DAVE FERGUSON'S VIEW ON MAMMILLARIA CLASSIFICATION
writted in 09-2000 Slightly modified in wording, but
with no rearrangement on Oct. 20, 2004
Here are some observations and opinions on Mammillaria
classification.
David Hunt's classification of the genus has been the most
accepted
for a number of years now. Hunt's classification of Mammillaria has
been
modified a bit here and there over the years, but not much, and I don't
agree with many things about it.
At the beginning of the new Pilbeam book on Mammillaria,
Luthy's
classification of Mammillaria into subgenera, sections, and series can
be
seen as compared with that of Hunt. In my opinion Luthy is getting
closer
to a logical and natural classification than Hunt did. There are,
however,
some problems with Luthy's classification as well. Some of these are
pointed out by John Pilbeam at the beginning of his book; some seem
just
too ingrained in tradition to go away easily.
One of the problem areas in Luthy's classification is the
separation
of the series Longiflorae (sensu Hunt, plus a few additions) into two
sections called Krainzia and Archiebnerella. Both of these sections in
Luthy's scheme include species which are not at all related to the
Longiflorae (i.e. M. barbata, M. wrightii, M. albiflora, M. herrerae,
& M.
humboldtii of Mamillopsis and the Lasiacanthae). The remainder of his
Archiebnerella are much to closely related to one another to be split
into
separate sections and should be in the series Longiflorae of his
section
Krainzia (= Longiflorae). This would leave his Archiebnerella with no
members of its own! One species, M. aurilanata, which is very close to
M.
zephyranthoides, is still left entirely out of this relationship by
Luthy.
Another problem is very closely related, and involves the
scattering
of various species of Mamillopsis (i.e. Dolicothele & kin)
among several
series and sections, when they should (at least in my opinion) all be
brought together under one very distinct subgenus. These include the M.
barbata/wrightii complex (which is lumped in with the unrelated M.
tetrancistra and M. guelzowiana by Luthy), M. oteroi (lumped with the
unrelated series Sphacelatae), M. glassii (into the unrelated series
Bombycinae), M. senilis (which remains in its own section), and M.
beneckei
(which remains in its own subgenus). After checking dates of
descriptions
it turns out that Mammillopsis is the oldest subgeneric name for
Dolicothele. [sort of like the Thelocactus into the Leuchtenbergia
situation], so all the species of Dolicothele would become Mammillopsis.
I mostly like the subgenus Cochemiea as recognized by Luthy,
but some
of the series and sections are still too close to Hunt's arrangement.
Also,
Luthy splits the subgenera Cochemiea and Mammillaria in an unnatural
way,
and many species which should be included in subgenus Cochemiea are
listed
under subgenus Mammillaria as Luthy's section Cylindricothelae.
The series under the Cylindricothelae still show strong
influence from
Hunt's classification. The species "groups" under the various series
are
mostly fairly natural, but these "groups" are composed themselves in
unnatural ways. Thus, the series Stylothelae [just what is the type
species of the Stylothelae anyway??????], Lasiacanthae, Pectiniferae
(should be a subset of Bombycinae), Bombycinae, and Proliferae
need some serious rearranging. The series Sphacelatae should actually
be
moved to the Section Cochemiea where the series Ancistracanthae is most
closely similar.
M. pottsii, which is actually close to M. bombycina, is still
misplaced in the series Leptocladodae, but the remainder of the
Leptocladodae species are finally placed in a reasonable position near
the
Heterochlorae and Polyacanthae.
Here is my approach to Mammillaria. I am hoping to receive
constructive feedback and/or criticism.
This is not meant to be an official publication, but rather is
a
working list, with the goal being eventually a more natural arrangement
of
the genus Mammillaria than those which have been previously published.
For the various infrageneric groupings higher than species, in
a few
instances priorities may be violated. I have not fully researched
publication dates and synonymies of sections, series, nor species
groups,
so some names may be incorrect, or may exist but remain unknown to me.
I
am following most closely Hunt's sectional and series names in
preference
to those of Luthy. I believe Hunt's are more recognizable, and I
suspect
they are properly and legally applied. As I don't know the type species
of
some of the sections and series, I could have assigned names to an
improper
group of species (but I hope no such goofs were made).
I have listed the majority of recognized names. A few with
which I am
not very familiar are bound to be incorrectly assigned, but I think
I've
gotten most of them placed where they really belong. The synonymies may
seem sweeping, but I think in most cases are reasonable. Even if this
"lumping" is not agreed with, it serves to show very close kinship of
the
plants involved.
A) MAMMILLARIA subgenus MAMMILLARIA -
Seeds brown, roughly
teardrop-shaped, with usually linear lateral hilum, with testa cells
tabular pitted and bordered by a very strongly undulating raised line;
body
of plant usually with some milky sap at least toward core of plant.
Flower
and fruit is generally characteristic, but minor variations are often
useful in identifying or grouping plants.
I) sect. Mammillaria
stems generally depressed globose,
sometimes
hemispheric, with relatively few spines and tubercles (stem usually
easily
visible through spines), plant body with milky sap to surface.
a) series Mammillaria (=
Macrothelae)
1) M. mammillaris group - Includes M. mammillaris (incl. ?
ekmanii, simplex
, etc.), M. nivosa. Placement of M. nivosa here may be artificial?
2) M. heyderi group - Includes M. coahuilensis (incl.
albiarmata), M.
heyderi (= applanata, incl. bullingtoniana, gaumeri, hemispherica), M.
peninsularis, M. roseoalba (often misapplied to plants of M. magnimamma
type).
3) M. uncinata group - Includes M. brandegeei (incl. arida,
baxteriana,
bellisiana, bocensis, craigii, evermanniana, dawsonii, gabbii, gatesii,
glareosa, johnstonii, lewisiana, macdougalii, marksiana, marshalliana,
movensis, pacifica, petrophila, sonorensis, tesopacensis, etc.), M.
gummifera (incl. crassa, durangicola, grusonii, pachycilindrica,
papasquiarensis, tortulospina, wagneriana, zeyeriana, etc.), M.
uncinata
(incl. boelderliana).
4) M. petterssonii group - Includes M. gigantea (incl. ?
antesbergeriana,
armatissima, hastifera, ocotillensis, ? saint-pieana), M. orcuttii, M.
petterssonii (incl. apozolensis, fulvaflora, hamiltonhoytea,
huiguerensis,
pilensis, saltensis), M. rubrograndis.
b) series Polyedrae
1) M. magnimamma group - Includes M. carnea (incl. sartorii,
tenampensis), M. lloydii, M. magnimamma (incl. angularis, aramberri,
bernalensis, bucareliensis, centralifera, centricirrha, compressa,
crassimammillis, esseriana, macracantha, priessnitzii, rioverdense,
saxicola, ?seitziana, tolimensis, vagaspina, vallensis, zuccariniana,
etc.), M. melanocentra(incl. euthele, freudenbergeri, meiacantha, ?
parrasensis, runyonii,
winterae, zahniana), M. mystax (incl. atroflorens, casoi, crispiseta,
erythra, huajuapensis, mixtecensis, ? multiseta, variaculeata, etc.),
M.
polyedra.
2) M. karwinskiana group - Includes M. karwinskiana (incl.
confusa,
conzattii, ebenacantha, eichlamii, jozef-bergeri, ?knippeliana [nomen
dubium], ? multiseta, neomystax, praelii, strobilina, voburnensis,
etc.).
II) sect. Supertextae (= sect. Leucocephalae)
Stems mostlyhemispherical to conical, with many tubercles;
often much
wool in areoles;
spines usually obscuring plant body (except in M. sempervivi), radials
often nearly white, typically with two or four, usually short, often
darker centrals, not hooked, one pointing up and one down (but number
and arrangement varies some); fruits red, clavate.
a) series
Leucocephalae - Stems usually milky to
surface. More northern. - Includes M. chionocephala
(incl. caerulea, ritteriana), M. geminispina (incl. albata, brauneana,
bravoae, giseliana, hahniana, klissingiana, leucocentra, mendeliana,
werdermanniana, woodsii ), M. formosa (incl. arroyensis,
avila-camachoi, cadereytensis, ? celsiana, infernillensis,
leucocentra, morganiana, microthele, muehlenpfordtii, parkinsonii,
perbella, queretarica), M. sempervivi (incl. ?amajacensis,
pseudocrucigera), M. standleyi (incl. auricantha, auritricha,
bellacantha, canalensis, cobrensis, floresii, hertrichiana, laneusumma,
lanisumma, lindsayi, mayensis, miegiana, montensis, nolascana,
taylorianum, ? xanthina).
b) series Supertextae
- Stems usually milky only in core. More southern.
Includes M. columbiana (incl. bogotensis, chiapensis, hennesii,
ruestii,
soehlemannii, yucatanensis), M. crucigera (incl. buchenaui, grandinosa,
huitzilopochtli), M. pseudoperbella, M. supertexta (incl.
acultzingensis,
albidula, albilanata, collina, conspicua, dealbata, dixanthocentron,
?donatii, dyckiana, elegans, fauxiana, flavicentra, haageana, ignota
Repp.,
igualensis, juxtlahuacensis, kunthii, lanata, lanigera, longicaudata,
lupina, martinezii, meissneri, monticola, niduliformis, noureddineana,
oaxacana, palmillensis, reppenhagenii, san-angelensis, schmollii,
tegelbergiana, vaupelii), M. tlolocii.
III) sect. Subhydrochylus
Plants usually columnar, with numerous
tubercles, with sap milky usually only in core, spines usually slender,
smooth, and glassy; centrals often (not always) hooked; fruits usually
not
bright red, but varying shades of green through brown, pink, or
purplish,
often translucent. The series in this section are weakly defined. Seed
is
characteristically lighter in color and more strongly sculptured than
in
previous sections (though this needs further documentation, and the
trait
varies from series to series in the previous sections).
a) series Elongatae
(= Leptocladodia sensu Hunt, minus its type species M.
pottsii) - smallish, mostly cylindrical; spines not hooked. Includes M.
densispina (incl. buxbaumiana), M. elongata (incl. amgiomea. densa,
echinaria, echinata, intertexta, mieheana, rufrocrocea, stella-aurata,
straminea, subcrocea, tenuis, var. viperina [not M. viperina]), M.
eriacantha, M. microhelia (incl. droegeana sensu Repp., miroheliopsis).
b) series Polyacanthae
- larger, tending toward cylindrical (sometimes
subglobose), central spines often (not always) hooked. Distinction of
species in this series is confused and they could conceivably ALL be
the
same species! Includes M. guerreronis (incl. aguilensis, aureispina,
backebergiana, claviformis, compacticaulis, duoformis, ernestii,
erythrocalix, hamata, heeriana, magnifica, matudae, minor, recta,
rectiformis, rossiana, subhamata, serpentiformis, virginis,
xaltianguensis,
xuchiapensis, zapilotensis), M. rekoi (incl. albrechtiana, leptantha,
krasuckae, mitlensis, pseudorekoi, pulliamata, rekoiana, sanjuanensis),
M.
spinosissima (incl. auricoma, bella, centraliplumosa, crassior,
gasterantha, gracilis, haasii, kladiwae, ? machucae, meyranii, nunezii,
pilcayensis, pitcayensis, tepoxtlana).
c) series Heterochlorae
- larger yet, otherwise much like the
Polyacanthae, except usually hemispherical to subglobose stems; spines
not
hooked. Very close to and probably not deserving of a separate series
ranking from the Polyacanthae. Includes M. discolor (incl.
esperanzaensis,
halbingeri, longispina, multispina, ochoterenae, pachyrhiza,
schmollii), M.
rhodantha (incl. apamensis, aureiceps, bonavitii, clacantha,
fera-rubra,
fuscata, mccartenii, mollendorffiana, mundtii, parensis, pratensis,
pringlei, sulphurea, verticealba, wiesingeri), M. polythele (incl.
durispina, hidalgensis, hoffmanniana, kelleriana, kewensis,
neophaeacantha,
subdurispina, tetracantha, xochipillii).
d) series Decipientes
- includes M. decipiens (incl. albescens,
camptotricha).
B) MAMMILLARIA subgenus NOV. (= vetula group)
Seeds and fruit like
subg. Mamillopsis, but seeds with undulating line along margin of testa
cells more as in subg. Mammillaria. Flowers and plants similar to
Stylothelae and to M. decipiens. Doesn't fit neatly anywhere. Includes
M.
vetula (incl. gracilis, kuentziana, magneticola, pulchella), ? M.
schwarzii
.
C) MAMMILLARIA subgenus MAMILLOPSIS (= DOLICOTHELE)
Characterized by
the soft stems with a "chin" above the areole; fruits very juicy with
thin
translucent walls, usually rounded; seeds roughly teardrop-shaped,
mostly
more laterally compressed than in subgenus Mammillaria, brown to nearly
black with well developed pits in testa pattern cells, the pattern
cells
bordered by nearly straight raised lines. In M. beneckei the seeds are
very irregular, but still conform to these criteria.
I) sect. Barbatae
Distinguished by shorter flower tube, often fringed
sepaloids; petaloids usually not rich yellow, but some other shade.
a) series Barbatae
- With fimbriate tepal margins. Includes M. barbata
(incl. chavezei, garessii, meridiorosei, morricalii, orestera,
santa-clarensis, viridiflora, wilcoxii, etc.), M. wrightii (incl.
wolfii).
b) series
Carretiiae [ser. nov.] - With tepal margins not noticeably
fimbriate. Includes M. carretii (incl. saffordii), M. glassii (incl. as
censionis, nominus-dulcis, siberiensis), ? M. oteroi. M. oteroi is
perhaps
(?) better placed in sect. Dolicothele, but the small flowers with a
shorter tube indicate placement here.
II) sect. Dolicothele (incl. Oehmea)
Distinguished by flower mostly
fragrant; usually long flower tube; petaloids usually rich yellow.
Includes
M. baumii, M. beneckei (has similarities in flower to M. senelis), M.
longimamma, M. melaleuca, M. sphaerica, M. surculosa.
III) sect. Mamillopsis
Weakly defined from Dolicothele by zygomorphic
structure of flower with tube possessing obvious "scales" (i.e. fleshy
reduced sepaloids); and by the apparent lack of fragrance in the
flower;
flower yellow (rare), or near white (rare) to pink or red. Otherwise
the
same. Includes only M. senilis.
D) MAMMILLARIA subgenus COCHEMIEA
I have at times tentatively split the
hummingbird pollinated Cochemiea sensu strictu from the remaining
"Bartschella" species, but have never thought this a valid separation
at
this ranking. The separation would be more of a compromise to
tradition.
It has always seemed to me that the Ancistracanthae plus Bartschella
[sensu
strictu] plus Sphacelatae were more closely akin to Cochemiea than the
resultant group as a whole is to other Mammillaria series. However, the
Lasiacanthae does show some strong similarities to the Ancistracanthae,
and
provides something of a bridge or link to other groups.
I) sect. Krainzia (= Longiflorae)
a) series Krainzia
(= Longiflorae) - Includes M. aureilanata, M.
guelzowiana, M. heidiae, M. longiflora (incl. stampferi), ? M.
tepexicensis, M. tetrancistra, M. zephyranthoides.
b) series Napinae
- Includes M. deherdtiana (incl. dodsonii), ? M.
hernandezii, M. napina.
c) series Saboae
- Includes M. saboae (incl. goldii, haudeana), ? M.
sanchez-mejoradae, M. theresae.
II ) sect. Solisia
Seeds brown to black with basal hilum, with testa
cell dividing lines undulate as in subgenus Mammillaria.
a) series Solisia
(= Bombycinae & Leptocladodia) - In many ways
intermediate to subgenus Cochemiea and subgenus Mammillaria. Seed shape
and habitus is most like subgenus Cochemiea. Plants with hard bodies
and
round-apices on tubercles, often (always?) with traces of milky sap or
the
internal structures associated with it; flowers characteristic; fruits
dimorphic, some green, never lengthening and eventually drying between
tubercle bases; some fruits lengthening but usually not exceeding
spines,
sticky, and hard to remove entire, usually white to pink. Includes M.
bombycina (incl. perezdelarosae), M. carmenae (incl. dasyacantha,
lauii,
subducta), ? M. marcosii, M. mercadensis (incl. berkiana,
brachytrichion,
fuscohamata, gilensis, guillauminiana, jaliscana, kleiniorum,
rettigiana,
posseltiana), M. moelleriana (incl. cowperae), M. pectinifera, M.
pottsii,
M. solisioides. Widespread east from Sierra Madre Occidental, south to
Oaxaca.
b) series Stylothelae
(proper usage??; what is type species?) - Plants
with soft bodies; ??? milky sap, or not ???; flowers more delicate in
structure; fruits elongating, mostly shades of green, pink, or
purplish, in
some (? all) with characteristic odor (similar to many subgenus
Mammillaria
fruit in smell). Includes M. bocosana (incl. ?aurihamata, eschauzieri,
knebeliana, multilanata, splendens), M. erythrosperma (incl.
multiformis),
M. fittkaui (incl. ?limonensis), M. glochidiata, M. zeilmanniana. This
is
a tentative, perhaps unnatural grouping. Central Mexico.
III) sect. Lasiacanthae
(incl. most of Crinitae & Proliferae) - Close to
the section Cochemiea, but smaller, mostly depressed globose plants
with
mostly shorter flowers with capitate stigmata. Spines less often
hooked,
often "pubescent" to "plumose". Concentrated east of Sierra Madre
Occidental and north of Trans-volcanic region.
a) series Lasiacanthae
- Plants mostly with body firm, mostly depressed
globose; fruits normally bright shades of orange, red, or pink,
elongate,
firm, with basal pore; seeds roughly lightbulb-shaped (i.e. narrowed at
hilum end). Includes M. gasseriana (incl. chica, lengdobleriana,
magallanii, neobertrandiana, roseocentra, viescensis), M. herrerae
(incl.
albiflora), M. lasiacantha (incl. denudata, egregia), M. lenta, ? M.
luethyi, M. nazasensis, M. pennispinosa, M. sinistrohamata (incl.
zacatecasensis).
b) series Mammilloydia
- Plants with body firm; with spines in axillary
areoles; fruits spindle-shaped, mostly firm, green to pinkish or
lavender
(but very soft and bright red in M. ortiz-rubiona; elongate and bright
orange-red in M. stella-de-tacubaya); seeds unusual for sbg. Cochemiea,
slightly flattened and rather smooth, with hilum relatively small, with
pitting often reduced (M. humboldtii) to nearly absent (M. candida
& M.
ortiz-rubiona). Includes M. candida (incl. estanzuelensis), M.
ortiz-rubiona (separated due to distinct fruit, slightly distinct seed,
distinct gypsum habitat, and distinct appearance of plant in habitat),
M.
humboldtii, ? M. plumosa, M. stella-de-tacubaya. Another possibly
unnatural grouping, but all except M. plumosa are at least grossly very
similar.
c) series Proliferae.
Plants soft; spines less densely covering body;
fruits mostly red, often not exceeding spines, soft and insubstantial;
seeds as in series Lasiacanthae. Includes M. albicoma, M. crinita
(incl.
aureoviridis, ? aurihamata [all specimens I have seen under this name
were
M. bocosana with yellow spines], calleana, duwei, erectohamata,
felipensis,
leucantha, mathildae, moeller-valdeziana, monancistracantha, nana,
painteri, purberula, pubispina, M. scheinvariana, schelhasei, ?
seideliana,
tezontle, trichacantha, variabilis, wildii) M. picta (not very distinct
from M. crinita; incl. anniana, aurisaeta, rayonensis, sanluisensis,
schieliana, subtilis, unihamata, viereckii, weingartiana), M. prolifera
(incl. arachnoidea, ?glomerata, grisea, haitiensis, multiceps,
perpusilla,
pusilla, texana, zubleri), M. schiedenana (incl. dumetorum, giselae).
IV) sect. Cochemiea
Plants mostly with cylindrical bodies (sometimes
other shapes), often clustering; central spines typically hooked; fruit
mostly shades of bright orange, red, or pink, usually elongate and
slightly
clavate, often (? always) with an open basal pore when pulled off
plant;
seeds grey to black with a basal hilum, in most narrower at hilum end
and
somewhat lightbulb-shaped, pitted. Concentrated west of Sierra Madre
Occidental, and also (Sphacelatae) Oaxaca/Puebla area.
a) series Sphacelatae
- Flowers small, usually magenta to purple; fruit
and seeds as in series Ancistracanthae and Lasiacanthae; southern
distribution. Includes ? M. kraehanbuehill, M. sphacelata (incl.
tonalensis), M. viperina.
b) series Bartschella
- Plants with soft epidermis, soft, gray-green,
usually depressed globose to depressed conical when growing in habitat;
flowers with long narrow spreading stigmata; fruits with large basal
pore;
seeds large, rounded, with obvious secondary sculpturing in shallow
pits.
Includes M. boolii (incl. insularis), M. schumannii. Baja and Sonora.
c) series Ancistracanthae
- Plants mostly relatively small with
cylindrical branching stems, but in some depressed globose or
geophytic;
hooked central spines common, and normal in most species; flowers with
long
narrow spreading stigmata; fruits with small basal pore; seeds smaller
with
light bulb shape, and with pits deep and not obviously sculptured.
Includes M. albicana (incl. dolorensis, sleviinii), M. armillata (incl.
cerralboa), M. blossfeldiana (incl. goodridgei, rectispina, shurliana),
M.
capensis, M.
dioica (incl. angelensis, estabanensis, multidigitata, neopalmeri,
phitauiana), M. grahamii (incl. arizonica, auricarpa, microcarpa,
milleri,
oliviae), M. louisae (distinction from M. mazatlanensis needs further
study; plants seem quite different), M. mainiae, M. mazatlanensis
(distinction from M. grahamii is dubius; incl. alamensis, albissima,
bullardiana, diabloa, gueldemanniana, guirocobensis, hutchisoniana,
inae,
littoralis, marierana, monocentra, occidentalis, patonii, sheldonii,
sinaloensis, swinglei), M. thornberi (incl. fraileana, yaquensis). Wide
distribution mostly along Pacific Slope.
d) series Cochemiea
- As in Ancistracanthae, except plants usually
larger; flowers zygomorphic with longer hypanthium and usually red;
stigmata relatively long but often not spreading. Includes M. halei,
M.pondii (incl. maritima), M. setispina, M. poselgeri. Baja California.
Dave