331 Amaryllis belladonna L. (Amaryllidaceae, Amaryllidoideae), first record as naturalized geophyte in Tunisia and continental North Africa Abstract Amaryllis belladonna L. is recorded for the first time as a naturalized non-native geophyte new to T unisian and continental North African flora. Additional information on its current distribution and habitat, a brief morphological description, as well as some taxonomic notes, are provided. Iz vleček Amaryllis belladonna L. je prvič zabeležena kot naturalizirani tujerodni geofit, nov za tunizijsko in celinsko severnoafriško floro. Predstavljene so dodatne informacije o trenutni distribuciji in habitatu, kratek morfološki opis, kot tudi nekaj taksonomskih zapiskov. Key words: New records, Aliens, Bulbs, Sejnane, North Africa. Ključne besede: novi zapisi, tujerodne vrste, čebulice, Sejnane, Severna Afrika. Received: 31. 3. 2020 Revision received: 29. 5. 2020 Accepted: 2. 6. 2020 1 University of Jendouba, Laboratory of Silvo-Pastoral Resources, Silvo-Pastoral Institute of Tabarka, BP. 345, Tabarka 8110, Tunisia. 2 University of Monastir, Laboratory of Botany, Cryptogamy and Plant Biology, Department of Pharmaceutical Sciences “A”, Faculty of Pharmacy of Monastir, Avenue Avicenna, 5000-Monastir, T unisia. Email: riridah@hotmail.com 3 University of Carthage, IRESA, Laboratory of Forest Ecology, INRGREF , BP . 10, Ariana 2080, T unisia. 4 Institute of Bioscience and BioResources (IBBR), Research Council (CNR), Unit of Palermo, Corso Calatafimi 414, 90129 Palermo, Italy. Ridha El Mokni1 , 2 , 3  , Salvatore Pasta4  & Davide Pacifico4  DOI: 10.2478/hacq-2020-0011 19/2 • 2020, 331–336 19/2 • 2020, 331–336 332 Ridha El Mokni, Salvatore Pasta & Davide Pacifico Amaryllis belladonna L. (Amaryllidaceae; Amaryllidoideae), first record as naturalized geophyte in T unisia and continental North Africa Introduction Extensive botanical surveys have been carried out by the first author between the years 2010 to 2020 and allowed to update and improve the botanical knowledge about Tunisian flora (e.g. El Mokni et al. 2010, El Mokni & El Aouni 2011, El Mokni et al. 2012, 2013a, 2013b, 2014, 2015a, 2015b, 2015c, El Mokni & Iamonico 2017 , 2018a, 2018b, 2019, El Mokni & Verloove 2019, El Mokni & Domina 2019, 2020, El Mokni 2018, 2020, Domina & El Mokni 2019). During one of these field surveys in 2016 a naturalized population of Amaryllis L., previously non recorded in the country, was found. Amaryllis L. is the only genus in the subtribe Amaryl- lidinae, tribe Amaryllideae (Brenzel 2001). It is a small genus of flowering bulbs, with only two species (A. bel- ladonna L. and A. paradisicola Snijman). In particular, A. belladonna L. is native to the Western Cape region of South Africa (Bond & Goldblatt 1984). This species occurs as a naturalized alien in several Mediterranean countries, where its degree of invasiveness is reported with “status unknown” (WCSP 2020). As for North Af- rica, the species has been previously reported only for the Canary Islands (Dobignard & Chatelain 2011, WCSP 2020, APD 2020). Our finding represents the first record for Tunisia and continental North Africa where it was found as naturalized in the Sejnane region (Northern Tunisia). This paper provides additional information on the naturalized population, as well as a review on the global distribution and ecology of the species. Moreover, a detailed morphological description of the plant, sup- ported by numerous detailed photographs, is provided. Material and methods The present work is based on field botanical surveys carried out by the first author (REM) in Northern Tunisia (North Africa), going with analysis of relevant literature and data about both known species of the genus (A. belladonna L. and A. paradisicola Snijman) available at https://www.wikizero.com/en/Amaryllis#cite_note- Biodiveristyexp-2 and https://www.wikizero.com/en/ Amaryllis_paradisicola and personal collection of the first author (REM) which is deposited in the Herbaria of the Faculty of Pharmacy of Monastir (not yet listed in Index Herbariorum, Thiers 2020). The description is in part original based on direct observations of Tunisian specimens. Research area The Sejnane area belongs to the Mogods Hills (Northern Tunisia), at the eastern boundary of the region Kabylia– Numidia– Kroumiria, recently shown to constitute a high hotspot of plant diversity within the Mediterranean basin (Véla & Benhouhou 2007). Its eroded hills are made of nummulitic sandstone and culminating around 400 m a.s.l. From a geologic point of view the area is mostly characterized by a succession of reddish sand- stone and silty clays (Crampon 1971). Each sandstone horizon is approximately 100 m thick and consists of fine to medium-grained sand with some lenses of coarse con- glomerate (250–300 mm in diameter). The interbedded clayey layers are generally greenish to greyish in colour and about 300 mm thick (Castany 1953, Rouvier 1977). The mean annual rainfall is comprised between 600 and 900 mm in the plains and reaches 1200 mm on the sur- rounding hills (Dimanche & Schoenenberger 1970). The woody vegetation of the Sejnane region is dominated by patches of degraded and discontinuous woodland domi- nated by cork oak woods (Quercus suber L.), intermin- gled with many other woody species typical to maquis and garrigue communities, such as Erica arborea L., Ar- butus unedo L., Pistacia lentiscus L., Myrtus communis L., Cistus salviifolius L., C. monspeliensis L., Lavandula stoe- chas L. Many species of Eucalyptus have been planted in the region, mostly for beekeeping purposes. Results and discussion A population counting more than one hundred indi- viduals was observed growing wild over an area of about 100 m 2 together with several thermo-xerophilous herbs and grasses forming the undergrowth of a plantation of Cupressus sempervirens L. Morphological description based on living plants (Figure 1): The plant has large brownish bulbs 5–10 cm in diameter (Figure 1-A) and bears long, distichous, bright green leaves of 30–50(–60) cm long and about 2–3 cm in width. The basal tuft of leaves can spread over 60 cm, when bulb is divided. Each bulb produces one or two leafless stems, red-brown (Figure 1-B), up to 75 cm high with ca. 2–12 crown flowers in its top. Flowers are trum- pet-shaped with pink-lilac or white with crimson-veined tepals, somehow recalling Lily flowers’ (Figure 1-D), gen- erally slightly bent towards the sun (Figure 1-C); corollas are about 6–10 cm long and 3–8 cm wide. Phenology: flowering starts in September whilst fruit- ing time ends at November. 19/2 • 2020, 331–336 333 Ridha El Mokni, Salvatore Pasta & Davide Pacifico Amaryllis belladonna L. (Amaryllidaceae; Amaryllidoideae), first record as naturalized geophyte in T unisia and continental North Africa Figure 1: A) Amaryllis belladonna L. in its habitat with flowering stems emerging from the bulbs of the extended population (Sejnane region, Northern T unisia), B) habit of the Belladonna Lily, C) detail of a pedicellate flower (lateral view), D) detail of flowers with pink-lilac tepals and medifixed anthers (top view). Photo credits: R. El Mokni (27.09.2016). Slika 1: A) Amaryllis belladonna L. v svojem habitatu s cvetočimi stebli, ki izhajajo iz čebulic v proučevani populaciji (regija Sejnane, Severna T unizija), B) habitus vrste Amaryllis belladonna, C) detajl cveta (stranski pogled), D) detajl cvetov z roza-lilastimi perionovimi listi in prašniki (pogled od zgoraj). Fotografija: R. El Mokni (27.09.2016). 19/2 • 2020, 331–336 334 Ridha El Mokni, Salvatore Pasta & Davide Pacifico Amaryllis belladonna L. (Amaryllidaceae; Amaryllidoideae), first record as naturalized geophyte in T unisia and continental North Africa Native habitat and distribution range (Figure 2): A. belladonna is endemic to the Western Cape region of South Africa, being particularly common on the rocky southwest area between the Olifants River Valley and Knysna (Bond & Goldblatt 1984). A. belladonna has been widely cultivated outside its natural habitat in several countries all over the world due to its ornamental value and responding to its climate needs (Johnson & Snijman 1996). In fact, the over-wintering foliage is not frost- tolerant; moreover, flowering in this species is strictly dependent on a dry resting period between leaf growth and flower production (Johnson & Snijman 1996). The species has become naturalized in many temperate countries, including the Mediterranean Basin, where it is reported for Portugal and Corse (WCSP 2020), Western and Southern Australia, California and New Zealand (Duncan 2004) where it is locally escaped from abandoned gardens and cut-flower cultivations (Duncan et al. 2016) and as casual to locally naturalized in Italy (cf. Galasso et al. 2018). Figure 2: Current distribution range of Amaryllis belladonna L. with new localities in T unisia, North Africa: A) global range, B) occurrence in Mediterranean countries (from http://plantsoftheworldonline.org/taxon/urn:lsid:ipni.org:names:62705-1, modified). Slika 2: T renutna razširjenost vrste Amaryllis belladonna L. z novimi nahajališči v T uniziji, Severna Afrika: A) globalna razširjenost, B) pojavljanje v Sredozemlju (od http://plantsoftheworldonline.org/taxon/urn:lsid:ipni.org:names:62705-1, spremenjeno). 19/2 • 2020, 331–336 335 Ridha El Mokni, Salvatore Pasta & Davide Pacifico Amaryllis belladonna L. (Amaryllidaceae; Amaryllidoideae), first record as naturalized geophyte in T unisia and continental North Africa Nomenclatural remarks Amaryllis belladonna L., Sp. Pl.: 293. 1753 ≡ Coburgia belladonna (L.) Herb., Bot. Mag. 47: t. 2113 (1819) ≡ Leopoldia belladonna (L.) M. Roem., Fam. Nat. Syn. Monogr. 4: 129 (1847) Since its creation in 1753, the genus Amaryllis L. has been object of several taxonomic revisions. From 1938 to 1984, controversy ensued over the lectotypification of the name A. belladonna, type of the genus Amaryllis L., and thus involving a lively debate on the correct application of the Linnean generic name. Goldblatt (1984) and Mee- row et al. (1997) reviewed the history of the controversy in detail. Despite prior general consensus that A. bella- donna applies to the species from South Africa known as the Cape Belladonna (Dandy & Fosberg 1954), Uphoff (1938) launched the contrary argument that the Linnean binomial must be applied to the neotropical south ameri- can Hippeastrum equestre Herb. [= H puniceum (Lam.) Kuntze]. This position was refuted by Sealy (1939). Up- hoff (1938, 1939) insisted that Amaryllis should stand for Hippeastrum, and his position was defended by T raub and his supporters (Traub & Moldenke 1949, Traub 1954, 1983, Tjaden 1979, 1981a, 1981b). Nomenclatural pro- posal supported by Jarvis (1984), and accepted at the 14 th International Botanical Congress held in 1987, that Amaryllis L. should be maintained, as a monotypic genus based on a specimen currently preserved in the Clifford Herbarium (the British Museum, Natural History). This specimen unquestionably represents the South African plant commonly known as the Cape Belladonna and agrees perfectly with Linnaeus definition of A. belladonna. Taxonomic notes: Besides Amaryllis belladonna L., the genus Amaryllis includes A. paradisicola Snijman, a spe- cies described from the succulent Karoo (Northern Cape) discovered in the Richtersveld National Park (Snijman & Williamson 1998). Although these two species have large, trumpet-shaped flowers, A. paradisicola generally bears more flowers per in- florescence than A. belladonna, and its flower stems spread in all directions. In A. belladonna the leaves are narrow, longer and hairless while being much wider and covered with hairs (especially when young) than in A. paradisicola. The color of the flowers varies in many shades from pink to dark pink or almost burgundy, rarely pure white, always with a creamy yellow throat, which are generally strongly oriented on one side in A. belladonna, whereas the color of flowers of A. paradisicola are light pale pink to dark pink, without a creamy yellow throat. Moreover, A. belladonna shows longer stamens and a more deeply divided trifid stig- ma (Snijman & Williamson 1998). In both species, the color of the flowers darkens with age and A. belladonna emits an overwhelming and sweet scent which intensifies at night, attracting the butterflies that visit them. The fra- grance of flowers is much less intense in A. paradisicola. Specimina visa: TUNISIA. Bizerta: Sejnane, Northern T unisia, coordinates 37°04’03.84’’ N, 09°07’40.94’’ E, co- occurring together with several thermos-xerophilous herbs and grasses in the undergrowth of an evergreen cypress plantation, at c. 120 m a.s.l., 27 September 2016, R. El Mokni s.n. (Herb. El Mokni!), R. El Mokni s.n. (Herb. Univ. Monastir). Some bulbs were brought and cultivated at the Faculty of Pharmacy of Monastir (Univ. Monastir). Acknowledgements Thanks are due to all colleagues and friends of forestry service of Bizerta (Sejnane delegation) for their support and their help during field prospections. Ridha El Mokni , https://orcid.org/0000-0003-3849-1039 Salvatore Pasta , https://orcid.org/0000-0003-3265-9072 Davide Pacifico , https://orcid.org/0000-0003-3620-9829 References APD. 2020: African Plant Database (version 3.4.0). Amaryllis belladonna L. – Conservatoire et Jardin botaniques; South African National Biodiversity Institute, Pretoria Genève, viewed on 30 March 2020, from http://www.ville-ge.ch/musinfo/bd/cjb/africa/details. php?langue=an&id=15978 Bond, P . & Goldblatt, P . 1984: Plants of the Cape Flora. Journal of South African Botany, 13 (Suppl.), pp. 1–455. Brenzel, K. N. (ed.) 2001: Sunset western garden book. 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