Abstract
The genus Andryala includes diploid plants distributed in the Mediterranean Basin, Macaronesia and in one isolated outpost in the Romanian Carpathians. We analysed nuclear genome size in a phylogenetic framework and assessed relationships between genome size and life form (perennials vs annuals/biennials) and the consequences of insular vs continental distribution. Absolute nuclear genome size of 18 species or subspecies was determined using propidium iodide flow cytometry. The evolution of genome size was investigated using the Brownian motion model with the tree scaling parameters λ, κ and δ. The mean 2C values differs up to 1.84-fold between species (from 2.69 to 5.01 pg). Chromosome numbers of six species are reported for the first time. The highest 2C values are present in two well supported basal lineages corresponding to the relict species A. laevitomentosa and A. agardhii. The rest of Andryala species that form a well-supported clade of closely related species (the ‘Major Radiation Group’), except for two populations of A. ragusina, have significantly smaller genome sizes. In the ‘Major Radiation Group’, genome size in perennial species is significantly greater compared to annual and biennial species. With a possible bias caused by A. dentata and A. integrifolia, insular species of the ‘Major Radiation Group’ have lower nuclear genomes than continental ones. The genome size variation contains strong phylogenetic signal, which could indicate that genome size in the group under study is not greatly influenced by selection and is probably a result of neutral evolution or genetic drift.
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Acknowledgements
We are grateful to I. Álvarez, M. Benedito, E. Bräutigam, S. Bräutigam, Z. Dočkalová, I. Ferreira, R. Jardim, K. Krak, L. Medina, P. Mráz, V. Mrázová, A. Pupo Correia, M. Puskás, M. Menezes de Sequeira, T. Urfus, K. Vazačová, P. Vít, J. Vítová, P. Zahradníčková and A. Zeddam for their collaboration with field sampling and plant collecting. Fred Rooks kindly improved our English. This work was supported by the Czech Science Foundation (P506/10/1353 to J.Ch.) and the long-term research development project No. RVO 67985939 of the Czech Academy of Sciences.
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Appendix 1
Appendix 1
List of localities of population samples. Records are presented as follows: species/subspecies (in alphabetical order), population code, locality description, elevation, geographic coordinates (WGS84), date of collection and collector.
A. agardhii
JC 2011_30 – Spain, Andalusia, province Granada, Sierra de Baza Mts: Calar de Santa Barbara, 7.5 km NE of the village of Las Juntas, 1,960 m a.s.l., 37°23′05″ N, 02°51′03″ W, 31 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_30 – Spain, Andalusia, province Granada, Sierra de Baza Mts: Calar del Desabezedo, 4.6 km E of the the village of Las Juntas, 1,995 m a.s.l., 37°19′49″ N, 02°51′20″ W, 31 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_33 – Spain, Andalusia, province Almería, Sierra de María Mts: María, near a path to Maimón 3.2 km SE of the village, 1,380 m a.s.l., 37°41′29″ N, 02°08′21″ W, 2 June 2011, leg. J. Chrtek and Z. Dočkalová; JZ 2011_3 – Spain, Andalusia, Sierra Mágina: north of Huelma, along the road to Mágina, 1,584 m a.s.l., 37°42′19″ N, 3°27′45.24″ W, 11 June 2011, leg. J. Zahradníček, K. Krak, P. Vít, J. Kalůsková, T. Urfus and M. Urfusová; JZ 2211 – Spain, Andalusia, Sierra de las Cabras: road from Canada de la Cruz, saddle west of Mt. Pinar Liano, 1,806 m a.s.l., 38°04′17.58″ N, 2°20′02″ W, 12 June 2011, leg. J. Zahradníček, K. Krak, P. Vít, J. Kalůsková, T. Urfus and M. Urfusová.
A. arenaria
JC 2011_3 – Spain, Andalusia, province Malaga, vicinity of Marbella: Artola, Urbanización Las Chapas, hill near the Avenida Uno, 25 m a.s.l., 36°29′38″ N, 04°45′36″ W, 25 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_7 – Spain, Andalusia, province Cádiz: Bornos, subruderal places near the main street 2 km WNW of the town, 237 m a.s.l., 36°49′01″ N, 05°46′01″ W, 26 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_9 – Spain, Andalusia, province Huelva: Almonte, near the road A-483, 5.9 km SSW of the town, 55 m a.s.l., 37°12′35″ N, 06°30′21″ W, 26 May 2011, leg. J. Chrtek and Z. Dočkalová.
A. atlanticola
10JZ_08 – Morocco, High Atlas Mts., region Marrakech-Tensift-El-Haouz: 2 km of Around, rocks along the road from Around to Tubkal, 1,960 m a.s.l., 31°06′59″ N, 7°55′11″ W, 20 June 2010, leg. J. Zahradníček and P. Schindlerová; JC 13_V – Morocco, High Atlas Mts., region Marrakech-Tensift-El-Haouz: Imlil, slopes near the road to Asni, 2.5 km NNW of the village, 1,585 m a.s.l., 31°09′29″ N, 07°55′29″ W, 19 May 2010, leg. J. Chrtek and Z. Dočkalová; Ixis – Morocco, High Atlas Mts., region Marrakech-Tensift-El-Haouz: Imlil, saddle above the town, 2,063 m a.s.l., 31°09′56″ N, 7°54′45″ W, 20 June 2010, leg. J. Zahradníček and P. Schindlerová.
A. cossyrensis
JC 3 – Morocco, region Oriental, province Berkane, Monts des Beni Snassen: Berkane, Zegzal (Zegzel) valley, along a road near a small bridge over the stream, 6.4 km SW of the town, 280 m a.s.l., 34°52′31″ N, 02°21′28″ W, 14 May 2010, leg. J. Chrtek and Z. Dočkalová.
A. crithmifolia
260_ZF – Portugal, Madeira, south coast: Pináculo, ca 280 m a.s.l., 32°38′N, 16°53′W, 12 August 2009, leg. Z. Ferreira; 259_ZF – Portugal: Madeira, south coast: Cabo Girão, 32°39′12.72″ N, 16° 59′ 36.06″ W, 09 August 2009, leg. Z. Ferreira and I. Ferreira; 301_ZF – Portugal, Madeira, south coast: Garajau, 32°38′15″ N 16°51′02″ W, 17 July 2012, leg. Z. Ferreira, A. Pupo Correia and R. Jardim.
A. dentata
JZ 0412 – Italy, Isola di Pantelleria: Rekhale, near the road 0.9 km ENE of the village, 305 m a.s.l., 36°45′41″ N, 12°00′20″ E, 17 May 2012, leg. J. Zahradníček and J. Chrtek; JZ 0512 – Italy, Isola di Pantelleria: Tracino, near the road to Casa Bono 2.4 km SSW of the village (square), disturbed place, 220 m a.s.l., 36°46′15″ N, 12°01′44″ E, 17 May 2012, leg. J. Zahradníček and J. Chrtek; JZ 0612 – Italy, Isola di Pantelleria: Siba-Roncone, near the road 0.9 km SE of the village, 450 m a.s.l., 36°46′58″ N, 11°59′11″ E, 17 May 2012, leg. J. Zahradníček and J. Chrtek.
A. glandulosa subsp. cheiranthifolia
248_ZF – Portugal: Madeira, Pico do Areeiro, 1,542 m a.s.l., 32° 43′N 16° 55′W, 26 July 2009, leg. Z. Ferreira., A. Pupo Correia and M. Sequeira; 271_ZF – Portugal, Madeira, SW coast: Jardim do Mar, ca 25 m a.s.l., 32°44′N, 17°12′W, 3 July 2010, leg. Z. Ferreira; 241_ZF – Portugal, Madeira: Encumeada, Pousada dos Vinháticos, ca 800 m a.s.l., 32°44′N, 17°01′W, 24 July 2009, leg. Z. Ferreira., A. Pupo Correia and M. Sequeira; 257_ZF – Portugal, Madeira: Fajã da Ovelha, 625 m a.s.l., 32°46′ N, 17°12′W, 29 July 2009, leg. Z. Ferreira, M. Benedito and M. Sequeira; 269B_ZF – Portugal, Madeira, south coast: Calheta, 5 m a.s.l., 32°43′N, 17°11′W, 3 July 2010, leg. Z. Ferreira.
A. glandulosa subsp. glandulosa
233_ZF – Portugal, Madeira, north coast: Seixal, 42 m a.s.l., 32°48′N, 17°04′W, 24 July 2009, leg. Z. Ferreira., A. Pupo and M. Sequeira; Glan_ZF – Portugal, Madeira, north coast: Porto Moniz, rocks near the road, 263 m a.s.l., 32°51′46″ N, 17°10′29″ W, 23 July 2011, leg. Z. Ferreira; 298_ZF – Portugal, Madeira, north coast: S. Jorge, 126 m a.s.l., 32°50′N, 16°54′W, 15 July 2012, leg. Z. Ferreira; ZF_278A – Portugal, Madeira, west coast: Ponta do Pargo, ca 400 m a.s.l., 32°48′N 17°15′W, 3 July 2010, leg. Z. Ferreira.
A. integrifolia
KV 4 – Spain, Canary Islands, Gran Canaria: Valleseco, 0.3 km W of the village, along the road, 1,060 m a.s.l., 28°02′59″ N, 15°34′43″ W, 15 April 2007, leg. K. Vazačová (seeds); SB_H12 – Spain: Canary Islands, El Hierro: Tiñor (3 km SW of Valverde), near the road to San Andrés, 0.2 km W of the village, 980 m a.s.l., 27°47′18″ N, 17°56′10″ W, 20 May 2010, leg. S. Bräutigam and E. Bräutigam; SB H15 – Spain, Canary Islands, El Hierro: San Andrés, near a road to El Pinar, 4 km SSW of the village, 1,125 m a.s.l., 27°44′08″ N, 17°58′28″ W, 22 May 2010, leg. S. Bräutigam and E. Bräutigam; SB H16i – Spain, Canary Islands, El Hierro: San Andrés, near the road to Frontera, 2 km WSW of the village, Chamaecytisus proliferus shrubs, 1,175 m a.s.l., 27°45′41″ N, 17°58′17″ W, 22 May 2010, leg. S. Bräutigam and E. Bräutigam; ZF LM5104 – Spain, Guadalix de la Sierra, 850 m a.s.l., 40°47′33″ N, 03°42′22″ W, 19 June 2010, leg. L. Medina and M. Sequeira; AZ_3 – Algeria, Alger: towm district of Kouba, 90 m a.s.l., 36°43′N, 03°05′E, 2 July 2011, leg. Abida Zeddam; AZ_4 – Algeria, Alger: town district of Le Caroubier, 2 m a.s.l., 36°44′N, 03°07′E, 3 July 2011, leg. Abida Zeddam.
A. laxiflora
JC 2011_15 – Spain, Andalusia, province Huelva: Encinasola, near a road H-211, 4.4 km SSE of the town, 260 m a.s.l., 38°05′56″ N, 06°51′16″ W, 27 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_19 – Spain, Andalusia, province Córdoba: Hornachuelos, near a road A-2212, 3.5 km NW of the town, 280 m a.s.l., 37°51′39″ N, 05°15′57″ W, 28 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_20 – Spain, Andalusia, province Jaén: Linares, near a road J-6030, 3 km WNW of the town center, 405 m a.s.l., 38°06′37″ N, 03°39′29″ W, 28 May 2011, leg. J. Chrtek and Z. Dočkalová.
A. laevitomentosa
A – Romania, Bukovina, Suceava county, Vatra Dornei: Mt. Pietrosul Bogolin, rocks near the top (northernmost population), 1,680 m a.s.l., 47°23′19″ N, 25°32′1″ E, 5 August 2011, leg. J. Chrtek, P. Mráz, V. Mrázová and M. Puskás; C – Romania, Bukovina, Suceava county, Vatra Dornei: Mt. Pietrosul Bogolin, rocks near the top, 1,740 m a.s.l., 47°23′08″ N, 25°32′19″ E, 5 August 2011, J. Chrtek, P. Mráz, V. Mrázová and M. Puskás; JC 2011/40/2P – Romania, Bukovina, Suceava county, Vatra Dornei: Mt. Pietrosul Brostenilor, rocks in the upper part, 1,780 m a.s.l., 47°22′31″ N, 25°32′29″ E, 5 August 2011, J. Chrtek, P. Mráz, V. Mrázová and M. Puskás.
A. maroccana
JZ 1711 – Morocco, region Oriental, rocks on northern seaside of peninsula near Melilla (Spain), 74 m a.s.l., 35° 26′ 4.62″ N, 2° 58′ 1.86″ W, 9 July 2011, J. Zahradníček and K. Krak.
A. mogadorensis
JC 15 – Morocco, region Sous-Masa-Drâa: Tamri, near the main road along the Atlantic coast, 4.7 km WSW of the village, 30 m a.s.l., 30°41′33″ N, 09°52′28″ W, 22 May 2010, leg. J. Chrtek and Z. Dočkalová; JC 18/V2 – Morocco, region Doukkala-Abda: Moulay Abdallah (SW of El Jadida), near the main road 13.5 km SW of the village, 12 m a.s.l., 33°05′09″ N, 08°39′16″ W, 22 May 2010, leg. J. Chrtek and Z. Dočkalová; JC 16N – Morocco, region Marrakech-Tensift-El-Haouz: Essaouira (Mogador), coastal dunes at the northern margin of the town, 8 m a.s.l., 31°31′40″ N, 09°45′01″ W, 22 May 2010, leg. J. Chrtek and Z. Dočkalová; JC 17 – Morocco, region Doukkala-Abda: Sidi Rosia (S of Safi), near the main road 3.4 km SSW of the village, 15 m a.s.l., 32°11′37″ N, 09°15′19″ W, 22 May 2010, leg. J. Chrtek and Z. Dočkalová; ZF 263A – Morocco, region Marrakech-Tensift-El-Haouz: Essaouira (Mogador), coastal dunes at the northern margin of the town, 31°32′40″ N, 09°45′00″ W, 22 May 2010, leg. Z. Ferreira and I. Álvarez.
A. mogadorensis subsp. jahandiezii
JC 14 – Morocco, region Sous-Masa-Drâa: Inezgane, near the road between the golf resort and King’s Palace, near the river of Sous, 7 m a.s.l., 30°21′46″ N, 09°34′27″ W, 21 May 2010, leg. J. Chrtek and Z. Dočkalová.
A. perezii
KV 51V – Spain, Canary Islands, Lanzarote: Haria, near the road 2 km SW of the village, 380 m a.s.l., 29°08′04″ N, 13°30′41″ W, 21 May 2008, leg. K. Vazačová; KV 52V – Spain, Canary Islands, Lanzarote: El Jurado, Guinate, near the road 0.7 km SW of the village, 440 m a.s.l., 29°10′33″ N, 13°29′57″ W, 20 May 2008, leg. K. Vazačová.
A. pinnatifida
KV 10 – Spain, Canary Islands, Gran Canaria: Barranco de Guayadeque, 4.5 km NW of the village of Agüimes, 540 m a.s.l., 27°55′59″ N, 15°28′44″ W, 13 May 2007, leg. K. Vazačová; KV 16 – Spain, Canary Islands, Tenerife: Cumbre de Bolico, near the road 2 km SE of the village of Las Portelas, 1,155 m a.s.l., 28°18′53″ N, 16° 49′48″ W, 4 June 2007, leg. K. Vazačová; KV 25 – Spain, Canary Islands, La Gomera: Alto de Garajonay, 1.5 km NE of the village of Igualero, 1,300 m a.s.l., 28°06′30″ N, 17°14′30″ W, 6 November 2007, leg. K. Vazačová; SB G5 – Spain, Canary Islands, La Gomera: Hermigua, El Curato, near the rocks Los Gemelos, 300 m a.s.l., 28°08′58″ N, 17°12′18″ W, 9 May 2010, leg. S. Bräutigam and E. Bräutigam; SB H13 – Spain, Canary Islands, El Hierro: San Andrés, near the road to Frontera, 4 km SW of the village, shrubs with prevailing Chamaecytisus proliferus, 980 m a.s.l., 27°44′46″ N, 17°59′08″ W, 20 May 2010, leg. S. Bräutigam and E. Bräutigam; SB G7b – Spain, Canary Islands, La Gomera: Arure, at the path to Montaña de los Manantiales 2.5 km ENE of the village, Erica arborea-Cistus monspeliensis shrubs, 940 m a.s.l., 28°08′32″ N, 17°18′06″ W, leg. S. Bräutigam and E. Bräutigam; SB G9 – Spain, Canary Islands, La Gomera: Vallehermoso, 3 km SSE of the village, at the road to Las Rosas, 980 m a.s.l., 28°09′05″ N, 17°15′13″ W, 11 May 2010, leg. S. Bräutigam and E. Bräutigam; SB T2 – Spain, Canary Islands, Tenerife: Puerto de la Cruz, above the village of Puerto de la Cruz, 7 km WSW of the town, disturbed places in the agricultural landscape, 780 m a.s.l., 28°22′31″ N, 16°36′37″ W, 3 May 2010, leg. S. Bräutigam and E. Bräutigam; SB H14 – Spain, Canary Islands, El Hierro: Sabinosa, near the Ermita Virgen de los Reyes, 3 km SW of the village, rocky places, nearby the Euphorbia-Kleinia shrubs, 720 m a.s.l., 27°43′51″ N, 18°07′16″ W, 21 May 2010, leg. S. Bräutigam and E. Bräutigam; SB H16p – Spain, Canary Islands, El Hierro: San Andrés, near the road to Frontera, 2 km WSW of the village, Chamaecytisus proliferus shrubs, 1,175 m a.s.l., 27°45′41″ N, 17°58′17″ W, 22 May 2010, leg. S. Bräutigam and E. Bräutigam.
A. pinnatifida subsp. teydensis
SB T4 – Spain, Canary Islands, Tenerife: Cañadas, at the road Arona – Vilaflor near La Escalona (km 13), 1,190 m a.s.l., 28°07′37″ N, 16°39′24″ W, 7 May 2010, leg. S. Bräutigam and E. Bräutigam; SB T18 – Spain, Canary Islands, Tenerife: Cañadas, near the road below El Portilla, rocks with Pinus canariensis forest, 1,950 m a.s.l., 28° 18′27″ N, 16°33′58″ W, 27 May 2010, leg. S. Bräutigam and E. Bräutigam; SB T17 – Spain, Canary Islands, Tenerife: Montañas de Anaga, 3 km E of Cruz del Carmen near Las Casas de Cumbre, road margin, “Fayal-Brezal” (Myrico fayae-Ericetum arboreae) vegetation, 800 m a.s.l., 28°32′02″ N, 16°15′12″ W, 26 May 2010, leg. S. Bräutigam and E. Bräutigam; SB T19 – Spain, Canary Islands, Tenerife: Cañadas, on the path 1.5 km E of Parador Nacional de la Cañadas, below Guajara, 2,200 m a.s.l., 28°13′32″ N, 16°36′46″ W, 28 May 2010, leg. S. Bräutigam and E. Bräutigam.
A. ragusina I
JC 2011_35 – Spain: Comunitat Valenciana: Sax, along the road to Catalla, 5.6 km NE of the town (city centre), 610 m a.s.l., 38°34′34″ N, 00°46′59″ W, 3 June 2011, leg. J. Chrtek; ZF LM5103 – Spain, province Madrid: Guadalix de la Sierra, 40°46′48″ N, 3°41′49″ W, 19 June 2010, leg. L. Medina and M. Sequeira.
A. ragusina II
JC 2011_2 – Spain, Andalusia, province Malaga: Alhaurin el Grande, near the old street to Alhaurin de la Torre, 5.7 km ESE of the town (near Golf Lauro), 230 m a.s.l., 36°38′52″ N, 04°37′17″ W, 24 May 2011, leg. J. Chrtek and Z. Dočkalová; JC 2011_27 – Spain, Andalusia, province Granada: Guadix, near the road to Jérez del Marquesado, 4.5 km of the town (centre), 1,075 m a.s.l., 37°15′25″ N, 03°08′16″ W, 31 May 2011, leg. J. Chrtek and Z. Dočkalová.
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Zahradníček, J., Chrtek, J., Ferreira, M.Z. et al. Genome size variation in the genus Andryala (Hieraciinae, Asteraceae). Folia Geobot 53, 429–447 (2018). https://doi.org/10.1007/s12224-018-9330-7
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DOI: https://doi.org/10.1007/s12224-018-9330-7