Review

Introduced and invasive cactus species: a global review

Ana 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 Africa 2 Centre for Invasion Biology, Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa 3 Invasive Species Programme, South African National Biodiversity Institute, Kirstenbosch Research Centre, Claremont 7735, South Africa Downloaded from 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 http://aobpla.oxfordjournals.org/

Abstract. Understanding which species are introduced and become invasive, and why, are central questions in invasion science. Comparative studies on model taxa have provided important insights, but much more needs to be done to unravel the context dependencies of these findings. The cactus family (Cactaceae), one of the most popular horticultural plant groups, is an interesting case study. Hundreds of cactus species have been introduced outside their native 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 current global distributions and modelled the potential global distributions based on distribution data of known invasive at University of Stellenosch on February 6, 2015 taxa. Finally, we identified whether invasiveness is phylogenetically clustered for cacti and whether particular traits are correlated with invasiveness. Only 57 of the 1922 cactus species recognized in this treatment have been recorded as invasive. There are three invasion hotspots: South Africa (35 invasive species recorded), Australia (26 species) and Spain (24 species). However, there are large areas of the world with climates suitable for cacti that are at risk of future invasion—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 tend to 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 for proactive management and risk assessments.

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

2009). Many of these introduced organisms have notable Introduction benefits to humans, but some have undesirable impacts The increased movement of humans around the world that can result in substantial monetary costs and/or has facilitated the intentional and accidental transporta- alterations to entire ecosystems and social systems tion of species far from their native ranges, often in a (McNeely 2006; Kumschick et al. 2012). Government manner that can facilitate invasions (Wilson et al. departments, 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 and among the most important alien species worldwide scientists are all facing escalating pressure to address (Weber 2003). and resolve a diversity of problems posed by invasive Studies of cactus invasions have shed light on crucial alien species (Hulme 2006). aspects of plant invasion ecology, e.g. the interaction Much work has recently focussed on reviewing the inva- of invasive plants with seed dispersers (Foxcroft and sive performance of particular genera or closely related Rejma´nek 2007; Padro´n et al. 2011), the role of propagule groups of organisms in different situations around the pressure in driving invasions (Foxcroft et al. 2004) and the world (e.g. Richardson et al. 2011; Moodley et al. 2013; role of herbivores in regulating some plant populations, Potgieter et al. 2013; Shackleton et al. 2014). Such studies with particularly striking examples from classical biologic- aim to update knowledge on the global occurrence and al control (Zimmermann et al. 2004; Paterson et al. 2011). potential range of these taxa and to understand the com- These studies have tended to focus on the genera Opuntia plex drivers of human-mediated introductions and inva- and Cylindropuntia, which contain most of the widely

sions. The findings of such studies are important for introduced, cultivated and invasive species in the family. Downloaded from developing protocols for preventing risky species introduc- However, hundreds of new cactus species are now being tions and for managing species that may become or have introduced all over the world, and many of them are already become invasive (Simberloff et al. 2009). However, becoming naturalized or invasive. For example, Cereus more comparative studies are needed to improve our hexagonus was included in a national list of regulated

understanding of the full suite of interacting factors that invasive species for the first time in South Africa in 2014. http://aobpla.oxfordjournals.org/ influence invasions and to unravel the context dependen- The current global distribution of the cactus family is cies of insights that emerge from particular studies being radically changed by humans, and no attempt (Kueffer et al. 2013). It is important to consider whether has been made to assess the status of each species in such comparisons yield broad generalities or whether terms of invasion or risk thereof. Consequently, a broad generalizations apply only to a subset of taxa. global assessment of the determinants of invasiveness The cactus family (Cactaceae; ‘cacti’) is an interesting of the family Cactaceae is an important requirement for case study. Cacti are a conspicuous component of the the formulation of control strategies. Moreover, reviewing arid regions of the New World and represent one of the the invasive performance of this family around the world at University of Stellenosch on February 6, 2015 world’s most spectacular desert radiations (Edwards may uncover new patterns, processes and invasion risks et al. 2005). The family is distributed from southern not seen in better-studied model groups. Patagonia in Argentina and Chile to Alberta and British This paper aims specifically to (i) compile a list of spe- Columbia in Canada (Edwards et al. 2005), with the only cies in the family Cactaceae, (ii) determine their current exception being Rhipsalis baccifera (mistletoe cactus), native and invasive ranges, and (iii) determine the poten- which is thought to have originated in tropical Americas, tial future ranges of invasive taxa. Using these lists we but was apparently dispersed across the Atlantic Ocean aim to answer the questions: (iv) how have cactus species by birds, reaching southern Africa, Madagascar and Sri been used inside and outside their native range?; and (vi) Lanka (Rebman and Pinkava 2001). are any traits correlated with invasiveness in the family? Cacti are among the first plants that were brought back from the Americas by European explorers in the 15th cen- Methods tury (Howard and Touw 1981) and soon became common in European collections and gardens (Anderson 2001). Defining a cactus The trade in horticultural cacti has developed over the Most taxa in the family Cactaceae are succulents with years into a substantial industry and is responsible for large, leafless, long-living, fleshy stems of different shapes the intercontinental spread of many species (Walters and sizes that often contain clusters of spines which arise et al. 2011). One of the earliest reasons for introduction, from areoles (Benson 1979, 1982; Eggli 1993). Areoles— however, was for use as drought-tolerant crops and for highly specialized axillary or lateral buds or short shoots hedging, with Opuntia ficus-indica (L.) Mill. being by far or branches—are unique to the family (Mauseth 1983; the most utilized (Walters et al. 2011). In an attempt to Gibson and Nobel 1986). However, cacti come in a wide minimize the risks of global climate change, land degrad- range of growth forms (Fig. 1). Succulent plants in other ation and diminishing food security, the Food and Agricul- families are often mistakenly called ‘cacti’ on nursery tural Organization has revived the interest in cactus labels and in popular publications. While it is usually easy cultivation for agricultural purposes in developing coun- to distinguish cacti from other succulents, some taxa look tries (Nefzaoui 2007). As a result of these human- very cactus-like due to convergent evolution, e.g. many mediated introductions, cactus species can be found all species in the genus Euphorbia of the family Euphorbia- over the world, and several members of the family are ceae (Anderson 2001).

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

Species list and invasive status to revise the taxonomy of the Cactaceae (Hunt et al. While the alpha taxonomy of the Cactaceae is well 2006), but concluded that this task is far from complete. known, and the clade is clearly a recent monophyletic ra- In this paper, we base our list on the classification sys- diation, a stabilized nomenclature has not yet been tem developed by the International Cactaceae Systema- achieved (Hunt and Taylor 1986, 1990; Nobel 2002). This tics Group and used by David Hunt in compiling both nomenclature instability can be attributed to inconsisten- editions of the Convention on International Trade in cies in previous taxonomies and to the generally poor re- Endangered Species of Wild Fauna and Flora’s CITES Cac- presentation of cacti in herbarium collections [their taceae Checklist (1992, 1999). This classification has been succulence and spines make them difficult to collect adopted by various sources (Walters 1989; Kubitzki et al. and curate (Walters et al. 2011)]. The International 1993; Anderson 2001). We updated the list to include Organization for Succulent Studies recently attempted 109 additional species, for which, since 2001, either an

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International Union for Conservation of Nature and Species distribution models were produced for each Natural Resources red list assessment has been made species using a simple envelope approach (implemented (http://www.iucnredlist.org/), or a new description has in R) which is equivalent to BIOCLIM’s marginal envelope been published in the scientific literature. None of these (Pearson and Dawson 2003). The following predictor vari- additional 109 species are recorded as invasive. ables were used: maximum temperature of warmest Information on the current distribution and invasive month, minimum temperature of coldest month, precipi- status was gathered from many sources [see Supporting tation of wettest quarter, precipitation of driest quarter, Information]. Each source uses different criteria for cat- precipitation of warmest quarter and precipitation of egorizing alien species. To record the invasive status of coldest quarter. These variables were obtained from the family Cactaceae, our list includes only cactus species WorldClim at a 10-min spatial resolution (Hijmans et al. where there is clear documented evidence of an invasion 2005) and were selected based on their success at pre- as per the definition in Richardson et al. (2000), i.e. plants dicting potential global distributions for other model inva-

spreading over considerable distances from original sive taxa (Richardson et al. 2011). Downloaded from planting or introduction sites. Two sets of models were produced using different ap- proaches. For the first approach, models were calibrated using native range records only. These models were then Potential distribution evaluated using invasive range records, where these were

Invasiveness elsewhere combined with broad-scale available. Sensitivity values (Fielding and Bell 1997) were http://aobpla.oxfordjournals.org/ climatic matching are the most widely used predictors calculated for each model based on the number of inva- of future invasiveness of introduced species (Rejma´nek sive range records that were predicted as present or ab- et al. 2005; Richardson et al. 2011; Petitpierre et al. sent by the model. Sensitivity values range between 2012). Determining which global areas are climatically zero and one, where values close to one indicate low suitable for invasive species establishment may aid future omission error (Fielding and Bell 1997). For the second ap- management efforts and/or preventative measures. We proach, models were calibrated using all available records therefore used bioclimatic models for invasive cacti to for the species (i.e. native and invasive range records) predict potential suitable ranges for individual species without evaluation of sensitivity. For both approaches at University of Stellenosch on February 6, 2015 onaglobalscale.Todothis,wecompiledadatasetof we produced models for species that had five or more na- occurrence records for invasive cactus species from sev- tive range records. Duplicate records per 10-min cell were eral different sources (e.g. Base de datos de ejemplares removed. de cacta´ceas de Norte y Centro Ame´rica, Herbario Virtual Maps of potential species richness were produced by da Flora e dos Fungos, Intermountain Regional Herbar- adding the maps of potential distribution for each of ium Network, Global Biodiversity Information Facility, In- the two approaches. stituto de Biologia de la UNAM, Southwest Environmental Information Network, Oregon Flora Project, San Diego Reasons for introduction and dissemination Natural History museum and CalFlora). For each species, Information on human uses of cacti both in their native the records were plotted on a map and climatic space and introduced ranges were extracted from many sources as defined by values of annual mean temperature and [see Supporting Information]. Five broad human-use annual precipitation extracted from 10-min resolution categories were defined: (i) ornamental (horticulture), WorldClim bioclimatic variables. A thorough data clean- (ii) food or fodder (i.e. for humans or livestock), (iii) medi- ing procedure was followed using the biogeo package cinal, (iv) hedging and (v) other (e.g. furniture or religious). in R (M. P. Robertson et al., in preparation): tests were per- Not all the species with a defined use in the native range formed on obvious outliers to determine whether the x- are introduced to other areas of the world for the same and y-coordinates had accidentally been transposed or reasons. To assess how the number of introduced species whether incorrect signs were used; records that were differs between uses, we compared the proportion of in- plotted in the ocean but directly adjacent to a coastal troduced and non-introduced species in each use cat- grid cell were assigned to that closest terrestrial grid egory with that of species in other use categories (using cell; and low-precision records were removed, e.g. when a Fisher Test in R). only degrees were available for the coordinate. Outliers in the environmental space were queried to identify where they lay in geographical space to identify any Correlates of invasiveness errors missed during the initial data cleaning, and A useful first step to improve our understanding of inva- removed or rectified if a particular cause of the error siveness is to identify the traits correlated with invasive- could be determined. ness (Pysˇek and Richardson 2007). Here, we looked at

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phylogeny, taxonomy (at the genus level), growth form good data on native range size for this genus (Dean and and metrics of native range size. Milton 2000; Erre et al. 2009; Padro´n et al. 2011; Lloyd and For phylogenetic reconstruction we collated genetic Reeves 2014). We compared the latitudinal ranges of in- data for the maturase K (matK) gene region for repre- vasive and non-invasive Opuntia species using a Student’s sentative taxa of all Cactaceae genera with available t-test in R. We also looked at Cactaceae listed on the IUCN data in the GenBank online repository (http://ncbi.nlm. Red List of Threatened Species (Rodrigues et al. 2006). For nih.gov). DNA sequence data were aligned in BioEdit ver- Cactaceae, most of the species considered at risk were so sion 7.0.5.3 (Hall 1999) and manually edited. Because of due to a small native range size (http://www.iucnredlist. differences in sequence lengths for different taxa we org/), and therefore, in this case, the list provides a trimmed flanking regions to avoid excessive missing rough proxy for native range size (as well as giving data. Our final dataset comprised 103 genera within some indication of population trends). Cactaceae. Phylogenetic relationships were estimated using Bayesian search criteria with parameter estimates Results Downloaded from obtained from the program jModelTest v2.1.3 (GTR + I + G; Species list and invasive status Darriba et al. 2012) in MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). For both datasets, MrBayes was The list of cacti assembled for this paper comprises 1922 run for 2 000 000 generations and trees sampled every species from 130 genera [see Supporting Information]. Genera differed widely in species richness, with several 1000 generations. Nodal support for the retrieved tree http://aobpla.oxfordjournals.org/ topology was determined as posterior probabilities in speciose genera [Opuntia (193 species), Echinopsis (133 MrBayes. To determine whether invasiveness within Cac- species) and Mammillaria (171 species)] and 35 monotypic taceae has a phylogenetic signal, we compared trait genera. Of the 1922 species we could definitively classify change (proportion of invasive taxa within a genus) only 57 species as invasive. with a null hypothesis of Brownian motion using Blom- The currently available distribution data are only ad- berg’s K statistic (Blomberg et al. 2003). Similarly, we equate for a country-level analysis (see Fig. 2). These ana- used Pagel’s lambda (l) statistic to determine the extent lyses show Mexico as the main ‘hot spot’ of native cactus to which branch length transformation explains the diversity (Fig. 2A). Three countries had notably more inva- at University of Stellenosch on February 6, 2015 distribution of trait states (proportion of invasive taxa/ sive taxa than the rest—Australia (39), South Africa (35 genus) at the tips of a phylogeny (Pagel 1999). Both species) and Spain (24)—while other countries had at tests are implemented in the function phylosig.R from most 13 (Fig. 2B, Table 1). The most widespread invasive the phytools package (Revell 2012). species is O. ficus-indica (22 different countries), with To assess how invasiveness differs at the genus level, other species invading 15 or fewer countries (Table 1). Un- we compared the number of invasive and non-invasive surprisingly, the origin of most of these invasive species is species in each genus with that in the rest of the family also Mexico (Fig. 2C). using a Fisher test in R. Potential distribution Based on information extracted from all sources, we We examined potential invasive distributions for only 39 also obtained information on the growth form of each of the invasive species [see Supporting Information] species. There are different classifications in the literature as none of the remaining 18 species had enough (i.e. (e.g. Barthlott and Hunt 1993; Anderson 2001; Lo´pez and five or more) records in their native ranges of sufficient Valdivia 2007; Ortega-Baes et al. 2010; Herna´ndez- accuracy. The median number of records per species Herna´ndez et al. 2011). Here, following discussions in modeled was 128 for the native range and 124 for the Anderson (2001) and Herna´ndez-Herna´ndez et al. (2011), invasive range. A large variation in sensitivity values we recognize 12 types: angled, cylindrical, cushion-like, was obtained. We found no significant differences be- leaf-like, flattened-padded, geophytic, globose, ovoid, tween the projected species richness maps for the two sprawling tree-like, tuberculate and prostrate growth modelling approaches, i.e. using native range occurrence forms (Fig. 1). records only or native and invasive occurrence records Available data on native range size are inadequate for (Fig. 3). The main known areas of invasion (Australia, an analysis of the relationship between native range size South Africa and Spain) were indicated as suitable, but and invasive status for the whole family. We were, how- there were also substantial regions that are suitable in ever, able to analyse the relationship between native central Africa, China and south-eastern Asia. range size and invasiveness for the genus Opuntia,be- cause species in the genus have been widely introduced Reasons for introduction and dissemination and disseminated around the world, there are many inva- A quarter of species recorded as being used for ornamen- sive and non-invasive taxa, and there are comparatively 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). Shading indicates 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 Supporting Information]. Downloaded from http://aobpla.oxfordjournals.org/ at University of Stellenosch on February 6, 2015

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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 one use category. Significance levels were determined by comparing the number of introduced vs. number of non-introduced species for any

category to all other taxa using Fisher’s exact test. Confidence intervals were determined for the percentage of introduced or http://aobpla.oxfordjournals.org/ non-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 at University of Stellenosch on February 6, 2015

elsewhere (Table 2). In contrast, only a seventh of the cac- phylogenetic signal for invasiveness (l ¼ 0.991; P , tus species used in their native ranges for food or fodder 0.01). This phylogenetic signal was not related to human have been introduced elsewhere, and we found no official usage, i.e. ornamental trade. records of species having been introduced for medicinal At the genus level, the 57 invasive species belong to just or other purposes (though Lophophora williamsii and 13 of the 130 genera (Fig. 5A). Opuntia, Cylindropuntia, Har- Echinopsis pachanoi have been introduced worldwide risia, Hylocereus and Austrocylindropuntia have a signifi- for their psycho-active uses). As an illustration of the cantly higher proportion of invasive species than other worldwide popularity of cacti as horticultural species, genera, while only Mammillaria has a significantly lower in- we give some examples of international cactus and cidence of invasiveness (it contains no invasive species). succulent journals and societies [see Supporting In terms of the 12 main growth forms we considered Information]. (Fig. 1), all of the invasive species were cylindrical, flattened-padded, sprawling, leaf-like or angled. The Correlates of invasiveness flattened-padded and angled growth forms stand out as Our phylogeny, representing 103 taxa (genera), retrieved having a significantly higher percentage of invasive spe- three main clades that differed substantially in the pro- cies than the other growth forms, while the globose portion of invasive taxa (Fig. 4). Clade 1 included mainly growth form is significantly underrepresented in terms of genera of the tribes of the subfamily Cactoideae (with invasiveness (there are no globose invasive taxa) (Fig. 5B). the exception of the tribe Cacteae), and some invasive The latitudinal ranges of invasive Opuntia species (218, taxa. Clade 2 comprised genera within the tribe Cacteae n ¼ 15) are significantly greater than those of non- (with the exception of Maihueniopsis from the tribe invasive species (8.78, n ¼ 42) (t-test, P , 0.009). As of Tephrocacteae) and does not include any invasive taxa. 2013, the IUCN had assessed 1409 cactus species. All in- Clade 3 (which includes the subfamily Opuntioideae) vasive taxa are in the Least Concern, Near Threatened or contains most of the invasive species. Using proportions Data Deficient categories and none of the known invasive of invasive taxa/genus, Blomberg’s K indicated no signifi- species are among the 303 taxa listed in the categories cant phylogenetic signal of invasiveness (K ¼ 0.260; P ¼ Critically Endangered (CR), Endangered (EN) or Vulnerable 0.187), whereas Pagel’s l indicated a significant (VU) (i.e. species that likely have smaller ranges, Fig. 5C).

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This provides preliminary evidence for a relationship be- Potential for further invasions tween native range size and invasiveness. Currently, most cactus invasions are recorded in Australia, South Africa and Spain. Unsurprisingly, the results of the Discussion broad-scale climatic matching identified these three countriesasbeingbioclimaticallyequivalenttoareas The results of this study allowed us to draw generaliza- within the range of a large number of cactus species. tions that are useful for managing future introductions A strong climatic match between native and recipient and invasions. Despite the extensive dissemination of ranges is recognized as a fundamental requirement for cactus species around the world, only a small proportion the success of introduced plants (Richardson et al. 2011; of the family is currently known to be invasive. However, Richardson and Pysˇek 2012). However, many other areas these invasive taxa have very large potential ranges glo- of the world where these species are currently absent are bally, including in areas where no cactus invasions have also highlighted as potentially suitable for invasion. This

yet been recorded. Cacti are introduced to new areas pattern probably reflects differences in introduction ef- Downloaded from mainly for ornamentation, but the selection of ornamen- fort, and suggests a substantial invasion debt (sensu tal species is not based on attributes that favour invasive- Essl et al. 2011) in agreement with the findings for ness. Invasive taxa are overrepresented in several genera, other model groups (e.g. the genera Casuarina and Proso- phylogenetic clades and growth forms. Species that are pis; Potgieter et al. 2013; Shackleton et al. 2014). Clearly, of conservation concern in their native ranges have not the natural experiment of plantings of cacti outside their http://aobpla.oxfordjournals.org/ become invasive. natural ranges is far from complete. Because efforts directed at prevention of new introduc- Proportion of invasive taxa in the family Cactaceae tions are the most cost-efficient component of invasive The main centres of cactus diversity are north-eastern species control strategies (Leung et al. 2002), our results Mexico, the eastern Andes of Bolivia and Argentina emphasize the importance of controlling the introduction and south-eastern Brazil (Mutke and Barthlott 2005). of cacti recorded as invasive in Australia, South Africa and However, species are distributed throughout a large var- Spain to other areas suitable for invasion.

iety of habitats, including hot deserts, sandy coastal at University of Stellenosch on February 6, 2015 stretches, scrublands, dry deciduous forests, high alpine Reasons for introduction and dissemination steppes and even tropical rain forests (Barthlott and Among the many uses of cacti, the main reason for Hunt 1993). Therefore, there are cactus species that are introductions of species to regions outside their native climatically suited to almost all habitats on Earth. How- ranges is the horticulture trade. Cultivation of ornamental ever, only 3 % of the species in the family are currently cactus species is very popular in temperate regions. There clearly invasive. It is difficult to say whether this reflects are more than 20 cactus and succulent journals and hun- the real extent of invasions or whether the pattern is dreds of societies around the world, as well as hundreds affected by different levels of reporting and the availabil- of cactus and succulent Facebook pages and groups. ity of accurate data, moreover, not all cactus species have Moreover, global introductions of new species are likely been afforded the same opportunities to become inva- to occur: just 23 % of the species considered to have or- sive. Nonetheless, this pattern has also been observed namental value in the native range have been introduced in other model groups. For example, only between 0.5 to other regions. and 0.7 % of the global pool of tree and shrub species The use of alien plants for ornamentation is an import- are currently clearly invasive outside their natural range ant driver of introductions and dissemination in many (Richardson and Rejma´nek 2011). plant groups, and several attributes associated with

Figure 4. Bayesian phylogeny based on matK DNA sequence data illustrating phylogenetic relationships among genera within Cactaceae. The aligned matK matrix contained 1354 characters and required 65 gaps (indels), ranging from 1 to 74 characters in size. Overall, the phylogeny yielded well-resolved relationships among all genera included. High nodal support (posterior probabilities .0.9) is indicated at nodes by yellow boxes. Genera with invasive taxa are indicated as red branches where red circles are proportional to the percentage of invasive taxa within each genus. 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 of non-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. Confidence intervals 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 species issimilartothatseeninconifers.Twenty-eightof for horticulture) are also important for invasiveness. For the known invasive conifer taxa belong to one family example, trees used for ornamentation are often selected (Pinaceae) and 21 of these are in a single genus—Pinus for their long-lasting displays of brightly coloured fleshy (Richardson and Rejma´nek 2004). fruits that are attractive to a wide range of generalist OnenoticeablefeatureofCactaceaeistherangeof seed dispersers (Richardson and Rejma´nek 2011). As growth forms within the family. Unlike most plant groups another example, Australian Acacia species used for studied to date, invasiveness in cacti is strongly asso- ornamentation have rapid growth rates and can survive ciated with particular growth forms. All invasive cacti and flourish in nutrient-poor, arid or degraded sites are angled, cylindrical, flattened-padded or sprawling. (Richardson et al. 2011; Donaldson et al. 2014). Ornamen- The reason for high levels of invasiveness in these growth tal cacti, on the other hand, appear to be selected for fea- forms probably relates to the strong ability of taxa in tures other than those that directly enhance invasiveness; these groups to grow vegetatively from cuttings which

in particular, species that survive without much input and can allow for rapid dispersal (Anderson 2001). Downloaded from grow slowly are favoured (i.e. more K-selected than It would seem that the same traits that allow some cac- r-selected). The most popular cactus species in the global tus species to become widespread in their native ranges ornamental trade belong to the genus Mammillaria contribute to their ability to overcome abiotic filters and (Novoa et al., unpubl. data); these species are valued for successfully establish in new regions. No cactus species

their globose growth form more than any other feature. that are of conservation concern in their native ranges http://aobpla.oxfordjournals.org/ As no Mammillaria spp. are invasive and no globose have been recorded as invasive, and there is a strong cor- taxa are invasive (Fig. 5A and B), it is likely that this relation between invasiveness and native range size in, for genus/growth form poses little risk of invasion or impact example, the genus Opuntia. A similar pattern has been due to its ecological strategy. observed for other model groups. For example, Australian Acacia species with large native ranges and low percola- Correlates of invasiveness tion exponents (i.e. high population increase rate) are Besides the past and current efforts directed at prevent- most likely to be introduced and become naturalized (Hui ing new introductions of species already known as inva- et al. 2011). Large native range size has been shown to be a at University of Stellenosch on February 6, 2015 sive elsewhere, additional protocols for regulating risk good predictor of invasiveness and invasion success in are needed. This is because most contemporary introduc- many, but not all, plant groups (e.g. Proches¸ et al. 2012; tions and dissemination of cacti are of ornamental taxa, Moodley et al. 2013; Potgieter et al. 2013). many of which do not have well-documented introduc- tion/invasion histories. Our results suggest that delimita- Conclusions tions based on membership to genera, position in the Cacti are already among the most widespread and dam- phylogeny, growth form and native range size need to aging of invasive alien plants in some parts of the world. be considered to produce objective and defendable The huge and growing interest in many cacti for ornamen- approaches for formal risk assessments. tation has created an important new pathway for the Primary attention with regard to invasiveness in cacti introduction and dissemination of a growing number of needs to be given to taxa in the 13 genera of Cactaceae cactus taxa around the world. Many new invasion events that consistently display invasive tendencies. These gen- are expected in the future. There is clearly a need to regu- era (comprising 538 species) share certain characteristics late the movement of cacti recorded as invasive elsewhere which include prolific fruiting, strong vegetative reproduc- (currently only 3 % of the species in the family) to areas tion and effective dispersal mechanisms (Walters et al. suitable for invasion, as well as taxa that pose a high risk 2011). This pattern is particularly seen in the ‘opuntoid of becoming invasive. Results from this study suggest that cacti’ (i.e. the genera Austrocylindropuntia, Cylindropuntia risk assessment protocols for cacti should evaluate taxa and Opuntia),whichhavebeenclassedtogetheras according to genera, position in the phylogeny of the fam- Weeds of National Significance in Australia (Lloyd and ily, growth form, and, potentially, native range size. Reeves 2014). Our phylogenetic analysis (Fig. 4) provides support for this approach. Invasive taxa are relatively common in the Opuntioideae clade, whereas the inci- Sources of Funding dence of invasiveness in the tribe Cacteae is zero. Inter- Funding for this work was provided by the Working estingly, genera from these clades are well represented for Water (WfW) Programme of the South African in the global horticultural trade, and presumably have Department of Environmental Affairs and the DST-NRF similar levels of dissemination and introduction effort Centre of Excellence for Invasion Biology (C.I.B) as part (Novoa et al., unpubl. data). This pattern in Cactaceae of the C.I.B/WfW collaborative research programme on

12 AoB PLANTS www.aobplants.oxfordjournals.org & The Authors 2014 Novoa et al. — Cacti as alien species

‘Research for Integrated Management of Invasive Alien Edwards EJ, Nyffeler R, Donoghue MJ. 2005. Basal cactus phylogeny: Species’. D.M.R. acknowledges additional support from implications of Pereskia (Cactaceae) paraphyly for the the National Research Foundation (grant 85417) and transition to the cactus life form. American Journal of Botany 92: 1177–1188. the Oppenheimer Memorial Trust. Eggli U. 1993. Glossary of botanical terms with special reference to Contributions by the Authors succulent plants. Richmond: British Cactus and Succulent Society. Erre P, Chessa I, Nieddu G, Jones PG. 2009. Diversity and spatial dis- A.N., J.R.U.W. and D.M.R. conceived the idea. A.N. collected tribution of Opuntia spp. in the Mediterranean Basin. Journal of the data. A.N. and J.R.U.W. ran the statistics, J.J.L.R. built Arid Environments 73:1058–1066. the phylogeny and M.P.R. undertook climate matching. EsslF,DullingerS,RabitschW,HulmePE,Hu¨lber K, Jarosˇ´ıkV, ¨ ` A.N. led the writing with assistance from others. Kleinbauer I, Krausmann F, Kuhn I, Nentwig W, Vila M, Genovesi P, Gherardi F, Desprez-Loustau M, Roques A, Pysˇek P. 2011. Socioeconomic legacy yields an invasion debt. Proceedings Conflicts of Interest Statement of the National Academy of Sciences of the USA 108:203–207. Fielding AH, Bell JF. 1997. A review of methods for the assessment of Downloaded from None declared. prediction errors in conservation presence/absence models. Environmental Conservation 24:38–49. Acknowledgements Foxcroft LC, Rejma´nek M. 2007. What helps Opuntia stricta invade Kruger National Park, South Africa: baboons or elephants? Comments from two anonymous reviewers improved this

Applied Vegetation Science 10:265–270. http://aobpla.oxfordjournals.org/ manuscript. The authors also thank Rodrigo Pintos who Foxcroft LC, Rouget M, Richardson DM, Mac Fadyen S. 2004. helped with the design of Fig. 1. Reconstructing 50 years of Opuntia stricta invasion in the Kruger National Park, South Africa: environmental determi- Supporting Information nants and propagule pressure. Diversity and Distributions 10: 427–437. The following Supporting Information is available in the Gibson AC, Nobel PS. 1986. The cactus primer. Cambridge: H. U. online version of this article – Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment Table S1. Examples of sources of information on cactus editor and analysis program for Windows 95/98/NT. Nucleic Acids species. Symposium Series 41:95–98. at University of Stellenosch on February 6, 2015 Table S2. List of cactus species. *57 species recorded as Herna´ndez-Herna´ndez T, Herna´ndez HM, De-Nova JA, Puente R, Eguiarte LE, Magallo´n S. 2011. Phylogenetic relationships and invasive outside their native range. evolution of growth form in Cactaceae (Caryophyllales, Eudicoty- Figure S3. Potential species richness based on available ledoneae). American Journal of Botany 98:44–61. native records only of 39 invasive cactus species. Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A. 2005. Very high Table S4. Examples of cacti and succulents Journals. resolution interpolated climate surfaces for global land areas. Table S5. The date of foundation of each society is International Journal of Climatology 25:1965–1978. shown. Howard RA, Touw M. 1981. The cacti of the Lesser Antilles and the typification of the genus Opuntia Miller. Cactus and Succulent Journal 53:233–237. Literature Cited HuiC,RichardsonDM,RobertsonMP,WilsonJRU,YatesCJ.2011. Anderson EF. 2001. The cactus family. Portland: Timber Press. Macroecology meets invasion ecology: linking the native distribu- tions of Australian Acacias to invasiveness. Diversity and Distribu- Barthlott W, Hunt DR. 1993. Cactaceae. In: Kubitzki K, ed. The families tions 17:872–883. and genera of vascular plants Vol. II. Berlin: Springer, 161–197. Hulme PE. 2006. Beyond control: wider implications for the manage- Benson L. 1979. Plant classification. MA: D.C. Heath. ment of biological invasions. Journal of Applied Ecology 43: Benson L. 1982. The cacti of the United States and Canada. Stanford: 835–847. Stanford University Press. Hunt DR. 1992. CITES Cactaceae checklist. Richmond: Royal Botanic Blomberg SP, Garland T, Ives A. 2003. Testing for phylogenetic signal Gardens. in comparative data: behavioral traits are more labile. Evolution Hunt DR. 1999. CITES Cactaceae checklist, 2nd edn. Richmond: Royal 57:717–745. Botanic Gardens. Darriba D, Taboada GL, Doallo R, Posada D. 2012. jModelTest 2: more Hunt DR, Taylor NP. 1986. The genera of Cactaceae: towards a new models, new heuristics and parallel computing. Nature Methods consensus. Bradleya 4:65–78. 9:772–772. Hunt DR, Taylor NP. 1990. The genera of Cactaceae: progress towards Dean WR, Milton S. 2000. Directed dispersal of Opuntia species in the consensus. Bradleya 8:85–107. Karoo, South Africa: are crows the responsible agents? Journal of Arid Environments 45:305–314. Hunt DR, Taylor NP, Charles G. 2006. New cactus lexicon.Milborne Port: Dh books. Donaldson JE, Hui C, Richardson DM, Wilson JRU, Robertson MP, Webber BL. 2014. Invasion trajectory of alien trees: the role of Kubitzki K, Rohwer JG, Bittrich V. 1993. The families and genera of introduction pathway and planting history. Global Change Biol- vascular plants. Flowering plants: dicotyledons. magnoliid, hama- ogy 20:1527–1537. melid and caryophyllid families. Berlin: Springer.

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14 AoB PLANTS www.aobplants.oxfordjournals.org & The Authors 2014 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, South Africa 2014. Government Gazette of South Africa, vol. 559, issue 37886, pp. 3–80. Pretoria. 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 Australia Opuntia spp.) in Western Australia. Department of Agriculture and Food. Goverment of Western Australia Book Walters M, Figueiredo E, Crouch NR, Winter PJD, Smith GF, Zimmermann HG, Mashope BK. 2011. Naturalised and South Africa invasive succulents of southern Africa (Y Samyn, D VandenSpiegel, and J Degreef, Eds.). Abc Taxa 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 Argentina forms related to germination responses to light? A test using Echinopsis species. Acta Oecologica 36: 339–342 Peer-review article Essl F, Kobler J. 2009. Spiny invaders – Patterns and determinants of cacti invasion in Europe. Flora - Morphology, Europe Distribution, Functional Ecology of Plants 204: 485–494 Peer-review article Dean WR., Milton S. 2000. Directed dispersal of Opuntia species in the Karoo, South Africa: are crows the responsible South Africa agents? Journal of Arid Environments 45: 305–314 Peer-review article Vilà M, Gimeno I. 2003. Seed predation of two alien Opuntia species invading Mediterranean communities. Plant Spain Ecology 167: 1–8. 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

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

Acanthocalycium ferrarii Armatocereus rupicola Austrocylindropuntia shaferi Cephalocereus senilis Acanthocalycium klimpelianum Arrojadoa albiflora Austrocylindropuntia subulata* Cephalocereus totolapensis Acanthocalycium spiniflorum Arrojadoa bahiensis Austrocylindropuntia verschaffeltii Cephalocleistocactus chrysocephalus Acanthocereus baxaniensis Arrojadoa beateae Austrocylindropuntia vestita Cereus adelmarii Acanthocereus colombianus Arrojadoa dinae Austrocylindropuntia yanganucensis Cereus aethiops Acanthocereus horridus Arrojadoa eriocaulis Aztekium hintonii Cereus albicaulis Acanthocereus occidentalis Arrojadoa multiflora Aztekium ritteri Cereus argentinensis Acanthocereus subinermis Arrojadoa penicillata Bergerocactus emoryi Cereus bicolor Acanthocereus tetragonus* Arrojadoa rhodantha Blossfeldia liliputana Cereus braunii Acharagma aguirreana Arthrocereus glaziovii Brachycereus nesioticus Cereus cochabambensis Acharagma roseana Arthrocereus melanurus Brasilicereus estevesii Cereus comarapanus Ariocarpus agavoides Arthrocereus rondonianus Brasilicereus markgrafii Cereus estevesii Ariocarpus bravoanus Arthrocereus spinosissimus Brasilicereus phaeacanthus Cereus fernambucensis Ariocarpus fissuratus Astrophytum asterias Brasiliopuntia brasiliensis Cereus fricii Ariocarpus kotchebeyanus Astrophytum capricorne Browningia albiceps Cereus haageanus Ariocarpus retusus Astrophytum caput-medusae Browningia altissima Cereus hankeanus Ariocarpus scaphirostris Astrophytum coahuilense Browningia amstutziae Cereus hexagonus* Ariocarpus trigonus Astrophytum myriostigma Browningia caineana Cereus hildmannianus* Armatocereus arduus Astrophytum ornatum Browningia candelaris Cereus horrispinus Armatocereus brevispinus Austrocactus bertinii Browningia chlorocarpa Cereus huilunchu Armatocereus cartwrightianus Austrocactus coxii Browningia columnaris Cereus insularis Armatocereus godingianus Austrocactus patagonicus Browningia hertlingiana Cereus jamacaru* Armatocereus humilis Austrocactus philippii Browningia microsperma Cereus kroenleinii Armatocereus laetus Austrocactus spiniflorus Browningia pilleifera Cereus lamprospermus Armatocereus mataranus Austrocylindropuntia cylindrica* Browningia viridis Cereus lanosus Armatocereus matucanensis Austrocylindropuntia floccosa Calymmanthium substerile Cereus mirabella Armatocereus oligogonus Austrocylindropuntia hirschii Carnegiea gigantea Cereus mortensenii Armatocereus procerus Austrocylindropuntia lagopus Cephalocereus apicephalium Cereus pachyrhizus Armatocereus rauhii Austrocylindropuntia pachypus Cephalocereus columna-trajani Cereus phatnospermus Armatocereus riomajensis Austrocylindropuntia punta-caillan Cephalocereus nizandensis Cereus pierre-braunianus Cereus repandus Cleistocactus neoroezlii Coleocephalocereus goebelianus Copiapoa taltalensis Cereus roseiflorus Cleistocactus orthogonus Coleocephalocereus pluricostatus Copiapoa tenuissima Cereus saddianus Cleistocactus pachycladus Coleocephalocereus purpureus Copiapoa tocopillana Cereus spegazzinii Cleistocactus palhuayensis Coleocephalocereus uebelmanniorum Corryocactus apiciflorus Cereus stenogonus Cleistocactus paraguariensis Consolea corallicola Corryocactus aureus Cereus tacuaralensis Cleistocactus parapetiensis Consolea falcata Corryocactus ayacuchoensis Cereus trigonodendron Cleistocactus parviflorus Consolea macracantha Corryocactus brachypetalus Cereus validus Cleistocactus peculiaris Consolea millspaughii Corryocactus brevistylus Cereus vargasianus Cleistocactus piraymirensis Consolea moniliformis Corryocactus chachapoyensis Cintia knizei Cleistocactus plagiostoma Consolea nashii Corryocactus charazanensis Cipocereus bradei Cleistocactus pungens Consolea picardae Corryocactus erectus Cipocereus crassisepalus Cleistocactus reae Consolea rubescens Corryocactus huincoensis Cipocereus laniflorus Cleistocactus ritteri Consolea spinosissima Corryocactus melanotrichus Cipocereus minensis Cleistocactus roezlii Copiapoa ahremephiana Corryocactus pulquiensis Cipocereus pusilliflorus Cleistocactus samaipatanus Copiapoa angustiflora Corryocactus quadrangularis Cleistocactus acanthurus Cleistocactus sepium Copiapoa atacamensis Corryocactus squarrosus Cleistocactus baumannii Cleistocactus serpens Copiapoa bridgesii Corynopuntia aggeria Cleistocactus brookeae Cleistocactus sextonianus Copiapoa calderana Coryphanta compacta Cleistocactus buchtienii Cleistocactus smaragdiflorus Copiapoa cinerascens Coryphantha calipensis Cleistocactus candelilla Cleistocactus strausii Copiapoa cinerea Coryphantha calochlora Cleistocactus chotaensis Cleistocactus sulcifer Copiapoa conglomerata Coryphantha clavata Cleistocactus clavispinus Cleistocactus tarijensis Copiapoa coquimbana Coryphantha cornifera Cleistocactus x crassiserpens Cleistocactus tenuiserpens Copiapoa decorticans Coryphantha cornuta Cleistocactus dependens Cleistocactus tominensis Copiapoa echinoides Coryphantha delaetiana Cleistocactus ferrarii Cleistocactus tupizensis Copiapoa esmeraldana Coryphantha difficilis Cleistocactus fieldianus Cleistocactus varispinus Copiapoa fieldleriana Coryphantha duragensis Cleistocactus grossei Cleistocactus viridiflorus Copiapoa grandiflora Coryphantha echinoidea Cleistocactus hildegardiae Cleistocactus vulpis-cauda Copiapoa haseltoniana Coryphantha echinus Cleistocactus hoffmannii Cleistocactus winteri Copiapoa humilis Coryphantha elephantidens Cleistocactus hyalacanthus Cleistocactus xylorhizus Copiapoa hypogaea Coryphantha erecta Cleistocactus hystrix Cochemiea halei Copiapoa krainziana Coryphantha georgii Cleistocactus icosagonus Cochemiea maritima Copiapoa laui Coryphantha glanduligera Cleistocactus jajoanus Cochemiea pondii Copiapoa longistaminea Coryphantha glassii Cleistocactus laniceps Cochemiea poselgeri Copiapoa malletiana Coryphantha gracilis Cleistocactus longiserpens Cochemiea setispina Copiapoa marginata Coryphantha grata Cleistocactus luribayensis Coleocephalocereus aureus Copiapoa megarhiza Coryphantha guerkeana Cleistocactus micropetalus Coleocephalocereus braunii Copiapoa mollicula Coryphantha hintoniorum Cleistocactus morawetzianus Coleocephalocereus buxbaumianus Copiapoa serpentisulcata Coryphantha indensis Cleistocactus muyurinensis Coleocephalocereus fluminensis Copiapoa solaris Coryphantha jalpanensis Coryphantha kracikii Cumulopuntia crassicylindrica Cylindropuntia munzii Disocactus amazonicus Coryphantha longicornis Cumulopuntia dactylifera Cylindropuntia pallida Disocactus aurantiacus Coryphantha macromeris Cumulopuntia frigida Cylindropuntia prolifera Disocactus biformis Coryphantha maiz-tablasensis Cumulopuntia fulvicoma Cylindropuntia ramosissima Disocactus cinnabarinus Coryphantha maliterrarum Cumulopuntia galerasensis Cylindropuntia rosea* Disocactus eichlamii Coryphantha melleospina Cumulopuntia hystrix Cylindropuntia sanfelipensis Disocactus flagelliformis Coryphantha neglecta Cumulopuntia ignescens Cylindropuntia santamaria Disocactus kimnachii Coryphantha nickelsiae Cumulopuntia mistiensis Cylindropuntia spinosior* Disocactus macdougallii Coryphantha octacantha Cumulopuntia pentlandii Cylindropuntia tesajo Disocactus macranthus Coryphantha odorata Cumulopuntia pyrrhacantha Cylindropuntia thurberi Disocactus martianus Coryphantha ottonis Cumulopuntia rossiana Cylindropuntia tunicata* Disocactus nelsonii Coryphantha pallida Cumulopuntia sphaerica Cylindropuntia versicolor Disocactus phyllanthoides Coryphantha poselgeriana Cumulopuntia ticnamarensis Cylindropuntia whipplei Disocactus quezaltecus Coryphantha potosiana Cumulopuntia tortispina Cylindropuntia wolfii Disocactus schrankii Coryphantha pseudoechinus Cumulopuntia tumida Cylindropuntia x congesta Disocactus speciosus Coryphantha pseudonickelsiae Cumulopuntia unguispina Cylindropuntia x deserta Echinocactus grusonii Coryphantha pseudoradians Cumulopuntia zehnderi Cylindropuntia x fosbergii Echinocactus horizonthalonius Coryphantha pulleineana Cylindropuntia abyssi Cylindropuntia x kelvinensis Echinocactus parryi Coryphantha pusilliflora Cylindropuntia acanthocarpa Cylindropuntia x neoarbuscula Echinocactus platyacanthus Coryphantha pycnacantha Cylindropuntia alcahes* Cylindropuntia x tetracantha Echinocactus polycephalus Coryphantha radians Cylindropuntia anteojoensis Cylindropuntia x viridiflora Echinocactus texensis Coryphantha ramillosa Cylindropuntia arbuscula Cylindropuntia x vivipara Echinocereus adustus Coryphantha recurvata Cylindropuntia bigelovii Dendrodendus nudiflorus Echinocereus apachensis Coryphantha reduncispina Cylindropuntia californica Dendrodendus undulosus Echinocereus barthelowanus Coryphantha retusa Cylindropuntia calmalliana Denmoza rhodacantha Echinocereus berlandieri Coryphantha robustispina Cylindropuntia caribaea Discocactus bahiensis Echinocereus bonkerae Coryphantha salinensis Cylindropuntia cedrosensis Discocactus cangaensis Echinocereus boyce-thompsonii Coryphantha sulcata Cylindropuntia cholla Discocactus catingicola Echinocereus brandegeei Coryphantha sulcolanata Cylindropuntia davisii Discocactus cephaliaciculosus Echinocereus bristolii Coryphantha tripugionacantha Cylindropuntia echinocarpa Discocactus diersianus Echinocereus chisoensis Coryphantha unicornis Cylindropuntia fulgida* Discocactus ferricola Echinocereus cinerascens Coryphantha vaupeliana Cylindropuntia ganderi Discocactus hartmannii Echinocereus coccineus Coryphantha vogtherriana Cylindropuntia hystrix Discocactus heptacanthus Echinocereus dasyacanthus Coryphantha werdermannii Cylindropuntia imbricata* Discocactus horstii Echinocereus engelmannii Coryphantha wohlschlageri Cylindropuntia kleiniae* Discocactus placentiformis Echinocereus enneacanthus Cumarinia odorata Cylindropuntia leptocaulis* Discocactus pseudoinsignis Echinocereus fasciculatus Cumulopuntia boliviana Cylindropuntia lindsayi Discocactus subterraneo-proliferans Echinocereus fendleri Cumulopuntia chichensis Cylindropuntia molesta Discocactus zehntneri Echinocereus ferreirianus Cumulopuntia corotilla Cylindropuntia multigeniculata Disocactus ackermannii Echinocereus freudenbergeri Echinocereus grandis Echinocereus viereckii Echinopsis chamaecereus* Echinopsis lageniformis Echinocereus klapperi Echinocereus viridiflorus Echinopsis chiloensis Echinopsis lamprochlora Echinocereus knippelianus Echinocereus websterianus Echinopsis chrysantha Echinopsis lateritia Echinocereus laui Echinocereus x roetteri Echinopsis chrysochete Echinopsis leucantha Echinocereus ledingii Echinomastus erectocentrus Echinopsis cinnabarina Echinopsis litoralis Echinocereus leucanthus Echinomastus gautii Echinopsis clavata Echinopsis macrogona Echinocereus longisetus Echinomastus intertextus Echinopsis cochabambensis Echinopsis mamillosa Echinocereus mapimiensis Echinomastus johnsonii Echinopsis comarapana Echinopsis marsoneri Echinocereus maritimus Echinomastus mariposensis Echinopsis conaconensis Echinopsis mataranensis Echinocereus mojavensis Echinomastus unguispinus Echinopsis coquimbana Echinopsis maximiliana Echinocereus nicholii Echinomastus warnockii Echinopsis coronata Echinopsis meyeri Echinocereus nivosus Echinopsis adolfofriedrichii Echinopsis cotacajesii Echinopsis mieckleyi Echinocereus ortegae Echinopsis ancistrophora Echinopsis crassicaulis Echinopsis minuana Echinocereus palmeri Echinopsis angelesii Echinopsis cuzcoensis Echinopsis mirabilis Echinocereus pamanesiorum Echinopsis antezanae Echinopsis densispina Echinopsis molesta Echinocereus papillosus Echinopsis arboricola Echinopsis derenbergii Echinopsis nigra Echinocereus parkeri Echinopsis arebaloi Echinopsis deserticola Echinopsis obrepanda Echinocereus pectinatus Echinopsis atacamensis Echinopsis escayachensis Echinopsis oxygona Echinocereus pensilis Echinopsis aurea Echinopsis eyriesii Echinopsis pachanoi Echinocereus pentalophus Echinopsis backebergii Echinopsis fabrisii Echinopsis pampana Echinocereus polyacanthus Echinopsis baldiana Echinopsis famatimensis Echinopsis pentlandii Echinocereus poselgeri Echinopsis bertramiana Echinopsis ferox Echinopsis peruviana Echinocereus primolanatus Echinopsis bolligeriana Echinopsis formosa Echinopsis pojoensis Echinocereus pseudopectinatus Echinopsis boyuibensis Echinopsis friedrichii Echinopsis pseudomamillosa Echinocereus pulchellus Echinopsis brasiliensis Echinopsis glauca Echinopsis pugionacantha Echinocereus rayonesensis Echinopsis breviflora Echinopsis glaucina Echinopsis quadratiumbonata Echinocereus reichenbachii Echinopsis bridgesi Echinopsis haematantha Echinopsis rhodotricha Echinocereus rigidissimus Echinopsis bruchii Echinopsis hahniana Echinopsis riviere-de-caraltii Echinocereus russanthus Echinopsis x cabrerae Echinopsis hammerschmidii Echinopsis rojasii Echinocereus scheeri Echinopsis caineana Echinopsis hertrichiana Echinopsis saltensis Echinocereus schereri Echinopsis cajasensis Echinopsis huascha Echinopsis sanguiniflora Echinocereus schmollii Echinopsis calliantholilacina Echinopsis huotii Echinopsis santaensis Echinocereus sciurus Echinopsis callichroma Echinopsis hystrichoides Echinopsis schickendantzii* Echinocereus scopulorum Echinopsis calochlora Echinopsis ibicuatensis Echinopsis schieliana Echinocereus spinigemmatus Echinopsis camarguensis Echinopsis jajoana Echinopsis schoenii Echinocereus stoloniferus Echinopsis candicans Echinopsis kladiwaiana Echinopsis schrieteri Echinocereus stramineus Echinopsis cephalomacrostibas Echinopsis klingleriana Echinopsis scopulicola Echinocereus subinermis Echinopsis cerdana Echinopsis knuthiana Echinopsis silvestrii Echinocereus triglochidiatus Echinopsis chalaensis Echinopsis korethroides Echinopsis skottsbergii Echinopsis smrziana Epiphyllum oxypetalum Eriosyce tenebrica Espostoa mirabilis Echinopsis spachiana* Epiphyllum phyllanthus Eriosyce umadeave Espostoa nana Echinopsis spinibarbis Epiphyllum pittieri Eriosyce vertongenii Espostoa ritteri Echinopsis strigosa Epiphyllum pumilum Eriosyce villicumensis Espostoa ruficeps Echinopsis subdenudata Epiphyllum rubrocoronatum Eriosyce villosa Espostoa senilis Echinopsis sucrensis Epiphyllum thomasianum Escobaria albicolumnaria Espostoa superba Echinopsis tacaquirensis Epiphyllum trimetrale Escobaria alversonii Espostoa utcubambensis Echinopsis taratensis Epithelantha bokei Escobaria chihuahuensis Espostoopsis dybowskii Echinopsis tarijensis Epithelantha micromeris Escobaria cubensis Estevesia alex-bragae Echinopsis tarmaensis Eriosyce aericarpa Escobaria dasyacantha Eulychnia acida Echinopsis tegeleriana Eriosyce andreaeana Escobaria deserti Eulychnia breviflora Echinopsis terscheckii Eriosyce aspillagae Escobaria duncanii Eulychnia castanea Echinopsis thelegona Eriosyce aurata Escobaria emskoetteriana Eulychnia iquiquensis Echinopsis thelegonoides Eriosyce bulbocalyx Escobaria guadalupensis Eulychnia ritteri Echinopsis thionantha Eriosyce chilensis Escobaria hesteri Facheiroa cephaliomelana Echinopsis tiegeliana Eriosyce confinis Escobaria laredoi Facheiroa squamosa Echinopsis trichosa Eriosyce crispa Escobaria lloydii Facheiroa ulei Echinopsis tubiflora Eriosyce curvispina Escobaria minima Ferocactus alamosanus Echinopsis tulhuayacensis Eriosyce engleri Escobaria missouriensis Ferocactus chrysacanthus Echinopsis tunariensis Eriosyce esmeraldana Escobaria orcuttii Ferocactus cylindraceus Echinopsis uyupampensis Eriosyce garaventae Escobaria organensis Ferocactus diguetii Echinopsis vasquezii Eriosyce heinrichiana Escobaria robbinsorum Ferocactus eastwoodiae Echinopsis vatteri Eriosyce islayensis Escobaria sandbergii Ferocactus echidne Echinopsis volliana Eriosyce krausii Escobaria sneedii Ferocactus emoryi Echinopsis walteri Eriosyce kunzei Escobaria tuberculosa Ferocactus flavovirens Echinopsis werdermannii Eriosyce laui Escobaria villardii Ferocactus fordii Echinopsis yuquina Eriosyce limariensis Escobaria vivipara Ferocactus glaucescens Epiphyllum anguliger Eriosyce marksiana Escobaria zilziana Ferocactus gracilis Epiphyllum cartagense Eriosyce napina Escontria chiotilla Ferocactus haematacanthus Epiphyllum caudatum Eriosyce occulta Espostoa baumannii Ferocactus hamatacanthus Epiphyllum columbiense Eriosyce odieri Espostoa blossfeldiorum Ferocactus herrehae Epiphyllum costaricense Eriosyce omasensis Espostoa calva Ferocactus histrix Epiphyllum crenatum Eriosyce recondita Espostoa frutescens Ferocactus johnstonianus Epiphyllum floribundum Eriosyce rodentiophila Espostoa guentheri Ferocactus latispinus Epiphyllum grandilobum Eriosyce senilis Espostoa huanucoensis Ferocactus lindsayi Epiphyllum guatemalense Eriosyce sociabilis Espostoa hylaea Ferocactus macrodiscus Epiphyllum hookeri Eriosyce strausiana Espostoa lanata Ferocactus peninsulae Epiphyllum laui Eriosyce subgibbosa Espostoa lanianuligera Ferocactus pilosus Epiphyllum lepidocarpum Eriosyce taltalensis Espostoa melanostele Ferocactus pottsi Ferocactus robustus Grusonia reflexispina Gymnocalycium marianae Haageocereus acranthus Ferocactus santa-maria Grusonia robertsii Gymnocalycium marsoneri Haageocereus albispinus Ferocactus schwarzii Grusonia schottii Gymnocalycium megalothelon Haageocereus australis Ferocactus tiburonensis Grusonia vilis Gymnocalycium mesopotamicum Haageocereus chalaensis Ferocactus townsendianus Gymnocalycium albiareolatum Gymnocalycium mihanovichii Haageocereus chilensis Ferocactus viridescens Gymnocalycium ambatoense Gymnocalycium monvillei Haageocereus chryseus Ferocactus wislizeni Gymnocalycium amerhauseri Gymnocalycium mostii Haageocereus decumbens Frailea buenekeri Gymnocalycium andreae Gymnocalycium mucidum Haageocereus fascilularis Frailea buiningiana Gymnocalycium angelae Gymnocalycium netrelianum Haageocereus icensis Frailea castanea Gymnocalycium anisitsii Gymnocalycium neuhuberi Haageocereus icosagonoides Frailea cataphracta Gymnocalycium baldianum Gymnocalycium obductum Haageocereus lanugispinus Frailea chiquitana Gymnocalycium bayrianum Gymnocalycium ochoterenae Haageocereus pacalaensis Frailea curvispina Gymnocalycium berchtii Gymnocalycium oenanthemum Haageocereus platinospinus Frailea friedrichii Gymnocalycium bodenbenderianum Gymnocalycium paediophilum Haageocereus pluriflorus Frailea gracillima Gymnocalycium borthii Gymnocalycium paraguayense Haageocereus pseudomelanostele Frailea grahliana Gymnocalycium bruchii Gymnocalycium parvulum Haageocereus pseudoversicolor Frailea knippeliana Gymnocalycium buenekeri Gymnocalycium pflanzii Haageocereus subtilispinus Frailea mammifera Gymnocalycium calochlorum Gymnocalycium platense Haageocereus tenuis Frailea perumbilicata Gymnocalycium capillaense Gymnocalycium pugionacanthum Haageocereus versicolor Frailea phaeodisca Gymnocalycium carmianthum Gymnocalycium quehlianum Haageocereus vulpes Frailea pseudopulcherrima Gymnocalycium castellanosii Gymnocalycium ragonesei Haageocereus zangalensis Frailea pumila Gymnocalycium catamarcense Gymnocalycium rauschii xHaagespostoa albisetata Frailea pygmaea Gymnocalycium chiquitanum Gymnocalycium riojense xHaagespostoa climaxantha Frailea schilinzkyana Gymnocalycium deeszianum Gymnocalycium ritterianum Harrisia adscendens Geohintonia mexicana Gymnocalycium delaetii Gymnocalycium robustum Harrisia alboriginum Grusonia aggeria Gymnocalycium denudatum Gymnocalycium rosae Harrisia balansae* Grusonia agglomerata Gymnocalycium erinaceum Gymnocalycium saglionis Harrisia brookii Grusonia bradtiana Gymnocalycium eurypleurum Gymnocalycium schickendantzii Harrisia divaricata Grusonia bulbispina Gymnocalycium eytianum Gymnocalycium schroederianum Harrisia earlei Grusonia clavata Gymnocalycium gibbosum Gymnocalycium spegazzinii Harrisia eriophora Grusonia dumetorum Gymnocalycium horstii Gymnocalycium stellatum Harrisia fernowii Grusonia emoryi Gymnocalycium hossei Gymnocalycium stenopleurum Harrisia fragans Grusonia grahamii Gymnocalycium hybopleurum Gymnocalycium striglianum Harrisia gracilis Grusonia Invicta Gymnocalycium hyptiacanthum Gymnocalycium stuckertii Harrisia hurstii Grusonia kunzei Gymnocalycium kieslingii Gymnocalycium taningaense Harrisia martini* Grusonia marenae Gymnocalycium kroenleinii Gymnocalycium terweemeanum Harrisia nashii Grusonia moelleri Gymnocalycium leeanum Gymnocalycium tillianum Harrisia pomanensis* Grusonia parishii Gymnocalycium leptanthum Gymnocalycium uebelmannianum Harrisia portoricensis Grusonia pulchella Gymnocalycium mackieanum Gymnocalycium uruguayense Harrisia regelii Harrisia simpsonii Lepismium brevispinum Maihueniopsis crassispina Mammillaria chinocephala Harrisia taetra Lepismium crenatum Maihueniopsis darwinii Mammillaria coahuilensis Harrisia taylori Lepismium cruciforme Maihueniopsis domeykoensis Mammillaria columbiana Harrisia tetracantha Lepismium houlletianum Maihueniopsis glomerata Mammillaria compressa Harrisia tortuosa* Lepismium ianthothele Maihueniopsis grandiflora Mammillaria crinita Hatiora cylindrica Lepismium incachacanum Maihueniopsis minuta Mammillaria crucigera Hatiora epiphylloides Lepismium lorentzianum Maihueniopsis nigrispina Mammillaria decipiens Hatiora gaertneri Lepismium lumbricoides Maihueniopsis ovata Mammillaria deherdtiana Hatiora herminiae Lepismium micranthum Maihueniopsis rahmeri Mammillaria densispina Hatiora rosea Lepismium miyagawae Maihueniopsis subterranea Mammillaria dioica Hatiora salicornioides Lepismium monacanthum Maihueniopsis tarapacana Mammillaria discolor Hatiora x graeseri Lepismium paranganiense Maihueniopsis wagenknechtii Mammillaria dixanthocentron Hylocereus bronxensis Lepismium warmingianum Mammillaria albicans Mammillaria duoformis Hylocereus calcaratus Leptocereus arboreus Mammillaria albicoma Mammillaria ekmanii Hylocereus costaricensis* Leptocereus assurgens Mammillaria albiflora Mammillaria elongata Hylocereus escuintlensis Leptocereus carinatus Mammillaria albilanata Mammillaria eriacantha Hylocereus estebanensis Leptocereus ekmanii Mammillaria amajacensis Mammillaria erythrosperma Hylocereus extensus Leptocereus grantianus Mammillaria angelensis Mammillaria estebanensis Hylocereus guatemalensis Leptocereus leonii Mammillaria anniana Mammillaria evermanniana Hylocereus lemairei Leptocereus maxonii Mammillaria armillata Mammillaria fittkaui Hylocereus microcladus Leptocereus paniculatus Mammillaria aureilanata Mammillaria flavicentra Hylocereus minutiflorus Leptocereus prostratus Mammillaria backebergiana Mammillaria formosa Hylocereus monacanthus Leptocereus quadricostatus Mammillaria barbata Mammillaria fraileana Hylocereus ocamponis Leptocereus santamarinae Mammillaria baumii Mammillaria gasseriana Hylocereus peruvianus Leptocereus scopulophilus Mammillaria beneckei Mammillaria geminispina Hylocereus polyrhizus* Leptocereus sylvestris Mammillaria blossfeldiana Mammillaria gigantea Hylocereus purpusii Leptocereus weingartianus Mammillaria bocasana Mammillaria glassii Hylocereus scandens Leptocereus wrightii Mammillaria bocensis Mammillaria glochidiata Hylocereus stenopterus Leuchtenbergia principis Mammillaria boelderliana Mammillaria goodridgei Hylocereus triangularis* Lophophora diffusa Mammillaria bombycina Mammillaria grahamii Hylocereus trigonus Lophophora williamsii Mammillaria boolii Mammillaria grusonii Hylocereus undatus* Maihuenia patagonica Mammillaria brachytrichion Mammillaria guelzowiana Isolatocereus dumortieri Maihuenia poeppigii Mammillaria brandegeei Mammillaria guerreronis Jasminocereus thouarsii Maihueniopsis archiconoidea Mammillaria canelensis Mammillaria guillauminiana Lasiocereus fulvus Maihueniopsis atacamensis Mammillaria capensis Mammillaria haageana Lasiocereus rupicola Maihueniopsis bonnieae Mammillaria carmenae Mammillaria hahniana Leocereus bahiensis Maihueniopsis camachoi Mammillaria carnea Mammillaria halbingeri Lepismium aculeatum Maihueniopsis clavarioides Mammillaria carretii Mammillaria heidiae Lepismium bolivianum Maihueniopsis colorea Mammillaria cerralboa Mammillaria hernandezii Mammillaria herrerae Mammillaria mieheana Mammillaria senilis Matucana madisoniorum Mammillaria hertrichiana Mammillaria moelleriana Mammillaria sheldonii Matucana oreodoxa Mammillaria heyderi Mammillaria morganiana Mammillaria solisioides Matucana paucicostata Mammillaria huitzilopochtli Mammillaria muehlenfordtii Mammillaria sonorensis Matucana polzii Mammillaria humboldtii Mammillaria multidigitata Mammillaria sphacelata Matucana pujupatii Mammillaria hutchisoniana Mammillaria mystax Mammillaria sphaerica Matucana ritteri Mammillaria insularis Mammillaria napina Mammillaria spinosissima Matucana tuberculata Mammillaria jaliscana Mammillaria neopalmeri Mammillaria standleyi Matucana webwebaueri Mammillaria johnstonii Mammillaria nivosa Mammillaria stella-de-tacubaya Melocactus x albicephalus Mammillaria karwinskiana Mammillaria nunezii Mammillaria supertexta Melocactus andinus Mammillaria klissingiana Mammillaria orcuttii Mammillaria surculosa Melocactus azureus Mammillaria kraehenbuehlii Mammillaria oteroi Mammillaria tayloriorum Melocactus bahiensis Mammillaria lasiacantha Mammillaria parkinsonii Mammillaria tepexicensis Melocactus bellavistensis Mammillaria laui Mammillaria pectinifera Mammillaria tetrancistra Melocactus braunii Mammillaria lenta Mammillaria peninsularis Mammillaria theresae Melocactus broadwayi Mammillaria limonensis Mammillaria pennispinosa Mammillaria thornberi Melocactus caroli-linnaei Mammillaria linaresensis Mammillaria perbella Mammillaria tonalensis Melocactus concinnus Mammillaria lindsayi Mammillaria perezdelarosae Mammillaria uncinata Melocactus conoideus Mammillaria lloydii Mammillaria petrophila Mammillaria varieaculeata Melocactus curvispinus Mammillaria longiflora Mammillaria petterssonii Mammillaria vetula Melocactus deinacanthus Mammillaria longimamma Mammillaria phitauiana Mammillaria voburnensis Melocactus ernestii Mammillaria luethyi Mammillaria picta Mammillaria wagneriana Melocactus estevesii Mammillaria magallanii Mammillaria pilispina Mammillaria weingartiana Melocactus glaucescens Mammillaria magnifica Mammillaria plumosa Mammillaria wiesingeri Melocactus harlowii Mammillaria magnimamma Mammillaria polyedra Mammillaria winterae Melocactus x horridus Mammillaria mainiae Mammillaria polythele Mammillaria wrightii Melocactus intortus Mammillaria mammillaris Mammillaria pottsii Mammillaria xaltianguensis Melocactus lanssensianus Mammillaria manana Mammillaria prolifera Mammillaria zephyranthoides Melocactus lemairei Mammillaria marcosii Mammillaria rekoi Mammillaria zublerae Melocactus levitestatus Mammillaria marksiana Mammillaria rhodantha Mammilloydia candida Melocactus macracanthos Mammillaria mathildae Mammillaria roseoalba Matucana aurantiaca Melocactus matanzanus Mammillaria matudae Mammillaria saboae Matucana aureiflora Melocactus mazelianus Mammillaria mazatlanensis Mammillaria sanchez-mejoradae Matucana comacephala Melocactus neryi Mammillaria melaleuca Mammillaria sartorii Matucana formosa Melocactus oreas Mammillaria melanocentra Mammillaria schiedeana Matucana fruticosa Melocactus pachyacanthus Mammillaria mercadensis Mammillaria schumannii Matucana haynei Melocactus paucispinus Mammillaria meyranii Mammillaria schwarzii Matucana huagalensis Melocactus perezassoi Mammillaria microhelia Mammillaria scrippsiana Matucana intertexta Melocactus peruvianus Mammillaria miegiana Mammillaria sempervivi Matucana krahnii Melocactus praerupticola Melocactus salvadorensis Opuntia abjecta Opuntia cochabambensis Opuntia huajuapensis* Melocactus schatzlii Opuntia acaulis Opuntia cochenillifera* Opuntia humifusa* Melocactus smithii Opuntia aciculata Opuntia cognata Opuntia hyptiacantha Melocactus stramineus Opuntia alko-tuna Opuntia colubrina Opuntia inaequilateralis Melocactus violaceus Opuntia ammophila* Opuntia conjungens Opuntia inaperta Melocactus zehntneri Opuntia amyclaea* Opuntia crassa* Opuntia infesta Micranthocereus albicephalus Opuntia anacantha Opuntia crystalenia Opuntia insularis Micranthocereus auriazurensis Opuntia antillana Opuntia curassavica Opuntia jaliscana Micranthocereus dolichospermaticus Opuntia apurimacensis Opuntia cymochila Opuntia jamaicensis Micranthocereus estevesii Opuntia arcei Opuntia darrahiana Opuntia joconostle Micranthocereus flaviflorus Opuntia armata Opuntia deamii Opuntia karwinskiana Micranthocereus hofackerianus Opuntia assumptionis Opuntia decumbens Opuntia laevis Micranthocereus polyanthus Opuntia atrispina Opuntia dejecta* Opuntia lagunae Micranthocereus purpureus Opuntia atropes Opuntia delaetiana Opuntia larreyi Micranthocereus streckeri Opuntia atrovirens Opuntia depauperata Opuntia lasiacantha Micranthocereus violaciflorus Opuntia auberi* Opuntia depressa Opuntia lata Mila caespitosa Opuntia aurantiaca* Opuntia dillenii* Opuntia leucotricha* Miqueliopuntia miquelii Opuntia arechavaletae Opuntia discolor Opuntia lilae xMyrtgerocactus lindsayi Opuntia aurea Opuntia durangensis Opuntia limitata Myrtillocactus cochal Opuntia aureispina Opuntia echios Opuntia littoralis Myrtillocactus eichlamii Opuntia austrina Opuntia eichlamii Opuntia lutea Myrtillocactus geometrizans* Opuntia azurea Opuntia ekmanii Opuntia macrocentra Myrtillocactus schenckii Opuntia basilaris Opuntia elata* Opuntia macrorhiza Neobuxbaumia euphorbioides Opuntia bella Opuntia elatior* Opuntia martiniana Neobuxbaumia laui Opuntia bensonii Opuntia elizondoana Opuntia megacantha Neobuxbaumia macrocephala Opuntia bisetosa Opuntia ellisiana Opuntia megapotamica Neobuxbaumia mezcalaensis Opuntia boldinghii Opuntia engelmannii* Opuntia megarhiza Neobuxbaumia multiareolata Opuntia bonplandii Opuntia excelsa Opuntia megasperma Neobuxbaumia polylopha Opuntia borinquensis Opuntia feroacantha Opuntia microdasys* Neobuxbaumia scoparia Opuntia bravoana Opuntia ficus-indica* Opuntia mieckleyi Neobuxbaumia squamulosa Opuntia canterae Opuntia fragilis Opuntia monocantha* Neobuxbaumia tetetzo Opuntia caracassana Opuntia fuliginosa Opuntia montevideensis Neolloydia conoidea Opuntia cardiosperma Opuntia galapageia Opuntia nejapensis Neolloydia matehualensis Opuntia chaffeyi Opuntia gosseliniana Opuntia neochrysacantha Neoraimondia arequipensis Opuntia chakensis Opuntia guatemalensis Opuntia nuda Neoraimondia herzogiana Opuntia chavena Opuntia guilanchi Opuntia orbiculata Neowerdermannia chilensis Opuntia chihuahuensis Opuntia helleri Opuntia oricola Neowerdermannia vorwerkii Opuntia chisosensis* Opuntia hondurensis Opuntia pachona Obregonia denegrii Opuntia chlorotica Opuntia howeyi Opuntia pachyrrhiza Opuntia pailana Opuntia schumannii Oreocereus celsianus Parodia curvispina Opuntia pampeana Opuntia securigera Oreocereus doelzianus Parodia erinacea Opuntia paraguayensis Opuntia soederstromiana Oreocereus hempelianus Parodia erubescens Opuntia parviclada Opuntia spinulifera* Oreocereus leucotrichus Parodia formosa Opuntia penicilligera Opuntia spraguei Oreocereus pseudofossulatus Parodia fusca Opuntia pennellii Opuntia stenarthra Oreocereus ritteri Parodia gaucha Opuntia phaeacantha* Opuntia stenopetala Oreocereus tacnaensis Parodia glaucina Opuntia pilifera Opuntia streptacantha* Oreocereus trollii Parodia haselbergii Opuntia pinkavae Opuntia stricta* Oreocereus varicolor Parodia hausteiniana Opuntia pittieri Opuntia strigil Oroya borchersii Parodia herteri Opuntia pituitosa Opuntia subsphaerocarpa Oroya peruviana Parodia horstii Opuntia polyacantha Opuntia sulfurea Ortegocactus macdougallii Parodia langsdorfii Opuntia pottsii Opuntia tapona xPacherocactus orcuttii Parodia leninghausii Opuntia prasina Opuntia taylori Pachycereus fulviceps Parodia linkii Opuntia puberula Opuntia tehuacana Pachycereus gatesii Parodia maassii Opuntia pubescens* Opuntia tehuantepecana Pachycereus gaumeri Parodia magnifica Opuntia pumila Opuntia tenuiflora Pachycereus grandis Parodia mammulosa Opuntia pusilla Opuntia tomentella Pachycereus hollianus Parodia meonacantha Opuntia pycnantha Opuntia tomentosa* Pachycereus lepidanthus Parodia microsperma Opuntia pyriformis Opuntia triacantha Pachycereus marginatus Parodia mueller-melchersii Opuntia pyrrhantha Opuntia tuna* Pachycereus militaris Parodia muricata Opuntia quimilo Opuntia turbinata Pachycereus pecten-aboriginum Parodia neoarechavaletae Opuntia quitensis Opuntia undulata Pachycereus pringlei Parodia neohorstii Opuntia rastrera Opuntia urbaniana Pachycereus schottii Parodia nigrispina Opuntia repens Opuntia velutina Pachycereus tepamo Parodia nivosa Opuntia rileyi Opuntia viridirubra Pachycereus weberi Parodia nothominuscula Opuntia ritteri Opuntia vitelliniflora Parodia alacriportana Parodia nothorauschii Opuntia robinsonnii Opuntia wetmorei Parodia allosiphon Parodia ocampoi Opuntia roborensis Opuntia wilcoxii Parodia arnostiana Parodia ottonis Opuntia robusta* Opuntia x aequatorialis Parodia aureicentra Parodia oxycostata Opuntia rufida Opuntia x bakeri Parodia ayopayana Parodia penicillata Opuntia salagria Opuntia x columbiana Parodia buiningii Parodia permutata Opuntia salmiana* Opuntia x cubensis Parodia carambeiensis Parodia procera Opuntia salvadorensis Opuntia x curvospina Parodia chrysacanthion Parodia rechensis Opuntia sanguinea Opuntia x lucayana Parodia columnaris Parodia ritteri Opuntia santa-rita Opuntia x occidentalis Parodia comarapana Parodia rudibuenekeri Opuntia saxicola Opuntia x spinosibacca Parodia commutans Parodia rutilans Opuntia scheeri Opuntia x vaseyi Parodia concinna Parodia saint-pieana Opuntia schickendantzii Opuntia x wootonii Parodia crassigibba Parodia schumanniana Parodia schwebsiana Peniocereus occidentalis Pilosocereus brasiliensis Pseudorhipsalis acuminata Parodia scopa Peniocereus rosei Pilosocereus catingicola Pseudorhipsalis alata Parodia sellowii Peniocereus serpentinus* Pilosocereus chrysacanthus Pseudorhipsalis amazonica Parodia stockingeri Peniocereus striatus Pilosocereus chrysostele Pseudorhipsalis himantoclada Parodia stuemeri Peniocereus tepalcatepecanus Pilosocereus collinsii Pseudorhipsalis horichii Parodia subterranea Peniocereus viperinus Pilosocereus densiareolatus Pseudorhipsalis lankesteri Parodia tabularis Peniocereus zopilotensis Pilosocereus diersianus Pseudorhipsalis ramulosa Parodia taratensis Pereskia aculeata* Pilosocereus estevesii Pterocactus araucanus Parodia tenuicylindrica Pereskia aureiflora Pilosocereus flavipulvinatus Pterocactus australis Parodia tilcarensis Pereskia bahiensis Pilosocereus flexibilispinus Pterocactus fischeri Parodia tuberculata Pereskia bleo Pilosocereus floccosus Pterocactus gonjianii Parodia turbinata Pereskia diaz-romeroana Pilosocereus frewenii Pterocactus hickenii Parodia turecekiana Pereskia grandiflora Pilosocereus fulvilanatus Pterocactus megiolii Parodia warasii Pereskia guamacho Pilosocereus glaucochrous Pterocactus reticulatus Parodia werdermanniana Pereskia horrida Pilosocereus gounellei Pterocactus tuberosus Parodia werneri Pereskia lychnidiflora Pilosocereus lanuginosus Pterocactus valentinii Pediocactus bradyi Pereskia marcanoi Pilosocereus leucocephalus Pygmaeocereus bieblii Pediocactus despainii Pereskia nemorosa Pilosocereus machrisii Pygmaeocereus bylesianus Pediocactus hermannii Pereskia portulacifolia Pilosocereus magnificus Pygmaeocereus familiaris Pediocactus knowltonii Pereskia quinqueyana Pilosocereus multicostatus Quiabentia verticillata Pediocactus nigrispinus Pereskia sacharosa Pilosocereus occultiflorus Quiabentia zehntneri Pediocactus paradinei Pereskia stenantha Pilosocereus oligolepis Rauhocereus riosaniensis Pediocactus peeblesianus Pereskia weberiana Pilosocereus pachycladus Rebutia albiflora Pediocactus simpsonii Pereskia zinniiflora Pilosocereus pentaedrophorus Rebutia albipectinata Pediocactus winkleri Pereskiopsis aquosa Pilosocereus piauhyensis Rebutia arenacea Pelecyphora aselliformis Pereskiopsis blakeana Pilosocereus polygonus Rebutia aureiflora Pelecyphora strobiliformis Pereskiopsis diguetii Pilosocereus purpusii Rebutia brunescens Peniocereus castellae Pereskiopsis gatesii Pilosocereus quadricentralis Rebutia caineana Peniocereus chiapensis Pereskiopsis kellermanii Pilosocereus royenii Rebutia canigueralii Peniocereus cuixmalensis Pereskiopsis porteri Pilosocereus x subsimilis Rebutia cardenasiana Peniocereus fosterianus Pereskiopsis rotundifolia Pilosocereus tuberculatus Rebutia cintia Peniocereus greggii Pereskiopsis spathulata Pilosocereus ulei Rebutia cylindrica Peniocereus hirschtianus Pfeiffera asuntapatensis Pilosocereus vilaboensis Rebutia deminuta Peniocereus johnstonii Pilosocereus albisummus Polaskia chende Rebutia einsteinii Peniocereus lazaro-cardenasii Pilosocereus alensis Polaskia chichipe Rebutia fabrisii Peniocereus macdougallii Pilosocereus arrabidae Praecereus euchlorus Rebutia fidaiana Peniocereus maculatus Pilosocereus aureispinus Praecereus saxicola Rebutia fiebrigii Peniocereus marianus Pilosocereus aurisetus Pseudoacanthocereus brasiliensis Rebutia flavistyla Peniocereus oaxacensis Pilosocereus azulensis Pseudoacanthocereus sicariguensis Rebutia fulviseta Rebutia gonjianii Rhipsalis goebeliana Sclerocactus papyracanthus Siccobaccatus estevesii Rebutia heliosa Rhipsalis grandiflora Sclerocactus parviflorus Siccobaccatus insigniflorus Rebutia huasiensis Rhipsalis hoelleri Sclerocactus polyancistrus Stenocactus coptonogonus Rebutia krugerae Rhipsalis juengeri Sclerocactus pubispinus Stenocactus crispatus Rebutia leucanthema Rhipsalis lindbergiana Sclerocactus scheeri Stenocactus hastatus Rebutia marsoneri Rhipsalis mesembryanthemoides Sclerocactus sileri Stenocactus multicostatus Rebutia mentosa Rhipsalis micrantha Sclerocactus spinosior Stenocactus obvallatus Rebutia minuscula Rhipsalis neves-armondii Sclerocactus uncinatus Stenocactus ochoterenanus Rebutia narvaecensis Rhipsalis oblonga Sclerocactus wetlandicus Stenocactus phyllacanthus Rebutia neocumingii Rhipsalis occidentalis Sclerocactus whipplei Stenocactus rectispinus Rebutia neumanniana Rhipsalis olivifera Sclerocactus wrightiae Stenocactus sulfureus Rebutia nigricans Rhipsalis ormindoi Selenicereus anthonyanus Stenocactus vaupelianus Rebutia oligacantha Rhipsalis pacheco-leonis Selenicereus atropilosus Stenocereus alamosensis Rebutia padcayensis Rhipsalis pachyptera Selenicereus boeckmannii Stenocereus aragonii Rebutia perplexa Rhipsalis paradoxa Selenicereus brevispinus Stenocereus beneckei Rebutia pseudodeminuta Rhipsalis pentaptera Selenicereus chontalensis Stenocereus chacalapensis Rebutia pulvinosa Rhipsalis pilocarpa Selenicereus chrysocardium Stenocereus chrysocarpus Rebutia pygmaea Rhipsalis pulchra Selenicereus coniflorus Stenocereus eichlamii Rebutia ritteri Rhipsalis puniceodiscus Selenicereus donkelaari Stenocereus eruca Rebutia simoniana Rhipsalis russellii Selenicereus grandiflorus Stenocereus fimbriatus Rebutia spegazziniana Rhipsalis sulcata Selenicereus hamatus Stenocereus fricii Rebutia spinosissima Rhipsalis teres Selenicereus hondurensis Stenocereus griseus Rebutia steinbachii Rhipsalis triangularis Selenicereus inermis Stenocereus gummosus Rebutia steinmannii Rhipsalis trigona Selenicereus innesii Stenocereus kerberi Rebutia wessneriana Samaipaticereus corroanus Selenicereus macdonaldiae* Stenocereus laevigatus Rebutia xanthocarpa Schlumbergera kautskyi Selenicereus megalanthus Stenocereus martinezii Rhipsalis agudoensis Schlumbergera microsphaerica Selenicereus murrillii Stenocereus montanus Rhipsalis baccifera Schlumbergera opuntioides Selenicereus nelsonii Stenocereus pruinosus Rhipsalis burchellii Schlumbergera orssichiana Selenicereus pteranthus Stenocereus queretaroensis Rhipsalis campos-portoana Schlumbergera russelliana Selenicereus rubineus Stenocereus quevedonis Rhipsalis cereoides Schlumbergera truncate Selenicereus setaceus Stenocereus standleyi Rhipsalis cereuscula Schlumbergera x buckleyi Selenicereus spinulosus Stenocereus stellatus Rhipsalis clavata Schlumbergera x exotica Selenicereus testudo Stenocereus thurberi Rhipsalis crispata Schlumbergera x reginae Selenicereus tricae Stenocereus treleasei Rhipsalis cuneata Sclerocactus brevihamatus Selenicereus urbanianus Stenocereus yunckeri Rhipsalis dissimilis Sclerocactus brevispinus Selenicereus vagans Stephanocereus leucostele Rhipsalis elliptica Sclerocactus glaucus Selenicereus validus Stephanocereus luetzelburgii Rhipsalis ewaldiana Sclerocactus mesae-verdae Selenicereus wercklei Stetsonia coryne Rhipsalis floccosa Sclerocactus nyensis Selenicereus wittii Strombocactus disciformis Tacinga braunii Thelocactus lausseri Turbinicarpus jauernigii Weberbauerocereus cuzcoensis Tacinga estevesii Thelocactus leucacanthus Turbinicarpus knuthianus Weberbauerocereus johnsonii Tacinga funalis Thelocactus macdowellii Turbinicarpus laui Weberbauerocereus longicomus Tacinga inamoena Thelocactus rinconensis Turbinicarpus lophophoroides Weberbauerocereus rauhii Tacinga palmadora Thelocactus setispinus Turbinicarpus mandragora Weberbauerocereus torataensis Tacinga saxatilis Thelocactus tulensis Turbinicarpus x mombergii Weberbauerocereus weberbaueri Tacinga subcylindrica Tunilla albisetacens Turbinicarpus pailanus Weberbauerocereus winterianus Tacinga werneri Tunilla chilensis Turbinicarpus pseudomacrochele Weberocereus biolleyi Tacinga x quipa Tunilla corrugata Turbinicarpus pseudopectinatus Weberocereus bradei Tephrocactus alexanderi Tunilla erectoclada Turbinicarpus rioverdensis Weberocereus frohningiorum Tephrocactus aoracanthus Tunilla ianthinantha Turbinicarpus saueri Weberocereus glaber Tephrocactus articulatus* Tunilla minuscula Turbinicarpus schmiedickeanus Weberocereus imitans Tephrocactus geometricus Tunilla orurensis Turbinicarpus subterraneus Weberocereus panamensis Tephrocactus molinensis Tunilla silvestris Turbinicarpus swobodae Weberocereus rosei Tephrocactus weberi Tunilla soehrensii Turbinicarpus valdezianus Weberocereus tonduzii Thelocactus bicolor Turbinicarpus alonsoi Turbinicarpus viereckii Weberocereus trichophorus Thelocactus buekii Turbinicarpus beguinii Turbinicarpus ysabelae Weberocereus tunilla Thelocactus conothelos Turbinicarpus bonatzii Turbinicarpus zaragosae Yavia cryptocarpa Thelocactus garciae Turbinicarpus booleanus Uebelmannia gummifera Yungasocereus inquis Thelocactus hastifer Turbinicarpus gielsdorfianus Uebelmannia pectinifera Thelocactus heterochromus Turbinicarpus hoferi Uebemannia buiningii Thelocactus hexaedrophorus Turbinicarpus horripilus Weberbauerocereus churinensis

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.