Taxonomic Study of Linaria depauperata and L. supina Complexes in Eastern Spain

Annals of Botany 87: 157±177, 2001 doi:10.1006/anbo.2000.1311, available online at http://www.idealibrary.com on

Taxonomic Study of Linaria depauperata and L. supina Complexes in Eastern Spain JO SEÂ G A B R I E L S E G A R R A * and I S A B E L M AT E U Instituto Cavanilles de Biodiversidad y BiologõÂa Evolutiva, Dpt. de BiologõÂa Vegetal, Universitat de ValeÁncia, C/ Dr. Moliner 50, Burjassot, ValeÁncia E-46100, Spain Received: 11 February 2000 Returned for revision: 4 August 2000

Accepted: 2 October 2000

Morphological characteristics were studied in two groups of closely related taxa of section Supinae, Linaria depauperata and L. supina, from eastern Spain. Vegetative, ¯oral and seed characteristics were of little use in distinguishing between these taxa as the ranges for all the taxa studied overlapped. Flower colour could be used as a diagnostic character for some but not all of these taxa. Seed coat characters were of high taxonomic value, both when trying to distinguish among these two groups, and the taxa included within them, and also when establishing relationships among the taxa studied. Some di€erences were found in the indumentum distribution and density. Three types of trichomes were identi®ed, two of them eglandular and common to all taxa. The third type of trichome was a simple gland, that varied in the number of cells in the stalk and gland. Trichome length and wall width of stalk cells allowed several groups of taxa to be distinguished. Similarity relationships among the samples studied are shown in a UPGMA phenogram. According to these results, two new species, L. cardonica and L. sulphurea, are described. L. arabiniana is conspeci®c with L. depauperata subsp. hegelmaieri. A key to identify the di€erent taxa is # 2001 Annals of Botany Company provided. Key words: Scropulariaceae, Antirrhineae, taxonomy, morphological characters, seed-coat, trichomes, Linaria depauperata, Linaria supina.

I N T RO D U C T I O N Linaria Miller is the largest genus of the tribe Antirrhineae (Sutton, 1988). It is distributed throughout the northern hemisphere and its distribution centre is located in the Mediterranean Basin (Hong, 1983) as the majority of its species are found in Europe, south-western Asia and North Africa. Linaria species are annual or perennial herbs, with heteromorphic shoots. Flowers are arranged in bracteate racemes and have a calcarate personate corolla. Seed morphology is various, but two main groups have been traditionally distinguished according to the presence or absence of a marginal encircling wing (Dumortier, 1827; Lange, 1863; ValdeÂs, 1970; Chater et al., 1972; Sutton, 1988). According to Sutton (1988), winged species of Linaria are distributed among three sections: Linaria, Pelisserianae ValdeÂs and Supinae (Bentham) Wettst; section Supinae is the largest, including a third of all known species. Its distribution centre is placed in the Iberian Peninsula, where it shows a pattern of geographic diversi®cation (ValdeÂs, 1970). This pattern has led to the di€erentiation of a large number of narrow endemics, often with specialized ecology (ValdeÂs, 1970; Carretero and Boira, 1986; Sutton, 1988; Crespo et al., 1994; De la Torre et al., 1996; Laguna et al., 1998). * For correspondence at: Dpt. Agricultura y Economõ a Agraria, Facultad de Veterinaria, Universidad de Zaragoza, C/Miguel Servet 177, 50013 Zaragoza, Spain. Fax ‡34 976762488, e-mail jogasemo@ posta.unizar.es

0305-7364/01/020157+21 $35.00/00

Supinae is a complex section that forms groups of closely related taxa, dicult to separate according to their variable vegetative characters. L. arabiniana M.B. Crespo, De la Torre & Solanas, L. depauperata Leresche ex Lange subsp. depauperata, L. depauperata Leresche ex Lange subsp. hegelmaieri (Lange) De la Torre, Alcaraz & M.B. Crespo, and Linaria supina (L.) Chaz. var. cardonica Font Quer are restricted to the eastern part of the Iberian Peninsula. In the northern part of this area (CastelloÂn province and Catalonia) some populations of Linaria are controversial. Font Quer (1950) described a new variety as Linaria supina var. cardonica, while Fabregat (1995), Riera (1992) and Tirado (1995, 1998) considered it equivalent to L. supina s.l. Recently, PeÂrez-Dacosta (1998) considered some of these populations to be L. depauperata subsp. hegelmaieri [labelled by us as L. depauperata subsp. hegelmaieri (Cs)] while others were considered to be L. aeruginea (Gouan) Cav. var. cardonica Font Quer (nom. inval.), excluding L. supina from the ¯ora of eastern Spain (Table 1). L. supina is a polymorphic taxon, with one of the widest areas of distribution among the taxa studied here. It is distributed from the northern part of the Iberian Peninsula to several countries in Western Europe. As a result of this wide range, and due to the diversi®cation pattern shown by the members of this section, it is divided into many infraspeci®c taxa, all of which, bar the typical one, are considered as endemics of the Iberian Peninsula. Of these taxa, only L. supina subsp. supina and L. s. var. cardonica have been reported in the study area. These taxa are very dicult to separate from each other as they di€er in few characters and are mostly sympatric. # 2001 Annals of Botany Company

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Segarra and MateuÐTaxonomy of Linaria T A B L E 1. Di€erent circumscription of the taxa studied

ValdeÂs (1970)

Sutton (1988)

BoloÁs and Vigo (1995)

Mateo and Crespo (1998)

PeÂrez-Dacosta (1998)

L. aeruginea subsp. aeruginea

L. aeruginea subsp. aeruginea

L. supina subsp. aeruginea var. aeruginea

L. arabiniana

L. arabiniana

L. depauperata subsp. depauperata var. depauperata

L. depauperata subsp. L. supina subsp. aeruginea depauperata var. cardonica

L. aeruginea subsp. aeruginea

L. aeruginea subsp. aeruginea var. aeruginea

L. depauperata subsp. depauperata

L. aeruginea subsp. aeruginea var. cardonica

L. depauperata subsp. hegelmaieri

L. depauperata subsp. depauperata

L. supina subsp. supina

L. depauperata subsp. hegelmaieri

L. depauperata subsp. depauperata var. hegelmaieri L. supina subsp. supina

L. supina subsp. supina

L. supina subsp. depauperata var. depauperata L. supina subsp. depauperata var. hegelmaieri

For this reason, ValdeÂs (1970) included all of them within L. supina. Sutton (1988) also did the same, and pointed out the need for further research on this group to determine how this infraspeci®c variation would be most appropriately expressed. Despite this, he described another subspecies of L. supina as L. s. subsp. duriaeana Sutton, although he could not be sure that it was not simply an extreme of clinal variation. Taxa of the L. depauperata complex have been included in L. supina by some authors (Table 1). Taxa considered to belong in this group are mainly distributed throughout the eastern part of the Iberian Peninsula. They are also very variable with regard to vegetative morphological traits, even within populations. This led Crespo et al. (1994) to describe a new species, L. arabiniana. The taxonomic status of this species is controversial, as its diagnostic features overlap with the range of variability of L. depauperata subsp. hegelmaieri from the Alicante province [labelled by us as L. depauperata subsp. hegelmaieri (A)], with which it shares its distribution range. L. aeruginea is also divided into several morphologically distinct infraspeci®c taxa. Of these, only the typical subspecies is present in this area. This taxon is included in the present study in order to clarify the relationships proposed by ValdeÂs (1970) and Sutton (1988) between this taxon and the group of L. depauperata, or with L. s. var. cardonica as suggested by PeÂrez-Dacosta (1998). The aims of this study were to establish the taxonomic status of these populations of Linaria and the relations between them. For this reason, levels of variability in microand macromorphological traits of vegetative, ¯oral, seed and indumentum characters were studied. The relationship between L. supina var. cardonica and other morphologically similar taxa included in L. supina s.l. was also studied.

M AT E R I A L S A N D M E T H O D S Material for this study was collected from wild populations and herbarium specimens as shown in the Appendix. Voucher specimens are deposited at the Herbarium of the Department of Plant Biology (VAB) at the University of Valencia, Spain. Measurements of vegetative and ¯oral parts and also of seed characters were carried out under a stereomicroscope

Leika M3Z. For scanning electron microscopy, dry mature seeds were mounted directly on stubs using one-sided adhesive tape, and coated with gold/palladium in a sputtercoater. The terminology of seed-coat surface sculpturing mainly follows Sutton (1988). For light microscopic observations of the indumentum, aerial parts of Linaria specimens were kept in a saturated solution of Chloral hydrate for 24 h to facilitate epidermis separation. Cellular contents were then eliminated by keeping the material in a commercial bleach solution for at least 6 h. The epidermis was separated and stained with Bismarck brown (Johansen, 1940), and unpermanently mounted with glycerol gelatine on glass slides. Observations were carried out in an Olympus BH2 microscope. Trichomes were drawn using a camera lucida. For scanning electron microscopic observation of the indumentum, aerial parts of Linaria specimens were dehydrated in an ethanol series and critical point dried using carbon dioxide. Afterwards, samples were coated with gold/palladium in a sputter-coater. Morphological observations were carried out in a Hitachi S-4100 ®eld emission SEM. Terminology of indumentum density and trichome types follows Font Quer (1970), Payne (1978) and Stearn (1972). Data obtained from observations (qualitative data) were converted to a binary data matrix, which is available upon request. The similarity matrix was obtained with the SM index, and clustered with UPGMA algorithm (Sneath and Sokal, 1973). Both matrix and cluster were generated by the Ntsys version 2.0 program (Rolf, 1998). R E S U LT S Vegetative, ¯oral and fruit characters Values obtained for vegetative, ¯oral and fruit characters are shown in Table 2. The ranges obtained for the measured characters overlapped in all the taxa studied; only for corolla-tube width were two separate ranges distinguished. In all the taxa studied, the corolla-tube can be considered as broad (tube width 4 5 mm) but in L. supina subsp. supina it is narrower (up to 4.5 mm). As this was the only diagnostic character among those studied for this group, it was binarized and treated as qualitative data. Flower colour was variable among the taxa studied. The L. supina group showed less variability as its taxa all had

Perennial, biennial or annual 3.2±16.5 0.4±1.3 5±22 0.5±1.5 2±6 0.5±3.25 3.5±9 0.4±1.6 2.25±5.5 0.6±1.75 Dark red

17±28 7±14 1.33±2.86 4.75±8 1±4.25 0.5±1.5 3.5±5.4 2.5±5

Leaf length on sterile shoots (mm) Leaf width on sterile shoots (mm) Leaf length on fertile shoots (mm) Leaf width on fertile shoots (mm) Bract length (mm) Pedicel length (mm) Adaxial calyx-lobe length (mm) Adaxial calyx-lobe width (mm) Abaxial calyx-lobe length (mm) Abaxial calyx-lobe width (mm) Flower colour

Corolla length (mm) (A) Spur length (mm) (B) A/B ratio Corolla tube width (mm) Adaxial lip sinus (mm) Abaxial lip sinus (mm) Capsule's septum length (mm) Capsule's septum width (mm)

Duration

L. aeruginea subsp. aeruginea Perennial, biennial or annual 3.25±10 0.6±1.25 5.50±12 0.75±1.3 2.25±6.5 0.38±4 5±7 0.6±1.25 3.5±5 0.75±1.5 Violet, reddish or yellowish orange 18±31.5 8±12 1.92±2.74 5±7 1.45±2.75 0.6±1.1 4.25±4.75 4±5

L. arabiniana

4±14 0.5±0.9 5.5±20 0.5±1.5 1.75±5.5 0.5±3.0 3.5±7 0.5±1.75 3±5 0.7±1.5 White, often with yellowish spur 19±26 7±13 1.82±3.17 5±7 0.9±3.75 0.5±1.5 3.25±5.5 2.5±4.5

Biennial or annual

L. depauperata subsp. depauperata

L. depauperata group

Perennial, biennial or annual 4.5±7 0.75±0.95 8±12 1±1.5 1.8±5.0 0.5±4.75 4.75±7.7 0.8±1.2 4±5.5 0.95±1.1 Violet, reddish, yellowish orange 20±28 8±12 2±2.63 5±6 1.25±2.5 0.5±1.5 4.25±5.55 4.25±6.15

L. depauperata subsp. hegelmaieri (A)

18±28 8±13 1.83±2.66 5±9 1.10±3.50 0.6±2.75 2.50±6.75 3±5

4.25±10 0.5±1.25 5±21 0.5±1.5 2±6.5 0.5±5 2.75±8.75 0.5±1.8 1±4.75 0.6±1.75 Bright yellow

Biennial or annual

L. depauperata subsp. hegelmaieri (Cs)

11±26 7±13.5 1.1±2.71 3±4.5 1±3.25 0.5±1.5 3.5±5.5 2.75±5

Perennial, biennial or annual 2.75±14 0.3±2 5±32 0.4±3.25 1.8±8 0.5±5 2.75±8.5 0.3±1.5 2.25±5.5 0.5±1.5 Bright yellow

16±25.5 7±12.5 1.76±3.14 5±7 0.75±3 0.5±1.5 4.75±5.25 4±4.75

3.25±10 0.6±1.25 5±16 0.75±1.45 2.5±3.75 1±4 4.25±7 0.5±1.1 3±4.5 0.5±1.5 Bright yellow

Biennial or annual

L. supina var. cardonica

L. supina group L. supina subsp. supina

T A B L E 2. Quantitative and qualitative characters of vegetative organs, ¯owers and fruits for all the taxa studied

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pale yellow corollas. By contrast the L. depauperata group was more variable as ¯ower colour ranged from white to dark red or violet. Some taxa, such as L. aeruginea and L. depauperata subsp. depauperata, always had ¯owers of the same colour, although ¯ower colour varied in tone and intensity. On the other hand, L. arabiniana and L. depauperata subsp. hegelmaieri from Alicante (A), appeared to be very variable for this character, while L. depauperata subsp. hegelmaieri from CastelloÂn (Cs) consistently had bright yellow corollas.

Seed characters Seed morphology showed striking di€erences between both groups of taxa studied. As a result, two types were distinguished. Type I. Concolourous seeds. Black, often metallic shiny. Wing broad, ¯at or strongly concavo-convex, membranous and translucent or opaque. Disc smooth or covered by tubercles. Seed-coat is composed of irregularly polygonal cells, with raised, sometimes incrassate, anticlinal walls and smooth or verrucate tabular-concave periclinal walls. Tubercles are the same colour as the disc. Tubercles vary in shape and density and their periclinal cell walls may be smooth or densely verrucate. Taxa included in this type are: Linaria supina subsp. supina and L. supina var. cardonica. (Fig. 1G and H). Type II. Discolourous seeds. Disc colour is black and wing ranges from whitish to grey. Wing broad, usually concavo-convex, sometimes strongly so, or ¯at, membranous and translucent. Disc smooth or covered by tubercles. Seed-coat is composed of irregularly polygonal or rounded cells, with raised, sometimes incrassate, anticlinal walls and smooth, reticulate or verrucate tabular-concave periclinal walls. Tubercles vary in shape, colour and density. Taxa included in this type are: L. aeruginea subsp. aeruginea, L. arabiniana, L. depauperata subsp. depauperata and L. depauperata subsp. hegelmaieri (Fig. 1 A±F). Values obtained for seed characters are shown in Table 3. Seeds of all studied taxa are orbicular to suborbicular in outline. As shown in Table 3, data obtained from measurement of seed parts overlap among the taxa studied. Observations of qualitative characters also proved to be variable among the taxa studied and, sometimes, even within one taxon. Several features of the seed coat were found to be exclusive for certain taxa. The disc surface could be smooth or tuberculate. This feature was not always constant for one taxon, as L. aeruginea subsp. aeruginea, L. depauperata subsp. hegelmaieri (A), L. supina and L. supina var. cardonica had seeds of both types. L. arabiniana always had seeds with a smooth disc (Fig. 1D). Seeds of L. depauperata subsp. depauperata and L. depauperata subsp. hegelmaieri (Cs), on the other hand, were always tuberculate (Fig. 1A and E). Features of tubercles were much more speci®c for each taxon. L. supina var. cardonica had smooth tubercles varying in length, with a smooth periclinal wall

(Fig. 4A±C), while those of L. supina, when present, were much shorter and had a verrucate wall (Fig. 4D±F). Regarding the L. depauperata group, L. depauperata subsp. depauperata had long white tubercles, often fused together forming ridges (Fig. 1A; Fig. 2A and B), while those of L. depauperata subsp. hegelmaieri (A) were rounded, shorter and generally more lax (Fig. 1B; Fig. 2D and E), like those of L. aeruginea subsp. aeruginea (Fig. 2J and K). The tubercles of L. depauperata subsp. hegelmaieri (Cs) were similar to those of L. depauperata subsp. depauperata but they were not fused together (Fig. 1E, Fig. 2G and H). The shape of disc cells was generally irregular, but in L. arabiniana and L. depauperata subsp. hegelmaieri (A) it was rounded (Fig. 3D and F). Seed-coat cell sculpturing was very variable among the taxa studied ranging from smooth in L. supina var. cardonica (Fig. 4C) to verrucate in L. depauperata subsp. hegelmaieri (A) (Fig. 2F) and reticulate in both L. depauperata subsp. depauperata and L. depauperata subsp. hegelmaieri (Cs) (Fig. 2C and I).

Indumentum characters Trichomes were found only on the in¯orescence and ¯ower parts. Variation in indumentum characters for both groups of taxa studied is shown in Table 4. The arrangement and density of trichomes were constant in the L. supina group, while they were variable amongst the taxa studied in the L. depauperata group. L. arabiniana and L. depauperata subsp. hegelmaieri (A) had scattered trichomes, located mainly on the edges of the calyx-lobes (Fig. 6A), while the rest of the ¯ower parts and in¯orescence were entirely glabrous. The rest of the taxa studied had a more dense indumentum covering on the ¯ower parts and in¯orescence (Fig. 6B). Three types of trichome were identi®ed. Two types of eglandular hair were found on the inner epidermis of the corolla and one glandular type on the gynoecium, external epidermis of the ¯ower parts and in¯orescence. Type I (Fig. 6C) is an eglandular unicellular hair. It is conical, short, not exceeding 100 mm, and has a verrucate wall (Fig. 6D). This type occurs at the base of the stamen ®lament and on the inner surface of the spur. Type II (Fig. 6D) is also an eglandular, unicellular hair, but is cylindrical and longer, up to 600 mm, and covers both nectar ducts, from the edge of the corolla lip to the base of the corolla tube. Both types of eglandular hairs are common to all the taxa studied. Type III (Figs 5A±G, 6E and F) is a simple glandular hair formed by a uniseriate stalk and a gland. The length and number of cells found in the stalk and gland, and the thickness of the stalk cell walls were variable. Trichome length varied from 38.8 mm in L. arabiniana to 827.03 mm in L. aeruginea subsp. aeruginea, and the number of stalk cells ranged from one to seven. The stalk cells of some taxa had an incrassate wall, with a wall width ranging from 0.72 mm in L. arabiniana to 5.89 mm in L. aeruginea subsp. aeruginea (Fig. 5E). The number of cells in the gland ranged from two to eight.

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F I G . 1. Stereoscopic microscope photographs of the taxa studied. A, L. depauperata subsp. depauperata. B, L. depauperata subsp. hegelmaieri (A) with tuberculate disc. C, L. depauperata subsp. hegelmaieri (A) with smooth disc. D, L. arabiniana. E, L. depauperata subsp. hegelmaieri (Cs). F, L. aeruginea subsp. aeruginea. G, L. supina var. cardonica. H, L. supina s.l. Bars ˆ 1 mm.

Whitish-grey 0.55±1.1 Membranous Translucent Polygonal Raised, incrassate Tabular, reticulate, without papillae Smooth Rounded Raised, incrassate Tabular-concave, smooth, without papillae Ð

Whitish-grey

0.55±1.1 Membranous Translucent Polygonal Raised, incrassate Tabular, reticulate, without papillae Smooth-tuberculate Irregularly polygonal Raised, incrassate Tabular-concave, reticulate, without papillae Low

Rounded

Same as wing Raised, incrassate or shallow Tabular, reticulate

Seed body/Seed wing Wing shape

Wing colour

Wing width (mm) Wing texture Wing appearance Shape of wing cells Anticlinal walls of wing cells Periclinal walls of wing cells

Shape of tubercles/ridges

Colour of tubercles/ridges Anticlinal walls of cells of tubercles/ridges Periclinal walls of cells of tubercles/ridges

Density of tubercles/ridges

Disc surface Shape of disc cells Anticlinal walls of disc cells Periclinal walls of disc cells

White Raised, incrassate Tabular, reticulate

Ð

Acute, or ridge-like

High

0.4±1 Membranous Translucent Polygonal Raised, incrassate Tabular, reticulate, without papillae Tuberculate Ð Ð Ð

Whitish-grey

2±3.4 Suborbicularorbicular Discolourous Concave

L. depauperata subsp. depauperata

Ð Ð

Ð

2.2±3.5 Suborbicularorbicular Discolourous Flat

2.3±3.7 Suborbicularorbicular Discolourous Concave

Seed size (mm) Seed outline

L. arabiniana

L. aeruginea subsp. aeruginea

L. depauperata group

Tabular, reticulate

Rounded or sometimes acute Whitish-grey Raised, incrassate

0.6±0.75 Membranous Translucent Polygonal Raised, incrassate Tabular, reticulate, without papillae Smooth-tuberculate Rounded Raised, incrassate Tabular-concave, smooth or verrucate, without papillae Medium

Whitish-grey

2.2±2.9 Suborbicularorbicular Discolourous Flat or concave

L. depauperata subsp. hegelmaieri (A)

T A B L E 3. Seed characters of the taxa studied

Tabular, verrucate

Whitish-grey Raised, incrassate

Acute

Low

0.55.1.1 Membranous Translucent Polygonal Raised, incrassate Tabular, reticulate, without papillae Tuberculate Ð Ð Ð

Grey

2.1±3.7 Suborbicularorbicular Discolourous Concave

L. depauperata subsp. hegelmaieri (Cs)

Tabular, verrucate

Same as wing Raised, incrassate

Rounded or acute

1.6±3.25 Suborbicularorbicular Concolourous Flat or slightly concave Black, sometimes metallic shiny 0.3±1 Membranous Opaque Polygonal Raised, incrassate Tabular, verrucate, without papillae Smooth-tuberculate Irregularly polygonal Raised, incrassate Tabular-concave, smooth or verrucate, without papillae Low or medium

L. supina subsp. supina

Tabular, smooth

Same as wing Raised, incrassate

0.75±1.3 Membranous Translucent Polygonal Raised, incrassate Tabular, smooth, without papillae Smooth-tuberculate Irregularly polygonal Raised, incrassate Tabular-concave, smooth or verrucate, without papillae Low, medium or high Acute

Black, metallic shiny

2.6±4 Suborbicularorbicular Concolourous Strongly concave

L. supina var. cardonica

L. supina group

162 Segarra and MateuÐTaxonomy of Linaria

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163

F I G . 2. Scanning electron micrographs of tuberculate seeds of the L. depauperata group. A±C, L. depauperata subsp. depauperata. D±F, L. depauperata subsp. hegelmaieri (A). G±I, L. depauperata subsp. hegelmaieri (Cs). J and K, L. aeruginea subsp. aeruginea. Bars ˆ 25 mm (C, F, H and K); 50 mm (B, E and I); 200 mm (A and G); 250 mm (D and J).

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F I G . 3. Scanning electron micrographs of smooth seeds of L. depauperata group. A and B, L. aeruginea subsp. aeruginea. C and D, L. arabiniana. E and F, L. depauperata subsp. hegelmaieri (A). Bars ˆ 25 mm (B, D, E and F); 125 mm (C); 200 mm (A).

In our opinion, and according to Juan (1994), it is not possible to distinguish trichome types with reference to the number of cells in the stalk and gland in these taxa, but it is possible using trichome length and wall thickness of the stalk cells. We have distinguished two di€erent categories for each of these two characters: (1) short trichomes (trichomes up to 110 mm) (Fig. 5B and C); and (2) long trichomes (up to 830 mm) (Fig. 5A, D±G); and (1) thin walled cells (wall width up to 3.5 mm) (Fig. 5A±C, F±G); and (2) thick walled cells (wall width 5 4 mm) (Fig. 5D and E). Phenetic analysis Figure 7 shows the UPGMA cluster obtained with qualitative and discontinuous quantitative data (binarized) of all the characters studied that were considered to be informative (Table 5). In this cluster, only individuals with complete characters for the seed have been included as they constitute over half of the total characters considered in the binary data matrix. As shown in the cluster, two main groups joined at the 0.67 similarity level are distinguished. Group 1 includes individuals with tuberculate seeds and group 2, those with smooth seeds. Within group 1, the taxa forming the L. depauperata s.l. group (3) are separated from those forming the L. supina

group (4) at the 0.72 similarity level. The inclusion of L. aeruginea subsp. aeruginea in group 3 points out its higher similarity to L. depauperata. Group 5 includes individuals closer to typical L. depauperata. L. depauperata subsp. depauperata and L. depauperata subsp. hegelmaieri from Alicante populations are joined at an intraspeci®c variation level of 0.84. Individuals of L. depauperata subsp. hegelmaieri from CastelloÂn province appear to be joined together at the same level but are distinctly separated from those populations from Alicante. This separation justi®es the description of a new species which is supported by di€erences in ¯ower colour, several seed features and the presence of longer trichomes with thicker stalk cell walls in these samples. Group 4 includes samples of L. supina var. cardonica (7) that appear separated from all the other infraspeci®c taxa considered to be part of L. supina s.l. (8). This separation is supported by di€erences in ¯ower morphology and several seed features. Some taxa, including L. aeruginea subsp. aeruginea, L. supina s.l. and L. supina var. cardonica, are present at the same time on both branches of the cluster (1, 2) as they can have tuberculate and smooth seeds. L. arabiniana only appears in branch 2 as it always has smooth seeds and is mixed with samples of L. depauperata subsp. hegelmaieri from Alicante with smooth seeds. By contrast, other taxa

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F I G . 4. Scanning electron micrographs of L. supina group. A±C, L. supina var. cardonica. D±F, L. supina s.l. Bars ˆ 25 mm (B, E and F); 250 mm (A, C and D).

including L. depauperata subsp. depauperata and L. depauperata subsp. hegelmaieri from CastelloÂn always have tuberculate seeds. The presence of the same taxon in both branches of the cluster reinforces the hypotheses of Sutton (1988) and ValdeÂs (1970) that point out that the presence or absence of tubercles is not a reliable taxonomic character in these groups of taxa.

DISCUSSION Data obtained from measurement of vegetative, ¯oral, fruit and seed characters have proved to be of little use in distinguishing between these taxa as the values obtained overlap. Only the width of the corolla tube distinguishes L. supina subsp. supina from the rest of the taxa. Flower colour was found to be informative among the taxa studied belonging to the L. depauperata group. Some taxa of this group show diagnostic ¯ower colours such as L. aeruginea subsp. aeruginea and L. depauperata subsp. depauperata. Others, by contrast, show a wider range of ¯ower colours. L. depauperata subsp. hegelmaieri (A), reported to have orangish ¯owers more or less red tinged, showed a wide variety of colours, from the typical colouration to violet or lilac with a yellow palate. This range is also shared by L. arabiniana. Individuals of L. depauperata subsp. hegelmaieri from CastelloÂn, in

contrast, do not show this colour pattern at all as their ¯owers are always bright yellow. Seed characters are of paramount importance in the taxonomy of Linaria species. Chavannes (1833) pointed out the possibility of their use as a base for a classi®catory system within the tribe Antirrhineae, and Sutton (1988) based the revision of the whole genus mainly on seed characters. In the L. depauperata group, the disc is always black and the wing ranges from white to whitish-grey (Fig. 1A±F). The uniformly grey wing of individuals of L. depauperata subsp. hegelmaieri from CastelloÂn (Fig. 1E) is remarkable as it distinguishes them from individuals of populations from Alicante that have a whitish-grey wing (Fig. 1B±C). When seeds are tuberculate, the morphology and microsculpturing of the tubercles may be useful taxonomic characters as they vary among the di€erent taxa. L. depauperata subsp. depauperata has long acute tubercles which are often fused together and ridge-like (Fig. 2B). This distinctive feature is also present in L. amoi Campo ex Amo from western Andalucõ a (southern Spain). The microsculpturing of the periclinal wall of the tubercle is reticulate. In L. depauperata subsp. hegelmaieri from Alicante, tubercles are shorter (Fig. 2D) or absent (Fig. 3E and F), more lax and with a densely verrucate periclinal wall as in L. aeruginea subsp. aeruginea (Fig. 2K). By contrast, in the samples from CastelloÂn province, initially assigned to

Glandular trichome length (mm) Glandular trichome wall width (mm)

Indumentum density 54.44±827.03 5.41 + 0.47

Pubescent

L. aeruginea subsp. aeruginea Glabrous/ glabrescent 38.8±103.7 0.84 + 0.12

L. arabiniana

51.25±552 0.80 + 0.058

Puberulent

L. depauperata subsp. depauperata

L. depauperata group

Glabrous/ glabrescent 46.66±101 0.76 + 0.040

L. depauperata subsp. hegelmaieri (A)

T A B L E 4. Indumentum characters of the taxa studied

49.25±740.19 4.74 + 0.57

Pubescent

L. depauperata subsp. hegelmaieri (Cs)

67.40±738.9 1.99 + 0.47

Pubescent

L. supina subsp. supina

64.82±640.37 2.48 + 1.04

Pubescent

L. supina var. cardonica

L. supina group

166 Segarra and MateuÐTaxonomy of Linaria

Segarra and MateuÐTaxonomy of Linaria

167

F I G . 5. Camera lucida drawings of glandular trichomes of the taxa studied. A, L. depauperata subsp. depauperata. B, L. arabiniana. C, L. depauperata subsp. hegelmaieri (A). D, L. depauperata subsp. hegelmaieri (Cs). E, L. aeruginea subsp. aeruginea. F, L. supina s.l. G, L. supina var. cardonica. Bars ˆ 100 mm.

this taxon, seeds are always tuberculate, with long acute tubercles with a reticulate periclinal wall (Fig. 2H). The morphology and microsculpturing of testa cells of the seeds of L. arabiniana (Fig. 3C and D) make it indistinguishable from samples of L. depauperata subsp. hegelmaieri from Alicante populations (Fig. 3E and F) when tubercles are absent in the latter taxon. Di€erences in seed morphology have also been found in the L. supina group. The strongly concave wing of L. supina var. cardonica (Fig. 4A) distinguishes it from L. supina s.l. (Fig. 4D) whose wings are ¯at or slightly concave.

Tubercles of both taxa are also important in their di€erentiation. L. supina var. cardonica has tubercles with a smooth periclinal wall (Fig. 4B), while in L. supina s.l. they are strongly verrucate (Fig. 4E). In agreement with other authors, we consider that seed characters are essential for the taxonomy of these taxa, but sometimes other characters are required for positive identi®cation. Trichome characters have been employed by several authors in the systematics of the Scrophulariaceae (Brooker et al., 1976; Raman, 1987, 1989a,b, 1990a,b, 1991;

168

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F I G . 6. Scanning electron micrographs of indumentum, glandular and non-glandular trichomes of the taxa studied. A, L. arabiniana. Indumentum glabrescent. B, L. supina s.l. Indumentum pubescent. C, L. supina var. cardonica. Conical, non-glandular trichome from base of stamen ®lament. D, L. supina var. cardonica. Cylindrical, non-glandular trichome from nectar duct. E, L. arabiniana. Short glandular trichome. F, L. depauperata subsp. hegelmaieri (Cs). Long glandular trichome. Bars ˆ 10 mm (D); 25 mm (C, E and F); 200 mm (A); 250 mm (B).

T A B L E 5. Morphological characters and states considered in the phenetic analysis 1

Duration:

2

Leaf margin:

3

Trichome length:

4

Stalk cell wall width:

5

Covering of indumentum on the in¯orescence:

6

Flower colour:

7

Corolla tube:

8

Colour of the disc of the seed:

Annual Biennal Perennial Plane Revolute Long Short Thick Thin Glabrous Glabrescent Puberulent Pubescent White Bright yellow Orangish Dark red Broad Slender Black, metallic shiny Black Grey

9 Wing colour:

10 Wing shape: 11 Wing texture: 12 Colour of tubercles: 13 Shape of tubercles: 14 Fusion of tubercles: 15 Tubercles density:

Black, metallic shiny Black Grey White or whitish Flat Concave Translucent Opaque White Greyish Black Acute Rounded Free Some fused All fused High Medium Low Absent

Segarra and MateuÐTaxonomy of Linaria

169

F I G . 7. UPGMA phenogram of Linaria samples clustered on the basis of qualitative data. Identi®cation codes for the samples are: LARA, L. arabiniana; LA, L. aeruginea subsp. aeruginea; LDD, L. depauperata subsp. depauperata; LDHA, L. depauperata subsp. hegelmaieri (A); LDHCS, L. depauperata subsp. hegelmaieri (Cs); LCARD, L. supina var. cardonica; LSUP, L. supina s.l.

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F I G . 8. Habit of L. sulphurea. From the holotype. Bar ˆ 1 cm.

Canne-Hilliker and Kampny, 1991; Doaigey and Harkiss, 1991; Neumann et al., 1997), pointing out their usefulness, even for the taxonomy of Linaria (Juan, 1994; Raman, 1987, 1989a; Juan et al., 1999). Raman (1987) distinguished the di€erent types of trichomes according to the number and shape of cells of the stalk and the gland. According to his criteria, we would obtain several more trichome types, all of them present in all the taxa studied here. By contrast, other characters such as stalk length and the wall thickness of the stalk cells were not considered by Raman (1987). In our opinion, these latter characters are important in the L. depauperata group as L. arabiniana and L. depauperata subsp. hegelmaieri (A) (Fig. 5B and C) only have short hairs, while the rest of taxa have both short and long ones (Fig. 5A, D±F). This character was also used by Juan et al. (1996) to distinguish taxa of Antirrhinum from west Andalucõ a (south-western Spain). The thickness of the walls of the stalk cells, in addition, distinguishes L. depauperata subsp. hegelmaieri (Cs) and L. aeruginea subsp. aeruginea from the rest of the taxa studied. This character has also been reported as taxonomically useful for several taxa of Antirrhinum by Brooker et al. (1976) and Doaigey and Harkiss (1991). TA X O N O M IC S Y N T H E S I S A taxonomic synthesis based on the results shown in the phenogram is presented: six taxa are considered, with L. cardonica and L. sulphurea described as new species. This

latter taxon includes the populations initially labelled as L. depauperata subsp. hegelmaieri from CastelloÂn province. L. arabiniana is included in the synonymy of L. depauperata subsp. hegelmaieri from Alicante. The authors prefer not to distinguish between the rest of the di€erent infraspeci®c taxa included in L. supina according to ValdeÂs (1970) and Sutton (1988), as further research to ascertain their taxonomic position is required. Linaria aeruginea (Gouan) Cav. subsp. aeruginea, Elench. Pl. Horti Matrit.: 21, 1803. 0 Antirrhinum aerugineum Gouan, Obs. Bot.: 38, 1773. 0 Linaria tristis Benth. raza aeruginea (Gouan) Sampaio, Herb. Portug.: 110, 1913. 0 Linaria supina (L.) Chaz. subsp. aeruginea (Gouan) BoloÁs & Vigo var. aeruginea (Gouan) BoloÁs & Vigo, Flora dels PaõÈsos Catalans. 3: 426, 1995. ˆ Linaria melanantha Boiss. & Reuter, Pugillus, 85, 1852. Linaria cardonica (Font Quer) Segarra & Mateu, comb. et stat. nov. 0 Linaria supina (L.) Chaz var. cardonica Font Quer, Florula de Cardo 129, 1950. 0 Linaria supina (L.) Chaz. subsp. aeruginea (Gouan) BoloÁs & Vigo var. cardonica Font Quer, Flora dels PaõÈsos Catalans. 3: 426, 1995, nom. inval. 0 Linaria aeruginea (Gouan) Cav. subsp. aeruginea var. cardonica Font Quer, Flora Montiberica, 8: 51, 1998, nom. inval. Linaria depauperata Leresche ex Lange subsp. depauperata, Prodr. Fl. Hisp. 2: 569, 1870. ˆ Linaria alpina v. pauci¯ora Lange in Willk. & Lange, Prodr. Fl. Hisp. 2: 571, 1870. ˆ Linaria supina (L.) Chaz. subsp. depauperata (Leresche ex Lange) BoloÁs & Vigo var. depauperata, Flora dels PaõÈsos Catalans. 3: 426, 1995.

Segarra and MateuÐTaxonomy of Linaria

171

Arti®cial Key for the taxa 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6. 6.

Seeds concolourous black, sometimes metallic shiny Seeds discolourous, whitish-grey or grey Corolla tube broad, 5 mm or more in width, seeds strongly concave, metallic shiny Corolla tube narrow, less than 5 mm in width, seeds ¯at or slightly concave, metallic shiny or not Flowers dark red (sometimes with tube and spur yellow). Trichomes longer than 0.1 mm, with purple cell wall thickenings Flowers of di€erent colour, trichomes various Flowers whitish, or violet Flowers yellowish, orangish or reddish Flowers whitish, sometimes with yellowish spur. Seed disc covered by white tubercles, the tubercles sometimes fused and ridge-like. Trichomes longer than 0.1 mm, present at least on surface of calyx lobes Flowers purplish, at least on corolla lips. Seed disc smooth or not totally covered by tubercles, the tubercles acute or rounded. Indumentum glabrous or glabrescent. Trichomes shorter than 0.1 mm, present only on margin or calyx-lobes Flowers yellow. Wing grey. Seed dics covered by white or whitish-grey tubercles. Indumentum pubescent, trichomes longer than 0.1 mm, present even on in¯orescence. Plants living on stony areas Flowers orangish. Wing whitish. Seed disc smooth or not totally covered by tubercles. Indumentum glabrous or glabrescent, trichomes shorter than 0.1 mm, present only on margin or calyx-lobes. Plants living on coastal or inland sand dunes

Linaria depauperata Leresche ex Lange subsp. hegelmaieri (Lange) De la Torre, Alcaraz & M.B. Crespo, Lazaroa 16: 141±158, 1996. 0 Linaria hegelmaieri Lange, Vid. Meddel Dansk Naturh. Foren Kjobenhavn: 10, 1881. 0 Linaria depauperata Leresche ex Lange var. hegelmaieri (Lange) Willk., Prodr. Fl. Hisp. 2: 569, 1870. 0 Linaria supina (L.) Chaz. subsp. depauperata (Leresche ex Lange) BoloÁs & Vigo var. hegelmaieri (Lange) BoloÁs & Vigo, Flora dels PaõÈsos Catalans. 3: 426, 1995. ˆ Linaria depauperata f. major Porta & Rigo, Iter III Hisp., n. 365, 1891, nomen in schaed. ˆ Linaria arabiniana M.B. Crespo, de la Torre & Solanas, Bot. J. Linn. Soc 116(2):135±144, 1994. -L. supina subsp. maritima auct. Linaria sulphurea Segarra & Mateu sp. nov. -Linaria depauperata Leresche ex Lange subsp. hegelmaieri auct. non. (Lange) De la Torre, Alcaraz & M.B. Crespo -L. supina auct. non. (L.) Chaz. Holotype: CastelloÂn: Catõ , pr. Rambla de la Belluga, 650 m, UTM: 30TYK57, 29 Mar. 1992, C. Fabregat & LoÂpez UdõÂas. VAB 947021 (Fig. 8). Latin description: Herba glauca, in¯orescentiam glanduloso-pubescens pilis multicellularibus dispersis, parietibus transversalis rubris; caules fertiles numerosi, 20 cm longi, decumbentes vel ascendentes, caules steriles plerumque numerosi. Folia caulium fertilium 5±21  0.5± 1.5 mm, linearia, secunda. Folia caulium sterilium sed foliis caulium fertilium parviora, 3±4 in verticillo disposita. In¯orescentia ¯oribus 3±10 praedita, in statu ¯orenti fere densa, in statu fructifero elongata. Bracteae 2±6.5 mm longae, lineares. Pedicellus erectus 0.5±4.5 mm in statu ¯orenti fructiferoque longus. Calycis lobi lineari vel oblanceolati 2.8±8.5  0.5±1.75 mm lineares vel oblanceolati. Corolla 19±28 mm longa, lutea, tubi lato 5±8.5 mm, sino labii adaxialis 1.1±3.25 mm alto, sino labii abaxialis 0.6±2.75 mm, calcari 8±13 mm longo. Capsula 5±7 mm. Semina (2.1)3±3.75 mm longa, reniformo-orbiculata, plano-convexa vel concavo-convexa, hilum angusti, ala

2 3 L. cardonica L. supina s.l. L. aeruginea subsp. aeruginea 4 5 6 L. depauperata subsp. depauperata L. depauperata subsp. hegelmaieri p.p. L. sulphurea L. depauperata subsp. hegelmaieri p.p.

0.6±1.1 mm, membranacea, cineraceis, discus reniformis, dense tuberculatus, fuscus, tuberculis densis longis acutis, cineraceis vel albis, parietes periclinalis cellularum testae convexo-tabularis, parietes anticlinales cellularum testae elevatum, juncturis intercellularibus cristatis. Description: Glaucous, annual, biennial or rarely perennial. Glabrous except for the glandular pubescent in¯orescence, the trichomes multicullular with violet incrassate walls. Fertile shoots numerous, simple or 1±3 branched, procumbent or ascending, up to 20 cm long, densely foliaceous. Sterile shoots numerous, 1±10 cm, densely foliaceous. Leaves linear to narrowly oblanceolate, 5±21  0.5±1.5 mm, secund, glaucous, obtuse, generally 3- to 4-verticillate, opposite near shoot apex and the uppermost alternate. In¯orescence racemose, dense, with 3±10 ¯owers, densely glandular pubescent, elongating in fruit 4±6 cm. Flowering pedicels patent, short, 0.5±4.5 mm, not elongating in fruit. Bracts linear to oblong, mostly erect, longer than the pedicels, 2±6.5 mm. Calyx-lobes oblanceolate, densely covered by glandular trichomes; adaxial lobe longer, 2.8±8.5  0.5±1.75 mm. Corolla 19±28 mm long, bright yellow, with orangish palate. Tube and spur with dark veins. Adaxial lip with two lobes of 1.1±3.25 mm. Abaxial lip sinus 0.6±2.75 mm. Tube broad, 5±9 mm. Spur slightly curved, acute, 8±13 mm. Capsule globose, 5±7 mm, exceeding calyx-lobes. Seeds discoid, orbicular, (2.1)3±3.75 mm, generally concave-convex, hilum narrow, disc reinform, black, tuberculate. Tubercles grey, strongly outward pointing, very dense. Wing entire, thin, 0.6±1.1 mm wide, membranous, greyish or blackish near disc. Testa cells polygonal. Periclinal wall of testa cells tabular or tabular-convex, verrucose. Anticlinal wall angular, elevate. Flowering period: April±June. Ecology: Stony areas in dry river beds and clearings of shrublands on calcareous and siliceous soils.

172

Segarra and MateuÐTaxonomy of Linaria

Distribution: Iberian Peninsula: CastelloÂn and Teruel provinces. Linaria supina (L.) Chaz. subsp. supina Dict. Jard. Suppl. 39, n. 8, 1790 0 Antirrhinum supinum L. Sp. Pl.: 615, 1753. ˆ Linaria pyrenaica DC in Lam. & DC., Fl. Fr., ed. 3, 3: 587, 1805. 0 Linaria supina subsp. pyrenaica (DC.) Rouy, Naturaliste, 5: 366, 1883. ˆ Linaria masedae Merino, Mem. Soc. Esp. Hist. Nat., 2: 489, 1904. 0 Linaria supina (L.) Chaz. subsp. masedae (Merino) Laõ nz, Inst. For. Inv. Exp.: 26, 1968. AC K N OW L E D G E M E N T S This study was supported by a Life Project grant from the Consellerõ a de Medio Ambiente (Valencia, Spain). L I T E R AT U R E C I T E D BoloÁs O, Vigo J. 1995. GeÂnero Linaria en Flora dels PaõÈsos Catalans. Barcelona: Barcino, 413±440. Brooker SE, Doaigey AR, Harkiss KJ. 1976. Comparative anatomy of the aerial parts of Antirrhinum molle and A. mollissimum. Planta Medica 30: 319±327. Canne-Hilliker JM, Kampny CM. 1991. Taxonomic signi®cance of leaf and stem anatomy of Agalinis (Scrophulariaceae) from the USA and Canada. Canadian Journal of Botany 69: 1935±1950. Carretero JL, Boira H. 1986. Linaria orbensis Carretero & Boira, Sp. Nov. Anales del JardõÂn BotaÂnico de Madrid 42: 411±414. Chater AO, ValdeÂs B, Webb DA. 1972. Linaria. In: Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, Webb DA, eds. Flora Europaea. Cambridge: Cambridge University Press, 226±236. Chavannes E. 1833. Monographie des AntirrhineÂes. Paris & Laussane. Crespo MB, De la Torre A, Solanas JL. 1994. A new Spanish species of Linaria Miller (Scrophulariaceae). Botanical Journal of the Linnean Society 116: 135±144. De la Torre A, Alcaraz F, Crespo MB. 1996. AproximacioÂn a la biogeografõ a del sector setabense ( provincia Catalano-ValencianoProvenzal). Lazaroa 16: 141±158. Doaigey AR, Harkiss KJ. 1991. Application of epidermal characters to the taxonomy of European species of Antirrhinum (Scrophulariaceae). Nordic Journal of Botany 11: 513±524. Dumortier B. 1827. Florula Belgica. Tournaic. Fabregat C. 1995. Estudio ¯orõÂstico y ®togeogra®co de la comarca del Alto Maestrazgo (CastelloÂn). PhD Thesis, University of Valencia, Spain. Font Quer P. 1950. FloÂrula de CardoÂ. Barcelona. Font Quer P. 1970. Dicionario de BotaÂnica. Barcelona: Labor. Hong DY. 1983. The distribution of Scrophulariaceae in the Holartic with special reference to the ¯oristic relationships between eastern Asia and Eastern North America. Annals of the Missouri Botanical Garden 70: 701±712. Johansen DA. 1940. Plant microtechnique. New York & London: McGraw-Hill Book Company. Juan R. 1994. MorfologõÂa y anatomõÂa en frutos y semillas de Scrophulariaceae de AndalucõÂa Occidental. PhD Thesis, Sevilla, Spain.

APPENDIX The information given in the Appendix is taken directly from herbarium records. Specimens studied L. aeruginea subsp. aeruginea. SPAIN: AlmerõÂ a. MarõÂ a, laderas de El Gavar, 30SWG7779, 26 May 1997, calizas

Juan R, Pastor J, FernaÂndez I. 1996. Estudio de microcaracteres en frutos y semillas de Antirrhinum L. (Scrophulariaceae). Acta Botanica Gallica 143: 181±190. Juan R, Pastor J, FernaÂndez I. 1999. Morphological and anatomical studies of Linaria species from south-west Spain: Fruits. Annals of Botany 84: 21±31. Laguna E, Crespo MB, Mateo G, LoÂpez-Udõ as S, Fabregat C, Serra L, Herrero-BorgonÄoÂn JJ, Carretero JL, Aguilella A, Figuerola R. 1998. Flora EndeÂmica, rara o amenazada de la Comunidad Valenciana. Generalitat Valenciana, Consellerõ a de Medio Ambiente. ColeccioÂn Biodiversidad 1: 1±443. Lange J. 1863. Pugillus plantarum imprimis hispanicarum. Videnskabelige. Meddeleser fra den naturhistoriske forening i KjoÈbenhavn 1±4: 1±58. Mateo G, Crespo MB. 1998. Manual para la determinacioÂn de la ¯ora valenciana. Flora Montiberica, MonografõÂas 3: 1±493. Neumann U, Pare J, Raynal-Roques A, Salle G, Weber HChr. 1997. Characteristic trichomes and indumentum specialization in African and European parasitic Scrophulariaceae. Botanica Acta 110: 150±158. Payne W. 1978. A glossary of plant hair terminology. Brittonia 30: 239±255. PeÂrez-Dacosta JM. 1998. Avance del geÂnero Linaria Miller en la Comunidad Valenciana. Flora Montiberica 8: 50±54. Raman S. 1987. A code proposed for the classi®cation of trichomes as applied to the Scrophulariaceae. Beitrage zur Biologie der P¯anzen 62: 349±367. Raman S. 1989a. The trichomes on the corolla of the Scrophulariaceae V: Tribe Antirrhineae Chavannes. Beitrage zur Biologie der P¯anzen 64: 357±375. Raman S. 1989b. The trichomes on the corolla of the Scrophulariaceae VII: Tribe Pediculariaceae D. Don 1825 (Rhinantheae Benth. 1835). Beitrage zur Biologie der P¯anzen 64: 377±390. Raman S. 1990a. The trichomes on the corolla of the Scrophulariaceae VI: Tribe Cheloneae Benth. Beitrage zur Biologie der P¯anzen 65: 223±234. Raman S. 1990b. The trichomes on the corolla of the Scrophulariaceae VIII: Tribe Gerardieae Benth. Beitrage zur Biologie der P¯anzen 65: 235±248. Raman S. 1991. The trichomes on the corolla of the Scrophulariaceae X: Taxa of uncertain relationship to, or within, the family. Beitrage zur Biologie der P¯anzen 66: 127±143. Riera J. 1992. Aproximacio al coneiximent ¯orõÂstic de la serra de Pina. Tesis de Licenciatura, Universitat de ValeÁncia. Rolf FJ. 1998. Ntsys, Numerical taxonomy and multivariate analysis system. Version 2.0, User guide. New York: Exeter software. Sneath PHA, Sokal RR. 1973. Numerical taxonomy. The principles and practice of numerical classi®cation. San Francisco: Freeman. Stearn WT. 1972. Botanical Latin. London: David & Charles. Sutton DA. 1988. A revision of the tribe Antirrhineae. London & Oxford: Oxford University Press. Triado J. 1995. Flora vascular de la comarca de La Plana Alta (CastelloÂn). PhD Thesis, University of Valencia, Spain. Tirado J. 1998. Flora vascular de la comarca de La Plana Alta (CastelloÂn). CastelloÂn: Servei de Publicacions de CastelloÂ. ValdeÂs B. 1970. RevisioÂn de las especies europeas de Linaria con semillas aladas. Anales de la Universidad Hispalense. Publicaciones de la universidad de Sevilla No 7.

dolomõÂ ticas, J. GuÈemes 1278, VAL 37393. CaÂceres. Finca de la Alberca, roquedos calizos, 9 Apr. 1986, Laredo & Santos, VAL 14107. CastelloÂn. Villafranca hacia San Miguel de la Puebla, YK2981, 1220 m, 25 Jun. 1988, G. Mateo 457, VAB882148; CastelloÂn. BegõÂ s, Collado del Cascajar, XK92, 15 Jun. 1984, A. Aguilella, G. Mateo & R. Figuerola, VAL 09075; CastelloÂn. SacanÄet, pico Bellida, XK91, 11 Jun. 1984, A. Aguilella, VAL 09076. Cuenca. San MartõÂ n de Boniches

Segarra and MateuÐTaxonomy of Linaria hacia Henarejos, 30SXK21, 1200 m, 9 May 1995, G. Mateo & Torres 9654 GM, VAB951349; Cuenca. Solera del GabaldoÂn, WK80, 1100 m, 3 Jun. 1989, Ricarob Ninet, VAB913171. Cuenca. Mira, pr. CanÄavedõÂ ja, 30SXJ2795, 750 m, 23 Apr. 1993, G. Mateo 7498 GM, VAB930579; Cuenca. Talayuelas, Jun. 1979, 1000 m, G. Mateo, VAB79479; Cuenca. Campillos-Paravientos, XK22, 110 m, 4 Jun. 1985, G. Mateo, VAB852981; Cuenca. Aliaguilla, 1000 m, Jun. 1976, G. Mateo, VAB76564; Cuenca. Tragacete, 1500 m, Jul. 1975, G. Mateo, VAB75260; Cuenca. UcleÂs, riscos calcaÂreos junto al monasterio, 30SWK1125, 860 m, 14 Jun. 1997, Apr.J. AraÂn & Ma.J. TohaÂ, VAB973417; Cuenca. Talayuelas, Las Lomas, laderas del barranco, 14 May 1998, D. Ballesteros Bargues, VAB981298; Cuenca. Cuenca, hoz del Jucar, ladera rocosa, umbrõÂ a, WK7438, 910 m, Moreno-Valdeolivas, VAB973840; Cuenca. BuendõÂ a, laderas pedregosas calcaÂreas, junto a la presa del embalse de BuendõÂ a, WK1372, 700 m, 22 Apr. 1993, Apr.J. AraÂn & Ma.J. TohaÂ, VAB934202; Cuenca. Priego, alrededores del pueblo, 820 m. 7 May 1997, J. GuÈemes & J. Riera 1158, VAL 37125; Cuenca. Priego, alrededores del pueblo, 820 m. 7 May 1997, J. GuÈemes & J. Riera 1159, VAL 37126. Garanada. Sierra Nevada, penÄones de San Francisco, base del primer penÄoÂn, 2000 m. comunidades de ®suras de roca, 20 Apr. 1987, J.M. Losa, VAL 025130. Guadalajara. Traid, casa de la Bujeda, suelos calizos, XL0307, 1220 m, 22 Jun. 1997, MarõÂn & Roda 1210 FM, VAB972776; Guadalajara. Traid hacia Alcoroches 30TXL0101, 1400 m, 30 Apr. 1992, G. Mateo 6171, VAB921918; Guadalajara. Zafrilla, fuente del Berro, 30TXK2152, 1320 m, 9 Jul. 1995, G. Mateo, Hernandez-Viadel, VAB960516; Guadalajara. Taravilla, pr. laguna de la Parra, 30TWL8600, 1130 m, 20 Jun. 1995, G. Mateo, C. Fabregat & S. LoÂpez-UdõÂas 1084 GM, VAB952006; Guadalajara. Anguix, Castillo, 30TWK17, 750 m. 6 May 1997, paredes del castillo, J. GuÈemes & J. Riera 1157, VAL 37123. JaeÂn. subida al Jabalcuz, 1000 m, 30SVG2876, 8 Apr. 1987, C. Fernandez LoÂpez et al. VAB951151. LeoÂn. Castillo de Cabrera, 29TQG0390, 940 m, borde de encinar, 18 May 1992, A. Penas, M.E. GarcõÂa & L. Herrero, VAB950146. Murcia. Moratalla, sierra de la Muela, 30SWH93, 1200 m. 24 May 1997, J. GuÈemes 1222, VAL 37297. Soria. MaÂrgenes del rõÂ o Duero por Berlanga, WL09, 880 m, 25 Jul. 1985, G. Mateo, VAB851485; Soria. BeratoÂn, 1040 m, arenoso sobre calizo, 24 Jun. 1994, A. Segura-Zubizarreta 43255, VAB973104; Soria. Los RaÂbanos, por Sa del Picazo, Cueva del Asno, WM4519, 1050 m, en pastizales, 30 Jul. 1997, Montamarta, VAB980545; Soria. Berlanga de Duero, hacia LumõÂ as, WL0777, 1050 m, paÂramo calizo, 22 Aug. 1997, Montamarta, VAB980300; Soria. Ventosilla de San Juan, pr. Cerro TinÄso, 30TWM52, 1200 m, 23 Apr. 1995, calizas, Montamarta, VAB955007; Soria. Torrevicente, WL07, 1200 m, pedregales calizos, 14 Aug. 1984, G. Mateo, VAB842746. Teruel. Cuevas de CanÄart, 17 Apr. 1981, A. Aguilella, VAL 13144; Teruel. Cuevas de CanÄent, 5 Apr. 1981, A. Aguilella, VAB s/n; Teruel. LadrunÄan, Hoces del Guadalope, YL10, 800 m, 4 May 1985, A. Aguilella, VAB s/n; Teruel. Corbalan, 1500 m, 23 Jun. 1984, G. Mateo, VAB842748; Teruel. CastelseraÂs, hacia AlcanÄiz, YL44, 380 m, 27 Apr. 1989, G. Mateo 2463, VAB891203; Teruel. Cedrillas a

173

CorbalaÂn, XK77, 1500 m, 29 Apr. 1988, G. Mateo 435, VAB882126; Teruel. RoÂdenas, pr. balsa de los Pozuelos, 30TXK2897, 1380 m, 22 Jun. 1995, G. Mateo, LoÂpez-UdõÂas & C. Fabregat 10604 GM, VAB953503; Teruel. La Ginebrosa, entre el rõ o Bergantes y el barranco de Torre de Arcas, 30TYL41, 600 m, 23 Apr. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB932879; Teruel. Cantavieja, sierra de la Polamita, 1600 m, Jul. 1976, Mansanet, Puche & Mateo, VABs/n; Teruel. Peracense, alto del monte San GineÂs, 30TXK2998, 1600 m, 21 Jul. 1995, C. Fabregat & S. LoÂpezUdõÂas, VAB954408; Teruel. Sierra de Gudar, puerto de Cabigordo (CorbalaÂn a Cedrillas), XK7475, 1500 m, pastizal seco calcaÂreo, 4 Jul. 1995, J.M. Esteve, VAB963620; Teruel. Puebla de Valverde, barranco de la Cedilla, 30TXK7353, 1400 m, 7 Jul. 1995, calizas, G. Mateo, C. Fabregat & S. LoÂpez-UdõÂas, VAB953918; Teruel. Villarluengo, Masõ a del Nogueral, 30TYL1406, 1150 m, 30 Apr. 1992, C. Fabregat & S. LoÂpez-UdõÂas, VAB922484; Teruel. Calomadre, valle del rõ o de la Fuente del Berro, 30TXK2370, 1250 m, 7 Jul. 1995, G. Mateo, S. LoÂpezUdõÂas & C. Fabregat, VAB953600; Teruel. Camarena de la Sierra, altos del Javalambre, 30TXK6841, 1960 m, calizas, 7 Jul. 1995, G. Mateo, C. Fabregat & S. LoÂpez-UdõÂas 11032 GM, VAB953932; Teruel. Camarena de la Sierra, altos del Javalambre, 30TXK6841, 1900 m, calizas, Jul. 1998, Mateu & Segarra, VAB s/n; Teruel. Camarena de la Sierra, pr. Ermita de San Pablo, 30TXK6445, 1700 m, 13 Jul. 1996, C. Fabregat & S. LoÂpez-UdõÂas 1084 GM, VAB963201; Teruel. Camarena de la Sierra, pr. Ermita de San Pablo, 30TXK6445, 1700 m, Jul. 1998, Mateu & Segarra, VAB s/n; Aliaga, XL90, 1200 m, 3 Jun. 1987, J.L. Lagares, VAB871504; Teruel. Mosqueruela, rambla de las Truchas, YK2277, 1250 m, 1 Jul. 1987, G. Mateo & R. Figuerola, VAB870943; Teruel. Alto del Javalambre, 1900 m, 23 Jun. 1984, G. Mateo, VAB842747. Teruel. Villastar, pr. Barranco del Tornero, 30TXK56, 1150 m, 15 Jun. 1995, C. Fabregat & S. LoÂpez-UdõÂas, VAB954347. Valencia. El Toro, Cueva del Agua, XK82, 26 Apr. 1984, A. Aguilella, VAL 09072; Valencia. El Toro, Pozo Junco, XK82, 19 Sept. 1983, A. Aguilella, VAL 09090; Valencia. Viver, El Ragudo, YK02, 6 Apr. 1984, A. Aguilella, VAL 09069. Vallanca, XK43, 900 m, 12 Jun. 1985, G. Mateo & R. Figuerola, VAB85202; Valencia. Vallanca, XK43, 900 m, Jun. 1998, Mateu & Segarra, VAB s/n; Valencia. Vallanca, XK4136, 960 m, 9 Jun. 1988, G. Mateo 054 & M.B. Crespo, VABs/n; Valencia. Puebla de San Miguel, XK5833, 1200 m, terrenos muy arenosos, 30 May 1988, G. Mateo, VAB880748; Valencia. La Puebla de San Miguel, XK63, 1500 m, 19 Jun. 1985, G. Mateo & R. Figuerola, VAB85480; Valencia. La Puebla de San Miguel, Cerro CalderoÂn, XK63, 1500 m, 19 Jun. 1998, Atienza & Segarra, VAB s/n; Valencia. La Yesa, 1400 m, 22 Jun. 1984, G. Mateo, R. Figuerola & A. Aguilella, VAB842742; Valencia. La Puebla de San Miguel (El Raco d'AdemuÂs), Cerro CalderoÂn, 30TXK6238, 1820 m, 16 Oct. 1997, J. Riera & E. Estrelles, VAL 37228; Entre Alcotas y Abejuela, 17 Apr. 1984, A. Aguilella, VAL 09077; Valencia. Andilla, La Pobleta, XK81, 17 Apr. 1984, A. Aguilella, VAL 09071; Valencia. Andilla, La Pobleta, XK81, 6 Apr. 1984, A. Aguilella, VAL 09062; Valencia. Alpuente, Corcolilla, soÁls arenosos, XK72, 2 Jun. 1984,

174

Segarra and MateuÐTaxonomy of Linaria

A. Aguilella, VAL 09063; Valencia. Castielfabib a TormoÂn, XK4452, 1150 m, 23 Apr. 1988, rodenos, G. Mateo 710, VAB882402; Valencia. Castielfabib a TormoÂn, XK4452, 1150 m, 23 Apr. 1998, calizas, Mateu, Segarra & Paula, VAB s/n; Valencia. Vallanca, 1000 m, 6 Jun. 1984, G. Mateo & R. Figuerola, VAB842745; Valencia. Muela de Arroyo Cerezo, XK3444, 1450 m, 15 Jun. 1988, G. Mateo, J.R. Nebot & C. Fabregat, VAB882166. Zaragoza. Calatayud, ladera N.O. montanÄa, en Campiel, 23 Apr. 1987, M. Alconchel Pina, VAB940526. L. cardonica. SPAIN. Barcelona. Coteaux, F. Sennen, 22 May 1925 MA 425321; Barcelona. In locus incultis pr. Barcelona. Barcelona, Mar. 1880, C. Pau, MA 109987; Barcelona. Mas Nou de la Mina; Jul. 1919, C. Pau, MA 109970; Barcelona. Cerca de Sant Cugat del ValleÂs, 22 Apr. 1935, Rocha, MA 346888; Garraf, 13 Jun. 1970, FernandezCasas 545, MA 413488. CastelloÂn. Roquetas, fuente, 31TBF7822, 700±800 m, rocas y pedregales calcaÂreos, 15 Jun. 1997, J.M. PeÂrez-Dacosta & R. TamasoÂn Monte, VAB972934; CastelloÂn. De Bojar al Ballestar, BF6006, 1000 m, pedregales calizos, 8 Apr. 1988, G. Mateo, VAB880909; CastelloÂn. Fredes, 1150 m, Jun. 1979, J. Mansanet & G. Mateo, VAB s/n; CastelloÂn. Puebla de Benifassar, Coratxar, 31TBF50, 1200±1250 m, 11 Jul. 1997, pastos montanos con erizoÂn, sobre suelos pedregosos calizos, J.M. PeÂrez-Dacosta, VAB9729233; CastelloÂn. Traiguera, valle del rõÂ o CeÂrvol, 31TBE7493, 200 m, 24 Mar. 1996, pedregales calizos, G. Mateo, C. Fabregat & S. LoÂpezUdõÂas 11226 GM, VAB961038; CastelloÂn. Vallafranca hacia Castellfort, 30TYK38, 1250 m, 30 Apr. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB947034; CastelloÂn. Benifassar et inter Fredes, 24 Jun. 1917, C. Pau, MA109971; Fredes, barranc del Salt, 31TBF61, 1000 m, Jul. 1998, rambla pedregosa, Mateu & Paula, VAB s/n; Fredes, CastelloÂn. El Baix Maestrat, font del poble, 31TBF6110, 1000 m, 12 May 1998, murs, E. Estrelles, VAL 39013. Gerona. La Molina, pista de arrastre de Supermolina, hacia los 1850 m, 28 Jun. 1969, Fernandez Casas, MA 413502; Gerona. Sagran, en los pedregales debajo de San Quirico, Jun. 1933, M. Losa, MA109931; Gerona. Olot, VolcaÁ Montsacopa; 14 Sept. 1984, A. Aguilella 2575-AAP, VAL 14069; Gerona. Nuria, Puigmal, 19 Jul. 1983, A. Aguilella VAL 13126. Lerida. Rialp. LlavoroÂs, Montesclado (PirineÂo), Yermos pizarrosos Flores violaÂceas Font Quer, 25 July. 1912, MA109986; Lerida. Llers, champs, 2 Apr. 1907, F. Sennen-334. MA 109988; Lerida. Oliola, 31TCG4839, 400 m, campo de barbecho, 8 Apr. 1985, J. Pedrol-231. MA 418493; Lerida. Oliola, 31TCG4739, 400 m, campo de barbecho, 10 Apr. 1985, J. Pedrol-507. MA 418517; Lerida. AloÁs de Balaguer, Serra Carbonera, 31TCG2839, 700 m, prados secos, calizas, 13 Apr. 1984, J. Pedrol MA 418392; Lerida. Pallars, pr. Espot, 31TCH31, ad 1700 m, in abietis raris, 09 Jul. 1977; Fernandez Casas 2020, Leal, MunÄoz Garmendia, Pueche MA 330508; Lerida. Oliola, 31TCG4739, 400 m, campo de barbecho, 10 Jun. 1985, J. Pedrol 507, MA 313838. Tarragona. Tortosa, subida hacia monte Caro, 31TBF7822, 550 m, 18 Jun. 1992, G. Mateo 6384, VAB922166; Tarragona. Mas de Barberans, Barranc de les Coves, 31TBF72, 700 m, rambla pedregosa, Jul. 1998, Mateu & Segarra, VAB s/n; Tarragona. Roquetas, monte

Caro, 31TBF7620, 1300±1430 m, roquedos y crestones calizos, 15 Jun. 1997, J.M. PeÂrez-Dacosta & R. TamasoÂn Monte, VAB972935; Tarragona. La Cenia, antes de la Font del Teix, BF6516, 1080 m, 26 Jun. 1991, G. Mateo & L. Serra 4793, VAB912276; Tarragona. Barranco de Lloret, 31TBF8216, 140 m, ramblas, 15 Jun. 1997, J.M. PeÂrezDacosta & R. TamasoÂn Monte, VAB972932; Tarragona. Delta de Tordera, CatalunÄa. Arenales, Casellas, Apr. 1948. MA 154362; Tarragona. Barranc de SeneÂn, 31TBF1235, 200 m, lecho de torrente, suelo seco y pedregoso, 15 Dec. 1997, L. SaÂez, MA 597954. Teruel. PenÄarroya dels Tastavins, barranco Escalona, 31TBF41, 700 m, 18 May 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB932768; Teruel. Puertos de Beceite, rõÂ o MatarranÄa, BF22, 800 m, M.B. Crespo et al. VAB862092; Teruel. Valderrobres, pr. rõÂ o Pena, 31TBF51, 800 m, 18 Jul. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB933025; Teruel. Pitarque, YL00, 1000 m. 12 Jul. 1996, J. GuÈemes, VAL 04910. L. depauperata subsp. depauperata. SPAIN. Alicante. Alcoi, Carrascar de Alcoi, 30SYH28, ABH 3586; Alicante. Alcoi, Carrascar de Alcoi, 30SYH1382, ABH19899; Alicante. Alcoi, Sierra dels Plans, 30SYH2181, L. Serra, ABH 9445; Alicante. Alcoi, Carrascar de La Font Roja, subiendo al Menejador 30SYH28, Jun. 1998, Mateu, Segarra & Paula VAB s/n; Alicante. Altea, Pont del Mascarat, 31SBC0080, ABH 16854; Alicante. Benifato, Barranco de Alfafar, 30SYH4083, ABH 14616; Alicante. Benitxembla, 5 May 1999, campos de almendros, Segarra, VAB 990600; Alicante. Benifato, Sierra de Aitana, 30SYH3881, ABH 35867; Alicante. Benifato, Sierra de Aitana, Font del Forat, 30SYH3882, ABH 1089; Alicante. Benifato, Sierra de Aitana, Font del Forat, 30SYH3882, Mateu & Segarra, VAB s/n; Alicante. Benimantell, Ponoig, 30SYH4577, ABH 763; Alicante. Benimantell, Sierra de Aitana, 30SYH4481, ABH 14293; Alicante. Benissa, Sierra de Bernia, 30SYH5583, ABH 523; Alicante. Benissa, Sierra de BeÂrnia, umbrõÂ a subiendo al fort, 30SYH5683, ABH 598; Alicante. Cocentaina, Alto del Montcabrer, 30SYH19, ABH 6066; Alicante. Confrides, Sierra de Aitana, 30SYH38, ABH 14056; Alicante. Confrides, Sierra de Aitana, bajo los radares, 30SYH3782, ABH 6353; Alicante. Confrides, Sierra de Aitana, bajo los radares, 30SYH3782, Jun. 1998, Mateu, Segarra & Paula, VAB s/n, Alicante. Parcent, Coll de rates, 30SYH5590, ABH 17061; Alicante. Quatretondeta, Font del racoÂ, 30SYH3388, ABH 6351; Alicante. Quatretondeta, Hort dels Frares, 30SYH3488, ABH6352; Alicante. Marina Alta, XaÁbia, cabo de San antonio, 31SBC59, 100 m, pastizal litoral, 29 Mar. 1997, J.G. Segarra, VAB981332; Alicante. BenifallõÂ m, Serra del Rentonar, 30SYH2480, 1050 m, 1 Apr. 1989, L. Serra, VAB93164; Alicante. BenifallõÂ m, Serra dels Plans, 30SYH2479, 1250 m, 19 Jun. 1990, L. Serra, VAB931642; Alicante. Alcoi, Bc. de Les Coves, 30SYH2180, 900 m, 20 May 1992, L. Serra, VAB931643; Alicante. Vall de alcalaÂ, 30SYH39, 500 m, 23 Apr. 1993, J.Oct. Soler 131 JXS, VAB933223; Alicante. Finestrat, Puig Campana, 1000 m, pedregales calizos, 19 Jun. 1984, G. Mateo & RamoÂn Figuerola, VAB842739; Alicante. De Pego a Vall d'Ebo, 600 m, 12 Apr. 1982, Mansanet, Curras & Mateo, VAB821293; Alicante. Sierra de BeÂrnia, YH58, 600 m,

Segarra and MateuÐTaxonomy of Linaria 1 Apr. 1989, claros de matorral, G. Mateo, VAB890449; Alicante. Agres, morro del Contador, 30SYH1794, 1200 m, 25 Apr. 1987, G. Mateo et al., VAB901763; Alicante. Sa de Serrella, 1000 m, 2 Jul. 1984, G. Mateo & R. Figuerola, VAB842738; Alicante. Alcoi, al pie de la pista Font RojaMenejador, 1250 m, 31 Apr. 1990, E. Laguna et al., VAB901119; Alicante. Cocentaina, alt del Montcabrer, 30SYH19, 1200 m, 15 Apr. 1988. J.R. Nebot, VAB901781; Alicante. Vall d'Ebo, 600 m, May 1982, claros de matorral calcõ cola, J. Mansanet & G. Mateo, VAB821292; Alicante. Gaianes, solana del Benicadell, YJ20, 27 May 1986, J.R. Nebot, VAB900670; Alicante. Benifato, La Marina Baixa, pr. el pas de la Rabosa, 30SYH4182, 1250 m. 22 Jul. 1997, J. GuÈemes, J. Riera & F. Marco, VAL 37109; Alicante. Aitana, font de l'Arbre, 19 Jun. 1984, A. Aguilella, Jan. Mateu, F. Boisset & J. GuÈemes, VAL 09952. Valencia. Villalonga de la Safor, Bancal fondo, 30SYJ3905, ABH 17730; Valencia. Villalonga de la Safor, Circo de la Safor, 30SYJ3805, ABH 16875; Valencia. Villalonga, umbrõ a del Azafor, 600 m, May 1980, J. Mansanet & G. Mateo, VABs/ n; Valencia. Villalonga, umbrõ a del Azafor, 750 m, May 1980, J. Mansanet & G. Mateo, VAB80697; Valencia. Bocairent, nacimiento del rõ o VinalopoÂ, 30SYH0688, 820 m, 29 Apr. 1988, matorral helio®lo, Ma Amor Cambra, VAB881804. L. depauperata subsp. hegelmaieri. SPAIN. Albacete. MugroÂn por Alpera, 1100 m, 12 Jun. 1984, G. Mateo & R. Figuerola, VAB842744. Alicante. Petrel, Arenal de Petrel, 30SXH9466, ABH 1433; Alicante. Petrel, Arenal de Petrel, 30SXH9466, ABH 18725; Alicante. Villena, Arenales de La Virgen, 30SXH8076, ABH 12676; Alicante. Villena, Arenales de La Virgen, 30SXH8076, ABH 12676; Alicante. Villena, Arenales de la Virgen, 30SXH89, 7 Apr. 1998, Mateu, Segarra & Tortajada, VAB 990594; Alicante. Villena, La Cerrata, collado de La Cerrata, 30SXH8072, ABH 33879; Alicante. Villena, Sierra de Salinas, 30SXH89, 7 Apr. 1998; Mateu, Segarra & Tortajada, VAB 990593; Alicante. Villena, La Cerrata, collado de La Cerrata, 30SXH8072, ABH 34295; Alicante. Petrel, 30SXH9466, 550 m, 20 Apr. 1988, arenales, G. Mateo, VAB880825; Alicante. Petrer, Les Valls del VinalopoÂ, L'Arenal, 30SXH9465, 500 m. 25 Jun. 1997, J. Riera & F. Marco, VAL 037087; Alicante. Petrel, arenal de Petrel, 30SXH96, 14 May 1998, Segarra, VAB 9950595; Alicante. Petrel, arenal de Petrel, 30SXH96, 3 May 1999, Segarra, VAB 9950596. (L. arabiniana.) SPAIN. Alicante. Alfaz del Pi, PenÄas del Arabõ , (Serra Gelada), 30SYH5671, 50 m, in sabulosis maritimis, 15 Mar. 1993, J.L. Solanas; A. De la Torre & M.B. Crespo, ABH 0355; Alicante. Alfaz del Pi, PenÄas del Arabõ , (Serra Gelada), 30SYH5671, 50 m, in sabulosis maritimis, 12 Apr. 1992, J.L. Solanas; A. De la Torre & M.B. Crespo, ABH 5561; Alicante. Alfaz del Pi, PenÄas del Arabõ , (Serra Gelada), 30SYH5671, 50 m, in sabulosis maritimis, 14 Apr. 1992, J.L. Solanas; A. De la Torre, M.B. Crespo & J.C. Crsitobal, ABH 6562; Alicante. Alfaz del Põ , Sierra Helada, Penyes del Arabõ , 30SYH5671, ABH 0355; Alicante. Alfaz del Põ , Sierra Helada, Penyes del Arabõ , 30SYH5671, ABH 12870; Alicante. Alfaz del Põ , Sierra Helada, Penyes del Arabõ , 30SYH5671, ABH 6562;

175

Alicante. Alfaz del PõÂ , Sierra Helada. Penyes del ArabõÂ , 30SYH5671, ABH 5561; Alicante. Alfaz del PõÂ , Serra Gelada, 3 May 1999. Segarra & Sapena, VAB 99601; Alicante, Benidorm, Sierra Helada, 30SYH5570, ABH 16373; Campoamor, Playa de La Glea, 30SXG9897, ABH 337; Alicante. Elche, Dunas de La Marina, 30SYH0723, ABH 6067; Alicante. Guardamar, UrbanizacioÂn Pinomar Km 65.5, 30SYH0512, ABH 16303; Alicante. Guardamar del Segura, 30SYH01, ABH 6070; Alicante. Guardamar del Segura, 30SYH01, BC 655348; Alicante. Guardamar del Segura, Desembocadura del rõÂ o Segura, 30SYH0621, ABH 36828; Alicante. Guardamar del Segura, Desembocadura del rõÂ o Segura, 30SYH0621, 2 Apr. 1998; Segarra & Sapena, VAB 990599; Alicante. Guardamar del Segura, Playa del rebollo, 30SYH0722, ABH 16475; Alicante. Guardamar del Segura, Playa del rebollo, 30SYH0722, 2 Apr. 1998, Segarra & Sapena, VAB 990597; Alicante. Guardamar del Segura, Playa Moncayo, 30SYH0615, ABH 16579; Alicante. Orihuela, Cerca de Campoamor, Cala de la Glea, 30SXG9898, ABH 6089; Alicante. Petrer, Loma del Arenal, 30SXH9465, ABH 19976; Alicante. Pilar de la Horadada, Rio Nacimiento, 30SXG9498, ABH 17276; Alicante. San Miguel de Salinas, Barranco de Fayona, 30SXH9605, ABH 16470; Alicante. San Miguel de Salinas, Barranco de Fayona, 30SXH9605, ABH 6090; Alicante. Santa Pola, Arenales, 30SYH12, BC 652857; Santa Pola, Playa lisa, 30SYH1029, ABH 32192; Alicante. Santa Pola, Playa lisa, 30SYH1029, 3 May 1999, Segarra & Sapena, VAB 990598; Alicante. San Miguel de Salinas, Barranco Fayona, 30SXH9605, 100 m, 27 Mar. 1992, G. Mateo 5789, F. Alcaraz et al., VAB921097; Alicante. Orihuela, pr. Campoamor, cala de LA Glea, 30SXG9898, 27 Mar. 1992, G. Mateo 5785, F. Alcaraz et al., VAB921093; Alicante. Elche, dunas de La Marina, 30SYH13, 16 May 1989, communidades de Linarion pedunculatae, M.B. Crespo & M.L. Manso, VAB892751; Alicante. Guaradamar del Segura, YH01, 8 Mar. 1987, G. Mateo & col., VAB871149; Alicante. Santa Pola, 30SYH12, 23 May 1981, arenales ®jados de las dunas secundarias, J. Molero, MA338319. L. sulphurea. SPAIN. CastelloÂn. Cervera del Maestre, gravas del lecho del rõÂ o seco o rambla de Cervera, BE6682, BE6683 y BE6782, 240±250 m, Apr.J.AraÂn & Ma. J. TohaÂ, 9 May 1993, VAB934166; CastelloÂn. Ares del Maestre, ®nca Barranc dels Horts, zona basal, 30TYK4674, 680 m, 30 Apr. 1996, C. Fabregat & S. LoÂpez-UdõÂas, VAB962335; CastelloÂn. Tirig, serra del MolloÂ, Les Paraes, 31TBE58, 560 m, 22 Oct. 1994, C. Fabregat & S. LoÂpez-UdõÂas, VAB947033; CastelloÂn. Culla, alrededores, 30TYK46, 1050 m, 1 Apr. 1994, C. Fabregat & S. LoÂpez-UdõÂas, VAB947026; CastelloÂn. Crevera del Maestre, Rambla de Cervera, 31TBE68, ABH 15615; CastelloÂn. Benasal, pr. Barranco de AlbocaÂcer, 30TYK47, 800 m, 4 Jun. 1994, C. Fabregat & S. LoÂpez-UdõÂas, VAB947019; CastelloÂn. CatõÂ , L'AvellaÂ, 31TBE48, 950 m, 28 Jun. 1992, C. Fabregat & S. LoÂpez-UdõÂas, VAB947024; CastelloÂn. Tirig, La Morellana, 31TBE48, 600 m, 29 Apr. 1993, C. Fabregat & S. LoÂpezUdõÂas, MA947032; CastelloÂn. CatõÂ , L'AvellaÂ, 31TBE48, 950 m, 26 Jun. 1992, C. Fabregat & S. LoÂpez-UdõÂas, VAB947023; CastelloÂn. Puero de Cabrillas, pr. Portell de

176

Segarra and MateuÐTaxonomy of Linaria

Morella, YK2988, 1280 m, 25 Jun. 1988, G. Mateo 606, VAB882298; CastelloÂn. Culla, rambla Carbonera, pr. els Ibarsos, 30TYK45, 330 m, 20 Aug. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB947025; CastelloÂn. Ares del Maestre, Barranco dels Horts, 30TYK4675, 800 m, pastizal montano seco en ladera, 22 Apr. 1994, C. Vives, MA943929; CastelloÂn. Rambla Carbonera, pr. los Rosildos, 30TYK56, 420 m, 28 Mar. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB947014; CastelloÂn. Rambla de Cervera, pr. Cogullada, BE6682, 280 m, pastizal seco, 23 Mar. 1996, Torres, VAB963618; CastelloÂn. Sierra del Toro, 1100 m, Jun. 1982, Mateo & Aguilella, VAB821294; CastelloÂn. Pina de Montalgrao, los CaÂrcamos, 30SYK0331, 1050 m, suelos calizos, 1 Apr. 1996, MarõÂn Campos 870 FM, VAB963721; CastelloÂn. Cervera del Maestre, rambla de Cervera, pr. Cogullada, 31TBE6782, 280 m, pedregales calizos, 23 Mar. 1996, G. Mateo, C. Fabregat & S. LoÂpez-UdõÂas, 11220 GM, VAB961032; CastelloÂn. Vistabella del Maestrazgo, Barranco del Avellanar, YK2659, 1300 m, 25 Nov. 1990, M. Rubio, VAB913414; CastelloÂn. Vallivana, 700 m, pedregales calizos, 2 Jul. 1983, Mateo & Aguilella, VAB831355; CastelloÂn. CatõÂ , pr. Mas Cremat, 30TYK58, 750 m, 16 Apr. 1992, C. Fabregat & S. LoÂpez-UdõÂas, VAB947022; CastelloÂn. Rambla de la Valltorta, 31TBE57, 400 m, 19 Sept. 1992, C. Fabregat, S. LoÂpez-UdõÂas & G. LoÂpez-UdõÂas, VAB947028; CastelloÂn. Pr. Mas del camp del FalcË, 31TBE58, 500 m, 19 Sept. 1992, C. Fabregat, S. LoÂpez-UdõÂas & G. LoÂpez-UdõÂas, VAB947030; CastelloÂn. AlbocaÂrcer, pr. barranco de La Muela, 30TYK56, 500 m, 29 Apr. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB947015; CastelloÂn. El PlaÁ de VilafameÂs, YK54, 300 m, arenas, 29 Apr. 1988, J.R. Nebot & C. Fabregat, VAB901925; CastelloÂn. Tirig, serra del MolloÂ, Les Paraes, 31TBE5080, 550 m, 8 Aug. 1992. C. Fabregat, S. LoÂpez-UdõÂas & G. LoÂpez-UdõÂas, VAB947027; CastelloÂn. Tirig, rambla de la Valltorta, 31TBE57, 400 m, 19 Sept. 1992, C. Fabregat, S. LoÂpez-UdõÂas & G. LoÂpez-UdõÂas, VAB947029; CastelloÂn. Cervera del Maestre, rambla de Cervera, 31TBE6683, 300 m, 27 Jun. 1996, C. Fabregat & S. LoÂpez-UdõÂas, VAB963111; CastelloÂn. Cervera del Maestre, Torre Beltrans, 30TYK4575, 700 m, 18 May 1992, C. Fabregat, S. LoÂpez-UdõÂas & G. LoÂpez-UdõÂas, VAB947016; CastelloÂn. Tirig, PlaÁ d'en Mig, 31TBE5481, 400 m, 1 Apr. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB947031; CastelloÂn. Ares del Maestre, Barranc dels Horts, 30TYK4776, 750 m, 28 Jun. 1996, C. Fabregat & S. LoÂpez-UdõÂas, MA963119; CastelloÂn. De Canales a SacanÄet, 1250 m, 15 Jun. 1984, G. Mateo, R. Figuerola & A. Aguilella, VAB842737; CastelloÂn. CatõÂ , pr. rambla de la Belluga, 30TYK57, 650 m, 29 Mar. 1992, C. Fabregat & S. LoÂpez-UdõÂas, VAB947021 HOLOTIPO; CastelloÂn. Sierra de EspadaÂn, 19 May 1887, C. Pau, MA109954; sierra de El Toro, supra la Almorja, C. Pau, MA109958; CastelloÂn. Vistabella (L'AlcalateÂn), cim de Penyagolosa, 30TYK2556, 1810 m. 10 Sept. 1997, J. Riera, VAL 37097. Teruel. Aguaviva, pr. rõÂ o Bergantes, 30TYL3920, 550 m, 1 Apr. 1994. C. Fabregat & S. LoÂpez-UdõÂas, VAB942659; Teruel. Manzanera, Barranco del Pozo Junco, 30TXK8327, 1400 m, 22 Oct. 1995, C. Fabregat & LoÂpez-UdõÂas, VAB954580; Teruel. Abejuela, Barranco de Santa Margarita, 30SXK8121, 1350 m, 22 Oct.

1995, C. Fabregat & S. LoÂpez-UdõÂas, VAB960229; Teruel. Aguaviva, pr. rõÂ o Bergantes, 30TYL32, 600 m, 23 Apr. 1993, C. Fabregat & S. LoÂpez-UdõÂas, VAB932864. L. supina s.l. SPAIN. Asturias. Puerto Ventana, 18 Jun. 1979, Amich, Rico & SaÂnchez, MA 330520. Asturias. Puerto San Glorio, 14 Jul. 1983, Casaseca & Fernandez Diez, MA330536; Asturias. Puerto San Glorio, 14 Jul. 1983, Casaseca & Fernandez DõÂez, MA330535; Asturias. Terrains au pico de Arvas, 27 Apr. 1864, E. Bourgeau, MA 109985; Asturias. Puerto de San Glorio, 26 Jun. 1982, G. Mateo, A. Aguilella, F. Boisset & P. GarcõÂa-Fayos, VAL 13143. Burgos. Sierra de Neila, Jul. 1932, M. Losa, MA109978. Huesca. Valle del Hecho, hayedos de la Selva de Oza, Jul. 1976, G. Mateo, VAB76568; Huesca. Jaca, rstrojos al ®nal de la pista forestal en el monte de AtareÂs, 30TXN9512, 970 m, 10 Oct. 1969, P. Montserrat, VAB871121; Huesca. BanÄos de Benasque, CH02, 2000 m, 3 Aug. 1985, G. Mateo, VAB852985; Bielsa, Jul, C. del Campo, MA 109965; Huesca. Jaca, rastrojos al ®nal de la pista forestal en el Monte de AtareÂs, 30TXN9512, 970 m, 10 Oct. 1969, P. Montserrat, MA391770; Huesca. Arguis, 1045 m, 28 Jun. 1953, J. Malato Beliz 325, MA 303563; Huesca. Sallent, aguas limpias, 30TYN1841, 1750 m, laderas sobre calizas, 18 Sept. 1986, J. Ascaso 861017 JA, MA 427137; Huesca. Sallent, aguas limpias, 30TYN1841, 1750 m, laderas sobre calizas, 18 Sept. 1986, J. Ascaso 861017 JA, MA 427137. La Rioja. LogronÄo, Rivera, 10 Jul, Zubia, MA 109961; La Rioja. LogronÄo, Viniegra de Arriba, roquedos calizos y crestas carsti®cadas entre La Risca Y Ormazal, 30TWM1355, 1750 m, 24 Jul. 1988, M.B. Fernandez de BetonÄo & J.A. Alejandre, MA468169; La Rioja. LogronÄo, Viniegra de Arriba, roquedos calizos y crestas carsti®cadas entre La Risca Y Ormazal, 30TWM1355, 1750 m, 24 Jul. 1988. M.B. Fernandez de BetonÄo & J.A. Alejandre, MA468169. LeoÂn. Puerto de las SenÄales, 26 Jul. 1979, Casaseca, Fernandez DõÂez, Amich, Rico & Sanchez, MA 330521; LeoÂn. Riega de Mampodre, MaranÄa, 30TUN229670, 1460 m, 23 Jun. 1997, prados huÂmedos, E. Puente, E. Alonso & E. de Paz, VAB973566; LeoÂn. Puerto de San Isidro, 30TUN086695, 1610±1660 m, 15 Jun. 1997, pastizal silõÂ ceo, E. Puente, E. Alonso & E. de Paz, VAB973594; LeoÂn. Puerto de San Glorio, 1800 m, 28 Jun. 1982, G. Mateo et al. VAB821295; LeoÂn. CandõÂ n, puerto de Ancares a PenÄa CuinÄa, 29TPH74, 1800 m, 27 Jun. 1994, G. Mateo, C. Fabregat & S. LoÂpezUdõÂas 9185 GM, VAB942977; LeoÂn. Ponferrada, montes Aquilianos, vertiente Norte de la Guiana, 29TPH9702, 30 Apr. 1979, E. Temprano 242 ET, MA 279034; LeoÂn. Ponferrada, montes Aquilianos, carretera de Villar de los Barrios al Morredero, 29TQG0399, 1500 m, sobre calizas, 19 Jun. 1981, Alamillo, Castroviejo, Fernandez QuiroÂs & Nieto Feliner 39 GN, MA 279035; LeoÂn. Ponferrada, montes Aqulianos, San Pedro de Montes, 29TPH9902, roquedos, 30 Apr. 1979, E. Temprano 276, GN 279033; LeoÂn. Ponferrada, los Alperrhecianos, Castillo Cornatel prope Villavieja, in locis glareosis, 600 m, 13 Apr. 1933. W. Rothmaler 40, MA109979; LeoÂn. Ponferrada, montes Aquilianos, PenÄalba de Santiago, 29TQG0299, 1300 m, sobre calizas, 31 Oct. 1982, Lansac & Nieto Feliner 819 GN, MA 278996; LeoÂn. Ponferrada, montes Aquilianos, PenÄalba de Santiago, 29TQG0299, 1350 m, gleras de pizarra, 20 Nov.

Segarra and MateuÐTaxonomy of Linaria 1982, Lansac & Nieto Feliner 824 GN, MA 278995; LeoÂn. San Emiliano-Barrios de Luna, 12 Jun. 1970, Rivas-Goday & Izco, MA 330507; LeoÂn. Ponferrada, montes Aquilianos, PenÄalba de Santiago, 29TQG0297, 1900 m, sobre pizarras, 5 Jul. 1978, E. Temprano 62 ET, MA 278999; LeoÂn. Ponferrada, montes Aquilianos, PenÄalba de Santiago, 29TQG0299, 1600 m, sobre substrato calizo, 5 Jul. 1978, E. Temprano 91 ET, MA 278998; LeoÂn. Ponferrada, montes de Valdueza (montes Aquilianos), 29TPH9903, 1000 m, en roquedos, Apr. 1979, E. Temprano 212 ET, MA 279032; LeoÂn. Ponferrada, montes Aquilianos, PenÄalba de Santiago, 29TQG0299, 1300 m, sobre calizas, 31 Oct. 1982. E. Lansac & Nieto Feliner 814 GN, MA 278997; LeoÂn. Los ApoÂstoles la Guiana, 26 Jun. 1984, Fernandez-Alonso, G. Nieto, S. Castroviejo et al. MA519000; LeoÂn. Pr. Villafeliz de Babia, 30TTN55, 1180 m, roquedo calizo, 9 Apr. 1994, C. Aedo, J.J. Aldasoro & F. MunÄoz, A 325, MA538732; LeoÂn. Ponferrada, montes Aquilianos, San Pedro de Montes, 29TPH9902, roquedos, 30 Apr. 1979, E. Temprano, 276 GN, MA279033; LeoÂn. Puerto de las SenÄales, 26 Jul. 1979, Casaseca, Fernandez DõÂez, Amich, Rico & Sanchez, MA330521; LeoÂn. PenÄalba de Santiago, 29TQH00, 1200 m. 15 Aug. 1989, J. GuÈemes 177JGH, M.B. Crespo,

177

M.J. Sanz & J.L. Solanas, VAL 0153041. Palencia. Cordillera CantaÂbrica, puerto de Piedrasluengas, 24 Jun. 1954, C. Vicioso, MA 168084; Palencia. Puerto de Piedras Luengas, 5 Jul. 1953, J. Malato Beliz 978, MA 303564. Palencia. Laguna de Las Lomas, CardanÄo de Arriba, 30TUN5763, 2120 m, pie de roquedos, granitos, 11 Jul. 1995, C. Aedo et al. CA 3567, MA560417. Santander. C. a 2Km N of Puerto de San Glorio. NE slopes with ungrazed grassland with slopes with dense heath. Few streams and snowbeds. UN584704, 1650±1670 m 19 Jul. 1985, Frost-Olsen, P. MA 544561. Soria. Montenegro de Cameros, 1500 m, 7 Apr. 1994, suelo calizo arenoso y pedregoso, A. Segura Zubizarreta 43255, VAB971443; Soria. Oncala, altos del puerto, 30TW54, 1450 m, 12 Jun. 1993, G. Mateo 7763 GM, VAB930872; Soria. La PoÂveda, sierra Cebollera, 30TWM3450, 1700 m, 4 Jun. 1994, G. Mateo, 8877 GM, VAB941906; Soria. Vinuesa, pr. Vailenguas, 30TWM2035, 1140 m, 14 May 1995, G. Mateo 9664 GM, VAB951359; Soria. Montenegro, Jun. 1926, A. Caballero, MA109981; Montenegro, Jun. 1926, A. Caballero, MA109980; Soria. Montenegro de Cameros, calizo arenoso y pedregosos 1500 m, 7 Apr. 1994, A. Segura Zubizarreta 43255, MA580475.