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Austrocylindropuntia lagopus - clarification of nomenclature and observations in habitat

Authors:
  • International Organisation for Succulent Plant Study
Austrocylindropuntia lagopus - clarification of
nomenclature and observations in habitat
by Ivor Crook, Martin Lowry and John Arnold
38, Clarendon Road, Audenshaw, Manchester M34 5QB
Summary
A review of the relevant literature is
presented and the nomenclature of this
opuntioid species from the high Andes of
southern Peru is clarified. The distribution of
the species is extended and observations made
on its interaction with its habitat.
Introduction
Few people have ever travelled to the area
around Macusani with the specific intention of
observing its cactus flora. The town lies in a
remote part of the Peruvian Andes north of
Lake Titicaca at an altitude of well over 4000 m.
The surrounding mountains reach to over
6000 m and are snow-capped all year long.
The area has a very low population density
since the climate is too cold for most human
food crops. Human activity is mostly limited to
the area in and around the town with its
uranium mine and markets dealing in Alpaca
wool, known worldwide for its excellent thermal
insulating properties. Isolation of the
surrounding land is compounded by a lack of
good quality roads. The three of us visited the
region in late October 2002 specifically to see
the species currently known as Tephrocactus
malyanus.
Nomenclature
Although it is immediately obvious that the
plant in question is an Opuntia and is closely
related to Austrocylindropuntia floccosa there is
still some confusion regarding its correct name.
In 1903 Schumann described Opuntia lagopus
from specimens sent to him by August
Weberbauer, a German botanist on sabbatical
in Peru for 4 years between 1901 and 1905.
These plants must have endured several
months’ travel by land and sea before they
arrived in the hands of Schumann to be
described. It is not surprising that after such a
journey these desiccated, and perhaps
etiolated, specimens were difficult to tell apart
from the other opuntioid cactus known from
Peru, Austrocylindropuntia floccosa.
Unfortunately the type locality is quite vague
and quoted only as ‘in the neighbourhood of
Arequipa in the Andes at 4000m altitude’. The
only opuntioid species currently found growing
in this region, forms of Cumulopuntia boliviana
and C. sphaerica, bear very little resemblance
to Schumann’s description. What then is
Opuntia lagopus? We cannot examine the type
specimen since, along with many others
preserved at the Herbarium in Berlin; it was
destroyed during World War II. The clues come
from consideration of Weberbauer’s own
publication, ‘Die Pflanzenwelt der peruanischen
Anden’, written after his return to Peru and
employment as Director of the Parque
Zoológico y Jardín Botánico, Lima and
published in 1911. In this article Weberbauer
clearly illustrates as plate 14 (fig. 1) the plant he
considers to be O. lagopus, growing alongside
O. floccosa near a place he calls Poto in the
region northeast of Lake Titicaca. Two
decades later Weberbauer acted as guide and
local advisor for two expeditions, in 1935 and
1938, of the plant hunting team of the American
botanist T. Harper Goodspeed. The main
purpose of the expeditions was to search for
plants of the genus Nicotiana in the eastern
Andes but the expedition route shows that his
collectors travelled north of Lake Titicaca,
possibly to Poto. When Goodspeed
subsequently published his book in 1941
relating his trips to South America he illustrated,
opposite page 81 (fig. 2), a plant of Opuntia
lagopus. This identification was very likely
made by Weberbauer, the man who provided
the material for Schumann’s description. In
both these early illustrations the plant pictured
is unmistakably that which we currently call
Tephrocactus malyanus and which still can be
found in great numbers northeast of Lake
Titicaca around the town of Macusani.
Unfortunately many subsequent authors
have associated the name lagopus with O.
floccosa. Even before T. Harper Goodspeed’s
published his book Backeberg had transferred
the species to the newly erected genus
Tephrocactus. He followed this in 1957 by
publishing 5 further names as varieties of O.
lagopus, all of which are only forms of O.
floccosa. In fact, it is clear from his writings and
illustrations that Backeberg never saw the true
O. lagopus of Schumann.
The true species surfaced again in 1961
when Marnier-Lapostolle described, albeit
invalidly (ICBN Art. 37.1), a plant growing in his
collection as Tephrocactus floccosus var.
cardenasii. The material was sent to Europe by
Martin Cardenas and was said to emanate from
Achacachi in Department La Paz, Bolivia at
3800 m. The photograph shows the same
species illustrated by both Weberbauer and by
Goodspeed. Achacachi lies at the southeast
corner of Lake Titicaca at a much lower altitude
and further south than T. malyanus is currently
found. Thus there is now some doubt that
Cardenas’ specimen was actually found at this
place.
In 1971 Walter Rausch brought to light the
huge populations of this species around
Macusani by describing Tephrocactus
malyanus. This is the first use of the epithet
malyanus for these plants. Presumably,
Rausch was accepting the then current view
that O. lagopus was a species similar to O.
floccosa and hence published his find as a
species new to science. It was subsequently
transferred to Opuntia in 1979 by Werner Rauh.
In 1981 Friedrich Ritter recognised the true
affinity of the species and transferred the two
names O. lagopus and T. malyanus to
Backeberg’s 1938 genus Austrocylindropuntia.
Unfortunately in transferring O. lagopus, the
older and hence correct name, he failed to
quote the page number of the basionym
rendering his transfer invalid (ICBN Art. 33.2).
The most recent author to consider the
classification of this species was Roberto
Table 1. Synonyms of Austrocylindropuntia lagopus (Schumann) Arnold, Crook & Lowry
1903 Opuntia lagopus Schumann
1936 Tephrocactus lagopus (Schumann) Backeberg
1958 Tephrocactus floccosus var. lagopus (Schumann) Ritter nom. inval. (Art 33.2)
1961 Tephrocactus floccosus var. cardenasii Marnier-Lapostolle nom. inval. (Art 37.1)
1971 Tephrocactus malyanus Rausch
1973 Opuntia floccosa var. cardenasii Rowley
1979 Opuntia malyana (Rausch) Rauh
1981 Austrocylindropuntia lagopus (Schumann) Ritter nom. inval. (Art 33.2)
1981 Austrocylindropuntia malyana (Rausch) Ritter
1998 Maihueniopsis lagopus (Schumann) R. Kiesling
Figure 1. Weberbauer’s illustration from
’Die Pflanzenwelt der peruanischen Anden’’
Figure 2. Goodspeed’s illustration from
’Plant Hunters in the Andes’’
Kiesling who in 1998 included O. lagopus within
his expansion of Maihueniopsis.
Although, today, the species is only found
around Macusani and the mountains within
120km of the town, it is clear that Marnier-
Lapostolle's Tephrocactus floccosa var.
cardenasii and Rausch's Tephrocactus
malyanus are the same as Schumann's original
description of plants he thought were from near
Arequipa. Thus, the oldest name for the plant
applies and all these names are relegated to
the synonymy of O. lagopus Schumann. With
the re-division of Opuntia into its component
genera the characters of the species, and in
particular those of its seeds, place it squarely
within Austrocylindropuntia Backeberg,
however, since Ritter’s combination in that
genus is invalid a further combination is
required, thus:
Austrocylindropuntia lagopus
(Schumann) Arnold, Crook & Lowry.
Basionym: Opuntia lagopus Schumann,
Gesamtbeschreibung der Kakteen.
Nachträge 1898 bis 1902: pages 151 – 152,
1903.
A full synonymy is detailed in Table 1.
Further, since the original material no longer
exists and Rausch’s type for T. malyanus
cannot be found, we also designate here a
neotype: Webster 26 (K) collected 10 miles
northeast of Macusani, department Puno, Peru
in 1965. This specimen was first noticed by
James Iliff who records that the collection is
liberal and in excellent condition with much
annotation.
Description
The plants form large, dense mounds of
short, cylindrical, much branched segments.
Each mound can be well over a metre across
and up to 60 cm high (fig. 3). Larger mounds
are of a density and strength sufficient to
support the weight of an adult male standing
upon them! From examination of mounds that
are dead or have been removed from the
ground and upturned it can easily be seen that
each mound is a solitary plant (fig. 4). Stem
segments can be up to 45 cm or more in length
in mature plants. In seedling plants of 5 - 6
heads, the ends of the segments are almost
flush with the ground. The segments are
Figure 3. A large specimen of A. lagopus nearly 4 m in
diameter growing 25 km south of Macusani at 4680 m
Figure 4. Overturned plant of A. lagopus showing the
branching pattern and single root
Figure 5. Nascent buds and flowers
Figure 6. Small butterflies feeding on the nectar of A.
lagopus flowers.
densely hairy along their whole length. The hair
arises from the areoles along with sparse, well-
hidden glochids and frequently one 2 - 2.5 cm
long yellow spine near the tip of the segment.
Iliff has shown that the hairs are unusual in that
they consist of a single linear series of cells
unlike those of A. floccosa, which are
multiseriate and barbed. At the tip of each
segment, occluded by the hairs are several,
short, persistent leaves up to 7 mm long.
Flowers are golden yellow, 2 - 3 cm long with
the pericarpels hairy above. They appear from
areoles 2 – 3 cm below the tip of each segment
and remain flush with the surface of the mound
until anthesis (fig. 5). On several occasions we
observed small butterflies visiting the flowers
(fig. 6). The insects were obviously obtaining
significant quantities of nectar since they spent
a considerable time at each flower. Fruits are
ovoid, pale yellow-green in colour and partially
sunken within the mound until mature. At
maturity they expand longitudinally, turn pale
pink and are fairly easily detached from the
plant. Unlike the fruits of A. floccosa they are
thin walled. They contain 5 - 10 pale brown,
almost spherical seeds approximately 5 mm in
diameter each bearing an equatorial girdle of
funicular origin (fig. 7).
Habitat
The plant is found in two different habitats.
Near Rosario and Crucero, and also to the
north of Macusani, the ground is frequently very
dry and barren. Plants rarely reach a size of
more than 1 m diameter when mature.
However, between Rosario and Macusani, on
either side of the pass over the Nudo Aricoma
mountains, plants grow to within a few
kilometres of the pass. This habitat is higher,
colder and much damper as indicated by the
grasses covering the ground. This is where the
largest specimens can be found. We observed
A. lagopus at altitudes of up to 4686 m,
confirming Alfred Lau’s statement that this is
the highest growing cactus species. Plants in
this region were observed growing alongside
rivulets of running water; indeed some were in
direct contact with running water. Early in the
morning we observed several plants to be
covered in snow near to the pass following
overnight showers.
In the more barren areas at lower altitudes
of its range between 4100 and 4350 m it is
seen growing alongside Echinopsis
maximiliana. At all locations it can be seen
growing alongside A. floccosa. Even at high
altitudes amongst temporary snow the plants
often had several holes, around 2 – 3 cm
diameter near the base and sides of the plant.
Whilst no rodents were seen in the area we did
find one plant to be home to a small lizard that
emerged in the early morning to sun itself on
the upper stems of a plant.
Distribution
As previously stated the area around
Macusani has never been extensively explored
with regard to its cactus flora by either
professional or amateur botanists. Most of the
observations of this plant come from recent
trips to the area made by Klaus Gilmer (TG),
Brian Bates (BB), Hakan Sonnermo (HNSO)
and our own trip on 27th to 29th October 2002
(ACL). Observations have been recorded in the
eastern part of the Cordillera de Carabaya and
the Nudo Aricoma mountains at altitudes of
between 4110 and 4686 m. Also Alfred Lau
reports finding this species close to Ulla Ulla in
Bolivia. Observed sightings of the plant are
shown in Map 1.
Acknowledgements
We are extremely indebted to James Iliff for
bringing to light the specimen Webster 26 at
Kew and for many other helpful discussions.
We also thank Graham Charles, Paul Hoxey,
David Hunt, Beat Leuenberger, Roy Mottram
and Nigel Taylor for their significant
contributions to this paper.
Figure 7. Mature fruits of A. lagopus
References
BACKEBERG, C. & KNUTH, F.M. (1936)
Kaktus – ABC, p106.
BACKEBERG, C. (1957) Descriptiones
Cactacearum novarum, p7.
GOODSPEED, T H. (1941) Plant Hunters in the
Andes JJ Little and Ives, New York, p81.
ILIFF, J. (2002) ‘The Andean Opuntias’ in
Studies in the Oputioidea, Succulent Plant
Research 6 p205.
KIESLING, R. (1998) ‘Revision and transfer of
some Austrocylindropuntia’ in Tephrocactus
Study Group Newsletter 4(1) p239.
LAU, A. (1980) ‘South American Cactus Log
Part XIII’ in Cactus & Succulent Journal (US)
52 p289.
MARNIER-LAPOSTOLLE, J. (1961) ‘Notes du
Jardin Botanique des cedres: une variete
nouvelle de tephrocactus’ Cactus (Paris) 72
p137.
RAUH, W. (1979) Kakteen an ihren Standorten
p93.
RAUSCH, W. (1971) ‘Tephrocactus malyanus
sp. nov.’ Kakteen und anderen Sukkulenten
22 p43 – 44.
RITTER, F. (1981) Kakteen in Sudamerika 4
p1242 - 1244.
ROWLEY, G.D. (1973) Repertorum Plantarum
Succulentarum 22 p11.
SCHUMANN, K. (1903) Gesamtbeschreibung
der Kakteen. Nachträge 1898 bis 1902:
p151 – 152.
WEBERBAUER, A. (1911) ‘Die Pflanzenwelt
der peruanischen Anden’, plate 14, in Engler
& Drude, Die Vegetation der Erde 12.
A large colony of A. lagopus plants east of Crucero
A. floccosa found growing alongside A. lagopus near
Macusani
Article
Full-text available
A short history of the Cactaceae holdings of the Berlin herbarium and their fate is provided. While a considerable number of historical spirit specimens have escaped the destruction of the herbarium during World War II, the pre-1943 collection of pressed specimens was lost almost entirely. More than half of the present Cactaceae holdings of B were made on the base of cultivated material. The importance of these "Garden Herbarium" specimens is stressed, especially for succulent plants, where much of the type material, even of recently described taxa, is of insufficient quality or even fragmentary. A complete survey of all extant Cactaceae specimens at B (including the material recently transferred from the former KOELN herbarium) has resulted in a detailed and commented list of type specimens and specimens derived from cultivated descendants of type collections. Nomenclatural and typification problems encountered during this compilation are discussed. A list of photographs preserved at B depicting type material located in other herbaria, is also given. Lectotypes are designated for the names Cereus lamprospermus, C. rhodoleucanthus, C. stenogonus, C. xanthocarpus, Echinocactus umadeave, Echinopsis rhodotricha, Pilocereus setosus, Rebutia steinbachii and Turbinicarpus pseudomacrochele var. lausseri.
The Andean Opuntias' in Studies in the Oputioidea
  • J Iliff
ILIFF, J. (2002) 'The Andean Opuntias' in Studies in the Oputioidea, Succulent Plant Research 6 p205.
Revision and transfer of some Austrocylindropuntia
  • R Kiesling
KIESLING, R. (1998) 'Revision and transfer of some Austrocylindropuntia' in Tephrocactus Study Group Newsletter 4(1) p239.
South American Cactus Log Part XIII
  • A Lau
LAU, A. (1980) 'South American Cactus Log Part XIII' in Cactus & Succulent Journal (US) 52 p289.
Tephrocactus malyanus sp. nov
  • W Rausch
RAUSCH, W. (1971) 'Tephrocactus malyanus sp. nov.' Kakteen und anderen Sukkulenten 22 p43 -44.
Gesamtbeschreibung der Kakteen. Nachträge 1898 bis 1902
  • K Schumann
SCHUMANN, K. (1903) Gesamtbeschreibung der Kakteen. Nachträge 1898 bis 1902: p151 -152.
Notes du Jardin Botanique des cedres: une variete nouvelle de tephrocactus' Cactus
  • J Marnier-Lapostolle
MARNIER-LAPOSTOLLE, J. (1961) 'Notes du Jardin Botanique des cedres: une variete nouvelle de tephrocactus' Cactus (Paris) 72 p137.
Kakteen an ihren Standorten p93
  • W Rauh
RAUH, W. (1979) Kakteen an ihren Standorten p93.
  • F Ritter
RITTER, F. (1981) Kakteen in Sudamerika 4 p1242 -1244.
Die Pflanzenwelt der peruanischen Anden
  • A Weberbauer
WEBERBAUER, A. (1911) 'Die Pflanzenwelt der peruanischen Anden', plate 14, in Engler & Drude, Die Vegetation der Erde 12.