Gadoria (Antirrhineae, Plantaginaceae): A new genus, endemic from Sierra de Gádor, Almería, Spain Author Güemes, Jaime Jardí Botànic, Universitat de València, C / Quart 80, E- 46008 València, Spain Author Mota, Juan F. text Phytotaxa 2017 2017-03-14 298 3 201 221 http://dx.doi.org/10.11646/phytotaxa.298.3.1 journal article 10.11646/phytotaxa.298.3.1 1179-3163 13694368 Gadoria falukei Güemes & Mota , sp. nov. Brief description :—Perennial suffrutescent, densely glandular-pubescent, with homotrichous indument; glandular trichomes with 7–10 uniseriate cells form a stalk, and one spheroidal, secretory apical cell. Flowers 11–15 mm ; tube 6–7 × 3–3.8 mm . Seeds 0.8–1.1 × 0.7–1 mm . Type:— SPAIN . Almería: Dalías, Sierra de Gádor , barranco Bernal, 36º49’33.97’’N , 02º43’26.37’’W , elev. 580 m , paredones calizos con fósiles marinos (bivalvos), orientación S , zona de refugio de ganado, 12 May 2012 , L . Posadas, F . Rodríguez, J . Vílchez, F . Martínez-Hernández, J . F . Mota & J . Güemes JG-4305 ( holotype : VAL 208956 !; isotype : HUAL s/n!) Glandular-pubescent perennial suffrutescent herb with glandular multicellular, uniseriate, straight, patent trichomes, 0.15–1.35 mm long, formed by 7–10 cells at the stalk, and a apical, spherical secretory cell. Stems few, 12–25(33) cm long, procumbent, pendant, rigid, brittle. Leaves 2.5–3.7 × 1.1–2.3 cm , reniform to ovate-cordate, lobed, with entire lobes; petiole 9–15 mm , sulcate. Flowers with pedicels 7–9 mm long, slightly larger than the calyx, shorter than the petiole, arched, erect-patent in flower. Calyx-lobes accrescent, 5–6 × 1.5–2 mm in flower, to 7.5–8 × 4–4.5 mm in fruit, ovate, acute, glandular-pubescent. Corolla 11–15 mm long, entirely yellow, without purple veins; tube 6–7 × 3–3.8 mm , scarcely gibbous at base, the gibossity 0.5–1.5 mm , hidden by the calyx-lobes; adaxial lip with two lobes 4.5–5 mm wide and sinus 3.5–4 mm ; abaxial lip with central lobe 2–2.3 mm wide, and lateral lobes 2.5–3 mm wide, sinus 3–3.5 mm ; palate yellow. Capsule 8.5–9 × 7–7.5 mm , glandular-pubescent, with glandular multicellular, uniseriate, patent trichomes. Seeds 0.8–1.1 mm , dark brown. Etymology :—The new species is dedicated to Francisco Rodríguez (Faluke), a tireless explorer from Sierra de Gádor, and an amateur botanist, who understood the rarity of those plants that hung from the walls of a cave in the Sierra. Additional specimens examined ( Paratypes ) :— SPAIN . Almería : Dalías , Sierra de Gádor , barranco Bernal , 36º49’33.97’’N , 02º43’26.37’’W , elev. 580 m , paredones calizos con fósiles marinos (bivalvos), orientación S , zona de refugio de ganado, 22 January 2012 , L . Posadas, F . Rodríguez & J . Vílchez s/n ( VAL 206223 !) ; same locality, pr. Peñón de Bernal , 36º49’33.97’’N , 02º43’26.37’’W , elev. 585 m , covacha umbrosa de materiales calizos, 05 May 2012 , J . F . Mota, L . Posadas, F . Martínez-Hernández, F . Rodríguez & A . Ivorra s/n ( HUAL 25970 !) . Illustrations: Figs. 1–3 . Additional specimens examined ( Asarina procumbens ) :— SPAIN . Barcelona : Montseny , pr. Santa Fe , 20 July 1994 , G . Mateo ( VAL 85741 !) ; Gerona: Pau, camino entre Vilaür y las Torroelles, 42º17’8.59’’N , 03º07’5.21’’E , brecha entre bloques graníticos, 2 July 2014 , J . Güemes JG-5102 ( VAL 231560 !) ; Gerona : Nùria , 19 July 1983 , J . Vigo & G . Mateo ( VAL 111316 !) ; Lérida: Carretera N 260 , a 2 km de Martinet , de la Seu d´Urgell a Bellver de Cerdanya , 42º22’07.74’’N , 01º39’59.06’’E , 25 April 2006 , J . Güemes, P . Blasco & E . Carrió ( VAL 181277 !) . Phenology :—Flowering: from April to June; fruiting: from May to July. FIGURE 1. Gadoria falukei Güemes & Mota : A) flowering branch with mature capsules from last year; B) detail of stem tricomes; C) middle leaf; D) detail of leaf trichomes; E) flower in frontal view; F) flower in lateral view; G) flower in longitudinal section; H) stamens with anther partially open; I) gynoecium; J) ovary in cross section at medium portion; K) leafy bract and immature capsule; L) mature capsule with the pores opened; M) seed. Drawing by Vicky Schwarzer from VAL 208956. FIGURE 2. Gadoria falukei Güemes & Mota : A) flower in lateral view; B) flower in oblique view; C) flowering branch apex; D) mature capsules; E) rupiculous habit. Photos by Luis Posadas. TABLE 2. Gadoria falukei Güemes & Mota , chromosome measures and centromeric position. From abbreviations: C, chromosome; CFL, chromosomal full length in μm; LAL, long arm length in μm; SAL, short arm length in μm; r, ratio of arms; N, chromosome type according to Levan et al. (1964) [M, metacentric with arm ratio = 1; m, metacentric with arm ratio = 1-1.7; sm, submetacentric with arm ratio = 1.8–3.0].

C	CFL	LAL	LAL	r	N
01. I	1.7	1	0.7	1.42	m
02. Q	1.5	0.9	0.6	1.5	m
03. C	1.5	0.9	0.6	1.5	m
04. P	1.4	0.9	0.5	1.8	sm
05. K	1.4	0.9	0.5	1.8	sm
06. H	1.5	0.8	0.7	1.14	m
07. B	1.4	0.8	0.6	1.33	m
08. F	1.3	0.8	0.5	1.6	m
09. E	1.3	0.8	0.5	1.6	m
10. R	1.4	0.7	0.7	1	M
11. J	1.4	0.7	0.7	1	M
12. N	1.3	0.7	0.6	1.17	m
13. O	1.2	0.7	0.5	1.4	m
14. L	1.2	0.7	0.5	1.4	m
15. G	1.2	0.7	0.5	1.4	m
16. D	1.2	0.7	0.5	1.4	m
17. A	1.2	0.7	0.5	1.4	m
18. M	1.1	0.7	0.5	1.4	m

Chromosomal studies :— Chromosome number 2n = 2x = 18, with symmetrical karyotype ( A2 = 0.1) and small chromosomes of 1.1–1.7 μm ( Figure 4 ). Based on measurements taken, the karyotypic formula is 2 M + 14m + 2sm. The arm length values, overall dimensions, type of chromosome and r-value are shown in Table 2 . Homologous pairs could not be identified ; therefore, the ideogram has been prepared with all chromosomes ordered consecutively ( Figure 4 ). Pollen morphology :—Pollen grains prolate (P/E = 1.3), with equatorial axis 18.1 μm (16–19 μm) and polar axis 24.5 (22–25.5 μm); trizonocolporate, with perforated-tectate exine, with perforations smaller than 0.5 μm ( Figure 3 ). Phylogenetic analysis :—The BI and MP analysis of ITS sequences yielded similar topologies, with BI displaying higher support values ( Figure 5 ). The BI analysis using GTR + I + G as the simplest model of DNA evolution reached equilibrium after 360,000 generations.The MP analysis rendered 400 most-parsimonious trees of 951 steps [consistency index (CI) = 0.548; retention index (RI) = 0.678]. The results are consistent with those of Vargas et al. (2004) since the tree topology of the MP and BI analysis showed the same pattern to their MP and BI trees and revealed the same six major clades ( Linaria , Gambelia , Anarrhinum , Maurandya , Chaenorhinum and Antirrhinum ) with high bootstrap (BS) and Posterior Probability (PP) values (only exception for the MP analysis with less than 75% BS support for the Chaenorhinum group). The Maurandya group, including the genera Maurandella (A. Gray 1868: 375 ) Rothmaler (1943: 26) , Rhodochiton Zuccarini ex Otto & A. Dietrich (1833: 153), Lophospermum D. Don (1827: 351) , Cymbalaria Hill (1756: 113) and Asarina , is well defined with a 100% BS support and PP value of 79. This clade includes two well supported monophyletic groups: the first is composed of the two specimens of Gadoria falukei (100% BP, 100 PP); and the second group is formed by the ITS sequences of Maurandella antirrhiniflora , Rhodochiton atrosanguineum and Lophospermum erubescens (90% BP, 100 PP). Reproductive biology :—Tests conducted in 2013, 2014 and 2015 showed similar results highlighting the high spontaneous autogamy in G. falukei . In all cases, fruit production was close to 100% and seed production to 90% ( Table 3 ). Ecology and biogeography :— Gadoria falukei is a new Iberian endemic from Sierra de Gádor (Almería province), in the Gadorense district, Alpujarreño–Gadorense sector, Murcian–Almeriensian biogeographical province, in the Mediterranean region ( Rivas-Martínez 2007 ). The only known population of G. falukei is found in the semiarid thermomediterranean bioclimatic zone, at an elevation of about 580 m , on almost vertical rock faces and overhanging cliffs that are composed of coastal Miocene conglomerates (Vicar unit). These marine conglomerates, which transition to calcarenite of an ancient marine platform, are surrounded by Alpujarride’s dolomites that make up Bernal’s Peñon ( IGME 1983 ). Despite Triassic materials being predominant in the area, G. falukei lives exclusively on much more recent friable conglomerates, representing an edaphic island in the sense of Rajakaruna (2004) . TABLE 3. Results of tests of spontaneous autogamy in Gadoria falukei Güemes & Mota for years 2013, 2014 and 2015, and data of another autogamy test in Maurandyinae species. *The taxonomy and nomenclature of Elisens’ papers have been actualized according Sutton (1988) .

Specie*	Year	Flowers	Ovule per	Fruit set %	Seeds per	Seed set %	Reference
number	flower, mean	fruit, mean
examined	(range)	(range)
Asarina procumbens Mill.	pr. 1985	25	--	100	--	--	Elisens, 1985a
Asarina procumbens Mill.	2016	25	294.454 (198-316)	100	267.36 (175-296)	90.80	This work
Epixiphium wislizeni (A. Gray) Munz	pr. 1985	14	--	100	152.6 (117-193)	--	Elisens, 1985b
Gadoria falukei Güemes & Mota	2013	20	106.7 (80-125)	95	111.7 (98-124)	88.16	This work
Gadoria falukei Güemes & Mota	2014	20	109.8 (85-123)	100	117.9 (105-133)	90.83	This work
Gadoria falukei Güemes & Mota	2015	20	103.9 (83-113)	95	110.6 (99-121)	89.26	This work
Holmgrenanthe petrophila (Coville & C.V. Morton) Elisens	pr. 1985	6	--	100	47.7 (28-68)	--	Elisens, 1985b
Lophospermum erectum (Hemsley) Rothm.	pr. 1985	19	--	100	255.0 (158-329)	--	Elisens, 1985b
Lophospermum erubescens D. Don	pr. 1985	20	--	100	553.0 (451-642)	--	Elisens, 1985b
Lophospermum purpusii (Brandegee) Rothm.	pr. 1985	25	--	100	92.2 (72-140)	--	Elisens, 1985b
Lophospermum scandens D. Don	pr. 1985	25	--	42	83.0 (12-127)	--	Elisens, 1985b
Lophospermum turneri Elisens	pr. 1985	21	--	100	395.0 (140-197)	--	Elisens, 1985b
Mabrya acerifolia (Pennell) Elisens	pr. 1985	29	--	100	77.6 (24-120)	--	Elisens, 1985b
Mabrya coccinea (I.M. Johnst.) Elisens	pr. 1985	25	--	61	29.5 (10-41)	--	Elisens, 1985b
Mabrya geniculata (B.L. Rob. & Fernald) Elisens	pr. 1985	12	--	0	96.4 (72-114)	--	Elisens, 1985b
Mabrya rosei (Munz) Elisens	pr. 1985	25	--	100	168.2 (113-212)	--	Elisens, 1985b
Maurandella antirrhiniflora (Willd.) Rothm.	pr. 1985	25	--	100	92.6 (28-149)	--	Elisens, 1985b
Maurandya barclaiana Lindl.	pr. 1985	25	--	100	132.2 (60-236)	--	Elisens, 1985b
Maurandya scandens (Cav.) Pers.	pr. 1985	25	--	100	134.7 (48-215)	--	Elisens, 1985b
Rhodochiton atrosanguineum (Zucc.) Rothm.	pr. 1985	25	--	25	85.7 (17-169)	--	Elisens, 1985b

Gadoria falukei inhabits a rupicolous plant community characterized by both low species richness and low vegetative cover. The most abundant species is Lafuentea rotundifolia Lagasca (1816: 19) , which is characteristic of the thermophilous alliance Cosentinio-Lafuenteion rotundifoliae A. Asensi, Molero Mesa, Pérez Raya, Rivas-Martínez & F. Valle (1990: 85), and brings together the chasmophytic and chomo-chasmophytic communities of the southeast Iberian Peninsula. Adiantum capillus-veneris Linnaeus (1753: 1096) , a pteridophyte that typically inhabits moist, seeping crevices, is also present at the locality, but its abundance is low due to limited water availability, a factor that may also explain the rarity of G. falukei . In any case, floristic composition and vegetation coverage of the rock-dwelling communities inhabited by this species is very different from the much more widespread neighboring rupicolous communities on Triassic dolomites and limestones, which are dominated by Teucrium intricatum Lange (1864: 21) and Athamanta vayredana ( Font Quer 1926: 3 ) C. Pardo (1981: 165) . FIGURE 3. Comparative scanning electron micrographs. Gadoria falukei Güemes & Mota : A) capsule, aperture; C) pollen grains; E) seed, lateral view; G) glandular trichomes in adaxial surface leaf; Asarina procumbens Mill. : B) capsule, aperture; D) pollen grains; F) seed, lateral view; H) glandular trichomes in adaxial surface leaf. Photos by M. Pilar Blasco. FIGURE 4. Gadoria falukei Güemes & Mota. A ) metaphase plate, scale 50 μm; B) chromosome ideogram. Scale in μm. Photos and drawing by Luis Delgado. TABLE 4. Accessions for the ITS-sequence study of Antirrhineae including: natural distribution of taxa, locality of wild populations, collection voucher, GenBank accession numbers, and reference of publication data.