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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, 1999), 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, 1999: 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, 2000, 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, Tucumn, 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), Mlica Muoz-Schick (for SGO), Urs Eggli (for ZSS), and G. Thijsse (for L) is highly appreciated.

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