new records of naturalised and invasive cacti (cactaceae ......c/. eduardo zamacois, 13-3ºa, 38005...

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Summary: Recent field work in Gran Canaria and Tenerife (Canary Islands, Spain) yielded records for 30 alien taxa of Cactaceae that had not been reported before, either from the whole area, or from one of the islands. Out of these, 17 are con- sidered locally naturalised and/or potentially in- vasive: Cylindropuntia bigelovii, C. fulgida, C. pallida, C. prolifera, C. tunicata, Echinocereus rigidissimus, Haageocereus kagenekii, Hylocereus triangularis, Opuntia basilaris, O. elatior, O. ficus-indica × O. tomentosa, O. macrocentra, O. microdasys, O. pilifera, Oreocereus pseudofossu- latus, Tephrocactus articulatus and Trichocereus huascha. The same applies to a rather character- istic form of O. ficus-indica that sometimes is re- ferred to as f. amyclaea (syn.: O. megacantha). The presence of Opuntia monacantha and O. robusta, two species with a dubious status in the Canary Islands, is confirmed. Ten further taxa are con- sidered casuals, often relics of cultivation. All taxa are illustrated, and for the naturalised and/or po- tentially invasive taxa additional information is provided. Two new combinations are proposed for Cylindropuntia fulgida f. mamillata and Tephro- cactus articulatus f. papyracanthus. Zusammenfassung: Bei jüngsten Feldstudien auf Gran Canaria und Teneriffa (Kanarische Inseln, Spanien) wurden 30 nicht indigene Taxa von Cactaceae nachgewiesen, die bisher weder vom Gesamtarchipel noch von einer der beiden Inseln gemeldet waren. Von diesen werden 17 als lokal eingebürgert und / oder potenziell invasiv ange- sehen: Cylindropuntia bigelovii, C. fulgida, C. pal- lida, C. prolifera, C. tunicata, Echinocereus rigidissimus, Haageocereus kagenekii, Hylocereus triangularis, Opuntia basilaris, O. elatior, O. ficus-indica × O. tomentosa, O. macrocentra, O. microdasys, O. pilifera, Oreocereus pseudofossu- latus, Tephrocactus articulatus und Trichocereus huascha. Das Gleiche gilt für eine sehr charakte- ristische Form von O. ficus-indica, die manchmal als f. amyclaea (syn.: O. megacantha) bezeichnet wird. Zehn weitere Taxa, oft Anbaurelikte, gelten als nicht eingebürgert. Alle Taxa sind abgebildet, zu den eingebürgerten und / oder potenziell inva- siven Taxa erfolgen weitere Informationen. Zwei neue Kombinationen werden vorgeschlagen: Cy- lindropuntia fulgida f. mamillata und Tephro- cactus articulatus f. papyracanthus. 58 Bradleya 35/2017 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])

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Page 1: New records of naturalised and invasive cacti (Cactaceae ......C/. Eduardo Zamacois, 13-3ºA, 38005 Santa Cruz de Tenerife, Canary Islands, Spain. (email: makaronesia68@yahoo.es) Introduction

Summary: Recent field work in Gran Canaria andTenerife (Canary Islands, Spain) yielded recordsfor 30 alien taxa of Cactaceae that had not beenreported before, either from the whole area, orfrom one of the islands. Out of these, 17 are con-sidered locally naturalised and/or potentially in-vasive: Cylindropuntia bigelovii, C. fulgida, C.pallida, C. prolifera, C. tunicata, Echinocereusrigidissimus, Haageocereus kagenekii, Hylocereustriangularis, Opuntia basilaris, O. elatior, O.ficus-indica × O. tomentosa, O. macrocentra, O.microdasys, O. pilifera, Oreocereus pseudofossu-latus, Tephrocactus articulatus and Trichocereushuascha. The same applies to a rather character-istic form of O. ficus-indica that sometimes is re-ferred to as f. amyclaea (syn.: O. megacantha). Thepresence of Opuntia monacantha and O. robusta,two species with a dubious status in the CanaryIslands, is confirmed. Ten further taxa are con-sidered casuals, often relics of cultivation. All taxaare illustrated, and for the naturalised and/or po-tentially invasive taxa additional information isprovided. Two new combinations are proposed forCylindropuntia fulgida f. mamillata and Tephro-cactus articulatus f. papyracanthus.

Zusammenfassung: Bei jüngsten Feldstudien aufGran Canaria und Teneriffa (Kanarische Inseln,Spanien) wurden 30 nicht indigene Taxa vonCactaceae nachgewiesen, die bisher weder vomGesamtarchipel noch von einer der beiden Inselngemeldet waren. Von diesen werden 17 als lokaleingebürgert und / oder potenziell invasiv ange-sehen: Cylindropuntia bigelovii, C. fulgida, C. pal-lida, C. prolifera, C. tunicata, Echinocereusrigidissimus, Haageocereus kagenekii, Hylocereustriangularis, Opuntia basilaris, O. elatior, O.ficus-indica × O. tomentosa, O. macrocentra, O.microdasys, O. pilifera, Oreocereus pseudofossu-latus, Tephrocactus articulatus und Trichocereushuascha. Das Gleiche gilt für eine sehr charakte-ristische Form von O. ficus-indica, die manchmalals f. amyclaea (syn.: O. megacantha) bezeichnetwird. Zehn weitere Taxa, oft Anbaurelikte, geltenals nicht eingebürgert. Alle Taxa sind abgebildet,zu den eingebürgerten und / oder potenziell inva-siven Taxa erfolgen weitere Informationen. Zweineue Kombinationen werden vorgeschlagen: Cy-lindropuntia fulgida f. mamillata und Tephro-cactus articulatus f. papyracanthus.

58 Bradleya 35/2017

Bradleya 35/2017pages 58–79

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

Filip Verloove1, Elizabeth Ojeda-Land2, Gideon F. Smith3,4, AlessandroGuiggi5, Jorge Alfredo Reyes-Betancort6, Carlos Samarín2, AntonioGonzález Hernández7 & Rubén Barone81. 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. 38071Santa 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, 6031South 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])

Page 2: New records of naturalised and invasive cacti (Cactaceae ......C/. Eduardo Zamacois, 13-3ºA, 38005 Santa Cruz de Tenerife, Canary Islands, Spain. (email: makaronesia68@yahoo.es) Introduction

IntroductionThe Canary Islands, Spain, have a well-docu-

mented and rich native flora with many local,highly specialized endemic species (e.g.Bramwell, 1976; Beltrán et al., 1999). In addition,as a result of their subtropical climate, the is-lands are an outstanding cultivation ground forhundreds of ornamental subtropical plantspecies. An increasing number of these are notonly well-adapted to be cultivated in the local cli-mate, but are indeed reproducing locally withouthuman intervention, eventually naturalising,spreading, or even becoming invasive. In pastyears the knowledge about these alien specieshas considerably increased and numerous ac-counts have been published (e.g. Verloove &Reyes-Betancort, 2011; Santos-Guerra et al.,2013; Scholz et al., 2013; Siverio Núñez et al.,2013; Verloove, 2013; González Montelongo et al.,2014; Santos-Guerra et al., 2014; Santos-Guerra& Reyes-Betancort, 2014; Otto & Verloove, 2016).However, some taxonomic groups are still insuf-ficiently known. This certainly holds true for in-troduced succulents, despite the fact that theislands have an ideal climate that allows the per-sistence and acclimatization of numerous species,especially those from arid and hot habitats. Thesouthern parts of Tenerife, for instance, have cli-matological circumstances comparable with thosein Arizona, U.S.A. Unsurprisingly, the islandhouses the most important producer for cacti andsucculents worldwide (Canary Cactus S.A.).Moreover, Spain is considered one of the three in-vasion hot spots for cacti in the world (Novoa etal., 2015; see also Essl & Kobler, 2008).

Not a single species of the family Cactaceae isnative to the Canary Islands, although twospecies have become a very familiar sight. Opun-tia ficus-indica (L.) Mill. and O. dillenii (Ker-Gawl.) Haw. were introduced quite long ago, andnow have invaded large parts of the islands.While O. dillenii is mostly confined to the loweraltitudes, the former is omnipresent, even athigher altitudes. Eriksson et al. (1979), latermerely repeated with a nomenclatural update byAcebes Ginovés et al. (2010), cited the names ofnine cactus species for the Canary Islands: Aus-trocylindropuntia cylindrica (Lam.) Backeb., A.exaltata (Berg) Backeb., Hylocereus undatus(Haw.) Britton & Rose, Opuntia dillenii, O. max-ima Mill., O. robusta H.L. Wendland, O. tomen-tosa Salm-Dyck, O. tuna (L.) Mill., and O.vulgaris Mill. Of these, O. maxima auct. and O.vulgaris auct. non Mill. are now referred to O.ficus-indica [although the binomial O. vulgarishas also been associated with O. monacantha

(Willd.) Haw., also in the Canary Islands; seelater]. Two others, O. tuna and Austrocylindrop-untia cylindrica, probably refer to respectivelyerroneous records, or are only very locally natu-ralised. The remaining species have been con-firmed by the authors of the present paper. In thepast few years some additional species of cactihave been recorded: Cylindropuntia prolifera(Engelm.) F.M. Knuth and Opuntia microdasys(Lehm.) Pfeiff. from Fuerteventura (Verloove &Guiggi, 2013), O. leucotricha DC. from Gran Ca-naria, La Palma, and Tenerife (Verloove, 2013;Santos-Guerra et al., 2014), and Espostoamelanostele from Tenerife (Verloove, 2016).

In the past few years some of us discoveredseveral escaped cacti in Tenerife and Gran Ca-naria that had not previously been recorded inthe Canary Islands. Their identities were as-sessed in close collaboration with two of the au-thors (G.F. Smith and A. Guiggi). In the presentpaper 30 taxa are presented and illustrated,most of them not previously reported from theCanary Islands. Their degree of naturalisationvaries from ephemerals to locally, or potentially,invasive aliens. In the absence of species-specificpollinators, some of the species are able to spreadby vegetative means, and their establishmentcould be favoured by climate change (Capdevila-Argüelles et al., 2011; Del-Arco, 2008). Severalspecies of cacti are among the most damaging in-vasive plant species in the world (Novoa et al.,2015) and some, for instance the whole genusCylindropuntia, are included in Spanish legisla-tion on invasive alien species (B.O.E., 2013).Therefore, it is important to document any in-cipient invasion event of cacti, even in earlystages.

It is noteworthy that likely most, if not all, theplants catalogued in this paper have a cultivatedorigin, including from commercial nurseries withpropagation facilities on the Canary Islands. It istherefore possible that, at least in some in-stances, the plants may not perfectly agree withbotanical taxa as known from the wild (Roy Mot-tram, personal communication). In these circum-stances, hybridisation through open pollinationin fields may well have given rise to hybrids that,sometimes through introgression and backcross-ing, resemble one of the parents.

Materials and methodsOnce harvested, virtually all species of cacti

are awkward to preserve as dried specimens fordeposition in a herbarium, and most of the recordshere reported are only documented with photo-graphs, unless otherwise stated.

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The presence or absence on the islands of GranCanaria or Tenerife was each time compared withdata provided by Acebes Ginovés et al. (2010), aswell as other literature sources. The paper is di-vided in two parts in both of which the taxa arearranged in alphabetical sequence. Naturalisedand/or potentially invasive taxa are dealt with inthe first part; these are treated in more detail thanthe taxa recorded as casuals in the second part.

Each entry includes: [1.] the scientific name ofthe taxon (accompanied by one or more synonyms,if deemed desirable); [2.] type of chorological nov-elty and estimated degree of naturalisation (sensuRichardson et al., 2000); [3.] enumeration of local-ities; [4.] origin of the taxon; and [5.] details aboutits secondary distribution.

Nomenclature of the taxa presented is mostlyin accordance with recent insights, often based onmolecular phylogenetic studies. Authorities ofplant names usually follow Tropicos (www.tropi-cos.org).

Results

Part 1. Naturalised and/or potentially inva-sive taxa

In this part those taxa are presented that ei-ther are naturalised locally, or are likely to be-come so in the near future. Some have at presenta very limited distribution range, but have beenrecorded as invasive in climatologically similarareas of the world, or are found in, or near, vul-nerable natural habitats on the islands.

Cylindropuntia bigelovii (Engelm.) F.M.Knuth, Kaktus-ABC 125. 1935. (Figure 1).≡ Opuntia bigelovii Engelm., Proc. Amer. Acad.Arts 3: 307. 1856.

New to the flora of the Canary Islands.

TENERIFE: Güímar, 28°18’34.15”N,16°22’57.44”W, 113 m.s.m., Camino de SamarinesE of bridge over TF-1 motorway, roadside, a singleindividual, 23.11.2016, F. Verloove.

A species from the southwestern U.S.A. andadjacent parts of Mexico, Cylindropuntia bigeloviiis cultivated as an ornamental (Sánchez deLorenzo Cáceres, 2000). In Güímar a single indi-vidual was observed in a dry roadside close to theCanary Cactus S.A. establishment from where itobviously had escaped.

With its easily detaching stem segments thatare densely covered in spines, this species resem-bles C. fulgida (Engelm.) F.M. Knuth var. fulgida.It is distinguished from the latter by its shrubby

habit, to 1.5 m high, larger branches c. 5cm across,larger yellowish-green flowers, 3-4cm across andits smaller fruit, c. 2cm across. The fruit is denselytuberculate, dry when ripe, usually solitary, anddoes not form chains.

Cylindropuntia bigelovii is not known as an in-vasive species (Novoa et al., 2015), nor has it beenreported before from Europe (Essl & Kobler,2008). In Güímar, C. bigelovii obviously is a re-cent escape and cannot be classified as natu-ralised yet. However, like other species from thisgenus, it certainly has the potential to establishand spread to the nearby nature reserve of Mal-país de Güímar.

Cylindropuntia fulgida (Engelm.) F.M. Knuth,Kaktus-ABC 126. 1935. (Figure 2).f. mamillata (Schott ex Engelm.) Guiggi & Ver-loove comb. nov.Basionym: Opuntia mamillata Schott ex Engelm.,Proc. Amer. Acad. Arts 3: 308. 1856.

≡ C. fulgida var. mamillata (Schott ex Engelm.)Backeb., Cactaceae Handb. Kakteen. Pereskioi-deae Opuntioideae 1: 204. 1958.≡ Opuntia fulgida Engelm., Proc. Amer. Acad.Arts 3: 306–307. 1856. var. mamillata (Schott ex Engelm.) J.M. Coult.,Contr. U.S. Natl. Herb. 3(7): 449. 1896.

New to the flora of the Canary Islands.

TENERIFE: Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, numerous individuals,24.06.2016, F. Verloove.; idem, 07.05.2016, E.Ojeda-Land et al.; Santiago del Teide: Puerto deSantiago, 28°14’13.42”N, 16°50’23.34”W, 51m.s.m., barranco de Santiago close to calle Seis,rocky slope of ravine, c. 50 individuals,15.11.2016, F. Verloove; idem, 06.01.2017, F. Ver-loove 12711 (BR).

Cylindropuntia fulgida is a species originallynative to Mexico and adjacent parts of the south-western U.S.A. It is widely cultivated as an orna-mental (e.g. Sánchez de Lorenzo Cáceres, 2000)and readily escapes, its stem segments easily de-taching and rooting. It is often considered an un-desirable, highly invasive species, for instance inAustralia (Chinnock, 2015) and South Africa(Walters et al., 2011; see also Novoa et al., 2015).It had not been recorded before from Europe (Essl& Kobler, 2008).

In Tenerife two populations were recently de-tected. In Playa de San Juan, numerous individu-als have been observed on a dry, sun-exposed

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61Bradleya 35/2017

slope close to the sea cliffs. In this locality it obvi-ously persists from an abandoned plantation.However, it evidently reproduces and appearsvery prolific. Further local expansion of the popu-lation is very likely. A smaller population with c.50 individuals was discovered in Puerto de Santi-ago on volcanic rocks bordering a ravine. In thislocality, C. fulgida also looks well-established andwill probably start spreading in the near future. It

grows with C. tunicata (Lehm.) F.M. Knuth, an-other potentially invasive species.

The plants found in Tenerife have stem seg-ments appearing spineless or nearly so from afar,exposing strongly mamillate tubercles, a trait ofvar. mamillata (Pinkava, 2003a). A taxonomicrank lower than varietas for such minor variation,however, seems more appropriate. A new combi-nation, as forma, is proposed here.

Figure 3. Cylindropuntia pallida, El Médano(TF), stony slope of ravine, June 2016. Spines arewhite throughout. Photograph by F. Verloove.

Figure 4. Cylindropuntia prolifera, Punta del Hi-dalgo (TF), September 2010.

Photograph by F. Verloove.

Figure 1. Cylindropuntia bigelovii, Güímar (TF),dry roadside, November 2016.

Photograph by F. Verloove.

Figure 2. Cylindropuntia fulgida, Puerto de San-tiago (TF), slope of ravine, January 2017.

Photograph by F. Verloove.

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62 Bradleya 35/2017

Figure 7. Echinocereus rigidissimus, Playa deSan Juan (TF), stony slope close to sea cliffs, May2016. Photograph by J.A. Reyes-Betancort.

Figure 8. Haageocereus kagenekii, El Médano(TF), stony slope of ravine, January 2017.

Photograph by E. Ojeda-Land.

Figure 5. Cylindropuntia tunicata, Puerto deSantiago (TF), slope of ravine, January 2017.

Photograph by F. Verloove.

Figure 9. Hylocereus triangularis, Valle de SanLorenzo (TF), dry river bed, January 2017.

Photograph by F. Verloove.

Figure 6. Cylindropuntia tunicata, new growthwith typical pinkish-brownish spines, Puerto deSantiago (TF), slope of ravine, January 2017.

Photograph by F. Verloove.

Page 6: New records of naturalised and invasive cacti (Cactaceae ......C/. Eduardo Zamacois, 13-3ºA, 38005 Santa Cruz de Tenerife, Canary Islands, Spain. (email: makaronesia68@yahoo.es) Introduction

Most of the plants observed belong to a com-monly cultivated morphotype with crested stems,which is often referred to as the ‘boxing-glove cac-tus’.

Cylindropuntia pallida (Rose) F.M. Knuth,Kaktus-ABC 126. 1935. (Figure 3).≡ Opuntia pallida Rose, Smithsonian Misc. Col-lect. 50: 507. 1908.= Cylindropuntia rosea auct. non (DC.) Backeb.,Cactaceae Handb. Kakteen. Pereskioideae Opun-tioideae 1: 197. 1958.

New to the flora of the Canary Islands.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’43.29”N,16°31’44.87”W, 60 m.s.m. to 28°03’36.70”N,16°31’41.68”W, 54 m.s.m., 04.04.2015, E. Ojeda-Land et al.; Granadilla de Abona: La Mareta,28º01’55.36”N, 16º34’27.51”W, 11 m.s.m.,05.01.2017, E. Ojeda-Land et al.; Buenavista delNorte: Costa de El Rincón, 28º22’13.21”N,16º52’15.61”W, 15 m.s.m., 09.02.2017, E. Ojeda-Land et al.

In El Médano a huge population is naturalisedand spreading from an abandoned, not well de-limited cultivation experiment. In this locality nu-merous cacti have been observed (Opuntia spp.,Cylindropuntia spp., Cleistocactus spp., Oreo-cereus spp., etc.), some of them showing invasivebehavior. In La Mareta a few naturalised individ-uals were observed that escaped from a gardenclose to the road TF-643, and in similar circum-stances five individuals were recorded in Costa deEl Rincón in Buenavista del Norte.

Cylindropuntia pallida has long been knownas C. rosea. The type of the latter, however, wasshown to belong to C. imbricata (Haw.) F.M.Knuth (Laguna et al., 2013). This species is doubt-fully distinct from C. tunicata and its nomencla-ture is very confusing. The neotype of C. imbricatais the painting of Haworth’s plant by Duncansonat Kew, designated by Hunt & Taylor (2006),which may as well depict O. subulata. Haworth’sdescription of C. imbricata calls for it to be imbri-cate and tessellate, which C. imbricata auct. iscertainly not. Thus an early misapplication of thename took place, probably by Engelmann, whichmeans that C. rosea should have been the correctname for what is called C. imbricata today (RoyMottram, personal communication).

Cylindropuntia prolifera (Engelm.) F.M.Knuth, Kaktus-ABC 126. 1935. (Figure 4).≡ Opuntia prolifera Engelm., Amer. J. Sci. Arts,

ser. 2 14(42): 338. 1852.

New to the flora of Tenerife. Previously reportedfrom Fuerteventura (Verloove & Guiggi, 2013)and Lanzarote where a project of eradication hasbeen conducted in La Triguera, Tenegüime(http://www.cabildodelanzarote.com/tema.asp?sec=Noticias&idTema=17&idCont=12077). Othertarget species of this eradication program wereCylindropuntia kleiniae (DC.) F.M. Knuth, Opun-tia engelmannii Salm-Dyck ex Engelm., and O.leucotricha. All were deliberately introduced ini-tially, but only C. prolifera was able to spread.

TENERIFE: San Cristóbal de La Laguna: Puntadel Hidalgo, 15.09.2010, F. Verloove; Granadillade Abona: Hoyas de Doña María, Las Arenas-ElMédano, 28°03’43.29”N, 16°31’44.87”W, 60 m.s.m.to 28°03’36.70”N, 16°31’41.68”W, 54 m.s.m.,04.04.2015, E. Ojeda-Land et al.; San Miguel deAbona: Amarilla Golf, 28º01’18.62”N,16º37’03.9”W, 20 m.s.m., 01.03.2017, E. Ojeda-Land.

This species is restricted to a relatively smallarea in Mexico (Baja California) and the U.S.A.(California). It is said to be a hybrid of two Mexi-can species, Cylindropuntia alcahes (F.A.C.Weber) F.M. Knuth and C. cholla (F.A.C. Weber)F.M. Knuth (Pinkava, 2003a). Outside its nativearea it is planted as an ornamental, although per-haps not widely so; for instance, it is not men-tioned by Sánchez de Lorenzo Cáceres (2000).

In Punta del Hidalgo in Tenerife a small pop-ulation of Cylindropuntia prolifera was recordedin 2010. In El Médano, also in Tenerife, a hugepopulation has naturalised and is spreading froman abandoned, not well controlled cultivation ex-periment. Close by, in Amarilla Golf, only two in-dividuals were observed growing on a volcanicrocky slope near to the golf court.

Although not mentioned as an invasive speciesby Novoa et al. (2015), Cylindropuntia prolifera isknown as such, at least in southern Australiawhere it is quite widespread (Chinnock, 2015). It isreminiscent of C. spinosior (Engelm.) F.M. Knuth,and the two species may have been confused. Thelatter is distinguished by its white spines and yel-low to orange fruits, as opposed to the brownishspines and green fruits of C. prolifera.

Cylindropuntia tunicata (Lehm.) F.M. Knuth,Kaktus-ABC 126. 1936. (Figures 5–6).≡ Opuntia tunicata (Lehm.) Pfeiff., Enum. Diagn.Cact. 170. 1837.≡ Cactus tunicatus Lehm., Nova Acta Phys.-Med.Acad. Caes. Leop.-Carol. Nat. Cur. 16: 319. 1832.

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New to the flora of the Canary Islands.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’43.29”N,16°31’44.87”W, 60 m.s.m. to 28°03’36.70”N,16°31’41.68”W, 54 m.s.m., 04.04.2015, E. Ojeda-Land et al.; Santiago del Teide: Puerto de Santi-ago, 28°14’13.42”N, 16°50’23.34”W, 51 m.s.m.,barranco de Santiago close to calle Seis, rocky slopeof ravine, ca. 20–30 individuals, 15.11.2016, F. Ver-loove; Granadilla de Abona: San Isidro,28°4’9.69”N, 16°33’41.05”W, 170 m.s.m., calleCuevas de Choportada, rough ground, c. 10–15 in-dividuals, 20.11.2016, F. Verloove; Santiago delTeide: Puerto de Santiago, 28°14’13.42”N,16°50’23.34”W, 51 m.s.m., barranco de Santiagoclose to calle Seis, rocky slope of ravine, 06.01.2017,F. Verloove 12710 (BR); Granadilla de Abona: SanIsidro, 28°4’13.08”N, 16°33’40.15”W, 172 m.s.m., Sof calle el Almendro, dry stony slope, scattered in-dividuals, 08.01.2017, F. Verloove; Granadilla deAbona: San Isidro, 28°4’15.56”N, 16°33’43.07”W,181 m.s.m., on the verge of barranco del Conde, fewindividuals, 08.01.2017, F. Verloove.

Cylindropuntia tunicata is a native species ina small area in Texas, U.S.A., and adjacent partsof Mexico. It is also known from Chile andEcuador in South America (Pinkava, 2003a). Else-where it is widely grown as an ornamental, alsoin Europe (see for example Sánchez de LorenzoCáceres, 2000).

In Tenerife several small populations were re-cently detected. In El Médano this species is nat-uralised in a gravelly riverbed along with manyother cacti, resulting from an abandoned cultiva-tion experiment. In Puerto de Santiago the speciesgrows on volcanic rocks of a ravine close to houses,along with C. fulgida. In San Isidro it was foundin several subpopulations, in disturbed xeric aswell as in more natural habitats. In all these lo-calities C. tunicata appears well-established and afuture local expansion is predicted, especiallysince the stem segments easily become detachedand root where they fall. This species is a knowninvader, for instance in Australia (Chinnock,2015; Novoa et al., 2015), but it was not previouslyreported from Europe according to Essl & Kobler(2008). Guillot et al. (2008), however, have sincereported it from several localities in Spain and itis also known from Italy (Guiggi, 2008). Previousclaims in South Africa proved to be referable to C.pallida (Rose) F.M. Knuth (Walters et al., 2011).

Cylindropuntia tunicata closely resembles C.pallida and the two species may have been con-fused in the past. It is distinguished, however, byits spines that are pinkish or brownish in young

growth (vs white), by its yellowish brown corollas(vs rose-red) and by less numerous spines per are-ole (4–7 vs 7–14).

Echinocereus rigidissimus (Engelm.) F. Haage,Special Preisverz. 13. 1897. (Figure 7).

New to the flora of the Canary Islands.

TENERIFE: Guía de Isora: Playa de San Juan,28°10’31.79”N, 16°48’27.37”W, 27 m.s.m.,07.05.2016, J.A. Reyes-Betancort et al.

This species is native to Arizona and New Mexico (U.S.A.) and Chihuahua and Sonora (Mex-ico) (Zimmerman & Parfitt, 2003). It is widelygrown as an ornamental, also in mainland Spain(Sánchez de Lorenzo Cáceres, 2000).

In Tenerife, in Playa de San Juan, two indi-viduals were found growing on the edge of acoastal cliff, in gravelly, sandy soils. To our knowl-edge, this species has not yet been recorded in thewild beyond its native distribution range. It is,however, cultivated as an ornamental (especiallyits subsp. rubispinus (G. Frank & A.B. Lau) N.P.Taylor (Taylor, 1989).

Haageocereus kagenekii (C.C.Gmel) Mottram,The Cactician 7: 12. 2014. (Figure 8).≡ Cactus kagenekii C.C.Gmel., Hortus magniducis badensis carlsruhanus: 48. 1811. (Fig. 3)≡ Cereus kagenekii (C.C.Gmel.) K.Schum., Ge-samtbeschreibung der Kakteen (2): 66. (15 May)1897.= Cereus pseudomelanostele Werderm. & Backeb.,Neue Kakteen 74. 1931.= Haageocereus pseudomelanostele (Werderm. &Backeb.) Backeb., Kaktus-ABC 209. 1935.= Haageocereus multangularis (Haw.) F. Ritter,Kakteen Südamer. 4: 1401. 1981.

New to the flora of the Canary Islands.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’43.29”N,16°31’44.87”W, 60 m.s.m. to 28°03’36.70”N,16°31’41.68”W, 54 m.s.m., 04.04.2015, E. Ojeda-Land et al.

As noted above this species is associated withan abandoned, not well managed cultivation ex-periment in El Médano. Several individuals weredetected in the surrounding native vegetationthat is dominated by Euphorbia balsamiferaAiton and Lycium intricatum Boiss., which indi-cates a local naturalisation event.

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Haageocereus kagenekii is not known as an in-vasive species (Novoa et al., 2015), nor has it beenreported before from Europe (Essl & Kobler,2008). It is widely grown as an ornamental, also inmainland Spain (Sánchez de Lorenzo Cáceres,2000; as H. multangularis (Haw.) F. Ritter).

Note that this species is still sometimestreated as Haageocereus pseudomelanostele. How-ever, Mottram (2014) published the combination

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

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

Figure 14. Opuntia macrocentra, Playa de SanJuan (TF), stony slope, November 2016.

Photograph by F. Verloove.

Figure 10. Opuntia basilaris, Güímar (TF), vol-canic slope, January 2017.

Photograph by F. Verloove.

Figure 11. Opuntia elatior, Buenavista del Norte(TF), rough ground, March 2017. Flowering culti-vated individual, growing close to the escapednon-flowering plants.

Photograph by J.A. Reyes-Betancort.

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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(TF), rough ground, November 2014.

Photograph by F. Verloove.

Figure 20. Oreocereus pseudofossulatus, El Mé-dano (TF), dry riverbed, June 2016.

Photograph by F. Verloove.

Figure 15. Opuntia macrocentra, detail of clado-des, Playa de San Juan (TF), stony slope, Novem-ber 2016. Photograph by F. Verloove.

Figure 16. Opuntia microdasys, El Médano (TF),dry riverbed, June 2015.

Photograph by A. González.

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H. kagenekii for the species, the earliest name ofwhich is Cactus kagenekii C.C.Gmel.

Hylocereus triangularis (L.) Britton & Rose,Contr. U.S. Natl. Herb. 12(10): 429. 1909. (Figure9).≡ Cactus triangularis L., Sp. pl. 1: 468. 1753.

New to the flora of the Canary Islands.

TENERIFE: Arona: Valle de San Lorenzo, 28°5’30.88”N, 16°39’30.50”W, 402 m.s.m., barranco deChija N of TF-28, rocky slope of ravine, a singleindividual, 28.11.2016, F. Verloove.

A native of the Caribbean, Hylocereus trian-gularis is widely cultivated in the subtropics, ei-ther as an ornamental, or for food (‘dragon fruit’).This species, as well as several others from thisgenus, are grown in the Canary Islands (Hernán-dez et al., 2013), although H. undatus is probablyby far the most widespread. The latter is also theonly species of the genus that has so far beenrecorded outside of cultivation (Acebes Ginovés etal., 2010).

The single plant found in a ravine in Valle deSan Lorenzo in Tenerife differs from H. undatusin having stems that are not horny-angled withareoles with numerous spines (6–8), and itssmaller stature with stems only c. 3–4cm across(Britton & Rose, 1919–23). Hybridisation, eitherartificial or spontaneous, may have blurredspecies boundaries in this genus resulting inplants found in the wild in Europe exhibiting in-termediate traits.

To the best of our knowledge, Hylocereus tri-angularis has only been recorded before in Europein Sicily (Guiggi, 2010). Although only a single in-dividual was found in Tenerife, a future naturali-sation cannot be excluded since its ecology andbiology typically resemble that of H. undatus, aknown invasive species (Novoa et al., 2015).

Opuntia basilaris Engelm. & J.M. Bigelow,Proc. Amer. Acad. Arts 3: 298–299. 1856. (Figure10).

New to the flora of the Canary Islands.

TENERIFE: Güímar, 28°18’28.42”N,16°22’49.64”W, 94 m.s.m., on the verge of the na-ture reserve Malpaís de Güímar, close to the Ca-nary Cactus S.A. plant, on volcanic soil, a singleclone, 11.01.2017, F. Verloove 12747; Güímar,28°18’25.19”N, 16°22’51.91”W, 114 m.s.m., on theverge of the nature reserve Malpaís de Güímar,close to the Canary Cactus S.A. plant, on volcanic

soil, two small populations of c. 8 and 9m²,11.01.2017, F. Verloove.

Opuntia basilaris is native to a relativelysmall area in the southwestern U.S.A. and the ad-jacent Sonoran Desert in Mexico. It is grown as anornamental, although probably not very widely so(see for example Hunt, 1989). As far as we couldestablish, the species had not been recorded be-fore outside of cultivation. In Tenerife, scatteredsmall populations were discovered on volcanicslopes on the verge of the Reserva Natural Espe-cial del Malpaís de Güímar, close to the CanaryCactus S.A. plant, from where it probably es-caped.

Opuntia basilaris is somewhat reminiscent ofO. microdasys with which it shares a low-growinghabit, and the cladodes having numerous spine-less areoles densely packed with short glochidia(Smith et al., 2011). It differs, however, by itscladodes being bluish-green with reddish-brownglochids, and its purplish-red corollas. The popu-lations currently found in Tenerife may include oreven pertain to hybrids with the similar O. rufidaEngelm. (Roy Mottram, personal communication).

Opuntia elatior Mill., Gard. Dict. ed. 8, 4. 1768.(Figure 11).= O. bergeriana F.A.C.Weber ex A.Berger, Gard.Chron. III, 1904(1): 34. 1904.

New to the flora of the Canary Islands.

TENERIFE: Buenavista del Norte: road to Costade El Rincón, 28º22’05.41’’N 16º52’13.88’’W, 37m.s.m., 05.03.2017, J.A. Reyes-Betancort.

Opuntia elatior is native to Mesoamerica, fromVenezuela to the Antilles (Bravo-Hollis & Arias,2011). It has been recorded as an alien from con-tinental Spain in Valencia (Guillot et al., 2008;Guillot et al., 2014) and the Balearic Islands (Ser-apio et al., 2016), twice as O. bergeriana. In Bue-navista del Norte in Tenerife, only two smallindividuals were discovered near to the road, in awaste place that has been partially used as agarbage dump for cacti. Cladodes were only 15cmlong and flowers were red.

Opuntia elatior has been reported before as aninvasive alien, for instance from Australia (Novoaet al., 2015). In Indonesia, India, and South Africait was introduced in the 19th century and quicklyspread. As a result of biological control, however,these invasions have much decreased lately. It isapparently not very widespread in cultivation; itwas not mentioned, for instance, in Hunt (1989)or Sánchez de Lorenzo Cáceres (2000). In Europe,

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O. elatior has also been reported from Italy(Guiggi, 2008).

Opuntia ficus-indica (L.) Mill. × O. tomentosaSalm-Dyck. (Figure 12).

New to the flora of the Canary Islands.

GRAN CANARIA: Arucas: Masapeses, urban-ización El Caserío, 28° 5’54.99”N, 15°31’56.76”W,374 m.s.m., GC-43 between Teror and Visvique,half-shady roadside, under Eucalyptus canopy,frequent, 19.11.2015, F. Verloove. TENERIFE:San Cristóbal de La Laguna: La Laguna, barrancode Santos E of TF-13, 28°28’37.12”N,16°18’20.74”W, 448 m.s.m., gravelly riverbed, fre-quent, 03.07.2014, F. Verloove; Tegueste: El So-corro, barranco de Las Cuevas, 28°31’7.80”N,16°21’39.99”W, 282 m.s.m., woody slope, frequent,13.01.2017, F. Verloove; Santa Úrsula: La Quinta,28°26’7.43”N, 16°29’36.69”W, 185 m.s.m., stonyslope, few plants, 15.01.2017, F. Verloove; Bue-navista del Norte: Los Carrizales, stony slopeswith the two parental species, 28°19’34.84”N,16°51’24.93”W, 25.02.2017, J.A. Reyes-Betancort.

Putative hybrids of Opuntia ficus-indica andO. tomentosa are increasingly recorded in the Ca-nary Islands. It is uncertain whether or not theyarose spontaneously or as a result of a deliberate,artificial crossing. However, in all populations inGran Canaria and Tenerife where it was recordedrecently, both putative parental species werefound growing within the flight distance of an in-sect or bird. This hybrid is apparently known fromMexico from prehistorical times (R. Mottram, per-sonal communication), wherever the two specieswere cultivated together (see also Smith &Figueiredo, 2012 on the species in South Africa).

The hybrid plants seen in the Canary Islandsare more or less intermediate between both par-ent species, although in most characters they arecloser to O. tomentosa. With the latter they sharethe tree-like habit and the velvety epidermis ofthe cladodes. However, in the hybrid the cladodesare much less greyish in appearance, and they arevery spiny, features typical of forms of the otherputative parent, O. ficus-indica.

Opuntia ×elisae D. Guillot & Van der Meer exD. Guillot has been described from continentalSpain, and probably also has Opuntia ficus-indicaand O. tomentosa as parents (Guillot & Van derMeer, 2004; Ferrer-Gallego et al., 2014). However,we refrain from using this name for the plantscurrently found in the Canary Islands since thelatter markedly differ, for instance in having vel-vety cladodes. O. ×elisae, with glabrous stem seg-

ments, obviously is closer to O. ficus-indica; more-over, the other putative parent is O. tomentosavar. hernandezii (DC.) Bravo (syn.: O. hernandeziiDC.).

Several perfectly established populations werediscovered in the past years, both in Gran Canariaand Tenerife (see above). It usually grows onrocky, often wooded, slopes, and may well havebeen overlooked elsewhere in the Canary Islands.

In the Canary Islands Opuntia ficus-indicaand O. tomentosa are both considered invasivespecies (Acebes Ginovés et al., 2010) and the samemay apply to the hybrid between them.

Opuntia tomentosa and O. ficus-indica, bothpolyploids (Pinkava, 2002), are also likely to be ofallopolyploid origin (Majure et al., 2012).

Opuntia ficus-indica (L.) Mill., Gard. Dict. (ed.8) No. 2. 1768. (Variant illustrated, see Figure 13).= O. vulgaris Ten., Syll. Fl. Neap. 239. 1831, nonMill. = O. ficus-barbarica A. Berger, Monatsschr. Kak-teenk. 22: 181. 1912.

The taxonomy and nomenclature of this mostwidely spread and highly invasive species of Op-untia in the Canary Islands are rather compli-cated. Acebes Ginovés et al. (2010) accept both O.maxima Mill. and O. vulgaris Mill., while both areoften considered synonyms of, respectively, O.ficus-indica and O. monacantha, or even of O. hu-mifusa (Guiggi, 2008). O. vulgaris is the earliestreplacement name for Cactus opuntia L. in Opun-tia in order to avoid the tautonym. That in turn isconsidered to be the same as Cactus ficus-indicaL. Using O. vulgaris for the plant correctly calledO. humifusa was a misapplication, curiouslystarted by Miller himself who illustrated O. hu-mifusa under the wrong name in his Icones(Miller, 1768; Roy Mottram, personal communi-cation).

Opuntia ficus-indica itself is a variablespecies, perhaps rather a complex of closely re-lated entities of contested taxonomic validity. This‘species’ is technically a cultivar, having been bredfor centuries. Its variability in the Canary Islandshas not been studied yet in depth, but surely morethan one taxon is present, the most notable prob-ably being the following.

Opuntia ficus-indica f. amyclaea (Ten.) Schelle,Handb. Kakteenkult. 51. 1907. Nomenclature ac-cording to Kiesling (1998).= O. megacantha Salm-Dyck, Hort. Dyck. 363.1834. (Figure 12).= O. maxima Salm-Dyck, Cact. Hort. Dyck. 68.1850, non Mill.

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Large populations of this ‘species’ were dis-covered on a sunny slope near Ayagaures in GranCanaria in 2015 and in an abandoned quarry inSantiago del Teide in Tenerife in 2016. It is alsorelatively 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, distinctlyflattened cladodes that are always spiny. Molecu-lar data seem to suggest that it probably is noth-ing else than the wild progenitor of domesticated

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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.

Figure 25. Cereus hildmannianus, El Médano(TF), dry riverbed, June 2015.

Photograph by A. González.

Figure 22. Tephrocactus articulatus, Playa deSan Juan (TF), stony slope, November 2016.

Photograph by F. Verloove.

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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.Acad. Arts 3: 292. 1856. (Figures 14–15).≡ Opuntia violacea Engelm. ex B.D. Jacks., IndexKew. 2: 358. 1895.

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Figure 26. Cleistocactus hyalacanthus, Playa deSan Juan (TF), stony slope, June 2016.

Photograph by F. Verloove. Figure 27. Cleistocactus strausii, Playa de SanJuan (TF), stony slope, June 2016.

Photograph by F. Verloove.

Figure 28. Isolatocereus dumortieri, El Médano(TF), dry riverbed, June 2015.

Photograph by A. González.

Figure 29. Neogriseocereus pruinosus, Playa deSan Juan (TF), stony slope, November 2016.

Photograph by F. Verloove.

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var. macrocentra (Engelm.) L.D. Benson, CactiArizona (ed. 3): 21. 1969.

New to the flora of the Canary Islands.

TENERIFE: Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, numerous individuals,24.06.2016, F. Verloove; idem, 07.05.2016, E.Ojeda-Land et al.

Originally native to the southern UnitedStates (Arizona, New Mexico, Texas) and Mexico(Chihuahua, Coahuila, Sonora), Opuntia macro-centra is sometimes grown as an ornamental inother warm and dry parts of the world (see for ex-ample Hunt, 1989; Sánchez de Lorenzo Cáceres,2000). Although, with its purple stem segmentswith spines of up to 12 or even 17cm long(Pinkava, 2003b), it is visually very garden-wor-thy, this species is less frequent in cultivationthan other members of Opuntia series Phaeacan-thae Britton & Rose.

Opuntia macrocentra has not been recordedoutside of cultivation, apparently, although inSouth Africa the probability of a future naturali-sation was envisaged (Walters et al., 2011).

In Playa de San Juan in Tenerife, Opuntiamacrocentra occurs in numerous subpopulationsin a relatively small area where it appears wellestablished.

Opuntia violacea Engelm. (1848) is the correctname for this plant if the paper in which it waspublished by Emory is not to be considered provi-sional only. Hunt et al. (2006) unilaterally deemedit so, but the Emory paper has never actuallybeen proposed for inclusion in the list of sup-pressed works (Roy Mottram, personal communi-cation).

Opuntia microdasys (Lehm.) Pfeiff., Enum.Diagn. Cact. 154. 1837. (Figure 16).≡ Cactus microdasys Lehm., Index sem. (Ham-burg) 16. 1827.

New to the flora of Tenerife. Previously reportedfrom Fuerteventura (Verloove & Guiggi, 2013)and La Palma (Mirador del Time) (Salas Pascual,2010).

TENERIFE: Arico: Abades, 28°8’41.56”N,16°26’27.67”W, 9 m.s.m., barranco N of the vil-lage, stony slope close to the Iglesia del Sanatoriode Abades, a single individual, 21.11.2016, F. Ver-loove; Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’44.60”N,16°31’44.49”W, 63 m.s.m., 04.04.2015, E. Ojeda-

Land et al.; Buenavista del Norte, road to Costade El Rincón, 28º22’05.40”N, 16º52’13.87”W, 25–30 m.s.m., 05.03.2017., J.A. Reyes-Betancort.

A native of Mexico, Opuntia microdasys iswidely cultivated as a popular and attractive or-namental. It is often recorded as an escape fromcultivation, in Europe at least in France, Italy,Portugal, and Spain (Essl & Kobler, 2008; Novoaet al., 2015). In the Canary Islands it had onlybeen reported before from a single locality inFuerteventura (Verloove & Guiggi, 2013) andfrom La Palma where initially there was somedoubt with regard to its identity (Salas Pascual,2010). One of us (J.A. R.-B.) also knows O. micro-dasys from Lanzarote where two small popula-tions were naturalised near the road to PlayaQuemada and in the village of Nazaret. In Abadesin Tenerife a single individual was seen in 2016where it can hardly be classified as naturalised.However, in El Médano its naturalisation hasbeen observed and it could well become an invaderin the future, as is the case in Australia and SouthAfrica (Walters et al., 2011; Novoa et al., 2015). InBuenavista del Norte one individual was observedgrowing in the inclined ditch of the road. In thislocality other species of Opuntia (one individualof O. pilifera, one of O. lindheimeri Engelm., andtwo of O. lindheimeri var. linguiformis (Griffiths)L.D. Benson) compete with Arundo donax L.,colonising the slopes and the water course of theadjacent shallow ravine. None of these, however,can be classified yet as genuinely naturalised inthat location.

Opuntia microdasys is a variable species. In ElMédano two more or less distinct forms are pres-ent. In some plants glochids are yellow or whitish,while in others they are reddish-brown. The lat-ter is sometimes distinguished as subsp. rufida(Engelm.) U. Guzmán & Mandujano (syn.: O. ru-fida Engelm.). However, glochid colour in O. mi-crodasys s.str. is a variable character and genuineO. rufida is a closely related, but quite differentspecies (Pinkava, 2003b). Schumann was the ear-liest to make the combination O. microdasys var.rufida, with O. rufida Engelm. as its basionym. Itwas Berger who was the first to apply the name O.microdasys var. rufida to a brown-glochided formof O. microdasys in cultivation at La Mortola,Italy, and explicitly excluding the type of O. ru-fida Engelm. He therefore created a later homo-nym of Schumann’s name – a different entity, andtherefore illegitimate. He can, however, bedeemed to have established the cultivar name O.microdasys ‘Rufida’. The name subsp. rufida (En-gelm.) U. Guzmán & Mandujano is validly pub-lished, but it is another superfluous synonym of

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O. rufida Engelm. No valid and legitimate botan-ical name currently exists for the brown-spinedform of O. microdasys (Roy Mottram, personalcommunication).

Opuntia monacantha (Willd.) Haw., Suppl. pl.succ. 81. 1819. (Figure 17).≡ Cactus monacanthos Willd., Enum. pl. 33.1813[1814].≡ Opuntia vulgaris Britton & Rose non Mill., Cact.1: 156. 1919

Confirmation for the flora of the Canary Islands.

TENERIFE: San Cristóbal de La Laguna: Tejina,charcas de Tejina, 28°32’12.33”N, 16°21’31.02”W,155 m.s.m., talus slope, scattered individuals,09.01.2017, F. Verloove; idem., 06.05.2008, J.A.Reyes-Betancort; Puerto de la Cruz, La Mon-tañeta, 28°23’13.97”N, 16°33’41.77”W, 306m.s.m., rough ground, c. 10-15 individuals,12.01.2017, F. Verloove; Santa Úrsula: La Quinta,close to calle Codeso, 28°25’59.06”N,16°29’42.92”W, 191 m.s.m., shrubland, small pop-ulation, 15.01.2017, F. Verloove; Tacoronte: Mesadel Mar, calle Mesa del Mar, 28°30’1.98”N,16°25’11.71”W, 252 m.s.m., small population,18.01.2017, F. Verloove.

This South American species (Argentina andBrazil) is widely cultivated as an ornamental, alsoin the Canary Islands, although it was surpris-ingly omitted by Sánchez de Lorenzo Cáceres(2000).

Acebes Ginovés et al. (2010) do not cite Opun-tia monacantha for the Canarian flora. Their ci-tation of O. vulgaris, however, most likely refersto O. monacantha (see Kunkel, 1972). O. vulgarisis a confusing name; according to Leuenberger(1993) it is conspecific with O. ficus-indica, whileHunt et al. (2006) wrongly consider it a synonymof O. humifusa (Raf.) Raf. (see above).

In Tenerife, Opuntia monacantha was observedin several localities in the northern part of the is-land, usually in small numbers and in the vicinityof houses. Most populations probably representrelics of former cultivation, or are established fromdiscarded garden waste. The species seems muchless vigorous than the superficially similar, butmuch larger-growing, O. ficus-indica. However, itis one of the most widely spread invasive cactiworldwide (Novoa et al., 2015). For instance, it is aserious invader in Australia, India, Madagascar,Mauritius, South Africa, and Sri Lanka, although,at least in some countries, it has decreased sincethe introduction of a cochineal species, Dactylop-ius ceylonicus (Walters et al., 2011).

Opuntia monacantha is easily separated by itsrather thin, pliable, and shiny cladodes. It oftenhas a somewhat drooping appearance, doubtlesslyas a result of the large fruits.

Opuntia pilifera F.A.C. Weber, Dict. Hort. 894.1898. (Figure 18).

New to the flora of the Canary Islands.

GRAN CANARIA: San Bartolomé de Tirajana:San Agustín, 27°46’12.73”N, 15°33’5.47”W, 48m.s.m., urbanización Colegio Arenas Sur, drystony slope, a large clone, 22.11.2015, F. Verloove;San Bartolomé de Tirajana: Bahía Feliz,27°47’7.45”N, 15°30’51.28”W, 10 m.s.m., roadsideGC-500, a single clone (cultivated in central reser-vation,), 27.11.2015, F. Verloove; TENERIFE:Buenavista del Norte: Costa de El Rincón,28º22’13.21”N, 16º52’15.61”W, 15 m.s.m.,09.02.2017., E. Ojeda-Land et al.; idem,05.03.2017, J.A. Reyes-Betancort.

Opuntia pilifera is a Mexican species that hasbeen reported from Puebla and Oaxaca. Outsideof its native distribution range it is widely culti-vated as an ornamental in warm-temperate partsof the world, also in southern Europe, although itis surprisingly omitted by, for instance, Hunt(1989) and Sánchez de Lorenzo Cáceres (2000). Ithas been reported in the wild from France (Tison& de Foucault, 2014), Italy (Manni & Guiggi,2015), Spain (Guillot & Lodé, 2009; Sánchez-Gul-lón et al., 2014) and Malta (see:http://www.maltawildplants.com/CACT/Opun-tia_pilifera.php). It is here reported for the firsttime from the Canary Islands. In Gran Canariasingle clones were encountered in two nearby lo-calities, while in Tenerife six clones have becomeestablished on the slope of a ravine, as escapesfrom the adjacent living fence of a cactus nursery.

Opuntia pilifera is a characteristic species. Itsglabrous cladodes are obovate to suborbicular, andlarge (up to 35cm long); the areoles are coveredwith whitish or yellowish, silky trichomes; and itsflowers are pinkish, turning purplish (Arias et al.,2012).

Opuntia robusta J.C. Wendl., Cat. Hort. Her-renh. 1835. (Figure 19).

Confirmation for the flora of the Canary Islands.

TENERIFE: Puerto de la Cruz, SW of calleSarasate,  28°24’24.64”N, 16°32’15.61”W, 127m.s.m., rough ground, a single large clone of c. 20m², 12.11.2014 and 12.01.2017, F. Verloove;

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Granadilla de Abona: Hoyas de Doña María, LasArenas-El Médano, 28°03’44.59”N, 16°31’44.63”W, 62 m.s.m., 04.04.2015, E. Ojeda-Land etal.; 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 fromGran Canaria, La Palma, and Tenerife in the Ca-nary Islands (Acebes Ginovés et al., 2010). How-ever, it is rather rarely grown as an ornamental

73Bradleya 35/2017

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

Photograph by F. Verloove.

Figure 33. Trichocereus peruvianus, Puerto de laCruz (TF), roadside slope, January 2017.

Photograph by F. Verloove.

Figure 34. Trichocereus vollianus, Playa de SanJuan (TF), stony slope, November 2016.

Photograph by F. Verloove.

Figure 30. Pilosocereus pachycladus, Playa deSan Juan (TF), stony slope, November 2016.

Photograph by F. Verloove.

Figure 31. Thelocactus hastifer, Playa de SanJuan (TF), stony slope, November 2016.

Photograph by F. Verloove.

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and most claims may refer to O. ficus-indica f.amyclaea (syn.: O. megacantha) and other formsof this species. The authors have only seen it inthree naturalised populations in Tenerife. Com-pared with O. ficus-indica, cladodes in O. robustaare larger and very glaucous, almost orbicular andits areoles are widely scattered and somewhat de-pressed. In the El Médano locality, some clones ofa putative O. robusta hybrid have also been ob-served.

Opuntia robusta is an invasive weed in severalcountries, including Australia, Botswana, andSouth Africa (Walters et al., 2011; Novoa et al.,2015). In Europe it is only rarely reported in thewild. Guiggi (2014) provides details for Italy.

Oreocereus pseudofossulatus D.R. Hunt,Bradleya 9: 89. 1991. (Figures 20–21).= Cleistocactus fossulatus Mottram, Chileans13(43): 30. 1985.[non Pilocereus fossulatus Labour., Rev. Hort. ser.4 4(2): 24–25. 1855; Oreocereus fossulatus(Labour.) Backeb., Kakt.-Freund 3(2): 13–14. 1934= Oreocereus celsianus (Salm-Dyck) Riccob.]

New to the flora of the Canary Islands.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’39.22”N,16°31’43.06”W, 54 m.s.m. to 28°03’36.80”N,16°31’41.68”W, 50 m.s.m., 04.04.2015, 04.05.2015,21.05.2015, 09.06.2015, E. Ojeda-Land et al.

As mentioned above the presence of Oreo-cereus pseudofossulatus in El Médano arose froman abandoned, not well delimited cultivation ex-periment. Birds that feed on the fleshy fruitscould enhance its expansion, although, at present,it is only locally naturalised.

Oreocereus pseudofossulatus is not known asan invasive species (Novoa et al., 2015), nor has itbeen reported before from Europe (Essl & Kobler,2008). It is often grown for ornament.

Tephrocactus articulatus (Pfeiff.) Backeb.,Cactus (Paris) 38: 249. 1953. (Figures 22–23).≡ Opuntia articulata (Pfeiff.) D.R. Hunt, Bradleya5: 93. 1987.f. papyracanthus (Phil.) Guiggi & Verloovecomb. nov.Basionym:Opuntia papyracantha Phil., Gartenflora 21: 129.1872.Synonym:Tephrocactus articulatus (Pfeiff.) Backeb. var.papyracanthus (Phil.) Backeb., Cactus (Paris) 38:249. 1953.

New to the flora of the Canary Islands.

TENERIFE: Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, numerous individuals,24.06.2016, F. Verloove; idem., 07.05.2016, E.Ojeda-Land et al.

Tephrocactus articulatus, which is native toArgentina, is sometimes grown as an ornamentalin warm and dry areas of the world. Outside of itsnative area it has been reported as an invasivealien in South Africa and Namibia (Walters et al.,2011).

In Playa de San Juan numerous individuals ofTephrocactus articulatus are found growing onbare, sun-exposed slopes close to the sea cliffs. Itsstem segments easily detach, which enables rapidvegetative dispersal. The species is well estab-lished at this location.

Tephrocactus articulatus, the most widelygrown species of this genus, is very variable andseveral varieties have been described. The plantsfound in Tenerife have long, distinctly flattenedspines, a feature of T. articulatus var. papyracan-thus (Phil.) Backeb. (see for example Kiesling,1984). Although most recent workers refrain fromusing any infraspecific names (e.g. Gilmer &Thomas, 1998; Gorelick & Mahr, 2013), this par-ticular variation may merit at least forma status,for which a new combination is established here.

The generic limits of Tephrocactus and otheropuntoid cacti have long been under debate. Mo-lecular data, however, have shown that its mem-bers form a distinct subclade (Nyffeler & Eggli,2010; Ritz et al., 2012).

Trichocereus huascha (F.A.C. Weber) Britton& Rose, Cact. 2: 142f. 208–210. 1920. (Figure 24).≡ Echinopsis huascha (F.A.C. Weber) Friedrich &G.D. Rowley, I.O.S. Bull. 3(3): 95. 1974.

New to the flora of the Canary Islands.

TENERIFE: Granadilla de Abona: Hoyas deDoña María, Las Arenas-El Médano,28°03’43.29”N, 16°31’44.87”W, 60 m.s.m. to28°03’36.70”N, 16°31’41.68”W, 54 m.s.m.,04.04.2015, 04.05.2015, 21.05.2015, 09.06.2015,E. Ojeda-Land et al.

A native of Argentina, this attractive orna-mental species was found in a dry riverbed in ElMédano in Tenerife, apparently as a relic of anabandoned cultivation experiment. The specieslooks rather prolific and now also occurs on thesurrounding rocky slopes.

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Trichocereus huascha is not known as an in-vasive species (Novoa et al., 2015), nor has it beenreported before from Europe (Essl & Kobler,2008). It is widely grown for ornament, also incontinental Spain (Sánchez de Lorenzo Cáceres,2000).

Columnar cacti with large flowers that wereaccommodated in Trichocereus (including T.huascha) have been transferred to Echinopsisbased on molecular data (Schlumpberger & Ren-ner, 2012). However, pending additional data, Rit-ter’s concept of Trichocereus is here followed(Ritter, 1980).

Part 2. Casual taxa

In Part 2, cactus species are presented that ap-pear to be non-problematic, neither at present,nor in the future. None of these have beenrecorded before from the Canary Islands. In mostcases single or a few individuals only were en-countered of these species, most of which are prob-ably unable to spread sexually in the absence ofspecies-specific pollinators. These species maypersist for some time in these localities, but futureexpansion of their ranges is rather unlikely. De-spite being widely grown as ornamentals (see forexample Hunt, 1989; Sánchez de Lorenzo Cáceres,2000), none is known as an invasive species(Novoa et al., 2015). Hence, the entries for thesespecies are brief, but they are all illustrated.

Cereus hildmannianus K. Schum., Fl. Bras.4(2): 202. 1890. (Figure 25).Cereus peruvianus auct. non (L.) Mill.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’44.60”N,16°31’44.4987”W, 63 m.s.m., 04.04.2015, E. Ojeda-Land et al.

A few plants of the ‘Monstruosus’ form of thisSouth American species were seen in El Médanoin Tenerife where it persists from an abandonedcultivation experiment. This and several otherspecies of Cereus are reputed invasive species(Novoa et al., 2015). In Europe, Cereus hildman-nianus has been reported before from Italy(Guiggi, 2010).

Application of this name for the well-knownplant of cultivation is contentious. Hunt et al.(2006) have championed the use of this name, butit is doubtfully different from C. jamacaru DC., oreven from C. hexagonus (L.) Mill. This present-day cultivar is so widely cultivated from ancienttimes that its true origin is lost in time (Roy Mot-tram, personal communication).

Cleistocactus hyalacanthus (K.Schum.) Rol.-Goss., Bull. Mens. Soc. Centr. Agric. Hort. Acclim.Nice Alpes-Marit. 44: 33. 1904. (Figure 26).≡ Cereus hyalacanthus K. Schum., Gesamtbeschr.Kakt. 101. 1897.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28º03’38,50’’ N,16º31’42,67’’ W, 53 m.s.m., 04.04.2015, E. Ojeda-Land et al.; Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, scattered individuals,24.06.2016, F. Verloove.

Scattered individuals of this South Americanspecies (Argentina, Bolivia) were found growingon a stony slope close to the sea cliffs in Playa deSan Juan and in a dry riverbed in El Médano inTenerife. In both localities the plants persist froman abandoned plantation. They were found to-gether with the congeneric and superficially sim-ilar-looking Cleistocactus strausii (see below). Thelatter, however, has longer corollas (80–90 vs 30–40mm), more numerous costae (25–30 vs c. 20),and its spines are white, not brownish.

Cleistocactus strausii (Heese) Backeb., Kak-teen-freund 3: 121. 1934. (Figure 27).≡ Pilocereus strausii Heese, Gartenflora 56: 410.1907.

TENERIFE: Granadilla de Abona: Hoyas deDoña María, Las Arenas-El Médano,28°03’43.29”N, 16°31’44.87”W, 60 m.s.m. to28°03’36.70”N, 16°31’41.68”W, 54 m.s.m.,04.04.2015, E. Ojeda-Land et al.; Guía de Isora:Playa de San Juan, 28°10’41.38”N,16°48’33.98”W, 48 m.s.m., stony slope next to thesea cliffs, scattered individuals, 24.06.2016, F.Verloove.

Originally native in Bolivia, Cleistocactusstrausii is widely cultivated as an ornamental.Scattered individuals are found on a stony slope inPlaya de San Juan and in a dry riverbed in El Mé-dano in Tenerife. The plants grow together withC. hyalacanthus. Both persist from an abandonedplantation.

Isolatocereus dumortieri (Scheidw.) Backeb.,Cactaceae (Berlin) 1941(2): 47. 1942. (Figure 28)≡ Stenocereus dumortieri (Scheidw.) Buxb., Bot.Stud. 12: 92. 1961.≡ Lemaireocereus dumortieri (Scheidw.) Britton &Rose, Contr. U.S. Natl. Herb. 12(10): 425. 1909.≡ Rathbunia dumortieri (Scheidw.) P.V. Heath,Calyx 2: 104. 1992.

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≡ Cereus dumortieri Scheidw., Hort. Belge 4: 220.1837.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’44.60”N,16°31’44.4987”W, 63 m.s.m., 04.04.2015, E. Ojeda-Land et al.

A few plants of this Mexican species grow in adry riverbed in El Médano in Tenerife where theypersist after an abandoned cultivation experi-ment.

Isolatocereus dumortieri was long included inStenocereus, but flower morphology and molecu-lar data seem to support its segregation (Cota &Wallace, 1997).

Neogriseocereus pruinosus (Otto ex Pfeiff.)Guiggi, Cactology 3, Suppl. 4: 1. 2013. (Figure 29).≡ Griseocereus pruinosus (Otto ex Pfeiff.) Guiggi,Cactology 3: 8. 2012.≡ Stenocereus pruinosus (Otto ex Pfeiff.) Buxb.,Bot. Stud. 12: 101. 1961.≡ Echinocactus pruinosus Otto ex Pfeiff., Enum.Diagn. Cact. 54. 1837.

TENERIFE: Granadilla de Abona: Hoyas de DoñaMaría, Las Arenas-El Médano, 28°03’44.60”N,16°31’44.4987”W, 63 m.s.m., 04.04.2015, E. Ojeda-Land et al.; Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, scattered individuals,24.06.2016, F. Verloove.

Scattered individuals of this impressivespecies from Central America (Guatemala, Hon-duras, Mexico) grow on a sun-exposed stony slopein Playa de San Juan and in a dry riverbed in ElMédano in Tenerife, persisting from an aban-doned plantation and cultivation experiment.

This species was long known as Stenocereuspruinosus. However, the generic limits of Steno-cereus are debated and this species and some rel-atives are best segregated (Heath, 1996; Guiggi,2012, 2013).

Pilosocereus pachycladus F. Ritter, KakteenSüdamer. 1: 70. 1979. (Figure 30).

TENERIFE: Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, scattered individuals,24.06.2016, F. Verloove; idem, 07.05.2016, E.Ojeda-Land et al.

Scattered individuals of this attractive, widelycultivated Brazilian species grow on a sun-ex-

posed stony slope in Playa de San Juan in Tener-ife, where they persist from an abandoned plan-tation.

Thelocactus hastifer (Werderm. & Boed.) F.M.Knuth, Kaktus-ABC 360. 1935. (Figure 31).≡ Echinocactus hastifer Werderm. & Boed., No-tizbl. Bot. Gart. Berlin–Dahlem 11: 274. 1931.

TENERIFE: Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, a single clone,24.06.2016, F. Verloove.

Scattered individuals of this Mexican speciesgrow on a sun-exposed stony slope in Playa de SanJuan in Tenerife, persisting from an abandonedplantation.

Trichocereus bridgesii (Salm-Dyck) Britton &Rose, Cact. 2: 134. 1920. (Figure 32).≡ Cereus bridgesii Salm-Dyck, Cact. Hort. Dyck.208. 1849[1850].

TENERIFE: Arona: Chayofa, 28° 4’33.03”N,16°41’28.13”W, 293 m.s.m., dry slope adjacent tocalle Barranco del Verodal, three individuals,22.11.2016, F. Verloove; Arona: Chayofa, 28°4’28.99”N, 16°41’38.39”W, 280 m.s.m., on theverge of vacant lot in residential area, two indi-viduals, 22.11.2016, F. Verloove.

A few individuals of this Bolivian species growin two different localities in Chayofa in Tenerife;in both instances material probably became es-tablished from discarded garden waste.

Trichocereus peruvianus Britton & Rose, Cact.2: 136, f. 197. 1920. (Figure 33).≡ Echinopsis peruviana Friedrich & G.D. Rowley,I.O.S. Bull. 3(3): 97. 1974.

TENERIFE: Puerto de la Cruz, calle Chahorra,28°24’21.56”N, 16°33’56.77”W, 67 m.s.m., roadsideslope, five individuals, 16.01.2017, F. Verloove.

Five individuals of this Peruvian species growon a slope in Puerto de la Cruz in Tenerife, prob-ably from discarded garden waste.

Trichocereus vollianus Backeb., Kaktus-ABC205, 412. 1935. (Figure 34).

TENERIFE: Guía de Isora: Playa de San Juan,28°10’41.38”N, 16°48’33.98”W, 48 m.s.m., stonyslope next to the sea cliffs, scattered individuals,24.06.2016, F. Verloove.

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Scattered individuals of this Bolivian speciesgrow on a sun-exposed stony slope in Playa deSan Juan in Tenerife, persisting from an aban-doned plantation.

AcknowledgementsDaniela Zappi (Brazil), Daniel Guillot Ortiz

(Spain) and Emilio Laguna (Spain) are thankedfor providing valuable comments on Pilosocereuspachycladus and Opuntia ×elisae, respectively.Fabián Hernández and Beatriz Fariña accompa-nied one of the authors (R.B.) in Playa de SanJuan (Tenerife), when some cacti were discov-ered. Juana María González Mancebo andLeopoldo Moro Abad, alerted us to the presenceof some cacti in El Médano and Amarilla Golf, re-spectively.

Roy Mottram (United Kingdom) is thanked foruseful discussions on, and assistance with, theidentification of some of the cactus species re-ported here. He also contributed valuable com-ments on a first version of the manuscript.

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