Research Article |
Corresponding author: Carmen Benítez-Benítez ( karmenbenitez.93@gmail.com ) Academic editor: Marcin Nobis
© 2023 Carmen Benítez-Benítez, Pedro Jiménez-Mejías, Modesto Luceño, Santiago Martín-Bravo.
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:
Benítez-Benítez C, Jiménez-Mejías P, Luceño M, Martín-Bravo S (2023) Carex quixotiana (Cyperaceae), a new Iberian endemic from Don Quixote’s land (La Mancha, S Spain). PhytoKeys 221: 161-186. https://doi.org/10.3897/phytokeys.221.99234
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Despite centuries of work, the basic taxonomic knowledge of the flora of the Iberian Peninsula is still incomplete, especially for highly diverse and/or difficult genera such as Carex. In this study, we conducted an integrative systematic study based on molecular, morphological and cytogenetic data to elucidate the taxonomic status of several problematic Carex populations from La Mancha region (S Spain) belonging to Carex sect. Phacocystis. These populations have been traditionally considered of uncertain taxonomic adscription, but close to C. reuteriana due to their morphological appearance and ecological preferences. A detailed morphological and cytogenetic study was performed on 16 La Mancha’s problematic populations (Sierra Madrona and Montes de Toledo) to compare them with the other Iberian sect. Phacocystis species. In addition, a phylogenetic analysis was conducted using two nuclear (ITS, ETS) and two plastid (rpl32‐trnLUAG, ycf6‐psbM) DNA regions, including representatives from all species of sect. Phacocystis. We found a significant degree of molecular and morphological differentiation that supports the recognition of La Mancha’s problematic populations as a new Iberian endemic species, described here as Carex quixotiana Ben.Benítez, Martín-Bravo, Luceño & Jim.Mejías. Our results reveal that C. quixotiana, unexpectedly, is more closely related to C. nigra than to C. reuteriana on the basis of phylogenetic relationships and chromosome number. These contrasting patterns reflect the taxonomic complexity in sect. Phacocystis and highlight the need for integrative systematic approaches to disentangle such complicated evolutionary scenarios.
Cytogenetics, Iberian Peninsula, Mediterranean, morphometrics, new species, Phacocystis, phylogenetics, Quercus forest, taxonomy
The Iberian Peninsula is one of the three large peninsulas of Southern Europe projecting into the Mediterranean Sea, and as such, it harbours a high diversity of plant species and endemism relative to the rest of Europe (
Section Phacocystis Dumort. (subg. Carex) is among the largest sections of Carex (ca. 112 spp. in
La Mancha (Fig.
During the preparation of Flora Iberica (
Representative photos of Carex quixotiana A habitat (riparian forest with Alnus lusitanica and Fraxinus angustifolia: Ciudad Real, Solana del Pino, Robledillo River; 57SMB17, UPOS-8924; C. quixotiana plants are in the lower right corner) B habitat (stream in Quercus pyrenaica open forest: Ciudad Real, between El Viso del Marqués and San Lorenzo de Calatrava; 8CBB18, UPOS-16897; C. quixotiana are the tussocks along the stream) C inflorescence D, E detail of terminal male and lateral androgynous spikes F detail of lateral female spikes and utricles (Ciudad Real, Fuencaliente, Minas del Horcajo, 54SMB17, UPOS-8922).
We collected specimens from 10 problematic populations from La Mancha region (Sierra Madrona and Montes de Toledo; Fig.
Material from all 16 sampled populations (Suppl. material
To tackle the phylogenetic placement of La Mancha’s problematic populations, we included these samples in the molecular phylogeny of sect. Phacocystis (
DNA extraction and sequence amplification followed
To explore the systematic relationships of La Mancha’s problematic populations at a finer scale and with respect to the other Iberian representatives of sect. Phacocystis, a statistical parsimony analysis of plastid haplotypes was conducted with TCS v. 1.2.1 (
Chromosome counts for five La Mancha’s problematic populations (Suppl. material
Following the taxonomic recognition of La Mancha’s problematic populations (see below), we evaluated their conservation status at the global level following criteria, categories, and guidelines from
The detailed examination of diagnostic morphological characters for La Mancha’s problematic populations and its comparison with closely related species revealed qualitative and quantitative morphological differences regarding the other Iberian taxa of sect. Phacocystis (Table
Comparison of main morphological diagnostic characters, chromosome numbers, and habitat (on the Iberian Peninsula) among C. quixotiana and all the other Iberian Carex sect. Phacocystis taxa. Measurements from the other taxa have been taken from
C. quixotiana | C. acuta | C. cespitosa | C. elata subsp. elata | C. nigra | C. reuteriana | C. trinervis | |||
---|---|---|---|---|---|---|---|---|---|
subsp. nigra | subsp. intricata | subsp. reuteriana | subsp. mauritanica | ||||||
Habit | From tussock-forming to rhizomes elongated | From more or less caespitose to rhizomes elongated | Tussock-forming | From tussock-forming to laxly caespitose | Rhizomes elongated | Tussock-forming | Rhizomes elongated | ||
Basal sheaths (at the base of fertile stems) | Scale-like, sometimes elongated, creamy-yellow, rarely reddish-brown | Culm bases bearing brown old-leaf remains, without conspicuous scale-like basal sheaths | Scale-like, dark purple | Scale-like to elongated, creamy-yellow | Culm bases with or without conspicuous scale-like basal sheaths, dark brown when present | Scale-like, orange to reddish-brown | Culm bases bearing straw-colored old-leaf remains, without conspicuous scale-like basal sheaths | ||
Leaf section | Flat to keeled | Flat | Flat to keeled | Flat to keeled | Flat to keeled | Flat to keeled | Strongly plicate to canaliculate | ||
Leaf width (mm) | (1.8)2–3.2(4.7) | 3–5(7) | 2–4 | 3–6(7) | 3–6(10) | (3)3.5–5.5(6) | (3)4–8(10) | (0.5)1.5–2.5(3) | |
Stomata distribution | Amphistomatic | Hypostomatic | Hypostomatic | Hypostomatic | Epistomatic | Amphistomatic | Hypostomatic | Amphistomatic | |
Relative lowermost bract-inflorescence length | Longer to shorter than the inflorescence | Longer than the inflorescence | Shorter than the inflorescence, rarely equalling it | Shorter than the inflorescence, rarely equalling or exceeding it | Equalling to slightly shorter than the inflorescence, rarely exceeding it | Equalling the inflorescence | Longer than the inflorescence | ||
Terminal (male) spikes number | (1)2–3 | 2–4 | 1 | 1–2 | 1(2) | 1(2) | (1)2–4 | (1)2–3 | |
Terminal (male) spike length | (15)20–60(85) | (15)20–50(60) | (12)15–30 | 25–80 | (5)10–30 | (15)20–60(70) | (10)20–70 | (15)25–40 | |
Lateral androgynous spikes | (0)1–3(4) | 1–2 | 0 | 0–3(4) | (0)1 | 0–3(6) | (0)1–5 | 2–4 | |
Lateral female spikes | 0–2(3) | 2–4 | 1–2(3) | 0–2(3) | 2–4 | 0–3(4) | 0(1) | 2–4 | |
Utricle nerves | Nerved | Nerved | Nerveless | Nerved | Nerved | Nerved | Nerved | ||
Utricle indumentum | With high whitish papillae on the upper half or towards the apex, sometimes aculeolate at the apex | Almost entirely covered with high papillae, somewhat inflated | With whitish high papillae towards the apex | With whitish high papillae, at least towards the apex, not inflated | Covered with high papillae at least the upper half | Smooth, very rarely with some scattered, low papillae towards the apex | Covered with low papillae | ||
Chromosome number (2n) | 82–83 | 84–86 | 78–80 | 74–80 | (80)82–88(92?) | 73–76 | (72)74–76 | 82–85 | |
Habitat (Iberian Peninsula) | Small streams, rivers and springs in riparian forests (Alnus lusitanica, Fraxinus angustifolia, Salix spp.), humid meadows, in marcescent and sclerophyllous Quercus forests, on siliceous bedrock substrate | River shores, usually in deciduous forests, without clear edaphic preferences regarding bedrock substrate | River shores in deciduous forest, on granitic substrates | River shores (or lakes) in diverse types of vegetation, on calcareous bedrock substrates, rarely on siliceous ones, also on coastal swamps | Montane-alpine wet meadows, bogs, and swamps, also in river and lake shores, in diverse types of vegetation and without clear edaphic preferences regarding bedrock substrate | Stream and river shores in diverse types of vegetation, usually on siliceous bedrock substrates, rarely on calcareous ones | Stream and river shores in riparian forests, in marcescent and sclerophyllous Quercus forests, without clear edaphic preferences regarding bedrock substrate | Sandy coastal swamps | |
Altitude (m) | 400–800 | 0–1300 | 10–40 | 10–1750 | 1000–3300 | 300–1900 | 20–650 | 0–30 |
The query tree built (see Suppl. material
Phylogram of sect. Phacocystis s.l. based on the maximum likelihood phylogenetic reconstruction of the complete singletons tree (ITS, ETS, rpl32‐trnLUAG, ycf6‐psbM) by
The statistical parsimony analysis revealed 14 different haplotypes (Suppl. materials
The studied meiotic plates, representing five different La Mancha’s problematic populations (Suppl. material
Different sources of evidence (morphological, cytogenetic, molecular) support the systematic distinctiveness of the studied La Mancha’s problematic populations (Table
Remarkably, the DNA markers used in the phylogenetic reconstructions showed that La Mancha’s problematic populations do not group with C. reuteriana, to which they had been traditionally assigned (
According to the evidence presented, La Mancha’s problematic populations warrant taxonomic recognition, so we proceed to describe them as a new species.
Similar in appearance to C. reuteriana Boiss. & Reut., from which it differs by the creamy-yellow, rarely reddish-brown basal sheaths (vs. orange to reddish-brown), amphistomatic leaves (vs. hypostomatic), and utricles with high papillae (vs. smooth or rarely with low papillae). It is also similar to C. nigra (L.) Reichard, from which it can be distinguished also by the creamy-yellow, rarely reddish-brown basal sheaths (vs. absent or dark brown when present), narrower leaves (1.8)2–3.2(4.7) mm (vs. 3–6(10) mm), and longer terminal male spike (18)20–60(85) mm (vs. (5)10–30 mm).
Spain. Ciudad Real: Fuencaliente, Azor stream recreational area, stream edges in Quercus faginea forests, 733m, 38.44906944, -4.327163889, 10 May 2017, S. Martín-Bravo & C. Benítez-Benítez 41SMB17 (holotype!: UPOS-8925, 41SMB17(5); isotypes!: GDA, JAEN, K, MA, NY, P, SALA and UPOS).
(paratypes): Spain. Ciudad Real: Fuencaliente, Cereceda stream, with Alnus lusitanica, 695m, 38.42363889, -4.297472222, 10 May 2017, S. Martín-Bravo & C. Benítez-Benítez 47SMB17 (UPOS-8927); Fuente del Almírez, puddled meadows in Quercus pyrenaica forests, 800 m, 38.47152778, -4.344888889, 10 May 2017, S. Martín-Bravo & C. Benítez-Benítez 48SMB17 (UPOS-8926); Minas del Horcajo, gorge of Nacedero stream with Salix sp., 729 m, 38.51397222, -4.445750000, 11 May 2017, S. Martín-Bravo & C. Benítez-Benítez 54SMB17 (UPOS-8922); Solana del Pino, Robledillo River, riparian forests with Alnus lusitanica and Fraxinus angustifolia, 453 m, 38.41783333, -4.003388889, 11 May 2017, S. Martín-Bravo & C. Benítez-Benítez 57SMB17 (UPOS-8924); Solanilla del Tamaral, gorge of Jandula River (Hoz del Jándula) with Alnus lusitanica, 392 m, 38.39186111, -3.96333333, 11 May 2017, S. Martín-Bravo & C. Benítez-Benítez 59SMB17 (UPOS-8923); Valdemancos de Esteras, riverside of Esteras River with Fraxinus angustifolia and Salix sp., 453 m, 38.90733333, -4.794222222, 15 May 2018, C. Benítez-Benítez & S. Martín-Bravo 6CBB18 (UPOS-16896); Between Viso del Marqués and San Lorenzo de Calatrava, riverside of Ballesteros stream, in open forest of Quercus pyrenaica and Salix sp., 872 m, 38.44450000, -3.740444444, 16 May 2018, C. Benítez-Benítez & S. Martín-Bravo 8CBB18 (UPOS-16897); Puebla de Don Rodrigo, Sala del Halconcillo stream, 559 m, 39.10527778, -4.744722222, 15 June 2022, M. Sanz-Arnal, P. García-Moro & P. Jiménez-Mejías 13MSA22 (UPOS-16898); Horcajo de los Montes, Chorrera de Horcajo, 642 m, 39.36111111, -4.614999999, 15 June 2022, M. Sanz-Arnal, P. García-Moro & P. Jiménez-Mejías 16MSA22 (UPOS-16899); Viso del Marqués, Las Hoces, 780 m, 38.42303569, -3.721722958, 16 May 1991, C. Fernández García-Rojo (JAEN-914254); Puebla de Don Rodrigo, Río Frío mountain range near a birch forest, 600 m, 39.08243815, -4.503007649, 9 May 1992, Carrasco, Garrido & Martín-Blanco (MACB-68849); Solana del Pino, valley of Nacedero stream, 590 m, 38.482505309, -4.169451970, 26 April 1997, R. García Río (MA-596319); Hinojosas, valley of Montoro River with Alnus glutinosa, Cervigón, 700 m, 38.491048629, -4.215454806, 5 May 1997, R. García Río (MA-596320); Almodóvar del Campo, Guadalmez River with Alnus glutinosa, 510 m, 38.513064049, -4.628724543, 4 July 1997, R. García Río (MA-596324). Jaén: Andújar, Sierra Quintana, Valmayor River, 570 m, 38.37460677, -4.144817294, 27 June 1985, E. Cano & C. Fernández García-Rojo (JAEN-855238).
(see Suppl. material
Rhizomes from dense and tussock-forming to elongated. Stems (48)60–80(95) cm long, (0.8)1–1.2(1.3) mm wide below the inflorescence, sharply trigonous, smooth for most of its length, densely scabrid above. Basal sheaths scale-like, sometimes elongated, creamy-yellow, rarely reddish-brown, coriaceous. Leaves (1.8)2–3.2(4.7) mm wide, pale green to bluish-green, amphistomatic, flat to keeled, usually shorter or equalling the stems, antrorsely scabrid on the margins, especially distally, and on the mid-vein on the abaxial side to the apex; ligule (2)3–7(10) mm long, usually more than twice as long as wide, apex acute to obtuse, rarely rounded or truncate, hyaline, the margins brownish to orange-brown. Inflorescence (9)12–19(22) cm long, all spikes erect, exceptionally the lowermost slightly nodding. Lowest bract leaf-like, very rarely setaceous, longer to shorter than the inflorescence, sheathless, with hyaline, pale brownish to dark purplish-brown auricles at its base. Terminal male spikes (1)2–3; the uppermost spike (15)20–60(85) × 2–3.5 mm, subsessile to long pedunculate, oblong-cylindrical to narrowly fusiform, densely flowered; subterminal male spikes (0)1–2, (5)10–30(33) × (1)1.5–2.3(2.6) mm, similar in outline to the terminal one, basally overlapping with it or with an internode up to 20 mm. Lateral spikes female or androgynous with the flowers spirally arranged; female spikes 0–2(3), (19)25–50(79) × (2)2.8–4(5.2) mm, cylindrical, densely to more or less laxly flowered proximally; androgynous spikes (0)1–3(4), (9)20–50(62) × (1)1.7–2.7(5) mm, with the male portion (1)3–15(50) mm long and the female one (6)10–40(50) mm long. Male glumes (1.9)2.3–3.9(4.1) × (0.6)0.7–1.2(1.4) mm, oblong to obovate-oblong, apex rounded, usually dark purplish brown, with a green, 1-veined central band, with or without whitish hyaline margins, sometimes also becoming hyaline towards the base. Female glumes 1.3–2.3(2.5) × (0.6)0.7–0.9(1) mm, lanceolate to ovate, rarely elliptical, apex obtuse, acute or mucronate, usually shorter and narrower than the utricles, very rarely shortly exceeding them, dark purplish brown, rarely pale brown, with a green, 1–3 veined central band, with or without whitish hyaline margins, mainly in the distal part. Utricles (1.8)2–2.7(3) × (1.1)1.3–1.8(2) mm, plano-convex, widely elliptical to almost suborbicular, green to straw-coloured, distally whitish, sometimes purplish-dotted or purplish-tinged towards the apex, with high, whitish papillae on the upper half or towards the apex, sometimes also aculeolate at the upper margins, faintly to conspicuously nerved, shortly stipitate, more or less abruptly contracted into a short, cylindrical, truncate, more rarely emarginate beak (0)0.1–0.2(0.3) mm long, whitish, sometimes brown-tinged. Achenes 1.5–2.1 × (1.1)1.3–1.5(1.7) mm, widely elliptical to suborbicular, straw-coloured to pale brown, biconvex, more or less stipitate; style base terete to slightly conical, up to 0.3 mm. Stigmas 2.
(Fig.
Small streams, rivers and springs in riparian forests (Alnus lusitanica, Fraxinus angustifolia, Salix spp.), and humid meadows, in marcescent and sclerophyllous Quercus forests, on siliceous bedrock substrate. 400–800 m.
(April) May-June (July).
2n=82, 83.
So far known from 16 populations (subpopulations according to IUCN; Fig.
The species epithet, quixotiana (pronounced kee·how·tee·a·na in English) is derived from Miguel de Cervantes’s (1547–1616) masterpiece Don Quixote (
Carex quixotiana populations have been consistently assigned to C. reuteriana. Local floristic studies have predominantly identified them as C. reuteriana subsp. reuteriana (e.g.,
These problematic populations clearly illustrate the taxonomic complexity of sect. Phacocystis. Their overall morphological appearance and ecological preferences suggested that the populations were conspecific with C. reuteriana (
The chromosome number has been used as a biosystematic tool for sect. Phacocystis in the Iberian Peninsula (
Considering the new species described here, the Iberian Peninsula has 12 endemic Carex taxa (C. asturica Boiss., C. camposii subsp. camposii, C. camposii subsp. tejedensis R. Sánchez-Villegas, M.Escudero & Luceño, C. caudata (Kük.) Pereda & Laínz, C. durieui Steud. & Kunze, C. furva Webb, C. lainzii Luceño, E.Rico & T.Romero, C. lucennoiberica Maguilla & M. Escudero, C. nevadensis Boiss. & Reut., C. quixotiana, C. reuteriana ssp. reuteriana and C. rorulenta Porta). However, this may appear as a relatively small number of endemics in proportion to the total number of Carex Iberian native taxa (108; Luceño et al. unpublished data), especially when compared with other species-rich genera (e.g., Alchemilla, Armeria, Centaurea, Limonium, Teucrium) with much larger numbers of Iberian endemics (>40 each;
As explained above, known populations of C. quixotiana are mostly located in La Mancha limits, in Sierra Madrona and, to a lesser extent, southern Montes de Toledo. Sierra Madrona is one of the secondary mountain ranges within the larger Sierra Morena range, which stretches for about 450 kms from W to E across the S Iberian Peninsula, separating the southern half of the Central Plateau from the Guadalquivir Valley. Sierra Madrona runs almost in parallel (NW–SE) along the northern side of the main range for about 80 kms. It includes the highest altitudes of all Sierra Morena (Bañuela peak, 1332 m). As the whole Sierra Morena, it is mainly composed by old Paleozoic siliceous rocks, especially quartzites. On the other hand, Montes de Toledo is a mountain range entirely belonging to the southern half of the Central Plateau, separating the Tajo and Guadiana River basins. It stretches for about 350 kms from E to W and is composed by quartzite ridges of relatively uniform elevation, around 1400 m, and a generally eroded relief (Muñoz Jiménez 1976). The immediate foothill landscapes are highly anthropised, with the vast majority of La Mancha dedicated to non-irrigated crops, mostly cereal (wheat, barley and oat), and vineyards.
Sierra Madrona is currently protected as a natural park established in 2011 (Valle de Alcudia and Sierra Madrona). Several restricted plant endemics from Sierra Morena are found in this range and its surroundings (i.e., Armeria pauana (Bernis) Nieto Fel., Coincya longirostra (Boiss.) Greuter and Burdet;
The other known populations of C. quixotiana occur mostly in rivers and creeks in the southern foothills of Montes de Toledo in W Ciudad Real province. These populations are located within or close to remarkable habitats, such as relictual peatbogs (e.g., Bonales de Puebla de Don Rodrigo) and birch forests (e.g., Abedular de Ríofrío), which are on protected land and considered to be of extraordinary conservation value. In addition, C. quixotiana marginally reaches NE Jaén province in Sierra Quintana, a small southern subrange of Sierra Madrona province (
1 | Leaves strongly plicate to canaliculate, rigid, up to 2.5(3) mm wide; stems obtusely trigonous; plants bluish-green | C. trinervis |
– | Leaves flat to carinate, soft to medium rigid, (1.8)2–8(10) mm wide; stems sharply trigonous; plants green to bluish-green | 2 |
2 | Utricles not veined; basal sheaths dark purple | C. cespitosa |
– | Utricles faintly to prominently veined; basal sheaths yellowish-brown to reddish-brown | 3 |
3 | Utricles smooth, rarely with some scattered, green, low papillae | 4 |
– | Utricles densely papillose, at least on the apex, with whitish, more or less high papillae | 5 |
4 | Male spike usually solitary, rarely 2; lowest spikes usually female; leaves up to 6 mm wide | C. reuteriana subsp. reuteriana |
– | Male spikes (1)2–4; lowest spikes usually androgynous; leaves (3)4–8(10) mm wide | C. reuteriana subsp. mauritanica |
5 | Leaves epistomatic or amphistomatic | 6 |
– | Leaves hypostomatic | 8 |
6 | Male spikes (1)2–3, the terminal one (15)20–60(85) mm long; leaves (1.8)2–3.2(4.7) mm wide; plants usually tussock-forming; basal sheaths scale-like, sometimes elongated, creamy yellow, rarely reddish-brown | C. quixotiana |
– | Male spike solitary, rarely 2, the terminal one (5)10–30 mm long; leaves 3–6(10) mm wide; plants rarely tussock-forming; scale-like basal sheaths absent or dark brown when present | 7 |
7 | Leaves densely stomatic on both sides (amphistomatic) | C. nigra subsp. intricata |
– | Leaves epistomatic or densely stomatic on the upper surface and with a few scattered stomata on the lower surface | C. nigra subsp. nigra |
8 | Lowest bract much longer than the inflorescence; utricles strongly biconvex, somewhat inflated; culm bases bearing brown old leaf remains, without conspicuous scale-like basal sheaths | C. acuta |
– | Lowest bract always shorter than the inflorescence; utricles plano-convex to slightly biconvex, not inflated; basal sheaths scale-like to elongated, creamy-yellow | C. elata subsp. elata |
Integrative approaches based on different sources of evidence are required to unveil complex systematic scenarios. Our study of some problematic Iberian populations belonging to the taxonomically difficult sect. Phacocystis revealed that they display a congruent set of diagnostic morphological, molecular, ecological and cytogenetic characters that allow their distinction with respect to close relatives (e.g., C. reuteriana, C. nigra). Therefore, we here described a new species to science (C. quixotiana) for these populations, which is endemic to the Iberian Peninsula.
C. Salazar (Univ. Jaén), M. Sanz-Arnal, P. García-Moro (Univ. Autónoma de Madrid), F. García de Leániz, and C. Fernández García-Rojo for help during fieldwork; R. Sánchez-Villegas, B. Quirós de la Peña, R. Medina (Univ. Complutense de Madrid) for helping us in the cytogenetic and/or herbarium work; and M. Sánchez-Villegas for the analytic drawing of C. quixotiana. The curators and staff of JAEN, MA, MACB and SALA herbaria for the loan or providing photos of critical material, and of UPOS herbarium, especially C. Barciela and P. Pineda, for technical support during the processing of herbarium specimens. Project DANZ (Ref. PID2020-113897GB-I00, Spanish Ministry of Science and Innovation) towards P. Jiménez-Mejías and S. Martín-Bravo, and PAIDI BIO-305 research group, are acknowledged for financial support.
List of studied material of C. quixotiana, including the 16 known populations highlighted in red
Data type: occurrence data (excel document)
List of specimens newly sequenced in the present study
Data type: Occurrence data (excel document)
Explanation note: Including taxon, sample code, locality, voucher and/or herbarium number, and NCBI accession numbers for each molecular marker (ITS, ETS, rpl32‐trnLUAG, ycf6‐psbM).
List of studied material, with their accession numbers included, which were previously published by
Data type: occurrence data (excel document)
Explanation note: Tip codes with “spm” denote samples previously published by Global Carex Group works (
Cladogram of sect. Phacocystis, including newly sampled sequences
Data type: phylogenetic tree (PDF file)
Explanation note: The cladogram is based on the maximum likelihood phylogenetic reconstruction of complete singleton’s tree (ITS, ETS, rpl32-trnL, ycf6-psbM) following a scaffolding approach previously implemented by
List of studied material for haplotype network analysis, displaying the corresponding haplotype number obtained for each sample
Data type: table (excel document)
Haplotype network obtained from the statistical parsimony analysis of the combined plastid sequences of Iberian Carex sect. Phacocystis species, including C. quixotiana.
Data type: image (PDF file)
Explanation note: Small circles represent inferred extinct or not sampled haplotypes and lines mutational connections.