Ortegocactus type

Abrupt groove formation with no transitional stage with grooves getting longer, floriferous before or after production of grooves. Nectary glands and trichomes in the axil may occur before groove formation, and later also in the groove.

5. Protomammillaria type:

Like type 4, but with obligatory tubercle groove when reaching the adult stage. Nectary glands and woolly hairs (trichomes) in the groove.

6. Escobaria type:

Production of longer complete grooves extending to the axil within a short period (less than 1 year), without persistence of shortened grooves. Flowering only after attainment of maximum length grooves to the axil. Reduced capacity for producing nectary glands and, if present, only directly behind the spiniferous areole.

7. Mammillaria type:

Flowering only after the spiniferous areole stops growing, from a flower base of the meristem in axils of the previous year. Communicating meristem between areole and axil completely inactive and without formation of grooves, trichomes or nectary glands. Offsetting capacity transposed to the axil.

Phylogenetic Interpretation of the Genus Coryphantha

The Ferocactus line was confirmed by Zimmerman as an evolutionary tribe and the Coryphantha and Neobesseya lines and the Mammillaria line respectively as another phylad which differs from Ferocacti by having ribs resolved into tubercles, indehiscent, juicy fruits and loss of bract scales.

This interpretation of the two phylads does not address an important, phylogenetically special feature: production of extrafloral nectary glands. Depending on the weight given to this feature, the resulting systematics are very different, because as a consequence Cory-phantha s.str. with its capacity to produce nectary glands might well need to be included in the Ferocactus phylad.

Development of extrafloral nectary glands is a very original and complex feature. Production of sugar-containing nectar attracts ants, which, by their markings, keep off plantand bud-eating animals. This is especially important immediately before and during the flowering period. Within the Mammillaria phylad, species with concave to tabular-concave testa cells (foveolate seeds), such nectary glands can be found only in Ferocactus and Coryphantha, but never in any of the other genera. Gland development in Coryphantha s.str. clearly shows an evolutionary tendency: in the more primitive Lepidocoryphanthas (C. macromeris) and also in Protocoryphantha (C. poselgeriana and C. robustispina), it is limited to the whole length of the groove, whereas in the Protomammillaria and Orte-gocactus types it extends to the axil and is finally reduced in the Escobaria type, where, around the flowering time, (perifloral) glands if present are only produced near the spinifer-ous areole.

Since the production of nectary glands is a very complicated process comparable to the Latex syndrome in the genus Mammillaria (J. LÜTHY 1995), it has a much greater phyloge-netic significance than earlier authors had attributed to it (ZIMMERMAN 1985). Consideration of this leads to the division of the Mammillaria phylad into two main branches, the gland-producing genera (Ferocactus, Coryphantha) and the non-gland-producing genera (Acharagma, Cumarinia, Escobaria, Mammillaria).

With its groove development the gland-producing branch represents a continuum having a common ancestor. We observe the formation of very short (Ferocactus), half-long (Macromeris) and finally complete (Coryphantha) tubercle grooves with varying areole developments (Protomammillaria, Ortegocactus,Escobaria areole types).

The non-gland-producing genera follow a parallel development with the formation of shortened grooves (Cumarinia), complete grooves with varying development (Ortego-cactus, Escobaría, Protomammillaria) und finally the highest developed form with the absence of groove formation (Mammillaria).

This phylogenetic interpretation is ascertained by testa morphology and directs the genus Escobaria away from Coryphantha and closer to Mammillaria. Thus. the differentiation between Escobaria and Coryphantha is no longer confined to the testa structure differences (HUNT & TAYLOR 1990), but is extended to the consideration of nectary glands and allocates the two genera into different phylads (Ferocactus/Coryphantha line and Escobaria/Mammillaria line respectively).

The cladograms in Figs. 31 and 32 show this new interpretation.

Fig. 31. Phylogenetic interpretation of the genus Coryphantha (bold areole types). 1 Foveolate seeds;

2 potency to produce extrafloral nectary glands;

3 absence of extrafloral nectary glands; 4 tubercles instead of ribs; 5 Ferocactus-areole type; 6 Acharagma-areole type; 7 Macromeris-areole type; 8 Protoco-ryphantha areole type; 9 Protomammillaria areole type; 10 Ortegocactus areole type; 11 Escobaria areole type; 12 Mammillaria areole type; 13 smooth, reticulate seeds; 14 dark,"pitted" seeds; 15 apical (central) flowers; 16 subapical flowers; 17 outer perianth segments fimbriate; 18 outer perianth segments entire; 19 extrafloral nectary glands in grooves and/or axils; 20 absence of extrafloral nectary glands except for optional glands close to the areoles; 21 reniform seeds with narrow hilum; 22 globose seeds with large hilum

Genus Coryphantha Subgenus Neocoryphantha Section Lepidocoryphantha

C. macromeris ssp. runyonii

12,17,15

12,17,15

Genus Coryphantha Subgenus Neocoryphantha Section Lepidocoryphantha

C. macromeris ssp. runyonii

Section Robustispina C.poselgeriana

C. robustispina ssp. robustispin C. robustispina ssp. scheeri

Section Neocoryphantha Series Echinoideae C. wohlschlageri C. vaupeliana C.glanduligera C. echinoidea

Series Clavatae C. octacantha C.jalpanensis C. clavata

C. clavata ssp. stipitata C. glassii C. erecta C.potosiana

Subgenus Coryphantha Section Coryphantha Series Retusae C. elephantidens C. elephantidens ssp. bumamma C. elephantidens ssp. greenwoodii C. retusa

Series Pycnacanthae C.pycnacantha C. tripugionacantha

Series Salinenses C. kracikii C. salinensis C. difficilis C. durangensjs

C. durangensis ssp. cuencamensis

C. Iongicornis

C.pallida

C.pallida ssp. calipensis

Series Coryphantha

C. maiz-tablasensis C. sulcata

C. hintoniorum ssp. hintoniorum C. hintoniorum ssp. geoffreyi

Series Corniferae Subseries Corniferae C. nickelsiae C.pseudonickelsiae C. compacta C. cornifera

C. recurvata ssp. recurvata C. recurvata ssp. canatlanensis

C. delicata C. neglect a

C.pseudoechinus ssp.pseudoechinus C.pseudoechinus ssp. laui

Subseries Delaetianae C. delaetiana C. ramillosa

C. ramillosa ssp. santarosa C.pulleineana C. werdermannii C. echinus

Section Gracilicoryphantha C. gracilis

Fig. 32. Phylogenetic interpretation of the genus Coryphantha (bold areole and gland types). 1 Foveolate seeds; 2 potency to produce extrafloral nectary glands; 3 tubercles; 4 fruits green,juicy, seeds reniform; 5 smooth, reticulate seeds; 6 extrafloral nectary glands in grooves and/or axils; 7 Ferocactus areole type; 8 Macromeris areole type; 9 Protocoryphantha areole type; 10 Protomammillaria areole type; 11 Ortegocactus areole type; 12 Escobaria areole type; 13 fimbriate outer perianth segments; 14 tubercle grooves complete; 15 cortex watery; 16 cortex mucilaginous; 17 reduction of extrafloral nectary glands except for optional glands close to the areole; 18 majority of radial spines subulate; 19 majority of radial spines needle-like; 20 three or more central spines always present, none porrect, all ap-pressed; 21 central spines 0-4, one dominating (if present) porrect; 22 tubercles appressed ("pineapple"); 23 tubercles upright; 24 tubercles porrect; 25 central spines 0-1; 26 central spines always 3-4; 27 tubercles more than 12 mm wide at the base; 28 tubercles up to 12 mm wide at the base; 29 seeds globose, fruits turning red and drying soon

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