Carex quixotiana Ben. Benitez , Martin-Bravo , Luceno & Jim. Mejias, 2023

Benitez-Benitez, Carmen, Jimenez-Mejias, Pedro, Luceno, Modesto & Martin-Bravo, Santiago, 2023, Carex quixotiana (Cyperaceae), a new Iberian endemic from Don Quixote's land (La Mancha, S Spain), PhytoKeys 221, pp. 161-186 : 161

Carex quixotiana Ben. Benitez, Martin-Bravo, Luceno & Jim. Mejias sp. nov.

Diagnosis.

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).

Type.

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) .

Selected material examined

(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 Almirez , 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 Jandula ) 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 Marques 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 Marques , Las Hoces, 780 m, 38.42303569, -3.721722958, 16 May 1991, C. Fernández García-Rojo (JAEN-914254); Puebla de Don Rodrigo , Rio Frio 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); Almodovar 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) .

Other material.

(see Suppl. material 1).

Morphological description.

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.

Distribution.

(Fig. 1 ) Endemic to South-Central Spain (Ciudad Real and marginally Jaén provinces). So far known from 16 populations (Suppl. material 1), mostly located in the Sierra Madrona range, but also extending north reaching the southern foothills of the Montes de Toledo range. Since it is a medium-size sedge rarely collected, there might be additional populations in these areas.

Habitat.

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.

Phenology.

(April) May-June (July).

Chromosome number.

2n=82, 83.

Iconography.

Fig. 5 .

Conservation status.

So far known from 16 populations (subpopulations according to IUCN; Fig. 1 ), deduced from the studied material (Suppl. material 1). This implies a relatively restricted distribution range enclosed in an extent of occurrence (EOO) of 4.920 km2, and an area of occupancy (AOO) of just 64 km2. This would point to the application of criteria B1 and B2 of the Endangered (EN) category (threshold of 5.000 km2 and 500 km2 for EOO and AOO, respectively; IUCN 2017). The species’ overall demographic tendency and number of mature individuals are unknown. In addition, the number of locations (16; n> 5) prevents the application of EN category since two conditions of criteria B must be fulfilled. Some populations are located in protected land (see below) so they should not be likely submitted to plausible threats in the short term (but see García Río 2007). With the currently available data, and taking into account the restricted EOO and AOO, we hypothesize that C. quixotiana 's formal IUCN conservation category at the global level would be Data Defficient (DD). As it is an Iberian endemic with a relatively small number of populations and distribution range, it would benefit from legal protection and inclusion in in-situ/ex-situ conservation programmes, at least at the regional level (Castilla-La Mancha, Andalucía).

Etymology.

The species epithet, Carex quixotiana (pronounced kee·how·tee·a·na in English) is derived from Miguel de Cervantes’s (1547-1616) masterpiece Don Quixote ( de Cervantes 1605, 1615), globally considered one of the best works in the history of literature, and whose number of editions and translations is only surpassed by the Bible. The setting of Don Quixote is La Mancha, the region of Spain where almost all populations of Carex quixotiana occur. We would like this epithet to serve as a double tribute: (1) First to Cervantes and his novel Don Quixote, flagship of Spanish culture. And (2) To Pedro Jiménez-Mejías’s father, Pedro Jiménez Gilabert, an enthusiastic reader who always enjoys reading Don Quixote above all other books, and who always transmitted Pedro’s curiosity and love for nature. In contrast to the first words in Don Quixote, ("En un lugar de La Mancha, de cuyo nombre no quiero acordarme (...)" [In a village of La Mancha, the name of which I have no desire to call to mind ( …)]), we desire to remember and commemorate the dedication of this epithet we are coining.

Systematic notes.

Carex quixotiana populations have been consistently assigned to C. reuteriana . Local floristic studies have predominantly identified them as C. reuteriana subsp. reuteriana (e.g., García Río 2004, 2007; Martín Blanco and Carrasco 2005; Fernández García-Rojo 2015; Fernández García-Rojo and Salazar-Mendías 2019), but also as C. reuteriana subsp. mauritanica ( García Río 1999, 2004). Specialized taxonomic treatments have considered the populations as morphologically intermediate between the two C. reuteriana subspecies, which has been attributed to hybridisation in a putative contact zone between the two C. reuteriana subspecies ( Luceño and Jiménez-Mejías 2008).

These problematic populations clearly illustrate the taxonomic complexity of sect. Carex Phacocystis . Their overall morphological appearance and ecological preferences suggested that the populations were conspecific with C. reuteriana ( Luceño and Jiménez-Mejías 2008). On the other hand, phylogenetic relationships (Fig. 2 ) and the relatively high chromosome number (see below) indicate closer genetic affinity with C. nigra . These misleading patterns denote the necessity of integrative approaches comprising different data sources to unmask complex systematic scenarios. Further phylogenomic studies based on High-Throughput-Sequencing techniques like GBS might provide a greater number of loci and therefore might help to shed more light on the systematic placement of C. quixotiana ( Benítez-Benítez et al. unpublished data).

The chromosome number has been used as a biosystematic tool for sect. Carex Phacocystis in the Iberian Peninsula ( Luceño and Aedo 1994; Luceño and Jiménez-Mejías 2008). Our chromosome counts indicate that C. quixotiana (2n=82-83) has a greater cytogenetic affinity with C. nigra (2n=(80)82-86) than with C. reuteriana (2n=(72)73-76).

Biogeographic and conservation issues.

Considering the new species described here, the Iberian Peninsula has 12 endemic Carex taxa ( C. asturica Boiss., C. camposii Boiss. & Reut. 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; Aedo et al. 2017; Buira et al. 2020). This highlights the novelty of the description of an additional Iberian endemic in Carex . Indeed, C. quixotiana , with a highly restricted distribution in southern-central Spain, would be the first Iberian endemic species belonging to sect. Carex Phacocystis , although one subspecies ( C. reuteriana ssp. reuteriana ) is also endemic to C-NW parts of the Iberian Peninsula. Other species of sect. Carex Phacocystis with limited distribution range in the Mediterranean basin are C. reuteriana ssp. mauritanica (S Iberian Peninsula-NW Africa), C. nigra ssp. intricata (Western Mediterranean mountains), and C. panormitana Guss. (Sicily, Sardinia and Tunisia).

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; García Río 2004). It also includes many different habitats protected at European level (Directive “Habitats” 92/43/CEE; see García Río 2004, 2007), including some considered of high conservation priority. Carex quixotiana habitats in Sierra Madrona, although included in a protected area, are not free of threats, so their biodiversity has been considered as highly valuable and sensitive ( García Río 2007). They include populations of other endangered and protected plant species at regional (e.g., Erica tetralix L., Drosera rotundifolia L.) or even at national level (e.g., Myrica gale L., Narcissus muñozii-garmendiae Fern.Casas).

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 ( Cano-Carmona and Valle-Tendero 1996), which separates Castilla-La Mancha from Andalucía (Fig. 1 ). Sierra Madrona and Montes de Toledo are considered biogeographically close ( García Río 2007). Interestingly, C. reuteriana subsp. reuteriana occurs in northern parts of Montes de Toledo, whereas C. reuteriana subsp. mauritanica is widespread in the main Sierra Morena range, therefore flanking C. quixotiana 's distribution ( Luceño and Jiménez-Mejías 2008; see maps in Benítez-Benítez et al. 2018). It would be interesting to precisely delimit these species’ range and to explore potential contact zones in detail in order to investigate the possible co-occurrence and/or hybridisation processes.

Figures

Figure 1. Distribution map representing the studied populations of Carex quixotiana.

Figure 2. Representative photos of Carex quixotiana A habitat (riparian forest with Alnus lusitanica and Fraxinus angustifolia: Ciudad Real, Solana del Pino, Robledillo River; 57 SMB 17, 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 Marques and San Lorenzo de Calatrava; 8 CBB 18, 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, 54 SMB 17, UPOS- 8922).

Figure 5. Analytical drawing of Carex quixotiana (Spain, Ciudad Real, Sierra Madrona) A habit B plant C androgynous spike D female glume E male glume F achene G utricle H details of papillae in the upper part of the utricle. Drawing by M. Sanchez-Villegas.