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Pilosella argyrogaliciana, P. argyrolegionensis and P. megargyrocoma (Asteraceae): three new species from the Cantabrian Mountains (N. Spain)/Pilosella argyrogaliciana, P. argyrolegionensis y P. megargyrocoma (Asteraceae): tres nuevas especies de la Cordillera Cantdbrica (N Espana).

INTRODUCTION

This work is a continuation of the intense study we are making in the genus Pilosella in the area of the Cantabrian Mountains (Mateo & del Egido, 2007, 2010, 2011, 2012a, 2012b; del Egido & Mateo, 2012) and takes place in the context of the monographic revision of this genus in the Iberian Peninsula started years ago (Mateo, 1988). This monographic revision reached a relatively mature synthesis in more recent times (Mateo, 2006) and its results are included in the large volume 2 of Med-Checklist (Greuter & Raabe-Straube, 2008).

In our exhaustive tours through the Cantabrian Mountains del Egido & Puente, 2011a, b; del EGIDO & al., 2012 in search of the diverse representation of species of Pilosella, we detected some important populations of P. argyrocoma and two of its intermediate species: P. nevadensis (argyrocoma <> saussureoides) and P. subulatissima (argyrocoma <> pseudopilosella) (Mateo & del Egido, 2011). In the environment of P. argyrocoma we also detected three new intermediate species that we describe and comment here.

Pilosella argyrocoma is a basic species of the sect. Pilosellina and it is well characterized basically by having monocephalic scapes, short and relatively thick (or absent) stolons, dense white stellate trichomes on both adaxial and abaxial surfaces of the leaves and abundant long simple eglandular and stellate trichomes on involucral bracts and scapes. It is an endemic species to the S, C and NW Iberian Peninsula and N Africa (MATEO & DEL EGIDO, 2011). There are only three intermediate species described which are thought to have resulted from hybridisation of P. argyrocoma: P. aranii (argyrocoma <> vahlii), only known in its classic locality in the Sierra de Guadarrama (Mateo 1997; 2006); P. nevadensis (argyrocoma <> saussureoides), known in the Sierra Nevada and its surroundings, the Central System and a pair of localities in the Cantabrian Mountains (Mateo, 1997; 2006; Mateo & del Egido, 2011); and P. subulatissima (argyrocoma <> pseudopilosella), only known in its classic locality in the Sierra del Cuarto (Granada), the Sierra de Guadarrama and three localities in the Cantabrian Mountains (Mateo, 2006; Mateo & del Egido, 2011).

The taxonomic treatment of the genus follows BRAUTIGAM & GREUTER (2007; 2008) (see also DEL EGIDO & MATEO, 2012) and the nomenclature of taxa cited can be consulted in BRAUTIGAM & GREUTER (2008). After the names of the intermediate taxa, we indicate (in parentheses) the species that are most probably responsible for their origin.

RESULTS

Pilosella argyrogaliciana Mateo & Egido spec. nova (argyrocoma <> galiciana) (Figure 1)

Holotypus: Spain. Leon, Carmenes, Canseco, pr. Pico Morala, 30TTN974136415, 2100 m, pastizal psicroxerofilo acidofilo, 28.VI.2009, F. del Egido, (holotype: LEB 103233).--Paratype: Spain. Leon, Valdelugueros, Redilluera, Sierra de Portillas, 30TTN9888863868, 1985 m, pastizal psicroxerofilo en zona de mezcla de sustratos, 28.VI.2009, F. del Egido, LEB 103229.

Diagnosis: Plantae rosulatae, stolonibus brevibus vel nullis. Folia ad 2-6 x 0.4-1.7 cm, elliptica oblonga vel oblaceolata, obtuso-mucronata, attenuata, subcoriacea, supra viridia, laxe subrigido-pilosa et floccosa, subtus laxe vel dense cano-floccosa,pilosa, eglandulosa. Scapi (2.5) 615 (18) cm x 1-2 mm alti, monocephali, modice floccosi, ad apicem pilosi et glandulosi. Involucra 7-12 x 6-11 mm. Bracteae lanceolato-lineares, 1 mm latae, acutae, dense pilosiae, modice floccosae et glandulosae.

Description: Perennial herb. Phyllopodous. Stolons absent or if present, scarce, short (up to 4 cm in studied specimens) and stout, with leaves smaller than those of the rosette. Rosette-leaves 2-6 x 0.4-1.7 cm; entire; elliptical, oblong, oblanceolate or linear-oblanceolate; rounded-obtuse or slightly mucronate at apex and gradually narrowing towards base; adaxial surface green or lightly greyish, with some subrigid long simple eglandular trichomes and, in some leaves, with few to numerous stellate trichomes (absent in others); abaxial surface green to greyish-green or whitish, with few (or even absent in some leaves) to numerous (variable from one leaf to another) stellate trichomes and some long simple eglandular trichomes thinner than those of the adaxial surface; without glandular trichomes. Scapes (2.5) 6-15 (18) cm tall and 1-2 mm in diameter, each with a single capitulum in studied specimens (it may have more), with stellate, glandular and simple eglandular trichomes in variable quantities, the most abundant being the stellate ones along the whole length of the stem whereas the simple eglandular and glandular ones are generally scarce towards base becoming more abundant towards apex. Involucre 7-12 x 6-11 mm. Involucral bracts linear-lanceolate, acute, [+ or -] 1 mm wide; with stellate, glandular and long simple eglandular trichomes in more or less similar proportions (generally, the eglandular trichomes are the most abundant). Ligules yellow, the outer with a dark red stripe on outer face.

Affinities: P. argyrogaliciana occupies an intermediate morphological position between P. argyrocoma and P. galiciana, and the influence of both species is very clear and unequivocal. Its principal differences from those are presented in Table 1. It is evident that P. argyrogaliciana has resulted from hybridisation of a species of the sect. Pilosellina (it has monocephalic scapes, stellate trichomes on abaxial surface of leaves, etc.) and a species of the sect. Auriculina (it has some leaves with very few, or absent, stellate trichomes on abaxial surface of leaves). The species of the sect. Pilosellina must have stellate trichomes also on adaxial surface of leaves, stolons absent or, if present, scarce, short and relatively thick and thin involucral bracts with abundant and dominant long simple eglandular trichomes and more or less abundant stellate trichomes: in this area it can only be P. argyrocoma. The species of the sect. Auriculina must have stolons absent or, if present, scarce, short and stout, leaves without glandular trichomes, and thin involucral bracts with glandular (generally the most abundant), stellate and simple eglandular trichomes in similar proportions: in this area it can only be P. galiciana.

Furthermore, as P. galiciana and P. vahlii are very similar, the most similar species to P. argyrogaliciana is P. aranii (argyrocoma <> vahlii). The basic difference between these two species is that P. aranii presents some glandular trichomes on the leaves due to the influence of P. vahli. The other species which are thought to have resulted from hybridisation of P. argyrocoma: P. nevadensis (argyrocoma <> saussureoides), P. subulatissima (argyrocoma <> pseudopilosella) and the two species described below in this paper, are less similar. P. argyrogaliciana differs clearly from all of them, among other characters, because it has abaxial surface of some leaves with few or absent stellate trichomes due to the influence of a species of the sect. Auriculina (these other four species have abaxial surface of all leaves with dense stellate trichomes).

Distribution: P. argyrogaliciana is endemic to the NW Iberian Peninsula. It must be very rare because P. galiciana is a very local and scarce plant only present in mountainous areas of the NW Iberian Peninsula and P. argyrocoma is very rare in the NW Iberian Peninsula.

Pilosella argyrolegionensis Mateo & Egido spec. nova (argyrocoma <> officinarum) (Figure 2)

Holotypus: Spain. Leon, Valdelugueros, Redi lluera, Sierra de Portillas, 30TTN9888863868, 1985 m, pastizal psicroxerofilo en zona de mezcla de sustratos, 28.VI.2009, F. del Egido (holotype LEB 103231; isotype VAL).--Paratype: Spain. Leon, San Emiliano-Sena de Luna, Riolago de Babia, Pico La Ferrera, 29TQH3717253588, 2110 m, pastizal psicroxerofilo acidofilo, 28.VII.2009, F. del Egido & E. Puente (LEB 82159; VAL).

Diagnosis: Plantae rosulatae, stolonibus brevibus vel [+ or -] elongatis (0.5-8 cm), foliisparvis ob siti. Folia ad (1.5) 3-5 (7) x 0.6-2 cm, elliptica oblonga vel oblaceolata, obtuso-mucronata, attenuata, supra viridia, laxe subrigido-pilosa et floccosa, subtus laxe vel dense cano-floccosa, pilosa, eglandulosa. Scapi 4-20 cm x 1-1.5 mm alti, monocephali, modice floccosi, ad apicem pilosi et glandulosi. Involucra 8-13 x 6-11 mm. Bracteae lanceolato-lineares, 1 mm latae, acutae, modice pilosae floccosae et glandulosae.

Description: Perennial herb. Phyllopodous. Stolons scarce, very short to relatively long (0.5-8 cm in the studied specimens), with leaves smaller than those of the rosette. Rosette-leaves (1.5) 3-5 (7) x 0.6-2 cm; entire; elliptical, oblong, oblanceolate or linear-oblanceolate; rounded-obtuse and slightly mucronate at apex, gradually narrowing towards base; adaxial surface green or greyish, with some subrigid long simple eglandular trichomes and, in some leaves, with few to numerous stellate trichomes (absent in others); abaxial surface whitish, with dense stellate trichomes and some long simple eglandular trichomes thinner than those of the adaxial surface. Scapes thin (1-1.5 mm in diameter); 4-20 cm tall; each with a single capitulum; with stellate, glandular and long simple eglandular trichomes in variable quantities (the stellate trichomes are the most abundant along whole length of the stem followed by the glandular trichomes, which become more abundant towards apex, whereas the simple trichomes appear more or less scattered along whole length of the stem but are more abundant at base and apex). Involucre 8-13 x 6-11 mm. Involucral bracts linear-lanceolate, acute, [+ or -] 1 mm wide; with stellate, glandular and simple eglandular trichomes in more or less similar proportions. Ligules yellow, the outer with a dark red stripe on outer face.

Affinities: P. argyrolegionensis occupies an intermediate morphological position between P. argyrocoma and P. officinarum, and the influence of both species is very clear and unequivocal. Its principal differences from those are presented in Table 2. It is evident that P. argyrogaliciana has resulted from hybridisation of two species of the sect. Pilosellina (it has monocephalic scapes, dense stellate trichomes on abaxial surface of all leaves, etc.). One of them must have stellate trichomes also on adaxial surface of leaves, stolons absent or, if present, scarce, short and relatively thick and thin involucral bracts with abundant and dominant long simple eglandular trichomes and more or less abundant stellate trichomes: in this area it can only again be P. argyrocoma. The other species must have abundant, elongated and thin stolons and thin involucral bracts with abundant and dominant black glandular trichomes: in this area it can only be P. officinarum.

P. argyrolegionensis is similar to the other species which are thought to have resulted from hybridisation of P. argyrocoma and other species of the sect. Pilosellina: P. nevadensis (argyrocoma <> saussureoides), and P. subulatissima (argyrocoma <> pseudopilosella). P. argyrolegionensis differs from both of them basically by the involucral bracts and apex of scapes with abundant black glandular trichomes due to the influence of P. officinarum (the involucral bracts and apex of scapes of these other two species have very scarce glandular trichomes, or they may even be absent). The abaxial surface with dense stellate trichomes on all leaves, among other characters, distinguishes this species from P. argyrogaliciana (described above) and from P. aranii (argyrocoma <> vahlii).

Distribution: P. argyrolegionensis is only known in two localities in the Cantabrian Mountains. It can also occur in some areas of the C and S Iberian Peninsula although P. officinarum is very rare in these areas.

Pilosella megargyrocoma Mateo & Egido spec. nova (argyrocoma <> peleteriana) (Figure 3)

Holotypus: Spain. Leon, Carrocera, Piedrase cha, Collado del Fito, 30TTN7443747896, 1670 m, brezal, 14.VII.2009, F. del Egido (holotype LEB 103251; isotype VAL 202513).--Paratype: Spain. Leon, San Emiliano-Sena de Luna, Riolago de Babia, La Ferrera peak, 29TQH371535, 2110 m, pastizal psicroxerofilo acidofilo, 28.VII.2009, F. del Egido & E. Puente, LEB 107301.

Diagnosis: Plantae rosulatae, stolonibus brevibus vel nullis. Folia ad4-11 x 0.8-2.5 cm, elliptica oblonga vel oblanceolata, obtusa vel obtuso-mucronata, attenuata, supra modice vel laxe floccosa, laxe subrigido-pilosa, subtus densiore cano-floccosa, modicepilosa. Scapi 30-40 (55) cm x 1.5-2 mm alti, monocephali, modice floccosi pilosi et glandulosi. Involucra magna, 13-15 x 10-14 mm. Bracteae lanceolato-lineares, 1.2-1.6 mm latae, acutae, dense pilosae, modice floccosae, laxe glandulosae.

Description: Perennial herb. Phyllopodous. Stolons absent or if present, scarce, relatively short (up to 9 cm in studied specimens) and stout, with leaves a little smaller than those of the rosette. Rosette-leaves 4-11 x 0.8-2.5 cm; entire; elliptical, oblong or oblanceolate; rounded-obtuse or slightly mucronate at apex, gradually narrowing towards base; adaxial surface green or greyish, with some very long subrigid (4-7 mm) simple eglandular trichomes and, in some leaves, with few to numerous stellate trichomes (absent in others); abaxial surface whitish, with dense stellate trichomes and some long simple eglandular trichomes thinner than those of the adaxial surface. Scapes stout (1.52 mm in diameter); 30-40 (55) cm tall; each with a single capitulum; with stellate, glandular and very long simple eglandular trichomes in more or less similar proportions. Involucre 13-15 x 10-14 mm. Involucral bracts linear-lanceolate, acute, 1.21.6 mm wide; with dominant long greyish (black at base) simple eglandular trichomes that cover other quite abundant stellate trichomes and some occasional glandular trichomes.

Affinities: P. megargyrocoma occupies an intermediate morphological position between P. argyrocoma and P. peleteriana, and the influence of both species is very clear and unequivocal. Its principal differences from those are presented in Table 3. As with P. argyrolegionensis, it is evident that P. megargyrocoma has resulted from hybridisation of two species of the sect. Pilosellina and one of them must again be P. argyrocoma for the same reasons. The other species must have scarce, short or not very long and stout stolons with leaves similar than those of the rosette, thicker scapes and capitula, and wider involucral bracts with dominant long simple eglandular trichomes: in this area it can only be P. peleteriana.

P. megargyrocoma differs clearly from all the other species which are thought to have resulted from hybridisation of P. argyrocoma because it is more robust, with thicker scapes and capitula, wider involucral bracts, larger leaves of the stolons, among other characters.

Distribution: P. megargyrocoma is only known in two localities in the Cantabrian Mountains. It can also occur in some areas of the center of the Iberian Peninsula although P. peleteriana is very rare in these areas.

doi: 10.5209/rev_LAZA.2013.v34.n1.40357

Received: 25 October 2012

Accepted: 8 April 2013

REFERENCES

Brautigam, S. & Greuter, W.--2007--A new treatment of Pilosella for the Euro-Mediterranean flora [Notulae ad floram euro-mediterraneam pertinentes No. 24]--Willdenow. 37: 123-137.

Brautigam, S. & Greuter, W.--2008--Pilosella Vaill.--In: Greuter, W. & Raabe-Straube E. (Eds.). Med-Checklist. A critical inventory of vascular plants of the circum-mediterranean countries, 2. Dicotyledones (Compositae). Pp. 559-674. OPTIMA, Palermo, Geneve & Berlin.

Del Egido, F. & Puente, E.--2011a--Valeriano apulae-Potentilletum nivalis y Festuco eskiae-Cytisetum oromediterranei: dos nuevas comunidades vegetales de la alta montana cantabrica--Lazaroa 32: 91-99.

Del Egido, F. del & Puente, E.--2011b--Sobre los nombres correctos de dos asociaciones homonimas: Epipactido palustris-Eriophoretum latifolii--Lazaroa 32: 181-182.

Del Egido, F., Fernandez, M., Ferreras, N., Puente, E. & Lopez, M.J.--2012--Notas sobre flora leonesa amenazada, II--Lazaroa 33: 207-216.

Del Egido, F. & Mateo, G.--2012--Three new species of Pilosella (Asteraceae) from the Cantabrian Mountains, N Spain--Ann. Bot. Fenn 49: 271-278.

Greuter, W. & Raabe-Straube, E. (eds.)--2008--Med-Checklist. A critical inventory of vascular plants of the circum-mediterranean countries, 2. Dicotyledones (Compositae)--OPTIMA, Palermo, Geneve & Berlin.

Mateo, G.--1988--Sobre las especies del genero Hieracium sect. Pilosellina Fr.--Lagascalia 15 (extra): 369-376.

Mateo, G.--1997--Sobre Pilosella argyrocoma (Fr.) F.W. Schultz & Schultz Bip. y especies relacionadas--Flora Montiberica 7: 67-71.

Mateo, G.--2006--Aportaciones al conocimiento del ge nero Pilosella Hill en Espana, VII. Revision sintetica--Flora Montiberica 32: 51-71.

Mateo, G. & del Egido, F.--2007--Novedades del genero Pilosella Hill (Compositae) en los montes astur-leoneses --Flora Montiberica 37: 3-16.

Mateo, G. & del Egido, F.--2010--Novedades del genero Pilosella Hill (Compositae) en los montes astur-leoneses, II--Flora Montiberica 44: 72-79.

Mateo, G. & del Egido, F.--2011--Novedades del genero Pilosella Hill (Compositae) en los montes astur-leoneses, III--Flora Montiberica 48: 38-51.

Mateo, G. & del Egido, F.--2012a--Novedades del genero Pilosella Hill (Compositae) en los montes astur-leoneses, IV--Flora Montiberica 52: 78-84.

Mateo, G. & del Egido, F.--2012b--Pilosella lactocanta brica (Asteraceae, Cichorieae), a new species from the Cantabrian Mountains (N Spain), and two new related species--Willdenowia 42: 181-189.

Fermin del Egido Mazuelas, Departamento de Biodiversidad y Gestion Ambiental (Area de Botanica). Facultad de CC Biologicas y Ambientales. Campus de Vegazana. Universidad de Leon. E-24071 Leon. Spain. E-mail: fegim@unileon.es

Gonzalo Mateo Sanz, Jardin Botanico e Instituto Cavanilles de Biodiversidad y Biologia Evolutiva. Universidad de Valencia. C/Quart, 80. E46008-Valencia. Spain. E-mail: gonzalo.mateo@uv.es.

Table 1
Principal morphological differences between Pilosella argyrocoma,
P. argyrogaliciana sp. nova and P. galiciana.

Characters           P. argyrocoma             P. argyrogaliciana

Number of capitula   1                         1
Stellate trichomes   abundant on both          few to numerous on
on leaves            surfaces of all leaves    adaxial surface of some
                                               leaves and absent on
                                               others; few to numerous
                                               on abaxial surface of
                                               some leaves and absent
                                               on others

Indumentum of        very dense, with          very dense, with
involucral bracts    dominant long simple      stellate, glandular and
                     eglandular trichomes,     long simple eglandular
                     [+ or -] abundant         trichomes in more or
                     stellate trichomes and    less similar proportions
                     very scarce or absent     (generally simple
                     glandular trichomes       eglandular ones most
                                               abundant)

Characters           P. galiciana

Number of capitula   1-4 (6)
Stellate trichomes   absent
on leaves

Indumentum of        not very dense, with
involucral bracts    glandular, stellate and
                     simple eglandular
                     trichomes in [+ or -] similar
                     proportions (generally
                     glandular ones most
                     abundant)

Table 2
Principal differences between Pilosella argyrocoma, P.
argyrolegionensis sp. nova and P. officinarum.

Characters           P. argyrocoma             P. argyrolegionensis

Stolons              absent or, if present,    scarce, very short to
                     scarce, short and         relatively long
                     relatively thick

Stellate trichomes   abundant on both          few to numerous on
on leaves            surfaces of all leaves    adaxial surface of some
                                               leaves and absent on
                                               others; very dense on
                                               abaxial surface of all
                                               leaves

Indumentum of        very dense, with          very dense, with
involucral bracts    dominant long simple      stellate, glandular and
                     eglandular trichomes,     long simple eglandular
                     [+ or -] abundant         trichomes in [+ or -]
                     stellate trichomes and    similar proportions
                     very scarce or absent
                     glandular trichomes

Characters           P. officinarum

Stolons              abundant, elongated and
                     thin

Stellate trichomes   absent on adaxial surface
on leaves            of all leaves and very
                     dense on abaxial surface
                     of all leaves

Indumentum of        very dense, with abundant
involucral bracts    black glandular trichomes,
                     scarce to moderate stellate
                     trichomes and scarce or
                     absent simple eglandular
                     trichomes

Table 3
Principal differences between Pilosella argyrocoma, P. megargyrocoma
sp. nova and P. peleteriana.

Characters           P. argyrocoma             P. megargyrocoma

Stellate trichomes   abundant on both          few to numerous on
on leaves            surfaces of all leaves    adaxial surface of some
                                               leaves and absent on
                                               others; very dense on
                                               abaxial surface of all
                                               leaves

Diameter of scapes   1-1.5 mm                  1.5-2 mm
Width of             [+ or -] 1 mm             1.2-1.6 mm
involucral bracts

Indumentum of        very dense, with          very dense, with
involucral bracts    dominant long simple      dominant long simple
                     eglandular trichomes,     eglandular trichomes,
                     [+ or -] abundant         quite abundant stellate
                     stellate trichomes and    trichomes and very
                     very scarce or absent     scarce glandular
                     glandular trichomes       trichomes

Characters           P. peleteriana

Stellate trichomes   absent on adaxial surface of
on leaves            all leaves and very dense
                     on abaxial surface

Diameter of scapes   [+ or -] 2 mm
Width of             up to 2-3 mm
involucral bracts

Indumentum of        very dense, with dominant
involucral bracts    very long simple eglandular
                     trichomes and very scarce
                     or absent glandular
                     trichomes and/or stellate
                     trichomes
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Title Annotation:articulo en ingles
Author:del Egido Mazuelas, Fermin; Sanz, Gonzalo Mateo
Publication:Lazaroa
Date:Jan 1, 2013
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