Morphometric analysis and bioclimatic distribution of Glebionis coronaria s.l. (Asteraceae) in the Mediterranean area.

We present a revision of Glebionis coronaria in the Mediterranean area based on: a) micro-morphology of the disc floret cypselas observed with a high-resolution confocal microscopy; b) measurements of the disc cypsela with a stereoscopic microscope - duly scaled; c) its distribution in several bioclimatic belts; d) field observations; e) comparisons of herbarium samples. Because of this study, we propose the elevation of Glebionis coronaria var. discolor to the rank of species, as Glebionis discolorcomb. & stat. nov., based on morphological and ecological characteristics such as the disposition of the intercostal glands, the size of the disc cypsela wings and its distribution according to the bioclimatic belts. Glebionis coronaria, with totally yellow ray florets and intercostal glands aligned, is exclusive to the thermo-Mediterranean bioclimatic belt, while Glebionis discolor, with white ray florets on a yellow base and intercostal glands arranged randomly, is found in the thermo- and meso-Mediterranean belt. Illustrations of micromorphological characteristics of the cypselas, an identification key, a taxonomic synopsis including information on nomenclatural types, synonyms, descriptions of the taxa, and, as supplementary information, a list of the specimens examined and bioclimatic classification of samples localities are also presented.

Introduction

The genus Cass. ex Spach is present in the Mediterranean area with two species: (L.) Cass. ex Spach (= L.) and (L.) Fourr. (= L.).

For the first species, d’Urville (1822) described the variety with yellow ray florets as d’Urv., and the other with white ray florets with a yellow base as d’Urv. The only character used by d’Urville to distinguish the two varieties was the colour of the ray florets.

Cassini (1826) gave the first description of the genus based on the species Desf., and published the new combination based on , wich was described later by Spach (1841). Subsequently, Pau described a new species under the name of for the island of Ibiza with the following diagnosis: “Intermedio entre el y el , pero más afine del primero, del cual difiere por las hojas simplemente pinado-cortadas; los aquenios son muy parecidos, pero carecen de alas tan pronunciadas, y sólo llevan una. ..... lígulas blanquecinas, en la base amarillas, apenas festonadas en la terminación;.....” (Pau 1899). Recently, Rosselló and Sáez (2001) designated a lectotype of Pau (MA 128240) from a specimen collected by Pau on the island of Ibiza, emphasizing that the type material is indistinguishable from other Balearic and Spanish accessions of L.

Many authors recognize these two different entities (Fiori 1923, Rechinger 1936, Valdés et al. 1987, Vogt and Aparicio 1999, Bacchetta 2006, Sell 2006, Abd El-Twab et al. 2008, Chilton and Turland 2008, Cano et al. 2012, 2013). Turland (2004) proposes to maintain the name L. as the conserved name to designate the type of L. [Typus: Greece, Kriti (Crete): Nomos Irakliou, Eparhia Kenourgiou, 500 m E of Gangales, E side of road to Vali (), 250 m, large field with crop, 13 Apr 2003, Kyriakopoulos & Turland sub Turland 1166 (UPA; isotypi: B, BM, MO), typ. cons. Humphries (in Jarvis et al., Regnum Veg. 127: 33. 1993)] previously proposed a lectotype of after the lectotypification of Dillon (Herb. Clifford: 416, no. 1, fol. 1 – BM). However, this specimen cannot be used for the lectotypification as it clearly presents ray florets with a darker base.

Turland (l.c.) also confirmed the differentiation of the two varieties and proposed a new combination under the name of (d’Urv.) Turland (Basionym: d’Urv. in Mém. Soc. Linn. Paris 1: 368. 1822). Turland (l.c.) notes that the two entities appear to be widespread in the Mediterranean region and show no obvious correlation with geographic distribution.

From the karyological point of view the two varieties of are both diploid, with 2n = 18 (Pavone et al. 1981 Strother and Watson 1997, Vogt and Aparicio 1999, Inceer and Hayirlioglu-Ayaz 2007, Paciolla et al. 2010, Lograda et al. 2013). Abd El-Twab et al. (2008) confirm this account and point out that the chromosome complement of consists of 18 median-centromeric chromosomes, while consists of 16 median- and 2 sub-median-centromeric chromosomes.

The aims of this paper were: (a) to highlight and compare some important micromorphological characters of the two entities of ; (b) to relate their taxonomic differences with their bioclimatic characteristics; (c) to indicate new informative characters for identification of these two taxa; (d) to prepare a key, make a more complete description and provide notes on ecology and distribution of these two entities.

Methods

Sampling areas

To clarify the morphological and ecological characters of the two varieties, we carried out several samplings in different areas of the Mediterranean basin: Sicily, southern Italian Peninsula (Calabria), and Iberian Peninsula (southern Spain and Portugal) (Fig. 1).

Figure 1.

Sampling areas.

The sampling was on bioclimatic criteria and according to the climate classification of Rivas-Martínez and Rivas-Saenz (1996-2009). A statistical analysis was performed with T-Student to establish a possible relationship between the two entities and bioclimatic belts.

Plant material

A micro- and macro-morphological study was made of sampled plants from pure non mixed populations. All the specimens collected in the field are conserved in the herbaria of Jaén (JAEN) and Reggio Calabria (REGGIO). We have also consulted the following herbaria which have specimens proceeding from eastern Mediterranean regions, the source location of the species originally described by Linnaeus: REGGIO, JAEN, FI, MS, CAT, SEV, VAL, COFC, MA. All 194 examined specimens are listed alphabetically by country in Appendix 1.

Seeds of obtained from pure populations in southern Portugal and Sicily and seeds of obtained from pure populations in Jaén (Spain) were cultivated for three years. Both specimens were cultivated in the thermo-Mediterranean town of Andújar (Spain) and in the meso-Mediterranean town of Jaén, where they were grown separately and together to determine their vigour and the permanence of the characters.

High-resolution confocal microscopy was used to study the micro-morphology of the disc floret cypselas. A total of 880 cypselas (322 of the entity with yellow ray florets and PageBreak558 of the entity of white ray florets) were measured by taking images with a stereoscopic microscope –duly scaled– of both entities from different populations of plants cultivated in Portugal, Spain and Italy. The measurements were based on several observations ranging from 296 for the variety with yellow ray florets, to 425 for the variety with white ray florets; a statistical treatment was then applied using the XLSTAT programme.

Using these samples, measurements were taken of the length and width of the disc cypselas (excluding ventral and dorsal wings) and the width of the ventral wings (Table 1). We added a measure of the glands dispersion in each cavity formed between the ribs of the disc cypselas. To measure the degree of glands dispersion, a linearity coefficient (Lc) is proposed. A two-pixel wide straight line was drawn on the image between the two most separated glands in length within the group. The glands in contact with the straight line (A) were counted, and these glands were related to all the glands occupying the cavity (T). For cypselas whose morphology was not straight, but whose glands were aligned, two or more lines were used to count the aligned glands, applying a correction factor depending on the number of lines used (C). The formula and its correction are as follows:

Table 1.

Disc cypsela measurements of and comb. & stat. nov.

Characters	Parameters	Species
		G. coronaria	G. discolor
Wing Width	No. observations	298	425
	Mean (mm)	0.741	0.557
	Int. for the mean of 95% (mm)	(0.719; 0.762)	(0.543;0.572)
	Student’s test p value	< 0.01
	Z test p value	< 0.01
Disc Cypsela Width Without Wing	No. observations	315	425
	Mean (mm)	1.960	1.932
	Int. for the mean of 95% (mm)	(1.905; 2.015)	(1.856; 2.007)
	Student’s test p value	0.552
	Z test p value	0.552
Disc Cypsela Length	No. observations	313	424
	Mean (mm)	2.740	2.830
	Int. for the mean of 95% (mm)	(2.678; 2.803)	(2.792; 2.868)
	Student’s test p value	0.016
	Z test p value	0.016
Linearity Coefficient (Lc)	No. observations	193	356
	Mean	0.683	0.473
	Int. for the mean of 95%	(0.661; 0.706)	(0.455; 0.490)
	Student’s test p value	< 0.01
	Z test p value	< 0.01
Ratio Cypsela-Wing Width	No. observations	296	425
	Mean	2.771	3.740
	Int. for the mean of 95%	(2.676; 2.866)	(3.556; 3.923)
	Student’s test p value	< 0.01
	Z test p value	< 0.01

Lc= (A-1)/T – (C-1)/A,

where (C-1)/A is the correction factor. If only one line is used, it is = 0.

Lc Linearity coefficient

A Aligned glands

T Number of glands in the valley

C Number of straight lines used

Results

To verify the observations made in the field, both varieties (from pure populations in different regions) were cultivated from seeds in the two bioclimatic belts for three years. In the thermo-Mediterranean belt, the seeds of both entities sprouted and produced plants that maintained their characters unchanged from year to year. In the meso-Mediterranean belt both seed entities sprouted initially; however only the white floret variety completed its life cycle and maintained its characters.

According to Heywood (1976), sessile non-mucilaginous glands are present between the ribs of cypselas in both varieties. However, after careful observation (Tab. 1), we noticed that in the variety with yellow ray florets these glands were neatly arranged between the ribs (Fig. 2a), while they were disordered in the variety with white ray florets (Fig. 2b).

Figure 2.

Disc cypsela of (a) and (b) photographed with high-resolution confocal microscopy.

Other characters that differentiate the two entities are the width and shape of the abaxial wing of the disc floret cypselas. In the yellow floret variety, this wing is wider and the distal tip is facing upward, while in the white floret variety it is narrower and not facing upward (Table 1, Fig. 2a–b).

Both the arrangement of the glands in the intercostal spaces and the wing width are good characters –among others– for differentiating the two entities, as can be seen from the statistical study (Figs 3, 4). The linearity coefficient was used to measure objectively the arrangement of the glands in the intercostal spaces.

Figure 3.

Box plot of alignment of glands distributed along the cypselas of and (Lc = Linearity coefficient).

Figure 4.

Statistical analysis by box plot of cypselas wing width of and .

In the boxplot (Fig. 3), the Linearity coefficient of the glands present in the intercostal valleys of the inner cypselas can be observed. In both species, they do not overlap, so it is an important differentiator character: it is therefore that both taxa present morphological differences in the arrangement of the glands.

As for the boxplot analysis of the wing width measurements of the cypsela (Fig. 4), it is observed as this character is also different in both taxa, by not overlapping measures significantly and having a bounded variance.

However, the ratio cypsela-wing width (Fig. 5), the measures of width (Fig. 6) and length (Fig. 7) of the disc cypselas, are not adequate parameters to differentiate both taxa, since the overlap of the measurements is evident. Although the cypsela length is statistically different between both taxa, as can be seen in Table 1.

Figure 5.

Statistical analysis by box plot of ratio cypsela-wing width of and .

Figure 6.

Statistical analysis by box plot of disc cypselas width of and .

Figure 7.

Statistical analysis by box plot of disc cypselas length of and .

An average confidence interval of 95% was used in the statistical treatment. A parametric distribution analysis was applied and gave a P-value with a significance of less than 0.05 in the Student’s T test and the Z test. The margin of error is < 1.62 % in the case of the length of the disc cypsela, and < 0.01% for the arrangement of glands (linearity) and the ratio cypsela-wing width of the disc cypsela (Table 1).

In the analysis of the width of the disc cypsela for the two species, the P value is > 0.05, The character of width and length of disc cypsela therefore does not have much strength in differentiating the species (Figs 6, 7).

According to the Worldwide Bioclimatic Classification System proposed by Rivas-Martínez and Rivas-Saenz (1996-2009), the localities in which the two entities were sampled, fall in two bioclimatic belts: thermo-Mediterranean and meso-Mediterranean [Fig. 8 and Appendix 2].

Figure 8.

Thermoclimatic distribution of (thermo-Mediterranean) and (thermo and meso-Mediterranean) selected samples studied.

The thermo-Mediterranean belt is differentiated into the lower (with 400

Specifically, was sampled in 84% of the 32 stations in the thermo-Mediterranean belt, while was sampled in 34% (Table 2).

Table 2.

Distribution of and comb. & stat. nov. selected samples studied, related to the different bioclimatic belts.

Bioclimatic belts	N. localities	G. coronaria		G. discolor		Total n. of samples
		N. of samples	%	N. of samples	%
Upper Infra-Mediterranean	11	10	91%	2	18%	12
Lower thermo-Mediterranean	32	27	84%	11	34%	38
Upper thermo-Mediterranean	50	25	50%	28	56%	53
Lower meso-Mediterranean	25	8	32%	19	76%	27
Upper meso-Mediterranean	4	1	25%	3	75%	4
Lower meso-Temperate	1	0	0%	1	100%	1
Total	123	71		64		135

The two entities are more or less equally distributed in 50 stations in the upper thermo-Mediterranean belt: was sampled in 50% and was sampled in 56% (Table 2).

was sampled in 32% of the 25 stations in the lower meso-Mediterranean bioclimatic belt (285

Only was sampled in the single station in the lower meso-temperate belt (Table 2).

On this basis, the application of the X2 test (=0,00247) highlighted the high significance of the preferential distribution of samples in the warmer belts (infra- and lower thermomediterranean), while was observed to have a significantly greater presence in cooler belts (meso- and upper thermomediterranean) than .

Discussion

D’Urville (1822) describes two varieties of – and – taking into consideration only the external female ray floret colour.

Specimens with totally yellow ray florets are now treated as (L.) Cass. ex Spach. Also, Turland (2004) considers these two entities as distinct taxa treated at the rank of variety, and proposes a new combination in for var. ( (d’Urv.) Turland, comb. nov. – Basionym: d’Urv. in Mém. Soc. Linn. Paris 1: 368. 1822). This author also maintains that the two varieties may appear in independent or mixed populations, with no difference in distribution. We cannot agree with this author, as our sampling carried out in Sicily, southern Italy, Spain and Portugal, and our observations of specimens from Great Britain (Gibraltar), France, Croatia, Greece, Turkey, Cyprus, Malta, Israel, Egypt, Morocco and Libya reveal that the is distributed exclusively throughout the whole of the thermo-Mediterranean belt with thermo-climatic values of It/Itc = 350–450; while is found throughout the thermo- and meso-Mediterranean belt with values of It/Itc =220-350 (Tab. 2), but it is more represented in percentage terms in stations in the meso-Mediterranean belt.

An entity at the specific level of with bicolour ray florets was previously described by Pau (1899) as . In our opinion, this species is different from d’Urville. According to the analysis of the herbarium sample (MA 128240) and from the description given by Pau (1899): “Intermedio entre el y …. lígulas blanquecinas...; aquenios calvos, los externos trigonos con una sola ala…”, Pau is a probable hybrid of and . In fact, lacks the characters of and has external cypselas with two wings and two dorsal ribs. , however, has only one wing on the cypsela and leaves that are clearly like those of .

Moreover, our studies on the morphology of disc cypselas using high-resolution confocal microscopy, morphometric analysis and statistical techniques have revealed sufficient differences to justify raising the variety to a higher rank. Since two subspecies cannot coexist in the same geographic area and even less in the same habitat (criterion of allopatry), we consider them to be two distinct species.

For all these reasons, we propose a lectotypification and a change in rank for d’Urville. The two species are listed below, with their differential characteristics highlighted.

Conclusions

The two entities traditionally included in (L.) Cass. ex Spach based on external female ray floret colour have differences in their morphological and ecological features that enable them to be attributed to two different species.

In the study of the material collected in the Mediterranean area, we can confirm that the two varieties given by d’Urville (1822) present major differences in their micro- and macro-morphological characters and their distribution. Moreover, the aforementioned characters of the cypselas are very important for the determination of herbarium specimens, as the colours of the ray florets do not persist when the plants are dried.

Since is conserved in the form of plants with yellow ray florets, corresponding to d’Urv. and necessarily to , we establish a change of rank for the var. d’Urv. Both entities present clear differences in the colour of their ray florets, the shape and size of their disc cypselas and in the disposition of their glands. For this reason, based strictly on the ICN (McNeill et al. 2012), we maintain the species and propose comb. & stat. nov.

Taxonomic treatment

Taxonomic synopsis

(L.) Cass. ex Spach, Hist. Nat. Vég. 10: 181. 1841

≡

Note.

Glabrous plant. Stems branched, tall 20–80 cm. Leaves semi-amplexicaul, oblong or obovate, 2-pinnatisect with oblong or lanceolate segments. Involucre 10–20 mm long; outer bracts ovate, with brownish marginal bands with a whitish scarious margin; inner bracts without marginal bands but with wider scarious margins. Female ray florets with completely yellow limb. Disc cypselas 2.6–2.8 mm long, with a pronounced wing (average width 0.71–0.76 mm) and intercostal glands aligned (Table 1, Fig. 2a). 2n = 18.

Habitat.

Cultivated grounds, along the ways and waste places.

Bioclimatic distribution.

Species distributed mainly throughout the thermo-Mediterranean bioclimatic belt.

(d’Urv.) Cano, Musarella, Cano-Ortiz, Piñar Fuentes, Spampinato & Pinto Gomes comb. & stat. nov.

urn:lsid:ipni.org:names:77163641-1

Basionym:

Like but plants frequently puberulous. Female ray florets white with a yellow base. Disc cypselas 2.8–2.9 mm long, with poorly pronounced wings (average width 0.54–0.57 mm) and intercostal glands arranged randomly (Table 1, Fig. 2b). 2n = 18.

Cultivated grounds, along the ways and waste places.

Species distributed throughout the thermo-Mediterranean and meso-Mediterranean bioclimatic belt.

1	Glabrous plant. Female ray florets with completely yellow limb. Disc cypselas 2.6-2.8 mm long, with a pronounced wing (average width 0.71–0.76 mm) and intercostal glands aligned. Species distributed mainly throughout the thermo-Mediterranean bioclimatic belt	G. coronaria
–	Plants frequently puberulous. Female ray florets white with a yellow base. Disc cypselas 2.8–2.9 mm long with poorly pronounced wings (average width 0.54–0.57 mm) and intercostal glands arranged randomly. Species distributed throughout the thermo-Mediterranean and meso-Mediterranean bioclimatic belt	G. discolor

Bioclimatic classification of samples localities and realted distribution of and comb. & stat. nov.

Locality	Glebionis coronaria	Glebionis discolor	Climate	Ombrotype
Abuhir	x		Lower Mesomediterranean	Upper Arid
Agadir		x	Upper Inframediterranean	Lower Semiarid
Alahurín de la Torre		x	Lower Thermomediterranean	Lower Dry
Alcalá de Guadaira	x		Upper Thermomediterranean	Lower Dry
Alicudi	x		Lower Thermomediterranean	Lower Dry
Almería 5 km Campohermoso	x		Lower Thermomediterranean	Lower Semiarid
Almería el Ejido	x		Lower Thermomediterranean	Lower Semiarid
Almodóvar del río	x	x	Upper Thermomediterranean	Upper Dry
Almuñécar	x	x	Lower Thermomediterranean	Lower Dry
Alrededores Puente Genil	x		Upper Thermomediterranean	Upper Dry
Altea	x	x	Lower Thermomediterranean	Lower Dry
Arjona hacia Porcuna		x	Lower Mesomediterranean	Lower Dry
Arroyo de Guadalbaida posadas		x	Upper Thermomediterranean	Upper Dry
Arroyo salado		x	Lower Mesomediterranean	Upper Dry
Atenas	x		Upper Thermomediterranean	Upper Semiarid
Atenas (acropolis)	x	x	Upper Thermomediterranean	Upper Semiarid
Badajoz		x	Lower Mesomediterranean	Lower Dry
Baeza Puente Mazuecos		x	Lower Mesomediterranean	Upper Dry
Bagaladi	x		Lower Mesomediterranean	Lower Subhumid
Bu giarrar	x		Upper Inframediterranean	Lower Semiarid
Caltanissetta	x		Lower Mesomediterranean	Lower Dry
Caria	x		Upper Thermomediterranean	Lower Subhumid
Carretea Santaella-Montalbán		x	Upper Thermomediterranean	Upper Dry
Carretera Córdoba-Sevilla		x	Upper Thermomediterranean	Upper Dry
Carretera las infantas		x	Upper Thermomediterranean	Lower Dry
Carretera Lora û constantina		x	Upper Thermomediterranean	Upper Dry
Castelló de la plana		x	Upper Thermomediterranean	Lower Dry
Catalan bay	x	x	Lower Thermomediterranean	Lower Subhumid
Catanzaro	x	x	Lower Mesomediterranean	Lower Subhumid
Catanzaro	x	x	Lower Mesomediterranean	Lower Subhumid
Catarroja	x		Lower Thermomediterranean	Lower Dry
Cerca de Cerro Molina (Úbeda)		x	Lower Mesomediterranean	Lower Dry
Cossyra	x		Lower Thermomediterranean	Lower Dry
Cruce del Bembezar con arroyo Guadalora		x	Upper Thermomediterranean	Upper Dry
Dehesa de Campamor	x		Upper Thermomediterranean	Lower Semiarid
Driana	x		Upper Inframediterranean	Lower Semiarid
El Abiar	x		Upper Thermomediterranean	Upper Arid
El Gandul (Se)	x		Upper Thermomediterranean	Lower Dry
El Pla Carcaixent		x	Upper Thermomediterranean	Lower Dry
El Zumbel		x	Upper Mesomediterranean	Upper Dry
Entre Arcos y Bornos.	x		Upper Thermomediterranean	Upper Dry
Entre Écija y Herrera		x	Upper Thermomediterranean	Lower Dry
Entre el Viso del Alcor y Carmona		x	Upper Thermomediterranean	Upper Dry
Entre Morón y Montellano		x	Upper Thermomediterranean	Upper Dry
Entre Motril y Almuñecar		x	Lower Thermomediterranean	Lower Semiarid
Entre Pilas y Aznalcázar	x		Upper Thermomediterranean	Lower Dry
Epidaura	x		Upper Mesomediterranean	Lower Subhumid
Estación de Hornachuleos		x	Upper Thermomediterranean	Upper Dry
Favignana	x		Lower Thermomediterranean	Lower Dry
Filicudi	x		Lower Thermomediterranean	Lower Dry
Fiume Ferro (Ct),	x		Upper Thermomediterranean	Lower Dry
Gibraleón cercanías		x	Upper Thermomediterranean	Lower Dry
Gozo	x		Lower Thermomediterranean	Lower Dry
Hornachuleos		x	Upper Thermomediterranean	Upper Dry
Illes Balears Mallorca		x	Lower Thermomediterranean	Upper Semiarid
Insula Comino	x		Lower Thermomediterranean	Lower Dry
Insula Linosa (Aethusa)	x	x	Lower Thermomediterranean	Lower Semiarid
Insula Pianosa		x	Upper Thermomediterranean	Lower Dry
Isole Eolie (Me)	x		Upper Thermomediterranean	Upper Dry
Jabalquinto (Jaén)		x	Lower Mesomediterranean	Lower Dry
Jerez		x	Upper Thermomediterranean	Upper Dry
Kalampta	x		Upper Thermomediterranean	Lower Subhumid
Kalfa (Me)	x		Lower Mesomediterranean	Lower Subhumid
La Nucia la marina baixa		x	Upper Thermomediterranean	Lower Dry
Lampedusa	x		Upper Inframediterranean	Lower Semiarid
Larnaka	x		Lower Thermomediterranean	Lower Semiarid
Las Cabezas	x		Upper Thermomediterranean	Lower Dry
Lesina	x		Lower Mesomediterranean	Upper Dry
Linares Ciudad Camino		x	Lower Mesomediterranean	Lower Dry
Linosa	x	x	Lower Thermomediterranean	Lower Semiarid
Lipari	x		Lower Thermomediterranean	Upper Dry
Lopadusa	x	x	Upper Inframediterranean	Lower Semiarid
Los Yesares		x	Upper Thermomediterranean	Lower Semiarid
M. Grasso (Sr),	x		Upper Thermomediterranean	Lower Dry
M. Mela (Ag)	x		Upper Thermomediterranean	Lower Dry
Mallorca Algaida		x	Lower Mesomediterranean	Upper Dry
Marettimo	x		Lower Thermomediterranean	Upper Semiarid
Marmolejo		x	Upper Thermomediterranean	Lower Dry
Mogador (actual Esauria)	x		Lower Thermomediterranean	Upper Semiarid
Monte Kalfa (Me)		x	Lower Mesomediterranean	Lower Subhumid
Morón de la Frontera		x	Upper Thermomediterranean	Upper Dry
Mota del Cuervo		x	Upper Mesomediterranean	Lower Dry
Nador	x		Lower Thermomediterranean	Lower Semiarid
Nafplio	x		Upper Thermomediterranean	Lower Dry
Noto (Sr)	x		Upper Thermomediterranean	Lower Dry
Novelda	x		Upper Thermomediterranean	Upper Semiarid
Oia (Santorini)		x	Lower Mesomediterranean	Lower Dry
Oropesa la plana alta		x	Upper Thermomediterranean	Lower Dry
Oujda		x	Upper Thermomediterranean	Upper Semiarid
Pantano Bruno, Pozzallo (Rg)		x	Lower Thermomediterranean	Lower Dry
Pantano Longarini	x		Lower Thermomediterranean	Lower Dry
Paterna	x		Upper Thermomediterranean	Lower Dry
Pellaro		x	Lower Thermomediterranean	Lower Subhumid
Pellaro	x		Lower Thermomediterranean	Lower Subhumid
Pentidattilo		x	Lower Mesomediterranean	Lower Subhumid
Petra – Olimpa	x		Upper Thermomediterranean	Upper Semiarid
Piana di Catania	x		Upper Thermomediterranean	Lower Dry
Piana di Soluq û ana	x		Lower Thermomediterranean	Lower Semiarid
Porcuna San Pantaleón		x	Lower Mesomediterranean	Lower Dry
Ramath-Gan	x		Upper Inframediterranean	Lower Dry
Riba-Roja camp de túria		x	Upper Thermomediterranean	Lower Dry
Río Anzur		x	Lower Mesomediterranean	Upper Dry
Río Lucena		x	Upper Mesomediterranean	Upper Dry
S. Pedro do Estoril		x	Upper Thermomediterranean	Lower Subhumid
S.Nicola da Crissa, Serre (VV),	x		Upper Thermomediterranean	Lower Subhumid
Sagunto (V)		x	Upper Thermomediterranean	Lower Dry
Sebchet Bu Giarrar	x		Upper Inframediterranean	Lower Semiarid
Tarifa	x		Lower Thermomediterranean	Lower Subhumid
Terme di San Calogero	x		Lower Thermomediterranean	Upper Dry
Termini Imerese	x		Lower Thermomediterranean	Lower Dry
Thyra (Santorini)		x	Lower Mesomediterranean	Lower Dry
Tocra	x		Upper Inframediterranean	Lower Semiarid
Tolmeta	x		Upper Inframediterranean	Lower Semiarid
Tolmeta	x		Upper Inframediterranean	Lower Semiarid
Úbeda		x	Lower Mesomediterranean	Lower Dry
Valle del Guadiato	x		Lower Mesomediterranean	Upper Dry
Varazze		x	Lower Mesotemperate	Lower Subhumid
Vejer	x		Lower Thermomediterranean	Upper Dry
Villaviciosa de Córdoba		x	Lower Mesomediterranean	Upper Dry
Wadi El û bab	x		Upper Inframediterranean	Upper Semiarid
Xabia	x	x	Lower Thermomediterranean	Lower Dry
Xátiva	x	x	Upper Thermomediterranean	Upper Dry
Yávota		x	Lower Mesomediterranean	Lower Dry
TOTAL	71	64