PHYLOGENETIC RELATIONSHIPS WITHIN
NOTOCACTUS - PARODIA - PUZZLE
A. B. Doweld
National Institute of Carpology
(Gaertnerian Institution), Moscow
Dedicated to 100th Anniversary of
Franz Buxbaum (1900-1979),
Eminent Austrian cactologist
Recently after detailed analysis of available literature data together
with some additional original observations on seed anatomy and morphology of the
representatives of the genera Notocactus (K. Schum.) Fric, Eriocactus, and Parodia
(Doweld, 1999а), the interrelationships of
these taxa remained unclear yet, and the creation of a new genus Ritterocactus
Doweld to accommodate species of notocacti, formerly ascribed to a distinct subgenus Neonotocactus
Backeb. (Notocactus), did not solve the problem of generic limits within the group.
As we pointed out earlier (Doweld, 1999а: 24),
the whole picture of the phylogeny and systematics of Notocactus-Parodia puzzle
might be solved only with additional seed survey of the genus Parodia. The data on
general seed morphology, published by Brandt (1982, 1986, 1989), are insufficient to use
them in modern biosystematic analysis of these taxa, because there are no SEM
illustrations of the ultrasculpturing of the seed surface, which is used as a leading
character in taxonomic constructions within the cactus family (Doweld, 1995, 1999, 2000,
etc.).
We attempt to elucidate phylogenetic relationships of above mentioned
genera with the assistance of additional original seed study of some species of Parodia,
Eriocactus, Brasilicactus, Notocactus, and Wigginsia (methods after Doweld,
1999b; vouchers were housed in the Carpotheca of the National Institute of
Carpology (Gaertnerian Institution), Moscow). Other data sets of vegetative and
reproductive characters were obtained from original observations in various horticultural
collections and literature sources (Backeberg, 1959; Brandt, 1989; Gerloff et al.,
1995; Glaetzle & Prestle, 1986; Neduchal, 2000а, 2000b; Stuchlik, 1993).
Delimitation of Notocactus and Ritterocactus
In the proposing to delimit two genera, Notocactus (as 'Peronocactus')
and Ritterocactus, we were based on the quite different seed and cuticular
sculpturing within two genera. In addition to first observations of Glaetzle and Prestle
(1986), the incomplete data were also provided by Stuchlik (1993) and Gerloff et al.
(1995). With the addition of some original observations, we have found several
intermediate forms which have been inadequately interpreted by Glaetzle and Prestle
(1986).
These authors proposed to relate Notocactus horstii and N.
fuscus with N. scopa-concinnus-Group (belonging to Notocactus s.str.).
However, this proposal might not be adopted and confirmed by seed ultrasculpturing. The
cuticular seed sculpturing of N. horstii is longitudinally-oriented striate, not a
radially-oriented striate or reticulate-striate which is characterized for N.
scopa-concinnus-Group. The species N. horstii, although formerly included into Notocactus
(Doweld, 1999a: 21) follow these authors, should be related to Wigginsia (W.
nothohorstii Doweld nom. nov.) with a similar cuticular seed sculpturing
and similar construction of spination, areoles, and rib number (cf. Wigginsia for
more details).
The same is true for N. fuscus: the species has a peculiar
longitudinally-oriented striate cuticular sculpturing which is typical of Wigginsia
and some Ritterocactus species, but not to N. scopa-complex. In addition, N.
fuscus shared a similar 4-6-merous construction of radial spines complex and mostly
4-merous central spination with Ritterocactus, but not with Wigginsia which
is characterised by mostly 5-merous system of radial spines and mostly solitary (rarely 4)
central spine. Therefore, this species should be included into Ritterocactus (Ritterocactus
fuscus (F. Ritter) Doweld, comb. nov.) as opposed to our former opinion
(Doweld, 1999a: 21).
With the addition of our original observations of seed anatomy,
morphology and ultrasculpturing of Wigginsia horstii F. Ritter (= Notocactus
neohorstii Theunissen, = Peronocactus neohorstii (Theunissen) Doweld), we came
to conclusion to exclude the species either from the genus Wigginsia or genus Notocactus
[in contrary to our former opinion, based on data of Glaetzle and Prestle (1986)]. The
species has a peculiar simple longitudinally-oriented striate cuticular sculpturing (like Ritterocactus),
and is characterised in addition by 4-6-merous construction of radial spination complex
(in contrast to mostly 5-merous of Wigginsia). By spination construction, the
species is undoubtedly related to Ritterocactus (R. horstii (F.
Ritter) Doweld comb. nov.), sharing a similar number of ribs (18-24) and central
spines (mostly 4-6).
With the addition of these two species, the genus Ritterocactus
become dividable into three distinct infrageneric groups, ranked below as sections. The
most archaic section is Herteria sect. nov. which is comprised of
species with a relatively low number of radial spines (8-16), 4-merous construction of
central spination complex, relatively higher number of ribs (16-24), and papillate seed
sculpturing with a simple longitudinally-oriented striate cuticular sculpturing (sometimes
alike radially-oriented cuticular sculpturing). The new section Uebelmannia
sect. nov. consists of only 3 species which is characterised in contrast by a
relatively low number of ribs (10-16), mostly solitary central spine, and preserving in
addition a low number of radial spines (8-12) of 4-merous basic construction. The seed
sculpturing of this section is apapillate (smooth - derivative of papillate type), but
with again longitudinally-oriented cuticular sculpturing. The type-section, Ritterocactus,
possesses a higher number of ribs (18-24), advanced number of radial spines (mostly 12-18)
and reduced (up to 1-3) central spination complex; the cuticular sculpturing is
longitudinally-oriented striate, but seeds possess a peculiar papillate seed sculpturing,
easily differing the section from any other sections of the genus. The similar
construction of seed sculpturing might be observed only within Parodia subgen. Obtextosperma
F. Buxb. which is also included into genus Ritterocactus as a distinct section, Pseudoparodia
sect. nov. (cf. Obtextosperma for more details).
With the exclusion of these species, the genus Notocactus
becomes more homogeneous. The genus might be divided into two gross groups (ranked as
subgenera) with some additional infrageneric entities within each group. The most
primitive subgenus, Notocactus, consists of species with papillate seed sculpturing
with a smooth or radially-oriented striate or finely reticulate-striate cuticular
sculpturing. They are characterised in addition by 4-merous construction of radial
spination complex (4n: 16-24), mostly also 4-merous system of central spines (4, rarely
less), and relatively low number of stem ribs (never more than 18: 6-18). The basic number
in the evolution of number of ribs is probably 6 (6-7) and is preserved only within N.
oxycostatus having a very low number of radial spines (4 only) and a solitary central
spine in addition to practically smooth cuticular sculpturing. The combination of these
characters leads us to hypothesise the derivation of smooth type of cuticular sculpturing
from that of striate one in primitive notocacti.
The subgenus is divided into 3 series: ser. Notocactus (with
almost smooth cuticular sculpturing), ser. Linkianae ser. nov. (with a fine
striate cuticular sculpturing, and low number of radials [mostly 10]), and ser. Praeparodia
ser. nov. (with a specific fine reticulate-striate cuticular sculpturing, higher
number of radial spines [14-24]). In Praeparodia (T.: N. minimus Fric &
Kreuzing.) we could find a specific reticulate-striate cuticular sculpturing which is a
basic type for the origin and evolution of that of Parodia s. str. (excl. subgen. Obtextosperma
and Protoparodia!).
The subgenus Scopacactus subgen. nov. (T.: N. scopa)
is characterised by a higher number of ribs (18-36, never less), mostly 4-merous central
spination complex, and preserving 4-merous construction of radials (8-12 in N.
concinnus to 40 in N. scopa ssp. scopa). The subgenus might have a
common origin with ser. Notocactus in particular by similar vegetative
characteristics, but this phyletic lineage diverged very early and it seems have had a
long distinct evolutionary history. In the representatives of subgenus, there is a
progressive evident development of reticulate-striate cuticular sculpturing, which was an
archetype of cuticular sculpturing in Parodia s. str.
Relationships of Eriocactus
After nomenclatural elucidation of the lectotypification of Notocactus,
the genus Eriocactus is nomenclaturally rehabilitated and become available for use
in current plant taxonomy (Doweld, 2000a). The formal proposal to conserve this
generic name over its earlier nomenclatural synonym, Eriocephala Backeb., is
already published (Doweld, 2000a) and awaited a special decision of Nomenclatural
Committee and forthcoming International Botanical Congress.
In our former paper (Doweld, 1999a: 23) this genus has been
treated as Notocactus, and now its taxonomic infrageneric system is corrected by
formal transferring of established taxonomic entities from Notocactus to Eriocactus
(Doweld, in press). We found (as formerly Glaetzle & Prestle, 1986) the genus
heterogeneous by seed cuticular sculpturing, being consisted of species with smooth
cuticular sculpturing (E. magnificus F. Ritter) and striate one (all other
species). A distinct subgenus, Eonotocactus Doweld, has been created to emphasise
the distinctness of E. magnificus. The seeds of the genus indicate of close
phylogenetic relationships with Notocactus, especially with ser. Notocactus,
having a similar smooth cuticular sculpturing. Moreover, the exotestal cells of Eriocactus
are elongated, rectangular, with a finely longitudinally-oriented reticulate-striate
cuticular sculpturing like in advanced taxa of Notocactus and primitive members of Parodia
(sect. Sulcatae Weskamp). It seems that Brandt (1982, 1986, 1989) was quite right
in proposing a close phylogenetic relationships between Eriocactus and his subgen. Parodia,
but nevertheless, the occurrence of smooth cuticular sculpturing is indicative of somewhat
another evolutionary direction of Eriocactus historical development. In addition to
seed characters, close phylogenetic relationships between Eriocactus and Notocactus
subgen. Notocactus are revealed by possessing a similar 4-merous construction of
radial spination (4n: from 4-8 in E. schumannianus ssp. claviceps to 16-20
in E. leninghausii), mostly 3-4-merous system of central spines, 6n basic number of
stem ribs. The origin of higher number of ribs (45) and 16-20 radials (from 4-8)
illustrates the distinctness of this phyletic line from Notocactus, confirming its
taxonomic distinctness.
Relationships of Brasilicactus
The proposal to conserve the later nomenclatural synonym, Brasilicactus
Backeb., against its earlier one, Acanthocephala Backeb. (Doweld, 2000b),
rehabilitates the name in current use. Recently we amplified the genus (Doweld, 1999a:
24) to include the genus Brasiliparodia F. Ritter; the amplification has been based
on results of study of seed morphology (Glaetzle & Prestle, 1986). An additional
argument pro has been later received from molecular studies of Nyffeler (1999). Our
original observations (this issue) confirmed the naturalness of the uniting of two genera,
however, the border between them is remained in number of ribs (Brasilicactus has
always more than 30, Brasiliparodia: 16-24) and radial spines (Brasilicactus:
more 24; Brasiliparodia: from 4 to 20-24, never more). This distinctness of two
complexes should be emphasised in establishing of two series, ser. Brasilicactus
and ser. Ritterocactoides ser. nov.
The phylogenetic relationships of the genus are revealed by occurrence
of specific longitudinally-oriented fine striate cuticular sculpturing, which is
indicative of close phylogenetic relationships with Ritterocactus (sect. Herteria)
and less close with Wigginsia (sect. Eowigginsia). The relationships with Wigginsia
are less evident if to take into consideration the construction and number of radial
spines (always 4n-merous, and number varying from 4-8 [in B. rechensis] to 16-24
[in B. catarinensis] to indefinitely numerous [in B. graessneri] in contrast
to 5-merous one of Wigginsia) and always 4-6-merous system of central spination (in
contrast to mostly solitary in Wigginsia). Therefore, the phylogenetic
relationships of Brasilicactus with Wigginsia are relatively remote,
although the occurrence of similar cuticular seed sculpturing are clearly indicative of
common origin of both taxa.
The phylogenetic relationships of Brasilicactus with sect. Herteria
(Ritterocactus) are more evident: both taxa have a similar construction and number
of radial spines (4n, mostly 4-16 in ser. Ritterocactoides and 8-16 in sect. Herteria)
and central spines (4-6-merous construction), number of ribs (16-22 in Ritterocactoides
& Herteria). The development of specialised types (ser. Brasilicactus)
points to a parallel and distinct evolutionary history of Brasilicactus and Ritterocactus.
With the inclusion of Parodia subgen. Obtextosperma into Ritterocactus, the
hypothesis of Brandt (1982, 1986, 1989) on the close relationships of Brasilicactus
and Obtextosperma receives a new partial support in present study of seeds and
other gross exomorphic characters, although resulted in a quite different formal taxonomic
arrangement.
Status of Parodia subgen. Obtextosperma
Buxbaum (1966) erected a distinct subgenus of Parodia, Obtextosperma,
to accommodate P. ayopayana, a peculiar species of the genus differing in seed
sculpturing and general morphology as well as in floral construction, fruit, and general
habit. Later Brandt (1982) erected two series within the subgenus, Calyptospermae
and Intectospermae, and amplified the subgenus by adding P. miguillensis and
other closely related forms. Brandt followed Buxbaum in regarding this subgenus most
primitive within Parodia, and thereby providing a missing phylogenetic link between
Parodia and Notocactus s.l. in their classification schemes.
In our present study we confirmed the distinctness of P. ayopayana
and P. miguillensis from the rest of Parodia in having a peculiar
hilum-micropylar structure, longitudinally-oriented striate cuticular sculpturing
reminiscent that of highly specialised Ritterocactus (sect. Ritterocactus).
The close phylogenetic relationships is supported by similar 4-merous system of radial
spines (12-18 in sect. Ritterocactus & subgen. Obtextosperma), but
differing in prevalence of 4 (or duplicate 8: P. miguillensis) central spines
within Obtextosperma, and reduced number of centrals within sect. Ritterocactus
(1-3; although in more primitive sect. Herteria the number of centrals is
constantly 4 or more, to 6-8). The number of stem ribs is nearly similar: 8-24 in P.
miguillensis and 18-24 in sect. Ritterocactus (16-22 in sect. Herteria
and 8-16 in sect. Uebelmannia). The occurrence of prevailing number of radial
spines (10) in P. ayopayana points to a somewhat likeness of the species with the
species of Wigginsia which are characterised by 5-merous system of radial
spination. Taking into consideration the basic similarity in seed construction, as well as
in other vegetative characters, we conclude to transfer the subgen. Obtextosperma
to Ritterocactus in the rank of a distinct section, Pseudoparodia sect.
nov.; two species only combined into Ritterocactus, R. ayopayanus
(Cardenas) Doweld comb. nov. and R. miguillensis (Cardenas) Doweld comb.
nov. (others are temporary treated as synonyms pending further studies of the group).
The series of Brandt (1982) are not recognised in present study.
Amplification of Wigginsia
Recently we established a distinct section of Notocactus (= Peronocactus),
sect. Eowigginsia (Doweld, 1999a: 24), based on Notocactus horstii F.
Ritter. The species is differing from the rest of Notocactus by peculiar 5-merous
construction of radial spines (10-15) which is typical of Wigginsia. In addition,
the clearly differentiated longitudinally-oriented striate cuticular sculpturing of seeds
points to a somewhat resemblance with sect. Herteria (Ritterocactus). By the
combination of these characters, the species should be included into Wigginsia (W.
nothohorstii Doweld nom. nov.) as a possible archaic, living missing link
between highly specialised Wigginsia and more primitive Ritterocactus and Brasilicactus.
The section Eowigginsia is transferred to Wigginsia: W. sect. Eowigginsia
(Doweld) Doweld comb. nov.
The species W. horstii F. Ritter (= Notocactus neohorstii Theunissen)
has been placed into Ritterocactus (sect. Herteria) on the base of 4-merous
construction of radial spination, 4-6-partite central spines, similar
longitudinally-oriented fine striate cuticular sculpturing.
Another species of Notocactus, N. werdermannianus, should
be also affiliated with Wigginsia in having similar 5-merous radial spination
complex (15-20) and 4-partite central spines, and a distinctive longitudinally-oriented
fine striate cuticular sculpturing of seeds (vide Glaetzle & Prestle, 1986).
The species is tentatively placed into sect. Eowigginsia, pending further revision
of the genus.
As amplified above, the genus Wigginsia becomes a more
definitive generic entity, characterising by 5-merous radial spination complex and
distinctive papillate crest-like seed sculpturing with a longitudinally-oriented striate
cuticular sculpturing. The phylogenetic roots of the genus are with that of Ritterocactus
(in similar distinctive cuticular seed sculpturing), but nonetheless, Wigginsia
represents itself as an easily recognised distinct genus.
Relationships of Parodia subgen. Protoparodia
In erecting a distinct subgenus of Parodia, Buxbaum (1966)
regarded it a possible notocactoid part of Parodia by a very similar structure of
seeds. In the studies of Brandt (1982, 1986, 1989), subgen. Protoparodia become a
central core of the evolution and differentiation of other higher taxa of Parodia,
being originated within subgen. Obtextosperma and giving off 11 taxonomic entities
(incl. subgen. Parodia s.str.). However, the numerous series created by Brandt
(1982) in addition to those established by Buxbaum (1966), could not be supported in
present limited seed morphological study. Leaving the infrastructure of this taxonomic
entity beyond the scope of present paper (pending extensive filed and morphological
studies), we ascertained the phylogenetic closeness of Protoparodia to Wigginsia
and Ritterocactus.
The seeds of Protoparodia are variable in uniting advanced and
archaic types of seed cuticular sculpturing. The seeds of P. maassii are somewhat
advanced in possessing practically radially striate cuticular sculpturing with smooth
apexes of the cupolas of exotestal cells, on which there are no rudiments of
longitudinally striateness. In contrary, the seeds of P. mairanana, P. hausteiniana
are characterised by clearly differentiated longitudinally striate cuticular sculpturing
pointing to phylogenetic closeness to sect. Herteria (Ritterocactus).
Moreover, sometimes observed cuticular 'straps' of longitudinally elongated striate
structures at the cells borders are also indicative of morphogenetic affinity of these
species with sect. Herteria. In advanced members of Protoparodia the
longitudinal striateness disappears, and the seeds are characterised by these 'straps'
with unclear striate surface of exotestal cells (P. procera, P. compressa). In
addition, Protoparodia has a similar structure of central spination (usually 4
central spines, sometimes duplicated in 8 or having an additional splitting of spine
meristems resulting in appearance of 6 spines) with that of sect. Herteria (4-6);
the radial spines (counts after Brandt, 1989) perhaps have a 4-partite structure: 8-24 (or
even up to 30 in P. laui), although Brandt (1989) listed for some strictly limited
species (P. neglecta, P. chaetocarpa, and P. schwebsiana) several counts
supposing a 5-merous system of radial spines (25, 10-15, and 10 respectively). At present
I am inclined to regard its as erroneous counts of rare extremal variation of radials
within these species (f.e. 25 might be corrected as well to 24). Although the appearance
of 5-merous counts of radial spination might confirm the existence of close phylogenetic
relationships of Protoparodia as well as with sect. Eowigginsia (Wigginsia),
which is closely allied to primitive sect. Herteria (Ritterocactus).
In contrary to subgen. Parodia, in the seeds of subgen. Protoparodia
there are no reticulate-striate characteristics of cuticular sculpturing. The cuticular
sculpturing of Protoparodia is clearly indicative of phylogenetic affinities with
archaic elements of Ritterocactus (sect. Herteria) and Wigginsia
(sect. Eowigginsia). On the basis of similar seed characteristics, radial and
central spination, and similar floral features (vide Buxbaum, 1966), the subgen. Protoparodia
is excluded from the genus Parodia and raised to generic status by erecting of a
distinct genus Bolivicactus gen. nov. The genus is related to Ritterocactus
and Wigginsia, but not to Parodia s.str. and allied genera Notocactus
s.str. and Eriocactus.
Phylogenetic roots of Parodia s.str.
The genus Parodia, re-circumscribed above to exclude two
aberrant subgenera Obtextosperma and Protoparodia, become a more natural
entity. The lack of resemblance of the Parodia seeds with that of Obtextosperma
and Protoparodia points to another phylogenetic stock to which genus Parodia
in a limited circumscription should be affiliated. I noticed within primitive (after
Brandt, 1989) species of his subgen. Parodia (ser. Eriospermae: P.
chrysacanthion, P. nivosa) the occurrence of the rudiments of specific
reticulate-striate seed cuticular sculpturing, which has been observed only within the
genera Eriocactus and Notocactus s.str. (in present redaction). It looks
very interesting to find a partial support of the hypothesis of Brandt (1982, 1989) on the
close relationships between eriocacti and parodias in strict sense. In addition to gross
morphological characters (Brandt, 1982, 1986), soundly pointing to a common stock of these
taxa, at present the somewhat similarity in the cuticular sculpturing should be added.
However, it should be noted that the seeds of both taxa are not identical, even in
cuticular sculpturing, and a certain morphogenetic gap between these two types of seeds
might not be overlooked. However, in the advanced members of Parodia (P.
penicillata-P. formosa), the reticulate-striate sculpturing are gradually disappeared
resulting in the nearly smooth seeds of P. microsperma.
In the seeds of ser. Praeparodia ser. nov. (Notocactus:
N. minimus), the peculiar reticulate-striate cuticular sculpturing might be also
observed, as well as in the representatives of the subgen. Scopacactus subgen.
nov., which has a long independent history of evolution within Notocactus s.str.
It seems that the reticulate-striate cuticular sculpturing is typical for a specific
phylogenetic phylum of Notocacteae, comprising of three distinct genera,
Eriocactus, Notocactus, and Parodia s.str. The other taxa (Brasilicactus,
Ritterocactus, Bolivicactus, and Wigginsia) have a common origin with this
branch in originating from the ancestral complex characterising by common smooth/striate
cuticular sculpturing. In present seed survey, the distinctness of the genus Parodia
from Eriocactus and Notocactus is completely supported and no grounds for
the unification of all these genera were found.
Lack of phylogenetic relationships with Frailea
In contrast to previous studies of Barthlott (1988: undocumented) and
proposals to synonymize the genus with Parodia s. lat. (Nyffeler & Eggli,
1998), we could not find any substantiative grounds to support the phylogenetic
relationships of Frailea with the genera of Notocactus-Parodia puzzle,
except for Blossfeldia (see below). The seeds of this genus (F. pumila, cf.
Figs) are characterised by trichomatous seed sculpturing and smooth (lacking) cuticular
sculpturing. The combination of such characters is very peculiar in the whole tribe
(similar seeds were observed in Blossfeldia only) and are far morphogenetically
from that of other genera of Notocactus-Parodia phylogenetic branch. Except for
minute sizes (similar to Parodia microsperma) and similar smooth cuticular
sculpturing misleading previous students of these seeds, no more similarities in the seeds
of Frailea and Parodia s.str. have been noted: the hilum-micropylar zone is
quite different, seed sculpturing is different (trichomatous and apapillate/finely
papillate), the anatomy of spermoderm is also different. After molecular studies of Notocactinae,
recently Nyffeler (1999) disavowed his former opinion, found that the genus is not related
to other Notocactinae at all. In my opinion, by seed anatomy and morphology Frailea
revealed close phylogenetic affinities with the genera of Neoporteriinae, and Copiapoa
in particular, in having a similar hilum-micropylar part, smooth cuticular sculpturing,
and papillate (ancestral for trichomatous) type of seed sculpturing. In any case, Frailea
should be excluded from the core of Notocactinae and placed to another branch of
the tribe Notocacteae; the formal proposal to conserve Parodia over Frailea
should be withdrawn as superfluous and not grounded in terms of plant phylogeny.
Lack of phylogenetic relationships with Blossfeldia
Analogous to Frailea, recently Blossfeldia has been
sunken into highly expanded (and thus become polyphyletic) genus Parodia s. lat.
(Barthlott, 1988; Hunt & Taylor, 1986). The seeds of Blossfeldia are alike
those of Frailea: minute obovoid seeds with trichomatous seed sculpturing, smooth
(lacking) cuticular sculpturing, but with hilum-micropylar zone (expanded micropyle,
strophiolate) like that with Parodia microsperma. The seeds are essentially
different from that of Parodia s.str. and no reliable morphogenetic possibilities
could be proposed to unite these distinct seed types. Therefore, the genus Blossfeldia
should be excluded from the core of Notocactinae as not having any close
relationships with none of the genus considered. Judging by seed anatomy and morphology, Blossfeldia
is a derivative of similar copiapoid line of evolution within Notocacteae, having a
somewhat similar seed type with that of Copiapoa.
Amplification of the subtribe Notocactinae
Buxbaum (1975) subdivided the tribe Notocacteae into several
subtribes, Corryocactinae, Gymnocalyciinae, Melocactinae, Neoporteriinae, and
Notocactinae. In the Notocactinae, he included 6 genera: Notocactus,
Frailea, Parodia, Blossfeldia, Uebelmannia, and Astrophytum. After our present
critical seed re-study, the genera Frailea and Blossfeldia should be
excluded from the Notocactinae and placed possibly into Neoporteriinae, in
the neighbourhood of Copiapoa. The relationships of the genus Astrophytum,
erroneously included into the tribe by Buxbaum (1958), is fully discussed in our previous
works (Doweld, 1999c, 2000c), in which the affinities of Astrophytum
with the genera of the subtribe Echinocactinae of Cacteae have been
confirmed. The seed morphology of Uebelmannia, recently studied by Nyffeler (1998),
does not show any resemblance to the seeds of other Notocactinae differing in seed
form (obovoid), hilum-micropylar part, smooth cuticular sculpturing and nearly apapillate
(smooth) seed sculpturing. The genus revealed a close affinity by seed morphology with the
genus Copiapoa and should be excluded from the Notocactinae and placed into Neoporteriinae.
Therefore, the Notocactinae should be re-circumscribed as to include 7 revised
genera, Notocactus, Eriocactus, Parodia, Brasilicactus, Ritterocactus, Bolivicactus, and
Wigginsia. The relationships of this subtribe with other Notocacteae would be
elucidated in forthcoming papers.
TAXONOMIC TREATMENT
Tribus Notocacteae F. Buxb., Madrono 14: 181. 1958.
Subtribus Notocactinae F. Buxb. ex A. Doweld, subtrib. nov. - Plantae magnae, globulares, elongatae, raro cereoideae, costis angustis, obtusis acutatisve, aculeis centralibus 4-meris, aculeis radialibus 4-5-meris (pro more (10-)15/12-24), floribus receptaculo plerumque infundibuliformis, pericarpellum lanatum, setoceum, rarius acutum spinulosum; semina reniformia, hilo magno, plano, poro mycropilario centrali prominenti vel strophiolati, testa papillata (raro secundarie apapillata), ornamentatione cuticularis striatibus vel reticulato-striatibus (raro secundarie (?) nullibus).
Type: Notocactus (K. Schum.) A. V. Fric. 7 genera.
Notocactus (K. Schum.) Fric, Cacti Coming Fash.: unpag. [3].
1928. - Echinocactus
subgen. Notocactus K. Schum., Gesamtb. Kakt.: 292. 1898. Echinocactus sect. Microgoni
Salm-Dyck in Walpers, Repert. Bot. Syst. 2(2): 273. 1843. -
Peronocactus Doweld, Sukkulenty 1999(2): 20. 1999, nom.
inval.
Lectotype (Backeb., BlStt. Kakteenforsch. 1938(6): unpag. [16]. 1938):
Notocactus ottonis (Lehmann) A. Berger, Kakteen: 212, 343. 1929. Cactus ottonis Lehm., Ind.
Sem. Hamburg.: 16. 1827.
Subgen. 1. Notocactus - Notocactus subgen.
Eunotocactus Backeb., Cact. Succ. J. (Los Angeles) 22: 153. 1950, nom.
inval. -
Notocactus subgen. Gymnocephalus Backeb. ex ?'da, Minimus 22(1-3): 29-30.
1991, nom. inval. -
Notocactus sect. Infundibuliflorales Havl'cek, Kaktusz Vilag 18(4): 73.
1989 ('1988'), nom. nud. - Echinocactus subsect. Ottonidei Lawr. in Loudon,
Gard. Mag. Ser. 3, 6: 317. 1841 («Ottonideae»).
Type: as in genus.
Ser. 1. Notocactus - Plantae magnae, costis 6-7
multiplibus (6-7 ad 14-18), aculeis radialibus 8-12(-14) (4 multiplibus!), aculeis
centralibus plerumque 4, seminibus papillatis, ornamentatione nullis (levis).
Type: as in genus.
1. Notocactus oxycostatus Buining & Brederoo in Krainz,
Kakteen 53: unpag. [CVIc]. 1973. Holotype: Buining 299 [Horst & Uebelmann 299]
(U!).
Notocactus oxycostatus ssp. oxycostatus
Notocactus oxycostatus ssp. gracilis (F.
Ritter) A. Doweld, comb. et stat. nov. - Notocactus glaucinus var. gracilis F. Ritter, Kakt.
Sudamer. 1: 168-169. 1979. Holotype: F. Ritter 1378 (U).
2. Notocactus ottonis (Lehm.) A. Berger, Kakteen: 212, 343.
1929. Lectotype (designated here): tab. 15 in Lehmann, Nov. Act.
Acad. Caes. Leopold.-Carol. 16(1): 316. 1828 (as Cactus ottonis).
Notocactus ottonis ssp. ottonis
Notocactus ottonis ssp. horstii (F. Ritter) A.
Doweld, comb. et stat. nov. -
Notocactus arechavaletae var. horstii F. Ritter,
Kakt. SYdamer. 1: 166. 1979. Holotype: F. Ritter 1027c (U).
3. Notocactus muricatus (Otto ex Pfeiffer) A. Berger, Kakteen:
210, 343. 1929. Neotype (designated here): «Cerro Branco Nr. 5, Rio
Grande do Sul, Brazil», F. Ritter 1381 (SGO 125294).
4. Notocactus minusculus Hofacker & Herm, Internoto 17: 75.
1996. Holotype: Hofacker 71 (ZSS).
Ser. 2. Linkianae A. Doweld, ser. nov. - Costae
12-14, aculei radiales plerumque 10, aculei centrales 4, semina papillata, ornamentatione
parcissime striatis (reticulato-striatis?).
Type: N. linkii (Lehmann) Herter.
5. Notocactus linkii (Lehmann) Herter, Cactus (Paris) 42: 120.
1954. Lectotype (designated here): tab. 14 in Lehmann, Nov. Act.
Acad. Caes. Leopold.-Carol. 16(1): 316. 1828 (as Cactus linkii).
6. Notocactus carambeiensis Buining & Brederoo, Kakteen
Sukk. 24: 1. 1973. Holotype: Buining 140a [Horst & Uebelmann 140a] (U!).
Ser. 3. Praeparodia A. Doweld, ser. nov. - Plantae
subglobosae vel elongatae, costis 12-18, aculeis radialibus 14-24, centralibus 4,
seminibus ornamentatione cuticularis longitudinaliter reticulato-striatibus.
Type: N. minimus Fric & Kreuzing.
7. Notocactus minimus Fric & Kreuzing., Succulenta 22:
90. 1940. Lectotype (designated here): fig. [unnumbered] in Fric
& Kreuzing. l.c. : 87.
8. Notocactus stockingeri Prestle, Succulenta 64: 226. 1985.
Holotype: Stockinger 141 (U).
Subgen. 2. Scopacactus A. Doweld, subgen. nov.
Echinocactus sect. Scopaea
Lawr. in Loudon, Gard. Mag. Ser. 3, 6: 316. 1841 («Scopaeae»). - Plantae
cereoideae, elongatae, costis 16-36, aculeis radialibus 10-40, centralibus plerumque 3-4,
seminibus papillatis, ornamentatione cuticularis longitudinaliter reticulato-striatibus.
Type: N. scopa (Sprengel) A. Berger.
9. Notocactus concinnus (Monville) A. Berger, Kakteen: 210, 343.
1929. Neotype (required).
Notocactus concinnus ssp. concinnus
Notocactus concinnus ssp. agnetae (van Vliet)
A. Doweld, stat. nov. -
Notocactus agnetae van Vliet, Succulenta 54: 6.
1975. Holotype: van Vliet 8.5 (U - ?).
Notocactus concinnus ssp. multicostatus
(Buining & Brederoo) A. Doweld, stat. nov. -
Notocactus multicostatus Buining & Brederoo in
Krainz, Kakteen 55-56: unpag. [CVIc]. 1974 ('1973'). Holotype: Horst 100
[Horst & Uebelmann 100] (U!).
10. Notocactus tabularis (Cels ex RYmpler) A. Berger, Kakteen:
211, 343. 1929. Neotype (required).
Notocactus tabularis ssp. tabularis
Notocactus tabularis ssp. muricatus (Doweld)
Doweld, comb. nov. -
Peronocactus tabularis ssp. muricatus Doweld,
Sukkulenty 1999(2): 21-22. -
Echinocactus muricatus K. Schum., Gesamtb. Kakt. 386. 1898
non Otto ex Pfeiff. 1837. -
Notocactus bommeljei van Vliet, Succulenta 47: 7. 1968. Holotype:
tab. 50, fig. 2 in Mart. Fl. Brasil. 4(2). 1890.
11. Notocactus rudibuenekeri Abraham, Succulenta 67:
133-134. 1988. Holotype: Abraham 355 (KOELN).
Notocactus rudibuenekeri ssp. rudibuenekeri
Notocactus rudibuenekeri ssp. glomeratus
(Gerloff) A. Doweld, stat. nov. -
Notocactus glomeratus Gerloff, Internoto 12: 10.
1991. Holotype: Stockinger 249 (ZSS).
12. Notocactus scopa (Sprengel) A. Berger, Kakteen: 208, 343.
1929. Neotype (designated here): «Rincon Segredo, Rio Grande do Sul,
Brazil», F. Ritter 1393 (SGO 125297).
Notocactus scopa ssp. scopa
Notocactus scopa ssp. marchesii (Abraham) A.
Doweld, stat. nov. -
Notocactus scopa var. marchesii Abraham, Kakteen
Sukk. 40: 176. 1989. Holotype: Abraham 34 (KOELN).
Notocactus scopa ssp. neobuenekeri (F.
Ritter) A. Doweld, stat. nov. -
Notocactus neobuenekeri F. Ritter, Kakt. SYdamer. 1:
181-182. 1979. Holotype: F. Ritter 1397 (U).
Notocactus scopa ssp. succineus (F. Ritter)
A. Doweld, stat. nov. -
Notocactus succineus ('sucineus') F. Ritter,
Succulenta 49: 109. 1970. Holotype: F. Ritter 1399 (U).
Incertae sedis:
13. Notocactus erubescens (Osten) Marchesi, Bol. Soc.
Argentina Bot. 14: 248. 1972. Lectotype (Marchesi, l.c.): Osten 16884a (MVM).
14. Notocactus memorialis Prestle, Internoto 9: 7. 1988.
Holotype: Schlosser s.n. (U).
Eriocactus Backeb., Cactaceae (Berlin) 1941(2): 37, 76. 1942, nom.
cons. prop. - Eriocephala
Backeb., BlStt. Kakteenforsch. 1938(6): unpag. [11, 21]. 1938, nom. prop. rejic.
non Eriocephalus L., Sp. Pl.: 926. 1753 (Asteraceae).
Chrysocactus Y. Ito, Bull.
Takarazuka Insect. 71: 18. 1950, nom. nud. -
Notocactus subgen. Eriocactus (Backeb.) F. Buxb. in
Krainz, Kakteen 35: sine pag. [CVIc]. 1967. -
Parodia subgen. Eriocactus (Backeb.) F. Brandt,
Kakt. Orch.-Rundsch. 7(4): 58. 1982 («Eriocacteae»). -
Notocactus subgen. Eriocephalus Backeb. in Backeb.
& F. M. Knuth, Kaktus ABC: 68, 256. 1936 ('1935'), nom. nud.
Notocactus subgen. Eriocephala
(Backeb.) Havl'cek, Kaktusz Vilag 18(4): 73. 1989 ('1988'), nom. superfl. et
invalid.
Type: Echinocactus schumannianus Nicolai ( -
Eriocactus schumannianus (Nicolai) Backeb.).
Subgen. 1. Eonotocactus (Doweld) Doweld, Turczaninowia 2000(3):
xx. 2000. - Notocactus
subgen. Eonotocactus Doweld, Sukkulenty 1999(2): 23. 1999.
Notocactus ser. Krainzianae
Havl'cek, Kaktusz Vilag 18(4): 78. 1989 ('1988'), nom. nud. - A subgen.
Eriocacto costis minoribus (11-15), aculeis radialibus plerumque 12, aculeis centralibus
4, seminibus papillatis, ornamentatione cuticularis nullis differt.
Type: Eriocactus magnificus F. Ritter.
Eriocactus magnificus F. Ritter, Succulenta 45: 60. 1966. Holotype: F. Ritter
1270 (U).
Subgen. 2. Eriocactus -
Notocactus sect. Protoseminiformia Havl'cek,
Kaktusz Vilag, 18(4): 78. 1989 ('1988'), nom. nud. - A subgen. Eonotocacto
costis pluribus (15-45) et seminibus ornamentatione cuticularis reticulato-striatibus
differt.
Type: as in genus.
Ser. 1. Pampocactus (Doweld) Doweld, Turczaninowia 2000(3): xx.
2000. - Notocactus
ser. Pampocactus Doweld, Sukkulenty 1999(2): 23. 1999. - A ser. Eriocacto costis
minoribus (15-16), aculeis pluribus (16-20) et ornamentatione cuticularis parum
reticulato-striatibus differt.
Type: Eriocactus warasii F. Ritter.
Eriocactus warasii F. Ritter, Bradea 1(34): 353. 1973. Lectotype (designated
here): «Rio Grande do Sul, Brazil», R. W. Bueneker s.n. [fl. cult. XI
1973] (HB 67286).
Ser. 2. Eriocactus -
Notocactus ser. Cephalioideae (Fric) Havlicek,
Kaktusz Vilag 18(4): 77. 1989 ('1988'), nom. invalid. - Plantae
elongatae, caespitosae, costis numerosis (20 vel multis), aculeis radialibus 4-8(-20) (4
multiplibus!) et centralibus plerumque 3-4, seminibus papillatis, reticulato-striatibus.
Type: as in genus.
Eriocactus schumannianus (Nicolai) Backeb., Cactaceae (Berlin) 1941(2): 37.
1942. Neotype (required).
Eriocactus leninghausii (F. Haage) Backeb. ex Schaff, Beitr. Sukkulentenk.
-Pflege 1942: 38. 1942. Neotype (designated here): «Corvo, Rio
Grande do Sul, Brazil», F. Ritter 1274 (SGO 125065).
Parodia Spegazz., Anales Soc. Ci. Argent. 96: 70. 1923, emend. A. Doweld.
Hickenia Britt. & Rose,
Cactaceae 3: 207. 1922 non Lillo, Physis (Buenos Aires) 4: 422. 1919 [Asclepiadaceae].
- Microspermia A.
V. Fric, Moller's Deutsche Gartn.-Zeit. 45: 43. 1930, nom. illegit.
Type: P. microsperma (F. A. C. Weber) Spegazz. ( -
Echinocactus microspermus F. A. C. Weber).
Parodia chrysacanthion (K. Schum.) Backeb., BlStt.
Kakteenforsch. 1935(3): unpag. [2]. 1935. Neotype (designated here):
«Volcan, Sudwestgebirge, Jujuy, Argentina», F. Ritter 45 (SGO 125383).
Parodia formosa F. Ritter, Succulenta 43: 57. 1964. Holotype: F. Ritter 735
(U).
Parodia microsperma (F. A. C. Weber) Spegazz., Anales Soc. Ci.
Argent. 96: 70. 1923. Neotype (designated here): «Trancas,
Tucumбn, Argentina», F. Ritter 918 (SGO
125402).
Parodia nivosa (A. V. Fric) Backeb., Blatt. Kakteenforsch.
1934(12): unpag. [3]. 1934. -
Microspermia nivosa A. V. Fric, Kaktusar 3(11):
101. 1932 (nom. valid.) Neotype (designated here):
«Chorillos, Salta, Argentina», F. Ritter 933 (SGO 125419).
Parodia penicillata Fechser & van der Steeg, Succulenta 39:
77. 1960. Holotype: [ex plants of van der Steeg, Wageningen, ex hort.] B. K.
Boom 45730 in 19 VI 1968 (sic!) (L!).
Brasilicactus Backeb., Cactaceae (Berlin) 1941(2): 36, 76, 1942,
nom. cons. prop. -
Notocactus subgen. Brasilicactus (Backeb.) F. Buxb. in Krainz, Kakteen, 35:
unpag. [CVIc]. 1967. - Parodia
subgen. Brasilicactus («Brasilicactea») (Backeb.) F. Brandt, Kakt.
Orch.-Rundsch. 7(4): 53, 66. 1982. -
Acanthocephala Backeb., BlStt. Kakteenf. 1938(6): unpag.
[7, 21]. 1938. non Acanthocephalus Karelin et Kirilov, Bull. Soc. Imp. Nat. Moscou
15: 127. 1842 [Asteraceae]. -
Dactylanthocactus Y. Ito, Explanat. Diagr.
Austroechinocact. : 294. 1957, nom. inval. = Sericocactus Y. Ito, l.c. :
223, 293. 1957, nom. inval. = Brasiliparodia F. Ritter, Kakt. SYdamer. 1 :
144. 1979. - Notocactus
subgen. Notobrasilia Havl'cek, Kaktusz Vilag 18(4): 79-80. 1989 ('1988').
Notocactus sect. Notobrasilia
Havl'cek in Gerloff, Neduchal & Stuchl'k, Notokakteen: 130-131. 1995, nom.
inval.
Type: B. graessneri (K. Schum.) Backeb. ( - Echinocactus graessneri K. Schum.).
Ser. 1. Ritterocactoides A. Doweld, ser. nov. - A
ser. Brasilicacto costis minoribus (16-24), aculeis radialibus quoque minoribus (4-20; 4
multiplibus!) differt.
Type: B. alacriportanus (Backeb. & Voll) A. Doweld.
1. Brasilicactus alacriportanus (Backeb. & Voll) A. Doweld,
Sukkulenty 1999(2): 24. 1999. Holotype: Berger s.n. (RB - ?).
2. Brasilicactus brevihamatus (W. Haage ex Backeb.) A. Doweld,
Sukkulenty 1999 (2): 24. 1999. Lectotype (designated here): fig. 1536
in Backeb., Cactaceae 3: 1599. 1959. Epitype (designated here):
«Jaquirana, Rio Grande do Sul, Brazil», F. Ritter 1277 (SGO 124992).
3. Brasilicactus buenekeri (Buining) A. Doweld, Sukkulenty
1999(2): 24. 1999. Lectotype (designated here): fig. 49 in Buining,
Succulenta 41: 99. 1961.
4. Brasilicactus catarinensis A. Doweld, Sukkulenty 1999(2): 24.
1999. Holotype: F. Ritter 1401a (U).
5. Brasilicactus rechensis (Buining) A. Doweld, Sukkulenty
1999(2): 24. 1999. Lectotype (designated here): fig. 2 in Buining,
Kakteen Sukk. 19(2): 24. 1968.
Ser. 2. Brasilicactus - Costae multae 30-60, aculei
radiales pluri (20 vel multi), aculei centrales plerumque 4-6.
Type: as in the genus.
6. Brasilicactus graessneri (K. Schum.) Backeb., Cactaceae
(Berlin) 1941(2): 36. 1942. Neotype (required).
7. Brasilicactus haselbergii (F. Haage ex Rumpler) Backeb. ex
Schaff, Beitr. Sukkulentenk. -Pflege 1942: 38. 1942. Neotype (required).
Ritterocactus Doweld, Sukkulenty 1999(2): 22. 1999.
Notocactus subgen. Neonotocactus
Backeb., Cact. Succ. J. (Los Angeles) 22: 153. 1950. -
Notocactus sect. Campanulatiflorales Havl'cek,
Kaktusz Vilag 18(4): 78. 1989 ('1988'), nom. nud. = Notocactus sect. Interflorales
Havl'cek, l.c.: 78. 1989 ('1988'), nom. nud.
T y p u s: R. mammulosus (Lem.) Doweld (
Echinocactus mammulosus
Lem.).
Sect. 1. Herteria A. Doweld, sect. nov. - Costae
16-24, aculei radiales 8-24, aculei centrales plerumque 4-6.
In honorem cl. Guillermo Gustavo Francisco Herter (1884-1958),
botanici Austro-Americani nominatur.
Type: R. herteri (Werderm.) Doweld.
1. Ritterocactus allosiphon (Marchesi) Doweld, Sukkulenty 1999(2): 22. 1999.
Holotype: Marchesi CHN 649 (MVFA).
2. Ritterocactus buiningii (F. Buxb.) A. Doweld, Sukkulenty
1999(2): 22. 1999. Holotype: Horst & Uebelmann 90 (U - ?).
3. Ritterocactus herteri (Werderm.) A. Doweld, Sukkulenty
1999(2): 22. 1999. Lectotype (designated here): tab. 3 in Werdermann,
Revista Sudamer. Bot. 3(4/6): 143. 1936.
4. Ritterocactus fuscus (F. Ritter) A. Doweld, comb.
nov. - Notocactus
fuscus F. Ritter, Kakt. SYdamer. 1 : 178. 1979. Lectotype (designated
here): fig. 121 in F. Ritter, l.c.
5. Ritterocactus horstii (F. Ritter) A. Doweld, comb.
nov. - Wigginsia
horstii F. Ritter, Kakt. SYdamer. 1: 199. 1979. Holotype: F. Ritter 1455
(U).
Ritterocactus horstii (F. Ritter) A. Doweld f. horstii
Ritterocactus horstii (F. Ritter) A. Doweld f. juvenaliformis
(F. Ritter) A. Doweld, comb. et stat. nov. -
Wigginsia horstii F. Ritter var. juvenaliformis F.
Ritter, Kakt. SYudamer. 1: 200. 1979. Holotype: F. Ritter 1402 (U).
6. Ritterocactus langsdorfii (Lehmann) A. Doweld, comb.
nov. - Cactus
langsdorfii Lehmann, Ind. Sem. Hamburg.: 17. 1827. Lectotype (designated
here): tab. 13 in Lehmann, Nov. Act. Acad. Caes. Leopold.-Carol. 16(1): 316. 1828
(as Cactus langsdorfii).
Ritterocactus langsdorfii (Lehmann) A. Doweld ssp. langsdorfii
Ritterocactus langsdorfii (Lehmann) A. Doweld ssp. multiceps
(Hofacker & Herm) A. Doweld, comb. nov. -
Parodia langsdorfii ssp. multiceps Hofacker &
Herm, Kakteen Sukk. 50: 62-63. 1999. Holotype: Hofacker 169 (ZSS).
Ritterocactus langsdorfii (Lehmann) A. Doweld ssp. pulvinatus
(van Vliet) A. Doweld, comb. et stat. nov. -
Notocactus pulvinatus van Vliet, Succulenta 49: 50. 1970. Holotype:
van Vliet 2 [DV 25] (U - ?).
Sect. 2. Uebelmannia A. Doweld, sect. nov. - A
sect. Ritterocacto costis minoribus (10-16), aculeis radialibus quoque minoribus (6-12),
aculeis centralibus solitariis, seminibus apapillatis differt.
In honorem Cactacearum Austro-Americanae exploratoris cl. Werner
Uebelmann nominatur.
Type: R. uebelmannianus (Buining) Doweld.
7. Ritterocactus crassigibbus (F. Ritter) A. Doweld, Sukkulenty
1999(2): 22. 1999. Holotype: F. Ritter 1394 (U).
8. Ritterocactus meonacanthus (Prestle) A. Doweld, Sukkulenty
1999(2): 22. 1999. Holotype: Prestle 318 (U).
9. Ritterocactus uebelmannianus (Buining) A. Doweld, Sukkulenty
1999(2): 23. 1999. Holotype: Horst & Uebelmann 78 (U - ?).
Ritterocactus uebelmannianus (Buining) A. Doweld ssp. uebelmannianus
Ritterocactus uebelmannianus (Buining) A. Doweld ssp. pleiocephalus
(Gerloff & Koenigs) A. Doweld, Sukkulenty 1999(2): 23. 1999. Holotype: Gerloff 52a
(STU).
Sect. 3. Ritterocactus -
Echinocactus subsect. Mammulosae Lawr. in Loudon,
Gard. Mag. Ser. 3, 6: 316. 1841. - Costae 18-24, aculeis radialibus 12-18, centralibus
plerumque 1-3, semina papillata cum ornamentatione longitudinaliter striatibus.
Type: as in genus.
10. Ritterocactus mammulosus (Lem.) A. Doweld, Sukkulenty
1999(2): 22. 1999. Neotype (designated here): «Strasse S?o
Francisco-Cerro Branco (23 km), Nostl. Sao Francisco, Rio Grande do Sul, Brazil», F.
Ritter 1382 (SGO 125295).
Ritterocactus mammulosus (Lem.) A. Doweld ssp. mammulosus
Ritterocactus mammulosus (Lem.) A. Doweld ssp. submammulosus
(Lem.) A. Doweld, Sukkulenty 1999(2): 22. 1999. Neotype (designated here):
«North of Villa Mercedes, Juan Llerena, San Luis, Argentina», F. Ritter 4 (ZSS
T4288).
Ritterocactus mammulosus (Lem.) A. Doweld ssp. erythracanthus
(Schlosser & Brederoo) A. Doweld, Sukkulenty 1999(2): 22. 1999. Holotype: Schlosser
165 (MVM).
Ritterocactus mammulosus (Lem.) A. Doweld ssp. eugeniae
(van Vliet) A. Doweld, Sukkulenty 1999(2): 22. 1999. Holotype: van Vliet 41.9 (U -
?).
Ritterocactus mammulosus (Lem.) A. Doweld ssp. brasiliensis
(Havl'cek) A. Doweld, Sukkulenty 1999(2): 22. 1999. Holotype: Havl'cek HAV VII/31
(Herb. Fac. Med. Univ. Carol. Plzen - ?).
11. Ritterocactus megalanthus (Schlosser & Brederoo) A.
Doweld, Sukkulenty 1999(2): 22. 1999. Holotype: Schlosser 163 (MVD).
12. Ritterocactus mueller-melchersii (Fric ex Backeb.) A.
Doweld, Sukkulenty 1999(2): 22. 1999. Lectotype (designated here):
fig. 1577 in Backeb., Cactaceae 3: 1647. 1959.
Ritterocactus mueller-melchersii (Fric ex Backeb.) A. Doweld
ssp. mueller-melchersii
Ritterocactus mueller-melchersii (Fric ex Backeb.) A. Doweld ssp. gutierrezii
(Abraham) A. Doweld, Sukkulenty 1999(2): 22. 1999. Holotype: Abraham 253 (KOELN).
Ritterocactus mueller-melchersii (Fric ex Backeb.) A. Doweld
ssp. winkleri (van Vliet) A. Doweld, Sukkulenty 1999(2): 23. 1999. Holotype: van
Vliet 3311 (U - ?).
13. Ritterocactus rutilans (Daeniker & Krainz) A.
Doweld, Sukkulenty 1999(2): 23. 1999. Holotype: «Cerro Largo ad finem Brasiliae,
Uruguay», Mueller-Melchers s.n. (ZSS TP 50-52).
Ritterocactus rutilans (Daeniker & Krainz) A. Doweld
ssp. rutilans
Ritterocactus rutilans (Daeniker & Krainz) A. Doweld
ssp. veenianus (van Vliet) A. Doweld, Sukkulenty 1999(2): 23. 1999. Holotype: van
Vliet 40.7 (U - ?).
14. Ritterocactus permutatus (F. Ritter) A. Doweld, Sukkulenty
1999(2): 23. 1999. Lectotype (designated here): fig. 142 in F.
Ritter, Kakt. SYdamer. 1: 188-189. 1979.
Sect. 4. Pseudoparodia A. Doweld, sect. nov.
Parodia subgen. Obtextosperma
F. Buxb. in Krainz, Kakteen 34: CVIc/2. -
Parodia ser. Calyptospermae F. Brandt, Kakt.-Orch.
Rundsch. 7(4): 52. 1982. = Parodia ser. Intectospermae F. Brandt, l.c. :
52-53. 1982. - Costae 11-21, aculeis radialibus 8-18, centralibus 4-6(-8), semina
papillata, ornamentatio cuticularis longitudinaliter striata.
Type: R. ayopayanus (Cardenas) Doweld.
15. Ritterocactus ayopayanus (C?rdenas) A. Doweld, comb.
nov. - Parodia
ayopayana C?rdenas, Cact. Succ. J. (Los Angeles) 23: 98. 1951. Holotype: M.
C?rdenas 4398 (LIL).
16. Ritterocactus miguillensis (C?rdenas) A. Doweld, comb.
nov. - Parodia
miguillensis C?rdenas, Cact. Succ. J. (Los Angeles) 33: 109-110. 1961. Holotype:
M. C?rdenas 5539 (LIL).
Incertae sedis:
Ritterocactus arnostianus (L'sal & Kolarik) A. Doweld,
Sukkulenty 1999(2): 22. 1999. Holotype: Horst & Uebelmann 338 (herb. Arb. Mus.
Siles. Opava-Novy Dur, Czech Rep. - ?) (ad sect. Ritterocacto?).
Ritterocactus curvispinus (F. Ritter) A. Doweld, Sukkulenty
1999(2): 22. 1999. Neotype (required): no holotype has been found in U, ZSS or SGO
(F. Ritter 1479) (ad sect. Ritterocacto?).
Ritterocactus rauschii (van Vliet) A. Doweld, Sukkulenty
1999(2): 22. 1999. Holotype: van Vliet 34 (U - ?) (ad sect. Herterio?).
Wigginsia D. M. Porter, Taxon 13: 210. 1964.
Malacocarpus Salm-Dyck, Cact.
Hort. Dyck. 1849: 24. 1850 non F. E. L. Fischer & C. A. Meyer, Index Sem. Hort.
Bot. Petropol. 9: 78. 1843 (Zygophyllaceae). -
Echinocactus subgen. Malacocarpus (Salm-Dyck) K.
Schum., Gesamtb. Kakt.: 291, 295. 1898. -
Notocactus subgen. Malacocarpus (Salm-Dyck) F. Buxb.
in Krainz, Kakteen 35: unpag. [CVIc]. 1967. -
Echinocactus sect. Crateranthi Lem., Cact. Gen. Nov.
Sp. Nov. : 87, [92]. 1839. -
Echinocactus sect. Gymnocarpi Salm-Dyck, Cact. Hort.
Dyck. : 17. 1842.
Lectotype (Britt. & Rose, Cactaceae 3: 187. 1922): W.
corynodes (Pfeiff.) D. M. Porter ( -
Echinocactus corynodes Pfeiff.) = W. tephracantha
(Link & Otto) D. M. Porter ( -
Echinocactus tephracanthus Link & Otto).
Sect. 1. Eowigginsia (A. Doweld) A. Doweld, comb. nov.
- Peronocactus
sect. Eowigginsia A. Doweld, Sukkulenty 1999(2): 24. 1999. - A sect. Wigginsio
aculeis radialibus pluribus (10-20, haud minoribus) et aculeis centralibus plerumque 4
differt.
Type: W. nothohorstii A. Doweld ( - Notocactus horstii F. Ritter).
1. Wigginsia nothohorstii A. Doweld, nom. nov.
Notocactus horstii F. Ritter,
Succulenta 45: 3-4. 1966 non Wigginsia horstii F. Ritter, Kakt. Sudamer. 1: 199.
1979. Holotype: F. Ritter 1269 (U).
2. Wigginsia werdermanniana (Herter) A. Doweld, comb.
nov. - Notocactus
werdermannianus Herter, Revista Sudamer. Bot. 7: 75. 1942. Lectotype (designated
here): fig. [unnumbered] in Herter, l.c.
Sect. 2. Wigginsia = Echinocactus sect. Erinacei
Lawr. in Loudon, Gard. Mag. Ser. 3, 6: 317 («Erinaceae»). - Costae 12-18,
aculeis radialibus plerumque 5-10 (semper 5 multiplum!), aculeis centralibus plerumque 1
(raro 2-3).
Type: as in genus.
3. W. erinacea (Haw.) D. M. Porter, Taxon 13: 210. 1964.
Neotype (required).
4. W. arechavaletae (K. Schum. ex Spegazz.) D. M. Porter, Taxon
13: 211. 1964. Neotype (required).
5. W. tephracantha (Link & Otto) D. M. Porter, Taxon 13:
210. 1964. Lectotype (designated here): tab. 14, fig. 2 in Link &
Otto, Verh. Ver. Beford. Gartenb. 3(7): 422. 1828 (as Melocactus tephracanthus).
6. W. turbinata (Arechavaleta) D. M. Porter, Taxon 13: 211.
1964. Lectotype (designated here): tab. 21 in Anales Mus. Nac.
Montevideo, ser. 2, 5: 235. 1905 (as 'Echinocactus (Malacocarus) sellowii
var. turbinata').
Bolivicactus A. Doweld, gen. nov.
Parodia subgen. Protoparodia
F. Buxb. in Krainz, Kakteen 34: unpag. [CVIc/6]. 1966.
Plantae magnae, habitu Parodiae, costis plerumque 13-21, aculeis
radialibus variantibus (plerumque 8-16, raro multi ad 20-30), aculeis centralibus 4-6(-8),
floribus acrotonibus, lanatibus, seminibus papillatis, ornamentatione longitudinaliter
striatibus.
Type: B. maassii (Heese) A. Doweld (
Echinocactus maassii Heese).
1. Bolivicactus aureicentrus (Backeb.) A.
Doweld, comb. nov. -
Parodia aureicentra Backeb., Cactus (Belg.) 4(4): 57. 1934.
Neotype (designated here): «vom Typstandorte bei Cachi, Salta,
Argentina», F. Ritter 916 (U 117675B).
2. Bolivicactus hausteinianus (Rausch) A.
Doweld, comb. nov. -
Parodia hausteiniana Rausch, Kakteen Sukk. 21(3): 45. 1970.
Holotype: Rausch 192 (ZSS).
3. Bolivicactus columnaris (C?rdenas) A.
Doweld, comb. nov. -
Parodia columnaris C?rdenas, Cact. Succ. J. (Los Angeles)
23: 95. 1951. Holotype: M. C?rdenas 4396 (LIL).
4. Bolivicactus comarapanus (C?rdenas) A.
Doweld, comb. nov. -
Parodia comarapana C?rdenas, Rev. Agric. Cochabamba 7(6):
24. 1951. Holotype: M. C?rdenas 4570 (LIL).
5. Bolivicactus commutans (F. Ritter) A.
Doweld, comb. nov. -
Parodia commutans F. Ritter, Succulenta 43: 22. 1964. Holotype:
F. Ritter 729 (U).
6. Bolivicactus maassii (Heese) A. Doweld, comb.
nov. - Parodia
maassii Heese, Gartenflora 56: 410. 1907. Lectotype (designated here):
fig. 50 in Heese, l.c. Epitype (designated here): «Escayachi,
Mendez, Tarija, Bolivia, F. Ritter 46h in 1963 (U 160345B).
7. Bolivicactus mairananus (C?rdenas) A.
Doweld, comb. nov. -
Parodia mairanana C?rdenas, Natl. Cact. Succ. J. 12(4):
84-85. 1957. Holotype: M. C?rdenas 5085 (LIL).
8. Bolivicactus ocampoi (C?rdenas) A.
Doweld, comb. nov. -
Parodia ocampoi C?rdenas, Kakteen Sukk. 6: 101-102. 1955. Holotype:
M. C?rdenas 5044 (LIL).
9. Bolivicactus procerus (F. Ritter) A.
Doweld, comb. nov. -
Parodia procera F. Ritter, Taxon 13: 117. 1964. Holotype:
F. Ritter 742 (U).
10. Bolivicactus ritteri (Buining) A.
Doweld, comb. nov. -
Parodia ritteri Buining, Succulenta 38: 17. 1959. Lectotype
(designated here): «El Puente, Mendez, Tarija, Bolivia», F. Ritter 85
in 1953 (ZSS T4467).
11. Bolivicactus saint-pieanus (Backeb.) A.
Doweld, comb. nov. -
Parodia saint-pieana Backeb., Descr. Cact. Nov. [1]: 31.
1957 ('1956'). Lectotype (designated here): fig. 1550 in Backeb.,
Cactaceae 3: 1611. 1959.
12. Bolivicactus schwebsianus (Werderm.) A.
Doweld, comb. nov. -
Echinocactus schwebsianus Werderm., Monatsschr. Deutsch.
Kakt. Ges. 2: 189. 1930. Lectotype (designated here): fig.
[unnumbered] in Werdermann, l.c.
13. Bolivicactus stuemeri (Werderm.) A.
Doweld, comb. nov. -
Echinocactus stuemeri Werderm., Monatsschr. Deutsch. Kakt.
Ges. 3: 122. 1931. Lectotype (designated here): fig. [unnumbered] in
Werdermann, l.c.
14. Bolivicactus subterraneus (F. Ritter)
A. Doweld, comb. nov. -
Parodia subterranea F. Ritter, Succulenta 43: 43. 1964. Holotype:
F. Ritter 731 (U).
15. Bolivicactus taratensis (C?rdenas) A.
Doweld, comb. nov. -
Parodia taratensis C?rdenas, Cact. Succ. J. (Los Angeles)
36: 24. 1964. Holotype: M. C?rdenas 5552 (LIL; US, iso).
16. Bolivicactus tilcarensis (Werderm. &
Backeb.) A. Doweld, comb. et stat. nov. -
Echinocactus stuemeri ('stu eri') var.
tilcarensis Werderm. & Backeb. in Backeb., Neue Kakt. 89. 1931. Lectotype
(designated here): fig. [unnumbered] in Backeb. l.c. : 59.
17. Bolivicactus tuberculatus (C?rdenas) A.
Doweld, comb. nov. -
Parodia tuberculata C?rdenas, Cact. Succ. J. (Los Angeles)
23: 97. 1951. Holotype: M. C?rdenas 4397 (LIL).
Genera excluded:
Tribus Notocacteae F. Buxb., Madro?o 14: 181. 1958.
Subtribus Copiapoinae A. Doweld, subtrib. nov. - A
subtr. Notocactino corporis globularibus vel cylindricibus, floribus vulgo campanulatibus,
seminibus obovoidibus, papillatis vel apapillatis cum ornamentatione cuticularis nullis
vel granularibus differt.
Type: Copiapoa Britt. & Rose. 4 genera.
Frailea Britt. & Rose, Cactaceae 3: 208. 1922.
Blossfeldia Werderm., Kakteenk. 1937: 162.1937.
Uebelmannia Buining, Succulenta 46: 159. 1967.
Copiapoa Britt. & Rose, Cactaceae 3: 85. 1922.
Tribus Cacteae Reichenb., Fl. Germ. Excurs. 2(2): 561. 1832.
Subtribus Astrophytinae (Horaninow) A. Doweld, Novitates Syst. Pl. Vasc. 32:
115. 2000. - Tribus Astrophyteae
Horaninow, Char. Ess. Fam. : 143. 1847.
Type: Astrophytum Lem. Monotypic.
Astrophytum Lem., Cact. Gen. Nov. Sp. Nov. 3. 1839.
Acknowledgements
The author is indebted to J. Smit-Reesink, Urs Eggli, J. Neduchal, N.
Gerloff, R. Nyffeler, A. Hofacker, D. Metzing, I. Sinev, S. Stuchlik, , W. Verheulpen, M.
Kimnach, J. Zazvorka, V. Zlotin, M. Lowry, and M.Veverka for invaluable support with seed
materials and literature sources used in the study. Special thanks are go to Prof. Werner
Greuter and Dr. Gea Zijlstra for valuable nomenclatural consultations. The assistance of
Gea Zijlstra (for U), J. Fontella Pereira ( for HB), Fernand Jacquemoud (for G), Mйlica
Muсoz-Schick (for SGO), Urs Eggli (for ZSS), and G. Thijsse (for L) is highly
appreciated.
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