Research Article |
Corresponding author: Paulo Muñoz-Schüler ( paulomschuler@gmail.com ) Academic editor: Muthama Muasya
© 2024 Paulo Muñoz-Schüler, Ana Morales-Alonso, José Ignacio Márquez-Corro, Mary T. K. Arroyo, Santiago Martín-Bravo, Pedro Jiménez-Mejías.
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.
Citation:
Muñoz-Schüler P, Morales-Alonso A, Márquez-Corro JI, Arroyo MTK, Martín-Bravo S, Jiménez-Mejías P (2024) Carex recondita Muñoz-Schüler, Martín-Bravo & Jim.Mejías (Carex section Junciformes Kük., Cyperaceae), a new sedge species from the Andes of central Chile. PhytoKeys 243: 15-30. https://doi.org/10.3897/phytokeys.243.115991
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Carex section Junciformes is one of the most diverse groups of the genus in South America, consisting of approximately 30 species. Here we describe a new species, Carex recondita, belonging to this section. We studied its placement within a molecular phylogeny of the group and found it to constitute an independent lineage. The new species is morphologically very close to C. austroamericana, from southern Patagonia, despite being phylogenetically divergent to the rest of Patagonian species of sect. Junciformes. So far, this species is known only from a few specimens recently collected in its type locality, despite growing in a well-collected area in the Andes of Metropolitana Region of Santiago, the most populated administrative region of Chile. We provide a detailed morphological description, comments on its relationship with other Southern Cone species of sect. Junciformes and relevant ecological notes.
Andes, Carex, Chile, Cyperaceae, South America, taxonomy
With about 96 species, Carex is among the three most biodiverse genera of the flora of Chile (
Carex section Junciformes Kük (subgenus Psyllophorae), with over 25 species in the continent (
During a fieldtrip in Chile in January 2023 we collected material of an undetermined acaulescent Carex sect. Junciformes taxon in the Andes of Metropolitana region of Santiago (see Taxonomic Treatment for details on the collection). In a preliminary assessment using the available identification keys (
Silica-dried leaves of the undetermined taxon collected on the field in January 2023 were used for DNA extraction and PCR amplification.
The DNA extraction was performed using a modified CTAB procedure (
The phylogenetic reconstructions analysis were run employing Maximum Likelihood (ML) in RAxML v.8.2.12 (
Three collections of the undetermined taxon were located and studied: the original collection from the 2023 fieldwork campaign, a second collection by M.T.K.A. in 2022, and a last voucher located in CONC collection, determined as Carex andina Phil., and collected in 2007. These vouchers were determined using the available keys and thoroughly compared against descriptions (
Direct comparisons were also carried out against material (including type material -herbarium specimens or high-resolution images of them-) belonging to the other three Junciformes clade acaulescent species: C. argentina Barros, C. austroamericana, and C. nelmesiana. In total, we studied 17 vouchers housed at BAB, CONC, SI, and UPOS.
Since our results showed that the undetermined taxon constitutes an undescribed species with a very restricted known distribution, a preliminary conservation status for it was assessed. We followed the criteria provided in the IUCN Red List Categories and Criteria Version 13 (
The multiaccession matrix consisted of 606 bp for ETS, 624 bp for ITS, 572 bp for matK, and 937 bp for rps16. The length of obtained sequences for the problematic specimen were 493 bp for ETS, 603 bp for ITS, 503 bp for matK, while for rps16 marker only 285 bp could be obtained. The obtained BI tree topology (Fig.
The sample of the undetermined taxon (Fig.
The samples of the undetermined specimen share a number of morphological affinities with C. austroamericana, such as an acaulescent growing habit, stiff leaves that surpass the entire inflorescence, and stipitate utricles. However, it can be distinguished from this and any other species of sect. Junciformes by having the following combination of characters: leaves up to 9 cm long, spikes ≤ 3.5 mm wide, and utricles ellipsoid, ovoid or sub-obovoid, glabrous to sparsely hispidulous, with -usually weakly- raised nerves on the faces and a stipitate base often flattened. A detailed morphological comparison between these problematic specimens, C. austroamericana and the other three acaulescent species of sect. Junciformes from the Southern Cone is provided in Table
Variation in diagnostic morphological characters of C. recondita and morphologically related Patagonian species of sect. Junciformes. Characters listed for C. austroamericana and C. nelmesiana are based largely on
Character | Carex austroamericana | Carex nelmesiana | Carex argentina | Carex recondita |
---|---|---|---|---|
Leaf length | 0.5–5 cm | 1–5 cm | 8–12 cm | 2.9–9 cm |
Spike width | 3.5–6 mm | 5–6 mm | 5 mm | 2.8–3.5 mm |
Number of female flowers/utricles | Four to six | Four to seven | Five to seven | Four to five |
Utricle shape | Obovoid | Sub-globose | Ovate to lanceolate | Elliptical, sub-fusiform or sub-obovate |
Utricle indurnenturn | Glabrous, but scaberulent on the beak and the margins of the distal third of the body | Puberulent, margins hispidulous | Flocculent | Glabrous to hispidulous, the margins scaberulent on the distal half of the body or beyond |
Utricle venation | Faintly or prominently veined, up to 6–8 prominent veins abaxially and 4–6 veined adaxially | Faintly veined or with 4–7 prominent veins in the proximal third of the abaxial side | Veinless | Weakly 2–5 veined adaxially and 1–4 veined abaxially |
Utricle base | Stipitate, rounded or flattened in cross section | Shortly stipitate | Truncate to stipitate | Stipitate and flattened in cross section |
Utricle margins | Rounded | Rounded | Rounded | Flattened |
Beak | Conical, bidentate | Shortly conical and obscurely bidentate | Longly conical and strongly bidentate | Conical, short and bidentate |
Rachilla | Subulate | Ovate | Lanceolate | Linear |
From the other three Junciformes clade species not included in the phylogeny (C. boelckeiana, C. moorei and C. nelmesiana) the undetermined taxon is readily distinguished because the two first species have well-developed stems, longer than leaves, as well as larger utricles (see
The set of evidence presented here makes it clear that the studied specimens should be treated as a new species. Even though it is morphologically very similar to C. austroamericana, the molecular data by itself is compelling enough to support its status as a different taxon (Fig.
Andean-Patagonian disjunctions are not uncommon within and between closely related species of Carex (e.g. South American sect. Racemosae,
Carex recondita is similar to C. austroamericana, from which it is distinguished by its longer leaves up to 9 cm long (versus 5 cm long in C. austroamericana), narrower spikes up to 3.5 mm wide (versus 3.5–6 mm wide), and weakly veined ellipsoid, obovoid or suborbicular utricles, with a flattened stipitate base (versus strongly veined obovoid utricles, with a rounded or flattened stipitate base).
Chile. Región Metropolitana de Santiago: Provincia de Santiago, Farellones, camino al centro de ski Valle Nevado, Tres Puntas, 2750 m.a.s.l., 33°21.39183'S, 70°15.45538'W, 16 January 2023, P. Jiménez Mejías, J.I. Márquez Corro, S. Martín Bravo & P. Muñoz Schüler 12PJM-CL23 (holotype: CONC 193519; isotypes: EIF 17307, UPOS).
Plants low-growing, densely caespitose 3–10.2 cm tall from the base of the shoots to the tip of the leaves, acaulescent or nearly so. Leaves numerous, up to eight per shoot, much longer than the flowering shoots and concealing them, 2–8.2 cm long, canaliculate but flattish distally, stiff to slightly flexuose. Flowering shoots elevated on erect stalks of 1.2–4.4 cm long formed by the basal leaf sheaths, with the portion between the insertion of the distal-most leaf and the proximal-most bract inconspicuous, up to 1.2 mm long. Inflorescence a single, terminal, androgynous, subglobose spike, 4.3–6.2 mm long and 2.8–3.5 mm wide, subtended by an involucral bract, with a glumaceous base and prolonged into a 5.8–9.1 mm long antrorsely scabrid or smooth setaceous portion. Staminate part concealed by the pistillate portion, 2-flowered. Pistillate part 4–5 flowered, sometimes with an extra infertile flower borne by the involucral bract; glumes ovate to widely elliptical, 3.1–4.1 mm long and 1.4–2.3 mm wide, glabrous, with wide hyaline margins contrasting with a narrow green middle strip, veinless, attenuated distally into an awn, the 2-proximalmost glumes with a 1.9–3.3 mm long awned portion that usually surpasses the whole inflorescence, distalmost glumes with a 0.6–0.9 mm long mucronate apex. Utricles 2.4–3 mm long and 1.2–1.8 mm wide, ellipsoid, suborbicular, ovoid or sub-obovoid, obscurely trigonous in cross section, green to pale green, with 2 prominent lateral veins dark greenish, contrasting with the rest of the body, veinless or 2–5 weakly veined adaxially and 1–4 veined abaxially, glabrous to sparsely hispidulous on its distal half, with the lateral veins hispidulous to ciliolate on its distal ⅔ portion, attenuated or constricted proximally into a sub-stipitate base 0.4–0.8 mm long, often flattened in cross section, constricted distally into a short, bidentate, and pale beak 0.3–0.6 mm long. Achenes 1.9–2.2 long and 1.2–1.6 mm wide, broadly elliptical to sub-obovoid, obscurely trigonous, often flattened, greenish, more or less constricted proximally into a sub-stipitate base and attenuated distally into a short beak. Rachilla often absent, when present 0.6–0.9 mm long, linear, margins smooth. Stigmas 3. Anthers 3.
The phenology of this species is poorly understood. A fully flowering specimen with some ripe utricles was collected in mid-late December, while a specimen with entirely ripe utricles bearing some female flowers was collected in mid-January. According to these observations we infer that the flowering period for the new species is likely to range from mid-November to late December with mature individuals bearing ripe utricles from mid-late December onwards. This agrees with the plant community peak flowering period at 2935 m.a.s.l. (late-December to early-January) in an adjacent high Andean valley (
The specific epithet recondita (female) refers to the Latin word recondito (male), meaning something hidden or occult, apropos the acaulescent inflorescence and low-growing habit of this species.
Carex recondita occurs in the area of La Parva and Farellones-Valle Nevado ski circuits, in the Andes of the Metropolitana region of Santiago (see Fig.
Carex recondita Muñoz-Schüler, Martín-Bravo & Jim.Mejías A, B aerial and frontal view of the plant and a spike C, D habit E two of the authors looking for more specimens of C. recondita on its habitat, the road can be seen at the upper-left portion of the picture F habitat of C. recondita, showing dominant Azorella ruizii (green cushions) and Anarthrophyllum cumingii (prostrate shrub with orange-yellow flowers). Photos by P M-S (A–E) and MTKA (F).
Detailed comparison between C. recondita and C. austroamericana A spikes of C. recondita (Aa) and C. austroamericana (Ab) B utricles of C. recondita (Ba) and C. austroamericana (Bb) C glumes of C. recondita D rachillas of C. recondita (Da) and C. austroamericana (Db). Note that the rachilla of C. recondita is attached to the utricle base E achenes of C. recondita (Ea) and C. austroamericana (Eb).
Phylogram resulting from BI analysis using MrBayes of the multiaccession matrix for Carex subg. Psyllophorae. Branch support is not indicated when PP=1.00/BS = 100. Posterior probability (PP) support is given above branches while ML BS support is given below branches. Asterisks above branches indicate BI 100 > PP ≥ 0.95. Bold thick branches indicate nodes supported by ML BS ≥ 80 and/or BI PP ≥ 0.95. ML support BS ≤ 80 is given below branches. The newly sequenced voucher of C. recondita is displayed in bold red letters. Tip labels include the geographical origin of the specimen using TDWG level 3 regions abbreviations (“botanical countries”;
Occurrence map of Carex sect. Junciformes species cited in this work A occurrence map of five Patagonian species of Carex sect. Junciformes. Species locations are depicted in colored circles and some cities are represented as black squares B detailed map of the area of occurrence of C. recondita (light-green circles). Another species occurring in the area, C. argentina, is depicted by red circles. The highlighted region corresponds to the Yerba Loca Nature Sanctuary. Altitude was represented by a color palette and contour lines, representing elevation every 200 meters. For orientation purposes, some landmarks were depicted on the map.
We propose the common name for this species to be “Carex de las nieves”.
C. recondita has an area of occupancy (AOO) smaller than 10 km2, and only a single population (three subsets) of approximately < 200 individuals has been reported. Although the entire known individuals of this species are located in protected land (Yerba Loca Nature Sanctuary), the management of the area that is effectively being protected only includes the watershed of the La Yerba Loca stream, which is located 4 km west (in straight line) and approximately 1000 m below the area in which C. recondita grows. Notwithstanding this, the new species grows within one of the most crowded ski circuits of Chile (La Parva and Farellones-Valle Nevado circuit), on an area contiguous to one of its main roads, which has also been affected by exotic plant invasions (
C. recondita is a tuft-forming species with stiff leaves and an acaulescent spike burrowed between the leaves, a type of growth form shared with many other Patagonian species of sect. Junciformes and recurrent in South American Carex species (see
Chile. Región Metropolitana de Santiago: Provincia de Santiago, camino entre Farellones y Valle Nevado, laderas por debajo de la curva 14, piso altoandino, 2853 m a.s.l., 33°21.46188'S, 70°15.32352'W, 20 December 2022, M.T.K. Arroyo, V. Robles, K. Robles, M. Acevedo & L. Retamal 29576 (CONC). Provincia de Santiago, Farellones, a 1 km de la Parva. 2674 m a.s.l., 33°20'S, 70°17'W, 1 March 2007, M. Mihoc 777 (CONC 178536).
The authors would like to thank Alicia Marticorena, María Monsalvo and Manuel Belgrano, curators of CONC, BAB and SI, respectively, for their help in accessing the Carex collections of those herbaria. We would also like to acknowledge Sebastian Teillier and Aron Cádiz-Véliz for their help with the discussion on some elements of this work, such as the suggested vernacular name, the conservation assessment, and the epithet of the new species.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was financially supported by DANZ Spanish Research Agency of the Ministry of Science and Innovation (DANZ, award# PID2020-113897GB-I00 to P.J.-M. and S.M-B.), the Ramón y Cajal program (award# RYC2021-031238-I) to P.J.-M, and the IAPT Research Grant 2022 awarded to JIM-C. Fieldwork made by MTKA was funded by Fondecyt 1221540, CHIC- ANID PIA/BASAL PFB 210018 and IEB- ANID PIA/BASAL 210006.
Methodology: AM-A, PJ-M, PM-S. Writing – original draft: PM-S, PJ-M. Review and editing: SM-B, PJ-M, JIM-C, AM-A, MTKA, PM-S.
Paulo Muñoz-Schüler https://orcid.org/0009-0001-7070-4121
Ana Morales-Alonso https://orcid.org/0000-0003-1497-1070
José Ignacio Márquez-Corro https://orcid.org/0000-0003-4277-2933
Mary T. K. Arroyo https://orcid.org/0000-0001-6315-0426
Santiago Martín-Bravo https://orcid.org/0000-0003-0626-0770
Pedro Jiménez-Mejías https://orcid.org/0000-0003-2815-4477
All of the data that support the findings of this study are available in the main text.