Additional of (/Agavaceae, sensu lat.) introduced and naturalising in Tunisia and North Africa

Authors: Mokni, Ridha El, and Verloove, Filip Source: Bradleya, 2021(39) : 221-235 Published By: British Cactus and Succulent Society URL: https://doi.org/10.25223/brad.n39.2021.a23

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Additional species of Agave (Agavoideae /Agavaceae, Asparagaceae sensu lat.) introduced and naturalising in Tunisia and North Africa

Ridha El Mokni1,2,3 and Filip Verloove4 1. University of Monastir, Laboratory of Botany, Cryptogamy and Biology, Faculty of Pharmacy of Monastir, Avenue Avicenna, 5000-Monastir, Tunisia. (email: [email protected]) 2.University of Jendouba, Laboratory of Silvo-Pastoral Resources, Silvo-Pastoral Institute of Tabarka, B P. 345, 8110-Tabarka, Tunisia. 3. University of Carthage, Laboratory of Forest Ecology, National Research Institute of Rural Engi- neering, Water and Forests, Ariana 2080, Tunisia. 4. Meise Botanic Garden, Nieuwelaan 38, B-1860 Meise, Belgium. (email: filip.verloove@botanicgar- denmeise.be)

Summary: Recent fieldwork in Tunisia yielded Introduction several new records of non-native species of the Non-native , known also as exotic, intro- Agave. var. marginata, A. duced, alien, or non-indigenous, are those taxa fourcroydes and A. sisalana are locally naturalised whose presence in a given area is due to inten- and reported for the first time from the country. To tional or unintentional human involvement, or our knowledge, naturalised populations of these which have arrived there without the help of peo- three taxa were also unknown elsewhere in the ple from an area in which they are native (Pyšek et Maghreb and North Africa. We also draw attention al., 2004). Such species are widely recognised as to A. attenuata subsp. attenuata, a widely grown a significant component of human-caused global ornamental that is expected to become naturalised environmental change, resulting often in a sig- in the near future. Updated nomenclature, brief nificant loss in biological diversity and function descriptions, as well as general and national dis- of invaded communities and ecosystems (Mooney tributions are provided for each of the naturalised & Hobbs, 2000; Mack et al., 2000; Pimentel et al., . A key to all the species, subspecies, and 2001). varieties of Agave that occur in Tunisia is present- In North Africa, published information re- ed, and all the taxa are illustrated. garding the naturalisation and expansion of alien plants has a substantial history. About 340 alien Zusammenfassung: Jüngste Feldforschungen in species have been recorded so Tunesien ergaben mehrere interessante neue far (i.e. c. 5% of the flora) (cf. Vilà et al., 1999). Nachweise nicht-heimischer Arten der Gattung From the viewpoint of the geographic origin of Agave. A. angustifolia var. marginata, A. fourcroy- the alien invading species, almost 50% come from des und A. sisalana sind lokal eingebürgert und the New World. Most of the expanding species werden erstmals für dieses Land dokumentiert. are ornamentals, originally deliberately intro- Nach unserem Kenntnisstand waren eingebürgerte duced by humans (Le Floc’h et al., 1990; Vilà et Populationen dieser drei Taxa auch anderswo al., 1999). In the past two decades, the knowledge im Maghreb und in Nordafrika unbekannt. Wir of the non-native flora of Tunisia and North Af- machen auch auf A. attenuata subsp. attenuata rica significantly increased (see e.g. El Mokni & El aufmerksam, eine mehr oder weniger weit verbre- Aouni, 2011a,b, 2012, 2013; El Mokni & Domina, itete Zierpflanze, von der zu erwarten ist, dass sie 2018, 2020; El Mokni & Verloove, 2019a, 2019b; in naher Zukunft eingebürgert sein wird. Für jedes El Mokni et al., 2012, 2016; El Mokni & Iamonico, Taxon der eingebürgerten Agaven werden die ak- 2018a, 2019; El Mokni, 2018, 2019; Iamonico & El tualisierte Nomenklatur, kurze Beschreibungen Mokni, 2016). However, information about alien sowie die allgemeine und nationale Verbreitung succulent plants is mostly recent and still scarce präsentiert. Ein Bestimmungsschlüssel zu allen in (see e.g. El Mokni & Iamonico, 2018a; El Mokni & Tunesien vorkommenden Arten, Unterarten und Sáez, 2019; El Mokni et al., 2018, 2019a, 2019b, Sorten der Agave wird vorgestellt und alle Taxa 2020). North Africa includes Morocco, the North- werden abgebildet. ern part of Mauritania, the Northern part of Mali,

Bradleya 39/2021 221 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Tibesti Mountains of Chad, Algeria, Tunisia, Lib- In this paper all taxa (subgenera, sections, species, ya and Egypt with Canary and Madeira Islands, and infraspecific taxa) are arranged alphabetical- whereas Maghrebian countries are restricted to ly. Nomenclature of the taxa presented is mostly Mauritania, Morocco, Algeria, Tunisia and Libya in accordance with recent phylogenetic classifica- (cf. Dobignard & Chatelain, 2010). tions of the family Agavaceae (see e.g. Thiede et Agave is the largest genus in the family Aga- al., 2019; Verloove et al., 2019). Newly reported vaceae. As presently circumscribed, this family taxa are indicated in bold in the key. is entirely confined to New World temperate and tropical areas, with eight genera and more than Results 276 named taxa (see e.g. Verhoek, 1998; Smith & Agavaceae Dumort., Anal. Fam. Pl. 57, 58. 1829. Figueiredo, 2014). The genus, comprising species nom. cons. Cronquist: 1217 (1981). of rosulate and succulent-leaved perennials, is na- Agave L., Sp. Pl. 1: 323. 1753. tive to the southern parts of the United States of Plants perennial and monocarpic. Stems very America, , Central America and the north- short or indistinct. Leaves in a basal rosette, large, ern parts of South America (Gentry, 1982; Mab- stout, leathery-fleshy or somewhat woody, con- berley, 1997). A number of species has, since the taining many fibers, margin usually spiny, rarely early 1860s, been introduced in Africa and even- entire, apex tipped with a spine. Flowering stems tually became established, among which two are branched or simple, tall, stout. Inflorescence ter- widely distributed: A. americana L. var. ameri- minal, a spike or panicle, mostly very large. Peri- cana and A. sisalana Perrine (Smith & Mössmer, anth tube short; lobes narrow, subequal. Stamens 1996). Both are currently included in catalogues inserted at throat or in tube of perianth; filament of problem plants in southern Africa (Wells et al., filiform, usually longer than perianth; anther ver- 1986; Henderson, 1995). satile. Ovary with many ovules. Style slender; Agave has been traditionally divided into two stigma 3-lobed. Fruit a capsule, oblong, 3-valved, subgenera (Berger, 1921; Gentry, 1982), defined on loculicidal. Seeds numerous, black, thin, flat- the basis of the inflorescence type: A. subg. Littaea tened. (Tagl.) Baker, which has an unbranched spike or Within Agavaceae subfam. Agavoideae five racemose inflorescence and contains about fifty- species and two varieties are here reported from three species with a more restricted distribution, Tunisia. For three of them, Agave angustifolia Haw. primarily in Mexico, and A. subg. Agave, whose var. marginata Hort. ex Gentry and A. fourcroydes species possess branched (paniculate) inflores- Lem., naturalised populations are reported for cences, i.e. large umbelliferous aggregates of flow- the first time in the Maghreb and North Africa. ers (Gentry, 1982), and includes c. 102 species. Salm-Dyck subsp. attenuata (the The present contribution aims at improving only taxon in A. subg. Littaea) is also included. the gaps in the knowledge on taxa of the alien ge- Its escape from cultivation is expected but not yet nus Agave L. that are either new to the Tunisian or observed in Tunisia. An updated key to all species Maghrebian allochthonous flora in North Africa. and varieties of Agave occurring in Tunisia is pro- vided. Material and Methods Floristic field investigations in Central and North- ern Tunisia (North Africa), mostly between 2010 Agave subg. Agave A. sect. Agave and 2020, have revealed new populations of Agave species. Some taxa, observed frequently in L. Sp. Pl.: 323. 1753. small colonies, were not previously reported from = Aloe americana (L.) Crantz, Inst. Rei Herb. 1: Tunisia or North Africa. Records here reported are 466. 1766. documented by a brief description of each taxon Agave americana is the type species of the genus and infraspecific taxa. The actual status (casual/ Agave. The species is divided into two subspecies, naturalised) for each taxon is assessed based on one of which has four varieties, namely Agave our field observations. For the identification nu- americana subsp. americana var. americana, A. merous sources were consulted, for instance americana subsp. americana var. expansa (Ja- Smith & Mössmer (1996), Smith & Steyn (1999a), cobi) Gentry, A. americana subsp. americana var. Thiede (2001), Smith (2011), Franck (2012), oaxacensis Gentry, A. americana subsp. ameri- Verloove et al. (2019), Flora of North America cana var. marginata Trel. and A. americana subsp. (2020) (retrieved from http://www.efloras.org/flo- protamericana Gentry. Of these, only A. ameri- rataxon.aspx?flora_id=1&taxon_id=10019), and cana subsp. americana var. americana is known Flora of China (2020) (retrieved from http://www. as widely naturalised in Tunisia and North Africa efloras.org/florataxon.aspx?flora_id=2&taxon_ (APD, 2020). id=100796). Information on flowering/fruiting We here report the presence of two varieties period, distribution, habitats occupied and field of A. americana subsp. americana in the wild in photographs are also presented. Tunisia. Both are commonly cultivated and have

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Figure 1. Agave americana var. americana on a sun-exposed clayey slope in Monastir (C. E. Tunisia, North Africa, 11 July and 5 October 2019.

spread locally in many regions. For example, cana var. marginata Trel., which has reflexed, var- individuals of this species were formerly mas- iegated leaves and has been observed to produce sively planted all along the Sousse-Tunis highway capsules (Figure 2.). for erosion control and soon started escaping. The first variety is A. americana var. americana, Agave americana L., Sp. Pl. 323. 1753 var. mar- known by its reflexed, glaucous leaves (Figure ginata Trel., in L. H. Bailey, Stand. Cycl. Hort. 1: 1) which is more frequent and widely present 235. 1914. (Figure 2). TUNISIA: Monastir, scat- throughout Tunisia. The other variety is A. ameri- tered individuals growing on a sun-exposed clayey

Bradleya 39/2021 223 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Figure 2. Agave americana var. marginata on a sun-exposed clayey slope in Monastir (C. E. Tunisia, North Africa), 30 May 2019.

slope in the Monastir coastal area, 35°46’50”N, similar in size to the typical variety, with a yellow 10°49’20”E, 10 m a.s.l., 30.05.2019, R. El Mokni sub-marginal band in the center of the leaf, some- s.c. times with yellow stripes; lateral bands 15mm Morphology: (Figure 2.) For a detailed descrip- wide, obovate. Apical spine black, 27-30 mm long tion, see Thiede (2001). Robust plant, with flat- H 2–3mm wide, ribbed; marginal spines 8–9 mm tened bluish-green to glaucous leaves, smaller or H 9–10mm wide, curved, with the apex facing the

224 Bradleya 39/2021 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Bradleya 39/2021 Inflorescence paniculate, flowers in umbellate clusters on lateral branches ...... A. subg.pages Agave 221–235

Inflorescence spicate or racemose………...... …… A. subg. Littaea

Agave subg. Agave

1 Leaves narrow and sword-shaped (ensiform), c. 10–20 as long as wide with almost straight mar- gins (hardly narrowed towards the base), very rigid and radiately spreading. Apical leaf spine not or hardly decurrent. Inflorescence nearly always bulbiliferous ……………..……….…...... 2

1 Leaves 100–200 H 15–25cm, markedly narrowed towards the base, never ensiform and with straight margins, pliable and often drooping to one side. Apical leaf spine strongly decurrent. Inflorescence never bulbiliferous……….………...…...... …………...... …. A. americana

2 Leaves 40–60(–100)cm long, light green to glaucous grey, often with white or yellow margins (var. marginata). Stem absent or short. Inflorescence 3–5m long ...... A. angustifolia

2 Leaves ensiform, fleshy, 90–130 H 9–12cm, never with white or yellow margins. Stem present, short or distinct. Inflorescence generally longer, up to 6m…………...... …...... ……….. 3

3 Leaf margins unarmed (exceptionally with well-developed triangular teeth but then not or only very weakly glaucous and with smaller flowers). Plant usually green, either yellowish or dark green. Flowers relatively small, 55–65mm long...... A. sisalana

3 Leaf margins with regularly spaced and distinct teeth. Plant often glaucous or greyish green. Flowers longer, 60–70(–80)mm long…...... A. fourcroydes

Key to Agave subgenera and species occurring in Tunisia.

base and the leaf apex, black (cf. Guillot-Ortiz et al., nisia (Dobignard & Chatelain, 2010; APD, 2020). 2009). In Tunisia, Agave americana var. marginata was Flowering period: July to September(-Octo- hitherto considered to be cultivated as an orna- ber). mental only; however, it can clearly be considered Agave americana sensu lat. is native to Mexico as naturalised locally. In fact, it was observed for and some parts of the southern United States of many years in several subpopulations in the cen- America. It is widely cultivated and has become tral and northern parts of the country. Like the naturalised in many parts of South Africa and other known variety, it certainly has the potential other temperate, sub-tropical and semi-arid re- to spread and become an invasive alien. gions of the world. The taxon is well established in the flora of North Africa, amongst others, in Tu- Agave subg. Agave A. sect. Rigidae (Baker) R.H.Webb & G.D.Starr, Cact. Succ. J. (US) 87(2): 85. 2015. Leaves glaucous, usually drooping to one side, thickly succulent; marginal teeth vari- Agave angustifolia Haw. Syn. Pl. Succ. 1: 72. able, generally hard and big; plants generally 1812 var. marginata Hort. ex Gentry, Agaves Con- massive ………....…………...var. americana tinental N America 564. 1982. (Figure 3). First record for the non-native flora of Tuni- Leaves variegated, almost drooping to one sia and the Maghreb in North Africa. About six side, less succulent and less large; marginal individuals surviving in the ‘wild’ on the margin teeth generally smaller; plants sometimes of a coastal pine forest. The species was prob- medium sized...... var. marginata ably planted for ornament in the area (or origi- nated from a nearby abandoned plant nursery?) Key to distinguish between the two varieties of Agave and now seems to be escaping, along with many americana subsp. americana occurring in the wild in young specimens of A. americana var. americana. Tunisia. Also, five individuals surviving on the margin

Bradleya 39/2021 225 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Figure 3. Agave angustifolia var. marginata on a sun-exposed field within the margin of a coastal pine forest in Biz- erta (N. E. Tunisia, North Africa), 12 July 2019.

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Figure 4. on a sun-exposed sandy field within the margin of a coastal pine forest with cultivated Eucalyptus spp. (N. E. Tunisia, North Africa), 7 December 2018.

Bradleya 39/2021 227 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Figure 5. Agave sisalana on a sun-exposed slope in Monastir, 30 May 2018 and Sousse, 4 June 2019 (C. E. Tunisia, North Africa).

228 Bradleya 39/2021 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise of a private garden in Monastir, on the left side linear, rigid, thickly roundedBradleya at the base, 39/2021 acumi- of the road towards Marina. TUNISIA: Bizerta, nate, guttered, 120–180cm long × 8–12cmpages 221–235 wide, scattered individuals growing on the margin of tipped by black spine; margins straight; margins a coastal pine forest on sandy field, 37°18’29”N, with small reddish-brown spines (3–6mm), regu- 09°51’51”E, 26 m a.s.l., 12.07.2019, R. El Mokni larly spaced; terminal spine conical, stout, openly s.c.; Monastir, five individuals surviving on the short-grooved above, mostly 2–3cm long, dark margin of a private garden at the left side of the brown; Inflorescence 5–6 m, bulbiliferous; Flow- road towards Marina, 35°46’27”N, 10°49’52”E, 20 ers 60–70 mm long; Tepals greenish-yellow, tube m a.s.l., 07.02.2020, R. El Mokni s.c. urceolate, 12–16 mm long, lobes subequal, 16– Morphology: (Figure 3). Rosette dense, with 18mm; Ovary fusiform, roundly trigonous, 35–40 stiffly erect, short, sword-shaped leaves. Blades mm, neck briefly constricted (see Thiede, 2001). greyish-green in colour with prominent white bor- Flowering period: June to September (–October). der and little red spines. Apex ending in a sharp Native to Mexico where its origin was attribut- brown spine (see Thiede, 2001 under A. vivipara ed to A. angustifolia (Gentry, 1982), the only wild ‘Variegata’). species of Agave growing in Yucatan Peninsula. Flowering period: Not seen in flower yet in Agave fourcroydes is widely cultivated as an orna- Tunisia. mental in tropical and temperate areas and pro- Agave angustifolia, a native of Central America, duces a large number of bulbils on the inflores- was considered to be a variable species with seven cence (Figure 4). Thus, the species easily estab- varieties (Gentry, 1982). It is known to be the most lishes itself wherever introduced and blooming. widespread species of the genus and ranges from In the Euro+Med Plantbase, Agave fourcroydes northern Mexico to Panama. The species is com- was only reported from the Canary Islands where monly cultivated as an ornamental and very easily it is naturalised and invasive (Acebes Ginovés escapes. Thus, it was seen naturalised in Florida et al., 2010; Santos-Guerra et al., 2013; Verloove (Franck, 2012), India (Drummond & Prain, 1906) et al., 2019). However, the species has also been and western Australia (Forster, 1996). In Europe it recorded from other countries, especially in the has been recorded in the Valencia area and in the Mediterranean area, for instance from Spain and Balearic Islands in Spain (Moragues & Rita, 2005; Italy (see e.g. Bacchetta et al. 2009; Guillot et al. Guillot Ortiz et al., 2009; Sáez et al., 2016) as well 2009). In North Africa, the species was reported as in Italy (Manni, 2015). In Africa, it has natu- from Morocco as cultivated (APD, 2020). It is here ralised in South Africa (Steyn & Smith, 2000; Wal- first recorded as naturalised only from Tunisia for ters et al., 2011) and the Canary Islands (Verloove the whole Maghreb in North Africa (APD, 2020). et al., 2019). The plant is here reported for the first time as naturalising in Tunisia and the Maghreb in Agave sisalana Perrine. Trop. Pl. [U.S. House Rep- North Africa (APD, 2020, under A. vivipara L. var. resentatives Rep. no. 564] 87 (1838); ex Englem., vivipara). Trans. Acad. Sci. St. Louis 3: 314, t. 2-4 (1875). For quite some time, this species was consid- (Figure 5). ered to be conspecific with A. vivipara (see e.g. = A. rigida var. sisalana (Perrine) Engelm. Trans. Wijnands, 1983; Forster, 1992; Smith & Steyn, Acad. Sci. St. Louis 3: 316, t. 2-4 (1875) 1999; Thiede, 2001). However, García-Mendoza & First record as naturalised for the non-native Chiang (2003) showed that these two species are flora of Tunisia and for the entire Maghreb in very distinct (see also Verloove et al., 2019). North Africa. More than 100 individuals surviv- ing in the wild in four distant populations in the Agave fourcroydes Lem. (III. Hort. 11 (Misc.): 65. central and northwestern parts of the country (Mo- 1864. (Figure 4). nastir, Sousse, and Jendouba). The species was in- = A. rigida var. elongata Baker, Bull. Misc. Inform. troduced as an agricultural crop (for the extraction Kew 1892: 23. 1893 of leaf fibers). Since it produces a lot of bulbils, First record for the non-native flora of Tunisia it was able to establish self-sustaining popula- and the Maghreb in North Africa, as naturalised. tions. TUNISIA: Monastir, coastal road towards More than thirty individuals surviving in the wild Sousse, the most extended population with scat- on the margin of a coastal pine forest with culti- tered individuals (78–80 in an area of about 16ha) vated Eucalyptus spp. The species escaped from a growing on a coastal, sun-exposed clayey slope, nearby plant nursery. TUNISIA: Bizerta, Menzel- 35°46’49”N, 10°49’26”E, 10 m a.s.l., 19.12.2018, Jemil-laâzib, one population with scattered indi- R. El Mokni s.c.; Ksar-Hellal, a small population viduals growing on the margin of a coastal pine with few individuals (10–12 in an area of about forest with cultivated Eucalyptus, 37°12’42”N, 100 m sq.) with hundreds of bulbils growing in an 09°58’24”E, 34 m a.s.l., 19.06.2018, R. El Mokni olive plantation, 35°37’49”N, 10°51’33”E, 5m a.s.l., s.c.; ibidem, 07.12.2019, R. El Mokni s.c. 11.04.2018, R. El Mokni s.c.; Sousse, road towards Morphology: (Figure 4). Stem thick, 1–1.7(–2.5) Sahloul, scattered individuals (about twenty in m long; Rosettes large, suckering; Leaves straight, an area of about 10ha) in an abandoned field near

Bradleya 39/2021 229 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Figure 6. Agave attenuata subsp. attenuata on the margin of a private garden within Ariana region (N. E. Tunisia, North Africa), 6 January 2020.

the ENISo School, 35°49’27”N, 10°35’30”E, 36 m bases. Leaves 90–160cm long, c. 8–10(–15)cm a.s.l., 04.06.2019, R. El Mokni s.c.; Jendouba, Ta- wide, widest about the middle, erect, neck not barka, coastal road in the touristic area, scattered sharply constricted, greyish green, often glabrous, individuals (more than 30 in an area about 1ha) in channeled to about ⅔ its length, the basal ⅓ flat dune vegetation of spontaneous Quercus coccifera above, margin not or rarely spiny, apex straight communities, 36°57’22”N, 08°48’01”E, 10m a.s.l., and tipped with a red brown spine 2–3cm. Inflo- 07.01.2019, R. El Mokni s.c. rescence a large panicle up to 6m, stout, usually Morphology: (Figure 5). A sterile pentaploid bearing bulblets after anthesis. Flowers with a (Doughty, 1936). Stems short, covered by leaf strong smell arranged on the terminal portion in

230 Bradleya 39/2021 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise dense clusters, sessile, 4–5 cm long. Perianth seg- Agave subg. Littaea (Tagl.)Bradleya Baker, Handb. 39/2021 Ama- ments spreading, equaling the ovary segment. Sta- ryll. 164. 1888. (Figure 6). pages 221–235 mens longer than the perianth segment, 2.5–3(–4) Inflorescence often dense, elongated panicles cm long. Style exserted; stigma 3-lobed, slightly with ± short-stalked partial inflorescences mostly capitate. Capsule (not seen in North Africa) ob- with paired or few clustered flowers (‘spicate’). long, c. 6cm long × 2–2.5cm wide (cf. Flora of China, 2000; Thiede, 2001). Agave sect. Inermes Salm-Dyck, Wochenschr. Flowering period: May to August (–September). Vereines Beförd. Gartenbaues Königl. Preuss. Agave sisalana can be distinguished from A. Staaten 4(23): 182. [6 June] 1861. fourcroydes mainly by its greener leaves and un- Agave sect. Inermes includes species with soft, armed adult leaf margins (versus regularly toothed unarmed, non-filiferous leaves and inflorescences leaf margins throughout). The species is known with densely arranged flowers. A few species have as a cultivated plant only. It was likely first do- leathery, unarmed leaves and all are endemic to mesticated in the Yucatan Peninsula in Mexico Mexico, occurring mainly in the Sierra Madre Oc- (Trejo-Torres et al., 2018). The species is widely cidental. cultivated as an ornamental, and for its medicinal value. It has the ability to grow in a wide vari- Agave attenuata Salm-Dyck subsp. attenuata, ety of habitats and often escapes from plantations Hortus Dyckensis: 303, 1834. into adjacent natural areas. It was cultivated as a = Ghiesbreghtia mollis Roezl, Gartenflora 10(4): fiber plant and reported naturalised to invasive in 122, IV.1861. North and South America, Central Africa (Ethio- = Agave glaucescens Hooker, Curtis’s Bot. Mag. pia, Kenya, Malawi, and Tanzania; CABI, 2020), 88: pl. 5333, 1.I.1862. (see more Ullrich, 2006) South Africa, Europe, East Asia, and eastern Aus- A. attenuata is more or less widely grown as tralia (Hochstätter, 2015; Nobel, 1988). In Austra- an ornamental, but probably less so in North Af- lia, Agave sisalana is listed as a priority environ- rica. Three individuals with only rosettes of large mental weed in the state of Queensland where it is leaves were observed in a private garden in cen- ranked among the top 200 most invasive plant spe- tral Tunisia, whereas five individuals were seen cies. It is also listed as one of the thirty-five most in a private garden in Ariana towards INSAT insti- troublesome weed species occurring on sandy tute, among them one with the typical spike-like beaches and dunes along Queensland’s east coast inflorescence (Figure 6). (Australian Weeds Committee, 2013). This species Morphology: (Figure 6). A perennial succulent occurs in disturbed habitats such as abandoned plant. After many years, it develops one or more fields, roadsides, and waste areas. It produces a upright, thick trunks up to 1.5m tall, at the tip of large number of bulbils on the flowering stem which a leaf rosette spreads. The numerous, egg- (pseudovivipary) (Elmqvist & Cox, 1996), so it eas- shaped and pointed leaves are somewhat softer ily spreads to the surroundings of the areas where than most agave species, they are light glaucous it was introduced. In addition the species is able gray or pale yellowish-green and spiky. They are to develop dense monospecific stands which may 50–70cm long and 12–16cm wide. The leaf mar- prevent the establishment of native vegetation gin is smooth or (very rarely) finely serrulate. Af- (Badano & Pugnaire, 2004). In the Mediterranean ter many years, the often-curved inflorescence area, it was reported as naturalised to invasive in develops, which grows up to 3.5m high and bears the Balearic Islands in Spain (Badano & Pugnaire, dense, greenish-yellow flowers. Flowers up to 50 2004; Moragues & Rita, 2005), in Albania (Domina mm long; tepals 16–24 long × 5 mm wide; ovary et al., 2018), in peninsular Italy, Sardinia, Sicily, 15–25mm long; filaments 35–45 mm long; an- Morocco, Algeria, and Libya (APD, 2020; CABI, thers 15–20 mm in length. Capsule 20–30 mm × 2020; DAISIE, 2020; Euro+Med Plantbase, 2020). 10 mm. Seeds c. 3–3.5mm long × 2–2.5mm wide It is here reported for the first time as an estab- (see Thiede, 2001 under Agave chamelensis; Ull- lished (naturalised) plant in Tunisia. rich, 2006). Flowering period: December to February (– March). Agave subg. Littaea Agave attenuata subsp. attenuata is native to central Mexico and tropical America. Unlike most Only one species of A. subg. Littaea has been re- species of Agave, it has a typical curved flower corded as a cultivated ornamental in Tunisia. spike. The plant is widely cultivated in warm- Leaves always with smooth to (very rarely) temperate regions of the world where it is com- serrulate margins and soft apical spine (subsp. monly grown as a garden plant and increasingly attenuata). Inflorescence spicate, very densely reported as escaping, also in natural habitats. It flowered and arching, not bulbiliferous; tepals is known, for instance, from the Kruger National greenish-yellow. Stem always present at maturity, Park in South Africa (Foxcroft et al., 2008), Italy, up to 100cm long, A. attenuata including Sardinia and Sicily (Podda et al., 2012;

Bradleya 39/2021 231 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Galasso et al., 2018) and many parts of Australia Cronquist, A. (1981) An Integrated System of Clas- (Randall, 2017). In Macaronesia it is also known sification of Flowering Plants. New York, Co- from Madeira, Fuerteventura, Gran Canaria, La lumbia University Press. 1262 pp. Palma and Tenerife (see e.g. Brandes & Fritzsch, DAISIE (2020) Delivering Alien Invasive Species 2002; Vieira, 2002; Borges et al., 2008; Verloove, Inventories for Europe. http://www.europeali- 2013; Otto & Verloove, 2016; Verloove et al., 2019) ens.org. Accessed 20 April 2020. where it is locally naturalised. In North Africa, the Dobignard, A. & Chatelain, C. (2010) Index syno- species is reported only from Libya as alien with nymique de la flore d’Afrique du Nord. Volume ‘unknown status’ (Euro+Med Plantbase, 2020). 1: Pteridophyta, Gymnospermae, Monocotyle- The plant was also reported more recently from doneae. Genève, Conservatoire et jardin bota- New Zealand (Thomson, 2020). niques, 455 pp. Although so far only observed as a cultivated Domina, G., Meço, M. & Mullaj, A. (2018) Agave si- ornamental in Tunisia, we here report this species salana Perrine (Asparagaceae). p 9. - In: Nobis, because it is likely to escape in the near future. M., Domina, G., Meço, M., Mullaj, A., Bazan, G., Ebel, A.L., Király, G., Erst, A., Nowak, A., References Sukhorukov, A.P., Pospelova E.B., Pospelov, I.N., Acebes Ginovés, J.R., León Arencibia, M.C., Ro- Vasjukov, V.M., Piwowarczyk, R., Seregin, A.P., dríguez Navarro, M.L., del Arco Aguilar, M., Király, A., Kushunina, M., Liu B., Molnár, A.V., García Gallo A., Pérez de Paz, P.L., Rodríguez Olonova, M., Óvári, M., Paszko, B., You-Sheng, Delgado, O., Martín Osorio, V.E., Wildpret de la Ch., Verkhozina, A.V., Zykova, E.Y., Klichowska, Torre, W. (2010) Pteridophyta, Spermatophy- E., Nobis, A., Wróbel, A., Aydın, Z.U., Dönmez, ta. In: Arechavaleta M., Rodríguez S., Zurita N., A.A., Garakhani, P., Koopman, J., Korolyuk, García A. (eds.) Lista de especies silvestres de A., Oklejewicz, K., Qasimova, T., Wang, W., Canarias (hongos, plantas y animales terrestres Więcław, H., Wolanin, M. & Xiang, K. (2018) (second edition). 119–172. La Laguna, Gobier- Contribution to the flora of Asian and Europe- no de Canarias. an countries: new national and regional vascu- APD (African Plant Database) (2020) Agave L. – In: lar plant records, 7, Botany Letters. https://doi. African Plant Database. – Published at http:// org/10.1080/23818107.2017.1415817. www.ville-ge.ch/musinfo/bd/cjb/africa/details. Doughty, L.R. (1936) Chromosome behaviour in php?langue=an&id=187877. Accessed 13 relation to genetics of Agave. I. Seven species April 2020. of fiber Agave. Journal of Genetics 33:197–205. Australian Weeds Committee (2013) Weeds of Aus- Drummond, J.R. & Prain, D. (1906) Notes on Agave tralia. Canberra, Australia: Australian Weeds and in India. Agricultural Ledger 7: Committee. http://www.weeds.org.au/. 127–321. Bacchetta, G., Mayoral, O., Podda, L. (2009) Dumortier, B.C.J. (1829) Agavineae. Analyse des Catálogo de la flora exótica de la isla de Cer- familles des plantes. J. Casterman, Tournay. deña (Italia). Flora Montiberica 41: 35–61. pp. 57-58. DOI: https://doi. org/10.5962/bhl. Badano, E.I., Pugnaire, F.I., (2004) Invasion of Aga- title.443 ve species (Agavaceae) in south-east Spain: El Mokni, R. & Domina, G. (2020) Additions to ter- invader demographic parameters and impacts restrial flora of Tunisia: occurrence and taxo- on native species. Diversity and Distributions nomic notes. Check List 16(3): 553–561. https:// 10(5/6): 493-500. doi.org/10.15560/16.3.553 Berger, A. (1921) Die Agaven. Jena, Springer-Ver- El Mokni, R. & Domina, G. (2018) Spirodela polyrhi- lag, 285 pp. za (L.) Schleid. & Wolffia arrhiza (L.) Wimm.– Borges, P.A.V., Abreu, C., Aguiar, A.M.F., Carval- Pp. 197-198. In: Raab-Straube E. von & Raus ho, P., Jardim, R., Melo, I., Oliveira, P., Sérgio, Th. (ed.) 2018: Euro+Med-Checklist Notulae, C., Serrano, A.R.M., Vieira, P. (Ed.) (2008) A 9 [Notulae ad floram euromediterraneam perti- list of the terrestrial fungi, flora and fauna of nentes No. 38]. Willdenowia 48: 195–220. Madeira and Selvagens archipelagos. Funchal El Mokni, R. & El Aouni, M.H. (2011a) Découverte and Angra do Heroísmo, Direcção Regional do de la grande camomille, Tanacetum parthe- Ambiente da Madeira and Universidade dos nium (Asteraceae) pour la flore de Tunisie : Açores. une adventice naturalisée. Flora Mediterranea Brandes, D. & Fritzsch K. (2002) Alien plants of 21: 299–303. Fuerteventura, Canary Islands. Plantas ex- El Mokni, R. & El Aouni, M.H. (2011b) Découverte tranjeras de Fuerteventura, Islas Canarias. de Sparaxis tricolore, Sparaxis tricolor (Curt.) Rev. Version. 25 pp. http://www.digibib.tu-bs. Ker-Gawl. (Iridaceae Juss.) pour la flore de Tu- de/?docid=00001079. nisie : une adventice naturalisée. Le Monde des CABI (2020) Agave sisalana. In: Invasive Species Plantes 505: 11-14. Compendium. Wallingford, UK: CAB Interna- El Mokni, R. & El Aouni, M.H. (2012) Zantedeschia tional. www.cabi.org/isc. aethiopica (Araceae) a new species naturalized

232 Bradleya 39/2021 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise in the Northwest of Tunisia. Flora Mediterra- El Mokni, R., Elaissi, A. & VeBradleyarloove, F. (2019b) 39/2021 New nea 22: 191-196. DOI : 10.7320/FlMedit22.191. succulents for the Tunisian and Northpages African221–235 El Mokni, R. & El Aouni, M.H. (2013) Le sésame, alien flora. Haseltonia 26: 68–77. Sesamum indicum L. (Pedaliaceae) une adven- El Mokni, R., Elaissi, A. & El Aouni, M.H. (2016) tice récemment naturalisée en Tunisie. Poiretia Cuscuta campestris (Cuscutaceae) un holopa- 5: 6–14. rasite nouveau et envahissant pour la flore de El Mokni, R. & Iamonico, D. (2018a) A new record Tunisie. Flora Mediterranea 26: 179–189. Doi : for the non-native flora of Tunisia, Eclipta 10.7320/FlMedit26.179. prostrata (Asteraceae), and a note on the na- El Mokni, R., Hamdi, N., De Bélair, G. & El Aouni, tional status of Erigeron bonariensis, Sym- M.H. (2012) Découverte d’Ibicella lutea (Lindl.) phyotrichum squamatum (Asteraceae), and Van Eselt. (Martyniaceae) en Kroumirie (Nord- Lepidium didymum (Brassicaceae). Flora Med- Ouest de la Tunisie). Poiretia 4: 1–6. iterranea 28: 145–153. El Mokni, R., Kasri, M. & El Aouni, M.H. (2013) El Mokni, R. & Iamonico, D. (2018b) Three new Volkameria inermis (Lamiaceae) a new alien records of Senecioneae (Asteraceae) for the al- species naturalized to the Tunisian coast, first lochthonous Tunisian flora: occurrence and tax- record for North-Africa. Flora Mediterranea onomic notes. Flora Mediterranea 28: 385–392. 23: 117–122. El Mokni, R. & Iamonico, D. (2019) Bassia scoparia El Mokni, R., Ardenghi, N.M.G. & Verloove, F. and Sesuvium portulacastrum, two new natu- (2019a). Opuntia elatior Mill. (Cactaceae). In: ralized aliens to the Tunisian flora. Flora Medi- Sukhorukov, A.P., Kushunina, M., El Mokni, R., terranea 29: 191–196. Ardenghi N.M.G., Verloove, F., Uotila, P., Bai- El Mokni, R. & Sáez, L. (2019) Kalanchoe marmo- der, C., Bruyns, P.V.& Klak, C. (2019) Chorolo- rata Baker & K. sexangularis N. E. Br. var. sex- gical and taxonomic notes on African plants, angularis – Pp. 99-100. In: Raab-Straube E. von 4: Caryophyllales, Botany Letters 166(4): 401– & Raus Th. (Ed.) 2019, Euro+Med-Checklist 416. DOI: 10.1080/23818107.2019.1652848. Notulae, 10 [Notulae ad floram euro mediter- El Mokni, R., Sáez Goñalons, L., El Aouni, M.H. raneam pertinentes No. 39]. Willdenowia 49: (2018) The genus Kalanchoe (Crassulaceae) 95–115. in Tunisia. In: Sukhorukov, A.P., Kushuni- El Mokni, R. & Verloove, F. (2019a) New records, na, M., El Mokni, R., Goñalons L.S., El Aou- distribution and taxonomic notes for non-na- ni M.H. & Daniel, T.F. (2018) Chorological tive vascular flora of Tunisia – I. Poaceae. Flora and taxonomic notes on African plants, Mediterranea 29: 45–53. 3. Botany Letters 165(2): 228–240, DOI: El Mokni, R. & Verloove, F. (2019b) Cyperus al- 10.1080/23818107.2018.1465467. ternifolius subsp. flabelliformis Kük. (Cypera- El Mokni, R., Verloove, F., Guiggi, A. & El Aouni, ceae). – Pp. 430–431. In: Raab-Straube E. von M.H. (2020) New records of cacti (Opuntioi- & Raus Th. (Ed.) 2019, Euro+Med-Checklist deae & Cactoideae, Cactaceae) from Tunisia. Notulae, 11 [Notulae ad floram euro-mediter- Bradleya 38: 35–50. raneam pertinentes No. 40]. Willdenowia 49: Elmqvist, T., Cox, P.A. (1996) The evolution of vi- 421–445. https:// doi.org/10.3372/wi.49.49312. vipary in flowering plants. Oikos 77: 3–9. El Mokni, R. (2018) Catharanthus roseus (L.) G. Euro+Med Plantbase. (2020) The Information Re- Don., Vinca difformis Pourr., V. major L., V. mi- source for Euro-Mediterranean Plant Diversity. nor L., Lonicera japonica Thunb., Canna indi- Accessed 28 March 2017. http://www.emplant- ca L., Tradescantia fluminensis Vell. & T. palli- base.org. da (Rose) D. R. Hunt., Ipomoea indica (Burm.) Flora of China (2000) AGAVE Linnaeus. 24: 270– Merr., Lablab purpureus (L.) Sweet., Polygo- 271. num argyrocoleon Kunze. & Solanum rostra- Forster, P.I. (1992) New varietal combinations in tum Dunal. – P. 197-213. In: Raab-Straube E. Agave vivipara (Agavaceae). Brittonia 44(1): von & Raus Th. (eds.) 2018: Euro+Med-Check- 74–75. https://doi.org/10.2307/2807447. list Notulae, 9 [Notulae ad floram euromedi- Forster, P.I. (1996) Naturalized succulents in the terraneam pertinentes No. 38]. Willdenowia 48: Australian flora. Haseltonia 4: 57–65. 195–220. Foxcroft, L.C., Richardson D.M., Wilson J.R. El Mokni, R. (2019) Lithraea molleoides (Vell.) (2008) Ornamental plants as invasive aliens: Engl., Schinus molle L., Schinus terebinthifolius problems and solutions in Kruger National Raddi (Anacardiaceae)., Nicotiana alata Link Park, South Africa. Environmental Manage- & Otto (Solanaceae). pp. 422–440. In: Raab- ment 41(1): 32–51. https://doi.org/10.1007/ Straube E. von & Raus Th. (ed.), Euro+Med- s00267-007-9027-9. Checklist Notulae, 11 [Notulae ad floram Franck, A.R. (2012) Guide to Agave, Cinnamo- euro-mediterraneam pertinentes No. 40]. Will- mum, Corymbia, Eucalyptus, Pandanus, and denowia 49: 421–445. https:// doi.org/10.3372/ Sansevieria in the flora of Florida. Phytoneu- wi.49.49312. ron 102: 1–23.

Bradleya 39/2021 233 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Galasso, G., Conti, F., Peruzzi, L., Ardenghi, N.M.G., Mooney, H.A. & Hobbs, R. J. (2000) Invasive species Banfi, E., Celesti Grapow, L., Albano, A., Ales- in a changing world. Washington, DC: Island sandrini, A., Bacchetta, G., Ballelli, S., Bandini Press. Mazzanti, M., Barberis, G., Bernardo, L., Blasi, Moragues, E. & Rita, J. (2005) Els vegetals in- C., Bouvet, D., Bovio, M., Cecchi, L., Del Guac- troduïts a les Illes Balears. Conselleria de Medi chio, E., Domina, G., Fascetti, S., Gallo, L., Gu- Ambient, Palma de Mallorca, Govern de les Il- bellini, L., Guiggi, A., Iamonico, D., Iberite, M., les Balears. Jiménez-Mejías, P., Lattanzi, E., Marchetti, D., Nobel, P.S. (1988) Environmental biology of Agaves Martinetto, E., Masin, R.R., Medagli, P., Passa- and cacti. Cambridge, Cambridge University lacqua, N.G., Peccenini, S., Pennesi, R., Pierini, Press. B., Podda, L., Poldini, L., Prosser, F., Raimondo, Otto, R. & Verloove, F. (2016) New xenophytes F.M., Roma-Marzio, F., Rosati, L., Santangelo, from La Palma (Canary Islands, Spain), with A., Scoppola, A., Scortegagna, S., Selvaggi, A., emphasis on naturalized and (potentially) in- Selvi F., Soldano, A., Stinca, A., Wagensommer, vasive species. Collectanea Botanica 35: e001. R.P., Wilhalm, T., Bartolucci, F. (2018) An up- doi: 10.3989/collectbot.2016.v35.001. dated checklist of the vascular flora alien to Pimentel, D., McNair, S., Janecka, J., Wightman, J., Italy. Plant Biosystems 152(3): 556–592. https:// Simmonds, C., O’Connell, C., Wong, E., Russel, doi.org/10.1080/11263504.2018.1441197. L., Zern, J., Aquino, T. & Tsomondo, T. (2001) García-Mendoza, A., Chiang, F. (2003) The confu- Economic and environmental threats of alien sion of Agave vivipara L. and A. angustifolia plant, animal, and microbe invasions. Agroeco- Haw., two distinct taxa. Brittonia 55(1): 82–87. systems and Environment 84: 1–20. https://doi.org/10.1663/0007-196X(2003)055[0 Pyšek, P., Richardson, D.M. & Williamson, M. (2004) 082:TCOAVL]2.0.CO;2. Predicting and explaining plant invasions Gentry, H.S. (1982) Agaves of Continental North through analysis of source area floras: some America. Tucson, The University of Arizona critical considerations. Diversity and Distribu- Press, pp. 270–290. tions 10: 179–187. Guillot-Ortiz, D., van der Meer, P., Laguna-Lum- Podda, L., Lazzeri, V., Mayoral, O. & Bacchetta, breras, E., Rosselló-Picornell, J.A. (2009) El G. (2012) The checklist of the Sardinian alien género Agave L. en la flora alóctona valenciana flora: an update. Notulae Botanicae Horti Agro- (Monografías de la revista Bouteloua, 3). Jol- botanici Cluj-Napoca 40(2): 14–21. https://doi. ube Consultor y Editor Ambiental. Teruel y org/10.15835/nbha4028225. Jaca, España. 94 pp. Randall, R.P. (2017) A Global Compendium of Henderson, L. (1995) Plant invaders of southern Af- Weeds. Third edition. Perth, Western Australia. rica. Pretoria, Plant Protection Research Insti- Sáez, L., Serapio, J., Gómez-Bellver, C., Ardenghi, tute. Agricultural Research Council. N.M.G., Guillot, D. & Rita, J. (2016) New re- Hochstätter, F. (2015) Agave Linné (Agavaceae). cords in vascular plants alien to the Balearic Privately published on the web. Available Islands. Orsis 30: 101–131. at https://issuu.com/fhnavajo/docs/agave. Santos-Guerra, A., Padrón Mederos, M.A., Mesa Acessed 20 April 2020. Coello, R., Ojeda Land, E. & Reyes-Betancort, Iamonico, D. & El Mokni, R. (2016) Patellifolia pro- J.A. (2013) 244. Establecimiento de plantas cumbens (Amaranthaceae), new to the Flora of intoducidas en la flora vascular silvestre ca- Tunisia in Raab-Straube & Raus: Euro+Med- naria I (Helechos, Gimnospermas Y Mono- Checklist Notulae, 6. Willdenowia 46(3): 423– cotiledóneas). Acta Botanica Malacitana 38: 442. 176–182. Le Floc’h, E., Le Houerou, H.N., Mathez, J. (1990) Smith, G. & Figueiredo, E. (2014) State of our knowl- History and patterns of plant invasion in edge of online resources of the types of names Northern Africa. In: di Castri, F., Hansen, A.J., of Agave L. (Agavaceae/Asparagaceae) with a Debussche, M. (Eds.). Biological invasions in Eu- list of names attributed to William Trelease rope and the . Dordrecht, with notes on types, online resources and ety- Kluwer Academic Pub, pp. 105–133. mology. Phytotaxa 170(4): 221–249. Mabberley, D.J. (1997) The Plant-Book. A portable Smith, G.F. & Mössmer, R.M. (1996) FSA contribu- dictionary of the vascular plants (secod edi- tions 4: Agavaceae. Bothalia 26: 31–35. tion). Cambridge: Cambridge University Press. Smith, G.F. & Steyn, E.M.A. (1999a) Agave vivip- Mack, R.N., Simberloff, D., Lonsdale, W.M., Evans, ara: the correct name for Agave angustifolia. H., Clout, M., & Bazzaz, F.A. (2000) Biotic in- Bothalia 29: 100. https://doi.org/10.4102/abc. vasions: Causes, epidemiology, global conse- v29i1.577. quences and control. Ecological Applications Smith, G.F. (2011) Agavaceae Dumort. In: Walters, 10: 689–710. M., Figueiredo, E., Crouch, N.R., Winter, P.J.D., Manni, Q.G. (2015) Noterella: 0165. Agave angusti- Smith, G.F., Zimmermann, H.G. & Mashope, B.K. folia Haw. Acta Plantarum Notes 3: 104. Naturalised and invasive succulents of south-

234 Bradleya 39/2021 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise ern Africa, pp. 34–63. ABC Taxa 11. Brussels, Verloove, F. (2013) New xenophytesBradleya from 39/2021 Gran Ca- The Belgian Development Cooperation. naria (Canary Islands, Spain), withpages emphasis 221–235 Steyn, E.M.A. & Smith, G.F. (2000) Agave vivipara: on naturalized and (potentially) invasive spe- a naturalised alien in Southern Africa. Bo- cies. Collectanea Botanica 32: 59-82. thalia 30: 43–55. https://doi.org/10.4102/abc. Verloove, F., Thiede, J., Marrero Rodríguez, Á., Sa- v30i1.537. las-Pascual, M., Reyes-Betancort, J.A., Ojeda- Thiede, J. (2001) Agave. In Eggli, U. (ed.), Illustrat- Land, E. & Smith, G.F. (2019) A synopsis of feral ed handbook of succulent plants. Monocotyle- Agave and Furcraea (Agavaceae, Asparagaceae dons. Berlin, Springer-Verlag, pp. 6–76. s. lat.) in the Canary Islands (Spain). Plant Thiede, J., Smith, G.F. & Eggli, U. (2019) Infrage- Ecology and Evolution 152(3): 470–498. https:// neric classification of Agave L. (Asparagaceae: doi. org/10.5091/plecevo.2019.1634. Agavoideae / Agavaceae): a nomenclatural Vieira, R. (2002) Flora da Madeira. Plantas vascu- assessment and updated classification at the lares naturalizadas no arquipélago da Madeira. rank of section, with new combinations. Brad- Boletim do Museu Municipal do Funchal, Sup- leya 37: 240–264. https://doi. org/10.25223/ plement 8: 5–281. brad.n37.2019.a22. Vilà, M., Meggaro, Y., & Weber, E. (1999) Prelimi- Thomson G. (2020) New Zealand’s endemic, nary analysis of the naturalized flora of north- indigenous, naturalised and casual succu- ern Africa. Orsis 14: 9–20. lents. Bradleya 38: 170–186. Walters, M., Figueiredo, E., Crouch, N.R., Winter, Trejo-Torres, J.C., Gann, G.D. & Christenhusz, P.J.D., Smith, G.F., Zimmermann, H.G. & Mashope, M.J.M. (2018) The Yucatan Peninsula is the B.K. (2011) Naturalised and invasive succu- place of origin of (Agave sisalana, As- lents of southern Africa. ABC Taxa 11: 37. paragaceae): historical accounts, phytogeog- Wells, M.J., Balsinhas, A.A., Joffe, H., Engelbrecht, raphy and current populations. Botanical Sci- V.M., Harding, G. & Stirton, C.H. (1986) A cat- ences 96(2): 366–379. https://doi.org/10.17129/ alogue of problem plants in southern Africa. botsci.1928. Memoirs of the Botanical Survey of South Af- Ullrich, B. (2006) Agave attenuata ssp dentata rica 53. (Roezl) Ullrich. Haseltonia 12: 22–30. Wijnands, D.O. (1983) The botany of the Comme- Verhoek, S. (1998) Agavaceae. In: Kubitzki, K. The lins. Rotterdam, Balkema. families and genera of vascular plants. III. Flow- ering plants. . Lilianae (except Orchidaceae): 60–70. Berlin, Springer-Verlag.

Editor’s addendum. Agaves were well known long for before the of Linnaeus. This illustrations of ‘Aloe Americana’ (= Agave americana) is from Basilius Besler’s Hortus Eyestettensis published in 1613.

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