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The genus Mammillaria is one of the largest in the cactus family (Cactaceae), with currently 171 known species and varieties recognized.^1] Most of the mammillarias are native to Mexico, but some come from the southwest USA, the Caribbean, Colombia, Venezuela, Guatemala and Honduras.

The first was described by Carolus Linnaeus as Cactus mammillaris in 1753, deriving name from Latin mammilla, "nipple", referring to the tubercles that are one of the plant's specific features. In 1812, the cactus specialist Adrian Haworth described the genus Mammillaria to contain this and related species. Numerous species are commonly known as nipple cactus, fishhook cactus or pincushion though such terms may also be used for related taxa, namely Escobaria

Mammillarias have extremely variable spination from species to species, and attractive flowers, making them specific ally attractive for cactus hobbyists. Most mammillarias plants are considered easy to cultivate, though some species are among the hardest cacti to grow. Several taxa are threatened with extinction at least in the wild, due to habitat destruction and especially overcollecting for the pot plant trade. Cactus fanciers can assist conservation of these rare plants by choosing nursery-bred specimens (wild-collected ones are illegal to possess for the rarest species anyway). Besides helping to preserve rare plants, one can gain experience in growing and breeding cacti in general with nursery-bred rare mammillarias: several mammillarias are quite easy (for cacti) to grow from seeds. One such species, popular and widely available from nursery stock but endangered in the wild, is Mammillaria zeilmanniana.

Description

The distinctive feature of the genus is the specific development of an areole, that is split into two clearly separated parts, one occurring at the tubercle's apex, the other at its base. The apex part is spine bearing, and the base part is always spineless, but usually bearing some bristles or wool. The base part of the areole bears the flowers and fruits, and is a branching point. The apex part of the areole does not carry flowers, but in certain conditions can function as a branching point as well.

The plants are usually small, globose to elongated, the stems from 1 cm to 20 cm in diameter and from 1 cm to 40 cm tall, clearly tuberc ulate, solitary to clumping forming mounds of up to 100 heads and possess radial symmetry. Tubercles can be conical, cylindrical, pyramidal or round. The roots are fibrous, fleshy or tuberous. The flowers are funnel-shaped and range from 7 mm to 40 mm and more in length and in diameter, from white and greenish to yellow, pink and red in color, often with a darker mid-stripe; the reddish hues are due to betalain pigments as usual for Caryophyllales. The fruit is berry-like, club-shaped or elongated, usually red but sometimes white, yellow or green. Some species have the fruit embedded into the plant body. The seeds are black or brown, from 1 to 3 mm in size.

Systematics

The large and diverse genus Mammillaria has seen multiple attempts to subdivide the species in it into smaller groups within the genus or attempts to split it to multiple genera for better understanding of the plants' relationship. Early classifications were performed by Pfeiffer (1837)^{[citation needed]}, Salm-Dick (1845)^{[citation needed]} and Engelmann (1856)^{[citation needed]}. The genus Mammillaria included members of some modern genera like Coryphantha and Ariocarpus at that time. Classifications by Schumann (1898)^{[citation needed]}, Britton and Rose (1923)^{[citation needed]}, Berger (1929)^{[citation needed]} Buxbaum (1951?56)^{[citation needed]} and Moran (1953)^{[citation needed]} followed, splitting the genus in parts and combining it back together again.

Later classification was performed by the cactus specialists Hunt, Reppenhagen and Luthy^{[citation needed]}, with a lot of work focusing on searching the meanings and value of the original plant descriptions, synchronizing them with modern taxonomic requirements and studying the morphology of plants and seeds, as well as ecological aspects of the genus. These works helped to expand the understanding of Mammillaria taxa.

Currently the classification of Mammillaria is in a state where few newly discovered species are likely, though some new species may yet be found when the chaos of names created earlier by commercial plant collectors is sorted out. Many names that were introduced for plants barely differentiated by a shade of flower color or variation in spination were eliminated in attempt to make the use of names consistent with the rest of the botanical world. The number of taxa, which at one time numbered above 500, is now below 200. Some genera (Dolichothele, Mammillopsis, Krainzia and others) have been merged back into Mammillaria, and others like Coryphantha, Escobaria and Mammilloydia were confirmed as separate.

Intense studies of DNA of the genus are being conducted, with preliminary results published for over a hundred taxa, and this promising approach might soon end the arguments. Based on DNA research results, the genus does not seem to be monophyletic and is likely to be split into two large genera, one of them possibly including certain species of other closely related genera like Coryphantha, Ortegocactus and Neolloydia.

Selected species

As noted above, some might not belong into this genus.

shhook cactus or pincushion though such terms may also be used for related taxa, namely Escobaria

Mammillarias have extremely variable spination from species to species, and attractive flowers, making them specifically attractive for cactus hobbyists. Most mammillarias plants are considered easy to cultivate, though some species are among the hardest cacti to grow. Several taxa are threatene d with extinction at least in the wild, due to habitat destruction and especially overcollecting for the pot plant trade. Cactus fanciers can assist conservation of these rare plants by choosing nursery-bred specimens (wild-collected ones are illegal to possess for the rarest species anyway). Besides helping to preserve rare plants, one can gain experience in growing and breeding cacti in general with nursery-bred rare mammillarias: several mammillarias are quite easy (for cacti) to grow from seeds. One such species, popular and widely available from nursery stock but endangered in the wild, is Mammillaria zeilmanniana.

Description

The distinctive feature of the genus is the specific development of an areole, that is split into two clearly separated parts, one occurring at the tubercle's apex, the other at its base. The apex part is spine bearing, and the base part is always spineless, but usually bearing some bristles or wool. The base part of the areole bears the flowers and fruits, and is a branching point. The apex part of the areole does not carry flowers, but in certain conditions can function as a branching point as well.

The plants are usually small, globose to elongated, the stems from 1 cm to 20 cm in diameter and from 1 cm to 40 cm tall, clearly tuberculate, solitary to clumping forming mounds of up to 100 heads and possess radial symmetry. Tubercles can be conical, cylindrical, pyramidal or round. The roots are fibrous, fleshy o r tuberous. The flowers are funnel-shaped and range from 7 mm to 40 mm and more in length and in diameter, from white and greenish to yellow, pink and red in color, often with a darker mid-stripe; the reddish hues are due to betalain pigments as usual for Caryophyllales. The fruit is berry-like, club-shaped or elongated, usually red but sometimes white, yellow or green. Some species have the fruit embedded into the plant body. The seeds are black or brown, from 1 to 3 mm in size.

Systematics

The large and diverse genus Mammillaria has seen multiple attempts to subdivide the species in it into smaller groups within the genus or attempts to split it to multiple genera for better understanding of the plants' relationship. Early classifications were performed by Pfeiffer (1837)^{[citation needed]}, Salm-Dick (1845)^{[citation needed]} and Engelmann (1856)^{[citation needed]}. The genus Mammillaria included members of some modern genera like Coryphantha and Ariocarpus at that time. Classifications by Schumann (1898)^{[citation needed]}, Britton and Rose (1923)^{[citation needed]}, Berger (1929)^{[citation needed]} Buxbaum (1951?56)^{[citation needed]} and Moran (1953)^{[citation needed]} followed, splitting the genus in parts and combining it back together again.

Later classification was performed by the cactus specialists Hunt, Reppenhagen and Luthy^{[citation needed]}, with a lot of work focusing on searching the meanings and value of the original plant descriptions, synchronizing them with modern taxonomic requirements and studying the morphology of plants and seeds, as well as ecological aspects of the genus . These works helped to expand the understanding of Mammillaria taxa.

Currently the classification of Mammillaria is in a state where few newly discovered species are likely, though some new species may yet be found when the chaos of names created earlier by commercial plant collectors is sorted out. Many names that were introduced for plants barely differentiated by a shade of flower color or variation in spination were eliminated in attempt to make the use of names consistent with the rest of the botanical world. The number of taxa, which at one time numbered above 500, is now below 200. Some genera (Dolichothele, Mammillopsis, Krainzia and others) have been merged back into Mammillaria, and others like Coryphantha, Escobaria and Mammilloydia were confirmed as separate.

Intense studies of DNA of the genus are being conducted, with preliminary results published for over a hundred taxa, and this promising approach might soon end the arguments. Based on DNA research results, the genus does not seem to be monophyletic and is likely to be split into two large genera, one of them possibly including certain species of other closely related genera like Coryphantha, Ortegocactus and Neolloydia.

Selected species

As noted above, some might not belong into this genus.

References

• Anderson, Edward F. (1991): The Cactus Family
• Butterworth, Charles A. & Wallace, Robert S. (2004): Phylogenetic studies of Mammillaria (Cactaceae) - insights from chloroplast sequence variation and hypothesis testing using the parametric bootstrap. Am. J. Bot. 91(7): 1086-1098. PDF fulltext Supplementary data

External links

• cactiguide.com is the main source for the species list, and in turn sourced from several books which are listed on that site.
• mammillarias.net is the main up-to-date internet resource, with complete species and varieties description, distribution maps and a large selection of photographs of all Mammillaria species both in nature and cultivated.
• SucculentCity Mammillaria Page: Cultivation Data and Photographs