Botanical Journal of the Linnean Society, 2005, 148, 229–244. With 23 figures
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 148, 229–244 229
Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2005? 2005
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Original Article
A REVISION OF
L. VERTICILLATA
GROUP
L. SÁEZ and M. B. CRESPO
*Corresponding author. E-mail: llorens.saez@uab.es
A taxonomic revision of the Linaria verticillata group
(Antirrhineae, Scrophulariaceae)
LLORENÇ SÁEZ
1
* and MANUEL B. CRESPO
2
1
Unitat de Botànica, Facultat de Ciències, Universitat Autònoma de Barcelona. E-08193, Bellaterra,
Barcelona, Spain
2
Instituto de la Biodiversidad (CIBIO), Universidad de Alicante, E-03080 Alicante, Spain
Received February 2003; accepted for publication November 2004
A taxonomic revision of the Linaria verticillata group is presented here. Taxonomic, nomenclatural, morphological,
geographical and ecological data are recorded for each taxon. Four taxa (one species with four subspecies) are
recognized, and the status of some related taxa is discussed. In addition, some lectotypifications are made and a
taxonomic key, descriptions and illustrations of all the accepted taxa are given. The following new combinations are
proposed: L. verticillata ssp. anticaria (Boiss. & Reut.) L. Sáez & M.B. Crespo comb. & stat. nov., L. verticillata ssp.
cuartanensis (Degen & Hervier) L. Sáez & M.B. Crespo, comb. nov., and L. verticillata ssp. lilacina (Lange) L. Sáez
& M.B. Crespo, comb. & stat. nov. © 2005 The Linnean Society of London, Botanical Journal of the Linnean
Society, 2005, 148, 229–244.
ADDITIONAL KEYWORDS: – endemism – Iberian Peninsula – infraspecific variation – taxonomy.
INTRODUCTION
The Linaria verticillata group comprises closely
related plants inhabiting the Betic and Subbetic
mountains of the southern Iberian Peninsula. They
are perennial herbs with erect to procumbent stems,
predominantly verticillate linear-oblong to elliptical
leaves, and mainly yellow to blue-violet or whitish
flowers. Taxa in the group show very close similarities
in seed-coat sculpturing (Sutton, 1988) and reliable
diagnostic characters are based on features of flowers
(mainly colour and shape) and indumentum.
Three species with remarkable macromorphological
similarities have usually been ascribed to the group
(Valdés, 1970; Sutton, 1988). The first is L. verticillata
Boiss., described from Sierra Nevada and other neigh-
bouring Betic ranges (Boissier, 1841). It was charac-
terized by glandular-pubescent stems throughout and
large yellow flowers, sometimes with dark stripes.
Later, L. anticaria Boiss. & Reut. was described from
Torcal de Antequera (Málaga province) in southern
Spain (Boissier & Reuter, 1852). Its main characters
were the glaucous, mostly glabrous stems, fruiting
inflorescence glandular-pubescent and usually rather
lax and greyish-lilac flowers, typically with a deep
bluish-violet palate. Boissier & Reuter (1852) also
recognized a variant of L. anticaria (var. angustifolia
Boiss. & Reut.) with narrower leaves. Finally,
L. lilacina Lange was described from the surroundings
of Jaén (Lange, 1854). It differed in having glandular-
pubescent stems throughout, dense inflorescence and
flowers with orange palate and shorter spur.
Morphological attributes underlying the separation
of the three taxa are scanty. This has sometimes pro-
duced confusion in plant identification (Sánchez &
Alcaraz, 1993). Sutton (1988) suggested that further
investigation of the limits of L. anticaria was required.
The delimitation of L. anticaria and L. lilacina is
sometimes controversial and several taxa have been
described (cf. Willkomm, 1859; Coincy, 1895; Hervier,
1905; Cuatrecasas, 1929) in order to accommodate
morphological variants which differ in some way from
the typical forms. Many have no taxonomic sig-
nificance, while others appear to be intermediate
forms (perhaps hybrid aggregates) with taxa of the
L. aeruginea (Gouan) Cav. group, as discussed below.
However, Linaria anticaria ssp. cuartanensis Degen &
Hervier is perhaps the most remarkable, since it is
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