Bradleya 35/2017 pages 58–79

New records of naturalised and invasive cacti (Cactaceae) from Gran Canaria and Tenerife, Canary Islands, Spain

Filip Verloove 1, Elizabeth Ojeda-Land 2, Gideon F. Smith 3,4 , Alessandro Guiggi 5, Jorge Alfredo Reyes-Betancort 6, Carlos Samarín 2, Antonio González Hernández 7 & Rubén Barone 8 1. Botanic Garden of Meise, Nieuwelaan 38, B-1860 Meise, Belgium. (email: [email protected]) 2. Viceconsejería de Medio Ambiente. Gobierno de Canarias. C/ Avda. de Anaga, 35. Planta 11. 38071 Santa Cruz de Tenerife, Canary Islands, Spain. (email: [email protected]; [email protected]) 3. Centre for Functional Ecology, Departamento de Ciências da Vida, Universidade de Coimbra, 3001- 455 Coimbra, Portugal. (email: [email protected]) 4. Department of Botany, P.O. Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth, 6031 South Africa. 5. DISTAV, Polo Botanico, Università degli Studi di Genova, International Cactaceae Research Center (ICRC), Corso Dogali, 1M - 16136 Genova, Italy. (email: [email protected]) 6. Jardín de Aclimatación de La Orotava (ICIA). C/ Retama 2, 38400 Puerto de la Cruz, Canary Islands, Spain. (email: [email protected]) 7. Servicio de Sanidad Vegetal. Consejería de Agricultura, Ganadería y Pesca del Gobierno de Canarias, Avda. José Manuel Guimerá 10, 3ª planta 38071 Santa Cruz de Tenerife, Canary Islands, Spain. (email: [email protected]) 8. C/. Eduardo Zamacois, 13-3ºA, 38005 Santa Cruz de Tenerife, Canary Islands, Spain. (email: [email protected])

Summary : Recent field work in Gran Canaria and Zusammenfassung : Bei jüngsten Feldstudien auf Tenerife (Canary Islands, Spain) yielded records Gran Canaria und Teneriffa (Kanarische Inseln, for 30 alien taxa of Cactaceae that had not been Spanien) wurden 30 nicht indigene Taxa von reported before, either from the whole area, or Cactaceae nachgewiesen, die bisher weder vom from one of the islands. Out of these, 17 are con - Gesamtarchipel noch von einer der beiden Inseln sidered locally naturalised and/or potentially in - gemeldet waren. Von diesen werden 17 als lokal vasive: bigelovii , C. fulgida , C. eingebürgert und / oder potenziell invasiv ange - pallida , C. prolifera , C. tunicata , Echinocereus sehen: Cylindropuntia bigelovii , C. fulgida , C. pal - rigidissimus , Haageocereus kagenekii , Hylocereus lida , C. prolifera , C. tunicata , Echinocereus triangularis , Opuntia basilaris , O. elatior, O. rigidissimus , Haageocereus kagenekii , Hylocereus ficus-indica × O. tomentosa , O. macrocentra , O. triangularis , Opuntia basilaris , O. elatior , O. microdasys , O. pilifera , Oreocereus pseudofossu - ficus-indica × O. tomentosa , O. macrocentra , O. latus , articulatus and Trichocereus microdasys , O. pilifera , Oreocereus pseudofossu - huascha . The same applies to a rather character - latus , Tephrocactus articulatus und Trichocereus istic form of O. ficus-indica that sometimes is re - huascha . Das Gleiche gilt für eine sehr charakte - ferred to as f. amyclaea (syn.: O. megacantha ). The ristische Form von O. ficus-indica , die manchmal presence of Opuntia monacantha and O. robusta , als f. amyclaea (syn.: O. megacantha ) bezeichnet two with a dubious status in the Canary wird. Zehn weitere Taxa, oft Anbaurelikte, gelten Islands, is confirmed. Ten further taxa are con - als nicht eingebürgert. Alle Taxa sind abgebildet, sidered casuals, often relics of cultivation. All taxa zu den eingebürgerten und / oder potenziell inva - are illustrated, and for the naturalised and/or po - siven Taxa erfolgen weitere Informationen. Zwei tentially invasive taxa additional information is neue Kombinationen werden vorgeschlagen: Cy - provided. Two new combinations are proposed for lindropuntia fulgida f. mamillata und Tephro - f. mamillata and Tephro - articulatus f. papyracanthus . cactus articulatus f. papyracanthus .

58 Bradleya 35/2017 Introduction (Willd.) Haw., also in the Canary Islands; see The Canary Islands, Spain, have a well-docu - later]. Two others, O. tuna and Austrocylindrop - mented and rich native flora with many local, untia cylindrica , probably refer to respectively highly specialized endemic species (e.g. erroneous records, or are only very locally natu - Bramwell, 1976; Beltrán et al ., 1999). In addition, ralised. The remaining species have been con - as a result of their subtropical climate, the is - firmed by the authors of the present paper. In the lands are an outstanding cultivation ground for past few years some additional species of cacti hundreds of ornamental subtropical have been recorded: Cylindropuntia prolifera species. An increasing number of these are not (Engelm.) F.M. Knuth and Opuntia microdasys only well-adapted to be cultivated in the local cli - (Lehm.) Pfeiff. from Fuerteventura (Verloove & mate, but are indeed reproducing locally without Guiggi, 2013), O. leucotricha DC. from Gran Ca - human intervention, eventually naturalising, naria, La Palma, and Tenerife (Verloove, 2013; spreading, or even becoming invasive. In past Santos-Guerra et al ., 2014), and Espostoa years the knowledge about these alien species melanostele from Tenerife (Verloove, 2016). has considerably increased and numerous ac - In the past few years some of us discovered counts have been published (e.g. Verloove & several escaped cacti in Tenerife and Gran Ca - Reyes-Betancort, 2011; Santos-Guerra et al ., naria that had not previously been recorded in 2013; Scholz et al ., 2013; Siverio Núñez et al ., the Canary Islands. Their identities were as - 2013; Verloove, 2013; González Montelongo et al ., sessed in close collaboration with two of the au - 2014; Santos-Guerra et al ., 2014; Santos-Guerra thors (G.F. Smith and A. Guiggi) . In the present & Reyes-Betancort, 2014; Otto & Verloove, 2016). paper 30 taxa are presented and illustrated, However, some taxonomic groups are still insuf - most of them not previously reported from the ficiently known. This certainly holds true for in - Canary Islands. Their degree of naturalisation troduced succulents, despite the fact that the varies from ephemerals to locally, or potentially, islands have an ideal climate that allows the per - invasive aliens. In the absence of species-specific sistence and acclimatization of numerous species, pollinators, some of the species are able to spread especially those from arid and hot habitats. The by vegetative means, and their establishment southern parts of Tenerife, for instance, have cli - could be favoured by climate change (Capdevila- matological circumstances comparable with those Argüelles et al. , 2011; Del-Arco, 2008). Several in , U.S.A. Unsurprisingly, the island species of cacti are among the most damaging in - houses the most important producer for cacti and vasive plant species in the world (Novoa et al ., succulents worldwide (Canary Cactus S.A.). 2015) and some, for instance the whole Moreover, Spain is considered one of the three in - Cylindropuntia , are included in Spanish legisla - vasion hot spots for cacti in the world (Novoa et tion on invasive alien species (B.O.E., 2013). al ., 2015; see also Essl & Kobler, 2008). Therefore, it is important to document any in - Not a single species of the family Cactaceae is cipient invasion event of cacti, even in early native to the Canary Islands, although two stages. species have become a very familiar sight. Opun - It is noteworthy that likely most, if not all, the tia ficus-indica (L.) Mill. and O. dillenii (Ker- catalogued in this paper have a cultivated Gawl.) Haw. were introduced quite long ago, and origin, including from commercial nurseries with now have invaded large parts of the islands. propagation facilities on the Canary Islands. It is While O. dillenii is mostly confined to the lower therefore possible that, at least in some in - altitudes, the former is omnipresent, even at stances, the plants may not perfectly agree with higher altitudes. Eriksson et al . (1979), later botanical taxa as known from the wild (Roy Mot - merely repeated with a nomenclatural update by tram, personal communication). In these circum - Acebes Ginovés et al . (2010), cited the names of stances, hybridisation through open pollination nine cactus species for the Canary Islands: Aus - in fields may well have given rise to hybrids that, trocylindropuntia cylindrica (Lam.) Backeb., A. sometimes through introgression and backcross - exaltata (Berg) Backeb., Hylocereus undatus ing, resemble one of the parents. (Haw.) Britton & Rose, Opuntia dillenii , O. max - ima Mill., O. robusta H.L. Wendland, O. tomen - Materials and methods tosa Salm-Dyck, O. tuna (L.) Mill., and O. Once harvested, virtually all species of cacti vulgaris Mill. Of these, O. maxima auct. and O. are awkward to preserve as dried specimens for vulgaris auct. non Mill. are now referred to O. deposition in a herbarium, and most of the records ficus-indica [although the binomial O. vulgaris here reported are only documented with photo - has also been associated with O. monacantha graphs, unless otherwise stated.

Bradleya 35/2017 59 The presence or absence on the islands of Gran habit, to 1.5 m high, larger branches c. 5cm across, Canaria or Tenerife was each time compared with larger yellowish-green flowers, 3-4cm across and data provided by Acebes Ginovés et al . (2010), as its smaller fruit, c. 2cm across. The fruit is densely well as other literature sources. The paper is di - tuberculate, dry when ripe, usually solitary, and vided in two parts in both of which the taxa are does not form chains. arranged in alphabetical sequence. Naturalised Cylindropuntia bigelovii is not known as an in - and/or potentially invasive taxa are dealt with in vasive species (Novoa et al ., 2015), nor has it been the first part; these are treated in more detail than reported before from Europe (Essl & Kobler, the taxa recorded as casuals in the second part. 2008). In Güímar, C. bigelovii obviously is a re - Each entry includes: [1.] the scientific name of cent escape and cannot be classified as natu - the taxon (accompanied by one or more synonyms, ralised yet. However, like other species from this if deemed desirable); [2.] type of chorological nov - genus, it certainly has the potential to establish elty and estimated degree of naturalisation ( sensu and spread to the nearby nature reserve of Mal - Richardson et al ., 2000); [3.] enumeration of local - país de Güímar. ities; [4.] origin of the taxon; and [5.] details about its secondary distribution. Cylindropuntia fulgida (Engelm.) F.M. Knuth, Nomenclature of the taxa presented is mostly Kaktus-ABC 126. 1935. (Figure 2). in accordance with recent insights, often based on f. mamillata (Schott ex Engelm.) Guiggi & Ver - molecular phylogenetic studies. Authorities of loove comb. nov. plant names usually follow Tropicos (www.tropi - Basionym: Opuntia mamillata Schott ex Engelm., cos.org). Proc. Amer. Acad. Arts 3: 308. 1856.

Results ≡ C. fulgida var. mamillata (Schott ex Engelm.) Backeb., Cactaceae Handb. Kakteen. Pereskioi - Part 1. Naturalised and/or potentially inva - deae 1: 204. 1958. sive taxa ≡ Opuntia fulgida Engelm., Proc. Amer. Acad. In this part those taxa are presented that ei - Arts 3: 306–307. 1856. ther are naturalised locally, or are likely to be - var. mamillata (Schott ex Engelm.) J.M. Coult., come so in the near future. Some have at present Contr. U.S. Natl. Herb . 3(7): 449. 1896. a very limited distribution range, but have been recorded as invasive in climatologically similar New to the flora of the Canary Islands. areas of the world, or are found in, or near, vul - nerable natural habitats on the islands. TENERIFE: Guía de Isora: Playa de San Juan, 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony Cylindropuntia bigelovii (Engelm.) F.M. slope next to the sea cliffs, numerous individuals, Knuth, Kaktus-ABC 125. 1935. (Figure 1). 24.06.2016, F. Verloove.; idem, 07.05.2016, E. ≡ Opuntia bigelovii Engelm., Proc. Amer. Acad. Ojeda-Land et al. ; Santiago del Teide: Puerto de Arts 3: 307. 1856. Santiago, 28°14’13.42”N, 16°50’23.34”W, 51 m.s.m. , barranco de Santiago close to calle Seis, New to the flora of the Canary Islands. rocky slope of ravine, c. 50 individuals, 15.11.2016, F. Verloove; idem, 06.01.2017, F. Ver - TENERIFE: Güímar, 28°18’34.15”N, loove 12711 (BR). 16°22’57.44”W, 113 m.s.m. , Camino de Samarines E of bridge over TF-1 motorway, roadside, a single Cylindropuntia fulgida is a species originally individual, 23.11.2016, F. Verloove. native to Mexico and adjacent parts of the south - western U.S.A. It is widely cultivated as an orna - A species from the southwestern U.S.A. and mental (e.g. Sánchez de Lorenzo Cáceres, 2000) adjacent parts of Mexico, Cylindropuntia bigelovii and readily escapes, its stem segments easily de - is cultivated as an ornamental (Sánchez de taching and rooting. It is often considered an un - Lorenzo Cáceres, 2000). In Güímar a single indi - desirable, highly invasive species, for instance in vidual was observed in a dry roadside close to the Australia (Chinnock, 2015) and South Africa Canary Cactus S.A. establishment from where it (Walters et al ., 2011; see also Novoa et al ., 2015). obviously had escaped. It had not been recorded before from Europe (Essl With its easily detaching stem segments that & Kobler, 2008). are densely covered in spines, this species resem - In Tenerife two populations were recently de - bles C. fulgida (Engelm.) F.M. Knuth var. fulgida . tected. In Playa de San Juan, numerous individu - It is distinguished from the latter by its shrubby als have been observed on a dry, sun-exposed

60 Bradleya 35/2017 Figure 1. Cylindropuntia bigelovii , Güímar (TF), Figure 2. Cylindropuntia fulgida , Puerto de San - dry roadside, November 2016. tiago (TF), slope of ravine, January 2017. Photograph by F. Verloove. Photograph by F. Verloove.

Figure 3. Cylindropuntia pallida , El Médano Figure 4. Cylindropuntia prolifera , Punta del Hi - (TF), stony slope of ravine, June 2016. Spines are dalgo (TF), September 2010. white throughout. Photograph by F. Verloove. Photograph by F. Verloove. slope close to the sea cliffs. In this locality it obvi - grows with C. tunicata (Lehm.) F.M. Knuth, an - ously persists from an abandoned plantation. other potentially invasive species. However, it evidently reproduces and appears The plants found in Tenerife have stem seg - very prolific. Further local expansion of the popu - ments appearing spineless or nearly so from afar, lation is very likely. A smaller population with c. exposing strongly mamillate tubercles, a trait of 50 individuals was discovered in Puerto de Santi - var. mamillata (Pinkava, 2003a). A taxonomic ago on volcanic rocks bordering a ravine. In this rank lower than varietas for such minor variation, locality, C. fulgida also looks well-established and however, seems more appropriate. A new combi - will probably start spreading in the near future. It nation, as forma, is proposed here.

Bradleya 35/2017 61 Figure 6. , new growth with typical pinkish-brownish spines, Puerto de Santiago (TF), slope of ravine, January 2017. Photograph by F. Verloove.

Figure 5. Cylindropuntia tunicata , Puerto de Santiago (TF), slope of ravine, January 2017. Photograph by F. Verloove.

Figure 7. Echinocereus rigidissimus , Playa de San Juan (TF), stony slope close to sea cliffs, May 2016. Photograph by J.A. Reyes-Betancort.

Figure 9. Hylocereus triangularis , Valle de San Figure 8. Haageocereus kagenekii , El Médano Lorenzo (TF), dry river bed, January 2017. (TF), stony slope of ravine, January 2017. Photograph by F. Verloove. Photograph by E. Ojeda-Land.

62 Bradleya 35/2017 Most of the plants observed belong to a com - ser. 2 14 (42): 338. 1852. monly cultivated morphotype with crested stems, which is often referred to as the ‘boxing-glove cac - New to the flora of Tenerife. Previously reported tus’. from Fuerteventura (Verloove & Guiggi, 2013) and Lanzarote where a project of eradication has Cylindropuntia pallida (Rose) F.M. Knuth, been conducted in La Triguera, Tenegüime Kaktus-ABC 126. 1935. (Figure 3). (http://www.cabildodelanzarote.com/tema.asp?sec ≡ Opuntia pallida Rose, Smithsonian Misc. Col - =Noticias&idTema=17&idCont=12077 ). Other lect. 50 : 507. 1908. target species of this eradication program were = auct. non (DC.) Backeb., Cylindropuntia kleiniae (DC.) F.M. Knuth, Opun - Cactaceae Handb. Kakteen. Pereskioideae Opun - tia engelmannii Salm-Dyck ex Engelm., and O. tioideae 1: 197. 1958. leucotricha . All were deliberately introduced ini - tially, but only C. prolifera was able to spread. New to the flora of the Canary Islands. TENERIFE: San Cristóbal de La Laguna: Punta TENERIFE: Granadilla de Abona: Hoyas de Doña del Hidalgo, 15.09.2010, F. Verloove; Granadilla María, Las Arenas-El Médano, 2 8°03’43.29”N, de Abona: Hoyas de Doña María, Las Arenas-El 16°31’44.87”W, 60 m.s.m. to 28°03’36.70”N, Médano, 2 8°03’43.29”N, 16°31’44.87”W, 60 m.s.m. 16°31’41.68”W, 54 m.s.m. , 04.04.2015, E. Ojeda- to 28°03’36.70”N, 16°31’41.68”W, 54 m.s.m. , Land et al .; Granadilla de Abona: La Mareta, 04.04.2015, E. Ojeda-Land et al. ; San Miguel de 28º01’55.36”N, 16º34’27.51”W, 11 m.s.m. , Abona: Amarilla Golf, 28º01’18.62”N, 05.01.2017, E. Ojeda-Land et al .; Buenavista del 16º37’03.9”W, 20 m.s.m. , 01.03.2017, E. Ojeda- Norte: Costa de El Rincón, 28º22’13.21”N, Land. 16º52’15.61”W, 15 m.s.m. , 09.02.2017, E. Ojeda- Land et al . This species is restricted to a relatively small area in Mexico (Baja ) and the U.S.A. In El Médano a huge population is naturalised (California). It is said to be a hybrid of two Mexi - and spreading from an abandoned, not well de - can species, Cylindropuntia alcahes (F.A.C. limited cultivation experiment. In this locality nu - Weber) F.M. Knuth and C. cholla (F.A.C. Weber) merous cacti have been observed ( Opuntia spp., F.M. Knuth (Pinkava, 2003a). Outside its native Cylindropuntia spp., Cleistocactus spp., Oreo - area it is planted as an ornamental, although per - cereus spp., etc.), some of them showing invasive haps not widely so; for instance, it is not men - behavior. In La Mareta a few naturalised individ - tioned by Sánchez de Lorenzo Cáceres (2000). uals were observed that escaped from a garden In Punta del Hidalgo in Tenerife a small pop - close to the road TF-643, and in similar circum - ulation of Cylindropuntia prolifera was recorded stances five individuals were recorded in Costa de in 2010. In El Médano, also in Tenerife, a huge El Rincón in Buenavista del Norte. population has naturalised and is spreading from Cylindropuntia pallida has long been known an abandoned, not well controlled cultivation ex - as C. rosea . The type of the latter, however, was periment . Close by, in Amarilla Golf, only two in - shown to belong to C. imbricata (Haw.) F.M. dividuals were observed growing on a volcanic Knuth (Laguna et al ., 2013). This species is doubt - rocky slope near to the golf court. fully distinct from C. tunicata and its nomencla - Although not mentioned as an invasive species ture is very confusing. The neotype of C. imbricata by Novoa et al . (2015), Cylindropuntia prolifera is is the painting of Haworth’s plant by Duncanson known as such, at least in southern Australia at Kew, designated by Hunt & Taylor (2006), where it is quite widespread (Chinnock, 2015). It is which may as well depict O. subulata . Haworth’s reminiscent of C. spinosior (Engelm.) F.M. Knuth, description of C. imbricata calls for it to be imbri - and the two species may have been confused. The cate and tessellate, which C. imbricata auct. is latter is distinguished by its white spines and yel - certainly not. Thus an early misapplication of the low to orange fruits, as opposed to the brownish name took place, probably by Engelmann, which spines and green fruits of C. prolifera . means that C. rosea should have been the correct name for what is called C. imbricata today (Roy Cylindropuntia tunicata (Lehm.) F.M. Knuth, Mottram, personal communication). Kaktus-ABC 126. 1936. (Figures 5–6). ≡ Opuntia tunicata (Lehm.) Pfeiff., Enum. Diagn. Cylindropuntia prolifera (Engelm.) F.M. Cact . 170 . 1837. Knuth, Kaktus-ABC 126. 1935. (Figure 4). ≡ Cactus tunicatus Lehm., Nova Acta Phys.-Med. ≡ Opuntia prolifera Engelm., Amer. J. Sci. Arts , Acad. Caes. Leop.-Carol. Nat. Cur . 16 : 319. 1832.

Bradleya 35/2017 63 New to the flora of the Canary Islands. growth (vs white), by its yellowish brown corollas (vs rose-red) and by less numerous spines per are - TENERIFE: Granadilla de Abona: Hoyas de Doña ole (4–7 vs 7–14). María, Las Arenas-El Médano, 28°03’43.29”N, 16°31’44.87”W, 60 m.s.m. to 28°03’36.70”N, Echinocereus rigidissimus (Engelm.) F. Haage, 16°31’41.68”W, 54 m.s.m. , 04.04.2015, E. Ojeda- Special Preisverz . 13. 1897. (Figure 7). Land et al .; Santiago del Teide: Puerto de Santi - ago, 28°14’13.42”N, 16°50’23.34”W, 51 m.s.m. , New to the flora of the Canary Islands. barranco de Santiago close to calle Seis, rocky slope of ravine, ca. 20–30 individuals, 15.11.2016, F. Ver - TENERIFE: Guía de Isora: Playa de San Juan, loove; Granadilla de Abona: San Isidro, 28°10’31.79”N, 16°48’27.37”W, 27 m.s.m. , 28°4’9.69”N, 16°33’41.05”W, 170 m.s.m. , calle 07.05.2016, J.A. Reyes-Betancort et al. Cuevas de Choportada, rough ground, c. 10–15 in - dividuals, 20.11.2016, F. Verloove; Santiago del This species is native to Arizona and New Teide: Puerto de Santiago, 28°14’13.42”N, Mexico (U.S.A.) and Chihuahua and Sonora (Mex- 16°50’23.34”W, 51 m.s.m. , barranco de Santiago ico) (Zimmerman & Parfitt, 2003). It is widely close to calle Seis, rocky slope of ravine, 06.01.2017, grown as an ornamental, also in mainland Spain F. Verloove 12710 (BR); Granadilla de Abona: San (Sánchez de Lorenzo Cáceres, 2000). Isidro, 28°4’13.08”N, 16°33’40.15”W, 172 m.s.m. , S of calle el Almendro, dry stony slope, scattered in - In Tenerife, in Playa de San Juan, two indi - dividuals, 08.01.2017, F. Verloove; Granadilla de viduals were found growing on the edge of a Abona: San Isidro, 28°4’15.56”N, 16°33’43.07”W, coastal cliff, in gravelly, sandy soils. To our knowl - 181 m.s.m. , on the verge of barranco del Conde, few edge, this species has not yet been recorded in the individuals, 08.01.2017, F. Verloove. wild beyond its native distribution range. It is, however, cultivated as an ornamental (especially Cylindropuntia tunicata is a native species in its subsp. rubispinus (G. Frank & A.B. Lau) N.P. a small area in Texas, U.S.A., and adjacent parts Taylor (Taylor, 1989). of Mexico. It is also known from Chile and Ecuador in South America (Pinkava, 2003a). Else - Haageocereus kagenekii (C.C.Gmel) Mottram, where it is widely grown as an ornamental, also The Cactician 7: 12. 2014. (Figure 8). in Europe (see for example Sánchez de Lorenzo ≡ Cactus kagenekii C.C.Gmel., Hortus magni Cáceres, 2000). ducis badensis carlsruhanus : 48. 1811. (Fig. 3) In Tenerife several small populations were re - ≡ Cereus kagenekii (C.C.Gmel.) K.Schum., Ge - cently detected. In El Médano this species is nat - samtbeschreibung der Kakteen (2): 66. (15 May) uralised in a gravelly riverbed along with many 1897. other cacti, resulting from an abandoned cultiva - = Cereus pseudomelanostele Werderm. & Backeb., tion experiment. In Puerto de Santiago the species Neue Kakteen 74. 1931. grows on volcanic rocks of a ravine close to houses, = Haageocereus pseudomelanostele (Werderm. & along with C. fulgida . In San Isidro it was found Backeb.) Backeb., Kaktus-ABC 209. 1935. in several subpopulations, in disturbed xeric as = Haageocereus multangularis (Haw.) F. Ritter, well as in more natural habitats. In all these lo - Kakteen Südamer . 4: 1401. 1981. calities C. tunicata appears well-established and a future local expansion is predicted, especially New to the flora of the Canary Islands. since the stem segments easily become detached and root where they fall. This species is a known TENERIFE: Granadilla de Abona: Hoyas de Doña invader, for instance in Australia (Chinnock, María, Las Arenas-El Médano, 2 8°03’43.29”N, 2015; Novoa et al ., 2015), but it was not previously 16°31’44.87”W, 60 m.s.m. to 28°03’36.70”N, reported from Europe according to Essl & Kobler 16°31’41.68”W, 54 m.s.m. , 04.04.2015, E. Ojeda- (2008). Guillot et al . (2008), however, have since Land et al. reported it from several localities in Spain and it is also known from Italy (Guiggi, 2008). Previous As noted above this species is associated with claims in South Africa proved to be referable to C. an abandoned, not well managed cultivation ex - pallida (Rose) F.M. Knuth (Walters et al ., 2011). periment in El Médano. Several individuals were Cylindropuntia tunicata closely resembles C. detected in the surrounding native vegetation pallida and the two species may have been con - that is dominated by Euphorbia balsamifera fused in the past. It is distinguished, however, by Aiton and Lycium intricatum Boiss., which indi - its spines that are pinkish or brownish in young cates a local naturalisation event .

64 Bradleya 35/2017 Figure 10. Opuntia basilaris , Güímar (TF), vol - Figure 11. Opuntia elatior , Buenavista del Norte canic slope, January 2017. (TF), rough ground, March 2017. Flowering culti - Photograph by F. Verloove . vated individual, growing close to the escaped non-flowering plants. Photograph by J.A. Reyes-Betancort.

Figure 13. Opuntia ficus-indica f. amyclaea (O. megacantha ), Ayagaures (GC), stony slope, Nov- ember 2015. Photograph by F. Verloove.

Figure 12. Opuntia ficus-indica × O. tomentosa , Masapeses (GC), under Eucalyptus canopy, No - vember 2015. Photograph by F. Verloove.

Haageocereus kagenekii is not known as an in - vasive species (Novoa et al ., 2015), nor has it been reported before from Europe (Essl & Kobler, 2008). It is widely grown as an ornamental, also in mainland Spain (Sánchez de Lorenzo Cáceres, 2000; as H. multangularis (Haw.) F. Ritter). Note that this species is still sometimes Figure 14. Opuntia macrocentra , Playa de San treated as Haageocereus pseudomelanostele . How - Juan (TF), stony slope, November 2016. ever, Mottram (2014) published the combination Photograph by F. Verloove.

Bradleya 35/2017 65 Figure 15. Opuntia macrocentra , detail of clado - Figure 16. Opuntia microdasys , El Médano (TF), des, Playa de San Juan (TF), stony slope, Novem - dry riverbed, June 2015. ber 2016. Photograph by F. Verloove. Photograph by A. González.

Figure 17. Opuntia monacantha , Mesa de Mar (TF), roadside, January 2017. Photograph by F. Verloove.

Figure 18. Opuntia pilifera , San Agustín (GC), stony slope, November 2015. Photograph by F. Verloove.

Figure 19. Opuntia robusta , Puerto de la Cruz Figure 20. Oreocereus pseudofossulatus , El Mé - (TF), rough ground, November 2014. dano (TF), dry riverbed, June 2016. Photograph by F. Verloove. Photograph by F. Verloove.

66 Bradleya 35/2017 H. kagenekii for the species, the earliest name of soil, two small populations of c. 8 and 9m², which is Cactus kagenekii C.C.Gmel. 11.01.2017, F. Verloove.

Hylocereus triangularis (L.) Britton & Rose, Opuntia basilaris is native to a relatively Contr. U.S. Natl. Herb . 12 (10): 429. 1909. (Figure small area in the southwestern U.S.A. and the ad - 9). jacent in Mexico. It is grown as an ≡ Cactus triangularis L., Sp. pl . 1: 468. 1753. ornamental, although probably not very widely so (see for example Hunt, 1989). As far as we could New to the flora of the Canary Islands. establish, the species had not been recorded be - fore outside of cultivation. In Tenerife, scattered TENERIFE: Arona: Valle de San Lorenzo, 28° small populations were discovered on volcanic 5’30.88”N, 16°39’30.50”W, 402 m.s.m. , barranco de slopes on the verge of the Reserva Natural Espe - Chija N of TF-28, rocky slope of ravine, a single cial del Malpaís de Güímar, close to the Canary individual, 28.11.2016, F. Verloove. Cactus S.A. plant, from where it probably es - caped. A native of the Caribbean, Hylocereus trian - Opuntia basilaris is somewhat reminiscent of gularis is widely cultivated in the subtropics, ei - O. microdasys with which it shares a low-growing ther as an ornamental, or for food (‘dragon fruit’). habit, and the cladodes having numerous spine - This species, as well as several others from this less areoles densely packed with short glochidia genus, are grown in the Canary Islands (Hernán - (Smith et al ., 2011). It differs, however, by its dez et al ., 2013), although H. undatus is probably cladodes being bluish-green with reddish-brown by far the most widespread. The latter is also the glochids, and its purplish-red corollas. The popu - only species of the genus that has so far been lations currently found in Tenerife may include or recorded outside of cultivation (Acebes Ginovés et even pertain to hybrids with the similar O. rufida al ., 2010). Engelm. (Roy Mottram, personal communication). The single plant found in a ravine in Valle de San Lorenzo in Tenerife differs from H. undatus Opuntia elatior Mill., Gard. Dict. ed. 8 , 4. 1768. in having stems that are not horny-angled with (Figure 11). areoles with numerous spines (6–8), and its = O. bergeriana F.A.C.Weber ex A.Berger, Gard. smaller stature with stems only c. 3–4cm across Chron. III, 1904 (1): 34. 1904. (Britton & Rose, 1919–23). Hybridisation, either artificial or spontaneous, may have blurred New to the flora of the Canary Islands. species boundaries in this genus resulting in plants found in the wild in Europe exhibiting in - TENERIFE: Buenavista del Norte: road to Costa termediate traits. de El Rincón, 28º22’05.41’’N 16º52’13.88’’W, 37 To the best of our knowledge, Hylocereus tri - m.s.m. , 05.03.2017, J.A. Reyes-Betancort. angularis has only been recorded before in Europe in Sicily (Guiggi, 2010). Although only a single in - Opuntia elatior is native to Mesoamerica, from dividual was found in Tenerife, a future naturali - Venezuela to the Antilles (Bravo-Hollis & Arias, sation cannot be excluded since its ecology and 2011). It has been recorded as an alien from con - biology typically resemble that of H. undatus , a tinental Spain in Valencia (Guillot et al. , 2008; known invasive species (Novoa et al ., 2015). Guillot et al. , 2014) and the Balearic Islands (Ser - apio et al. , 2016), twice as O. bergeriana . In Bue - Opuntia basilaris Engelm. & J.M. Bigelow, navista del Norte in Tenerife, only two small Proc. Amer. Acad. Arts 3: 298–299. 1856. (Figure individuals were discovered near to the road, in a 10). waste place that has been partially used as a garbage dump for cacti. Cladodes were only 15cm New to the flora of the Canary Islands. long and flowers were red. Opuntia elatior has been reported before as an TENERIFE: Güímar, 28°18’28.42”N, invasive alien, for instance from Australia (Novoa 16°22’49.64”W, 94 m.s.m. , on the verge of the na - et al. , 2015). In Indonesia, India, and South Africa ture reserve Malpaís de Güímar, close to the Ca - it was introduced in the 19 th century and quickly nary Cactus S.A. plant, on volcanic soil, a single spread. As a result of biological control, however, clone, 11.01.2017, F. Verloove 12747; Güímar, these invasions have much decreased lately. It is 28°18’25.19”N, 16°22’51.91”W, 114 m.s.m. , on the apparently not very widespread in cultivation; it verge of the nature reserve Malpaís de Güímar, was not mentioned, for instance, in Hunt (1989) close to the Canary Cactus S.A. plant, on volcanic or Sánchez de Lorenzo Cáceres (2000). In Europe,

Bradleya 35/2017 67 O. elatior has also been reported from Italy ments, obviously is closer to O. ficus-indica ; more - (Guiggi, 2008). over, the other putative parent is O. tomentosa var. hernandezii (DC.) Bravo (syn.: O. hernandezii Opuntia ficus-indica (L.) Mill. × O. tomentosa DC.). Salm-Dyck. (Figure 12). Several perfectly established populations were discovered in the past years, both in Gran Canaria New to the flora of the Canary Islands. and Tenerife (see above). It usually grows on rocky, often wooded, slopes, and may well have GRAN CANARIA: Arucas: Masapeses, urban - been overlooked elsewhere in the Canary Islands. ización El Caserío, 28° 5’54.99”N, 15°31’56.76”W, In the Canary Islands Opuntia ficus-indica 374 m.s.m. , GC-43 between Teror and Visvique, and O. tomentosa are both considered invasive half-shady roadside, under Eucalyptus canopy, species (Acebes Ginovés et al ., 2010) and the same frequent, 19.11.2015, F. Verloove. TENERIFE: may apply to the hybrid between them. San Cristóbal de La Laguna: La Laguna, barranco Opuntia tomentosa and O. ficus-indica , both de Santos E of TF-13, 28°28’37.12”N, polyploids (Pinkava, 2002), are also likely to be of 16°18’20.74”W, 448 m.s.m. , gravelly riverbed, fre - allopolyploid origin (Majure et al ., 2012). quent, 03.07.2014, F. Verloove; Tegueste: El So - corro, barranco de Las Cuevas, 28°31’7.80”N, Opuntia ficus-indica (L.) Mill., Gard. Dict . (ed. 16°21’39.99”W, 282 m.s.m. , woody slope, frequent, 8) No. 2. 1768. (Variant illustrated, see Figure 13). 13.01.2017, F. Verloove; Santa Úrsula: La Quinta, = O. vulgaris Ten., Syll. Fl. Neap. 239. 1831, non 28°26’7.43”N, 16°29’36.69”W, 185 m.s.m. , stony Mill. slope, few plants, 15.01.2017, F. Verloove; Bue - = O. ficus-barbarica A. Berger, Monatsschr. Kak - navista del Norte: Los Carrizales, stony slopes teenk. 22 : 181. 1912. with the two parental species, 28°19’34.84”N, 16°51’24.93”W, 25.02.2017, J.A. Reyes-Betancort. The and nomenclature of this most widely spread and highly invasive species of Op - Putative hybrids of Opuntia ficus-indica and untia in the Canary Islands are rather compli - O. tomentosa are increasingly recorded in the Ca - cated. Acebes Ginovés et al . (2010) accept both O. nary Islands. It is uncertain whether or not they maxima Mill. and O. vulgaris Mill., while both are arose spontaneously or as a result of a deliberate, often considered synonyms of, respectively, O. artificial crossing. However, in all populations in ficus-indica and O. monacantha , or even of O. hu - Gran Canaria and Tenerife where it was recorded mifusa (Guiggi, 2008). O. vulgaris is the earliest recently, both putative parental species were replacement name for Cactus opuntia L. in Opun - found growing within the flight distance of an in - tia in order to avoid the tautonym. That in turn is sect or bird. This hybrid is apparently known from considered to be the same as Cactus ficus-indica Mexico from prehistorical times (R. Mottram, per - L. Using O. vulgaris for the plant correctly called sonal communication), wherever the two species O. humifusa was a misapplication, curiously were cultivated together (see also Smith & started by Miller himself who illustrated O. hu - Figueiredo, 2012 on the species in South Africa). mifusa under the wrong name in his Icones The hybrid plants seen in the Canary Islands (Miller, 1768; Roy Mottram, personal communi - are more or less intermediate between both par - cation). ent species, although in most characters they are Opuntia ficus-indica itself is a variable closer to O. tomentosa . With the latter they share species, perhaps rather a complex of closely re - the tree-like habit and the velvety epidermis of lated entities of contested taxonomic validity. This the cladodes. However, in the hybrid the cladodes ‘species’ is technically a cultivar, having been bred are much less greyish in appearance, and they are for centuries. Its variability in the Canary Islands very spiny, features typical of forms of the other has not been studied yet in depth, but surely more putative parent, O. ficus-indica . than one taxon is present, the most notable prob - Opuntia ×elisae D. Guillot & Van der Meer ex ably being the following. D. Guillot has been described from continental Spain, and probably also has Opuntia ficus-indica Opuntia ficus-indica f. amyclaea (Ten.) Schelle, and O. tomentosa as parents (Guillot & Van der Handb. Kakteenkult. 51. 1907. Nomenclature ac - Meer, 2004; Ferrer-Gallego et al ., 2014). However, cording to Kiesling (1998). we refrain from using this name for the plants = O. megacantha Salm-Dyck, Hort. Dyck . 363. currently found in the Canary Islands since the 1834. (Figure 12). latter markedly differ, for instance in having vel - = O. maxima Salm-Dyck, Cact. Hort. Dyck. 68. vety cladodes. O. ×elisae , with glabrous stem seg - 1850, non Mill.

68 Bradleya 35/2017 Figure 22. Tephrocactus articulatus , Playa de San Juan (TF), stony slope, November 2016. Photograph by F. Verloove.

Figure 21. Oreocereus pseudofossulatus in flower, El Médano (TF), dry riverbed, June 2016. Photograph by F. Verloove.

Figure 24. Trichocereus huascha (in the back - ground Cylindropuntia pallida ), El Médano (TF), stony slope of ravine, June 2016. Photograph by F. Verloove.

Figure 23. Tephrocactus articulatus , in flower, Playa de San Juan (TF), stony slope, June 2016. Photograph by F. Verloove.

Large populations of this ‘species’ were dis - covered on a sunny slope near Ayagaures in Gran Canaria in 2015 and in an abandoned quarry in Santiago del Teide in Tenerife in 2016. It is also relatively widespread in the Anaga mountains. These records seem to confirm the previous as - sumption of Marrero et al. (1995). Compared with Opuntia ficus-indica it has ob - long to suborbicular, glaucous-pruinose, distinctly flattened cladodes that are always spiny. Molecu - Figure 25. Cereus hildmannianus , El Médano lar data seem to suggest that it probably is noth - (TF), dry riverbed, June 2015. ing else than the wild progenitor of domesticated Photograph by A. González.

Bradleya 35/2017 69 Figure 26. Cleistocactus hyalacanthus , Playa de San Juan (TF), stony slope, June 2016. Photograph by F. Verloove. Figure 27. Cleistocactus strausii, Playa de San Juan (TF), stony slope, June 2016. Photograph by F. Verloove.

Figure 29. Neogriseocereus pruinosus , Playa de San Juan (TF), stony slope, November 2016. Photograph by F. Verloove.

O. ficus-indica (Labra et al ., 2003). Other authors, however, accept it as a species distinct from O. ficus-indica , based on morphometric analyses (Reyes-Agüero et al ., 2005).

Opuntia macrocentra Engelm., Proc. Amer. Figure 28. Isolatocereus dumortieri , El Médano Acad. Arts 3: 292. 1856. (Figures 14–15). (TF), dry riverbed, June 2015. ≡ Opuntia violacea Engelm. ex B.D. Jacks., Index Photograph by A. González. Kew. 2: 358. 1895.

70 Bradleya 35/2017 var. macrocentra (Engelm.) L.D. Benson, Cacti Land et al. ; Buenavista del Norte, road to Costa Arizona (ed. 3): 21. 1969. de El Rincón, 28º22’05.40”N, 16º52’13.87”W, 25– 30 m.s.m. , 05.03.2017., J.A. Reyes-Betancort . New to the flora of the Canary Islands. A native of Mexico, Opuntia microdasys is TENERIFE: Guía de Isora: Playa de San Juan, widely cultivated as a popular and attractive or - 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony namental. It is often recorded as an escape from slope next to the sea cliffs, numerous individuals, cultivation, in Europe at least in France, Italy, 24.06.2016, F. Verloove; idem, 07.05.2016, E. Portugal, and Spain (Essl & Kobler, 2008; Novoa Ojeda-Land et al. et al ., 2015). In the Canary Islands it had only been reported before from a single locality in Originally native to the southern United Fuerteventura (Verloove & Guiggi, 2013) and States (Arizona, New Mexico, Texas) and Mexico from La Palma where initially there was some (Chihuahua, Coahuila, Sonora), Opuntia macro - doubt with regard to its identity (Salas Pascual, centra is sometimes grown as an ornamental in 2010). One of us (J.A. R.-B.) also knows O. micro - other warm and dry parts of the world (see for ex - dasys from Lanzarote where two small popula - ample Hunt, 1989; Sánchez de Lorenzo Cáceres, tions were naturalised near the road to Playa 2000). Although, with its purple stem segments Quemada and in the village of Nazaret. In Abades with spines of up to 12 or even 17cm long in Tenerife a single individual was seen in 2016 (Pinkava, 2003b), it is visually very garden-wor - where it can hardly be classified as naturalised. thy, this species is less frequent in cultivation However, in El Médano its naturalisation has than other members of Opuntia series Phaeacan - been observed and it could well become an invader thae Britton & Rose. in the future, as is the case in Australia and South Opuntia macrocentra has not been recorded Africa (Walters et al ., 2011; Novoa et al ., 2015). In outside of cultivation, apparently, although in Buenavista del Norte one individual was observed South Africa the probability of a future naturali - growing in the inclined ditch of the road. In this sation was envisaged (Walters et al ., 2011). locality other species of Opuntia (one individual In Playa de San Juan in Tenerife, Opuntia of O. pilifera , one of O. lindheimeri Engelm., and macrocentra occurs in numerous subpopulations two of O. lindheimeri var. linguiformis (Griffiths) in a relatively small area where it appears well L.D. Benson) compete with Arundo donax L., established. colonising the slopes and the water course of the Opuntia violacea Engelm. (1848) is the correct adjacent shallow ravine. None of these, however, name for this plant if the paper in which it was can be classified yet as genuinely naturalised in published by Emory is not to be considered provi - that location. sional only. Hunt et al. (2006) unilaterally deemed Opuntia microdasys is a variable species. In El it so, but the Emory paper has never actually Médano two more or less distinct forms are pres - been proposed for inclusion in the list of sup - ent. In some plants glochids are yellow or whitish, pressed works (Roy Mottram, personal communi - while in others they are reddish-brown. The lat - cation). ter is sometimes distinguished as subsp. rufida (Engelm.) U. Guzmán & Mandujano (syn.: O. ru - Opuntia microdasys (Lehm.) Pfeiff., Enum. fida Engelm.). However, glochid colour in O. mi - Diagn. Cact . 154. 1837. (Figure 16). crodasys s.str. is a variable character and genuine ≡ Cactus microdasys Lehm., Index sem. (Ham - O. rufida is a closely related, but quite different burg) 16. 1827. species (Pinkava, 2003b). Schumann was the ear - liest to make the combination O. microdasys var. New to the flora of Tenerife. Previously reported rufida , with O. rufida Engelm. as its basionym. It from Fuerteventura (Verloove & Guiggi, 2013) was Berger who was the first to apply the name O. and La Palma (Mirador del Time) (Salas Pascual, microdasys var. rufida to a brown-glochided form 2010). of O. microdasys in cultivation at La Mortola, Italy, and explicitly excluding the type of O. ru - TENERIFE: Arico: Abades, 28°8’41.56”N, fida Engelm. He therefore created a later homo - 16°26’27.67”W, 9 m.s.m. , barranco N of the vil - nym of Schumann’s name – a different entity, and lage, stony slope close to the Iglesia del Sanatorio therefore illegitimate. He can, however, be de Abades, a single individual, 21.11.2016, F. Ver - deemed to have established the cultivar name O. loove; Granadilla de Abona: Hoyas de Doña microdasys ‘Rufida’. The name subsp. rufida (En - María, Las Arenas-El Médano, 28°03’44.60”N, gelm.) U. Guzmán & Mandujano is validly pub - 16°31’44.49”W, 63 m.s.m. , 04.04.2015, E. Ojeda- lished, but it is another superfluous synonym of

Bradleya 35/2017 71 O. rufida Engelm. No valid and legitimate botan - Opuntia monacantha is easily separated by its ical name currently exists for the brown-spined rather thin, pliable, and shiny cladodes. It often form of O. microdasys (Roy Mottram, personal has a somewhat drooping appearance, doubtlessly communication). as a result of the large fruits.

Opuntia monacantha (Willd.) Haw., Suppl. pl. Opuntia pilifera F.A.C. Weber, Dict. Hort . 894. succ . 81. 1819. (Figure 17). 1898. (Figure 18). ≡ Cactus monacanthos Willd., Enum. pl . 33. 1813[1814]. New to the flora of the Canary Islands. ≡ Opuntia vulgaris Britton & Rose non Mill., Cact. 1: 156. 1919 GRAN CANARIA: San Bartolomé de Tirajana: San Agustín, 27°46’12.73”N, 15°33’5.47”W, 48 Confirmation for the flora of the Canary Islands. m.s.m. , urbanización Colegio Arenas Sur, dry stony slope, a large clone, 22.11.2015, F. Verloove; TENERIFE: San Cristóbal de La Laguna: Tejina, San Bartolomé de Tirajana: Bahía Feliz, charcas de Tejina, 28°32’12.33”N, 16°21’31.02”W, 27°47’7.45”N, 15°30’51.28”W, 10 m.s.m. , roadside 155 m.s.m. , talus slope, scattered individuals, GC-500, a single clone (cultivated in central reser - 09.01.2017, F. Verloove; idem., 06.05.2008, J.A. vation,), 27.11.2015, F. Verloove; TENERIFE: Reyes-Betancort; Puerto de la Cruz, La Mon - Buenavista del Norte: Costa de El Rincón, tañeta, 28°23’13.97”N, 16°33’41.77”W, 306 28º22’13.21”N, 16º52’15.61”W, 15 m.s.m. , m.s.m. , rough ground, c. 10-15 individuals, 09.02.2017., E. Ojeda-Land et al .; idem, 12.01.2017, F. Verloove; Santa Úrsula: La Quinta, 05.03.2017, J.A. Reyes-Betancort . close to calle Codeso, 28°25’59.06”N, 16°29’42.92”W, 191 m.s.m. , shrubland, small pop - Opuntia pilifera is a Mexican species that has ulation, 15.01.2017, F. Verloove; Tacoronte: Mesa been reported from Puebla and Oaxaca. Outside del Mar, calle Mesa del Mar, 28°30’1.98”N, of its native distribution range it is widely culti - 16°25’11.71”W, 252 m.s.m. , small population, vated as an ornamental in warm-temperate parts 18.01.2017, F. Verloove. of the world, also in southern Europe, although it is surprisingly omitted by, for instance, Hunt This South American species (Argentina and (1989) and Sánchez de Lorenzo Cáceres (2000). It Brazil) is widely cultivated as an ornamental, also has been reported in the wild from France (Tison in the Canary Islands, although it was surpris - & de Foucault, 2014), Italy (Manni & Guiggi, ingly omitted by Sánchez de Lorenzo Cáceres 2015), Spain (Guillot & Lodé, 2009; Sánchez-Gul - (2000). lón et al ., 2014) and Malta (see: Acebes Ginovés et al . (2010) do not cite Opun - http://www.maltawildplants.com/CACT/Opun - tia monacantha for the Canarian flora. Their ci - tia_pilifera.php ). It is here reported for the first tation of O. vulgaris , however, most likely refers time from the Canary Islands. In Gran Canaria to O. monacantha (see Kunkel, 1972). O. vulgaris single clones were encountered in two nearby lo - is a confusing name; according to Leuenberger calities, while in Tenerife six clones have become (1993) it is conspecific with O. ficus-indica , while established on the slope of a ravine, as escapes Hunt et al. (2006) wrongly consider it a synonym from the adjacent living fence of a cactus nursery . of O. humifusa (Raf.) Raf. (see above). Opuntia pilifera is a characteristic species. Its In Tenerife, Opuntia monacantha was observed glabrous cladodes are obovate to suborbicular, and in several localities in the northern part of the is - large (up to 35cm long); the areoles are covered land, usually in small numbers and in the vicinity with whitish or yellowish, silky trichomes; and its of houses. Most populations probably represent flowers are pinkish, turning purplish (Arias et al ., relics of former cultivation, or are established from 2012). discarded garden waste. The species seems much less vigorous than the superficially similar, but Opuntia robusta J.C. Wendl., Cat. Hort. Her - much larger-growing, O. ficus-indica . However, it renh . 1835. (Figure 19). is one of the most widely spread invasive cacti worldwide (Novoa et al ., 2015). For instance, it is a Confirmation for the flora of the Canary Islands. serious invader in Australia, India, Madagascar, Mauritius, South Africa, and Sri Lanka, although, TENERIFE: Puerto de la Cruz, SW of calle at least in some countries, it has decreased since Sarasate, 28°24’24.64”N, 16°32’15.61”W, 127 the introduction of a cochineal species, Dactylop - m.s.m. , rough ground, a single large clone of c. 20 ius ceylonicus (Walters et al ., 2011). m², 12.11.2014 and 12.01.2017, F. Verloove;

72 Bradleya 35/2017 Figure 30. Pilosocereus pachycladus , Playa de Figure 31. Thelocactus hastifer , Playa de San San Juan (TF), stony slope, November 2016. Juan (TF), stony slope, November 2016. Photograph by F. Verloove. Photograph by F. Verloove.

Figure 33. Trichocereus peruvianus , Puerto de la Cruz (TF), roadside slope, January 2017. Photograph by F. Verloove.

Figure 32. Trichocereus bridgesii , Chayofa (TF), vacant lot in residential area, November 2016. Photograph by F. Verloove.

Granadilla de Abona: Hoyas de Doña María, Las Arenas-El Médano, 28°03’44.59”N, 16°31’ 44.63”W, 62 m.s.m. , 04.04.2015, E. Ojeda-Land et al. ; Buenavista del Norte: Costa de El Rincón, 28º22’13.21”N, 16º52’15.61”W, 15 m.s.m. , 05.03.2017, J.A. Reyes-Betancort .

This Mexican species has been reported from Gran Canaria, La Palma, and Tenerife in the Ca - Figure 34. Trichocereus vollianus , Playa de San nary Islands (Acebes Ginovés et al. , 2010). How - Juan (TF), stony slope, November 2016. ever, it is rather rarely grown as an ornamental Photograph by F. Verloove.

Bradleya 35/2017 73 and most claims may refer to O. ficus-indica f. New to the flora of the Canary Islands. amyclaea (syn.: O. megacantha ) and other forms of this species. The authors have only seen it in TENERIFE: Guía de Isora: Playa de San Juan, three naturalised populations in Tenerife. Com - 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony pared with O. ficus-indica , cladodes in O. robusta slope next to the sea cliffs, numerous individuals, are larger and very glaucous, almost orbicular and 24.06.2016, F. Verloove; idem., 07.05.2016, E. its areoles are widely scattered and somewhat de - Ojeda-Land et al. pressed. In the El Médano locality, some clones of a putative O. robusta hybrid have also been ob - Tephrocactus articulatus , which is native to served. Argentina, is sometimes grown as an ornamental Opuntia robusta is an invasive weed in several in warm and dry areas of the world. Outside of its countries, including Australia, Botswana, and native area it has been reported as an invasive South Africa (Walters et al ., 2011; Novoa et al ., alien in South Africa and Namibia (Walters et al ., 2015). In Europe it is only rarely reported in the 2011). wild. Guiggi (2014) provides details for Italy. In Playa de San Juan numerous individuals of Tephrocactus articulatus are found growing on Oreocereus pseudofossulatus D.R. Hunt, bare, sun-exposed slopes close to the sea cliffs. Its Bradleya 9: 89. 1991. (Figures 20–21). stem segments easily detach, which enables rapid = Cleistocactus fossulatus Mottram, Chileans vegetative dispersal. The species is well estab - 13 (43): 30. 1985. lished at this location. [non Pilocereus fossulatus Labour., Rev. Hort. ser. Tephrocactus articulatus , the most widely 4 4(2): 24–25. 1855; Oreocereus fossulatus grown species of this genus, is very variable and (Labour.) Backeb., Kakt.-Freund 3(2): 13–14. 1934 several varieties have been described. The plants = Oreocereus celsianus (Salm-Dyck) Riccob.] found in Tenerife have long, distinctly flattened spines, a feature of T. articulatus var. papyracan - New to the flora of the Canary Islands. thus (Phil.) Backeb. (see for example Kiesling, 1984). Although most recent workers refrain from TENERIFE: Granadilla de Abona: Hoyas de Doña using any infraspecific names (e.g. Gilmer & María, Las Arenas-El Médano, 28°03’39.22”N, Thomas, 1998; Gorelick & Mahr, 2013), this par - 16°31’43.06”W, 54 m.s.m. to 28°03’36.80”N, ticular variation may merit at least forma status, 16°31’41.68”W, 50 m.s.m. , 04.04.2015, 04.05.2015, for which a new combination is established here. 21.05.2015, 09.06.2015, E. Ojeda-Land et al. The generic limits of Tephrocactus and other opuntoid cacti have long been under debate. Mo - As mentioned above the presence of Oreo - lecular data, however, have shown that its mem - cereus pseudofossulatus in El Médano arose from bers form a distinct subclade (Nyffeler & Eggli, an abandoned, not well delimited cultivation ex - 2010; Ritz et al ., 2012). periment. Birds that feed on t he fleshy fruits could enhance its expansion, although, at present, Trichocereus huascha (F.A.C. Weber) Britton it is only locally naturalised. & Rose, Cact. 2: 142f. 208–210. 1920. (Figure 24). Oreocereus pseudofossulatus is not known as ≡ huascha (F.A.C. Weber) Friedrich & an invasive species (Novoa et al ., 2015), nor has it G.D. Rowley, I.O.S. Bull. 3(3) : 95. 1974. been reported before from Europe (Essl & Kobler, 2008). It is often grown for ornament. New to the flora of the Canary Islands.

Tephrocactus articulatus (Pfeiff.) Backeb., TENERIFE: Granadilla de Abona: Hoyas de Cactus (Paris) 38 : 249. 1953. (Figures 22–23). Doña María, Las Arenas-El Médano, ≡ Opuntia articulata (Pfeiff.) D.R. Hunt, Bradleya 28°03’43.29”N, 16°31’44.87”W, 60 m.s.m. to 5: 93. 1987. 28°03’36.70”N, 16°31’41.68”W, 54 m.s.m. , f. papyracanthus (Phil.) Guiggi & Verloove 04.04.2015, 04.05.2015, 21.05.2015, 09.06.2015, comb. nov. E. Ojeda-Land et al. Basionym: Opuntia papyracantha Phil., Gartenflora 21 : 129. A native of Argentina, this attractive orna - 1872. mental species was found in a dry riverbed in El Synonym: Médano in Tenerife, apparently as a relic of an Tephrocactus articulatus (Pfeiff.) Backeb. var. abandoned cultivation experiment. The species papyracanthus (Phil.) Backeb., Cactus (Paris) 38 : looks rather prolific and now also occurs on the 249. 1953. surrounding rocky slopes.

74 Bradleya 35/2017 Trichocereus huascha is not known as an in - Cleistocactus hyalacanthus (K.Schum.) Rol.- vasive species (Novoa et al ., 2015), nor has it been Goss., Bull. Mens. Soc. Centr. Agric. Hort. Acclim. reported before from Europe (Essl & Kobler, Nice Alpes-Marit . 44 : 33. 1904. (Figure 26) . 2008). It is widely grown for ornament, also in ≡ Cereus hyalacanthus K. Schum., Gesamtbeschr. continental Spain (Sánchez de Lorenzo Cáceres, Kakt . 101. 1897. 2000). Columnar cacti with large flowers that were TENERIFE: Granadilla de Abona: Hoyas de Doña accommodated in Trichocereus (including T. María, Las Arenas-El Médano, 28º03’38,50’’ N, huascha ) have been transferred to Echinopsis 16º31’42,67’’ W, 53 m.s.m. , 04.04.2015, E. Ojeda- based on molecular data (Schlumpberger & Ren - Land et al. ; Guía de Isora: Playa de San Juan, ner, 2012). However, pending additional data, Rit - 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony ter’s concept of Trichocereus is here followed slope next to the sea cliffs, scattered individuals, (Ritter, 1980). 24.06.2016, F. Verloove.

Part 2. Casual taxa Scattered individuals of this South American species (Argentina, Bolivia) were found growing In Part 2, cactus species are presented that ap - on a stony slope close to the sea cliffs in Playa de pear to be non-problematic, neither at present, San Juan and in a dry riverbed in El Médano in nor in the future. None of these have been Tenerife. In both localities the plants persist from recorded before from the Canary Islands. In most an abandoned plantation. They were found to - cases single or a few individuals only were en - gether with the congeneric and superficially sim - countered of these species, most of which are prob - ilar-looking Cleistocactus strausii (see below). The ably unable to spread sexually in the absence of latter, however, has longer corollas (80–90 vs 30– species-specific pollinators. These species may 40mm), more numerous costae (25–30 vs c. 20), persist for some time in these localities, but future and its spines are white, not brownish. expansion of their ranges is rather unlikely. De - spite being widely grown as ornamentals (see for Cleistocactus strausii (Heese) Backeb., Kak - example Hunt, 1989; Sánchez de Lorenzo Cáceres, teen-freund 3: 121. 1934. (Figure 27). 2000), none is known as an invasive species ≡ Pilocereus strausii Heese, Gartenflora 56 : 410. (Novoa et al ., 2015). Hence, the entries for these 1907. species are brief, but they are all illustrated. TENERIFE: Granadilla de Abona: Hoyas de Cereus hildmannianus K. Schum., Fl. Bras. Doña María, Las Arenas-El Médano, 4(2) : 202. 1890. (Figure 25). 28°03’43.29”N, 16°31’44.87”W, 60 m.s.m. to Cereus peruvianus auct. non (L.) Mill. 28°03’36.70”N, 16°31’41.68”W, 54 m.s.m. , 04.04.2015, E. Ojeda-Land et al .; Guía de Isora: TENERIFE: Granadilla de Abona: Hoyas de Doña Playa de San Juan, 28°10’41.38”N, María, Las Arenas-El Médano, 28°03’44.60”N, 16°48’33.98”W, 48 m.s.m. , stony slope next to the 16°31’44.4987”W, 63 m.s.m. , 04.04.2015, E. Ojeda- sea cliffs, scattered individuals, 24.06.2016, F. Land et al. Verloove.

A few plants of the ‘Monstruosus’ form of this Originally native in Bolivia, Cleistocactus South American species were seen in El Médano strausii is widely cultivated as an ornamental. in Tenerife where it persists from an abandoned Scattered individuals are found on a stony slope in cultivation experiment. This and several other Playa de San Juan and in a dry riverbed in El Mé - species of Cereus are reputed invasive species dano in Tenerife. The plants grow together with (Novoa et al. , 2015). In Europe, Cereus hildman - C. hyalacanthus . Both persist from an abandoned nianus has been reported before from Italy plantation. (Guiggi, 2010). Application of this name for the well-known Isolatocereus dumortieri (Scheidw.) Backeb., plant of cultivation is contentious. Hunt et al. Cactaceae (Berlin) 1941(2) : 47. 1942. (Figure 28) (2006) have championed the use of this name, but ≡ dumortieri (Scheidw.) Buxb., Bot. it is doubtfully different from C. jamacaru DC., or Stud. 12 : 92. 1961. even from C. hexagonus (L.) Mill. This present- ≡ Lemaireocereus dumortieri (Scheidw.) Britton & day cultivar is so widely cultivated from ancient Rose, Contr. U.S. Natl. Herb. 12(10) : 425. 1909. times that its true origin is lost in time (Roy Mot - ≡ Rathbunia dumortieri (Scheidw.) P.V. Heath, tram, personal communication). Calyx 2: 104. 1992.

Bradleya 35/2017 75 ≡ Cereus dumortieri Scheidw., Hort. Belge 4: 220. posed stony slope in Playa de San Juan in Tener - 1837. ife, where they persist from an abandoned plan - tation. TENERIFE: Granadilla de Abona: Hoyas de Doña María, Las Arenas-El Médano, 28°03’44.60”N, Thelocactus hastifer (Werderm. & Boed.) F.M. 16°31’44.4987”W, 63 m.s.m. , 04.04.2015, E. Ojeda- Knuth, Kaktus-ABC 360. 1935. (Figure 31). Land et al . ≡ Echinocactus hastifer Werderm. & Boed., No - tizbl. Bot. Gart. Berlin–Dahlem 11 : 274. 1931. A few plants of this Mexican species grow in a dry riverbed in El Médano in Tenerife where they TENERIFE: Guía de Isora: Playa de San Juan, persist after an abandoned cultivation experi - 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony ment. slope next to the sea cliffs, a single clone, Isolatocereus dumortieri was long included in 24.06.2016, F. Verloove. Stenocereus , but flower morphology and molecu - lar data seem to support its segregation (Cota & Scattered individuals of this Mexican species Wallace, 1997). grow on a sun-exposed stony slope in Playa de San Juan in Tenerife, persisting from an abandoned Neogriseocereus pruinosus (Otto ex Pfeiff.) plantation. Guiggi, Cactology 3, Suppl. 4: 1. 2013. (Figure 29). ≡ Griseocereus pruinosus (Otto ex Pfeiff.) Guiggi, Trichocereus bridgesii (Salm-Dyck) Britton & Cactology 3: 8. 2012. Rose, Cact . 2: 134. 1920. (Figure 32). ≡ Stenocereus pruinosus (Otto ex Pfeiff.) Buxb., ≡ Cereus bridgesii Salm-Dyck, Cact. Hort. Dyck . Bot. Stud . 12 : 101. 1961. 208. 1849[1850]. ≡ Echinocactus pruinosus Otto ex Pfeiff., Enum. Diagn. Cact . 54. 1837. TENERIFE: Arona: Chayofa, 28° 4’33.03”N, 16°41’28.13”W, 293 m.s.m. , dry slope adjacent to TENERIFE: Granadilla de Abona: Hoyas de Doña calle Barranco del Verodal, three individuals, María, Las Arenas-El Médano, 28°03’44.60”N, 22.11.2016, F. Verloove; Arona: Chayofa, 28° 16°31’44.4987”W, 63 m.s.m. , 04.04.2015, E. Ojeda- 4’28.99”N, 16°41’38.39”W, 280 m.s.m. , on the Land et al .; Guía de Isora: Playa de San Juan, verge of vacant lot in residential area, two indi - 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony viduals, 22.11.2016, F. Verloove. slope next to the sea cliffs, scattered individuals, 24.06.2016, F. Verloove. A few individuals of this Bolivian species grow in two different localities in Chayofa in Tenerife; Scattered individuals of this impressive in both instances material probably became es - species from Central America (Guatemala, Hon - tablished from discarded garden waste. duras, Mexico) grow on a sun-exposed stony slope in Playa de San Juan and in a dry riverbed in El Trichocereus peruvianus Britton & Rose, Cact . Médano in Tenerife, persisting from an aban - 2: 136, f. 197. 1920. (Figure 33). doned plantation and cultivation experiment. ≡ Echinopsis peruviana Friedrich & G.D. Rowley, This species was long known as Stenocereus I.O.S. Bull . 3(3): 97. 1974. pruinosus . However, the generic limits of Steno - cereus are debated and this species and some rel - TENERIFE: Puerto de la Cruz, calle Chahorra, atives are best segregated (Heath, 1996; Guiggi, 28°24’21.56”N, 16°33’56.77”W, 67 m.s.m. , roadside 2012, 2013). slope, five individuals, 16.01.2017, F. Verloove.

Pilosocereus pachycladus F. Ritter, Kakteen Five individuals of this Peruvian species grow Südamer . 1: 70. 1979. (Figure 30). on a slope in Puerto de la Cruz in Tenerife, prob - ably from discarded garden waste. TENERIFE: Guía de Isora: Playa de San Juan, 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony Trichocereus vollianus Backeb., Kaktus-ABC slope next to the sea cliffs, scattered individuals, 205, 412. 1935. (Figure 34). 24.06.2016, F. Verloove; idem, 07.05.2016, E. Ojeda-Land et al . TENERIFE: Guía de Isora: Playa de San Juan, 28°10’41.38”N, 16°48’33.98”W, 48 m.s.m. , stony Scattered individuals of this attractive, widely slope next to the sea cliffs, scattered individuals, cultivated Brazilian species grow on a sun-ex - 24.06.2016, F. Verloove.

76 Bradleya 35/2017 Scattered individuals of this Bolivian species pecies exóticas invasoras en España. Diag - grow on a sun-exposed stony slope in Playa de nóstico preliminar y bases de conocimiento San Juan in Tenerife, persisting from an aban - sobre impacto y vulnerabilidad. Oficina Espa - doned plantation. ñola de Cambio Climático, Ministerio de Medio Ambiente y Medio Rural y Marino. Ma - Acknowledgements drid, 146 pp. [accessed at http://www.ma - Daniela Zappi (Brazil), Daniel Guillot Ortiz grama.gob.es/es/cambio-climatico/temas/impa (Spain) and Emilio Laguna (Spain) are thanked ctosvulnerabilidad-yadaptacion/Especies_in - for providing valuable comments on Pilosocereus vasoras_tcm7-197788_tcm7-217991.pdf ; 4 pachycladus and Opuntia ×elisae , respectively. April 2016]. Fabián Hernández and Beatriz Fariña accompa - CHINNOCK , R.J. (2015). Feral opuntioid cacti in nied one of the authors (R.B.) in Playa de San Australia: Part I. Cylindrical-stemmed ge - Juan (Tenerife), when some cacti were discov - nera: Austrocylindropuntia , Cylindropuntia ered. Juana María González Mancebo and and Corynopuntia . Journal of the Adelaide Leopoldo Moro Abad, alerted us to the presence Botanic Gardens Supplement 3: 1–69. of some cacti in El Médano and Amarilla Golf, re - COTA , J.H. & W ALLACE , R.S. (1997). Chloroplast spectively. DNA evidence for divergence in Ferocactus Roy Mottram (United Kingdom) is thanked for and its relationships to North American co - useful discussions on, and assistance with, the lumnar cacti (Cactaceae: ). Sys - identification of some of the cactus species re - tematic Botany 22 : 529–542. ported here. He also contributed valuable com - DEL -A RCO , M. (2008). 4. La flora y la vegetación ments on a first version of the manuscript. canaria ante el cambio climático actual. In: AFONSO -C ARRILLO , J. (ED.). Naturaleza ame - References nazada por los cambios en el clima. Pp. 105– ACEBES GINOVéS , J.R., L EóN ARENCIBIA , M.C., R O- 140. Actas III Semana Científica Telesforo DRíGUEZ NAVARRO , M.L . et al . (2010). Pterido - Bravo. Instituto de Estudios Hispánicos de phyta, Spermatophyta. In: ARECHAVALETA , M., Canarias. RODRíGUEZ , S., Z URITA , N. & G ARCíA , A. (EDS ), ERIKSSON , O., H ANSEN , A. & S UNDING , P. (1979). Lista de especies silvestres de Canarias (hon - Flora of Macaronesia: checklist of vascular gos, plantas y animales terrestres) . (2 nd ed.). plants (2 nd ed.). Botanical Garden and Mu - Pp. 119–172. Gobierno de Canarias, La La - seum, University of Oslo, Oslo. guna. ESSL , F. & K OBLER , J. (2008). Spiny invaders – ARIAS , S., G AMA , S., V áZQUEZ , B. & G UZMáN , L.U. Patterns and determinants of cacti invasion (2012). Flora del Valle de Tehuacán-Cuicatlán in Europe. Flora - Morphology Distribution 95 : 1–235. Functional Ecology of Plants 204(7) : 485–494. BELTRáN TEJERA , E., W ILDPRET DE LA TORRE , W., FERRER -G ALLEGO , P.P., L AGUNA LUMBRERAS , E. & LEóN ARENCIBIA , M.C., G ARCíA GALLO , A. & GUILLOT ORTIZ , D. (2014). Sobre el híbrido REyES HERNáNDEZ , J. (1999). Libro Rojo de la Opuntia ×elisae D. Guillot & Van der Meer ex Flora Canaria contenida en la Directiva-Há - D. Guillot (Cactaceae). Bouteloua 17 : 42–48. bitats Europea. Organismo Autónomo Par - GILMER , K. & T HOMAS , H.-P. (1998). Die Gattung ques Nacionales . Madrid. Tephrocactus Lemaire s. str. – Taxonomie, B.O.E. (2013). Real Decreto 630/2013, de 2 de Ökologie und Kultur. Schumannia 2: 85–141. agosto, por el que se regula el Catálogo español GONZáLEZ MONTELONGO , C., L EóN -A RENCIBIA , de especies exóticas invasoras [accessed at: M.C., A CEBES GINOVéS , J.R. & L OSADA -L IMA , http://www.boe.es/boe/dias/2013/08/03/pdfs/B A. (2014). Comentarios para la actualización OE-A-2013-8565.pdf ; 12 December 2016]. del catálogo de plantas vasculares de las islas BRAMWELL , D. (1976). The endemic flora of the Canarias. Vieraea 42 : 207–218. Canary Islands. In: KUNKEL , W.H. (ED.), Bio - GORELICK , R. & M AHR , D. (2013). Tephrocactus . geography and ecology in the Canary Islands . Cact. Succ. J. (US) 85(6) : 262–276. Dr. W. Junk, The Hague. GUIGGI , A. (2008). Catalogo delle Cactaceae natu - BRAVO -H OLLIS , H. & A RIAS , S. (2011). Flora Me - ralizzate in Italia con osservazioni tassonomi - soamericana , Volumen 2(1) . Universidad Na - che, nomenclaturali e corologiche. Rivista cional Autónoma de México, México D.F. Piemontese di Storia Naturale 29 : 103–140. BRITTON , N.L. & R OSE , J.N. (1919–23). The Cac - GUIGGI , A. (2010). Aggiunte e correzioni al Cata - taceae , vols. 1–4. Carnegie Inst. Publ. 248. logo delle Cactaceae naturalizzate in Italia. CAPDEVILA -A RGüELLES L., Z ILLETTI , B. & S UáREZ Rivista Piemontese di Storia Naturale 31 : 35– áLVAREZ , V.A. (2011). Cambio climático y es - 54 .

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