Linaria Argillicola (Plantaginaceae), a New Species of L

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Phytotaxa 343 (2): 127–138 ISSN 1179-3155 (print edition) http://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2018 Magnolia Press Article ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.343.2.3

Linaria argillicola (Plantaginaceae), a new species of L. sect. Supinae from the southern Iberian Peninsula

ANA JUAN1, GABRIEL BLANCA2*, MIGUEL CUETO3, JULIÁN FUENTES4 & LLORENÇ SÁEZ5 1Departamento de Ciencias Ambientales y Recursos Naturales, Universidad de Alicante, PO Box 99, ES-03080 Alicante, Spain; email: [email protected] 2Departamento de Botánica, Facultad de Ciencias, Universidad de Granada, C/ Fuentenueva s/n, ES-18001 Granada, Spain; email: [email protected] 3Departamento de Biología y Geología, CECOUAL, Universidad de Almería, ES-04120 Almería, Spain; email: [email protected] 4C/ Castillo 5, bajo F, ES-18140 La Zubia, Granada, Spain; email: [email protected] 5Unitat de Botànica, Facultat de Biociències, Universitat Autònoma de Barcelona, ES-08193, Bellaterra, Barcelona, Spain; email: [email protected] *author for correspondence

Abstract

A new species of the genus Linaria is described, illustrated and compared with its morphologically closest relatives from L. sect. Supinae: L. accitensis, L. aeruginea, L. tristis and L. badalii. A principal component analysis (PCA), correspondence analysis (CA), and linear discriminant analysis (LDA) were carried out for morphological differentiation. The species is characterized by long abaxial sepals, corolla mostly yellow or rarely reddish with wide tube, wide winged seeds with a tuberculate seminal disc, and a non-continuous fruiting inflorescence. Linaria argillicola is an edapho-endemic species, growing on the marly gypsiferous deposits from the Guadiana Menor river basin, on the border of Granada and Jaén provinces (Andalusia, Spain).

Key words: Antirrhineae, endemic plants, Spain, taxonomy

Introduction

Linaria Miller (1754) is the largest genus (c. 150 species) within the tribe Antirrhineae Dumortier (1827: 34) (Plantaginaceae sensu APG IV 2016). This genus includes annual or perennial herbs, mostly with heteromorphic stems; flowers arranged in bracteate racemes, with calcarate personate corollas; sepals equal or unequal, fused at their base; ovary bilocular, style simple or bifid; capsules ovoid to globose, dehiscent by 3–5 valves; seed morphology variable (discoidal, trigone, kidney-shaped), with the presence or absence of a marginal encircling visible wing (Sáez & Bernal 2009). It is highly diversified around the Mediterranean basin, especially L. sect Supinae (Bentham 1846: 280) Wettstein (1891: 59) (cf. Sutton 1988, Blanco-Pastor et al. 2012). This section comprises hermaphroditic annual and perennial plants differentiated from other sections by their laterally compressed winged seeds and horizontal arrangement of globose capsules. It is the most complex taxonomic section of the genus and the Iberian Peninsula is considered the centre of diversity with 44 species (Sutton 1988). According to Blanco-Pastor et al. (2012) this section is monophyletic and corolla size, seed wing shape, and life form were treated as diagnostic characters for winged-seeded Linaria species (Valdés 1970, Blanco-Pastor et al. 2012). Among the three subsections recognised by Blanco-Pastor et al. (2012), L. sect. Supinae subsect. Supinae includes perennial to rarely annual herbs, with membranous seed wings. Saéz & Bernal (2009) mentioned 46 taxa (including subspecies) for the Iberian Peninsula and the Balearic Islands, of which 18 taxa are found in Eastern Andalusia (Saéz & Sainz 2011). Many of these taxa have been described as endemic with narrow southern Iberian distributions such as L. accitensis L. Sáez, Juan, M.B. Crespo, F.B. Navarro, J. Peñas & Roquet (in Sáez & Bernal 2009: 623), L. saturejoides Boissier (1841: 463), L. verticillata Boissier (1841: 462) sensu lato, L. glacialis Boissier (1838: 70) and L. aeruginea subsp. nevadensis (Boissier) Malagarriga (1978: 1428), among others (Valdés 1970, Sutton 1988, Sáez & Crespo 2005, Sáez & Bernal 2009, Sáez & Sainz 2011).

Accepted by Manuel B. Crespo: 18 Jan. 2018; published: 9 Mar. 2018 127 Our fieldwork in Eastern Andalusia (southern Spain), specifically around the area of Guadix basin (Granada province) revealed several populations of Linaria of unknown identity growing in marly gypsiferous areas. An initial identification, based on Flora iberica (Sáez & Bernal, 2009), led us to consider this plant as the northern Iberian species L. badalii Loscos y Bernal (1885: 58), but this latter species had not been previously confirmed in Andalusia (Sáez & Bernal 2009, Sáez & Sainz 2011) so far. Furthermore, a detailed comparison with closely related taxa of L. sect. Supinae subsect. Supinae revealed constant and conspicuous morphological differences in both vegetative and floral features. The micromorphology of seeds also showed differences for these plants, thus supporting its taxonomic distinctiveness. The combination of morphological and seed-coat characters with diagnostic value, and the isolated distribution of these Linaria populations suggest that the plants belong to a different taxon, which is named here L. argillicola. Data on its morphological characteristics, distribution, and habitat are discussed with regard to other closely related Linaria taxa.

Material and methods

Morphological observations and measurements of vegetative and reproductive parameters were undertaken on plant material kept in the herbaria ABH, BC, GDA, MA, SEV, and VAL (acronyms according to Thiers 2017), as provided in Annex 1. For the morphological observations, an Olympus SZX12 binocular microscope was used with objective DFPL-1X-PF and incorporated micrometre, and seeds were metalized for micrographs to be taken with a GEMINI (FESEM) CARL ZEISS Scanning Electron Microscope. For taxonomic identification, Spanish floras and specific scientific works were revised (Sutton 1988, Sáez & Bernal 2009, Sáez & Sainz 2011). Diagnostic characters reported by Sutton (1988) and Sáez & Bernal (2009) and other key features based on our field experience were included. The descriptive terminology follows Sutton (1988). For statistical studies, a preliminary principal component analysis (PCA) was firstly carried out to identify those vegetative and floral characters that have the highest potential to differentiate the closest species, i.e. L. accitensis, L. aeruginea (Gouan 1773: 38) Cavanilles (1803: 21) subsp. aeruginea, L. tristis (Linnaeus 1753: 613) Miller (1768: 8) subsp. tristis and L. badalii plus the newly described species. Subsequently, 13 morphological characters were analysed in detail, nine of wich were quantitative (Table 1). For 96 specimens (Annex 1), the morphometric data were scored and subsequently used for analysis. The specimens were chosen among those showing all the characters to be measured. Five randomly selected leaves, flowers, capsules, and seeds from fertile stems were measured and the average values used. A principal component analysis (PCA), correspondence analysis (CA), and linear discriminant analysis (LDA) were applied to the morphological data. Prior to the multivariate analyses, characters not satisfying the assumption of normal distribution were log-transformed. PCA and CA analyses were conducted in order to visualize the multivariate morphometric character distribution among the taxa using both qualitative and quantitative characters. The components found to have the highest variance were plotted along two axes. In addition, an LDA was performed also to determine a classifier for the morphological distinction among the geographical coexisting taxa (L. badalii not included), using continuous variables, and the classification was evaluated by cross validation. Statistical analyses were performed using the package PAST v. 3.14 (Hammer et al. 2001).

TABLE 1. Comparison of Linaria argillicola with its closest relatives in L. sect. Supinae. (Abbreviations: fl.: flower; fr.: fruit). Linaria argillicola Linaria accitensis Linaria badalii Linaria aeruginea Linaria tristis subsp. aeruginea subsp. tristis Habit Perennial Perennial Annual (rarely Perennial (rarely Perennial perennial) annual) Fertile stems Number 4–13 10–20 1–24 1–10 1–16 Length (cm) 7–25(–31) 8–22 6–40 6–70 9–50 Leaves (fertile stems) Size (mm) 6.5–20(–23) × 0.3– 7–22(–25) × 0.2–1.2 3–22 × 0.5–1.5(–2) 4.5–25(–38) × 0.4–1.5 6–40 × 1–3(–4) 1.2(–1.4) ...continued on the next page

128 • Phytotaxa 343 (2) © 2018 Magnolia Press JUAN et al. TABLE 1. (Continued) Linaria argillicola Linaria accitensis Linaria badalii Linaria aeruginea Linaria tristis subsp. aeruginea subsp. tristis Arrangement alternate, sometimes alternate, lowermost verticillate 3–5 up to lowermost verticillate, generally unilateral, lowermost verticillate verticillate 3–4 half of stems, superior generally unilateral lowest verticillate 3–4 alternate 4–6 Inflorescence Axis glabrous (rarely with glabrous (sometimes glabrous, glaucous pilose-glandulose pilose-glandulose scarce and small with small glanduliferous hairs) glanduliferous hairs) Hair length 0.1–0.2 0.1–0.2 - 0.3–1.2 0.3–0.8 (mm) Nº flowers/ 3–15(–18) 5–16 2–30 3–34 2–25 inflorescence Length (2–)3–11 3.5–7 6–26 5–21 2.5–10 inflorescence in flower (cm) Inflorescence Simple or branched Simple, sometimes Simple Simple Simple branching branched Size bract (1–)1.5–5.8 × 0.1–0.9 1.5–6(–8.6) × 0.1–0.7 2–8 × 0.4–1 2.5–10 × 0.5–1.2 3–9 × 1–1.7 (mm) Flower (measures in mm) Pedicel length 0.3–1.8(–2.3) 0.4–2.5 0.5–2.5 (0.3–)0.5–2.5(3) 0.5–4.0 fl. Pedicel length 0.6–3.5(–4.5) 0.8–3.5 0.5–4 1–4 2.5–6 fr. Calyx adaxial 4–9 × 0.6–1 5.5–8 × 0.7–1.5 2.5–6 × 0.4–0.9 3.5–7.5(–9) × 0.5–1.2 5–7.8 × 0.6–1.1 lobe fl. Calyx abaxial 3–6 × 0.5–1 2.5–4.5 × 1–2.3 2.5–5.5 × 0.4–1 2.5–5 × 0.6–1.9 4–5.5 × 1.2–1.6 lobe fl. Calyx adaxial 5–10 × 0.5–12 7–9 × 1–2 3–6.5 × 0.5–1 4–8.5(–10) × 0.5–1.7 7.5–9 × 1.2–1.6 lobe fr. Calyx abaxial 4–9 × 0.5–2 4–6 × 1–2 3–6 × 0.6–1.1 3–6 × 0.7–2 5–6.5 × 1.3–2.8 lobe fr. Corolla colour mostly yellow or white with reddish, white, whitish-pinkish, red or purple to yellow, whitish or orangish, rarely violet veins, (palate yellowish, purple yellow (palate white-yellowish reddish yellow, sometimes veins (palate yellow to orangish or reddish) purple) orangish) Corolla length (12–)14.5–25 (7–)10–23 12–22 15–26(–29) 18–24 Width tube 3–6.5 2.5–5 2.2–4 (3–)4–6(–7) 6–6.7 Spur length (4.5–)5–10 3.2–9 5–9 (4.5–)6–11.5 8.5–11.5 Spur width 0.8–2(–2.3) 0.4–2.4 1–2 (0.5–)0.9–3(4) 2–3 Capsule Size (mm) 3.2–7.1 × 3–7.2 2.4–6 × 2.5–6 3–5 × 3–5 3.5–7 × 4–7 3.5–8 × 3.4–8 Indument glabrous glabrous glabrous glabrous or hairy glabrous or hairy Seed Size (mm) 1.8–3.2 × 2.1–3.8 1.6–2.6 × 1.2–2.5 1.5–2.4 × 1.5–2.3 2.2–3 × 2–3 2.4–3 × 2.3–2.7 Disc surface tuberculate smooth papillose tubeculate or smooth tuberculate or smooth Wing width 0.4–1.4 0.3–0.8 0.3–0.6 0.5–1.2 0.6–1 (mm) Habitat marl-gypsiferous volcanic soils (andesites calcareous, siliceous, calcareous soils limestone and no- susbtrates and alkaline basalts) marly, gypsiferous calcareous basic soils substrates

Linaria argillicola Juan, Blanca, Cueto, J. Fuentes & L. Sáez, sp. nov. (Figs. 1–3) Type:—SPAIN. Granada: Dehesas de Guadix, Collado de la Higuera, laderas abruptas de litología margosa con yesos, en suelos poco evolucionados, 30SVG9159, 745 m, 21 May 2011, J. Fuentes (holo-: GDA 62646!).

Diagnosis:—It differs from L. accitensis L. Sáez et al. in having flowers with abaxial calyx lobes longer (up to 6 and 9 mm long in flower and fruit, respectively), corolla mostly yellow or orangish, rarely reddish, larger (up to 25 mm long, including the spur), and with a wider tube (3.0–6.5 mm width); seeds larger (1.8–3.8 × 2.1–3.8 mm), with a wider wing (up to 1.4 mm broad), and tuberculate disc; and inflorescence non-continuous in fruiting stage. Description:—Herbaceous perennial (rarely annual) plant, glaucous, glabrous; fertile stems, up to 14–25(–31) cm long, decumbent to suberect, simple or sometimes branched in the upper part; sterile stems, up to 6–7 cm long. Leaves of fertile stems 6.5–20.0 × 0.3–1.2(–1.4) mm, linear, narrowly oblong, flat, acute (sometimes obtuse), alternate, sometimes the lowermost in whorls of 3–4; leaves of sterile stems (3.0–)5.0–12.0 × 0.5–1.0 mm, similar to the leaves of fertile stems, flat, acute (sometimes obtuse), mostly alternate, sometimes basal leaves in whorls of 3–4. Inflorescence simple or clearly branched, (2–)3–11 cm long [2–7(–8.5) cm long in fruit], with 3–15(–18) flowers, glabrous (rarely with sparse hairs 0.1–0.2 mm long on the axis); aborted fruits sometimes appearing among well-developed flowers and fruits in the same inflorescence. Bracts (1.0–)1.5–5.8 × 0.1–0.9 mm, oblanceolate or oblong, acute or obtuse, glabrous. Pedicels 0.3–1.8(–2.3) mm long in flower [0.6–3.5(–4.5) mm long in fruit], erect. Calyx lobes unequal, glabrous, obovate, oblong-obovate or oblanceolate, acute or subacute in flower, oblong, obtuse in fruit; adaxial lobe 4.0–9.0 × 0.6–1.0 mm in flower (5.0–10.0 × 0.5–12.0 mm in fruit); abaxial lobes 3.0–6.0 × 0.5–1.0 mm in flower (4.0–9.0 × 0.5–2.0 mm in fruit). Corolla (12.0–)14.5–25.0 mm long, mostly yellow or orangish, rarely reddish; tube 3.0–6.5 mm broad in dorsiventral section; adaxial lip sinus 4–6 mm; abaxial lip sinus 6–10(–12) mm; spur (4.5–)5.0–10.0 × 0.8–2.0(–2.3) mm, stout, straight or slightly curved at the apical part, shorter than or slightly subequalling the rest of the corolla. Capsule 3.2–7.1 × 3.0–7.2 mm, globose, subglobose or obovate-globose, glabrous. Seeds 1.8–3.2 × 2.1–3.8 mm, suborbicular, discoid, slightly concavo-convex to flat; wing 0.4–1.4 mm broad, dark greyish to brownish, membranous, entire; disc 0.7–1.8 mm, reniform, greyish to blackish-brown, densely tuberculate. Etymology:—The specific epithet refers to the type of substrate on which the new species typically grows. Distribution and ecology:—Only four populations of L. argillicola are known so far, including about 9000– 10000 reproductive individuals in total, with an extent of occurrence of 87 km2, and an occupancy area of 2.05 km2. This taxon would be considered an edapho-endemic species from the southern Iberian Peninsula (Fig. 4), growing on the marly gypsiferous deposits from the Guadiana Menor river basin, on the border of Granada and Jaén provinces. The populations appear on steep hillsides, between 600–800 m elevation, with poorly evolved soils of marls with gypsum deposits, belonging to the Keuper (Triassic origin). Linaria argillicola typically participates in ephemeral therophyte pastures characterized by scant plant biomass and cover. The populations are found under a xeric oceanic Mediterranean Ombroclime with a Mesomediterranean Thermotype and semiarid Ombrotype (Marchal et al. 2011). Phenology:—The flowering period occurs from mid-March to early June, and fruiting time from April to the end of June. Conservation status:—The distribution area of L. argillicola is not included within any Site of Community Importance (SCI) and hence remains outside any protected areas of the Nature 2000 Network. Its natural habitat shows remarkable environmental stochasticity, with strong interannual fluctuations during different phenological states. Potential causes of threat include the erosion and disturbance of the habitat; the low environmental valuation of gypsiferous and semiarid sites, since they are traditionally considered to be uncultivated lands, and thus apt for agricultural and forestry uses; the cattle overload in critical drought years; the direct predation of the individuals by coleopteran larvae (Fam. Chrysomelidae), which sometimes consume nearly entire populations; future urban expansion; and, finally, climatic change. According to the IUCN categories (2012) and recommendations provided by IUCN (2017), we suggest labelling L. argillicola as Endangered (EN), with the following criteria B1ac(i,ii,iv) + B2ac(i,ii,iv). Further monitoring of the evolution of the known populations is recommended to more accurately evaluate the conservation status of the newly described species. Linaria argillicola may require conservation actions and management plan, and therefore it should be included in the Spanish and Andalusian Red Lists of vascular plants (i.e. Cabezudo et al. 2005).

130 • Phytotaxa 343 (2) © 2018 Magnolia Press JUAN et al. FIGURE 1. Linaria argillicola (from holotype). A, Habit; B, leaves detail (fertile stems); C, flower lateral view; D, flower front view; E, capsule; F, seed front view; G, seed lateral view.

Linaria argillicola (Plantaginaceae) Phytotaxa 343 (2) © 2018 Magnolia Press • 131 FIGURE 2. Linaria argillicola (A, Habit; B, inflorescence detail; C, idem with red corollas; A‒B: GDA 62648; C: GDA 62649). Linaria accitensis (D; GDA 62645). Linaria badalii (E; GDA 62650). Linaria aeruginea subsp. aeruginea (F; Jaén province, Torredelcampo, G. Blanca personal collection of photographs). Linaria tristis subsp. tristis (G; Málaga province, Benaoján, G. Blanca personal collection of photographs).

132 • Phytotaxa 343 (2) © 2018 Magnolia Press JUAN et al. FIGURE 3. Scanning-electron micrographs of seeds of Linaria argillicola (A, abaxial view; B, adaxial view; GDA 62648), Linaria accitensis (C; GDA 62645), and Linaria badalii (D; GDA 62650).

FIGURE 4. Distribution map of the localities studied.

Linaria argillicola (Plantaginaceae) Phytotaxa 343 (2) © 2018 Magnolia Press • 133 Multivariate analyses Multivariate statistical analyses performed on all the Linaria material studied gave similar results, and hence only PCA data are presented (Fig. 5). The two first principal-component axes explained a total of 83.0% of the variance (74.8 and 8.2%, respectively). The taxa studied were separated from one another by certain floral and seed characters such as length of corolla and spur (both first and second axes), width of corolla (first axis), and seed-disc surface (second axis). The first axis separated between L. argillicola from L. accitensis, and L. badalii from L. aeruginea subsp. aeruginea-L. tristis subsp. tristis, whereas the second one differentiated L. argillicola-L. accitensis, L. aeruginea subsp. aeruginea- L. tristis subsp. tristis, and L. badalii. Among the species studied, L. badalii proved to be morphologically the most different, showing unique qualitative features such an annual habit, white-yellowish flowers and papillate seeds, and differs from the new species by its smaller flowers and seeds (Table 2).

FIGURE 5. First two axes of the principal component analysis (PCA) of the Linaria species studied. First and second principal components explain 74.8% and 8.2% of the variance, respectively.

TABLE 2. Eigenvalues of the principal component analysis (PCA) for the first (PC1) and second (PC2) components PC 1 PC 2 Width of corolla 0.2779 0.0116 Width of seed 0.0753 -0.1786 Width of leaf of fertile stems 0.0798 0.1818 Length of fruiting pedicels -0.0080 -0.1492 Length of corolla 0.8107 -0.4099 Length of spur 0.4765 0.7240 Length of seed 0.0520 -0.0774 Width of seed wing 0.0219 -0.0291 Length of hairs 0.0255 0.0393 Habit 0.0460 -0.2125 Pubescence 0.1026 0.0976 Main corolla 0.0972 -0.2189 Disc surface 0.0198 0.3308

134 • Phytotaxa 343 (2) © 2018 Magnolia Press JUAN et al. The LDA scattered plot displays noteworthy differentiation among the four geographically coexisting Linaria taxa studied (Fig. 6). The first axis (49.3% of variation) is most correlated with the length of spur, and the second (41.9% of variation) with the length of corolla. Cross-validated classification results of this analysis show that 98.5 % of the specimens that were assigned a priori to the four groups were classified correctly. The statistical analyses indicated that there is enough information in quantitative characters for their notable separation.

FIGURE 6. Linear discriminant analysis (LDA) of the four Linaria species studied in Eastern Andalusia.

Discussion: Taxonomic relationships

Morphological features such as the presence of a globose or ovoid-globose fruit, dehiscing from the apex to the half of its length or even more, plus discoid, reniform, and broadly winged seeds, strongly suggest that Linaria argillicola should be included in Linaria sect. Supinae (Benth.) Wettst. subsect. Supinae. Most taxa of this subsection are narrowly endemics from the Iberian Peninsula, mainly in southern Spain (Sutton 1988, Sáez & Bernal 2009). The new species and other morphologically related species are compared in detail in Table 1. Based on morphological grounds, L. argillicola is clearly related to the Andalusian endemic L. accitensis. The two taxa share many vegetative and floral characters, although a careful comparison of their morphological features reveals identifying differences (Table 1, Figs. 3–4). Linaria argillicola differs from L. accitensis by its larger seeds (up to 3.2 × 3.8 mm), with wider wing seed (up to 1.4 mm) and tuberculate seminal discs (Fig. 3), a lower number of stems (up to 13), longer abaxial sepals (up to 6 mm in flower), larger and wider corollas (up to 26 × 6.5 mm), and a different colour of the corolla (yellow, rarely reddish; Fig. 2). In addition, most of the inflorescences of L. argillicola usually showed a non-continuous appearance, since aborted fruits are found between well-developed basal fruits and apical flowers on the same branch. This peculiar character was commonly observed in different individuals within each population studied. Ecologically, L. accitensis and L. argillicola are also restricted to several subpopulations occupying a small area in Granada and Jaén provinces (Fig. 4), but the former grows on local and scattered volcanic soils (andesites and alkaline basalts) and the latter typically occurs on marly gypsiferous substrates. The new species is also related to L. tristis and L. aeruginea groups, sharing with them most of the vegetative characters (e.g. fertile stems decumbent to suberect, flat or slightly revolute leaves) and some flower features (spur usually shorter than the rest of the corolla). Nevertheless, L. argillicola is easily distinguished from L. aeruginea s.l. (a species widespread throughout the Iberian Peninsula) by its yellowish or rarely reddish corollas and glabrous

Linaria argillicola (Plantaginaceae) Phytotaxa 343 (2) © 2018 Magnolia Press • 135 inflorescence and calyx (Figs. 1–2). The typical subspecies of L. tristis, endemic to south-western Spain, differs from L. argillicola by its inflorescence axis and calyx lobes densely glandular-pubescent (Fig. 2), longer spur (up to 12 mm long) and wider leaves [1–3(–4) mm width]. Finally, L. badalii (a species widespread in northern and north-eastern Iberian Peninsula, recently found by us in Andalusia, see Annex 1) differs from the new species by its smaller seeds (1.5–2.4 × 1.5–2.3 mm), which are commonly papillate (Fig. 3). In addition, L. badalii is an annual plant, with leaves on fertile stems grouped in whorls of 3–5 up to the half of stem length and narrower corolla tube (Table 1, Fig. 2). According to our data, the mention of L. badalii on volcanic soils from Granada province reported by Baena et al. (2003) would correspond to L. accitensis.

Acknowledgements

We express our gratitude to the herbaria of ABH, BC, GDA, MA, SEV and VAL for the loan of material. We greatly appreciated the constructive comments by Prof. Dr. M.B. Crespo; we also thank Noelia Zarza for her help in the laboratory; the Centro de Instrumentación Científica (University of Granada) for the use of the SEM; the Andalusian Regional Government (Consejería de Medio Ambiente y Ordenación del Territorio de la Junta de Andalucía) for permission to herborise throughout the Andalusian territory; David Belchí for the line drawings; E. López-Carrique for assistance with Fig. 4; and David Nesbitt for the language editing.

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Linaria argillicola (Plantaginaceae) Phytotaxa 343 (2) © 2018 Magnolia Press • 137 ANNEX 1. Studied selected material in alphabetical order: Linaria accitensis:—SPAIN. Granada: Alamedilla, Cortijo de Cornelio, laderas abruptas de litología volcánica (andesitas y basaltos alcalinos, canchales y pedregales, 30SVG8161, 765 m, 22 May 2011, J. Fuentes (GDA 62645!); Alicún de Ortega, Loma de la Solana, arenas volcánicas, 30SVG8262, 24 April 1997, F.B. Navarro, H. Tenorio & J. Muñoz (GDA 45541–1!, GDA 45541–2!, sub L. badalii); Alicún de Ortega, Los Colladicos, 30SVG8163, 890 m, 17 May 2007, F.B. Navarro, J. Peñas, A. Juan & M.B. Crespo (ABH 51419–1!, ABH 51419–2!, ABH 51419–3!, ABH 51419–4!, ABH 51435–1!, ABH 51435–2!). Linaria aeruginea subsp. aeruginea:—SPAIN. Albacete: alrededores de la Laguna Ojos de Villaverde, 30SWH5495, 920 m, 21 June 1986, J.M. Herranz (MA 361056–1!, MA 361056–2!); entre Viano y Riópar, Monte Cerrolloso, 28 June 1988, A. Aparicio, F. García & S. Silvestre (MA 490842!). Almería: Bacares, Sierra de los Filabres, sobre calizas del Calar del Gallinero, 2000 m, 1 June 1994, A. Pallarés (MA 542258!). Ávila: Puerto de Menga, Valle de Amblés, 30 May 1976, E. Fuertes & M. Ladero (MA 333299–1!, MA 333299–2!). Castellón: La Pobla de Benifassà, pr. presa del Embalse de Ulldecona, 40.66N 0.23W, 18 April 2000, V.J. Arán & M.J. Tohá (ABH 43502!). Granada: Sierra Elvira, roquedos calizos, 28 April 1980, M. Ladero & J. Hurtado (MA 330370!); Sierra Nevada, Las Víboras, 1500 m, 2 June 1966, Valdés (MA194813!); Sierra de la Sagra, roquedos calizos, 1900 m, 13 July 1978, E. Fuertes, M. Ladero & C. Navarro (MA 211093–1!, MA 211093–2!). Cuenca: Alto Júcar, 16 June 1969, S. Rivas Goday, J. Izco & M. Ladero (MA 330377–1!, MA 330377–2!). Guadalajara: Labros, Carramayas, 41.05 N -1.96 W, 22 June 1995, L. Serra, A. Juan & J.C. Cristóbal (ABH 13350!); Taravilla, Laguna de la Parra, 30TWL8600, 1120 m, 20 June 1995, L. Serra, A. Juan & J.C. Cristóbal (ABH 13414!); Taravilla, laguna de la Parra, 30TWL8600, 1120 m, 20 June 1995, M. Carrasco et al. 608FCL (MA 558736–1!, MA 558736–2!); Barriopedro, 26 June 1970, Bellot, Carballal & Ron (MA 194119!). Jaén: Sierra Seca, El Chaparral, 1800 m, 29 June 1988, Valdés et al. It 2753/88 (SEV 160549!). León: Sierra de la Cabrera, camino de la Baña al Lago, 29TPG9182, 1150 m, 10 July 1981, Lansac & Nieto Feliner 240GN (MA 279030!). Madrid: El Escorial, encima de la Fuente de la Reina, suelo ácido, 21 June 1966, F. Getliffe, M. Gilbert & B. Valdés 911 (MA 194816!). Murcia: Moratalla, Sierra de la Muela, 30SWH9035, 1250 m, 24 April 1997, C. Aedo et al. 1091 IA (MA 591462!); Sierra Espuña, 1200 m, 2 May 1970, J. Fernández Casas (MA 408138!). Teruel: El Pobo, Sierra del Pobo, Alto de Castelfrío, ladera E, XK7782, 1600 m, 5 July 1995, V.J. Arán (MA 561212–1!, MA 561212–2!). Zamora: Villaciervos, 21 June 1973, B. Casaseca (MA 197749!). Linaria argillicola (paratypes):—SPAIN. Granada: Villanueva de las Torres, bco. del Caballo, taludes margosos ricos en yesos, 30SVG9757, 650 m, 25 May 2012, J. Fuentes (GDA 62648!). Jaén: Cabra de Santo Cristo, próx. Alicún de Ortega, entre Tollo de la Hierba y Cueva de Juan y Pedro, laderas abruptas de litología margosa con yesos, en suelos poco evolucionados, 30SVG8763, 735 m, 22 May 2011, J. Fuentes (GDA 62647!); Pozo Alcón, Los Collejares, balsa de las Bombas, taludes margo-yesíferos en orientación norte, 30SVG8784, 490 m, 31 March 2017, J. Fuentes, M. Cueto & G. Blanca (GDA 62649!). Linaria badalii:—SPAIN. Álava: Valdegovía, Guinea, monte Pilistrones, roquedo calizo, 30TVN9844, 850 m, 20 June 1985, Alejandre 1352–85 (MA 333493!). Córdoba: Zuheros, Parque Natural Sierras Subbéticas, hacia Fuente de la Zarza, pastizales terofíticos en sustrato calizo, 30SUG8554, 1000 m, 13 April 2017, J. Fuentes & G. Blanca (GDA 62650!). Huesca: Guara, Tozal de Guara, 1900 m, L. Sáez (L. Sáez-Herb. pers.!). La Rioja: Logroño, sobre roquedos silúricos, Barranco de Cambrones, Mansilla, 7 September 1976, Bote et al. (MA 330387! sub f. odoratissima). Palencia: Santibáñez de la Peña, Santuario de Nuestra Señora del Brezo, 30TUN6045, 1400–1600 m, 10 July 1995, X. Giráldez et al. (SALA 101842–1!, SALA 101842–2!). Segovia: Sepúlveda, 18 July 1983, T. Romero (SALA 39437!); Fuentidueña, 17 June 1984, T. Romero (SALA 39436–1!, SALA 39436–2!); Espirdo, La Higuera, calizas, 30TVL0944, 1000 m, 2 July 1988, R. García 5055RG (MA 560166!); Pedraza, Las Torcas, 30TVL3052, 1160 m, calizas, 24 May 1987, P. Egido & R. García 3400RG (MA 560163!). Soria: Cañón Río Lobos, VM9225, 16 June 1982, A. Buades (MA 541513! sub f. odoratissima). Teruel: Torrijas, Barranco de Domingo, XK7235, 1620 m, 2 July 1988, G. Mateo (ABH 6079!); Orihuela del Tremedal, a Alusante, prox. Cerro S.M., XK1592, 1430 m, G. Mateo (ABH 6080!). Linaria tristis subsp. tristis:—SPAIN. Cádiz: Algeciras, Puerto de Comadres, 20 April 1962, B. Casaseca (SEV 189092!); Puerto de Comadres, Algeciras, 20 April 1962, B. Casaseca (SEV 180922!); Grazalema, Puerto de las Palomas, 27 May 1990, Martin (MGC 27259!); Jerez de la Frontera, de El Picacho al Puerto de Cádiz, TF6448, 17 May 1985, A. Asensi et al. (MGC 19306!); Grazalema, el Pinsapar, 2 July 1990, A. Asensi (MGC 27293!); Grazalema, Puerto de las Palomas, TF8770, 1100 m, 29 April 1983, A. Aparicio & S. Silvestre (SEV 160563!); El Gastor, Sierra del Gastor, TF9280, 1000 m, 29 May 1983, A. Aparicio et al. (SEV 160580!). Gibraltar: Catalan Bay, arenas con rocas calizas, 16 May 1985, J. Bensusan et al. (SEV 124973!); Punta del Pasagre, 16 May 1985, J. Bensuan et al. (SEV 124908!). Málaga: subida a Benahavis, cunetas removidas, 28 April 1978, Luque et al. 2324/78 (SEV 160750!).