Chronology

The first Coryphantha sp. reached Europe in the 1820s. Some of the central Mexican species had been collected by that time in the mining region of the Valle de Mexico by Thomas Coulter (1793-1843) and sent to Geneva to Augustin Pyramus De Candolle (1778-1841), one of the best-known botanists of the period. Coulter worked in the mining region of Real del Monte (today Mineral del Monte) until 1825 and from 1827 at Zimapan. The plants for De Candolle were shipped on, or prior to, 1 May 1828 from Veracruz. Type localities given for these plants were usually "Mexico" or sometimes "Real del Monte" or "Zimapan".

By the time De Candolle had received the plants, his important work Prodromus had already been published and he thus described them under the generic name Mammillaria in a lesser known journal in 1828. Unfortunately, De Candolle did not know then which diagnostic criteria for differentiation within the Cactaceae were important and sometimes his descriptions remained so short that it is hard to decide which species Coulter had actually sent him. He also made the mistake of describing several young plants as separate species. Furthermore, none of the Coryphantha sp. described were illustrated and no herbarium material exists.

Consequently, De Candolle's descriptions remain quite dubious, even if well-known species such as octacantha, radians and cornifera are among them and these names have been used for more than a century. Some of the descriptions by De Candolle belonging to the genus Mammillaria had to be rejected due to insufficient documentation or justified doubts as nomina dubia (HUNT 1981).

Due to De Candolle's shaky base, the chaos of future Coryphantha nomenclature was, in a way, preprogrammed.

The first Coryphantha species were all subsumed into the genus Mammillaria (or Mamillaria), which had been proposed by the English expert on succulents Adrian Hardy HAWORTH (1772-1833). He, therefore, separated all cacti with tubercles from all the other globular cacti included at the time in the genus Cactus.

In 1827 Heinrich Friedrich LINK (17671851) and Friedrich Christian OTTO (17831856) proposed Echinocactus as a separate genus of certain globular cacti from the similar genus Melocactus. However, they did not point out any differentiating criteria to the genus Mammillaria and, therefore, some Coryphantha sp. were also described as Echi-nocactus.

Until 1848 all publications concerning the genus Coryphantha were published in Europe, mainly by Charles Lemaire (18011871), Michael Josef Scheidweiler (17991861), Friedrich Scheer (1793-1869), Friedr. Chr. Otto (1783-1856),Albert Dietrich (17951856), Louis Pfeiffer (1805-1878), Fr.Muhlen-pfordt, J. Gerhard Zuccharini (1797-1848), Carl Friedrich Philipp von Martius (17941868) and Willem Henrik De Vriese (18061862). Most of these descriptions were based on living plants in European greenhouses, often lacking any collection data. Few of these plants were conserved and most of these taxa are thus as poorly typified as the rudimentary descriptions of De Candolle. Even at this time, none of the above botanists clearly knew what to do with De Candolle's descriptions, hence, new names were published without reference to the earlier names. Most of these new descriptions appeared in the gardening journals of different countries, and because the exchange of literature was quite difficult at that time, all the information was scattered in a manner which made it very difficult for anyone to obtain an overview.

Important information about the distribution area of several Mexican species was given by Karl August EHRENBERG (1801-1849), who lived and studied the botany of Mineral del Monte from 1831 to 1840 and who sent many plants to Europe. His field activities together with plant location details are contained in an article which appeared in the journal Linnaea in 1846. This paper remained the most reliable source of locations for many of the plants collected before 1840.

In PFEIFFER's publications (1837a), the Coryphantha sp. were still mixed up with Eumamillaria sp. in the series Conothelae and Brachythelae. In 1839 LEMAIRE first published the epithet Aulacothelae as an infra-generic taxon in which he included those "Mammillarias, whose tubercles show a groove on the upper surface". Lemaire also gave a Latin diagnosis and compared the new taxon with Echinocactus stenogoni.

At first, the taxon Aulacothelae was regarded as a subgenus of Mammillaria (G. LAWRENCE 1841).Although Lawrence,in his Catalogue of the cacti in the collection of Rev. Theodore Williams, at Hendon Vicarage, Middlesex did not mention Lemaire's name, the list of names clearly relates to Lemaire.

In July 1846 the famous collection of exotic plants of the Baron de MONVILLE (Hippolyte Boissel) was sold by auction. The plants offered for sale were listed in a catalogue, using essentially Lemaire's system.

Here, for the first time, we meet Aulacothelae as a separate genus with the spelling Aula-cothele. Monville did not mention Lemaire's name either, but his list also clearly relates to that of Lemaire.

In 1850 the work of Prince Josef von SALM-DYCK (1773-1861) was published in Europe, entitled Cacteae in horto Dyckensi cultae anno 1849. For the first time, a scientifically based difference between the genera Mammillaria and Echinocactus was established: in Mammillaria, which he allocated to the tribe Melocacteae, the fruit is at first within the plant body and only protrudes after maturity. The fruit is longish, smooth and crowned by dried perianth segments. In the genus Echinocactus, which he allocated to the tribe Echinocacteae, the fruit protrudes from the beginning and the dried perianth segments are discarded. Since in some of those species, which, because their bodies were covered by tubercles, were counted among Mammillaria, and the fruit protrudes from the beginning as in Echinocactus, the Prince of Salm-Dyck took LEMAIRE's (1839) Aulacothelae and separated it as a subgroup from Mammillaria. Certain members of the subgroup were then removed and included within the subgenus Glanduliferae. The remainder were left within Aulacothelae. By doing this, however, he made the mistake of listing Mammillaria aulacothele Lemaire, the type species of Lemaire's Aulacothelae, under Glanduliferae, thus making this name invalid (ICBN Art. 52.1).

In 1853, Hermann POSELGER (1818-1883) put Glanduliferae Salm-Dyck and Aulacothelae Lemaire into the genus Echinocactus. He did this because the flowers of the reallocated plants are mostly solitary and arise from the top of the plant or at least from younger parts of the top as in Echinocacti, as opposed to Mammillarias in which the flowers protrude between the tubercles from axils of the year before, or older axils and are mostly arranged in a circle around the top.

From 1848 onwards, publications from George Engelmann (1804-1884) concerning the cacti of the United States followed. Up to this time, all Coryphantha sp. were allocated to Mammillaria, the generic name being Aulacothele (Lemaire) Monville. Although valid, this was not generally recognised and was not referred to after 1846. Engelmann was faced with the nomenclatural chaos described above. In 1856 he suggested the name Coryphantha (Coryphantha [Greek] = flowering from the top) as a subgenus of Mammillaria for a group of about 16 species. Incidentally, three of them (papyracantha, pottsii and conoidea) had to be excluded again later. The remainder were from the southern United States and the adjacent regions of Mexico. All other Coryphantha species described at that time had originated from Mexico. They were not mentioned by ENGELMANN, which is probably why he did not discuss the Glanduliferae of Salm-Dyck, which, of course, should also have been included within Coryphantha. Even so, Engel-mann's publication is superior to any written about Coryphantha at that time. Once again, however, herbarium material was not preserved.

In 1868 the group of plants named Coryphantha was raised to generic level by Charles LEMAIRE. In his Iconographie descriptive des Cactées (1853), the generic name Aulacothelae can still be found. He proposed Mammillaria sulcolanata as the type species of this genus ("Typumque sat completum praebet generis Aulacothelis, jam ab auctoribus propositum"). In 1868 he finally adopted the name Coryphantha from Engelmann, but as a genus and no longer as a subgenus of Mam-millaria. However, the species listed by the two authors only partly overlapped. Engel-mann's list mainly contained species from the United States or the bordering regions of Mexico, whereas Lemaire in his publications mentions mainly Mexican species. In all, Lemaire mentioned 25 names of species (24

species and 1 variety). This list was valid for a long period; 27 years later only one additional species was listed in the Index Kewensis. Today, however, only eight of Lemaire's Coryphantha sp. are still in use. All the others are now either synonyms or nomina dubia.

As the lectotype of the genus Coryphantha, Lemaire chose Coryphantha sulcolanata and this, as we will see later, remained valid until 1976.

Before Mammillaria was generally accepted as nomen conservandum, John Merle COULTER (1851-1928) in 1894 included all known Coryphantha species within the genus Cactus as a parallel section to "Eumammil-laria". Lemaire's work was probably unknown to Coulter, because in his list of synonyms not a single Coryphantha name of Lemaire appears.

In 1898 Karl Moritz SCHUMANN's (18511904) Gesamtbeschreibung der Kakteen appeared. This is a detailed monograph of the Cactaceae, in which 22 species of Cory-phantha appeared, considered, however, as a subgenus of Mammillaria and divided between the series Aulacothelae Lemaire and Glanduliferae Salm-Dyck. It was obvious that by this time Lemaire's work was not highly esteemed. Schumann's work contains detailed descriptions, Latin diagnoses and synonyms, but confuses readers by using many catalogue names as first publications, and cultivators of the plants, who had given them any name, as the original species names. Schumann's plants had been preserved in herbaria. However, few specimens survived the World War II (among them not one Coryphantha) and, therefore, many of the new names introduced by Schumann must now be regarded as not identifiable nomina dubia.

Post-1900 more first descriptions of Coryphantha followed. These mostly originated from German authors and appeared in German journals. To name a few: Leopold Quehl (1849-1922), Walter Mundt (18531927), Jos. Anton Purpus (1860-1933), Fried-

rich Bodeker (1867-1937) and Friedrich Vau-pel (1876-1927).

In 1919-1923, an important new work appeared, which became an important basis for Coryphantha specialists until today: The Cactaceae by Nathaniel BRITTON (18581934) and Joseph Nelson ROSE (1862-1928). It is the first complete monograph on cactus existing in English. Thanks to their fieldwork and the intensive use of herbaria located worldwide, the fourth volume gives much greater insight into the genus Coryphantha than Schumann's Gesamtbeschreibung der Kakteen. It is generally known that it was J.N. Rose who studied the small globular cacti of northern America, although both authors, Britton and Rose, are named.

Britton and Rose studied all the existing cactus literature diligently and listed a great number of the earlier Coryphantha publications in their work. As "splitters" they did not hesitate to follow Lemaire in accepting Coryphantha as a genus. From the former subgenus of Engelmann, they separated three more genera, Toumeya, Neobesseya und Escobaria.

In total, Britton and Rose described four new Coryphantha species and formally added 16 more to the genus Coryphantha, thereby listing 50 species under Coryphantha, this number includes four Neobesseyas und eight Escobarias.

Most of the type plants of Britton und Rose are correctly named and associated with herbarium material, but it is a pity that most of Rose's descriptions are far too short in general and sometimes superficial or inexact. This led to further confusion later on. They also misapplied several names (e.g., C. pyc-nacantha, C. salmdyckiana, C. sulcolanata, C. difficilis etc.).

After Britton and Rose, the next important publication was that of Alwin BERGER (1871-1931). In his Monographie der Kakteen (1929) he gave the infrageneric range within the genus Coryphantha to three former gen era of Britton and Rose (Escobaria, Neo-besseya and Neolloydia). His work lists over 50 species. Among them are many new descriptions of Friedrich Bodeker, who, in the period 1920-1930, described many new cactus species, including some Coryphantha sp. In his Mammillarien-Vergleichs-Schlussel, BODEKER (1933) mentioned 64 Coryphantha species, and he also treated Neobesseya and Escobaria as separate genera.

After the World War II, the Austrian Franz Buxbaum (1900-1979) published several important scientific works on cacti.As a starting point, he studied Alwin BERGER's Die Entwicklungslinien der Kakteen of 1926 where Berger set out to clarify the supposed phylogenetic interrelationship between all the cactus genera acknowledged by Britton and Rose. He did so by using dendrograms.

Buxbaum emphasised the polarisation of characteristics (primitive versus highly derived) and contributed greatly to the analysis of the homology of different taxonomi-cally important features of the Cactaceae. He reclassified several plant groups including Coryphantha and Mammillaria at the levels of genus/subgenus. He was the first author who transferred the species group Coryphantha vivipara from Coryphantha sensu stricto to Escobaria.

During the same time period, Curt BACKEBERG (1894-1966) worked on his detailed monograph Die Cactaceae. In the fifth volume, published in 1961, he devoted 131 pages to the genus Coryphantha and its relatives. He considered all publications since Britton and Rose, including seven original descriptions of Coryphantha sp.. These had been published in lesser known gardening journals and were introduced to a larger audience by their incorporation into the Cactaceae.

Among all the great cactus monographers, Backeberg was unique in one aspect in that he completely neglected to make herbarium sheets. He did not conserve one single plant; all his descriptions and taxonomic decisions were based on living plants, which unfortunately no longer exist.

Since most of Backeberg's first descriptions occurred before 1 January 1958, the day when valid nomenclatural rules were introduced, necessitating the indication of a type to achieve a valid description, they are still valid nowadays, but his descriptions after 1 January 1958 are, due to the lack of typification, invalid.

However, Backeberg's work provides a valuable overview of the genus Coryphantha and includes all the important publications of the nineteenth and twentieth centuries.

In the years after 1970 Lew Bremer wrote many publications, including 15 first descriptions of Coryphantha sp. At present, only three are regarded as good species and two as subspecies, all the others being considered as synonyms only. Many of his species are new descriptions of long-known taxa, but with new names, others are also incorrect applications of names to long-recognised species. It is true that Bremer published excellent descriptions, very detailed and usually photographically well documented. However, on the whole, his work has caused more confusion to taxonomy than contributing to science.

In 1982 a publication appeared, which is regarded as a standard work by the American government and also by most American botanists, namely Lyman BENSON's The Cacti of the United States and Canada. Benson accepts 33 species of Coryphantha in the USA, although today only 15 can be regarded as good species. In his work some names are incorrectly applied, and some good species are not mentioned.

In 1940 BENSON had retraced the US Coryphantha species back to Mammillaria. This was at a time when the elevation of Coryphantha to genus level by BRITTON & ROSE (1923) was being accepted worldwide. Up to this time, the criteria separating Cory-phantha from Mammillaria contained two characteristics only, namely the flower position and the areole morphology.

In 1969 BENSON came back to the concept of Coryphantha being a natural plant group which had to be separated from Mam-millaria. Therefore, he raised it to the level of genus and at the same time decided to include all Escobaria species within Coryphantha.

Over the same period, Del WENIGER (1970) published a book in which he applied Benson's old classification, thus lumping all Coryphantha sp. into Mammillaria. This classification was followed in Texas only.

It has to be said that the taxonomic work of both Bremer and Benson outlined above caused a lot of confusion among botanists concerned with the classification of the Coryphantha species. On the one hand, taxonomy was influenced by Bremer's "splitting" of the species endemic to Mexico, on the other hand, by Benson's "lumping" of the species from USA.

The next large-scale publication is that of Helia BRAVO and Sanchez MEJORADA entitled Las Cactaceas de Mexico (1991). Bremer's "splitting" and Benson's "lumping" are presented with equal weight in a non-critical manner in the hope of achieving a compromise. Unfortunately, the outcome is that no progress has been made with classification matters relating to the genus. BREMER's misapplication of names naturally confused matters further.

Allan Dale ZIMMERMAN's dissertation of 1985 entitled Systematics of the Genus Coryphantha (Cactaceae), which was never validly published, proved to be an extremely valuable work on the genus Coryphantha. In this work, many of the systematic errors and parallelisms are highlighted and corrected for the first time. Zimmerman evolved an amplified theory of areole development with the differentiation of different areole-types. He recognises 46 species of Coryphantha.

The latest complete publication on cacti by E. ANDERSON (2001) widely follows the work of BRAVO (1991) concerning Coryphantha.

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