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Bradl eya 18/2000 29

Summary: A pheneti c anal ysi s of the genus


Thelocactus i s presented. The col l ected data, 65,
and the 34 Operati onal Taxonomi c Uni ts used for
the study were processed by a pri nci pal compo-
nents anal ysi s method. The ordi nati ons made
l ead us to concl ude that Hamatocactus setispinus
i s not congeneri c wi th Thelocactus from whi ch i t
has to be separ ated. Some nomencl atur al
changes are al so made i n order to adjust the tax-
onomy of the genus to the resul ts obtai ned.
Zusammenfassung: Es wi rd ei ne phneti sche
Anal yse der Gattung Thelocactus vorgestel l t. Di e
fr di e Studi e gesammel ten Daten (65) und di e
34 operati onel l en taxonomi schen Ei nhei ten wer-
den ei ner Hauptkomponentenanal yse unterzo-
gen. Di e Ordi nati onen fhren uns zur Fol gerung,
dass Hamatocactus setispinus ni cht i n di e
Gattung Thelocactus gehrt und separi ert wer-
den muss. Ei ni ge nomenkl atori sche nderungen
bri ngen di e Taxonomi e der Gattung mi t den
gefundenen Resul taten i n Ei nkl ang.
Introduction
The genus Thelocactus, as we know i t nowadays,
was proposed by Bri tton and Rose i n 1922,
i ncl udi ng i n i t twel ve speci es that the ori gi nal
descri pti on of the genus uni tes for bei ng cacti of
medi um si ze wi th few, l ow or i ndi sti nct ri bs di vi d-
ed i nto tubercl es, a scal ed ovary, the frui ts dehi s-
cent by a basal pore and the seeds wi th a l arge
basal hi l um. The name Thelocactus appeared for
the fi rst ti me i n the l i terature i n the year 1898 i n
Gesamtbeschr ei bung der Kakteen, wher e
Schumann used i t as a name for a subgenus of
Echinocactus, arrangi ng i n i t, besi des those taxa
that now bel ong to Thelocactus, al so many other
speci es that l ater have been moved to other gen-
era.
After the work of Bri tton and Rose, many
other speci es have been added to the ori gi nal
group, both as new taxa and as new combi na-
ti ons. Some of the speci es combi ned i n
Thelocactus (T. saueri, T. knuthianus, T.
viereckii, T. subterraneus) have been l ater segre-
gated by Backeberg (1938, 1951) i n an al l i ed
genus, Gymnocactus, now i ncl uded i n
Turbinicarpus. I n 1978 Anderson, on the basi s of
the compari son of some characters, proposed to
i ncl ude the speci es bel ongi ng to Gymnocactus i n
Thelocactus, but i n 1986 he reassessed hi s previ -
ous proposal and put Gymnocactus i n Neolloydia,
extendi ng the l i mi ts of the l atter genus. The
Ei ghti es saw the descri pti on of some new taxa,
both at vari ety and speci es rank, and, sti l l by
Anderson (1987), a new del i mi tati on of the genus.
I n thi s r evi si on the author i ncl uded
Hamatocactus setispinus i n Thelocactus, ground-
i ng hi s proposal on the si mi l ari ti es observed
between Thelocactus bicolor and H. setispinus for
several of the characters he chose to compare
these two speci es. Thi s new del i mi tati on of the
genus Thelocactus l ed to a total upset of i ts l i m-
i ts, as two essenti al characters on whi ch Bri tton
and Rose based the genus, the presence of tuber-
cl es, a character to whi ch the generi c name
refers, and the basal l y dehi scent frui ts, suddenl y
fai l ed, H. setispinus l acki ng tubercl es and havi ng
fl eshy i ndehi scent fr ui ts. Fr om the ti me of
Andersons revi si on, other speci es have been
descri bed, Thelocactus garciae, Thelocactus mul-
ticephalus, Thelocactus panarottoanus and
Thelocactus flavus, whi l e others have been segre-
gated from i t.
Recentl y Dowel d (1998) has proposed to seg-
regate Thelocactus conothelos and i ts vari eti es
from Thelocactus and to i ncl ude them i n a new
genus, Torreycactus, the whol e on the basi s of just
the di fference i n the mi cromorphol ogy of the sec-
ondary scul pture of the seed cuti cl e that i n these
speci es i s smooth rather then mi cro-papi l l ate.
Anderson regarded the genus Thelocactus as con-
si sti ng of several l oosel y rel ated speci es and
speci es groups... The starti ng poi nt for our work
was thi s statement. We have tri ed to establ i sh
how and how much the di fferent taxa are i nter-
rel ated, wi th the ai m to veri fy i f the present sys-
temati c treatment i s congruent wi th the morpho-
Bradleya 18/2000
pages 29 54
A phenetic analysis of the genus Thelocactus
Alessandro Mosco
1
and Carlo Zanovello
2
1
Vi a Moreri 152, I -34135 Tri este, I tal y
2
Pi azza Mercato 9, I -36040 Brendol a (VI ), I tal y
Bradl eya 18/2000 30
T. bicolor ssp. bicolor, General Cepeda, Coahui l a.
Photo Jauerni g.
T. bicolor ssp. bicolor (commodus), Montemorel os, Nuevo
Len. Photo Jauerni g.
T. bicolor ssp. bolaensis (Pedri cena), San Pedro de l as
Col oni as, Coahui l a.
T. bicolor ssp. bolaensis (wagnerianus), Hi pol i to,
Coahui l a.
T. bicolor ssp. bicolor (schottii), SB567 Brewster Co.,
Texas.
T. bicolor ssp. bolaensis, Cerro Bol a, Coahui l a.
Photo Jantschgi .
Bradl eya 18/2000 31
T. bicolor ssp. flavidispinus, Marathon, Brewster Co.,
Texas. Photo Bercht.
T. bicolor ssp. flavidispinus, SB424 Brewster Co., Texas.
T. bicolor ssp. schwarzii, Cal l es, Tamaul i pas.
Photo Jauerni g.
T. buekii ssp. buekii, La Sol edad, Nuevo Len.
T. bicolor ssp. heterochromus, Ri o Nazas, Durango. T. bicolor ssp. heterochromus (pottsii), Hi dal go del Parral ,
Chi huahua. Photo Lausser.
Bradl eya 18/2000 32
l ogi cal data we have col l ected. I ndeed, there i s
sti l l not a consensus ei ther on the speci es to be
i ncl uded i n the genus (Dowel d, besi des excl udi ng
conothelos and i ts vari eti es, retai ns the monotyp-
i c genus Hamatocactus) or for the subspeci fi c
tr eatment for the taxa bel ongi ng to the
Thelocactus rinconensis compl ex (Ander son,
1999; Gl ass, 1997; Lthy, 1999, Mosco &
Zanovel l o, 1999). To reach thi s goal we have cho-
sen a pheneti c approach, based on the observa-
ti on of many morphol ogi cal characters. The phe-
neti c anal ysi s we made has al l owed us to reach,
we thi nk, a better comprehensi on of the studi ed
speci es and to formul ate some resul tant taxo-
nomi c proposal s.
Materials and methods
The majori ty of the data used i n our study have
been obtai ned by obser vati ons made on the
pl ants i n our col l ecti ons. Most of these pl ants are
provi ded wi th fi el d data and have been grown
from seed. Some data ori gi nate from observati ons
Figure 1. Pri nci pal components anal ysi s of Thelocactus. H. setispinus i ncl uded i n the ordi nati on.
Characters: stem shape I ; stem shape I I ; maxi mum stem di ameter; maxi mum stem hei ght; head number; epi dermi s
col our I ; epi dermi s col our I I ; epi dermi s col our I I I ; number of hypodermal l ayers; crystal s; seedl i ng shape; ri bs; orthos-
ti chy number; tubercl es; tubercl e or ri b wi dth; tubercl e or ri b hei ght; areol ar gl ands; mi ni mum total spi ne number;
maxi mum total spi ne number; central and radi al spi nes di sti ngui shabl e; hooked spi nes; spi ne col our I ; spi ne col our
I I ; spi ne col our I I I ; upper spi nes fl attened; shredded spi nes; mi ni mum fl ower di ameter; maxi mum fl ower di ameter;
mi ni mum petal oi d l ength/wi dth rati o; maxi mum petal oi d l ength/wi dth rati o; shape of the petal oi d apex; margi n of the
petal oi d apex; fl ower col our I ; fl ower col our I I ; fl ower col our I I I ; fl ower col our I V; styl e col our I ; styl e col our I I ; styl e
col our I I I ; sti gma col our I ; sti gma col our I I ; fi l ament col our I ; fi l ament col our I I ; pri mary fi l aments i nserti on; pol l en;
scented fl owers; earl y bud devel opment; fl oweri ng peri od; frui t col our; frui t dehi scence; frui t succul ence; seed average
di ameter; seed average l ength; l arge HMR; mi cropyl e i nsi de the hi l um; mi cropyl e on the edge of the hi l um; mi cropy-
l e outsi de the hi l um; appendages on the hi l um edge; funi cl e rests conspi cuous; testa cel l s shape; si nuate anti cl i nal
cel l wal l s; mi cro-papi l l ate seedcoat; stri ate seedcoat.
Factor axes 1 and 2
1.35E+00 OTUs: 34
Characters: 63
Cumulative variation
in the first two axes: 36%
+
+ * +
+* + ++
+
+ + + + +
+ *

-4.27E-01 ++ + 2.11E+00
axis 1




++*+

*+ BICOLORES
+
+





setispinus +

-1.12E+00
Bradl eya 18/2000 33
we made i n the fi el d duri ng our travel s i n Mexi co,
whi l e sti l l others have been taken from l i tera-
ture. Compl ete l i sts of the speci mens exami ned
al ong wi th the characters used and thei r scal i ng
are gi ven i n an appendi x. The materi al for l i ght
mi croscope anatomi cal studi es was fi xed i n for-
mal i n/aceti c aci d/ethanol (10:5:85) and then
processed for the usual paraffi n embeddi ng pro-
cedure. I t was secti oned at 30 m and stai ned
wi th haematoxyl i n-eosi n. Freshl y col l ected pol l en
has been embedded i n bal sam pri or to observa-
ti on. Seeds under goi ng Scanni ng El ectr on
Mi croscope (SEM) anal ysi s were washed wi th
di sti l l ed water, ai r dri ed and then gol d coated.
For SEM anal ysi s we used Phi l i ps 500 and Lei ca
Stereoscan mi croscopes. We used MULVA-4 pro-
Factor axes 1 and 2
OTUs: 33 9.19E-01
Characters:57
Cumulative variation
in the first two axes: 45%


lab
+
+ mul
+ rin
+ phy
+ hin
+ nid
+ loy
+ hex + fre
BICOLORES

kra+ mat+ + bue

+ + schm
-7.88E-01 * + leu 9.39E-01
+ + tul axis 1
+* +
++ + lau
+ mac



+ has

+ con
+ gar


flv ++ arg

+ aur

-7.61E-01
Figure 2. Pri nci pal components anal ysi s of Thelocactus. H. setispinus excl uded from the ordi nati on.
OTUs: arg = argenteus; aur = aurantiacus; bue = buekii; con = conothelos; fl v = flavus; fre = freudenbergeri; gar = gar-
ciae; has = hastifer; hex = hexaedrophorus; hi n = hintonii; kra = krainzianus; l ab = La Bol sa; l au = lausseri; l eu =
leucacanthus; l oy = lloydii; mac = macdowellii; mat = matudae; mul = multicephalus; ni d = nidulans; phy = phyma-
tothelos; ri n = rinconensis; schm = schmollii; tul = tulensis.
Characters: stem shape I ; stem shape I I ; maxi mum stem di ameter; maxi mum stem hei ght; head number; epi dermi s
col our I ; epi dermi s col our I I ; epi dermi s col our I I I ; number of hypodermal l ayers; seedl i ng shape; ri bs; orthosti chy
number; tubercl e shape; grooved tubercl es; tubercl e or ri b wi dth; tubercl e or ri b hei ght; areol ar gl ands; mi ni mum total
spi ne number; maxi mum total supi ne number; central and radi al spi nes di sti ngui shabl e; spi ne col our I ; spi ne col our
I I ; spi ne col our I I I ; upper spi nes fl attened; shredded spi nes; mi ni mum fl ower di ameter; maxi mum fl ower di ameter;
mi ni mum petal oi d l ength/wi dth rati o; maxi mum petal oi d l ength/wi dth rati o; shape of the petal oi d apex; margi n of the
petal oi d apex; fl ower col our I ; fl ower col our I I ; fl ower col our I I I ; fl ower col our I V; styl e col our I ; styl e col our I I ; styl e
col our I I I ; sti gma col our I ; sti gma col our I I ; fi l ament col our I ; fi l ament col our I I ; pri mary fi l aments i nserti on; pol l en;
earl y bud devel opment; fl oweri ng peri od; seed average di ameter; seed average l ength; l arge HMR; mi cropyl e i nsi de
the hi l um; mi cropyl e on the edge of the hi l um; mi cropyl e outsi de the hi l um; appendages on the hi l um edge; funi cl e
rests conspi cuous; testa cel l s shape; si nuate anti cl i nal cel l wal l s; mi cro-papi l l ate seedcoat.
Bradl eya 18/2000 34
T. buekii ssp. matudae, Gal eana, Nuevo Len. T. conothelos ssp. conothelos, Dr Arroyo, Nuevo Len.
T. conothelos ssp. flavus, El Hui zache, San Lui s Potos .
Photo Nodari .
T. conothelos ssp. garciae, Bustamante, Tamaul i pas.
T. conothelos ssp. argenteus, Ascensi on, Nuevo Len. T. conothelos ssp. aurantiacus, La Escondi da,
Nuevo Len.
Bradl eya 18/2000 35
T. hastifer, east of Cadereyta, Quertaro. T. hexaedrophorus, Si erra El Azul , Nuevo Len.
T. leucacanthus ssp. leucacanthus, Mesa del Len,
Quertaro.
T. leucacanthus ssp. schmollii, Vi zarrn, Quertaro.
T. hexaedrophorus (lloydii), SB113 Sal i nas,
San Lui s Potos .
T. lausseri.
Bradl eya 18/2000 36
gramme for the mul ti vari ate anal ysi s. Character
vectors were normal i sed and the resembl ance
matri x was obtai ned by a si mi l ari ty al gori thm.
The ordi nati on techni que used was a pri nci pal
components anal ysi s (PCA).
Results
From the ordi nati on pl ot (Fi gure 1) obtai ned
usi ng al l the Oper ati onal Taxonomi c Uni ts
(OTU), i t becomes i mmedi atel y evi dent that
Hamatocactus setispinus di verges strongl y from
al l other OTUs. I t di ffers from the other taxa i n
l acki ng crystal s and tubercl es, for havi ng hooked
central spi nes, scented fl owers, bri l l i ant red,
fl eshy, i ndehi scent frui ts and smal l er seeds wi th
a stri ate seedcoat.
I n the next ordi nati on made excl udi ng H.
setispinus, the defi ni ti on of the pheneti c struc-
ture of the genus Thelocactus i s much better
(Fi gure 2). Two OTU groups separate neatl y from
the other enti ti es of the genus: one i s made up of
Thelocactus conothelos and i ts subspeci es togeth-
er wi th Thelocactus garciae and Thelocactus
flavus, the other by Thelocactus bicolor and al l i ed
taxa. The taxa bel ongi ng to the T. conothelos
group di ffer from al l the other Thelocactus i n two
characters, the pecul i ar fl ower tube morphol ogy
that i n these speci es bears the i nserti on of the
pri mary fi l aments wel l above the basi s of the nec-
tar chamber, and the mi cromorphol ogy of the
Factor axes 1 and 2
1.47E+00 OTUs: 10
Characters: 30
Cumulative variation
in the first two axes: 53%









+ flavidispinus



+ schwarzii

+ commodus

+ schottii

+ bicolor
-9.52E-01 + pottsii 1.43E+00
axis 1
+ pedricena

+ wagnerianus


heterochromus +


+ bolaensis

-8.41E-01
Figure 3. Pri nci pal components anal ysi s of the bicolor compl ex.
Characters: stem shape I ; maxi mum stem di ameter; maxi mum stem hei ght; head number; epi dermi s col our I ; orthos-
ti chy number; tubercl e or ri b wi dth; tubercl e or ri b hei ght; areol ar gl ands; mi ni mum total spi ne number; maxi mum
total spi ne number; spi ne col our I ; spi ne col our I I ; upper spi ne fl attened; mi ni mum fl ower di ameter; maxi mum fl ower
di ameter; mi ni mum petal oi d l ength/wi dth rati o; maxi mum petal oi d l ength/wi dth rati o; shape of the petal oi d apex;
margi n of the petal oi d apex; styl e col our I I ; sti gma col our I ; sti gma col our I I ; fi l ament col our I I ; fl oweri ng peri od; seed
average di ameter; seed average l ength; mi cropyl e i nsi de the hi l um; mi cropyl e outsi de the hi l um; appendages on the
HMR.
Bradl eya 18/2000 37
seeds whose testa cel l s are coni cal wi th a smooth
surface. Thelocactus garciaei s very si mi l ar, mor-
phol ogi cal l y, to T. conothelos, fr om whi ch i t
di verges by the di fferent shape of the tubercl es,
that are rounded, whereas i n conothelos, at l east
when young, they are angl ed; by the epi dermi s
col our, but, mai nl y, by the mi cromorphol ogy of the
seeds that i n thi s speci es have not such a l arge
hi l um mi cropi l ar regi on as i n conothelos and
i nstead have conspi cuous funi cl e rests. I n thi s i t
si mi l ar to Thelocactus tulensis, the onl y other
speci es, al ong wi th T. flavus, to have thi s charac-
ter. Thelocactus flavus i s the l atest speci es
bel ongi ng to thi s group. I t i s a cl ose al l y of T. gar-
ciae, from whi ch i t di ffers i n the cl usteri ng habi t,
the di fferent spi ne number and the fl ower col our.
The bicolor group i s the ri chest i n enti ti es and
to assess the rel ati onshi ps i nsi de i t we made a
Factor axes 1 and 2
OTUs: 18 1.34E+00
Characters: 47
Cumulative variation
in the first two axes: 45%



+ kra






+ leu
+ schm




+ hex
+ lab + loy


mul ++ hin
rin + + + fre 1.46E+00
-6.84E-01 phy + mat axis 1

+ nid + bue



+ tul

has +

+ lau
+ mac

-7.45E-01
Figure 4. Pri nci pal components anal ysi s of Thelocactus: bicolor and conothelos compl exes excl uded from the ordi -
nati on.
OTUs: bue = buekii; fre = freudenbergeri; has = hastifer; hex = hexaedrophorus; hi n = hintonii; kra = krainzianus; l ab
= La Bol sa; l au = lausseri; l eu = leucacanthus; l oy = lloydii; mac = macdowellii; mat = matudae; mul = multicephalus;
ni d = nidulans; phy = phymatothelos; ri n = rinconensis; schm = schmollii; tul = tulensis.
Characters: stem shape I ; stem shape I I ; maxi mum stem di ameter; maxi mum stem hei ght; head number; epi dermi s
col our I ; epi dermi s col our I I ; epi dermi s col our I I I ; number of hypodermal l ayers; seedl i ng shape; ri bs; orthosti chy
number; tubercl e shape; grooved tubercl es; tubercl e or ri b wi dth; tubercl e or ri b hei ght; areol ar gl ands; mi ni mum total
spi ne number; maxi mum total spi ne number; central and radi al spi nes di sti ngui shabl e; spi ne col our I ; spi ne col our
I I ; spi ne col our I I I ; shredded spi nes; mi ni mum fl ower di ameter; maxi mum fl ower di ameter; mi ni mum petal oi d
l ength/wi dth rati o; maxi mum petal oi d l ength/wi dth rati o; margi n of the petal oi d apex; fl ower col our I ; fl ower col our
I I ; fl ower col our I I I ; styl e col our I ; styl e col our I I ; fi l ament col our I I ; pol l en; earl y bud devel opment; fl oweri ng peri od;
seed average di ameter; seed average l ength; mi cropyl e i nsi de the hi l um; mi cropyl e on the edge of the hi l um; mi cropy-
l e outsi de the hi l um; appendages on the hi l um edge; funi cl e rests conspi cuous; testa cel l s shape; si nuate anti cl i nal
cel l wal l s.
Bradl eya 18/2000 38
T. leucacanthus ssp. schmollii (krainzianus), Pea Mi l l er,
Quertaro.
T. macdowellii, Hi gueras, Coahui l a.
T. multicephalus, Sandi a, Nuevo Len. T. multicephalus (La Bol sa), El Desi erto, Nuevo Len.
separate ordi nati on (Fi gure 3). The taxa of thi s
compl ex di ffer from the other taxa of the genus i n
two characters: the fl ower that has al ways a more
or l ess deep red throat, and the pol l en whose
grai ns are pol ycol pate, a character that we fi nd
el sewhere onl y i n Thelocactus macdowellii. The
PCA resul ts poi nt out the affi ni ty of T. hete-
rochromus wi th T. bicolor, so that i t stands apart
from al l the enti ti es of the bicolor compl ex.
Heterochromus di ffers from bicolor i n the stem
shape, whi ch i s depressed, by havi ng more robust
spi nes and for the absence of areol ar gl ands. Lack
of areol ar gl ands i s a character common al so to
other popul ati ons, scattered through Durango
and Chi huahua, that have been ti l l now consi d-
ered bicolor, whi ch they morphol ogi cal l y resem-
bl e, but from whi ch they di ffer i n havi ng rather
more robust spi nes. We have associ ated, i n our
anal ysi s, these popul ati ons wi th the pl ants di s-
tri buted by Brack as Thelocactus bicolor var.
pottsii. These or i gi nate fr om an ar ea near
Ji menez, i n the state of Chi huahua, and have an
i nteresti ng feature: they have the l ower central
spi ne hooked. We fol l owed Brack i n the use of the
epi thet pottsii to name thi s i ntermedi ate form,
because, wi th regard to the spi ne number and
stoutness, they fi t the descri pti on by Sal m-Dyck.
I nsi de the bicolor group three OTUs al l segregate
together: bolaensis, wagnerianus and Pedri cena.
Thi s l ast i ncl udes some enti ti es di stri buted by
Brack as red-spi ned bolaensis and nati ve of
Coahui l a, between San Pedro and Boqui l l as. Al l
these three enti ti es are uni ted, and di sti ngui sh
from bicolor, for havi ng a cyl i ndri cal stem, often
sl i ghtl y caespi tose, wi th 813 ri bs and a greater
radi al spi ne number.
Leavi ng out the ordi nati on the groups of
bicolor and conothelos, we reached an opti mum
Bradl eya 18/2000 39
T. rinconensis ssp. rinconensis, Ramos Ari zpe, Coahui l a. T. rinconensis ssp. rinconensis (phymatothelos), Arteaga,
Coahui l a.
T. rinconensis ssp. freudenbergeri. T. rinconensis ssp. hintonii, Rayones, Nuevo Len.
Photo Jantschgi .
T. rinconensis ssp. nidulans, Si erra Pai l a, Coahui l a. T. tulensis, El Hui zache, San Lui s Potos .
Bradl eya 18/2000 40
resol uti on of the remai ni ng OTUs (Fi gure 4). Two
cl usters are recogni sabl e here: the fi rst i s formed
by Thelocactus rinconensis and subspeci es,
Thelocactus hexaedrophorus and subspeci es,
Thelocactus multicephalus, Thelocactus buekii
and Thelocactus matudae; the other by
Thelocactus leucacanthus and subspeci es. The
l atter group of three rel ated enti ti es i s made up
by leucacanthus, schmollii and krainzianus
sensu Nagl . The OTUs that are not i ncl uded
i nsi de any cl uster are formed by four speci es:
Thelocactus lausseri, Thelocactus hastifer,
Thelocactus macdowellii and Thelocactus tulen-
sis, and are not cl osel y rel ated to any other
speci es.
The group wi th the greatest number of OTUs
i s formed by rinconensis, nidulans, freudenberg-
eri, hintonii, phymatothelos, multicephalus, La
Bol sa, hexaedrophorus, lloydii, buekii and matu-
dae, and to obtai n a good resol uti on i nsi de i t, we
made a further ordi nati on wi th onl y these OTUs.
I n the rel ated pl ot (Fi gure 5) these OTUs sepa-
rate i nto four cl usters. The l argest i s formed by T.
rinconensis and i ts subspeci es, that share a
uni que char acter : they have a mul ti -l ayer ed
hypodermi s, a feature typi cal of these enti ti es
and present onl y i n thi s compl ex i n the whol e
genus. T. multicephalus, i n i ts more southern
Figure 5. Pri nci pal components anal ysi s of Thelocactus: the group formed by buekii, hexaedrophorus, multicephalus
and rinconensis resol ved.
OTUs: bue = buekii; fre = freudenbergeri; hex = hexaedrophorus; hi n = hintonii; l ab = La Bol sa; l oy = lloydii; mat =
matudae; mul = multicephalus; ni d = nidulans; phy = phymatothelos; ri n = rinconensis.
Characters: stem shape I ; maxi mum stem di ameter; maxi mum stem hei ght; head number; epi dermi s col our I ; epi -
dermi s col our I I ; epi dermi s col our I I I ; number of hypodermal l ayers; ri bs; orthosti chy number; tubercl e shape; tuber-
cl e or ri b wi dth; tubercl e or ri b hei ght; mi ni mum total spi ne number; maxi mum total spi ne number; central and radi -
al spi nes di sti ngui shabl e; spi ne col our I I ; spi ne col our I I I ; shredded spi nes; mi ni mum fl ower di ameter; maxi mum
fl ower di ameter; mi ni mum petal oi d l ength/wi dth rati o; maxi mum petal oi d l ength/wi dth rati o; margi n of the petal oi d
apex; fl ower col our I ; fl ower col our I I ; fl ower col our I I I ; styl e col our I ; styl e col our I I ; fl oweri ng peri od; seed average
di ameter; seed average l ength; mi cropyl e i nsi de the hi l um; mi cropyl e on the edge of the hi l um; mi cropyl e outsi de the
hi l um; appendages on the hi l um edge; testa cel l s shape.
6.60E-01

+ mat + bue


+ nid




+ fre

hin +


-1.01E+00 phy ++ rin 7.84E-01
axis 1



mul +




+ loy lab +









Factor axes 1 and 2
OTUs: 11
+ hex Characters: 37
Cumulative variation
-1.08E+00 in the first two axes: 42%
Bradl eya 18/2000 41
forms (OTU La Bol sa) that usual l y have si ngl e
depressed stems, resembl es a rinconensis, but
di ffers from i t by two characters: the l ack of ri bs
and the presence of onl y one hypodermal l ayer
(Fi gure 6), whi l e rinconensis and al l i ed taxa
al ways have the stem di vi ded i nto ri bs and two or
three hypodermal l ayers (Fi gure 7). Thelocactus
hexaedrophorus i s di stri buted over a rather l arge
area and has numerous l ocal forms sl i ghtl y di f-
feri ng from each other and al l i ncl uded i n the
OTU hexaedrophorus. The pl ants ori gi nati ng
from the area near Fresni l l o have been i denti fi ed
by Anderson as the Thelocactus lloydii of Bri tton
and Rose, so we i ncl uded them i n the OTU lloy-
dii. The l ast two taxa we have taken i nto consi d-
erati on for thi s ordi nati on are Thelocactus buekii
and Thelocactus matudae.
Discussion
The pheneti c structure of the genus Thelocactus,
as i t appears from the PCA, does not match that
proposed by Anderson (1986) i n hi s revi si on of
the genus. Thi s i s due to the fact that we have
used for our study a much greater number of
characters, 65 versus 19, to establ i sh the rel a-
ti onshi ps i nsi de the genus. From the compari son
made by Anderson among Hamatocactus setispi-
nus, Thelocactus bicolor and Thelocactus hexae-
drophorus, i t emerged that setispinus and bicolor
share ten of the ni neteen characters used, whi l e
onl y four are common to setispinus and hexae-
drophorus. The scanti ness of the shared charac-
ters between H. setispinus and T. hexaedropho-
rus (that i s, the type speci es of the genus) made
Anderson uncertai n i f to i ncl ude setispinus i n
Thelocactus or not. Eventual l y H. setispinus was
i ncl uded i n Thelocactus, suppressi ng the mono-
typi c genus Hamatocactus, as consi dered troubl e-
some, but i n the meanti me maki ng the del i mi ta-
ti on of Thelocactus awkward.
We thi nk that Anderson has underesti mated
some characters that are basi c to the ci rcum-
scri pti on of thi s genus, namel y the presence of
tubercl es and a semi -fl eshy frui t dehi sci ng at
maturi ty. The seed shape, pyri form and wi th a
basal hi l um, excl udes any rel ati onshi p of setispi-
nus wi th Ferocactus, a rel ati onshi p once uphel d
by Benson (1982), but advocates that i t bel ongs to
the same phyl ogeneti c l i ne as Thelocactus.
Accordi ng to us H. setispinus, for the si ze of the
seeds, that are smal l er than those of Thelocactus,
and for the secondary scul pture of the seedcoat,
stri ate rather than mi cro-papi l l ate, i s, cl oser
al l i ed to Turbinicarpus, whose seeds have a si m-
i l ar si ze and, i n the majori ty of the speci es, a stri -
ate seedcoat. An anal ogous stand has been
al ready taken by Dowel d i n 1998, wi th the i ncl u-
si on of Hamatocactus i n hi s new subtr i be
Turbi ni carpi nae to whi ch Thelocactus does not
bel ong. To concl ude, i n vi ew of the PCA resul ts
and of the morphol ogi cal di fferences regardi ng
some characters presence of tubercl es, frui t
morphol ogy and seed mi cromorphol ogy, accordi ng
to us fundamental for the ci rcumscri pti on of the
genus Thelocactus we thi nk that H. setispinus
i s not to be i ncl uded i n Thelocactus and therefore
the monotypi c genus Hamatocactus i s to be
retai ned.
The cuti cl e mi cromorphol ogy of Thelocactus
conothelos and i ts subspeci es l ed Dowel d (1998)
to propose a new genus, Torreycactus, for them,
pl aci ng i t i n a di sti nct phyl ogeneti c l i ne, together
wi th Kadenicarpus and Bravocactus, two other
new monotypi c genera, i ncl udi ng i n the fi rst
Turbinicarpus horripilus and i n the second
Turbinicarpus pseudomacrochele, both havi ng
seeds wi th a smooth cuti cl e. Our opi ni on i s that
segr egati ng conothelos fr om the genus
Thelocactus onl y on the basi s of thi s si ngl e char-
acter i s wrong. Moreover, when al l the other char-
acters are congruent wi th those of the other
Thelocactus speci es, we do not agr ee wi th
Figure 6. Outer cel l l ayers of Thelocactus multi-
cephalus, HO 809 Sandi a, Nuevo Len, onl y one hypo-
dermal l ayer i s present. The bar 100 m l ong.
Figure 7. Outer cel l l ayers of Thelocactus rinconensis
ssp. freudenbergeri, CSD173 Grutas de Garci a, Nuevo
Len, a several l ayered hypodermi s i s present.
Bradl eya 18/2000 42
Dowel ds proposal .
The typi cal form of T. conothelos ssp. auranti-
acus i s morphol ogi cal l y suffi ci entl y di sti nct from
the type speci es, both for body shape and spi na-
ti on, but there are popul ati ons, si tuated at the
begi nni ng of the Aramberri val l ey and more
southerl y too, that we have observed near La
Escondi da and Zaragoza, that are morphol ogi cal -
l y i denti cal to the ssp. conothelos, the sol e di ffer-
ence bei ng the fl ower col our that vari es from a
deep yel l ow to orange i n the popul ati ons near
Zaragoza. We thi nk that these enti ti es fal l wi thi n
the normal vari abi l i ty of thi s subspeci es that di f-
fers from the type speci es onl y for the di fferent
fl ower col our. I t i s our opi ni on that thi s character
i s suffi ci ent to mai ntai n as di sti nct these two
taxa as the pi gments that account for the two
col ours, betaxanthi ns for the yel l ow and beta-
cyani ns for the magenta, are the product of two
di fferent bi osyntheti c pathways (Gi bson & Nobel ,
1986). Of the subspeci es argenteus are known
onl y some popul ati ons nati ve of the Ascensi on
val l ey. They are al l morphol ogi cal l y si mi l ar wi th
each other and di sti nct from the type speci es for
the spi nati on, that i s denser, and for the di fferent
col our of the central spi nes, whi te i nstead of grey-
i sh.
The mor phol ogi cal di ffer ences between T.
conothelos and T. garciaeare scanty and the di f-
ferences founded i n seed mi cromorphol ogy seem
not suffi ci ent to justi fy the mai ntenance of garci-
ae, that i n the l atest CI TES Cactaceae Checkl i st
has been l i sted as provi si onal l y accepted at the
speci es rank, so we thi nk that i s more appropri -
ate to treat i t as a subspeci es of conothelos. Al so
i f flavus di ffers from conothelos for some more
characters, spi ne number and the cl usteri ng
habi t, than garciae does, due to i ts cl ose rel ati on-
shi p wi th garciae based on seed mi cromorphol o-
gy, i t has to be treated as a subspeci es of conoth-
elos too. A taxon that can be referred to thi s group
i s Thelocactus panarottoanus Hal da. Hal da
(1998), i n hi s descri pti on ful l of gaps and not
accompani ed by any photograph, compared thi s
speci es wi th Thelocactus rinconensis and T.
conothelos. The i ndi cati ons of the si mi l ari ty of the
seeds of T. panarottoanus wi th those of T. conoth-
elos, al ong wi th the fl ower col our and the generi c
type l ocal i ty, make us suppose that i t i s conspe-
ci fi c wi th T. flavus. The author al so states i t i s
rel ated to T. rinconensis. Therefore T. panarot-
toanus i s compri sed i n the same OTU of T. flavus,
and the two epi thets are synonymous.
Thelocactus bicolor i s the speci es wi th the
greatest di stri buti on area that extends from
Texas, north, to San Lui s Potos , south, and from
Tamaul i pas, east, to Durango, west. I t i s then
natural to fi nd popul ati ons that di ffer from each
other more or l ess neatl y and that many of them
have been formal l y descri bed. Of al l the vari eti es
once descr i bed, today onl y two ar e for mal l y
accepted, al l the others have i ncreased the syn-
onym l i st of T. bicolor. Moreover the val i di ty of
the systemati c posi ti on of Thelocactus heterochro-
mus, at speci es rank, has been recentl y ques-
ti oned, thi s taxon bei ng l i sted as provi si onal l y
accepted i n the new CI TES Cactaceae Checkl i st.
Due to the weak di fferences, depressed stem,
l ack of areol ar gl ands and stronger spi nes, found
between heterochromus and bicolor, we thi nk
that i t i s not justi fi ed to use speci es rank for het-
erochromus, but that i t shoul d be consi dered a
subspeci es of bicolor. We bel i eve that the enti ti es
i ncl uded i n the OTU pottsii, for the l ack of areo-
l ar gl ands, are more cl osel y al l i ed wi th hete-
rochromus than wi th bicolor, so the l i mi ts of the
former shoul d be extended to i ncl ude al so these
forms. Thi s i s, at present, the best choi ce, wai ti ng
for more data on the di stri buti on of these enti ti es
and of heterochromus i n thi s area, data that
ascertai n or not the opportuni ty to create a new
subspeci es to accommodate those enti ti es mor-
phol ogi cal l y i ntermedi ate between bicolor and
heterochromus. Anderson (1987) di d not consi der
i t appropri ate to recogni se ei ther bolaensis or
wagnerianus at vari etal rank, i ncl udi ng both i n
the synonym l i st of T. bicolor.
We dont agree wi th thi s l i ne, but bel i eve that
these three enti ti es shoul d be, i nstead, i ncl uded
i n a subspeci es of bicolor. Thi s woul d be congru-
ent wi th the taxonomi c treatment reserved for
two other enti ti es, flavidispinus and schwarzii,
whose taxonomi c rank we feel di sposed to accept,
but that surel y are not pheneti cal l y so far from
bicolor as are bolaensis or wagnerianus. To con-
cl ude the anal ysi s of the bicolor compl ex, we have
to comment on two more OTUs: schottii and com-
modus. Wi th the name schottii are i denti fi ed
some Texas popul ati ons whose pl ants have the
characteri sti c of beari ng a fl attened, upper radi al
spi ne up to 7 cm l ong, whi l e commodus i s
descri bed as havi ng onl y one central spi ne.
At thi s poi nt i t i s necessary to open a paren-
thesi s. I n T. bicolor, as i n many other speci es of
the genus, the di scri mi nati on between central
and radi al spi nes i s awkward. Typi cal l y bicolor
has three central spi nes, of whi ch the l owest i s
the l ongest and porrect, whi l e the other two are
erect, general l y sl i ghtl y more robust and l onger
than the radi al s, wi th whi ch they can often be
mi staken by bei ng appressed agai nst the stem as
the radi al s are. The typi cal upper spi ne, fl attened
and erect, has been consi dered by us as a radi al
spi ne. Based on these premi ses, commodus has
three central spi nes, as does the type speci es, and
the mai ntenance of thi s vari ety or of var. schottii,
whose onl y di fference wi th bicolor consi sts i n the
l ong upper spi ne, i s not justi fi ed.
Bradl eya 18/2000 43
T. leucacanthus and subsp. schmollii di ffer
onl y i n the fl ower col our: yel l ow i n the former and
magenta i n the l atter. As for T. conothelos subsp.
aurantiacus, we thi nk thi s character i s suffi ci ent
to mai ntai n these two taxa separate. For many
years the i denti ty of Thelocactus krainzianus has
remai ned uncertai n, and after the descri pti on of
Thelocactus matudae i t has been thought possi -
bl e to i denti fy krainzianus wi th i t (Chvastek,
1985; Anderson, 1987). Nagl , i n 1991, after he vi s-
i ted some leucacanthus popul ati ons i n the area
near Pea Mi l l er, Quertaro, bel i eved that i t was
possi bl e to i denti fy these pl ants wi th the
krainzianus of Oehme.
The pl ants of thi s area fi t wel l the descri pti on
gi ven by Oehme, have 8 ri bs, are cl usteri ng and,
above al l , have the tubercl es cross-grooved at the
base, characters that do not bel ong to matudae.
The presence of a cross-groove on the tubercl es i s
a typi cal character for thi s enti ty, that we found
between Pea Bl anca and Pea Mi l l er, but can
occasi onal l y appear both i n leucacanthus and
schmollii. The l atter taxon we came across a l i t-
tl e north of Vi zarron, di fferi ng from krainzianus
al so i n the si ze of the tubercl es, that are smal l er,
and i n the greater number of radi al spi nes. We
bel i eve that these di fferences are not suffi ci ent to
separate krainzianus from schmollii, as Nagl has
just done, pr oposi ng the r ank of for ma for
krainzianus. Therefore we thi nk i t i s more appro-
pri ate to consi der these two taxa synonyms,
mai ntai ni ng the name schmollii that i s more
wi despread.
Al l the taxa bel ongi ng to the rinconensis
group are cl earl y cl osel y rel ated, but ti l l now a
consensus has not been reached on the accep-
tance of subspeci es r ank for these enti ti es.
Anderson (1999) thi nks i t i s not justi fi ed to sub-
di vi de T. rinconensis i nto i nfr aspeci fi c taxa
because there i s a conti nuum from very spi ny to
spi nel ess forms. Gl ass (1997), i nstead, bel i eved i t
appropri ate to rai se nidulans and phymatothelos
to subspeci fi c l evel . As regards freudenbergeri
and nidulans we feel i t i s justi fi ed to use sub-
speci es rank as they di ffer si gni fi cantl y from rin-
conensis. T. nidulans i s wel l di sti ngui shabl e for
havi ng a greater number of spi nes, di vi ded i nto
central and radi al spi nes, and freudenbergeri
both for havi ng a greater spi ne count, wi th cen-
tral and radi al spi nes di sti nct, and for the di ffer-
ent fl ower col our that i n thi s enti ty i s magenta.
We can say very l i ttl e about T. rinconensis
ssp. hintonii, as we know thi s taxon onl y from the
l i terature and so the data we have are i ncom-
pl ete. Our acceptance of thi s enti ty at subspeci es
rank i s based, above al l , on the di fferent fl ower
col our, yel l ow accordi ng to the descri pti on. I f thi s
character proves i n the future to be i nconsi stent
(i n the photographs publ i shed al ong wi th the
descri pti on the yel l ow col our of the fl ower i s
rather fai nt), then thi s enti ty, whose spi ne num-
ber i s vari abl e from 3 to 10, bei ng so i ntermedi ate
between rinconensis and subspeci es freudenberg-
eri and nidulans, shoul d be treated as a l ocal
form of rinconensis and the status of the other
subspeci es, whose acceptance i s based mai nl y on
the spi ne count, shoul d be re-eval uated. Among
the vari ous taxa that form thi s compl ex we have
been i n doubt onl y on the acceptance of phyma-
tothelos, whi ch i s the enti ty that separates l east
from rinconensis, di sti ngui shed from i t for havi ng
a more depressed stem and the spi nes general l y
shorter and recurved. We have chosen, eventual -
l y, to fol l ow Anderson i n consi deri ng i nappropri -
ate the recogni ti on of phymatothelos at sub-
speci es rank, as the di fferences found are too
weak.
We do not agree wi th the proposal by Lthy to
i ncl ude Thelocactus multicephalus i n rinconen-
sis, because, i f multicephalus morphol ogi cal l y
resembl es a rinconensis, i t di ffers from i t by hav-
i ng i ndi sti nct ri bs and onl y one hypodermal l ayer
whi l e al l the enti ti es bel ongi ng to the rinconensis
group have a mul ti -l ayered hypodermi s. Thi s l at-
ter character i s for us suffi ci ent to separate these
two taxa. T. multicephalus has a di stri buti on
area much l arger than that reported by Lthy: i t
actual l y extends south as far as Matehual a. The
type form can be found near Sandi a, Nuevo Len,
whi l e i n the mor e souther l y l ocal i ti es i t i s
repl aced by forms wi th a si ngl e depressed stem
wi th shorter spi nes that we have i ncl uded i n the
OTU La Bol sa and that we bel i eve not suffi -
ci entl y di sti nct to justi fy recogni ti on at i nfraspe-
ci fi c rank.
The ori gi nal descri pti on of T. hexaedrophorus
reports that thi s speci es bears one central spi ne,
whose absence, i n the majori ty of the popul ati ons
we observed i n habi tat, i s constant. Thi s i s not
true for the pl ants comi ng from Sal i nas, San Lui s
Potos , (SB113) and Fresni l l o, Zacatecas (the
l ocal i ty we vi si ted i s si tuated about 10 km north
of Fresni l l o, on the hi ghway to Sal ti l l o), therefore
we are i ncl i ned to bel i eve that the ori gi nal pl ant
coul d come fr om thi s ar ea. We thi nk that
Anderson was wrong i n i denti fyi ng them wi th T.
lloydii, a speci es that was descri bed as havi ng
ei ght radi al spi nes, but for whi ch no menti on was
made on the presence of a central spi ne.
Recentl y a photogr aph was publ i shed
(Succul enta, 1995) of a T. hexaedrophorus comi ng
from La Mancha Durango, near the northern bor-
der of Zacatecas and then from an area corre-
spondi ng to that gi ven i n the ori gi nal descri pti on
(northern Zacatecas), wi th heavy spi nes and
wi thout central s, that fi ts wel l the descri pti on
made by Bri tton and Rose. Due to speci es vari -
abi l i ty we thi nk i t i s not justi fi ed to recogni se thi s
Bradl eya 18/2000 44
Key to the species
1a Fl owers magenta wi th a red throat .......................................................................................T. bicolor
A Areol ar gl ands absent .................................................................................subsp. heterochromus
AA Areol ar gl ands present
B Central spi nes 0(1) ........................................................................................subsp. schwarzii
BB Central spi nes 34
C Stem sl i ghtl y cl usteri ng, cyl i ndri cal , spi nes 1829 ..................................subsp. bolaensis
subspeci es, or other enti ti es, at a formal rank.
Thelocactus buekii and Thelocactus matudae
are cl osel y rel ated and di sti ngui sh from each
other by the di fferent shape of the tubercl es, that
are hi gher i n matudae, for the di fferent number
of spi nes, that are more numerous i n matudae,
and for the di fferent fl ower si ze, that i s l arger i n
matudae. We feel that we can accept the sub-
speci es matudae provi si onal l y, because the fi nd-
i ng of new l ocal i ti es and the presence of i nterme-
di ate forms, the densel y spi ned forms typi cal of
Rayones i ntergradi ng southerl y, near Gal eana,
wi th l ess spi ny forms beari ng onl y one central
spi ne, coul d make superfl uous the di vi si on of
these two enti ti es.
Anderson consi dered both these two taxa as
vari eti es of T. tulensis. Accordi ng to us, however,
they have nothi ng i n common wi th thi s speci es,
whose seed mi cromorphol ogy i s compl etel y di ffer-
ent. The seeds of buekii and matudae have the
testa cel l s convex, the anti cl i nal wal l s strai ght
and the funi cl e remai ns not vi si bl e (Fi gure 8),
whi l e the seeds of tulensis have tabul ar testa
cel l s, undul ate anti cl i nal wal l s and the funi cl e
remai ns conspi cuous (Fi gure 9). We thi nk that
these di fferences are suffi ci ent to consi der T.
buekii unrel ated to T. tulensis and then to mai n-
tai n the former taxon at speci es rank, whi l e
matudaeshoul d be treated as a subspeci es of i t.
Thelocactus hastifer, Thelocactus lausseri,
Thelocactus macdowelli and Thelocactus tulensis,
are not cl osel y rel ated to any other speci es. None
of the morphol ogi cal characters i s useful to estab-
l i sh a rel ati onshi p among these four speci es and
the other thel ocacti , but some suggesti ons can be
obtai ned from seed mi cromorphol ogy. The fl at
testa cel l s of the seeds of hastifer and tulensis
suggest a rel ati onshi p wi th leucacanthus or hexa-
edrophorus, whi l e lausseri and macdowelli, hav-
i ng convex testa cel l s, can be compared to any
other speci es havi ng these characters, except T.
conothelos whose seeds have coni cal testa cel l s
wi th a smooth cuti cl e.
Systematic treatment
Thel ocactus Bri tton & Rose, Bul l . Torrey Bot.
Cl ub 49: 251 (Aug. 1922). Type: Echinocactus
hexeadrophorus Lemai re.
Torreycactus Dowel d, Sukkul enty 1: 19
(1998).
Stem si ngl e or cl usteri ng, depressed, gl obose,
ovoi d or cyl i ndri cal , 220 cm di ameter, 340 cm
hi gh. Ribs present or i ndi sti nct. Tubercles pre-
sent, rounded to coni cal . Areoles on the api ces of
the tubercl es, someti mes wi th a short adaxi al
groove, wi th or wi thout gl ands. Spines usual l y
strai ght, vari abl e i n di ameter, l ength and col our,
central and radi al spi nes general l y di sti ngui sh-
abl e. Flowers api cal , funnel -shaped, wi th a scal ed
tube, whi te, yel l ow or magenta. Fruits greeni sh to
reddi sh, semi -fl eshy, scal y wi th the peri anth rem-
nants persi stent, openi ng at maturi ty by a basal
pore. Seeds pyri form, wi th a basal hi l um, bl ack,
testa cel l s fl at, convex or coni cal , cuti cl e mi cro-
papi l l ate or smooth.
Figure 8. Seed of Thelocactus buekii ssp. matudae,
SB973 Gal eana, Nuevo Len. (x 40).
Figure 9. Seed of Thelocactus tulensis, s. n. Tul a,
Tamaul i pas. (x 40).
CC Stem si ngl e, ovoi d, spi nes 1123
D Stem up to 20 cm hi gh, ri bs 8, spi nes 1122 ...........................................subsp. bicolor
DD Stem up to 9 cm hi gh, ri bs 13, spi nes 1523................................subsp. flavidispinus
1b Fl owers whi te, magenta or yel l ow wi thout a red throat
2a Pri mary fi l aments i nserted above the base of the nectar chamber, testa cel l s coni cal , cuti cl e
smooth ........................................................................................................................T. conothelos
A Fl owers whi te to magenta
B Stem green, tubercl es angl ed, fl owers magenta (rarel y whi te), seeds wi thout conspi cuous
funi cl e remai ns
C Central spi nes whi te and shreddi ng, spi nes 2429 ...........................subsp. argenteus
CC Central spi nes greyi sh, not shreddi ng, spi nes 1124.......................subsp. conothelos
BB Stem ol i ve-green or reddi sh, tubercl es rounded, fl owers whi te to pal e magenta, seeds
wi th conspi cuous funi cl e remai ns .................................................................subsp. garciae
AA Fl owers yel l ow
B Stem si ngl e, tubercl es angl ed, spi nes 1227........................................subsp. aurantiacus
BB Stem cl usteri ng, tubercl es rounded, spi nes 79..............................................subsp. flavus
2b Pri mary fi l aments i nserted near the base of the nectar chamber, testa cel l s fl at or convex, cuti -
cl e mi cro-papi l l ate
3a Central and radi al spi nes whi te........................................................................T. macdowellii
3b Central spi nes col oured, radi al spi nes whi te or col oured
4a Fl owers smal l , 2545 mm di am., whi ti sh wi th a magenta mi dstri pe..................................
...............................................................................................................................T. lausseri
4b Fl owers 30100 mm di am., whi te, magenta or yel l ow
5a Areol ar gl ands present
6a Stem erect or decumbent, cyl i ndri cal , up to 40 cm hi gh, ri bs 1318, spi nes 2430
....................................................................................................................T. hastifer
6b Stem erect, gl obose, up to 15 cm hi gh, ri bs 8, spi nes 921...........T. leucacanthus
A Fl owers yel l ow ....................................................................subsp. leucacanthus
AA Fl owers magenta ........................................................................subsp. schmollii
5b Areol ar gl ands absent
7a Stem depressed to gl obose, ri bs 1321, tubercl es angl ed, hypodermi s wi th two or
three l ayers ........................................................................................T. rinconensis
A Fl owers yel l ow..............................................................................subsp. hintonii
AA Fl owers whi te to magenta
B Central and radi al spi nes not di sti ngui shabl e ..............subsp. rinconensis
BB Central and radi al spi nes di sti nct
C Stem gl aucous, spi nes heavy, shredded, 817, fl owers whi te to magenta
..........................................................................................subsp. nidulans
CC Stem green, spi nes not heavy or shredded, 810, fl owers magenta.........
..............................................................................subsp. freudenbergeri
7b Stem depressed, gl obose or short cyl i ndri cal , ri bs 821, tubercl es angl ed or round-
ed, hypodermi s wi th one l ayer
8a Ri bs i ndi sti nct, tubercl es angl ed, central and radi al spi nes i ndi sti ngui shabl e..
..................................................................................................T. multicephalus
8b Ri bs present or i ndi sti nct, tubercl es rounded, central and radi al spi nes di sti nct
9a Stem depressed to gl obose, gl aucous, tubercl es rounded, central spi nes gen-
eral l y mi ssi ng, fl owers whi te to l i ght magenta ............T. hexaedrophorus
9b Stem depressed to short cyl i ndri cal , not gl aucous, tubercl es coni cal , central
spi nes present, fl owers whi te to magenta
10a Stem gl obose to short cyl i ndri cal , ol i ve-green, ri bs present, fl owers
whi te to pal e pi nk, seeds wi th fl at testa cel l s .......................T. tulensis
Bradl eya 18/2000 45
Bradl eya 18/2000 46
List of accepted taxa
Thel ocactus bi col or (Gal eotti ex Pfei ffer )
Bri tton & Rose, Bul l . Torrey Bot. Cl ub 49: 251
(1922). Lectotype: Abbi l d. Beschr. Cact. 2: Pl .
25 (1848).
Echinocactus bicolor Gal eotti ex Pfei ffer ,
Abbi l d. Beschr . Cact. 2: pl . 25 (1848).
Ferocactus bicolor (Gal eotti ex Pfei ffer) N. P.
Tayl or, Cact. Succ. J. Gr .Bri t. 41: 30 (1979).
Echinocactus rhodophthalmus Hooker, Bot.
Mag. 76: pl . 4486 (1850). Echinocactus
rhodophthalmus var. ellipticus Hooker, Bot.
Mag. 78: pl . 4634 (1852). Echinocactus ellipti-
cus Lemai re, Jard. Fl eur. 3: pl . 270 (1853).
Echinocactus bicolor var. schottii Engel m.,
Syn. Cact. U. S. 21 (1856). Echinocactus schot-
tii Smal l , Fl . Southeast U. S. 814 (1903).
Thelocactus bicolor var. schottii Krai nz, Di e
Kakteen, Lfg. 18 (1961). Echinocactus bicolor
var. tricolor Schumann, Gesambt. Kakt. 303
(1898). Thelocactus bicolor var. commodus
Haas, Kakt. and. Sukk. 39: 86 (1988).
Thelocactus bicolor ssp. commodus (Haas)
Dowel d, Sukkul enty 1: 30 (1999).
ssp. bol aensi s (Runge) Dowel d, Sukkul enty 1:
2731 (1999). Lectotype: Gartenfl ora 38: 106,
Abb. 21 (1889)
Echinocactus bolaensis Runge (bolansis),
Gar tenfl or a 38: 106 (1889). Echinocactus
bicolor var. bolansis Schumann, Gesambt.
Kakt. 303 (1898). Thelocactus bicolor var.
bolansis Ber ger , Kakteen, 256 (1929).
Ferocactus bicolor var. bolaensis N. P. Taylor,
Cact. Succ. J. Gr . Br i t. 41: 30 (1979).
Echinocactus wagnerianus Berger, Kakteen,
256 (1929). Thelocactus wagnerianus Berger,
Kakteen, 346 (1929). Thelocactus bicolor var.
wagnerianus Krai nz, Di e Kakteen, Lfg. 18
(1961).
ssp. fl avi di spi nus (Backeberg) N. P. Tayl or, CCI
5: 14 (1998). Lectotype: Bei tr. Sukk.-Kunde
Pfl ege 1941: 6 (1941) Thelocactus bicolor var.
flavidispinus Backeb., Bei tr. Sukk.-Kunde
Pfl ege 1941: 6 (1941). Thelocactus flavidispi-
nus Backeb., Cact. Succ. J. (US) 23: 150
(1951). Echinocactus flavidispinus Weniger,
Cacti S. W. 87 (1970) nom. nud. Ferocactus
bicolor var. flavidispinus N. P. Taylor, Cact.
Succ. J. Gr. Bri t. 41: 30 (1979).
Thel ocactus bi col or ssp. heterochromus
(Weber) Mosco & Zanovel l o comb. et stat.
nov. Basionym: Echinocactus heterochromus
F. A. C. Weber, Boi s Di ct. Hort. 466 (1896).
Neotype: Durango, 3.5 km al Oeste de l a
Sol edad, R. D. Worthington 10902 (TEX).
Thelocactus heterochromus (F. A. C. Weber)
van Oosten, Kakteenkunde 58 (1940).
Ferocactus heterochromus N. P. Taylor, Cact.
Succ. J. Gr. Bri t. 41: 90 (1979). Echinocactus
bicolor var. pottsii Sal m-Dyck, Cact. Hort.
Dyck. 173 (1850).
ssp. schwarzi i (Backeberg) N. P. Tayl or, CCI 5:
14 (1998). Lectotype: Cact. Succ. J. Gr. Bri t.
12: 84 (1950). Thelocactus schwarzii Backeb.,
Cact. Succ. J. Gr . Br i t. 12: 81 (1950).
Ferocactus bicolor var. schwarzii N. P. Taylor,
Cact. Succ. J. Gr . Br i t. 41: 30 (1979).
Thelocactus bicolor var. schwarzii E. F.
Anderson, Bradl eya 5: 61 (1987).
Thel ocactus bueki i (Kl ei n) Br. & R., Cact. 4: 8
(1923). Lectotype: Gartenfl ora 8: 257, Taf. 266
(1859).
Echinocactus buekii Kl ei n (buckii),
Gartenfl ora 8: 257 (1859). Thelocactus tulen-
sis var. buekii (Kl ei n) E. F. Ander son,
Bradl eya 5: 65 (1987). Thelocactus tulensis
ssp. buekii (Kl ei n) N. P. Tayl or, CCI 5:14
(1998).
Thel ocactus bueki i ssp. matudae (Sanchez-
Mej. & Lau) Mosco & Zanovel l o comb. nov.
Basionym: Thelocactus matudae Sanchez-
Mej. & Lau, Cact. Suc. Mex. 23: 5152 (1978).
Type: Nuevo Len, near Rayones, A. Lau
Rubens s. n. (MEXU).
Thelocactus buekii var. matudae (Sanchez-
Mej. & Lau) E. F. Anderson, Bradl eya 5: 66
(1987). Thelocactus tulensis ssp. matudae
(Sanchez-Mej. & Lau) N. P. Tayl or, CCI 5: 14
(1998).
Thel ocactus conothel os (Regel & Kl ei n)
Backeb. & F. Knuth, Kaktus-ABC, 385 (1935).
Neotype: Tamaul i pas, 1 mi l e south-west of La
Perdi da, 22 Jan. 1961, E. F. Anderson 1725
(POM).
Echinocactus conothelos Reg. & Kl ei n, I nd.
Sem. Hort. Petrop. 48 (1860). Gymnocactus
conothelos Backeb., Di e Cactaceae V: 2859
(1961). Torreycactus conothele (Regel & Kl ei n)
Dowel d, Sukkul enty 1: 1530 (1998).
Echinocactus smithii Muehl enpf. nom. rej.
prop., Al l g. Gartenzei tung 14: 370 (1846).
Echinocactus saussieri Weber , Boi s Di ct.
Hor t., 468 (1896). Thelocactus saussieri
Berger, Kakteen, 257 (1929). Gymnocactus
10b Stem depressed to gl obose, ol i ve-green to reddi sh, ri bs general l y i ndi s-
ti nct, fl owers magenta, seeds wi th convex testa cel l s...............T. buekii
A Tubercl es l ow, poi nted, spi nes 512, fl owers 3545 mm di am............
.........................................................................................subsp. buekii
AA Tubercl es hi gh, coni cal , spi nes 1019, fl owers 4080 mm di am. .......
....................................................................................subsp. matudae
Bradl eya 18/2000 47
saussieri Backeb., Cact. Succ. J. (US) 23: 151
(1951).
ssp. argenteus (Gl ass & Foster) Gl ass, I dent.
Gui de Threatened Cacti of Mexi co 1: TH/CON
(1997). Type: Nuevo Len, about 7 mi l es west
of Ascensi on, 12 Feb. 1971, Glass & Foster
3176 (ZSS).
Thelocactus conothelos var. argenteus Gl ass &
Foster, Cact. Succ. J. (US) 44: 48 (1972).
Torreycactus conothelevar. argenteus (Gl ass &
Foster) Dowel d, Sukkul enty 1: 19 (1998).
ssp. auranti acus (Gl ass & Foster) Gl ass, I dent.
Gui de Threatened Cacti of Mexi co 1: TH/CON
(1997). Type: Nuevo Len, about 9 mi l es east
of La Escondi da, 12 Feb. 1971, Glass & Foster
3183 (ZSS).
Thelocactus conothelos var. aurantiacus Gl ass
& Foster, Cact. Succ. J. (US) 44: 48 (1972).
Torreycactus conothele var. aurantiacus
(Gl ass & Foster) Dowel d, Sukkul enty 1: 19
(1998).
Thel ocactus conothel os ssp. fl avus (Mosco &
Zanovel l o) Mosco & Zanovel l o comb. et stat.
nov. Basionym: Thelocactus flavus Mosco &
Zanovel l o Cactus & Co., 3: 20 (1999). Type:
San Lui s Potos , Hui zache, Kuenzler 362, cul t.
A. Mosco, Nov. 1998 (HG-PAD).
Thelocactus panarottoanus Hal da, Acta Mus.
Ri chnov. sect. Nat. 5: 161 (1998).
Thel ocactus conothel os ssp. garci ae (Gl ass)
Mosco & Zanovel l o comb. et stat. nov.
Basionym: Thelocactus garciae Gl ass, I dent.
Gui de Threatened Cacti of Mexi co 1: TH/GA
(1997). Type: Tamaul i pas, Muni ci pi o de
Bustamante, J . A. Garcia Luna 383, (CANTE).
Thel ocactus hasti fer (Werderm. & Boedeker) F.
Knuth, i n Backeb. & F. Knuth, Kaktus-ABC,
360 (1935). Type: (B). Echinocactus hastifer
Werdeman. & Boedeker, Noti zbl . Bot. Gart.
Mus. Ber l i n-Dahl em 11: 274 (1931).
Ferocactus hastifer N. P. Taylor, Cact. Succ. J.
Gr. Bri t. 41: 90 (1979).
Thel oactus hexaedrophorus (Lemai re) Bri tton
& Rose, Bul l . Torrey Bot. Cl ub 49: 251 (1922).
Neotype: I conogr aphi e descr i pti ve des
Cactees, Pl . 2 (Dec. 1841).
Echinocactus hexaedrophorus Lemai re, Cact.
Gen. Nov. Sp. 27 (1839). Echinocactus
droegeanus Hi l dm. ex Schumann, Gesambt.
Kakt. 438 (1898). Echinocactus hexaedropho-
rus var. droegeanus R. Meyer, Monatsschr.
Kakt.-Kunde 27: 40 (1917). Echinocactus fos-
sulatus Schei dw., Al l g. Gartenz. 9: 49 (1841).
Echinocactus hexaedrophorus var. fossulatus
Sal m-Dyck ex Labouret, Monogr. Cact. 251
(1853). Thelocactus fossulatus Br. & R., Cact.
4: 10 (1923). Thelocactus hexaedrophorus var.
fossulatus Backeb., Di e Cactaceae V: 2800
(1961). Echinocactus hexaedrophorus var.
labouretianus Schumann, Gesambt. Kakt.
438 (1898). Thelocactus hexaedrophorus var.
labouretianus A. Berger, Kakteen 253 (1929).
Echinocactus hexaedrophorus var. major
Quehl , Monatsschr. Kakt.-Kunde 4: 29 (1894).
Thelocactus hexaedrophorus var. major A.
Berger, Kakteen 253 (1929). Echinocactus
hexaedrophorus [var.] roseus Lemai r e ex
Labour et, Monogr . Cact. 251 (1853).
Echinocactus hexaedrophorus [var.] subcosta-
tus Sal m-Dyck, Cact. Hort. Dyck. 1849, 34
(1850). Thelocactus hexaedrophorus var.
decipiens A. Ber ger , Kakteen 253 (1929).
Thelocactus lloydii Bri tton & Rose, Cact. 4: 11
(1923). Thelocactus hexaedrophorus var. lloy-
dii (Br. & R.) Kl adi wa & Fi ttkau, i n Krai nz,
Di e Kakteen Lfg. 61 (1975). Thelocactus hexa-
edrophorus ssp. loydii (Br. & R.) N. P. Taylor,
CCI 5: 14 (1998).
Thel ocactus l ausseri J. Ri ha & J. Busek, Kakt.
and. Sukk. 37: 162164 (1986). Type:
Coahui l a, Si erra de l as Ovejas, A. Lausser s. n.
(PR).
Thel ocactus l eucacanthus (Zucc. ex Pfei ffer )
Bri tton & Rose, Cact. 4: 8 (1923). Lectotype:
Abb. Bayer. Akad. Wi ss. Mnchen 2: Tab. I I
(1837).
Echinocactus leucacanthus Zucc. ex Pfei ffer,
Enum. Cact. 66 (1837). Ferocactus leucacan-
thus N. P. Tayl or Cact. Succ. J. Gr. Bri t. 41: 90
(1979). Cereus maelenii Pfei ffer, Al l g. Gartenz.
5: 378 (1837). Echinocactus maelenii Sal m-
Dyck, Cact. Hor t. Dyck. 18 (1842).
Mammillaria maelenii Sal m-Dyck, Cact.
Hort. Dyck. 14 (1845). Echinocactus maelenii
Hemsl ey (macleanii), Bi ol . Centr. Amer. Bot.
1: 534 (1880). Cereus tuberosus Pfei ffer ,
Enum. Cact. 102 (1837). Echinocactus tubero-
sus Sal m-Dyck ex Foerster, Handb. Cact. 287
(1846). Echinocactus leucacanthus var.
tuberosus Foerster, Handb. Cact. 287 (1846).
Echinocactus ehrenbergii Pfei ffer , Al l g.
Gartenz. 6: 275 (1838). Thelocactus ehren-
bergii F. Knuth, i n Backeb. & F. Knuth,
Kaktus-ABC, 359 (1935). Echinocactus leuca-
canthus var. crassior Sal m-Dyck, Cact. Hort.
Dyck. 35 (1850). Echinocactus porrectus
Lemai r e, Cact. Al i q. Nov. 17 (1838).
Thelocactus porrectus F. Knuth, i n Backeb. &
F. Knuth, Kaktus-ABC, 361 (1935).
Thelocactus leucacanthus var. porrectus
Backeb., Di e Cactaceae V: 2818 (1961).
Echinocactus subporrectus Lemai r e, Cact.
Al i q. Nov. 25 (1838). Echinocactus tuberosus
var. subporrectus Foerster, Handb. Cact. 523
(1846). Echinocactus theloideus Sal m-Dyck,
Al l g. Gartenz. 18: 396 (1850).
ssp. schmol l i i (Werder.) Mosco & Zanovel l o, CCI
7: 18 (1999). Lectotype: Kakt. and. Sukk. Pfl .
Bradl eya 18/2000 48
3: Taf. 160 (1939).
Thelocactus leucacanthus var. schmollii
Werderm., Bl h. Kakt. and. Sukk. Pfl . 3: Taf.
160 (1939). Thelocactus krainzianus Oehme,
Bei tr . Sukk-Kunde Pfl ege 1; 1 (1940).
Thelocactus leucacanthus var. schmollii fa.
krainzianus (Oehme) Nagl , Kakt. and. Sukk.
42: 183 (1991). Thelocactus sanchezmejoradai
Meyr n, Cact. Suc. Mex. 3: 77 (1958).
Thelocactus leucacanthus var. sanchezmejo-
radai Backeb., Di e Cactaceae V: 2817 (1961).
Thel ocactus macdowel l i i (Rebut ex Quehl )
Gl ass, Cact. Suc. Mex. 14: 4 (1969). Neotype:
Coahui l a, 33 km northeast of Sal ti l l o on hy. 40
to Monterrey, 22 Jul y 1972, E. F. Anderson
3182 (US).
Echinocactus macdowellii Rebut ex Quehl
(Mc. Dowellii), Monatsschr. Kakt.-Kunde 4:
133134 (1894). Echinomastus macdowellii
Br. & R., Cact. 3: 151 (1922). Neolloydia mac-
dowellii H. E. Moore, Bai l eya 19: 166 (1975).
Thelocactus conothelos var. macdowellii Gl ass
& Foster (mcdowellii), Cact. Succ. J. (US) 49:
220 (1977).
Thel ocactus mul ti cephal us Hal da &
Panarotto, Acta Mus. Ri chnov. sect. Nat. 5: 40
(1998). Type: Feb. 1985, J . J . Halda s. n. (PR).
Thelocactus rinconensis ssp. multicephalus
(Hal da & Panarotto) Luethy, Kakt. and Sukk.
50: 80 (1999).
Thel ocactus ri nconensi s (Posel ger) Bri tton &
Rose, Cact. 4: 7 (1923). Neotype: About 20 km
northeast of Sal ti l l o, 22 Jul y 1972, E. F.
Anderson 3180 (US).
Echinocactus rinconensis Posel ger, Al l g.
Gartenz. 23: 18 (1855). Echinocactus rin-
conadensis Schumann, Gesambt. Kakt. 433
(1898). ?Echinocactus lophothele Sal m-Dyck,
Al l g. Gartenz. 18: 395 (1850). Thelocactus
lophotheleBr. & R., Bul l . Torrey Bot. Cl ub 49:
251 (1922). Echinocactus phymatothelos
Posel ger ex Ruempl er, i n Frster, Handb.
Cact., ed. 2, 602 (1885). Thelocactus phyma-
tothelos Br. & R. (phymatothele), Cact. 4: 8
(1923). Thelocactus rinconensis var. phyma-
tothelos Gl ass & Foster, Cact. Succ. J. (US) 49:
246 (1977). Thelocactus rinconensis ssp. phy-
matothele (Posel ger ) Gl ass, I dent. Gui de
Threatened Cacti of Mexi co 1: TH/RI N (1997).
Thelocactus rinconensis ssp. phymatothelos
(Posel ger) Dowel d nom. superfl ., Sukkul enty
1: 30 (1999).
ssp. freudenbergeri (Haas) Mosco & Zanovel l o,
CCI 7: 18 (1999). Type: Freudenberger s. n. i n
Col l . Haas Nr. 157 (ZSS).
Thelocactus rinconensis var. freudenbergeri
Haas, Kakt. and. Sukk. 43: 9698 (1992).
ssp. hi ntoni i Luethy, Kakt. and. Sukk. 48: 39
(1997). Type: Nuevo Len, Muni ci pi o Gal eana,
Weg von Ci enega del Toro nach Santa Rosa,
1610 m, 3 Oct. 1995, Hinton et al . 25762
(Herbari um G. B. Hi nton).
ssp. ni dul ans (Quehl ) Gl ass, I dent. Gui de
Threatened Cacti of Mexi co 1: TH/RI N (1997).
Type: (B). Echinocactus nidulans Quehl ,
Monatsschr . Kakt.-Kunde 21: 119 (1911).
Thelocactus nidulans Br. & R., Cact. 4: 9
(1923). Thelocactus lophothele var. nidulans
Kl adi wa & Fi ttkau, i n Krai nz, Di e Kakteen,
Lfg. 61 (1975). Thelocactus rinconensis var.
nidulans Gl ass & Foster, Cact. Succ. J. (US)
49: 245 (1977). Thelocactus rinconensis ssp.
nidulans (Quehl ) Dowel d nom. super fl .,
Sukkul enty 1: 30 (1999).
Thel ocactus tul ensi s (Posel ger) Bri tton & Rose,
Cact. 4: 11 (1923). Neotype: Tamaul i pas, at
KM 14 on hi ghway 101 between Tul a and the
juncti on at hy. 80, 30 Jul y 1979, E. F.
Anderson 3202 (US). Echinocatus tulensis
Posel ger, Al l g. Gartenz. 21: 125 (1853).
Specimens examined
I f not di fferentl y speci fi ed, al l the data concern-
i ng the fol l owi ng OTUs refer to pl ants grown by
A. Mosco.
bicolor speci mens obser ved: Chi huahua:
HK74-2170; Coahui l a: s.n. Moncl ova, SB287
Sal ti l l o, SB563 Cuesta l a Mural l a, Z37 Ojo
Cal i ente; Nuevo Len: s.n. Tanqueci l l os; San
Lui s Potos : s.n. Hui zache, SB278 Hui zache,
PAN194 El Coyote; Texas: SB866 Starr Co.
Li terature: stem di ameter and hei ght, Bravo-
Hol l i s & Sanchez-Mejorada (1991).
bolaensis speci mens observed: cul ti vated ori -
gi n, col l ecti on number 54; Coahui l a: SB281
Cerro Bol a. Li terature: stem di ameter and
hei ght, Runge (1889); head number, Pi l beam
(1996).
commodus speci mens observed: Nuevo Len:
s.n. Montemorel os. Li terature: stem di ameter
and hei ght, tubercl e hei ght and wi dth, fl ower
maxi mum di ameter, Haas (1988).
flavidispinus speci mens obser ved: Texas:
SB424 Brewster Co. Li terature: stem di ame-
ter, tubercl es hei ght and wi dth, radi al spi nes
mi ni mum number, Anderson (1987); stem
hei ght, Benson (1982); radi al spi nes maxi -
mum number, Weni ger (1988).
Pedri cena speci mens observed: Coahui l a:
SB607 San Pedro, SB1715 South Boqui l l as.
pottsii speci mens observed: Chi huahua: SB77
Ji menez; Durango: SB1433 se Cuencam, s.n.
Mapi mi . Li terature: stem di ameter, Pi l beam
(1996).
schottii speci mens observed: Texas: SB567
Brewster Co. Li terature: stem di ameter and
hei ght, Benson (1982); tubercl e hei ght and
Bradl eya 18/2000 49
wi dth, Weni ger (1988).
schwarzii speci mens observed: Tamaul i pas:
LAU684 Zaragoza, Z107 Cal l es. Li terature:
stem di ameter and hei ght, head number,
Pi l beam (1996); tubercl e hei ght, Bravo-Hol l i s
& Sanchez-Mejorada (1991); radi al spi nes
maxi mum number, Anderson (1987).
wagnerianus speci mens observed: cul ti vated
ori gi n: col l ecti on number 314; Coahui l a: Z57
El Dorado. Li terature: stem di ameter and
hei ght, head number, Berger (1929).
heterochromus speci mens observed: cul ti vated
or i gi n: col l ecti on number s 63, 266;
Chi huahua: SB564 Matamor os, SB608
Hi dal go del Parral , SB1879 south Satevo.
Li terature: stem di ameter and hei ght, fl ower
maxi mum di ameter, Pi l beam (1996); radi al
spi nes mi ni mum number, Anderson (1987).
conothelos speci mens observed: Nuevo Len:
s.n. Dr. Arroyo, s.n. east Dr. Arroyo; San Lui s
Potos : SB302 Matehual a, CH186 San
Antoni o, CSD109 Guer r er o, CSD115 La
sol edad; Tamaul i pas: s.n. Mi qui huana, CH237
Tul a. Li terature: stem di ameter and hei ght,
tubercl e hei ght and wi dth, Anderson (1987).
argenteus speci mens observed: Nuevo Len:
SB311 Ascensi on. Li terature: stem di ameter
and hei ght, Anderson (1987); tubercl e hei ght
and wi dth, Gl ass (1972).
aurantiacus speci mens observed: cul ti vated
ori gi n: col l ecti on numbers 75, 136; Nuevo
Len: s.n. Zaragoza, GL758 Zaragoza, SB329
Ar amber r i , LAU1009 Ar amber r i , CH194
Aramberri , CSD149 Marmol ejo. Li terature:
stem di ameter and hei ght, Anderson (1987).
garciae speci mens observed: Tamaul i pas: s.n.
Bustamante. Pl ants obser ved i n habi tat:
Tamaul i pas: s.n. near Bustamante.
Li terature: orthosti chy number, radi al spi nes
maxi mum number , head number , Gl ass
(1997).
flavus speci mens observed: San Lui s Potos :
HK362 Hui zache, s.n. La Hi ncada. pl ants
observed i n habi tat: San Lui s Potos : s.n.
Hui zache.
buekii speci mens observed: garden ori gi n: col -
l ecti on number 57; Nuevo Len: CH195
Ar amber r i , CSD133 Escondi da, CSD145
Lampaci tos, CSD155 Ascensi on. Li terature:
al l the data regardi ng morphol ogi cal charac-
ters have been taken from Anderson (1987)
and Pi l beam (1996).
matudae speci mens observed: garden ori gi n:
col l ecti on number 71; Nuevo Len: LAU744
Rayones, SB973 Gal eana, SB1339 Rayones.
Li terature: stem di ameter and hei ght, tuber-
cl e hei ght and wi dth, spi ne count, Anderson
(1987); fl ower maxi mum di ameter, Pi l beam
(1996).
rinconensis speci mens observed: garden ori -
gi n: col l ecti on number 69; Coahui l a: s.n. San
Jose de l os Nunci os, s.n. Ramos Ar i spe,
HO969 El Chi fl on, SB301 Ri nconada; Nuevo
Len: s.n. Santa Catar i na, CSD185 Casa
Bl anca. Li terature: stem di ameter and hei ght,
Pi l beam (1996); tubercl e hei ght and wi dth,
Bravo-Hol l i s & Sanchez-Mejorada (1991).
freudenbergeri speci mens observed: Nuevo
Len: s.n. Grutas de Garci a, CSD173 Grutas
de Garci a. Li terature: stem di ameter and
hei ght, tuber cl e hei ght and wi dth, Haas
(1992).
hintonii we have been unabl e to observe any
pl ant of thi s taxon, therefore al l the morpho-
l ogi cal data have been taken from the l i tera-
ture, Gl ass (1997) and Lthy (1997), and the
mi ssi ng data have been consi dered equal to
those of T. rinconensis.
nidulans speci mens observed: Coahui l a: s.n.
Si erra de l a Pai l a. Li terature: stem di ameter
and hei ght, Bravo-Hol l i s & Sanchez-Mejorada
(1991); tuber cl e hei ght and wi dth, Quehl
(1911), spi ne number, Lthy (1997).
phymatothelos speci mens observed: garden
ori gi n: col l ecti on numbers 41, 64; Coahui l a:
CH205 Arteaga. Li terature: stem di ameter,
Br avo-Hol l i s & Sanchez-Mejor ada (1991),
stem hei ght, Schumann (1898).
multicephalus speci mens obser ved: Nuevo
Len: HO809 Sandi a. Pl ants observed i n habi -
tat: Nuevo Len: s.n. Tri ni dad. Li terature:
stem di ameter and hei ght, tubercl e hei ght
and wi dth, head number, Hal da (1998).
La Bol sa speci mens observed: Nuevo Len:
s.n. La Bol sa. Pl ants observed i n habi tat:
Nuevo Len: s.n. El Desi erto; San Lui s Potos :
s.n. south of Matehual a.
hexaedrophorus speci mens observed: San Lui s
Potos : s.n. Buenavi sta, s.n. San Lui s Potos ,
CH230 Char co Bl anco, Hayek63 Vi l l ar ,
Hayek71 Moctezuma, PAN137 Ri o Ver de,
SB892 Guaxcama, SB1074 Charco Bl anco.
Nuevo Len: BZ44 west of Dr Arroyo, SB291
Dr Ar r oyo; Tamaul i pas: s.n. La Per di da,
H2313 Mi qui huana. Li terature: stem di ame-
ter and hei ght, Pi l beam (1996); tuber cl e
hei ght and wi dth, central spi nes maxi mum
number, Anderson (1987).
lloydii speci mens observed: San Lui s Potos :
SB113 Sal i nas. Li terature: stem di ameter and
hei ght, tubercl e hei ght and wi dth, spi ne num-
ber, Anderson (1987).
tulensis speci mens observed: Tamaul i pas: s.n.
Tul a, BZ28 Tul a, CH236 Tul a, CSD69 Mi guel
Hi dal go; San Lui s Potos : s.n. Hui zache, s.n.
La Li bertad, CSD103 Hui zache; Nuevo Len:
CSD127 Dr Ar r oyo/Mi er y Nor i ega.
Li ter atur e: stem di ameter and hei ght,
Bradl eya 18/2000 50
Pi l beam (1996); spi ne number , Ander son
(1987).
macdowellii speci mens observed: garden ori -
gi n: col l ecti on numbers 121, 143; Coahui l a:
s.n. Hi gueras, CSD192 Arteaga. Li terature:
stem di ameter and hei ght, Pi l beam (1996);
tubercl e hei ght and wi dth, spi ne number, sty-
l us col our, head number, Anderson (1987).
lausseri speci mens observed: garden ori gi n:
col l ecti on numbers 127, 198. Li terature: stem
di ameter and hei ght Pi l beam (1996); spi ne
number, Anderson (1987).
leucacanthus speci mens observed: Hi dal go:
s.n. San Fr anci sco, SB514 I xmi qui l pan;
Quertaro: s.n. Mesa del Len. Li terature:
stem di ameter and hei ght, spi ne number,
fl ower di ameter , Br avo-Hol l i s & Sanchez-
Mejorada (1991); tubercl e hei ght and wi dth,
Anderson (1987).
schmollii speci mens obser ved: Quer tar o:
SB579 Vi zarrn. Li terature: stem di ameter
and hei ght, spi ne number, Bravo-Hol l i s &
Sanchez-Mejorada (1991); tubercl e hei ght and
wi dth, Meyran (1958).
krainzianus speci mens observed: garden ori -
gi n: col l ecti on number 108; Quertaro: s.n.
west of Pea Mi l l er , Z185 Pea Bl anca.
Li terature: stem di ameter and hei ght, Oehme
(1940).
hastifer speci mens observed: garden ori gi n:
col l ecti on numbers 115, 255. Li terature: stem
di ameter and hei ght, Guzman (1992); tuber-
cl e hei ght and wi dth, spi ne number, styl e
col our , Ander son (1987); fl ower di ameter ,
Pi l beam (1996).
setispinus speci mens observed: Texas: s.n.
Bexar Co., SB851 Ji m Hogg Co., SB858
Cameron Co. Li terature: stem di ameter and
hei ght, head number, Weni ger (1988); ri b
hei ght and wi dth, spi ne number, petal oi d
l ength and wi dth, Anderson (1987); fl ower
di ameter, Pi l beam (1996).
Acknowledgements
Thanks ar e due to Mr T. Ubal di ni , Centr o
Pol i val ente Servi zi di Ateneo, Tri este Uni versi ty,
for the preparati on of the materi al to be observed
by SEM, and to our fr i ends, both of the
Di parti mento di Bi ol ogi a, Tri este Uni versi ty, Dr
P. Gi ul i ani ni for hi s support i n the executi on of
the l i ght mi croscope anal ysi s and Mr F. Bersan
for hi s advi ce i n the use of the software MULVA-
4. We are grateful to the many fri ends, credi ted
al ong thei r photos, who send us some beauti ful
transparenci es to compl ete those we l acked.
References
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Bradl eya 18/2000 51
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Nomencl atural adjustments i n Thelocactus
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Appendix
Characters and their scaling
Stem shape I: 0 fl attened, 1 fl attened to gl obose,
2 gl obose to ovoi dal , 3 cyl i ndri cal . Stem shape
II: 0 erect, 1 decumbent. Maximum stem diam-
eter (mm): 0 100, 1 150, 2 > 150. Maximum
stem height (mm): 0 50, 1 100, 2 150, 3
200, 4 > 200. Clustering: 0 si ngl e stems, 1 10
heads, 2 20 heads, 3 > 20 heads. Epidermis
colour I: 0 gr een, 1 ol i ve-gr een r eddi sh.
Epidermis colour II: glaucous, 0 no, 1 yes.
Epidermis colour III: blotched, 0 no, 1 yes.
Number of hypodermal layers: 0 one l ayer, 1
two or more l ayers. Crystals: 0 absent, 1 present.
Seedling shape: 0 gl obose, 1 cyl i ndri cal . Ribs: 0
absent, 1 present. Orthostichy number: the
orthosti chy number corresponds to the ri b num-
ber or, for the pl ants wi th i ndi sti nct ri bs, to the
sum of the orthogonal parasti chi es that, accord-
i ng to the phyl l otaxi s theory of Thomas (1975), i s
equal to the number of the present recti seri al
orthosti chi es. 0 813, 1 1321, 2 > 21. Tubercles:
0 absent, 1 present. Tubercle shape: 0 rounded,
1 angl ed. Grooved tubercles: 0 no, 1 yes.
Tubercle or rib width (mm): 0 5, 1 10, 2
15, 3 20, 4 > 20. Tubercle or rib height (mm):
0 10, 1 15, 2 20, 3 > 20. Areolar glands: 0
absent, 1 present. Minimum total spine num-
ber: 0 5, 1 10, 2 15, 3 20, 4 > 20.
Maximum total spine number: 0 10, 1 15,
2 20, 3 25, 4 > 25. Central and radial spines
distinguishable: 0 no, 1 yes. Central spines
hooked: 0 no, 1 yes. One or more upper
spines flattened: 0 no, 1 yes. Shredded
spines: 0 no, 1 yes. Spine colour I: white: 0 no,
1 yes. Spine colour II: ochre to reddish: 0 no,
1 yes. Spine colour III: ochre to greyish: 0 no,
1, yes. Minimum flower diameter (mm): 0
40, 1 50, 2 60, 3 > 60. Maximum flower
diameter (mm): 0 50, 1 60, 2 70, 3 80, 4
> 80. Minimum ratio between length and
width of the inner petaloids: we have chosen
to use a r ati o i nstead of l ength and wi dth
absol ute val ues to normal i se the vari ati ons due
to envi ronmental or growth reasons. 0 2, 1 3,
2 4, 3 5, 4 > 5. Maximum ratio between
length and width of the inner petaloids: 0
4, 1 5, 2 6, 3 7, 4 8, 5 > 8. Shape of the
petaloid apex: 0 acute, 1 acute or obtuse, 2
obtuse. Margin of the petaloid apex: 0 enti re,
1 enti re or erose, 2 erose. Flower colour I:
white: 0 no, 1 yes. Flower colour II: magenta:
0 no, 1 yes. Flower colour III: yellow-orange:
0 no, 1 yes. Flower colour IV: red throat: 0 no,
1 yes. Style colour I: white: 0 no, 1 yes. Style
colour II: magenta: 0 no, 1 yes. Style colour
III: yellow: 0 no, 1 yes. Stigma colour I:
creamy white: 0 no, 1 yes. Stigma colour II:
orange: 0 no, 1 yes. I f thi s and the previ ous char-
acter are both zero, then the sti gma col our i s yel -
low. Filament colour I: white: 0 no, 1 yes.
Filament colour II: magenta: 0 no, 1 yes. I f
thi s and the previ ous character are both zero,
then the fi l ament col our i s yel l ow. Primary fila-
ments insertion: 0 l ow, 1 hi gh. Pollen: 0 tri col -
pate, 1 pol ycol pate. Scented flowers:0 no, 1 yes.
Early bud development: 0 no, 1 yes.
Flowering period: 0 spri ng (I I I V), 1 spri ng-
summer (I VX), summer (VI X). Fruit colour: 0
reddi sh-green, 1 bri l l i ant red. Fruit succulence:
0 no, 1 yes. Fruit dehiscence: 0 no, 1 yes. Seed
average diameter (mm): 0 1.2, 1 1.5, 2 >
1.5. Seed average length (mm): 0 1.4, 1 1.8,
2 > 1.8. Hilum micropylar region width: 0
smal l er than testa, 1 l ar ger then testa.
Micropyle position I: micropyle inside the
hilum: 0 no, 1 yes. Micropyle position II:
micropyle on the edge of the hilum: 0 no, 1
yes. Micropyle position III: micropyle out-
side the hilum: 0 no, 1 yes. Appendages on
the hilum edge: 0 no, 1 yes. Funicle rests con-
spicuous: 0 no, 1 yes. Testa cell shape: 0 fl at, 1
fl at or sl i ghtl y convex, 2 convex, 3 coni cal .
Anticlinal cell walls: 0 strai ght, 1 si nuate.
Secondary sculpture of the seedcoat I:
micro-papillate: 0 no, 1 yes. Secondary sculp-
ture of the seedcoat II: striate: 0 no, 1 yes.
Bradl eya 18/2000 52
bic bol com fla ped pot scho schw wag het con arg
Stem shape I ov cy ov ov cy gl ov ov cy gl ov ov
Stem shape II erect erect erect erect erect erect erect erect erect erect erect erect
Maximum stem diameter (mm) 125 100 80 70 60 150 120 55 60 150 180 130
Maximum stem height (mm) 200 400 70 90 >150 60 130 90 >200 70 450 180
Head number 1 5 1 1 1 1 1 5 5 1 1 1
Epidermis colour I gr gr gr gr gr gr gr gr gr gr/red gr gr
Epidermis colour II: glaucous - - - - - - - - - - - -
Epidermis colour III: blotched - - - - - - - - - - - -
Number of hypodermal layers 1 1 1 1 1 1 1 1 1 1 1 1
Crystals + + + + + + + + + + + +
Seedling shape gl gl gl gl gl gl gl gl gl gl gl gl
Ribs + + + + + + + + + + - -
Orthostichy number 8 13 8 13 8-13 8 8 13 13 8 21 21
Tubercles + + + + + + + + + + + +
Tubercle shape rd rd rd rd rd rd rd rd rd rd ad ad
Grooved tubercles - - - - - - - - - - - -
Tubercle or rib width (mm) 25 18 18 8 15 25 18 15 14 37 18 15
Tubercle or rib height (mm) 15 9 14 5 6 14 12 13 7 17 16 10
Areolar glands + + + + + - + + + - + +
Minimum total spine number 11 20 14 15 19 12 15 10 18 7 11 24
Maximum total spine number 22 28 16 23 24 16 22 13 29 14 24 29
Central and radial spines distinct + + + + + + + + + + + +
Hooked spines - - - - - - - - - - - -
Upper spine flattened + + + + + + + + - + - -
Shredded spines - - - - - - - - - - - +
Spine colour I: white - + - - - - - - - - + +
Spine colour II: ochre to reddish + - + + + + + + + + - -
Spine colour III: ochre to greyish - - - - - - - - - - + +
Minimum flower diameter (mm) 55 60 50 60 55 60 92 65 60 45 45 50
Maximum flower diameter (mm) 90 80 100 80 70 90 115 80 65 82 60 55
Min petaloid length/width ratio 3 2.9 3 1.9 3.4 3.5 3.6 2.8 3.4 2.8 4.1 3.2
Max petaloid length/width ratio 5 6.6 4.3 3.5 4.4 4.2 5.3 3.9 3.6 4.7 6.6 5.2
Shape of the petaloid apex a/o a/o a/o a a a/o a/o a/o o a/o a a
Margin of the petaloid apex er er er en/er er en/er er er er er en en
Flower colour I: white - - - - - - - - - - w -
Flower colour II: magenta mg mg mg mg mg mg mg mg mg mg mg mg
Flower colour III: yellow-orange - - - - - - - - - - - -
Flower colour IV: red throat + + + + + + + + + + - -
Style colour I: white w w w w w w w w w w w -
Style colour II: magenta - - - mg - - mg - - mg mg mg
Style colour III: yellow - - - - - - - - - - - -
Stigma colour I: creamy white w w - - w w - - w w w w
Stigma colour II: orange or - or or - - or or - or - -
Filament colour I: white w w w w w w w w w w w w
Filament colour II: magenta mg - - mg - - - mg - - - -
Primary filaments insertion l l l l l l l l l l h h
Pollen pol y pol y pol y pol y pol y pol y pol y pol y pol y pol y tri tri
Scented flowers - - - - - - - - - - - -
Early bud development - - - - - - - - - - - -
Flowering period I V-I X VI -I X I I I -VI I V-VI I I V-I X V-VI I VI V-VI I I V-X V-I X I I I -VI I I I -I V
Fruit colour gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red
Fruit dehiscence + + + + + + + + + + + +
Fruit succulence - - - - - - - - - - - -
Seed average diameter (mm) 1.3 1.31 1.33 1.41 1.3 1.4 1.53 1.33 1.42 1.56 1.57 1.46
Seed average length (mm) 1.68 1.75 2.06 1.74 1.7 1.71 1.84 1.58 1.89 1.75 1.83 1.87
Large hilum micropylar region - - - - - - - - - - + +
Micropyle inside the hilum - - - - - - - - - + + +
Micropyle on the edge of the hilum + + + + + + + + + + - -
Micropyle outside the hilum + + - - + - - - + + - -
Appendages on the hilum edge + + - - - - - + - - - -
Funicle remains conspicuous - - - - - - - - - - - -
Testa cells shape cvx cvx cvx cvx cvx cvx cvx cvx cvx cvx con con
Sinuate anticlinal cell walls - - - - - - - - - - - -
Micro-papillate seedcoat + + + + + + + + + + - -
Striate seedcoat - - - - - - - - - - - -
Table I.
Key: a = acute; ad = angl ed; con = coni cal ; ct = tabul ar or sl i ghtl y convex; cvx = convex; cy = cyl i ndri c; de = depressed; en
= enti re; er = erose; gl = gl obose; gr = green; h = hi gh; l = l ow; mg = magenta; o = obtuse; or = orange; ov = ovoi d; pol y =
pol ycol pate; rd = rounded; tab = tabul ar; tri = tri col pate; w = whi te; y = yel l ow.
bic = bicolor; bol = bolaensis; com= commodus; fla = flavidispinus; ped = Pedri cena; pot = pottsii; scho = schottii;
schw= schwarzii; wag= wagnerianus; het = heterochromus; con = conothelos; arg= argenteus.
Bradl eya 18/2000 53
aur gar flv bue mat rin fre hin nid phy mul lab
Stem shape I ov ov ov gl gl gl gl gl gl de gl de
Stem shape II erect erect erect erect erect erect erect erect erect erect erect erect
Maximum stem diameter (mm) 110 140 120 180 150 200 150 200 200 150 200 100
Maximum stem height (mm) 100 100 120 60 140 80 80 80 100 30 150 40
Head number 1 10 11 1 1 1 1 1 5 1 65 1
Epidermis colour I gr gr/red gr/red gr/red gr/red gr gr gr gr gr gr gr
Epidermis colour II: glaucous - - - - - + - + + + + +
Epidermis colour III: blotched - - - - - - - - - - + +
Number of hypodermal layers 1 1 1 1 1 2 2 2 2 2 1 1
Crystals + + + + + + + + + + + +
Seedling shape gl gl gl gl gl gl gl gl gl gl gl gl
Ribs - - - - + + + + + + - -
Orthostichy number 21 21 13 13-21 13-21 25 21 24 25 13 13-21 13-21
Tubercles + + + + + + + + + + + +
Tubercle shape ad rd rd rd rd ad ad ad ad ad ad ad
Grooved tubercles - - - - - - - - - - - -
Tubercle or rib width (mm) 19 15 18 25 20 25 20 20 20 17 20 18
Tubercle or rib height (mm) 14 20 15 24 25 20 20 25 20 13 20 12
Areolar glands + + + - - - - - - - - -
Minimum total spine number 12 12 7 5 10 3 8 3 8 1 3 3
Maximum total spine number 27 18 9 12 19 5 10 10 17 5 5 5
Central and radial spines distinct + + + + + - + + + - - -
Hooked spines - - - - - - - - - - - -
Upper spine flattened - - - - - - - - - - - -
Shredded spines - - - - - - - - + - - -
Spine colour I: white + + - - - - - - - - - -
Spine colour II: ochre to reddish - - - - - - - - - - - -
Spine colour III: ochre to greyish + + + + + + + + + + + +
Minimum flower diameter (mm) 40 26 30 35 40 40 50 45 45 40 30 55
Maximum flower diameter (mm) 60 45 45 45 80 70 60 70 50 65 58 60
Min petaloid length/width ratio 4 2.4 3.3 5 4.4 4 5.3 4 5.5 4.7 3.5 3.6
Max petaloid length/width ratio 4.9 3.5 6 5 6.2 5.3 7 5.3 6.3 6 6.1 6.1
Shape of the petaloid apex a a/o a a a a a a a a a a
Margin of the petaloid apex en en er en en/er en en en en en en en
Flower colour I: white - w - - - w - - w - w w
Flower colour II: magenta - mg - mg mg mg mg - mg mg - -
Flower colour III: yellow-orange y - y - - - - y - - - -
Flower colour IV: red throat - - - - - - - - - - - -
Style colour I: white - w - w w w - w w w w w
Style colour II: magenta - mg mg mg mg - mg - - - - -
Style colour III: yellow y - y - - - - - - - - -
Stigma colour I: creamy white - w w w w w w w w w w w
Stigma colour II: orange - - - - - - - - - - - -
Filament colour I: white - w - w w w w w w w w w
Filament colour II: magenta - - - - - - - - - - - -
Primary filaments insertion h h h l l l l l l l l l
Pollen tri tri tri tri tri tri tri tri tri tri tri tri
Scented flowers - - - - - - - - - - - -
Early bud development - - - - - - - - - - - -
Flowering period I I I -I V I I -I I I I I I -I V I I I - I I I -I X I I -I X I V I I I - I I -V I I -VI I I I I I -I V I I I -I X
Fruit colour gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red
Fruit dehiscence + + + + + + + + + + + +
Fruit succulence - - - - - - - - - - - -
Seed average diameter (mm) 1.58 1.54 1.69 1.45 1.52 1.5 1.56 1.5 1.6 1.57 1.65 1.58
Seed average length (mm) 1.85 2.3 2.21 1.89 2.14 1.9 2.31 1.9 2.07 1.93 2.19 2.04
Large hilum micropylar region + - - - - - - - - - - -
Micropyle inside the hilum + + + - - - - - - - - -
Micropyle on the edge of the hilum + - + + + + - + + + + +
Micropyle outside the hilum - - - + + + + + - - + +
Appendages on the hilum edge - - - + + - - - - - + +
Funicle remains conspicuous - + + - - - - - - - - -
Testa cells shape con con con cvx cvx ct tab ct ct ct tab tab
Sinuate anticlinal cell walls - - - - - - - - - - - -
Micro-papillate seedcoat - - - + + + + + + + + +
Striate seedcoat - - - - - - - - - - - -
Table I. (Continued)
Key: a = acute; ad = angl ed; con = coni cal ; ct = tabul ar or sl i ghtl y convex; cvx = convex; cy = cyl i ndri c; de = depressed; en
= enti re; er = erose; gl = gl obose; gr = green; h = hi gh; l = l ow; mg = magenta; o = obtuse; or = orange; ov = ovoi d; pol y =
pol ycol pate; rd = rounded; tab = tabul ar; tri = tri col pate; w = whi te; y = yel l ow.
aur = aurantiacus; gar = garci ae; flv = flavus; bue= buekii; mat = matudae; rin = rinconensis; fre= freudenbergeri; hin
= hintonii; nid = nidulans; phy = phymatothelos; mul = multicephalus; lab = La Bol sa.
Bradl eya 18/2000 54
hex loy tul mac lau leu schm kra has set
Stem shape I gl gl ov gl ov ov ov ov cy cy
Stem shape II erect erect erect erect erect erect erect erect dec erect
Maximum stem diameter (mm) 150 120 80 150 85 70 70 60 55 125
Maximum stem height (mm) 75 60 250 95 100 150 100 80 400 300
Head number 1 1 7 5 1 >50 35 23 28 8
Epidermis colour I gr gr gr/red gr gr gr gr gr gr gr
Epidermis colour II: glaucous + + - - - - - - - -
Epidermis colour III: blotched - - - - - - - - - -
Number of hypodermal layers 1 1 1 1 1 1 1 1 1 1
Crystals + + + + + + + + + -
Seedling shape gl gl gl gl gl gl gl gl cy gl
Ribs + + + - + + + + + +
Orthostichy number 8-13 8 8-13 21 8 8 8 8 18 13
Tubercles + + + + + + + + + -
Tubercle shape rd rd rd rd rd rd rd rd rd
Grooved tubercles - - - - - - - + -
Tubercle or rib width (mm) 33 18 20 6 17 14 12 17 5 2
Tubercle or rib height (mm) 11 9 21 4 14 11 10 14 6 18
Areolar glands - - - - - + + + + +
Minimum total spine number 4 7 6 17 24 9 14 9 24 10
Maximum total spine number 9 11 16 29 31 24 21 16 30 20
Central and radial spines distinct + + + + + + + + + +
Hooked spines - - - - - - - - - +
Upper spine flattened - - - - - - - - - -
Shredded spines - - - - - - - - - -
Spine colour I: white - - - + + - - - + -
Spine colour II: ochre to reddish + + - - - - - - - -
Spine colour III: ochre to greyish - - + - + + + + + +
Minimum flower diameter (mm) 50 50 30 40 25 35 40 53 35 30
Maximum flower diameter (mm) 100 60 60 65 45 45 65 90 50 42
Min petaloid length/width ratio 3.1 5.5 4.8 4.6 5 3.5 5.4 5.4 4 3.3
Max petaloid length/width ratio 7.2 6.3 6 7 8.4 5.2 6.8 7.1 4.6 3.6
Shape of the petaloid apex a a a a a a a a a a
Margin of the petaloid apex en/er en en en en en/er en/er en/er en en
Flower colour I: white w w w - - - - - - -
Flower colour II: magenta mg - mg mg mg - mg mg mg -
Flower colour III: yellow-orange - - - - - y - - - y
Flower colour IV: red throat - - - - - - - - - +
Style colour I: white w w w w w w w w w -
Style colour II: magenta mg - mg mg mg mg mg mg mg -
Style colour III: yellow - - - - - - - - - y
Stigma colour I: creamy white w w w w w w w w w -
Stigma colour II: orange - - - - - - - - - -
Filament colour I: white w w w w w w w w w w
Filament colour II: magenta - - - - - mg mg mg - -
Primary filaments insertion l l l l l l l l l h
Pollen tri tri tri pol y tri tri tri tri tri tri
Scented flowers - - - - - - - - - +
Early bud development - - - - + - - - + -
Flowering period I I I -VI I I V-VI I I I -I X I I I -I V I I -VI VI -I X V-I X VI -I X I I I -I V VI -I X
Fruit colour gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red gr/red red
Fruit dehiscence + + + + + + + + + -
Fruit succulence - - - - - - - - - +
Seed average diameter (mm) 1.52 1.38 1.38 1.43 1.23 1.38 1.34 1.46 1.53 1.13
Seed average length (mm) 1.72 1.68 1.68 2.04 1.71 1.6 1.55 1.96 1.69 1.39
Large hilum micropylar region - - - - - - - - - -
Micropyle inside the hilum + - + + - - - - - -
Micropyle on the edge of the hilum + - + + + + - - - +
Micropyle outside the hilum + + + + - + + + + -
Appendages on the hilum edge - - - - - - - - - +
Funicle remains conspicuous - - + - - - - - - -
Testa cells shape tab ct tab cvx cvx tab tab tab tab cvx
Sinuate anticlinal cell walls - - + - - - - - - -
Micro-papillate seedcoat + + + + + + + + + -
Striate seedcoat - - - - - - - - - +
Table I. (Continued)
Key: a = acute; ad = angl ed; con = coni cal ; ct = tabul ar or sl i ghtl y convex; cvx = convex; cy = cyl i ndri c; de =
depressed; en = enti re; er = erose; gl = gl obose; gr = green; h = hi gh; l = l ow; mg = magenta; o = obtuse; or
= orange; ov = ovoi d; pol y = pol ycol pate; rd = rounded; tab = tabul ar; tri = tri col pate; w = whi te; y = yel l ow.
hex = hexaedrophorus; loy = lloydii; tul = tulensis; mac = macdowellii; lau = lausseri; leu = leucacanthus;
schm= schmollii; kra = krainzianus; has = hastifer; set = setispinus.

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