Allan Dale Zimmerman
Systematics of the genus Coryphantha (Cactaceae), The University of Texas at Austin, 1985
(UMI Dissertation Information Service)
A dissertation of A. D. Zimmerman with a provisional division of the Coryphanta genus including the Escobaria subgenus (Britton et Rose) A. Berger and the Protomammillaria subgenus (John et Říha) A. Zimmerman. The respective dissertation targets in detail the Pseudocoryphantha section (Buxbaum) Moran.
Genus Coryphantha
Subgenus
|
Escobaria |
Coryphantha (Engelmann) Lemaire subgenus Escobaria (Britton et Rose) A. Berger Section Pseudocoryphantha (Buxbaum) Moran, 1953 |
|
| Diagnosis: The section Pseudocoryphantha is a monophyletic group of taxa, sharing a unique synapomorphy: presence of unusually large lenticular druses (up to nearly 1 mm wide) in the pith and cortex of every taxon in the group, a feature absent from the rest of the tribe Cacteae. | ||
|
Species and varieties: |
||
| 1. Coryphantha alversonii (J. M. Coulter) Orcutt, 1926 | ||
| Synonym: | Cactus radiosus [var.] alversonii J. M. Coulter, 1894 | |
| Mammillaria alversonii (J. M. Coulter) Coulter ex Zeissold, 1895 | ||
| Mammillaria radiosa [var.] alversonii (J. M. Coulter) K. Schumann, 1898 | ||
| Mammillaria radiosa f. alversonii (J. M. Coulter) Schelle, 1907 | ||
| Mammillaria arizonica var. alversonii (J. M. Coulter) Coulter ex Davidson et Moxley, 1923 | ||
| Mammillaria vivipara var. alversonii (J. M. Coulter) L. Benson, 1950 | ||
| Coryphantha vivipara var. alversonii (J. M. Coulter) L. Benson, 1969 | ||
| Escobaria vivipara var. alversonii (J. M. Coulter) D. R. Hunt, 1978 | ||
| 2. Coryphantha chlorantha (Engelmann) Briton et Rose, 1923 | ||
| Synonym: | Mammillaria chlorantha Engelmann, 1878 | |
| Cactus radiosus [var.] chloranthus (Engelm.) J. M. Coulter, 1894 | ||
| Mammillaria radiosa var. chlorantha (Engelm.) Engelm. ex K. Schumann, 1898 | ||
| Mammillaria vivipara var. chlorantha (Engelm.) L. Benson, 1950 | ||
| Escobaria chlorantha (Engelm.) F. Buxbaum, 1951 | ||
| [Coryphantha deserti var. chlorantha (Engelm.) Backeberg ex P. C. Fischer, 1971] | ||
| Mammillaria deserti Engelm., 1880 | ||
| Cactus radiosus [var.] deserti (Engelm.) J. M. Coulter, 1894 | ||
| Mammillaria radiosa var. deserti (Engelm.) K. Schumann, 1898 | ||
| Mammillaria radiosa f. deserti (Engelm.) Schelle, 1907 | ||
| Coryphantha deserti (Engelm.) Britton et Rose, 1923 | ||
| Mammillaria arizonica var. deserti (Engelm.) Engelm. ex Davidson et Moxley, 1923 | ||
| Mammillaria vivipara var. deserti (Engelm.) L. Benson, 1944 | ||
| Coryphantha vivipara var. deserti (Engelm.) W. T. Marshall, 1950 | ||
| Escobaria deserti (Engelm.) F. Buxbaum, 1951 | ||
| Coryphantha chlorantha var. deserti (Engelm.) Backeberg, 1961 | ||
| ? Mammillaria utahensis Hildmann ex K. Schumann, 1898 | ||
| Coryphantha vivipara var. buoflama P. C. Fischer, 1980 | ||
| Escobaria vivipara var. buoflama (P. C. Fischer) N. P. Taylor, 1983 | ||
| 3a. Coryphantha vivipara var. vivipara (Nutt.) Britton et Rose, 1923 | ||
| Synonym: | Cactus viviparus Nutt., 1813 | |
| Mammillaria vivipara (Nutt.) Haw., 1819 | ||
| Echinocactus viviparus (Nutt.) Poselger, 1853 | ||
| Mammillaria vivipara var. vera Engelm., 1857 | ||
| Mammillaria radiosa f. vivipara (Nutt.) Schelle, 1907 | ||
| Neomammillaria vivipara (Nutt.) Britton et Rose ex Rydb., 1932 | ||
| Escobaria vivipara (Nutt.) F. Buxbaum, 1951 | ||
| [Coryphantha vivipara (Nutt.) Engelm. ex Backeberg, 1961] | ||
| ? Mammillaria montana Blanc, 1891 | ||
| ? Mammillaria hirschtiana Haage f. ex Hirscht, 1896 | ||
| Coryphantha columnaris Lahman, 1934 | ||
| Coryphantha oklahomensis Lahman, 1949 | ||
| Escobaria oklahomensis (Lahman) F. Buxbaum, 1951 | ||
| 3b. Coryphantha vivipara var. radiosa (Engelm.) Backeberg, 1961 | ||
| Synonym: | Mammillaria radiosa Engelm., 1850 | |
| Echinocactus radiosus (Engelm.) Poselger, 1853 | ||
| Mammillaria vivipara var. radiosa (Engelm.) Engelm., 1857 (preprint, 1856) | ||
| Mammillaria radiosa var. radiosa subvar. texana Engelm., 1857 (preprint, 1856) | ||
| Mammillaria vivipara subsp. radiosa (Engelm.) Engelm., 1859 | ||
| Mammillaria vivipara subsp. radiosa var. texana (Engelm.) Engelm., 1859 | ||
| Mammillaria radiosa var. texana (Engelm.) Engelm., 1859 | ||
| Cactus radiosus (Engelm.) Coulter, 1894 | ||
| Mammillaria radiosa f. texensis (Engelm.) Schelle, 1907 | ||
| Coryphantha radiosa (Engelm.) Rydb., 1917 | ||
| Neomammillaria radiosa (Engelm.) Britton et Rose ex Rydb., 1932 | ||
| Escobaria radiosa (Engelm.) Frank, 1960 | ||
| Escobaria vivipara var. radiosa (Engelm.) D. R. Hunt, 1978 | ||
| 3c. Coryphantha vivipara var. neomexicana (Engelmann) Backeberg, 1961 | ||
| Synonym: | Mammillaria vivipara var. radiosa subvar. neomexicana Engelm., 1857 (preprint, 1856) | |
| Mammillaria vivipara [var.] neomexicana (Engelm.) Engelm., 1857 | ||
| Mammillaria vivipara subsp. radiosa var. neomexicana (Engelm.) Engelm., 1859 | ||
| Cactus radiosus [var.] neomexicana (Engelm.) J. M. Coulter, 1894 | ||
| Cactus neomexicanus (Engelm.) Small, 1903 | ||
| Mammillaria radiosa f. neomexicana (Engelm.) Schelle, 1907 | ||
| Mammillaria neomexicana (Engelm.) A. Nels, 1909 | ||
| Coryphantha neomexicana (Engelm.) Britton et Rose, 1923 | ||
| Coryphantha radiosa var. neomexicana (Engelm.) Schelle, 1926 | ||
| Escobaria neomexicana (Engelm.) F. Buxbaum, 1951 | ||
| Escobaria vivipara var. neomexicana (Engelm.) F. Buxbaum ex.? Krainz, 1951 | ||
| Mammillaria vivipara var. radiosa subvar. borealis Engelm., 1857 (preprint, 1856) | ||
| Mammillaria vivipara subsp. radiosa var. borealis (Engelm.) Engelm., 1859 | ||
| Mammillaria radiosa var. borealis (Engelm.) Engelm., 1859 | ||
| Mammillaria borealis (Engelm.) Engelm. ex Britton et Rose, 1923 | ||
| "Mammillaria vivipara var. borealis (Engelm.) Engelm. [sic]" Weniger, 1970 | ||
| Coryphantha fragrans Hester, 1941 | ||
| "Mammillaria fragans (Hester)" Weniger, 1970 | ||
| 3d. Coryphantha vivipara var. arizonica (Engelmann) W. T. Marshall, 1950 | ||
| Synonym: | Mammillaria arizonica Engelmann, 1876 | |
| Cactus radiosus [var.] arizonicus (Engelm.) J. M. Coulter, 1894 | ||
| Mammillaria radiosa var. arizonica (Engelm.) Engelm. ex K. Schumann, 1898 | ||
| Mammillaria radiosa f. arizonica (Engelm.) Schelle, 1907 | ||
| Coryphantha arizonica (Engelm.) Britton et Rose, 1923 | ||
| Mammillaria vivipara var. arizonica (Engelm.) L. Benson, 1944 | ||
| Escobaria arizonica (Engelm.) F. Buxbaum, 1951 | ||
| Escobaria vivipara var. arizonica (Engelm.) D. R. Hunt, 1978 | ||
| Coryphantha marstonii E. U. Clover, 1938 | ||
| Coryphantha missouriensis var. marstonii (E. U. Clover) L. Benson, 1969; pro parte, as to nomenclature | ||
| Escobaria missouriensis var. marstonii (E. U. Clover) D. R. Hunt, 1978; pro parte, as to nomenclature | ||
| 3e. Coryphantha vivipara var. bisbeeana (Orcutt) L. Benson, 1969 | ||
| Synonym: | Coryphantha bisbeeana Orcutt, 1926 | |
| Escobaria bisbeeana (Orcutt) Borg, 1937 | ||
| [Escobaria bisbeeana (Orcutt) W. T. Marshall, 1941; cited by numerous subsequent authors] | ||
| Mammillaria bisbeeana (Orcutt) Orcutt ex Backeberg, 1961 | ||
| Escobaria vivipara var. bisbeeana (Orcutt) D. R. Hunt, 1926 | ||
| Coryphantha vivipara var. bisbeeana f. sonorensis P. C. Fischer, 1980 | ||
| Mammillaria aggregata auct. non Engelm., 1923 | ||
| Coryphantha aggregata (Engelm.) Britton et Rose, 1923; pro parte, as to the populations treated but not as to nomenclature | ||
| Mammillaria vivipara var. aggregata (Engelm.) L. Benson, 1944; pro parte | ||
| Coryphantha vivipara var. aggregata (Engelm.) W. T. Marshall, 1950; pro parte | ||
| Escobaria aggregata (Engelm.) F. Buxbaum, 1950; pro parte | ||
| ? Mammillaria vivipara var. deserti Engelm. ex Weniger, 1984; nomen nudum | ||
| 3f. Coryphantha vivipara var. kaibabensis P. C. Fischer, 1979 | ||
| Synonym: | Escobaria vivipara var. kaibabensis (P. C. Fischer) N.P.Taylor, 1983 | |
| 3g. Coryphantha vivipara var. rosea (Clokey) L. Benson, 1969 | ||
| Synonym: | Coryphantha rosea Clokey, 1943 | |
| Escobaria vivipara var. rosea (Clokey) D. R. Hunt, 1978 | ||
| Coryphantha alversonii var. exaltissima Wiegand et Backeberg, 1961 | ||
| 4a. Coryphantha sneedii var. sneedii (Britton et Rose) A. Berger, 1929 | ||
| Synonym: | Escobaria sneedii Britton et Rose, 1923 | |
| [Coryphantha sneedii (Britton et Rose) Fosberg, 1931] | ||
| Mammillaria sneedii (Britton et Rose) Cory, 1936 | ||
| Coryphantha pygmaea Frič, 1924 | ||
| 4b. Coryphantha sneedii var. leei (Rose ex Boedeker) L. Benson, 1969 | ||
| Synonym: | Escobaria leei Rose ex Boedeker, 1933 | |
| "Mammillaria leei" (Boedeker) Weniger, 1970 ; invalid | ||
| Escobaria sneedii var. leei (Boedeker) D. R. Hunt, 1978 | ||
| 4c. Coryphantha sneedii var. nov. #1 A. Zimmerman, 1985, ined. | ||
| Comments: |
Mexico: Chihuahua, ca. 21 mi. w. of Ojinaga, 0.4 mi. downstream from the highway on a side road, near
e. bajada of the Sierra Peguis, 850 m elev., on very rocky limestone aliuvium dissected by steep-walled
arroyos. Rare, on ridgetops and small hills, with Echinomastus warnockii and Coryphantha echinus.
A. Zimmerman 1536, with T. L. Burgess and T. R. VanDevender, 12 August 1978; flowers and fruits obtained
later in cultivation. Stem, flowers, and fruits pressed but still unaccessioned as of June 1985.
|
|
| 4d. Coryphantha sneedii var. orcuttii (Boedeker) A. Zimmerman, 1985, comb. nov. ined. | ||
| Synonym: |
Escobaria orcuttii Rose, unpublished manuscript name, ca. 1925 (Non Neolloydia orcuttii Rose, unpublished manuscript name for Neolloydia conoidea var. stuetzlei; non Neolloydia orcuttii Rose ex Fric, Moellers Deutsch. Gaertner-Zeitung 142. 1926, pro syn., nom. nud., and not accepted by its author, also with reference to N. c. stuetzlei.) |
|
| ? Escobaria (or Neolloydia?) orcuttii Rose ex Orcutt, 1926; nomen nudum | ||
| Escobaria orcuttii Hort. of 1925 ex Boedeker, 1933 | ||
| Coryphantha strobiliformis var. orcuttii (Boedeker) L. Benson, 1969 | ||
| Coryphantha orcuttii (Boedeker) D. Zimmerman, 1972 | ||
| Escobaria arizonica Hester, 1941, nomen nudum | ||
| Escobaria orcuttii var. macraxina Castetter, Pierce et Schwerin, 1975 | ||
| Escobaria orcuttii var. koenigii Castetter, Pierce et Schwerin, 1975 | ||
| 4e. Coryphantha sneedii var. albicolumnaria (Hester) A. Zimmerman, 1985, comb. nov. ined. | ||
| Synonym: | Escobaria albicolumnaria Hester, 1941 | |
| Escobesseya albicolumnaria (as albocolumnaria) Hester ex L. Benson, 1969; nom. nud., pro syn. | ||
| "Mammillaria albicolumnaria (Hester)" Weniger, 1970, comb. nud. | ||
| Coryphantha albicolumneria (Hester) D. Zimmerman, 1972 | ||
| Escobaria tuberculosa var. durispina auct. non Quehl: Backeberg, 1961 | ||
| Coryphantha strobiliformis var. durispina auct. non Quehl: L. Benson, 1969 | ||
| Escobaria strobiliformis var. durispina auct. non Quehl: Bravo, 1982 | ||
| Escobaria durispina Hort., nomen nudum, probably dating from Horst Kunzler's seed catalogs | ||
| 4f. Coryphantha sneedii var. nov. #2 A. Zimmerman, 1985, ined. | ||
| Comments: |
Mexico: Chihuahua: Sierra Mojina, along Highway 10, 5.9 road miles (9.5 km) east of Ricardo Flores Magon
(measured from the dry Rio Carmen [= Santa Clara], in town). About half-way up a northeast-facing limestone
slope. John B. Iverson, s.n., prior to May 1978. Flower and fruits preserved later, in cultivation; whole
specimen pressed but unaccessioned as of June 1985.
|
|
| 4g. Coryphantha sneedii var. organensis (D. Zimmerman) A. Zimmerman, 1985, comb. nov. ined. | ||
| Synonym: |
Escobaria organensis Hort., nomen nudum, in seed catalogs dating from the late 1960's,
probably from a provisional epithet assigned by Castetter
|
|
| Coryphantha organensis D. Zimmerman, 1972 | ||
| Escobaria organensis (D. Zimmerman) Castetter, Pierce et Schwerin, 1975 | ||
| 4h. Coryphantha sneedii var. villardii (Castetter, Pierce et Schwerin) A. Zimmerman, 1985, comb. nov. ined. | ||
| Synonym: | Escobaria villardii Hort., Castetter, Pierce et Schwerin, 1975 | |
| 4i. Coryphantha sneedii var. sandbergii (Castetter, Pierce et Schwerin) A. Zimmerman, 1985, comb. nov. ined. | ||
| Synonym: |
Escobaria sandbergii Hort., Castetter, Pierce et Schwerin, 1975
|
|
| Others sections of the subgenus Escobaria. | ||
|
Species and varieties: |
||
| 5. Coryphantha hesteri Y. Wright | ||
| 6a. Coryphantha tuberculosa var. tuberculosa (Engelmann) Berger | ||
| 6b. Coryphantha tuberculosa var. varicolor (Tiegel) A. Zimmerman, 1985, comb. nov. ined. | ||
| 6c. Coryphantha tuberculosa var. nov. A. Zimmerman, 1985, ined. (se. Chihuahua) | ||
| 7a. Coryphantha pottsiana var. pottsiana (Poselger) A. Zimmerman, 1985, comb. nov. ined. | ||
| Comments: |
including, as synonyms Escobaria runyonii Britton et Rose Escobaria bella Britton et Rose Coryphantha robertii Berger Coryphantha eskoetteriana (Quehl) Berger Coryphantha muehlbaueriana Boedeker |
|
| 7b. Coryphantha pottsiana var. nov. A. Zimmerman, 1985, ined. Nuevo León (Bustamante) | ||
| 8. Coryphantha sp. nov. A. Zimmerman, 1985, ined. (Sierra de la Madera, Coah.) | ||
| 9. Coryphantha laredoi Glass et R. A. Foster | ||
| 10. Coryphantha chaffeyi (Britton et Rose) | ||
| 11. Coryphantha dasyacantha (Engelmann) Orcutt | ||
| 12. Coryphantha duncanii (Hester) L. Benson | ||
| 13. Coryphantha sp. nov. A. Zimmerman, 1985, ined. (n. of Marte, Coahuila) | ||
| 14. Coryphantha lloydii (Britton et Rose) Fosberg | ||
| 15. Coryphantha zilziana Boedeker | ||
| 16. Coryphantha robbinsorum (Earle) A. Zimmerman | ||
| 17a. Coryphantha missouriensis var. missouriensis (Sweet) Britton et Rose | ||
| 17b. Coryphantha missouriensis var. similis (Engelmann) A. Zimmerman 1985, comb. nov. ined. | ||
| 17c. Coryphantha missouriensis var. asperispina (Boedeker) A. Zimmerman 1985, comb. nov. ined. | ||
| 18. Coryphantha cubensis Britton et Rose | ||
| 19. Coryphantha minima Baird | ||
|
Protomammillaria |
Coryphantha (Engelmann) Lemaire subgenus Protomammillaria (John et Říha) A. Zimmerman, 1985, comb. nov. ined. | |
| basionym: | Escobaria Britton et Rose, 1923 | |
| subg. Protomammillaria John et Říha, 1981 | ||
|
Species: |
||
| 20. Coryphantha strobiliformis (Poselger) Moran | ||
| Comments: | [= Coryphantha chihuahuensis (Britton et Rose) Berger, ] | |
| 21. Coryphantha henricksonii (Glass et Foster) Glass et Foster | ||
|
Subgenus novum |
Coryphantha (Engelmann) Lemaire subgenus novum A. Zimmerman, 1985, comb. nov. ined. | |
|
Species: |
||
| 22. Coryphantha gracilis Bremer et Lau | ||
|
Coryphantha |
Coryphantha (Engelmann) Lemaire subgenus Coryphantha | |
|
Species and varieties: |
||
|
All other taxa of the Coryphanta genus are included into this subgenus (51 taxa, 45 species). See "Allan Dale Zimmerman, Systematics of the genus Coryphantha (Cactaceae), The University of Texas at Austin, 1985 (UMI Dissertation Information Service)". |
||
Genus Ortegocactus
| Comments: Ortegocactus is not a Neobesseya. Contrary to John and Riha, there is no evidence that it is an Escobaria of any kind. Its large yellow flowers and general vegetative appearance suggest Coryphantha sensu stricto or Mammillaria subgenus Dolichothele, but it has no putative relatives within those groups, either. Ortegocactus resultantly is left as a valid monotypic genus. It is a restricted endemic in Oaxaca. | |
|
Species: |
|
| Ortegocactus macdougallii Alexander, 1961 | |
| Synonym: | Neobesseya macdougallii (Alexander) Kladiwa, 1974 |
| Escobaria macdougallii (Alexander) John et Říha, 1981 | |
Generic Rank Acharagma
| Comments: Taxa of Acharagma are excluded from Coryphantha. Acharagma is newly raised to generic rank. Foveolate seeds and fimbriate sepaloid bracts, formerly viewed as synapomorphies linking Escobaria to primitive taxa of Mammillaria rather than to Coryphantha sensu stricto, are re-interpreted as plesiomorphies. | |
|
Species: |
|
| Acharagma roseana | |
| Acharagma aguirreana |
Last modified February 10, 2008