Eriosyce. The Genus revised, amplified

172 Pages·1994·49.96 MB·English

by  Kattermann F.

Succulent Plant Research
Volume One
Eriosyce
( Cactaceae)
The genus revised and amplified
by
Fred Kattermann
with contributions by
Jonas M. Liithy, Robert S. Wallace
David Hunt, Nigel Taylor and Daniela Zappi
edited by
David Hunt & Nigel Taylor
Royal Botanic Gardens
Kew
1994
Eriosyce (Cactaceae): The genua revined and amplified
AUTHOR'S I want to thank all those who have helped me in the
ACKNOWLEDGMENTS preparation of this book. I especially want to thank Heriberto
and Adela Nichterlein and their family, of Vina del Mar, Chile.
They provided my home away from home. It was always with
enthusiasm that they put up with "el fanatico". Special
mention must also be made of Hans Nichterlein, once an
exchange student in our house, on whom I could always
depend, just like a son. I am also very grateful to Sr Mario
Pino, for many years the president of the Sociedad Cientffica
de Valparaiso, a naturalist without equal: Sra Adriana
Hoffmann, botanist, author, friend and travel companion, for
sharing her knowledge and smoothing the way through
Chilean bureaucracy; to Herr Walter Jung, cactophile, guide
and friend, who made my first field expeditions possible; to Dr
David Hunt and Nigel Taylor (Royal Botanic Gardens, Kew) for
their editorial help and to Dr Danicla Zappi (Kew) for
redrawing the distribution maps; to Prof. Otto Zoellner, Dept
of Botany, Universidad Catolica de Valparaiso, who
accompanied me on many of my trips; to Dr E.F. Anderson,
formerly of Whitman College, Walla Walla, Washington, and
now at the Desert Botanical Garden, Phoenix, Arizona, for his
advice and assistance in preparing most of the seed, plant and
pollen SEM pictures, and especially to Christine Thomson at
Whitman for her technical help; to Dr Steven Koppe from
Montclair State College, New Jersey, for allowing me use of the
SEM facility at that college; and last but not least to Dr Robert
Wallace of Iowa State University for preparing and writing the
cladistic analysis.
Fred Kattermann
719 Route 519, Wantage,
© 1994 Fred Kattermann NJ 04761, U.S.A.
and contributors
Alt rights reserved. No part of
this publication may be
reproduced, in any form or by
any means, without prior
permission in writing from the
Publisher.
ISBN 0 9517234 2 1
Published by
David Hunt
54 Priory Road
Richmond
Surrey TW9 3DH
England
Printed in England
Succulent Plant Research
Volume One
CONTENTS
Summary - Zusammenfassung - Resumen 6
Erioayce (Cactaceae): The genus revised and amplified
Fred Kattermann
Introduction 7
Previous study 7
Historical landmarks 8
Scope of present study 10
General conclusions 10
Outlline of proposed classification 11
Characters used in classification 12
Reproductive system 17
Allied genera 17
Uses and economic importance 20
Threats to the survival of Eriosyce species 20
Cultivation 21
Selected bibliography 23
Systematic treatment 25
APPENDICES
I. Validation of new names 117
II. Eriosyce laui J. Luthy sp. nov 120
Jonas M. Liithy
III. Phylogenetic analysis of Erioayce 125
Robert S. Wallace
IV. Documentation of accepted names and
synonyms 131
David Hunt, Nigel Taylor & Daniela Zappi
Alphabetical list of synonyms 170
Index of accepted taxa 174
Index of illustrations 175
В
Eriosyce (Cactaceae): The genus revised and amplified
Summary. The genus Eriosyce Phil. (Cactaceae-NotocacteacJ is revised and amplified to include 33 species
and various infraspecific taxa previously classified in Neoporteria sens. lat. As circumscribed here Eriosyce
ranges along the western aide of South America, from central Chile to south-western Peru, and on the east
side of the Andes in western Argentina, from Mendoza to Salta. The data upon which this study is based
derive from field observations and herbarium collections made by the principal author (Kattermann).
Bibliographical and nomenclatural details (Appendix IV» and a pbylogeneLic analysis (Appendix III) are
supplied by various collaborating authors, and a new species, E. laui J. Luthy, is described in Appendix II.
A study of seed-morphology utilizing the Scanning Electron Microscope is presented and taxa at all ranks
are keyed out, described and illustrated by means of colour photographs and/or line drawings. The genus is
divided into two sections and several subsections and all new names and combinations required are
published and listed ' Appendix 11: sect. Eriosyce with subsect. Eriosyce \E. aurata. E. rodent iophila). subsect.
Pyrrhocactus <E. bulbocalyx. E. strausiana, E. umadeave. E. andreaeana, E. villicumensis) and subsect. Islaya
fE. islayensis); and sect. Neoporteria with subsect. Horridocactus (E. recondita, E. vurvispina, E. kunzei, E.
limariensis, E. confinis. E. crispa, E. garaventae. E. aspdlagae. E. marksiana. E. engleri, E. heinrichiana, E.
napina, E. odieri, E. taltalensis, E. occidta). subsect. Chileosycc <E. esmeraldana, E. aerocarpa, E. krausii.
E. tenehrica. E. laui) and subsect. Neoporteria IE. chilensis, E. subgibbosa, E. sociabilis, E. senilis, E. villosa).
Zusammenfaasung. Die Gattung Eriosyce Phil. ' Cactaceae-Notocacteae ( wird revidiert und erweitert und
umfasst nun 33 Arten und verschiedene infraspezifische Taxa. die fruher zu Neoporteria sens. lat. geslelll
worden sind. In der hier vorgeschlagenen Umschreibung ist die Gattung Eriosyce auf der westlichen Spite
von Sudamerika zwischen Zentral-Chile und dem siidwestlichen Peru verbreitet, sowie dstlich der Andcn im
westlichen Argentinien von Mendoza bis Salta. Die dieser Studie zugrundeliegenden Duten wurden im Feld
und aulgrund der Herbarbclege des Hauptautnrs < Kattermann) erarbeitet. Hibliographische und nomenkla-
torische Details iAnhang IV) sowie die phylogenetische Analyse lAnhang Ш) wurden von verschiedenen
Mitarbeitern beigesteuert, und die neue Art E. laui J. Luthy wird beschrieben i Anhang III. Zusatzlich wird
eine SEM-Studie der Samenmorphologie vorgestellt. Die Taxa aller Rangslufen werden aulgeschlusselt.
beschrieben und mit Fnrbbildern nder Zeichnungen illustriert. Die Gattung wird in zwei Sektionen und
verschiedene Subsektionen eingeteilt. und zahlreiche neue Namen und Kumbinationen werden publiziert und
aufgelistet (Anhang I): Sektion Eriosyce mit Suhsektion Eriosyce <E. aurata. E. rodentiophilu), Subsektion
Pyrrhocactus <E. bulbocalyx. E. strausiana. E. umadeave, E. andreaeana, E. vitlicumensisl und Subsektion
Islaya (E. islayensis); und die Sektion Neoporteria mit der Subsektion Horridocactus <E. recondita, E.
curvispina. E. kunzei. E. limariensis, E. confinis, E. crispa. E. garaventae. E. aspdlagae, E. marksiana, E.
engleri, E. heinrichiana. E. napina. E. odien. E. taltalensis. E. occulta*, der Subsektion Chileosyce IE.
esmeraldana, E. aerocarpa. E. krausii. E. tenebrica. E. laui) und mit der Subsektion Neoporteria >E. chilensis,
E. subgibbosa. E. sociabilis. E. senilis, E. villosa).
Resumen. El genero Eriosyce Phil. (("actaceae-Notocacteaei ha sido revisado у ampliado, aclualmente incluye
33 especies у diversas categories infraespecificas que anteriormente fueron clasificadas en Neoporteria sens,
lat. Eriosyce se encuentra distribuido a lo largo del oeste de America del Sur. de la parte central de Chile
hasta el Sudoeste de Peru, tambien en el lado este de la Cordillera de los Andes, oeste de Argentina, de
Mendoza a Sella. Los datos fundamenlales en el presente estudio fueron obtenidos por el principal autor
< Kattermann i en el cam no. у documenlados con materia) de herbano. Detalles de bibliugrafiay nomenclatura
lApendice IV) у la filogenia del grupo lApendice III) fueron contribuidos por varios eolaboradores. Se describe
una especie nueva, E. laui J. Luthy lApendice II). Se efectuaron estudios de morfologia de las semillas
utilizandn Microscopio Electronic*) de barrido. Se presentan claves de identifkaciun para todos los niveles.
descripciones у folografias a color y/o dibujos. El genero se subdivide en dos secciones con varias
subsccciones. los nombres у combinaciones nuevas se encuentran en el Apendice I: secc. Eriosyce con subsecc.
Eriosyce IE. aurata. E. rodentiophilai, subsecc. Pyrrhocactus >E. bulbocalyx. E. strausiana, E. umadeave, E.
andreaeana, E. villicumensis) у subsecc. Islaya <E. islayensis); у secc. Neoporteria con subsecc. Harridwactus
IE. recondita. E. curvispina, E. kunzei. E. limariensis, E. confinis, E. crispa. E. garaventae, E. aspdlagae, E.
marksiana, E. engleri. E. heinrichiana. E. napina. E. (xlieri. E. taltalensis, E. occulta), subsecc. ChUemyce
iE. esmeralduna, E. aerocarpa. E. krausn, E. tenehrica, E. laui) у subsecc. Neoporteria <E. chilensis. E.
subgibbosa. E. sociabilis. E. senilis. E. villosal.
Succulent Plant Research
Volume One
Eriosyce (Cactaceae):
The genus revised and amplified
Fred Kattermann
Introduction
Eriosyce, as here redefined, is a genus of 33 species of subglobular, globular, elongated
or rarely almost columnar form. It can be recognized by the presence of wool and
bristles on the pericarpel and tube (except for Eriosyce laui J. Luthy sp. nov., in which
the flower is naked: see Appendix II), a hollow, dehiscent fruit, which has a basal
opening, the seeds loose within and seeds with a small hilum (hilum-micropylar region
or HMRi. The hilum rim at the micropyle of the seed is often modified by a thickening,
forming a partial to complete annular structure around the micropyle. Floral
characters vary somewhat within the genus. The flower-tube is funnelform and usually
has some wool or bristles in the axils of the bract-scales. Some flowers are adapted for
hummingbird pollination, and in some of these the flower is almost tubular.
Eriosyce belongs to tribe Notocacteae, and occurs from central Chile, with its
southernmost limit near Concepcion at 37°S, to southern Peru, where it reaches the
Rio Omas at 14CS, and eastward into north-west Argentina, from Mendoza to Salta
Map 1).
Previous study
The most recent taxonomic treatment of this group of plants was that of Backeberg
1958-62). The later treatment by F. Ritter (1980) can only be viewed as a catalogue
at population descriptions because it lacks a key to species. Backeberg's treatment
separated these plants into the genera Eriosyce, Islaya, Horridwactus, Neochilenia,
Neoporteria and Pyrrhocactus. Donald & Rowley (1966) consolidated the above genera
in Neoporteria, except for Eriosyce. In the past Eriosyce was always excluded from any
revision of this South American group, as in the work of Donald & Rowley, although
they did suggest that Eriosyce was closely related. In a complete study one cannot
exclude Eriosyce. Nevertheless, their work laid the foundation for the present
treatment.
The group as a whole has had a chequered taxonomic and nomenclatural history
since the first of its species were discovered more than a century and a half ago. Some
:>f the chief landmarks are as follows:
7
Kattermann: Eriosyce
Historical landmarks
1829. Cactus curvispinus, the earliest known species now referred to Eriosyce, was
named by Bertero. It was described by Colla in 1833. Meanwhile, in 1831, Haworth
described Echinocactus subgibbosus, later to be the type of Neoporteria. Various other
species, including the type of Eriosyce itself, Echinocactus sandillon Gay (= E. auratus
Pfeiffer, described a few months earlier), were named before 1850.
1872. Dr R.A. Philippi (Descr. pi. nuev., 63. 1872) proposed the genus Eriosyce for E.
sandillon, separating it from Echinocactus by reason of the floral tube, produced
beyond the ovary and spiny at the apex, and the dry fruit Ctubo elongato supra
ovarium producto, apice spinas cum petalis gerente; bacca exsucca').
1922. Apart from Eriosyce sandillon, all known plants of this South American group
had been included in Echinocactus Link & Otto until Britton & Rose's major sub­
division of the latter genus in vol. 3 of their monograph The Cactaceae. Britton & Rose
recognized Eriosyce as a monotypic genus, using the name E. ceratistes (now considered
misapplied) for the only species and referred all but one of the other relevant species
to two other genera. Those with funnel-shaped flowers and bristles on the flower-tube
were placed in the new genus Neoporteria, with Echinocactus subgibbosus Haworth as
the type, and those with campanulate flowers in Malacocarpus Salm-Dyck (nom.
illegit., ICBN Art. 64), which is now for the most part included in Parodia Speg. sens,
lat.). Echinocactus villosus Monville was listed as an uncertain species, perhaps
referable to Matucana. The exact identity of the original E. villosus remains uncertain,
along with several of the seven species Britton & Rose recognized in Neoporteria.
Relevant species placed by Britton and Rose in Malacocarpus were M. reichei
(Schumann) B. & R., M. napinus (Philippi) B. & R., M. catamarcensis (Spegazzini) B.
& R., M. islayensis (Forster) B. & R., M. strausianus (Schumann) B. & R., M.
tuberisulcatus (Jacobi) B. & R. and M. curvispinus (Bertero ex Colla) B. & R. Their
broadly based concept of Malacocarpus included plants from Bolivia. Peru, Brazil,
Paraguay and Uruguay, as well as from Argentina and Chile. They distinguished it
from Eriosyce by the absence of spines at the top of the fruit.
1929. Alwin Berger, in his book Kakteen, divided Malacocarpus, referring the hollow-
fruited species toNotocact'us (flowers yellow) and Pyrrhocactus (flowers flame-coloured
or reddish yellow), with Echinocactus strausianus Schumann as its type. Berger also
included E. catamarcensis Speg., E. tuberisulcatus Jacobi and Cactus curvispinus
Bertero ex Colla. This classification distinguished the Argentinian forms now referred
to Eriosyce from other Notocacteae in Argentina, like Parodia (Notocactus), whilst
confirming the close relationship of plants from both sides of the Andes.
1934. Curt Backeberg proposed the genus Islaya, with Echinocactus islayensis Forster
as type species, separating it from Malacocarpus on account of its hollow fruit.
1935. Backeberg (in Backeberg & F. Knuth, Kaktus-ABC, 299, without Latin diagn.)
proposed the genus Chilenia for part of Neoporteria B. & R., but overlooked that one
В
Kattermann: Eriosyce
of the species he included, Echinocactus subgibbosus, was the type of the latter name.
After abortive attempts by Backeberg to undo this mistake himself, it was eventually
rectified in 1942 by B. Doelz (in Fedde, Repert. Sp. Nov. 51: 60) who published Neo-
chilenia Backeberg ex Doelz for the part of Neoporteria with funnelform, rotate flowers
(though it is arguable that the name Nichelia Bullock (published in 1938), should take
precedence).
Meanwhile, Backeberg (Blatter f. Kakteenforschung 1938(6): [16, 21J) proposed
another new genus, Horridocactus Backeberg, based on Cactus horrid us Colla, for
Chilean species allied to the Argentinian Pyrrhocactus but with the ovary nearly
naked. His proposal drew attention to the differences between plants growing on the
two sides of the Andes, which he argued were a barrier between the two groups, but
the basis for separating the Chilean plants into Horridocactus and Neochilenia (the
amount of bristles etc on the floral tube) now seems inadequate.
1957. Y. Ito {Explanatory diagram of the Austroechinocactinae, 148, 292) proposed the
genus Thelocephala for Neoporteria napina and another species with prominently
tubercled stems.
1959. F. Ritter (in Cactus (Paris) 14(63): Suppl. [5]. 1959) took up Frit's proposed but
undescribed genus Chileorebutia, based on Echinocactus reichei Schumann, for various
species with tuberous roots and a stem reminiscent of Rebutia spp. These included
Neoporteria napina, the type of Ito's Thelocephala, and so the name Chileorebutia Fric
ex Ritter is illegitimate.
In the same year Ritter (in Succulenta 1959( 10): 129-131) also rejected Backeberg's
division of Pyrrhocactus, broadening the genus to include Horridocactus and
Neochilenia.
1962. Backeberg's monumental work. Die Cactaceae, completed publication. He
continued to place the plants from the east and west side of the Andes in different
generic groups and restricted Pyrrhocactus to species from Argentina. His 'Pacific'
genera included Eriosyce, Horridocactus, Islaya, Neochilenia (incl. Chileorebutia and
Thelocephala), Neoporteria and Rodentiophila (an invalid name).
In the 1962 version of his 'phylogenetic' classification, F. Buxbaum (in Krainz, Die
Kakteen, Lfg. 21) accepted Pyrrhocactus, Islaya and Neoporteria (incl. Chilenia,
Chileorebutia, Horridocactus, Neochilenia and the misplacedReicheocactus) and placed
them in his tribe Notocacteae. At that time, Buxbaum regarded the relationships of
Eriosyce and Rodentiophila as uncertain, grouping them next to Eulychnia in tribe
Leptocereeae, but by the time the final version of his system appeared in 1974 (in
Endler, J. & Buxbaum, F.. Die Pflanzenfamilien der Kakteen, ed. 3), he had realized
:hat Eriosyce and Rodentiophila should be placed in the Notocacteae (I.e., page 161,
note no. 48).
1966. Donald & Rowley (in Cact. Succ. J. Gr. Brit. 28: 54-58, 74-77, 83) reunited
Hurridocactus, Islaya, Neochilenia, Pyrrhocactus and Thelocephala under Neoporteria.
Their work was based on a statistical^analysis of floral and other characters. Plants
9
Kattermann; Eriosyce
growing on the eastern side of the Andes were once again classified with plants
growing on the western side. Islaya from Peru was also included, but Eriosyce was
excluded, largely, it seems because of the nomenclatural implications: lNotocactus... and
Eriosyce run so close that they can be separated by combinations of characters, no one
of which is absolute. Artificial considerations, like convenience of use and desire to
retain old familiar names tip the balance here.'
1980. In his monograph, Ritter (Kakteen in Sudamerika, vol. 3) retained the narrower
generic concepts proposed in his earlier work, recognizing£>/osycc, Islaya, Neoporteria,
Pyrrhocactus (including Horridocactus and Neochilenia) and Thelocephala as genera.
He also described many new species. His locality data greatly helped the present
author to collect the material used in this revision.
1986. For its 'consensus' list of accepted genera (Hunt & Taylor, eds., in Bradleya 4:
65-78), the IOS Working Party adopted the Donald & Rowley circumscription of
Neoporteria, noting that Eriosyce is 'An older name than Neoporteria with which it
might otherwise be combined'. The IOS list has been adopted for various subsequent
treatments, including Adriana Hoffmann's work on the cacti of Chile (Hoffmann 1989).
Scope of present study
The writer's conclusions are based on more than a decade of studies of about 500
collections or population samples made during many field trips to Chile and Argentina.
Observations in the field and greenhouse were documented with photographs of plants,
flowers and fruits, herbarium vouchers and many notes and analytical drawings. A
detailed study of seed-morphology was carried out using both the light microscope and
Scanning Electron Microscope (SEM). Numerous vegetative, floral, fruit and seed
characters were considered, and all the recognizable species scored for 58 characters
considered to be of taxonomic value. The resulting data-matrix was subjected to a
cladistic analysis to test my intuitive conclusions concerning the classification and
phylogeny of the group.
General conclusions
As data accumulated, it became increasingly apparent to me that all the species under
discussion should be included in a single genus. The data also suggested that at the
species level a much broader approach was necessary than had been adopted of late.
Many of the described taxa are local populations representative of much larger species
complexes. Number of spines and their length, number of ribs, quantitative covering
of the flower tube and ovary by bract-scales and wool etc. are environmentally
influenced and are therefore of limited use for classification. Past misunderstandings,
incomplete data, ignorance of the rules of nomenclature, and a tendency among authors
to describe individual plant populations as separate species, have led to the publication
of a large number of unacceptable or doubtful names and consequent taxonomic
problems. Many of the names must now be consigned to synonymy or discarded for lack
of adequate typification.
10
Kattermann: Eriosyce
Outline of proposed classification
Fieldwork carried out from 1977 to 1986 convinced me that Eriosyce in its narrow
original sense is only part of a larger and closely related group which includes plants
growing in north-western Argentina, central to northern Chile and southern Peru.
Consequently I am proposing to enlarge Eriosyce by merging it with Neoporteria sensu
Donald & Rowley. Unfortunately the rules of nomenclature prescribe that the larger
group lNeoporteria) must be sunk into the smaller (Eriosyce), because Eriosyce is the
older name. In defence of Eriosyce, however, it can be argued that it has been a well-
known name, used with a consistent circumscription, since Britton & Rose's mono­
graph, whereas the circumscription of Neoporteria has varied widely, from few to many
species. Included in the expanded concept of Eriosyce are Chileorebutia Ritter, Islaya
Backeberg, Horridocactus Backeberg, Neochilenia Backeberg, Neoporteria Britton &
Rose, Pyrrhocactus A. Berger and Thelocephala Y. Ito.
I am further proposing to divide the expanded genus into two sections, sect. Erio­
syce, and sect. Neoporteria. Sect. Eriosyce is split into subsection Eriosyce, subsect.
Islaya and subsect. Pyrrhocactus. Sect. Neoporteria is divided into subsect. Chileosyce
<Chileorebutia Ritter pro parte), subsect. Horridocactus and subsect. Neoporteria.
The species recognized in subsect. Eriosyce are E. aurata and E. rodentiophila.
These are often very large globular cacti, reaching up to 50 cm in diameter. The
bristles on the floral tube are spine-like. The fruit is greenish to red, dehiscing by a
basal opening after complete abscission from the plant. The subsection is distributed
along the western side of the Andes from 28-33c'S. In subsection Pyrrhocactus we have
E. umadeave, E. strausiana, E. andreaeana, E. bulbocalyx and E. villicumensis. The
bristles on the floral tube are thin, not thick and spine-like. Fruit dehiscence is by a
partial circumscissile splitting at the base of the fruit, except for E. bulbocalyx where
the fruit is indehiscent. Abscission of the fruit follows shortly thereafter. These plants
grow on the eastern side of the Andes, almost paralleling the distribution of subsect.
Eriosyce. Subsection Islaya includes only E. islayensis. The fruit in this plant elongates
and enlarges, becoming balloon-like. Its distribution is along the Pacific coast from
northern Chile to central Peru.
Section Neoporteria also grows on the western side of the Andes. The fruit is red,
nften elongating at the time of maturity. Dehiscence is by a basal opening. Abscission
is variable, since in some species the fruit does not free itself from the plant and can
be removed only with some force. For part of its distribution it overlaps that of subsect.
Eriosyce, where in many localities plants from these two taxa grow side by side.
However, the distribution of sect. Neoporteria extends further south, to 37°S. Section
Neoporteria is further divided into subsections: Subsect. Chileosyce comprises E. aero­
carpa, E. esmeraldana, E. krausii, E. tenehrica and E. laui, which is here described for
the first time. Excepting the last-named, these species have porrect bristles on the
flower-tube and fruit, with an early abscission of the fruit and usually different seeds.
Subsect. Horridocactus comprises E. confinis, E. crispa, E. curvispina, E. engleri, E.
kunzei, E. aspillagae, E. garaventae, E. heinrichiana, E. limariensis, E. napina, E.
marksiana, E. odieri, E. occulta, E. taltalensis and E. recondita. These plants have a
more fleshy fruit wall, abscission is incomplete, since the fruit can be removed only
with some force, and the flowers have the perianth-segments curving outward. Lastly,
11
Kattermann: Eriosyce
subsect. Neoporteria, comprising E. chilensis, E. senilis, E. sociabilis, E. subgibbosa and
E. villosa, has flowers which have incurved perianth-segments and are adapted for
hummingbird pollination.
Characters used in classification
Previous classifications of Eriosyce and allied genera have largely relied on a relatively
few characters, such as flower shape, style and stigma colour, tube armature, and
whether the fruit is fleshy or dry (cf. Donald & Rowley 1966). In the course of my work
I have identified useful additional characters, notably in the form of the nectary, the
junction of style and stigma, the attitude of the stigma-lobes, and the mode of
dehiscence of the fruit. The Scanning Electron Microscope has facilitated examination
and comparison of the seeds, and several characters of taxonomic importance can now
be recognized.
On the basis of my fieldwork and observations of documented material in
cultivation, I was able to distinguish 33 species. To test my ideas on their classification
and inter-relationships, I scored them for numerous characters, eventually reduced to
58. After various refinements, the resulting data-matrix was subjected to a cladistic
analysis by Dr Robert Wallace (see Appendix III). The characters, and the character-
states observable for each, are listed in Table 1 (pages 126-128), which is in itself a
summary of the morphological diversity present in the genus. Brief notes are given
below on the vegetative and floral characters employed in the analysis, followed by a
systematic overview of fruit and seed characters in the genus.
Vegetative characters (Matrix nos. 1-19)
1. Stem elongation. Most species of Eriosyce are globular or subglobular in habit but
the stem may also be elongate or columnar, as in some forms of E. subgibbosa. 2. Stem
diameter varies from 1.5—40 cm. 3. Stem firmness. In most species the stem-surface is
not easily depressed, but soft stem tissue has been observed in E. aspillagae, E. crispa
and E. villosa. 4. Taking the genus as a whole, rib number varies from 7-30 or more
but the range in individual groups and species overlaps to such as extent that it is of
little use as a 'key' character. In many populations the number of ribs increases with
age. 5. Rib shape. In all species the ribs are somewhat notched or depressed between
the areoles and in some the ribs are dissolved into elongate or isodiametric tubercles.
6. When the ribs are deeply notched, one can often observe a chin-like protrusion below
the areole. 7. The areole itself may be superficial (slightly raised to almost flat) or
sunken to a varying degree. 8. Areolar wool is always present, and may be very short
or longer, in some species 3 mm or more and floccose. 10. Apex. Depending on the
development of areolar wool, this varies from naked to densely woolly.
The only spine characters used in the analysis were as follows. Radial spines:
10. Thickness, 11. Attitude and 12. Curvature. Central spines: 13. Presence I absence, 14.
Thickness, 15. Attitude and 16. Curvature. Spine number is very variable within taxa
and is of little value as a diagnostic character.
17. Roots. The root system in plants of this genus varies from fibrous to turbinate
or tuberous. 18. In thick-rooted species, the presence or absence of a neck-like
constriction where the root joins the plant-body is diagnostic. 19. The root-epidermis
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