeb. (a synonym of the Western Palearctic C. nigra (L.) Reichard), due to the superficial similitude between both species. On the other hand, C. antucensisKunth was described in 1837 (Kunth 1837) based on material collected by E. Poeppig in Antuco. The inspection of the protologue and type material of C. antucensis revealed that the original material of that latter is indistinguishable from that of C. chillanensis, which makes C. antucensis the prior name for the species. Remarkably, Philippi already noticed in the protologue of C. chillanensis the similitude between it and C. antucensis. Despite this, he still proceeded with the formal description of the new species after mistaking the number of stigmas between them (reporting three in C. chillanensis vs. two in C. antucensis). To help stabilize the correct identity of the two names, we proceed to lectotypify C. antucensis here.

Supplementary Material. Regional distribution of Carex species in Chile (Excel file).

ACKNOWLEDGEMENT

We would like to thank Conaf and its park rangers for granting us collecting permission and assistance during fieldwork in the National Parks and Reserves of the SNASPE that we visited during our fieldtrip expedition in January 2023. We would also like to thank Alicia Marticorena, curator of CONC herbarium in Concepción, for her unconditional support regarding the access of information, material and for her help in logistical issues before, during and after the field trip expedition. In the same line, we would like to thank Gina Arancio, curator of ULS herbarium in La Serena, and her helpful assistant Matías Herrera, as well as Nicolás García, curator of EIF herbarium in Santiago, for letting us revise the Carex collection of both herbaria. Also, to Sebastián Teillier and Diego Alarcón, for providing us relevant information about sampling localities, historical aspects of some old collections, and keep interest in the update of Carex in Chile. Thanks to Nicholas Turland for a useful discussion on the homonymy of C. berteroana and C. berteroniana. Finally, we would like to thank María Ismenia Barría, Diego Penneckamp’s grandmother, for her hospitality while visiting their home at Pitriuco, in Southern Chile, as well as Begoña Peceño, professor of Universidad Católica del Norte, and her husband Juan, for their friendly hospitality while we stayed at their home in the city of Coquimbo.

This research has been funded by project PID2020-113897GB-I00 (DANZ, to P.J.-M. and S.M.-B.) and a Ramón y Cajal postdoctoral contract (ref: RYC2021-031238-I), both from the the Ministerio de Ciencia e Innovación from Spain, and by an IAPT Research Grant 2022 (to J.I.M.-C.).

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IDENTIFICATION KEY TO THE SPECIES OF CAREX GROWING IN CHILE

Received: July 26, 2023; Accepted: November 02, 2023

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