Content uploaded by Martin Lowry
Author content
All content in this area was uploaded by Martin Lowry on Dec 02, 2017
Content may be subject to copyright.
89
Bradleya 31/2013
Summary: The taxonomic and nomenclatural his-
tory of Lobivia chrysochete (Werdermann) Wess-
ner is reviewed and a lectotype established.
Biogeographic, morphological and genetic infor-
mation is provided to support the assertion that
one member of the group, Lobivia chrysochete var.
minutiflora Rausch, should be considered a dis-
tinct species. Illustrations of both species in habi-
tat and cultivation are presented.
Zusammenfassung: Die taxonomische und nomen-
klatorische Geschichte von Lobivia chrysochete
(Werdermann) Wessner wird zusammengefasst
und ein Lektotypus definiert. Biogeographische,
morphologische und genetische Merkmale werden
aufgeführt um zu belegen, dass ein Vertreter der
Verwandtschaftsgruppe, Lobivia chrysochete var.
minutiflora Rausch als eigene Art anerkannt wer-
den sollte. Abbildungen zeigen beide Arten im
Habitat und in Kultur.
Introduction
In 1936 Werdermann published the name
Echinopsis chrysochete for a plant found by Mar-
soner in 1934 in the high mountains of the eastern
border of Jujuy with Salta, Argentina (Werder-
mann, 1936). In Argentina these mountains are
known as the Sierra de Santa Victoria and extend
Bradleya 31/2013
pages 89-95
Lobivia minutiflora: a cryptic species finally recognised
M. Lowry1and B.O. Schlumpberger2
183, Seaton Road, Hessle, Hull, East Yorkshire, UK email: m.lowry@hull.ac.uk
2Herrenhausen Gardens, Herrenhäuser Strasse 4, 30419 Hannover, Germany.
email: boris.schlumpberger@hannover-stadt.de
Photographs: M. Lowry
Figure 1. Lobivia minutiflora BLMT795, Palacio Tambo, Chuquisaca, Bolivia
90 Bradleya 31/2013
from just north of Jujuy city in the south to the
Bolivian border in the north. The range continues
north into Bolivia as the Serrania de Sama in De-
partment Tarija and the Sierra Mandinga in
Chuquisaca. They are some of the highest of the
easternmost Andean ranges and frequently reach
altitudes of over 4000m.
The name Echinopsis chrysochete was subse-
quently transferred to Lobivia by Wessner in 1938
(Wessner, 1938) and the new combination was
used by Rausch (Rausch, 1975) as the species
name for a range of forms found by him and oth-
ers along the same mountain chain. In the most
recent synoptic treatments of Echinopsis (Ander-
son & Eggli, 2005; Hunt et al., 2006) all these
Figure 2. Lobivia chrysocheteBLMT730, Inca
Huasi, Chuquisaca, Bolivia
Figure 4. Lobivia chrysochete BLMT731, Inca
Huasi, Chuquisaca, Bolivia
Figure 3. Lobivia chrysocheteBLMT731, Inca
Huasi, Chuquisaca, Bolivia
Figure 5.Lobivia chrysocheteWR215, Volcan,
Jujuy, Argentina, the form sometimes labelled
var. markusii.
Figure 7. Lobivia chrysochete BLMT155, Cuesta
de Sama, Tarija, Bolivia, the form published by
Ritter as Lobivia hystrix.
Figure 6. Lobivia chrysochete BLMT155, Cuesta
de Sama, Tarija, Bolivia, showing the extremely
woolly buds 3 weeks before anthesis.
91
Bradleya 31/2013
names were considered synonyms of Echinopsis
chrysochete Werdermann.
Over the course of several visits to South
America the authors have travelled through the
afore mentioned mountains to find these plants.
Of particular interest was the road from La
Quiaca over the Sierra de Santa Victoria to the
eponymous village, Santa Victoria. After passing
the village of Suripujio on the Altiplano the road
climbs first to the Abra Lizoite at just over 4500m,
descends a little and crosses a dry high plateau
before reaching another ridge on the eastern side,
the Abra Patahuasi.
It is here, at around 4300m altitude, that we
found, independently, very large Lobivia plants
that we considered to be Lobivia chrysochete var.
minutiflora Rausch. These plants can be distin-
guished from the type variety primarily on the
basis of flower form and size (unusually small for
such a large plant, hence the epithet), but also by
differences in body size and spination. In the same
general area however, we have both also found
plants with the typical large flowers of var.
chrysochete.
Their proximity raised the obvious question as
to whether these two forms are truly distinct, a
situation which prompted the current investiga-
tion. From several lines of evidence we eventually
concluded that they are in fact distinct species and
hence published, in brief, a new combination, Lo-
bivia minutiflora (Rausch) Schlumpberger &
Lowry, at species level for the small-flowered
taxon (Schlumpberger, 2012). In this article, we
present our data and discuss the ecology of the
two species.
Methods
Fieldwork
Over the course of several expeditions to the
Andes of Bolivia and Argentina we photographed
and recorded the locations and altitudes of each
population of this species alliance which we en-
countered. This information was used to produce
distribution maps using GIS software (DIVA-GIS,
version 7.5, http://www.diva-gis.org/). At several
locations we collected fruits containing ripe seeds
that were subsequently used to provide seedlings
for comparative morphological and genetic analy-
sis.
Figure 8. Lobivia minutiflora BLMT795, Palacio Tambo, Chuquisaca, Bolivia, illustrating the small, al-
most glabrous, tubular red flower.
92 Bradleya 31/2013
Morphological comparison
In addition to the seedlings grown from wild-
collected seed one of us (M.L.) acquired specimens
of known provenance from specialized amateur
collectors. These have been grown under identical
conditions for several years eventually reaching
maturity and producing flowers for comparison.
Genetic comparison
Chloroplast markers of both taxa were se-
quenced as part of a recently published molecular
phylogenetic study (Schlumpberger & Renner,
Figure 11. Lobivia minutiflora PM374, Santa
Victoria, Salta, Argentina.
Map 1. Geographic distribution of Lobivia chryso-
chete (red) and Lobivia minutiflora (yellow). Cir-
cles identify locations of confirmed findings of the
taxa whilst squares mark the approximate posi-
tions of the type localities of the varieties included
within L. chrysochete and L. minutiflora. Several
important population centres mentioned in the
text are shown. The international border between
Argentina and Bolivia is indicated by the dark
line transecting the map at the latitude of La
Quiaca.
Figure 9. Lobivia chrysochete BLMT441,
Nazareno, Salta, Argentina, the form sometimes
labelled var. subtilis, showing the large campan-
ulate red flowers.
Figure 10. Lobivia minutiflora BLMT335, Abra
Patahuasi, Salta, Argentina, showing the short
red buds 2 days before anthesis.
93
Bradleya 31/2013
2012). For the purpose of this taxonomic study,
the sequence characteristics of the trnS-G inter-
genic spacer and the trnL region of L. chrysochete
and L. minutiflora were compared. For details of
primers, DNA extraction, amplification and se-
quencing as well as voucher specimens and Gen-
Bank accession numbers see Schlumpberger and
Renner (2012).
Results and Discussion
Geographic distribution
Populations of plants currently accepted as L.
chrysochete were found over a large geographic
range stretching 420km from northern Argentina
to southern Bolivia (Map 1). Additional confirmed
observations (Sierra Mandinga, Chuquisaca, Bo-
livia, J. Carr, pers. comm.) and locations of type
localities for L. chrysochete var. subtilis (Santa
Ana, Salta, Argentina), L. chrysochete var.
markusii (Volcan, Jujuy, Argentina) and L. minu-
tiflora (Iruya, Salta, Argentina) extend the range
both north and south by a further 60km to about
480km.
In contrast, the west-to-east extent of the dis-
tribution is only 60km. Interestingly, the small-
flowered plants are found in two disjunct areas
separated by about 300km, one in central Bolivia
at ~20°S, the other in northern Argentina at
~22.5°S. All populations were found at altitudes
greater than 2800m. There is, however, a signifi-
cant difference in the altitudinal range occupied
by the large- and small-flowered plants (P<0.001,
Chart 1). The large flowered plants (L. chryso-
chete) occur over the range 2900-4100m (median
3440m), whilst the small-flowered plants (L.
minutiflora) are found at higher altitudes of 3900-
4400m (median 4160m). A preference of L. minu-
tiflora for the cooler climate at higher altitudes,
where average temperatures are about 4°C lower
throughout the year, could explain its disjunct dis-
tribution since the mountains reach these eleva-
tions at relatively few places along the
north/south transect of the distribution.
Morphological characters
Both L. chrysochete and L. minutiflora are ap-
planate plants with umbilicate centres and can
grow to quite sizable specimens of 25-40cm diam-
eter (Figures 1 & 2) They both have 20-40 spi-
ralling ribs broken into hatchet-shaped tubercles
with the areoles placed in the depressions. Each
areole carries 8-30 pale yellow to dark brown
Table 1. Summary of the main distinguishing morphological characteristics of L. chrysochete and
L. minutiflora. Observations were made from multiple specimens both in habitat and in cultivation.
Character L. chrysochete L. minutiflora
Maximum body diameter
(cm) 25 40
Spines (number per areole,
nature, orientation) 6-20, stiff, porrect ~30, flexible, adpressed
Receptacle Very woolly, green/brown Almost glabrous, red
Flowers (shape, size,
colour)
Campanulate, 4cm tall x 5cm
diameter, yellow, orange or red
Tubular, 3cm tall x 2.5cm
diameter, red
Chart 1. Scatter plot demonstrating the differ-
ence in altitude (metres) at which the populations
of the two taxa where discovered. Although there
is some overlap, the two distributions are signifi-
cantly different (P<0.001 by Wilcoxon’s non-para-
metric test). Red dots: L.chrysochete. Yellow dots:
L. minutiflora.
94 Bradleya 31/2013
spines that may be flexible or stiff. Flowers are
produced profusely in a ring at the shoulder of the
plant and are generally orange-red, but yellow is
also known from some populations (Figures 3 &
4). Fruits mature relatively quickly and are large,
up to 2cm x 3cm, and tuberculate with a thick
wall. They contain many small, shiny black oval
seeds embedded in a white funicular pulp.
Thus, at first sight, they can appear similar;
however, careful comparison of these characters
shows consistent differences between the two
species (Table 1). Mature specimens of L. chryso-
chete are generally a little smaller than those of
L. minutiflora and have stiffer, more porrect
spines, whereas the adpressed spine disposition of
L. minutiflora is quite characteristic being remi-
niscent of that in Eriosyce umadeave (Werder-
mann) Katterman.
The biggest differences are seen in the flowers;
those of L. chrysochete are large, flamboyant and
campanulate (Figures 3, 5, 7 & 9), with many dark
hairs on the receptacle, whilst those of L. minuti-
flora are much smaller, almost tubular and carry
only a few white hairs (Figure 12). In contrast to
L. chrysochete, the exterior of the floral tube of L.
minutiflora is bright red (brownish to greenish in
L. chrysochete Figure 10, 11 & 12). These floral
characteristics may be consistent with a differ-
ence in pollinator preference between the two
species, with L. chrysochete flowers being more at-
tractive to insects (entomophilous), specifically
bees, and those of L. minutiflora potentially better
adapted to hummingbirds (ornithophilous). This
dichotomy of pollinators is not unknown in the
genus since L. pentlandii (Hooker) Britton & Rose
and L. maximiliana (Heyder ex A. Dietrich)
Backeberg are entomophilous and ornithophilous,
respectively (Schlumpberger, unpublished data)
and frequently occur sympatrically (Lowry, un-
published data)
Figure 12.Sectioned flowers from Lobivia chryso-
cheteWR215 (left) and Lobivia minutiflora
PM374 (right) demonstrating the marked differ-
ences in flower size and receptacle covering. Scale
divisions are 1cm apart.
Figure 13. Lobivia minutiflora PM374, Santa
Victoria, Salta, Argentina. Flower section from
flower in Figure 11. Scale divisions are 1cm apart.
Figure 14. Lobivia chrysochete WR215, Volcan,
Jujuy, Argentina. Flower section. Scale divisions
are 1cm apart.
Figure 15. Sectioned flowers from Lobivia minu-
tiflora PM374 (left) and Lobivia chrysochete
WR215 (right) demonstrating the marked differ-
ences in flower size and form. Scale divisions are
1cm apart.
95
Bradleya 31/2013
Genetic characters
In the molecular phylogenetic analysis on
Echinopsis s.l. and the Trichocereeae using three
chloroplast markers (Schlumpberger & Renner,
2012), L. chrysochete and L. minutiflora did not
cluster together. This finding underlines the idea
that the two taxa may indeed be different species.
The trnS-G sequences diverge by seven muta-
tions, i.e. one three nucleotide indel and six sin-
gle-nucleotide polymorphisms (SNPs). The trnL
sequences differ by 3 SNPs. While L. chrysochete
shares a number of sequence characteristics with
L. cinnabarina, L. ferox and others, L. minutiflora
has a number of sequence similarities in common
with the L. ancistrophora alliance.
Taxonomy & Nomenclature
Lobivia chrysochete (Werdermann) Wessner.
Beitr. Sukkulentenk. Sukkulentenpflege (1938) III:
71. Basionym: Echinopsis chrysochete Werder-
mann Repert. Spec. Nov. Regni Veg. (1936) 39:
271-272. Type: Marsoner s.n. 1934, Northern Ar-
gentina, province Jujuy, approx 2800m altitude.
Herbarium: B†. Lectotype: (designated here), the
unnumbered sheet at B identified by Eggli &
Leuenberger (2008) and containing two photo-
graphic prints and a negative. Synonyms: Lobivia
hystrix Ritter Succulenta (1966) 45: 84; Lobivia
chrysochete var. hystrix (Ritter) Ullmann Kaktusy
(Brno) (1992) 28: 34; Lobivia markusii Rausch
Kakteen And. Sukk. (1966) 17: 121 (nom. inval.;
no type stated); Lobivia markusii Rausch Kakteen
And. Sukk. (1970) 21: 45; Lobivia chrysochete var.
markusii (Rausch) Rausch Lobivia (1975) 2: 96
(nom. inval.; incorrect basionym citation); Lobivia
chrysochete var. subtilis Rausch Succulenta
(Netherlands) (1980) 59: 54; Lobivia tenuispina
Ritter (1966) Succulenta 45: 85; Lobivia chryso-
chete var. tenuispina (Ritter) Rausch Lobivia
(1975) 2: 96.
The sheet at the Berlin herbarium, here se-
lected as lectotype, was considered by Eggli &
Leuenberger (2008) to be of the type collection and
is identified on an original label as the “gepfropfte
Köpfe und getrocknete Import Typ” in Werder-
mann’s handwriting. Proof that the plant pho-
tographed was part of Marsoner’s original
collection is clear from the protologue, where Wer-
dermann explicitly states that one head from a
many-headed specimen was grafted on “Spachi-
anus” and grown in the Botanic Garden. It is prob-
able that one of the prints was subsequently
reproduced by Backeberg in his magnum opus Die
Cactaceae as Abb. 1351, since the illustration
shows an apparently grafted plant and the legend
gives the source as “Botan. Garten Berlin-
Dahlem”.
Lobivia minutiflora (Rausch) Schlumpberger
& Lowry Cact. Syst. Init. (2012) 28: 30. Basionym:
Lobivia chrysochete var. minutiflora Rausch Kak-
teen And. Sukk. (1977) 28: 74. Type: Rausch 512,
Argentina, Salta, near Iruya at 3500m. Herbar-
ium: ZSS.
Conclusion
The combined information from biogeographic,
morphologic and genetic evidence presented here
considerably reinforces our opinion that L. minu-
tiflora (Rausch) Schlumpberger & Lowry should
be considered a species distinct from L. chryso-
chete Werdermann.
Acknowledgement
We are very grateful to Mats Hjertson of the
Museum of Evolution, Uppsala University for ob-
taining for us a copy of the protologue of Echinop-
sis chrysochete. Boris Schlumpberger was
supported by the Deutsche Forschungsgemein-
schaft (grants SCHL 1820/1-1 & SCHL 1820/1-2)
References
ANDERSON, E.F. & EGGLI, U. (ED.) (2005) Das
grosse Kakteen-Lexikon. Stuttgart (GE):
Ulmer.
EGGLI U. & LEUENBERGER, B.E. (2008) Type spec-
imens of Cactaceae names in the Berlin
Herbarium (B). Willdenowia 38: 213-280.
HUNT, D.R., TAYLOR, N. & CHARLES, G. (2006) The
New Cactus Lexicon. Milborne Port (GB): dh
books. Text and atlas volumes.
RAUSCH, W. (1975) Lobivia, the day flowering
Echinopsidinae from a geographical distribu-
tion point of view. 2: 96.
SCHLUMPBERGER, B.O. (2012) New combinations in
the Echinopsis alliance. Cact. Syst. Init. 28: 29.
SCHLUMPBERGER, B.O. & RENNER, S.S. (2012) Mo-
lecular phylogenetics of Echinopsis (Cac-
taceae): Polyphyly at all levels and convergent
evolution of pollination modes and growth
forms. Am. J. Bot. 99: 1335-1349.
WERDERMANN, E. (1936) Neue Sukkulenten.
Repert. Spec. Nov. Regni Veg. 39: 271-272.
WESSNER, W. (1938) Lobivia chrysochete Werd.
nov. spec. 1936. Beitr. Sukkulentenk. Sukku-
lentenpflege (III): 71.