Leuenberger on Pereskia

Haseltonia.14:.54–93..2007. 54

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IntroductionPereskia is one of the earliest cacti known to botanists, first collected in the late 17th cen-tury and described as genus at the beginning of the 18th century. It is a genus of mostly tropi-

cal distribution, of leaf bearing, only slightly succulent, deciduous, spiny trees, shrubs, or vines. Plants belonging to Maihuenia were found quite early in the 19th century but, due to their habit were first classified as species of

Abstract: The taxonomic history of the three genera Pereskia, Maihuenia, and Blossfeldia is reviewed and discussed in the context of general historical aspects of cactus taxonomy and in view of ongoing discussion on evolution and phylogenetic position of the three genera. Some updates and additional observations complementing earlier monographic treatments of Pereskia and Maihuenia and other papers on all three genera are provided. The appearance of these cacti since 2004 on websites is briefly analysed with selected data retrieved from Google searches. All recognized species are illustrated.Key words: Cactaceae, taxonomic history, nomenclature, Pereskia, Maihuenia, Blossfeldia

Pereskia, Maihuenia, and Blossfeldia—taxonomiC history, uPdates, and notes

BEAT ERNST LEUENBERGERBotanischer Garten and Botanisches Museum Berlin-Dahlem,

Freie Universität Berlin, Berlin, Germany; [email protected]

Figure 1. Pereskia lychnidiflora, tree in dry forest near Tehuantepec, Oaxaca, Mexico. Figure 2. Maihuenia pata-gonica, a large cushion in habitat on black volcanic ash in the Payunia region southwest of Malargüe, Mendoza, Argentina. Figure 3. Blossfeldia liliputana, individual tiny plants growing in fissures of a weathered granitic rock face near Cacheuta, Mendoza, Argentina (scale in cm).

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Opuntia and later as Pereskia before being rec-ognized as a separate genus at the end of the 19th century. They are low, caespitose, spiny succulents with small but long lasting leaves. Their geographical range is predominantly Pa-tagonian. The rock-fissure-dwelling, tiny, glob-

ular cactus Blossfeldia was far off the path of the earliest botanical explorers, due to its range from the Andes of central Bolivia to western Argentina, but it was also overlooked by early collectors reaching this area. It was found and recognized as new only in the first half of the

Distribution of Pereskia.(dots, open circles refer to cultivated or subspontaneus records), Blossfeldia (rhom-boids) and Maihuenia (inverted triangles), modified from Leuenberger (1986, 1997) and Leuenberger and Eggli (1999), with updates. Map base from Flora Neotropica base map, University of Utrecht (1979), southern part added from (modified) Goode Base Map Series No. 103, University of Chicago (1961).

56. LEUENBERGER—Pereskia, MaiHuenia, aND Blossfeldia

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20th century, at a time when cactus botany was much influenced by commercial activities. The three genera thus belong to quite differ-ent eras of exploration and botanical taxon-omy, with the Pereskia story starting in pre-Linnaean time, in about 1689, Maihuenia in 1828, and Blossfeldia only in 1936.

Between 1986 and 1999 I contributed sev-eral papers on the three genera. Pereskia and Maihuenia were treated in classical taxonomic monographs (Leuenberger 1986, 1997) and in additional papers (Leuenberger 1992a, b, c, d, among others). Blossfeldia liliputana was also studied in the field in Argentina, and a bibliographic survey and review of the genus appeared in this journal (Leuenberger and Eggli 1999). It could hardly be foreseen at that time that all three genera would come into the particular focus of DNA studies relevant for basic and rather revolutionary questions of cactus evolution and systemat-ics (Wallace 1995b; Butterworth and Wal-lace 2005; Crozier 2004; Butterworth 2006; Nyffeler 2002; Edwards and 2005; Edwards and Donoghue 2006) The purpose of the present paper is primarily to give a sum-

mary of the taxonomic history of the three genera, including some updates and obser-vations, complemented by illustrations of some selected morphological features of these plants of widely differing habit and newly discussed phylogenetic and taxonomic po-sition (Figs 1–3).

Taxonomic history of PereskiaPereskia is one of the earliest cactus genera, already established in pre-Linnaean times by Plumier (1703). It was described from mate-rial collected by Charles Plumier, a French friar, naturalist, explorer, and botanist under King Louis XIV of France. Plumier’s herbar-ium collections of Pereskia were lost, but it can be assumed that they were collected be-tween 1689 and 1695 on the Caribbean isles. According to Mottram (2002: 83, 94), who reproduced the first description and pub-lished illustration of genus Pereskia by Plum-ier (1703) and Plumier’s unpublished plate of “Pereskia aculeata, flore albo, fructu flaves-cente” from the manuscript “Botanicon Amer-icanum,” the plate was drawn between 1695 and 1697. This first drawing is much more

Table 1. Taxonomic history of Pereskia (incl. Rhodocactus)

Author YearAccepted

species

Accepted infraspecific

taxa

Total of published names at species rank

in the same yearMiller 1754 1 0 1de Candolle 1828 9 0 9Pfeiffer 1837a 12 3 12Mittler 1844 8 3 12Salm-Dyck 1850 8 3 12Schumann 1898 11

( + 4 insufficiently known)

0 38(incl. nom. nud.)

Britton and Rose 1919–23 19 0 50Vaupel 1925 18 6 47

(incl. nom. nud.)Berger 1929 ca. 18 1 (23)

(listed in Berger 1929)Backeberg 1966 24

(8 Pereskia, 16 Rhodocactus)

1 53(35 Pereskia,

18 Rhodocactus)Leuenberger 1986 16 ( + 1 unnamed) 4 ca. 50

(ca. 59 in Index Kewensis incl. nom. nud.)

Hunt 1999 16 4 (37 listed by Hunt 1999)Anderson 2001 17 4Anderson 2005 17 4Hunt, Taylor and Charles 2006 17 4 ca. 63

(in IPNI incl. nom. nud.)


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representative of Pereskia aculeata than the illustration published by Plumier (1703).

In his grand work Species plantarum, which marks the starting point of modern botanical nomenclature, Linnaeus (1753) placed all cacti in the genus Cactus. Two of his species belong to the leafy cacti, Cactus pereskia and Cactus portulacifolius, but the connection of C. portu-lacifolius with Pereskia, as circumscribed later on, had not yet become evident. The British horticulturist Philip Miller did not follow the Linnaean one-genus concept for cacti and re-instated the genus Pereskia only one year later in his Gardners dictionary (Miller 1754). The genus Pereskia is therefore nomenclaturally valid not from Plumier (1703) but only from Miller (1754) as explained by Stafleu (1969) (Table 1). Miller (1754) mentioned only one species, here still with the pre-Linnaean phrase name “Pereskia aculeata, flore albo, fructu flaves-cente…” cited from Plumier (1703). The cur-rent name Pereskia aculeata was established by Miller (1768), avoiding the tautonym Pereskia pereskia (Figs 4–6).

For nearly two centuries, the genus Pereskia was not questioned again, though its circum-scription changed with the time. The major

“instability” concerned the spelling of the name. The name was explicitly published as Pereskia

Table 2. Pereskia and its orthographic variants found by Google searches in the WWW, 18. 10. 2007.

Name Image1

Pereskia 80900 32002

Peireskia 589 393Peirescia 241 69Perescia 353 60Pereskya 62 293

1 Usually only about 10% of the search results actually display an image showing a plant of the genus.

2 Less than 300 show a pereskia.3 Only two show a pereskia.

by Plumier (1703) and mentioned three times in the paper. The same spelling was used by Linnaeus in the epithet of Cactus pereskia and by Miller (1754) in genus Pereskia. The orig-inal spelling Pereskia was also maintained by Linnaeus (1738), Haworth (1812), de Can-dolle (1828), Förster (1846), Britton and Rose (1919) and Werdermann (1933, 1937a). It was modified by various other authors to “Peires-cia” (Zuccarini 1837; Pfeiffer 1837a, 1837b; Salm-Dyck 1850; Rümpler 1886), “Perescia” (Lemaire 1838), and “Peireskia” (Steudel 1841; Schumann 1890, 1898; Vaupel 1925; Berger 1929), based on the assumption that “Pereskia” was an orthographical error. Backeberg (1934)

Table 3. Taxonomic history of Cactaceae: number of genera and species.

Author Year Genera SpeciesLinnaeus 1753 1 22Candolle 1828 7 174Pfeiffer 1837 10 422Schumann 1898 21 578Britton and Rose 1919–23 124 1325Berger 1929 41 —Krainz 1960 119 —Backeberg 1966 236 3100Krainz 1967 152 —Hunt 1967 84 —Buxbaum in

Endler and Buxbaum

1974 159 —

Hunt and Taylor 1986 86 —Hunt 1999 114 1942Anderson 2001 125 1810Anderson 2005 126 19001

Hunt, Taylor and Charles

2006 124 14382

1 Plus about 500 subspecies and varieties.2 Plus 378 subspecies.

Figures 4–6. Pereskia aculeata. Figure 4. Young leaf-bearing stem with broad leaves, and an old spiny stem with one areole producing new spines. Figure 5. Recurved spines in the leaf axils of the young shoot and addi-tional straight spines formed on the areole. Figure 6. Mature fruit.

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and Backeberg and Knuth (1936) also cited it as “Peireskia Plum.”

The debate on the correct spelling acquired a political touch in the 1930s in Germany, when the disagreement stirred emotions among German and Austrian cactus specialists. In a brief and concise nomenclatural analysis of the case, Werdermann (1937a) explained that the spelling Pereskia by Plumier (1703) was apparently intentional and correct as based on Miller (1754) and in accordance with the International Code of Botanical Nomencla-ture. In the same year Dölz (1937) published an entirely opposite view on this and simi-lar nomenclatural issues, favoring Peireskia. The highly political article appeared—rather surprisingly—in Friedrich Fedde’s private

and purely taxonomical journal Repertorium Specierum Novarum, edited in Berlin. One can only speculate on the unusual indication of authorship as “Landgerichtsrat Dölz” (Dis-trict court advisor Dölz). Was this the editors’ intention or imposed by the author? Dölz was a lawyer by profession and amateur cac-tologist, president of the German cactus so-ciety from 1934 to 1945 in Berlin, and head of the “Zentralforschungsstelle” (central re-search office) of the German Cactus Society. Fedde’s botanist co-editor Otto Schwarz re-sponded simultaneously in a semi-editorial article (Schwarz 1937). It is a rather uncom-mon case of political opposition expressed in a botanical journal and can only be fully un-derstood in the light of the divergent biogra-phies of the two authors. For political reasons Schwarz had lost his position in a national bi-ological research institute in 1934 with the rise of the “Third Reich” but found tempo-rary employment with the private publisher Friedrich Fedde. Dölz followed and used the official policy until his death at the end of the war in Berlin in 1945 to foment cactological progress in close cooperation with Curt Backe-berg, an activity later depicted as purely ben-eficial by Backeberg (1958: VIII) and Scholz and Stützel (1999).

In an article on the morphology of the flower of Pereskia sacharosa, the Austrian bot-anist Franz Buxbaum, also a member of the

“Zentralforschungsstelle,” followed the nomen-clatural rules but added a “declaration of pro-test” in favor of Peireskia in a footnote (Bux-baum 1940). A formal proposal to conserve

Figure 7. Pereskia portulacifolia in dry forest habitat near Jimaní in the eastern part of the Dominican Republic. Figure 8. Pereskia portulacifolia. Leafy twig at the end of the growing season. This year’s growth with leaves sub-tending the spiny areoles, older stem parts bear some areoles with two or three leaves indicating the beginning brachyblast formation.

Figure 9. Pereskia guamacho. Leafy twig with reddish growing tip and new primary leaves, young, still reddish spines on the areoles. The basal part is of the previous year, with brachyblast leaves formed on the areoles.

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nPeireskia, following Steudel (1841) against Pereskia Miller (1754) was even submitted to the International Botanical Congress by Janchen (1943). The proposal was not success-ful (Lanjouw 1952). Having been confronted with this issue during the preparation of the monograph of Pereskia, Leuenberger (1987) analysed the case in some more detail, repro-ducing Plumier’s first description and the rel-evant comments by Weber (1898a: 163) in support of the original spelling Pereskia. In-dependent of nomenclatural rules, Weber had already concluded that, “respecting Plum-ier and Linnaeus who certainly did not need lessons of Latin,” one ought to maintain the original orthography.

Not surprisingly, the orthographic variants of Pereskia (excepting for Pereskea) persist to some degree, particularly in amateur literature, and a search in the internet demonstrates the current situation (Table 2). Another, entirely new variant “Pereskya” has recently turned up on various websites in Mexico and the Do-minican Republic. The persistent confusion on the correct spelling reaches straight into the newest scientific publications. Even But-terworth and Wallace (2005: 800), presumably by a slip of the pen, repeat the popular myth that Peireskia was the original spelling.

Leaving the rather unfruitful discussions on spelling aside, let us turn to the history of discovery and naming of the species. Among his 21 species of cacti, Linnaeus (1753) distin-guished two species today belonging to Pereskia, namely Cactus pereskia L., and C. portulaci-folius L. This means that in his work, nearly 10% of recognized cacti were pereskias (com-pared to the little more than 1% today). The simple reason for this: pereskias were easier to collect as herbarium specimens than the more highly succulent other cacti, which are nor-mally underrepresented in herbaria.

Although both species had originally been collected by Plumier, the second one (today Pereskia portulacifolia, Figs 7, 8), was not pub-lished by Plumier himself. It was already men-tioned and illustrated in Plumier’s manuscript

“Botanicon Americanum,” of which incom-plete copies were distributed (Hunt 1984). The original set of illustrations of cacti kept in the Natural History Museum in Paris was published by Mottram (2002). Surprisingly, Plumier did not yet relate the small tree with purslane like leaves with his own genus Pereskia. He noted similarities of the flower with roses, but the fruit led him to classify it as an arbo-rescent Opuntia. This plate and the text were also reproduced by Mottram (2002: 116). The

details are remarkable and include a leafy and spiny stem, one staminate and two pistillate flowers, four fruits, and a transverse section of the fruit under the phrase name “Opuntia arborescens spinosissima foliis portulaca corda-tis.” It was apparently not in cultivation and for a long time remained little known. Cac-tus pereskia, subsequently named Pereskia acu-leata, was a more robust plant in cultivation and already widespread as a pot plant in Eu-ropean gardens in the 18th century.

A closer analysis of the bibliographic cita-tions in Linné’s Species plantarum shows that these really contain another, third species, which had in fact been collected and illustrated in the 17th century but was not recognized as dis-tinct from P. aculeata. According to Wijnands (1983) these pre-Linnaean records and illus-trations under phrase names (Plukenet 1692, 1696; Commelin 1697), the latter introduced before 1690/1691 from Isla Margarita, Vene-zuela, show an early introduction of this third species, which Linnaeus (1753) still cited under Cactus pereskia. Remarkably, it took nearly 150 years until this third species, hidden in the Linnaean classification of Cactus, arrived again in Europe and was described as Pereskia guamacho by Weber (1898b). Weber was quite unaware of the much earlier illustrations and the herbarium records of P. guamacho, which are in fact among the earliest still-extant her-barium specimens of a pereskia (two present in Herb. Sloane at the British Museum [Nat-ural History] London). Weber referred to a plant with the local name “Guamacho” from the Orinoco basin in Venezuela but did not cite a herbarium specimen. A neotype speci-men from this area was therefore designated by Leuenberger (1986; Fig 9).

Table 4. “Pereskia” name and images in the WWW found by Google search between 2004 and 2007.

Date

Pereskia WWW entries:

Name

Pereskia WWW entries:Images

16 Jul 2004 5650 2369 Feb 2005 17800 45214 Jul 2005 20700 57628 Dec 2005 52900 51210 May 2006 92100 51818 Oct 2006 85200 52416 Jul 2007 91200 3760*18 Oct 2007 80900 3200*

* Usually only about 10% of search results actually display an image showing a plant of the genus. In July 2007 only ca. 300 of 3760 images showed a pereskia. In October 2007, less than 300 of 3200 images showed a pereskia.


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n In 1812 Haworth named two species, of which Pereskia grandifolia, with its flowers re-sembling a wild rose, is today a well known species in gardens (Figs 10–15). Besides P. acu-leata it is the most widespread species in cul-tivation worldwide and is easily grown from seed, flowering rapidly on cuttings rooted

from flowering branches. The other species of Haworth, P. longispina, an untypified name doubtfully from South America and said to be in cultivation in England before 1808, was treated by de Candolle (1828) with doubts as a variety of P. aculeata, with the comment that it could be an insufficiently known spe-

Figures 10–15. Pereskia grandifolia ssp grandifolia. Figure 10. Leafy shoots of flowering specimen in the display glasshouse of the Berlin-Dahlem Botanic Garden. Figure 11. Pruned stem with new lateral branch-es also subsequently pruned. On the main stem the large pith, weak wood cylinder, and thin green cortex are conspicuous. Figure 12. Flowering branch with terminal passed flower, one open flower, and numerous flow-er buds. Figure 13. One branch of a proliferating inflorescence in lateral view. Figure 14. Infructescence with only distal fruit developed. Figure 15. Longitudinal section of nearly mature fruit, with seeds still attached to placenta, flower remnants partly removed except for the dry pistil standing on the ovary roof surrounded by the bulging receptacle rim. Two seeds in lateral view.

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cies. The scanty description and unknown or-igin does not allow for a clear identification, though the lanose areoles and fleshy leaves could point to Pereskia guamacho. It seems appropriate to leave it under doubtful names (Leuenberger 1986)

Two new species were discovered by Hum-boldt and Bonpland, Pereskia bleo in Colom-bia in April 1801 and P. horrida in Peru in August 1802, published (with this spelling!) as

“Cactus (Pereskea) bleo” and “Cactus (Pereskea) horridus” (Kunth in Humboldt, Bonpland and Kunth 1823). Herbarium material of these spe-cies extant at Paris and Berlin was discussed in detail by Leuenberger (2002, 2004). The two names were formally transferred to Pereskia by de Candolle (1828). Neither species was in cultivation until much later (Figs 16–24, P. bleo, and Figs 25–29, P. horrida).

Four new names were published by de Can-

Currently accepted Pereskia taxa indicating changes since my 1986 monograph.

1. Uupdated numerical list following the informal infrageneric groupings of Leuenberger (1986). Deviating names used by Leuenberger (1986) are added in parentheses.

1. P. aculeata Miller1a. P. aculeata Miller ‘Godseffiana’2. P. horrida DC.2a. P. horrida DC. ssp horrida

( = P. humboldtii Britton & Rose var humboldtii)

2b. P. horrida ssp rauhii (Backeb.) Os-tolaza ( = P. humboldtii var rauhii (Backeb.) Leuenberger)

3. P. diaz-romeroana Cárdenas4. P. weberiana K. Schum.5. P. lychnidiflora DC.6. P. aureiflora F. Ritter7. P. guamacho F. A. C. Weber8. P. zinniiflora DC.

8′. P. marcanoi Areces (1992)9. P. portulacifolia (L.) DC.10. P. quisqueyana Liogier11. P. bleo (Kunth) DC.12. P. nemorosa Rojas Acosta13. P. sacharosa Griseb.14. P. grandifolia Haw.14a. P. grandifolia Haw. ssp grandifolia

( = P. grandifolia Haw. var grandifolia)14b. P. grandifolia Haw. ssp viola-

cea (Leuenb.) N. P. Taylor & Zappi ( = P. grandifolia Haw. var violacea Leuenb.)

15. P. bahiensis Gürke16. P. stenantha F. Ritter

2. Updated alphabetical list of currently accepted names. Deviating names used by Leuenberger (1986) are added in parentheses.

P. aculeata MillerP. aculeata Miller ‘Godseffiana’P. aureiflora F. RitterP. bahiensis GürkeP. bleo (Kunth) DC.P. diaz-romeroana CárdenasP. grandifolia Haw. ssp grandifolia

( = P. grandifolia Haw. var grandifolia)P. grandifolia Haw. ssp violacea (Leuenb.)

N. P. Taylor & Zappi ( = P. grandifolia Haw. var violacea Leuenb.)

P. guamacho F. A. C. WeberP. horrida DC.P. horrida DC. ssp horrida ( = P. humboldtii

Britton & Rose var humboldtii)

P. horrida ssp rauhii (Backeb.) Ostolaza ( = P. humboldtii var rauhii (Backeb.) Leuenb.)

P. lychnidiflora DC.P. marcanoi Areces

( = P. sp A of Leuenberger 1986)P. nemorosa Rojas AcostaP. portulacifolia (L.) DC.P. quisqueyana LiogierP. sacharosa Griseb.P. stenantha F. RitterP. weberiana K. SchumannP. zinniiflora DC.

3. Frequent synonyms of Pereskia species with cross-reference to accepted names.

Pereskia aculeata var godseffiana = P. aculeata ‘Godseffiana’

Pereskia colombiana = P. guamachoPereskia corrugata = P. bleo

Pereskia cubensis = P. zinniifloraPereskia godseffiana = P. aculeata ‘Godseffiana’Rhodocactus antonianus = Pereskia weberianaRhodocactus cubensis = Pereskia zinniiflora


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n dolle (1829), solely from the plates of Moci-ño’s unpublished Flora Mexicana. The dra-matic fate of these plates has been elucidated by McVaugh (1980, 1982). The species de-scribed there have long been assumed to be from Mexico, whereas all but one were re-ally from other countries, including Nicara-gua (Pereskia opuntiiflora, P. lychnidiflora) and Cuba (P. zinniiflora), as explained by Leuen-berger (1986, 1988). The first two represent Pereskia lychnidiflora, a species ranging from western Mexico to the Pacific coast of Costa Rica (Figs 1, 30–34). Pereskia zinniiflora is the oldest name for the endemic species of Cuba (Figs 35–40). The fourth species, P. rotundi-folia, is a leaf bearing cactus from Mexico, but due to the glochids on its areoles (visible in the drawing) it belongs to a different genus Pereskiopsis. The Sessé and Mociño plates are now kept at the Hunt Institute for Botani-cal Documentation at Pittsburgh. The cac-tus plates were reproduced in color with com-ments by Rowley (1994).

It has to be taken into account that the early treatments of Pereskia did not exclusively con-tain plants today belonging to the genus. Pfei-ffer (1837a, 1837b), Mittler (1844), Förster (1846), Salm-Dyck (1850), and Rümpler (1886) still included several leaf-bearing species in Pereskia that were excluded from the genus by later monographers. Some were excluded temporarily and later recognized as belonging to the genus again, for instance, Pereskia pitit-ache Karw. ex Pfeiffer (1837a) from Mexico, for a long time placed in Pereskiopsis following Britton and Rose (1919), was reinstated as a true Pereskia by Boke (1963a). It was actually a re-description of P. lychnidiflora.

Early illustrations of Pereskia in cultiva-tion have caused some confusion. Up to 1850 probably only three species were really in cul-tivation in Europe: Pereskia aculeata, P. gran-difolia, and P. lychnidiflora, the latter under the name P. pititache, of which a seedling was illustrated by Zuccarini (1837). This plant does not seem to have established in cultiva-tion, which is not surprising. Our experience over 25 years with this species in the botan-ical garden calls for very high summer tem-peratures and hothouse conditions in win-ter, necessary for successful cultivation from seed to flowering specimens (Figs 30–34). The early records of P. bleo in cultivation are probably all incorrect, and Britton and Rose (1919) noted that all previous illustration in fact showed P. grandifolia, including that by Schumann 1890 in the Flora brasiliensis. Brit-ton and Rose (1919) and Weber (1898b) re-

solved the case, but the confusion persisted in many collections. Pereskia bleo was pro-vided with two additional names, now listed in synonymy. One is, remarkably, by Weber (1898b) himself, Pereskia panamensis, due to erroneous information on flower color. The other is P. corrugata by Cutak (1951), clearly a re-description of P. bleo, caused by misla-belled plants of P. grandifolia in private and public collections.

In the second half of the 19th century, new species were described mainly from south-ern South America. The first was Pereskia sa-charosa Grisebach (1879) from the western Chaco region, named for its rose-like flowers. Its vernacular name, sacha rosa, means either “false rose” or “wild rose” (Figs 41–44). The name is often misspelt but has nothing to do with sugar. As pointed out by Leuenberger (1986), the species had in fact been collected more than 100 years earlier (between 1747 and 1755) in Bolivia by Joseph de Jussieu, but the specimen in the Jussieu herbarium at Paris had remained there under the name Cac-tus pereskia unnoticed, though already men-tioned as dried specimen (from “Perou”) under Pereskia by Jussieu (1789). Pereskia sacharosa is one of the hardier species, as also reported by Mauseth (2001), who found this species and P. aculeata to endure low temperatures. A closely related species, sometimes still confused with P. sacharosa, is P. nemorosa Rojas Acosta (1897), which has a more eastern distribution in the Chaco region (Figs 45–48).

Only one year later P. nemorosa was given two more names, P. amapola and P. argentina, by Weber (1898b), who was unaware of the publication of Rojas Acosta.

The modern circumscription of the genus Pereskia dates back to Schumann (1898), who used primarily seed and spine characters rather than leaf characters to classify the subfamilies. He placed Pereskia (flat leaves) and Maihuenia (terete leaves), both without glochids, in sub-family Pereskioideae. The subfamily is often attributed to Schumann, but according to Re-veal (2003) and Thorne and Reveal (2007) it had already been formally established by En-gelmann in 1876. Regardless of leaf shape, Schumann placed all species with glochids and having seeds provided with bony aril in subfamily Opuntioideae, with the only genus Opuntia. Schumann (1898) described only one new species, based on a single herbarium specimen from Bolivia. He dedicated Pereskia weberiana to his contemporary, the French (Alsacian born) medical doctor FAC Weber, author of the Cactaceae entries in the French


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nDictionary of Horticulture by Bois (1893–1899) and other papers on cacti (Figs 49–50). Weber (1898a) described Pereskia tampicana based on a photograph and live plant in the Berlin Botanical Garden, but this was clearly a re-discription of P. grandifolia, as already recognized by Britton and Rose (1919). It is the first record of this Brazilian species grow-ing as a neophyte in Mexico.

In the 20th century, Berlin botanist Gürke (1908) published Pereskia bahiensis based on an herbarium collection of Ule. It remained for a long time a little-known species and is curiously missing from the Index Kewensis (Figs 51–54). Britton and Rose (1912, 1919, 1923) added five supposedly new species to Pereskia, but four are now considered synonyms of earlier names, and one was removed from the genus. Pereskia colombiana is the same as P. guamacho from Venezuela. P. cubensis is a synonym of P. zinniiflora. Pereskia zehntneri was later placed in Quiabentia by its authors. Pereskia conzattii is the same as P. lychnidiflora, and this species, ranging over several coun-tries from Mexico to Central America, was provided with further names by other authors at the beginning of the 20th century. It was described as P. nicoyana F. A. C. Weber from Costa Rica and P. autumnalis Eichlam from Guatemala. Pereskia moorei Britton & Rose

(1919) from the Matto Grosso is now consid-ered a synonym of P. sacharosa, a species with a wide distribution along the eastern margin of the Andes from Bolivia to northern Argen-tina and reaching into Brazil and Paraguay. Britton and Rose (1923) coined a new name (P. humboldtii) for Pereskia horrida (Kunth) DC, on the assumption that the latter was il-legitimate. The name Pereskia humboldtii was used by Leuenberger (1986), but following the correction by Brako and Zarucchi (1993), P. horrida DC (considered as new name, not combination) has been generally accepted in accordance with the ICBN.

Britton and Rose (1919) reduced the sub-families to the rank of tribes. Two new genera, Pereskiopsis and Quiabentia, were created by them to accommodate some former Pereskia species with glochids and bony seeds (Britton and Rose 1919, 1923).

Berger (1926, 1929) had a phylogenetic approach to taxonomy, discussing evolution-ary lines and levels within the genus based on characters of the ovary. He placed Pereskia aculeata and P. sacharosa in Pereskia subge-nus Pereskia (as “Eupeireskia”) and all others in a newly proposed subgenus, Rhodocactus. In morphological terms, his distinction of ovules at the base of the pistil versus ovules in a true locule (interpreted as superior ver-

Table 5. Names and images of selected Pereskia species found in the WWW (including major synonyms and frequent orthographic variants).

Name, synonym or orthographic variant (“nn”)

WWWentries

20.5.2005

WWWimages

20.5.2005

WWWentries

19.10.2006

WWWimages

19.10.2006

WWWentries

18.10.2007

WWWimages

18.10.2007Pereskia aculeata 788 87 22500 121 19600 13601

Pereskia aureiflora 62 0 280 0 277 881,2

Pereskia grandifolia 696 74 860 140 860 4531

Rhodocactus grandifolius 59 2 185 10 268 551

“Pereskia grandiflora” 201 25 830 41 1520* 105“Rhodocactus grandiflorus” 8 0 28 0 38 821

Pereskia nemorosa 57 0 330 2 332 1551,2

“Pereskia nemerosa” 2 0 4 0 6 0Pereskia sacharosa 337 17 1040 22 1180 991

Rhodocactus sacharosus 10 0 2 0“Pereskia saccharosa” 51 3 286 3 61 101

“Peireskia sacharosa” 12 0 41 0 35 31,2

“Peireskia saccharosa” 0 0 0 0 2 0Pereskia zinniiflora 61 5 228 4 198 1591

Rhodocactus zinniiflorus 10 0 9 0 7 0Pereskia cubensis 55 1 182 1 120 151

Rhodocactus cubensis 34 3 51 2 47 1411,3

1 Usually only about 10% of the search results actually display an image showing a plant of the species; 2 None correct; 3 Only one is Pereskia.


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n sus inferior ovary) is rather questionable and not supported by later studies (for instance, Boke 1964). But Berger’s cladistical-looking

graph (Berger 1926: 4, scheme 1) can be in-terpreted as an informal proposal of differ-ent genera, although he maintained a single

Figures 16–24. Pereskia bleo. Figure 16. Vigorous growth of a new shoot, a leafy and spiny stem grown in the same year is visible in the background. Figure 17. Strong main shoot several months old, large leaves subtending the spiny areoles (on such shoots a maximum leaf length of 42 cm was observed in cultivation). Figure 18. Older main stem after the leaves have been shed, with areoles producing new spines every year. Figure 19. Shoot with three secondary branches (two with flowers), developed from the uppermost leaf axils of the terminal inflores-cence (remnants hidden by the leaves). Figure 20. Flowering branch developed laterally from near the tip of the primary flowering branch (pedicel scars of fallen flowers still visible). Figure 21. Terminal flower in postfloral stage and three flower buds in the axils of the uppermost leaves. Figure 22. Longitudinal section of closed flow-er exposing the inferior ovary, flat ovary roof, bulging filament bases over the nectary, erect stamens, and pistil. Figure 23. Young fruit in lateral position in leaf axil of a few-flowered inflorescence exposing pedicel scar of a lateral flower. Figure 24. Twig with mature fruit. The twig at right without fruit but showing the conspicuous pedicel scars of fallen flowers.

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ngenus Pereskia and formally only established a division into two subgenera (Berger 1929), treating two names (P. aculeata and P. sa-charosa) under subgenus Pereskia, and four names (Pereskia bleo, P. grandifolia, P. autum-nalis, and P. conzattii) under subgenus Rho-docactus. Edwards and others (2005) refer to Berger’s idea, but Berger’s informal Rhodocac-tus was actually poorly defined and is hetero-geneous in terms of Edward’s clades, due to the placement of P. grandifolia.

The most detailed anatomical investiga-tions, addressing evolutionary theories as well, are by Boke (1963b, 1964), who presented a new interpretation of the cactus gynoecium, explaining the relative position of carpellary tissue and receptacular tissue in ontogenetic development. Berger’s interpretations were later opposed by Ritter (1979: 20), based on examination of longitudinal sections of flow-ers of several South American species. Rit-ter found the ovary in P. aculeata, P. sacha-rosa and the Andean species, compared to P. bahiensis and P. stenantha of the P. gran-difolia group representing Berger’s subgenera Pereskia and Rhodocactus, to be neither su-perior nor inferior but intermediate, and he noted transitional stages, making a clear dis-tinction impossible.

Formally it was Knuth, in Backeberg and Knuth (1936: 48, 96), who elevated Berg-er’s subgenus Rhodocactus to the rank of genus, recombining some of the names, ap-parently based on supposed, rather than ob-served, characters of the gynoecium. Backe-berg (1958: 105, 113) only repeated Bergers’s illustrations, and his reclassification of some species is neither based on investigations of the flowers. This becomes significant be-cause Backeberg (1958) selected Rhodocac-tus grandifolius as type of the genus, but in fact only the Caribbean species and P. bleo have clearly epigynous flowers (Leuenberger 1986). Backeberg (1957) made further sub-divisions and classified the remaining species of Pereskia into two subgenera by creating a subgenus Neopeireskia for the Andean spe-cies with small leaves. Later authors ques-tioned the justification of the generic status of Rhodocactus and other subdivisions and upheld only one genus, Pereskia (Hunt 1967; Bravo-Hollis 1978; Ritter 1979).

In the same decades, some remarkable dis-coveries of new species were made by Cárde-nas (1950, 1964), Ritter (1979), and Liogier (1980), and the number of published names increased to about 50 by 1979. The names still accepted today, as well as some of the

names subsequently reduced to synonymy, are briefly discussed.

Pereskia diaz-romeroana Cárdenas (1950) is now sometimes written “diazromeroana” in newer literature (and, for instance, in TROP-ICOS), but as the epithet commemorates the Bolivian naturalist Belisario Díaz Romero, the original spelling is appropriate (Fig 55). From the same group of Andean species, all occurring in isolated valleys, a new species and two new varieties were described from Peru: Pereskia vargasii H. W. Johnson (1952), P. vargasii var longispina Rauh & Backeb., and P. vargasii var rauhii Backeb. (in Backeberg 1957). They were illustrated by Rauh (1958) and are now considered to belong to the two subspecies of Pereskia horrida.

Pereskia higuerana (Cárdenas 1964) is a cu-rious case, because from the description, men-tioning detachable spines in the axils, and from the illustration it is clearly a Pereski-opsis, a mainly Mexican genus not native in Bolivia (but often cultivated in cactus collec-tions and in the past much used as grafting stock for seedlings). The cited type and iso-type specimens of P. higuerana, collected near La Higuera in “April 1954” by D Candia, a student of Cárdenas, were never located, and this yellow flowered plant remains enigmatic. Some doubts prevail if this might have been an error or even a hoax based on a plant from cultivation. This hypothesis has some argument based on the existence of a plant identified as Pereskiopsis by Leuenberger (1986), received as Pereskia higuerana from the Schönbrunn collection in Vienna and said to have origi-nated from Cárdenas from Bolivia. On the other hand it is perhaps significant that Ritter (1980) mentioned and illustrated (though in black and white) a yellow flowering (unnamed) variety of Pereskia diaz-romeroana from Bolivia, which has not been reported again and is nei-ther found among the material of FR 904 or other numbers (Eggli and others 1996). Kies-ling (1999) mentions that he found P. diaz-romeroana near La Higuera.

The most remarkable novelties, Pereskia aureiflora and P. stenantha, based on Ritter’s field research made in 1964 in eastern Brazil, were published by him 1979. The two spe-cies were apparently not in cultivation until 1983 (Leuenberger 1989a, b). Pereskia aure-iflora from Minas Gerais is the only purely yellow-flowered species besides P. guamacho (Figs 56, 57). Pereskia stenantha is a remark-able case of flower evolution (Figs 58–60). It is evidently the sister species of P. bahiensis, but it has a remarkable adaptation in its hum-


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n mingbird flowers (Ritter 1979; Leuenberger 1989a). The upright tepals form a flower of tubular shape (but not a receptacular tube like that of other Cactaceae). Another feature is its conspicuous nectar chamber. This species is from the interior of Bahia (Ritter 1979). Col-lection details were given by Eggli and oth-ers (1996), who also confirmed that the type specimens of both names are missing. Only seed samples could be located.

One year later, Ritter (1980) published P. sparsiflora, a plant introduced to cultiva-tion in Europe by Ritter’s 1958 and 1959 collections from Bolivia. The name is clearly a synonym of P. sacharosa, based on FR 640 material still in cultivation, but Ritter’s field number was found to be heterogeneous and also included plants from Paraguay belong-ing rather to P. nemorosa according to Leuen-berger (1986). (For an explanation of Ritter’s complex field number system see also Eggli and others 1996).

A most remarkable discovery was Pereskia quisqueyana, a narrowly endemic, dioecious spe-

cies of the P. portulacifolia alliance, described by Liogier (1980) from a clone of staminate plants at Bayahibe near La Romana, today a well-known holiday resort on the south coast of the Dominican Republic (Figs 61–66).

Looking at the taxonomic status of Pereskia prior to my 1986 revision, the number of Pereskia species published over different eras merits comment. The early increase of species numbers in Pereskia until Britton and Rose (1919) is notable as compared to the situation within the family as a whole (Tables 1, 3). The greatest increase in species numbers in Cacta-ceae falls into an era of collectors and authors with commercial interests. It took place between 1925 and 1966, in particular due to publish-ing activities of Curt Backeberg in Germany. In contrast, the leafy cacti were never particularly attractive for amateurs and no special increase of names in Pereskia can be related with com-mercial activities. Backeberg (1963) published Rhodocactus antonianus based on a plant of un-known origin in his own collection, an invalid name for lack of a preserved type (Eggli 1985;

Figures 25–29. Pereskia horrida ssp horrida. Figure 25. New flowering branches with terminal and lateral flowers. Figure 26. Thicket of leafless but flowering older branches. Figure 27. Young leafy twig with long hairs on areoles, green epidermis with stomata, and transverse section of stem showing comparatively large succulent pith and cortex. Figure 28. Twig with terminal flower, flowers, young fruit and flower l. s. Figure 29. Leafy twig with mature fruit and flower bud with hairy areoles (Photo C Hillmann-Huber BGBM).

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nEggli and Taylor 1991). He later modified the vague ori-gin data “Northeastern Peru” (Backeberg 1966), but the taxon is undoubtedly iden-tical with Pereskia weberiana, a species from Bolivia. Taxo-nomically, the cactus family was expanded by Britton and Rose with over 900 new names (Table 3). As noted by Mitich (1979), Britton was conserva-tive in delimiting species but tended to separate genera liber-ally. In the case of Pereskia, the situation is rather the reverse, since Britton and Rose (1919) published five new Pereskia species. The effect of Britton and Rose, and subsequently of Backeberg, on names in Cac-taceae was characterized by Hunt (1991). The dominant splitter in Cactaceae—both in genera and species—was Backeberg, with over 1400 species, but he published only one new species and one vari-ety of leafy cacti, both under the generic name Rhodocactus. Ritter contributed names for over 900 taxa of cacti, among them three new species of Pereskia. Many of these names are now considered synonyms, but many still remain insuffi-ciently known.

My Pereskia monograph (Leuenberger 1986) recog-nized 16 ( + 1 unnamed) species, based on 450 her-barium specimens from 50 herbaria, field studies of 8 species (Mexico, Domini-can Republic, Brazil), and the study of live material of all species cultivated at the Berlin-Dahlem Botanic Gar-den between 1976 and 1986 (75 accessions, the majority still extant). In it, based on my own field studies in 1983 in Minas Gerais and Bahia, I described a plant already observed in cul-tivation and originally introduced by Horst and Uebelmann from more than one locality under number HU226 as new variety Pereskia grandifolia var violaca (Figs 67–70).

In the past 20 years only one new species of Pereskia has been published: Pereskia mar-canoi Areces (1992) from the Dominican Republic, a taxon closely related to Pereskia zinniiflora and which I had already recog-

Figures 30–24. Pereskia lychnidiflora. Figure 30. Shoot apex with whorl of branches and some brachyblast leaves. Figure 31. Longitudinal section of stem of lateral branch less than one year old, with succulent green pith and cortex, leaves subtending areoles, mature spine with still-fleshy base. Figure 32. Flowering twig, the lower part one year old with brachyblast leaves, the new leafy growth (with reddish colored epidermis) terminating in a flower with green fleshy outer tepals and orange, thin, fimbriate inner tepals. The pistil is concealed by the numerous stamens. Figure 33. Longitudinal section of flower showing gynoecium, elongated flower axis, large mucilage cells in pith and cortex as well as in the base of the outer tepals. Figure 34. Longitudinal section of withered flower after a few days, with receptacle, outer tepals, and ovary roof still fleshy, other flower parts including the ovules, pistil, stamens, and outer tepals shriv-elled or dry.

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n nized as probably new (Leuenberger 1986), treating it provisionally under “Pereskia sp A,” for lack of sufficient data. A small plant observed in cultivation over many years in Berlin-Dahlem had flowered only once in 1984, producing a pistillate flower. It flow-ered again in 1993 and 1998 and once pro-duced a fruit with some seeds germinating within the fruit (Figs 71–72). Remarkably, an old but sterile herbarium material of a plant cultivated at the Jardin des Plantes, proba-bly of the same taxon, exists in the AL Jus-sieu Herbarium (13546A) and was cited by Leuenberger (1986). Unfortunately it bears no further information and can only be vaguely dated between ca. 1789 and 1850. No further hints were found in the informa-tion on the Jussieu herbarium provided by Stafleu (1964). Perhaps molecular methods will some day permit us to confirm its geo-graphical origin and identification.

Since the publication of my 1986 mono-graph only a few changes have occurred. The taxonomic circumscription of the genus re-mained stable, and Pereskia is one of the gen-era of Cactaceae with the highest stability of names in the past 20 years (see also Hunt and

others 2006: 5). Some nomenclatural adjust-ments and some status modifications have been made. As noted, the most significant change affecting a formerly well-established name is a nomenclatural correction, initiated by Brako and Zarucchi (1993): the replacement of the well established name Pereskia humboldtii Britton & Rose (1919) by the available older name Pereskia horrida DC in accordance with the ICBN (Art. 58.1 Ex0.1), where an analo-gous example is specifically mentioned (Greuter and others 1994; McNeill and others 2006). Following earlier practice common in Cacta-ceae, particularly in American botanical liter-ature, I treated infraspecific taxa of P. grandi-folia and P. horrida (under P. humboldtii) as varieties (Leuenberger 1986). In the past de-cades the treatment as subspecies has become more common and promoted (Hunt and oth-ers 2006: 4). Further data on the geographi-cal distribution led to the new combination Pereskia grandifolia ssp violacea (Leuenb.) N. P. Taylor & Zappi (1997). Ostolaza (1998) formally gave P. horrida ssp rauhii this new status, but data on this taxon are still scarce. In current standard works on Cactaceae (An-derson 2001, 2005; Hunt and others 2006)

Figures 35–40. Pereskia zinniiflora. Figure 35. Older pruned branches exposing thick wood cylinder, and flowering twig, only the green, leafy part of this season’s growth, with staminate flower. Figure 36. Young branch at the end of the growing season, with primary leaves and spiny areoles, transverse section showing massive wood cylinder compared to pith and cortex. Figure 37. Several-year-old branch with numerous old stumps of flow-ering branchlets and one flowering branchlet with bud of staminate flower. Figure 38. Twig with bud of pistil-late flower, with conspicuous larger receptacle (pericarpel). Figure 39. Several year old branch with numerous old and dead flowering branchlet stumps and one open staminate flower, flower buds, and withered flower. Figure 40. Staminate flower (left), and pistillate flower (right) with incomplete tepals.

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nthe 16 + 1 species of Pereskia distinguished by me in 1986 are accepted as 17 species, with the same infraspecific taxa and with the up-dated nomenclature outlined here.

It should be noted that some illustrations in these standard works bear wrong captions. In Anderson (2001), Pereskia aculeata on p 566 is obviously P. grandifolia ssp grandifolia, while P. grandifolia ssp grandifolia on p 568 is really ssp violacea. In the German edition of the same book (Anderson 2005), edited and comple-mented by U Eggli, the eleven illustrations of Pereskia are all correctly identified. The New Cactus Lexicon (Hunt and others 2006) illus-trates all 17 species, but unfortunately one photograph has the wrong caption: Figure 451.2 (captioned P. diaz-romeroana), is cer-tainly P. weberiana (the clone formerly known in cultivation as Rhodocactus antonianus).

A number of papers contributing data on

various aspects of Pereskia have been pub-lished in the past 20 years. Only few can be mentioned here. Much useful information on various aspects of morphology, anatomy, and biology can be found in Gibson and Nobel (1986). Stem and leaf anatomy was treated by Sajeva and Mauseth (1991) and Maus-eth and Landrum (1997), and leaf anatomy of four northern South American and Carib-bean species by García and others (2000). In regional studies, the most significant recent work including a detailed treatment of five species of Pereskia is a book on cacti of east-ern Brazil by Taylor and Zappi (2004). This includes numerous new localities documented by vouchers and cited in detail, distribution maps, photographic illustrations, and a con-servation assessment of each species. Studies on individual species treated the distribution biology of P. aculeata in Brazil (Pedroni and

Figures 41–44. Pereskia sacharosa. Figure 41. Grown as a hedge in the display glasshouse of the Berlin-Dahlem Botanic Garden. The cactus on the right, planted as a fence, is Pachycereus marginatus Figure 42. Young developing shoot with new spine growth. Figure 43. Pruned branch with new side branch developing from an areole of this year’s growth (still subtended by a primary leaf ). The older part of the branch is more than one year old and shows brachyblast leaves formed between the spines. Figure 44. Flowering twig with spineless areoles and brachyblast leaves. Flower fully open.

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Sanchez 1997), invasive potential of P. acu-leata in South Africa (Moran and Zimmer-mann 1991), conservation aspects and new re-cords of the narrow endemic and endangered species of the Dominican Republic, Pereskia quisqueyana (García and others 1999; Mejía and others 2001; Corman 2003).

Doweld (2004) proposed and named new infrageneric groups resulting in three sec-tions, two subsections, and five series, based on the informal groups outlined by Leuen-berger (1986). These named entities are of questionable purpose. The hierarchy does not agree with the cladograms of Nyffeler (2002), Butterworth and Wallace (2005), and the new entities thus seem to be little more than a pro-spective name game. The groups do not agree with the ideas of Edwards and others (2005) of two main clades.

Pereskia in the WWW (2004–2007)Since about 2004 numerous websites world-wide have provided scattered information on, and an ever increasing number of images of, Pereskia. The rapid increase is shown in the results of a casual Google search at irregular intervals (Table 4). The increase of images (or rather image links) found from summer 2006 onward is striking, but the actual number of images displaying the species searched for is much smaller. In this genus and for many other plants often only about 10% of images are of the searched-for species. Before this date, the percentage of images actually showing the plant was also variable, and often also lower, but details were not recorded.

In any case, any search result must be in-terpreted with caution and some knowledge on the group, because search engines cannot

Figures 45–48. Pereskia nemorosa. Figure 45. Pruned stem with new spines, new lateral branches emerging from the uppermost areoles, main stem exposing weak wood cylinder, large pith and thin green cortex. Epider-mis without stomata but with late periderm formation. Figure 46. Older stem with striate periderm formation, transverse section of wood cylinder, and large pith. Figure 47. Flowering branch with passed terminal flower, one opening flower, and two flower buds with characteristic long hairs in the axils of the receptacular bracts. Figure 48. Flowering twig with young terminal fruit and lateral flower arising from receptacle

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interpret the content of the image. Even when the image is clearly connected with the correct caption, the identification can still be wrong. This is more often the case than one might ex-pect, and not only for plants with unsettled taxonomy. The problem is naturally widespread in taxonomic problem groups, when numer-ous images are put on the WWW by non-ex-perts. The search results could not be analysed in detail, but there seems to be a tendency to-ward an increase in correctly identified images, possibly by some sort of self-cleaning effect stimulated by well-identified plants on reli-able websites. However, in some cases there is a proportional increase of wrongly named material (Tables 4, 5).

Selected updates to the 1986 monograph on Pereskia

Pereskia aculeata. Since 1986 numerous new records from within the previously known range of P. aculeata have been added by col-lectors, particularly in Paraguay and NE Ar-gentina (see TROPICOS) and also from Bra-zil (Taylor and Zappi 2004). In 1996 Pereskia aculeata was reported for the first time from the lower Rio Grande valley in Texas (Ideker 1996). Pereskia aculeata had earlier been re-ported from continental US only from Florida (Benson 1982). More recently it has arrived and in 2001 been recorded as a potentially invasive species in Hawaii according to Staples (pers. comm. Oct 2001) and Hawaiian Ecosystems at Risk project (HEAR; Anonymous 2007). The species is invasive in various regions and was declared a weed in 1979 in southern Af-rica, where it has been well studied (Moran and

Zimmermann 1991; Henderson 1995). New herbarium re-cords from Peru and Bolivia need criti-cal evaluation and verification in hab-itat, as the species is also often culti-vated, and escapes are possible.

Pereskia bleo. Maximum leaf length was given by Britton and Rose (1919) as 19 cm, and by Leuenberger (1986) as 26 cm based on herbar-ium specimens and live material. New

measurements are documented by a garden herbarium specimen (Cubr 38344 at B) col-lected from a specimen planted in a display glasshouse under hot and humid conditions, where the plant developed on a vigorous long shoot in summer 2000. A main shoot reached about 3 m height in one summer season, and the leaves attained a maximum length of 42 cm, certainly an impressive record for a cactus (Fig 73).

Pereskia diaz-romeroana. In 2003, holo-type material of this species was located in the Cárdenas collection of Cactaceae deposited in the herbarium of the Instituto Miguel Lillo, Tucumán, Argentina (LIL), not accessible at the time of the monograph but now fully cata-logued by Eggli and Leuenberger (2005). The holotype specimen, Cárdenas 4388, located there supersedes the lectotype (a duplicate at US) I had selected (Leuenberger 1986).

Pereskia guamacho. At the time of my-monograph’s publication, I had no informa-tion on the underground part of Pereskia gua-macho. The only specimen available to me, at Berlin-Dahlem, did not show any particular features. Only recently could a root tuber be observed on a seed-grown specimen. Obser-vations made in Venezuela, documented by photographs kindly provided by NP Taylor several years ago, show that this species pro-duces very large root tubers developed prob-ably by the primary root (Fig 74). These can attain a length of more than one meter and are about 15 cm in diameter on a still-young tree observed with its root system exposed on a roadside cut. Such a massive water stor-ing root system could be ecologically impor-

Figures 49, 50. Pereskia weberiana. Figure 49. Leafy twig with terminal flower bud. Figure 50. Transverse section of young stem, showing conspicuous green cortex, epidermis with stomata.

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n Figure 51. Pereskia bahiensis planted as a hedge in Bahia, Brazil. Figure 52. Pereskia bahiensis flower in culti-vation (Braun 534). Figure 53. Pereskia bahiensis, stem of plant in bonsai-like growth, with brachyblast leaves formed on areoles and on a condensed lateral shoot several years old. Figure 54. Pereskia bahiensis stem of plant in bonsai-like growth with brachyblast leaves formed on the areoles. The short lateral shoot several years old and very condensed. Figure 55. Pereskia diaz-romeroana flower surrounded by long hairs of flowering areoles. Figure 56. Pereskia aureiflora flowering in cultivation. Figure 57. Pereskia aureiflora. Older twig with brachy-blast leaves formed on the areoles. Figure 58. Pereskia stenantha inflorescence with flower bud, open flower and passed flower, showing the fleshy erect outer tepals characteristic of a hummingbird flower. The flowers of this clone (Leuenberger 3081) barely open at the tip. Figure 59. Pereskia stenantha. Longitudinal section of termi-nal, passed flower with wilted inner floral parts and axillary flower in late flowering stage. The intermediate posi-tion of the ovary and the comparatively large nectary zone inside the rim of the receptacular cup are conspicuous in both stages. Figure 60. Pereskia stenantha. Longitudinal section of flower with fleshy outer and inner tepals, intermediate position of ovary in receptacular cup, placentae, and conspicuous nectary.

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ntant. This massive underground storage sys-tem was not mentioned in the recent studies by Edwards (2006) and Edwards and Diaz (2006), who noted that leaf production is not so clearly related to drought and rain cycles as hitherto suggested by data extracted from herbarium labels. The phenology of this spe-cies may deserve further examination, taking into account the underground water reservoir of the trees. In cultivation, unfortunately, no flowers could be observed hitherto, even after various experiments (though only with one single larger specimen) to stimulate flowering. Prolonged drought in summer did not give any result. It is possible that this species needs a drought period with high temperatures, pos-sibly combined with shorter daylight hours, to stimulate flowering—impossible to achieve under our conditions. Even though herbarium data (some 40 samples studied) suggest that flowering occurs mainly from March to May, normally while plants are in a leafless stage or with their first flush of brachyblast leaves, and

that fruit development occurs when the plants are in leaf, the species has also been observed with flowers on leafy shoots during the rainy season, as documented by a photograph taken by Jason Grant in Venezuela in 1998.

Pereskia zinniiflora. A special feature of this species is the presence of large root tu-bers, first observed late in 1994. In his paper dealing in detail with the morphological char-acters of the species, Areces (1984) did not yet mention the roots of P. zinniiflora. I had found only fibrous roots in juvenile plants of Pereskia zinniiflora (Leuenberger 1986) and could not study this species in the field. But I did note the possible presence of root tubers with a question mark in Table 1 on p 36 of the monograph. The diagram illustrating evolu-tionary groups within the genus already sug-gested the possible occurrence of root tubers in this taxon. Areces, in a later publication, used the character “root tubers absent” in the key to distinguish P. zinniiflora and P. marcanoi from P. quisqueyana (Areces 1992).

Figures 61–66. Pereskia quisqueyana. Figure 61. in habitat on coral limestone at the type locality in Bay-hahibe, La Romana, Dominican Republic. Figure 62. Branch about one year old with persistent primary leaves. Figure 63. Young stem with leaves subtending spiny areoles. Early stage of periderm/cork formation. Figure 64. Older, leafless stem with continuing formation of new spines. Figure 65. Succulent root tuber of small potted plant. Figure 66. Staminate flower in cultivation.

61 62 63

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The presence of root tubers could be con-firmed for P. zinniiflora on a plant cultivated in the Botanical Garden Berlin-Dahlem. This specimen, originally received from the Jar-din de Plantes in Paris under number G.563, leg. Alain, originally collected in Cuba, and first grown in a pot, was planted in the bro-meliad house under hothouse conditions. When it had to be removed in 1994, it was found that it had produced three large root tubers of remarkable size, the largest fusiform and 60 cm long and to 14 cm in diameter. The main stem of this plant, fiercely spiny and only about 3 cm in diameter, is illus-trated together with one of the three root tu-bers three years (!) after uprooting (Fig 75). In September 1997 the slightly shrivelled root tuber was still fleshy inside, as demon-strated by a small piece cut from the tuber. Kept on a windowsill at room temperature in the laboratory, it took almost two addi-tional years to dry out completely. Although formation of new root tips was observed at the distal end of the rood tuber while it was kept dry, no shoot regeneration occurred. This agrees with an earlier observation on detached root tubers of Pereskia weberiana (Fig 76). These root tubers are apparently unable to produce buds and new stems even if planted in soil, as already noted (Leuen-berger 1986, 1992d).

Taxonomic history of Maihuenia

The genus Maihuenia was proposed by Philippi (1883) and formally described by Schumann (1898). The first reports of plants belonging to this genus date back much earlier. Due to their superficial similarity with some sym-patric, cushion-forming opuntias, they were first described as species of Opuntia. The taxonomic history is therefore more com-plex than in Pereskia. Remarkably, maihue-nias were not yet collected or illustrated by the very first naturalists surrounding south-ern South America, perhaps due to lack of flowers at the time, or because they had a bad reputation caused by the first contacts with opuntias. One sympatric cactus was recorded before any maihuenia from east-ern Patagonia in today’s Province of Santa Cruz, Argentina, but remained undescribed. It was found on the Malaspina expedition in 1789. According to the illustration by José Guio based on a drawing by Née, an-notated only as “Cactus” (Anonymous 1989: 168 and pl. 14), it was Austrocactus bertinii. Another cactus, found after but described before the first maihuenia, was collected on Charles Darwin’s journey in 1833 on the eastern coast of Patagonia (Port Desire, today Puerto Deseado, Santa Cruz, Argen-tina) and described as Opuntia darwinii by

Figures 67–70. Pereskia grandifolia ssp violacea. Figure 67. Branch with dark foliage, young inflorescence; flower buds with conspicuous purplish pink bracts. Figure 68. Inflorescence with open terminal flower and flower buds. Figure 69. Flower and flower bud of a different clone (also received as HU226) with broader tepals and deeper flower color but the same flower bud characters. Figure 70. Branched infructescence/inflorescence of the same, showing a terminal fruit with repeated branching from the receptacle, on lower branch producing a lateral fruit, receptacles of additional flowers without fruit development, one open flower, and flower buds.

67 68

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eppig 1835). In his travelogue, Poeppig de-scribed the plant observed and collected be-tween Yumbel and Antuco in today’s Province of Bio Bío, Chile, very briefly in a single sen-tence within the text (translated from German:

“…and a peculiar cactus (Opuntia caespitosa. Poepp.) extends in broad, light green mats at the driest sites, immediately conspicuous by its sulphur yellow flowers and the silvery white

Henslow (1837), is now classified as Mai-hueniopsis darwinii, superficially similar to but never considered to be a Maihuenia.

The oldest collection of Maihuenia located hitherto can be attributed to the German nat-uralist Eduard Poeppig, who visited Chile in 1828. Poeppig’s account on his voyage was published seven years later. The new cactus was named “Opuntia caespitosa Poepp.” (Po-

Figure 71. Pereskia marcanoi. Pistillate flower in longitudinal section showing globular receptacle, large ovary and pistil, dwarfed stamens. Figure 72. Pereskia marcanoi. Longitudinal section of mature fruit with seeds em-bedded in mucilaginous pulp formed by the funicles; seed and viviparous plantlet with cotyledons, found to have germinated within the fruit (scale in cm). Figure 73. Pereskia bleo. Maximum leaf size of a cultivated specimen grown in 2000 at the Berlin-Dahlem Botanic Garden, kept as garden herbarium specimen. Figure 74. Pereskia guamacho root thickening exposed on a road side cut in Venezuela (photo NP Taylor). Figure 75. Pereskia zinni-iflora, one of three large succulent storage roots and main stem of plant three years after removal from the display glasshouse at Berlin-Dahlem (photo C Hillmann-Huber). Figure 76. Pereskia weberiana. Uprooted root thick-enings from a large potted plant.

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spines, which stick out everywhere from the short club-shaped stem segments.” This brief description very well depicts the species, which cannot be confused with any other cactus in the area (Figs 77–86).

The name Opuntia caespitosa Poepp., how-ever, cannot be used because the North Amer-ican Opuntia cespitosa Rafinesque (1830) pre-dates it. The first valid and legitimate name for a maihuenia was formally published by Pfei-ffer (1837a). He named Poeppig’s discovery independently as “Opuntia poeppigii Otto” based on seed-grown material. Pfeiffer attrib-uted only the name to Otto. The description was provided by himself, hence the author-ship remains with Pfeiffer (1837a). The plant had in fact already been mentioned earlier as a provisional name, without description, in a list of the 292 cacti of the Royal Botanic Gar-den of Berlin compiled by the head gardener,

Otto (1833), with the entry “208. O. Poeppi-gii.* Chili.” This indicates that it was already grown in Germany from seed sent to Berlin by Poeppig before the actual publication of Poeppig’s work. The species must have been quite widespread in cultivation according to Mittler (1844), who mentions it for seven pri-vate and public collections, including the bo-tanical gardens of Berlin, Munich, the Haage nursery, and the Salm-Dyck collection.

The same species was described once more as Opuntia maihuen Gay (1847). Gay used the vernacular name (maihuen) as the spe-cific epithet and noted that it was closely re-lated to Opuntia poeppigii. When Salm-Dyck (1850) published a new systematic subdivi-sion of the family Cactaceae, he distinguished seven tribes, among them the Opuntieae and Pereskieae. He transferred Opuntia poeppigii Pfeiffer to the genus Pereskia together with some other leaf-bearing cacti (several now ex-cluded from Pereskia and attributed either to Pereskiopsis or to Opuntia).

Curiously, a specimen located in the garden herbarium of the Missouri Botanical Garden (MO) originating from “hort. Dyck,” dated August 1857 and labelled “Peirescia poeppigii” is not Maihuenia poeppigii. The long spines (to 3 cm long) and conspicuous spur shoots identify it clearly as Maihuenia patagonica. It represents the oldest, though still mysteri-ous, record now known of a plant belonging to the second species recognized today: Mai-huenia patagonica (Figs 87–98). But this spe-cies was also first published as a member of a different genus: Opuntia patagonica Philippi (1863). Its history is interesting but compli-cated. The first collection, though fragmentary,

Table 6. Taxonomic history of Maihuenia.

Author Year Accepted speciesAccepted

subspecific taxa

Published combinations at species rank

Schumann 1898 3 3Britton and Rose 1919–23 5 0 6Vaupel 1925 5 0 6Berger 1929 5 0 7Backeberg 1966 5 0 8Ritter 1980 7 1 11Kiesling 1988 2 0 11Leuenberger 1997 2 0 11Hunt 1999 2 0 11Anderson 2001 2 0 11Anderson 2005 2 0 11Hunt, Taylor and Charles 2006 2 0 11

Table 7. Maihuenia, name and images in the WWW (2005–2007).

Maihuenia WWW entries

Maihuenia WWW images

23 May 2005 884 11219 Jul 2005 847 10924 Nov 2005 908 13118 Oct 2006 17500 9220 Feb 2007 13400 10426 Jun 2007 13600 574*16 Jul 2007 13100 633*18 Oct 2007 13800 751*

* Usually only about 10% of search results actually display an image showing a plant of the genus. In June 2007, only 40 of 574 images showed Maihuenia.


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nwas made by a traveller who came—quite re-markably—from the western side of the Andes to explore the dry interior of Patagonia. In 1862–1863, the Chilean physician Guillermo Cox was the first explorer to bring back bo-tanical collections from this part of the inte-rior of Patagonia, territory of the Pehuenche Indians, giving the origin as “Pampa de Pa-tagonia” (Cox 1863a, b; Middleditch 1988; Philippi 1863, 1864). The difficult interpre-tation of the specimen and its probable geo-graphical origin were discussed in more de-tail by Leuenberger and Arroyo-Leuenberger (1992). In more recent years a search in part of the area visited by Cox West of the Río Collon Curá in Neuquén was in vain. The area east of the river towards the Río Limay remains the most probable site of origin, and the epitype I selected (Leuenberger 1997) is in fact from this area.

None of these early introductions (herbar-ium specimens and plants grown from seeds) were illustrated in contemporary literature. The first illustration of a maihuenia is in a horticultural source in Germany. It is a wood cut of an imported, potted plant, published in 1881 in the journal Gartenflora under the name “Peireskia Pöppigii Salm” (Regel 1881). According to Philippi (1883), this illustra-tion was taken from an earlier publication in a catalogue of the German cactus nurs-ery Haage and Schmidt. It was only in 1883 that German born botanist RA Philippi in Chile, using the vernacular name maihuén, proposed the genus Maihuenia in an article on “Opuntia poeppigii Otto.” He based his description on plants he observed in its nat-ural habitat in Chile, at the Volcan Chillán,

along the Itata river and in the Cordillera de Talca (Philippi 1883). His color plate il-lustrates vegetative characters, a flower bud, and a flower of a plant from Chillán. Un-fortunately, Philippi did not formally pub-lish the new genus and only stated that

“one could perhaps make a genus of its own” (based on Opuntia poeppigii), which “one could name Maihuenia.” This means that he did not formally accept it himself. Ac-cording to the International Code of Botan-ical Nomenclature, ICBN Art. 34.1.a and b (McNeill and others 2006), the genus was therefore not validly published by Philippi. Many subsequent authors nevertheless at-tributed the genus to Philippi (1883), but it was only formally established 15 years later by Schumann (1898). It can be cited with the authorship “Philippi ex K. Schum.” to give credit to Philippi, who recognized the difference from Opuntia and who proposed the generic name based on the vernacular name maihuén (the significance of which is not quite clear).

Opinions on the acceptance of the genus varied at the time. FAC Weber (1898b), for instance, did not accept Maihuenia as genus. He treated two species under Pereskia, sub-genus Maihuenia, but stated that it should rather form a distinct genus. He attributed the name to Philippi. Of the little-known spe-cies described earlier, Weber recognized only Pereskia poeppigii. Simultaneously, he described a supposedly new species, Pereskia philippii (prospectively adding Maihuenia philippii as synonym!) from Chile, dedicating it to the author of the proposed genus.

This ambivalent treatment by Weber and

Table 8. Maihuenia names and images of species in the WWW (2005–2007).

Maihuenia poeppigii Maihuenia patagonicaWWW entries WWW images WWW entries WWW images

23 May 2005 202 43 181 3719 Jul 2005 313 40 261 3824 Nov 2005 258 52 213 4518 Oct 2006 1450 43 521 2620 Feb 2007 2010 40 1490 2626 Jun 2007 2150 122

(only ca. 20)801 69

(only ca. 20)16 Jul 2007 2100 90

(only ca. 20)711 66

(only ca. 22)28 Sep 2007 1840 125

(only ca. 23)1550 107

(only ca. 18)18 Oct 2007 1840 116* 1370 123*

* Usually only about 10% of the search results actually display an image showing a plant of the species.


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n tioideae), and before his genus no. 20, Pereskia (Pereskioideae). Schumann (1898) accepted three species: M. poeppigii, M. brachydelphys (new), and M. philippii. Whereas Schumann (1899) maintained the placement of the genus in the subfamily Pereskioideae, based on the seed characters and the lack of glochids, his later treatment in the appendix to the 1897–98 monograph is again ambivalent (Schumann 1903: 15). Here he affords Maihuenia an in-termediate position and places the genus in Opuntioideae, but he calls it an intermediate form due to the black, brittle seed coat. This may have been influenced also by his (errone-ous) indication of the presence of glochids.

In the 20th century, a series of new taxa and

by Schumann has caused confusion on the authorship. When Schumann began to pub-lish his comprehensive monograph of Cac-taceae, issued first in several instalments in 1897–1898, he did not yet consider Maihue-nia a genus: it does not appear in the generic key on p 31 at the beginning of his mono-graph (Schumann 1897). However, he later inserted Maihuenia in the treatment of the subfamily Pereskioideae in the text on p 754, where he stated that Maihuenia was justified as genus, and where he formally established it (Schumann 1898). Schumann placed Maihue-nia in the subfamily Pereskioideae with a re-mark on the transitional position, as genus no. 19a after his genus no. 19, Pterocactus (Opun-

Figures 77–83. Maihuenia poeppigii. Figure 77. Giant cushion at La Invernada, Maule Valley in Chile. Figure 78. Flowering cushion on a basaltic outcrop at Primeros Pinos, Neuquén, Argentina, with Baccharis sp and Araucaria araucana in the background. Figure 79. Dense cushion with flower at early anthesis in Neu-quén, Argentina (Leuenberger and Arroyo 3433). Figure 80. Interior of flower, cultivated (Correa s.n., from Neuquén). Figure 81. Stem and flower in longitudinal section (Leuenberger and Arroyo 3433) showing the large mucilaginous pith of the stem. Figure 82. Flower and fruit sample showing different shapes according to stage of maturity and variation. Río Guaqui, Chile. Figure 83. Dense cushion with mature fruit protruding from the cushion surface.

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re-interpretations of species names began with the publications of the Argentine botanist Spegazzini (1902). His descriptions of Mai-huenia tehuelches from Santa Cruz and M. val-entinii from Chubut are each based on two or three herbarium collections of contempo-raneous collectors. They are meticulous, giv-ing detailed measurements of specimens with flowers and immature fruit. Spegazzini did not know the plants from the field. The type specimens were illustrated by Leuenberger (1997). The moderate increase of names is shown in Table 6.

Britton and Rose (1919: 8) placed the genus Maihuenia in their tribe Opuntieae (not in Pereskieae!) although they noted the similarity of the seeds with Pereskia and the lack of glochids (an exception within Opun-tieae) but considered it “otherwise very dif-ferent [from Pereskia]”. They recognized five species, among them Maihuenia patagonica. Their treatment is somewhat confusing be-cause the illustration of Maihuenia valenti-nii, a photograph received from Spegazzini, in fact shows M. poeppigii. Britton and Rose also erroneously placed M. philippii in the syn-onymy of M. patagonica. The nomenclatural problems resulting from this mix-up were dis-cussed and resolved only much later by Pichi-Sermolli and Bizzarri (1978).

In a more exhaustive revision, though still based exclusively on herbarium material and following much the same line of Britton and Rose, the Berlin-Dahlem botanist Vaupel (1925) also distinguished five species. He accurately illustrated M. poeppigii and M. tehuelches but excluded the older name Opuntia patagonica, stating that the description was insufficient. Vaupel tried to introduce a new tribal classifi-

cation solely based on seed characters, in which he placed Maihuenia in a tribe comprising all cacti with “soft seeds” in contrast to the ones with a bony seed coat (the latter coinciding with the opuntioids of Schumann and tribe Opuntieae of Britton and Rose). He accepted Maihuenia as second genus after Pereskia in his monographic series. Vaupel (1925, 1926) was able to publish only two volumes (includ-ing the genera Pereskia, Maihuenia, Rhipsalis in volume 1; and Rhipsalis, Wittia, Epiphyl-lum, and Aporocactus in volume 2) before his tragic early death. His classification was not followed by later authors.

Berger (1926, 1929) placed Maihuenia in subfamily Pereskioideae and distinguished five species but treated in some detail only the two that he knew from cultivation, M. poeppigii and M. patagonica. Backeberg and Knuth (1936) listed five species and provided the first pho-tograph of a flower of Maihuenia patagonica. Marshall and Bock (1941) mentioned Mai-huenia poeppigii as desirable for cultivation and obtainable in trade.

Castellanos and Lelong (1943) published only a general account, without details on the species, but theirs is accompanied by a fairly accurate distribution map. Buxbaum (1951) never saw maihuenias in the field, nor proba-bly in cultivation, but he quite accurately rec-ognized from published illustrations the two growth habits representing the two species of Maihuenia accepted today. In 1954 and 1955 Friedrich Ritter collected seeds of Maihuenia in Chile and Argentina. These were distributed through the seed catalogues of Ritter’s sister Hildegard Winter in Germany (Winter 1955, 1956, 1957). In her catalogue of 1957 two new taxa were mentioned as nomina nuda (naked

Figures 84–86. Maihuenia poeppigii. Figure 84. Germinating seedlings of one to few days (germination took place in May following a one year delay in the seed tray) (Leuenberger and Arroyo-L. 4180). Figure 85. Two year old seedling with incipient ramification. Figure 86. Seedling plant in winter at temperature below freezing; note the shrivelled stem but fully turgescent and persistent leaves (Figs 82–86 from Leuenberger and Arroyo-L. 4180).

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n names): “M. albolanata Ritter sp. nov.” and its variety “viridulispina Ritter var. nov.” These were validly published much later (Ritter 1980) but seem to represent no more than variations in spine color (Leuenberger 1997).

Significant information on Maihuenia is found in publications of Argentine and Chil-ean botanists, for instance, by Ruiz Leal (1956) on habitats of Maihuenia in Mendoza, on drought resistance, seed germination, and pol-len; by Muñoz Pizarro (1959, 1960, 1966) on the vernacular names of Maihuenia in Chile and the early herbarium specimen of Cox at

SGO; and by Roig (1963), who discussed the northern limit of the geographical distribu-tion of the genus in Mendoza.

Hunt (1967) treated Maihuenia as second genus of Tribe Pereskieae in Hutchinsons’ Genera of Flowering Plants. Buxbaum (1969) concluded that Maihuenia was very close to Pereskia in seed and flower structure but dis-tinct in growth habit. Kiesling (1975) pro-vided key characters for the genus, a generic description, and illustrations of a branchlet of Maihuenia sp (M. patagonica) including flower bud and internal flower structure. An-

Figures 87–90. Maihuenia patagonica. Figure 87. In habitat in the Portezuelo del Viento in southern Men-doza, Argentina, Figure 88. Habit with conspicuous spur shoots at Coipo Lauquén in southern Mendoza (Leuenberger and others 3989). Figure 89. Newly growing stem with leaves and young developing spines, stem showing early periderm formation, lower stem part of the previous year with spur shoots, old and new leaves hardly distinguishable (cultivated, Bertolami s.n., from Chubut, Argentina). Figure 90. Newly grown stem, leaves, and nearly mature spines, first axillary leaves of new spur shoots present. Lower half of stem with spur shoots of the previous year, the oldest leaves (one year old) turning yellow.

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natomical characters of two species were stud-ied in detail by Gibson (1976, 1977). In the second, most substantial paper on the subject, Gibson also discussed the systematic position of Maihuenia and concluded that vegetative features negate its inclusion in the subfamily Opuntioideae.

The number of published species increased again when Ritter (1980), in the second vol-ume of his book on the cacti of South Amer-ica, described three new species and one new form from Argentina, all based on his collec-tions made in 1955 (listed by Eggli and oth-ers 1996).

Earlier detailed biological observations are scarce. Only Jung (1982) gave an accurate and well-illustrated account on the habitat, roots, and fruit of Maihuenia poeppigii in Chile. In the Flora Patagonica Kiesling (1988, 1990) re-duced the ten species names available at the time to only two, M. patagonica and M. po-eppigii. The treatment was based on herbar-ium collections made for the Flora Patagon-ica project and personal field experience. Both species are illustrated in line drawings with-out flower, fruit, or seed. Maihuenia poeppigii was illustrated in a more representative color plate including flower and fruit by Hoffmann (1989). Hunt and Taylor (1990), and Hunt (1992) listed Maihuenia as a genus with two accepted species.

Field observations and notes on the morphol-ogy and ecology in Argentina and Chile were provided by Leuenberger (1992a, 1992b) and Kraus (1993, 1994, 1995). Wallace (1995a, b) analyzed molecular data suggesting Maihue-nia should be treated as a monogeneric sub-family of Cactaceae. This prompted Fearn (1996, unpaged) to publish the new subfam-ily Maihuenioideae. The new subfamily was supported by later molecular studies (Nyffeler 2002) and accepted in standard works (An-derson 2001, 2005).

In my 1997 monograph, the genus was completely revised on the basis of 275 herbar-ium collections from 32 herbaria, including 55 of my own collections, field studies made between 1985 and 1992 in Argentina and Chile, and observations of seed-grown plants in cultivation (Leuenberger 1997).

Notes on the distribution of Maihuenia

A purely hypothetical and most certainly er-roneous report of a southernmost occurrence of Maihuenia poeppigii at Torres del Paine Na-tional Park, some 1400 km south of its known distribution, was published by Jonkers (1998),

somewhat allusively as “Maihuenia poeppigii’s best kept secret.” Jonkers, who never saw the plant there, just as presumably no one else ever did, asks “how did the plant get that far south?” The answer is still lacking, and the report, with attractive (but in terms of the geographical distribution, misleading) illus-trations, is apparently based on hearsay rather than on facts. Substantial information was contributed by Mauseth (1999), who studied anatomical adaptations to xeric conditions in Maihuenia. A well-illustrated account on mai-huenias in the field was prepared by Kümmel and Klügling (2005).

Altitudinal distribution is sometimes erro-neously reported as Patagonia and High Andes, but this is not quite correct, as Maihuenia oc-curs predominantly in lowland and medium altitudes, rarely occurring over 2000 m in the northernmost part of the range (where the Andes reach 3000–5000 m altitude). I anal-ysed the myths surrouning high-Andean lo-calities (Leuenberger 1997), which date back to Backeberg and freely copied later. The dis-crepancy concerning the altitude between the herbarium labels of the lost type specimen (Ar-gentina, [Mendoza], Paso Cruz, 2600 m, Kun-tze s.n. (B)) and the neotype from the same area (“Paso Cruz 34° Cordillera,” 1500 m, 01. 1892, Kuntze 109”) may be also significant.

Both species occur on a variety of substrates. It was only in 2003 that I had an opportunity to visit the impressive habitat in the Payunia vol-

Synonyms of Maihuenia with reference to accepted name.

Maihuenia

M. albolanata = M. patagonicaM. albolanata forma viridulispina = M. patagonica

M. andicola = M. patagonicaM. brachydelphys = M. patagonica

M. cumulata = M. patagonicaM. latispina = M. patagonicaM. philippii = M. poeppigii

M. tehuelches = M. patagonicaM. valentinii = M. patagonica

Opuntia

O. caespitosa = M. poeppigiiO. maihuen = M. poeppigii

O. patagonica = M. patagonicaO. philippii = M. poeppigiiO. poeppigii = M. poeppigii

Pereskia

P. philippii = M. poeppigiiP. poeppigii = M. poeppigii


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canic region in Mendoza, Argentina, where vast areas covered by lava pebbles (lapilli) bear only very sparse vegetation but are dominated by large cushions of Maihuenia patagonica (Fig 2).

Maihuenia in the WWW (2005–2007)

Maihuenia is still moderately well represented in the WWW, surprising for a relatively little-known genus that is not widespread in culti-vation. But the quality of some illustrations exceeds that in other published sources, and the number of good images from habitat is rapidly increasing (Tables 7, 8).

Taxonomic history of BlossfeldiaWhen the first plants of this genus were dis-covered by the German collector Harry Bloss-

feld and the Argentine collector Oreste Mar-soner in 1936, it was hardly conceivable that this taxon would cause so much discussion on evolutionary lineages in Cactaceae seventy years later. Blossfeldia liliputana was published by the Berlin-Dahlem botanist Werdermann (1937b) based on a single collection from Tumbaya, Province of Jujuy, Argentina. Bloss-feld considered the plant to be exceptionally rare (Werdermann 1937b). Blossfeldias grow well-camouflaged in rock fissures and are cer-tainly very difficult to find in the natural hab-itat, but Ritter (1980) found it to be one of the most widespread cacti of Bolivia, and we now know it to be far from rare. Its double reputation as a threatened and non-threat-ened plant is evident in conservation liter-ature, where Blossfeldia serves as an illustra-

Figures 91–98. Maihuenia patagonica. Figure 91. Stem with leaves in winter at temperature far below freez-ing point (–15° C). Figure 92. Transverse section of stem, note the mucilaginous pith, green cortex and thick brown periderm. Figure 93. Stem with flower bud surrounded by new growth of three lateral branches in the axils of the spines. Figure 94. Flower buds and dry flower remnants of the previous year. Figure 95. Three flower buds opening in the morning. Figure 96. Three fully open flowers in the afternoon, anthers dehisc-ing, stigma lobes still connivent. Figure 97. Flower in late stage of anthesis, stigma lobes widely expanded. Figure 98. Longitudinal section of flowering stem, receptacular cup with green cortex, fleshy base of tepals and filament insertion zone, locule immersed in the receptacular cup but ovary roof broad.

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tion but is simultaneously considered as not threatened in the relevant geographical chap-ter (Oldfield 1997: 115, 116.). Where it is found, it occurs usually in large numbers of individuals, and the species has a remarkably wide range of distribution (Leuenberger and Eggli 1999) (Figs 99–107).

Due to its special habitat, the late discovery of this tiny cactus is not too surprising. The range of the genus is considerable, covering an area along the eastern side of the Andes over 1800 km in N–S direction from the depart-ment Santa Cruz in central Bolivia to the Prov-ince of Mendoza in western Argentina. Com-paratively few collectors traveled in this vast area before the end of the 19th century, and most of them did not pay much attention to cacti, mainly because they were difficult to handle for the herbarium. The first collectors with an interest in cacti presumably missed the plants because of their very peculiar hab-itat, and perhaps because the plants were not seen in flower. For example, John Gillies col-lected several of the more conspicuous cacti in Mendoza. Gillies was a Scottish physician and botanist who lived in Argentina from 1820 to 1828 and sent herbarium material, plants, and seeds to England and Scotland (Dawe 1988). In 1824 Gillies made a trip to the foothills of the Andes and sent to Hooker a box of plants (herbarium specimens) and seeds, including cacti and tillandsias (Dawe 1988: 44). Seeds or live plants of some cacti (but apparently not blossfeldias) reached other gardens. Some-what cryptic but unambiguous origin data are found in a published list of the cacti cultivated in 1829 in the Royal Botanical Garden of Ber-lin, namely Cereus strigosus and C. candicans with the indication “H. Angl. Gillies Men-dez,” and three opuntias (including Opuntia ovata published later by Pfeiffer 1837a) also with the indication “Gillies Mendez,” point-ing to the province of Mendoza, Argentina as the origin and Gillies as the collector (Anony-mous 1830). In an earlier list, not yet accom-panied by data of origin (Anonymous 1827, apparently written by Link and Otto), these names were not yet mentioned. The year of arrival of these cacti via England or Scotland

to gardens in central Europe can thus be nar-rowed to 1829 onward.

Figures 99–101. Blossfeldia liliputana. Figure 99. Habitat in the Quebrada de Humahuaca, Jujuy, Ar-gentina, between the type locality and Humahuaca. Figure 100. Habitat, coarse granitic weathered rock faces at Cuesta de la Miranda, La Rioja, Argentina. The cliff-dwelling bromeliad is a Deuterocohnia species. Figure 101. Habitat, steep schist (phyllite?) rock faces with fissures, near Tupiza, Potosí, Bolivia.

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n Some other, also small and rather inconspic-uous cacti growing within the range of Bloss-feldia were found and described much earlier than Blossfeldia, perhaps due to their habitat and the more showy red flowers. Examples are Re-butia pygmaea and Opuntia subterranea, found by the Swedish botanist RE Fries in 1901 on the Altiplano of Jujuy in northern Argentina. Another small cactus, Rebutia steinmannii, was collected by Steinmann around 1902 in Bolivia, but Blossfeldia apparently escaped the attention of these and other early collectors. Some very earlier travellers in NW Argentina and Bolivia probably came close to (or even stepped over) Blossfeldia habitats. For Bolivia one could men-tion D’Orbigny around 1842, Weddell 1845–1848, Fiebrig 1903–1904, and Buchtien from 1906 onward. In Argentina, besides Gillies, it was Tschudi in 1859 in Catamarca, and Claren in 1901 in Jujuy, who collected in the area. Information on early botanical collectors was compiled by Fries (1905) and Funk and Mori (1989). I am not aware of any unrecognized older herbarium record of Blossfeldia, and it appears that it is a true twentieth century dis-covery by commercial plant explorers.

The first local botanists to provide more in-formation on Blossfeldia after 1937 and citing additional herbarium material in Argentina were Castellanos and Lelong (1943). Krainz (1949a) gave an account of successful culti-vation and described and illustrated the fruit and seed. Notes on the introduction and cul-tivation soon appeared in cactophile journals (Johnson 1954; Krainz 1941, 1946, 1949b; Steeg 1955).

German collectors with commercial interests then discovered more populations of Blossfeldia in other regions of Argentina. Between 1954 and 1962 Fechser sent living plants to Curt Backeberg and reported on it in the US cac-tus journal (Fechser 1960). Backeberg (1959a, 1959b, 1962) treated the known and new taxa. It is remarkable that the 50 year cumu-lative index of the US journal indicates only five entries on Blossfeldia between 1929 and 1979. Before 1960, Blossfeldia was mentioned and illustrated there only twice (Anonymous 1949, Johnson 1954).

The first records from Bolivia are rather late, from 1953 onward, thanks to the travels of Friedrich Ritter. Ritter (1980) noted that he had also missed them on his first trip to Bolivia. A new variety, B. liliputana var caineana was published by the Bolivian botanist Cárdenas (1964). Blossfeldia is apparently quite wide-spread and frequent in Bolivia but equally re-stricted to the special habitat in narrow can-

yons and in rock crevices on steep hillsides. Friedrich Ritter collected numerous herbarium samples for his private herbarium and seeds for sale through the seed catalogues issued in Frankfurt-Fechenheim, Germany, by his sister Hildegard Winter (Winter 1954, 1959). The first ones were distributed as Blossfeldia lili-putana in 1959—three from Bolivia, as sup-posed new species, at first under unpublished names (see also Ritter 1980).

The commercial interest prompted the de-scription of further insufficiently known new taxa (Table 9). Blossfeldia campaniflora Backe-berg (1959a, 1959b) was based on a Fechser collection received in 1954 with the vague in-dication of origin “North of Argentina“, and Blossfeldia fechseri Backeberg (1962),whose indication of origin was “ca. 800 km further to the south than the type of the genus.” The reason for this intentionally vague data was evidently the commercial interest to withhold exact locality data. Both names remained in-valid according to the International Code of Botanical Nomenclature, ICBN Art. 37.1 (McNeill and others 2006) for lack of indi-cation of the type collection (see also Eggli 1985). Blossfeldia fechseri was later mentioned by Fechser (1965) for the province of Cata-marca, again without locality, in a note on the cultivation of imported plants.

Ritter’s numerous collections made between 1953 and 1959 in Argentina (Prov. Salta, Jujuy) and Bolivia (Departments of Potosí, Chuqui-saca, and Cochabamba) were well documented. The localities of his field numbers and some new taxa were published by him (Ritter 1965, 1980). All of these are currently considered to be synonyms of Blossfeldia liliputana. Locality data for all of Ritter’s often very fragmentary herbarium samples were catalogued in detail by Eggli and others (1996).

The first fairly detailed morphological study of the flower, fruit, and seeds of Blossfeldia lili-putana was made by Buxbaum (1964), who also summarized the scattered information available on the genus. Doubts on the jus-tification of the other five published species, which are hardly distinguishable in cultivation, led Krainz (1975) to reduce them to varieties of Blossfeldia liliputana. Krainz did not know the plants from the field but had at least five accessions with different names in cultivation at the Zurich Municipal Succulent Collection (Krainz 1967), all acquired after 1957.

The southernmost blossfeldias in Men-doza were already mentioned by Buining (1969), who had observed them on the way from Mendoza to Uspallata thanks to a local


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ncontact, Dr Gomez Ruepel of Mendoza. Rit-ter (1980) comments on them as well, and the same habitat was reported again as new by Méndez (1983), who added detailed and new observations on the biology of the plant. Both Ritter (1980: 433) and Méndez (1983) think that the large strophiole of the seed per-mits wind gusts to transport the very small seeds along rocky cliffs, and that the papil-lae of the seed coat enable them to cling to the substrate. Both authors also comment on the ability of blossfeldias to form new stems within the plant body, which apparently break through the epidermis.

Since names once established (whether no-menclaturally valid or not) prove to be rather persistent, particularly in the amateur litera-ture and on commercial lists, it is not surpris-ing that Ritter’s names, as well as provisional names listed by Knize (1987), are sometimes still found in collections and commercial cat-alogues today, at least in Europe (and duly re-cycled on the internet).

Further published accounts on seed, pol-len, and chromosome number of Blossfeldia were summarized by Leuenberger and Eggli (1999). For over forty years, the genus was not questioned taxonomically except by Hunt and Taylor (1986), who placed Blossfeldia in small print under genus nr. 61. Parodia (of a total number of 86 of the “consensus list”), stating that it could be referred either to Par-odia or to Frailea. Taylor (1987, 1989) made and used the combination Parodia liliputana. But soon thereafter Blossfeldia was reinstated as a genus following discussion in the IOS Working Party (Hunt and Taylor 1990). Hunt (1992, 1999) listed Blossfeldia with one spe-cies, B. liliputana, in the CITES Cactaceae Checklist. Both Anderson (2001, 2005) and Hunt and others (2006) maintain Blossfeldia as a monotypic genus. The status as genus is supported by newer data from molecular stud-ies (Nyffeler 2002), which surprisingly sug-gest a basal position of this seemingly highly evolved and reduced plant. Mauseth (2006) studied the anatomy in detail and showed that anatomical data neither prove nor disprove the molecular findings. These have been sup-ported further by Crozier (2004) and Butter-worth (2006) with the consequences of pro-posing of a new subfamily name by Crozier (2004) and new tribe Blossfeldieae by Crozier (2004) and again (but illegitimatly) by But-terworth (2006).

Mottram (2007) compares the flowers of Blossfeldia with those of Rhipsalis and thinks a relationship more probable, while speculating

Figures 102–104. Blossfeldia liliputana. Figure 102. Several small plants on fine soil accumulation in fis-sures of the schist. Damaged plant with exposed root in the center. Figure 103. Small heads (seed-lings or regenerated heads) in a moist rock fissure. Figure 104. Rock fissure with root buds on a partially decayed plant, and seedling.

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n simultaneously that Blossfeldia might be a de-rived member of the Maihuenioideae, an op-tion already discussed by Butterworth (2006) but not supported by additional data. The lack or presence of a true receptacular tube in Blossfeldia has caused discussion based on still insufficient data. Having observed only recep-tacular tubes of up to 0.5 mm length, Ritter (1980) questioned the details of the illustra-tion of a longitudinal section of the flower by Buxbaum (1964), which shows a tube at least 1 mm long. The variation in receptacle shape has not yet been studied and, considering the wide differences in flower tube length found in some other globular cacti like Rebutia and Sulcorebutia, the question needs serious ana-tomical study more than theoretical discus-sion. Preliminary observations may indicate considerable differences between clones or even on the same plant. Ritter (1980) also criticized the production of new species by Backeberg based on uncritically interpreted flower characters.

Notes on the distribution and biology of Blossfeldia

Since the general distribution map published by Leuenberger and Eggli (1999), some new records from San Juan (Kiesling 2003) and a number of records and sightings from Bo-livia add to the overall picture of Blossfeldia distribution, showing a surprisingly large area of geographical distribution though ap-parently in widely disjoint rock fissure habi-tats along the eastern edge of the Andes, that is in the eastern Cordillera and the western and northern Sierras Pampeanas of Argentina and Bolivia. The altitudinal range is limited to about 1000–2550 m in Argentina and to over 3000 m in Bolivia (according to Ritter (1980) up to 3400 m).

How are the seeds of Blossfeldia trans-ported, and how did blossfeldias reach their widely scattered and—at least today as far as explored—widely disjoint habitats? Theories could not be more divergent. From the obser-vations of Ritter (1980) and Méndez (1983) on dispersal of the seeds by wind one could imagine long-distance displacement due to exceptional meteorological events. By con-trast, Barthlott and Porembski (1996), after an experiment in the laboratory, believe ants to be the active agents transporting the seeds. My own chance observation of ants and fruit-ing Blossfeldia near Fiambalá in Catamarca do not support (though neither do they dis-prove) the latter theory. In a single case ob-served, an ant coming by a fresh, manually

Figures 105–107. Blossfeldia liliputana. Figure 105. Uprooted specimen showing underground ramifica-tions, the massive succulent roots representing the major volume of the whole plant. Figure 106. Old head of a cultivated specimen, with flower. Plant body showing effect of drought (Leuenberger and Arroyo 3579). Figure 107. Whole plant of the same, with fruit. Oldest heads are ca. 20 years old. All photos by the author unless otherwise stated.

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split fruit of Blossfeldia eagerly absorbed the moisture available from the still fleshy funi-cles but did not carry away any seeds. Much still remains to be studied on the biology of Blossfeldia. Longer distance transport by birds does not seem entirely impossible (compare the slovenly plover mechanism, a remarkable case of long distance dispersal, with which Larrea reached the northern hemisphere 4–8 million years ago, according to Wells and Hunziker (1976) and Lia and others (2001). Still purely hypothetical, an exozoochorous transport, by seeds with their minute papil-late processes clinging to feathers, seems pos-sible as well. This has not yet been analysed, and near to nothing is known on flower and fruit biology in the genus.

Barthlott and Porembski (1996) state that Blossfeldia is a poikilohydrous plant. There

have been no reports on such experiences with Blossfeldia in cultivation, presumably be-cause nobody takes the risk of ending up with non-resurrected plants. Experience with some ferns, selaginellas, and Myrothamnus flabelli-folius shows that they do not like to dry out entirely in cultivation, and the special fea-ture of true resurrection of the whole plant is not “on display” in the flower pot in gardens. The report of poikilohydry for Blossfeldia still merits a critical analysis and further investi-gation. The comparatively enormous under-ground amount (often more than 90%) of fleshy root and stem tissue would seem to be an argument against complete drying out of Blossfeldia in habitat. What can be observed in nature (Ritter 1980) and also in cultiva-tion under drought conditions is the marked shrinking of the green part of the plant. This

Table 9. Taxonomic history of Blossfeldia.

Author YearAccepted

speciesAccepted

subspecific taxa

Total of published combinations at species rank in the same year

Werdermann 1937 1 0 1Backeberg 1959 2 0 2Backeberg 1962 3 0 3Backeberg 1966 5 1 5Krainz 1975 1 6 5Ritter 1980 4 4 6Eggli and Leuenberger 1999 1 0 6Hunt 1999 1 0 6Anderson 2001 1 0 6Anderson 2005 1 0 7Hunt, Taylor and Charles 2006 1 0 7

Table 10. Blossfeldia names and images in the WWW (2005–2007).

DateBlossfeldia

WWWBlossfeldia

WWW Images

Blossfeldia liliputanaWWW

Blossfeldia liliputana

WWW images19 Jan 2005 508 192 513 7717 May 2005 571 185 450 9315 Jun 2005 4160 205 883 9619 Jul 2005 3930 205 904 9619 Dec 2005 13400 197 526 8510 May 2006 33800 206 634 8726 Jun 2007 21900 1750* 2670 346*28 Sep 2007 24700 1960 2030 40618 Oct 2007 23800 1930* 2150 435*

* Usually only about 10% of the search results actually display an image showing a plant of the genus or species. In July 2007 less than 60 of 346 B. liliputana images were correct. In September 2007 only about 64 of 406 were correct, while only 80 of 1960 images for a search on “Blossfel-dia” were correct.


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can also happen when blossfeldias lose part of their roots due to excessive watering and, unable to take up water, are affected by sub-sequent drought. The remaining plant body then loses much water, regeneration is slow, and part of the cortical tissue does not re-cover. The new growth then appears within the sunken part of the older stem apex (Fig 106). Blossfeldias have an impressive capac-ity of regeneration of lost stem parts and ap-parently produce new shoots not only from damaged stem parts but also from exposed roots, as reported by Méndez (1983). In any regard a remarkable plant.

Blossfeldia in the WWWBlossfeldia is comparatively well represented on the internet due to the interest of amateur cactus growers (Table 10). No repeated searches were made of synonyms and nomina nuda, but the overall pattern is much the same as for other popular cacti, according to one search made for

a selection of names on 28 Sept 2007 (Bloss-feldia campaniflora, name entries 137 / images 187; Blossfeldia fechseri, 180 / 32; Blossfeldia ped-icellata 146 / 31, Blossfeldia minima 265 /28; Blossfeldia flocculosa 69 / 4).

A postcript on Pereskia, Maihuenia, and Blossfeldia in the WWW

More restricted and substantial information is found by Google scholar search, made for all three genera at a late stage of this study (24 October 2007). Numerous links to sci-entific papers were found, 746 referring to Pereskia, 106 to Maihuenia, and 51 to Bloss-feldia. The image links are apparently not more restricted and thus not more reliable than general Google searches. The result was: Pereskia 3540, Maihuenia 811, Blossfel-dia 1910, with very varied image links and the same low percentage of actual and cor-rect images.

AcknowledgmentsI wish to thank Michael Rodewald for image scans, the horticulturists of the Berlin-Dahlem Botanic Garden for taking care of the live plants over many years, Mrs Marion Cubr for preparation of garden herbarium specimens, Dr Nigel Taylor (Kew) for photographs of the root system of Pereskia guamacho, an anon-ymous reviewer for extensive detailed com-ments, and Dr Silvia Arroyo-Leuenberger for help with the manuscript.

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Blossfeldia atroviridisB. campaniflora

B. fechseriB. flocculosaB. minima

B. pedicellataParodia liliputana


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Leuenberger on Pereskia