Aa Series Retusae Dicht A Lthy

Type: Coryphantha elephantidens (Lemaire)

Lemaire.

Definition: Radial spines mostly subulate.

17 a. Coryphantha elephantidens (Lemaire)

Lemaire subsp. elephantidens Cactées 35,1868

Basionym: Mammillaria elephantidens Lemaire, Cact.Aliq. Nov. 1,1838. Lectotype: Lemaire, Iconogr. descr. des Cactées, Part 5,t.9,1841 (Dicht & A. Lüthy, CSI 11: 14,2001).

Synonyms: Mammillaria retusa Pfeiffer, Allg. Gartenz. 5: 369,1837 (nom. rejic. prop.); Aula-cothele elephantidens (Lemaire) Monville, Cat. Pl.Exot.,21,1846; Echinocactus elephantidens (Lemaire) Poselger,Allg.Gartenz.21:102, 1853; Cactus elephantidens (Lemaire) Kuntze, Rev. Gen. Pl. 1: 260, 1891; Coryphantha ele-phantidens var. barciae Bremer, Cact. Suc. Mex. 18: 55, 1973; Mammillaria sulcolanata Lemaire, Cact. aliqu. nov. 2,1838; Aulacothele sulcolanata (Lemaire) Monville, Cat. Pl. Exot., 21,1846; Echinocactus sulcolanatus (Lemaire) Poselger, Allg. Gartenzeitung 21: 102, 1853; Cactus sulcolanatus (Lemaire) Kuntze, Rev. Gen. Pl. 1: 261, 1891; Mammillaria re-curvispina De Vriese, Tijdscht. Nat. Gesch. 6: 53, 1839; Cactus recurvispinus (De Vriese) Kuntze, Rev. Gen. Pl. 1:261,1891; Coryphantha recurvispina (De Vriese) Bremer, Cact. Suc. Mex. 21: 12, 1976; Coryphantha garessii Bremer, Cact. Succ.J.(US) 52:82,1980. Body solitary, depressed-globose to globose, apex flattened and strongly white-woolly, about 8 cm high and 10 cm diameter, dull dark-green. Roots napiform, deep, secondary roots fibrous. Tubercles in 5 and 8 or 8 and 13

series, broad conical, rounded, upper surface flattened, cut at edges, at the base 22-30 mm wide, 15-20 mm high, length on upper surface 14-20 mm, on lower surface 15-20 mm, with a deep, white-woolly groove in youth. Axils recessed, longish, 8 x 4 mm, in youth white-woolly. Radial spines 8-10,2-3 of them on each side and 1 below pointing downwards, all subulate, straight, slightly curved to the body, 18-22 mm long, in the upper part of the areole 1-4 much thinner, set back, straight or slightly curved to the side, 12-14 mm long, all radial spines horn-coloured to brown, darker towards the tips, dull, later becoming grey from the base. Flowers 5-8 (-10) cm diameter, 5 cm long, light purple or yellow. Outer perianth segments broad lanceolate, acute, purple with light margins or yellow with red-brown midstripe dorsally, inner perianth segments broad lanceolate, margins entire, dentate towards the tip, acute, light purple, lighter to nearly white towards the throat, sometimes yellowish or yellow, in the throat sometimes reddish, filaments purple or whitish or whitish with purple base, anthers yellow, stigma whitish-yellow, stigma lobes small, whitish-yellow. Fruit longish, 3.5 cm long, 1 cm diameter, green, juicy berry with attached flower remnants. Seeds longish, reniform, 3.5 mm long, 1.5 mm wide, brown, testa reticulate.

Distribution: Mexico: Morelos, Puebla, Oax-

aca, Querétaro, Jalisco, Guanajuato, Aguas-

calientes and Zacatecas.

Habitat: Lava soil in plains and low ridges in grassland, sometimes protected by bushes or

Opuntias.

Locations checked: Mexico: Morelos: Zacate-pec, Tlatizapan,Yautepec, Las Estacas. Puebla: Tepexco, Izucar de Matamoros, Tejalapa. Oax-aca: Etla. Querétaro: Humilpán. Guanajuato: Irapuato-Cueramaro. Jalisco: Lagos de Moreno, Huejucar. Zacatecas: Tepetongo. Incidence: Least concern. At some localities systematically removed from pastures by farmers.

Illustrations: Distribution see Plate 11,map 1; plant portrait see Plates 31 and 32.

17 b. Coryphantha elephantidens subsp. bumamma (Ehrenberg) Dicht & A. Luthy

Cact.Syst.Init. 11: 14,2001. Basionym: Mammillaria bumamma Ehrenberg, Allg. Gartenz. 17: 243,1849. Typ: not designated.

Synonyms: Mammillaria elephantidens bumamma Schumann, Keys Monogr. Cact. 43, 1903; Coryphantha bumamma (Ehrenberg) Britton & Rose, Cact. 4: 33,1923. Body globose, somewhat flattened, 13 cm diameter, epidermis slightly bluish-green. Forming large groups of more than 50 cm diameter. Tubercles in 8 and 13 series, conical, rounded, upper part flattened, at the base 27 mm wide, 16 mm high, length of upper surface 13 mm, of lower surface 27 mm. Axils in youth very woolly, later naked. Areoles longish, 5 x 3 mm, slightly woolly. Radial spines 3 strong ones to the sides, 16-20 mm long, brown with dark tips, then greyish-brown with dark tips, in upper part of areole 2 more and below 1 more 11 mm long, all subulate, the lower one and the upper ones somewhat thinner than the lateral ones. Flowers 5-6 cm diameter, outer and inner perianth segments as for subsp. greenwoodii (see 17.c). Filaments reddish at the base, lobes greenish, stigma longer than filaments. Fruit claviform, 38 mm long and 13 mm diameter, whitish-green with a reddish tinge. Seeds 4 mm long and about 1 mm diameter, light brown. Distribution: Mexico: Oaxaca and Michoa-can.

Habitat: Deep, fine lava-soil with grass, sometimes with Cereus forests. Locations checked: Mexico: Oaxaca: Toto-lapan. Michoacan: El Paradero. Incidence: Least concern. Illustrations: Distribution see Plate 11,map 1; plant portrait see colour Plate 33.

17 c. Coryphantha elephantidens subsp.

greenwoodii (Bravo) Dicht & A. Luthy Cact. Syst. Init. 11: 14,2001. Basionym: Coryphantha greenwoodii Bravo, Cact. Suc.Mex. 15: 27,1970. Type: Mexico, Veracruz, near Acultzingo, 1600 m, Greenwood s.n. (MEXU). Body solitary or sprouting, group-forming, flat-globose, about 9 cm diameter, mostly under groundlevel, above ground 5-6 cm high, apex depressed, with much white wool. Tubercles in 5 and 8 or 8 and 13 series, obliquely conical, 18-22 mm wide, 17-19 mm high, length of upper surface 11-12 mm, of lower surface 14-20 mm. Radial spines somewhat protruding and recurved, 9-10, among them 5 strong ones to the sides, 1 thinner below oblique and 3-4 thinner ones bundled at top. The thinner ones nearly white with dark tip, the stronger ones brown with dark tips. Flowers fragrant, 5 cm in height and diameter, outer perianth-segments lanceolate, yellow with reddish midstripe dorsally, inner perianth-segments lanceolate, margins entire, acute, 5 mm wide, yellow, filaments dark yellow, stigma whitish with 4 very small lobes. Fruits as for subsp. elephantidens (see 17.a). Seeds reniform, 1.2 mm wide, 2 mm long, brown, testa reticulate. Distribution/location checked: Veracruz in the region of Acultzingo, at 1600 m above sea level, on grassy ground. Habitat: Lava soil with grass and small bushes.

Incidence: Endangered. Small range of distribution, one single location, constant threat by human activities like farming, road construction, clearing of pastures by fire observed. Comments: This subspecies was named in honour of Ing. Edward W. Greenwood, a cactus and orchid enthusiast and photographer who accompanied H. Bravo on many excursions and who was the first one to discover this taxon.

At first sight, ssp. greenwoodii looks quite similar to Coryphantha pycnacantha. How ever, it has the typical tubercle form of Coryphantha elephantidens which is always wider than long (C. pycnacantha longer than wide) and therefore was allocated to Coryphantha elephantidens by us. Illustrations: Distribution see Plate 11,map 1; plant portrait see Plate 34.

18. Coryphantha retusa Britton & Rose Cactaceae 4: 38,1923, nom. cons. prop. Lectotype: Britton & Rose, Cact.4:38 (Fig. 36), 1923, Dicht & A. Lüthy, Cact.Syst.Init. 11:11, 2001).

Synonyms: Melocactus mammillariaeformis Salm-Dyck, Allg. Gartenz. 4: 192, 1836 (nom. rejic. prop.); Mammillaria cephalophora Salm-Dyck, Cact. Hort. Dyck 1849: 137 1850 (nom. illegit.); Echinocactus cephalophorus (Salm-Dyck) Poselger, Allg. Gartenz. 21: 102, 1853; Cactus cephalophorus (Salm-Dyck) Kuntze, Rev. Gen. Pl. 1: 260, 1891; Coryphantha melleospina Bravo, An. Inst. Biol. Mex. 25: 526,1954; Coryphantha retusa (Pfeiffer) Britton & Rose var. melleospina (Bravo) Bravo, Cact. Suc.Mex.27: 17,1982. Body solitary, flattened-semiglobose to globose with narrowed base, 5-10 cm diameter, 3-8 cm high, dark green, apex depressed,with much white wool, roots fibrous. Tubercles in 8 and 13 or 13 and 21 series, conical, rounded, flattened above, below slightly bulging,

13-18 mm wide, 7-12 mm high, upper surface 9-12 mm long, lower surface 12-17 mm long, with deep, complete groove. Axils in youth with copious white wool. Areoles oblong, 4 mm long, 2 mm wide, in youth woolly. Radial spines 10-14 (-19), 3-4 of them on each side, strong, subulate, curved to the body and slightly downwards, radiating, the lowest one repositioned, straight downwards, 3-5 thinner needle-like spines in the upper part of the areole, repositioned, grouped together, directed upwards and slightly to one side, all

14-18 mm long, in new growth pale yellow, then white with darker tips, later turning grey. Central spine: occasionally and on some areoles 1 porrect central spine curved downwards, more rarely upwards, subulate, 18 mm long, colour as radial spines. Flowers 4 cm in length and diameter, yellow, all perianth segments lanceolate, acute, filaments reddish, anthers yellow, stigma and lobes whitish-yellow. Fruit olive-green, juicy berry, oblong, 28 mm long, 9 mm wide. Seeds light brown, oblong, date-shaped, 2.5 x 1 mm, reticulate. Distribution: Mexico: Puebla and Oaxaca. Habitat: Grassland on lava-soil. Locations checked: Mexico: Puebla: Tehuit-zingo, Acatlan, Petlalcingo, Xayacatlan. Oaxaca: Saltenango, Etla, 15 km south of Oaxaca, Buenavista, Mitla, Santiago Matatlan, Hua-juapan de Leon, Ocotepec, El Molino, Chazumba, Ocotlan, Cerro Verde. Incidence: Least concern. Comments: BACKEBERG described a variety pallidispina (Cactaceae VI: 3874, 1962) with the following differences: spines 15, among them 12 more rigid with thickened bases, yellowish, the other 3 much thinner, close to each other and porrect, from horn-coloured to pale grey, with dark tips and yellowish, thickened base. In 1954 BRAVO published a Coryphantha melleospina, which she combined to Coryphantha retusa var. melleospina in 1982. As our field-studies have shown, this is just one form of Coryphantha retusa, but with some more radial spines of a golden-yellow colour and somewhat smaller dimensions. Illustrations: Distribution see Plate 11,map 1; plant portrait see Plates 35 and 36.

Comments relating to the Series Retusae When BRITTON & ROSE (1923-1938) allocated Mammillaria retusa Pfeiffer to Coryphantha, they obviously misinterpreted the first description. They used this epithet for small-tubercled plants with 12 radial spines, which they had received from various collectors from Oaxaca in 1920, among them Solis,whose plant they illustrated (l.c. Fig. 36). However, Pfeiffer's first description was of a plant with much larger tubercles of more than 1 in. width and with fewer, but much longer radial spines. It undoubtedly corresponded with what is known as Coryphantha elephantidens today. Indeed, Mammillaria retusa had been cited as a synonym of Mam-millaria elephantidens by all authors of the nineteenth century. Thus, following the rules of ICBN the name Coryphantha retusa as the oldest and therefore, valid name should be used for Coryphantha elephantidens.

A plant as widely distributed as Coryphan-tha retusa would surely have been collected and described early in the nineteenth century. There is, in fact, such a description which matches the plant in all respects: Melocactus mammillariiformis Salm-Dyck, Allg. Gartenz. 4: 148, 1836. This first description by SalmDyck was very precise and even mentioned the single strong central spine appearing on some areoles only. The very woolly apex of this plant had led Salm-Dyck to regard it as a Melocactus, but with a plain cephalium ("cephalium planum"). However, Lemaire and Pfeiffer indicated to him that this would be an error. In consequence, Salm-Dyck redescribed this species, but this time as Mammillaria cephalophora in Cact. Hort. Dyck 1849,1850.

According to the ICBN rules, Coryphantha retusa (Pfeiffer) Br. & R. should now be named Coryphantha mammillariaeformis and Coryphantha elephantidens (Lemaire) Lemaire renamed Coryphantha retusa.

However, following Britton & Rose, the name Coryphantha retusa has been used by all authors in the sense of Britton & Rose and thus strict application of ICBN rules would lead to negative nomenclatural changes. Therefore, we made a proposal to the Committee for Spermatophytes to conserve Coryphantha retusa in the sense of Britton & Rose and to reject Melocactus mammillariiformis Salm-Dyck (CSI 10: 19, 2000). This was the only way to ensure continuity of the present nomenclature.

II.A.b Series Pycnacanthae Dicht & A.Lüthy

Type: Coryphantha pycnacantha (Martius)

Lemaire.

Definition: 3 or more central spines always present, but none porrect, all appressed.

19. Coryphantha pycnacantha (Martius)

Lemaire Cactées 35,1868

Basionym: Mammillaria pycnacantha Mar-tius, Nov. Act. Nat. Cur. 16: 325,1832. Lectotype: Martius, Nov. Act. Nat. Cur.16: 325, 1832 (Dicht & A. Lüthy, CSI 11: 15,2001). Synonyms: Mammillaria pycnacantha spin-osior Monville ex Salm-Dyck,Hort.Dyck 1844: 4, 1845; Aulacothele pycnacantha (Martius) Monville, Cat. Pl. Exot., 21,1846; Echinocactus pycnacanthus Poselger, Allg. Gartenz. 21: 102, 1853; Cactus pycnacanthus Kuntze, Rev. Gen. Pl. 1: 261,1891; Mammillaria acanthostephes Lehmann, Allg. Gartenz. 3: 228, 1835; Aula-cothele acanthostephes (Lehmann) Monville, Cat. Pl. Exot., 21, 1846; Mammillaria acan-thostephes recta Hort ex Labouret, Monogr. Cact. 138,1853; Cactus acanthostephes Kuntze, Rev. Gen. Pl. 1: 260,1891; Mammillaria magn-imamma Otto, Allg. Gartenz. 29: 228, 1835 [non Haworth, 1824]; Mammillaria magni-mamma lutescens Salm-Dyck, Cact. Hort. Dyck 1849: 121, 1850; Mammillaria arietina Lemaire, Cact.Aliq.Nov. 10,1838; Mammillaria arietina spinosior Lemaire, Cact. Gen. Nov.Sp. 94,1839; Mammillaria scepontocentra Lemaire, Cact. Gen. Nov. Sp. 43, 1839; Cactus scepontocentrus Kuntze, Rev. Gen. Pl. 1: 261 1891; Mammillaria winkleri Foerster, Allg. Gartenz. 15: 50, 1853; Echinocactus winkleri Poselger, Allg. Gartenz. 21: 102, 1853; Cactus winkleri Kuntze, Rev. Gen. Pl. 1: 261, 1891; Coryphantha connivens Br. & R., Cactaceae 4: 34, 1923; Coryphantha andreae Purpus & Bödeker, Zeitschr. Sukk.-Kunde 3: 251,1928.

Body solitary, flat-globose to semiglobose, 9.5 cm diameter, 3.5 cm high, blue-grey green, apex depressed, woolly. Central part of root nearly a taproot, with lateral fibrous roots. Tubercles in 5 and 8 (rarely 8 and 13) series, compressed conical, flattened above, bulging underneath, edges obliquely cut, at the base

17 mm wide, 12 mm high, upper surface

18 mm long, lower surface 22 mm, with deep, woolly groove in youth. Axils white-woolly in youth. Areoles elliptical, 4 mm long, 2 mm wide, woolly in youth. Radial spines 6-15, irregularly radiating, but bundled in the upper part, or merely bundled above, lower and lateral spines 9 mm long, straight, needlelike, the bundled ones above in two layers, some thicker and longer, up to 16 mm, all grey-white with dark tips. Central spines 5-7, 2-3 of them on each side, somewhat spreading, curved to the body, thick subulate, 15-21 mm long, 1 downwards, a bit thinner and shorter, in a subcentral position, all pale brown, then greyish with black tips. Flowers 30 mm long, 45 mm diameter, outer perianth-segments narrow-lanceolate, acute, 15 mm long, 3 mm wide, lemon-yellow with brown-red dorsal midstripe. Inner perianth segments broad-lanceolate, dentate towards the tip, acute, 24 mm long, 5 mm wide, lemon-yellow. Filaments short, 12 mm long, yellow, anthers yolk-yellow, stigma yellowish, with 5-7 whitish-yellow lobes. Fruit green, juicy, oblong berry, 24 mm long, 13 mm wide with attached flower remnants. Seeds reniform, 2.5 mm long, 1.5 mm wide, brown, testa reticulate.

Distribution: Mexico, Hidalgo, Puebla, Veracruz.

Habitat: Deep and flat lava soil in gras or besides anthills, together with Opuntias. Locations checked: Mexico: Mexico: Otumba, Cd. Shahogun. Hidalgo: Barranca de Metztit-lan, San Miguel Regla, Singuilucan, Ato-tonilco, Zempoala. Puebla: Yaltepec.Veracruz: Perote.

Incidence: Endangered. The locations of the plants are deep, plane lava soils, best arable land, and the range of distribution east of

Mexico City is the area of most intensive agricultural production in Mexico. Nearly all potential habitats are cultivated and the populations known are mostly reduced to very small areas along railway tracks, garbage dumps, and so on.

Differentiation: Coryphantha pycnacantha is reminiscent of Coryphantha elephantidens (Lemaire) Lemaire. However, it has more radial spines, several central spines and smaller rounder, more loosely arranged, longer than broad tubercles than the latter species. Moreover, it has flowers of pure yellow, while the filaments do not have any redness. Finally, the seeds are rounder and reni-form.

Comments: Mammillaria pycnacantha Mar-tius was imported from Mexico by the Baron L.B. von Karwinski and first described in detail by MARTIUS (1832), with informative illustrations, including a young plant, a flower and the spine-arrangement. Therefore, there are no doubts about the characteristics of the described species, which, contrary to many other descriptions of that time, can clearly be verified. The first description by Martius includes two minor errors: the indicated height of the plant and its origin Oaxaca, where, as we know today, no comparable Coryphanthas can be found. Comparing the indicated measurements of the plant with the illustration of Martius, re-measurement shows that the indication of the height applies to the complete length of the plant including the roots. The origin of the species had already been extended in 1843 by PFEIFFER and OTTO by the indication "prope Oajaca & Pachuca: Karw." In the same publication, these authors stated that this species was very close to the more flat-globose Mammillaria acanthostephes Lehmann (1835), which, consequently, was regarded as a synonym of Mammillaria pycnacantha Martius in the whole literature of the nineteenth century.

The most important hint as to the effective distribution area of Coryphantha pycnacan tha was ultimately given by EHRENBERG, who studied the botany of Mineral del Monte from 1831 to 1840. His article appeared in the journal Linnaea in 1846 and is regarded as the most reliable source of the locations of many plants collected before 1840. Ehrenberg found Coryphantha pycnacantha near the capital and also mainly in the state of Hidalgo in the plains near Pachuca, near San Mateo, Atotonilco el Grande, Regla and other locations.

The identity of the species, which was allocated to Coryphantha by Lemaire in 1868, seems to have been clear in the nineteenth century, although the descriptions vary slightly (Pfeiffer und Otto, Karwinski, SalmDyck, Schumann etc.).

The confusion surrounding this taxon only began with BRITTON & ROSE (1923). They reproduced the illustrations of Martius, but in their text they mentioned a completely different plant from the region of Oaxaca, which they had obtained from Prof. Conzatti in 1920. In contrast to the basionym, this plant, besides having 15-20 white radial spines, had 2-3 black, porrect, curved central spines and much smaller tubercles of about 10 mm diameter. In the same publication, Britton and Rose redescribed Coryphantha connivens from near the capital, Mexico City, without realising, that this very plant was identical to Martius' Coryphantha pycnacantha. However, they did mention that it was very close to Coryphantha pycnacantha (Br. & R. Cactaceae V: 34,1923). The error of Britton & Rose persisted throughout the twentieth century with the result that the name Coryphantha pyc-nacantha was, if ever, used for the wrong species. BODEKER (1933), however, discovered this mistake: in 1932 he received from Halbinger in Mexico a plant identical to that received from Prof. Conzatti by Britton & Rose. However, he noticed the discrepancy compared with the first description by Mar-tius and described this plant, which was erroneously named Coryphantha pycnacantha by

Britton & Rose, as Coryphantha reduncuspina Bodeker. This latter plant today is now included within the variety of Coryphantha pallida Br. & R.

In 1928, BODEKER was the co-author of a first description which is practically identical to that of Mammillaria pycnacantha Martius: Coryphantha andreae Purpus & Bodeker. It has to be regarded as merely a younger synonym. This name has been used until today for most specimens of Coryphantha pyc-nacantha in collections.

The first author to rectify the confusion surrounding Coryphantha pycnacantha was ZIMMERMAN (1985) who rightly treated Coryphantha connivens Br. & R. and Coryphantha andreae Purpus & Bodeker as younger synonyms of Coryphantha pyc-nacantha.

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