The genus Orobanche in Andalusia (southern Spain); taxonomical, chorological and ecological aspects A. Pujadas-Salvá 1' 2, E. Hernández-Bermejo

l,2

and J.A.R. Oliveira-Velloso l

Abstract In order to determine the Orobanche species living in Andalusia, as well as their geographical distribution, ecological characteristics and hosts, a survey was carried out, together with a revision of the main regional and national herbaria. Twenty-two different taxa were identified, but only bibliographical references can be given about seven other taxa that were not found. The main hosts that have been identified are also indicated. Continuation of botanical surveying and taxonomic research is required in order to complete the information on this genus. Introduction Although there are several floristic catalogues and regional floras dealing with the genus Orobanche (Valdés, 1986; Sagredo, 1987; Molero-Mesa and Pérez-Raya, 1987) found in Andalusia, an exhaustive catalogue has not been produced yet. Besides, the different works mentioned above are incomplete and in general not very satisfactory at the taxonomic level. We have to go back to Willkomm (1870) for a more reliable reference for the Iberian Peninsula, or to some other works for areas close to Andalusia such as Guimaraes (1903), besides the obligatory reference Beck von Mannagetta (1890, 1930). This encouraged us in 1985 to start cataloguing and analysing the Orobanche spp. of Andalusia. We consider that an accurate knowledge of our floristic resources is necessary, especially concerning groups with a remarkable economic interest. Some of these species are well-known as crop parasites (O. crenata, O. cernua, O. minor) and are easily identified. Our work aims to achieve an accurate taxonomic catalogue of all the Orobanche species of this region, because any of them could eventually evolve into a crop parasite in the future, as has already happened with O. foetida on faba bean (Vicia faba) and winter chickpea (Cicer arietinum) in North Africa (Kharrat et al., 1992). Orobanche is probably one of the most controversial genera as far as its specific characterisation is concerned. Many authors have expressed the difficulties that

1 2

Departamento de Ciencias y Recursos Agrícolas y Forestales, E.T.S.I.A.M., Apdo. 3048, Universidad de Córdoba, Córdoba, Spain Jardín Botánico de Córdoba, Avda. de Linneo s/n, Córdoba, Spain

Biology and management of Orobanche, Proceedings of the Third International Workshop on Orobanche and related Striga research. A.H. Pieterse, J.A.C. Verkleij and S.J. ter Borg (eds.). Amsterdam, The Netherlands, Royal Tropical Institute, 1994.

Figure

I

Andalusia geographical

location

appear when studying dried herbarium specimens, because significant differences, if any, are hardly appreciable (Chater and Webb, 1972; Musselman, 1986; OliveiraVelloso, 1990). Thus, the use of fresh material is essential for an accurate identification. To these problems we must also add that frequently when collecting, most of the species are taken for O. crenata, and due to this confusion are not collected. For this reason, in some of the herbaria, we have found fewer specimens than expected. We are aware that this is the first survey, but the important task of additional surveys and taxonomic research still has to be done. Material and methods Physiographical conditions Andalusia is located in the south of the Iberian Peninsula (Fig. 1) between the ' ' ' parallels 38°44 and 35°59 ' N and meridians 7°33 and 1°37 W. This region is an Autonomous Community whose management is divided into eight provinces: Córdoba, Jaén, Almería, Granada, Málaga, Cádiz, Sevilla and Huelva. It has an area of 87268 sq. km, that is, 17.3% of the total area of Spain. The orography is very polymorphous, and we can distinguish three large areas: Sierras Béticas (South Andalusian Mountains), Guadalquivir Depression and Sierra Morena Mountains. Due to its latitude, it has an extreme Mediterranean climate, although with a considerable Atlantic influence as regards rainfall: very hot summers and temperate winters (with average temperatures over 12°C at the seaside and below

The genus Orobanche in Andalusia 133

2°C on the Sierras Béticas in January, and over 28°C in the central area of the Guadalquivir Valley and below 20°C in some areas of the Sierras Béticas in July). The rainfall pattern presents a strong seasonal irregularity, ranging from average quantities below 200 mm in Almería to over 1500 mm in Grazalema. Moreover, there are high levels of insolation. Altogether, this Mediterranean climate on the border of the temperate zone presages the proximity of the North African subtropical desert band. The typical Mediterranean vegetation has been seriously damaged by man. In many areas, the original plant communities have been replaced by croplands. As a consequence, the forest formations (basically Quercus rotundifolia) and shrublands have taken refuge in less productive areas. Revision and floristic survey We have been carrying out since 1986 a taxonomic and chorological study of the Orobanchaceae in the Iberian Peninsula, concentrating on Andalusia. In this study, we have accomplished the following: 1. A critical revision of the bibliography on existing species and their chorology, based mainly on the works of Willkomm (1870), Beck von Mannagetta (1890, 1930), Valdés (1986), Sagredo (1987) and Molero-Mesa and Pérez-Raya (1987). 2. The revision of the existing herbarium material in the main regional and national herbaria: Córdoba (COA), Granada (GDA), Jaén (JAEN), Madrid (MA), Málaga (MGC) and Sevilla (SEV). 3. A floristic survey of the entire region in order to collect fresh material of Orobanche for easier and more accurate identification and the determination of their corresponding hosts. The samples from these collections are in the Herbarium (COA) of the Departamento de Ciencias y Recursos Agrícolas y Forestales, University of Córdoba, in the Córdoba Botanical Garden. For the identification and nomenclature of the species, we have mainly followed Beck von Mannagetta (1890, 1930), Tutin et al. (1964-1980) and Valdés et al. (1986). Results and discussion Floristic composition and chorology Twenty-two different taxa of Orobanche have been identified in the region. Their relation and geographical distribution are shown in Table 1. Whenever their existence was not verifiable with herbarium material, the first existing bibliographical reference is indicated. In Table 2, seven species are shown for which there are bibliographical references but no herbarium material. Some of them, such as O. coerulescens, O. purpurea and O. reticulata, are very unlikely to be found here because they have an Euroasiatic distribution and have not penetrated the Iberian Peninsula (Beck von Mannagetta, 1930). O. variegata has a Middle Europe distribution that extends to the Pyrenees, so its presence in our region is doubtful. As regards O. lavandulacea and O. trichocalyx, two very closely related species, and O. clausonis, it is quite possible that they will be found some day.

134 Pujadas-Salvá et al.

Table I.

Distribution of Orobanche genus AL

CA

CO

GR

JA

HU

MA

SE

Sect. Trionychon x

0 ramosa L. subsp. ramosa 0 ramoso L. subsp. mutelii (F.W. Schultz) Coutinho x

x

x

0 ramoso L. subsp. nana (Reuter) Coutinho

x

x

[S]

x

x

x

x

x [B1]

O. schultzii Mutel 0. arenarla Borkh

[ ]

x

x

x

S

x

x

x

x

x

x

Sect. Orobanche (=Osproleon) Subsect Coerulescentes O. cernua Loefl. var. cernua 0. cernua Loefl. var. cumana (Wallr.) G. Beck

x

x

x

x

x

x x

Subsect Galeatae O. latisquama (F.W. Schultz) Batt

[S]

x

x

x

x

x

x

x

[S]

[V]

x

x

x

x

x

O. foetida Poiret

x

x

x

O. sanguinea C. Presl

[B1]

x

[ ] x

[V]

x x

x

x

x

x

x

x

x

x

S

x

x

x

[v ]

x

x

x

S

x

x

x

x

x

x

x

x

O. caryophyllacea Sm. Subsect Arcuatae O. rapum-genistae Thuill .

B [ 2]

Subsect Cruentae O. gracilis Sm.

x

x

x

x

x

x

x

[

B2

]

Subsect Glandulosae O. alba Stephan ex Willd.

S

0. haenseleri Reuter

x [W ]

Subsect Speciosae O. crenata Forskál Subsect Minores O. amethystea Thuill. 0. loricataReichnb

[ ]

.

[ ] x

0. minor Sm.

x

x x

x

0. calendulae Pomel O. densiflora Salzm. ex Reuter

x

O. hederae DubY

x

AL = Almeria CA = Cádiz CO = Córdoba GR = Granada HU = Huelva JA = Jaén MA = Málaga SE =

X [ ] [B 1] [B2] V

[ ] [W [S]

I

x

x x

[

B2

]

x

x

= Herbarium reference = Just bibliographical references = Beck von Mannagetta (1890) = Beck von Mannagetta (1930) = Valdés (1986) = Willkomm (1870) = Sagredo (1987)

Sevilla

The genus Orobanche in Andalusia 135

Table 2.

Bibliographical references to species for which no herbarium material was found

Species

References

0. coerulescens Stephan

Molero-Mesa and Pérez-Raya (1987) Sagredo (1987)

0. clausonis Pomel

Beck von Mannagetta (1930) Sagredo (1987)

O. lavandulacea Reichenb.

Willkomm (1870)

0. purpurea Jacq.

Sagredo (1987) Willkomm (1870)

O. reticulata Wallr .

Sagredo (1987) Valdés (1986) Willkomm (1870)

0. trichocalyx (Webb & Berth)

G. Beck von Mannagetta (I 890)

O. variegata Wallr.

Sagredo (1987)

Taxonomy O. ramosa subsp. nana and O. ramosa subsp. mutelii have been separated as

different taxa in Table 1. However, we prefer to include them under the same taxon, because the characteristics that distinguish the subspecies (size of the corolla and stem branching) are not consistent; there is a continuity as regards the size of the corolla among the specimens analysed, and there are even specimens with very different sizes of corolla and with stems that may or may not be branched or do not coexist in the same population. According to Beck von Mannagetta (1930), we can distinguish two varieties of 0. cernua: var. cernua, with dense spikes and flowers generally smaller than 15 mm, parasitizes Artemisia species in our region, and var. cumana, with lax spikes and flowers generally larger than 15 mm, which parasitizes sunflower ( Helianthus annuus). In some cases, the taxonomic separation of the subsect. Minores is complicated, and a more detailed study on the relations among these species is required. Ecology There are many hosts for the different species of Orobanche. The hosts and their corresponding parasites are given in Table 3. The diversity of hosts parasitized by 0. crenata and O. minor is remarkable, in contrast to most of the remaining Orobanche, for which only one or two hosts have been identified. Some species present a particular ecology, as is the case with O. densiflora which frequently occurs in the sandy soils of the seashore, although it has also been found in the interior more than 100 km away. Other species live in mountain areas over 1500 m altitude such as O. sanguinea in humid meadows, O. haenseleri which has been collected over 1600 m and O. alba collected over 2500 m in the Sierra Nevada. O. gracilis can be found at very different altitudes, from the seashore to over 2500 m. Most of the species are found in the Mediterranean forest and shrubs. Some appear occasionally in ruderal ecosystems (taluses, ditches, roadsides): O. ramosa subsp. mutelii, O. cernua, O. latisquama , O. gracilis, O. foetida, O. crenata, O. amethystea,

136 Pujadas-Salvá et al.

O. minor. Only three species parasitize cultivated plants: O. crenata, O. ramosa subsp. ramosa and O. cernua var. cumana. Table 3.

Hosts of the different Orobanche species

Parasites

Hosts found

0. ramoso subsp. ramosa

Nicotiana tabacum (Oliveira-Velloso 1990)

0. ramosa subsp. mutelii

Trifolium campestre, Tropaeolum majus, Artemisia absinthium, Asteriscus maritimus, Coleostephus myconis

0. ramosa subsp. nano

Malva hispanica, Hedypnois cretica, Sonchus oleraceus

0. schultzii 0. arenaria

Artemisia glutinosa

0. cernua var. cernua

Artemisia barrelieri

0. cernua var. cumana

Helianthus annuus (Oliveira-Velloso, 1990)

0. latisquama

Rosmarinus officinalis

0. caryophyllacea 0. rapum-genistae 0. gracilis

Genista lobelii, Ulex parviflorus, Phlomis purpurea

0. foetida

Astragalus lusitanicus, Scorpiurus muricatus, Picris echioides

0. sanguinea

Trifolium repens, T. pratense

0. alba

Thymus serphyllioides

0. haenseleri

Helleborus foetidus

0. crenata

Leguminous crops, Carthamus tinctoreus, Daucus carota subsp. sativus and

0. amethystea

Foeniculum vulgare, Eryngium campestre, Anthyllis vulneraria

0. loricata

Eryngium campestre, Santolina elegans, Carduus tenuiflorus

0. minor

Trifolium angustifolium, T. glomeratus, T. stellatum, T. pratense, Vicio sativo,

other wild species (Oliveira-Velloso, 1990)

Daucus carota subsp. maxima, Eryngium campestre, Ballota hirsuta, Plantago lagopus, Tolpis barbata, Hypochoeris glabra, Picris echioides 0. calendulae

Eryngium campestre

0. densiflora

Lotus creticus

O. hederae

Hedera helix

References Beck von Mannagetta, G. (1890). Monographie der Gattung Orobanche. Bibliotheca Botanica 19: 1-275. Beck von Mannagetta, G. (1930). Orobanchaceae. In: A. Engler (ed.), Das Pflanzenreich 96 (IV.261). Germany, Wilhelm Engelmann, pp. 1-348. Chater, A.O. and D.A. Webb (1972). Orobanche. Flora Europaea 3: 286-293. Guimaraes, J.A. (1903). Monografia das Orobanchaceas. Broteria: 1-188. Kharrat, M., M.11. Halila , K.H. Linke and T. Iladdar (1992). First report of Orobanche foetida Poiret on faba bean in Tunisia. FABIS Newsletter 30: 46-47.

The genus Orobanche in Andalusia 137

Molero-Mesa, J. and F. Pérez-Raya (1987). La Flora de Sierra Nevada. Avance sobre el catálogo florístico nevadense. Granada, Spain, Universidad de Granada. Musselmann, L.J. (1986). Taxonomy of Orobanche. In: S.J. ter Borg (ed.), Proceedings of a Workshop on Biology and Control of Orobanche. Wageningen, The Netherlands, LH/VPO, pp. 2-10. Oliveira-Velloso, J.A.R. (1990). El género Orobanche en los cultivos en el sur de España, taxonomía, corología y ecología. (PhD Thesis, Universidad de Córdoba, Spain). Sagredo, R. (1987). Flora de Almería. Almería, Spain, Instituto de Estudios Almerienses. Tutin, T.G., V.H. Heywood, N.A. Burges, D.M. Moore, D.H. Valentine, S.M. Walters and D.A. Webb (eds.), (1964-1980). Flora Europaea, Vols. 1-5. Cambridge, UK, Cambridge University Press Valdés, B. (1986). Orobanchaceae. Flora Vascular de Andalucia Occidental 2: 550-558. Valdés, B., S. Talavera and E. Fernández-Galiano (eds.) (1986). Flora Vascular de Andalucía Occidental, Vols. 1-3. Barcelona, Spain, Ketrés. Willkomm, M. (1870). Orobanchaceae. Prodromus Florae Hispanicae 2: 620-632.

138 Pujadas-Salvá et al.