Notes on Genuine Agave Vivipara (Agavaceae), a Poorly Known Caribbean Species, Recently Introduced in the Canary Islands (Spain)

Notes on genuine Agave vivipara (Agavaceae), a poorly known Caribbean species, recently introduced in the Canary Islands (Spain)

Authors: Verloove, Filip, and Pascual, Marcos Salas Source: Bradleya, 2021(39) : 259-264 Published By: British Cactus and Succulent Society URL: https://doi.org/10.25223/brad.n39.2021.a28

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Notes on genuine Agave vivipara (Agavaceae), a poorly known Carib- bean species, recently introduced in the Canary Islands (Spain)

Filip Verloove1 and Marcos Salas Pascual2

1 Meise Botanic Garden, Nieuwelaan 38, B-1860 Meise, Belgium. (email: filip.verloove@botanicgar- denmeise.be) 2 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. (email: [email protected]) Photographs: Figure 1 and 6 Marcos Salas Pascual, figures 2–5 and 7–9 Filip Verloove

Summary: The Caribbean species Agave vivipara One of these species is Agave vivipara L. It was confused during several decades with the has a very restricted natural distribution that is widely cultivated ornamental species A. angusti- limited to the Leeward Islands (Aruba, Bonaire folia. Both are in fact quite dissimilar, the former and Curaçao) in the Caribbean. There, however, being rarely offered for sale in the horticultural it is locally frequent and doubtlessly the most trade. However, in recent years it was rather fre- common representative of the genus (Wagenaar quently planted in Gran Canaria in the Canary Is- Hummelinck, 1938). It has also been reported on lands (Spain) and it soon started to escape. Since various occasions from Margarita, also by the lat- it is very poorly known, it is copiously illustrated ter author, but these populations in fact belong to here and an extensive description, based on native the similar-looking A. cocui Trel. (Hoyos, 1985; and introduced plant material, is also presented. Thiede, 2020). Although the name A. vivipara frequently pops up in the horticultural literature, it is Zusammenfassung: Die karibische Art Agave vi- in fact a poorly known and much-misunderstood vipara wurde mehrere Jahrzehnte lang mit der species. Wijnands (1983) erroneously considered häufig kultivierten Zierpflanze A. angustifolia it to be conspecific with (and having nomencla- verwechselt. Beide sind jedoch recht verschieden, tural priority over!) the very popular and wide- und die erste Art wird sehr selten im Pflanzenhan- spread garden ornamental A. angustifolia Haw. As del zum Verkauf angeboten. In den letzten Jahren a consequence, since then the latter was known wurde sie jedoch häufig auf der kanarischen Insel under the erroneous name A. vivipara, not only in Gran Canaria (Spanien) gepflanzt und begann dort the horticultural trade but also in flora accounts bald auch zu verwildern. Da sie kaum bekannt ist, and scientific papers (e.g. Forster, 1992; Smith & wird sie hier ausgiebig im Bild dargestellt und, ba- Steyn, 1999a; Thiede, 2001). García-Mendoza & sierend auf heimischem und eingeführtem Pflan- Chiang (2003) pointed out that A. angustifolia and zenmaterial, ausführlich beschrieben. A. vivipara are in fact quite different species that belong to separate sections (Rigidae and Viviparae Introduction respectively). Moreover, they are geographically Species of Agave are native to the New World but isolated, the former being distributed in conti- extensively planted (usually but not exclusively) as nental America, ranging from northern Mexico to ornamentals in climatologically suitable areas else- Panama. where in the world. The Canary Islands, locally Although very attractive, genuine Agave vivi- blessed with a hot desert (BWh) or hot semi-arid para appears to be a rare species in the ornamental climate type (BSh) (Kottek et al., 2006), are partic- trade. It is not mentioned in standard works such ularly suitable for the cultivation of agaves. As a as Huxley (1999), Irish & Irish (2000) or Couper & matter of fact, circumstances are so favorable that Cullen (2011), nor was it included in a thorough several species not only easily survive in ornamen- overview of the genus in cultivation in Spain and tal plantations but also escape from them. In a re- the Balearic Islands (Guillot Ortiz & van der Meer, cent survey, fourteen species and several additional 2006). It is here copiously illustrated and an ex- infraspecific taxa of Agave have been identified as tensive description, based on Caribbean and Ca- occurring in the wild there (Verloove et al., 2019). narian individuals, is presented.

Bradleya 39/2021 259 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Identification and similar taxa type localities on Aruba (Wagenaar Hummelinck, Agave vivipara is easily recognizable on a num- 1936), where their presence was recently con- ber of characteristic features (Figures 1–7). Its ro- firmed (Thomson, 2014). Both are only margin- sette is less than 100cm in diameter and 50–60cm ally separated from A. vivipara but yet considered high. The leaves are all abruptly tapered towards distinct (Thiede, 2020). Anyway, it is likely that the apex, canaliculate and curved outwards (often far too many species of Agave have been described distinctly sigmoidal). They are broadly lanceolate, in the past in the Caribbean. For instance, in the widest at about or just above mid length and nar- Lesser Antilles Rogers (2000) reduced the number rowed above base to c. ½ leaf width (Figure 4); of species from 12 to a single, A. karatto Mill. they are on average 50×15cm (c. 3.5 × as long Despite being very characteristic, the identifi- as wide). Leaf margins are closely and regularly cation of Agave vivipara in the Canary Islands was toothed with short teeth (c. 4–5mm long) on very not straightforward. Canarian plants were initially distinct lunate bases (Figure 5). The apical spine thought to be A. desmetiana Jacobi, a species that is c. 25mm long. The colour of the plant is green to is unknown in the wild but frequently cultivated glaucous, sometimes slightly pruinose. The inflor- as an ornamental. In general appearance it close- escence is relatively short (c. 3m tall) and flowers ly resembles A. vivipara but its leaf margins are are bright yellow (Figure 6). either smooth or only have weak or small teeth It is in fact incomprehensible that Wijnands near the leaf base. In addition, its leaves are lon- (1983) considered this species to be conspecific ger (50–80cm) and have a larger length to width with Agave angustifolia, both species being very ratio. Also, its flowers are a paler yellow (Irish dissimilar. The latter has much narrower, sword- & Irish, 2000). It is locally naturalised in Florida shaped (ensiform) leaves that are c. 10-20× as (Franck, 2012). Agave wercklei F.A.C. Weber ex long as wide. Also, leaves are very rigid and radi- Wercklé, a species from Costa Rica, is also similar ately spreading, margins are straight and hardly but it is non-surculose, has a much taller inflo- narrowed towards the base (Figures 8–9). Flowers rescence with very short peduncle, paler, rough are always greenish-yellow. (at least when young) leaves with marginal teeth Although quite characteristic and well defined, not on distinct protuberances (García-Mendoza & Agave vivipara itself is a variable species. Plants Lott, 1994). It has been recorded as an escape in with deviating features are regularly observed South Africa (Smith & Steyn, 2002; Walters et al., in their area of origin (Wagenaar Hummelinck, 2011). Finally, A. cocui is also very similar and in 1936). Even though the species predominantly fact the only species of those mentioned here that reproduces clonally (from suckers and bulbils), is closely related to A. vivipara, both being mem- hybridisation probably occurs in places where re- bers of section Viviparae. It is native to Venezuela lated species grow in close proximity. According and Colombia but also occurs on Margarita island, to Wagenaar Hummelinck (1993) putative hybrids where it probably is a naturalised introduction with A. cocui are most notable and on Margarita (Wagenaar Hummelinck, 1936; Hoyos, 1985; Wa- island these two species are very hard to distin- genaar Hummelinck, 1993; Thiede, 2020). It is a guish. This, however, is very unlikely since A. coarser, non-surculose species with slightly lon- vivipara is absent from Margarita (Hoyos, 1985; ger leaves and a much taller inflorescence reach- Thiede 2020). In the Can-arian plants leaf mar- ing 5–10m in length (Thiede, 2020). It is, however, ginal teeth are more closely placed than usually not always clearly separated from A. vivipara (Wa- recorded. This may indicate introgression or even genaar Hummelinck, 1938) and may be a taller, hybridization although this character also seems more robust expression of it. to be very variable in autochthonous populations as shown in N.L.Britton & J.A.Shafer 3060 from Nomenclature Willemstad, Curaçao, March 1913 (U.1048296, Agave vivipara L., Species Plantarum 1: 323. herb. U). 1753. (Figures 1–7) Trelease (1913) described the very similar Aga- (Agave foliis dentatis, staminibus corollam ae- ve vicina Trel. from Aruba. It was differentiated quantibus) by its dull green (vs. transiently glaucous) leaves non A. vivipara Arruda, Diss. Pl. Brazil 23. 1810, with heavily triangular marginal teeth from large nom. illegit. lunate bases (vs. more slender teeth from smaller Type: Wijnands (1983) lectotypified Agave vi- lunate bases). Also, marginal teeth are slightly vipara and relegated A. angustifolia to synonymy. less closely placed in A. vicina than in A. vivipara. Even though he referred to Trelease (1913), Berger According to Wagenaar Hummelinck (1993) and (1915), and Wagenaar Hummelinck (1936), he ap- Álvarez de Zayas (1995), A. vicina is best consid- parently disagreed with these authors in treating ered conspecific with A. vivipara, an opinion not A. vivipara and A. angustifolia as distinct species. shared by recent authors (Thiede 2020). Two fur- He designated Commelin’s illustration of Aloe ther species, A. rutteniae Hummelinck and A. aru- americana polygona (Commelin, 1703, t. 15), bensis Hummelinck, are only known from their wrongly cited as t. 65 by Linnaeus (1753) in the

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Figure 1. (left above) Agave vivipara in an ornamental plantation in Las Palmas de Gran Canaria. This is a relatively small-statured species. Figure 2. (left below) Agave vivipara in Las Palmas de Gran Canaria (escaped individual). At first flowering this species grows solitary. Suckering only starts later. Figure 3. (above) Agave vivipara in Arucas (abandoned plantation). This species is soon suckering after flowering .

protologue, as lectotype of the name A. vivipara, a viewpoint subsequently approved by García- Mendoza & Chiang (2003; see also Jarvis, 2007). In fact, neither the protologue nor the lectotype make any approach for a secure definition of the species, as was already stated before by Wagenaar Hummelinck (1993). Description: This description is based on Trelease (1913) and Wagenaar Hummelinck (1936, 1993), adjusted based on plants currently found in Gran Canaria (cultivated as well as wild). Floral details were measured in ten flowers from three different localities. Leaf measurements were as- sessed from 15 leaves originating in five different localities. Our own data (in bold) were checked against and found to be in agreement with the descriptions presented by Trelease l.c. and Wa- genaar Hummelinck l.c., except for the number of leaf marginal teeth which usually is slightly higher in Canarian plant material (see before). In his short differential diagnosis, Wagenaar Hum- melinck (1936) states that leaf marginal prickles in A. vivipara may range between 6 and 22 per 10 cm (vs. 18 to 26 according to our own measurements). Also, measurements of stamens are (very) slightly divergent. Medium-sized, herbaceous, monocarpic, succulent, (nearly) trunkless, multi-annual; rosettes with few to many leaves, solitary at first but soon suckering after flowering, c. 1m in diameter, 0.5– 0.6m tall. Leaves erect to spreading and gracefully arched, all abruptly tapered towards the apex, canaliculate and curved outwards (often distinctly sigmoidal), broadly lanceolate, widest at about

Bradleya 39/2021 261 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise long as wide, about ½ or more of total length, with c. 20–30 (25 on average) ascending side-branches, freely bulbiferous after flowering; peduncle lax- ly bracteate with broadly triangular, imbricated bracts that are weakly toothed but with pungent apical spine; pedicels 4–8mm long (5mm on average). Flowers 50–60 per partial inflorescence, narrowly cylindrical, 40–50mm long (on average 37–44mm: corolla tube 24–28mm, corolla lobes 13–16mm), the lobes bright yellow. Ovary 20– 25mm long, about equaling the perianth, oblong; tube open, about 4mm deep; segments 4 by 15mm, shorter than the ovary; filaments inserted a little below the throat, 30–36mm long (or slightly longer: 38mm on average), whitish to pale yellow, an- thers 16–20mm long (or slightly shorter: 15mm on Figure 4. Agave vivipara in Arucas (rosette leaves). average), bright yellow. Capsules broadly oblong, Leaves are broadly lanceolate, widest at about or just 25 × 30mm, very shortly stipitate and beaked, above mid length and narrowed above base to about ½ usually dropped before ripening. Seeds not pro- leaf width. duced or empty and non-viable (Figures 1–7). Illustrations: Illustrations of genuine Agave vivipara are rare. Representative photos and line drawings are provided by Trelease (1913), Wage- naar Hummelinck (1936, 1938, 1993), Verloove et al. (2019) and in this paper (Figures 1-7). Distribution: Agave vivipara is originally restricted to the Leeward Islands (Aruba, Bonaire and Curaçao) (Wagenaar Hummelinck, 1993). Claims from Margarita are erroneous and refer to A. cocui (Thiede, 2020). In addition and outside its natural distribution range it is grown as an ornamental, although probably rarely so. It has been known at least since 2012 from Gran Can-aria in the Canary Islands where it easily reproduces (from bulbils) and escapes in the vicinity of planted individuals (Verloove et al., 2019). It is regularly planted there in public green in the wide Las Palmas de Gran Canaria area but it was also noticed in other mu- Figure 5. Agave vivipara in Arucas. Leaf margins are nicipalities (e.g. in Arucas and Telde). Its presence closely and regularly toothed. The short teeth are on – solely as a garden ornamental, not yet in the wild very distinct lunate bases. – was also observed on the island of Tenerife (pers. obs. first author in December 2018 in La Mareta, Granadilla de Abona). A. vivipara was also reported mid length and narrowed above base to c. ½ leaf before from South Africa (Walters et al., 2011; see width (on average 7.29cm at narrowest width), also Smith & Steyn, 1999b, sub A. decipiens Baker). 40–60 × 12–20cm (on average 49.07 × 14.29cm), According to Berger (1915) the species was intro- c. 3.5 × as long as wide; green to glaucous, some- duced to the Mortola Botanic Garden in Italy (now times slightly pruinose; margins closely and regu- Hanbury Botanical Gardens). However, the name larly toothed (about 2 teeth per centimeter in the no longer is included in the actual alphabetical middle of the blade; ranging between 9 and 13 catalogue of plants growing in the Gardens. per 5cm at mid leaf length) with short blackish- Flowering time and phenology: In the Canary brown teeth 3–9mm long (on average c. 4.5mm Islands the inflorescence of Agave vivipara starts long, ranging between 2 and 8mm at mid leaf to develop shortly after Christmas (end of Decem- length) on much paler lunate bases, commonly ber, beginning of January). The lower branches of upcurved above and recurved below; apical spine the inflorescence open at the beginning of April, pungent, 17–28mm long (c. 24–25mm long) and c. and the last ones in May. This occurs both in irri- 6mm wide at base, not or very slightly decurrent, gated and cultivated specimens and in feral speci- grooved, blackish-brown, dull, slightly rough to mens and does not seem to depend on irrigation the touch. Inflorescence paniculate, erect, 3m tall but on the photoperiod. In the Canaries the flowers (rarely more), the pole narrowly oblong, c. 3 × as fall off soon (they remain 20–25 days on the plant)

262 Bradleya 39/2021 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Acknowledgements Bradleya 39/2021 John de Freitas (Curaçao) and André pagesvan Proosdij 259–264 (the Netherlands) are thanked for confirming the identity of Agave vivipara in the Canary Islands and for providing useful additional information. The authors are grateful to Joachim Thiede (Ger- many) who provided useful comments on a first draft of this paper.

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Bradleya 39/2021 263 Downloaded From: https://bioone.org/journals/Bradleya on 18 Jun 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Botanic Garden Meise Figure 9. Agave angustifolia in Mogán. Leaf margins are straight and hardly narrowed towards the base.

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