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Article

Sideritis royoi (Lamiaceae): A New Orophilous Species from Northeastern Spain †

by
Llorenç Sáez
1,*,
Rafel Curto
2 and
Manuel B. Crespo
3
1
Systematics and Evolution of Vascular Plants (UAB)—Associated Unit to CSIC, Department BABVE, Autonomous University of Barcelona, ES-08193 Bellaterra, Spain
2
Grup EbreRecerca, ES-43500 Tortosa, Spain
3
Department of Environment Sciences and Natural Resources (dCARN), University of Alicante, P.O. Box 99, ES-03080 Alicante, Spain
*
Author to whom correspondence should be addressed.
†
IPNI: urn:lsid:ipni.org:names:77334707-1.
Taxonomy 2024, 4(1), 112-125; https://doi.org/10.3390/taxonomy4010006
Submission received: 11 December 2023 / Revised: 5 January 2024 / Accepted: 16 January 2024 / Published: 18 January 2024
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Abstract

:
Sideritis royoi is found in the rocky limestone habitats of the Port Massif (southern Catalonia, Spain). The species was first collected by the local botanist Lluís de Torres in the late part of the 20th century, but the specimens have remained unidentified positively in herbaria for over 40 years. Sideritis royoi likely belongs to section Sideritis subsection Hyssopifoliae and shows some morphological affinities with the relatively widespread South European species S. hyssopifolia L., but it differs from this species because it has subspinescent upper leaves, the main surfaces of its leaves are glabrous or glabrescent, the main abaxial surface of its bracts is without eglandular hairs, and due to the fact that it has shorter inflorescences. Weaker similarities have also been observed with some species belonging to S. subsection Fruticulosae Obón & D.Rivera. In this paper, a description for the new orophilous species is provided, along with a detailed illustration, field photographs, and a comparison with closely related species. We include an assessment of its conservation status and a dichotomous key for the identification of all the species of Sideritis subsection Hyssopifoliae.

1. Introduction

Sideritis L. is a large genus of family Lamiaceae Martynov, nom. cons., that includes nearly 150 species that are mainly distributed in Europe, North Africa, Macaronesia, and Western and Central Asia [1,2,3,4], though some of these species have been introduced or naturalised in other areas. The Port Massif (northeastern Spain, Tarragona Province) exhibits high plant species richness and endemism [5]. During our field work and subsequent study of herbarium materials from this mountainous area, some specimens of Sideritis L. caught our attention due to a combination of characteristics that do not match those of any described species within the genus. In the upper areas of the Port Massif, the presence of plants of the genus Sideritis, whose identity is uncertain, dates back more than 40 years. These plants had initially been collected by the local botanist Lluís de Torres in 1982 at the base of Caro (the highest mountain in the area), but strikingly, it was not listed in the floristic catalogue of the Port Massif [6]. Subsequently, Buira et al. [7], including two of the authors of this paper (L.S. and R.C.), tentatively attributed those plants to S. hyssopifolia L. while noting some discordant morphological characters. The most recent reference for the Sideritis occurring in the upper parts of the Port Massif is the work of Sáez and Aymerich [8], who, commenting on S. hyssopifolia, indicated that the taxonomic identity of those populations was unclear.
In order to clarify that point, during field trips conducted between 2021 and 2023, we gathered new specimens of the Sideritis from the Port Massif which had doubtfully been attributed to S. hyssopifolia. After studying the material collected, herbarium specimens, the existing literature, and photographs, as well as conducting field observations, it was revealed that the Port Massif plants do not fit with any currently accepted species within the genus and represent a new species that is described in this paper.

2. Materials and Methods

The present morphological and comparative study is based on the examination of herbarium specimens. In addition, field observations were carried out on individuals from the populations of the Port Massif area. Morphological characters recognised as taxonomically discriminant within Sideritis were studied according to Obón and Rivera [1], Rivera et al. [9,10], and Ríos et al. [11]. Morphological observations of materials were carried out under a binocular stereoscopic microscope Zeiss Stemi DV4. We examined the material of morphologically related species held mostly at the herbaria ABH, BC, and MUB. Authors of the taxa cited in the text follow IPNI [12], and the herbarium acronyms used are in line with the work of Thiers [13], except for MMA (Museu de les Terres de l’Ebre, Amposta, Tarragona). The representative herbarium material examined is listed in the Appendix A. Bioclimatic classification conformed to Rivas-Martínez [14], and the way in which we approached biogeographical data followed the approach of Rivas-Martínez [15]. To evaluate the conservation status of the new species, the IUCN Red List Categories and Criteria [16,17] were used.

3. Results

The combination of morphological characters with diagnostic values, the biogeographical data, and a review of the foremost regional floras and taxonomic revisions [1,11,18] suggested that the plants collected in the Port Massif area are distinct enough to merit recognition as a new taxon for which we believe the rank of species to be the most appropriate and which we describe herein.

Taxonomic Treatment

Sideritis royoi L.Sáez, R.Curto & M.B.Crespo, sp. nov. Figure 1 and Figure 2.
Holotype: SPAIN. Catalonia, Tarragona province: Baix Ebre, Roquetes, La Barcina, 31TBF7721, limestone rocks, 1250 m a.s.l., 16 July 2023, R. Curto s.n. (ABH 83741!). Isotype: BC 849997!.
Diagnosis: Sideritis royoi differs from S. hyssopifolia in that its lower leaves (main surface) are glabrous or sometimes glabrescent (vs. hairy); the upper leaves are subspinescent and narrower than the lower leaves (vs. non-subspinescent and similar to the lower); the adaxial and abaxial surfaces of the bracts lack eglandular hairs (vs. eglandular hairs present); and it has shorter inflorescences with 1–2(–3) verticillasters (vs. longer inflorescences with (1–)2–13 verticillasters).
Description: Dwarf shrub. Woody basal parts 7–14 cm, including branches, decumbent to suberect. Non-woody branches 8–30 cm long, ascending to erect. Branchlets with goniotrichous and homotrichous trichomes, with scarce to abundant glandular hairs up to 0.1 mm long and lacking glands; trichomes scarce, antrorse, 0.2–1 mm long, with 2–3(4) cells cylindrical, the apical one conical. Leaves patent to suberect, sessile to shortly petiolate (petiole up to 7 mm long), greenish, flat, subspatulate, obovate–lanceolate or sublinear, attenuate at base, with 0-4 secondary veins; main surfaces glabrous or glabrescent, with very scarce eglandular hairs 0.2–0.7 mm long, patent mainly at the margins and at the midrib of the adaxial surface, with scarce glandular hairs up to 0.1 mm long and always lacking sessile glands; lower and middle leaves 8–32 × 3–12 mm, 2–4-dentate, apiculate or mucronate at the apex; upper leaves 6–15 × 2–4 mm, subspinescent, usually 2-dentate, sometimes entirely; axillary fascicles of leaves usually absent at the flowering time. Inflorescence commonly yellowish or greenish, ovoid to globose, 0.7–2.5(–4) × 0.7–1.8 cm, with 1–2(–3) verticillasters, the central 3–3.5 mm apart; axis usually yellowish or greenish, with sparse sessile glands and glandular hairs up to 0.1 mm long abundant antrorse eglandular hairs 0.2–0.8 mm long. Bracts greenish to yellowish, erect, ovate, with the widest part towards its basal third; divided into 1/3–1/2 of its width; adaxial surface glabrous, abaxial with sparse (sometimes abundant) glandular hairs up to 0.1 mm long and always lacking sessile glands; eglandular hairs usually absent, sometimes present (very scarce) at the margins, 0.2–0.5 mm long; teeth 1–3(–4) long, narrowly triangular to subulate; lower bracts 7–9 × 6–10 mm, with 3–8 teeth on each side; middle bracts 8–10 × 9–12 mm, divided to 1/3–1/2 of its width, with 6–10 teeth on each side. Six-flowered verticillasters. Calyx campanulate, 8–11 mm long (calyx tube 4–5.2 mm long), with five subequal divergent teeth, 4–8 mm long, ending in spines 0.7–2 mm long; the outer surface with scarce to abundant glandular hairs up to 0.15 mm long, sessile glands and antrorse eglandular hairs 0.5–1.8 mm long; carpostegium (ring of hairs within calyx) discontinuous. Corolla yellowish, 6–9 mm long, with tube 3–4, 4 mm long, subcylindrical; upper lip bifid up to 1/5–1/3 of its length; stamens included in the corolla tube, with short filaments 0.3–0.5 mm long; style 1.8–2.5 mm long. Nutlets ovoid, c. 2 × 1.5 mm.
Eponymy: The new species is dedicated to the late Ferran Royo Pla (1969–2016) for his outstanding work that contributed to improving the botanical knowledge of the Port Massif.
Phenology: Flowering is from June to early July; fruiting is from late July to late August.
Distribution and ecology: Sideritis royoi is an orophilous species endemic to a small area in the Port Massif (Figure 3) in the northeastern Iberian Peninsula. The new species is found in the rocky slopes and in patches in the understory of the open Pinus sylvestris L. forests at the northern face of the Caro-La Barcina mountain (Figure 4), between 1080 and 1325 m a.s.l. The known sites of the new species show a pluviseasonal Mediterranean bioclimate, mostly within the Supramediterranean thermotype and with a Subhumid or locally Lower-Humid ombrotype [14]. Biogeographically, this area belongs to the Puertobeceitan-Morellan District of the Western-Catalanid subsector (Valencian-Tarraconensian Sector, Catalan-Provencian-Balear Province, Mediterranean Region), according to Rivas-Martínez [15]. The Caro-La Barcina mountain hosts a rich flora including a high number of plant species that are endemic to the northeastern Iberian Peninsula [5], and some of them are restricted to the Port Massif. Font Quer [19] first noticed that several plant species occurring in the Port Massif usually showed conspicuous morphological differences compared with other related taxa occurring in other areas of the northeastern Iberian Peninsula. The new species grows together with several Puertobeceitan-Morellan endemics, such as Aquilegia paui Font Quer, Arenaria conimbricensis subsp. viridis (Font Quer) Font Quer, Knautia rupicola (Willk.) Font Quer, or Thymus willkommii Ronniger, as well as more widely distributed orophilous species such as Arctostaphylos uva-ursi (L.) Spreng., Erinacea anthyllis Link subsp. anthyllis, Festuca trichophylla (Gaudin) K.Richt., and Teucrium aureum Schreb. subsp. aureum.
Conservation status: Based on the present state of knowledge, due to its low population size (c. 100 individuals), S. royoi should be listed as “Endangered” (EN), according to IUCN criterion D [16]. This species has an extent of occurrence and an area of occupancy of 0.75 km2, calculated on a 0.5 × 0.5 km grid. Based on the data currently available, we have no evidence of a population decline. However, further field studies are needed to define a more accurate conservation category for this species.
Additional specimens examined: SPAIN. Tarragona Province: Alfara de Carles, prop de les Clotes, 1080 m a.s.l., 10 July 1982, L.Torres (MMA); per davall del Pas de la Barcina, cara Nord, 1200 m a.s.l., 14 July 2009, Aparicio, Beltran, Curto, Mesa & Royo 6085 (MMA); La Barcina, matollar, 31TBF72, 1270 m a.s.l., 25 July 2021, R.Curto (L. Sáez pers. herb.).

4. Discussion

Sideritis royoi shows morphological affinities with taxa of S. sect. Sideritis subsect. Hyssopifoliae Obón & D.Rivera and is assumed to belong here based on its hair covering at the base of its goniotrichous (or slightly holotrichous) branchlets with antrorse trichomes [1]. Most of the taxa belonging to this group are found in mountain areas of the Iberian Peninsula and Northern Africa, with a higher concentration in Southern and Eastern Spain. Some of these species are orophyte specialists that are narrowly endemic to a reduced mountain range [1,11]. A comparison of S. royoi with morphologically related species belonging to S. subsect. Hyssopifoliae is shown in Table 1.
The type species of this subsection, S. hyssopifolia, is quite variable, particularly in habit, as well as in the size of the stems, leaves, and flowers. Several variants are treated at species or infraspecific ranks [1,18,20], or even included in synonymy [18]. This is the case for S. brachycalyx Pau (≡ S. hyssopifolia var. brachycalyx (Pau) Font Quer), an entity often regarded as a distinct species based mostly on the lack of carpostegium. However, this is also a variable character that is even inconstant in S. brachycalyx. Therefore, for practical purposes, in this work, S. hyssopifolia is considered, in a broad sense, to include S. brachycalyx, a taxon that, in our opinion, is best to treat as a subspecies or variety of the former [21].
Sideritis hyssopifolia is apparently closely related to S. royoi, with which it shares the goniotrichous and homotrichous hair covering at base of branchlets, the general habit and calyx shape, size, and indumentum. However, S. royoi differs from S. hyssopifolia in several striking characteristics (see Table 1), namely the glabrous nature of the main surfaces of the leaves, the subspinescent upper leaves, the absence of eglandular hairs on the main surfaces of the bracts (Figure 1A), and shorter inflorescences, while S. hyssopifolia has usually hairy leaves, non subspinescent upper leaves, eglandular hairs on the main surfaces of the bracts, and longer inflorescences.
Weaker similarities were also observed with other taxa in S. subsection Hyssopifoliae. Sideritis pungens (including subsp. vigoi Peris et al. and also, for practicity, S. javalambrensis Pau ≡ S. pungens subsp. javalambrensis (Pau) Obón & D.Rivera) differ from the new species mainly due to the heterotrichous hair covering the base of their branchlets, the entirety of the lower leaves, the fact that they have cylindrical inflorescences, hairy adaxial surfaces with respect to the middle bracts, and shorter calyces with a continuous carpostegium. The differences between S. pungens and S. javalambrensis and other related orophilous taxa have been highlighted by López-Udias [22].
Sideritis carbonellii Socorro and S. tugiensis S. Ríos et al., two species closely related to each other that occur in southern Spain, are much more different morphologically. They are easily separated from S. royoi by the base of their branchlets having holotrichous hairs and glands, the main surfaces of their leaves being hairy, the fact that they have longer inflorescences, the fact that they have fewer teeth on middle bracts, and the fact that they have shorter calyces. See Ríos et al. [11], López-Udias [22], and Table 1 for the differences between S. carbonellii and S. tugiensis.
Finally, the morphological relationships with species belonging to S. subsect. Fruticulosae Obón & D.Rivera seem to be remote. It was suggested that the plants now described as S. royoi might be the result of introgression with S. spinulosa Barnades ex Asso [7] because of the presence of subespinescent upper leaves in S. royoi. Although this genus is very rich in hybrids and hybrid swarms in the Iberian Peninsula [23,24,25,26], our results indicate that the morphological differences between S. spinulosa and S. royoi are notable, allowing for a clear separation between those two species. Sideritis spinulosa differs by having holotrichous hair covering the base of branchlets, leaves with scarce glands and abundant trichomes 1.5–2.0 mm long, cylindrical inflorescences with 3–15 verticillasters, the abaxial surface of its middle bracts with scarce glands and scarce abundant trichomes (0.5–1.5 mm long), and calyx with a continuous carpostegium. Another species of S. subsect. Fruticulosae occurring in the Port Massif but in lower areas (up to 750 m a.s.l. altitude) is S. fruticulosa Pourr. This species differs from S. royoi by having holotrichous hair covering the base of its branchlets, leaves with scarce glands and abundant trichomes 0.3–1.5 mm long, cylindrical inflorescences with 3–7 verticillasters, the abaxial surfaces of middle bracts with scarce glands and scarce abundant trichomes (1.0 mm long), and calyx with a continuous carpostegium. In a recent and detailed taxonomic revision of S. fruticulosa [27], four subspecies were recognised, but none of them come close to having the morphological characteristics of S. royoi. Interestingly, Roselló et al. [27], who were very sensitive to detecting small morphological variations and hybrids, indicate no morphological relationship or existence of hybrids with S. hyssopifolia. Further, we have not observed S. spinulosa or S. fruticulosa in the locations where S. royoi occurs. Finally, regarding the suggested possible hybrid origin, S. hyssopifolia is widespread in the northern part of the Iberian Peninsula, mainly in the Pyrenees [11,28], but the species is not known to originate from the Port Massif [6,28,29], and thus, no range overlap with S. royoi occurs (Figure 3). Further molecular and cytogenetic work will help to elucidate this point.

5. Identification Key for Sideritis subsect. Hyssopifoliae

The following key for species of S. sect. Sideritis subsect. Hyssopifoliae has been adapted from Obón and Rivera [1] in order to accommodate the new described species. As stated before, for practical purposes, S. hyssopifolia is considered in this work in a broad sense (i.e., including S. brachycalyx). Similarly, herein, S. pungens includes S. javalambrensis.
1 .
Calyces with carpostegium continuous …………………………………………………………………………………………………………….….2
1′.
Calyces with carpostegium discontinuous ……………………………………………………………………………………………………………7
2 .
Central part of verticillasters 2–10 mm apart …………………………………………………………………………………………………………3
2′.
Central part of verticillasters 15–35 mm apart ………………………………………………………………………………………………………11
3 .
Inflorescence axis without glands ………………………………………………………………………………………………………………………4
3′.
Inflorescence axis with glands ………………………………………………………………………………………………………………………….5
4 .
Base of branchlets with glands, trichomes 0.3–0.7 mm long; inflorescence axis trichomes 0.2–0.4(–1) mm long ……………… S. getula Batt.
4′.
Base of branchlets without glands, trichomes 0.5–2.0 mm long; inflorescence axis trichomes 0.8–1.5 mm long ………… S. hyssopifolia (s.l.)
5 .
Leaves without glands; lower bracts 5–7 × 8–10 mm, with 2–4 teeth on each side; corollas 6–7 mm long ………………… S. maura de Noé
5′.
Leaves with glands; lower bracts 7–12 × 5–12 mm, with 3-8 teeth on each side; corollas 8–10 mm long ……………………………………6
6 .
Middle bracts 7–12 mm; calyx teeth 3 mm long, trichomes 0.6–1.3 mm long …………………………………………………… S. pungens (s.l.)
6′.
Middle bracts 5–6 mm; calyx teeth 1–2 mm long, trichomes 2 mm long ……………………………………… S. algarviensis D. Rivera & Obón
7 .
Branchlets with glands at base …………………………………………………………………………………………………………………………8
7′.
Branchlets without glands at base ……………………………………………………………………………………………………………………10
8 .
Base of branchlets goniotrichous to holotrichous, covered with hairs 0.8–1.5 mm ……………………………………………… S. pungens (s.l.)
8′.
Base of branchlets holotrichous, covered with very short hairs up to 0.8 mm ………………………………………………………………… 9
9 .
Lower leaves entire, 10–25 × 1–1.5 mm, axillary fascicles absent at flowering time; uppermost leaves similar to the lower, entire; calyx with scarce trichomes ……………………………………………………………………………………………………………………… S. carbonellii
9′.
Lower leaves dentate, 12–17 × 2–3 mm, axillary fascicles commonly present at flowering time; uppermost leaves bract-like, with 0–4 teeth on each side; calyx with abundant trichomes ……………………………………………………………………………………………… S. tugiensis
10.
Main surfaces of the leaves with eglandular hairs; upper leaves non subspinescent; main abaxial surface of the bracts hairy; inflorescences with (1–)2–13 verticillasters ……………………………………………………………………………………… S. hyssopifolia (s.l.)
10′.
Main surfaces of the leaves glabrous or glabrescent; upper leaves subspinescent; main abaxial surface of the bracts without eglandular hairs; inflorescences with 1–2(–3) verticillasters …………………………………………………………………………………………. S. royoi
11.
Leaves without glands; lower bracts 6–8 × 3–6 mm, middle bracts 4–5 × 7–8 mm ………………………………………… S. ochroleuca Willk.
11′.
Leaves with glands; lower bracts (8–)10–17 × 3–9(–10) mm, middle bracts 7–10 × 8–10 mm ……………………………… S. hyssopifolia (s.l.)

Author Contributions

Conceptualisation, L.S., R.C. and M.B.C.; methodology, L.S. and M.B.C.; investigation, L.S., R.C. and M.B.C.; resources, L.S., R.C. and M.B.C.; data curation, L.S., R.C. and M.B.C.; writing—original draft preparation, L.S. and M.B.C.; writing—review and editing, L.S., R.C. and M.B.C.; visualisation, L.S., R.C. and M.B.C.; supervision, L.S., R.C. and M.B.C. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Data are contained within the article.

Acknowledgments

We thank the staff of the cited herbaria for their help in the study of the herbarium sheets.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A. Representative Specimens

Sideritis carbonellii Socorro: SPAIN. Granada: Huéscar, pico de La Sagra, embudo de la ladera N, 30SWH385014, 1910 m a.s.l., 13 July 2011, S.Patino, E.Miguel & V.J.Arán 7732 (ABH 80790, MA 00868149); Huéscar, Sierra de La Sagra, 30SWH30, 2100 m a.s.l., 29 August 1995, pedregales calizos, J.L.Solanas, L.Serra, F.Alcaraz & S.Ríos (ABH 14172); ibidem, La Sagra, 30SWG39, 27 July 1984, F.Alcaraz (ABH 43817, MUB); Baza, Sierra de Baza, Calar de Santa Bárbara, 30SWG13, September 1992, D.Rivera & C.Obón (ABH 43819, MUB).
Sideritis fruticulosa Pourr.: SPAIN. Huesca: Candasnos, Valle de la Valcuerna, 31TBF5396, 3 May 1981, cerros de margas yesíferas, Alamillo, S.Castroviejo, Fernández Quirós & Nieto (ABH 53472, MA 436752). Lérida: Castelldans, Lo Timorell, plataforma superior, 31TCF1496, 420 m a.s.l., 2 June 2007, matorral calcícola, V.J.Arán, J.Rebull & R.Valls 6579 (ABH 52005). Navarra: Cizur, Astráin, Sierra de Erreniega, Las Bordas, 30TXN028324, 755 m a.s.l., 18 July 2013, matorral-pasto mesoxerófilo, sur-suroeste, L.Romeo, M.Imas & R.Ibáñez (ABH 70291). Tarragona: Hospitalet de l’Infant, 16 June 1916, P.Font Quer (BC 73560). Zaragoza: Tauste, Balsa Tres Montes, 30TXM4950, 280, 13 July 1997, M.B.Crespo & E.Camuñas (ABH 35770).
Sideritis hyssopifolia L. (s.l.): FRANCE. Alpibus delphiniensibus, [Dépt. 05] Coll de Montginebre, 1750 m a.s.l., 22 July 1973, O.Bolòs (BC 609226); Dépt. 01, entre Sur Thoiry et le Reculet, 24 September 1978, C.Defferrard, P.Braun & A.-M.Frei CDF-837 (ABH 63405); ibidem, entre Sur Thoiry et le Reculet, 28 August 1992, C.Defferrard & P.Braun CDF-7413 (ABH 63406); Dépt. 11, Dulhac, Rouffiac-des-Corbières, Castell de Perapertusa, 31TDH64, 804 m a.s.l., 20 July 2006, sobre les roques calcàries poc inclinades, J.L.Solanas (ABH 80011); Dépt. 16, Cognac-Angoulême, Vignac, 100 m a.s.l., 6 July 1990, pelouse sèche, D.Masson CDF-DM-1394 (ABH 69098, S. hyssopifolia subsp. guillonii (Timb.-Lagr.) Nyman). SPAIN. Asturias, Puerto de Pajares, 1800 m a.s.l., 29 July 1935, W. Rothmaler (BC 650529, S. hyssopifolia subsp. somedana Obón & D.Rivera); Peña Ubiña, 2300 m a.s.l., 10 August 1935, W. Rothmaler (BC 650529, S. hyssopifolia subsp. somedana Obón & D.Rivera); cerca de Celorio, 1952 M.Laínz (BC 123573, S. brachycalyx); Caso, Gobezanes, 30TUN079868, 780 m a.s.l., 19 July 2002, calizas de la Formación Escalada, E.Alonso & E.de Paz (ABH 47415, LEB); Onís, Picos de Europa, 30TUN4290, 1500 m a.s.l., 27 June 2003, M.Mart.Azorín (ABH 48241). Barcelona: Cabrera, pr. Berga, 1600 m a.s.l., 1 August 1912, Sennen (BC 73975, S. hyssopifolia subsp. peyrei (Timb.-Lagr.) Briq.). Barcelona: La Clusa, 1620 m a.s.l., N Font del Tudó, 5 October 1975, A.Rosell (BC 622077); Vallcebre, La Foranca, 1640 m a.s.l., 20 July 2020, D.Pérez (BC 982974). Cantabria: Sant Vicente de la Barquera, August 1920, E. Jiménez (BC 73959, S. brachycalyx); Piélagos, Liencres-Playa de Valdearenas, 30TVP2212, 10 m a.s.l., 24 August 2010, acantilados, E.Camuñas & M.B.Crespo (ABH 57387). Huesca: Torla-Ordesa, Valle de Ordesa, 30TYN42, 1300 m a.s.l., 27 July 1970, A.Rigual (ABH 20728); Torla-Ordesa, Valle de Otal, río Otal, 30TYN33, 1600 m a.s.l., 06 August 1994, A.Juan (ABH 16078); Torla-Ordesa, Ordesa, Valle de Bujaruelo, 30TYN3832, 1900 m a.s.l., 09 August 1996, A.Juan, E.Camuñas & M.B.Crespo (ABH 18406); Broto, prox., 30TYN32, 1000 m a.s.l., 27 July 1970, A.Rigual (ABH 20699); Bielsa, Sobrarbe, Chisagüés, 31TBH6529, 1700 m a.s.l., 20 June 1996, M.B.Crespo, L.Serra, A.Juan, J.C.Cristóbal & al. (ABH 19210); Aísa, Collado de la Magdalena, 30TXN9933, 2020 m a.s.l., 17 July 1997, M.B.Crespo, E.Camuñas, A.Juan, J.L.Solanas, J.L.Benito & A.Barber (ABH 68950, S. hyssopifolia subsp. eynensis (Sennen) Malag.). Panticosa, Ibón de los Asnos, hacia mirador de los Valles, 30TYN2329, 2180 m a.s.l., 22 August 2012, M.B.Crespo & E.Camuñas (ABH 59850, S. hyssopifolia subsp. aranensis (Font Quer) Malag.). Lleida: Pr. Salardú, Vall d’Aran, 18 September 1922, Gros (ABH 59850, lectotype of S. hyssopifolia subsp. aranensis); Espot, estación invernal Super Espot, 31TCH41, August 1991, M.A.Jover (ABH 4772). León: Cueto Ancino, a 2 km al N de Nocedo,1200 m a.s.l., 11 August 1996, F.Gómiz (BC 827328, subsp. nocedoi Obón & D.Rivera); La Pola de Gordón, Santa Lucía de Gordon, 30TTN8450, 1200 m a.s.l., 14 October 1993, A.Juan, M.Vicedo & M.Á.Alonso (ABH 6417); San Emiliano, Peña Ubiña, vert. SE, 30TTN5967, 2200 m a.s.l., 30 July 2001, V.J.Arán & M.J.Tohá (ABH 45583, S. hyssopifolia subsp. somedana Obón & D.Rivera); Cabrillanes, Cabrillanes, La Cueta, Collado la Fontanina, 29TQH2868, 1638 m a.s.l., 16 July 2021, roquedos calizos y matorral sobre sustrato silíceo, A.Buira, L.Medina, S.Andrés Sánchez, J.Güemes & al. SA2016 (ABH 82071); San Emiliano, San Emiliano, Torrestío, subida al puerto de la Farrapona, valle Sañedo por encima del puente del río Traspando, 29TQH3870, 1583 m a.s.l., 15 July 2021, calizas, A.Buira, L.Medina, S.Andrés Sánchez, J.Güemes & al. LM11342 (ABH 82023). Lugo: Folgoso do Courel, Visuña, salida S del pueblo, 29TPH5819, 1050 m a.s.l., 8 July 2000, J.Amigo (ABH 44442, S. hyssopifolia subsp. caureliana Obón & D.Rivera). Navarra: Isaba, Rincón de Belagua, 30TXN7655, 1000 m a.s.l., 29 July 1991, P.M.Uribe-Echebarría (ABH 46453, VIT 5653); Isaba, Larra, 30TXN818597, 1700 m a.s.l., 04 August 2005, pastos pedregosos subalpinos, N.Jáuregui & R.Ibáñez (ABH 49905); Isaba, Macizo de Larra-Belagua, cercanías de la estación de esquí nórdico, sector La Contienda, hacia el Collado de La Piedra de San Martín, 42°57.820′ N 0°46.392′ W, 3 July 2022, taludes y rocas calizas del karst, M.Martínez Ortega, M.Á.Alonso, M.Mart.Azorín & al. MO6326 (ABH 83077). Orense: Carballeda de Valdeorras, Sierra de Campo Romo, Fonte da Cova, 29TPG8687, 1780 m a.s.l., 29 June 1994, M.B.Crespo, M.D.Lledó, L.Serra, A.Juan & J.C.Cristóbal (ABH 13291). Palencia: Peña Redonda, ca. Cervera de Pisuerga, 9 August 1914, P. Font Quer (BC 73925, subsp. santanderina D. Rivera & Obón); Velilla del Río Carrión, Cardaño de Arriba, prox., Pico Espigüete, sima del Anillo, 30TUN5357, 1890 m a.s.l., 12 July 2006, suelos calcáreos karstificados, V.J.Arán & G.Arán 6353 (ABH 52045); Velilla del Río Carrión, Sierra del Brezo, Peña Cueto, 30TUN5646, 1670 m a.s.l., 11 August 1991, pastizales psicroxerófilos calizos, A.Penas, M.E.García & L.Herrero (ABH 74617). Vitoria: Vitoria: Bernedo, Markinez, de Alto Raposeras a Ermita de Beolarra, 30TWN3528, 750 m a.s.l., 23 August 1999, P.M.Uribe-Echebarría (ABH 46454, VIT 61486, S. hyssopifolia subsp. castellana (Sennen & Pau) Malag.:); Lagrán, Sierra de Cantabria, Cruz del Castillo, 30TWN3316, 1375 m a.s.l., 18 July 2006, grietas de peñascos calizos, en la solana, P.M.Uribe-Echebarría (ABH 51864).
Sideritis pungens Benth. (s.l.): SPAIN. Burgos: Briviesca, 30TVN7308, 725 m a.s.l., 23 June 1998, M.B.Crespo, J.C.Cristóbal & al. (ABH 48708); ibidem, 30TVN7409, 780 m a.s.l., 25 July 1999, S.Patino (ABH 42818). Castellón: Vistabella del Maestrat, Penyagolosa, en la base, 30TYK2757, 1500 m a.s.l., 11 July 1995, E.Laguna (ABH 16549); Ares del Maestre, 30TYK4288, 1000 m a.s.l., 24 July 1995, A.de la Torre, M.Vicedo & M.Á.Alonso (ABH 16922); La Pobla de Benifassà, El Coratxà, Tossal de Mitjavila, 31TBF5310, 1340 m a.s.l., 20 June 2001, J.Riera & E.Estrelles (ABH 46363); Xodos (Alcalatén), Massís de Penyagolosa, Roca del Migdia, 30TYK2656, 1400 m a.s.l., 17 July 2018, matorrales calizos sobre suelos pedregosos, J.Riera & F.J.Fabado JRV-9012 (ABH 81808). Granada: Iznalloz, Iznalloz-estación del Piñar, 30SVG6038, 960 m a.s.l., 8 June 2000, V.J.Arán (ABH 69853). Logroño: Los Ábalos, 580 m a.s.l., 29 July 1996, R. Auriault (BC 837198). Navarra: Larraga, Larraga-Tafalla, 30TXN01, 12 July 1993, D.Rivera & C.Obón (ABH 43820, ABH 43821, MUB). Palencia: Hontoria de Cerrato, El Raposillo, 30TUM829426, 827 m a.s.l., 26 June 2019, matorral aclarado en cerro yesoso, S. Andrés-Sánchez & al. SA1573 (ABH 80216). Soria: pr. Numancia 1050 m a.s.l., 10 July 1935, P.Font Quer & W.Rothmaler (BC 638196); Almarza, Portelárbol, 30TWM480385, 1175 m a.s.l., 16 July 2020, encinar abierto en calizas, J.Güemes, C.Molina, A.Prunell, E.Rico, E.Sahuquillo & C.Urones ER8570 (ABH 82270). San Felices, bajando al río Alhama, 30TWM8043, 775 m a.s.l., 13 August 2014, calizas, M.B.Crespo & E.Camuñas (ABH 70704); Cueva de Ágreda, hacia Beratón, 30TWM9521, 1320 m a.s.l., 24 July 1997, M.B.Crespo & E.Camuñas (ABH 36427). Teruel: In monte Javalambre 1923, Pau (subsp. javalambrensis (Pau) Obón & D.Rivera, BC 74006); Teruel, pr. Masada Cociero, 30TXK6170, 935 m a.s.l., 9 July 2008, M.B.Crespo (ABH 53733); La Puebla de Valverde, Altos del Buitre, 30TXK7243, 1650 m a.s.l., 20 July 1995, E. Laguna (ABH16552, subsp. javalambrensis); Cerro de Javalambre, 30TXK6840, 2000 m a.s.l., 8 August 1996, C.Fabregat & S.López (ABH 44512, subsp. javalambrensis); Perales del Alfambra, hacia Visiedo, prox. del pueblo, 30TXK6800, 1160 m a.s.l., 24 July 2005, suelos arcillosos áridos, V.J.Arán & M.J.Tohá 6182 (ABH 51957). Valencia: Alpuente, Muela del Buitre, 30SXK6424, 1460 m a.s.l., 20 July 1996, J.J.Herrero-Borgoñón (ABH 30425). Zaragoza: Tauste, Balsa Tres Montes, 30TXM4950, 280 m a.s.l., 13 July 1997, M.B.Crespo & E.Camuñas (ABH 35771); La Zaida, 30TYL17, 30 July 1985, D.Rivera & C.Obón (ABH 43822, MUB).
Sideritis spinulosa Barnades ex Asso subsp. spinulosa: SPAIN. Guadalajara: Tartanedo, La Aguarrosa, 30TWL9137, 1180 m a.s.l., 22 June 1995, L.Serra, A.Juan & J.C.Cristóbal (ABH 13336). Palencia: Alba de Cerrato, 30TUM8727, 800 m a.s.l., 16 July 1980, F.Amich, E.Rico & J.Sánchez (ABH 30820, SALA). Teruel: Aguaviva, 3 July 1919, Rubió (BC 73596); Teruel, pr. Masada Cociero, 30TXK6170, 935 m a.s.l., 9 July 2008, M.B.Crespo (ABH 53735). Zaragoza: Calcena, hacia Ermita San Cristóbal, 30TXM0711, 900 m a.s.l., 23 July 1997, M.B.Crespo & E.Camuñas (ABH 39876); Borja, La Muela, Cerro del Boquerón, encinar, 30TXM1536, 750 m a.s.l., 18 June 2002, V.J.Arán 5216 (ABH 46677); Pedrola, Barranco de Juán Gastón, 30TXM446247, 289 m a.s.l., 22 May 2014, A.Terrones & A.Vicente (ABH 73535); Ibdes, río Mesa, matorrales próximos, 30TWL96, 12 August 1992, J.L.Solanas (ABH 1847).
Sideritis spinulosa subsp. subspinosa (Cav.) Molero: SPAIN. Castellón, Ares del Maestre, 30TYK4288, 1000 m a.s.l., 24 July 1995, A.de la Torre, M.Vicedo & M.Á.Alonso (ABH 16921); Zorita del Maestrazgo, Cerros de la Gallinera, 30TYL4015, 700 m a.s.l., 1 June 2004, A.Juan, M.Á.Alonso & B.Coca (ABH 55250). Teruel: Castellote, Cuevas de Cañart, El Batán, al pie del Salto de San Juan, 30TYL163158, 900 m a.s.l., 13 August 2015, M.Mart.Azorín & Á.Ortiz Lledó (ABH 72452); Pitarque, camino de la Ermita de San Cristóbal y del nacimiento del río Pitarque, 30TYL00, 1082 m a.s.l., 14 June 2016, borde del sendero, M.Velayos, M.Á.Alonso & al. MV13764 (ABH 75815). Tarragona: inter Cenia et Alcanar, 14 July 1921, P.Font Quer (BC 73662); Gandesa, 450 m a.s.l., 17 July 1921, P.Font Quer (BC 73671).
Sideritis tugiensis S.Ríos, M.B.Crespo & D.Rivera: SPAIN. Granada: Castril, Cerro Laguna-Sierra Seca (Sierra de Segura), 30SWG2799, 1980 m a.s.l., 27 July 1998, S.Ríos, M.B.Crespo, J.L. Solanas & E. Camuñas (holotype: ABH 43003; isotypes: ABH 43004, 43005, 43006, MA, MUB); Castril, Sierra Seca, pr. Cerro Laguna, 30SWH2698, 1900 m a.s.l., 21 August 2000, S.Jury, M.B.Crespo, S.Ríos & J.L.Solanas (ABH 45333); Castril, Cañada de la Sabina, 30SWH2800, 1800 m a.s.l., 13 July 2000, S. Ríos, J.L. Solanas & M.B. Crespo (ABH 43660); ibidem, 24 July 1999, J.L.Solanas, S.Ríos, M.B.Crespo & A.Juan (MA 00805836); Castril, Morro del Pocico o de los Cánovas, 30SWG2697, 2030 m a.s.l., 13 July 2000, S.Ríos, J.L.Solanas & M.B.Crespo (ABH 43658); ibidem, 21-08-00, S.Jury, M.B.Crespo, S.Ríos & J.L.Solanas (ABH 45334); Castril, Morro del Buitre, 30SWG2595, 2130 m a.s.l., 13 July 2000, S.Ríos, J.L.Solanas & M.B.Crespo (ABH 43662); ibidem, 21 August 2000, M.B.Crespo, S.Ríos & J.L.Solanas (ABH 45331); Huéscar, Mojón Alto o Tornajuelos, Sierra Seca, 30SWG2696, 2100 m a.s.l., 13 July 2000, S.Ríos, J.L.Solanas & M.B.Crespo (ABH 43659, MA 00779845, MA 00805995); Huéscar, Torca de la Nieve, 30SWG2595, 2060 m a.s.l., 13 July 2000, S.Ríos, J.L.Solanas & M.B.Crespo (ABH 43661).

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Taxonomy 04 00006 g001
Figure 1. Sideritis royoi (holotype: ABH 83741). (A) Habit; (B) leaves; (C) middle stem indumentum; (D) middle bract; (E) calyx; (F) corolla (frontal and dorsal views). Drawing: L. Sáez.
Figure 1. Sideritis royoi (holotype: ABH 83741). (A) Habit; (B) leaves; (C) middle stem indumentum; (D) middle bract; (E) calyx; (F) corolla (frontal and dorsal views). Drawing: L. Sáez.
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Taxonomy 04 00006 g002
Figure 2. Sideritis royoi (field images in habitat from Caro-La Barcina mountain). Habit (left); inflorescence (above right); basal and middle leaves (bottom right). Photos: R. Curto.
Figure 2. Sideritis royoi (field images in habitat from Caro-La Barcina mountain). Habit (left); inflorescence (above right); basal and middle leaves (bottom right). Photos: R. Curto.
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Taxonomy 04 00006 g003
Figure 3. Distribution map of Sideritis royoi and related species belonging to S. subsect. Hyssopifoliae in the Iberian Peninsula. The areas were established based on bibliographic information [1,11] and herbarium specimens.
Figure 3. Distribution map of Sideritis royoi and related species belonging to S. subsect. Hyssopifoliae in the Iberian Peninsula. The areas were established based on bibliographic information [1,11] and herbarium specimens.
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Taxonomy 04 00006 g004
Figure 4. Habitat for Sideritis royoi in the Caro-La Barcina mountain. Photo: R. Curto.
Figure 4. Habitat for Sideritis royoi in the Caro-La Barcina mountain. Photo: R. Curto.
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Table 1. Main diagnostic morphological characters of Sideritis royoi and related species belonging to Sideritis subsect. Hyssopifoliae.
Table 1. Main diagnostic morphological characters of Sideritis royoi and related species belonging to Sideritis subsect. Hyssopifoliae.
CharacterS. royoiS. hyssopifoliaS. pungensS. tugiensisS. carbonellii
Arrangement of hair covering at base of branchletsGoniotrichousHolotrichous to GoniotrichousGoniotrichous to holotrichousHolotrichousHolotrichous
Base of branchlets: hair length (mm)0.2–1.00.2–1.9(–2.0)0.2–1.5(0.2–)0.4–0.8(0.4–)0.6(–0.8)
Lower leaves: size (mm)6–32 × 3–127–50 × 2–118–50 × 2–412–17 × 2–310–25 × 1–1.5
Lower leaves: shapeSubspatulate to sublinearSpatulate to lanceolateLinearOblanceolateLinear
Lower leaves: marginDentateEntire to serrateEntireDentateEntire
Lower leaves: density of trichomes at main surfaceGlabrous or almost glabrousScarceScarceScarceScarce to very scarce
Lower leaves: arrangement of trichomes at marginClearly distinct when presentClearly distinctClearly distinctIndistinct, except in the basal of leavesClearly distinct
Uppermost leavesNarrower than the lower leaves, subspinescentSimilar to the lowerSimilar to the lower or narrower leavesBract-like, with 0–4 teethSimilar to the lower leaves, exceptionally with 1 tooth
Axillary fascicles during flowering timeUsually absentOccasionally presentOccasionally presentVery frequentAbsent
Number of verticillasters1–2(–3)(1–)2–133–121–2(–4)1–3(–5)
Distance of internode in central verticillasters (mm)2–3.52–123–73–65–6
Shape of the inflorescenceOvoid or globoseCylindrical to globoseCylindricalOvoid or globoseOvoid or globose
Inflorescence length (cm)0.7–2.5(–4)0.8–121–70.7–2.50.5–1.5(–2)
Inflorescence axis: density of glandsAbsentAbsentVery scarceVery abundantAbsent
Lower bract size (mm)7–9 × 6–104–17 × 3–127–12 × 9–124.5–5.5 × 6–85–6 × 4–7
Teeth on each side of the lower bracts3–80–8(–9)4–72–4(–5)2–3(–5)
Lower bracts: greatest width1/3 above the base1/5–1/2 above the base1/8–1/3 above the base1/3 above the base1/2 above the base
Middle bracts size (mm)8–10 × 9–123–11 × 6–116–12 × 8–134–5 × 6–74–5 × 5–6
Number of teeth on each side of middle bracts6–103–8(–9)4–83–62–3(–5)
Abaxial surface of middle bracts: hairinessGlabrousHairyHairyGlabrousGlabrous
Middle bracts: hair length (mm)–0.8–1.50.1–0.50.2–0.70.3–0.5
Calyx length (mm)8–11(5–)6–10(–11)7–86–77–8
Calyx: density of glandsScarceScarce or absentScarceAbundantScarce or abundant
Calyx: density of trichomesScarceScarceVery scarceAbundantScarce
Calyx: length of hairs (mm)0.5–1.81–2.20.6–1.30.5–1.50.9–1.1
Corolla length (mm)6–96–127–98–97–10
Corolla: upper lipEmarginateEntire to emarginateEmarginate or notchedBifidEmarginate or notched
Style length (mm)1.8–2.52–42–53–3.11.8–2.2
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MDPI and ACS Style

Sáez, L.; Curto, R.; Crespo, M.B. Sideritis royoi (Lamiaceae): A New Orophilous Species from Northeastern Spain. Taxonomy 2024, 4, 112-125. https://doi.org/10.3390/taxonomy4010006

AMA Style

Sáez L, Curto R, Crespo MB. Sideritis royoi (Lamiaceae): A New Orophilous Species from Northeastern Spain. Taxonomy. 2024; 4(1):112-125. https://doi.org/10.3390/taxonomy4010006

Chicago/Turabian Style

Sáez, Llorenç, Rafel Curto, and Manuel B. Crespo. 2024. "Sideritis royoi (Lamiaceae): A New Orophilous Species from Northeastern Spain" Taxonomy 4, no. 1: 112-125. https://doi.org/10.3390/taxonomy4010006

APA Style

Sáez, L., Curto, R., & Crespo, M. B. (2024). Sideritis royoi (Lamiaceae): A New Orophilous Species from Northeastern Spain. Taxonomy, 4(1), 112-125. https://doi.org/10.3390/taxonomy4010006

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