IB Section Robustispina Dicht A Luthy

Cact. Syst. Init. 11:9,2001. Type: Coryphantha robustispina (Schott ex Engelmann) Britton & Rose. Definition: Areoles Protocoryphantha type: grooves becoming slowly longer on each newly formed tubercle during the ontogenesis of the individual plant, this process being interrupted for a longer period (years) when the grooves reach about three-quarters of the way to the tubercle axils and incomplete grooves persist on old tubercles, flowering only after attainment of maximum length areolar grooves reaching the axils. Cortex watery.

2. Coryphantha poselgeriana (Dietrich)

Britton & Rose Cactaceae 4: 28,1923

Basionym: Echinocactus poselgerianus Dietrich, Allg. Gartenz.19: 346,1851. Type: not designated.

Synonyms: Mammillaria salm-dyckiana Scheer in Salm-Dyck, Cact. Hort. Dyck 1849:134,1850 nom. rejic. (Dicht 2000); Mammillaria salm-dyckiana brunnea Salm-Dyck, Allg. Gartenz. 18:394,1850 nom. rejic.; Echinocactus salm-dyckianus Poselger,Allg. Gartenz. 21:102, 1853, nom. rejic.; Cactus salm-dyck-ianus Kuntze, Rev. Gen. Pl. 1: 261, 1891, nom. rejic.; Coryphantha salm-dyckiana (Scheer) Britton & Rose, Cactaceae 4: 28, 1923, nom. rejic.; Coryphantha salm-dyckiana (Scheer) Britton & Rose var. brunnea (Salm-Dyck) Unger, Kakt. and. Sukk. 37(5):85, 1986 nom. rejic.; Echinocactus saltillensis Poselger, Allg.Gartenz. 21: 101, 1853; Mammillaria saltillensis Bödeker, Zeitschr. Sukk-Kunde 3: 268, 1928; Coryphantha poselgeriana var. saltillensis (Poselger) Bremer, Cact. Suc. Mex. 22:16, 1977; Mammillaria valida Purpus, Monatsschr. Kakt. 21: 97, 1911, nom.illeg. ICBN Art.53 [Non Mammillaria valida Weber, Dict. Hort. Bois 806, 1898. Nec Mammillaria scheeri var. valida Engelmann, Proc. Amer. Acad.3: 265, 1857 (=Coryphantha robustispina subsp. scheeri (Lemaire) Taylor)]; Coryphantha valida (Purpus) Bremer, Cact. Suc. Mex. 22: 14, 1977 (see ICBN Art. 58.3); Coryphantha poselgeriana (Dietrich) Britton & Rose var. valida (Purpus) Heinrich ex Backeberg, Cactaceae 5: 3050, 1961 (see ICBN Art. 58.3) [Non Coryphantha scheeri (Kuntze) Benson var. valida (Engelmann) Benson, The Cacti of Arizona 25, 1969 (=Coryphantha robustispina subsp. scheeri (Lemaire) Taylor)]; Mammillaria kieferiana

Hort. ex Bodeker, Zeitschr. Sukk.-Kunde 3: 270, 1928; Coryphantha kieferiana Berger, Kakteen 276, 1929; Echinocactus salinensis Poselger 1853 sensu Britton & Rose, Cactaceae 4:28, 1923; Mammillaria difficilis Quehl 1908 sensu Britton & Rose,Cactaceae 4: 28,1923.

Body solitary, semi-globose to short cylindrical, 10-30 cm high and 13-18 cm in diameter, dull grey-bluish-green, apex rounded, woolly, central thin taproot. Tubercles in 8 and 13 series, near the base rounded 4-6-edged, delineated by a groove, near the areole convex-conical, somewhat flattened on the upper surface, rounded at the end, up to 30 mm high and 50 mm wide, younger ones narrower, up to 25 mm long, with a deep, woolly groove with 1-5 yellow-orange glands over the whole length. Axils woolly, with gland. Areoles round, 7 mm diameter, slightly woolly. Radial spines 9-12, with 7-8 of them radiating sideways and downwards, slightly protruding, straight or somewhat curved, strong, thick subulate, the strongest 3-4 below, flattened, edged laterally, 30-40 mm long, in youth whitish, later reddish-brown with dark tips, ultimately grey, blackish on the upper side. Upper 4-5 spines thinner, ascending, slightly curved, in a bundle, slightly shorter, whitish with black tips, ultimately greyish. Central spine 1, straight, porrect, in shape, thickness, colour and length similar to lower radial spines. Flower funnelform, 6-7 cm diameter, outer perianth segments oblanceolate, margin entire,dentate towards the tip,tips usually acute, light rose to carmine-red with darker, sometimes greenish midstripe. Inner perianth segments oblanceolate, margin entire, dentate towards the tip, pale rose with carmine-red throat. Flower at some locations also yellow. Filaments carmin-red, anthers dark yellow, style reddish, stigma lobes 8-10, 3-5 mm long, greenish-yellow. Fruit green, juicy berry with attached flower remnants, oblong to large ovoid, 25-50 mm long and 7-18 mm in diameter. Seeds reniform,

2-2.5 mm long and 1.5 mm wide, testa shiny, reticulate, red-brown.

Development: At first, the tubercles are flat and separated from each other by a deep groove only ("sea-urchin"). Later, they are extended and groove development starts. Before the plant is completely adult, sterile faint flowers appear, which never open. Distribution: Mexico: Coahuila, San Luis Potosí, Durango and Zacatecas. Habitat: Alluvial plains with sandy-gypsum or gravelly soil, nearly bare. Locations checked: Mexico: Coahuila: Hipolito, Estacion Marte, Saltillo, Cuatro-cienegas, El Papalote, Zona de Minas, La Rosa, Nueva Atalaya, Arteaga, Ocampo. San Luis Potosí: Villa de Ramos, Vanegas, Tolosa. Durango: Santa Clara, Cuencamé, Yerbanis. Zacatecas: Rio Grande, Juan Aldama. Incidence: Least concern. Comments: This strongly spined species, reminiscent of an Echinocactus, has a long nomen-clatural history,like Coryphantha robustispina ssp. scheeri Br. & R, with which it was confused again and again and, in fact, it should be named Coryphantha salm-dyckiana.

As UNGER (1986) correctly stated, this species was first described as Mammillaria salm-dyckiana by SCHEER (1850) 1 year before DIETRICH (1851) described it as Echinocactus poselgerianus.

Mammillaria salm-dyckiana Scheer ex Salm-Dyck, Cact. Hort. Dyck 1849:134, 1850 was the first description of a globose plant with very large tubercles of 27 mm (one thumb) width (in later years depressed and even up to 40.5 mm [1.5 thumbs] diameter), 8-10 radial spines of 27-40 mm (1-1.5 thumbs) in length and one single central spine of 54 mm (2 thumbs) in length. This, however, is the older description of Echinocactus posel-gerianus Dietrich 1851, the basionym of Coryphantha poselgeriana (Dietrich) Britton & Rose 1923 (UNGER 1986,DICHT 1996).

When Britton and Rose published the new combination Coryphantha salm-dyckiana

(Scheer) Britton & Rose in Cactaceae 4:39, 1923 they erroneously applied the name to a different plant with small tubercles (15 mm width), with 15 radial spines of 10-15 mm length and 1-4 central spines (the longest of them 20-25 mm long) corresponding to Coryphantha delaetiana (Quehl 1908) Berger 1929, a plant which was listed by BRITTON & ROSE (1923) as one of the synonyms of Coryphantha salm-dyckiana (Scheer) Britton & Rose.

Since BRITTON & ROSE (1923), Coryphantha salm-dyckiana (Scheer) Britton & Rose has continuously been used in its incorrect sense as a synonym of Coryphantha delaetiana (Quehl) Berger (BERGER 1929, BORG 1937, BACKEBERG 1961,GLASS 1975,BRAVO & SANCHEZ-MEJORADA 1991, PRESTON-MAFHAM 1991) and never as the older, but valid synonym of Coryphantha poselgeriana (Dietrich) Britton & Rose 1923.

The confusion that would be caused if Coryphantha salm-dyckiana (Scheer) were to displace C. poselgeriana (Dietrich) Britton & Rose, the lack of an extant type, the difficulty of typifying the name and the evident misapplication of the name by Britton & Rose are sufficient grounds to justify a proposal for rejection of Scheer's basionym Mammillaria salm-dyckiana, thus removing it from contention with Echinocactus poselgerianus Dietrich, and adopting Coryphantha delaetiana (Quehl) Berger for Coryphantha salm-dyck-iana Britton & Rose.

This proposal was submitted to the Committee for Spermatophytes in 2000. Illustrations: Distribution see Plate 9, map 1; plant portrait see Plates 15 and 16.

3 a. Coryphantha robustispina

(Schott ex Engelmann) Britton & Rose subsp. robustispina Cactaceae IV: 33,1923

Basionym: Mammillaria robustispina Schott ex Engelmann, Proc. Amer. Acad. 3: 265,1856.

Type: Mexico: Llanos on the southern side of Baboquivaria mountain, presumably on the Sonora side of the boundary (Schott 4, MO 01443835, holo; F 42679, iso.). Synonyms: Cactus robustispinus (Schott) Kuntze, Rev.Gen.Pl.1:261,1891; Mammillaria robustissima Schott, West. Am. Scient. 21, 1900; Coryphantha muehlenpfordtii (Posel-ger) Britton & Rose var. robustispina (Schott ex Engelmann) Marshall, Ariz. Cact. 94, 1953; Coryphantha muehlenpfordtii Britton & Rose subsp. robustispina (Engelmann) Dicht, Kakt. and. Sukk. 47 (12): 252, 1996; Coryphantha scheeri (Kuntze) Benson var. robustispina (Schott ex Engelmann) Benson, Cact. Arizona 25, 1969; Coryphantha scheeri Lemaire var. robustispina Benson, cacti of the United States and Canada 820, 1982; Mammillaria brownii Toumey, Bot. Gaz. 22: 253, 1896; Cactus brownii Toumey, Bot. Gaz. 22: 253,1896. Body solitary or group-forming, globose or slightly higher than wide, 5-15 (-20) cm high, dull grey-green, apex flattened. Tubercles in 8 and 13 series, conical, rounded, upper surface somewhat flattened, older ones rhomboid, base 20 mm wide, 17 mm high, length of upper surface 22 mm, of lower surface 25 mm, groove on upper surface from spiniferous areole to axil, in youth white-woolly, with small, hidden glands. Areoles round, 7 mm in diameter, in youth white-woolly. Radial spines 8-12, lower and lateral 6-8 radial, horizontal, straight, subulate, stiff, 30 mm long, upper ones in two layers closely set, somewhat thinner and longer, all brownish in youth, later becoming grey from the base, uppermost lighter with dark tips. Central spine 1, more robust, straight, porrect, tip curved to hooked, 30 mm long, colour as for lower radial spines. Flowers 5-7 cm diameter, 5-6 cm long, funnelform, outer perianth segments lanceolate, margin fimbriate, acute, dentate towards the tip, yellow with reddish dorsal midstripe, inner perianth segments lanceolate, margin entire, dentate towards the tip, acute, yellow to bronze-coloured, some times reddish at the base, filaments reddish, anthers yellow, stigma and stigma lobes light yellow. Fruit green, juicy, longish berry, 30-55 mm long, 12-20 mm wide. Seeds reni-form, 3.2 mm long, 2.5 mm wide, shiny, brown, testa reticulate.

Distribution: South of Arizona in a defined area spreading about 70 km east-west and 80 km north-south and in the adjacent region of the Mexican State Sonora. Habitat: Flat and sparsely vegetated plains or on flat hilltops in semi-desert grasslands. Location checked: USA: Arizona: Pima County.

Incidence: Endangered, mainly by urbanisation, constructions, burning off, Lehman's love-grass, illegal collection, off-road vehicles.

This species, popularly known as the Pima Pineapple Cactus, was included in the List of Endangered Species of the USA on 20 April 1992.

Comments: Inclusion of Coryphantha robustispina Br. & R as a variety of Coryphantha muehlenpfordtii (MARSHALL 1953) and Coryphantha scheeri (BENSON 1969), respectively, was not accepted by everybody, since in all the literature, contrary to Coryphantha scheeri, it was mentioned as being without any glands. Therefore, it was allocated to the series Aulacothelae Lemaire (Sulcolanatae Britton & Rose) and not to Glanduliferae Salm-Dyck (Recurvatae Britton & Rose, Neo-coryphantha Backeberg).

Our own observations showed that C. muehlenpfordtii subsp. robustispina clearly does make glands. These are much smaller than those of the typical subspecies and are often hidden by felt in the grooves. Therefore, this species often shows sooty moulds ("black fungus").

Illustrations: Distribution see Plate 9, map 1; plant portrait see Plate 17, photo 1.

3 b. Coryphantha robustispina subsp. scheeri

(Lemaire) N.P. Taylor Cact. Cons. Init. 6: 18,1998. Basionym: Coryphantha scheeri Lemaire, Cactées 35,1868.

Lectotype: Allg.Gartenz.15 (3): t.2,1847 (Taylor, CCI 6:18,1998).

Synonyms: Mammillaria scheeri Müh-lenpfordt,Allg. Gartenz. 5:97,1847 [non Mam-millaria scheeri Mühlenpfordt, Allg. Gartenz. 13: 346,1845 (= Neolloydia conoidea)]; Mam-millaria scheeri Mühlenpfordt var. valida Engelmann, Proc. Amer. Acad.3: 265, 1856 [non Mammillaria valida Weber, Dict. Hort. Bois 806, 1898, nec Mammillaria valida, Purpus, Monatsschr. Kakt.21: 97, 1911, nec Coryphantha valida (Purpus) Bremer,Cact.Suc.Mex.22: 14, 1977]; Cactus scheeri (Mühlenpfordt) Kuntze, Rev. Gen. Pl.1: 261,1891; Coryphantha scheeri (Kuntze) Benson, Cact. Succ. J. (US) 41: 234,1969; Coryphantha scheeri (Kuntze) Benson var. valida (Engelmann) Benson,Cact.Ari-zona 25, 1969; Coryphantha scheeri (Kuntze) Benson var. uncinata Benson, Cact. Succ. J.(US) 41: 234, 1969; Coryphantha scheeri Lemaire var. uncinata Benson, Cacti of the United States and Canada: 820, 1982; Echino-cactus muehlenpfordtii Poselger,Allg. Gartenz. 21: 102, 1853 [non Mammillaria muehlenpfordtii Förster, ex Otto & Dietr.,Allg. Gartenz. 15:49,1847, nec Echinocactus muehlenpfordtii Fennel,Allg.Gartenz.9:65,1847 (=Thelocactus setispinus)]; Coryphantha muehlenpfordtii (Poselger) Britton & Rose, Cactaceae 4 :28, 1923; Coryphantha muehlenpfordtii Britton & Rose subsp. muehlenpfordtii Dicht, Kakt. and. Sukk. 47 (5): 98,1996; Coryphantha muehlenpfordtii subsp. uncinata (Benson) Dicht, Kakt. and. Sukk. 47(5): 98,1996; Mammillaria engel-mannii Cory, Rhodora 8: 405, 1936 [non Coryphantha engelmannii Lemaire, Cactées 34,1868, nec Cactus engelmannii Kuntze, Rev. Gen. Pl.1: 261,1891]; Coryphantha neoscheeri Backeberg, Cactaceae 5: 3051, 1961; Cory-phantha robustispina subsp. uncinata (Benson) N.P.Taylor, CCI 6:18,1998.

Body solitary, rarely sprouting at the base, globose to egg-shaped, then short cylindrical or somewhat conical, 10-20 cm high and 12 cm in diameter, dull green, apex rounded, slightly woolly. Tubercles in 5 and 8 series, prisma-shaped at the base, then cylindrical, at the base 15-19 mm wide, 12 mm high, length of upper and lower surface 30-35 mm, with deep, complete groove with white wool and up to four variously separated reddish-brown glands. Axils in youth with white wool, then naked and somewhat broadened. Areoles oval, 3.5 mm wide, 5 mm long, in youth woolly, later naked. Radial spines 8-11, 3 thinner ones in the upper part of the areole, closer set, the other ones horizontal and radially arranged, straight, stiff needle-like, 20-25 mm long, yellowish-white with dark tips, lower ones sometimes brownish. Central spines 1-4, the dominating one straight, por-rect, thin-subulate, up to 35 mm long, the others straight upwards and slightly directed outwards, somewhat thinner and shorter, all somewhat darker and more intensely coloured than radials. Flowers, fruits and seeds: as C. robustispina subsp. robustispina. Distribution: USA: south of New Mexico and Texas. Mexico: in the adjacent regions of Chihuahua.

Habitat: Flat and sparsely vegetated plains or lower hill slopes or on flat hilltops in semi-desert grasslands.

Locations checked: USA: New Mexico: Las Cruces, Carlsbad, Eddy County. Texas: Pecos County. Mexico: Chihuahua: Ciudad Chihuahua, Villa Ahumada, El Sueco und Benito Juarez.

Incidence: Least concern. Comments: From the beginning, the nomen-clatural history of this large-growing, long-tubercled species with its bronze-coloured flowers was characterised by vagueness and confusion. The first description (with illustration) as Mammillaria scheeri by MUH-LENPFORDT 1847 was an illegitimate homonym, because under the same name,

MÜHLENPFORDT himself had already described Neolloydia conoidea in 1845.

This circumstance may have induced POSELGER in 1853 to publish anew Mammil-laria scheeri as Echinocactus muehlenpfordtii. However, by so doing, he made a similar mistake as Mühlenpfordt before, since this name was in use already: FENNEL in 1839 had described an Echinocactus muehlenpfordtii (syn. Thelocactus setispinus). Poselger's new name should have replaced Mammillaria scheeri Mühlenpfordt as a synonym, but Poselger failed to state which of the two relevant publications of Mühlenpfordt he was referring to in his publication. For this reason, the species name used by Poselger must be considered invalid.

In 1856, ENGELMANN described Mammil-laria scheeri var. valida from Texas without having seen Mammillaria scheeri Mühlenpfordt. His knowledge of this species was taken from the description of Prince SALMDYCK 1850. Engelmann had presumed that his var. valida might be just a somewhat larger and more heavily spined northern form of Mammillaria scheeri from Chihuahua. When visiting the collection of Prince Salm-Dyck,he found original plants of Mammillaria scheeri which corresponded completely with his var. valida, thus confirming his initial assumption (cf. J.COULTER 1894).

As Engelmann himself had realised already, var. valida does not contain enough distinct criteria to delineate a variety or subspecies, a view which, among others, was also shared by Coulter, Quehl, Britton & Rose und Backeberg.

When LEMAIRE (1868) included Mammil-laria scheeri in the genus Coryphantha, he did not mention Mühlenpfordt as its author, nor did he specify which one of the earlier two versions of Mammillaria scheeri he meant.

LEMAIRE (in Cactées 35 ff., 1868) pointed out that he referred to SALM-DYCK (1850). In Salm-Dyck's book, Mammillaria scheeri

Muehlenpfordt 1847 belonged to the group of species to which, among others, Mammillaria aulacothelae also belonged and which Lemaire raised to genus level as Coryphantha (Engelmann) Lemaire, while Mammillaria (Neoloydia) conoidea (syn. M. scheeri Muehlenpfordt 1845) was put into the same infrageneric group of Mammillaria (viz. §6, Centrispinae) by both authors. This fact was overseen by a series of later authors, since it makes Coryphantha scheeri Lemaire a valid nomen novum (avowed substitute).

KUNTZE (1891) removed Mammillaria scheeri Muehlenpfordt to Cactus scheeri, naming its author, but without indicating whether the description of 1845 or 1847 was meant. Therefore, this name was also invalid.

In 1923 in Cactaceae IV: 28 by BRITTON & ROSE, for the first time Coryphantha muehlenpfordtii Britton & Rose appeared as a new combination. These two authors had realised the problems around Mammillaria scheeri Muehlenpfordt and, therefore, avoided the name Coryphantha scheeri, but obviously they were not aware that Echinocactus muehlenpfordtii was also an invalid name and, therefore, the combination Coryphantha muehlenpfordtii was inadmissible.

As a valid name we find Mammillaria engelmannii Cory 1936, which was used by BENSON (1950) as a synonym of Coryphantha muehlenpfordtii sensu Britton & Rose in The Cacti of Arizona.

In BACKEBERG's Cactaceae V:3051 (1961) one more epithet appears: Coryphantha neoscheeri. Probably Backeberg intended to escape the confusion surrounding the name Coryphantha scheeri. He had noticed that Mammillaria scheeri and Mammillaria muehlenpfordtii were both homonyms and he also believed that Lemaire's Coryphantha scheeri was invalid, too. In fact, Coryphantha neoscheeri is a valid description, but has to be regarded as superfluous.

In 1969, the new combination Coryphantha scheeri (Kuntze) L. Benson was published in

Cact. Succ. J. US 41:234 with the contention that following article 72 of the International Code of Botanical Nomenclature Cactus scheeri Kuntze,Rev.Gen.Pl. 1:261,1891 should be regarded as the first valid publication of this species. However, as shown above, the name Cactus scheeri Kuntze is invalid because of Muhlenpfordt's imprecise indication of its publication and,therefore,Benson's new combination has to be regarded as invalid, too.

BENSON himself for the first time mentioned the circumstances which make Coryphantha scheeri Lemaire a valid nomen novum in 1982 (Cacti of the United States & Canada: 959) and consequently recombined the varieties valida, uncinata and robustispina. By doing so, he, like DICHT (1996), also overlooked the fact that Mammillaria robustispina Engelmann (1856) is an older name than Coryphantha scheeri Lemaire (1868) and thus cannot be a subspecies, but must be the real, valid name of the species with Lemaire's Coryphantha scheeri as a subspecies. This context was finally rectified by Nigel P. TAYLOR in 1998 (Cactaceae Consensus Initiatives 6: 18) by recombining the two subspecies scheeri and uncinata. The subspecies valida had already been retracted by DICHT (1996).

Finally, the present nomenclature completely corresponds with the rules of the ICBN.

Illustrations: Distribution see Plate 9, map 1; plant portrait see Plate 17, photos 2 and 3.

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