Article

Sideritis royoi (Lamiaceae): A New Orophilous Species from

Northeastern Spain †
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; [email protected]
3 Department of Environment Sciences and Natural Resources (dCARN), University of Alicante, P.O. Box 99,
ES-03080 Alicante, Spain; [email protected]
* Correspondence: [email protected]
† IPNI: urn:lsid:ipni.org:names:77334707-1.

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.

Keywords: endemism; orophytes; taxonomy; morphology; Mediterranean Basin; Iberian Peninsula

Citation: Sáez, L.; Curto, R.; Crespo,

M.B. Sideritis royoi (Lamiaceae): A New 1. Introduction
Orophilous Species from Northeastern Sideritis L. is a large genus of family Lamiaceae Martynov, nom. cons., that includes
Spain. Taxonomy 2024, 4, 112–125. nearly 150 species that are mainly distributed in Europe, North Africa, Macaronesia, and
https://doi.org/10.3390/ Western and Central Asia [1–4], though some of these species have been introduced or
taxonomy4010006 naturalised in other areas. The Port Massif (northeastern Spain, Tarragona Province) ex-
Academic Editor: Germinal Rouhan hibits 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.
Received: 11 December 2023 caught our attention due to a combination of characteristics that do not match those of any
Revised: 5 January 2024
described species within the genus. In the upper areas of the Port Massif, the presence of
Accepted: 16 January 2024
plants of the genus Sideritis, whose identity is uncertain, dates back more than 40 years.
Published: 18 January 2024
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
Copyright: © 2024 by the authors.
authors of this paper (L.S. and R.C.), tentatively attributed those plants to S. hyssopifolia
Licensee MDPI, Basel, Switzerland. L. while noting some discordant morphological characters. The most recent reference
This article is an open access article for the Sideritis occurring in the upper parts of the Port Massif is the work of Sáez and
distributed under the terms and Aymerich [8], who, commenting on S. hyssopifolia, indicated that the taxonomic identity of
conditions of the Creative Commons those populations was unclear.
Attribution (CC BY) license (https:// In order to clarify that point, during field trips conducted between 2021 and 2023, we
creativecommons.org/licenses/by/ gathered new specimens of the Sideritis from the Port Massif which had doubtfully been
4.0/). attributed to S. hyssopifolia. After studying the material collected, herbarium specimens,

Taxonomy 2024, 4, 112–125. https://doi.org/10.3390/taxonomy4010006 https://www.mdpi.com/journal/taxonomy

Taxonomy 2024, 4 113

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 biogeograph-
ical 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 recog-
nition 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. Figures 1 and 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
 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
Taxonomy 2024, 4 114
greenish, with sparse sessile glands and glandular hairs up to 0.1 mm long abundant an-
trorse 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
basal third; divided into 1/3–1/2 of its width; adaxial surface glabrous, abaxial with sparse
glabrous, abaxial with sparse (sometimes abundant) glandular hairs up to 0.1 mm long
(sometimes abundant) glandular hairs up to 0.1 mm long and always lacking sessile glands;
and always lacking sessile glands; eglandular hairs usually absent, sometimes present
eglandular hairs usually absent, sometimes present (very scarce) at the margins, 0.2–0.5 mm
(very scarce) at the margins, 0.2–0.5 mm long; teeth 1–3(–4) long, narrowly triangular to
long; teeth 1–3(–4) long, narrowly triangular to subulate; lower bracts 7–9 × 6–10 mm, with
subulate; lower bracts 7–9 × 6–10 mm, with 3–8 teeth on each side; middle bracts 8–10 × 9–
3–8 teeth on each side; middle bracts 8–10 × 9–12 mm, divided to 1/3–1/2 of its width,
12 mm, divided to 1/3–1/2 of its width, with 6–10 teeth on each side. Six-flowered verticil-
with 6–10 teeth on each side. Six-flowered verticillasters. Calyx campanulate, 8–11 mm
lasters. Calyx campanulate, 8–11 mm long (calyx tube 4–5.2 mm long), with five subequal
long (calyx tube 4–5.2 mm long), with five subequal divergent teeth, 4–8 mm long, ending
divergent teeth, 4–8 mm long, ending in spines 0.7–2 mm long; the outer surface with
in spines 0.7–2 mm long; the outer surface with scarce to abundant glandular hairs up to
scarce to abundant glandular hairs up to 0.15 mm long, sessile glands and antrorse eglan-
0.15 mm long, sessile glands and antrorse eglandular hairs 0.5–1.8 mm long; carpostegium
dular hairs 0.5–1.8 mm long; carpostegium (ring of hairs within calyx) discontinuous. Co-
(ring of hairs within calyx) discontinuous. Corolla yellowish, 6–9 mm long, with tube 3–4,
rolla yellowish, 6–9 mm long, with tube 3–4, 4 mm long, subcylindrical; upper lip bifid up
4 mm long, subcylindrical; upper lip bifid up to 1/5–1/3 of its length; stamens included
to the
in 1/5–1/3 of its
corolla length;
tube, withstamens included0.3–0.5
short filaments in the mm
corolla tube,
long; with
style shortmm
1.8–2.5 filaments 0.3–0.5
long. Nutlets
mm long; style 1.8–2.5
ovoid, c. 2 × 1.5 mm. mm long. Nutlets ovoid, c. 2 × 1.5 mm.

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.
4, FOR PEER REVIEW 4

Taxonomy 2024, 4 115

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 2. Sideritis royoi (field images in habitat from Caro-La Barcina mountain). Habit (left); inflo-
Figure 2. Sideritis royoi (field images in habitat from Caro-La Barcina mountain). Habit (left);
rescence (above right); basal and middle leaves (bottom right). Photos: R. Curto.
inflorescence (above right); basal and middle leaves (bottom right). Photos: R. Curto.

Eponymy: The new speciesThe

Eponymy: is dedicated
new speciesto is
the late Ferran
dedicated Royo
to the Pla (1969−2016)
late Ferran for his
Royo Pla (1969–2016) for his
outstanding workoutstanding
that contributed to improving
work that contributed to theimproving
botanicalthe knowledge of the PortofMas-
botanical knowledge the Port Massif.
sif. Phenology: Flowering is from June to early July; fruiting is from late July to late August.
Phenology: FloweringDistribution
is from and ecology:
June Sideritis
to early royoi
July; is an orophilous
fruiting species
is from late Julyendemic to a small area
to late Au-
gust. 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
Distribution and ecology: Sideritis royoi is an orophilous species endemic to a small area
the northern face of the Caro-La Barcina mountain (Figure 4), between 1080 and 1325 m a.s.l.
in the Port Massif (Figure 3) in the northeastern Iberian Peninsula. The new species is
The known sites of the new species show a pluviseasonal Mediterranean bioclimate, mostly
found in the rocky slopes
within theand in patches in the
Supramediterranean understory
thermotype andof theaopen
with Pinusorsylvestris
Subhumid L.
locally Lower-Humid
forests at the northern face of the Caro-La Barcina mountain (Figure 4), between 1080 and
ombrotype [14]. Biogeographically, this area belongs to the Puertobeceitan-Morellan District
1325 m a.s.l. The known sites of the newsubsector
of the Western-Catalanid species show a pluviseasonal Mediterranean
(Valencian-Tarraconensian bi-
Sector, Catalan-Provencian-
Balear Province,
oclimate, mostly within Mediterranean Region),
the Supramediterranean according
thermotype andtowithRivas-Martínez
a Subhumid[15]. The Caro-La
or lo-
cally Lower-Humid ombrotype [14]. Biogeographically, this area belongs to the Puerto- that are
Barcina mountain hosts a rich flora including a high number of plant species
beceitan-Morellanendemic
Districttoofthe
thenortheastern Iberian Peninsula
Western-Catalanid subsector [5], and some of them are restricted to the
(Valencian-Tarraconensian
Port Massif. Font Quer [19] first noticed that several plant species occurring in the Port
Sector, Catalan-Provencian-Balear Province, Mediterranean Region), according to Rivas-
Massif usually showed conspicuous morphological differences compared with other related
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
 Taxonomy 2024, 4 116

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
Taxonomy 2024, 4, FOR PEER REVIEW such as Arctostaphylos uva-ursi (L.) Spreng., Erinacea anthyllis Link subsp. anthyllis, Festuca 5
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
compared
criterionwith other
D [16]. related
This speciestaxa
hasoccurring
an extent ofinoccurrence
other areas of an
and thearea
northeastern
of occupancyIberian
of Pen-
insula. km2 ,new
0.75The calculated
species 0.5 × 0.5
on agrows km grid.with
together Based on the data
several currently available, we have
Puertobeceitan-Morellan endemics,
suchnoasevidence
Aquilegiaof apaui
population decline.Arenaria
Font Quer, However,conimbricensis
further field studies areviridis
subsp. needed(Font
to define a
Quer) Font
more accurate conservation category for this species.
Quer, Knautia rupicola (Willk.) Font Quer, or Thymus willkommii Ronniger, as well as more
Additional specimens examined: SPAIN. Tarragona Province: Alfara de Carles, prop de
widely
les Clotes, 1080 morophilous
distributed a.s.l., 10 Julyspecies such(MMA);
1982, L.Torres as Arctostaphylos uva-ursi
per davall del Pas de (L.) Spreng.,
la Barcina, caraErinacea
anthyllis
Nord,Link
1200subsp.
m a.s.l.,anthyllis, Festuca
14 July 2009, trichophylla
Aparicio, (Gaudin)
Beltran, Curto, MesaK.Richt., and(MMA);
& Royo 6085 TeucriumLa aureum
Schreb. subsp.
Barcina, aureum.
matollar, 31TBF72, 1270 m a.s.l., 25 July 2021, R.Curto (L. Sáez pers. herb.).

Figure 3. Distribution
Figure map
3. Distribution mapofofSideritis royoiand
Sideritis royoi and related
related species
species belonging
belonging to S. subsect.
to S. subsect. Hyssopifoliae
Hyssopifoliae
in the
in Iberian Peninsula.
the Iberian Peninsula.The
Theareas wereestablished
areas were established based
based on bibliographic
on bibliographic information
information [1,11] and[1,11] and
herbarium specimens.
herbarium specimens.

Conservation status: Based on the present state of knowledge, due to its low popula-
tion 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
Taxonomy 2024, 4,
Taxonomy FOR
2024, 4 PEER REVIEW 117 6

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.

4. Discussion
4. Discussion
Sideritis
Sideritis royoishows
royoi shows morphological
morphological affinities
affinitieswith
withtaxa
taxaof S.
of sect. Sideritis
S. sect. subsect.
Sideritis subsect.
Hyssopifoliae Obón & D.Rivera and is assumed to belong here based on its
Hyssopifoliae Obón & D.Rivera and is assumed to belong here based on its hair covering hair covering at at
the base of its goniotrichous (or slightly holotrichous) branchlets with antrorse trichomes [1].
the base of its goniotrichous (or slightly holotrichous) branchlets with antrorse trichomes
Most of the taxa belonging to this group are found in mountain areas of the Iberian
[1].Peninsula
Most of and
the Northern
taxa belonging to this group are found in mountain areas of the Iberian
Africa, with a higher concentration in Southern and Eastern Spain.
Peninsula and Northern Africa,
Some of these species are orophyte with a higher
specialists concentration
that in Southern
are narrowly endemic and Eastern
to a reduced
Spain. Some range
mountain of these species
[1,11]. are orophyte
A comparison of S.specialists
royoi with that are narrowly
morphologically endemic
related to a re-
species
duced mountain
belonging range [1,11].
to S. subsect. A comparison
Hyssopifoliae is shown ofin S. royoi
Table 1. with morphologically related spe-
cies belonging to S. subsect. Hyssopifoliae is shown in Table 1.
Table 1. Main diagnostic morphological characters of Sideritis royoi and related species belonging to
Sideritis
Table subsect.
1. Main Hyssopifoliae.
diagnostic morphological characters of Sideritis royoi and related species belonging to
Sideritis subsect. Hyssopifoliae.
Character S. royoi S. hyssopifolia S. pungens S. tugiensis S. carbonellii
Character
Arrangement of S. royoi S. hyssopifolia S. pungens S. tugiensis S. carbonellii
Holotrichous to Goniotrichous to
Arrangement of hairatcoveringGoniotrichous
hair covering Holotrichous to Goniotrichous to Holotrichous Holotrichous
base of branchlets
Goniotrichous Goniotrichous holotrichous Holotrichous Holotrichous
at base of branchlets Goniotrichous holotrichous
Base of Base
branchlets: hair length
of branchlets:
0.2–1.0
0.2–1.0 0.2–1.9(–2.0)
0.2–1.9(–2.0) 0.2–1.5
0.2–1.5 (0.2–)0.4–0.8
(0.2–)0.4–0.8 (0.4–)0.6(–0.8)
(0.4–)0.6(–0.8)
(mm) hair length (mm)
Lower leaves: size (mm)
Lower leaves: size 6–32 × 3–12
6–32 × 3–12
7–50 × 2–11
7–50 × 2–11
8–50 × 2–4
8–50 × 2–4
12–17 × 2–3 10–2510–25
12–17 × 2–3 × 1–1.5
× 1–1.5
(mm) Subspatulate to sub- Spatulate to lanceo-
Lower leaves: shape Linear Oblanceolate Linear
Lower leaves: Subspatulate to
linear Spatulate late
to
Linear Oblanceolate Linear
shape margin
Lower leaves: sublinear
Dentate lanceolate
Entire to serrate Entire Dentate Entire
Lower leaves:
Lower leaves: density of tri- GlabrousDentate or almost
Entire to serrate
Scarce EntireScarce Dentate
Scarce Entireto very scarce
Scarce
chomesmargin
at main surface glabrous
Indistinct, except
Lower leaves: arrangement of Clearly distinct
Clearly distinct Clearly distinct in the basal of Clearly distinct
trichomes at margin when present
leaves
Narrower than the Similar to the Similar to the lower
Bract-like, with 0–4
Uppermost leaves lower leaves, subspi- Similar to the lower lower or narrower leaves, exceptionally
teeth
nescent leaves with 1 tooth
Axillary fascicles during flow- Occasionally pre- Occasionally pre-
Usually absent Very frequent Absent
Taxonomy 2024, 4 118

Table 1. Cont.

Character S. royoi S. hyssopifolia S. pungens S. tugiensis S. carbonellii

Lower leaves:
density of Glabrous or almost Scarce to very
Scarce Scarce Scarce
trichomes at main glabrous scarce
surface
Lower leaves:
Indistinct, except
arrangement of Clearly distinct
Clearly distinct Clearly distinct in the basal of Clearly distinct
trichomes at when present
leaves
margin
Similar to the
Narrower than the Similar to the
Similar to the Bract-like, with 0–4 lower leaves,
Uppermost leaves lower leaves, lower or narrower
lower teeth exceptionally with
subspinescent leaves
1 tooth
Axillary fascicles
Occasionally Occasionally
during flowering Usually absent Very frequent Absent
present present
time
Number of
1–2(–3) (1–)2–13 3–12 1–2(–4) 1–3(–5)
verticillasters
Distance of
internode in
2–3.5 2–12 3–7 3–6 5–6
central
verticillasters (mm)
Shape of the Cylindrical to
Ovoid or globose Cylindrical Ovoid or globose Ovoid or globose
inflorescence globose
Inflorescence
0.7–2.5(–4) 0.8–12 1–7 0.7–2.5 0.5–1.5(–2)
length (cm)
Inflorescence axis:
Absent Absent Very scarce Very abundant Absent
density of glands
Lower bract size
7–9 × 6–10 4–17 × 3–12 7–12 × 9–12 4.5–5.5 × 6–8 5–6 × 4–7
(mm)
Teeth on each side
3–8 0–8(–9) 4–7 2–4(–5) 2–3(–5)
of the lower bracts
Lower bracts: 1/5–1/2 above the 1/8–1/3 above the
1/3 above the base 1/3 above the base 1/2 above the base
greatest width base base
Middle bracts size
8–10 × 9–12 3–11 × 6–11 6–12 × 8–13 4–5 × 6–7 4–5 × 5–6
(mm)
Number of teeth
on each side of 6–10 3–8(–9) 4–8 3–6 2–3(–5)
middle bracts
Abaxial surface of
middle bracts: Glabrous Hairy Hairy Glabrous Glabrous
hairiness
Middle bracts: hair
– 0.8–1.5 0.1–0.5 0.2–0.7 0.3–0.5
length (mm)
Taxonomy 2024, 4 119

Table 1. Cont.

Character S. royoi S. hyssopifolia S. pungens S. tugiensis S. carbonellii

Calyx length (mm) 8–11 (5–)6–10(–11) 7–8 6–7 7–8
Calyx: density of
Scarce Scarce or absent Scarce Abundant Scarce or abundant
glands
Calyx: density of
Scarce Scarce Very scarce Abundant Scarce
trichomes
Calyx: length of
0.5–1.8 1–2.2 0.6–1.3 0.5–1.5 0.9–1.1
hairs (mm)
Corolla length
6–9 6–12 7–9 8–9 7–10
(mm)
Entire to Emarginate or Emarginate or
Corolla: upper lip Emarginate Bifid
emarginate notched notched
Style length (mm) 1.8–2.5 2–4 2–5 3–3.1 1.8–2.2

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. Fruticu-
losae 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 [23–26], our results indicate that the
Taxonomy 2024, 4 120

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 inflores-
cences 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 recog-
nised, 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 tri-
chomes 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
Taxonomy 2024, 4 121

9. Lower leaves entire, 10–25 × 1–1.5 mm, axillary fascicles absent at flowering time; up-
permost 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 flower-
ing 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.
Taxonomy 2024, 4 122

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,
Taxonomy 2024, 4 123

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: Tar-
tanedo, 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
Taxonomy 2024, 4 124

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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