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A synopsis of feral Agave and Furcraea (Agavaceae, Asparagaceae s. lat.) in the

Canary Islands (Spain)

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Plant Ecology and Evolution 152 (3): 470–498, 2019
https://doi.org/10.5091/plecevo.2019.1634

REGULAR PAPER

A synopsis of feral Agave and Furcraea (Agavaceae, Asparagaceae s. lat.)

in the Canary Islands (Spain)
Filip Verloove1,*, Joachim Thiede2, Águedo Marrero Rodríguez3, Marcos Salas-Pascual4,
Jorge Alfredo Reyes-Betancort5, Elizabeth Ojeda-Land6 & Gideon F. Smith7

1
Meise Botanic Garden, Nieuwelaan 38, B-1860 Meise, Belgium
2
Schenefelder Holt 3, 22589 Hamburg, Germany
3
Jardín Botánico Canario Viera y Clavijo, Unidad Asociada al CSIC, C/ El Palmeral nº 15, Tafira Baja, E-35017 Las Palmas de Gran
Canaria, Gran Canaria, Canary Islands, Spain
4
Instituto de Estudios Ambientales y Recursos Naturales (i-UNAT), Campus Universitario de Tafira, Universidad de las Palmas de Gran
Canaria, E-35017 Las Palmas de Gran Canaria, Gran Canaria, Canary Islands, Spain
5
Jardín de Aclimatación de La Orotava (ICIA). C/ Retama 2, 38400 Puerto de la Cruz, Canary Islands, Spain
6
Viceconsejería de Medio Ambiente. Gobierno de Canarias. C/ Avda. de Anaga, 35. Planta 11. 38071 Santa Cruz de Tenerife, Canary
Islands, Spain
7
Department of Botany, P.O. Box 77000, Nelson Mandela University, Port Elizabeth, 6031 South Africa / Centre for Functional Ecology,
Departamento de Ciências da Vida, Calçada Martim de Freitas, Universidade de Coimbra, 3001-455 Coimbra, Portugal
*Corresponding author: filip.verloove@meisebotanicgarden.be

Background – Species of Agave and Furcraea (Agavaceae, Asparagaceae s. lat.) are widely cultivated as
ornamentals in Mediterranean climates. An increasing number is escaping and naturalising, also in natural
habitats in the Canary Islands (Spain). However, a detailed treatment of variously naturalised and invasive
species found in the wild in the Canary Islands is not available and, as a result, species identification is
often problematic.
Methods – The present study is based on many years of fieldwork on the islands of Fuerteventura, Gran
Canaria, Lanzarote and Tenerife.
Results – Fourteen species and several additional infraspecific taxa of Agave have been identified. In
addition to the widely naturalised and invasive Agave americana, A. fourcroydes and A. sisalana (incl. the
not previously reported var. armata), the following species of Agave were observed: A. angustifolia (incl.
var. marginata), A. attenuata, A. filifera, A. franzosinii, A. lechuguilla, A. macroacantha, A. murpheyi,
A. oteroi, A. salmiana (var. ferox and var. salmiana), A. aff. tequilana and A. vivipara. From the genus
Furcraea three species were found: F. foetida, F. hexapetala and F. selloana. Several of these newly
detected species of Agave and Furcraea are at least locally naturalised. A key for the identification of the
representatives of these genera in the study area is presented and all species are illustrated. Additional
nomenclatural, taxonomic and ecological notes are also provided.

Keywords – Agavaceae; Agave; Canary Islands; Fuerteventura; Furcraea; Gran Canaria; Lanzarote;
nomenclature; Spain; taxonomy; Tenerife.

© 2019 The Authors. This article is published and distributed in Open Access under the terms of the Creative Commons Attribution
License (CC BY 4.0), which permits use, distribution, and reproduction in any medium, provided the original work (author and source)
is properly cited.
Plant Ecology and Evolution is published by Meise Botanic Garden and Royal Botanical Society of Belgium
ISSN: 2032-3913 (print) – 2032-3921 (online)
 Verloove et al., Feral Agave and Furcraea in Canary Islands

INTRODUCTION invasive species. According to Acebes Ginovés et al. (2010)

this applies at least to Agave americana L. and Furcraea
Agave L. and Furcraea Vent. are two genera from the Agava- foetida (L.) Haw. The identity, however, of the species cur-
ceae (s. str.), a well-known succulent plant family of the New rently found in the wild there required critical re-assessment:
World, that now is usually included in Asparagaceae, based some morphologically similar species had been overlooked
on molecular studies (Bogler et al. 2006; Chase et al. 2009). so far, whereas several additional distinct species have been
The adherence to traditional concepts, such as the Agavace- recorded in recent years, some of them at least locally natu-
ae s. str., represents a permissible and alternative, narrower ralised.
family concept (Thiede 2016). The exact number of species
In this paper a synopsis of the genera Agave and Furcraea
in Agave is uncertain and varies from one author to another,
in Gran Canaria, Tenerife, Lanzarote and Fuerteventura is
depending on species delimitation, and new species are still
presented. A key for the identification of all species currently
regularly described (e.g. Giraldo-Cañas 2017; Arzaba-Villal-
found in the wild is presented and all species are illustrated.
ba et al. 2018; Starr et al. 2018; García Mendoza et al. 2019).
Further nomenclatural, taxonomic and ecological comments
A recent synopsis (Thiede in press) covers some 265 species
are also provided.
which range on the mainland from the southern U.S.A. to
Panama and Venezuela. Several additional species are found
in the Caribbean. The center of diversity is in Mexico where MATERIALS AND METHODS
at least 125 species occur (García-Mendoza & Galván 1995;
García-Mendoza 2002). The majority of species occur in arid The present study is based on many years of fieldwork on the
to semiarid habitats in or near deserts, especially in desert islands of Fuerteventura, Gran Canaria, Lanzarote and Ten-
grasslands and oak-pine woodlands. The genus is economi- erife by most of the authors (FV, AMR, MSP, JARB, EOL).
cally important, either as agricultural crops for the produc- The other authors (JT, GFS) assisted with the identification
tion of fibres (sisal, henequen, ixtle) or beverages (tequila, of some of the more critical taxa and provided further useful
pulque, mescal), or as ornamentals. Furcraea is a smaller information on the genera Agave and Furcraea. In addition,
genus with ca. 23 species (Thiede in press) that are distrib- numerous literature references (including original descrip-
uted throughout tropical America. Although they usually oc- tions and protologues) were consulted in order to better un-
cur in less arid climate types, they are more frost-sensitive derstand the distinguishing features used for the separation
than most agave species. Several species are economically of the relevant taxa.
important: they are grown as agricultural crops, mostly for Each entry includes: (1) the scientific name of the taxon
the production of fibres, or as ornamentals. (accompanied by one or more synonyms, if useful); (2) a list
The generic boundaries of Agave have long been and to of localities (including geographical coordinates which en-
some extent still are controversial. Most molecular sequence able local governments to eradicate plants or populations, if
data thus far available (e.g. Bogler et al. 2006; Good-Ávi- deemed desirable) and voucher specimens when available;
la et al. 2006; Flores-Abreu et al. 2019) demonstrated that, (3) the distribution of the taxon (native as well as introduced
as traditionally circumscribed, the genus is paraphyletic. In area); (4) ecology and habitat data; and (5) additional useful
order to render Agave monophyletic, the genera Manfreda comments, usually about its invasiveness (for definitions, see
Salisb., Polianthes L. and Prochnyanthes S.Watson needed Blackburn et al. 2011), identification and/or nomenclatural
to be merged with it (Thiede 2012). If traditional generic or taxonomic issues. Photographs are presented for all taxa
concepts for these genera were retained, these would be in (figs 1–7). Taxa nomenclature is mostly in accordance with
conflict with molecular phylogenetic data and derived classi- recent insights. Authorities of plant names follow the Inter-
fication concepts (Thiede 2016). However, AFLP molecular national Plant Names Index (IPNI 2019).
markers showed these genera to group separate from and not As a rule, all records were documented by photographs
nested within Agave (Gil-Vega et al. 2007). Furcraea is mor- only (deposited with the authors). Herbarium specimens were
phologically distinguished from Agave in several respects. exceptionally collected and these were deposited in the her-
Its inflorescence always is a massive pyramidal panicle (vs. baria of Meise Botanic Garden, Belgium (BR) and the Bo-
narrowly elongate to ± broad panicles or elongated spikes tanic Garden Viera y Clavijo in Las Palmas de Gran Canaria
or racemes) with pendent white or whitish flowers (vs. erect (LPA). Herbarium acronyms follow Thiers (continuously up-
flowers that are rarely whitish). Leaves are often more flex- dated).
ible with a much less prominent terminal spine and stamens
are only half as long as tepals (vs. usually longer than the RESULTS AND DISCUSSION
tepals). Also, the tepals are (almost) free (vs. basally fused to
form a ± long tube). According to recent molecular and mor- Agave L. (Linnaeus 1753: 323).
phological phylogenies, Furcraea is sister to Beschorneria
Kunth, and both together are sister to Agave in most analyses Taxonomic synopsis of Agave in the Canary Islands
(Hernández-Sandoval 1995; Bogler & Simpson 1996; Bogler (Fuerteventura, Gran Canaria, Lanzarote, Tenerife) (based on
et al. 2006). Thiede in press and Thiede et al. 2019).
While evidently no indigenous representatives of Agave Subgenus Agave
and Furcraea exist in the Canary Islands, a number of spe- Section Agave
cies have become naturalised in the archipelago. Some even Agave americana L.
reproduce prolifically in natural habitats and are considered Agave franzosinii (Sprenger) P.Sewell

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Pl. Ecol. Evol. 152 (3), 2019

Section Ditepalae Hochstätter across a greater or lesser spectrum of habitats and extent of
Agave murpheyi F.Gibson occurrence).
Section Rigidae (Baker) R.H.Webb & G.D.Starr Agave americana is a highly polymorphic species from the
southern U.S.A. and Mexico and includes several infraspe-
Agave angustifolia Haw.
cific taxa. In addition to cultivars with variegated leaves,
Agave fourcroydes Lem. four varieties or subspecies can be distinguished according to
Agave macroacantha Zucc. Gentry (1982): var. americana, var. expansa (Jacobi) Gentry
Agave sisalana Perrine (syn.: subsp. expansa (Jacobi) Hochstätter), var. oaxacen-
sis Gentry and subsp. protamericana Gentry. In the study
Agave aff. tequilana F.A.C.Weber
area A. americana also occurs in many forms. The invasive
Section Salmianae (A.Berger) Verloove & Thiede form is usually a relatively small plant with short, erect and
Agave salmiana Otto ex Salm-Dyck stiff leaves. It somehow resembles var. expansa, a more or
Agave salmiana var. ferox (K.Koch) Gentry less distinct ‘race’ that is naturalised in parts of South Af-
rica (Smith & Figueiredo 2011) and Australia (Forster 1986).
Agave salmiana var. salmiana
This variety is medium-sized to large with leaves erect to
Section Viviparae (Baker) Verloove & Thiede stiffly spreading, adaxially deeply channeled and distinctly
Agave vivipara L. widened towards the tip, less succulent and with an apical
leaf spine 20–30 mm long. Seeds are never produced. Al-
Subgenus Littaea (Tagl.) Baker though var. expansa was an early introduction in continental
Section Heteracanthae Salm-Dyck Europe (Jacobi 1869) it apparently was not able to escape,
Agave lechuguilla Torr. unless it remained unnoticed. The invasive Canarian plants
Agave oteroi G.D.Starr & T.J.Davis probably do not belong to this variety since they lack the
distinctly widened upper part of the leaf and the terminal
Section Inermes Salm-Dyck spine is longer (fig. 1A). Another expression of A. americana
Agave attenuata Salm-Dyck found in the study area is a massive plant with thickly suc-
Section Littaea (Tagl.) Bentham culent leaves that are usually drooping to one side, adaxially
Agave filifera Salm-Dyck not deeply channeled and with an apical leaf spine 30–50
mm long (fig. 1B). Such plants probably correspond with
var. americana. This taxon is extremely variable, especially
Subgenus Agave in leaf characters. Leaf margins sometimes are remarkably
Inflorescence paniculate (often several times compound) sinuate, especially in young individuals or in re-growth (after
with flowers in large umbellate clusters on lateral peduncles. cutting). Also, marginal teeth can considerably vary in length
and form, even in a single individual. All these glaucous-
leaved forms of A. americana are believed to represent two
Section Agave (= Group Americanae Baker nom. inval.) or more clones but are all assigned to a single taxon, var.
This section, formerly referred to as Group Americanae americana.
Baker, includes the type of the genus, Agave americana. The In addition to plants with glaucous leaves several culti-
autonym section Agave should hence be used (art. 22.2 & vars with variegated leaves are widely grown as ornamentals
art. 22 ex. 2 and 4, art. 37.1 of the ICN; Turland et al. 2018). in the study area. One such form is particularly common in
Plants are usually medium- to large-sized, have long and lan- cultivation and is locally naturalised as well, at least in Gran
ceolate, glaucous leaves, a usually strong surculose habit and Canaria and Tenerife. It has leaves with broad pale yellow-
tall, open panicles with yellow flowers with long tepals and ish margins and is here assigned to var. marginata Trel. (fig.
stamens (Gentry 1982; Thiede in press). In the study area it 1C). Like var. americana it has a suckering habit and easily
is reportedly represented by the very widely distributed and escapes. It has been recorded on rather numerous occasions.
highly invasive species A. americana (Acebes Ginovés et al. However, naturalised populations were only rarely observed,
2010). The latter, however, is extremely variable and records for instance in the following localities:
of it may hide related or similar cryptic taxa (see below). Observation records
During our field studies we noticed that a second species, A. Gran Canaria
franzosinii (Sprenger) P.Sewell, also escapes from cultiva- Las Palmas de Gran Canaria, La Matanza, Calle El Mon-
tion, at least in Gran Canaria. dalon, 28°2′9.92″N, 15°26′21.23″W, 193 m a.s.l., abundantly
naturalised in roadside and edges of fields, 16 Apr. 2018,
Agave americana L. (Linnaeus 1753: 323) obs. F. Verloove.
Las Palmas de Gran Canaria, Monte Lentiscal, Montaña de
Fig. 1A–C Tafira, 28°3′8.79″N, 15°28′28.12″W, 474 m a.s.l., volcanic
Known in the Canary Islands from all islands – naturalised slope, frequent, 24 Apr. 2018, obs. F. Verloove.
and invasive (Acebes Ginovés et al. 2010). Category E sensu Las Palmas de Gran Canaria, GC-4 at Barranco del Salto del
Blackburn et al. (2011) (Fully invasive species, with individ- Negro, 28°3′16.8″N, 15°26′36.19″W, 230–260 m a.s.l., sev-
uals dispersing, surviving and reproducing at multiple sites eral groups, 1 Nov. 2018, obs. Á. Marrero.

472
 Verloove et al., Feral Agave and Furcraea in Canary Islands

Identification key for the species of Agave occurring in the wild

in Fuerteventura, Gran Canaria, Lanzarote and Tenerife

The key is primarily based on vegetative characters since non-flowering individuals are encountered more often than flowering
ones. Only races and forms occurring in the study area are taken into account. For each species the known occurrence in the
Canary Islands is provided between brackets (abbreviations: H = El Hierro; P = La Palma; G = La Gomera; T = Tenerife; C =
Gran Canaria; F = Fuerteventura; L = Lanzarote).

1. Leaf margins filiferous (i.e., margins separating as hair-like threads), unarmed........ Agave filifera (T)
1’. Leaf margins never filiferous, armed or not.......................................................................................... 2
2. Leaves narrow and sword-shaped (ensiform), c. 10–20 × as long as wide with almost straight margins
(hardly narrowed towards the base), very rigid and radiately spreading. Flowers always greenish-
yellow and in a paniculate inflorescence. Plants surculose and very often bulbiliferous (A. angustifolia
complex)............................................................................................................................................... 3
2’. Leaves wider; if narrow then not very rigid and straight-margined and inflorescence not paniculate.
Flowers variable. Plants surculose or not, bulbiliferous or not............................................................. 7
3. Leaves usually (much) less than 100 cm long, sometimes with white or yellow margins. Stem absent
or short.................................................................................................................................................. 4
3’. Leaves usually (much) longer, never with white or yellow margins [from A. sisalana and A. tequilana,
cultivars with variegated margins are known, but these are much less commonly cultivated than those
of A. angustifolia, and appear to be absent from the Canaries]. Stem present, short or distinct.......... 5
4. Leaves 40–60(–100) cm long, light green to glaucous grey, often with white or yellow margins (var.
marginata). Inflorescence 3–5 m..................................................................... A. angustifolia (C, F, T)
4’. Leaves 25–35 cm long, bluish-grey glaucous. Inflorescence ca. 2 m........... A. macroacantha (F, P, T)
5. Leaf margin unarmed (exceptionally with well-developed triangular teeth but then not or only very
weakly glaucous leaved and with smaller flowers). Plant usually green, either yellowish or dark green.
Flowers relatively small, 55–65 mm long...................................................... A. sisalana (C, F, L, P, T)
5’. Leaf margin with regularly spaced and distinct teeth. Plant often glaucous or greyish green. Flowers
usually longer, 60–80 mm long............................................................................................................. 6
6. Apical leaf spine short, 10–20 mm long. Leaves glaucous, with almost parallel margins, rarely
exceeding 120 cm in length. Inflorescence with 20–25 partial inflorescences.......A. aff. tequilana (C)
6’. Apical leaf spine usually longer, 20–30 mm long. Leaves usually greyish-green, tapering towards both
ends, often longer. Inflorescence with 10–18 partial inflorescences... A. fourcroydes (C, F, G, L, P, T)
7. Leaves with conspicuous horny margin. Inflorescences narrowly elongate panicles........................... 8
7’. Leaves without horny margin. Inflorescence narrowly elongate or broad panicles.............................. 9
8. Leaves narrow, 10–20 × as long as wide, marginal teeth typically deflected, weak and friable, regular
in size, 2–6 mm. Inflorescence laxly flowered, with linear bracts.............................A. lechuguilla (C)
8’. Leaves broadly-ovate, sometimes not much longer than wide (especially the innermost),
marginal teeth deltoid or irregular, (10–) 20–40 mm. Inflorescence densely flowered, with deltoid
bracts...................................................................................................................................A. oteroi (C)
9. Leaves large to massive, often at least 100 cm long in normally developed plants (except in A. salmiana
var. ferox). Leaf margins with very distinct teeth. Inflorescence never bulbiliferous......................... 10
9’. Leaves medium-sized (usually not exceeding 70 cm in length). Leaf margins smooth or with short
teeth ca. 1–4 mm long. Inflorescence bulbiliferous or not.................................................................. 12
10. Leaves green. Inflorescence pole short, hardly longer than wide.............................A. salmiana (C, T)
10’. Leaves glaucous. Inflorescence pole elongate, at least twice as long as wide.....................................11
11. Leaf surface smooth. Plant glaucous, usually not distinctly patterned with green (variegated in var.
marginata).........................................................................................A. americana (C, F, G, H, L, P, T)
11’. Leaf surface rough. Plant glaucous to whitish, often patterned with dark green where the glaucous
covering was rubbed off (never variegated)................................................................A. franzosinii (C)

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Pl. Ecol. Evol. 152 (3), 2019

12 Leaves ovate-acuminate, often arching or sigmoid. Margins either smooth or with closely and regularly
spaced teeth......................................................................................................................................... 13
12’. Leaves lanceolate, arching, not sigmoid. Margins regularly toothed, small, mostly 3–4 mm................
.......................................................................................................................................A. murpheyi (C)
13 Leaves always with smooth margins and soft apical spine. Inflorescence spicate, very densely flowered
and arching, not bulbiliferous; tepals greenish-yellow. Stem always present at maturity, up to 100 cm
long...................................................................................................................A. attenuata (C, F, P, T)
13’. Leaves with margins with closely-spaced short teeth 3–7 mm and distinct apical spine. Inflorescence
paniculate, erect, bulbuliferous; tepals bright yellow. Stem absent................................ A. vivipara (C)

Tenerife americana by most contemporary authors (Gentry 1982;

Candelaria, Cuesta de Las Tablas, TF-28 road close to Em- Thiede in press) and this taxonomy is followed here.
balse, 28°23′26.36″N, 16°21′8.37″W, 220 m a.s.l., five in- At least two additional glaucous-leaved species have a
dividuals with A. fourcroydes, 7 Feb. 2010, obs. E. Ojeda- similar general appearance and also attain very large rosette
Land. and inflorescence dimensions: Agave lurida Aiton and A. we-
San Cristóbal de La Laguna, Bajamar, 28°33′23.40″N, beri J.F.Cels ex J.Poiss. Both have been recorded as escapes
16°20′4.42″W, 70 m a.s.l., three young individuals in a field from cultivation (e.g. Guillot Ortiz & van der Meer 2008b;
near the road together with Furcraea selloana, 2 May 2011, Franck 2012; Sáez & Guillot Ortiz 2015; Smith & Figueiredo
obs. E. Ojeda-Land. 2015; Verloove et al. 2018) and can be expected to occur in
San Cristóbal de La Laguna, La Cuesta, Barranco Los the study area as well. Agave lurida differs from A. america-
Menceyes, 28°28′14.00″N, 16°17′44.86″W, 375 m a.s.l., na in being only sparingly surculose (if at all) with leaf mar-
three established individuals on anthropised slopes of ravine,
gins with more regularly and closely spaced, slightly shorter
18 May 2017, obs. E. Ojeda-Land.
teeth (1–2 cm apart and 5–7 mm long vs. 2–5 cm apart and
La Matanza de Acentejo, Carretera El Caletón, at several
4–10 mm long) (Gentry 1982). Agave weberi was placed in
points (for instance at 28°27′16.26″N, 16°27′43.18″W;
Group Sisalanae by Gentry (1982) but more likely belongs
28°27′17.32″N, 16°27′42.83″W; 28°27′28.87″N,
in section Agave (Ullrich 1990; Smith & Figueiredo 2015;
16°27′54.67″W), roadside, between 50 and 250 m a.s.l.,18
Thiede in press). Compared with A. americana it has leaves
Apr. 2018, obs. E. Ojeda-Land.
Santa Úrsula, La Quinta, 28°26′1.62″N, 16°29′40.75″W, 195 with teeth either absent or minute to small and then closely
m a.s.l., four individuals near houses and another five near spaced. The inflorescence pole often leans or droops as op-
the roundabout of the TF-5, 28°25′38.28″N, 16°29′37.12″W, posed to the rigidly erect inflorescence pole of A. americana.
255 m a.s.l., 25 Apr. 2018, obs. E. Ojeda-Land.
Specimen collected Agave franzosinii (Sprenger) P.Sewell (Sewell 1889: 639)
Gran Canaria Figs 1D–F, 2A
Santa Lucía de Tirajana, bajada a La Sorrueda, Llanos de la
Piedra, 27º53′32.8″N, 15º32′06.5″W, 550 m a.s.l, antiguos Agave americana L. var. franzosinii Sprenger (Sprenger
bancales, subespontánea, estolonífera, 29 Jun. 2019, Á. Mar- 1885: 130). – Type: [icon in] Curtis’s Botanical Magazine
rero & R. López González 37495-37496 (LPA). 136: t. 8317 (Berger 1910) (Thiede 2017).
Similar plants with leaf margins with intercalary teeth (i.e., Agave beaulueriana Jacobi (Jacobi 1869: 150). – Type: [icon
with a mixture of smaller and larger teeth) are sometimes in] Curtis’s Botanical Magazine 136: t. 8317 (Berger 1910)
distinguished as Agave ingens A.Berger var. picta (Salm- (Thiede 2017).
Dyck) A.Berger (syn.: A. picta Salm-Dyck) (Guillot Ortiz &
New to the flora of the Canary Islands – category C2 sensu
van der Meer 2003, 2010, 2013a; Guillot Ortiz et al. 2012;
Blackburn et al. (2011) (Individuals surviving in the wild in
Rubal et al. 2013). However, this taxon probably is of limited
taxonomic value. In Gran Canaria (for instance in La Ma- location where introduced, reproduction occurring, but popu-
tanza, Las Palmas de Gran Canaria) plants with variegated lation not self-sustaining).
leaves and intercalary teeth have been observed but on the Observation records
same plant normally developed teeth are also found. Some Gran Canaria
plants from Tenerife are yet different and even more closely Las Palmas de Gran Canaria, Tamaraceite, 28°5′32.46″N,
approach A. ingens var. picta. Apart from the presence of 15°28′45.95″W, 193 m a.s.l., roadside, a single plant (es-
intercalary teeth in most or all leaves they differ in having caped from an ornamental plantation ca. 250 m further
leaves with a brighter, almost shiny green part. It is obvious north), 18 Apr. 2018, obs. F. Verloove (fig. 1D–E).
that the variegated plants show some degree of variation, Las Palmas de Gran Canaria, Tafira Baja, close to Finca La
and may possibly be of multiple origins. However, A. ingens Palmita, 28°4′6.78″N, 15°27′25.61″W, 327 m a.s.l., aban-
(incl. var. picta) is considered a heterotypic synonym of A. doned area, scattered individuals (relic of former planta-

474
 Verloove et al., Feral Agave and Furcraea in Canary Islands

Figure 1 – A. Agave americana, Adeje, Taucho (T), July 2016. This is a very variable species in the study area. This form looks like but is
not identical with var. expansa. It is a relatively small plant with shorter leaves that are held stiffly upright. B. Idem, Santa Brigida (C), April
2018. Another expression of the same species is massive with very large, rather pliable leaves. This form is reminiscent of A. franzosinii. C.
Idem. Var. marginata, Santa Lucía de Tirajana (C), April 2018. D. A. franzosinii, Las Palmas de Gran Canaria (Tamaraceite) (C), April 2018.
E. Idem. The glaucous covering of the leaves is easily rubbed off. F. Idem. Comparison of leaf colour and spine orientation in A. franzosinii
(upper) and A. americana. Photographs: A–E by F. Verloove; F by M. Salas Pascual.

475
Pl. Ecol. Evol. 152 (3), 2019

Figure 2 – A. Agave franzosinii, Las Palmas de Gran Canaria (La Paterna) (C), May 2018. This species has a somewhat laxer inflorescence
than A. americana. B. A. murpheyi, Santa Lucía de Tirajana (C), April 2018. C. Idem. Detail of leaf marginal spines. D. A. angustifolia,
Mogán (C), December 2017. Inflorescences are nearly always bulbiliferous in this species. E. Idem. Detail of leaf spination of green form.
F. Idem, Güímar (T), January 2017. Photographs: A by Á. Marrero; B–F by F. Verloove.

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tion?), 24 Apr. 2018, obs. F. Verloove, Á. Marrero Rodríguez Section Ditepalae Hochstätter (= Group Ditepalae Gentry
& M. Salas Pascual. nom. inval.)
Specimen collected Gentry (1982) first published this name but it was not valid
Gran Canaria since the type of the name was not indicated (Art. 37.1 of the
Las Palmas de Gran Canaria, Tafira Baja, entornos del Cam- ICN; Turland et al. 2018). This section includes 17 species
pus, 28º04′12.2″N, 15º27′16.0″W, 320 m a.s.l., zonas ajar- from the southern U.S.A. (Arizona, New Mexico) and Mex-
dinadas algo abandonadas, subespontánea, estolonífera, 27 ico (Thiede in press). Plants are usually small to medium-
Jun. 2019, Á. Marrero 37476-37477 (LPA). sized (leaves usually well below 100 cm in length), generally
Agave franzosinii is not known in the wild. It is frequently surculose and tepals are strongly dimorphic with the outer
grown as an ornamental, for instance in parts of the Mediter- lobes conspicuously larger (Gentry 1982; Thiede in press). A
ranean area, the West Indies and – more recently – also in single species has been recorded in the study area.
the Canary Islands. It is very closely similar to A. americana
and, like the latter, is highly surculose. As a result, it eas- Agave murpheyi F.Gibson (Gibson 1935: 83–85, f. 1)
ily escapes and is increasingly observed in the wild, also in
continental Europe, for instance in Spain (Guillot Ortiz et al. Fig. 2B–C
2008; Sáez et al. 2014; Mesquida et al. 2016) and Italy, incl. New to the flora of the Canary Islands – category C3 sensu
Sardinia and Sicily (Celesti-Grapow et al. 2010; Galasso et Blackburn et al. (2011) (Individuals surviving in the wild in
al. 2018). It is expected to be present also in Portugal (Smith location where introduced, reproduction occurring, and pop-
& Figueiredo 2007) and is here reported for the first time ulation self-sustaining).
as an escape in the Canary Islands (Gran Canaria). Escaped Specimens collected
individuals are likely found wherever this species has been Gran Canaria
planted in the past. Santa Lucía de Tirajana, La Sorrueda, 27°53′32.10″N,
With its diameter of 3.5–4 m, Agave franzosinii is one of 15°32′6.48″W, 554 m a.s.l., stony sun-exposed slope at long-
the largest agaves (fig. 1D). Smaller or less glaucous individ- abandoned finca, several individuals in two discrete subpop-
uals can be confused with A. americana. However, A. fran- ulations, 28 Apr. 2018; F. Verloove 13205 (BR) (fig. 2B–C).
zosinii is distinguished by its more intense bluish to almost Santa Lucía de Tirajana bajada a La Sorrueda, Llanos de la
whitish, pruinose leaves with a rough upper surface. The Piedra, 27º53′32.1″N, 15º32′06.6″W, 545 m a.s.l., ambientes
lower portion of the leaf is often patterned with dark green del tabaibal xerófilo-termoesclerófilo, en antiguas terrazas de
where the glaucous covering was rubbed off (fig. 1E). The cultivos, subespontánea, estolonífera, 26 May 2018, Á. Mar-
leaves have large dark teeth along the margins that contrast rero & C. Santiago 37295-37297 (LPA).
a lot with the pale surface (fig. 1F). In addition, leaves are In the wild Agave murpheyi is only known from relatively
much thinner in cross-section and hence very easily bend, es- few localities in central and southern Arizona in the U.S.A.
pecially in the upper third. Finally, A. franzosinii appears to and in northwest Sonora in Mexico (Gentry 1982; Thiede in
have a somewhat laxer inflorescence (fig. 2A). press). It is known only from around pre-Columbian agricul-
The taxon was first described by Sprenger (1885) as A. tural and settlement areas and thought to have been a very
americana var. franzosinii. The description of A. franzosi- important food source (Irish & Irish 2000; Reveal & Hodg-
nii by Sewell (1889), dated 30 Nov. 1889, is seen as a new son 2002). It is in fact much more widely distributed in horti-
combination at species rank based on Sprenger’s A. ameri- culture. In a long-abandoned finca in Santa Lucía de Tirajana
cana var. franzosinii (see Thiede 2017), and antedates the de- in Gran Canaria several individuals persist for many decades
scription by Watson (1889). At Villa Hanbury the plant was in two small subpopulations.
thought to be a hybrid of A. americana with another uniden- Agave murpheyi is characterised by its dark glaucous-
tified species. It was in fact validly described 20 years earlier green, spatulate, acuminate leaves (ca. 63 × 9 cm in mature
by Jacobi (1869), as A. beaulueriana. The correct name of plants in the Canarian population) with light cross-banding
the species would therefore be A. beaulueriana Jacobi (as and a clear bud imprint (fig. 2B). Leaf margins have short
pointed out by Howard 1979), a binomial that is rarely used brownish-reddish teeth that are more numerous in the lower
except perhaps in the Caribbean where the species is fre- half of the blade (fig. 2C). The apical spine is non-decurrent
quently grown as an ornamental (e.g. Wagenaar Hummelinck and short (less than 2 cm long). The species is moderately
1987; Fournet 2002; Proctor & Acevedo-Rodríguez 2005; surculose and produces a lot of bulbils (seed set is very rare).
see also Hochstätter 2015). A proposal to conserve the name From closely similar ornamental agaves like A. chrysantha
A. franzosinii against the name A. beaulueriana was pub- Peebles and A. palmeri Engelm., it is readily separated by
lished (Thiede 2017) in order to foster nomenclatural stabil- its very short terminal spine (Gentry 1982; Thiede in press).
ity. We here adhere to the binomial A. franzosinii following
Although relatively widely available in the horticultural
Recommendation 14A.1. of the ICN applicable to proposals
trade Agave murpheyi had – to our knowledge – not been
for the conservation of names (Turland et al. 2018): “When
recorded before in the wild outside of its native distribution
a proposal for the conservation (Art. 14) or protection (Art.
range.
F.2) of a name has been referred to the appropriate specialist
committee for study, authors should follow existing usage of
names as far as possible pending the General Committee’s Section Rigidae (Baker) R.H.Webb & G.D.Starr (≡ Group
recommendation on the proposal.” Rigidae Baker [“Gentry”]; = Group Sisalanae Trel., incl.

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Pl. Ecol. Evol. 152 (3), 2019

sect. Sisalanae Thiede & Gideon F.Sm.; = Unterreihe Agave angustifolia Haw. (Haworth 1812: 72)
Sisalanae A.Berger)
Figs 2D–F, 3A–B
Plants from this section have linear, ensiform and rigid leaves New to the flora of the Canary Islands – category C3 sensu
with almost straight margins and a usually non-decurrent (or Blackburn et al. (2011) (Individuals surviving in the wild in
rarely thinly decurrent) terminal spine. All have a surculose location where introduced, reproduction occurring, and pop-
habit, greenish-yellow flowers and frequently bulbiliferous ulation self-sustaining).
inflorescences (Gentry 1982; Thiede in press). The group Observation records
includes 14 species that are mainly distributed in Mexico. Gran Canaria
Several are only known from cultivation. Five have been re- Mogán (Las Casillas), road GC-200, 27°52′5.65″N,
corded in the study area; several of these are quite expansive 15°44′12.02″W, 168 m a.s.l., roadside, a single large clone
and can be considered invasive species. with many rosettes, 31 Mar. and 14 Dec. 2017, obs. F. Ver-
This section – and more precisely those species closely loove (fig. 2D–E).
related to or derived from Agave angustifolia – is by far the San Bartolomé de Tirajana, Monte León, 27°48′50.50″N,
15°37′0.99″W, 382 m a.s.l., sun-exposed rocky slope facing
most critical one in the study area. It is composed of Gentry’s
Barranco de Chamoriscán, by the dozen, 22 Dec. 2017 and
Rigidae Group and evidently also includes A. sisalana from
19 Apr. 2018, obs. F. Verloove (fig. 3B).
the Sisalanae Group (Ullrich 1990). All species recorded San Bartolomé de Tirajana, Monte León, Calle Mozart, close
in the area under study are morphologically similar. While to Calle Bach, 27°49′20.14″N, 15°36′54.18″W, 531 m a.s.l.,
their extreme expressions are relatively easily told apart, in- foot of steep slope, a single individual, 19 Apr. 2018, obs. F.
termediate forms are often found. Three of the species here Verloove.
recognised (A. fourcroydes, A. sisalana and A. tequilana) are Mogán, Barranco del Cura, 27°48′42.26″N, 15°43′49.36″W,
sexually sterile clones that were derived from A. angustifo- 81 m a.s.l., rocky slope, a single individual, 25 Apr. 2018,
lia for commercial purposes (mostly for fiber or beverages). obs. F. Verloove. 
They are not known from natural populations. Gentry (1982) Tenerife
already admitted that they are “maintained as species more Güímar (Puertito de Güímar), 28°17′58.74″N,
for taxonomic convenience than founded on basic morpho- 16°22′27.13″W, 17 m a.s.l., long-abandoned plant nursery,
logical and biological knowledge”. Indeed, based on our small population, 11 Jan. 2017, obs. F. Verloove (figs 2F,
field observations all these species seem to be distinguished 3A).
only on quantitative characters. Agave angustifolia tends to El Rosario, Tabaiba Alta, 28°24′39.34″N, 16°20′07.72″W,
have shorter leaves (although large-leaved forms also exist, 348 m a.s.l., in abandoned lot, a solitary flowering individu-
including in the area under study) and a shorter stem than its al, together with Opuntia ficus-indica, Austrocylindropuntia
derivative species but there is considerable overlap. Agave subulata, etc., 3 Jun. 2018, obs. J.A. Reyes-Betancort.
sisalana (var. sisalana) has smooth leaf margins but a form Puerto de la Cruz, La Paz above Camino de La Cos-
with well-developed triangular teeth (A. sisalana var. armata ta, 28°24′58.62″N, 16°31′49.60″W and 28°24′54.57″N,
Trel.) occurs throughout the species’ range of occurrence. 16°31′52.37″W, 71 and 82 m a.s.l., in abandoned lot close
Agave fourcroydes is much reminiscent of A. sisalana (habit, to urbanisations, at least 50 individuals, escaped and natural-
robustness of leaves and stems) and mostly seems to differ ised from plantation, with A. fourcroydes, 1 Aug. 2018, obs.
J.A. Reyes-Betancort; idem, 18 Jan. 2019, obs. F. Verloove
by the constant presence of regularly spaced marginal teeth.
& J.A. Reyes-Betancort.
Similarly, A. tequilana resembles A. fourcroydes and merely
differs in having a shorter terminal leaf spine and a shorter Fuerteventura
stem. La Oliva, Corralejo, 28°43′42.31″N, 13°52′15.20″W, 24 m
In this account we follow traditional species circumscrip- a.s.l.to 28°43′35.50″N, 13°52′17.79″W, 26 m a.s.l., very nu-
merous individuals growing on the abandoned grounds of the
tions, also because combinations at a lower taxonomic rank
former Acua Water Park, 26 May 2018, obs. E. Ojeda-Land.
under the progenitor species are mostly lacking. However, it
is clear that maintaining A. fourcroydes, A. sisalana and A. Agave angustifolia is the most widespread species of the ge-
tequilana at species rank is untenable (Thiede 2016). All are nus and ranges from northern Mexico to Panama. In addition,
it is commonly cultivated as an ornamental and very easily
preferably subsumed under A. angustifolia, either as mere
escapes. It is naturalised in, for instance, Florida (Franck
cultivars of the latter or as varieties or subspecies (Thiede in
2012), South Africa (Steyn & Smith 2000, sub A. vivipara;
press). Walters et al. 2011), India (Drummond & Prain 1906; sub A.
This section was referred to as Group Viviparae by wightii J.R.Drumm. & Prain) and western Australia (Forster
Thiede (2001) since A. angustifolia and A. vivipara were 1996). In Europe it has been recorded in the Valencia area
considered conspecific, based on Wijnands (1983). These and in the Balearic Islands in Spain (Guillot Ortiz et al. 2008;
are, in fact, two very distinct species (García-Mendoza & Sáez et al. 2016) as well as in Italy (Manni 2015).
Chiang 2003) and belong to different sections, section Rigi- This species is extremely variable and counts many in-
dae and Viviparae respectively. The traditional infrageneric fraspecific taxa and synonyms. Gentry (1982) distinguished
classification (Trelease 1913; Berger 1915; Gentry 1982) for six varieties in addition to the nominal variety: var. dew-
these species should therefore be retained. eyana (Trel.) Gentry, var. letonae (F.W.Taylor) Gentry, var.

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Figure 3 – A. Agave angustifolia, Güímar (T), January 2017. This form with variegated leaves (var. marginata) is most frequently cultivated
as ornamental. B. Idem, San Bartolomé de Tirajana (Monte León) (C), December 2017. In this population inflorescences are not bulbiliferous
although they usually are in this species. C. A. fourcroydes, Santa Cruz de Tenerife, Santa Maria del Mar (T), November 2014. Inflorescences
are very bulbiliferous in this species. D. Idem, Güímar (T), January 2017. Detail of leaf spination. E. A. macroacantha, La Oliva (F), May
2018. The blackish spines contrast a lot with the pale foliage. F. A. sisalana, Santa Lucía de Tirajana, Pozo Izquierdo (C), November 2015.
Leaf margins are smooth in the typical variety of this species. Photographs: A–D, F by F. Verloove; E by E. Ojeda-Land.

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Pl. Ecol. Evol. 152 (3), 2019

marginata Hort. ex Gentry, var. nivea (Trel.) Gentry, var. invasive (Acebes Ginovés et al. 2010; Santos-Guerra et al.
rubescens (Salm-Dyck) Gentry and var. sargentii Trel. Most 2013). Category E sensu Blackburn et al. (2011) (Fully in-
of these, however, are mere cultivated selections and culti- vasive species, with individuals dispersing, surviving and
vars and should better be named as such (Thiede in press). reproducing at multiple sites across a greater or lesser spec-
Compared with most other members of this section (except trum of habitats and extent of occurrence).
A. macroacantha; see there), most plants of A. angustifolia Agave fourcroydes is known as a cultivated plant only.
currently found in the wild in the study area are distinctive It is a sexually sterile clone that was probably derived from
in being only medium-sized with less parallel-sided, flatter A. angustifolia. It is cultivated nearly worldwide in tropical
leaves (fig. 2D–F). Even in the tallest mature individuals, regions, either for the production of hard fibre (henequen) or
leaves are rarely more than 60 cm long (see below, how- as an ornamental. Like other members of this group it copi-
ever), much less than in A. fourcroydes, A. sisalana and A. ously produces bulbils on the inflorescence (fig. 3C). These
aff. tequilana. Although bulbils are usually produced in the often already root whilst still attached to the plant and other-
inflorescences of this species, these appear to be absent in the wise quickly do so once dropped to the ground. As a result,
naturalised population in Monte León in Gran Canaria (fig. A. fourcroydes easily establishes itself wherever introduced.
3B). Trelease (1913) already noticed that this species indeed In hilly landscapes like those in the study area it naturalises
sometimes fails to produce bulbils. In ornamental plantings a or even becomes invasive, penetrating natural habitats (lad-
cultivar with white to pale yellow leaf margins is frequently eras, barrancos).
observed (var. marginata Trel.) (fig. 3A). Small populations
of this variety are established in a long-abandoned plant In its most typical form this species is easily distinguished
nursery in Puertito de Güímar in Tenerife as well as on the from Agave angustifolia and A. sisalana. Compared with the
abandoned grounds of the former Acua Water Park in Cor- former it is a much coarser, often gigantic plant with leaves
ralejo in Fuerteventura. The genuinely wild form with un- sometimes attaining 2 m in length. With age it also develops
variegated leaves is much less frequently seen in cultivation. a distinct trunk although this is only evident if lower leaves
Escaped and naturalised plants are only known from Mogán are cut. From A. sisalana it differs in its leaf margins that
(Las Casillas) (fig. 2D–E) and San Bartolomé de Tirajana are distinctly and regularly toothed throughout (fig. 3D). The
(Monte León) in Gran Canaria. separation of A. aff. tequilana and A. fourcroydes in the study
area is much less clear-cut (see under A. aff. tequilana).
Another extreme expression that we refer to this species
is naturalised in several places in Tenerife. It has markedly
longer mid-green leaves up to 100 cm long with regularly Agave macroacantha Zucc. (Zuccarini 1833: 676)
toothed margins. From a distance, especially based on stature Fig. 3E
and leaf length and colour, it is somehow reminiscent of A.
sisalana (var. armata). However, it has a much stronger and New to the flora of Fuerteventura and Tenerife. Previously
longer terminal spine that is thinly but very distinctly decur- reported from La Palma (Otto & Verloove 2018) – category
rent, a feature only known to occur in A. angustifolia in this C1 sensu Blackburn et al. (2011) (Individuals surviving in
section (Gentry 1982). Such plants are locally naturalised in the wild (i.e. outside of captivity or cultivation) in location
an abandoned lot close to a residential area in Puerto de la where introduced, no reproduction).
Cruz (La Paz). Identical plants have also been observed in Observation records
Granadilla de Abona (El Médano, San Isidro) and in Arona Fuerteventura
(Cho). La Oliva, Corralejo, 28°43′42.31″N, 13°52′15.20″W, 24 m
Wijnands (1983) considered Agave angustifolia to be a.s.l. to 28°43′35.50″N, 13°52′17.79″W, 26 m a.s.l., escaping
conspecific with A. vivipara and it was accepted under this from ornamental plantation on the abandoned grounds of the
name by several subsequent students of the genus (e.g. For- former Acua Water Park, 26 May 2018, obs. E. Ojeda-Land
ster 1992; Smith & Steyn 1999a; Thiede 2001). However, (fig. 3E).
García-Mendoza & Chiang (2003) showed that these are in Tenerife
fact two very distinct species with non-overlapping distribu- Güímar, Puertito de Güímar, 28°17′52.07″N, 16°22′31.43″W,
tion ranges. Agave angustifolia is a species from continen- long-abandoned plant nursery, now rough ground, persisting
tal America that ranges from Mexico to Panama whereas A. and clonally reproducing, 3 Dec. 2018, obs. F. Verloove.
vivipara originates in the West Indies (Curaçao, Aruba, Bo- Agave macroacantha is a Mexican endemic that naturally oc-
naire and other islands off the Venezuelan coast). Although curs in Puebla and Oaxaca (Gentry 1982; Thiede in press). It
A. angustifolia is much more frequent as an ornamental (and is commonly grown as an ornamental, in Spain for instance
as an escape), genuine A. vivipara is also available in the or- in Castellón and Valencia provinces (Guillot Ortiz & van der
namental trade and has been introduced outside of its native Meer 2006b). Although frequently cultivated, this species
distribution area, albeit very rarely so (see further). is very rarely reported as an escape. According to Randall
(2017) it has not yet been recorded as a weed outside of its
Agave fourcroydes Lem. (Lemaire 1864: 66) native range. Otto & Verloove (2018) cited a single casual
record from La Palma (Canary Islands). In Tenerife this spe-
Fig. 3C–D cies persists on the grounds of a long-abandoned plant nurs-
Known in the Canary Islands from numerous localities ery, along with A. filifera and other ornamental succulents. In
(none enumerated here) in Fuerteventura, Gran Canaria, La Fuerteventura A. macroacantha was formerly planted as an
Gomera, La Palma, Lanzarote and Tenerife – naturalised and ornamental in the Acua Water park and now reproduces from

480
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suckers in the abandoned area, although less prolifically than taxonomic value and often included in the synonymy of A.
A. angustifolia. In Gran Canaria it also freely reproduces in sisalana by contemporary authors (e.g. Thiede in press).
numerous localities where it has been planted in the past. A Observation record
future naturalization in the Canary Islands is imminent. Lanzarote
Agave angustifolia and A. macroacantha are morpho- Haría, Barranco Chafariz, 29º07′23.53″ N, 13º30′25.91″, 288
logically similar. Agave macroacantha, however, is much m a.s.l., in the transformed bed of the ravine (abandoned cul-
smaller in stature and leaf size and further distinguished by tivated terraces) and on adjacent slopes, ca. 100 individuals,
its blue-gray leaves with dark, nearly black contrasting teeth 24 July 2018, obs. A. Reyes-Betancort (fig. 4A–B).
and spine (fig. 3E). It also has a much stouter terminal spine Agave sisalana Perrine is often wrongly considered to be an
that is c. 10% of the total leaf length. invalid name (e.g. Tropicos 2018). On page 8 Perrine (1838)
Agave macroacantha can sexually reproduce by seeds wrote: “Two varieties … which I take the liberty to christen
and propagate vegetatively by aerial bulbils and ground-lev- Agave Sisalana ...”, indeed without any further description
el basal shoots and rhizomes. Ground-level vegetative off- of the new species. However, on pages 87–88 a detailed de-
spring and bulbils, however, showed much higher survival scription is provided.
rates than seedlings (Arizaga & Ezcurra 2002).
Agave aff. tequilana F.A.C.Weber (Weber 1902: 220–223,
Agave sisalana Perrine (Perrine 1838: 87–88) figs. 1–2)
Figs 3F, 4A–B Fig. 4C–D
Known in the Canary Islands from numerous localities (none New to the flora of the Canary Islands – category C3 sensu
enumerated here) from Fuerteventura, Gran Canaria, La Blackburn et al. (2011) (Individuals surviving in the wild in
Palma, Lanzarote and Tenerife – naturalised and invasive location where introduced, reproduction occurring, and pop-
(Acebes Ginovés et al. 2010; Santos-Guerra et al. 2013). ulation self-sustaining).
Category E sensu Blackburn et al. (2011) (Fully invasive Observation record
species, with individuals dispersing, surviving and reproduc- Gran Canaria
ing at multiple sites across a greater or lesser spectrum of Santa Lucía de Tirajana, La Sorrueda, 27°53′31.61″N,
habitats and extent of occurrence). 15°32′13.77″W, 552 m a.s.l., stony slope, sun-exposed, many
Agave sisalana is known as a cultivated plant only. It is individuals, widely dispersed, 23 Jul. 2016, obs. Á. Marrero
most likely that it was first domesticated in the Yucatan Pen- and 18 Dec. 2017, obs. F. Verloove (fig. 4C–D).
insula in Mexico (Trejo-Torres et al. 2018). It is a sexually Specimens collected
sterile, pentaploid clone that was most likely derived from Gran Canaria
A. angustifolia. It is cultivated nearly worldwide in tropical Santa Lucía de Tirajana, bajada a La Sorrueda, Llanos de la
regions, either for the production of hard fibre (sisal) or as Piedra, 27º53′33.0″N, 15º32′13.0″W, 550 m a.s.l., ambientes
an ornamental. It persists indefinitely in and near abandoned del tabaibal xerófilo-termoesclerófilo, en antiguas terrazas de
plantations. Each plant produces thousands of bulbils that cultivos, subespontánea, estolonífera, 26 May 2018, Á. Mar-
easily drop to the ground and root. The bulbils are in fact rero & C. Santiago 37288-37290 (LPA).
perfectly formed plantlets that are carried far and wide, es- Ibidem, Llanos de la Piedra, 27º53′30.4″N, 15º32′14.6″W,
pecially in hilly habitats like in the study area. Although A. 540 m a.s.l., setos de bordes de bancales de antiguos cultivos
sisalana was merely considered a naturalised introduced spe- en zona termoesclerófila, subespontánea, estolonífera, 29
cies in the Canary Islands (Acebes Ginovés et al. 2010) it has Jun. 2019, Á. Marrero & R. López González 37492-37494
become in fact an invasive species there (at least locally) that (LPA).
penetrates natural habitats, for instance barrancos. Agave tequilana is unknown as a wild species although
Plants of Agave sisalana are usually readily distinguished closely similar plants have been found in the wild in Mexico
from the other members of the section on their unarmed (Gentry 1982). It is cultivated in large plantations in Mexi-
leaf margins (fig. 3F). However, in general habit and from co, especially near the town of Tequila in Jalisco, and is an
a distance it is often indistinguishable from A. fourcroydes. important source of the distilled liquor tequila (Thiede in
It tends to have greener leaves as compared with the more press). At present, it is also more or less widely grown as
glaucous leaves of A. fourcroydes and its flowers are usually an ornamental in climatologically suitable areas. However,
a trifle smaller (55–65 mm vs. 60–75 mm). The existence of it has only exceptionally been recorded as an escape outside
a form with short but well-developed triangular teeth, var. ar- its native area. In Queensland, Australia, it is considered a
mata Trel. (fig. 4A–B), further blurs the boundaries between ‘Non-Declared Weed of Agriculture’ (Randall 2017). In Gran
these two species. Such plants occur sporadically and at ran- Canaria a population with several dozens of individuals is
dom in widely scattered localities, in cultivation but also in naturalised on the grounds of a long-abandoned finca in La
the wild (Gentry 1982). In Europe it has been reported as an Sorrueda (fig. 4C). In this locality the reputable German
escape in Spain (Guillot Ortiz & van der Meer 2006a). In the botanist Günther Kunkel (1928–2007) introduced numer-
Canary Islands a well-naturalised population of A. sisalana ous non-indigenous plants for xero-gardening purposes. The
var. armata was recently discovered on the slopes of a bar- site is abandoned since the 1970’s and suffered from several
ranco in Lanzarote (see below). This taxon had not been re- severe fires. Nonetheless, many species established them-
ported before from the study area. It probably is of limited selves, including cacti like Cylindropuntia imbricata (Haw.)

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Pl. Ecol. Evol. 152 (3), 2019

F.M.Knuth, Opuntia leucotricha DC. and O. microdasys infraspecific taxa, var. ferox and var. salmiana have been re-
(Lehm.) Pfeiff. (Verloove et al. 2017) or agaves like A. lechu- corded in the study area. They are contrasted in the following
guilla and A. murpheyi (this paper). couplet:
The plants naturalised in La Sorrueda most closely re- 1 Leaves usually massive, 1–2 m long, linear-lanceolate
semble Agave tequilana and more or less correspond with (L/W ratio ca. 5/1), often distinctly sigmoid, dull greyish
the plants that are cultivated as ornamentals under this name green, with small or prominent marginal teeth. Flowers
in Europe. However, they are not identical with the plants large, 90–110 mm long................................var. salmiana
from Jalisco (A. Valenzuela-Zapata, Belgium, pers. comm., 1’. Leaves often not exceeding 1 m in length, broadly-oblan-
March 2018). They are much reminiscent of A. fourcroydes ceolate (L/W ratio ca. 3/1), recurved at tip, often a bright-
but are less massive with shorter, distinctly glaucous leaves er, shiny green, with very prominent marginal teeth.
(ca. 107 × 10 cm on average), a shorter terminal spine (10– Flowers smaller, 70–90 mm long...................... var. ferox
20 mm) and inflorescences with 20–25 umbels (rather than
10–18 in A. fourcroydes and A. angustifolia) (fig. 4D). The
material from La Sorrueda also tends to have a shorter stem. Agave salmiana var. salmiana
Another entity that roughly resembles these plants is A. ×ca- Fig. 4F, 5A–C
vanillesii Guillot & van der Meer (Guillot Ortiz & van der
Meer 2004), a putative hybrid of A. decipiens Baker and A. New to the flora of the Canary Islands – category C3 sensu
fourcroydes that spontaneously arose in the Valencia area Blackburn et al. (2011) (Individuals surviving in the wild in
in Spain. It shares many features with A. tequilana, includ- location where introduced, reproduction occurring, and pop-
ing the shorter leaves with smaller marginal teeth and apical ulation self-sustaining). Previous records of ‘Agave ferox’
spine, but it has green leaves (D. Guillot & P. van der Meer, from Gran Canaria (Kunkel 1972) doubtlessly belong here,
Valencia, Spain, pers. comm., May 2018). at least for the most part.
Although the exact identity of the population from La Observation records
Sorrueda remains somewhat uncertain it is included in this Gran Canaria
account because it is obviously well-established in this local- Las Palmas de Gran Canaria, Tafira Alta, Calle Lomo de En-
ity and sufficiently different from Agave fourcroydes to be medio, 28°3′11.76″N, 15°27′20.13″W, 319 m a.s.l., by dirt
treated as a distinct entity. Similar (but identical?) plants also track, on the verge of barranco, scattered individuals amidst
have been observed elsewhere in the study area, for instance numerous A. americana, 6 Dec. 2017, obs. F. Verloove.
in Las Palmas de Gran Canaria (Tafira Alta) and in Grana- Telde, Hornos del Rey, abandoned finca adjacent to Barranco
dilla de Abona (El Médano) in Tenerife. de Las Goteras, 28°2′5.81″N, 15°26′1.93″W, 133 m a.s.l.,
small population, 16 Apr. 2018, obs. F. Verloove.
Section Salmianae (A.Berger) Verloove & Thiede, in Thiede Telde, Camino a La Sima, 28°1′47.46″N, 15°26′1.82″W, 164
et al. (2019: 257) m a.s.l., roadside, scattered individuals, 16 Apr. 2018, obs. F.
Verloove.
Agave [unranked infragen. “Reihe”] Salmianae A.Berger Las Palmas de Gran Canaria, Los Hoyos, Camino Sabro and
Gentry (1982: xi) explicitly stated that his informal groups Cuesta de los Espinos, between 28°3′2.23″N, 15°27′38.66″W
are “tantamount to sections”, and he used the term “group” and 28°2′53.83″N, 15°27′41.25″W, 340–365 m a.s.l., natu-
throughout the taxonomic treatment except for the synoptical ralised from hedges on edges of crops, roads and paths, very
classification on p. 269 given as “Sectional list of species”. common, 20 Mar. 2018, obs. Á. Marrero Rodríguez and 17
Mottram (2015) used the latter page as indication of section- Apr. 2018, obs. F. Verloove (figs 4F, 5A–B).
al rank, albeit otherwise the term “group” is used throughout. Santa Brígida, Camino Las Arenillas, 28°2′6.56″N,
We treat the “sectional list” as erroneous and treat all infor- 15°28′5.64″W, 525 m a.s.l., roadside, few plants, 17 Apr.
mal groupings as informal groups. Thus, a new combination 2018, obs. F. Verloove.
for “Salmianae” was needed which was proposed in Thiede Las Palmas de Gran Canaria, GC-802 road, W slope of
et al. (2019). Pico de Bandama, 28°2′27.59″N, 15°27′33.56″W and
This group includes three species (Thiede in press) and 28°2′26.60″N, 15°27′38.47″W, 410–412 m a.s.l., roadside,
is characterised by large to massive green leaves, a densely numerous plants, 17 Apr. 2018, obs. F. Verloove.
surculose habit and a pyramidal inflorescence with fleshy, Las Palmas de Gran Canaria, GC-800 road at San Francisco
densely imbricate bracts (Gentry 1982). A single variable de Paula, 28°3′21.55″N, 15°27′11.88″W, 359 m a.s.l., one in-
species with two varieties is naturalised in the study area. dividual, 18 Apr. 2018, obs. F. Verloove.
Las Palmas de Gran Canaria, Tafira Baja, Montaña del So-
corro, 28°4′5.36″N, 15°27′11.02″W, 350–380 m a.s.l., hilly
Agave salmiana Otto ex Salm-Dyck (Salm-Dyck 1859: 88) shrubland with Pistacia, Olea, etc., by the dozen, 20 Mar.
Figs 4E–F, 5A–C 2018, obs. Á. Marrero and 24 Apr. 2018, obs. F. Verloove, Á.
Agave salmiana, a Mexican endemic, is a variable species Marrero Rodríguez & M. Salas Pascual.
and includes several infraspecific taxa (Gentry 1982). In ad- Tenerife
dition to the nominal species the following are most notable: Arafo, road down to Arafo, 28°21′32.99″N, 16°25′57.54″W,
subsp. crassispina (Trel.) Gentry (syn.: A. crassispina Trel.) 955 m a.s.l., border of an abandoned crop lot, a single sur-
and var. ferox (K.Koch) Gentry (syn.: A. ferox K.Koch). Two culose clone with old inflorescences, 11 Jul. 2018, obs. J.A.

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Figure 4 – A. Agave sisalana var. armata, Haria (L), July 2018. B. Idem. In this variety leaf margins are minutely denticulate. C. A. aff.
tequilana, Santa Lucía de Tirajana (C), April 2018. D. Idem, December 2017. Detail of leaf spination. E. A. salmiana var. ferox, Los Realejos
(T), July 2018. F. A. salmiana var. salmiana, Las Palmas de Gran Canaria (Los Hoyos) (C), August 2018. Compared with A. americana
leaves are greenish, not glaucous. Photographs: A, B, E by A. Reyes-Betancort; C, D by F. Verloove; F by M. Salas Pascual.

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Pl. Ecol. Evol. 152 (3), 2019

Reyes-Betancort (fig. 5C); idem, 04 Dec. 2018, obs. F. Ver- a naturalised escape in South Africa (Smith & Figueiredo
loove. 2012a).
Specimen collected Records of Agave atrovirens Karw. ex Salm-Dyck in
Gran Canaria Europe (Webb 1980) turned out to refer to A. salmiana var.
Las Palmas de Gran Canaria, Tafira Baja, Montaña del So- salmiana (Smith & Figueiredo 2007). However, the identity
corro, 28º04′02.1″N, 15º27′10.8″W, 380 m a.s.l., cono vol- of the plant naturalised under this name in Madeira requires
cánico con vegetación termoesclerífila del acebuchal y len- further study. According to Vieira (2002) it is highly bulbilif-
tiscal, subespontánea, estolonífera, 27 Jun. 2019, Á. Marrero erous, a feature compatible with neither A. atrovirens nor A.
37481-37482 (LPA). salmiana var. salmiana.
Agave salmiana var. salmiana is endemic to Mexico where
many forms are cultivated for the tapping of agua miel for Agave salmiana var. ferox (K.Koch) Gentry (Gentry 1982:
the production of pulque, a fermented, lightly alcoholic bev- 611–612)
erage. In other parts of the world it is widely grown as an
ornamental. It is particularly common in parts of the Medi- Fig. 4E
terranean basin. A. ferox K.Koch (Koch 1860: 23). –A. salmiana subsp. ferox
In stature and habit A. salmiana var. salmiana is much (K.Koch) Hochstätter (Hochstätter 2015: 23). – Type: La
reminiscent of forms of A. americana and these taxa certain- Mórtola, Mr. Hanbury’s garden, 2 and 18 Jul. 1896, Anon.
ly have been confused by local botanists. Both (and especial- s.n. (K).
ly the latter) are variable and have several features in com- New to the flora of Tenerife. Previous records of ‘Agave fer-
mon: a heavily surculose habit, massive, ± curved leaves, ox’ from Gran Canaria (Kunkel 1972) doubtlessly belong to
etc. Non-flowering individuals of A. salmiana var. salmiana var. salmiana, at least for the most part – category C1 sensu
can be separated based on leaf colour: leaves are dull green Blackburn et al. (2011) (Individuals surviving in the wild
(fig. 4F) while they are always glaucous in A. americana. (i.e. outside of captivity or cultivation) in location where in-
Typically, leaves are also sigmoidally curved (fig. 4F). The troduced, no reproduction).
inflorescence pole, however, is most characteristic: it is hard- Observation records
ly longer than wide and more or less pyramidal in outline Tenerife
with the longest branches located in the lower half of the Los Realejos, La Montañeta, in the SSW slope of the vol-
pole (fig. 5A). Peduncular bracts tend to be very prominent cano, 28°23′27.72″N, 16°33′36.54″W, 308 m a.s.l., gravelly
and fleshier in texture (fig. 5B). The naturalised plants found slope, 3 Jun. 2018, obs. J.A. Reyes-Betancort (fig. 4E); idem,
in Gran Canaria and Tenerife are fairly uniform and further 18 Jan. 2019, obs. F. Verloove & J.A. Reyes-Betancort.
differ from var. ferox in being usually massive with much Tegueste, road down to El Socorro, 28°30′34.37″N,
narrower leaves up to or even exceeding 200 cm in length 16°21′37.35″W, 342 m a.s.l., ravine slope, 3 Jun. 2018, obs.
with long-acuminate sigmoidal tips and usually (not always) J.A. Reyes-Betancort; idem, 18 Jul. 2018, obs. E. Ojeda-
less pronounced marginal teeth. Land; idem, 18 Jan. 2019, obs. F. Verloove.
In Gran Canaria Agave salmiana var. salmiana is rare Compared with var. salmiana this variety usually has a more
although it is locally naturalised in relative abundance. It is urn-shaped (urceolate) rosette with smaller (70–90 × 23–30
most frequent near Los Hoyos, exactly the place from where cm), broadly oblanceolate, outcurving and often more shiny,
‘Agave ferox’ was reported by Kunkel (1972). There is no greener leaves with marginal teeth on more prominent protu-
doubt that Kunkel also saw A. salmiana var. salmiana and berances (fig. 4E). It is easily recognisable but of uncertain
that this species is naturalised for half a century or longer systematic status (Thiede in press). At present, A. salmiana
in this area. All recently observed populations are from the var. ferox is more frequently grown in private or public gar-
northeastern part of the island and often obviously refer to dens or as a roadside ornamental in the Canary Islands than
relics of former cultivation (near old fincas, etc.). At present, var. salmiana. On several occasions young plants have been
however, var. salmiana is only exceptionally grown as an observed close to the planted individuals, doubtlessly a result
ornamental in Gran Canaria, var. ferox being more frequent of its surculose habit, but genuinely wild plants have rarely
now in the horticultural trade. In Tenerife A. salmiana var. been recorded so far, and not at all in Gran Canaria. This is
salmiana only occurs in a single locality in Arafo, in a rocky, of interest since it was previously reported from Gran Ca-
arid upland locality, at an altitude of nearly 1,000 m. naria and La Gomera in the study area (Kunkel 1972, 1975).
Agave salmiana var. salmiana is a frequent ornamental However, all records from Gran Canaria that could be veri-
in parts of the Mediterranean area and in climatologically fied belong to var. salmiana (see before).
similar areas elsewhere in the world. It readily escapes and In Tenerife A. salmiana var. ferox has recently been re-
is reportedly naturalised in several countries, for instance in corded on two occasions. In Los Realejos and in Tegueste it
southern France (Tison & de Foucault 2014), Italy (Celesti is escaping from an ornamental plantation. Obviously self-
Grapow et al. 2010), Portugal (Smith & Figueiredo 2007; sown plants have established themselves in suitable habitats
Silva et al. 2015) and Spain (Guillot Ortiz & van der Meer close to the plantations. In the first locality A. salmiana var.
2005, 2008a; Guillot Ortiz et al. 2008; Peña & Sánchez ferox is spreading on the slope of the volcano (together with
2016). A. salmiana var. salmiana is considered an invasive the similarly spreading A. americana var. marginata) where-
species in Sardinia (Podda et al. 2012). It is also known as as in Tegueste it is colonising the slope and the dried-out riv-

484
 Verloove et al., Feral Agave and Furcraea in Canary Islands

erbed of a barranco adjacent to the road where it had been viduals, 12 Apr. 2018, obs. Á. Marrero; idem, 18 Apr. 2018,
planted in the past. obs. F. Verloove (fig. 5 D-F).
Agave salmiana var. ferox has naturalised in the Medi- Specimen collected
terranean area, although often less commonly so than var. Gran Canaria
salmiana. It is well known from Spain and the Balearic Is- Las Palmas de Gran Canaria, Tafira Baja, entornos del Cam-
lands where it has been recorded on several occasions (Guil- pus; 28º04′20.8″N, 15º27′07.1″W; 315 m a.s.l., bordes de zo-
lot Ortiz et al. 2008). Still in the Iberian Peninsula its pres- nas ajardinadas, subespontánea, estolonífera; 27 Jun. 2019,
ence has been reported from Portugal (Smith & Figueiredo Á. Marrero, 37474-37475 (LPA).
2007) whereas Podda et al. (2012) consider it as naturalised Agave vivipara is the first-named Caribbean species of the
in Sardinia. genus. It was originally restricted to the Leeward Islands
(Aruba, Bonaire, Curaçao and Margarita) (Thiede in press).
Section Viviparae (Baker) Verloove & Thiede, in Thiede et In recent years, however, it apparently was introduced in the
al. (2019: 260) horticultural trade. In Gran Canaria it has been relatively
frequently planted as an ornamental for several years. In
Agave [unranked infragen. “Group”] Viviparae Baker recent years escaped individuals were seen in scattered lo-
calities where A. vivipara was formerly planted. The species
A new name for this assemblage (Group Vicinae Thiede)
produces numerous bulbils (fig. 6B) and these easily drop
was introduced by Thiede (2001) since the type of Group
to the ground and root. A future naturalisation (as for other
Viviparae (Agave vivipara) was thought to be conspecific
species with a similar biology, especially those from section
with A. angustifolia (based on Wijnands 1983). These are,
Rigidae) is very likely. Agave vivipara also reproduces from
however, quite different species that belong to separate sec-
suckers. The latter are usually produced only after flowering
tions (García-Mendoza & Chiang 2003). Their traditional in-
whereas in the vegetative phase plants often appear solitary
frageneric classification (Trelease 1913; Berger 1915; Gentry
(figs. 5D, 6A).
1982) should be retained.
This species is easily recognisable. Its rosette is less than
Section Viviparae includes eight species from northern 100 cm in diameter and 50 to 60 cm high. The leaves are all
South America (Venezuela and Colombia) and the Leeward abruptly tapered towards the apex, canaliculate and curved
Islands (Trelease 1913; Wagenaar Hummelinck 1993; Thiede outwards (often distinctly sigmoidal) (figs 5F, 6A). They are
in press). Plants are always surculose (at least after flower- broadly lanceolate, widest at about mid length and narrowed
ing). Leaves are fleshy (not hard), green to slightly glaucous, above base to ca. ½ leaf width; they are on average 50 × 15
very broadly lanceolate and curved (sigmoid) with relatively cm (ca. 3.5 × as long as wide). Leaf margins are closely and
short, regularly spaced marginal teeth. Inflorescences are regularly toothed with short teeth (ca. 4–5 mm long) on lu-
freely bulbiliferous (Trelease 1913; Álvarez de Zayas 1995). nate bases (fig. 5F). The apical spine is ca. 25 mm long. The
In the Caribbean species this combination of features only colour of the plant is green to glaucous, sometimes slightly
occurs in section Viviparae. A single species is increasingly pruinose (fig. 6A). The inflorescence is relatively short (ca.
cultivated as an ornamental in the study area and easily es- 3 m tall) and flowers are bright yellow (fig. 5E). In the Ca-
capes. narian plants leaf marginal teeth are more closely placed
than usually recorded. This may indicate introgression or
Agave vivipara L. (Linnaeus 1753: 323) even hybridisation. However, this character also seems to
be very variable in autochthonous populations as shown in
Figs 5D–F, 6A–B a specimen collected in Curaçao (Willemstad, March 1913,
New to the flora of the Canary Islands – category C1 sensu N.L.Britton & J.A.Shafer 3060 U.1048296, herb. U).
Blackburn et al. (2011) (Individuals surviving in the wild Trelease (1913) described the very similar Agave vicina
(i.e. outside of captivity or cultivation) in location where in- Trel. from Aruba. It was differentiated by its dull green (vs.
troduced, no reproduction). transiently glaucous) leaves with heavily triangular marginal
Observation records teeth from large lunate bases (vs. slender teeth from small
Gran Canaria lunate bases). Also, marginal teeth are slightly less closely
Las Palmas de Gran Canaria, Parque Central, 28°6′2.54″N, placed in A. vicina than in A. vivipara. According to Wage-
15°26′45.10″W, 171 m a.s.l., escaping from ornamental plan- naar Hummelinck (1993) and Álvarez de Zayas (1995), A.
tation, 15 Mar. 2012 and onwards, obs. M. Salas Pascual. vicina is best considered conspecific with A. vivipara, a point
Las Palmas de Gran Canaria, San José del Álamo, of view with which we disagree (Thiede in press). Nonethe-
28°4′54.02″N, 15°30′12.59″W, 423 m a.s.l., 5 Apr. 2016, less, it is likely that far too many species of Agave have been
obs. M. Salas Pascual. described in the past in the Caribbean. For instance, in the
Las Palmas de Gran Canaria, Campus Facultad Medicina, Lesser Antilles Rogers (2000) reduced the number of species
San Cristóbal, 28°4′32.67″N, 15°24′59.19″W, 19 m a.s.l., es- from 12 to a single one, A. karatto Mill.
caping from ornamental plantation, Mar. 2018, obs. M. Salas The identification of A. vivipara was not straightforward.
Pascual. Canarian plants were initially thought to be Agave desme-
Las Palmas de Gran Canaria, Lomo de la Cruz, 28°5′28.81″N, tiana Jacobi, a species that is unknown in the wild but fre-
15°26′51.07″W, 207 m a.s.l., on top of slope along GC-23 quently cultivated as an ornamental. In general appearance
motorway, escaped from ornamental plantation, several indi- it closely resembles A. vivipara but its leaf margins are ei-

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Pl. Ecol. Evol. 152 (3), 2019

Figure 5 – A. Agave salmiana var. salmiana, Las Palmas de Gran Canaria (Los Hoyos) (C), August 2018. The inflorescence in this species
is hardly longer than wide and more or less pyramidal. B. Idem, April 2018. Inflorescence bracts in this species are triangular and fleshy. C.
Idem, Arafo (T), July 2018. In this species leaves are green with a long-decurrent terminal spine. D. A. vivipara, Las Palmas de Gran Canaria,
Lomo La Cruz (C), April 2018. E. Idem, May 2018. Flowers of this species are bright yellow. F. Idem, April 2018. Detail of leaf spination.
Photographs: A by M. Salas Pascual; B, D, F by F. Verloove; C by A. Reyes-Betancort; E by Á. Marrero.

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 Verloove et al., Feral Agave and Furcraea in Canary Islands

Figure 6 – A. Agave vivipara, Arucas (C), April 2018. Leaves are abruptly narrowed towards base. B. Idem. Inflorescences are bulbiliferous.
C. A. lechuguilla, La Aldea de San Nicolás (C), April 2018. D. Idem. The inflorescence of this species is spicate, not racemose. Bracts
are linear. E. A. oteroi, Santa Lucía de Tirajana (C), April 2018. F. A. attenuata, Firgas (Cambalud) (C), April 2018. The arching, spicate
inflorescence is very characteristic and leaf margins are unarmed. Photographs: A–F by F. Verloove.

487
Pl. Ecol. Evol. 152 (3), 2019

ther smooth or only have weak or small teeth near the leaf cultivos, subespontánea, estolonífera, 26 May 2018, Á. Mar-
base. In addition, its leaves are longer (50–80 cm) and have a rero & C. Santiago 37291-37294 (LPA).
larger length to width ratio. Also, its flowers are a paler yel- La Aldea de San Nicolás (Tocodomán), 27°57′34.84″N,
low (Irish & Irish 2000; Thiede in press). It is locally natural- 15°46′52.38″W, 296 m a.s.l., on top of stony slope, a small
ised in Florida (Franck 2012). Agave wercklei F.A.C.Weber colony, 25 Apr. 2018, F. Verloove 13231 (BR) (fig. 6 C–D).
ex Wercklé, a species from Costa Rica, is also similar but it A native of the southern U.S.A. (New Mexico, Texas) and
is non-surculose, has a much taller inflorescence with very Mexico, Agave lechuguilla is a characteristic species of the
short peduncle, paler, rough (at least when young) leaves Chihuahuan Desert. It is restricted to desert habitats and pre-
with marginal teeth not on distinct protuberances (García- fers limestone soils. It is readily recognisable by its ‘spicate’
Mendoza & Lott 1994). It has been recorded as an escape inflorescence, widely suckering habit and narrow, deeply
in South Africa (Smith & Steyn 2002; Walters et al. 2011). convex leaves with down-slanted teeth on straight margins
Finally, A. cocui Trel. is also very similar and in fact the (Thiede in press) (fig. 6C–D).
only species of those mentioned here that is closely related Two small populations were recently detected in Gran
to A. vivipara (both belong to section Viviparae). It is na- Canaria. In La Sorrueda it grows on the grounds of a long-
tive to Venezuela and Colombia and also widely occurs in abandoned finca, whereas in Tocodomán a few individuals
the Leeward Islands where it probably is a naturalised intro- at the beginning of flowering (and with some remains of in-
duction (Wagenaar Hummelinck 1936, 1993). It is a coarser, fructescences of the previous year) were noticed close to the
non-surculose species with slightly longer leaves and a much Cactualdea cactus park. The plants grow on top of a slope
taller inflorescence reaching 5–10 m in length (Thiede in with various kinds of debris and garden waste and most like-
press). It is, however, not always clearly separated from A. ly can be considered as having become established from dis-
vivipara (Wagenaar Hummelinck 1938) and may be a taller, carded garden material.
more robust expression of it. To our knowledge and although widely grown as an or-
To our knowledge, true Agave vivipara is very rarely seen namental, Agave lechuguilla has only exceptionally been
outside of its native distribution area. It was recorded from reported in the wild outside of its native distribution range.
South Africa (Walters et al. 2011; also Smith & Steyn 1999b, Guillot Ortiz & van der Meer (2005) cite a single record
sub A. decipiens Baker) but the records require confirmation. from El Saler (Valencia) in Spain. Although further records
from northeastern Spain (Maranges 2011; Aymerich & Gus-
tamante 2016) have been considered erroneous (Mesquida et
Subgenus Littaea (Tagl.) Baker al. 2016), its presence was confirmed lately from Tarragona
Inflorescence often dense, elongated panicles with ± short- province (Aymerich 2017). Holm et al. (1979) and Randall
stalked partial inflorescences mostly with paired or few clus- (2017) report it as a weed in North America.
tered flowers (‘spicate’). The morphologically similar and related A. difformis
A.Berger was recently also recorded in Spain (López-Pujol
et al. 2016). This is usually a more robust species with a
Section Heteracanthae Salm-Dyck (= Group Marginatae much longer inflorescence, wider and longer leaves (often
Baker [“Gentry”]) with more or less sinuous margins) and with marginal teeth
Species from this section have leaves with conspicuous horny less frequently down-slanted. Both species, however, are
margins and moderate to strong marginal teeth. The section very polymorphic and they are known to intergrade (Gen-
accommodates 22 species in Mexico, the southern U.S.A. try 1982). The plants currently known from Gran Canaria
(New Mexico, Texas) and Guatemala (Thiede in press). Two have relatively short leaves: they are on average ca. 35 cm
species are known to occur as escapes in the study area. long and 2–3 cm wide in flowering individuals. Leaf mar-
gins are straight and all teeth are deflected (fig. 6C). Inflo-
rescences are small, not exceeding 2 m in length. All these
Agave lechuguilla Torr. (Torrey 1859: 213–214) characters point at A. lechuguilla. Agave funkiana K.Koch
Fig. 6C–D & C.D.Bouché and A. lophantha Schiede ex Kunth (syn.:
A. univittata Haw.; see, however, Smith et al. 2018) are also
New to the flora of the Canary Islands – category C1 sensu related species that were found as escapes in Spain (Guillot
Blackburn et al. (2011) (Individuals surviving in the wild Ortiz & van der Meer 2008b, 2013b). Both are coarser spe-
(i.e. outside of captivity or cultivation) in location where in- cies with wider and longer leaves and more densely flowered
troduced, no reproduction). inflorescences. The latter further differs in having sinuous to
Specimens collected undulate leaf margins with at least part of the teeth double set
Gran Canaria (Gentry 1982).
Santa Lucía de Tirajana, La Sorrueda, 27°53′31.61″N, Agave lechuguilla was originally described as ‘Agave le-
15°32′13.77″W, 552 m a.s.l., stony slope, sun-exposed, sev- cheguilla’ by Torrey (1859) and this spelling is still adopted
eral individuals, widely dispersed, 28 Apr. 2018, F. Verloove by databases such as Tropicos (Tropicos 2018). The specific
13239 (BR). epithet, however, refers to ‘lechuguilla’, a common name in
Santa Lucía de Tirajana, bajada a La Sorrueda, Llanos de la Chihuahua meaning “small lettuce”. Engelmann (1875) al-
Piedra, 27º53′33.1″N, 15º32′08.2″W, 550 m a.s.l., ambientes ready considered this to be a misprint and its spelling was
del tabaibal xerófilo-termoesclerófilo, en antiguas terrazas de corrected by Gentry (1982).

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Agave oteroi G.D.Starr & T.J.Davis (Starr & Davis 2019: Observation records
134–136) Tenerife (selected records)
Tegueste (El Socorro), Barranco de Las Cuevas, dry gravelly
Fig. 6E
riverbed, 27 Jun. and 5 Nov. 2014, 13 Jan. 2017, obs. F. Ver-
New to the flora of the Canary Islands – category C1 sensu loove.
Blackburn et al. (2011) (Individuals surviving in the wild Candelaria, Barranco de Aroba, 28°21′37.11″N,
(i.e. outside of captivity or cultivation) in location where in- 16°22′8.02″W, 29 m a.s.l., dry, gravelly riverbed, several in-
troduced, no reproduction). dividuals in close proximity (looks ± established), 11 Nov.
Observation record 2014, obs. F. Verloove.
Gran Canaria Palm-Mar, rough ground near habitations, 21 Feb. 2015, obs.
Santa Lucía de Tirajana, La Sorrueda, 27°53′17.15″N, F. Verloove.
15°32′3.77″W, 481 m a.s.l., stony roadside, a single individ- Arona (La Camella towards Valle de San Lorenzo), roadside
ual, 28 Apr. 2018, obs. F. Verloove (fig. 6E). slope, 21 Jun. 2016, obs. F. Verloove.
Gentry (1982) described Agave titanota from Rancho Tam- Santa Cruz de Tenerife, Barranco de Santos near La Ermita,
bor and for quite a long time it was only known in the wild slope of ravine, 13 Nov. 2016, obs. F. Verloove.
from the type locality in northern Oaxaca in Mexico. In Granadilla de Abona (La Tejina near Urbanización Sotaven-
recent years it was shown to be more widespread in north- to), by track, 20 Nov. 2016, obs. F. Verloove.
eastern Oaxaca (Pilbeam 2013), and it was also collected in Granadilla de Abona (San Isidro), Calle Imade, on the verge
southernmost Puebla (García-Mendoza 2011). In the horti- of dry riverbed, 20 Nov. 2016, obs. F. Verloove.
cultural trade, however, A. titanota was widely distributed Arona (Chayofa), Barranco del Verodal, 22 Nov. 2016, obs.
from very soon after its discovery. The same applies to simi- F. Verloove.
lar-looking plants found in 1984 by Felipe Otero in the Sierra Güímar, abandoned dump alongside TF-28, 11 Jan. 2017,
Mixteca and distributed under his collection number FO- obs. F. Verloove.
076. Several forms and cultivars are offered for sale in the Santa Cruz de Tenerife, La Cuesta, 28°27′37.22″N,
horticultural trade (see Guillot Ortiz & van der Meer 2014 16°16′57.61″W, 241 m a.s.l., a clone on the rocky slope of
for an overview). the road, probably escaped from a garden, 19 Apr. 2018, obs.
Agave titanota is characterised by its ovate, spatulate to E. Ojeda-Land.
broadly lanceolate, glaucous white or blue-white leaves with La Victoria de Acentejo, Risco La Sardina at Urbanization
large, irregular, whitish-grey teeth, the terminal spine form- La Palmita, 28°26′21.49″N, 16°28′55.28″W, 167 m a.s.l.,
ing a broad robust plate on the lower face. The only plant hillside near abandoned farms, a clone with Hylocereus un-
seen in the wild in the study area differs from Gentry’s origi- datus, 25 Apr. 2018, obs. E. Ojeda-Land.
nal description, for instance in leaves not being alabaster- or In the Canary Islands this ornamental from Central Mexico
glaucous-white-coloured. They are referable to what was had previously been recorded from Fuerteventura, Gran Ca-
long known as ‘Agave FO-076’ or Agave ‘Felipe Otero’ and naria and La Palma. In recent years it has been increasingly
were recently described as a separate species, A. oteroi (Starr found on these islands and it is here reported for the first time
& Davis 2019). It differs from A. titanota by its green, most- from Tenerife as well. In most instances the occurrences con-
ly obovate leaves, broader woody margin which is extreme- sist of solitary individuals and Agave attenuata is definitely
ly pronounced below the terminal spine on the back of the much less expansive than many of its congeners. In some
leaves (fig. 6E). Both species are also separated geographi- places in Tenerife, however, small but apparently established
cally and inhabit different substrates (Starr & Davis 2019). populations were detected (including flowering individuals),
for instance in a ravine in Candelaria (Barranco de Aroba) or
in La Victoria de Acentejo where it grows on the cliff that is
Section Inermes Salm-Dyck (≡ Group Amolae Gentry nom.
part of the Protected Natural Area of La Costa de Acentejo.
inval. Art. 37.1.; Turland et al. 2018)
A future naturalisation on a wider scale is not unlikely. Also,
This section includes species with unarmed, non-filiferous in Gran Canaria A. attenuata was detected in numerous addi-
soft leaves and inflorescences with densely arranged flow- tional localities in recent years. At least in some of these, for
ers. Ten species are usually recognised and all are endemic instance on the slopes of a ravine in Arucas (Santidad), small
to Mexico, mainly occurring in the Sierra Madre Occidental but apparently established populations were seen with flow-
(Thiede in press). A single species has been observed in the ering and fruiting individuals. Although very widespread in
study area. cultivation A. attenuata has a very limited native distribution
range that remained unknown for quite a long time (Chazaro
et al. 1998).
Agave attenuata Salm-Dyck (Salm-Dyck 1834: 303)
Agave attenuata is widely cultivated in warm-temperate re-
Fig. 6F gions of the world and increasingly reported as escaping,
New to the flora of Tenerife. Previously reported from also in natural habitats. It is known, for instance, from the
Fuerteventura, Gran Canaria and La Palma (Brandes & Kruger National Park in South Africa (Foxcroft et al. 2008),
Fritzsch 2002; Verloove 2013; Otto & Verloove 2016) – cat- Italy, including Sardinia and Sicily (Podda et al. 2012; Galas-
egory C2 or C3 sensu Blackburn et al. (2011) (Individuals so et al. 2018) and many parts of Australia (Randall 2017).
surviving in the wild in location where introduced, reproduc- In Macaronesia it is also known from Madeira (Vieira 2002;
tion occurring, population ± self-sustaining or not). Borges et al. 2008) where it is locally naturalised.

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Figure 7 – A. Agave filifera, Güímar (T), December 2018. Leaf margins unroll as hair-like threads. B. Furcraea foetida, Adeje, Playa de las
Americas (T), November 2016. Leaf margins are either smooth or prickly only in the lower half of the blade. C. F. hexapetala, San Cristóbal
de La Laguna (Valle de Guerra) (T), August 2014. D. Idem. Detail of leaves. E. F. selloana, Adeje, Playa de las Américas (T), March 2016.
Nearly all plants seen in the study area belong to a variegated form, var. marginata. Leaf margins are spiny throughout and the red spines
contrast a lot with the pale greenish blade. F. Idem, Fasnia (T), December 2018. In Furcraea, contrary to Agave, flowers are pendent and
tepals whitish. Photographs: A, B, E, F by F. Verloove; C, D by A. Reyes-Betancort.

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 Verloove et al., Feral Agave and Furcraea in Canary Islands

Section Littaea (Tagl.) Bentham (= Group Filiferae Baker 1’. Leaves wider, distinctly narrowed above base and with
[“Gentry”]) a lesser L/W ratio, usually paler in colour (often varie-
gated)...............................................................................2
This section includes species with unarmed, filiferous, white
bud-printed leaves (Gentry 1982). Eight species are usually 2. Leaves with smooth margins or occasionally with a few
recognised and all are endemic to Mexico, mainly occurring hooked teeth towards the base, the lower surface smooth,
in the Sierra Madre Occidental (Thiede in press). A single green or variegated. Stem usually absent or very short ....
species has been observed in the study area. ........................................................................... F. foetida
2’. Leaves with margins spiny throughout, the lower surface
predominantly rough, nearly always variegated (in the
Agave filifera Salm-Dyck (Salm-Dyck 1834: 309) study area). Stem distinct at maturity, up to 1.5 m long ...
Fig. 7A .........................................................................F. selloana
New to the flora of the Canary Islands – category C1 sensu
Blackburn et al. (2011) (Individuals surviving in the wild Furcraea foetida (L.) Haw. (Haworth 1812: 73)
(i.e. outside of captivity or cultivation) in location where in-
Fig. 7B
troduced, no reproduction).
F. gigantea Vent. (Ventenat 1793: 65). Type – not designated
Specimen collected (?).
Tenerife
Güímar, Puertito de Güímar, 28°17′52.07″N, 16°22′31.43″W, [According to Wagenaar Hummelinck (1987) Furcraea
14 m a.s.l., long-abandoned plant nursery, now rough ground; gigantea is a synonym of Agave scheuermaniana Trel.,
persisting and clonally reproducing, 3 Dec. 2018, F. Verloove a species related to and probably conspecific with Agave
13430 (BR) (fig. 7A). karatto. Furcraea foetida and A. scheuermaniana in fact
look quite dissimilar.]
Agave filifera is an endemic of Mexico where it naturally
occurs in the states of Aguascalientes, Jalisco, Guanajuato, New to the flora of Gran Canaria. Previously reported from
Querétaro, Hidalgo and México. It grows on rocks, in desert Fuerteventura, La Palma and Tenerife (Acebes Ginovés et al.
scrub, thorn, oak and oak-juniper forests, at high altitudes 2010; Santos-Guerra et al. 2013) – category E sensu Black-
ranging between 2,340 and 3,100 m (Thiede in press). It is burn et al. (2011) (Fully invasive species, with individuals
well known in European horticulture as a clonal species, dispersing, surviving and reproducing at multiple sites across
stemming from only few introductions (Gentry 1982). a greater or lesser spectrum of habitats and extent of occur-
In the area under study, it is the only species with leaf rence).
margins that unroll as hair-like threads (fig. 7A). The plants Observation records
found in Tenerife grow in large clumps of small green dense Gran Canaria
rosettes with many thick, slightly curved leaves, resembling Santa Brígida, El Tejar, 28°2′52.71″N, 15°29′40.05″W, 476
a cultivar named ‘Prolifera’. The species grows in several m a.s.l., 18 Dec. 2012, obs. M. Salas Pascual.
patches on the grounds of a long-abandoned plant nursery Valsequillo, Valle de San Roque, 28°0′27.29″N,
where it prolifically reproduces, although probably only 15°28′2.55″W, 349 m a.s.l., escaping from ornamental plan-
clonally. tation, 26 May 2017, obs. M. Salas Pascual.
Agaete, valle de Agaete, Los Pasitos, 28°4′28.21″N,
Agave filifera is much cultivated as ornamental but has 15°39′59.55″W, 263 m a.s.l., dry riverbed, a single individu-
rarely been reported as an escape so far. It is known as an al, 10 Dec. 2017, obs. F. Verloove.
alien species in Algeria (Randall 2017) and Italy in Calabria Santa Brígida, cuesta Caraballo near Calle El Drago,
(Musarella et al. submitted). 28°1′49.60″N, 15°28′14.09″W, 530 m a.s.l., alongside trail, a
single individual, 17 Apr. 2018, obs. F. Verloove.
Furcraea Vent. (Ventenat 1793: 65) Santa Lucía de Tirajana, La Sorrueda, 27°53′32.39″N,
15°32′6.19″W, 549 m a.s.l., abandoned finca, 21 Apr. 2018,
All species of Furcraea currently found in the wild in the obs. F. Verloove.
study area belong to section Furcraea. Stems are either ab-
Tenerife (invasive: see comments below)
sent or short (0.7–1.5 m), leaf margins have conspicuous
Parque Rural de Anaga, between La Fortaleza (near
teeth or are smooth, bulbils are bracteate or foliose and seed-
to Catalanes) and la Hoya Las Colmenas (for instance
lings have small cotyledons. Species of section Serrulatae
at 28°31’10.17”N, 16°15’34.95”W; 28°31’10.12”N,
Drumm. are mostly arborescent with conspicuous stems (1–9 16°15’34.15”W; 28°31’12.29”N, 16°15’39.14”W;
m), leaf margins are closely minutely denticulate, bulbils fo- 28°31’15.08”N, 16°15’44.81”W), ranging in altitude be-
liose, with chartaceous outer leaves and seedlings have large tween 600 and 700 m a.s.l., invading slopes of the basins that
cotyledons (Thiede in press). flow into El Tomadero and El Barranco of Tahodio, along
1. Leaves linear-lanceolate, ca. 10–20 × as long as wide with Agave americana, 20 Apr. 2010, obs. E. Ojeda-Land.
and of more or less uniform width (i.e., not pronouncedly In the lower part of the Barranco de Tahodio, from Hoya de
sword-shaped), dull green in colour, very rigid and radi- La Cantina, 28°30’30.40”N, 16°16’3.99”W, 250 m a.s.l.,
ately spreading.............................................F. hexapetala to 28°30’25.09”N, 16°15’58.14”W, 190 m a.s.l., dozens of

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plants in the bed and on ravine slopes, 8 Oct. 2010, obs. E. Observation record
Ojeda-Land. Tenerife
Buenavista del Norte, Camino El Rincón, 28°22’6.05”N, San Cristóbal de La Laguna, Valle de Guerra, Finca Isa-
16°51’52.42”W, 65 m a.s.l., together with Furcraea selloa- mar, 28°30′36.90″N, 16°23′11.66″W and 28°30′38.45″N,
na, Agave sisalana and A. attenuata, 1 Mar. 2017, obs. E. 16°23′18.14″W, initially cultivated as a curiosity, now local-
Ojeda-Land. ly spreading by bulbils, 3 Jun. 2018, obs. J.A. Reyes-Betan-
Furcraea foetida is a very variable species. Plants seen in cort (fig. 7 C–D).
the wild in the study area range from fairly small to mas- Specimen collected
sive, with either fresh bright green to creamy or variously Tenerife
variegated leaves. Compared with the similar F. selloana it is San Cristóbal de La Laguna, Valle de Guerra, Finca Isamar,
distinguished by its smooth (not rough) leaves with unarmed 28°30′38.45″N, 16°23′18.14″W, 18 Jan. 2019, F. Verloove &
margins or at most with a few marginal teeth in the lower J.A. Reyes-Betancort 13456 (BR).
half (fig. 7B). The occasional presence of such marginal Furcraea hexapetala is a native species in Cuba with the
teeth is sometimes suggested to be the result of introgression type from La Havana. It probably also occurs as a native
or hybridisation (Fontaine 2016). Originally, F. foetida was species elsewhere in the Caribbean and Antilles, for instance
described as having entirely smooth leaf margins (“foliis in- in Bahamas, Bermuda, Hispaniola and Jamaica (Álvarez de
tegerrimis”, sub Agave foetida L.) (Linnaeus 1753). Zayas 1996).
Furcraea foetida is widely cultivated and naturalised in This species is closely related to F. selloana yet mor-
climatologically suitable areas. It is considered an invasive phologically quite different. It is acaulescent and has dull
species in many areas, often on islands (e.g. Hawaii, Mada- green and narrower leaves that are not markedly narrowed
gascar, New Zealand, Réunion), but also in, for instance, towards the base (fig. 7C–D). It is further characterised by
South Africa (Crouch & Smith 2011) and Brazil (Barbosa large and remote teeth along the whole leaf margin, a very
et al. 2017). In Brazil it was shown that F. foetida may be small terminal conical spine ca. 1 mm long, brown-reddish
favoured in a scenario of climate change, increasing its nega- rhomboidal inflorescences with few partial inflorescences in
tive effects on biodiversity of coastal ecosystems (Barbosa et the upper 3/4 to nearly the whole inflorescence, flowers with
al. 2018). very narrow tepals and a very short ovary much shorter than
In the Canary Islands it was previously reported from the tepals (García-Mendoza 2001). Out of the species that
Fuerteventura, La Palma and Tenerife (Acebes Ginovés et al. are more or less widely grown as ornamentals, F. tuberosa
2010; Santos-Guerra et al. 2013), although at least some re- (Mill.) Aiton is probably the most reminiscent of F. hexa-
cords probably refer to F. selloana. In Tenerife F. foetida is a petala and both are often confused. The latter has upcurved
locally invasive species. The largest populations are known single teeth (vs. recurved double teeth) and scabrous lower
in Parque Rural de Anaga, between La Fortaleza (near to leaf surfaces (vs. smooth). In the population from Tener-
Catalanes) and la Hoya Las Colmenas. In this area it invades ife young plants have rough lower leaf surfaces whereas at
slopes of the basins that flow into El Tomadero and El Bar- maturity leaves are almost completely smooth except for a
ranco of Tahodio with hundreds of individuals, along with small basal portion. Also, leaves tend to be a bit narrower in
Agave americana (see obs. E. Ojeda-Land, 20 Apr. 2010). F. hexapetala: 8–10(–15) cm vs. 10–15(–17) in F. tuberosa
It is also found in the lower part of the Barranco de Tahodio and ovaries (and other floral characters) are slightly smaller
(see obs. E. Ojeda-Land, 8 Oct. 2010). In similar circum- as well (respectively 17–21 mm vs. 20–25 mm) (Thiede in
stances it is also known from Buenavista del Norte, Camino press). Furcraea tuberosa is locally naturalised in South Af-
El Rincón (see obs. E. Ojeda-Land, 1 Mar. 2017). rica (Smith & Figueiredo 2012b). Furcraea guatemalensis
Elsewhere in Macaronesia Furcraea foetida is also known Trel. is also very similar and often confused with F. hexa-
petala. It has, however, a distinct stem (vs. acaulescent) and
from Cabo Verde and Madeira (Vieira 2002; Sánchez-Pinto
leaf marginal teeth are more numerous (60–90 per side vs.
et al. 2005) where it is naturalised. In continental Europe it
20–50).
is surprisingly rare as an escape, perhaps for climatological
reasons. It is reportedly known from Portugal (Almeida & Furcraea hexapetala is here treated as understood now-
Freitas 2006). adays in the horticultural trade (e.g. Huxley 1999; Irish &
Irish 2000; Couper 2011). It is very similar to F. cabuya
Trel., a species from southeastern Mexico and Central Amer-
Furcraea hexapetala (Jacq.) Urb. (Urban 1903: 152) ica with type from Costa Rica (Thiede in press). Both species
Fig. 7C–D are rather similar in their vegetative features and differ main-
ly in the larger flowers, fruits and seeds of F. cabuya: flowers
Agave cubensis Jacq. (Jacquin 1763: 100). – Furcraea
(37–)40–55(–62) mm vs. 40–50(–55) mm, fruits 55–60(–75)
cubensis (Jacq.) Vent. (Ventenat 1793: 66). – Type: not
× (35–)40–45 mm vs. 30–50 × 25–40 mm, seeds winged for
designated (?).
3 mm, 15–17 × 9–10 mm vs. winged for 2 mm, 11–13 × 6–7
New to the flora of the Canary Islands – category C3 sensu mm (García-Mendoza 2001; Thiede in press). It is not im-
Blackburn et al. (2011) (Individuals surviving in the wild in possible that some plants in cultivation as F. hexapetala in
location where introduced, reproduction occurring, and pop- fact refer to F. cabuya. We were unable to check this possi-
ulation self-sustaining). bility in the Canarian populations.

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Furcraea hexapetala is known from a single locality in individuals in roadside TF-5, 25 Apr. 2018, obs. E. Ojeda-
Tenerife where it is firmly established in two small popu- Land.
lations. This species is considered invasive, especially on La Quinta, Avenida Los Pesqueros, 28°25′44.40″N,
islands in the subtropics, for instance in New Caledonia, 16°29′20.67″W, two individuals, 25 Apr. 2018, obs. E. Oje-
Galapagos and Pacific Islands and Samoa (Lundh 2006; Gar- da-Land.
dener et al. 2013; Randall 2017). It is one of the most aggres- Icod de Los Vinos, La Capellanía, 28°22′58.05″N,
sive invaders in Galapagos Islands (Tye et al. 2012). 16°40′36.54″W, 205 m a.s.l., several naturalised plants on
the slope of a cliff and on the margins of a path, 25 Jun.
2018, obs. E. Ojeda-Land.
Furcraea selloana K.Koch (Koch 1860: 22) Barranco de las Ánimas, 28°22′56.80″N, 16°40′35.89″W,
Fig. 7E–F 205 m a.s.l., five large plants, 25 Jun. 2018, obs. E. Ojeda
New to the flora of the Canary Islands – category C3 sensu Land.
Blackburn et al. (2011) (Individuals surviving in the wild in Icod de Los Vinos, Los Barbusanos, 28°22′54.48″N,
location where introduced, reproduction occurring, and pop- 16°40′32.66″W, 225 m a.s.l., some individuals in roadside,
ulation self-sustaining). 25 Jun. 2018, obs. E. Ojeda-Land.
Observation records Gran Canaria
Tenerife (selection of records) Santa Brígida (La Atalaya, Los Veroles), 28°1′55.43″N,
San Cristóbal de La Laguna, Bajamar, 28°33′21.80″N, 15°29′6.78″W, 541 m a.s.l., roadside, slope of ravine, a few
16°20′4.59″W, 60 m a.s.l., a single flowering individual on individuals, 5 Dec. 2017 and 17 Apr. 2018, obs. F. Verloove.
slope of ravine and others in a plot near the road, escaped San Bartolomé de Tirajana, Bahía Feliz, 27°47′5.86″N,
from a garden, 2 May 2011, obs. E. Ojeda-Land. 15°31′28.19″W, 41 m a.s.l., stony slope N of GC-1 motor-
San Cristóbal de La Laguna, Punta del Hidalgo, Barranco de way, several individuals in two distinct populations (prob-
la Hoya, 28°33′42.18″N, 16°19′53.54″W, 40 m a.s.l., a single ably long-abandoned plantation, with numerous Agave si-
flowering individual on slope of ravine, together with Aus- salana), 21 Apr. 2018, obs. F. Verloove.
trocylindropuntia subulata, 2 May 2011, obs. E. Ojeda-Land. Teror, La Molineta, 28°4′38.53″N, 15°31′10.73″W, 372 m
Arona, Valle de San Lorenzo, 28°5′30.88″N, 16°39′30.50″W, a.s.l., on the verge of GC-21 road, 24 Apr. 2018, obs. F. Ver-
409 m a.s.l., Barranco de Chija near TF-28 road, a single in- loove, Á. Marrero Rodríguez & M. Salas Pascual.
dividual, 22 Jun. 2014, obs. F. Verloove. Telde, Barranco de Telde, 27°59′56.27″N, 15°25′14.58″W,
Puerto de la Cruz, Malpaís, calle Chinyero, 28°24′38.21″N, 367 m a.s.l., slope of ravine, two individuals, 26 Apr. 2018,
16°33′1.20″W, 81 m a.s.l., ruderalized slope, common (by obs. F. Verloove.
the dozen), 4 Nov. 2014, obs. F. Verloove. Arucas (Santidad), slope parallel to Calle Alhucemas,
Arafo, La Hidalga, 28°20′11.94″N, 16°23′37.79″W, 190 m 28°6′45.19″N, 15°30′30.99″W, 238 m a.s.l., steep stony
a.s.l., Barranco de la Madre at TF-28 road, a single individu- slope, sun-exposed, dozens escaped from plantation, 27 Apr.
al, 10 Nov. 2014 and 19 Jan. 2017, obs. F. Verloove. 2018, obs. F. Verloove.
Tegueste, El Socorro, 28°31′7.80″N, 16°21′39.99″W, 292 Specimens collected
m a.s.l., slope of ravine, relatively frequent (naturalised), 13 Tenerife
Jan. 2017, obs. F. Verloove. San Miguel de Abona, Golf del Sur, Barranco del Saltadero,
Puerto de la Cruz, 28°24′19.79″N, 16°34′4.24″W, 79 m a.s.l., 28°1′58.62″N, 16°36′46.06″W, 56 m a.s.l., rough ground
grassy slope, scattered individuals, 17 Jan. 2017, obs. F. Ver- adjacent to golf court, scattered young individuals, 30 Oct.
loove. 2014 and 12 Jun. 2015, F. Verloove 11530 (BR).
La Matanza de Acentejo, auxiliary road of TF-5 near Arona, La Camella, S of the village, 28°4′37.09″N,
road down to El Caletón, scattered individuals from 16°40′54.53″W, 337 m a.s.l., dry rough ground, a single ro-
28°27′11.29″N, 16°27′32.46″W, to 28°26′56.13″N, sette, 11 Mar. 2016, F. Verloove 12462 (BR).
16°27′44.09″W, 280 m a.s.l., 20–30 individuals, together Gran Canaria
with Austrocylindropuntia subulata and Furcraea foetida, 17 Las Palmas de Gran Canaria, Tafira Baja, close to Finca La
Apr. 2018, obs. E. Ojeda-Land. Palmita, 28°4′7.82″N, 15°27′26.47″W, 321 m a.s.l., rocky
La Victoria de Acentejo, Risco La Sardina, at Urbanization slope; scattered individuals (relic of former plantation?), 24
La Palmita, 28°26′21.09″N, 16°28′59.13″W, 150 m a.s.l., Apr. 2018, F. Verloove 13203 (BR).
three naturalised plants (escaped from a nearby garden), 25 Tafira Baja, entornos del Campus, sobre el Guiniguada,
Apr. 2018, obs. E. Ojeda-Land. 28º04′08.2″N, 15º27′26.8″W, 310–315 m a.s.l., laderas y
Santa Úrsula, La Quinta, several points on the cliff, near Calle antiguas terrazas de cultivo, subespontánea, estolonífera, 27
Codeso, 28°25′54.56″N, 16°29′50.89″W, 180 m a.s.l., five Jun. 2019, Á. Marrero 37487-37489 (LPA).
plants together with Aloe arborescens, from 28°25′58.59″N, Furcraea selloana is native to Mexico and parts of Central
16°29′52.49″W to 28°26′0.23″N, 16°29′52.63″W, 170 and South America. It is widely cultivated as an ornamental
m.a.s.l, 10–15 naturalised individuals, 25 Apr. 2018, obs. E. in dry warm-temperate and subtropical areas of the world. It
Ojeda-Land. is increasingly reported as a naturalised or even (potentially)
Santa Úrsula, La Quinta, near the roundabout of the TF- invasive escape, for instance in Australia and New Zealand,
5, 28°25′35.62″N, 16°29′37.66″W and 28°25′37.55″N, South Africa and the U.S.A. (e.g. Forster 1996; Batianoff &
16°29′37.82″W, 255 m a.s.l., numerous naturalised plants Butler 2002; Verhoek 2002; Smith & Figueiredo 2016; Ran-
and 28°25′34.43″N, 16°29′49.55″W, 71 m a.s.l., dozens of dall 2017). In Europe, F. selloana has been reported from

493
Pl. Ecol. Evol. 152 (3), 2019

Portugal (Aedo 2013; Silva et al. 2015) and Spain (Sánchez reviewers are acknowledged for providing useful comments
Gullón 2013; Guillot Ortiz et al. 2016) although it may have on an earlier version of this paper and Sven Bellanger (Meise
been widely neglected elsewhere. In Tenerife and Gran Ca- Botanic Garden, Belgium) for preparing the figures.
naria, it is widely cultivated as an ornamental and occurs on
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