Leuenberger On Pereskia

Haseltonia 14: 54–93.
2007 54
Pereskia, Maihuenia, and Blossfeldia—

early cactus evolution
taxonomic history, updates, and notes

Beat Ernst Leuenberger
Botanischer Garten and Botanisches Museum Berlin-Dahlem,
Freie Universität Berlin, Berlin, Germany; b.leuenberger@bgbm.org
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
Introduction cal distribution, of leaf bearing, only slightly

Pereskia is one of the earliest cacti known to succulent, deciduous, spiny trees, shrubs, or
botanists, first collected in the late 17th cen- vines. Plants belonging to Maihuenia were
tury and described as genus at the beginning of found quite early in the 19th century but, due
the 18th century. It is a genus of mostly tropi- to their habit were first classified as species of
1 2
igure 1. Pereskia lychnidiflora, tree in dry forest near Tehuantepec, Oaxaca, Mexico. Figure 2. Maihuenia pata-
F
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).
HASELTONIA 14, 2008 55
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).
Opuntia and later as Pereskia before being rec- ular cactus Blossfeldia was far off the path of
ognized as a separate genus at the end of the the earliest botanical explorers, due to its range
19th century. They are low, caespitose, spiny from the Andes of central Bolivia to western
succulents with small but long lasting leaves. Argentina, but it was also overlooked by early
Their geographical range is predominantly Pa- collectors reaching this area. It was found and
tagonian. The rock-fissure-dwelling, tiny, glob- recognized as new only in the first half of the
56 Leuenberger—Pereskia, Maihuenia, and Blossfeldia
Table 1. Taxonomic history of Pereskia (incl. Rhodocactus)

Accepted Total of published

Accepted infraspecific names at species rank
Author Year species taxa in the same year
Miller 1754 1 0 1
de Candolle 1828 9 0 9
Pfeiffer 1837a 12 3 12
Mittler 1844 8 3 12
Salm-Dyck 1850 8 3 12
Schumann 1898 11 0 38
( + 4 insufficiently (incl. nom. nud.)
known)
Britton and Rose 1919–23 19 0 50
Vaupel 1925 18 6 47
(incl. nom. nud.)
Berger 1929 ca. 18 1 (23)
(listed in Berger 1929)
Backeberg 1966 24 1 53
(8 Pereskia, (35 Pereskia,
16 Rhodocactus) 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 4
Anderson 2005 17 4
Hunt, Taylor and Charles 2006 17 4 ca. 63
(in IPNI incl. nom. nud.)
20th century, at a time when cactus botany mary of the taxonomic history of the three
was much influenced by commercial activities. genera, including some updates and obser-
The three genera thus belong to quite differ- vations, complemented by illustrations of
ent eras of exploration and botanical taxon- some selected morphological features of these
omy, with the Pereskia story starting in pre- plants of widely differing habit and newly
Linnaean time, in about 1689, Maihuenia in discussed phylogenetic and taxonomic po-
1828, and Blossfeldia only in 1936. sition (Figs 1–3).
Between 1986 and 1999 I contributed sev-
eral papers on the three genera. Pereskia and Taxonomic history of Pereskia
Maihuenia were treated in classical taxonomic Pereskia is one of the earliest cactus genera,
monographs (Leuenberger 1986, 1997) and already established in pre-Linnaean times by
in additional papers (Leuenberger 1992a, b, Plumier (1703). It was described from mate-
c, d, among others). Blossfeldia liliputana was rial collected by Charles Plumier, a French
also studied in the field in Argentina, and a friar, naturalist, explorer, and botanist under
bibliographic survey and review of the genus King Louis XIV of France. Plumier’s herbar-
appeared in this journal (Leuenberger and ium collections of Pereskia were lost, but it
Eggli 1999). It could hardly be foreseen at can be assumed that they were collected be-
that time that all three genera would come tween 1689 and 1695 on the Caribbean isles.
into the particular focus of DNA studies According to Mottram (2002: 83, 94), who
relevant for basic and rather revolutionary reproduced the first description and pub-
questions of cactus evolution and systemat- lished illustration of genus Pereskia by Plum-
ics (Wallace 1995b; Butterworth and Wal- ier (1703) and Plumier’s unpublished plate
lace 2005; Crozier 2004; Butterworth 2006; of “Pereskia aculeata, flore albo, fructu flaves-
Nyffeler 2002; Edwards and 2005; Edwards cente” from the manuscript “Botanicon Amer-
and Donoghue 2006) The purpose of the icanum,” the plate was drawn between 1695
present paper is primarily to give a sum- and 1697. This first drawing is much more
Table 2. Pereskia and its orthographic variants found Table 3. Taxonomic history of Cactaceae: number

by Google searches in the WWW, 18. 10. 2007. of genera and species.
Name Image1 Author Year Genera Species

Pereskia 80900 32002 Linnaeus 1753 1 22
Peireskia 589 393 Candolle 1828 7 174
Peirescia 241 69 Pfeiffer 1837 10 422
Perescia 353 60 Schumann 1898 21 578
Pereskya 62 293 Britton and Rose 1919–23 124 1325
1
Usually only about 10% of the search results actually display an Berger 1929 41 —
image showing a plant of the genus.
2
Less than 300 show a pereskia. Krainz 1960 119 —
3
Only two show a pereskia. Backeberg 1966 236 3100
Krainz 1967 152 —
representative of Pereskia aculeata than the Hunt 1967 84 —
illustration published by Plumier (1703). Buxbaum in 1974 159 —
In his grand work Species plantarum, which Endler and
marks the starting point of modern botanical Buxbaum
nomenclature, Linnaeus (1753) placed all cacti Hunt and Taylor 1986 86 —
in the genus Cactus. Two of his species belong Hunt 1999 114 1942
to the leafy cacti, Cactus pereskia and Cactus Anderson 2001 125 1810
portulacifolius, but the connection of C. portu- Anderson 2005 126 19001
lacifolius with Pereskia, as circumscribed later
Hunt, Taylor 2006 124 14382
on, had not yet become evident. The British and Charles
horticulturist Philip Miller did not follow the 1
Plus about 500 subspecies and varieties.
Linnaean one-genus concept for cacti and re- 2
Plus 378 subspecies.
instated the genus Pereskia only one year later
in his Gardners dictionary (Miller 1754). The
genus Pereskia is therefore nomenclaturally by Plumier (1703) and mentioned three times
valid not from Plumier (1703) but only from in the paper. The same spelling was used by
Miller (1754) as explained by Stafleu (1969) Linnaeus in the epithet of Cactus pereskia and
(Table 1). Miller (1754) mentioned only one by Miller (1754) in genus Pereskia. The orig-
species, here still with the pre-Linnaean phrase inal spelling Pereskia was also maintained by
name “Pereskia aculeata, flore albo, fructu flaves- Linnaeus (1738), Haworth (1812), de Can-
cente…” cited from Plumier (1703). The cur- dolle (1828), Förster (1846), Britton and Rose
rent name Pereskia aculeata was established by (1919) and Werdermann (1933, 1937a). It was
Miller (1768), avoiding the tautonym Pereskia modified by various other authors to “Peires-
pereskia (Figs 4–6). cia” (Zuccarini 1837; Pfeiffer 1837a, 1837b;
For nearly two centuries, the genus Pereskia Salm-Dyck 1850; Rümpler 1886), “Perescia”
was not questioned again, though its circum- (Lemaire 1838), and “Peireskia” (Steudel 1841;
scription changed with the time. The major Schumann 1890, 1898; Vaupel 1925; Berger
“instability” concerned the spelling of the name. 1929), based on the assumption that “Pereskia”
The name was explicitly published as Pereskia was an orthographical error. Backeberg (1934)
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.
4 5 6
7 8
igure 7. Pereskia portulacifolia in dry forest habitat near Jimaní in the eastern part of the Dominican Republic.
F
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.
and Backeberg and Knuth (1936) also cited and purely taxonomical journal Repertorium
it as “Peireskia Plum.” Specierum Novarum, edited in Berlin. One
The debate on the correct spelling acquired can only speculate on the unusual indication
a political touch in the 1930s in Germany, of authorship as “Landgerichtsrat Dölz” (Dis-
when the disagreement stirred emotions among trict court advisor Dölz). Was this the editors’
German and Austrian cactus specialists. In a intention or imposed by the author? Dölz
brief and concise nomenclatural analysis of was a lawyer by profession and amateur cac-
the case, Werdermann (1937a) explained that tologist, president of the German cactus so-
the spelling Pereskia by Plumier (1703) was ciety from 1934 to 1945 in Berlin, and head
apparently intentional and correct as based of the “Zentralforschungsstelle” (central re-
on Miller (1754) and in accordance with the search office) of the German Cactus Society.
International Code of Botanical Nomencla- Fedde’s botanist co-editor Otto Schwarz re-
ture. In the same year Dölz (1937) published sponded simultaneously in a semi-editorial
an entirely opposite view on this and simi- article (Schwarz 1937). It is a rather uncom-
lar nomenclatural issues, favoring Peireskia. mon case of political opposition expressed in
The highly political article appeared—rather a botanical journal and can only be fully un-
surprisingly—in Friedrich Fedde’s private derstood in the light of the divergent biogra-
phies of the two authors. For political reasons
Figure 9. Pereskia guamacho. Leafy twig with reddish Schwarz had lost his position in a national bi-
growing tip and new primary leaves, young, still reddish ological research institute in 1934 with the
spines on the areoles. The basal part is of the previous rise of the “Third Reich” but found tempo-
year, with brachyblast leaves formed on the areoles. 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
Peireskia, following Steudel (1841) against Table 4. “Pereskia” name and images in the WWW

Pereskia Miller (1754) was even submitted found by Google search between 2004 and 2007.
to the International Botanical Congress by Pereskia Pereskia WWW
Janchen (1943). The proposal was not success- WWW entries: entries:
ful (Lanjouw 1952). Having been confronted Date Name Images
with this issue during the preparation of the 16 Jul 2004 5650 236
monograph of Pereskia, Leuenberger (1987)
analysed the case in some more detail, repro- 9 Feb 2005 17800 452
ducing Plumier’s first description and the rel- 14 Jul 2005 20700 576
evant comments by Weber (1898a: 163) in 28 Dec 2005 52900 512
support of the original spelling Pereskia. In- 10 May 2006 92100 518
dependent of nomenclatural rules, Weber had 18 Oct 2006 85200 524
already concluded that, “respecting Plum- 16 Jul 2007 91200 3760*
ier and Linnaeus who certainly did not need
18 Oct 2007 80900 3200*
lessons of Latin,” one ought to maintain the
* Usually only about 10% of search results actually display an image
original orthography. showing a plant of the genus. In July 2007 only ca. 300 of 3760
Not surprisingly, the orthographic variants images showed a pereskia. In October 2007, less than 300 of 3200
of Pereskia (excepting for Pereskea) persist to images showed a pereskia.
some degree, particularly in amateur literature,
and a search in the internet demonstrates the details are remarkable and include a leafy and
current situation (Table 2). Another, entirely spiny stem, one staminate and two pistillate
new variant “Pereskya” has recently turned up flowers, four fruits, and a transverse section
on various websites in Mexico and the Do- of the fruit under the phrase name “Opuntia
minican Republic. The persistent confusion arborescens spinosissima foliis portulaca corda-
on the correct spelling reaches straight into tis.” It was apparently not in cultivation and
the newest scientific publications. Even But- for a long time remained little known. Cac-
terworth and Wallace (2005: 800), presumably tus pereskia, subsequently named Pereskia acu-
by a slip of the pen, repeat the popular myth leata, was a more robust plant in cultivation
that Peireskia was the original spelling. and already widespread as a pot plant in Eu-
Leaving the rather unfruitful discussions ropean gardens in the 18th century.
on spelling aside, let us turn to the history of A closer analysis of the bibliographic cita-
discovery and naming of the species. Among tions in Linné’s Species plantarum shows that
his 21 species of cacti, Linnaeus (1753) distin- these really contain another, third species, which
guished two species today belonging to Pereskia, had in fact been collected and illustrated in
namely Cactus pereskia L., and C. portulaci- the 17th century but was not recognized as dis-
folius L. This means that in his work, nearly tinct from P. aculeata. According to Wijnands
10% of recognized cacti were pereskias (com- (1983) these pre-Linnaean records and illus-
pared to the little more than 1% today). The trations under phrase names (Plukenet 1692,
simple reason for this: pereskias were easier to 1696; Commelin 1697), the latter introduced
collect as herbarium specimens than the more before 1690/1691 from Isla Margarita, Vene-
highly succulent other cacti, which are nor- zuela, show an early introduction of this third
mally underrepresented in herbaria. species, which Linnaeus (1753) still cited under
Although both species had originally been Cactus pereskia. Remarkably, it took nearly
collected by Plumier, the second one (today 150 years until this third species, hidden in
Pereskia portulacifolia, Figs 7, 8), was not pub- the Linnaean classification of Cactus, arrived
lished by Plumier himself. It was already men- again in Europe and was described as Pereskia
tioned and illustrated in Plumier’s manuscript guamacho by Weber (1898b). Weber was quite
“Botanicon Americanum,” of which incom- unaware of the much earlier illustrations and
plete copies were distributed (Hunt 1984). the herbarium records of P. guamacho, which
The original set of illustrations of cacti kept are in fact among the earliest still-extant her-
in the Natural History Museum in Paris was barium specimens of a pereskia (two present
published by Mottram (2002). Surprisingly, in Herb. Sloane at the British Museum [Nat-
Plumier did not yet relate the small tree with ural History] London). Weber referred to a
purslane like leaves with his own genus Pereskia. plant with the local name “Guamacho” from
He noted similarities of the flower with roses, the Orinoco basin in Venezuela but did not
but the fruit led him to classify it as an arbo- cite a herbarium specimen. A neotype speci-
rescent Opuntia. This plate and the text were men from this area was therefore designated
also reproduced by Mottram (2002: 116). The by Leuenberger (1986; Fig 9).
In 1812 Haworth named two species, of from flowering branches. The other species of
which Pereskia grandifolia, with its flowers re- Haworth, P. longispina, an untypified name
sembling a wild rose, is today a well known doubtfully from South America and said to
species in gardens (Figs 10–15). Besides P. acu- be in cultivation in England before 1808, was
leata it is the most widespread species in cul- treated by de Candolle (1828) with doubts
tivation worldwide and is easily grown from as a variety of P. aculeata, with the comment
seed, flowering rapidly on cuttings rooted 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.
10 11
12
13 14 15

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 Miller 8′. P. marcanoi Areces (1992)

1a. P. aculeata Miller ‘Godseffiana’ 9. P. portulacifolia (L.) DC.
2. P. horrida DC. 10. P. quisqueyana Liogier
2a. P. horrida DC. ssp horrida 11. P. bleo (Kunth) DC.
( = P. humboldtii Britton & Rose 12. P. nemorosa Rojas Acosta
var humboldtii) 13. P. sacharosa Griseb.
2b. P. horrida ssp rauhii (Backeb.) Os- 14. P. grandifolia Haw.
tolaza ( = P. humboldtii var rauhii 14a. P. grandifolia Haw. ssp grandifolia
(Backeb.) Leuenberger) ( = P. grandifolia Haw. var grandifolia)
3. P. diaz-romeroana Cárdenas 14b. P. grandifolia Haw. ssp viola-
4. P. weberiana K. Schum. cea (Leuenb.) N. P. Taylor & Zappi
5. P. lychnidiflora DC. ( = P. grandifolia Haw. var violacea
6. P. aureiflora F. Ritter Leuenb.)
7. P. guamacho F. A. C. Weber 15. P. bahiensis Gürke
8. P. zinniiflora DC. 16. P. stenantha F. Ritter
2. Updated alphabetical list of currently accepted names.

Deviating names used by Leuenberger (1986) are added in parentheses.
P. aculeata Miller P. horrida ssp rauhii (Backeb.) Ostolaza
P. aculeata Miller ‘Godseffiana’ ( = P. humboldtii var rauhii (Backeb.)
P. aureiflora F. Ritter Leuenb.)
P. bahiensis Gürke P. lychnidiflora DC.
P. bleo (Kunth) DC. P. marcanoi Areces
P. diaz-romeroana Cárdenas ( = P. sp A of Leuenberger 1986)
P. grandifolia Haw. ssp grandifolia P. nemorosa Rojas Acosta
( = P. grandifolia Haw. var grandifolia) P. portulacifolia (L.) DC.
P. grandifolia Haw. ssp violacea (Leuenb.) P. quisqueyana Liogier
N. P. Taylor & Zappi ( = P. grandifolia P. sacharosa Griseb.
Haw. var violacea Leuenb.) P. stenantha F. Ritter
P. guamacho F. A. C. Weber P. weberiana K. Schumann
P. horrida DC. P. zinniiflora DC.
P. horrida DC. ssp horrida ( = P. humboldtii
Britton & Rose var humboldtii)
3. Frequent synonyms of Pereskia species with cross-reference to accepted names.
Pereskia aculeata var godseffiana Pereskia cubensis = P. zinniiflora
= P. aculeata ‘Godseffiana’ Pereskia godseffiana = P. aculeata ‘Godseffiana’
Pereskia colombiana = P. guamacho Rhodocactus antonianus = Pereskia weberiana
Pereskia corrugata = P. bleo Rhodocactus cubensis = Pereskia zinniiflora
cies. The scanty description and unknown or- “Cactus (Pereskea) bleo” and “Cactus (Pereskea)
igin does not allow for a clear identification, horridus” (Kunth in Humboldt, Bonpland and
though the lanose areoles and fleshy leaves Kunth 1823). Herbarium material of these spe-
could point to Pereskia guamacho. It seems cies extant at Paris and Berlin was discussed in
appropriate to leave it under doubtful names detail by Leuenberger (2002, 2004). The two
(Leuenberger 1986) names were formally transferred to Pereskia
Two new species were discovered by Hum- by de Candolle (1828). Neither species was
boldt and Bonpland, Pereskia bleo in Colom- in cultivation until much later (Figs 16–24,
bia in April 1801 and P. horrida in Peru in P. bleo, and Figs 25–29, P. horrida).
August 1802, published (with this spelling!) as Four new names were published by de Can-
dolle (1829), solely from the plates of Moci- solved the case, but the confusion persisted
ño’s unpublished Flora Mexicana. The dra- in many collections. Pereskia bleo was pro-
matic fate of these plates has been elucidated vided with two additional names, now listed
by McVaugh (1980, 1982). The species de- in synonymy. One is, remarkably, by Weber
scribed there have long been assumed to be (1898b) himself, Pereskia panamensis, due to
from Mexico, whereas all but one were re- erroneous information on flower color. The
ally from other countries, including Nicara- other is P. corrugata by Cutak (1951), clearly
gua (Pereskia opuntiiflora, P. lychnidiflora) and a re-description of P. bleo, caused by misla-
Cuba (P. zinniiflora), as explained by Leuen- belled plants of P. grandifolia in private and
berger (1986, 1988). The first two represent public collections.
Pereskia lychnidiflora, a species ranging from In the second half of the 19th century, new
western Mexico to the Pacific coast of Costa species were described mainly from south-
Rica (Figs 1, 30–34). Pereskia zinniiflora is the ern South America. The first was Pereskia sa-
oldest name for the endemic species of Cuba charosa Grisebach (1879) from the western
(Figs 35–40). The fourth species, P. rotundi- Chaco region, named for its rose-like flowers.
folia, is a leaf bearing cactus from Mexico, but Its vernacular name, sacha rosa, means either
due to the glochids on its areoles (visible in “false rose” or “wild rose” (Figs 41–44). The
the drawing) it belongs to a different genus name is often misspelt but has nothing to do
Pereskiopsis. The Sessé and Mociño plates are with sugar. As pointed out by Leuenberger
now kept at the Hunt Institute for Botani- (1986), the species had in fact been collected
cal Documentation at Pittsburgh. The cac- more than 100 years earlier (between 1747
tus plates were reproduced in color with com- and 1755) in Bolivia by Joseph de Jussieu,
ments by Rowley (1994). but the specimen in the Jussieu herbarium at
It has to be taken into account that the early Paris had remained there under the name Cac-
treatments of Pereskia did not exclusively con- tus pereskia unnoticed, though already men-
tain plants today belonging to the genus. Pfei- tioned as dried specimen (from “Perou”) under
ffer (1837a, 1837b), Mittler (1844), Förster Pereskia by Jussieu (1789). Pereskia sacharosa
(1846), Salm‑Dyck (1850), and Rümpler is one of the hardier species, as also reported
(1886) still included several leaf-bearing species by Mauseth (2001), who found this species
in Pereskia that were excluded from the genus and P. aculeata to endure low temperatures. A
by later monographers. Some were excluded closely related species, sometimes still confused
temporarily and later recognized as belonging with P. sacharosa, is P. nemorosa Rojas Acosta
to the genus again, for instance, Pereskia pitit- (1897), which has a more eastern distribution
ache Karw. ex Pfeiffer (1837a) from Mexico, in the Chaco region (Figs 45–48).
for a long time placed in Pereskiopsis following Only one year later P. nemorosa was given
Britton and Rose (1919), was reinstated as a two more names, P. amapola and P. argentina,
true Pereskia by Boke (1963a). It was actually by Weber (1898b), who was unaware of the
a re-description of P. lychnidiflora. publication of Rojas Acosta.
Early illustrations of Pereskia in cultiva- The modern circumscription of the genus
tion have caused some confusion. Up to 1850 Pereskia dates back to Schumann (1898), who
probably only three species were really in cul- used primarily seed and spine characters rather
tivation in Europe: Pereskia aculeata, P. gran- than leaf characters to classify the subfamilies.
difolia, and P. lychnidiflora, the latter under He placed Pereskia (flat leaves) and Maihuenia
the name P. pititache, of which a seedling was (terete leaves), both without glochids, in sub-
illustrated by Zuccarini (1837). This plant family Pereskioideae. The subfamily is often
does not seem to have established in cultiva- attributed to Schumann, but according to Re-
tion, which is not surprising. Our experience veal (2003) and Thorne and Reveal (2007) it
over 25 years with this species in the botan- had already been formally established by En-
ical garden calls for very high summer tem- gelmann in 1876. Regardless of leaf shape,
peratures and hothouse conditions in win- Schumann placed all species with glochids
ter, necessary for successful cultivation from and having seeds provided with bony aril in
seed to flowering specimens (Figs 30–34). subfamily Opuntioideae, with the only genus
The early records of P. bleo in cultivation are Opuntia. Schumann (1898) described only
probably all incorrect, and Britton and Rose one new species, based on a single herbarium
(1919) noted that all previous illustration in specimen from Bolivia. He dedicated Pereskia
fact showed P. grandifolia, including that by weberiana to his contemporary, the French
Schumann 1890 in the Flora brasiliensis. Brit- (Alsacian born) medical doctor FAC Weber,
ton and Rose (1919) and Weber (1898b) re- author of the Cactaceae entries in the French
Dictionary of Horticulture by Bois (1893– (1919) from the Matto Grosso is now consid-

1899) and other papers on cacti (Figs 49–50). ered a synonym of P. sacharosa, a species with
Weber (1898a) described Pereskia tampicana a wide distribution along the eastern margin
based on a photograph and live plant in the of the Andes from Bolivia to northern Argen-
Berlin Botanical Garden, but this was clearly tina and reaching into Brazil and Paraguay.
a re-discription of P. grandifolia, as already Britton and Rose (1923) coined a new name
recognized by Britton and Rose (1919). It is (P. humboldtii) for Pereskia horrida (Kunth)
the first record of this Brazilian species grow- DC, on the assumption that the latter was il-
ing as a neophyte in Mexico. legitimate. The name Pereskia humboldtii was
In the 20th century, Berlin botanist Gürke used by Leuenberger (1986), but following
(1908) published Pereskia bahiensis based on the correction by Brako and Zarucchi (1993),
an herbarium collection of Ule. It remained P. horrida DC (considered as new name, not
for a long time a little-known species and is combination) has been generally accepted in
curiously missing from the Index Kewensis accordance with the ICBN.
(Figs 51–54). Britton and Rose (1912, 1919, Britton and Rose (1919) reduced the sub-
1923) added five supposedly new species to families to the rank of tribes. Two new genera,
Pereskia, but four are now considered synonyms Pereskiopsis and Quiabentia, were created by
of earlier names, and one was removed from them to accommodate some former Pereskia
the genus. Pereskia colombiana is the same as species with glochids and bony seeds (Britton
P. guamacho from Venezuela. P. cubensis is a and Rose 1919, 1923).
synonym of P. zinniiflora. Pereskia zehntneri Berger (1926, 1929) had a phylogenetic
was later placed in Quiabentia by its authors. approach to taxonomy, discussing evolution-
Pereskia conzattii is the same as P. lychnidiflora, ary lines and levels within the genus based
and this species, ranging over several coun- on characters of the ovary. He placed Pereskia
tries from Mexico to Central America, was aculeata and P. sacharosa in Pereskia subge-
provided with further names by other authors nus Pereskia (as “Eupeireskia”) and all others
at the beginning of the 20th century. It was in a newly proposed subgenus, Rhodocactus.
described as P. nicoyana F. A. C. Weber from In morphological terms, his distinction of
Costa Rica and P. autumnalis Eichlam from ovules at the base of the pistil versus ovules
Guatemala. Pereskia moorei Britton & Rose 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 WWW WWW WWW WWW WWW WWW

or orthographic entries images entries images entries images
variant (“nn”) 20.5.2005 20.5.2005 19.10.2006 19.10.2006 18.10.2007 18.10.2007
Pereskia 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 0
Pereskia 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 0
Pereskia zinniiflora 61 5 228 4 198 1591
Rhodocactus zinniiflorus 10 0 9 0 7 0
Pereskia 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.
sus inferior ovary) is rather questionable and graph (Berger 1926: 4, scheme 1) can be in-
not supported by later studies (for instance, terpreted as an informal proposal of differ-
Boke 1964). But Berger’s cladistical-looking 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.
16 17 18
19 20 21
22 23 24
genus Pereskia and formally only established names subsequently reduced to synonymy,

a division into two subgenera (Berger 1929), are briefly discussed.
treating two names (P. aculeata and P. sa- Pereskia diaz-romeroana Cárdenas (1950)
charosa) under subgenus Pereskia, and four is now sometimes written “diazromeroana” in
names (Pereskia bleo, P. grandifolia, P. autum- newer literature (and, for instance, in TROP-
nalis, and P. conzattii) under subgenus Rho- ICOS), but as the epithet commemorates the
docactus. Edwards and others (2005) refer to Bolivian naturalist Belisario Díaz Romero,
Berger’s idea, but Berger’s informal Rhodocac- the original spelling is appropriate (Fig 55).
tus was actually poorly defined and is hetero- From the same group of Andean species, all
geneous in terms of Edward’s clades, due to occurring in isolated valleys, a new species
the placement of P. grandifolia. and two new varieties were described from
The most detailed anatomical investiga- Peru: Pereskia vargasii H. W. Johnson (1952),
tions, addressing evolutionary theories as well, P. vargasii var longispina Rauh & Backeb., and
are by Boke (1963b, 1964), who presented a P. vargasii var rauhii Backeb. (in Backeberg
new interpretation of the cactus gynoecium, 1957). They were illustrated by Rauh (1958)
explaining the relative position of carpellary and are now considered to belong to the two
tissue and receptacular tissue in ontogenetic subspecies of Pereskia horrida.
development. Berger’s interpretations were Pereskia higuerana (Cárdenas 1964) is a cu-
later opposed by Ritter (1979: 20), based on rious case, because from the description, men-
examination of longitudinal sections of flow- tioning detachable spines in the axils, and
ers of several South American species. Rit- from the illustration it is clearly a Pereski-
ter found the ovary in P. aculeata, P. sacha- opsis, a mainly Mexican genus not native in
rosa and the Andean species, compared to Bolivia (but often cultivated in cactus collec-
P. bahiensis and P. stenantha of the P. gran- tions and in the past much used as grafting
difolia group representing Berger’s subgenera stock for seedlings). The cited type and iso-
Pereskia and Rhodocactus, to be neither su- type specimens of P. higuerana, collected near
perior nor inferior but intermediate, and he La Higuera in “April 1954” by D Candia, a
noted transitional stages, making a clear dis- student of Cárdenas, were never located, and
tinction impossible. this yellow flowered plant remains enigmatic.
Formally it was Knuth, in Backeberg and Some doubts prevail if this might have been
Knuth (1936: 48, 96), who elevated Berg- an error or even a hoax based on a plant from
er’s subgenus Rhodocactus to the rank of cultivation. This hypothesis has some argument
genus, recombining some of the names, ap- based on the existence of a plant identified as
parently based on supposed, rather than ob- Pereskiopsis by Leuenberger (1986), received
served, characters of the gynoecium. Backe- as Pereskia higuerana from the Schönbrunn
berg (1958: 105, 113) only repeated Bergers’s collection in Vienna and said to have origi-
illustrations, and his reclassification of some nated from Cárdenas from Bolivia. On the
species is neither based on investigations of other hand it is perhaps significant that Ritter
the flowers. This becomes significant be- (1980) mentioned and illustrated (though in
cause Backeberg (1958) selected Rhodocac- black and white) a yellow flowering (unnamed)
tus grandifolius as type of the genus, but in variety of Pereskia diaz-romeroana from Bolivia,
fact only the Caribbean species and P. bleo which has not been reported again and is nei-
have clearly epigynous flowers (Leuenberger ther found among the material of FR 904 or
1986). Backeberg (1957) made further sub- other numbers (Eggli and others 1996). Kies-
divisions and classified the remaining species ling (1999) mentions that he found P. diaz-
of Pereskia into two subgenera by creating a romeroana near La Higuera.
subgenus Neopeireskia for the Andean spe- The most remarkable novelties, Pereskia
cies with small leaves. Later authors ques- aureiflora and P. stenantha, based on Ritter’s
tioned the justification of the generic status field research made in 1964 in eastern Brazil,
of Rhodocactus and other subdivisions and were published by him 1979. The two spe-
upheld only one genus, Pereskia (Hunt 1967; cies were apparently not in cultivation until
Bravo-Hollis 1978; Ritter 1979). 1983 (Leuenberger 1989a, b). Pereskia aure-
In the same decades, some remarkable dis- iflora from Minas Gerais is the only purely
coveries of new species were made by Cárde- yellow-flowered species besides P. guamacho
nas (1950, 1964), Ritter (1979), and Liogier (Figs 56, 57). Pereskia stenantha is a remark-
(1980), and the number of published names able case of flower evolution (Figs 58–60). It
increased to about 50 by 1979. The names is evidently the sister species of P. bahiensis,
still accepted today, as well as some of the but it has a remarkable adaptation in its hum-
mingbird flowers (Ritter 1979; Leuenberger cies of the P. portulacifolia alliance, described
1989a). The upright tepals form a flower of by Liogier (1980) from a clone of staminate
tubular shape (but not a receptacular tube like plants at Bayahibe near La Romana, today a
that of other Cactaceae). Another feature is its well-known holiday resort on the south coast
conspicuous nectar chamber. This species is of the Dominican Republic (Figs 61–66).
from the interior of Bahia (Ritter 1979). Col- Looking at the taxonomic status of Pereskia
lection details were given by Eggli and oth- prior to my 1986 revision, the number of
ers (1996), who also confirmed that the type Pereskia species published over different eras
specimens of both names are missing. Only merits comment. The early increase of species
seed samples could be located. numbers in Pereskia until Britton and Rose
One year later, Ritter (1980) published (1919) is notable as compared to the situation
P. sparsiflora, a plant introduced to cultiva- within the family as a whole (Tables 1, 3). The
tion in Europe by Ritter’s 1958 and 1959 greatest increase in species numbers in Cacta-
collections from Bolivia. The name is clearly ceae falls into an era of collectors and authors
a synonym of P. sacharosa, based on FR 640 with commercial interests. It took place between
material still in cultivation, but Ritter’s field 1925 and 1966, in particular due to publish-
number was found to be heterogeneous and ing activities of Curt Backeberg in Germany. In
also included plants from Paraguay belong- contrast, the leafy cacti were never particularly
ing rather to P. nemorosa according to Leuen- attractive for amateurs and no special increase
berger (1986). (For an explanation of Ritter’s of names in Pereskia can be related with com-
complex field number system see also Eggli mercial activities. Backeberg (1963) published
and others 1996). Rhodocactus antonianus based on a plant of un-
A most remarkable discovery was Pereskia known origin in his own collection, an invalid
quisqueyana, a narrowly endemic, dioecious spe- 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).
25 26
27
28 29
Eggli and Taylor 1991). He Figures 30–24. Pereskia lychnidiflora. Figure 30. Shoot apex with

later modified the vague ori- whorl of branches and some brachyblast leaves. Figure 31. Longitudinal
gin data “Northeastern Peru” section of stem of lateral branch less than one year old, with succulent
(Backeberg 1966), but the green pith and cortex, leaves subtending areoles, mature spine with still-
fleshy base. Figure 32. Flowering twig, the lower part one year old with
taxon is undoubtedly iden- brachyblast leaves, the new leafy growth (with reddish colored epidermis)
tical with Pereskia weberiana, terminating in a flower with green fleshy outer tepals and orange, thin,
a species from Bolivia. Taxo- fimbriate inner tepals. The pistil is concealed by the numerous stamens.
nomically, the cactus family Figure 33. Longitudinal section of flower showing gynoecium, elongated
was expanded by Britton and flower axis, large mucilage cells in pith and cortex as well as in the base of
Rose with over 900 new names the outer tepals. Figure 34. Longitudinal section of withered flower after
(Table 3). As noted by Mitich a few days, with receptacle, outer tepals, and ovary roof still fleshy, other
flower parts including the ovules, pistil, stamens, and outer tepals shriv-
(1979), Britton was conserva- elled or dry.
tive in delimiting species but
tended to separate genera liber- 30
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
31 32
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
33 34
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- In the past 20 years only one new species
tivation and originally introduced by Horst of Pereskia has been published: Pereskia mar-
and Uebelmann from more than one locality canoi Areces (1992) from the Dominican
under number HU226 as new variety Pereskia Republic, a taxon closely related to Pereskia
grandifolia var violaca (Figs 67–70). zinniiflora and which I had already recog-
nized as probably new (Leuenberger 1986), others 2006: 5). Some nomenclatural adjust-
treating it provisionally under “Pereskia sp ments and some status modifications have been
A,” for lack of sufficient data. A small plant made. As noted, the most significant change
observed in cultivation over many years in affecting a formerly well-established name is a
Berlin-Dahlem had flowered only once in nomenclatural correction, initiated by Brako
1984, producing a pistillate flower. It flow- and Zarucchi (1993): the replacement of the
ered again in 1993 and 1998 and once pro- well established name Pereskia humboldtii
duced a fruit with some seeds germinating Britton & Rose (1919) by the available older
within the fruit (Figs 71–72). Remarkably, an name Pereskia horrida DC in accordance with
old but sterile herbarium material of a plant the ICBN (Art. 58.1 Ex0.1), where an analo-
cultivated at the Jardin des Plantes, proba- gous example is specifically mentioned (Greuter
bly of the same taxon, exists in the AL Jus- and others 1994; McNeill and others 2006).
sieu Herbarium (13546A) and was cited by Following earlier practice common in Cacta-
Leuenberger (1986). Unfortunately it bears ceae, particularly in American botanical liter-
no further information and can only be ature, I treated infraspecific taxa of P. grandi-
vaguely dated between ca. 1789 and 1850. folia and P. horrida (under P. humboldtii) as
No further hints were found in the informa- varieties (Leuenberger 1986). In the past de-
tion on the Jussieu herbarium provided by cades the treatment as subspecies has become
Stafleu (1964). Perhaps molecular methods more common and promoted (Hunt and oth-
will some day permit us to confirm its geo- ers 2006: 4). Further data on the geographi-
graphical origin and identification. cal distribution led to the new combination
Since the publication of my 1986 mono- Pereskia grandifolia ssp violacea (Leuenb.)
graph only a few changes have occurred. The N. P. Taylor & Zappi (1997). Ostolaza (1998)
taxonomic circumscription of the genus re- formally gave P. horrida ssp rauhii this new
mained stable, and Pereskia is one of the gen- status, but data on this taxon are still scarce.
era of Cactaceae with the highest stability of In current standard works on Cactaceae (An-
names in the past 20 years (see also Hunt and 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.
35 36 37
38 39 40
the 16 + 1 species of Pereskia distinguished by various aspects of Pereskia have been pub-

me in 1986 are accepted as 17 species, with lished in the past 20 years. Only few can be
the same infraspecific taxa and with the up- mentioned here. Much useful information on
dated nomenclature outlined here. various aspects of morphology, anatomy, and
It should be noted that some illustrations in biology can be found in Gibson and Nobel
these standard works bear wrong captions. In (1986). Stem and leaf anatomy was treated
Anderson (2001), Pereskia aculeata on p 566 is by Sajeva and Mauseth (1991) and Maus-
obviously P. grandifolia ssp grandifolia, while eth and Landrum (1997), and leaf anatomy
P. grandifolia ssp grandifolia on p 568 is really of four northern South American and Carib-
ssp violacea. In the German edition of the same bean species by García and others (2000). In
book (Anderson 2005), edited and comple- regional studies, the most significant recent
mented by U Eggli, the eleven illustrations of work including a detailed treatment of five
Pereskia are all correctly identified. The New species of Pereskia is a book on cacti of east-
Cactus Lexicon (Hunt and others 2006) illus- ern Brazil by Taylor and Zappi (2004). This
trates all 17 species, but unfortunately one includes numerous new localities documented
photograph has the wrong caption: Figure by vouchers and cited in detail, distribution
451.2 (captioned P. diaz-romeroana), is cer- maps, photographic illustrations, and a con-
tainly P. weberiana (the clone formerly known servation assessment of each species. Studies
in cultivation as Rhodocactus antonianus). on individual species treated the distribution
A number of papers contributing data on biology of P. aculeata in Brazil (Pedroni and
41 42
43
44
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.
igures 45–48. Pereskia nemorosa. Figure 45. Pruned stem with new spines, new lateral branches emerging
F
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
45 46
47 48
Sanchez 1997), invasive potential of P. acu- Pereskia in the WWW (2004–2007)

leata in South Africa (Moran and Zimmer- Since about 2004 numerous websites world-
mann 1991), conservation aspects and new re- wide have provided scattered information on,
cords of the narrow endemic and endangered and an ever increasing number of images of,
species of the Dominican Republic, Pereskia Pereskia. The rapid increase is shown in the
quisqueyana (García and others 1999; Mejía results of a casual Google search at irregular
and others 2001; Corman 2003). intervals (Table 4). The increase of images (or
Doweld (2004) proposed and named new rather image links) found from summer 2006
infrageneric groups resulting in three sec- onward is striking, but the actual number of
tions, two subsections, and five series, based images displaying the species searched for is
on the informal groups outlined by Leuen- much smaller. In this genus and for many
berger (1986). These named entities are of other plants often only about 10% of images
questionable purpose. The hierarchy does not are of the searched-for species. Before this date,
agree with the cladograms of Nyffeler (2002), the percentage of images actually showing the
Butterworth and Wallace (2005), and the new plant was also variable, and often also lower,
entities thus seem to be little more than a pro- but details were not recorded.
spective name game. The groups do not agree In any case, any search result must be in-
with the ideas of Edwards and others (2005) terpreted with caution and some knowledge
of two main clades. on the group, because search engines cannot
Zimmermann 1991;

49 50
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
Figures 49, 50. Pereskia weberiana. Figure 49. Leafy twig with terminal flower (1986) as 26 cm
bud. Figure 50. Transverse section of young stem, showing conspicuous green cortex, based on herbar-
epidermis with stomata. ium specimens and
live material. New
interpret the content of the image. Even when measurements are documented by a garden
the image is clearly connected with the correct herbarium specimen (Cubr 38344 at B) col-
caption, the identification can still be wrong. lected from a specimen planted in a display
This is more often the case than one might ex- glasshouse under hot and humid conditions,
pect, and not only for plants with unsettled where the plant developed on a vigorous long
taxonomy. The problem is naturally widespread shoot in summer 2000. A main shoot reached
in taxonomic problem groups, when numer- about 3 m height in one summer season, and
ous images are put on the WWW by non-ex- the leaves attained a maximum length of
perts. The search results could not be analysed 42 cm, certainly an impressive record for a
in detail, but there seems to be a tendency to- cactus (Fig 73).
ward an increase in correctly identified images, Pereskia diaz-romeroana. In 2003, holo-
possibly by some sort of self-cleaning effect type material of this species was located in the
stimulated by well-identified plants on reli- Cárdenas collection of Cactaceae deposited in
able websites. However, in some cases there the herbarium of the Instituto Miguel Lillo,
is a proportional increase of wrongly named Tucumán, Argentina (LIL), not accessible at
material (Tables 4, 5). the time of the monograph but now fully cata-
logued by Eggli and Leuenberger (2005). The
Selected updates to the 1986 holotype specimen, Cárdenas 4388, located
monograph on Pereskia there supersedes the lectotype (a duplicate at
Pereskia aculeata. Since 1986 numerous new US) I had selected (Leuenberger 1986).
records from within the previously known Pereskia guamacho. At the time of my-
range of P. aculeata have been added by col- monograph’s publication, I had no informa-
lectors, particularly in Paraguay and NE Ar- tion on the underground part of Pereskia gua-
gentina (see TROPICOS) and also from Bra- macho. The only specimen available to me, at
zil (Taylor and Zappi 2004). In 1996 Pereskia Berlin-Dahlem, did not show any particular
aculeata was reported for the first time from features. Only recently could a root tuber be
the lower Rio Grande valley in Texas (Ideker observed on a seed-grown specimen. Obser-
1996). Pereskia aculeata had earlier been re- vations made in Venezuela, documented by
ported from continental US only from Florida photographs kindly provided by NP Taylor
(Benson 1982). More recently it has arrived and several years ago, show that this species pro-
in 2001 been recorded as a potentially invasive duces very large root tubers developed prob-
species in Hawaii according to Staples (pers. ably by the primary root (Fig 74). These can
comm. Oct 2001) and Hawaiian Ecosystems attain a length of more than one meter and
at Risk project (HEAR; Anonymous 2007). are about 15 cm in diameter on a still-young
The species is invasive in various regions and tree observed with its root system exposed
was declared a weed in 1979 in southern Af- on a roadside cut. Such a massive water stor-
rica, where it has been well studied (Moran and ing root system could be ecologically impor-
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.
51 52 53
54 55 57
56
58 59 60
tant. This massive underground storage sys- that fruit development occurs when the plants

tem was not mentioned in the recent studies are in leaf, the species has also been observed
by Edwards (2006) and Edwards and Diaz with flowers on leafy shoots during the rainy
(2006), who noted that leaf production is not season, as documented by a photograph taken
so clearly related to drought and rain cycles by Jason Grant in Venezuela in 1998.
as hitherto suggested by data extracted from Pereskia zinniiflora. A special feature of
herbarium labels. The phenology of this spe- this species is the presence of large root tu-
cies may deserve further examination, taking bers, first observed late in 1994. In his paper
into account the underground water reservoir dealing in detail with the morphological char-
of the trees. In cultivation, unfortunately, no acters of the species, Areces (1984) did not
flowers could be observed hitherto, even after yet mention the roots of P. zinniiflora. I had
various experiments (though only with one found only fibrous roots in juvenile plants of
single larger specimen) to stimulate flowering. Pereskia zinniiflora (Leuenberger 1986) and
Prolonged drought in summer did not give any could not study this species in the field. But
result. It is possible that this species needs a I did note the possible presence of root tubers
drought period with high temperatures, pos- with a question mark in Table 1 on p 36 of the
sibly combined with shorter daylight hours, monograph. The diagram illustrating evolu-
to stimulate flowering—impossible to achieve tionary groups within the genus already sug-
under our conditions. Even though herbarium gested the possible occurrence of root tubers in
data (some 40 samples studied) suggest that this taxon. Areces, in a later publication, used
flowering occurs mainly from March to May, the character “root tubers absent” in the key
normally while plants are in a leafless stage or to distinguish P. zinniiflora and P. marcanoi
with their first flush of brachyblast leaves, and from P. quisqueyana (Areces 1992).
igures 61–66. Pereskia quisqueyana. Figure 61. in habitat on coral limestone at the type locality in Bay-
F
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
64 65 66
67 68 70
69
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.
The presence of root tubers could be con- Taxonomic history

firmed for P. zinniiflora on a plant cultivated of Maihuenia
in the Botanical Garden Berlin-Dahlem. This The genus Maihuenia was proposed by Philippi
specimen, originally received from the Jar- (1883) and formally described by Schumann
din de Plantes in Paris under number G.563, (1898). The first reports of plants belonging
leg. Alain, originally collected in Cuba, and to this genus date back much earlier. Due to
first grown in a pot, was planted in the bro- their superficial similarity with some sym-
meliad house under hothouse conditions. patric, cushion-forming opuntias, they were
When it had to be removed in 1994, it was first described as species of Opuntia. The
found that it had produced three large root taxonomic history is therefore more com-
tubers of remarkable size, the largest fusiform plex than in Pereskia. Remarkably, maihue-
and 60 cm long and to 14 cm in diameter. nias were not yet collected or illustrated by
The main stem of this plant, fiercely spiny the very first naturalists surrounding south-
and only about 3 cm in diameter, is illus- ern South America, perhaps due to lack of
trated together with one of the three root tu- flowers at the time, or because they had a
bers three years (!) after uprooting (Fig 75). bad reputation caused by the first contacts
In September 1997 the slightly shrivelled with opuntias. One sympatric cactus was
root tuber was still fleshy inside, as demon- recorded before any maihuenia from east-
strated by a small piece cut from the tuber. ern Patagonia in today’s Province of Santa
Kept on a windowsill at room temperature Cruz, Argentina, but remained undescribed.
in the laboratory, it took almost two addi- It was found on the Malaspina expedition
tional years to dry out completely. Although in 1789. According to the illustration by
formation of new root tips was observed at José Guio based on a drawing by Née, an-
the distal end of the rood tuber while it was notated only as “Cactus” (Anonymous 1989:
kept dry, no shoot regeneration occurred. 168 and pl. 14), it was Austrocactus bertinii.
This agrees with an earlier observation on Another cactus, found after but described
detached root tubers of Pereskia weberiana before the first maihuenia, was collected
(Fig 76). These root tubers are apparently on Charles Darwin’s journey in 1833 on
unable to produce buds and new stems even the eastern coast of Patagonia (Port Desire,
if planted in soil, as already noted (Leuen- today Puerto Deseado, Santa Cruz, Argen-
berger 1986, 1992d). tina) and described as Opuntia darwinii by

71
72 73
74 75 76
igure 71. Pereskia marcanoi. Pistillate flower in longitudinal section showing globular receptacle, large ovary
F
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.
Henslow (1837), is now classified as Mai- eppig 1835). In his travelogue, Poeppig de-
hueniopsis darwinii, superficially similar to scribed the plant observed and collected be-
but never considered to be a Maihuenia. tween Yumbel and Antuco in today’s Province
The oldest collection of Maihuenia located of Bio Bío, Chile, very briefly in a single sen-
hitherto can be attributed to the German nat- tence within the text (translated from German:
uralist Eduard Poeppig, who visited Chile in “…and a peculiar cactus (Opuntia caespitosa.
1828. Poeppig’s account on his voyage was Poepp.) extends in broad, light green mats at
published seven years later. The new cactus the driest sites, immediately conspicuous by
was named “Opuntia caespitosa Poepp.” (Po- its sulphur yellow flowers and the silvery white
Table 6. Taxonomic history of Maihuenia.
Published
Accepted combinations
Author Year Accepted species subspecific taxa at species rank
Schumann 1898 3 3
Britton and Rose 1919–23 5 0 6
Vaupel 1925 5 0 6
Berger 1929 5 0 7
Ritter 1980 7 1 11
Kiesling 1988 2 0 11
Leuenberger 1997 2 0 11
Hunt 1999 2 0 11
Anderson 2001 2 0 11
Anderson 2005 2 0 11
Hunt, Taylor and Charles 2006 2 0 11
spines, which stick out everywhere from the Otto (1833), with the entry “208. O. Poeppi-
short club-shaped stem segments.” This brief gii.* Chili.” This indicates that it was already
description very well depicts the species, which grown in Germany from seed sent to Berlin
cannot be confused with any other cactus in by Poeppig before the actual publication of
the area (Figs 77–86). Poeppig’s work. The species must have been
The name Opuntia caespitosa Poepp., how- quite widespread in cultivation according to
ever, cannot be used because the North Amer- Mittler (1844), who mentions it for seven pri-
ican Opuntia cespitosa Rafinesque (1830) pre- vate and public collections, including the bo-
dates it. The first valid and legitimate name for tanical gardens of Berlin, Munich, the Haage
a maihuenia was formally published by Pfei- nursery, and the Salm-Dyck collection.
ffer (1837a). He named Poeppig’s discovery The same species was described once more
independently as “Opuntia poeppigii Otto” as Opuntia maihuen Gay (1847). Gay used
based on seed-grown material. Pfeiffer attrib- the vernacular name (maihuen) as the spe-
uted only the name to Otto. The description cific epithet and noted that it was closely re-
was provided by himself, hence the author- lated to Opuntia poeppigii. When Salm-Dyck
ship remains with Pfeiffer (1837a). The plant (1850) published a new systematic subdivi-
had in fact already been mentioned earlier as sion of the family Cactaceae, he distinguished
a provisional name, without description, in a seven tribes, among them the Opuntieae and
list of the 292 cacti of the Royal Botanic Gar- Pereskieae. He transferred Opuntia poeppigii
den of Berlin compiled by the head gardener, Pfeiffer to the genus Pereskia together with
some other leaf-bearing cacti (several now ex-
cluded from Pereskia and attributed either to
Table 7. Maihuenia, name and images in the WWW Pereskiopsis or to Opuntia).
(2005–2007).
Curiously, a specimen located in the garden
Maihuenia Maihuenia herbarium of the Missouri Botanical Garden
WWW entries WWW images (MO) originating from “hort. Dyck,” dated
23 May 2005 884 112 August 1857 and labelled “Peirescia poeppigii”
is not Maihuenia poeppigii. The long spines
19 Jul 2005 847 109
(to 3 cm long) and conspicuous spur shoots
24 Nov 2005 908 131 identify it clearly as Maihuenia patagonica.
18 Oct 2006 17500 92 It represents the oldest, though still mysteri-
20 Feb 2007 13400 104 ous, record now known of a plant belonging
26 Jun 2007 13600 574* to the second species recognized today: Mai-
16 Jul 2007 13100 633* huenia patagonica (Figs 87–98). But this spe-
18 Oct 2007 13800 751*
cies was also first published as a member of a
different genus: Opuntia patagonica Philippi
* 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 (1863). Its history is interesting but compli-
showed Maihuenia. cated. The first collection, though fragmentary,
was made by a traveller who came—quite re- along the Itata river and in the Cordillera

markably—from the western side of the Andes de Talca (Philippi 1883). His color plate il-
to explore the dry interior of Patagonia. In lustrates vegetative characters, a flower bud,
1862–1863, the Chilean physician Guillermo and a flower of a plant from Chillán. Un-
Cox was the first explorer to bring back bo- fortunately, Philippi did not formally pub-
tanical collections from this part of the inte- lish the new genus and only stated that
rior of Patagonia, territory of the Pehuenche “one could perhaps make a genus of its own”
Indians, giving the origin as “Pampa de Pa- (based on Opuntia poeppigii), which “one
tagonia” (Cox 1863a, b; Middleditch 1988; could name Maihuenia.” This means that
Philippi 1863, 1864). The difficult interpre- he did not formally accept it himself. Ac-
tation of the specimen and its probable geo- cording to the International Code of Botan-
graphical origin were discussed in more de- ical Nomenclature, ICBN Art. 34.1.a and b
tail by Leuenberger and Arroyo-Leuenberger (McNeill and others 2006), the genus was
(1992). In more recent years a search in part therefore not validly published by Philippi.
of the area visited by Cox West of the Río Many subsequent authors nevertheless at-
Collon Curá in Neuquén was in vain. The tributed the genus to Philippi (1883), but it
area east of the river towards the Río Limay was only formally established 15 years later
remains the most probable site of origin, and by Schumann (1898). It can be cited with
the epitype I selected (Leuenberger 1997) is the authorship “Philippi ex K. Schum.” to
in fact from this area. give credit to Philippi, who recognized the
None of these early introductions (herbar- difference from Opuntia and who proposed
ium specimens and plants grown from seeds) the generic name based on the vernacular
were illustrated in contemporary literature. name maihuén (the significance of which
The first illustration of a maihuenia is in a is not quite clear).
horticultural source in Germany. It is a wood Opinions on the acceptance of the genus
cut of an imported, potted plant, published varied at the time. FAC Weber (1898b), for
in 1881 in the journal Gartenflora under the instance, did not accept Maihuenia as genus.
name “Peireskia Pöppigii Salm” (Regel 1881). He treated two species under Pereskia, sub-
According to Philippi (1883), this illustra- genus Maihuenia, but stated that it should
tion was taken from an earlier publication rather form a distinct genus. He attributed
in a catalogue of the German cactus nurs- the name to Philippi. Of the little-known spe-
ery Haage and Schmidt. It was only in 1883 cies described earlier, Weber recognized only
that German born botanist RA Philippi in Pereskia poeppigii. Simultaneously, he described
Chile, using the vernacular name maihuén, a supposedly new species, Pereskia philippii
proposed the genus Maihuenia in an article (prospectively adding Maihuenia philippii as
on “Opuntia poeppigii Otto.” He based his synonym!) from Chile, dedicating it to the
description on plants he observed in its nat- author of the proposed genus.
ural habitat in Chile, at the Volcan Chillán, This ambivalent treatment by Weber and
Table 8. Maihuenia names and images of species in the WWW (2005–2007).
Maihuenia poeppigii Maihuenia patagonica

WWW entries WWW images WWW entries WWW images
23 May 2005 202 43 181 37
19 Jul 2005 313 40 261 38
24 Nov 2005 258 52 213 45
18 Oct 2006 1450 43 521 26
20 Feb 2007 2010 40 1490 26
26 Jun 2007 2150 122 801 69
(only ca. 20) (only ca. 20)
16 Jul 2007 2100 90 711 66
28 Sep 2007 1840 125 1550 107
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.
by Schumann has caused confusion on the tioideae), and before his genus no. 20, Pereskia
authorship. When Schumann began to pub- (Pereskioideae). Schumann (1898) accepted

lish his comprehensive monograph of Cac- three species: M. poeppigii, M. brachydelphys
taceae, issued first in several instalments in (new), and M. philippii. Whereas Schumann
1897–1898, he did not yet consider Maihue- (1899) maintained the placement of the genus
nia a genus: it does not appear in the generic in the subfamily Pereskioideae, based on the
key on p 31 at the beginning of his mono- seed characters and the lack of glochids, his
graph (Schumann 1897). However, he later later treatment in the appendix to the 1897–
inserted Maihuenia in the treatment of the 98 monograph is again ambivalent (Schumann
subfamily Pereskioideae in the text on p 754, 1903: 15). Here he affords Maihuenia an in-
where he stated that Maihuenia was justified termediate position and places the genus in
as genus, and where he formally established it Opuntioideae, but he calls it an intermediate
(Schumann 1898). Schumann placed Maihue- form due to the black, brittle seed coat. This
nia in the subfamily Pereskioideae with a re- may have been influenced also by his (errone-
mark on the transitional position, as genus no. ous) indication of the presence of glochids.
19a after his genus no. 19, Pterocactus (Opun- In the 20th century, a series of new taxa and
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.
77 78
80 81
79 82 83

84 85 86
igures 84–86. Maihuenia poeppigii. Figure 84. Germinating seedlings of one to few days (germination took
F
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).
re-interpretations of species names began with cation solely based on seed characters, in which
the publications of the Argentine botanist he placed Maihuenia in a tribe comprising all
Spegazzini (1902). His descriptions of Mai- cacti with “soft seeds” in contrast to the ones
huenia tehuelches from Santa Cruz and M. val- with a bony seed coat (the latter coinciding
entinii from Chubut are each based on two with the opuntioids of Schumann and tribe
or three herbarium collections of contempo- Opuntieae of Britton and Rose). He accepted
raneous collectors. They are meticulous, giv- Maihuenia as second genus after Pereskia in
ing detailed measurements of specimens with his monographic series. Vaupel (1925, 1926)
flowers and immature fruit. Spegazzini did was able to publish only two volumes (includ-
not know the plants from the field. The type ing the genera Pereskia, Maihuenia, Rhipsalis
specimens were illustrated by Leuenberger in volume 1; and Rhipsalis, Wittia, Epiphyl-
(1997). The moderate increase of names is lum, and Aporocactus in volume 2) before his
shown in Table 6. tragic early death. His classification was not
Britton and Rose (1919: 8) placed the followed by later authors.
genus Maihuenia in their tribe Opuntieae Berger (1926, 1929) placed Maihuenia in
(not in Pereskieae!) although they noted the subfamily Pereskioideae and distinguished five
similarity of the seeds with Pereskia and the species but treated in some detail only the two
lack of glochids (an exception within Opun- that he knew from cultivation, M. poeppigii and
tieae) but considered it “otherwise very dif- M. patagonica. Backeberg and Knuth (1936)
ferent [from Pereskia]”. They recognized five listed five species and provided the first pho-
species, among them Maihuenia patagonica. tograph of a flower of Maihuenia patagonica.
Their treatment is somewhat confusing be- Marshall and Bock (1941) mentioned Mai-
cause the illustration of Maihuenia valenti- huenia poeppigii as desirable for cultivation
nii, a photograph received from Spegazzini, and obtainable in trade.
in fact shows M. poeppigii. Britton and Rose Castellanos and Lelong (1943) published
also erroneously placed M. philippii in the syn- only a general account, without details on the
onymy of M. patagonica. The nomenclatural species, but theirs is accompanied by a fairly
problems resulting from this mix-up were dis- accurate distribution map. Buxbaum (1951)
cussed and resolved only much later by Pichi- never saw maihuenias in the field, nor proba-
Sermolli and Bizzarri (1978). bly in cultivation, but he quite accurately rec-
In a more exhaustive revision, though still ognized from published illustrations the two
based exclusively on herbarium material and growth habits representing the two species of
following much the same line of Britton and Maihuenia accepted today. In 1954 and 1955
Rose, the Berlin-Dahlem botanist Vaupel (1925) Friedrich Ritter collected seeds of Maihuenia
also distinguished five species. He accurately in Chile and Argentina. These were distributed
illustrated M. poeppigii and M. tehuelches but through the seed catalogues of Ritter’s sister
excluded the older name Opuntia patagonica, Hildegard Winter in Germany (Winter 1955,
stating that the description was insufficient. 1956, 1957). In her catalogue of 1957 two new
Vaupel tried to introduce a new tribal classifi- taxa were mentioned as nomina nuda (naked
names): “M. albolanata Ritter sp. nov.” and SGO; and by Roig (1963), who discussed the
its variety “viridulispina Ritter var. nov.” These northern limit of the geographical distribu-
were validly published much later (Ritter 1980) tion of the genus in Mendoza.
but seem to represent no more than variations Hunt (1967) treated Maihuenia as second
in spine color (Leuenberger 1997). genus of Tribe Pereskieae in Hutchinsons’
Significant information on Maihuenia is Genera of Flowering Plants. Buxbaum (1969)
found in publications of Argentine and Chil- concluded that Maihuenia was very close to
ean botanists, for instance, by Ruiz Leal (1956) Pereskia in seed and flower structure but dis-
on habitats of Maihuenia in Mendoza, on tinct in growth habit. Kiesling (1975) pro-
drought resistance, seed germination, and pol- vided key characters for the genus, a generic
len; by Muñoz Pizarro (1959, 1960, 1966) on description, and illustrations of a branchlet
the vernacular names of Maihuenia in Chile of Maihuenia sp (M. patagonica) including
and the early herbarium specimen of Cox at flower bud and internal flower structure. An-
igures 87–90. Maihuenia patagonica. Figure 87. In habitat in the Portezuelo del Viento in southern Men-
F
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.
87
88 89 90
atomical characters of two species were stud- somewhat allusively as “Maihuenia poeppigii’s

ied in detail by Gibson (1976, 1977). In the best kept secret.” Jonkers, who never saw the
second, most substantial paper on the subject, plant there, just as presumably no one else
Gibson also discussed the systematic position ever did, asks “how did the plant get that far
of Maihuenia and concluded that vegetative south?” The answer is still lacking, and the
features negate its inclusion in the subfamily report, with attractive (but in terms of the
Opuntioideae. geographical distribution, misleading) illus-
The number of published species increased trations, is apparently based on hearsay rather
again when Ritter (1980), in the second vol- than on facts. Substantial information was
ume of his book on the cacti of South Amer- contributed by Mauseth (1999), who studied
ica, described three new species and one new anatomical adaptations to xeric conditions in
form from Argentina, all based on his collec- Maihuenia. A well-illustrated account on mai-
tions made in 1955 (listed by Eggli and oth- huenias in the field was prepared by Kümmel
ers 1996). and Klügling (2005).
Earlier detailed biological observations are Altitudinal distribution is sometimes erro-
scarce. Only Jung (1982) gave an accurate and neously reported as Patagonia and High Andes,
well-illustrated account on the habitat, roots, but this is not quite correct, as Maihuenia oc-
and fruit of Maihuenia poeppigii in Chile. In curs predominantly in lowland and medium
the Flora Patagonica Kiesling (1988, 1990) re- altitudes, rarely occurring over 2000 m in the
duced the ten species names available at the northernmost part of the range (where the
time to only two, M. patagonica and M. po- Andes reach 3000–5000 m altitude). I anal-
eppigii. The treatment was based on herbar- ysed the myths surrouning high-Andean lo-
ium collections made for the Flora Patagon- calities (Leuenberger 1997), which date back
ica project and personal field experience. Both to Backeberg and freely copied later. The dis-
species are illustrated in line drawings with- crepancy concerning the altitude between the
out flower, fruit, or seed. Maihuenia poeppigii herbarium labels of the lost type specimen (Ar-
was illustrated in a more representative color gentina, [Mendoza], Paso Cruz, 2600 m, Kun-
plate including flower and fruit by Hoffmann tze s.n. (B)) and the neotype from the same
(1989). Hunt and Taylor (1990), and Hunt area (“Paso Cruz 34° Cordillera,” 1500 m, 01.
(1992) listed Maihuenia as a genus with two 1892, Kuntze 109”) may be also significant.
accepted species. Both species occur on a variety of substrates.
Field observations and notes on the morphol- It was only in 2003 that I had an opportunity to
ogy and ecology in Argentina and Chile were visit the impressive habitat in the Payunia vol-
provided by Leuenberger (1992a, 1992b) and
Kraus (1993, 1994, 1995). Wallace (1995a, b)
analyzed molecular data suggesting Maihue- Synonyms of Maihuenia with reference
nia should be treated as a monogeneric sub- to accepted name.
family of Cactaceae. This prompted Fearn
(1996, unpaged) to publish the new subfam- Maihuenia
ily Maihuenioideae. The new subfamily was M. albolanata = M. patagonica
supported by later molecular studies (Nyffeler M. albolanata forma viridulispina = M. patagonica
2002) and accepted in standard works (An- M. andicola = M. patagonica
derson 2001, 2005). M. brachydelphys = M. patagonica
In my 1997 monograph, the genus was M. cumulata = M. patagonica
completely revised on the basis of 275 herbar- M. latispina = M. patagonica
M. philippii = M. poeppigii
ium collections from 32 herbaria, including
M. tehuelches = M. patagonica
55 of my own collections, field studies made M. valentinii = M. patagonica
between 1985 and 1992 in Argentina and
Chile, and observations of seed-grown plants Opuntia
in cultivation (Leuenberger 1997). O. caespitosa = M. poeppigii
O. maihuen = M. poeppigii
Notes on the distribution O. patagonica = M. patagonica
of Maihuenia O. philippii = M. poeppigii
A purely hypothetical and most certainly er- O. poeppigii = M. poeppigii
roneous report of a southernmost occurrence Pereskia
of Maihuenia poeppigii at Torres del Paine Na-
tional Park, some 1400 km south of its known P. philippii = M. poeppigii
P. poeppigii = M. poeppigii
distribution, was published by Jonkers (1998),
91 92 93 94
95 97 98
96
igures 91–98. Maihuenia patagonica. Figure 91. Stem with leaves in winter at temperature far below freez-
F
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.
canic region in Mendoza, Argentina, where vast feld and the Argentine collector Oreste Mar-
areas covered by lava pebbles (lapilli) bear only soner in 1936, it was hardly conceivable that
very sparse vegetation but are dominated by large this taxon would cause so much discussion
cushions of Maihuenia patagonica (Fig 2). on evolutionary lineages in Cactaceae seventy
years later. Blossfeldia liliputana was published
Maihuenia in the by the Berlin-Dahlem botanist Werdermann
WWW (2005–2007) (1937b) based on a single collection from
Maihuenia is still moderately well represented Tumbaya, Province of Jujuy, Argentina. Bloss-
in the WWW, surprising for a relatively little- feld considered the plant to be exceptionally
known genus that is not widespread in culti- rare (Werdermann 1937b). Blossfeldias grow
vation. But the quality of some illustrations well-camouflaged in rock fissures and are cer-
exceeds that in other published sources, and tainly very difficult to find in the natural hab-
the number of good images from habitat is itat, but Ritter (1980) found it to be one of
rapidly increasing (Tables 7, 8). the most widespread cacti of Bolivia, and we
now know it to be far from rare. Its double
Taxonomic history of Blossfeldia reputation as a threatened and non-threat-
When the first plants of this genus were dis- ened plant is evident in conservation liter-
covered by the German collector Harry Bloss- ature, where Blossfeldia serves as an illustra-
Figures 99–101. Blossfeldia liliputana. Figure 99.

Habitat in the Quebrada de Humahuaca, Jujuy, Ar- 99
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.
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 100
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 101
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 to gardens in central Europe can thus be nar-
arrival of these cacti via England or Scotland rowed to 1829 onward.
Some other, also small and rather inconspic- yons and in rock crevices on steep hillsides.
uous cacti growing within the range of Bloss- Friedrich Ritter collected numerous herbarium
feldia were found and described much earlier samples for his private herbarium and seeds
than Blossfeldia, perhaps due to their habitat and for sale through the seed catalogues issued in
the more showy red flowers. Examples are Re- Frankfurt-Fechenheim, Germany, by his sister
butia pygmaea and Opuntia subterranea, found Hildegard Winter (Winter 1954, 1959). The
by the Swedish botanist RE Fries in 1901 on first ones were distributed as Blossfeldia lili-
the Altiplano of Jujuy in northern Argentina. putana in 1959—three from Bolivia, as sup-
Another small cactus, Rebutia steinmannii, was posed new species, at first under unpublished
collected by Steinmann around 1902 in Bolivia, names (see also Ritter 1980).
but Blossfeldia apparently escaped the attention The commercial interest prompted the de-
of these and other early collectors. Some very scription of further insufficiently known new
earlier travellers in NW Argentina and Bolivia taxa (Table 9). Blossfeldia campaniflora Backe-
probably came close to (or even stepped over) berg (1959a, 1959b) was based on a Fechser
Blossfeldia habitats. For Bolivia one could men- collection received in 1954 with the vague in-
tion D’Orbigny around 1842, Weddell 1845– dication of origin “North of Argentina“, and
1848, Fiebrig 1903–1904, and Buchtien from Blossfeldia fechseri Backeberg (1962),whose
1906 onward. In Argentina, besides Gillies, it indication of origin was “ca. 800 km further
was Tschudi in 1859 in Catamarca, and Claren to the south than the type of the genus.” The
in 1901 in Jujuy, who collected in the area. reason for this intentionally vague data was
Information on early botanical collectors was evidently the commercial interest to withhold
compiled by Fries (1905) and Funk and Mori exact locality data. Both names remained in-
(1989). I am not aware of any unrecognized valid according to the International Code
older herbarium record of Blossfeldia, and it of Botanical Nomenclature, ICBN Art. 37.1
appears that it is a true twentieth century dis- (McNeill and others 2006) for lack of indi-
covery by commercial plant explorers. cation of the type collection (see also Eggli
The first local botanists to provide more in- 1985). Blossfeldia fechseri was later mentioned
formation on Blossfeldia after 1937 and citing by Fechser (1965) for the province of Cata-
additional herbarium material in Argentina marca, again without locality, in a note on the
were Castellanos and Lelong (1943). Krainz cultivation of imported plants.
(1949a) gave an account of successful culti- Ritter’s numerous collections made between
vation and described and illustrated the fruit 1953 and 1959 in Argentina (Prov. Salta, Jujuy)
and seed. Notes on the introduction and cul- and Bolivia (Departments of Potosí, Chuqui-
tivation soon appeared in cactophile journals saca, and Cochabamba) were well documented.
(Johnson 1954; Krainz 1941, 1946, 1949b; The localities of his field numbers and some
Steeg 1955). new taxa were published by him (Ritter 1965,
German collectors with commercial interests 1980). All of these are currently considered to
then discovered more populations of Blossfeldia be synonyms of Blossfeldia liliputana. Locality
in other regions of Argentina. Between 1954 data for all of Ritter’s often very fragmentary
and 1962 Fechser sent living plants to Curt herbarium samples were catalogued in detail
Backeberg and reported on it in the US cac- by Eggli and others (1996).
tus journal (Fechser 1960). Backeberg (1959a, The first fairly detailed morphological study
1959b, 1962) treated the known and new of the flower, fruit, and seeds of Blossfeldia lili-
taxa. It is remarkable that the 50 year cumu- putana was made by Buxbaum (1964), who
lative index of the US journal indicates only also summarized the scattered information
five entries on Blossfeldia between 1929 and available on the genus. Doubts on the jus-
1979. Before 1960, Blossfeldia was mentioned tification of the other five published species,
and illustrated there only twice (Anonymous which are hardly distinguishable in cultivation,
1949, Johnson 1954). led Krainz (1975) to reduce them to varieties
The first records from Bolivia are rather late, of Blossfeldia liliputana. Krainz did not know
from 1953 onward, thanks to the travels of the plants from the field but had at least five
Friedrich Ritter. Ritter (1980) noted that he accessions with different names in cultivation
had also missed them on his first trip to Bolivia. at the Zurich Municipal Succulent Collection
A new variety, B. liliputana var caineana was (Krainz 1967), all acquired after 1957.
published by the Bolivian botanist Cárdenas The southernmost blossfeldias in Men-
(1964). Blossfeldia is apparently quite wide- doza were already mentioned by Buining
spread and frequent in Bolivia but equally re- (1969), who had observed them on the way
stricted to the special habitat in narrow can- from Mendoza to Uspallata thanks to a local
igures 102–104. Blossfeldia liliputana. Figure 102.

F contact, Dr Gomez Ruepel of Mendoza. Rit-

Several small plants on fine soil accumulation in fis- ter (1980) comments on them as well, and
sures of the schist. Damaged plant with exposed the same habitat was reported again as new
root in the center. Figure 103. Small heads (seed- by Méndez (1983), who added detailed and
lings or regenerated heads) in a moist rock fissure.
Figure 104. Rock fissure with root buds on a partially new observations on the biology of the plant.
decayed plant, and seedling. Both Ritter (1980: 433) and Méndez (1983)
think that the large strophiole of the seed per-
102 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-
103 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)
104 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
simultaneously that Blossfeldia might be a de-

rived member of the Maihuenioideae, an op-

105
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- 106
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 107
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 Figures 105–107. Blossfeldia liliputana. Figure 105.
an experiment in the laboratory, believe ants Uprooted specimen showing underground ramifica-
to be the active agents transporting the seeds. tions, the massive succulent roots representing the
My own chance observation of ants and fruit- major volume of the whole plant. Figure 106. Old
head of a cultivated specimen, with flower. Plant body
ing Blossfeldia near Fiambalá in Catamarca showing effect of drought (Leuenberger and Arroyo
do not support (though neither do they dis- 3579). Figure 107. Whole plant of the same, with
prove) the latter theory. In a single case ob- fruit. Oldest heads are ca. 20 years old. All photos by
served, an ant coming by a fresh, manually the author unless otherwise stated.
Table 9. Taxonomic history of Blossfeldia.

Total of published
combinations at
Accepted Accepted species rank in
Author Year species subspecific taxa the same year
Werdermann 1937 1 0 1
Krainz 1975 1 6 5
Ritter 1980 4 4 6
Eggli and Leuenberger 1999 1 0 6
Hunt 1999 1 0 6
Anderson 2001 1 0 6
Anderson 2005 1 0 7
Hunt, Taylor and Charles 2006 1 0 7
split fruit of Blossfeldia eagerly absorbed the have been no reports on such experiences
moisture available from the still fleshy funi- with Blossfeldia in cultivation, presumably be-
cles but did not carry away any seeds. Much cause nobody takes the risk of ending up with
still remains to be studied on the biology of non-resurrected plants. Experience with some
Blossfeldia. Longer distance transport by birds ferns, selaginellas, and Myrothamnus flabelli-
does not seem entirely impossible (compare folius shows that they do not like to dry out
the slovenly plover mechanism, a remarkable entirely in cultivation, and the special fea-
case of long distance dispersal, with which ture of true resurrection of the whole plant is
Larrea reached the northern hemisphere 4– not “on display” in the flower pot in gardens.
8 million years ago, according to Wells and The report of poikilohydry for Blossfeldia still
Hunziker (1976) and Lia and others (2001). merits a critical analysis and further investi-
Still purely hypothetical, an exozoochorous gation. The comparatively enormous under-
transport, by seeds with their minute papil- ground amount (often more than 90%) of
late processes clinging to feathers, seems pos- fleshy root and stem tissue would seem to be
sible as well. This has not yet been analysed, an argument against complete drying out of
and near to nothing is known on flower and Blossfeldia in habitat. What can be observed
fruit biology in the genus. in nature (Ritter 1980) and also in cultiva-
Barthlott and Porembski (1996) state that tion under drought conditions is the marked
Blossfeldia is a poikilohydrous plant. There shrinking of the green part of the plant. This
Table 10. Blossfeldia names and images in the WWW (2005–2007).
Blossfeldia Blossfeldia
Blossfeldia Blossfeldia liliputana liliputana
Date WWW WWW Images WWW WWW images
19 Jan 2005 508 192 513 77
17 May 2005 571 185 450 93
15 Jun 2005 4160 205 883 96
19 Jul 2005 3930 205 904 96
19 Dec 2005 13400 197 526 85
10 May 2006 33800 206 634 87
26 Jun 2007 21900 1750* 2670 346*
28 Sep 2007 24700 1960 2030 406
18 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.
Synonyms of Blossfeldia liliputana. a selection of names on 28 Sept 2007 (Bloss-

feldia campaniflora, name entries 137 / images

Blossfeldia atroviridis 187; Blossfeldia fechseri, 180 / 32; Blossfeldia ped-
B. campaniflora icellata 146 / 31, Blossfeldia minima 265 /28;
B. fechseri Blossfeldia flocculosa 69 / 4).
B. flocculosa
A postcript on Pereskia, Maihuenia,
B. minima
and Blossfeldia in the WWW
B. pedicellata
Parodia liliputana More restricted and substantial information
is found by Google scholar search, made for
all three genera at a late stage of this study
can also happen when blossfeldias lose part (24 October 2007). Numerous links to sci-
of their roots due to excessive watering and, entific papers were found, 746 referring to
unable to take up water, are affected by sub- Pereskia, 106 to Maihuenia, and 51 to Bloss-
sequent drought. The remaining plant body feldia. The image links are apparently not
then loses much water, regeneration is slow, more restricted and thus not more reliable
and part of the cortical tissue does not re- than general Google searches. The result
cover. The new growth then appears within was: Pereskia 3540, Maihuenia 811, Blossfel-
the sunken part of the older stem apex (Fig dia 1910, with very varied image links and
106). Blossfeldias have an impressive capac- the same low percentage of actual and cor-
ity of regeneration of lost stem parts and ap- rect images.
parently produce new shoots not only from
damaged stem parts but also from exposed Acknowledgments
roots, as reported by Méndez (1983). In any I wish to thank Michael Rodewald for image
regard a remarkable plant. scans, the horticulturists of the Berlin-Dahlem
Botanic Garden for taking care of the live
Blossfeldia in the WWW plants over many years, Mrs Marion Cubr for
Blossfeldia is comparatively well represented on preparation of garden herbarium specimens,
the internet due to the interest of amateur cactus Dr Nigel Taylor (Kew) for photographs of the
growers (Table 10). No repeated searches were root system of Pereskia guamacho, an anon-
made of synonyms and nomina nuda, but the ymous reviewer for extensive detailed com-
overall pattern is much the same as for other ments, and Dr Silvia Arroyo-Leuenberger for
popular cacti, according to one search made for help with the manuscript.
Literature Cited
Anderson EF. 2001. The cactus family. Timber Areces AE. 1984. Un raro caso de diclinia en
Press, Portland. Rhodocactus cubensis. Rev Jard Bot Nac Univ
Anderson EF. 2005. Das große Kakteen-Lexikon. Habana 5(2): 151–171.
Ulmer. Stuttgart. Areces Mallea AE. 1992. Pereskia marcanoi,
Anonymous. 1827. Cacti Lin. Index specierum a new species of Cactaceae from Hispaniola.
horti botanici Berolinensis. Verh. Vereins Be- Brittonia 44(4): 423–428.
förd. Gartenbaues Königl Preuss Staaten 3: Backeberg C. 1934. Cactaceae. Systematische
430–432. Übersicht–Systematic Synopsis. Blätter für Kak-
Anonymous. 1830. Cacti Lin. Index specierum teenforschung 1934–3 (unpaged, 4 pp).
horti botanici Berolinensis 1829. Verh. Ver- Backeberg C. 1957. (“1956”). Descriptiones Cac-
eins Beförd. Gartenbaues Königl Preuss Staaten tacearum novarum. Gustav Fischer, Jena.
6: 429–435. Backeberg C. 1958. Die Cactaceae. Handbuch der
Anonymous. 1949. From H. Krainz, Zurich. Cact Kakteenkunde, vol 1. G. Fischer, Jena
Succ J (US) 21: 103–104. Backeberg C. 1959a. A new Blossfeldia (B. cam-
Anonymous. [1989]. La Botánica en la Expedición paniflora Backeb.). Cact Succ J Great Brit. 21:
Malaspina 1789–1794. Colección Encuentros, 32, 36.
Turner, Madrid. Backeberg C. 1959b. Die Cactaceae. Handbuch
Anonymous. 1999. Se descubre planta hembra de der Kakteenkunde, vol 3. G. Fischer, Jena.
Rosa de Bayahibe. Noti-Flora 4: 1. Backeberg C. 1962. Die Cactaceae. Handbuch der
Anonymous. 2007. HEAR mystery plants, Hawai- Kakteenkunde, vol 6. G. Fischer, Jena.
ian Ecosystems at Risk (HEAR Project). Pub- Backeberg C. 1963. Descriptiones Cactacearum
lished on the internet: http://www.hear.org/ novarum III. Gustav Fischer, Jena.
mysteryplants/autogendhtml/mysteryplant5. Backeberg C. 1966. Das Kakteenlexikon. G. Fischer,
htm (accessed 29 Oct 2007). Jena.
Backeberg C, Knuth FM. 1936. (“1935”). Kak- Castellanos A, Lelong HV. 1943. Opuntia-

tus-ABC. Nordisk Forlag, Copenhagen. les vel Cactales. Cactaceae. In Descole H, ed-
Barthlott W, Porembski S. 1996. Ecology and itor. Genera et Species Plantarum Argentina-
morphology of Blossfeldia liliputana (Cactaceae): rum 1: 47–142 and pl. 1–62. Guillermo Kraft,
a poikilohydric and almost astomate succulent. Buenos Aires.
Bot Acta 109: 161–166. Commelin J. 1697. Horti medici Amstelodamen-
Berger A. 1926. Die Entwicklungslinien der Kak- sis rariorum plantarum descriptio et icones. Pars
teen. G. Fischer, Jena. prima. Amstelodami.
Berger A. 1929. Illustrierte Handbücher sukkul- Corman PH. 2003. Bayahibe: quelques cactus au
enter Pflanzen: Kakteen. Ulmer, Stuttgart. pays du tourisme. Succulentes 2003(3): 3–9.
Bois D. 1893–99. Dictionnaire d’Horticulture. Cox GE. 1863a. Jeografía de Chile. Viaje a las re-
Klincksieck, Paris. jiones septentrionales de la Patagonia. Anales
Boke NH. 1963a. The genus Pereskia in Mexico. Univ. Chile 23: 1–103, 151–238, 437–509.
Cact Succ J (US) 35: 3–10. Cox GE. 1863b. Viaje en las rejiones septentrio-
Boke NH. 1963b. Anatomy and development of nales de la Patagonia (1862–1863). Imprenta
the flower and fruit of Pereskia pititache. Amer Nacional. Santiago de Chile (reprinted by El
J Bot 50: 843–858. Elefante Blanco, Buenos Aires 1999).
Boke N. 1964. The cactus gynoecium: a new in- Crozier BS. 2004. Subfamilies of Cactaceae Juss.,
terpretation. Amer J Bot 51: 598–610. including Blossfeldioideae subfam. nov. Phy-
Brako L, Zarucchi JL. 1993. Catalogue of the tologia 86: 52–64.
flowering plants and gymnosperms of Peru. Cutak L. 1951. A new scarlet Pereskia. Cact Succ
Monogr Syst Bot Missouri Bot Gard 45. J (US) 23: 171–173.
Bravo-Hollis H. 1978. Las Cactáceas de México, Dawe O. 1988. Piedras de Afilar. The unful-
1. UNAM, México. filled dream of a 19 th century Scottish bota-
Britton NL, Rose JN. 1912. Undescribed spe- nist in South America. Charles Skilton Ltd.,
cies of Cuban cacti. Torreya 12: 13–14. Haddington.
Britton NL, Rose JN. 1919. The Cactaceae, Dölz B. 1937. Botanische Nomenklaturregeln im
Vol 1. Carnegie Institution, Publ. no. 248, Spiegel unserer Weltanschauung. Repert Spec
Washington. Nov Regni Veg 42: 185–198.
Britton NL, Rose JN. 1923. The Cactaceae, Doweld AB. 2004. A system of classification of
Vol 4. Carnegie Institution, Publ. no. 248, the genus Pereskia Mill. (Pereskioideae, Cac-
Washington. taceae). Byull Moskovsk Obshch Isp Prir, Otd
Buining AFH. 1969. Bei den Kakteen Argenti- Biol 109(4): 44–49.
niens. Kakteen Sukk 20(9): 169–172. Edwards EJ. 2006. Correlated evolution of stem
Butterworth CA. 2006. Resolving “Nyffeler’s and leaf hydraulic traits in Pereskia (Cactaceae).
Puzzle”—the intriguing taxonomic position of New Phytol. 172(3): 479–789.
Blossfeldia. Haseltonia 12: 3–10. Edwards EJ, Diaz M. 2006. Ecological physiol-
Butterworth CA, Wallace RS. 2005. Molecular ogy of Pereskia guamacho, a cactus with leaves.
phylogenetics of the leafy cactus genus Pereskia Plant, Cell Environ 29: 247–256.
(Cactaceae). Syst Bot 30: 800–808. Edwards EJ, Donoghue MJ. 2006. Pereskia
Buxbaum F. 1940. Blütenmorphologische Einze- and the origin of the cactus life form. Ameri-
luntersuchungen. Pereskia sacharosa. Cacta- can Naturalist 167: 777–793.
ceae 1940: 1–3. Edwards EJ, Nyffeler R, Donoghue MJ. 2005.
Buxbaum F. 1951. Morphology of cacti. Section I. Roots Basal cactus phylogeny: Implications of Pereskia
and stems. Abbey Garden Press, Pasadena. paraphyly for the transition to the cactus life
Buxbaum F. 1964. Gattung Blossfeldia. In Krainz form. Am J Bot 92: 1177–1188.
H, editor. Die Kakteen CVIc, 1. 11. 1964. Eggli U. 1985. Backeberg’s invalid cactus names.
Stuttgart. Bradleya 3: 97–102.
Buxbaum F. 1969. Die Entwicklungswege der Eggli U, Leuenberger BE. 2005. The Cárdenas
Kakteen in Südamerika. pp 583–623. In Fitt- type specimens of Cactaceae names in the her-
kau EJ, Illies J, Klinge H, Schwabe GH, H barium of the Instituto Miguel Lillo, Tucumán,
Sioli, editors. Biogeography and Ecology in South Argentina (LIL). Willdenowia 35: 179–192.
America 2. Monographiae Biologicae 19. Junk, Eggli U, Muñoz Schick M, Leuenberger BE.
The Hague. 1996. (“1995”). Cactaceae of South America:
Cárdenas M. 1950. Cactáceas nuevas de Bolivia. The Ritter Collections. Englera 16: 1–646.
Lilloa 23: 15–27. Eggli U, Taylor NP, editors. 1991. IOS Index
Cárdenas M. 1964. Blossfeldia liliputana Werd. of Names of Cactaceae published 1950–1990
Cactus (Paris) 82: 53. from Repertorium Plantarum Succulenta-
De Candolle AP. 1828. Cacteae. pp 457–476. In rum. Kew.
deCandolle AP. Prodromus systematis naturalis Endler J, Buxbaum F. 1974. Die Pflanzenfami-
regni vegetabilis 3. lie der Kakteen. Philler, Minden.
De Candolle, AP. 1829. (“1828”): Revue de la Fearn PJ. 1996. A review of the origins of the cac-
Famille des Cactées. Mem Mus Hist Nat Paris tus family and the search for a system of classifi-
17: 1–119. cation. Privately published. Wimborne.
Fechser H. 1960. Blossfeldia liliputana. The ti- Hunt DR. 1984. The Cactaceae of Plumier’s Bo-
niest cactus. Cact Succ J (US) 32: 123–125. tanicum Americanum. Bradleya 2: 39–64.
Förster CF. 1846. Handbuch der Cacteenkunde. Hunt DR. 1991. Stabilization of names in suc-
Wöller, Leipzig. culent plants. pp 151–155 in Hawksworth DL,
Fries RE. 1905. Zur Kenntnis der alpinen Flora editor. Improving the stability of names: needs
im nördlichen Argentinien. Nova acta Reg Soc and options. Regnum Veg 123.
Scient Upsaliensis (Ser 4) 1: 1–205. Hunt DR. 1992. CITES Cactaceae checklist. Royal
Funk VA, Mori SA. 1989. A Bibliography of Botanic Garden, Kew.
Plant Collectors in Bolivia. Smithson Con- Hunt DR. 1999. CITES Cactaceae checklist. Ed.
trib Bot 70. Smithsonian Institute, Wash- 2. Royal Botanic Gardens, Kew.
ington DC. Hunt DR, Taylor NP, editors. 1986. The gen-
García M, Lapp M, Castro M, Torrecilla era of Cactaceae: towards a new consensus.
P. 2000. Anatomia foliar comparada de cu- Bradleya 4: 65–78.
atro especies del genero Pereskia (Plum.) Miller Hunt DR, Taylor NP, editors. 1990. The gen-
(Cactaceae). Ernstia 10(1): 27–41. era of Cactaceae: progress towards consensus.
García M, Mejía M, Rodríguez S. 1999. La Report of the IOS Working Party, 1987–90.
Rosa de Bayahibe, salvamento de una espe- Bradleya 8: 85–107.
cie. Boletín Jard Bot Nacional Dr Rafael Ma. Hunt D, Taylor N, Charles G, editors. 2006.
Moscoso 8(6): 12–13. 1999. The New Cactus Lexicon, Vol I and II: Descrip-
Gay C. 1847. Historia Física y Política de Chile. tions and Illustrations of the Cactus Family.
Botánica 3. Paris. DH Books, Milborne Port.
Gibson AC. 1976. Vascular organization in shoots Ideker J. 1996. Pereskia aculeata (Cactaceae), in
of Cactaceae. I. Development and morphology the Lower Rio Grande Valley of Texas. Sida
of primary vasculature in Pereskioideae and 17(2): 527–528.
Opuntioideae. Amer J Bot 63: 414–426. Janchen E. 1943. Zur Nomenklatur der Gat-
Gibson AC. 1977. Vegetative anatomy of Mai- tungsnamen II. Repert Sp Nov Regni Veg 52(2):
huenia (Cactaceae) with some theoretical dis- 144–161.
cussions of ontogenetic changes in xylem cell Johnson H. 1952. Notes on a new group of
types. Bull Torrey Bot Club 104: 35–48. Pereskias. Cact Succ J (US) 24: 114–115.
Gibson AC, Nobel PS. 1986. The Cactus Primer. Johnson H. 1954. The tiniest cactus. Cact Succ
Harvard University Press, Cambridge. J (US) 26: 182.
Greuter W, Barrie FR, Burdet HM, Chaloner Jonkers B. 1998. Maihuenia poeppigii—the south-
WG, Demoulin V, Hawksworth DL, Nicol- ernmost cactus? Brit Cact Succ J 16: 146–153.
son DH, Silva PC, Trehane P, Mcneill J, Jung W. 1982. Beobachtungen an den Standorten
editors. 1994. International Code of Botanical von Maihuenia poeppigii und Maihuenia philip-
Nomenclature (Tokyo Code), adopted by the pii. Kakteen Sukk (Dresden) 17: 74–78.
Fifteenth International Botanical Congress, Jussieu AL. 1789. Genera Plantarum secundum
Yokohama, August September 1993. Koeltz, Ordines Naturales disposita. Hérissant et Bar-
Koenigstein/Taunus Regnum Veg 131. rois, Paris.
Grisebach A. 1879. Symbolae ad Floram Ar- Kiesling R. 1975. Los géneros de Cactaceae de
gentinam. Dietrich’sche Verlags-Buchhand- Argentina. Bol Soc Argent Bot 16: 197–227.
lung, Göttingen. Kiesling R. 1988. Cactaceae. pp 218–243. In
Gürke M. 1908. Neue Kakteen-Arten aus Brasil- Correa MN, editor. Flora Patagónica 5, Dico-
ien. Monatsschr Kakteenk 18: 84–89. tiledoneas dialipétalas (Oxalidaceae a Corna-
Haworth AH. 1812. Synopsis plantarum suc- ceae). Col. Ci. INTA 8. Buenos Aires.
culentarum. Richard Taylor, London. Kiesling R. 1990. Cactus de la Patagonia. Edición
Henderson L. 1995. Plant invaders of southern del autor. Buenos Aires.
Africa. Plant Protection Research Institute Kiesling R. 1999. Les Cactées de Bolivie. Succu-
Handbook No. 5, Agriculture Research Coun- lentes (France) 22.
cil, ARC/LNR, Pretoria, South Africa. Kiesling R, editor. 2003. Flora de San Juan,
Henslow JS. 1837. Opuntia darwinii. Mag Zool Republica Argentina, Vol II. Estudio Sigma,
Bot 1: 466–467. Buenos Aires (and Missouri Botanical Garden
Hofacker A. 1995. Pereskia stenantha Ritter. Press , St. Louis).
Kakeen Sukk. 46(3): center page pullout no. Knize K. 1987. KK-Index 1967–1987. Lima
08(2p.). (Peru).
Hoffmann A. 1989. Cactáceas en la flora silves- Krainz H. 1941. Blossfeldia liliputana Werd. Sch-
tre de Chile. Fundación Claudio Gay. Santi- weizer Garten 11(4): 113–115.
ago de Chile. Krainz H. 1946. Neue und seltene Sukkulenten.
Humboldt ADE, Bonpland A, Kunth C. 1823. Mitteilungen aus der Städtischen Sukkulenten
Nova genera et species plantarum 6. Lutetiae sammlung Zürich. (Sonderdruck aus Sch-
Parisiorum. weizer Garten).
Hunt DR. 1967. Cactaceae. pp 427– 467, In Krainz H. 1949a. Neue Beobachtungen an Bloss-
Hutchinson J, editor. The Genera of Flowering feldia liliputana Werd. Sukkulentenkunde 3:
Plants. Vol 1. Clarendon Press, Oxford. 32–35.
Kr ainz H. 1949b. Blossfeldia liliputana Werd. 64(5): 247–263.

Cact Succ J (US) 21: 103–104. Leuenberger BE. 1997. Maihuenia—monograph
Krainz H. 1960. Fam. Cactaceae, Kakteen. pp 98– of a Patagonian genus of Cactaceae. Bot Jahrb
156 In Encke F, editor. Pareys Blumengärtnerei, Syst 119: 1–92.
Vol 2. Parey Verlag, Berlin and Hamburg. Leuenberger BE. 2002. Humboldt and Bon-
Kr ainz H. 1967. Städtische Sukkulentensam- pland’s Cactaceae in the herbaria at Paris and
mlung Zürich, Katalog der in Kultur stehen- Berlin. Willdenowia 32(1): 137–153.
den Arten. Zürich. Stuttgart. Leuenberger BE. 2004. The Cactaceae of the
Krainz H. 1975. Blossfeldia liliputana. In Krainz Willdenow herbarium, and of Willdenow
H, editor. Die Kakteen CVIe, 1. 10. 1975. (1813). De Herbario Berolinensi Notulae No.
Kr aus R. 1993. Ökophysiologie Chilenischer Kak- 45. Willdenowia 34: 309–322.
teen. Technischen Universität München (Doc- Leuenberger BE, Arroyo-Leuenberger SC.
toral Thesis). 1992. Nota sobre el espécimen tipo de Mai-
Kraus R. 1994. Die Standorte von Maihuenia po- huenia patagonica (Philippi) Britton and Rose
eppigii (Otto ex F. A. C. Weber) K. Schumann (Cactaceae). Parodiana 7(1–2): 25–30.
in Südchile-Eine ökologische Charakterisier- Leuenberger BE, Eggli U. 1999. (“1998”). Notes
ung. Kakteen Sukk 45: 218–221. on the genus Blossfeldia (Cactaceae) in Argen-
Kr aus R. 1995. The environmental conditions of tina. Haseltonia 6: 2–13.
Cactaceae in Chile. Haseltonia 3: 110–129. Lia V V, Confalonieri VA, Comas CI, Hun-
Kümmel F, Klügling K. 2005. Maihuenia po- ziker JH. 2001. Molecular phylogeny of Lar-
eppigii und M. patagonica—zwei südameri- rea and its allies (Zygophyllaceae): reticulate
kanische Kakteenarten für die Freilandkul- evolution and the probable time of creosote
tur. Kakteen Sukk 56: 85–92. bush arrival to North America. Molecular Phy-
Kunth CS. 1823. Opuntiaceae. pp 64–70. In logenetics and Evolution 21: 309–320.
Humboldt A de, Bonpland A, Kunth CS. Linnaeus C. 1738. Hortus Cliffortianus. Privately
Nova genera et species plantarum 6. Lutetiae published, Amsterdam.
Parisiorum. Linnaeus C. 1753. Species plantarum. Holmiae.
Lanjouw J and others, editors 1952. Inter- Liogier A. 1980. Novitates Antillanae, VIII. Phy-
national code of botanical nomenclature ad- tologia 47: 167–198.
opted by the Seventh International Botani- Marshall WT, Bock TM. 1941. Cactaceae.
cal Congress, Stockholm, July 1950. Regnum Abbey Garden Press, Pasadena.
Veg 3: 1–228. Mauseth JD. 1999. Anatomical adaptations to
Lemaire C. 1838. Cactearum aliquot novarum xeric conditions in Maihuenia (Cactaceae), a
ac insuetarum in horto Monvilliano cultarum relictual, leaf-bearing cactus. The Journal of
accurata descriptio. Lutetia Parisiorum. Plant Research 112: 307–315.
Leuenberger BE. 1986. Pereskia (Cactaceae). Mauseth JD. 2001. Pereskias survive frosty win-
Mem New York Bot Gard 41: 1–141. ters in Austin. Cact Succ J (US) 73: 122–126.
Leuenberger BE. 1987. Pereskia oder Peireskia- Mauseth JD. 2006. Blossfeldia lacks cortical bun-
eine immer noch umstrittene Frage? Kakteen dles and persistent epidermis; is it basal within
Sukk 38(4): 96–100. Cactoideae? Bradleya 24: 73–82.
Leuenberger BE. 1988. Zur Geschichte der Tax- Mauseth JD, Landrum JV. 1997. Relictual veg-
onomie von Pereskia lychnidiflora DC. P. zin- etative anatomical characters in Cactaceae:
niiflora DC. (Cactaceae) und ihrer Synonyme. the genus Pereskia. Journal of Plant Research
Beitr Biol Pflanzen 63: 179–198. 110: 55–64.
Leuenberger BE. 1989a. Pereskia stenantha Rit- Mcneill J, Barrie FR, Burdet HM, Demoulin
ter-eine bemerkenswerte Art aus Bahia, Brasil- V, Hawksworth DL, Marhold K, Nicol-
ien. Kakteen Sukk 40(2): 29–31. son DH, Pr ado J, Silva PC, Skog JE, Wi-
Leuenberger BE. 1989b. Pereskia aureiflora er- ersema JH. 2006. International Code of Botan-
stmals farbig abgebildet. Mit einem Nachtrag ical Nomenclature 2005 (Vienna Code). ARG
zur Typisierung des Namens. Kakteen Sukk Gantner, Ruggell (Regnum Veg. 146).
40(11): 280–282. McVaugh R. 1980. Botanical Results of the Sessé
Leuenberger BE. 1992a. Observations on Mai- and Mociño expedition 1787–1803. II. The
huenia (Cactaceae) in Argentina and Chile. Icones Florae mexicanae. Contr Univ Michi-
Part 1. Introduction and notes on root sys- gan Herb 14: 99–140.
tems. Cact Succ J (US) 64(2): 71–79. McVaugh R. 1982. The lost paintings of the Sessé
Leuenberger BE. 1992b. Observations on Mai- and Mociño expedition: a newly available re-
huenia (Cactaceae) in Argentina and Chile. Part source. Taxon 31: 691–692, 1982.
2. Notes on flowers, temperature, and climate. Mejía M, García R, Rodríguez S, Salazar
Cact Succ J (US) 64(3): 125–130. J. Figure 2001. Pereskia quisqueyana Alain
Leuenberger BE. 1992c. Observations on the (Cactaceae) historia y conservación. Mosco-
growth of seedlings of Pereskia (Cactaceae). soa 12: 45–53.
Cact Succ J (US) 64(4): 237–241. Méndez E. 1983. Observaciones sobre Blossfel-
Leuenberger BE. 1992d. Leaf-bearing cacti dia liliputana Werd., nueva cactácea para Men-
(Pereskia) in cultivation. Cact Succ J (US) doza. Deserta 7: 145–155.
Middleditch H. 1988. A trip to the northern Chile. Viaje a las rejiones septentrionales de
regions of Patagonia 1862–1863 by G Cox, la Patagonia, tercera parte. Anales Univ Chile
translated and condensed by H Middleditch. 23: 437–509.
Chileans 14: 6–8. Philippi R A. 1864. Plantarum novarum Chilen-
Miller P. 1754. The gardeners dictionary, abr. ed sium Centuriae, inclusis quibusdam Mendoci-
4. Rivington, London. nis et Patagonicis. Linnaea 33: 1–308.
Miller P. 1768. The gardeners dictionary, ed 8. Philippi R A. 1883. Opuntia poeppigii Otto und
Rivington, Millar and Payne, London. O. segethi Philippi. Gartenflora 32: 259–261
Mitich LM. 1979. The great Nathaniel Lord Brit- pl. 1129.
ton. Cact Succ J (US) 51: 118–121, 172–175. Pichi-Sermolli, REG, Bizzarri MP. 1978. The
Mittler L. 1844. Taschenbuch für den Cactusli- botanical collections (Pteridophyta and Sper-
ebhaber. Schreck, Leipzig. matophyta) of the AMF Mares-G. R. S. T. S. Ex-
Mor an VC, Zimmermann HG. 1991. Biologi- pedition to Patagonia, Tierra del Fuego and
cal control of cactus weeds of minor impor- Antarctica. Webbia 32(2): 455–534.
tance in South Africa. Agriculture, Ecosystems Plukenet L. 1692. Phytographia sive stirpium…
and Environment 37: 37–55. icones. Londini.
Mottram R. 2002. Charles Plumier, the King’s Plukenet L. 1696. Almagestum Botanicum.
botanist: his life and work. With a facsimile Londini.
of the original cactus plates and text from Bo- Plumier C. 1703. Nova plantarum Americana-
tanicon Americanum (1689–1697). Bradleya rum genera. Parisiis.
20: 79–120. Poeppig E. 1835. Reise in Chile, Peru und auf
Mottram R. 2007. The Blossfeldia story. Cactus dem Amazonenstrome während der Jahre 1827–
World 25(3): 162. 1832, 1. Friedrich Fleischer, Leipzig.
Muñoz Pizarro C. 1959. Sinopsis de la Flora R afinesque CS. 1830. Extrait d’une lettre de
Chilena. Santiago de Chile. M. C. S. Rafinesque, professeur… Philadelphie,
Muñoz Pizarro C. 1960. Las especies de plantas adressée… M. le professeur De Candolle (Mai
descritas por R. A. Philippi en el siglo XIX. Univ. 1830). Bull Bot, Geneva 1: 214–216.
de Chile. Santiago de Chile. R auh. 1958. Beitrag zur Kenntnis der perua-
Muñoz Pizarro C. 1966. Sinopsis de la Flora nischen Kakteenvegetation. Sitzungsber Hei-
Chilena. Editorial Universidad de Chile, San- delberger Akad Wiss Math naturw Kl 1. (542pp).
tiago de Chile. Springer. Heidelberg.
Nyffeler R. 2002. Phylogenetic relationships Regel E. 1881. Neue und empfehlenswerte Zi-
in the cactus family (Cactaceae) based on evi- erpflanzen. C. Empfohlen von E. Regel und
dence from trnK/matK and trnL-trnF sequences. E. Schmidt. 4) Peireskia poeppigii Salm. Gar-
Amer J Bot 89: 312–326. tenflora 30: 412.
Oldfield S, editor. 1997. Cactus and Succulent Reveal JL. 2003. Indices Nominum Supragener-
Plants. Status Survey and Conservation Action icorum Plantarum Vascularium. Published on
Plan. IUCN/SSC Cactus and Succulent Spe- the internet; accessed 24.9.2007.
cialist Group. IUCN. Gland, Cambridge. Ritter F. 1965. Nieuwe cactussen uit Zuid
Ostolaza Nano C. 1995. Cacti in the vicinity Amerika. Succulenta 44(2): 23–26.
of Lima, Peru, and their conservation status. Ritter F. 1979. Kakteen in Südamerika 1. Brasil-
Cact Succ J (US) 67(1): 6–13 ien / Uruguay / Paraguay. Spangenberg.
Ostolaza Nano C. 1996. A closer look at the con- Ritter F. 1980. Kakteen in Südamerika 2. Argen-
servation status of cacti in the vicinity of Lima, tinien / Bolivien. Spangenberg.
Peru. Brit Cact Succ J 14(4): 158–174. Roig FA. 1963. Bosquejo fitogeográfico de las pro-
Ostolaza Nano C. 1998. Nomenclatural ad- vincias de Cuyo. IDIA 188: 74–86.
justments in Peruvian Cactaceae. Cact Cons Rojas Acosta N. 1897. Historia natural de Cor-
Init 6: 9. rientes: catálogo. Buenos Aires.
Ostolaza C. 2005. Humboldt y los cactus pe- Rowley GD. 1994. The Sessé and Mociño cac-
ruanos. Quepo 19: 17–25. tus plates. Bradleya 12: 8–31.
Otto F. Figure 1833. Über die Cactus-Pflanzen Rümpler T. 1886. Carl Friedrich Förster’s Handbuch
des hiesigen botanischen Gartens. Allg Gar- der Cacteenkunde. Wöller, Leipzig.
tenz 1 (46): 361–368. Ruiz Leal A. 1956. Observaciones sobre la bi-
Pedroni F, Sanchez M. Figure 1997. Dispersão ología de tres Cactáceas argentinas. Rev Fac
de sementes de Pereskia aculeata Miller (Cac- Ci Agrar Univ Nac Cuyo 6(1): 29–62.
taceae) num fragmento florestal no sudeste do Sajeva M, Mauseth JD. 1991. Leaflike structure
Brasil. Rev Brasil Biol 57(3): 479–486. in the photosynthetic, succulent stems of cacti.
Pfeiffer L. 1837a. Enumeratio diagnostica Cactearum Ann Bot 68: 405–411.
hucusque cognitarum. Oehmigke, Berolini. Salm-Dyck J. 1850. Cacteae in horto dyckensi cul-
Pfeiffer L. 1837b. Beschreibung und Synonymik tae anno 1849. Henry and Cohen, Bonnae.
der in deutschen Gärten lebend vorkommenden Scholz V, Stützel H. 1999. Chronik der Deutschen
Cacteen. Oehmigke, Berlin. Kakteen-Gesellschaft. Würzburg.
Philippi R A. 1863. Catálogo de las plantas re- Schumann K. 1890. Cactaceae. In Martius
cojidas. pp 448–474. In Cox GE, Jeografía de CFP, editor. Flora brasiliensis 4(2): 185–334.
F Fleischer, Monachii. for taxonomic realignments. IOS Bull 6: 13.

Schumann K. 1897. 1898. Gesamtbeschreibung Wallace RS. 1995b. Molecular systematic study
der Kakteen. Neumann, Neudamm. of the Cactaceae: Using chloroplast DNA vari-
Schumann K. 1903. Gesamtbeschreibung der ations to elucidate cactus phylogeny. Bradleya
Kakteen. Nachträge 1898 bis 1902. Neumann, 13: 1–12.
Neudamm. Weber F. 1898a. Les Pereskia et Opuntia péreski-
Schwarz O. 1937. Gemeinschaftsgeist und Au- oides du Mexique. Bull Mus Hist Nat (Paris)
torenrechte in der Botanischen Nomenklatur. 4: 162–167.
Repert Sp Nov Regni Veg 42: 191–198. Weber FAC. 1898b. Pereskia. pp 937–939. In
Spegazzini C. 1902. Nova addenda ad Floram Bois D, editor. Dictionnaire d’Horticulture.
Patagonicam (Cactaceae). Anales Mus Nac Bue- Klincksieck, Paris, 1893–1899.
nos Aires 7: 285–292. Wells PV, Hunziker JH. 1976. Origin of the
Stafleu FA. 1964. An introduction to Jussieu’s Creosote Bush (Larrea) Deserts of southwest-
Genera plantarum. Cramer, Weinheim. ern North America. Ann Missouri Bot Gard
Stafleu FA. 1969. Miller’s 1754 Gardeners Dic- 63: 843–861.
tionary. Taxon 18: 713–715. Werdermann E. 1933. Brasilien und seine Säu-
Steeg, MG van der. 1955. Blossfeldia lilipu- lenkakteen. Neumann, Neudamm.
tana Werd. De kleinste cactus? Succulenta Werdermann E. 1937a. Beiträge zur Nomenkla-
1955. 19–21. tur. Kakteenk 1937: 17–24.
Steudel ET. 1841. Nomenclator botanicus, Vol 2. Werdermann E. 1937b. Aus den Sammelergeb-
Stuttgartiae, Tubingae. nissen der Reisen von H. Blossfeld und O. Mar-
Taylor NP. 1987. New and unfamiliar names of soner durch Südamerika III. Kakteenk 1937:
Cactaceae to be used in the European Garden 161–165.
Flora. Bradleya 5: 91–94. Wijnands DO. 1983. The botany of the Comme-
Taylor NP. 1989. Parodia. pp 257–265. In Wal- lins. Balkema, Rotterdam.
ters SM and others, editors. The European Gar- Winter H. undated 1955. Kakteen, Cacti,
den Flora, 3. Cambridge. Cactées. Samen-seed-semence. Frankfurt a.
Taylor NP, Zappi D. 1997. Nomenclatural ad- M. Fechenheim.
justments and novelties in Brazilian Cactaceae. Winter H. undated 1956. Kakteen, Cacti,
Cact Cons Init 3: 7–8. Cactées. Samen-seed-semence. Frankfurt a.
Taylor NP, Zappi D. 2004. Cacti of eastern Brazil. M. Fechenheim.
Royal Botanic Gardens Kew, Richmond. Winter H. undated 1957. Kakteen, Cacti,
Thorne RF, Reveal J. 2007. An Updated Clas- Cactées. Samen-seed-semence. Frankfurt a.
sification of the Class Magnoliopsida (“Angio- M. Fechenheim.
spermae”). Bot Review 73: 67–182. Winter H. 1959. Kakteen, Cacti, Cactées. Samen-
Vaupel F. 1925. Die Kakteen. Monographie der seed-semence. Frankfurt a. M. Fechenheim.
Cactaceae. Lieferung 1. Berlin. Zuccarini JG. 1837. Plantarum novarum vel
Vaupel F. 1926. Die Kakteen. Monographie der minus cognitarum, quae in horto botanico
Cactaceae. Lieferung 2. Berlin. herbarioque regio monacensi servantur, fasc.
Wallace RS. 1995a. A family-wide phylogeny, sub- 3, Cactaceae. Abh Math Phys Cl Königl Bayer
familial and tribal relationships, and suggestions Akad Wiss 2: 597–742.

Leuenberger On Pereskia

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Haseltonia 14: 54–93.

Pereskia, Maihuenia, and Blossfeldia—

taxonomic history, updates, and notes

Introduction cal distribution, of leaf bearing, only slightly

early cactus evolution

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

Accepted Total of published

early cactus evolution

Name Image1 Author Year Genera Species

early cactus evolution

early cactus evolution

1. P. aculeata Miller 8′. P. marcanoi Areces (1992)

2. Updated alphabetical list of currently accepted names.

early cactus evolution

Name, synonym WWW WWW WWW WWW WWW WWW

early cactus evolution

early cactus evolution

early cactus evolution

Sanchez 1997), invasive potential of P. acu- Pereskia in the WWW (2004–2007)

early cactus evolution

early cactus evolution

The presence of root tubers could be con- Taxonomic history

early cactus evolution

Table 6. Taxonomic history of Maihuenia.

early cactus evolution

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

Maihuenia poeppigii Maihuenia patagonica

authorship. When Schumann began to pub- (Pereskioideae). Schumann (1898) accepted

early cactus evolution

early cactus evolution

­Figures 99–101. Blossfeldia liliputana. Figure 99.

early cactus evolution

tion but is simultaneously considered as not

­ igures 102–104. Blossfeldia liliputana. Figure 102.

early cactus evolution

simultaneously that Blossfeldia might be a de-

rived member of the Maihuenioideae, an op-

Table 9. Taxonomic history of Blossfeldia.

early cactus evolution

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

Synonyms of Blossfeldia liliputana. a selection of names on 28 Sept 2007 (Bloss-

feldia campaniflora, name entries 137 / images

early cactus evolution

Kr ainz H. 1949b. Blossfeldia liliputana Werd. 64(5): 247–263.

early cactus evolution

F Fleischer, Monachii. for taxonomic realignments. IOS Bull 6: 13.

early cactus evolution

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Figures 99–101. Blossfeldia liliputana. Figure 99.

igures 102–104. Blossfeldia liliputana. Figure 102.