Review - Invasive Species cactus speci… · de cacta´ceas de Norte y Centro Ame´rica, Herbario Virtual da Flora e dos Fungos, Intermountain Regional Herbar-ium Network, Global

Review

Introduced and invasive cactus species: a global reviewAna Novoa1*, Johannes J. Le Roux1, Mark P. Robertson2, John R.U. Wilson1,3 and David M. Richardson1

1 Centre for Invasion Biology, Department of Botany and Zoology, Stellenbosch University, Matieland 7602, South Africa2 Centre for Invasion Biology, Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa3 Invasive Species Programme, South African National Biodiversity Institute, Kirstenbosch Research Centre, Claremont 7735, South Africa

Received: 27 September 2014; Accepted: 13 November 2014; Published: 3 December 2014

Associate Editor: J. Hall Cushman

Citation: Novoa A, Le Roux JJ, Robertson MP, Wilson JRU, Richardson DM. 2015. Introduced and invasive cactus species: a global review.AoB PLANTS 7: plu078; doi:10.1093/aobpla/plu078

Abstract. Understanding which species are introduced and become invasive, and why, are central questions ininvasion science. Comparative studies on model taxa have provided important insights, but much more needs to bedone to unravel the context dependencies of these findings. The cactus family (Cactaceae), one of the most popularhorticultural plant groups, is an interesting case study. Hundreds of cactus species have been introduced outside theirnative ranges; a few of them are among the most damaging invasive plant species in the world. We reviewed the dri-vers of introductions and invasions in the family and seek insights that can be used to minimize future risks. We com-piled a list of species in the family and determined which have been recorded as invasive. We also mapped currentglobal distributions and modelled the potential global distributions based on distribution data of known invasivetaxa. Finally, we identified whether invasiveness is phylogenetically clustered for cacti and whether particular traitsare correlated with invasiveness. Only 57 of the 1922 cactus species recognized in this treatment have been recordedas invasive. There are three invasion hotspots: South Africa (35 invasive species recorded), Australia (26 species) andSpain (24 species). However, there are large areas of the world with climates suitable for cacti that are at risk of futureinvasion—in particular, parts of China, eastern Asia and central Africa. The invasive taxa represent an interesting sub-set of the total species pool. There is a significant phylogenetic signal: invasive species occur in 2 of the 3 major phylo-genetic clades and in 13 of the 130 genera. This phylogenetic signal is not driven by human preference, i.e.horticultural trade, but all invasive species are from 5 of the 12 cactus growth forms. Finally, invasive species tendto have significantly larger native ranges than non-invasive species, and none of the invasive species are of conser-vation concern in their native range. These results suggest fairly robust correlates of invasiveness that can be used forproactive management and risk assessments.

Keywords: Biological invasions; cactus invasions; climate suitability; introduction pathways; invasion debt; invasivespecies; phylogenetic signal.

IntroductionThe increased movement of humans around the worldhas facilitated the intentional and accidental transporta-tion of species far from their native ranges, often in amanner that can facilitate invasions (Wilson et al.

2009). Many of these introduced organisms have notablebenefits to humans, but some have undesirable impactsthat can result in substantial monetary costs and/oralterations to entire ecosystems and social systems(McNeely 2006; Kumschick et al. 2012). Governmentdepartments, non-governmental organizations, extension

* Corresponding author’s e-mail address: [email protected]

Published by Oxford University Press on behalf of the Annals of Botany Company.This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

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services, environmental managers, conservationists andscientists are all facing escalating pressure to addressand resolve a diversity of problems posed by invasivealien species (Hulme 2006).

Much work has recently focussed on reviewing the inva-sive performance of particular genera or closely relatedgroups of organisms in different situations around theworld (e.g. Richardson et al. 2011; Moodley et al. 2013;Potgieter et al. 2013; Shackleton et al. 2014). Such studiesaim to update knowledge on the global occurrence andpotential range of these taxa and to understand the com-plex drivers of human-mediated introductions and inva-sions. The findings of such studies are important fordeveloping protocols for preventing risky species introduc-tions and for managing species that may become or havealready become invasive (Simberloff et al. 2009). However,more comparative studies are needed to improve ourunderstanding of the full suite of interacting factors thatinfluence invasions and to unravel the context dependen-cies of insights that emerge from particular studies(Kueffer et al. 2013). It is important to consider whethersuch comparisons yield broad generalities or whethergeneralizations apply only to a subset of taxa.

The cactus family (Cactaceae; ‘cacti’) is an interestingcase study. Cacti are a conspicuous component of thearid regions of the New World and represent one of theworld’s most spectacular desert radiations (Edwardset al. 2005). The family is distributed from southernPatagonia in Argentina and Chile to Alberta and BritishColumbia in Canada (Edwards et al. 2005), with the onlyexception being Rhipsalis baccifera (mistletoe cactus),which is thought to have originated in tropical Americas,but was apparently dispersed across the Atlantic Oceanby birds, reaching southern Africa, Madagascar and SriLanka (Rebman and Pinkava 2001).

Cacti are among the first plants that were brought backfrom the Americas by European explorers in the 15th cen-tury (Howard and Touw 1981) and soon became commonin European collections and gardens (Anderson 2001).The trade in horticultural cacti has developed over theyears into a substantial industry and is responsible forthe intercontinental spread of many species (Walterset al. 2011). One of the earliest reasons for introduction,however, was for use as drought-tolerant crops and forhedging, with Opuntia ficus-indica (L.) Mill. being by farthe most utilized (Walters et al. 2011). In an attempt tominimize the risks of global climate change, land degrad-ation and diminishing food security, the Food and Agricul-tural Organization has revived the interest in cactuscultivation for agricultural purposes in developing coun-tries (Nefzaoui 2007). As a result of these human-mediated introductions, cactus species can be found allover the world, and several members of the family are

among the most important alien species worldwide(Weber 2003).

Studies of cactus invasions have shed light on crucialaspects of plant invasion ecology, e.g. the interactionof invasive plants with seed dispersers (Foxcroft andRejmanek 2007; Padron et al. 2011), the role of propagulepressure in driving invasions (Foxcroft et al. 2004) and therole of herbivores in regulating some plant populations,with particularly striking examples from classical biologic-al control (Zimmermann et al. 2004; Paterson et al. 2011).These studies have tended to focus on the genera Opuntiaand Cylindropuntia, which contain most of the widelyintroduced, cultivated and invasive species in the family.However, hundreds of new cactus species are now beingintroduced all over the world, and many of them arebecoming naturalized or invasive. For example, Cereushexagonus was included in a national list of regulatedinvasive species for the first time in South Africa in 2014.

The current global distribution of the cactus family isbeing radically changed by humans, and no attempthas been made to assess the status of each species interms of invasion or risk thereof. Consequently, a broadglobal assessment of the determinants of invasivenessof the family Cactaceae is an important requirement forthe formulation of control strategies. Moreover, reviewingthe invasive performance of this family around the worldmay uncover new patterns, processes and invasion risksnot seen in better-studied model groups.

This paper aims specifically to (i) compile a list of spe-cies in the family Cactaceae, (ii) determine their currentnative and invasive ranges, and (iii) determine the poten-tial future ranges of invasive taxa. Using these lists weaim to answer the questions: (iv) how have cactus speciesbeen used inside and outside their native range?; and (vi)are any traits correlated with invasiveness in the family?

Methods

Defining a cactus

Most taxa in the family Cactaceae are succulents withlarge, leafless, long-living, fleshy stems of different shapesand sizes that often contain clusters of spines which arisefrom areoles (Benson 1979, 1982; Eggli 1993). Areoles—highly specialized axillary or lateral buds or short shootsor branches—are unique to the family (Mauseth 1983;Gibson and Nobel 1986). However, cacti come in a widerange of growth forms (Fig. 1). Succulent plants in otherfamilies are often mistakenly called ‘cacti’ on nurserylabels and in popular publications. While it is usually easyto distinguish cacti from other succulents, some taxa lookvery cactus-like due to convergent evolution, e.g. manyspecies in the genus Euphorbia of the family Euphorbia-ceae (Anderson 2001).

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Species list and invasive status

While the alpha taxonomy of the Cactaceae is wellknown, and the clade is clearly a recent monophyletic ra-diation, a stabilized nomenclature has not yet beenachieved (Hunt and Taylor 1986, 1990; Nobel 2002). Thisnomenclature instability can be attributed to inconsisten-cies in previous taxonomies and to the generally poor re-presentation of cacti in herbarium collections [theirsucculence and spines make them difficult to collectand curate (Walters et al. 2011)]. The InternationalOrganization for Succulent Studies recently attempted

to revise the taxonomy of the Cactaceae (Hunt et al.2006), but concluded that this task is far from complete.

In this paper, we base our list on the classification sys-tem developed by the International Cactaceae Systema-tics Group and used by David Hunt in compiling botheditions of the Convention on International Trade inEndangered Species of Wild Fauna and Flora’s CITES Cac-taceae Checklist (1992, 1999). This classification has beenadopted by various sources (Walters 1989; Kubitzki et al.1993; Anderson 2001). We updated the list to include109 additional species, for which, since 2001, either an

Figure 1. The 12 growth-form categories of the family Cactaceae considered in this paper.





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International Union for Conservation of Nature andNatural Resources red list assessment has been made(http://www.iucnredlist.org/), or a new description hasbeen published in the scientific literature. None of theseadditional 109 species are recorded as invasive.

Information on the current distribution and invasivestatus was gathered from many sources [see SupportingInformation]. Each source uses different criteria for cat-egorizing alien species. To record the invasive status ofthe family Cactaceae, our list includes only cactus specieswhere there is clear documented evidence of an invasionas per the definition in Richardson et al. (2000), i.e. plantsspreading over considerable distances from originalplanting or introduction sites.

Potential distribution

Invasiveness elsewhere combined with broad-scaleclimatic matching are the most widely used predictorsof future invasiveness of introduced species (Rejmaneket al. 2005; Richardson et al. 2011; Petitpierre et al.2012). Determining which global areas are climaticallysuitable for invasive species establishment may aid futuremanagement efforts and/or preventative measures. Wetherefore used bioclimatic models for invasive cacti topredict potential suitable ranges for individual specieson a global scale. To do this, we compiled a dataset ofoccurrence records for invasive cactus species from sev-eral different sources (e.g. Base de datos de ejemplaresde cactaceas de Norte y Centro America, Herbario Virtualda Flora e dos Fungos, Intermountain Regional Herbar-ium Network, Global Biodiversity Information Facility, In-stituto de Biologia de la UNAM, Southwest EnvironmentalInformation Network, Oregon Flora Project, San DiegoNatural History museum and CalFlora). For each species,the records were plotted on a map and climatic spaceas defined by values of annual mean temperature andannual precipitation extracted from 10-min resolutionWorldClim bioclimatic variables. A thorough data clean-ing procedure was followed using the biogeo packagein R (M. P. Robertson et al., in preparation): tests were per-formed on obvious outliers to determine whether the x-and y-coordinates had accidentally been transposed orwhether incorrect signs were used; records that wereplotted in the ocean but directly adjacent to a coastalgrid cell were assigned to that closest terrestrial gridcell; and low-precision records were removed, e.g. whenonly degrees were available for the coordinate. Outliersin the environmental space were queried to identifywhere they lay in geographical space to identify anyerrors missed during the initial data cleaning, andremoved or rectified if a particular cause of the errorcould be determined.

Species distribution models were produced for eachspecies using a simple envelope approach (implementedin R) which is equivalent to BIOCLIM’s marginal envelope(Pearson and Dawson 2003). The following predictor vari-ables were used: maximum temperature of warmestmonth, minimum temperature of coldest month, precipi-tation of wettest quarter, precipitation of driest quarter,precipitation of warmest quarter and precipitation ofcoldest quarter. These variables were obtained fromWorldClim at a 10-min spatial resolution (Hijmans et al.2005) and were selected based on their success at pre-dicting potential global distributions for other model inva-sive taxa (Richardson et al. 2011).

Two sets of models were produced using different ap-proaches. For the first approach, models were calibratedusing native range records only. These models were thenevaluated using invasive range records, where these wereavailable. Sensitivity values (Fielding and Bell 1997) werecalculated for each model based on the number of inva-sive range records that were predicted as present or ab-sent by the model. Sensitivity values range betweenzero and one, where values close to one indicate lowomission error (Fielding and Bell 1997). For the second ap-proach, models were calibrated using all available recordsfor the species (i.e. native and invasive range records)without evaluation of sensitivity. For both approacheswe produced models for species that had five or more na-tive range records. Duplicate records per 10-min cell wereremoved.

Maps of potential species richness were produced byadding the maps of potential distribution for each ofthe two approaches.

Reasons for introduction and dissemination

Information on human uses of cacti both in their nativeand introduced ranges were extracted from many sources[see Supporting Information]. Five broad human-usecategories were defined: (i) ornamental (horticulture),(ii) food or fodder (i.e. for humans or livestock), (iii) medi-cinal, (iv) hedging and (v) other (e.g. furniture or religious).Not all the species with a defined use in the native rangeare introduced to other areas of the world for the samereasons. To assess how the number of introduced speciesdiffers between uses, we compared the proportion of in-troduced and non-introduced species in each use cat-egory with that of species in other use categories (usinga Fisher Test in R).

Correlates of invasiveness

A useful first step to improve our understanding of inva-siveness is to identify the traits correlated with invasive-ness (Pysek and Richardson 2007). Here, we looked at





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phylogeny, taxonomy (at the genus level), growth formand metrics of native range size.

For phylogenetic reconstruction we collated geneticdata for the maturase K (matK) gene region for repre-sentative taxa of all Cactaceae genera with availabledata in the GenBank online repository (http://ncbi.nlm.nih.gov). DNA sequence data were aligned in BioEdit ver-sion 7.0.5.3 (Hall 1999) and manually edited. Because ofdifferences in sequence lengths for different taxa wetrimmed flanking regions to avoid excessive missingdata. Our final dataset comprised 103 genera withinCactaceae. Phylogenetic relationships were estimatedusing Bayesian search criteria with parameter estimatesobtained from the program jModelTest v2.1.3 (GTR + I + G;Darriba et al. 2012) in MrBayes 3.1.2 (Ronquist andHuelsenbeck 2003). For both datasets, MrBayes wasrun for 2 000 000 generations and trees sampled every1000 generations. Nodal support for the retrieved treetopology was determined as posterior probabilities inMrBayes. To determine whether invasiveness within Cac-taceae has a phylogenetic signal, we compared traitchange (proportion of invasive taxa within a genus)with a null hypothesis of Brownian motion using Blom-berg’s K statistic (Blomberg et al. 2003). Similarly, weused Pagel’s lambda (l) statistic to determine the extentto which branch length transformation explains thedistribution of trait states (proportion of invasive taxa/genus) at the tips of a phylogeny (Pagel 1999). Bothtests are implemented in the function phylosig.R fromthe phytools package (Revell 2012).

To assess how invasiveness differs at the genus level,we compared the number of invasive and non-invasivespecies in each genus with that in the rest of the familyusing a Fisher test in R.

Based on information extracted from all sources, wealso obtained information on the growth form of eachspecies. There are different classifications in the literature(e.g. Barthlott and Hunt 1993; Anderson 2001; Lopez andValdivia 2007; Ortega-Baes et al. 2010; Hernandez-Hernandez et al. 2011). Here, following discussions inAnderson (2001) and Hernandez-Hernandez et al. (2011),we recognize 12 types: angled, cylindrical, cushion-like,leaf-like, flattened-padded, geophytic, globose, ovoid,sprawling tree-like, tuberculate and prostrate growthforms (Fig. 1).

Available data on native range size are inadequate foran analysis of the relationship between native range sizeand invasive status for the whole family. We were, how-ever, able to analyse the relationship between nativerange size and invasiveness for the genus Opuntia, be-cause species in the genus have been widely introducedand disseminated around the world, there are many inva-sive and non-invasive taxa, and there are comparatively

good data on native range size for this genus (Dean andMilton 2000; Erre et al. 2009; Padron et al. 2011; Lloyd andReeves 2014). We compared the latitudinal ranges of in-vasive and non-invasive Opuntia species using a Student’st-test in R. We also looked at Cactaceae listed on the IUCNRed List of Threatened Species (Rodrigues et al. 2006). ForCactaceae, most of the species considered at risk were sodue to a small native range size (http://www.iucnredlist.org/), and therefore, in this case, the list provides arough proxy for native range size (as well as givingsome indication of population trends).

Results

Species list and invasive status

The list of cacti assembled for this paper comprises 1922species from 130 genera [see Supporting Information].Genera differed widely in species richness, with severalspeciose genera [Opuntia (193 species), Echinopsis (133species) and Mammillaria (171 species)] and 35 monotypicgenera. Of the 1922 species we could definitively classifyonly 57 species as invasive.

The currently available distribution data are only ad-equate for a country-level analysis (see Fig. 2). These ana-lyses show Mexico as the main ‘hot spot’ of native cactusdiversity (Fig. 2A). Three countries had notably more inva-sive taxa than the rest—Australia (39), South Africa (35species) and Spain (24)—while other countries had atmost 13 (Fig. 2B, Table 1). The most widespread invasivespecies is O. ficus-indica (22 different countries), withother species invading 15 or fewer countries (Table 1). Un-surprisingly, the origin of most of these invasive species isalso Mexico (Fig. 2C).

Potential distribution

We examined potential invasive distributions for only 39of the invasive species [see Supporting Information]as none of the remaining 18 species had enough (i.e.five or more) records in their native ranges of sufficientaccuracy. The median number of records per speciesmodeled was 128 for the native range and 124 for theinvasive range. A large variation in sensitivity valueswas obtained. We found no significant differences be-tween the projected species richness maps for the twomodelling approaches, i.e. using native range occurrencerecords only or native and invasive occurrence records(Fig. 3). The main known areas of invasion (Australia,South Africa and Spain) were indicated as suitable, butthere were also substantial regions that are suitable incentral Africa, China and south-eastern Asia.


A quarter of species recorded as being used for ornamen-tation in their native ranges have been introduced





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Figure 2. Cactus species richness across the native (A) and invasive range (B) as well as the native distribution of invasive cacti (C). Shadingindicates the number of taxa per country. Lighter colors correspond to less taxa.





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Table 1. Distribution of invasive Cactaceae species outside their native range. Data were compiled from a range of sources [see SupportingInformation].





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elsewhere (Table 2). In contrast, only a seventh of the cac-tus species used in their native ranges for food or fodderhave been introduced elsewhere, and we found no officialrecords of species having been introduced for medicinalor other purposes (though Lophophora williamsii andEchinopsis pachanoi have been introduced worldwidefor their psycho-active uses). As an illustration of theworldwide popularity of cacti as horticultural species,we give some examples of international cactus andsucculent journals and societies [see SupportingInformation].


Our phylogeny, representing 103 taxa (genera), retrievedthree main clades that differed substantially in the pro-portion of invasive taxa (Fig. 4). Clade 1 included mainlygenera of the tribes of the subfamily Cactoideae (withthe exception of the tribe Cacteae), and some invasivetaxa. Clade 2 comprised genera within the tribe Cacteae(with the exception of Maihueniopsis from the tribeTephrocacteae) and does not include any invasive taxa.Clade 3 (which includes the subfamily Opuntioideae)contains most of the invasive species. Using proportionsof invasive taxa/genus, Blomberg’s K indicated no signifi-cant phylogenetic signal of invasiveness (K ¼ 0.260; P ¼0.187), whereas Pagel’s l indicated a significant

phylogenetic signal for invasiveness (l ¼ 0.991; P ,

0.01). This phylogenetic signal was not related to humanusage, i.e. ornamental trade.

At the genus level, the 57 invasive species belong to just13 of the 130 genera (Fig. 5A). Opuntia, Cylindropuntia, Har-risia, Hylocereus and Austrocylindropuntia have a signifi-cantly higher proportion of invasive species than othergenera, while only Mammillaria has a significantly lower in-cidence of invasiveness (it contains no invasive species).

In terms of the 12 main growth forms we considered(Fig. 1), all of the invasive species were cylindrical,flattened-padded, sprawling, leaf-like or angled. Theflattened-padded and angled growth forms stand out ashaving a significantly higher percentage of invasive spe-cies than the other growth forms, while the globosegrowth form is significantly underrepresented in terms ofinvasiveness (there are no globose invasive taxa) (Fig. 5B).

The latitudinal ranges of invasive Opuntia species (218,n ¼ 15) are significantly greater than those of non-invasive species (8.78, n ¼ 42) (t-test, P , 0.009). As of2013, the IUCN had assessed 1409 cactus species. All in-vasive taxa are in the Least Concern, Near Threatened orData Deficient categories and none of the known invasivespecies are among the 303 taxa listed in the categoriesCritically Endangered (CR), Endangered (EN) or Vulnerable(VU) (i.e. species that likely have smaller ranges, Fig. 5C).

Figure 3. Potential species richness based on all available records (native and invasive records) of 39 invasive cactus species.

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Table 2. The number of species inside and outside their native range across human uses. Note that one species can be included in more than oneuse category. Significance levels were determined by comparing the number of introduced vs. number of non-introduced species for anycategory to all other taxa using Fisher’s exact test. Confidence intervals were determined for the percentage of introduced ornon-introduced based on an assumption of binomial errors. Other uses include minor uses such as water source.

Ornamental Food Medicinal Hedging Other

Native range 837 261 345 15 10

Non native range 250 45 0 5 0

Percentage (95 % CIs) 23.0 % (20.5–25.6) 14.7 % (10.9–19.2) 0 % (0–10.3) 25.0 % (8.7–49.1) 0 % (0–30.8)

Significant P , 0.01 P , 0.01 P , 0.01 P , 0.58 P , 0.23





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This provides preliminary evidence for a relationship be-tween native range size and invasiveness.

DiscussionThe results of this study allowed us to draw generaliza-tions that are useful for managing future introductionsand invasions. Despite the extensive dissemination ofcactus species around the world, only a small proportionof the family is currently known to be invasive. However,these invasive taxa have very large potential ranges glo-bally, including in areas where no cactus invasions haveyet been recorded. Cacti are introduced to new areasmainly for ornamentation, but the selection of ornamen-tal species is not based on attributes that favour invasive-ness. Invasive taxa are overrepresented in several genera,phylogenetic clades and growth forms. Species that areof conservation concern in their native ranges have notbecome invasive.

Proportion of invasive taxa in the family Cactaceae

The main centres of cactus diversity are north-easternMexico, the eastern Andes of Bolivia and Argentinaand south-eastern Brazil (Mutke and Barthlott 2005).However, species are distributed throughout a large var-iety of habitats, including hot deserts, sandy coastalstretches, scrublands, dry deciduous forests, high alpinesteppes and even tropical rain forests (Barthlott andHunt 1993). Therefore, there are cactus species that areclimatically suited to almost all habitats on Earth. How-ever, only 3 % of the species in the family are currentlyclearly invasive. It is difficult to say whether this reflectsthe real extent of invasions or whether the pattern isaffected by different levels of reporting and the availabil-ity of accurate data, moreover, not all cactus species havebeen afforded the same opportunities to become inva-sive. Nonetheless, this pattern has also been observedin other model groups. For example, only between 0.5and 0.7 % of the global pool of tree and shrub speciesare currently clearly invasive outside their natural range(Richardson and Rejmanek 2011).

Potential for further invasions

Currently, most cactus invasions are recorded in Australia,South Africa and Spain. Unsurprisingly, the results of thebroad-scale climatic matching identified these threecountries as being bioclimatically equivalent to areaswithin the range of a large number of cactus species.A strong climatic match between native and recipientranges is recognized as a fundamental requirement forthe success of introduced plants (Richardson et al. 2011;Richardson and Pysek 2012). However, many other areasof the world where these species are currently absent arealso highlighted as potentially suitable for invasion. Thispattern probably reflects differences in introduction ef-fort, and suggests a substantial invasion debt (sensuEssl et al. 2011) in agreement with the findings forother model groups (e.g. the genera Casuarina and Proso-pis; Potgieter et al. 2013; Shackleton et al. 2014). Clearly,the natural experiment of plantings of cacti outside theirnatural ranges is far from complete.

Because efforts directed at prevention of new introduc-tions are the most cost-efficient component of invasivespecies control strategies (Leung et al. 2002), our resultsemphasize the importance of controlling the introductionof cacti recorded as invasive in Australia, South Africa andSpain to other areas suitable for invasion.


Among the many uses of cacti, the main reason forintroductions of species to regions outside their nativeranges is the horticulture trade. Cultivation of ornamentalcactus species is very popular in temperate regions. Thereare more than 20 cactus and succulent journals and hun-dreds of societies around the world, as well as hundredsof cactus and succulent Facebook pages and groups.Moreover, global introductions of new species are likelyto occur: just 23 % of the species considered to have or-namental value in the native range have been introducedto other regions.

The use of alien plants for ornamentation is an import-ant driver of introductions and dissemination in manyplant groups, and several attributes associated with

Figure 4. Bayesian phylogeny based on matK DNA sequence data illustrating phylogenetic relationships among genera within Cactaceae. Thealigned matK matrix contained 1354 characters and required 65 gaps (indels), ranging from 1 to 74 characters in size. Overall, the phylogenyyielded well-resolved relationships among all genera included. High nodal support (posterior probabilities .0.9) is indicated at nodes by yellowboxes. Genera with invasive taxa are indicated as red branches where red circles are proportional to the percentage of invasive taxa within eachgenus. Scale bar ¼ number of substitutions/site. The three main clades are indicated under the numbers 1, 2 and 3. CB: Subfamily Cactoideae,Tribe Browningieae; CC: Subfamily Cactoideae, Tribe Cacteae; CCE: Subfamily Cactoideae, Tribe Cereeae; CH: Subfamily Cactoideae, Tribe Hylo-cereeae; CN: Subfamily Cactoideae, Tribe Notocacteae; CP: Subfamily Cactoideae, Tribe Pachycereeae; CR: Subfamily Cactoideae, Tribe Rhipsa-lideae; CT: Subfamily Cactoideae, Tribe Trichocereeae; OA: Subfamily Opuntioideae, Tribe Austrocylindropuntieae; OC: Subfamily Opuntioideae,Tribe Cylindropuntieae; OO: Subfamily Opuntioideae, Tribe Opuntieae; OP: Subfamily Opuntioideae, Tribe Pterocacteae; OT: Subfamily Opuntioi-deae, Tribe Tephrocacteae; M: Subfamily Maihuenioideae. PE: Subfamily Pereskioideae. The bars in the left graph indicate the percentage ofnon-invasive species (black) against percentage of invasive species (red) per tribe or subfamily.





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Figure 5. The distribution of invasive cacti within (A) genera, (B) growth forms and (C) IUCN Red List categories. Significance levels were deter-mined by comparing the number of invasive vs. number of non-invasive for any group to all other taxa using Fisher’s exact test. Confidenceintervals were determined for the per cent invasive or introduced based on an assumption of binomial errors.





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attractiveness (and hence the popularity of the speciesfor horticulture) are also important for invasiveness. Forexample, trees used for ornamentation are often selectedfor their long-lasting displays of brightly coloured fleshyfruits that are attractive to a wide range of generalistseed dispersers (Richardson and Rejmanek 2011). Asanother example, Australian Acacia species used forornamentation have rapid growth rates and can surviveand flourish in nutrient-poor, arid or degraded sites(Richardson et al. 2011; Donaldson et al. 2014). Ornamen-tal cacti, on the other hand, appear to be selected for fea-tures other than those that directly enhance invasiveness;in particular, species that survive without much input andgrow slowly are favoured (i.e. more K-selected thanr-selected). The most popular cactus species in the globalornamental trade belong to the genus Mammillaria(Novoa et al., unpubl. data); these species are valued fortheir globose growth form more than any other feature.As no Mammillaria spp. are invasive and no globosetaxa are invasive (Fig. 5A and B), it is likely that thisgenus/growth form poses little risk of invasion or impactdue to its ecological strategy.


Besides the past and current efforts directed at prevent-ing new introductions of species already known as inva-sive elsewhere, additional protocols for regulating riskare needed. This is because most contemporary introduc-tions and dissemination of cacti are of ornamental taxa,many of which do not have well-documented introduc-tion/invasion histories. Our results suggest that delimita-tions based on membership to genera, position in thephylogeny, growth form and native range size need tobe considered to produce objective and defendableapproaches for formal risk assessments.

Primary attention with regard to invasiveness in cactineeds to be given to taxa in the 13 genera of Cactaceaethat consistently display invasive tendencies. These gen-era (comprising 538 species) share certain characteristicswhich include prolific fruiting, strong vegetative reproduc-tion and effective dispersal mechanisms (Walters et al.2011). This pattern is particularly seen in the ‘opuntoidcacti’ (i.e. the genera Austrocylindropuntia, Cylindropuntiaand Opuntia), which have been classed together asWeeds of National Significance in Australia (Lloyd andReeves 2014). Our phylogenetic analysis (Fig. 4) providessupport for this approach. Invasive taxa are relativelycommon in the Opuntioideae clade, whereas the inci-dence of invasiveness in the tribe Cacteae is zero. Inter-estingly, genera from these clades are well representedin the global horticultural trade, and presumably havesimilar levels of dissemination and introduction effort(Novoa et al., unpubl. data). This pattern in Cactaceae

is similar to that seen in conifers. Twenty-eight ofthe known invasive conifer taxa belong to one family(Pinaceae) and 21 of these are in a single genus—Pinus(Richardson and Rejmanek 2004).

One noticeable feature of Cactaceae is the range ofgrowth forms within the family. Unlike most plant groupsstudied to date, invasiveness in cacti is strongly asso-ciated with particular growth forms. All invasive cactiare angled, cylindrical, flattened-padded or sprawling.The reason for high levels of invasiveness in these growthforms probably relates to the strong ability of taxa inthese groups to grow vegetatively from cuttings whichcan allow for rapid dispersal (Anderson 2001).

It would seem that the same traits that allow some cac-tus species to become widespread in their native rangescontribute to their ability to overcome abiotic filters andsuccessfully establish in new regions. No cactus speciesthat are of conservation concern in their native rangeshave been recorded as invasive, and there is a strong cor-relation between invasiveness and native range size in, forexample, the genus Opuntia. A similar pattern has beenobserved for other model groups. For example, AustralianAcacia species with large native ranges and low percola-tion exponents (i.e. high population increase rate) aremost likely to be introduced and become naturalized (Huiet al. 2011). Large native range size has been shown to be agood predictor of invasiveness and invasion success inmany, but not all, plant groups (e.g. Proches et al. 2012;Moodley et al. 2013; Potgieter et al. 2013).

ConclusionsCacti are already among the most widespread and dam-aging of invasive alien plants in some parts of the world.The huge and growing interest in many cacti for ornamen-tation has created an important new pathway for theintroduction and dissemination of a growing number ofcactus taxa around the world. Many new invasion eventsare expected in the future. There is clearly a need to regu-late the movement of cacti recorded as invasive elsewhere(currently only 3 % of the species in the family) to areassuitable for invasion, as well as taxa that pose a high riskof becoming invasive. Results from this study suggest thatrisk assessment protocols for cacti should evaluate taxaaccording to genera, position in the phylogeny of the fam-ily, growth form, and, potentially, native range size.

Sources of FundingFunding for this work was provided by the Workingfor Water (WfW) Programme of the South AfricanDepartment of Environmental Affairs and the DST-NRFCentre of Excellence for Invasion Biology (C.I.B) as partof the C.I.B/WfW collaborative research programme on





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‘Research for Integrated Management of Invasive AlienSpecies’. D.M.R. acknowledges additional support fromthe National Research Foundation (grant 85417) andthe Oppenheimer Memorial Trust.

Contributions by the AuthorsA.N., J.R.U.W. and D.M.R. conceived the idea. A.N. collectedthe data. A.N. and J.R.U.W. ran the statistics, J.J.L.R. builtthe phylogeny and M.P.R. undertook climate matching.A.N. led the writing with assistance from others.

Conflicts of Interest StatementNone declared.

AcknowledgementsComments from two anonymous reviewers improved thismanuscript. The authors also thank Rodrigo Pintos whohelped with the design of Fig. 1.

Supporting InformationThe following Supporting Information is available in theonline version of this article –

Table S1. Examples of sources of information on cactusspecies.

Table S2. List of cactus species. *57 species recorded asinvasive outside their native range.

Figure S3. Potential species richness based on availablenative records only of 39 invasive cactus species.

Table S4. Examples of cacti and succulents Journals.Table S5. The date of foundation of each society is

shown.

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Supporting information. File 1. Examples of sources of information on cactus species.

Category Source Region/extent

Alien species list Weeds of National Significance (WoNS) Australia

Alien species list Decreto-Lei n.º 565/99 de 21 de Dezembro Portugal

Alien species list National Environmental Management: Biodiversity Act 2004 (Act No. 10 of 2004) Alien and Invasive Species Lists, 2014. Government Gazette of South Africa, vol. 559, issue 37886, pp. 3–80. Pretoria.

South Africa

Alien species list Real Decreto 630/2013, de 2 de Agosto, por el que se regula el catalogo español de especies exóticas invasoras Spain

Book Anderson EF. 2001. The cactus family. Timber Press Global

Book Lloyd S, Reeves A. 2014. Situation Statement on Opuntioid Cacti (Austrocylindropuntia spp ., Cylindropuntia spp . and Opuntia spp.) in Western Australia. Department of Agriculture and Food. Goverment of Western Australia

Australia

Book Walters M, Figueiredo E, Crouch NR, Winter PJD, Smith GF, Zimmermann HG, Mashope BK. 2011. Naturalised and invasive succulents of southern Africa (Y Samyn, D VandenSpiegel, and J Degreef, Eds.). Abc Taxa

South Africa

Book Benson L. 1982. The cacti of the United States and Canada. Standford: Stanford University Press United States and Canada

Cactus and succulents’ journal Cactusvrieden Belgium

Cactus and succulents’ journal Terra seca France

Cactus and succulents’ journal Succulenta Netherlands

Cactus and succulents’ journal Cactus-adventures Spain

Peer-review article Ortega-Baes P, Aparicio-González M, Galíndez G, del Fueyo P, Sühring S, Rojas-Aréchiga M. 2010. Are cactus growth forms related to germination responses to light? A test using Echinopsis species. Acta Oecologica 36: 339–342

Argentina

Peer-review article Essl F, Kobler J. 2009. Spiny invaders – Patterns and determinants of cacti invasion in Europe. Flora - Morphology, Distribution, Functional Ecology of Plants 204: 485–494

Europe

Peer-review article Dean WR., Milton S. 2000. Directed dispersal of Opuntia species in the Karoo, South Africa: are crows the responsible agents? Journal of Arid Environments 45: 305–314

South Africa

Peer-review article Vilà M, Gimeno I. 2003. Seed predation of two alien Opuntia species invading Mediterranean communities. Plant Ecology 167: 1–8.

Spain

Online http://cactusguide.com/ Global

Online http://cactus-art.biz/ Global

Online http://www.environment.gov.au/cgi-bin/biodiversity/invasive/weeds Australia

Online http://www.invasives.org.za/ South Africa

http://cactusguide.com/

http://cactus-art.biz/

http://www.environment.gov.au/cgi-bin/biodiversity/invasive/weeds

http://www.invasives.org.za/

Supporting information. File 2. List of Cactus species. *: 57species recorded as invasive outside their native range

Acanthocalycium ferrarii Acanthocalycium klimpelianum Acanthocalycium spiniflorum Acanthocereus baxaniensis Acanthocereus colombianus Acanthocereus horridus Acanthocereus occidentalis Acanthocereus subinermis Acanthocereus tetragonus* Acharagma aguirreana Acharagma roseana Ariocarpus agavoides Ariocarpus bravoanus Ariocarpus fissuratus Ariocarpus kotchebeyanus Ariocarpus retusus Ariocarpus scaphirostris Ariocarpus trigonus Armatocereus arduus Armatocereus brevispinus Armatocereus cartwrightianus Armatocereus godingianus Armatocereus humilis Armatocereus laetus Armatocereus mataranus Armatocereus matucanensis Armatocereus oligogonus Armatocereus procerus Armatocereus rauhii Armatocereus riomajensis

Armatocereus rupicola Arrojadoa albiflora Arrojadoa bahiensis Arrojadoa beateae Arrojadoa dinae Arrojadoa eriocaulis Arrojadoa multiflora Arrojadoa penicillata Arrojadoa rhodantha Arthrocereus glaziovii Arthrocereus melanurus Arthrocereus rondonianus Arthrocereus spinosissimus Astrophytum asterias Astrophytum capricorne Astrophytum caput-medusae Astrophytum coahuilense Astrophytum myriostigma Astrophytum ornatum Austrocactus bertinii Austrocactus coxii Austrocactus patagonicus Austrocactus philippii Austrocactus spiniflorus Austrocylindropuntia cylindrica* Austrocylindropuntia floccosa Austrocylindropuntia hirschii Austrocylindropuntia lagopus Austrocylindropuntia pachypus Austrocylindropuntia punta-caillan

Austrocylindropuntia shaferi Austrocylindropuntia subulata* Austrocylindropuntia verschaffeltii Austrocylindropuntia vestita Austrocylindropuntia yanganucensis Aztekium hintonii Aztekium ritteri Bergerocactus emoryi Blossfeldia liliputana Brachycereus nesioticus Brasilicereus estevesii Brasilicereus markgrafii Brasilicereus phaeacanthus Brasiliopuntia brasiliensis Browningia albiceps Browningia altissima Browningia amstutziae Browningia caineana Browningia candelaris Browningia chlorocarpa Browningia columnaris Browningia hertlingiana Browningia microsperma Browningia pilleifera Browningia viridis Calymmanthium substerile Carnegiea gigantea Cephalocereus apicephalium Cephalocereus columna-trajani Cephalocereus nizandensis

Cephalocereus senilis Cephalocereus totolapensis Cephalocleistocactus chrysocephalus Cereus adelmarii Cereus aethiops Cereus albicaulis Cereus argentinensis Cereus bicolor Cereus braunii Cereus cochabambensis Cereus comarapanus Cereus estevesii Cereus fernambucensis Cereus fricii Cereus haageanus Cereus hankeanus Cereus hexagonus* Cereus hildmannianus* Cereus horrispinus Cereus huilunchu Cereus insularis Cereus jamacaru* Cereus kroenleinii Cereus lamprospermus Cereus lanosus Cereus mirabella Cereus mortensenii Cereus pachyrhizus Cereus phatnospermus Cereus pierre-braunianus

Cereus repandus Cereus roseiflorus Cereus saddianus Cereus spegazzinii Cereus stenogonus Cereus tacuaralensis Cereus trigonodendron Cereus validus Cereus vargasianus Cintia knizei Cipocereus bradei Cipocereus crassisepalus Cipocereus laniflorus Cipocereus minensis Cipocereus pusilliflorus Cleistocactus acanthurus Cleistocactus baumannii Cleistocactus brookeae Cleistocactus buchtienii Cleistocactus candelilla Cleistocactus chotaensis Cleistocactus clavispinus Cleistocactus x crassiserpens Cleistocactus dependens Cleistocactus ferrarii Cleistocactus fieldianus Cleistocactus grossei Cleistocactus hildegardiae Cleistocactus hoffmannii Cleistocactus hyalacanthus Cleistocactus hystrix Cleistocactus icosagonus Cleistocactus jajoanus Cleistocactus laniceps Cleistocactus longiserpens Cleistocactus luribayensis Cleistocactus micropetalus Cleistocactus morawetzianus Cleistocactus muyurinensis

Cleistocactus neoroezlii Cleistocactus orthogonus Cleistocactus pachycladus Cleistocactus palhuayensis Cleistocactus paraguariensis Cleistocactus parapetiensis Cleistocactus parviflorus Cleistocactus peculiaris Cleistocactus piraymirensis Cleistocactus plagiostoma Cleistocactus pungens Cleistocactus reae Cleistocactus ritteri Cleistocactus roezlii Cleistocactus samaipatanus Cleistocactus sepium Cleistocactus serpens Cleistocactus sextonianus Cleistocactus smaragdiflorus Cleistocactus strausii Cleistocactus sulcifer Cleistocactus tarijensis Cleistocactus tenuiserpens Cleistocactus tominensis Cleistocactus tupizensis Cleistocactus varispinus Cleistocactus viridiflorus Cleistocactus vulpis-cauda Cleistocactus winteri Cleistocactus xylorhizus Cochemiea halei Cochemiea maritima Cochemiea pondii Cochemiea poselgeri Cochemiea setispina Coleocephalocereus aureus Coleocephalocereus braunii Coleocephalocereus buxbaumianus Coleocephalocereus fluminensis

Coleocephalocereus goebelianus Coleocephalocereus pluricostatus Coleocephalocereus purpureus Coleocephalocereus uebelmanniorum Consolea corallicola Consolea falcata Consolea macracantha Consolea millspaughii Consolea moniliformis Consolea nashii Consolea picardae Consolea rubescens Consolea spinosissima Copiapoa ahremephiana Copiapoa angustiflora Copiapoa atacamensis Copiapoa bridgesii Copiapoa calderana Copiapoa cinerascens Copiapoa cinerea Copiapoa conglomerata Copiapoa coquimbana Copiapoa decorticans Copiapoa echinoides Copiapoa esmeraldana Copiapoa fieldleriana Copiapoa grandiflora Copiapoa haseltoniana Copiapoa humilis Copiapoa hypogaea Copiapoa krainziana Copiapoa laui Copiapoa longistaminea Copiapoa malletiana Copiapoa marginata Copiapoa megarhiza Copiapoa mollicula Copiapoa serpentisulcata Copiapoa solaris

Copiapoa taltalensis Copiapoa tenuissima Copiapoa tocopillana Corryocactus apiciflorus Corryocactus aureus Corryocactus ayacuchoensis Corryocactus brachypetalus Corryocactus brevistylus Corryocactus chachapoyensis Corryocactus charazanensis Corryocactus erectus Corryocactus huincoensis Corryocactus melanotrichus Corryocactus pulquiensis Corryocactus quadrangularis Corryocactus squarrosus Corynopuntia aggeria Coryphanta compacta Coryphantha calipensis Coryphantha calochlora Coryphantha clavata Coryphantha cornifera Coryphantha cornuta Coryphantha delaetiana Coryphantha difficilis Coryphantha duragensis Coryphantha echinoidea Coryphantha echinus Coryphantha elephantidens Coryphantha erecta Coryphantha georgii Coryphantha glanduligera Coryphantha glassii Coryphantha gracilis Coryphantha grata Coryphantha guerkeana Coryphantha hintoniorum Coryphantha indensis Coryphantha jalpanensis

Coryphantha kracikii Coryphantha longicornis Coryphantha macromeris Coryphantha maiz-tablasensis Coryphantha maliterrarum Coryphantha melleospina Coryphantha neglecta Coryphantha nickelsiae Coryphantha octacantha Coryphantha odorata Coryphantha ottonis Coryphantha pallida Coryphantha poselgeriana Coryphantha potosiana Coryphantha pseudoechinus Coryphantha pseudonickelsiae Coryphantha pseudoradians Coryphantha pulleineana Coryphantha pusilliflora Coryphantha pycnacantha Coryphantha radians Coryphantha ramillosa Coryphantha recurvata Coryphantha reduncispina Coryphantha retusa Coryphantha robustispina Coryphantha salinensis Coryphantha sulcata Coryphantha sulcolanata Coryphantha tripugionacantha Coryphantha unicornis Coryphantha vaupeliana Coryphantha vogtherriana Coryphantha werdermannii Coryphantha wohlschlageri Cumarinia odorata Cumulopuntia boliviana Cumulopuntia chichensis Cumulopuntia corotilla

Cumulopuntia crassicylindrica Cumulopuntia dactylifera Cumulopuntia frigida Cumulopuntia fulvicoma Cumulopuntia galerasensis Cumulopuntia hystrix Cumulopuntia ignescens Cumulopuntia mistiensis Cumulopuntia pentlandii Cumulopuntia pyrrhacantha Cumulopuntia rossiana Cumulopuntia sphaerica Cumulopuntia ticnamarensis Cumulopuntia tortispina Cumulopuntia tumida Cumulopuntia unguispina Cumulopuntia zehnderi Cylindropuntia abyssi Cylindropuntia acanthocarpa Cylindropuntia alcahes* Cylindropuntia anteojoensis Cylindropuntia arbuscula Cylindropuntia bigelovii Cylindropuntia californica Cylindropuntia calmalliana Cylindropuntia caribaea Cylindropuntia cedrosensis Cylindropuntia cholla Cylindropuntia davisii Cylindropuntia echinocarpa Cylindropuntia fulgida* Cylindropuntia ganderi Cylindropuntia hystrix Cylindropuntia imbricata* Cylindropuntia kleiniae* Cylindropuntia leptocaulis* Cylindropuntia lindsayi Cylindropuntia molesta Cylindropuntia multigeniculata

Cylindropuntia munzii Cylindropuntia pallida Cylindropuntia prolifera Cylindropuntia ramosissima Cylindropuntia rosea* Cylindropuntia sanfelipensis Cylindropuntia santamaria Cylindropuntia spinosior* Cylindropuntia tesajo Cylindropuntia thurberi Cylindropuntia tunicata* Cylindropuntia versicolor Cylindropuntia whipplei Cylindropuntia wolfii Cylindropuntia x congesta Cylindropuntia x deserta Cylindropuntia x fosbergii Cylindropuntia x kelvinensis Cylindropuntia x neoarbuscula Cylindropuntia x tetracantha Cylindropuntia x viridiflora Cylindropuntia x vivipara Dendrodendus nudiflorus Dendrodendus undulosus Denmoza rhodacantha Discocactus bahiensis Discocactus cangaensis Discocactus catingicola Discocactus cephaliaciculosus Discocactus diersianus Discocactus ferricola Discocactus hartmannii Discocactus heptacanthus Discocactus horstii Discocactus placentiformis Discocactus pseudoinsignis Discocactus subterraneo-proliferans Discocactus zehntneri Disocactus ackermannii

Disocactus amazonicus Disocactus aurantiacus Disocactus biformis Disocactus cinnabarinus Disocactus eichlamii Disocactus flagelliformis Disocactus kimnachii Disocactus macdougallii Disocactus macranthus Disocactus martianus Disocactus nelsonii Disocactus phyllanthoides Disocactus quezaltecus Disocactus schrankii Disocactus speciosus Echinocactus grusonii Echinocactus horizonthalonius Echinocactus parryi Echinocactus platyacanthus Echinocactus polycephalus Echinocactus texensis Echinocereus adustus Echinocereus apachensis Echinocereus barthelowanus Echinocereus berlandieri Echinocereus bonkerae Echinocereus boyce-thompsonii Echinocereus brandegeei Echinocereus bristolii Echinocereus chisoensis Echinocereus cinerascens Echinocereus coccineus Echinocereus dasyacanthus Echinocereus engelmannii Echinocereus enneacanthus Echinocereus fasciculatus Echinocereus fendleri Echinocereus ferreirianus Echinocereus freudenbergeri

Echinocereus grandis Echinocereus klapperi Echinocereus knippelianus Echinocereus laui Echinocereus ledingii Echinocereus leucanthus Echinocereus longisetus Echinocereus mapimiensis Echinocereus maritimus Echinocereus mojavensis Echinocereus nicholii Echinocereus nivosus Echinocereus ortegae Echinocereus palmeri Echinocereus pamanesiorum Echinocereus papillosus Echinocereus parkeri Echinocereus pectinatus Echinocereus pensilis Echinocereus pentalophus Echinocereus polyacanthus Echinocereus poselgeri Echinocereus primolanatus Echinocereus pseudopectinatus Echinocereus pulchellus Echinocereus rayonesensis Echinocereus reichenbachii Echinocereus rigidissimus Echinocereus russanthus Echinocereus scheeri Echinocereus schereri Echinocereus schmollii Echinocereus sciurus Echinocereus scopulorum Echinocereus spinigemmatus Echinocereus stoloniferus Echinocereus stramineus Echinocereus subinermis Echinocereus triglochidiatus

Echinocereus viereckii Echinocereus viridiflorus Echinocereus websterianus Echinocereus x roetteri Echinomastus erectocentrus Echinomastus gautii Echinomastus intertextus Echinomastus johnsonii Echinomastus mariposensis Echinomastus unguispinus Echinomastus warnockii Echinopsis adolfofriedrichii Echinopsis ancistrophora Echinopsis angelesii Echinopsis antezanae Echinopsis arboricola Echinopsis arebaloi Echinopsis atacamensis Echinopsis aurea Echinopsis backebergii Echinopsis baldiana Echinopsis bertramiana Echinopsis bolligeriana Echinopsis boyuibensis Echinopsis brasiliensis Echinopsis breviflora Echinopsis bridgesi Echinopsis bruchii Echinopsis x cabrerae Echinopsis caineana Echinopsis cajasensis Echinopsis calliantholilacina Echinopsis callichroma Echinopsis calochlora Echinopsis camarguensis Echinopsis candicans Echinopsis cephalomacrostibas Echinopsis cerdana Echinopsis chalaensis

Echinopsis chamaecereus* Echinopsis chiloensis Echinopsis chrysantha Echinopsis chrysochete Echinopsis cinnabarina Echinopsis clavata Echinopsis cochabambensis Echinopsis comarapana Echinopsis conaconensis Echinopsis coquimbana Echinopsis coronata Echinopsis cotacajesii Echinopsis crassicaulis Echinopsis cuzcoensis Echinopsis densispina Echinopsis derenbergii Echinopsis deserticola Echinopsis escayachensis Echinopsis eyriesii Echinopsis fabrisii Echinopsis famatimensis Echinopsis ferox Echinopsis formosa Echinopsis friedrichii Echinopsis glauca Echinopsis glaucina Echinopsis haematantha Echinopsis hahniana Echinopsis hammerschmidii Echinopsis hertrichiana Echinopsis huascha Echinopsis huotii Echinopsis hystrichoides Echinopsis ibicuatensis Echinopsis jajoana Echinopsis kladiwaiana Echinopsis klingleriana Echinopsis knuthiana Echinopsis korethroides

Echinopsis lageniformis Echinopsis lamprochlora Echinopsis lateritia Echinopsis leucantha Echinopsis litoralis Echinopsis macrogona Echinopsis mamillosa Echinopsis marsoneri Echinopsis mataranensis Echinopsis maximiliana Echinopsis meyeri Echinopsis mieckleyi Echinopsis minuana Echinopsis mirabilis Echinopsis molesta Echinopsis nigra Echinopsis obrepanda Echinopsis oxygona Echinopsis pachanoi Echinopsis pampana Echinopsis pentlandii Echinopsis peruviana Echinopsis pojoensis Echinopsis pseudomamillosa Echinopsis pugionacantha Echinopsis quadratiumbonata Echinopsis rhodotricha Echinopsis riviere-de-caraltii Echinopsis rojasii Echinopsis saltensis Echinopsis sanguiniflora Echinopsis santaensis Echinopsis schickendantzii* Echinopsis schieliana Echinopsis schoenii Echinopsis schrieteri Echinopsis scopulicola Echinopsis silvestrii Echinopsis skottsbergii

Echinopsis smrziana Echinopsis spachiana* Echinopsis spinibarbis Echinopsis strigosa Echinopsis subdenudata Echinopsis sucrensis Echinopsis tacaquirensis Echinopsis taratensis Echinopsis tarijensis Echinopsis tarmaensis Echinopsis tegeleriana Echinopsis terscheckii Echinopsis thelegona Echinopsis thelegonoides Echinopsis thionantha Echinopsis tiegeliana Echinopsis trichosa Echinopsis tubiflora Echinopsis tulhuayacensis Echinopsis tunariensis Echinopsis uyupampensis Echinopsis vasquezii Echinopsis vatteri Echinopsis volliana Echinopsis walteri Echinopsis werdermannii Echinopsis yuquina Epiphyllum anguliger Epiphyllum cartagense Epiphyllum caudatum Epiphyllum columbiense Epiphyllum costaricense Epiphyllum crenatum Epiphyllum floribundum Epiphyllum grandilobum Epiphyllum guatemalense Epiphyllum hookeri Epiphyllum laui Epiphyllum lepidocarpum

Epiphyllum oxypetalum Epiphyllum phyllanthus Epiphyllum pittieri Epiphyllum pumilum Epiphyllum rubrocoronatum Epiphyllum thomasianum Epiphyllum trimetrale Epithelantha bokei Epithelantha micromeris Eriosyce aericarpa Eriosyce andreaeana Eriosyce aspillagae Eriosyce aurata Eriosyce bulbocalyx Eriosyce chilensis Eriosyce confinis Eriosyce crispa Eriosyce curvispina Eriosyce engleri Eriosyce esmeraldana Eriosyce garaventae Eriosyce heinrichiana Eriosyce islayensis Eriosyce krausii Eriosyce kunzei Eriosyce laui Eriosyce limariensis Eriosyce marksiana Eriosyce napina Eriosyce occulta Eriosyce odieri Eriosyce omasensis Eriosyce recondita Eriosyce rodentiophila Eriosyce senilis Eriosyce sociabilis Eriosyce strausiana Eriosyce subgibbosa Eriosyce taltalensis

Eriosyce tenebrica Eriosyce umadeave Eriosyce vertongenii Eriosyce villicumensis Eriosyce villosa Escobaria albicolumnaria Escobaria alversonii Escobaria chihuahuensis Escobaria cubensis Escobaria dasyacantha Escobaria deserti Escobaria duncanii Escobaria emskoetteriana Escobaria guadalupensis Escobaria hesteri Escobaria laredoi Escobaria lloydii Escobaria minima Escobaria missouriensis Escobaria orcuttii Escobaria organensis Escobaria robbinsorum Escobaria sandbergii Escobaria sneedii Escobaria tuberculosa Escobaria villardii Escobaria vivipara Escobaria zilziana Escontria chiotilla Espostoa baumannii Espostoa blossfeldiorum Espostoa calva Espostoa frutescens Espostoa guentheri Espostoa huanucoensis Espostoa hylaea Espostoa lanata Espostoa lanianuligera Espostoa melanostele

Espostoa mirabilis Espostoa nana Espostoa ritteri Espostoa ruficeps Espostoa senilis Espostoa superba Espostoa utcubambensis Espostoopsis dybowskii Estevesia alex-bragae Eulychnia acida Eulychnia breviflora Eulychnia castanea Eulychnia iquiquensis Eulychnia ritteri Facheiroa cephaliomelana Facheiroa squamosa Facheiroa ulei Ferocactus alamosanus Ferocactus chrysacanthus Ferocactus cylindraceus Ferocactus diguetii Ferocactus eastwoodiae Ferocactus echidne Ferocactus emoryi Ferocactus flavovirens Ferocactus fordii Ferocactus glaucescens Ferocactus gracilis Ferocactus haematacanthus Ferocactus hamatacanthus Ferocactus herrehae Ferocactus histrix Ferocactus johnstonianus Ferocactus latispinus Ferocactus lindsayi Ferocactus macrodiscus Ferocactus peninsulae Ferocactus pilosus Ferocactus pottsi

Ferocactus robustus Ferocactus santa-maria Ferocactus schwarzii Ferocactus tiburonensis Ferocactus townsendianus Ferocactus viridescens Ferocactus wislizeni Frailea buenekeri Frailea buiningiana Frailea castanea Frailea cataphracta Frailea chiquitana Frailea curvispina Frailea friedrichii Frailea gracillima Frailea grahliana Frailea knippeliana Frailea mammifera Frailea perumbilicata Frailea phaeodisca Frailea pseudopulcherrima Frailea pumila Frailea pygmaea Frailea schilinzkyana Geohintonia mexicana Grusonia aggeria Grusonia agglomerata Grusonia bradtiana Grusonia bulbispina Grusonia clavata Grusonia dumetorum Grusonia emoryi Grusonia grahamii Grusonia Invicta Grusonia kunzei Grusonia marenae Grusonia moelleri Grusonia parishii Grusonia pulchella

Grusonia reflexispina Grusonia robertsii Grusonia schottii Grusonia vilis Gymnocalycium albiareolatum Gymnocalycium ambatoense Gymnocalycium amerhauseri Gymnocalycium andreae Gymnocalycium angelae Gymnocalycium anisitsii Gymnocalycium baldianum Gymnocalycium bayrianum Gymnocalycium berchtii Gymnocalycium bodenbenderianum Gymnocalycium borthii Gymnocalycium bruchii Gymnocalycium buenekeri Gymnocalycium calochlorum Gymnocalycium capillaense Gymnocalycium carmianthum Gymnocalycium castellanosii Gymnocalycium catamarcense Gymnocalycium chiquitanum Gymnocalycium deeszianum Gymnocalycium delaetii Gymnocalycium denudatum Gymnocalycium erinaceum Gymnocalycium eurypleurum Gymnocalycium eytianum Gymnocalycium gibbosum Gymnocalycium horstii Gymnocalycium hossei Gymnocalycium hybopleurum Gymnocalycium hyptiacanthum Gymnocalycium kieslingii Gymnocalycium kroenleinii Gymnocalycium leeanum Gymnocalycium leptanthum Gymnocalycium mackieanum

Gymnocalycium marianae Gymnocalycium marsoneri Gymnocalycium megalothelon Gymnocalycium mesopotamicum Gymnocalycium mihanovichii Gymnocalycium monvillei Gymnocalycium mostii Gymnocalycium mucidum Gymnocalycium netrelianum Gymnocalycium neuhuberi Gymnocalycium obductum Gymnocalycium ochoterenae Gymnocalycium oenanthemum Gymnocalycium paediophilum Gymnocalycium paraguayense Gymnocalycium parvulum Gymnocalycium pflanzii Gymnocalycium platense Gymnocalycium pugionacanthum Gymnocalycium quehlianum Gymnocalycium ragonesei Gymnocalycium rauschii Gymnocalycium riojense Gymnocalycium ritterianum Gymnocalycium robustum Gymnocalycium rosae Gymnocalycium saglionis Gymnocalycium schickendantzii Gymnocalycium schroederianum Gymnocalycium spegazzinii Gymnocalycium stellatum Gymnocalycium stenopleurum Gymnocalycium striglianum Gymnocalycium stuckertii Gymnocalycium taningaense Gymnocalycium terweemeanum Gymnocalycium tillianum Gymnocalycium uebelmannianum Gymnocalycium uruguayense

Haageocereus acranthus Haageocereus albispinus Haageocereus australis Haageocereus chalaensis Haageocereus chilensis Haageocereus chryseus Haageocereus decumbens Haageocereus fascilularis Haageocereus icensis Haageocereus icosagonoides Haageocereus lanugispinus Haageocereus pacalaensis Haageocereus platinospinus Haageocereus pluriflorus Haageocereus pseudomelanostele Haageocereus pseudoversicolor Haageocereus subtilispinus Haageocereus tenuis Haageocereus versicolor Haageocereus vulpes Haageocereus zangalensis xHaagespostoa albisetata xHaagespostoa climaxantha Harrisia adscendens Harrisia alboriginum Harrisia balansae* Harrisia brookii Harrisia divaricata Harrisia earlei Harrisia eriophora Harrisia fernowii Harrisia fragans Harrisia gracilis Harrisia hurstii Harrisia martini* Harrisia nashii Harrisia pomanensis* Harrisia portoricensis Harrisia regelii

Harrisia simpsonii Harrisia taetra Harrisia taylori Harrisia tetracantha Harrisia tortuosa* Hatiora cylindrica Hatiora epiphylloides Hatiora gaertneri Hatiora herminiae Hatiora rosea Hatiora salicornioides Hatiora x graeseri Hylocereus bronxensis Hylocereus calcaratus Hylocereus costaricensis* Hylocereus escuintlensis Hylocereus estebanensis Hylocereus extensus Hylocereus guatemalensis Hylocereus lemairei Hylocereus microcladus Hylocereus minutiflorus Hylocereus monacanthus Hylocereus ocamponis Hylocereus peruvianus Hylocereus polyrhizus* Hylocereus purpusii Hylocereus scandens Hylocereus stenopterus Hylocereus triangularis* Hylocereus trigonus Hylocereus undatus* Isolatocereus dumortieri Jasminocereus thouarsii Lasiocereus fulvus Lasiocereus rupicola Leocereus bahiensis Lepismium aculeatum Lepismium bolivianum

Lepismium brevispinum Lepismium crenatum Lepismium cruciforme Lepismium houlletianum Lepismium ianthothele Lepismium incachacanum Lepismium lorentzianum Lepismium lumbricoides Lepismium micranthum Lepismium miyagawae Lepismium monacanthum Lepismium paranganiense Lepismium warmingianum Leptocereus arboreus Leptocereus assurgens Leptocereus carinatus Leptocereus ekmanii Leptocereus grantianus Leptocereus leonii Leptocereus maxonii Leptocereus paniculatus Leptocereus prostratus Leptocereus quadricostatus Leptocereus santamarinae Leptocereus scopulophilus Leptocereus sylvestris Leptocereus weingartianus Leptocereus wrightii Leuchtenbergia principis Lophophora diffusa Lophophora williamsii Maihuenia patagonica Maihuenia poeppigii Maihueniopsis archiconoidea Maihueniopsis atacamensis Maihueniopsis bonnieae Maihueniopsis camachoi Maihueniopsis clavarioides Maihueniopsis colorea

Maihueniopsis crassispina Maihueniopsis darwinii Maihueniopsis domeykoensis Maihueniopsis glomerata Maihueniopsis grandiflora Maihueniopsis minuta Maihueniopsis nigrispina Maihueniopsis ovata Maihueniopsis rahmeri Maihueniopsis subterranea Maihueniopsis tarapacana Maihueniopsis wagenknechtii Mammillaria albicans Mammillaria albicoma Mammillaria albiflora Mammillaria albilanata Mammillaria amajacensis Mammillaria angelensis Mammillaria anniana Mammillaria armillata Mammillaria aureilanata Mammillaria backebergiana Mammillaria barbata Mammillaria baumii Mammillaria beneckei Mammillaria blossfeldiana Mammillaria bocasana Mammillaria bocensis Mammillaria boelderliana Mammillaria bombycina Mammillaria boolii Mammillaria brachytrichion Mammillaria brandegeei Mammillaria canelensis Mammillaria capensis Mammillaria carmenae Mammillaria carnea Mammillaria carretii Mammillaria cerralboa

Mammillaria chinocephala Mammillaria coahuilensis Mammillaria columbiana Mammillaria compressa Mammillaria crinita Mammillaria crucigera Mammillaria decipiens Mammillaria deherdtiana Mammillaria densispina Mammillaria dioica Mammillaria discolor Mammillaria dixanthocentron Mammillaria duoformis Mammillaria ekmanii Mammillaria elongata Mammillaria eriacantha Mammillaria erythrosperma Mammillaria estebanensis Mammillaria evermanniana Mammillaria fittkaui Mammillaria flavicentra Mammillaria formosa Mammillaria fraileana Mammillaria gasseriana Mammillaria geminispina Mammillaria gigantea Mammillaria glassii Mammillaria glochidiata Mammillaria goodridgei Mammillaria grahamii Mammillaria grusonii Mammillaria guelzowiana Mammillaria guerreronis Mammillaria guillauminiana Mammillaria haageana Mammillaria hahniana Mammillaria halbingeri Mammillaria heidiae Mammillaria hernandezii

Mammillaria herrerae Mammillaria hertrichiana Mammillaria heyderi Mammillaria huitzilopochtli Mammillaria humboldtii Mammillaria hutchisoniana Mammillaria insularis Mammillaria jaliscana Mammillaria johnstonii Mammillaria karwinskiana Mammillaria klissingiana Mammillaria kraehenbuehlii Mammillaria lasiacantha Mammillaria laui Mammillaria lenta Mammillaria limonensis Mammillaria linaresensis Mammillaria lindsayi Mammillaria lloydii Mammillaria longiflora Mammillaria longimamma Mammillaria luethyi Mammillaria magallanii Mammillaria magnifica Mammillaria magnimamma Mammillaria mainiae Mammillaria mammillaris Mammillaria manana Mammillaria marcosii Mammillaria marksiana Mammillaria mathildae Mammillaria matudae Mammillaria mazatlanensis Mammillaria melaleuca Mammillaria melanocentra Mammillaria mercadensis Mammillaria meyranii Mammillaria microhelia Mammillaria miegiana

Mammillaria mieheana Mammillaria moelleriana Mammillaria morganiana Mammillaria muehlenfordtii Mammillaria multidigitata Mammillaria mystax Mammillaria napina Mammillaria neopalmeri Mammillaria nivosa Mammillaria nunezii Mammillaria orcuttii Mammillaria oteroi Mammillaria parkinsonii Mammillaria pectinifera Mammillaria peninsularis Mammillaria pennispinosa Mammillaria perbella Mammillaria perezdelarosae Mammillaria petrophila Mammillaria petterssonii Mammillaria phitauiana Mammillaria picta Mammillaria pilispina Mammillaria plumosa Mammillaria polyedra Mammillaria polythele Mammillaria pottsii Mammillaria prolifera Mammillaria rekoi Mammillaria rhodantha Mammillaria roseoalba Mammillaria saboae Mammillaria sanchez-mejoradae Mammillaria sartorii Mammillaria schiedeana Mammillaria schumannii Mammillaria schwarzii Mammillaria scrippsiana Mammillaria sempervivi

Mammillaria senilis Mammillaria sheldonii Mammillaria solisioides Mammillaria sonorensis Mammillaria sphacelata Mammillaria sphaerica Mammillaria spinosissima Mammillaria standleyi Mammillaria stella-de-tacubaya Mammillaria supertexta Mammillaria surculosa Mammillaria tayloriorum Mammillaria tepexicensis Mammillaria tetrancistra Mammillaria theresae Mammillaria thornberi Mammillaria tonalensis Mammillaria uncinata Mammillaria varieaculeata Mammillaria vetula Mammillaria voburnensis Mammillaria wagneriana Mammillaria weingartiana Mammillaria wiesingeri Mammillaria winterae Mammillaria wrightii Mammillaria xaltianguensis Mammillaria zephyranthoides Mammillaria zublerae Mammilloydia candida Matucana aurantiaca Matucana aureiflora Matucana comacephala Matucana formosa Matucana fruticosa Matucana haynei Matucana huagalensis Matucana intertexta Matucana krahnii

Matucana madisoniorum Matucana oreodoxa Matucana paucicostata Matucana polzii Matucana pujupatii Matucana ritteri Matucana tuberculata Matucana webwebaueri Melocactus x albicephalus Melocactus andinus Melocactus azureus Melocactus bahiensis Melocactus bellavistensis Melocactus braunii Melocactus broadwayi Melocactus caroli-linnaei Melocactus concinnus Melocactus conoideus Melocactus curvispinus Melocactus deinacanthus Melocactus ernestii Melocactus estevesii Melocactus glaucescens Melocactus harlowii Melocactus x horridus Melocactus intortus Melocactus lanssensianus Melocactus lemairei Melocactus levitestatus Melocactus macracanthos Melocactus matanzanus Melocactus mazelianus Melocactus neryi Melocactus oreas Melocactus pachyacanthus Melocactus paucispinus Melocactus perezassoi Melocactus peruvianus Melocactus praerupticola

Melocactus salvadorensis Melocactus schatzlii Melocactus smithii Melocactus stramineus Melocactus violaceus Melocactus zehntneri Micranthocereus albicephalus Micranthocereus auriazurensis Micranthocereus dolichospermaticus Micranthocereus estevesii Micranthocereus flaviflorus Micranthocereus hofackerianus Micranthocereus polyanthus Micranthocereus purpureus Micranthocereus streckeri Micranthocereus violaciflorus Mila caespitosa Miqueliopuntia miquelii xMyrtgerocactus lindsayi Myrtillocactus cochal Myrtillocactus eichlamii Myrtillocactus geometrizans* Myrtillocactus schenckii Neobuxbaumia euphorbioides Neobuxbaumia laui Neobuxbaumia macrocephala Neobuxbaumia mezcalaensis Neobuxbaumia multiareolata Neobuxbaumia polylopha Neobuxbaumia scoparia Neobuxbaumia squamulosa Neobuxbaumia tetetzo Neolloydia conoidea Neolloydia matehualensis Neoraimondia arequipensis Neoraimondia herzogiana Neowerdermannia chilensis Neowerdermannia vorwerkii Obregonia denegrii

Opuntia abjecta Opuntia acaulis Opuntia aciculata Opuntia alko-tuna Opuntia ammophila* Opuntia amyclaea* Opuntia anacantha Opuntia antillana Opuntia apurimacensis Opuntia arcei Opuntia armata Opuntia assumptionis Opuntia atrispina Opuntia atropes Opuntia atrovirens Opuntia auberi* Opuntia aurantiaca* Opuntia arechavaletae Opuntia aurea Opuntia aureispina Opuntia austrina Opuntia azurea Opuntia basilaris Opuntia bella Opuntia bensonii Opuntia bisetosa Opuntia boldinghii Opuntia bonplandii Opuntia borinquensis Opuntia bravoana Opuntia canterae Opuntia caracassana Opuntia cardiosperma Opuntia chaffeyi Opuntia chakensis Opuntia chavena Opuntia chihuahuensis Opuntia chisosensis* Opuntia chlorotica

Opuntia cochabambensis Opuntia cochenillifera* Opuntia cognata Opuntia colubrina Opuntia conjungens Opuntia crassa* Opuntia crystalenia Opuntia curassavica Opuntia cymochila Opuntia darrahiana Opuntia deamii Opuntia decumbens Opuntia dejecta* Opuntia delaetiana Opuntia depauperata Opuntia depressa Opuntia dillenii* Opuntia discolor Opuntia durangensis Opuntia echios Opuntia eichlamii Opuntia ekmanii Opuntia elata* Opuntia elatior* Opuntia elizondoana Opuntia ellisiana Opuntia engelmannii* Opuntia excelsa Opuntia feroacantha Opuntia ficus-indica* Opuntia fragilis Opuntia fuliginosa Opuntia galapageia Opuntia gosseliniana Opuntia guatemalensis Opuntia guilanchi Opuntia helleri Opuntia hondurensis Opuntia howeyi

Opuntia huajuapensis* Opuntia humifusa* Opuntia hyptiacantha Opuntia inaequilateralis Opuntia inaperta Opuntia infesta Opuntia insularis Opuntia jaliscana Opuntia jamaicensis Opuntia joconostle Opuntia karwinskiana Opuntia laevis Opuntia lagunae Opuntia larreyi Opuntia lasiacantha Opuntia lata Opuntia leucotricha* Opuntia lilae Opuntia limitata Opuntia littoralis Opuntia lutea Opuntia macrocentra Opuntia macrorhiza Opuntia martiniana Opuntia megacantha Opuntia megapotamica Opuntia megarhiza Opuntia megasperma Opuntia microdasys* Opuntia mieckleyi Opuntia monocantha* Opuntia montevideensis Opuntia nejapensis Opuntia neochrysacantha Opuntia nuda Opuntia orbiculata Opuntia oricola Opuntia pachona Opuntia pachyrrhiza

Opuntia pailana Opuntia pampeana Opuntia paraguayensis Opuntia parviclada Opuntia penicilligera Opuntia pennellii Opuntia phaeacantha* Opuntia pilifera Opuntia pinkavae Opuntia pittieri Opuntia pituitosa Opuntia polyacantha Opuntia pottsii Opuntia prasina Opuntia puberula Opuntia pubescens* Opuntia pumila Opuntia pusilla Opuntia pycnantha Opuntia pyriformis Opuntia pyrrhantha Opuntia quimilo Opuntia quitensis Opuntia rastrera Opuntia repens Opuntia rileyi Opuntia ritteri Opuntia robinsonnii Opuntia roborensis Opuntia robusta* Opuntia rufida Opuntia salagria Opuntia salmiana* Opuntia salvadorensis Opuntia sanguinea Opuntia santa-rita Opuntia saxicola Opuntia scheeri Opuntia schickendantzii

Opuntia schumannii Opuntia securigera Opuntia soederstromiana Opuntia spinulifera* Opuntia spraguei Opuntia stenarthra Opuntia stenopetala Opuntia streptacantha* Opuntia stricta* Opuntia strigil Opuntia subsphaerocarpa Opuntia sulfurea Opuntia tapona Opuntia taylori Opuntia tehuacana Opuntia tehuantepecana Opuntia tenuiflora Opuntia tomentella Opuntia tomentosa* Opuntia triacantha Opuntia tuna* Opuntia turbinata Opuntia undulata Opuntia urbaniana Opuntia velutina Opuntia viridirubra Opuntia vitelliniflora Opuntia wetmorei Opuntia wilcoxii Opuntia x aequatorialis Opuntia x bakeri Opuntia x columbiana Opuntia x cubensis Opuntia x curvospina Opuntia x lucayana Opuntia x occidentalis Opuntia x spinosibacca Opuntia x vaseyi Opuntia x wootonii

Oreocereus celsianus Oreocereus doelzianus Oreocereus hempelianus Oreocereus leucotrichus Oreocereus pseudofossulatus Oreocereus ritteri Oreocereus tacnaensis Oreocereus trollii Oreocereus varicolor Oroya borchersii Oroya peruviana Ortegocactus macdougallii xPacherocactus orcuttii Pachycereus fulviceps Pachycereus gatesii Pachycereus gaumeri Pachycereus grandis Pachycereus hollianus Pachycereus lepidanthus Pachycereus marginatus Pachycereus militaris Pachycereus pecten-aboriginum Pachycereus pringlei Pachycereus schottii Pachycereus tepamo Pachycereus weberi Parodia alacriportana Parodia allosiphon Parodia arnostiana Parodia aureicentra Parodia ayopayana Parodia buiningii Parodia carambeiensis Parodia chrysacanthion Parodia columnaris Parodia comarapana Parodia commutans Parodia concinna Parodia crassigibba

Parodia curvispina Parodia erinacea Parodia erubescens Parodia formosa Parodia fusca Parodia gaucha Parodia glaucina Parodia haselbergii Parodia hausteiniana Parodia herteri Parodia horstii Parodia langsdorfii Parodia leninghausii Parodia linkii Parodia maassii Parodia magnifica Parodia mammulosa Parodia meonacantha Parodia microsperma Parodia mueller-melchersii Parodia muricata Parodia neoarechavaletae Parodia neohorstii Parodia nigrispina Parodia nivosa Parodia nothominuscula Parodia nothorauschii Parodia ocampoi Parodia ottonis Parodia oxycostata Parodia penicillata Parodia permutata Parodia procera Parodia rechensis Parodia ritteri Parodia rudibuenekeri Parodia rutilans Parodia saint-pieana Parodia schumanniana

Parodia schwebsiana Parodia scopa Parodia sellowii Parodia stockingeri Parodia stuemeri Parodia subterranea Parodia tabularis Parodia taratensis Parodia tenuicylindrica Parodia tilcarensis Parodia tuberculata Parodia turbinata Parodia turecekiana Parodia warasii Parodia werdermanniana Parodia werneri Pediocactus bradyi Pediocactus despainii Pediocactus hermannii Pediocactus knowltonii Pediocactus nigrispinus Pediocactus paradinei Pediocactus peeblesianus Pediocactus simpsonii Pediocactus winkleri Pelecyphora aselliformis Pelecyphora strobiliformis Peniocereus castellae Peniocereus chiapensis Peniocereus cuixmalensis Peniocereus fosterianus Peniocereus greggii Peniocereus hirschtianus Peniocereus johnstonii Peniocereus lazaro-cardenasii Peniocereus macdougallii Peniocereus maculatus Peniocereus marianus Peniocereus oaxacensis

Peniocereus occidentalis Peniocereus rosei Peniocereus serpentinus* Peniocereus striatus Peniocereus tepalcatepecanus Peniocereus viperinus Peniocereus zopilotensis Pereskia aculeata* Pereskia aureiflora Pereskia bahiensis Pereskia bleo Pereskia diaz-romeroana Pereskia grandiflora Pereskia guamacho Pereskia horrida Pereskia lychnidiflora Pereskia marcanoi Pereskia nemorosa Pereskia portulacifolia Pereskia quinqueyana Pereskia sacharosa Pereskia stenantha Pereskia weberiana Pereskia zinniiflora Pereskiopsis aquosa Pereskiopsis blakeana Pereskiopsis diguetii Pereskiopsis gatesii Pereskiopsis kellermanii Pereskiopsis porteri Pereskiopsis rotundifolia Pereskiopsis spathulata Pfeiffera asuntapatensis Pilosocereus albisummus Pilosocereus alensis Pilosocereus arrabidae Pilosocereus aureispinus Pilosocereus aurisetus Pilosocereus azulensis

Pilosocereus brasiliensis Pilosocereus catingicola Pilosocereus chrysacanthus Pilosocereus chrysostele Pilosocereus collinsii Pilosocereus densiareolatus Pilosocereus diersianus Pilosocereus estevesii Pilosocereus flavipulvinatus Pilosocereus flexibilispinus Pilosocereus floccosus Pilosocereus frewenii Pilosocereus fulvilanatus Pilosocereus glaucochrous Pilosocereus gounellei Pilosocereus lanuginosus Pilosocereus leucocephalus Pilosocereus machrisii Pilosocereus magnificus Pilosocereus multicostatus Pilosocereus occultiflorus Pilosocereus oligolepis Pilosocereus pachycladus Pilosocereus pentaedrophorus Pilosocereus piauhyensis Pilosocereus polygonus Pilosocereus purpusii Pilosocereus quadricentralis Pilosocereus royenii Pilosocereus x subsimilis Pilosocereus tuberculatus Pilosocereus ulei Pilosocereus vilaboensis Polaskia chende Polaskia chichipe Praecereus euchlorus Praecereus saxicola Pseudoacanthocereus brasiliensis Pseudoacanthocereus sicariguensis

Pseudorhipsalis acuminata Pseudorhipsalis alata Pseudorhipsalis amazonica Pseudorhipsalis himantoclada Pseudorhipsalis horichii Pseudorhipsalis lankesteri Pseudorhipsalis ramulosa Pterocactus araucanus Pterocactus australis Pterocactus fischeri Pterocactus gonjianii Pterocactus hickenii Pterocactus megiolii Pterocactus reticulatus Pterocactus tuberosus Pterocactus valentinii Pygmaeocereus bieblii Pygmaeocereus bylesianus Pygmaeocereus familiaris Quiabentia verticillata Quiabentia zehntneri Rauhocereus riosaniensis Rebutia albiflora Rebutia albipectinata Rebutia arenacea Rebutia aureiflora Rebutia brunescens Rebutia caineana Rebutia canigueralii Rebutia cardenasiana Rebutia cintia Rebutia cylindrica Rebutia deminuta Rebutia einsteinii Rebutia fabrisii Rebutia fidaiana Rebutia fiebrigii Rebutia flavistyla Rebutia fulviseta

Rebutia gonjianii Rebutia heliosa Rebutia huasiensis Rebutia krugerae Rebutia leucanthema Rebutia marsoneri Rebutia mentosa Rebutia minuscula Rebutia narvaecensis Rebutia neocumingii Rebutia neumanniana Rebutia nigricans Rebutia oligacantha Rebutia padcayensis Rebutia perplexa Rebutia pseudodeminuta Rebutia pulvinosa Rebutia pygmaea Rebutia ritteri Rebutia simoniana Rebutia spegazziniana Rebutia spinosissima Rebutia steinbachii Rebutia steinmannii Rebutia wessneriana Rebutia xanthocarpa Rhipsalis agudoensis Rhipsalis baccifera Rhipsalis burchellii Rhipsalis campos-portoana Rhipsalis cereoides Rhipsalis cereuscula Rhipsalis clavata Rhipsalis crispata Rhipsalis cuneata Rhipsalis dissimilis Rhipsalis elliptica Rhipsalis ewaldiana Rhipsalis floccosa

Rhipsalis goebeliana Rhipsalis grandiflora Rhipsalis hoelleri Rhipsalis juengeri Rhipsalis lindbergiana Rhipsalis mesembryanthemoides Rhipsalis micrantha Rhipsalis neves-armondii Rhipsalis oblonga Rhipsalis occidentalis Rhipsalis olivifera Rhipsalis ormindoi Rhipsalis pacheco-leonis Rhipsalis pachyptera Rhipsalis paradoxa Rhipsalis pentaptera Rhipsalis pilocarpa Rhipsalis pulchra Rhipsalis puniceodiscus Rhipsalis russellii Rhipsalis sulcata Rhipsalis teres Rhipsalis triangularis Rhipsalis trigona Samaipaticereus corroanus Schlumbergera kautskyi Schlumbergera microsphaerica Schlumbergera opuntioides Schlumbergera orssichiana Schlumbergera russelliana Schlumbergera truncate Schlumbergera x buckleyi Schlumbergera x exotica Schlumbergera x reginae Sclerocactus brevihamatus Sclerocactus brevispinus Sclerocactus glaucus Sclerocactus mesae-verdae Sclerocactus nyensis

Sclerocactus papyracanthus Sclerocactus parviflorus Sclerocactus polyancistrus Sclerocactus pubispinus Sclerocactus scheeri Sclerocactus sileri Sclerocactus spinosior Sclerocactus uncinatus Sclerocactus wetlandicus Sclerocactus whipplei Sclerocactus wrightiae Selenicereus anthonyanus Selenicereus atropilosus Selenicereus boeckmannii Selenicereus brevispinus Selenicereus chontalensis Selenicereus chrysocardium Selenicereus coniflorus Selenicereus donkelaari Selenicereus grandiflorus Selenicereus hamatus Selenicereus hondurensis Selenicereus inermis Selenicereus innesii Selenicereus macdonaldiae* Selenicereus megalanthus Selenicereus murrillii Selenicereus nelsonii Selenicereus pteranthus Selenicereus rubineus Selenicereus setaceus Selenicereus spinulosus Selenicereus testudo Selenicereus tricae Selenicereus urbanianus Selenicereus vagans Selenicereus validus Selenicereus wercklei Selenicereus wittii

Siccobaccatus estevesii Siccobaccatus insigniflorus Stenocactus coptonogonus Stenocactus crispatus Stenocactus hastatus Stenocactus multicostatus Stenocactus obvallatus Stenocactus ochoterenanus Stenocactus phyllacanthus Stenocactus rectispinus Stenocactus sulfureus Stenocactus vaupelianus Stenocereus alamosensis Stenocereus aragonii Stenocereus beneckei Stenocereus chacalapensis Stenocereus chrysocarpus Stenocereus eichlamii Stenocereus eruca Stenocereus fimbriatus Stenocereus fricii Stenocereus griseus Stenocereus gummosus Stenocereus kerberi Stenocereus laevigatus Stenocereus martinezii Stenocereus montanus Stenocereus pruinosus Stenocereus queretaroensis Stenocereus quevedonis Stenocereus standleyi Stenocereus stellatus Stenocereus thurberi Stenocereus treleasei Stenocereus yunckeri Stephanocereus leucostele Stephanocereus luetzelburgii Stetsonia coryne Strombocactus disciformis

Tacinga braunii Tacinga estevesii Tacinga funalis Tacinga inamoena Tacinga palmadora Tacinga saxatilis Tacinga subcylindrica Tacinga werneri Tacinga x quipa Tephrocactus alexanderi Tephrocactus aoracanthus Tephrocactus articulatus* Tephrocactus geometricus Tephrocactus molinensis Tephrocactus weberi Thelocactus bicolor Thelocactus buekii Thelocactus conothelos Thelocactus garciae Thelocactus hastifer Thelocactus heterochromus Thelocactus hexaedrophorus

Thelocactus lausseri Thelocactus leucacanthus Thelocactus macdowellii Thelocactus rinconensis Thelocactus setispinus Thelocactus tulensis Tunilla albisetacens Tunilla chilensis Tunilla corrugata Tunilla erectoclada Tunilla ianthinantha Tunilla minuscula Tunilla orurensis Tunilla silvestris Tunilla soehrensii Turbinicarpus alonsoi Turbinicarpus beguinii Turbinicarpus bonatzii Turbinicarpus booleanus Turbinicarpus gielsdorfianus Turbinicarpus hoferi Turbinicarpus horripilus

Turbinicarpus jauernigii Turbinicarpus knuthianus Turbinicarpus laui Turbinicarpus lophophoroides Turbinicarpus mandragora Turbinicarpus x mombergii Turbinicarpus pailanus Turbinicarpus pseudomacrochele Turbinicarpus pseudopectinatus Turbinicarpus rioverdensis Turbinicarpus saueri Turbinicarpus schmiedickeanus Turbinicarpus subterraneus Turbinicarpus swobodae Turbinicarpus valdezianus Turbinicarpus viereckii Turbinicarpus ysabelae Turbinicarpus zaragosae Uebelmannia gummifera Uebelmannia pectinifera Uebemannia buiningii Weberbauerocereus churinensis

Weberbauerocereus cuzcoensis Weberbauerocereus johnsonii Weberbauerocereus longicomus Weberbauerocereus rauhii Weberbauerocereus torataensis Weberbauerocereus weberbaueri Weberbauerocereus winterianus Weberocereus biolleyi Weberocereus bradei Weberocereus frohningiorum Weberocereus glaber Weberocereus imitans Weberocereus panamensis Weberocereus rosei Weberocereus tonduzii Weberocereus trichophorus Weberocereus tunilla Yavia cryptocarpa Yungasocereus inquis

Supporting information. File 4. Examples of cacti and Succulents Journals

North America

The cactus and succulent journal. California.

Saguaroland Bulletin. Phoenix.

Cactaceas y Suculentas Mexicanas. Mexico, D. F.

Europe

Cactus. France.

Cactusvrieden. Belgium.

Succulenta. Netherlands

Stackelpost. Germany.

Sukkulentenkunde. Switzerland.

The Cactus Explorers Club Journal. UK

CACTUS-ADVENTURES. Spain.

Cactus & Co. Italy.

Australia and New Zealand

The Spine. Victoria.

Cactus and Succulent Journal. New South Wales.

New Zealand Cactus and Succulent Journal. Auckland.

Japan

Succulentarum Japonica. Tokyo.

Journal of the Cactus and Succulent Society of Japan. Kyoto.

The Study of Cactus. Kanagawaken.

Shaboten-sha. Kanagawa.

Supporting information. File 5. Examples of cactus and succulent societies. The date of foundation of each society is shown.

Australia

The Cactus and Succulent Society of Australia Inc (CSSA). It is the oldest Cactus and Succulent Society

in Australia, established in Melbourne in 1927.

The Cactus and Succulent Society of New South Wales Inc. 1957.

Cactus and Succulent Society of Queensland Inc. 1963.

Cactus and Succulent Society of South Australia. 1964.

Canada

Toronto Cactus and Succulent Club. 1977

Victoria Cactus & Succulent Society. 1992.

Europe

Kakteenfreunden Berliner. 1893.

Magyar Kaktuszgyűjtők Országos Egyesülete. 1971

Oslo sukkulentforening. 1980.

The Dublin and District Cactus and Succulent Society. 1992

Société Succulentophile Francilienne. 1994.

ASAC, Asociación dels amics dels cactus. 1996

Cactus & Co. International Society. 1996

India

The Indian Society of Cactus & Succulents (ISOCS). 1984.

Japan

Kagawa Cactus Club. 1965.

Mexico

La Sociedad Mexicana de Cactologia. 1951.

The Cactus and Succulent Society of New Mexico (CSSNM). 1955.

Sociedad de Cactáceas y Suculentas del Estado de Nuevo León. 2001.

New Zealand

Cactus & Succulent Society of New Zealand Inc. 1947.

Philippines

Cactus and Succulent Society of the Philippines. 1994.

South Africa

Succulent Society of South Africa. 1963.

South America and the Caribbaen

Sociedad Peruana de Cactus y Suculentas (SPECS). 1987.

Sociedad Latinoamericana y del Caribe de Cactáceas y otras Suculentas. 1989.

United States

The Cactus and Succulent Society of America (CSSA). 1929

The Cactus & Succulent Society of Hawaii. 1965.

The Cactus and Succulent Society of Southern Nevada (CSSSN). 1976.

Cactus and Succulent Society of Massachusetts. 1991.

Documents

Review - Invasive Species cactus speci… · de cacta´ceas de Norte y Centro Ame´rica, Herbario Virtual da Flora e dos Fungos, Intermountain Regional Herbar-ium Network, Global