Morphology of Cacti [I]
Morphology of Cacti Section I - Roots and stems
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Science QK 495 2011 B96
THE
.1 BIOEMON
UNIVERS
MICE
. CHIGAN
OF
MORPHOLOGY OF CACTI Section
I. Roots
and Stems
Ву DR . FRANZ BUXBAUM STYRIA
,
AUSTRIA
Edited by
EDWIN
B
. KURTZ , Jr . ;
Kerckhoff Laboratories of Biology California Institute of Technology Pasadena
4
, California ,
Original Drawings by
Mrs. Friedl
BUXBAUM
Published by
ABBEY GARDEN 132
West Union Street ,
Pasadena
PRESS 1
, California
Science Botany Wahr 7
-
5
- 51
75449
TABLE OF CONTENTS PREFACE INTRODUCTION
.
.
.
.
.
.
.
.
.
.
.
I
OF THE ROOTS
.
.
.
.
.
.
.
.
.
1
.
.
.
.
.
.
. .
.
.
.
.
.
. . . . .
. .
. .
. . .
.
.
. . .
·
.
. .
.
. .
. . . . .
. .
.
. . . . . .
. .
.
. . . .
. .
.
. .
·· ·· ··· · · · ·· · ·· ··
.
. .
.
.
.
.
. .
.
.
.
. .
. .
. .
periodical thickening
.
. .
stem
THE CACTACEAE
.
of
.
LITERATURE CITED
OF
··
.
. .
Cereeae
Habit
. .
B
Opuntieae
THE
.
A
Pereskieae
the Cactaceae
.
.
.
. .
and
factor and bilateral symmetry
Branching
PHYLOGENY
.
branching
OF
.
of
. . .
.
Light
G
of
.
.
.
B C
D
E
Angle
F
Longitudinal symmetry Acrotony and basitony
the primordial
41
30 24 22
.
.
.
. . .
. .
.
.
.
.
. . . . . .
.
.
.
.
.
.
.
. . . . . . . .
·
OF
mamilles
.
.
.
.
.
. . . .
.
.
. . . . .
· ··
spines
. . . ·
. . . ·
·
. .
)
.
of
·
·
.
( .
.
. . . . ·
of
OF
·
. .
. .
Habit THE CACTACEAE General factors Longitudinal growth
A
.
3
.
Secondary evolution
.
···
.
of
-
-
. . . .
D E
D
tubercles
E
B C
Ribs
Mamilles
C
4
. ···· .. ·· ·· ·· ··
THE CACTACEAE
Leaves Cotyledons and seedlings
A
.
areoles
the spines
Structure the areole Anatomy and morphology
SUCCULENCE
2
the
. . .
of
Leaf nature
plants
. . .
Bud nature
.
A B
of
dicotyledonous
Form
.
THE AREOLES
C
1 .
MORPHOLOGY
of
OF
III .
MORPHOLOGY OF THE STEM
.
II. MORPHOLOGY
.
. . . . .
1.
PREF
first of three treating with the morphology
familiarity
,
,
in
.
the many
or
the
for making
Taylor
W
Mr
thank
genus
the present systems
Dr
.
by
the names used
the editor that the inclusion
with
on
. of
,
literature
addi
species
of
of
course
confusion for the reader
following
these
classifica
my sincere
within thanks
parentheses
Mr
the end
suffice the
the text
.
be found
in
literature cited occurs
will
for
at
good dictionary
brief defini
of
,
in A
.
footnotes
.
express
list
the manuscript
to
to
.
wish
I
Scott Haselton
his
suggestions
.
JR
KUTRZ
,
E
. B .
.
this time
this may
as
terms have been added
and references
many valuable
1950
of
for
Arizona
scientific terms used
of
certain
A
these
clarify
to
,
At
wish
used
of
increase
those not otherwise defined
section
,
,
is
the hope
also
has
. to
of
of
In
for
help
to
.
Buxbaum
and
cacti
order
tions
needless
These occur within parentheses
It
.
.
Dr
synonyms will tion
avoid
this connection
,
by of
synonyms
given
scientific names
Phoenix
to “ translate ” and
alteration
I
his manuscript
Marshall Desert Botanical Garden tions
of
a
was deemed advisable
publication
the main generic and specific synonyms
in
Buxbaum
minimum
synonyms
.
by
including
so
the numerous
,
the Cactaceae
with
the task has been
in English
In
Because
it
of
desire
do
my
been
to
organize the original manuscript
.
editing this monograph
In
of
.
to
appear shortly
flowers , fruits , and seeds , will
on
it
the Cactaceae. The following two sections
of
,
is the
to
This SECTION on roots and stems
CE
A
MORPHOLOGY OF CACTI ROOTS AND STEMS
A
small
diameter deeper
of
echinocarpa less than two feet tall and one and one -half inches in root thirty - five feet long . This and another root almost as long were no than three inches beneath the ground and covered an area of about eight square feet around the base of the plant . plant
had
Opuntia
a main
I. INTRODUCTION
,
the
.
in
,
plants that have survived
the path
the evolutionary
.
be
How
Thus
,
,
its
, is
's
of
,
of
the plant can the plant
vital
a
of
the Cactaceae with
a
morphologi
guide and incentive
cacti
.
of
the purpose
of
will fulfill
MORPHOLOGY OF THE ROOTS The woody portion the mature roots like that the stem but more closely meshed and becomes creasingly dense toward the center This parenchyma tissue caused reduction from the periphery inward the center parenchyma Milk tubes may there found the cortical parenchyma the Sec Galactochylus Mammillarias
has shown He found that there are usually four six xylem points the stele growth early stage periderm At
Haehnel found that lateral roots Echinofossulocactus multicostatus have xy groups that lein and phloem not alter nate The xylem groups are separated by
in ,
of
is
.
of
.
be
.
In
of
by
).
do
of
.
20 ( )
(
parenchyma ]
1
-
well
[
a
.
is
.
in
,
a
of
so
a
in
.
to
an
no
is
in
,
as
.
) to
that the young roots have
.
is
,
of of
-
as
By
all
.
,
of
in
29
(
formed
is
of
or
formed near the base the stem and some genera roots are formed from other parts the stem the aerial roots most epiphytic species modification these basic forms the root types found the Cactaceae may be derived Generally the roots are anatomically simi lar other dicotyledonous plants Reiche also
corky layer
developed
more roots are
of
.
a
CACTUS seedlings usually possess less strong tap root Adventitious
-
-
II .
life beauty
understood
students
the evolution
the plant itself that
.
of be
of
,
provide
all
it
of
will
forms
chain
genera and species found
the primary habit
the author that this monograph
further exploration
phylogenetic
ana
both
evolutionary
.
an
to
knowledge
cal study and that to
understanding
struggle for existence
the hope
partially filled
plant has
phylogeny
the key
also
Only from its
.
is
It
is
Morphology power and
,
the key
links may
a
transformation
organs
and the true relationships
followed
is
morphology
to
changes may now
feature of
the intergrading
numerous missing links
the missing
be
,
the present time
and habit
many
comprehensive study and comparison
a
,
to
a
by
,
is
have been lost and the result
. Any
plant organ
evolutionary
of
.
Unfortunately
long
of
is
tomical and physiological
product
, of a
the
not arisen suddenly , but
ever
of each
the development of shape
must also show
descriptions
exact
of
true, morphology must encompass more than
. For this to be of plants ; morphology
phylogeny
to
of
that morphology is the key
BEEN STATED
of all
IT HAS
and
the phloem
groups
are
MORPHOLOGY
-
to
of
mistaken
.
,
)
( )
. .
cm 1
(
5
.
,
-
,
)
(
have
Melocactus intortus Cactus intortus ex hibits extensive lateral root system The roots this species grow only below the surface but extend laterally more than seven meters One specimen eighteen being transplanted months after had new length roots that were This readily explains why some species such Melocactus declare are difficult culti pots Cacti with lateral root systems vate quickly absorbing water are well adapted light from rains Also coastal forms Melocactus are able absorb the limited quantities rainfall because the ground too saline and cannot used by water the plant of
:
Schumann Carleton Preston observed that large plants have two types roots strong roots which pene
Schumann may
cause and effect however
an
to
of
33
According
system
.
,
lateral root
.
to
attached the xylem tips the whole struc being very the antaomy similar ture the stem
OF THE ROOTS
as
.
to
,
.
in
cm
75
so
of
,
.
is
be
of
to
.
IKE
to
.
in
,
..
H
of
set
a
,
Soul Level
are modified for the water and become rapiform especially and Aleshy This common extremely arid regions cacti that grow those that not have very fleshy stems Fleshy roots occur Mammillaria napina longimamma Dolichothele often
do
or
.
of
-
in
)
20
.
(
of
,
trate downward into the ground and anchor the plant and very long lateral roots which extend horizontally near the surface the ground Haehnel also mentions very long chord like roots for anchorage cer Opuntia tain species
species
RY ORS
For ex roots oc
the same genus
.
of
in
WON
ERGI
,
occur
,
-
Small species may possess lateral roots beet like roots and both forms may
deep
-
,
.)
, L .
33
(
Pseudolob
.)
.
L
[
.
L
aurea
,
.
,
L
eriana
of
)
a
of
(
cantha Schumann believed two forms roots are result vironment because according ).
helbundet
in
- (
, ( L .
.
L
); . )
.)
(
, [L L . .
,
in
,
,
in
the genus Lobivia tap cur especially those species with oblong stems Wesseriana Mediolob Pyga Haagei oruensis Mediolob grow brachyantha laterally maelob and ing adventitious roots are found species Tegel scoparia with globular stems ample
hamata that the the en
,
EMA From 1919
.
Rose
:
and
92 ,
.
Opuntia subterranea 1
Britton
,
impenetrable developed
well
. 2 .
and a
,
soil covering very hard termite nests and has
FIG
in
as
-
a
of
in
,
to
Hirscht deep Notocactus Schumannianus grows tap root where soils meadows and has Notocactus nigrispinus grows the thin
.
M
,
. of
roots
-
fleshy
-
phantha exhibit non
Species tap
.
and others
.
,
)
(
cephalophora
,
.
.
88 ,
:
7
.
.
in
.
. 1 .
cates the soil surface
. -
Mammillaria Dawsonii The dotted line indi From Houghton Cact Succ 1935 Jour
Fig
M
.
in
in
or
1 .5
X
in
.
is
in
of
Tap roots preservation
Cory
.
.
mandragora
.
during the winter and the saine covered with soil
normally
it is
;
,
x 2 . 5
subterraneus shrivelled the root has been uncovered
,
R
.
Right
:
young plant
,
subterraneus
of
of
:
Rapicactus
Rapicactus Three year old seedling three months later The upper part .
. 5 .
IG
'
eedling
Left
.4Fig .
OF THE ROOTS
.3Fig .
MORPHOLOGY
-
at
;
a
of ,
.
)
.
8
(F
its to
of
a
a
.
)
, .7
.
(
cm
a
,
in
8
O
.
12
cm
length clusters and attains and diameter Fig puntia Chaffeyi has long branched tuberous root which yearly produces new stem that decays the ground during the dry periods ig environment This the only cactus which does not per sist is
a
,
is
.
-
singly
OF THE ROOTS m
!
of
.
a
at
a
.
Other modifications occur also Some Rebutias have very thick root which bears large number the top small globular fleshy stems Lophophora Williamsii has tap root which larger than the stem and the stem appears above the soil only during damp periods Mammillaria Dawsonii pos similar growth habit the surface sesses
or
MORPHOLOGY
.
)
1
.
(F
its
similar
Br .
in
SED
found Oehme Backberg
et
.
A
is
.
)
.
(
2
a
subterranea derives name from growth habit Fig very interesting root type the new genus Rapicactus Buxb
. . 4 3 ) ) . ,
(F
(
is
it
of
.
R
of
.
in
is
a
of
.
R
in
.
R
ment
NES
to
is
by
-
.
In
R
).
. R.
&
in
.
R
(
Thelocactus mandragora and subterraneus the genus Gymnocactus this genus the beet like root connected the turbi very thin woody nate stem means especi This woody connector section ally long ig subterraneus but mandragora short Fig From seedlings subterraneus the develop 1941 places
the long neck was studied
.
In
.
the rapiform
).
5
of
a
or
in
,
to a
.
,
is
'
of
to
(
the fleshy and simi the hypocotyl seedlings lar other cacti The epicotyl however grows and elongates thin stem length without form several centimeters globular ing Fig thickened head Both the hypocotyl and primary root form seedlings
a
A
.
(
.)
is
in
of
. is .
to is
is
a
,
is
at
If
.
root the thin extension injured the apex lateral formed from this point which branch similar the primary stem The neck be tween the root and head therefore not ecological factors result similar root morphology Weingartia Speg found Bckbg Neoporteria napina azzinia and
the epicotyl
.
,
seedling
Schumann
.
Fig
.6 .
as
in
From
in
as
Rapi
are
Ancistro The thin Fig has )
10
.
.
)
of
to
.
of
,
at
,
roots which are tuberous thickened only the ends and are very similar the roots Dahlia Similar this Opuntia macror are the tuberous roots to
is
9
Fig
Wilcoxia Poselgeri
connected
in
found
(
stemmed
the stem
.
cactus megarhizus
of
the head also been
(
to
neck cactus have a
.
).
6
up5
-
is
to to to
(F
.
cm
20
. to
Long tuberous roots which
by
grows many lateral
the structure below the cotyle that structure above the cotyle
is
'
.
.
to in
15
-
in
Pterocactus tuberosus which
The hypocotyl dons The epicotyl
dons
Greggii
Peniocereus
is
In
. . cm . in it
root
of
cm
8
a
.
-
,
to
of
The preceding form root leads the large rooted species with thin stems Peniocereus Greggii and Pterocactus tuber already osus the first species the root large beet like root ig the seedling long and Later becomes diameter Much larger roots diameter have been known occur Similar this the big tuber like 60
;
a
it
,
a
very flat disc bear this species forms only ing the tubercles but underground has very large branched root ig Opuntia
,
form
confirmed
.
been
.
,
. G .
of
,
a
,
of
.
on
in
,
to
.
of
in
as
of
do
as
.
of
to
.
do
a
in
)
a
20
do
as
in
,
.
may also
proliferum
roots have been Schele aerial roots lateral stems but this has not from
.
Gymnocalycium
,
,
and
lateral stems that arise According observed
to
scopa
,
,
metrizans Notocactus Ottonis Notocactus
(
-
In
of
.
a
as
as
are thick the stems All Aleshy roots are covered with cork and have mucous cortical parenchyma which holds water Myrtillocactus geo some specimens
the intervals between the ribs and Zygocactus the articulated species roots arise from the base the joints The young aerial roots are able absorb water from the air the aerial roots other epiphytes Aerial roots that contact the soil grow other roots Aerial roots that re adhering main the air mature form incapable absorbing roots moisture Haehnel states that aerial roots have parted stele with cambium several layers Soon the young root the central pith becomes woody the phloem ele Finally the xylem groups ments soon after not grow further and become useless and from
.
. .
-
in
.
its
of
.
so
branched roots Cereus pachyrhizus was named because swollen tuberous roots Fleshy but non tuberous roots are also found Aporocactus flagriformis Here the roots
in as
.
.
the broad end genera also have Aeshy
many epiphytic climb The aerial roots ing creeping species are special form adventitious roots The roots are borne laterally different parts the stem but Usually they arise never from the areoles
-
Other
at
-
or
.
of a
of
's
to
, . )
(
14
roots
thin
Dodds Mendoza
, or
According Engelmann picture this species forms cluster tuberous roots from the base the stem These tubers are club like and bear one more hiza
Photograph
of
.
root and branched root
.
Simple
six
:
. 7 .
Pierocactus tuberosus
by L
OF THE ROOTS
do
Fig
MORPHOLOGY
.
,
, :
and Rose
1923
.
Britton
1919
,
From
213
,
megarbizus
and Rose
1
From Britton
4 :4
Ancistrocactus
Chaffeyi
.
. 9 .
Fig
Opuntia
.
.
FIG .8
MORPHOLOGY
OF THE ROOTS
MORPHOLOGY
OF THE ROOTS
broken by extension . The old adhering root now is non - conductive , and only fun tions as a tractive organ . Some specimens also have band - like adhering roots (Fig . 11 ) .
When this occurs , it usually is a result of a mineral deficiency or water stress . parts of most cacti , sometimes Severed even disjointed fruits of Opuntia , rapidly
form adventitious roots . These arise , as do aerial roots , from the central stele and they also must penetrate the cortex or callus of the wound . In Opuntia these roots are al ways formed below the areoles , as was shown for Pereskia . Old rootless cactus parts readily form new roots if the base is not corky , or if the corky layer is gently
10
.
,
2
, : .
.
,
Wilcoxia Poselgeri cluster tuberous roots From Britton and Rose 111 1920
.
of Fig
removed .
)
in
26
x
2
:
Z -
.
;
,
;
of
in
.
.
X -
E of - a
Selenicereus hamatus section flat adher stage the old woody endodermis xylem elements remains central woody part After Reiche and Haehnel
FIG 11
ing root
.
.
,
of
.
is
It
as
.
of
as
of
(
Leinfellner found aerial roots cultivated plants Pereskia aculeata that arose just below the areoles and disappeared they had penetrated the epidermis soon the stem not known whether this also occurs naturally Adventitious roots may also arise from many species stems even columnar forms
III .
OF
MORPHOLOGY
.
1
of
.
A
dicotyledonous
plants is
it
of
of
.
a
to
of
To clarify the morphology areoles comparison first necessary make the cactus stem with the ordinary dicotyledon ous stem The dicot plant constructed by Sachs shows the organization the plant
.
)
.
.
12
even
in
leaves
The apex
of
the axils the cotyledons
Fig
,
,
buds arise
the axils
and leaf organs
(
stem
,
All
of
into root
in
de -
Form
the
, ,
of
,
is
do
Rhodocactus just
,
leaves
as
sub genus
-
its
.
and
well developed
AREOLES
of
, - -
,
of
.
to
to
of
it
of
.
a
as
others that each species can fied cactus Because nature the Cactaceae has been difficult ascertain the exact relation this family plant kingdom the Nevertheless mem bers the Cactaceae are typical dicotyle donous plants of the Order Centrospermae and the primitive genera are obviously rived from the morphological type the the most primitive genus order That Pereskia possess
MORPHOLOGY OF THE
as
does the Cact distinct from readily classi this singular
of be
is
.
so
a
forms
Yet this family
aceae
plants that have
THE STEM
all
families
of
There are few variety such
of
MORPHOLOGY
OF THE AREOLES
be
.
other dicotyledonous plants broadly The Cactaceae may
of
a
M
-nb
.
branches
"}
4
,
of
;
of
a
2
; 3 , ;
,
1
,
:by
character the presence areoles bearing hairs and spines succulent stem with foliage leaves and green cortex lack lateral usually very sparse development
-
l-
.
,
nb -s
-
lb
, :
.
undeveloped
covered by The axillary
and
leaves
.
,
young
growing point
is
a
is
stem
of
13
or
,
.
dicotyle FIG Leaf plants donous leaf petiole base lamina tipules blade p
,
an
buds which usually remain dormant for scaly
,
of
period time are covered by beginning with two lateral which are the first leaves the
indefinite
,
leaves
The
of
of
prophylls
angiosperm
a
,
).
13
plants
After Sachs
the meristem
which
,
of
leaf organs
forms the epi
.
part
very characteristic of
"
of
That dermis
.
dicotyledonous
representation
.
Diagrammatic
.
. 12 .
.
arise Fig
It is ,
is
.
of
tip
?
,
In
of
(F
and
.
leaf base
an
.
part
basal
of or
leaf consists upper part consisting ig the petiole and blade the development the leaf leaf pri mordia are formed from the dermatogen and subepidermal region the stem The leaf base first formed from the pri mordium and then the petiole and blade bud
,
ized
PLANTS
OF DICOTYLEDONOUS
FORM
the
of greatest importance , that leaves have apical growing point and that they intercalary growth size crease
early stages of development the base of bud may also merge with the leaf base and
.
; B -
Leaf
after
C
;
of
forma
Leaf with
C to -
elongation
leaf podarium
podarium
of
:
longitudinal
elongation
A
downward
of
.
with
-
tion
podarium
primordia
(
14
Schematic representation
Growing point with
A
.
FIG
6X
in
by
-
in
no
and
is
as in
shaded
a
.
.
Troll
in
of
,
be
.
.
displaced
As
ium be
.
a
14
.
.
of
After
toward the top the podar will shown this occurs the Cactaceae
.
of
Euphorbia
a
of
15
.
of
Podaria
of
. 15 .
Fig
podarium with the stem form Fig shows such podaria succulent Euphorbia Here the leaf base and the rest the leaf are quite distinct Fig shows podarium the method formation to
enlarges
. ,
the Cactaceae and others
in
,
The leaf base
elongation
downward
.
region podarium
of ;
of
bud
;
axillary
-
K
,
)
; -
D
of
podar Leaf with longitudinal elongation the Cactaceae ium and displaced axillary bud Troll
. :
-
; A - of
Opuntia ficus indica
; V
.
Bud nature
of
.
c )
spines
.
. B
,
(b
( a ),
The areole developed
Troll
the areoles
.
a
,
,
of
of
One the most characteristic organs the cactus stem the areole has been sub ject controversy for many years After preliminary work the problem by Goebel on
.
Growth between the apex and base
After
of
is
,
.
if
by
can
,
between
'
and
in
arises the angle the leaf base and the stem also be displaced upward the leaf base During the growth stretched upward
Pereskia aculeata
-
The axillary bud which
prophyll spines
)
above
aculeata
Old branch
b
the
:
Fig
. 18 .
Right
the growing point
axillary bud meristem terminal meristem prophyll spines areole and and leaf
(a of
Pereskia
:
;
podarium
,
Center
.
.
Troll
.
After
section through
; of -P
spine primordium
.
.
( d )
-
B
scar
representation
Schematic
leaf primordium Fig After Goebel
D 17 -
Left
:
. 16 .
FIG
AR
ci ha
mm
are
the spines more hairs Between The whole differentiated lateral formed young areole bud with spines and hairs During the growth the lateral bud young leaf becomes dif the areole form spine
.
.
is a
an
to
of
26
a
In
- .
of )
(
),
18
on
his
to
-
(
co
Leinfellner workers and morphology clarify the was the first Opuntia Rafinesquei work areoles by this species the leaf primordia are organ large hemispherical growing ized about
OF THE AREOLES
MORPHOLOGY
.
his
10
. of
,
.
is
is
in
of
.
of
a
is
in
a
in
of
as
in
plants other dicotyledonous primor development leaf the the axil formed lateral branch dium The lateral point the young leaf the axil but growth not the bottom actually situated slightly above the leaf The leaf primordium grows and be base
point Early
on
of
of
of its
.
*
its
on a
long cylindrical acuminate leaf and the lower part adaxial side The lateral point the lateral growth branch growth now forms true hairs along
comes bears
this row and later outside tissue hairs several small acuminate cones
scheme
Pereskia
.
Prototype
of
. 20 .
Fig
of
,
margins
of
,
or
podar
.
)
.
(F
the leaf base
16
,
is
lower part which ig
ium
or
,
a
,
of
,
at
as
.
On
the
the leaf pri outside ferentiated the first spine the same level mordium appears groove which the lateral bud the separates the upper part leaf from
of no
At
.
,
of
in
).
17
.
(
of
a
is
is
.
. .
...
C .
-
of
on
Pereskia aculeata has also shown Work axil first areoles the bud nature present lary bud this the leaf axils only plant short hairs cluster There and the place the areole bears only two Fig spines These spines short lateral
a
,
of
If a
).
18
do
as
.
,
.
,
(
is
do
.
.
of
( is (
,
of
may
be
20
.
'
areoles are homologous
to
therefore concluded that cactus the axillary buds dicotyledonous plants other
Yareples
re
✓
;
-
e ;
to
.
as
'
to
the ventral side the stem axis The side next opposed the abaxial side away from the stem the dorsal side
or
of of
,
.
)
a
19
of
tip
at
.
of
new
severed
but
It ).
toward
cones spine these cones becomes Each tissue base the while They first harden the lengthens the and meristematic remains the leaf
of
These become development
portion not develop rather lateral branches are Fig formed bearing first squamous leaves like Pereskia Thus the growth Fig other dicotyledonous plants that the
spines
. of
by .
are formed
displaced
of
P of .
A
. ;
-
1
:
the
leaf leaf the shoot
.
the first leaves After Troll
aculeata showing
of
of
Pereskia
axil
of
in
be -
to
,
b
;
branch
the
of
Fig
. 19 .
a
Part lateral shoot areole
all
of
the are actually the two lateral prophylls other areole and precede organs the bud Later the bud develops strong long spines areoles cluster young stem other cacti Fig aculeata cut off the uppermost areoles
young bud
Weingart (42 )
spines
that
seem
are
of
,
and
calcium
oxalate
spines
modified leaves
in
spines
of Pereskia
undulata he even observed chlorophyll and leucoplasts . Weingart failed , however, to axillary buds and the describe the areoles
Rudolf
only
stated that spines are formed
epidermis
the
from
but Delbrouk
showed
of
of
to
as
.
but always the areole
, .
on
the axils
as
leaves
spines are
leaves
.
modified
,
,
Goebel therefore concluded that
.
in
lateral branches arise
of ,
in
of
a
as
If
the growing point
in
,
in
as
in
a
developed into leaves were formed the same position spines ordinary areoles the areole were the areoles the severed stem had spines then these were displaced lateral branch formed the axil each spine just
arise from
.
first first
spines never
of ,
ral branch branch and the lateral and the
the der
growing point leaves Ganong
the
)
(
.
P
this bud
aculeata
from
the formation has pointed out that new arise between the old spines
a
As
areole
in
or
bud
arise
from
,
of
's
21 ).
is
of
matogen Leaves similarly
grasses are formed
15
.
.
-
;
D
of
-p
V
:
Londitudinal section through areole oint growth young spine After Delbrouck
Opuntia
of
the
21 .
.Fig
leaves
. of
to
(
OL
.
IN
is
as
,
)
12
(
Opuntia are that spines formed from the dermatogen and periblem the usual method formation most Fig Rudolf statement not leaves spines opposed the leaf homology
of
(
)
a
a
off
many postulated this, and the following studies are presented as proof of this hypothesis . Goebel ( 18 ) cut stem Pereskia young grandifolia above Rhodocactus
as
it would
leaves , as is the case with thorns of (48 ) first dicot plants . Zuccarini
modified
.
supposition
(41 ) he correctly stated and glochids arise di rectly from the growing point of the areole , and that the hairs arise from the epidermis . He also found in the subepidermal cells of the spines of Pereskia autumnalis crystals of Earlier
that the spines
)
Next the nature of spines must be clari spines arise from areoles , and fied . Because areoles have been shown to be buds, by
also found vascular strands
.
in some spines
,
the spines
30
. Leaf-nature of
(
C
II
THE SPINE
OF
as
LEAF - NATURE
)
)
(
of
.
(
26
anatomical studies Leinfellner reached the same conclusion On very young lateral branches Echinopsis Eyri esii hybrid he found intergradations be From
22
;
. of
D
t-i
:
.
;
h -
.
After Wetter .
wald
Wetterwald by basal
en
.
do
)
meristem
as
a
.
is
in
of
.
is
ig
)
still
22
top the
47
hard
(
the
showed that spines leaves The the spine becomes hardened when spine first seen the areole and lengthening After attaining maturity the spine also becomes basal portion
large
of
- - -
as
meristem
.
.
it
,
.
of
.
of )
(
Eriocereus Martinii the lower part the spine The spiny acuminate scales and petals Acantho calycium and other genera may be transi tory leaves and spines Inasmuch trich omes never possess vascular strands unless they are glandular appears that spines are modified leaves in
.
,
some thick
Harrisia Martinii vascular strands occur
Immature spine arborescens ntercalary hair
(F
flowers
Fig Opuntia spine
of
the
.
,
and
,
of
spines
in
Opuntia monacantha
P
In
.
in
of
to
of
.
tween leaves and spines He also found vascular strands leading the base each spine areoles Selenicereus Macdon aldiae Pereskia grandifolia bleo
I2
MORPHOLOGY
OF THE AREOLES
develop modified leaves because leaves around the growing point , whereas spines arise only unilaterally on the outside mar gin
. Also
they
observed that
flowers and
lateral branches originate from the growing point and do not appear between the spines , but rather above them . These workers must be in error because Leinfellner ( 26 ) foun ! that on a part around the flower tube of Selenicereus Macdonaldiae there are always radial areoles with spines arising around the
growing point
. 23 .
FIG
147
as if they
were leaf organs.
----
of
an
),
D
l-
.
.
; -
b
( h )
in
;
)
( D
as it
(
.
Opuntia vulgaris The spine shows the basal leaf part occurs the axil with areole spines podarium with and hairs the spine After Kauffman vascular strands
in
DIE
-
existence intermediate forms both these organs Such spine leaf forms are pecially frequent Pereskia and Opuntia
!
of
es of
is
of
of .
of
as
proof Perhaps the most important point spines and leaves the similarity the
a
23 ).
)
24
.
a
of
(
in
(
. -
be
or
. ,
in
to
.
a
an
leafy shoot when areole grows into Some primordia which have begun form spines form leaf tissue thus resulting leaves with spiny tips Petals which are spine tipped may also intermediate gans Kaufmann showed such transi Opuntia Fig tions The basal part containing the spine forms podarium a
,
if
,
in
of
,
in
.
is
,
to a
in
strand and the axil small are Similarly formed the bud grows shoot Pereskia aculeata hairy areoles with spines are formed the axils the vascular
ole
: :
;
H -
.
;
D -s
a
( B )
of
of in
.
(
of A )
G -
and
believed that spines are not
24 .
.
FIG Radial areole the axil scale of the flower tube Selenicereus Macdonaldiae point rowing pine the areole hair vascular strand After Leinfellner
S -g
),
Velenovsky
40
of
the areole (
),
32
) (
(
Schumann Chorinsky
Structure 10
.
D
.
oldest hooked spines
the
.
a
in
,
of
all
in
by of
)
method
the spines
formed
in
are
Although
rapid
).
In
.
26 a g
.
(
of
of
,
in
in
some genera others the spines develop gradually and the order formation can be easily studied Lobivia cinnabarina the spines are formed slowly long period and over time Fig The areoles this species are first only succession
at
of
B
IK
an
. 25 .
D - FIG
Echinofossulocactus areoles the same
in
is
in
).
25
By
of
on
)
,
of 37
.
,
( of
Fig
.
(
genus
a
areoles usually are unilateral Troll has shown this symmetry the result more active growth the abaxial side the growing point the areole this abaxial growth the areole becomes oval outline along the median cussed
studied
and found that the different types each species were formed
.
As - of
of
.
to
his
-
,
.
a
of
in
or
vascular strand (Fig . 24 ) . Troll ( 37 ) cites an observation by Zuc carini that Pereskia saccharosa one two leaves arise the second year from the middle spine cluster These leaves are formed from the areole growing point and accord ing description the spines may also arise from the growing point With the exception the radial areole Selenicereus Macdonaldiae previously dis
Opuntia ramosissima , and others . The suc cession of spine formation may not be the genera same in the Cactaceae but further research will undoubtedly show that related genera spines are formed Tiegel and Oehme similar progression 36
each spine base he found a
to
(
leading
Also
THE AREOLE
OF
of
STRUCTURE
of
;
of
;
b of -
a
;
I.
.
)
of
a
.
to
is
to
as
the
.
,
of
.
of
,
in
all
In
of
.
.
of
In
.
These
also
Lobivia
,
and remain erect thus central spines the areole Be spines cause the formation slow the spines old become pectinate while the young ones are still growing Fig 26g that
time
of .
so
is
.
developed
a
and
conditions
at
is
spines
cultural
In
of
.
of
becoming
,
in
as
,
the areole
other species
spines arise
median
7
the
In
.
.
3
is
the two sides turn downward
so
in
as
, no
that more spines are formed areole Pereskiopsis pititache monacanthous
center
develop
8
) in
-
are
i
the These the the two pro
Pereskia aculeata where two spines were formed The next spine form the large spine the event
hooked to
.
spines and develop and must
abaxial
phylls
had overlooked 2
.
IV
Wetterwald
be
first
to
.
Fig most
25
in
.
to
of
of
Opuntia spines the formation According Ganong the arborescens spines are formed the series shown studied
. IV -
.
)
-
,
15
and
(
)
47
(
Wetterwald
Ganong
and other spines spine later this becomes the lowest and turned downward this species further spines arise laterally and alternately
formed
.
.
adaxial side Troll terms this form anisophylly
the
the growth
Finally pair very fine hair develop laterally like the medial line the areole These are the two pro phyll spines and because their delicate nature they decay rapidly and are usually appear not seen The next spine spines
is
least the organs
woolly
at
than
of
)
stages
ab
(
( at of
is
in
primordial
the
on
of
.
of
and these organs
this results the formation earlier and therefore lower
on in
the
bract Growth the organs part the areole also increased axial
III
an
of
G -
of ;
a
( all
of
.,
; ;
1, V 2,
B ; of II - - A a -
; ;
3,
of
Diagrammatic scheme Opuntia the development areole axillary growing point prophyll stem areole and spines bristle etc succession The areole primor development dium Theoretical radial areole Theoretical Diagram scheme bilateral areole after Troll areole Opuntia after Ganong bract spines
poor
these spines
re
MORPHOLOGY
14
OF THE AREOLES
IS
3
.
See
at 28
In
).
.
the
a
.
)
30
29
,
.
(
some genera may
also
remain
,
old areoles
or
.
on
no
(
other adaxial the areole At this point the side border the areole flower lateral Figs branch arises and The very
of
the
text
Fig areole forms glochids genera spines are formed
of
,
as
form
Rebutia Pygmaeo For correct classi
be
a
(
.
.
it
of
spine
cluster
,
whole
Trichocereus spachianus and the areoles produce once
but
the areoles
a
at
pasacana
,
.
such
as
,
In
forms
remain
In
.
pasacana
.
new areole wool
old
is an
spine from The new spine
a
of
Development
Trichocereus
by
.Fig . 27
of
of
.
)
.
27
of
In
(
a
is
.
old
,
-t
of
.
of
.
of
further growth Many years later new spines may be formed from the upper part the areole Trichocereus pasacana the areoles are the Lobivia ype with spines may be central spines and new formed from areoles Each the new spines always surrounded by cluster Fig new areole hairs Opuntia the adaxial portion the capable
of
T
others and some
but rather
cinnabarina
may necessary these plants use these relationships some cacti especially the columnar
fication to
Echinopsis
)
,
lobivia
,
lated genera not spines simultaneously
re - in
,
manner described do
the
of
.
.
,
as
long mained soft were not the others and did not turn outward Thus certain irregularities may occur Although many species Lobivia develop spine clusters
Lobivia
.
the spines
of
of
Development
of
. 26 .
Fig
3
areole
surrounded
STRUCTURE OF THE AREOLE
active lateral
not only spines , but branches and fowers . and form
also
itself ) and one, or rarely more , forming parts branch . In Thelocactus ( 15 ) the point of growth of the areole divides into two parts while forming spines . The abaxial portion degenerates and stops the areole
formation of spines , whereas the adaxial portion moves off and remains connected to the other part by a furrow filled with hairs . This adaxial portion becomes the
point of lateral branch ( Fig . 31 ) .
and flower formation
In Coryphantha the upper point of growth remains in the axil of the podarium , where as the spiny areole is raised up with the top
of the mamille (tubercle ) . A hairy furrow connects the two portions . Figs . 32 and 33 illustrate the different forms of areoles re sulting from such divisions of the growing point . These divisions may be explained as
the podarium
If
the podarium lengthened
,
basal part
.
its
follows . As has been shown , the lateral point of growth in the Cactaceae is always displaced from the axil of the bract to
side views
Fiebrigii
Rebutia
the tuber
Aylostera
)
flower bud
of di
in a
of
The outer part
. to
the
parts
the top
.
Areole with
. .
of
two
others
of
the
and
occur
(
of
between
may
of
;
Leuchtenbergia
lengthening
growing point now displaced
this
is
vided
.30
(
:
a
flower bud above the gibbosum Top and
podarium
.
-
.
of
-f
young with Gymnocalycium
.
in
Areole
a
to
a
to
.
29
.
Fig
spine cluster
Secondly
FIG
ab
on
Areoles that form axial part become extremely unilateral This ultimately leads split the growing point spine orming portion form the the
in
occurs
This
only
spines
the tubercle and Alowers top the tubercle
on
areole the top branches arise to
to
Glochids
Photographed
or
.
the center Cordes
by
puberula
.
Opuntia .
.
areole top and hairs
in of
An
FIG . the 28
.
:
;
a
to
becomes tubercle two things may occur First growth may occur below the whole areole This displaces the
AREOLES
.
to
a
the
in
lengthens
because
growth
of
ally
of
.
of
a
in
)
15
(
or
Be -
.
the
the
area
furrow
on
tubercle the axil later becomes succulent between two parts becomes the
cause the tubercle
OF THE
growing point remains axil without furrow connection the areole Ganong changes studied these morphological Coryphantha the meriste and found that matic region the young podarium actu
the upper
in
surface
or
the other part remains
and
of
,
cle
MORPHOLOGY
the
elongated young fiower Thelocactus hexaedro phorus
with
a
Areole bearing
.
.
FIG furrow bud
in
31 .
y
is
of
(
A -
:
)
.
,
;
; b (
of
-g
X
of
; B -p
to
;
is of
;
in
,
.
-
to
to
of
; c e -
;
-
b
V -g
F ) g - : -
;
)
-t
(
(C
; - f -
of
; d -E
in
to
elongated furrow displaced the growing point
areoles longated
,
Leafy sprout Bilateral areole Theoretical radial areole spines Areole type areole with pectinate arrangement Hamalocactus Areole with furrow connected the growing point which oryphantha type axil Areole without connecting furrow the displaced rowing point the spine bearing tissue black the axil Mammillaria ype furrow types
: a -
Schematic
of
32 .
.
FIG
Kdo
vtel
the most frequent type
of 1 .
,
-
is
-
IV -
) ;
:
of a
is
Il ; , -
;
-t
(
of
so
,
)
.
)
)
(
Schematic
.
.
a
Fig
.
(
In
development
mamille Mamille without division the areole growing point only the basal part elon ype gated Thelocactus III Mamille with furrow Coryphantha elongated the part below the areole type Cross section through the mamille III showing the tissue wall about the furrow basal part art below the areole lamina rudiment rowing point the areole shaded After Troll FIG
Mamille primordium
furrow 32e Mammillaria the division the grow ing point occurs very early that the spiny part areole becomes elevated the top upper part podarium the and the the of
33
,
so
is
of
,
as
a
In
Hamatocactus setispinus furrow just described but the upper part formed slightly re only the growing point proper moved from the areole that the Aower arises just above the spine areole and the part above the flower becomes short
ANATOMY AND MORPHOLOGY
OF SPINES
of
;
at
tip
;
of
tip
described the structure of spines , however . In Opuntia rhodantha the spine is formed from the meristem or growing point of the areole . The epidermis of the young spine near the areole consists of many small cells which become prismatic further up the spine. Progressing toward the the spine the prismatic cells become speckled near the the epidermis becomes striped obliquely and the top the spine the
.
.
The middle
.
has been shortened
After Weingart
.
.
)
(
glochid
of a
.
Fig
35
.
of
;
Schematic drawing section the glochid
of
Vi . of
,
v - ; S -
-
F
of
;
;
,
Va -
:
.-
of : M b -
M
:
of
M
,
of
The
to .
of
34 .
.
longitudinal sec Schematic representation FIG through tion the growing point of Coryphantha very show the development mamille primordia young growing point older mamille primordia growing point lamina rudiment the stem the areole outer spine bearing part inner flower bearing part furrow the divided growing point After the old mamille M2
the spine
at
matic speckled cells
tip
to
,
of
in
are
a
is
of
.
cells become scaled with barbed hooks Underlying the epidermis body cells spheroidal which the base the spine pris and which progress through cubic
.
Ganong
podarium
.
of
One end this elongated areole spines and the other end remains The furrow
division
In
).
34
.
the meristem
no
between the ends seedlings
Fig
occurs
so
of
only forms hairs
(
.
meristem
a
forms the
that the
lateral shoots from
form
the
of
of
in
.
a
whole growing point forms spiny areole Therefore young plants which the termi growth has been destroyed may nal point remainder
.
of
of
.
by
.
36
of
part the spine consists prosenchyma Weingart terms the spine the kernel The size
.
of
innermost long core
a
.
glochids
Cordes
of
.
of
this part the lumen these cells decreases gradu ally toward the center the spine The impregnated throughout with cal spine pectic substances cium carbonate and Weingart believes that the absorption
.
of
of
by
is
by
these
.
regulated
glochids which surround Two groups the spines above and below arise the are They quick succession formed in
in
first accurately
spines
of
He
defen
water pectins
are
.
,
Weingart
three
.
.
sive spines
the group
of
placed glochids
( 41 )
also
in
:
,
Ganong classified spines into groups defending soft and glandular
ole
spines
-
of
Anatomy and morphology ( 15 )
.
E
.
is
of
each
Photomicrograph Photograph
of
.
,
into several parts flower
.
a
divides
which forms
In
.
)
in
26
(
in
A
of
.
stems for times stress serial division without separation the growing point occurs some genera Lein Rhipsalis and fellner observed this Myrtillocactus the former genus the areole
FIG
.
a
of
a
to
by
of
.
the areole An old plant Mammillaria pusilla which frost was found had been injured produce spiny areole lateral shoot from The areoles are apparently potential meri
Aowers
Mar
Harrisia all
of
,
OF THE AREOLES
to
38
)
and.
. to
in
,
,
,
,
(
(
.
,
from
(
)
transitory Eriocereus Martinii tinii spines forms from short and thick soft forms spines and hairs Intermediate hairs Fig forms are straight Fisia hard and brown the browspines basal part but become like soft tip
-
of
.
in
are formed such large numbers Glochids have bulb like base small cells which contracts above the areole epidermis Above the base the glochid consists shaft with a
the areoles
in
by.
a
a
is
.
of in
be
in
of
or
groups less The formation twelve groups may serial division caused the glochid primordium Thus each glochid modified leaf even though they
of
MORPHOLOGY
of
of .
do do
as as
.
as
O
.
.
O
in
of
,
.
is
,
.
as
.
to
to
of
,
of
found
]
a
.
)
(
26
,
. -
)
(
.
Leinfellner
At -
.
-ap
.
a
,
.
,
soft spines
cylindrical Daumann states that mon acantha the glands are bottle shaped and are homologous glochids the top the gland the tissue becomes loose nectar Eventually accumulates underneath the epidermis ruptures the liquid extruded and then the stump becomes woody Wein gart states that under certain condi tions spines some Opuntias secrete juice spines thus indicating the relationship glands
forms arise between the first spines formed by the areole Long often hair like spines and hairs are termed
,
to
( u 11 )
( ) or
)
in
described
and
,
as
spines
as
of
intermediate glochids They are long glochids have similar pearance and are completely covered with Usually scaly cells these intermediate
forms twice
),
,
.
to to (
15
(
-
a
by
air
collector
41
) ).
Ganong
the
15
( (
from
25
.
in
as
,
dense water mechanism
a
is
.
-
,
of
a
.
is
in
,
of
)
.
36
35
.
- -
en
.
-
,
(F
of
.
of
,
a
of
.
single hair ward the which often also found soft spines that branch and two sometimes four fork like crossed fork These results indicate that soft spines groups epidermis cells The consists are true spines tirely scaly cells igs species and Be The extra floral nectar glands cylindrical Opuntia Hylocereus Hamatocactus and neath the epidermis are spotted Coryphantha cells with thin walls and the center Sect Glanduligerae are Ganong prosenchyma Essentially homologous oryphantus spines according the glochid spinta glochids are without cuticle and the plant cormólogo They also de and Daumann Daur through velop spines can therefore absorb water the the same manner same Ganong glandular glochids Opuntia spines was observed described the micro dasys by Weingart Spines may also con Opuntia arborescens and imbricata
He
1
e
cd
g h
inte
.
;
; -
h
; -c
; b -
; g -
an
; f-
-
of
k -
;
i-
-
; e -
: a -
of
areoles and areole parts acicular and conical spines curved spine hooked paper like spine tongue like spine plumose spine hair like spine bristle glochids isolated growing point areole
-
37 .
Diagrams
spine
; d -
.
FIG
f
CA MARO
ANATOMY AND MORPHOLOGY
OF
SPINES
Leinfellner ( 26 ) noted that the turbinate gland spines of Hamatocactus setispinus are formed from the growing point of the are ole , and in Coryphantha (Sect. Glanduli gerae ) the gland spines arise from the inner part of the growing point . On young speci mens of Coryphantha clava the glands oc cur both in the axils and on top of the above the spines of the areole .
of
remains
potential
.
point
a
growing
papyra
Opuntia
diademata
of
the
meristem
spines (
cantha
).
Paper like
-
. .41
FIG
tubercles
in
.
in
by
by cultural
is
of
a
of
of
of
of
descriptions spines The use the classification cacti has been considerably overlooked because the apparent vari species The ex ability spines within spines probably caused treme variation
.
an
,
is
, It
,
in
.
, h
, of
in to
(8 )
is
as
in
schemes
descriptions
this characteristic proposed several spine arrangements for use
Buxbaum
of
.
variable
used
in
spines could great advantage
be
.
of
,
as
,
,
an
have been formed from areole the growing point becomes very hard and woody thus firmly holding the spines Nevertheless has been shown
of
spines
.
typ
do
cultural factors not affect the arrange spines Therefore the arrangement ment
.
all
After
.
in
the
etc
.
to E )
of (A
and areole hair Eriocereus Martinii After Leinfellner ( F )
.
38
.
FIG
or
Hair spines
all too
,
., ing ,
tender spines culture Insufficient light ing too much organic matter the soil spine affect mffect the type extremely important fact fact however that
OAN
trichome
of
conditions rather than herent factors the plant Specimens that ordinarily possess strong spines often have
.
( N . )
-
hooked central spine and nectar spine tongue like central spine latis pinus with a
a
of
:
.
setispinus with Hamatocactus Right Areole Ferocactus
. of 40 .
Left Areole Schumann FIG
:
.
FIG
39 .
SUN
After
Mammillaria Schiedeana with Photograph Cordes by
Areoles plumose
THE AREOLES
OF
spines
.
:
of
43 .
.
of
.
of
:
greatly enlarged
of
,
,
;
a
.
)
of
in
.
41
;
, .
,
O
.
.
or
,
(
.
).
42
of
of
is
as
,
of
a
is
.
a
of
.
to
of
, (O
,
,
acicular
wiry forms
.
(
others Fig The soft spines
and
in
,
,
-
to
as
)
.
cereus senilis and Echinocereus De Laetii many spines resembling horse hair cephalias the hair like often soft and silky radial spines Mammillaria bocasana
45 ).
Fig
.
.
M
)
.
44
.
(F
,
in
of
(
of
mature areoles often seedlings are similar those seedlings Plumose spines occur Mam plumosa millaria and these are retained dur ing later growth ig Schiedeana grows similarly and Neolloydia Beguinii spines retains plumose about two years pusilla Mammillaria prolifera and related
.
)
to
thin
cinnabarina spines begins with the type progresses Alexible puntia leucotricha Cephalo
series
(
,
to
of
Lobivia
Another
,
,
.
F
of
)
.
40
;
case
to
, , -
,
.
,
,
of
- .
39
.
groups especially
areoles are densely covered by two ranks short and flat spines arranged like the spine ar This form comb teeth rangement derived from common pecti the radial spines nate arrangement
,
,
in
, ,
of
of
)
spines
large ones are annulated The ribbed Pelecyphora aselliformis are inter Fig estingly arranged Here the long
spines
in
of
.
a
in
in
on
. all
(
-
different
Many
of
.
in
,
in
cm
25
,
,
-
in
a
,
of
In
.
to
occurs
the same family
or
.
or to
an
up
of
(
shape
spine
shapes and multitude exist Browningia sizes candelaris and Corryo very brevistylus thin acicular spines cactus length and short conic spines that hardly appear above the areole Hooked the same plant wool occur Austro spines may occur Hamatocactus Mammillaria cactus Parodia Mamillopsis Phellosperma Ferocactus Ancistrocactus and others which are very different and unrelated genera Also interest are the flattened tongue shaped and curved central spines Ferocactus latispinus and strong spines the curved Wislizenii and others Figs and Modification tongue shaped spines leads the weak scarcely pricking spines Turbinicarpus Ariocarpus and others and further the Spines
Turpinii Toumeya and Fig Again certain
43 ) of . of
.
used
Spines usually arise symmetrically and ascending rank about the plant perpen Either they form the bract dicular figure elliptical above the areole circular the upper but below the growing point Usually part the areole remains spineless greater number the radial spines arise than central spines but the latter are more striking size and shape
diademata
Leuchtenbergia
(
they could readily
Opuntia sub papyracantha
parchment like paper spines tribe Tephrocactus Opuntia
.
entirely
0
).
(
not
are
to
schemes
,
these
correct morphologically be
Fig
and taxonomic studies
Although
.
systematic
37
.
.
Fig
:
Center aselliformis Fig Mammillaria plumosa with
Pelecyphora Areoles Right Areole
.
.
.
Left hair spines
44
42 .
FIG
MORPHOLOGY
,
20
ANATOMY AND MORPHOLOGY
21
OF SPINES
22
-- -
These spines are enclosed
.
fication
.
;
;
-
c
;
. tip
.
.
)
46
..
.
Saussieri
Na
)
15
(F
or
Thelocactus guelzowiana
colors As spines mature they gradu flect ally lose the color and become greyish Along with the color change the spine may become fragmented
yellowish ig white sheath Ganong believed that the sheath was formed by areole hairs which had grown together Weingart opposes this viewpoint
hudum
.
(
Poselgeri
Mammillaria
a
by
a
-
of
. .
to
retain
Cochemiea
ingens
.
short hairy spines which are very similar the acicular seedling spines North American Cylin The spines very interesting modi dropuntias exhibit
forms
guinii
; bf-
a
Be
Echinocactus
all
-
Neolloydia
; -
grabliana
Frailea
e
seedling spines d
Types
of :
of
. 45 .
Fig
e
and suggests the calyptra
the sheath formed like moss capsule the young way upward underneath the
a
as
of
is
that
sil
as
worthy
hulle
, of
is
It
layers above
it .
there are
no
is
.
be
its
spine forces areole layers This cannot correct the spine formed from the growing point and
.)
immature
are
Spines
especially
when
,
, .
or
(
,
as
,
mention that was the case with spines light etc under certain conditions low the sheaths become stunted are not even formed
)
31
.
at
a -
:
Opuntia iunicata Areole with sheathed Single spine with the sheath cut off the base show the spine After Schumann
.
FIG spines
to
,
re do
,
not
and
.
-
color was composed black yellow White spines actually any pigments contain but merely
,
red
b -
of
brown black
. ; 46 .
-
of
.
(
Sadovsky often strongly colored specimens Astrophytum cross bred with different colored spines and found that the
22
SUCCULENCE
OF THE CACTACEAE
leaf characteristics are especially except that the leaf usually very large and green and the
These
,
The leaf
discussed
.
now
.
reduced
be
will
,
leaf blade
,
a
)
.
a
cm
a
of
27
of (
)
18
(
. .
in
a
.
cm
10
,
of
of
,
.
of .
of
.
po
a
the branches are slender and The leaf blade however well developed these species and reaches Opuntia length more than
,
,
and
switch like
is
darium
in
.
cm
10
THE
Leaves
As
.
A
SUCCULENCE CACTACEAE
OF
.
in
or
18
.
is
it
to
in
aid
or
of
.
2
true
The stems these genera are slender and become woody rapidly There fore these plants are leaf succulents not stem succulents Other members the Opuntieae and also the genus Maihuenia the Pereskieae have fleshy cylindrical leaves which usually are small and decay rapidly The leaves the most primitive species Cylindropuntia not have distinct do
of
.by
to
”
"
so
In
Pereskia aculeata the first two slightly fastening the plant hooked spines growing upon the shrub tree which Spines epiphytic spe other climbing cies function the same manner
podarium
but the
in
off
)
(C
passing
these leaves are
of
to
of
.
as
Bigelovii break ylindropuntia and easily that the common name adhere jumping cholla be this plant has become jump onto one cause the joints seem
of ,
of
. of as
of
In
.
in
the species
Most
succulent and contain mucous cells leaf bases are not developed into
.
of
,
bya
as
to
or
.
is
propagation
vegetative
the lateral joints cling by the spines ani Opuntia they brush by The joints mals
width
.
.
There are apparently many functions spines The spine may act defense against being eaten mutilated animals but the ability condense water from the probably the most important function air Spines which are accompanied with dense areole wool may also decrease the rate transpiration many cases the spines aid
veloped leaves and some them even have distinct petioles These leaves may attain remarkable size Goebel found that the largest ones Pereskia Rhodocactus grandifolia reach length and
of
Backeberg
After
of
leaf
-
of
reduced
microsphaericus
a
with
Areole
Arubrocereus
an
of
.
a
in
.
of
to
of
.
a
,
of
16
of
Figure shows the growing point Opuntia with the blade part the leaf primordia distinctly separated from the leaf by contraction This contraction base must be homologous the petiole other dicotyledonous Early cessation plants leaf growth results reduced leaf The most primitive genera Pereskia the Pereskieae and Pereskiopsis and Quia bentia the Opuntieae have well de
. or
Fig
. 47 .
scale
or
,
,
greatly
leaf blade
is
is
base
the Cactaceae
in
true
a
48
.
of
:
of
.
;
D
-
)
(
;
spine leaf primordia axillary bud areole After Wetterwald
.
- B
,
)
-Bj(V
.
.
hair
of
podarium
Longitudinal section grow FIG ing point Cereus peruvianus B
do
,
leaves
but have intercalary the leaf base often covers the
h
tip ,
meristem
;
the
the
stem
-
form
,
)
and
the
.
stem
to
;
growth
at
grow
a
dermal cells
not
of
.
a
(
,
of
,
was shown earlier the dicotyledonous leaf consists leaf base and the laminar part blade the two often being separated by petiole Also leaves ggenerally are enereparated epidermal and subepi formed from the epidaves
-
LEAVES
Fig . 49. Longitudinal sections of stem primordia The leaf primordia B1
( a v B2 ) tip , ) v -s of of
a
no
( a )
,
B
.
.
(b , ) V -
a
at
.
is
reduced and there
region
.
,
Nevertheless some small scale like leaves below the very young areoles This occurs Hylocereus Rhipsalis Epiphyllum Pfeif in
,
,
. -
have
of ,
genera
Cereeae the
the
(
.
; c
-
a
; b -
in a
a
.
is
it
is
is
In
).
47
.
(
of
of
on
-
,
of
laminar part
great use because the scales acter not rapidly decay after appearing only very young shoots Figure shows the development the
of
the
of
In .
or
of
in
genera
.
.
podaria
primitive development
of
-
or
of
leaves
Mammillaria
microsphaericus Fig the latter spe Backeberg cies believes that the scale leaf specific but actually probably not only specific characteristic and stage the evolution the group Also the char
48
Thus the growth follows acro tony the blade whereas the more advanced genera the Cactaceae development follow basitony the leaf base
have hunch like
Cereus
,
,
leaves and podaria and the more Cylindropuntia species which
Opuntia
is
advanced
schemes
of
cylindrical
Prototype
of :
. 50 .
Fig
a
w
kwon
"
)
Nycto Acanthocereus some species cereus and Anthrocereus Arthrocereus
fera
,
.
is
Maihuenia which has small
always greatly
petiolar
,
illustrated
by
be
a
the leaf blade becomes reduced the leaf base becomes more distinct and well developed podarium formed This may
leaf
,
of
As
.
in
an
.
a
subulata These leaves persist for long time and play active role the physiology the plant
is
.
,
in
tip . , h -
Bz -
B -,
.tip
,
B
(
.
Mammillaria coronaria Mamille points bear axillary growing has spine producing part not divided and has divided into the top and secondary growing point the axil the leaf primordium Whole econdary axillary meristems stem leaf primordia hairs stem After Wetterwald
SUCCULENCE
24
OF THE CACTACEAE
in in
, ,
in
and only
the leaf base enlarges
.
blade
microscopic
a
a a
of
into
as
.
in
leaves Cereus peruvianus Here the leaf Opuntia but primordium develops but differentiating into leattia stead leaf base and distinct blade theting the laminar part remains
of
to
to
of ,
is
of
the tubercle and the tissue that usually grow ceases Thus to
-p
W
,
,
.
.
near the cotyledons which are Later the cotyledons drop off Rafinesquei compressa one cotyledon Fig also slightly shorter than the other
. (
.
),
of
53
.
in
the seed
.
a
.
of
-
,
of
,
.
of
.
-
and
each
of
is
small areole with min The primary shoot
.
)
.
(
Fig
54
ute spines
is
the first leaves has
exposed
,
now
seedlings
a
have large cotyledons slightly Pereskia but the hypocotyl
is
The epicotyl
51 ).
.
(
shows the growth
the seed coats The hard testa ruptured and the primary root hypocotyl and lower part the cotyle By gradual enlargement dons emerge the cotyledons the seed coats are shaken off
of
.
,
(
.
O 52 is
to
is
in
,
is
ling out becomes
,
in
as
Opuntia
Fig
.
of is
of
)
of
the seedling similar stems but the cotyledons may be well developed even species which the mature plant leafless The seed any other ling Pereskia like that dicotyledonous plant The slender hypo cotyl bears two large and non succulent cotyledons and the first true leaves have normal blade and petiole Fig leaves and
In
,
feshy
.
.
B
The evolution
that
rimary
After Troll
thickened
and seedlings
grandi
Pereskia
leaf
-
of
-
,
-
:
root
Cotyledons
seedlings
hypocotyl
P
cotyledons
H
Co
.Fig 51 .
in to
,
is
Different ages
of
A
.
(F
flora
.
.
Fig
the evolution from leafy shown tuberculate forms
50
,
leaftess
to
sentation
49 )
in
leaf blades are formed the Mammil ig diagrammatic repre
larieae
of .
no
forms the leaf blade
.
form
b
In
the first stage the development leaf primordia similar the other groups but during further growth the basal part the primordium enlarges the Mammillarieae
The lamina
)
testa
(
The seed coat
. .55
the epicotyl
FIG
. 53 .
Fig
.
size
of
Old seed the first leaves have already dropped
.
of
the
terete and ovate
unequal
Development
.
stem
cotyledons
of
that the primary
showing
the cotyledons
FIG
Rafinesquei
enlargement
. . 54 .
ling showing
Opuntia
.
removed
is
Seedling
of
FIG
becomes
by of
. 52 .
55
COTYLEDONS AND SEEDLINGS
.
)
.
the axils
and
of
.
),
After Berger in
areoles 56
of
cotyledonary buds the cotyledons
Hylocereils
.Figor
Seedling
if (
,
.
.
or
. 58 .
.
to
so
the epicotyl con The succulence that the hypocotyl downward even becomes swollen this structure spherical Sometimes the primary root also Most seedlings have succulent becomes stem
tinues
FIG
observed
has characteristics
and
that the seedling the mature plant
of of
( 22 )
.
Leinfellner
is
.
)
.
57
(
)
26
(
destroyed found that the epicotyl these cotyledonary buds develop normal shoots Fig
.
(
.
)
on 58
( 3 )
as
.
a
,
It
of
appears that the genera Cereanae are very primitive because they have large coty rather slender hypocotyl these dons and structures not differing greatly from the very primitive Pereskieae The Hylo especially cereanae have well developed Berger Fig cotyledons has shown Hylocereus also has quite large leaves
a
is
.
a
.
to
( 3 ) of
.
as
a
of
it
of
,, B
,
.
-
of
-
C
D -
.
,
,
,
S -
A - of
B2 -
-
:
.
.
56
seedling Longitudinal section cotyledons Macdonaldiae Selenicereus the cotyledons CA axillary areoles spine pri areole leaf primordia growing point After Leinfellner mordia Fig
a
(
(
of
.Fig . , , c ) ) .
to
.
of
Epi the young stem The seedlings phyllum are the only present day plants similar these Fig 590 Because seedlings Rhipsalis 59b have succulent hypocotyl and short half conical cotyle dons shows that this more recent primitive genus the Cereeae rather than Backeberg believes He states that group they are the most primitive members the According Berger sub family the genus Lemaireocereus has comparatively large cotyledons and succulent hypocotyl Backeberg also places this genus with the of
.
primitive forms At this point the seedling
Marginato
it
of
is
)
by
. (
marginatus cereus Lemaireocereus interest This species has been separated Backeberg because from Pachycereus
.
"
.
).
a
60
.
59d
. ) .
so
Eriocereus ig these
(
Harrisia quite similar
(F
are
to
seedlings
of
of a
( K )
of
The
Guelichii
.
.
.
,
Longitudinal
through section Macdonaldiae Selenicereus which the terminal growing point has been damaged The cotyledonary areoles form After Leinfellner lateral shoots seedling
.
57
Fig
mm
.
(
is
.
has flowers much different from the Pachy cereus type Not only the flower but also primi the seedling indicates that this plant tive Fig The seedling has succulent conical hypocotyl and very large cotyle dons During germination the cotyledons are not succulent but they become later
)
of
Jonsson stem
16
Ganong
seedling structure
(
is a
In
to
.
)
55
.Fig
succulent and shows many characteristics of The first leaves the mature plant ( ground rapidly dry and fall the great variability the Cereeae there
) ; , dH (
-h
;
)
f
.
. al )
,
-
it ,
of
( the
.
is a
,
,
an
As
it
a
.)
63 ).
.
,
a
in
.
,
) (
(
(
to
II .)
63
.
(F
-
as
,
to
in
ypo
ly
is
of
-
In
the short columnar and globular spe more the Cereanae the hypocotyl cies globular and the hypocotyl less ovate lengthens the short columnar species or
to
-
(
c
( d
a
-
(
).
,
59e
f
in
.
have Erio
also
Harrisia
a
.
as
(
,
)
may
center
the plant matures Thus the seedling ready shows the habit mature plant example Morawetzia Cleistocactus short columnar form Doelziana which comparatively thick hypocotyl but has highly evolved species because has Berger showed Fig small cotyledons Binghamia Borzicactus Seticereus Bckg be similar Copiapoa cinerea which re globular form for some time be mains forming thick short columns has near fore spherical seedlings with short cotyledons ig is
)
.
62 )
.
.
) (
)
.
. ,
( is F
.
)
of
44
(
Weisse described the seedlings some spherical and short cacti Hamato cactus hamatacanthus has long acute cotyle Fig spherical dons and the hypocotyl 64a Ferocactus pilosus Stainesii has
,
.
)
.
of
A
.
)
e
,
(
a
)
(
c
,
(
very small succulent cotyledons Fig 64b whereas Eriocactus Notocactus Len inghausii has globular hypocotyl bearing Fig broad and short succulent cotyledons longitudinal view 64d Eriocactus
)
elongated
to
is
be
of
).
or
the globular
hypocotyl
This occurs
slightly
in
Le .
distinguishable from
.
:
.
.
:
.
of(
. of
Seedling Marginatocereus Fig 61 Right Seedling Monvillea phainosperma
Left
maireocereus
)
. 60 .
FIG
so
(F
of
,
as
(
Notocactus Schumannianus shows clearly buds developed areoles and displaced onto the cotyledons the characteristic most the Cereeae ig 64f Finally the cotyledons may completely reduced that they are hardly cotyledonary
.
(F
(
J
-
.
-
or
a
of
Many the columnar Cereanae have cylindrical hypocotyl and more less more semi conical cotyledons Monvillea short semi conical Fig 61 and phatnosperma and Cephalocereus ig that TrollCephieves believes that illustrate this some columnar species shorter hypocotyl Fig cereus Bonplandii
THE CACTACEAE
Co -
;
b
)
, W -p
f-
;
and
OF
Rhipsalis Cassytha after Irmisch after Berger and cotyledons Eriocereus Bon plandii after Troll cotyl rimary root
(
Epiphyllum
Guelichii
e
(
)
Harrisia
-
Seedlings
Eriocereus
a
of :
. 59 .
Fig
SUCCULENCE
)
26
Melo
COTYLEDONS AND SEEDLINGS
FIG . 62. Different ages of seedlings
After C . Meritz , photograph
.
Doelziana ;
Copiapoa
I
cinerea
.
of : la, b, c. d- Morawetzia
-
Seedlings
by Cordes
m® ( * la
b
.
.
AB
ke
цк
.
.
; of
(
of
-c
; K , -
;
; b
e -
;
; h -
)
)
, f B - ) ; -c
Longitudinal section Hamatocactus hamatacanthus Ferocactus pilosus Stanesii Longitudinal section Leninghausii same Eriocactus Schumannianus Notocactus otyledons Kz leaf After Weisse Melocactus Maxonii Cactus Zuccarinii
-
Seedlings Eriocactus Melocactus
( of : a
same
(
.
FIG
g - ; 64 d . -
h
Ke
,
CMC
6
4
B2
.
--1
Ka
Fig . 63
Sartorianus
IIa ,
Om
of Cephalocereus
28
SUCCULENCE
OF THE CACTACEAE
;
-p
.
.
.
)
(
to
B
,
;
).
K
(p
of
.
in
.
55 )
(F
.
as
, ,
old
in
the
,
in
-
,
become
.
elevated
of
on
.
-
.
to
of
.
it
,
)
or
.
.
cm
10
noticed that
of
( 44 )
.
pilosus
Ferocactus
4
in
)
in
whorls
the same species
Echi
.
In
rows
parted
alter
-
five
found
3
he
in
seedlings
;
nating
of
,
Other
gib
Gymnocalycium
.
G
in
;
)
.
(F
-
.
4
parted var fexox Weisse observed alternating whorls ig 68a and
bosum
.
)
.
a
in
Saglionis the podaria disperse clockwise spiral Fig 68b The same formation was
(
.
a
.
of
In
.
of
.
, .
de -
of
,
is
it
ig
:
.
rows
nocactus Gerardii
,
flattened
Stainesii had verticillate podaria parted whorls thus forming eight vertical parte had areoles dispersed
of
- : a K2 - -
tip
of
In
.
of
Weisse
diameter seedlings
F
. -
.
)
of
dispersed
Quiabentia
Tacinga have not been probable that they resemble scribed Opuntia the small those Pereskia terete blade the first leaves the seed ling are much smaller than the cotyledons podarium The leaf base rapidly forms which bears the axillary bud The podaria
areoles
maturation these form ribs After further development two arrangements are possi ble the areoles may become whorled
Pereskia genera have seedlings like those Pereskiopsis Although the seedlings and
FIG
Post seedling development the Cereeae has been very incompletely studied Gen appears that axillary buds erally the cotyledons form the first areoles The leaves odaria the young shoot arise exactly alternate the cotyledons and form small bearing axillary areoles tubercles After
(p
-
-
.
of (F
,
of
,
67
.
(
the most primitive
After Weisse
Rebutia deminuala small humps which
miana the podaria soon become because they grow only laterally Thus the young joints rapidly smooth and succulent without any podaria
in
re
In
.
)
.
(
in
(
as
,
,
In
lar
In
the Opuntieae only
ongi si
Cylindropuntia are quite distinct which usually has hump like podaria plants Platyopuntia and perhaps some Opuntia sal the Cylindropuntias
(
).
.
.
in
,
to
of
,
as
.
of
of
,
a
is
of
(
.
h
,
to
of
its
of
to
in
a
)
).
the
,
b
Rebutia and related genera also reduced cotyledons thus show ing advanced evolution these genera Rebutia subgenus Aylostera the globu seedlings have such extremely reduced cotyledons that only the podaria and the Fig displaced areoles remain very
cotyledons
are reduced have cotyledonary
'
in
)
(
at
of
-
of
only Rapicactus border Thelo cactus subterraneus Fig 65 Mam millaria rhodantha the axillary areoles the cotyledons become slightly elevated ig 66a
cotyledons
The
cactus Cactus which the cotyledons are very short and rounded form two short lobes the top the seedling Fig 64g not the cotyledons The reduction primordia rather but leaf reduced result the cotyledonary blade the reduction nearly this the Because absence cotyledonary buds are displaced onto the Fig podaria 5ob the primordia hypocotyl Enlargement the causes the podarium become flattened and this the reduced cotyledons which form sults
have
:
67 .
. of
of
of
K
subter
cotyledons
.
; (
Thelocactus
raneus
)
Rapicactus
seedling
-
. 65 .
FIG
Upper part
Seedling
Mammillaria rhodantha tudinal section stem odaria first leaves mordia Right Seedling Aylostera
; b-L
. 66 .
FIG
NODARI
COTYLEDONS AND SEEDLINGS
O
old
FIG
10
of
-
.
.
, (F
in
.
.
).
in
in
In
.
in
to
.
(F
).
of
Melocactus babien
of
)
in
two cases Notocactus
(
,
.
hausii and
in
,
)
in
(
44
In
,
Hamatocactus hamatacan found dispersed tuber eight parallel rows but also
one case five and rows Eriocactus
thirteen
Lening
, 9 ,
8
of
in
or
in
(
.
)
Schumannianus have dispersed podaria the form ribs and the latter species ribs are E
Notocactus
,
-
( 5 - -de ) ).
,
seedlings
thus Weisse cles usually
.
E .
.
seedlings
of
arrangement the immature podaria are formed After Brückner
of
vo of
the types
of
of
Schemes
.
M
in
,
.
M
species with few large Negrii and depressus the second type the first areoles
va
4
in
of
-
- 4-
a
in
10
.
(F
-
4
In
.
.
. .69
FIG
In in
as
,
ribs
others
ig
species that are
the different varieties
seedling found three different forms velopment the first form he found regu larly alternating parted whorls ig 69a
This type occurs
This usually occurs
re
2
1
.
, 8 . , , . 9 , of . 7
.
in
In
Brückner
Melocactus
but
large with many ribs the third type the podaria form comparatively large humps irregularly dispersed form fashion many ribs ig 69c This type occurs
6
, , 5 , 4
3
)
(
to
)
(
69b
the rows become irregular studying
with the cotyledonary areoles arise four parallel rows
later areoles
u
podaria
and stand alternate the cotyledons The next are oles and are formed alternate with the cotyledons and first leaf areoles parted whorl Areoles and alter nate again with the first whorl This all eight vertical rows Toward the top sults leaves
alternate
an
.
G
.
Quehlianum
Jussieui seedlings
sis .
44
the first
G
In
of
in
observed Buxbaum the development
.
by
in a
b -
of
.
var
.
an
-
a . -
. 68 .
gib seedling Gymnocalycium Cross section through parted ferox The mamille primordia arise alternate Saglionis Top view plant Gymnocalycium The mamille whorls dispersed arrangement primordia arise After Weisse bosum
Melocactus
before
ribs
SUCCULENCE OF THE CACTACEAE
the
seedling Mam the dispersed
.
of
, .
of
-
In
.
)
.
,
. 70
N
,
-
;
K
to 9 )
are
the
form
the Mammil ig
.
.
of
podaria
Astrophytum asterias that seedlings ribs almost immediately especially the some the Opuntieae Cylindropuntia North American species
found
the first year
seedlings
larieae have dispersed
Ko
Weisse
(F
of
at
the end
all
,
In
formed only general
After
areoles
of
podaria
arrangement cotyledonary
( 1
millaria mutabilis showing
of
.
Fig . 71. Young lateral shoot of Piptanthocereus ( Cereus) argenti distinctly nensis showing separated podaria
of
part
Upper
of
70
.Fig
C
,
in
it
to
Although data insufficient state any definite conclusions appears that po daria are first formed whorls and become
is
.
,
,
of
In
.
dispersed later the advanced Cactaceae the dispersion begins early with the first podaria and the divergence the whorls great whereas the primitive genera have only slight divergence
by
in
grossly the
Cactus seedlings have been study and therefore
passed
data
and
.
C
.
to
of
on It
.
conclusions possible are greatly lacking seems probable that further research seedlings and their development may lend much the morphology and concepts the Cactaceae
Ribs
.
By
.
, As
.
at
or
of
,
in
,
48
in
)
48
(
Zuccarini stated that ribs arise from podaria arranged Fig vertical rows podaria separately shows the arise even ribbed species the growing point elongation the stem each podarium can
a
.
From Cact 60
.
.
The podaria Succ Jour
. .
Grusonia santamaria
into true ribs Am
.
of
Joints
are joined
., 6 :
)
37
(
,
Fig
.
at
In
of
of
71
a
of
.
(
)
.
shows the separate origin young shoot Piptanthocereus Cereus argentinensis the genus Echi nofossulocactus the plants remain tubercu late for least one year before forming many ribs On the other hand Troll podaria
. 72 .
.Fig
rib
so
,
to
become isolated form tubercles each may become stretched longitudi podarium nally that each vertical row becomes
RIBS
by
)
.
etc .
,
, . ,
to
is
%
-
.
5
a a
% 5
can
.
.
a
is
in
.
A
,
to
in
be
,
in
.
,
to
In
.
.
of
”
of
“
as
the forma tortuosa the spe Euphorbia Some columnar species show this characteristic and Werdermann Pip specimen has found such ig 74a tanthocereus Cereus peruvianus
known
,
.
(F
)
(
)
a
of
,
.
cies
Werdermann believed that this peculiar growth was caused by bending the plant some tension He prob is
a
.
by
of
of
ribs depends upon the
is is
of
er
%
( 4 )
only Bilhuber and Troll state that ribs formed when the leaves are dispersed angle consistent form exact vertical rows This usually occurs the Cactaceae but sometimes plants with irregular disper sion patterns form ribs these cases no vertical rows occur but the podaria join spiral ribs These abnormal plants are form
form
.
The number
myriostigma has whereas dispersion and forms five ribs
asterias
of
cent old ribs
di
is
is
is
,
,
.
)
.
(
Also
,
.
35
,
Thus the fifth leaf the last leaf the first spiral the sixth leaf above the first the seventh above the second etc T The he cond ribbed stem this dispersion result which common dispersion Another commopersion According Bilhuber forms eight ribs Astrophytum dispersion found the stem
46
of
,
or
-
- of
two
the formation may merely push aside the adja 73b
(
in
of
or
of
or
in
.
of
is
usually
.
. -
of
the termi of
by
is
to
rib
increased
and the formation
the whorl plants the This indi
podarium leaf rectly above the first leaf the whole series leaves having spiralled completely around
)
,
be
to
)
(
In
.
of
ribs
of
rib
leaves
sixth
a
of is
as
- of a
of
.
of
of
.
of
.
the number
nation one new ribs Fig a
factor
cates that the
(
in
of
,
to
of
, or or
of .
of
( 3 )
a
is
and some Rhipsalidanae The number ribs may also change because the nutri tion the plant Poor nutrition causes reduction the number ribs and good nutrition causes more ribs formed Fig 73a shows the lateral branch Pip argentinensis tanthocereus Cereus which formed more ribs after the application Epiphyllum fertilizer and some other genera two ribs may unite form one
new
the number
podaria
the leaves
many other dicotyledonous
dispersion
b
.
Berger states that young plants have ribs As the plant small number ribs increases becomes older the number by insertion the number ribs new two ribs three decreased ribs Epiphyllum the flat shoots the case
only
dispersion
.
-
).
72
in
do
.
as
(
to
in
Grusonia
As
upon
of
-
.
as
of
in
-
is
in
cal rows that unite form ribs Fig they The ribs form the Cereeae
angle
of
In
of
no
do
,
in
as
.
of
-
as
hump like projections the podaria remain throughout the life the plant some cases the podaria become flattened the joint enlarges but Opuntia species the podaria become joined ribs Gru sonia the only sub genus the Opuntieae which podaria are formed dense verti
of
of
of
rib . b -
Fig . 73 . Pipianthocereus ( Cereus ) argentinensis : a- The reduction of the number of Multiplication ribs by the sudden termination of one ribs the termina one rib and the formation two new ribs tion
SUCCULENCE
OF
THE CACTACEAE
though not so striking , spiral ribs also occur Ottonis and others . Many of the Cactaceae have the podaria arranged in whorls even in the old plant . in Notocactus
The whorls alternate so that the leaves of every second whorl are above each other . Mrs. Helia Bravo shows a fine picture of alternating whorls of Myrtillocactus geo metrizans in her book Las Cactaceas de Mexico . Because the whorls alternate there are twice as many ribs as there are parts of
each whorl . According to Bilhuber ( 4 ) , for example , Selenicereus Macdonaldiae has
-
3
.
75 )
.
,
(F
rib
alternate 3 - parted whorls and therefore six ribs ; Aporocactus flagelliformis has 6 -parted whorls and twelve ribs; and often plants with 4 -parted whorls and eight ribs occur . Rhipsalis paradoxa exhibits a very inter ig esting type of formation This parted whorls but the podaria species has only along one internode are elongated
.
rather than into the second whorl There each internode has three ribs which the three ribs
the next
.
alternate with internode
of
fore
to
,
.
25
of
.
55
35
Most
multicostatus has over ribs other species this genus have
E
14
,
to
10
22
100to
14
.
E
E
,
. .
of
Echinofossulocactus has the greatest num coptonogonus has ber ribs gladiatus ribs and ribs but has
76
ribs
are
ribs are inserted
new
old
or
of
as
, ).
(
.
ribs Fig As previously mentioned the number and dispersion ribs may become changed
sometimes
termi
Under poor cultivation some species have of
of
.
.
a
.
P
or
of
of
arted
whorls
.
the next whorl
of
3 -p
The alternating ribs which extend only After Troll
.
paradoxa
.
of
poor cultivation ribs four because Astrophytum myriostigma and other spe
to
alternating
Al -
.
of
in
Rhipsalis
gonus cereus vianus
(
of
or
of
a
Branch podaria result
.
.
Fig
75
of
is
to
.
ably mistaken because the top view the plant shows the ribs arising spirally from This abnormal growth the growing point likely change more due mutation hereditary makeup plant the the
(
.
vation
Many
of
changed
)
a
,
.
by
.
spiral ribs
ribs and the angle Under good culti the normal number arises again the garden plants Cereus tetra Piptantho are actually specimens peru Cereus argentinensis of number reduced which have
reduced number
dispersion
)
monstrous form abnormal angle become joined into photographed After Werdermann Kuhlmann
is
cactus
Cereus peruvianus dispersion the podaria
.
Turning
By", an a
"
.
74
.
of of Fig
.
nated
of
.
-
A 5
:
Astrophytum
but also the form
Echinofossulo
.
varied
,
;
;
of -
,
cactus has very thin sheet like often undu Echinocactanae late ribs many species have sharply angular ribs and Trichocereus
.
Most
of
ribs
candicans has rounded
, ),
.
(F
In
77
ig Astrophytum asterias Lophophora Williamsii
ribs nearly
the
have flat slightly rounded
,
Gymnocalyciums
and
the flat and are separated ribs are completely only intermediate fine furrows Also nature ribs exist forms
.
in
.
all
in
of by
of
do
be
is
, or
of
, . if .
of
The outline the ribs worthy men perfectly tion too The margin may straight the podaria the areoles not project Very often the podaria are greatly as
of
the number greatly
is
of -
12
,
is
of
be
.
areoles
only
ribs
.
of
,
of
a
in
,
If
ribbed specimen From Britton and Rose
so
(
).
(
is
on
it
,
(
)
examined the dispersion Weisse and the Rhipsalides some cacti especially mechanical caused by believed that mod view leaf position but affect morphology his concepts cannot with ern podaria the the dispersion stand these must but result shoots known flat well with the fat joints not confused Opuntia and Epiphyllanthus obtusangulus which have flattened joints covered with
.
Not
occur
also
43
-
ribbed
Fig
76
are
variety
ribbed
. 5-
which normally
as a 4
in
as in
-
or
.
,
elevated Ferocactus Stainesii also Malacocarpus thickened Vorwerck ianus Very frequently fine cross lines wrin
,
in
on
of .
.
in
of
as
,
-
be ,
of
In
aid
of
.
in
,
in
kles occur between the areoles both col umnar and short forms These aid shrink age the plant during dry periods and short species they basal shrinkage the latter case the wrinkles introduce the formation tubercles will be shown Fine cross lines may observed ribs Cephalocereus senilis many species Echi
.
,
and many the cross
in
.
a
,
.
asterias
Jour Bull
.
Succ
.
Cact
of
Flat ribs
From
.
.
77
.
FIG
others
of
of
species Notocactus By gradual deepening wrinkles flat tubercles arranged
nocereus
Astrophytum
,
cies
Right
4 to 8 ) . .
Many ribbed Echinofossulocactus Zacatecasensis myriostigma number ribs vary from of
Left
:
.
76
.
FIG
RIBS
straight
OF THE CACTACEAE
SUCCULENCE
type .
ium above the areole causes this third
The biological effect of furrowed and humped ribs may be that they enable the plant to shrink during dry periods . A longitudinal folding straight resists
, of
a
in
a
.
,
is
rib
of
of of
of
is
rib
,
-
as in
of
.
in
.
of
of
in
in
is
In In
)
winged
(
. ).
-
.
.
three wings are alated
the whole
,
platense
and
.
G
.
species
G
Some
,
have chin like tubercles and others
-
stellatum
,
.
them
as
-
in
.
.
,
G
.
82
by
-
a
G
.
-
.
G
in
G
.
.
,
.
or be
rib
of
.
G
.
fine cross wrinkles The ribs become Bodenbenderianum and other fissured Saglionis species Monvillei and others have the ribs cleft into tubercles The Gymnocalycium humps often are below the areoles with the cross wrinkle just above
growth hump
the odd also the cause the areoles Hylo angles and hooks which occur hamatus cereus calcaratus and Selenicereus ig the latter species these aid Hylocereus triangularis the climbing
(F
is
,
.
in
.
be
nigri and denudatum lar evolution humped slightly ribs distinct have areolum
with
at
is
; F -
: d -
.
are
clearly This evolution formed Discocactus Hartmannii which shown has rounded ribs cut by deep wrinkles into connected fat humps These may they may even small bridges tissue totally separated shows simi The genus Gymnocalycium
rows
and
to
of :
the
of
Fig to . ( 78 - .
a
mamilles schemes cross Neowerdermannia podaria wrinkle between
rapid
formed the the areole part the the part but sometimes growth favored and the areole below the areole the displaced the upper surface only hump Finally sections between the the areoles This results areoles grow the areole less occurring the axils was shown for Neowerder tubercles mannia the ribs below This increased growth just
Evolutionary Gymnocalycium
more
separation
Usually rib
of
the podarium
rib
furrows indicate
adjacent podaria
any
viewpoint the fine
morphological a
and
become folded
.
( a
From lines
is of
K
can
Fig
direction
rib 81 )
humped
.
a
but
a
rib
'
have projected tubercles bearing the areoles
.
the
ribs
.
78 )
.
of
of
ridges
Brückner
.
of
).
rib
. in
occurring
80
of (F
(5 )
(
.
79 )
in
to
(
in
in
of
.
on
the upper surface Further evolution re Neowerdermannia the tubercles the axils Fig which has the areoles Gym The transition this genus occurs Fig nocalycium multiflorum has discussed the different Brückner seedlings margins types Melo ig cactus The first type has areoles sults
with regular verticillate arrange always have this type areoles type part ment The second has the the that bears the areole elevated thus leav ing the part between the areoles concave above the The third type has the part that the areole the areole developed the lower side of the turned downward species
Fig
. .79
of
.
A
.
.
rib
of
the podar
Gymnocalycium The multiflorum Ribs lateral ribs are greatly humped between areoles seen arising from above the spine cluster shoot may the center be
appears that growth
It
.
hump
of
on
is
so
rib
of
.
,
rib
.
of
of
states that
. a
, E
.
Brückner
possibly
.
(
.
of
rib
of of
) rib
resembling
.
an
shoots calcaratus only the
the
Hylocereus
In
the general habit Epiphyllum
of
,
length
to
Right
Fig 81
profiles The different types Melocactus seedlings After Echinopsis oxygona The humped cultivated specimen hybrid which withers during dry periods
: c-
a
. . 80
. .
Fig
RIBS
rib
in
.
of
rib
-
a
to
the three ribs remain low and areas below the areoles grow wing like projection The same type growth occurs Selenicereus hamatus ex cept that the protruding parts are turned
.
backward Flat shoots
tour
”
at
)
5
.
12
.
to
/5
UMA
).
-
3
in
.
) ) ) ) )"
)
28
(
new
observed
of
latent buds
shoot
cerei the
.
woody Epiphyllum plants basal part This usually occurred under poor cultural certainly indicates conditions This that species with bilateral flat shoots arose
2
ancestors
calcaratus
Selenicereus
-
a
ribs
. of :
. 82 .
FIG
Hooked
Hylocereus
-
b
a
a
to a
5
.
of
c
,
.
shows the transition from hybrid Epiphyllum ribbed shoot bilateral form Here the lowest terete part 83b
AMIN
cereiform
.
from Fig
)
(
.
.
From
Britton
hamatus
;
an
).
has
on
Buxbaum
a
,
in
. if
areole has grown into
83a
shoots arising from
form
"
of -
2
is
(n
, old
in
(
ally Fig
DDDD
it
,
is
radial but later becomes ovate and reduced from Very often intermediate three angled shoots hybrids Epiphyllum occur especially angled and Heliocereus ormally Also plants the shoots become ribbed Massart states that this commonly oc Nopalxochia phyllanthoides especi curs dispersion
and Rose
.
first the
are
of
are
(u
-
“
-
so
some Epiphyllums and the leaf cacti also formed from sually plants which nor few ribbed mally cereiform The growing point called
SUCCULENCE
THE CACTACEAE
OF
a
" C "
In
-
.
3
rib
Fig . 83 . The loss of ribs in species of the Epiphyllanae : a-Shoot from an areole phyllanthoides which was first 5- ribbed with of an old branch of Nopalxochia spiny areoles and later became a flat shoot . One of the ribs is rapidly disappear ing ( after Troll ) ; b and c-Gradual loss of ribs of a shoot of an Epiphyllum hy . brid . In " b " the first two ribs ( of four ) joined to form one and the shoot be winged came the two other ribs joined and the branch became flat
.
R
in
,
)
R
in
2
.
is
is
,
to
)
and
small
of
. of
.
(F to
. .
linearis also
-
.
of
,
(
.
is In
.
.
is
.
R
of
in
.
cm
.
un
an
of
to
is
in
of
this species the margin the flat shoot serrate and the areoles appear the axils the teeth This form leaf like shoot caused by the preferential growth the part the podarium below the areole This may compared with the hooks of
. of
-
,
(F
.
do
.
to
-d
/5
2
The same
a
not protrude
of
The ribs
re
ribs
interest because the small scale like leaves are not deciduous but rather are included the formation the ribs These flat shoots become quite long and are not more than width This form not Fig like the flat shoots Houlletiana 84d which are several centimeters wide and very similar Epiphyllum those
of
.
R
of
in
.
).
84a
pentaptera
Rhipsalis have the number three and finally Fig 84c the divergence flat shoots result This species
duced
.
- -
of
.
have begun with Loef The young shoot this species has ivergence small leaves arranged podaria elongated and the are downward ig the next leaf the same vertical row
grenii
small
1 5
.
in
)
.
(
a
rib
of
,
of
of
like Rhipsalis may
occurs
but here the podaria are elevated but rather high ribs Some species
)
)
-
of
(
in
Fig 83a the sudden termination the The same development occurs Rhipsalidanae forming both the Subgenus Rhipsalis and the flat shoot forms Alatae Zygocactus The evolution the leaf
84b
be of
.
1
of
a
of a
.
shoot Sometimes the reduction ribs can proceed Piptantho the manner cereus argentinensis Cereus peruvianus by
also
ig
of
to
in
of
.
A
)
13
is
.
up ,
),
Fig
.
(
.
rib
(
the first two ribs merge Fig 83b form one The divergence now few internodes further two more ribs merge resulting divergence 83c Further development this plant was flat
divergence
leaf
(
In
the ribbed portion
in
vertical
and the fifth
disappears
.
soon
of
divergence
-
'5
a
has row
2
.
shoot
Hylo
RIBS
. 84 . The lines of evolution of Rhipsalis branches : a - Top of a young shoot of Rhipsalis Loefgrenii ; b - Joint of R. pentaplera ; c- Part of a branch of R . linearis ; d- Branch of R . Houlletiana ; e-Pseudowhorl of R. cereuscula
Fig
(a to c after Britton
tip
is
86
.
of
or
)
.
(F
is
)
of
.
of
.
!
es
all
All
a
.
of
or
joints
and older plants have
a
.
angled
,
later forms
.
, ,
is
as
. so in
five
.
84e
)
.
Fig
of (
to
of
, of of
,
, . R
termi areole covered with short wool nal flowers and new joints arise from this areole The origin these large areoles has been closely studied the related species Rhip salidopsis rosea Young plants have four in
.
- - or
,
,
,
,
.
A
ovate orbicular convergent evolution
of
In
.
of
,
es
R
.
broadly
.
of
.
is
,
in
genus also Epiphyllum branches occur from basal areoles whereas Rhipsalis branches from the uppermost every fat joint Mor even apical areoles phologically therefore Epiphyllum branch basitonically and Rhipsalis branches Rhipsalis Houl acrotonically The joints letiana are small lanceolate and rather long Rhipsalis The more typical form crispata pachyptera and others widely spreading that the joints become each
usually
at
length , sometimes up to nine meters . The shoots of Rhipsalis have a determinate growth the length being characteristic Lateral branching different each species
limited
(
85 ) . Others may have straight margins . The flat shoots of Rhipsalis Houlletiana and related species , as well as those of Epi phyllum , begin growth as a more or less long shoot which is terete , this resembling the shoot of any dicotyledonous shrub . The shoots of Epiphyllum grow to an almost un -
Rhipsalis leads to the characteristic jointed Alat shoots of Zygocactus and Schlum bergera . Also another relative of Rhipsalis , Pseudozygocactus epiphylloides (Compos Porto et Werderm .) Bckbg. (Syn . Rhip salis epiphylloides Compos - Porto et Wer in the same manner as derm . ) , is jointed Zygocactus . Zygocactus , the well known “ Christmas cactus, ” has jointed flat shoots which are ovate to oblong , and in some varieties the joints are serrate with the are oles only on the margin . The stem Marginal are ig the retuse apex oles never bear any lateral branches Rhipsalis This the opposite flowers Subgenus Platyrhipsalis which bears flow the joints After from areo flowering terminally the joint bears new joints beside the flower The retuse top Zygocactus bears very large the joints ers
cereus calcaratus and Selenicereus hamatus discussed above. Similar flat shoots occur in Epiphyllum , especially in E . Darrahii ( Fig .
and Rose ) .
is
.
.)
(
of
.
at
.
is
89 )
.
(
.
as
”
"
of
a
.
.
If
.
90
.
as
(
-
un
is
it
of
.
of
of
Zygo believed that the flat shoots except Epiphyllum cactus are the same that the former has determinate growth Although the flat jointed Rhipsalidanae in
as
.
growth habit the attain one extreme Subgenus Phyllorhipsalis and the related genus Zygocactus another evolutionary
,
,
.
are
.
E
In
.
in
as in
.
E
by
as
obovatus the joints are flattened and areoles formed Opuntia over the whole shoot These not true flat shoots but are are actually only flattened cylindrical shoots This obtusangulus which has terete shown
formed from these areoles they gradually arise further back from the center the cluster areoles From these plants comprehend why Leinfellner difficult
a
the flowers
coalesced into one areole Schlumbergera Gaertneri which shown Fig has many united areoles joints are destroyed they are the new
of
re
.
in
Zygocactus
with
terminal apical areoles This may also be the
to
).
88
or
as
.
Fig
(
a
born
genus Epiphyllanthus
plants show that may be group
These
areole
the
in
of
,
)
.
87
at
.
(F
at
and
some areoles the top are circumscribed by thickened border The joints only the apical are extremely acrotonical flowers areoles bear new joints Similar acrotony occurs the next lated
the these
,
on
.
)
(
of
,
The the the flat
26
39
as
of
has shown the angled joints occur ribs but only the areoles the top joint bear new joints ig The shoots have areoles along the margin areoles
joints but some areoles are grouped top and branching occurs only from Fig areoles This growth form Rhipsalido psis rosea same
(
,
and become flat like those joints also become
Zygocactus Grafted Vaupel bilateral
Zygocactus truncatus Branch var which exhibits acrotony and determinate longitudinal growth
)
below
.
. /12
of
divergence
THE CACTACEAE
in
the podaria the areoles of
enlargement
OF
.
.
The part between the areoles becomes wide angled by
86
Epiphyllum
of
.
Branch
)
Fig . 85 . Darrahii
FIG
SUCCULENCE
RIBS
00
of
.
be
of
It aby
is
of .
,
to
in
of .
has been shown that the Rhipsalis are alike habit type because they have cereus
on
Hylocereus
a
form
of
“
”
any the spiny areoles and tubercles which soon ribs The young shoots are like those
.
Later the basal lateral branches which are like the primary stem The primary stem therefore branches basitonically After lateral branches further development slender
grow
terete
and
These
also
form
.
of
of
-
In
.
to
During further growth
.
of
Large apical areole Schlumbergera Gaertneri The large clearly separated areole into single along small areoles the
right part
.
.
.
)
no
.
same vertical row
Fig
.
(
by
of
joint Epiphyllanthus obovatus view the apical areoles marked arrows are forming new shoots
Top
of
.
Two
. of 89
FIG
of is of
.
91 )
.
also interest this species podarium the each small scale like leaf elongates downward the next leaf the
is
,
three circumwalled areoles arrow has formed new shoot
ribless branches are formed protruding podaria have Rhipsalis The development
.90
of
fat
a
marked
Rhipsalidopsis
)
of
which
joint
.
.
Top view The top consists
by a of an
rosea one
FIG
ig
Loefgrenii (
.
88 of
(F
of
.
.
areoles
to
-
)
(
. E.
terete ribless branches
seedlings
seedlings
of
by
- -
of
in
in
, ,
in
).
The genera Erythrorhipsalis and Epiphyllanthus candidus also have
of
-
to
.
(
genetically
any doubt that these There cannot Rhipsalidanae are descendents plants ribbed cereiform This fact brought out study the development terete
.
from the primitive five ribbed those which have terete ribless forms branches These plants were classified Rhipsalis Subgenera Eurhip Schumann Lepismium Sub salis and Orhiorhipsalis genus Calamorhipsalis and the genus Hariota the present classification the Rhipsalidanae cannot be accepted phylo
trend leads
b -D
.
60
a
of
tip of a 4 -a
.
tip
FIG . 87 . Rhipsalidopsis rosea : a-Development of only one new shoot from the top of a evelopment two 5- angled joint. The new joint arises near the center of the ngled joint new shoots from the top The two shoots clearly arise near the degree angle between each other the and form center
OF THE CACTACEAE
SUCCULENCE
40
TORE
UN
AN
N
A
.
Co
;
;
.
S -
H -p
.
.
of
:
;
.
;
Right
of
:
.
Rhipsalis cassysha Basal part the young plant The primary Fig shoot has been removed and three lateral terete shoots are developing rimary root rimary stem lateral branch Three year old plant cotyledon After Irmisch Development
W -p
.
91b
:
.
Fig 91a Left epicotyledonary
is
.
”
“
in
is
.
a
at
a
of
.
so
of
the primary stem the podaria grow to gether that the stem becomes quite terete Hariota exhibits peculiar growth the joints large areole These joints have their tips and branching acrotonical Also Hariota the terminal large areole
)
Joints
by of
93 .
.
Fig
in
in
as
.
of
(
of
composed usually five several areoles which cluster near the top the shoot The young shoots arise the same manner
Hariota .
only one clarata formed basal internode
).
92a
basal internode becomes without thickening while the other internodes become succulent Because the uppermost internodes remain short the areoles this part the stem form the first
large apical areole
. of
of
,
.
Soon the lengthened
slight
.
(F
ly
other Rhipsalidanae with the podaria projecting between the scales ig
.
)
of
of
at In a
)
.
(F
. ,
a
a
of of
an
.
at ; b -
of
;
c -
In
.
of
of
to
a
at
a -
.
:
.
Development sali Hariota the joints Young shoot developing between four cornioides joints formed the top the branch Old stage joint growing the border show the scales the large areole Mature joint
. -
92
FIG
.
of
(F
ig 92b The podaria fleshy wall which this large areole form young shoots the surrounds the areole appear scale like leaves the border this wall mature shoots Hariota sali very tender cornioides the joints have basal internode and succulent upper part upturned bottle ig 92c much like
MAMILLES
of growth occurs the one basal inter node is greatly stretched and becomes Aeshy , whereas the other internodes remain short , succulent , and all the areoles are col lected together at the top of the joint to form a large areole ( Fig . 93 ) . It is not known whether the succulence is brought about by union of the podaria or by a thick ening of the green cortex , but nevertheless this plant undoubtedly shows the greatest reduction of ribs in the Cactaceae . of
which
on
;
-
;
.
3 .
-L
1
of
:
a
of
2 -L
,
,
.
in
,
at
.
is
.
to
very different types may occur
Three parts
of
,
.
-
,
-
in
is
-
.
be -
,
-
or
a
.
Usually however and Rhipsalis growth both longitudinal directions and altitudinally this resulting favored podaria overlapping form ribs Mamilles growth This favored altitudinal arise very striking even altitude increase Fig the growing point the mamille two After further growth (
.
,
a
directions
ongitudinally
ongitudinally downward Altitudinal
Lateral
Opuntia
Because the projections are usually elon gated the term mamille seems more appro priate than tubercle which generally implies But less knot like structure more cause Britton and Rose used the word tuber cle for all wart like organs this expression the English literature firm hold has cacti As explained above the mamilles are the
the
of 33 ) .
eve
mamille
.
the naked
upward
Diagram
is
by
seen
Ariocarpus
)
(
cases
growth
4
,
in be
at
is
.
be
to
small
some
. 94 .
of
-
of
too
of
The term mamille or tubercle applies to the wart - like , hump- like , or sometimes even leaf like succulent elevations the body usually some cacti An areole born the top each mamille but the areole may
of
(tubercles )
;
Mamilles
FIG
.
D
this form
in
in
clavata
if
.
in
The extreme
in H
or
)
,
or
the
pendicular
.
in
To
,
;
stem
2
; :
)
growth upward parallel growth downward paral axis lateral growth per
, 1,
axis
.
stem
(
94
of
top
,
The furrow
.
wool Photographed George Lindsay
by
.
elephantidens
or
Coryphantha
not bear hair
.
in
is
as
or
,
do
in
.
to
from
4
the stem
;
to
axis and altitudinal the base the the tubercle When growth occurs longitudi nally one both directions and laterally the stem becomes covered with leaf bases that not project shown some
growth
.Fig .95
lel
Fig
the to
to
tions
Growth these four different direc in
apex
the stem
tubercles may occur
3
of
point
at
in
or
as
.
to
as
.
to
so
as
,
of
in
(
.
or
podaria succulent leaf bases the cactus podaria leaves These same organs can Opuntia also form the humps found they can become the ribs ribbed cacti flattened not form any projection The question arises how the same organ may form ribs and mamilles clarify this point one must imagine that these organs arise primordially small the seedling the growing tubercles
does
SUCCULENCE
OF THE CACTACEAE
size before the areole has differentiated , the two parts of the areole become separated , one being elevated to the top of the mamille and the other remaining in the axil. A fur row may also be formed on the upper sur
face of the mamille , but because it contains no meristematic tissue this furrow does not bear wool or hairs (Fig . 95 ). The spini ferous part of the areole normally becomes non -meristematic at maturity , but the axil lary part remains active and forms the flow ers and lateral branches ( Fig . 96 ) .
by
.
etc
.,
up
set
Berger
.
to
and
in
to
included
of
97
.
(
a
of
in
is
of
.
of
to
,
”
,
(
“
to
)
),
to
of
(
by
is
.
)
in
.
,
to
).
is
(
of
, of
to In
.
to
,
to ,
in
to
,
.
to
,
do
.
,
,
of
)
.
)
.
08
of
in
of
in
,
if
:"
in
*
Roseocactus was Ariocarpus fissuratus Sch
apparent that the classification cacti some attention should be directed to ward the development the plant rather merely the superficial resemblances than the mamilles
.
,
as or -
.
the furrow
Coryphantha and Roseocactus On the other hand the primordium increases
is
, - -
on
from
34 so )
.
(
“ A ”
lateral branches arise
when mature but also for Dolichothele longimamma which has the Mammillaria largest mamilles all the Mammillarieae Fig
to
an
of
di -
.
inIf
it
of of
of
a
two by
nected
in
by
in
,
),
.
of
growth
of
is
,
to
in
”
,
is
(
“ B ”
so
of
in
nect
parts two parts
the areole remain con hairy furrow the upper sur Fig face the mamille Sometimes lengthens the basal part that the fur row does not reach the axil and flowers the
(
,
in
“ A In ”cam
part .
.
of
in
.
of
.
is In
lengthened by the increase the mamille that becomes vided serially into two growing points the mamille continues lengthening after after the axillary point growth hasening has differentiated size
succulent leaves The second line evolution occurs the Mammillarieae beginning with Thelocactus progressing Mamillopsis Coryphantha Mammil laria and Cochemiea these plants the cotyledons are reduced very small rudi ments The first mamilles the seedling protrude greatly remain small and not but later primordia greatly elongate form mamilles This development holds not only for species that have very small tubercles resemble
(
the is
,
so so
of
to
to
in
is
,
,
“
The second type characterized by part Fig 331 increase which bears the areole this case the growing point
The succession flower formation Leuchtenbergia flowers from the shown top the mamille and has well developed cotyledons Roseocactus flowers from consisting the furrows the Ariocarpi Ariocarpus and Encephalocereus flow ers from the axil The genus Obregonia Leuchtenbergia closely but has related more reduced cotyledons All these genera have rather peculiar mamilles that they
It
,
of
;
.
( B ) ) ;
(“
A ”
in
Fig 331 growing point which and the top the rudimentary blade the leaf the mamille the basal part first type length that ebasal the whole areole creases that ength top This the mamille the elevated levated type Leuchtenbergia Gymno found Berger calycium and according the Pseudomamillarieae the genera Thelo Neolloydia cactus Ancistrocactus and Mamillopsis
the areole
importance systematists The succession apical flowering mamilles flowers axillary flowering has occur the furrows red two convergent lines evolution The first line evolution characterized seedlings which the first mamilles are very long and form long and tender tuft Fig mamilles just below the cotyledons
of
be
"
:
)
of
(F
.
point
the displaced
distin
the basal part below growth the areole the part bearing this
331
proces of great
appears
in
.
the areole
Zurich
of
ig
that the evolutionary
of flower formation may be
of
. (
of
parts
Plüss
.
(
by
)
)
axil
and inner Photographed
the primordial mamille may
guished
melano
no
tubercles have
of
Mammillaria
The the fruits arising from the axils spine furrow connecting the outer
show
bearing
the body
of
Part
to
centra
of
. 96 .
Fig
It sion
MAMILLES
La
ць
voll
(
in a of
.
)
( H )
of
in
of
varied
obtuse
from
and round
thin
to
of
Wildii
Mammillaria
All
.
ed
tubercles
greatly
or .
mamilles
forms occur flat conical mamilles the
cm
All
is
The habit transitory
of
a
.
in
III :
FIG . 97 . I : Leuchtenbergia -type seedling , a young seedling ; b- old seedling . The cotyledons ( Co ) are quite well developed and readily distinguished in the old seedling . A cluster of large mamilles is formed between the cotyledons even in the young stage (after Troll ) ; II : Roseocactus ( Ariocarpus ) fissuratus ( a and b ) and R. Lloydii (c ) . a- First stage before development of mamilles , b- stage with first mamille , c -old seedling . Only in the first stage may the small cotyledons be easily distinguished . The cotyledons soon become reduced by growth of the hypocotyl . This genus also has very fine spines ( after A . Berger ) ; Three seedling stages Ario carpus retusus growth As Roseocactus the cotyledons are included the hypocotyl and later stage form mamilles after Troll border around the cluster
long and the diameter finger like tubercles longimamma Dolichothele Mammillaria
of
-
)
(
Fig
.
7
from
in
.
1 .5
occur
also
cm
forms
,
to
.
of
.
C
of
),
99
.
(
the blunt nearly hemispherical Coryphantha bumamma and mamilles elephantidens The tubercles Pelecyphora is
,
100
ig
of
-
and Solisia are sharp edged by lateral com pression and the color like that stones
,
).
.
,
is
of
of
)
.
(
,
(F
in
,
)
carpus
.
.
to
(F
These genera and some related Strombocactus are called mimicry forms The peak evolution tubercles ig reached the genera Roseocactus Fig 102 101 Obregonia and Encephalo
of
.
- (( .98
Fig
in
.
of
-
(
,
is
of
a
.
Co ))
.
type seedling The Dolichothele The cotyle dons Co dons are more reduced than the Leuchten bergia type long mamilles No cluster found between the cotyledons and the first podaria mamil les are very small and later ones gradually become larger After Troll .
,
)
a
by
a
).
The areoles are reduced
to
.
ened surface
.
(
is
of
-
,
,
.
A
.
In
-
of
in
as the
,
to
-
)
.
(F
ig 103 The mamilles Ario carpus are thickly three edged and are very similar leaves rosette forming leaf trigonus succulents Haworthia long especially the mamilles are three edged and leaf like Fig 104 The upper strengthened surface these hard
Fig
SUCCULENCE OF THE CACTACEAE
of
.
)
(
.
.
Ariocarpus fissuratus show 101 The mamilles ing the hard surface and prolongated areole furrow bearing wool
-
of
,
on
.
in
cm
12 .5
,
.
.
-
.
leaves
this group
,
plants
,
possibly belongs
is
interesting genus to
very
The old tubercles are decidu
most dicot
of
.
wool are
in
. ,
ar a .
a
of
:
.
P
.
.
: of
FIG
as
-
ab
in
-
on
,
at
.
of
,
do
.
Laterally depressed Above Pelecyphora aselliformis Right pseudo pectinata
100 mamilles
areole
ous A
-
or
or
of
, at
pointed projection upper sur the top face the mamille are completely young sent The areoles bear wool only specimens and spines not arise except seedlings Obregonia are The mamilles like these but spine areoles occur the top Encephalocarpus strobiliformis has very many thin leaf like tubercles which are keeled the back and are curved upward pine cone the whole plant resembling Only young tubercles this species have small oblong areole with pectinately
a
.
.
Dolichothele longimamma showing the long From Marshall and Bock
terete mamilles
.
.
Fig
.
99
in
)
.
.Fig
(
In
.
ranged spines the evolution leaf like principis attains tubercles Leuchtenbergia the greatest development 105 These mamilles are the largest found the Cact length aceae and may become and are flattened above and keeled the lower side The areoles bear long paper like spines and few very small conical ones which are completely covered by the
which
the
new
spine clusters the From Marshall and Bock
.
(
.
cm
,
.
.
to
.
the
young
of a
Ariocarpus
a
of
.
.
Casteneda
In
.
in
.
by M
Photograph
b
of
.
. is
tip
,
i
of
3 -5
agavoides
In
Neogomezia
.
at
the lower part
Neogomezia the mamille and flowers arise the axil the top each mamille and flowers arise from the areole
immature mamilles
.
.
,
a
a
,
is
arise
-
trigonus
b
is
fowers
;
-
a
of :
below
Ariocarpus
the inner juvenile
to
to
-
,
,
,
cm .
1
at
-
areole occurs about
of
.
tip to
or
-
in
-
104 Leaf like mamilles areole occurs the top
.
.Fig
The lower surface
,
.
40
(
)
at
a
of
.
6
,
small
areole
leaf
with
convex and slightly keeled and the acute mucronate and usually recurved upper On the surface about from the large usually spineless round there long gray wool areole filled with mass Occasionally the areole bears one three long spines subulate depressed mm From this growth habit the plant seems re lated the Strombocactus group The tip
ap . It , -
is
,
)
,
of
be
.
( to
bears several terminal tubercles mm long mm wide the base which look like the leaves leaf succulent because they are semi circular cross section the upper surface being flat somewhat con the areole and concave vex from the base the
strobiliformis The furrow first flowers surface bears plants the
upper
fusiform
trunk termed root stock but this mor phologically wrong Actually the trunk pears formed from the primary root hypocotyl the and part the epicotyl
from
Encephalocarpus 103 upturned mamilles
.
, .
.
like
)
fleshy
Fig
The
in on ,
.
-
.
(
of
plants
right
104
)
Neogomezia Fig this group have
genus
a
Denegrii
Obregonia
at
Leaf like mamilles with -
.
102
45
m
.
top
in
.
FIG
MAMILLES
large very
at
In
. .
In
.
. 3 :
Am
.
of
“ b ”
is
is
it
of is
“ B ”
mer
of
of
14
.
“ A ”
,
is
a
na
.
is
old
of
.
is
.
it
is
,
of
,
, ,
genus
cause
the
dense
rosette
ole and the flower bearing part the axil Cory lengthened Here part phantha type but elongates before the areole primordium has differentiated Since situated the upper surface the areole homologous the above must the
quite rough due granular hard mimicry epidermis Roseocactus also cactus Very similar these but without elongated areoles are the flat tubercles ig 106 Strombocactus disciformis
.
a
in
is
blunt to
.
This
,
a
is
to
.
top
on
are
a
and
These hardened tubercles lead
.
to
.
)
,
of
(F
,
.
to
be on
it
is
tip
part
is
is
- ). -
the
(
(
in
as
it
is
“ B ”
,
-
of
tip
its
,
a
-
)
placed Britton and Rose tubercles are deeply furrowed and the flowers arise not the axils but from these furrows The tubercles form
.
.
is
In
which
“ b ”
laminar part remains reduced Ario carpus the very small areole situated the upper surface and near the the growing point leaf like tubercle but spine bearing part the are divided into
it
by
be
-
the
in
fissuratus Ariocarpus
)
Ariocarpus
on
Roseocactus distinct from
is
.
.
tubercles are deciduous Also the indicating their leafy nature (
-
in
,
-
blade again
is
“ B ”
,
.
a
-
is
,
tip
.
In
the
identical with the laminar part tubercle actu the primordium Therefore ally leaf No other cactus has mamilles the leaf base and the leaf that consist
on
and invisible
The areole
the leaf primordium greatly also reduced the flower born from
not This shows that part areole lengthened and that the greater part the long tubercle formed from the primor The part above the areole dial base the the total length about which
.
ille
laminar part Ariocarpus Neogomezia
a
in
on
“ A ”
,
-
-
of
,
the mature mam therefore situated just and flowers arise above the spines the Coryphantha type and also the Roseo part cactus also lengthened and the
reduced
.
.
is
(
the
is
", the
“ b
" “A A ."”, tip
In
.
)
um
m de
be
de - by
-
of
In
.
of .
33
of
of
,
“ B ”
.
of
around the center the plant morphology The these leaf like mam Neogomezia may explained illes Fig these schemes the means velopment the three parts shown the area part below the part basal part below the areole part the which actually vasal and the very reduced blade the leaf primordi the Thelocactus type and also grows part only Leuchtenbergia markedly while the laminar part becomes
“ b ”
.
-
Leuchtenbergia
est areoles
THE CACTACEAE
OF
spine producing areole like long mamilles with principis From Cact Succ Jour 153
The Agave
of
.
105
the
top
FIG
SUCCULENCE
the
odd
to
as
at
.
by
arrow
gray
green
-
.”
-
Risteri bear top
another
surface
mimicry
mamilles
all
tubercles are morphologi organs there actually great difference among mamilles they are great phylogenetically examined has been
It
if
is a
,
.
en
- -
There to
tubercles
.
hard
fore the new mamilles are not able large except outward direction
Although
cally homologous
died
so
(
Secondary evolution
of
.
.
,
.
,
no
fastened
,
well
-
,
old
among
tubercles are formed
new
between ribs The plant
causes this species
)
.
(
is
).
,
slowly
Aztekium
be
shown
of
(6 )
or
This
tubercles
wool and spines but become hardened
pseudoribs Aztekium Ritteri plant grows exceedingly slowly the young lateral shoots have ribs and the ovate tubercles bear areoles with wool and several conical spines Fig 107 Above the areole the tubercle also hardened shown by arrow Because the plant grows ribs
107 New
of
.
of
.
Fig
106 . The flat, hard surfaced mamilles Strombocactus disciformis .
.
.
MAMILLES
E
Fig
OF
"
SECONDARY EVOLUTION
This
form
is
.
Aztekium Ritteri three fourths size -
is
.
.
.
FIG 108 Hardened tubercles form pseudoribs Illustration
of
.
).
.Fig
,
to
together
to .
be an
pressed
.
them
(
a
to
in
phyloge shown that the Mammillarieae contains highly evolved cactiMammhp which progressed from pseudorib Rolls between two pseudoribsribbed species tuberculate Thelocactus undoubtedly derived are caused by pressure between adjacent and that Rebutia tubercles 108 These have been called from ribbed Trichocerei Thus the mamilles causes
SUCCULENCE OF THE CACTACEAE
.)o ) o). 2
of
.
of in
-t
by
-
up
of
in
of
Lololo
)
loooo
volo
in
ob
of
to
.
.
.
These
en
.
-
.
-
-
furrows
).
1
.
of
to
so
(
-
).
-
deep
the areoles that the ribs very between the areoles are narrowed small strips Other specimens this very species may have the ribs com variable Fig pletely resolved into tubercles especially species Other related Strombo
,
.
),
.
of
-
of in
(
-
ribs separated large around
by
flat
,
as
.
-
Fig 110 the evolution Schemes lique oblique hawk like mamilles from cross wrinkles The Lobivia type
.
Boedeckeriana this genus spirally arranged Fig Rebutia
species
have ribs changed into they occur tubercles 109 shows the evolution tubercles from ribs with cross wrinkles Rebutia type oblique and Fig 110 shows the evolution hawk like tubercles from ribs with trans verse fissures Lobivia type
(
.
Como
. do
.
.
different angles
of
.
,
often has
!
L
.
.
of
ribs but other ,
Turbinocarpus
lophophoroides
,
(
,
-
In L.
E
L
. .
.
of
ly
of
because tension from the dispersion Mediolobivia also has
:
-
;
.
)
).
of .
wrinkles
-
-
cross
in
to
calochlora has flat distinct ribs Lobivia Pseudolobivia curea also has straight ribs and Backe bergii has slightly humped ribs cin nabarina and others the ribs are jointed by oblique fissures into hawk like humps The Neohaageana are set tubercles straight rows which often become spiral fissured
.
is
.
(F
.
with
Strombocactus
An analogous evolution may be seen Echinopsis the series from Rebutia Echinopsis oxygona has straight ribs which cannot fold during dry times but they become thin
d By
b -
;
in
-
forms
)
Gymnocal
-
genus
.
the
.
. -
of
.
in
-
In
for
been discussed ycium
ribs with straight
so
-
,
of
,
of
or
to
It
evolu that the different lines tuberculate species were ribbed the base with shrinkage associated the whole plant during dry periods even the more primitive species the ribs first become notched and the plant can there any direction Evolu fore be compressed tion tubercles along this line has already
(
appears
tion from
the
the base the tubercles the cross wrinkles become more and more supplanted the adjacent tubercles The tubercles the same verti displaced and also become cal row that finally the ribs are totally down transverse lost and the tubercles occur evolution This type rows Fig 109d the Rebutia type transversely arranged The evolution tubercles from ribs may also occur without
,
,
of
.
of
,
.
it
Also has been shown that tuberculate Gymnocalycium Rebutia and Mammillaria represent different convergent lines tubercle evolution
Tubercles which evolved from cross wrinkles first are arranged By growth ig rows 109c
of
not the
is
as
of
plants phylogenetically the primordial tubercles
(
are
of
mature the same seedlings
of
of
c -
;
of
-
a -
.
.
to
109 Schemes the secondary evolution mamilles from ribs with cross wrinkles Straight ribs with transverse cross wrinkles between the areoles Podaria becoming Development larged around the areoles growth mamilles vertical rows base the mamilles the regular arrangement mamilles lost The Rebutia ype
( F en )
ooooo
.
of
O
Fig
00000
100101010
ololololo
lao
1o
.). ).) )o ) . )o •) ). ) )
SECONDARY EVOLUTION
MAMILLES
OF
FIG . 111. The secondary evolution of tubercles from ribs in : a- Turbinicarpus ( Strombocactus ) lophophoroides : b - Thelocactus fossulatus . Photography by Backeberg .
two grow
he seen in The origin of this growth may be young plants they repeat the phylogeny Coryphantha the group the Sect long yellow Glanduliferae has red gland back the areole the axil Nor mally old plants this gland the axil . or
.
of
of
top
,
in
on is
in
is
it
young specimens
the
of of
to
Fig
.а
but
in
in
of
or
a
in
of
sis
.
.
as
Thelocactus to Mammillaria . T . tulen and others have small distinct connecting ribs between tubercles the same vertical row The podaria are greatly enlarged
from
evolution may be termed the Thelocactus type . Another evolution , originating with the Thelocactus - type , occurs in the Mammil larieae . Here the areole becomes elongated larieae Hertivides into two growing points . and finally divides
.
tubercles and the ribs are totally resolved into obliquely arranged tubercles ( Fig . 112 ) . the A similar evolution is shown by the Bckbg . , progressing Boreoechinocactanae
of
cactus disciformis , have lost all connections between
.
.
.
)
113 The old plants
.
-t
.
.
,
Jr .
,
.
A
by
of
.
,
showing FIG 113 Thelocactus tulensis sec ondary tubercles the Thelocactus ype with the tubercles connected small low ribs Photograph Haage Erfurt by F .
).
.
(
of
,
in
is
it
,
.
T
totally tulensis and lost most Thelocactus these having instead oblique rows tubercles Fig 114 This
typical
.
(
lost
of
often
Fig
in
is
into long tubercles
connecting
rib
altitude
-t
.
of
.
of
by
.
112 Schemes the evolution growth tubercles the basal part podaria the around the areoles No cross preceded wrinkles the formation mamilles The Thelocactus ype
HABIT OF THE CACTACEAE
of other dicotyledonous
affect the growth plants :
.
growth
The longitudinal
1
of the
pri
mary
stem . The longitudinal symmetry of the pri mary stem , including primordial thicken ing. 3. Acrotony and basitony of branching . of the 4 . The longitudinal symmetry branches , including periodical thickening . 2
*** il
18
SIDE
.
The angle of branching . 6 . Radial and bilateral symmetry . 5
.
the plant
of
factors are attributes
These
itself and are controlled by heredity . To these factors certain ecological factors must especially affect longi also be added which growth tudinal and bilateral symmetry :
. 8.
of branching
and the angle
Geotropism
.
(
be
is
.
B
than
primordial
's if
is
or
,
.
in
be
is
or
a
is
is
,
of
.
or
.
in
rapid
.
,
If
,
,
.
re -
a
of
as
the very slow terminal growth and lateral growth Thus the very large globular forms arise slowly but after course
,
to
all
of
)
a
of
of the
of
a
.
in
of
;
.;
m
3
of
. or
(
branching the exception loss great weight large and the the shoots these factors are the same those which
always short cacti there shrinking process which nearly compensates length Thus the forma for the increase globular short columnar habit tion has two very different causes globular forms The primary cause extremely
of
in
to
a 2 m
of of
18 m
of
;
of .
m
25
With
.
20
,
-
of
the Cact habit The great variety encompassing the small mm lilliputana Blossfeldia diameter forms Echinocactus with the giant forms the giant diameter and height height the Pachycereus more leafy trees and shrubs the Pereskia climbing Harrisias Eriocerei more than length and many others are only sult the influence few factors
aceae
.
of
factors
of
OF
General
the
the stem origi many cacti has indeterminate longitudinal growth and the growing point remains active until the death very the plant terminal growth and rapid the plant becomes columnar even slow the longitudinal growth surpassed by lateral growth the plant globular Also comes short cereiform
The primary stem that nating from the seedling
THE CACTACEAE
of
.
A
.
3
HABIT
is
stem
,
.
from the axils
the
in is ,
rather
,
mamillaris
Longitudinal growth
of
.
f
of
of
)
Mammillaria
it
in
this monograph may encourage future research along these lines
i
-
is
has
hope that
(
( 48 )
.
,
,
.
It
ly
in
of
. ,
the different lines
.
of
in
ly
and
on
of
factors and the effect them evolution will brief regrettable that these prob discussed lems have not been more completely studied before for they are exceedingly important phylogenetic studies and author These
tubercle above the spiny areole Apparent yet the division the areole has not yet not per occurred the young plants This per found lateral haps why Zuccarini shoots originating from the top growing point Coryphantha vivipara tubercles
.
affect of gravity
The
.)
of
-
.
in
as
.
114 Thelocactus type second ary tubercles without any connection hexaedrophorus Thelocactus Ribs and vertical rows are absent and oblique the tubercles are arranged rows.
Light,
the plant
on
Fig
7
LONGITUDINAL GROWTH OF THE PRIMORDIAL
the maximum
diameter
has been
STEM
reached
continue to grow slowly in height . This growth habit accounts for the speci mens of Echinocactus ingens which Kar they
winsky (23 ) observed
to be 1. 5 to 2 m . in
diameter , and Ehrenberg saw some speci mens of E . platyacanthus which were 3 m . in height and 2 m . in diameter . Other spe cies continue to grow in length after hav
short columns ) . As the plant becomes in portion ap creasingly old the shriveled proaches the top . The old parts become corky and are densely compressed , and the whole plant along with the closely spines becomes hardened .
packed
ing attained a maximum diameter . Echi nopsis is of this type , and Neoporteria sub gibbosa normally occurs as a globular or
,
is
.
( P
at
is
at
dia
to
.
115
.
Fig
in
grammatically
FIG . 116 . Scheme to the formation of the flat habit of the first the plant The point Discocactus growth the plant top and the plant the lower and finally displaced the side surface of
i
high ) ,
to
.
on Ps ( ) by P .)
30 cm
( P , )
to
.
(up
form
but it may also form columns over m . in length . This form of growth is shown
of
short columnar
Injuries received near the top are gradually
of
the plant the shriveled zone
to
displaced
of of
no
A
.
to
the
at
a
,
in
.
A
three year old there disappear elegans which had been Mammillaria jured by deep one centimeter long wound top was observed signs have specimen this wound after two years and
.
, .
111
. is
the
,
·
In
AM
attaining after the maximum gradually diameter height increased
in
growth width sur growth greatly that the terminal the plant spreads nearly disciform upon the soil Discocactus alteolens fine example growth Fig of this from shows point near how the shape changes and how the growing the young plant moved during this growth some species
Man
116
is
by
.
tip
of
a
.
.
of
.
is a
so
passes
The distinctly short cacti first grow similar manner that they enlarge rapidly during the first few years However soon the plant begins shrink the basal por rapidly tions the growing point adds
a
wwur
.
,
so
.
per of
of a
basal zone the top shrinkage
of
:
at
growth
the formation
ZZ zone and direction SZ -
Scheme
manent short form
, -
117
.
FIG
.
.
(
.
)
,
of
(
the plant Fig 117 Mammillaria growth whereas Cory exhibits this type phantha follows the first type matures into
to
in
as to
to as
height
a
a
, it
its
to
.
is
.
115 Scheme the formation the short columnar habit The specimen first globular and
,
at
of Fig
,
.
,
to
Mammillaria centricirrha which was cul tured under low light intensity grew long thin column Later after being placed formed conditions under optimum normally broad head and then by shrink ing the base became shorter and finally resembled normal globular habit Some
HABIT OF THE CACTACEAE
of
of
.
:
9
.,
Am
.
,
,
,
in
of
.
is
.
in
be
of
a
,
.
of
of
.
,
not occur
in
it
does
the
in
on
be
in
.
mary
pri
.
an
is
to
and limited the lateral branches Dendrocereus may be excep tion this rule stem
Echinocereus the color
of
in
of
.
a
the Pectinatae group periodic change is
there spines
These plants
of
.
”
of
.
are therefore called the the zones because cacti reddish and white spines lateral branches both longitudinal sym metry and periodical thickening commonly Zygo Opuntia Epiphyllum occur rainbow
,
,
in
,
In
all
,
.
at
a
joint
of
the Cactaceae
“
ap
growth
of -
of
term
at
certain
rhythm
the
and lateral branches Not the stem are active once but only parts bear lateral branches any
of
stem
buds
the periodical
to
plied the
is
Longitudinal symmetry
each
The longitudinal symmetry may also accompanied by periodic change the Yearly growth usually begins diameter with small diameter growth and later internodes become gradually more succu lent and the greater part the joint has the normal succulence Toward the end the period the internodes decrease size This growth period may seen the primary stems some succulent Euphorbia but
as
,
,
.
thickening
repeated
a
in
.
of
in
all
, .
.
as
,
,
of
in
.
periodical
and
there are always different distances between the areoles and the rhythmical sequence
to
as
If in
.
As
in
In
.
C
Longitudinal symmetry
Cereus
532
time The buds also arise specific equal distances apart most the Cactaceae but especially some Rhipsalides Hariota
one
In
, -
in
.
,
Sometimes the root also thickens does the stem such plants grow more length very flat forms result width than which may even shrink into the ground during dry periods does Lophophora These flat forms must not be confused with Discocactus the fat discs previously mentioned the primary indefinitely grow cacti stem does not many species the growth the shoot stops the seedling and lateral branches are formed instead This growth habit may be found Harrisia Guellichii Rhipsalis and some Opuntieae
.
.
at
,
If
.
,
as
Echinopsis plants such not shrink the base these are grown under poor conditions they become abnormally shaped optimum but after being placed condi tions normal stem parts are formed from the growing point
the braoches
in
do
.
Longitudinal symmetry and periodic growth peruvianus Jour Cact Suce Soc
.
118
.
.
FIG
29-187
mm
ACROTONY AND BASITONY m
53
m
cactus, Rhipsalis , and others . A fine example of longitudinal symmetry of branches is shown by the tree - like cactus Cereus peru vianus ( Fig . 18) . Longitudinal symmetry without change in diameter often occurs in the primordial stem .
branches arise from the cotyledonous are oles ( Fig . 119 ) . It is worthy of mention that different parts of the same plant may branch acro tonically and basitonically . The primary stem of Rhipsalis cassytha branches basi tonically , but these lateral branches are dis tinctly acrotonical informing secondary
lateral branches . In Lemaireocereus Weberi it appears that the stem branches acrotoni cally and the secondary lateral branches are formed from the first lateral branches basi tonically ( Fig . 143 ) . Large crowns may formed by acro branching tonical low growing shrubs may formed basitonical branching organ pipe cacti are formed Also basi or
.
by
long columns
.
of
Angle
of
.
E
tonical branching
the
”
“
22
-
be
by
,
be
Cok
AN
Cok
branching
, is
in
of
as
,
A
).
of
.
(
,
.
)
of
at
is
is
a
to
.
the geotropic
of is
In a
of
.
,
it
of of
of
,
in
classification
these
other
cacti
.
the first year and the first lateral
regard
or
(
- -
a
,
an
As
120
a
stem
Without further
may be branching affect the angle said that the angle branch formation study characteristic each species Rhipsalis this character the terete species distinguishing species but this was used fact has been completely overlooked the of
mary
,
)
.
of
.
-
Shrub like plants also arise by basitonical branching This means that the lateral branches arise near the base the latest growth example Harrisia Erio cereus Guelichii has well developed pri
the very the large branches extremely species fleshy columnar are weighty mini transverse growth apparently very Here geotropism mum strong since the lateral branches rapidly be short come perpendicular columns after growth horizontal
in
.
.
.
a
if
-
.
- - - -
O
,
O
as
If ,
.
)
(
such
121
Because
,
,
.
of
in
brasiliensis and some Cylindro Bige spinosissima and the young plant branches acrotoni lovii cally the plant becomes shrub like Tree like forms are derived the primary stem grows for long time before branching
Opuntia
Fig
.
,
is
in
often occurs only especially with branching mainly when this which acrotonical means that the lateral branches arise only This may the latest growth from the top Brasiliopuntia young be observed
connection
on
Longitudinal symmetry
puntias
Dendrocereus nudiflorus Fig nine year specimen shows the wide angle branching but the fully grown plant shows the wide crown and overhanging branches old
basitony
.
and
(
.
D
Acrotony
an a
,
In
.
a
After Troll
a
buds
.
a
, : )
-
,
angle effects slender growth especially succulents the great weight the spreading effect branches causes thick
.
cotyledonary
-
(
Harrisia Eriocereus branching basitonical ypocotyl Co cotyledons Cok lateral shoot from the
.
, ,
WA -p -p
rimary stem rimary root
of
Seedling showing
H -h
119
.
.
Fig
Guelichii
of
es
The angles between the stem and branch and the angles between two branches are obviously quite different for different plants growth habits That wide angle effects spreading habit and acute
. S .
.
E
121
Mature plant Dendrocereus nudiflorus
.
showing jointed overhanging branches
of
FIG
.
:
7
52 .
.
by
of
.,
Cact
.
Jour
Dendrocereus nudiflorus showing longitudinal sym growth Photograph Mrs de Gimenez Lanier Succ Soc Am
.
by
specimen periodic
.
in
.
.
120 Nine year old cultivated metry and jointed branches formed
.
FIG
HABIT OF THE CACTACEAE
a
net
the
in
as
, .
furround net near near the surface Although the Cylindr radial opuntia soon becomes ovate and the joints change from flattened radial base bearing form bearing form the areoles about itself with
with primordial meristem
a
to
.
all
a
-
a
of
The
to
the
”
"
,
, 2
and
large
the leaf cacti and Platy opuntia which
of
bilateral flat shoots the flattened joints
developed
: ; 1,
of
to
.
in
is
.
of
a
.
not
it
very striking radial cacti have being radial symmetry the shoots The beauty shoots e geometri part due the Mammillaria the geometri cally arranged mamilles There are are two two exceptions this radial symmetry the
aa
Most
is
these the midrib
factor and bilateral symmetry
and
nous
,
F
Light
SYMMETRY
is
LIGHT FACTOR AND BILATERAL
,
(
).
.
in
have leaves and areoles about the whole Epi The latter type also occurs surface phyllanthus opuntioides obovatus As previously shown the winged Alat
Epiphyllum
are derived
from
of
by
of
branches
al
of
.
to
of
a
radial shoots reduction the number ribs two The young branches are ways terete and the mature plants some
.
E
in
as
or
.
.
.
.
of
.
to
be
is
in
If
a
,
in
)
(
til
18
.
thes
an
.
in
.
,
the light factor However there must hereditary factor associated with flat joints because some Opuntias always form flat tened joints the dark
of
The most interesting effect the light factor occurs the two genera Stropho cactus and Deamia which not only become bilateral but also dorsiventral that the upper and lower sides the shoots are dif ferent Strophocactus Wittii Fig 122 has winged branches somewhat similar Epiphyllum those and the areoles occur only along the margin the two wings Because the two ribs enlarge the part to
)
.
(
of
;
is ,
,
.
in
of
In
of
.
a
are
the light There part caused by
of
to
. joints Platyopuntia totally different manner
flattened in
The formed
fore the bilateral growth
, ,
. in
be
,
of
It
.
the areoles
tened perpendicularly
in
-
of
.
R
is
a
species have terete midrib from which arise the lateral vascular strands leading
Schumann
joints These are therefore truly flattened But Goebel cultivated specimens Goebel Opuntia leucotricha the dark and found fleshy that they produced terete slightly joints strong light was applied from one direction the terete joints became flat
.
-
-
2
as
.
to R
.
of
,
in
it
environmental factors should noted that the bilateral branches these
Strophocactus
.
-
im
)
18
(
. or
of
.
.
by
122
Wittii After
-
as
as
far
come
it
are
as
.
In
in
the dark Rhipsalis Sect Phyllorhipsalis the winged flat joints Houlletiana and light does Epi others react similarly phyllum crispata However the species and others never have terete portions flat joints Zygocactus also never has terete bases when cultivated low light intensity appears that bilateral growth From this these species controlled by heredity not vated
FIG
of
a
in
a
In
)
.
18
, isis
dat if
of plant
(
of
if it
to
The fat shoot may become terete again begins grow toward the top new period growth study the Epiphyllum Aat branches was found grown under poor light that birazhe the plant conditions the newsighoots shoots are much nar rower and terete but they become flat soon the lighting conditions are proved Also Goebel found that Epiphyllum branches become terete without wings succulence when culti
2
. -
,
to 4
3
of
of
.
E
-
3
. ,
species have winged branches phyllanthus and lepidocarpum Because the basal part the branches and even the stem these plants are terete sometimes winged they show longitudinal sym metry
HABIT OF THE CACTACEAE
det
of on
a 4 -
At
to
in
its
in
, . A . rib
en
is
)
of
at
an
re
.
4
-
.
a
Alattened
the
in
the Cactaceae
the Cactaceae branch from the branch from the furrow the areole from the Myrtillocactus and axil Some species Notocactus have also been observed form adventitious stems from the roots Because quite small the growing point areole deeply laced connection between there
or
of
of
to
.
is
an
of
-
-
Branching
and only the Mammillarieae
is a
de
in
the ribbed
arise from 5
.
shoots
a
the
In
under
of .
as
the same manner
Aerial roots
Most
areoles
.
. ,
,
-
3
is
,
rib
and are appressed
Strophocactus side
becoming
of
.
G
is
,
,
's
stem
in
than the third
tree
low
in
in
shoot
lateral
light
.
is
of
- .
or In
a
to
veloped
.
(F
.
.
light intensity Aerial roots the duced the angle between the top and lateral arise young Fig 123 also shows ribbed ribs
.
to
,
stem
The two lateral ribs are covered the two uppermost ribs and become
each other
by
by
-
to
,
is
.
.
.
to
a
in
the support the surface next forms aerial roots which fasten the flat shoot the host stem Therefore this cactus bilateral and dorsiventral Even more striking the dorsiventral growth Deamia testudo another climbing cactus This plant occurs with three four five winged ribs but usually ribbed this case two ribs are more highly ing
Mexico
ig 123 one turned toward branch supporting the stem and becomes greatly The uppermost ribs become reduced angle larged and spread 180° from
.
, .
stem
home
.
at
a
in
are as
in
as
of
a
tree the cactus becomes closely spiral arrangement the The plant surface away from the
light that
testudo
S
Fig
.
of
.
Aat as
, of
.
.
of
stem
Deamia
Columbia
the much under the areoles nearly internodal region the shoots like the ribbon The shoots are not even terete the plant which receives The side base growth light surpasses the other side the plant grows along the this Because appressed
123a
.
in
rib
a
of - to
is
II -
;
. to
rib
FIG . 123. Deamia testudo : I- Part of a branch has viewed from above . A part of the left been removed show the adventitious roots angled lateral the right between the ribs shoot has begun flatten the side receiving the light Cross section mature branch The two wings the upper surface are well developed and the fifth the the middle very small After Goebel lower surface
IN THE CACTACEAE
BRANCHING
57
, ,
)
the
large areoles the joints
is
of
of
bow This pressure
.
a
by
be
to
shot from that they seem brought about effect may the walls surrounding because the rather thick base
a
be
so
(
In
.
,
,
,
salidopsis Rhipsalis cereuscula and other terete species Epiphyllanthus and Zygo cactus become too dry they also dehisce Epiphyllopsis Schlumbergera Gaert rapid neri the last joints are pulled down
.
crushed In
do of
Gymnocalycium some species and the shoots not appear from the areole but rather from the depth the plant body perforate the ribor These tubercle above the areole Apparently the point grown over growth the areole by the fleshy tissue the rib and the spiny is
of
by
,
)
of
of
(
.
it
.
of
.
( 7 )
,
In
,
of
bya
or
at
of
as
,
as
a
,
of
of
too
.
if
.
a
.
of
of
,
of .
of
.
and others often have Vaupel equal size
.
of
,
.
,
of
.
a
young seedling
Mam
,
rhodantha
but
it
top
to
of
as an
,
(
be
error for
formed three heads near appeared that this growth
an
of
injury the primordial The heads were not equal heads Also four heads apparently the development resulted
from
.
stem
of
as
,
.
of
heads
that these are lateral branches the top the plant and are the same the primordial plant This seems
millaria the
rhodantha
four
.
- of or -
-
of .
to
or
.
,
of
a
in
aid
do
,
frost
states
from size
M
. .
,
M
in
,
sonii
two
or
In
in
.
-
.
,
,
not remain vegetative ods but these propagate the species When the joints Rhip the Rhipsalidanae such some
disease
arise been
formed Sometimes isolated tubercles some the Mammillarieae may form roots especially longi those Dolichothele mamma Mammillaria Parkin Old specimens
)
.
11 .55
or
.
.
cm
#
corky abscission some evidence that joints but the falling this has not been proved Often roots are joints while the formed from the shoots joints are still attached the stem Opuntia The flattened leaf like joints brasiliensis are deciduous during dry peri layer may
parts
growing point may plant which has
new
capable cacti remain growth for long time Therefore various tissues will form new roots under the proper conditions This occurs only the corky surface layers the plant are not corky specimen hard For example Tephrocactus Tephrocactus papyracanthus did not form roots for two years but soon the corky layer was removed roots were rapidly or
)
35
.
(
..
cm cm
i
.
)
to
2
In
tree shrub like cacti the vascular strands connecting the stem and branches other become very strong and woody cases they remain soft and the young shoots adily fall off and function vegetative reproduction This occurs Chamaecereus caespi Silvestrii Mammillaria gracilis Bigelovii Opuntia titia and others There
All
38
all
.
or
of
Mammillaria fasciculata forms short stolons from the axils old corky tubercles near Fig slightly below the ground level long and 124 The stolons are about long and after the new young shoots are about bout long the stolons disappear
is
some species
Mammillaria sulco Echinopsis Eryiesii
and
a
.
.
the
According
Ottonis
glandulifera
from the depth mostly destroyed
(
species
.
lent
basal axils Cact Succ
succu Stockwell
and the lateral branch
in
stem
the surface
Notocactus
187
to
the
Am
:
.
Soc
3
.
of Mammillaria
.
stolons arising from fusciculata From Jour
.,
.
.
Fig
124
callus Lateral
the podara reports that from the wound the severed top
elevated
Schumann also appear
32
the areole
with shoots may
ium
is
of
part
,
of
of
.
.
of
,
others
HABIT OF THE CACTACEAE
58
formation of four equal parts , each of which grew to a normal head ( Fig . 125 ) . Thus it appears that the development of two of two or four equal heads in in Mamment Mammillaria is caused by a true divisies division of the growing point and a true by not lateral shoots . similar division may have occurred in Epiphyllum an branch ( 21 ) . The branch was first terete and then formed two grow ing points , each of which developed into a A
normal flat shoot . The two flat shoots were parallel . Other deformations of the growing point growths known as cause monstrous " cristate " forms. Such a growth may be hereditary , so it should be referred to taxo
may
of
in
,
an
to
or
.
to
or
)
.
126
(
.
be
-
.
Wm
Hertrich
.
Photographed
.
.
Normal and monstrous form Cereus peruvianus Huntington Botanical Garden of
126
.
.
Fig
by
10
in
.
in
.
if
as
in
to
in
be -
in
.
to
.
( 38 ) seems to have also misinter the cristate forms which occur Cereus peruvianus and some other columnar cacti Fig He believed that this form podaria occurred because the humps would not join form ribs Actually the irregular rolls and humps are never formed anything similar parallel vertical rows This form can only abnormal preted
without cause Mammillaria magnimamma was observed The growing point first came stretched one direction the plant were become cristate Later the lengthened growing point was distorted direction perpendicular the first elon gated point This finally resulted the
a
aberration or forma .
.ity
the head
Mammillaria magnimamma
as an
Vaupel
.
partition
of
The
4 -
125
.
.
Fig
nomically
IN THE CACTACEAE
at
,
)
19
,
,
,
of
(
.
to
,
or a
by
var
nivea
,
.
submamulosus hotographs
is
Parodia microsperma by Haselton
.
,
geminispina
,
-
nobilis Malacocarpus
) .
-
Mammillaria
,
ex
of
, -
top row Cephalocereus
.
.
H
a
,
:
of
crests
de to
to
(
is
by17 )
.
of
is
it
)
13
(
.
Types
Lower row Solisia pectinata
to
.
in
of
.
the
.
FIG
127
form monstrous partly cris after being minced Also seed lings from cristate plants were usually cris tate when mature According Grant true cristation
were found
tate growth
to
when
comes deformed
(p
the normal growing point one direction The two ends the growing point continue form new tissue whereas the middle section stops actually quite similar growth Thus quatri partition Mammillaria Vries and Georgesku state that the cristate growth caused recessive hereditary character but this does not plain sufficiently the occurrence this type caused
.
,
.
of
and
throughout the flowering plants . peculiar growth is probably caused by This stimulus from disease the plant top by parasitic animals but injury reports not limited these Grant that when thousands cacti were injured by crushing electricity chemicals and other means not one cristate plant was Only Melocactus intortus plants formed
.
the Cact -
of growth
(7 a is ),
than
aceae , such as Taraxacum , the tree the garden variety Celosia cristata
It
other
128 )
it
and
127
be - is
( Fig .
The cristate form may occur in plants
ash
BRANCHING
PHYLOGENY
60
OF THE HABIT
OF THE CACTACEAE
fixed by heredity in certain races , and ex ternal stimuli are necessary before the de formation will occur . It is not known whether or not the Mendelian laws apply to this characteristic , but they probably do not . Backeberg ( 2 ) believes that cristation may be similar to the dichotomous division of the growing point in Mammillaria , but this does not explain why cristation occurs in species
which do not divide
in this
man
ner . Linsbauer (27 ) has discussed the occur in of panachure in cacti , especially Opuntia . Because this phenomenon is his it will tological rather than morphological rence
not be
further .
discussed
FIG . 128. Crested Carnegiea gigantea . Haselton photo in Desert Botanical Garden , Papago Park , Arizona
. PHYLOGENY OF THE HABIT OF
is
or
are
.
)
.
,
is
es
a
to
.
in
a
,
is
all
of
is
in
,
of
as
of
.
m
10
,
the
of
to
3
a
of
.
scales
and
long
In
.
)
,
130
,
but later bear
old and apparently
.
(F
ig
the flowers
shoots
,
in
.
P
other spe cies Pereskia and Rhodocactus leaves occur along the entire length and they arise aculeata
of
is
.
of
These first bear
dense arrangement
of
.
A
Undoubtedly the Pereskieae the most primitive group the Cactaceae This
129
.
a
.
shoots
leaves Pereskieae
do
of
or
(
,
short
.
.
.
to
will serve stimulate further this interesting field
with two different kinds shoots many other woody plants Those the first type are long shoots with long inter length nodes reaching Aris ing each year from the axils the leaves the long shoots are the second type shrub
a
it
of
so
,
far
in
research
a
.
of
in
of of
a
be
discussion
brown cortex but sometimes the cortex remains green for considerable time The similarity other dicots pecially Pereskia aculeata This clear plant which Aoral characteristics the climbing most primitive cacti form
,
.-
as
a
an
of in
as
of
to
of
in
of
it
is
in
ous fashion different lines evolution great interest Thus follow the evolution the habit cacti occurs the different living species These studies great importance will later dis cussion the phylogeny the Cactaceae Also only fraction this problem has been thus studied that perhaps this
highly branch These plants the branches and twigs are slender and without succulence Usually they soon and
in
.
be
,
of
it
explained gradual Therefore can development which follows certain tenden analog cies the family and occurs analog
Fig
ed
,
of
aa
is
it
to
its
-
a
plant does not appear sud The habit without association ancestors always the result long but rather long evolution from more primitive ancestors denly
in
of
its
THE CACTACEAE
shown both by analysis of the flowers and the growth habit . All species of Pereskia subgenus Rhodocactus are leafy trees and shrubs and not differ greatly from the any dicotyledonous tree habit shrub
of
4
.
,
of
of
be if
P
.
Maihuenia
in
.
, In
but
tehuelches these
.
a
M
Calentinii has except that several erect stems length arise above the soil surface This certainly indicates that the stem branches basitonically just de
.
also become succulent in
.
cm
20
to
he
is
Philippii
.
M
scribed
of .
for
,
as
.
8
of
,
similar habit
.
is
M
quite similar habit tehuelches monstrous Platyopuntia found by Brass field this monstrous plant every areole the stem forms short lateral joint Each areole these joints also forms lateral joints and until the plant becomes monstrous and eventually kills itself Fig 132 The only difference habit between this plant and Maihuenia that the latter In
.
a
in
.
(
on
of
in
could
remain short and become succu species the areoles bear short
is
or
habit
Bahia
both
so
.
a
This
das Contas
Pereskia derived from the habit one imagines that the branches
aculeata
lent shoots
,
to
it
shoot
.
oblong
M
.
short and Aeshy stem and short globular
grown
to
is
has
a
in
.
to
areole to
ovate
131
)
.
(F
ig
each
aculeata
.
,
,
-
of
but the primitive flowers clearly show belong the Pereskieae There are two different habits this genus tehuelches
Rio
a
be -
In
to
no
is
.
to
.
is
in
.
of
,
to
.
the facts are incom does not seem systematic value here There exists the Pereskieae the genus Maihuenia which already has very Aeshy living species branches There link genus this Pereskia habit Maihuenia especially much like the Opuntieae some species the sub genus Tephrocactus
readily
to T
re
,
or
spectively Although plete this relationship
- of
.
a
is a
or
,
As a
.
.
of
these At the base above the areoles leaves are some scales Therefore each leaf and few scales arise from short shoot previously discussed the effect acrotony basitiony tree shrub
near Calderao and Rose
.
Britton
at
the Catinga
From
on
Pereskia bahiensis
).
.
.
129
at
FIG
PERESKIEAE
PHYLOGENY
OF THE HABIT OF THE CACTACEAE
genus
the areoles of the stem bear short shoots which do not repeat the branching . The other growth habit occurring in the genus Maihuenia is that shown by Vaupel for M . Poeppigii (Fig . 131 ) . This plant , similar in habit to a Tephrocactus , is a growing plant with jointed bushy , low branches . The joints are oblong , rounded , 1. 5 cm . thick , and up to 6 cm . long . The joints have a determinate growth and branch acrotonically . Thus this species is
like the morphological type of Opuntia , differs considerably from Maihuenia tehuelches, which is a shortened form of Pereskia .
of FIG .
and
After Vaupel
.
;
Maihuenia Poeppigii
-
tehuelches
b
-
a
Maihuenia
.
ta
UN 131
.
.
FIG
flowers
.
leaves and latent buds
-
S
shoot with
,
,
-y j
:
a , K -
.
to
130 Scheme the growth Pereskia saccharosa old long oung long shoot shoot short
OPUNTIEAE
).
Fig
.
as
in
of
is
is
.
flattened
Platyopuntia and bears flat joints The following year this flat seedling tender
,
the Opunti
a
to
it
,
of
-
of
the oldest genera
in
.
tened lateral from the top erect terete stem formed which branches joints acrotonical whorls flattened This stem grows indeterminately whereas all the flat joints have determinate growth The flat joints may form branches terete stems The flat joints differ from those
.
.
of
of
.
a
,
in
of
,
is
a
,
of
.
.
eae
,
of
its
be
one
as
non
Later the primitive nature flowers will be discussed but for the present the non succulent habit shows fleshy branches
,
slender
-
is
as other Opuntieae . shrub with
Tacinga
a
, just
es
Be
because some of the branches become broken and the symmetry is therefore lost . TK This type of branching occurs in Opuntia 134 alcahes and some other species ( Whorls also occur section Brasili opuntia but this genus primitive not flowers indicate Seedlings Brasili opuntia have primordial joint which its
.
The Opuntieae are descendants of the through the genus Pereskiopsis . Pereskieae genus has broad , Aeshy leaves as in This most species of Pereskia and Rhodocactus . Also the species in this genus are trees or shrubs with many long and slender branch -
try and branching is definitely acrotonical . Both of these characteristics occur in some of the Cylindropuntia . Also the branches are terete , green and fleshy , and are very easily detached from the stem . As the plant be branching is more irregular becomes old
as
B
Opuntieae
OPUNTIA Whatiaita
.
by
.
.
in
a
of
ole
in
132 Monstrous Plat yopuntia which each are the stem has formed shoot and this growth Photograph has been repeated each shoot Carl Brassfield
.
Fig
Another connecting
to
to
a
with
a
.
high
is
Zehntneri many branches Young specimens have central
. .
in
. . G .
by P
133 Quiabentia Zehnineri showing the acrotoni Photograph cal whorls Russell Britton and Rose
.
Fig
.
.
to
has up a
of
.
in
with nearly horizontal branches arising whorls The whorls are always developed near the top each growth unit The ten branches and may contain longitudinal symme tree very clearly
.
m
). to
2
.
(
Quiabentia
.
Cereeae
shrub Fig 133 erect stem
3
the
,
to
of .
in
is
link the Pereski eae shown the genus Quiabentia This genus shows some characteristics habit Cylindropuntia which lead and also
PHYLOGENY
OF THE HABIT
OF THE CACTACEAE
branch
)
(
.
.
later
Opuntia
first
inde
whorls
.
and
grow
in
is
described
.
.
.
.
a
,
young plant with acrotoni 134 Opuntia alcahes cal whorls From Britton and Rose
.
FIG
spines
sheathed
to
have
pachypus terminately
to
not
A
its
do
The species of Cylindropuntia which still have slender fleshy branches may be as sumed to be of the most primitive habit . 0 . leptocaules , O . ramosissima , and O . cary baea all are much branched shrubs , and the branches are not thicker than a pencil . Pos sibly as primitive as these are the species of Austrocylindropuntia ( Backeberg ) which , big have leaves such as A . (Opuntia ) subu lata and A . (Opuntia ) exaltata . Austro cylindindropuntia may be more primitive species than Cylindropuntia because
in
in
to
asso
of
.
determinate terete stems Indeterminate growth with acrotonical whorls which
form
is
of
the
opuntia the progressive tendency Alat joints determinate growth primitive character ciated with
of
.
or
,
Platyopuntia they because are flattened horizontally whereas the latter are flattened more less vertically Thus Brasili
by
Photograph Am
Otto
253
1 :
Soc
.,
.
Succ
.
.
Cact
.
Jour
.
Bigelovii
Opuntia
Roller
.
.
.
,
.
,
O
)
.
FIG 135
in
.
(
in
of
is
a
or
,
to
terete stem connected terete more less cylindrical joints Opuntia Bige determinate growth occurs spinosior and possibly lovii Fig 135 others
OPUNTIEAE
is
It
of
in
.
in
is
.
.
early
2
.
Mexico
454
:
Am
,
Coahuila
.,
Soc
,
Paila
Succ
.
Cact
de la
the Sierra
glochids have been
because
.
the Opuntieae
Jour
of
a
to
to
The third trend leads Grusonia which appears be highly developed genus
.
south
Count Knuth
is a species
.
.
,
Grusonia Bradtiana
Photograph
by
.
.
136
in of
is
all
in
,
are
in
.
-
in
the sub genus Tephrocactus and also Backeberg found that Cylindropuntia and Austrocylindropuntia the globular joints become lengthened when the plants grown under glass but Tephrocactus the globular habit fixed under conditions
the Opuntieae . Also with globular joints , worthy Hickenii notice that this Opuntia Chaffeii Same tendency occurs which has very tender low branches which drop during dry periods evolved very in this genus
P
power
From the habit of Cylindropuntia evolu tion proceeded in several different direc tions . One tendency was for the joints to become shortened into a globular form : Such globular jointed species of low growth generelth joininicpecies of globular greche are found in Ó . tunicata , O . Verschaffeltii ,
Fig
Another line of evolution leads to a habit or less slender and weak branches in which the succulence is found to the greatest extent in the hypocotyl and root. This leads to Pterocactus which apparently of more
.
Thus the primitive nature of Quiabentia .
repeated
OF THE HABIT
OF THE CACTACEAE
.
.
.
in
&
Right
Jour Cact Succ Am
:
",
., 7 by : J 77 . . . R .
,
of
Young plant the Consolea rubescens showing the very regular Lorraine Photograph Sons Pirtle
.
"
Croix branching
. de
.
Fig
137
PHYLOGENY
Typical
Platyopuntia
the po distinctly elevated more less Austrocylindropuntia Opuntia Sal
are
.
to
.
,
)
(
.
37
of
's
if
.
-
)
(
, .
”
de
-
“
-
,
of of
It
.
a
of
.
,
O
in
.
low
of
.
O
of ,
as
in
of
.
of
.
to
a
of
.
, -
-
a
,
5
or m .
.
cm
of
a
a
.
.
O
as
,
.
as
in
.
a
by
is
of
so
,
in
an
of
.
in
of
,
are
of
.
in
O
.
of
of
.
.
there are also dwarf species such microdisca well the large species the branches size and the joints the stem increase firmly attached that the end become marked only fine furrow each joint course
as
.
-
in
indeterminate growing flat stem which also shows longitudinal symmetry and periodical thickening The flat branches the edge this arise acrotonically from not the type Aat stem Also these joints found Opuntia for they branch clearly
height
over and have more the trunk diameter large whole plant appearing tree like with Opuntia ficus indica and others crown form rather tall shrubs and small shrubs phaeacantha and many others occur Of cantha attain
the stem occurs Indeterminate growth the flat jointed Opuntia ammophila The genus Consolea must also be placed the group having indeterminate growth the Young plants this genus have stem
30
the section
.
as
.
of
members
a
. -
.
,
.
in
as
-
is
,
has both terete and Therefore these species must be considered the most primitive also
joints
flattened
leucotricha the shrub like shown basitony but rather not result habit stops growing longitudi the primary stem early nally very then branches acrotoni Acrotony also causes the habit cally Opuntias The joints branches determinate growth and form these have This gives rise new joints from the top Opuntia the typical habit The greatest variety form occurs Opuntia mega Some species such is
in to
.
of
of
to
of
-
.
of
evolution leads Platyopuntia Also the char thë Alat jointed Opuntieae some acteristics the primitive genera reoccur The tendency form flat joints must be very old for the primary stem the seed ling first terete but soon becomes flat just tened the very primitive section Brasiliopuntia Opuntia aurantiaca first has terete joints and later small flattened joints are formed and Opuntia Chaffeii has very joints Opuntia small slightly flattened Schickendantzii
Aattened
.
.
of
.
The last direction
thg fattened joints
acrotonically and the new shoots transversely the first The name Lorraine was derived from this Croix Fig 137 cross jointed characteristics the genus Opuntia tree like shrubby and low prostrate species occur The tree the primary stem grow like habit occurs time before for considerable length branching acrotonically has As Troll again
In
in
as
in
the
)
),
Grusonia miana has terete joints podaria are connected into true ribs Fig 136 the Cereeae
(
-
)
(
(
only
or
daria
Cylindropuntia
in
Whereas are
.
lost
CEREEAE genera of different evolutionary lines , al though similar in habit , differ in the mor phology of the Aower . Although Grusonia is similar in habit to
or
,
e
For
.
cm
.
,
.
. .
1 5
1
,
to
is a
in in
.
of
in
a
3
.
.
prostratus
.
adscendens 156
and
are
more ,
,
fewer branches
and others are ascending species
.
2 )
,
in
.
is 8 -
.
as
of
L
as (
.
succulent
Harrisia 138 Eriocereus From Britton and Rose .
.
to
have
:
by
is
FIG
the
inin
.
L 5
.
of
,
as
of
Many
the Pereskieae
genus
6
a
,
of
I
is
of
evolved habit This indicates that the differ ent lines evolution were separated long ago and this also shown the absence connecting links between the groups
well
corky
ribbed branches The trunk this only diameter and has cortex Some other species this
tree
.
as
as
arid
.
adapted
It
habit
climates well tropical climates are found here This sub tribe divided into several evolutionary evolutionary development and each these lines primitive lines begins with morphologically highly genera and species and reaches Plants
cm
,
is
m
others
Cereeae
variety
to
-tribe , the
Cereus
in
sub
.
which exceeds
Cereeae
of
third
the
to all
It is
.
in
C
, there is no doubt today that no connection between the Opun tieae and the Cereeae. The most primitive of the Cereeae are probably those which are branched like other dicotyledonous , trees and shrubs have tender branches with Species few ribs and are climbers that especi branch like other dicot trees occu occur trees especi dicot ally genus Leptocereus the genus For example ally Leonii much brandeus branched shrub tree height much and has thick height any
there
the
of the smaller Opuntias , especially , the joints drop off up to the microdasys O. poor under environmental conditions . stem This same tendency occurs as previously discussed in the section Brasiliopuntia and in the Rhipsalidanae . In some
appears that strength
OF THE HABIT OF THE CACTACEAE
.
)
.
-
or
:
;
Well branched slightly thickened reeping half
d -C
(
)
-
,
-
,
,
;
)
(t
Browningia
are
to
is
,
.
a
columnar species the ascending species more primitive Lepto clearly related canthocereus
.
at
with mechanical tissue Therefore genus containing both ascending and erect stem
in
the
first did not support the stem
A
ening
of
.
.
a
tree with determinate candelaris the trunk and extreme acrotonical branching From Britton and Rose
,
.
.
140 growth
of
Fig
-
c -
(
)
;
(
b -
,
of
to
.
.
tree and shrub like forms weak slender branched the evolution Schemes 139 gracilis type Harrisia Few branched habit the branches habit with slender branches type Acanthocereus type Harrisia Nashii Shrub with drooping slender branches ype Eriocereus adscendens ascending shrub
a -
Fig
PHYLOGENY
CEREEAE
Fig . 141. Schemes to the evolution from a very succulent well -branched tree to a large candelabra and simple column by mechanical thickening and loss of branching : a-Highly branched tree, jointed , non - sturdy branches ( type Dendrocereus nudiflorus ) ; b - Tree with determinate growth of the stem and few acrotonically formed branches , a secondary evolu tionary form ( type Browningia candelaris ) ; c- Well -branched tree with mesotonic branching and strong branches (type Escontria ) ; d- Tree with basitonical branching and high columnar branches ( type Lemaireocereus Weberi ) ; e- Typical chandelier ( type Carnegiea gigantea ) .
,
.
,
M
.
4
or
3
.
(
a
in
.
M
so
all
to
)
45
.
A
,
.
is
a
)
(
a
to
of
in
,
with more less succulent branches the plants being trees shrubs depending upon
,
or
half erect
hanging
or
also
,
Nyctocereus
,
related
of
,
Heliocereus
shows these changes
to
.
of
-
.
H
of
-
to
-
resemblance sticks that they lose cactus Two evolutionary lines are shown these primitive genera First the development mechanical tissue without the loss the highly branched habit This forms plants
.
(F
in
as
,
of
.Figin
,
.
ascending branches
genus with partly heterogenous group This erect and partly hanging branches pencillata about especially Arrojadoa writes that old which Werdermann dry parts similar this shrub become
,
.
.
H
(
. eri
in
in
in
,
),
).
to
.
forms
shortly
ribs The same shape oc Monvillea especially Cavendishii Spegazzinii has only three ribs whereas very There also the Cephalocerei curs
in
,
to
),
5
or
by
(
or
The genus Harrisia also contains spe thin branches cies with crowns species with few and slender gracilis and branches 139 shows this evolution habit from the richly but thin branched trees the few branched the ascending 138
climbing , and with only
is
.
higher degree
The ribs reduced three the plants are less branched and the branches are not sturdy but droop leaning against other plants climb branching and climbing columnar Similar stems with many ribs also occur the genus Nyctocereus partly Harrisia ig ophora and Eriocereus adscendens 4
evolution rarely
a
of
of
habit and shows
.
in
are
cereus
PHYLOGENY
HABIT OF THE CACTACEAE
OF THE
large , somewhat erector branches which usually do not further branch . The stem obviously grows for a long time without branching , but growth of the stem is finally terminated and branches arise in a whorl acrotonically . The pseudowhorls may arise , as in Rhipsalis cereuscula , if the growing point of the pri mary stem is injured before it finishes normal growth the branches true whorl not develop normal manner Backeberg has written that Brown simple spiny ingia has columnar stem the juvenal stem Towards the top this column becomes gradually smaller and has fewer spines and the top there
whorls many
of
. of
,
at
no
,
it
(
).
a
,
(2 )
in
a
a
if
do
, or
its
hanging
different habit
.
a
of
,
of
of
by
lateral branches are that the long shoots
Second by loss later globular
of
branching
columnar plants are formed
,
the form branching
.
of
.
,
of
.
Photograph 142 Cereus peruvianus Janes courtesy the Botanical Institute University Vienna
.
by .Fig
,
of
of
.
be
,
,
to
, .
is
.
by
spines and the stem does not grow Spineless branches now arise further Backeberg probably extreme acrotony correct therefore call the spiny stem the juvenal stem and the spineless branching part the old stem This growth may compared with the formation the long Rhipsalis Saglionis which branches shoots only after finishing apical growth and the are
.
and
is
a
,
.
is
of es ,
is a
in
.
,
a
.
of
,
,
.
,
of
a
.
well Here
in of
development shown Neoabbottia and Den big trunk has drocereus much very succulent branched top composed highly ribbed branches The branches are longitudinal symmetry jointed and show are not sturdy and are irregular and hang downward Plants Dendrocereus grown long stem from seeds become trees with and highly branched crown whereas plants grown vegetatively from lateral branches soon become branched and shrubby This polarity shows that there the branch capable for only the primordial stem lengthening without branching Jasmino cereus also has jointed somewhat irregular The first line
exceedingly
,
a
Lemaireocereus Weberi tree with From Britton and Rose
.
basitonical branching
.
143
.
Fig
.
.
.
)
or
in
its
at
a
(
A
in
.
ly
branched species very peculiar growth habit occurs Browningia candelaris Fig 140 This tree has thick slightly conical trunk which top pseudo bears only whorls
than
mi
CEREEAE
all
are formed
of
lines evolution certain In nearly species already have developed strong col
.
.
.
branches
.
the columnar
118
., 8 :
Basitonical branching Jour Cact Succ Soc Am
of
umnar branches with rather rich branching
grandis
From
.
Pachycereus
shrubby forms the branches acrotonically .
.
.
144
.
.Fig
Lemaireocereus laetus is similar in habit to Jasminocereus in that it has jointed branch es , but this species also can become shrub like whenever the stem does not develop . Nevertheless , it appears that even in these
€
PHYLOGENY
OF THE HABIT
OF THE CACTACEAE
If branching of the stems becomes basi tonical the branching of the primary branches is also displaced to their basal part . In this way trees result with a large crown consisting of many erect columns . This habit exists along many different lines of evolution , and is especially striking in Le maireocereus Weberi ( Fig . 143 ) , which has a very broad crown formed of many col umns . Also Pachycereus chrysomallus has a more narrow but very high crown ( up to 12 to 18 m . ) . Pachycereus grandis var . gigas is more complex ( Fig. 144 ) . It is not known whether or not the primary stem of this species branches acrotonically be parts occur just cause many large stem above the soil surface . The branches defi nitely branch basitonically and the crown is quite similar to that of Lemaireocereus Weberi. In Pachycereus Pringlei ( Fig . 145 ) the number of branches is reduced to a few , sometimes even a single column . The branches reach a diameter of up to 0 .5 m . This habit leads to further reduction of lateral branching and greater increase of the diameter of the lateral branches . This is shown in Carnegiea gigantea ( Fig . 146 ) .
Pringlei, near Punta Prieta , Fig . 145. Pachycereus Lower California , showing the large short stem and few basitonically formed columns . From Britton and Rose .
of
.
146 Carnegiea gigantea Typical candelabra large columnar stem and few large branches
, a
very
.
anoi.
.
Pilocereus Ulei , P. Robinii , Myrtillocactus , few -branched forms of Espostoa lanata , and in Trichocereus peruvianus and T . Pach
Fig
(Fig . 141 ). When the stem branches late acrotonically , a tree with a very big and densely branched crown is formed . If branching begins at an early stage , columnar shrubs are formed . This is shown by the shrubby specimens ( 5 to 7 m . high ) of Lemaireocereus chende , and in the tree -like Escontria . Here the branches re -branch and the crown consists of many rather short columns ( Fig . 1410 ) . This habit also occurs in Cereus Hildmannianus , C . peruvianus ( Fig . 142 ) and others , and in Stetsonia ,
CEREEAE
by
,
is
-
.
in
is
as
is
It
.
)
. of .
(
)
.
(
on
Swett
;
.
Chas
, ),
(
in
is
as
.
Photo
W
Viscaino Expeditions
.
.”
,
of
.
lie
do
,
in
.
,
a
is
of
of
of
.
of
In
the Desierto
Allan Hancock
by
gummosus Courtesy
,
Muchaerocereus
on
.
.
in
,
147
.
.
Fig
be
to
in
of
.
-
in
of
) ",
.
m
4
been discussed but these habits were formed differently the Trichocerei this group the slender stem first became succulent and columnar before becoming woody with mechanical tissue
.
- .
,
and the largest
columns this type occur Neorai mondia macrostibas The development the crowns and simple columns found the Pachycerei has high
ilocereus Gounelli Fig 148 shows the evolution one the strangest habits plant the Cactaceae This plant low with few branches which usually form from the lower parts The lowest branches length sometimes become several meters strong and because they not have me chanical tissue they the soil without bearing lateral roots This habit may ex plain the habit Machaerocereus eruca Creeping Devil the This plant forms long and thick creeping columns Fig 149 but the development these not known may although Pilocereus the same
.
Cacti
( up
“
of
pipe
Heliocereus
it
-
)
or
If,
of
.
(
Neobuxbaumia trajani during the evolution branching the trunk becomes suppressed along with very early acrotonical basitonical branching many erect columns are formed shrubs Organ These are commonly called the
de
in
,
as
)
.
This habit unbranched columns Pachycereus columna
In the Heliocerei , which may belong to the ancestors of the Trichocerei, thickening the branches forms half erect but still highly branched shrubs This shown Bergerocactus and Machaerocereus gum Fig 147 probable that the mosus branching these plants basitonical of
to
. of
m
“
24
P
12 to
is
years
200 to
to
height of
have a circumference greatest age estimated 34
Its
over 2 m .
from 150 finally leads
a
may
be
reaches
(
plant
This
and the stem
PHYLOGENY
Gounellei , Catinga of Parahyba , Brazil. A candelabra with basitonical branching
.
T
,
, . T
in
.
of
,
to to
a
of
"
.
E
of
,
,
.
-
in
Another line evolution occurs Echi nocereus This development begins with the slender low much branched plants
the
Trichocereus huascha candicans and strigosus results more highly evolved group this habit occurs Cleistocactus anguineus and even higher Oreocereus Celsianus Further development few basitonically formed branches leads single columns similar shrubs and then Carnegiea gigantea but not great such height Trichocereus chiloensis represents to
of
.
-
“
In
.
to
-
from
with
.
.
of these cacti
the young parts the soil and absorb nutri chor the plant creep ents this way the plants actually along the soil surface with one end decay ing and the other end growing roots
long and prostrate
.
parts
,
The old
an die
lowest branches becoming
in
Pilocereus
Gounelli . but lateral
THE CACTACEAE
OF
a
148.
In
.
Fig
OF THE HABIT
. to
(
.
C
It
).
(F
a
arise
seen
be the effect
that
of
be
.
.
.
it
may
very
similar
different This fact has
very
.
be in of
(
).
of
to
be
in
it
.
Fig 151 lines evolution sufficiently not been considered cactus great research and will shown phylo importance later discussions the geny the Cactaceae
of
of
-
If
can
,
to
in
Sil -
in
as
-
In
.
T
,
is
,
of
.
.
vestrii the basal branches become strengthened the small lawn like habit
Thus habits
of
to
.
creeping habit somewhat similar that Machaerocereus eruca found coquim Trichocereus thelegonus and poor culture conditions this habit banus becomes snake like Chamaecereus A
)
- -
.
E
.
E
of
the
be
E
of
to
-
.
E
! of
-
the.
to to
)
(5
)
10
.
sis
to
trunk and columnar branches which basitonically
ribs
(8
rigidissimus and even columns nearly globular plants Standleyi
7
gigantea well the shrubby group and Fig represents the single column type 150b Trichocereus Terscheckii appears ig 150a intermediate has short
to
ribs and Scheeri Scheeri and progresses the more succu many hegresses lent and many headed plants mohaven This leads finally the single short
Blanckii
to
or
,
.
of
-
of
.
of
,
.
; -
b
.
.
is
-
,
,
showing the low candelabra with Terscheckii Carnegiea gigantes lower part the stem of
California
cactus ingens and others the large globular plants Echinocactus Grusonii The globular many headed habit Re butia also belongs this line evolution was observed that species which become short columnar forms when the plant to
.
to
to
,
, as
in
Trichocereus
-
a
.
150
Lower
and Rose
,
in
in
,
,
Fig
.
-
A
,
further evolution from short creep columnar forms may result short globular habits columnar and finally This development which tends form large short col columns can also lead umns Ferocactus acanthodes Echino ing
near Santa Domingo
,
Devil
",
Creeping
From Britton
It
the
"
eruca
,
149 Machaerocereus
.
.
Fig
CEREEAE
branches
arising
from
the
OF THE HABIT OF THE CACTACEAE CACTACEAE THE
PHYLOGENY
,
, to
of
-s
(
,
: a -C
of
or
or
by
to
.
.
Fig 151 Schemes the evolution from very succulent weak shrub like forms the prostrate habit low candelabras means basitonical mesotonical branch reep ing the primary stems ing non turdy columns type b -
,
(
) ; d (t -
)
;
c - (
-
) ;
Pros Machaerocereus eruca ascending trate like habit with type branches Trichocereus Huascha Prostrate with often drooping mesotonically formed ype Chamaecereus Sil branches vestrii Basitonical branching permanent type Re low form of
.
in ,
f-
;
es ) (
)
e -
;
Groups butia minuscula basitonically formed erect branch type Neoraimondia macrosti usually bas Candelabra form
some other authors This extension the genus Lepismium based flowers arising
.
in
.
,
of
's
,
-
,
sub
by
be
,
on
.
of
.
,
in
ium
is
in
be
.
of
to
is
of
a
of
a
,
a
,
in
.
to
.
,
of
,
is
is
is
for
in
of
a
,
a
of
was shown previously
,
-
as
the
.
of
to
.
.
,
- .
- -
of
,
of
of
be
on
subject
's
do
or
is
a
.
by
is
no
a
monograph
complete discussion almost im Nevertheless because only few species are used the division the genus discussion these not essential the present study that possible
As
,
of ,
it
, .
of
to
of
A
would involve
deepened
work
a
is
It
of
.
.
A
,
,
, ,
,
like the Cereeae detailed explanation the habit the Rhipsalides will not presented confirm systematic placement this new This dis cussion means complete that
this
in
is
,
.
,
to
at
,
as
and that the primordial shoot cereiform This means according the Law Re capitulation that the ancestors Rhip salides were already quite succulent few ribbed and basitonical branching habit
accepted areole will not for the generic character very short flower tube with very tuft long hairs arising from the flowering are Rhipsalidanae oles The whole tribe not yet clarified sufficiently warrant such change the genera Therefore this work the genus Lepismium will discussed the old sense containing only Backe Finally material berg Eulepismium very difficult some Rhipsalides obtain from
so
the columnar ancestors and the true globular forms have this habit fixed Commonly the group Rhipsalides placed the beginning the Sub family Berger Cereeae did assumed many that the slender ribless branches Rhipsalides the simple Aowers and the short and unbranched funiculi the seeds are primitive characteristics Actually these characters are not primitive for has been Rhipsalis shown that seedlings not cotyledons have primitive hypocotyls related
a
,
is
.
in
a
,
fully seedlings elongated mature have whereas species which have true globular habit are already globular the seedling Therefore the first group more closely
is
-
incomplete and au because the literature disagreement Even the divi thors are sions the genus Rhipsalis are not definite nor are the species clearly distinct For ex Rhip ample Backeberg places Schumann Epallagon Calamorhipsalis genera salis Trigonorhipsalis and the genus Lepismium This has also been followed
of
(t
)
.
in
of
as
broader and lower than the first Figure 141 evolution line ype Trichocereus Terscheckii
the highly
de
CEREEAE
.
This species also leads to terete species
Rhipsalis
,
floccosa
R
.
chrysocarpa
,
and
R
. .
the .
10
to
Pacheco -Leonii are also said to branch laterally . The latter species has different branches , the first of which are cereiform , 6 to 9 - ribbed , and have a length of 5 to 15 and diameter mm Later 8
.
its
third year , typical shoots are formed which branch extremely acrotonically . Also the genus Hariota grows for a long time cerei formly before forming typical branches
mm
of
-
cereu
a
.
are
R
.
Rhipsalis , such as
seedlings and branch scula , have cereiform ing is basitonical , but later , sometimes in the
cm
of
rived species
be in
.
to
,
as
,
to
-
6
to
4
angled angles branches and alternate from areole areole occurs Rhipsalis paradoxa The last branches
is
,
of
.
the genus
Lepismium
,
This
.
in
as
-
a
in
in
of
.
formed are nearly cylindrical This plant thus clearly shows the tendencies evolu tion different directions The most striking lateral branching oc Rhipsalis which sub genus curs by Phyllorhipsalis termed Schumann and genus must
.
FIG
2
. ,
a
,
.
.
Rhipsalis leiophloea Phyllorhipsalis 152 showing terete long shoots and only lateral flat shoots .
After Vaupel
be
In
mesotonical
whereas acro
later characteristic
or
.
a
tony
is
basitonical
,
or
is
or
of
,
it
of
.
a
The cereiform branches have definite agreement with the acrotonical growth Recapitulation Law must assumed that these members the Rhipsalides are the most primitive because they have ribbed angular branches and lateral branching
Acanthorhipsalis
micrantha has three branches shows mesotonical branching and the flowers have areoles and least scales the pericarp Rhipsalis rigida also branches the ovary laterally This species also vegetatively very primitive because areoleless flowers
Fig
a , ;
:
.
.
.
b -
,
to
,
of
K -
, ,
B -
.
Rhipsalis mesembryanthemoides 153 Young long shoot with indeterminate growth axillary bud which later forms short scale lateral terete shoots which because de terminate longitudinal growth are similar succulent leaves Mature branch with the short shoots After Troll ;
15 a
to
and reaches 8
stem
diameter
of
and
a
cm
. 5-
angled
.
of is
)
erect 60
It
has length
of an
.
(
.
at
on
,
ribbed
,
angled
or
more
PHYLOGENY
are
of
,
to
In
of
.
.
be
-
.
.
.
.
flat
,
are
of
,
in
.
in
is
of
(F
.
.
P
do
is
or
of
.
R
in
.
.
a
in
al
.
R
3
of
.
R
,
.
of
in
as
, by
a
in
This plant has indefinitely
and
of
).
-
in
as
of it
.
(
Fig 153 long terete
154 This species has small flattened branches and these bear lateral branches which are flattened the same plane the branching main branches This type Warmingiana but most shown the branches this species are sharply ngled Houlletiana also has lateral branches but this species flat shoots ways begin with terete portion Also the long shoots often are jointed and each joint begins with terete part From descriptions appears that platycarpa also belongs this group The third group the Phyllorhipsalides
.
-
.
an to
.
cm
40
to
bryanthemoides
up
Vaupel
After
places this group the Phyllo rhipsalides the two species Pseudorhipsalis himantoclada but these also alata and have long shoots which are flattened and therefore not belong this group Phyllorhipsalides The second group ig characterized best by Rhipsalis linearis
, .
or
is
in
A
.
.
in
,
be
is
.
At
this point should be mentioned branching that the same type this Rhipsalis mesem first group also occurs
ence
shoots
).
be
-
of
,
from the areoles arise flat lateral shoots determinate growth Sometimes two more arise from one areole habit like this occurs the Epiphyllanae Disocactus biformis but this plant not primitive and cannot related the Rhipsalides This similarity only evolutionary converg
small
also
-a
by
(
an
top
,
at
)
.
to
an
or
.
.
a
to
be
,
on
is
,
Ramulosae which characterized scales and therefore must the pericarp primitive group assumed The habit also shows primitive characteristics The angular plants have terete trunk only Rhipsalis Wercklei seems excep be tion and biform branches The long shoots only are terete Aattened the and
4
XII
,
group
the branches
it
.
Vaupel
All
linearis
's
belongs
Rhipsalis
.
.
FIG
species
154
to
2
.
(
)
is
.
by
shoots a
.
L
in
or
-
3
In
Phyllorhipsalis there are three distinct The first group best characterized Rhipsalis leiophloea Fig 152 This
types
. These not fleshy and very short oblong shoot from opposition each areole those Berger Phyllorhipsalis sub genus these short shoots are terete The phylogeny Vaupel this species will discussed later growing
they bear
.
be primitive because the stems
to
-
assumed
winged angled are often caver nosum whenever flat shoots are formed
's
be
OF THE HABIT OF THE CACTACEAE
this
.
.
.
R
in
of 3-
.
in
(
in
is
in
at . .
.
.
R
of
R
of
.
as
-
3
.
to
a
is
In
,
FIG
.
.
3 -
of
,
.
.
, .
Rhipsalis crispimargi 155 Phyllorhipsalis with de nata growth terminate the joints winged which are sometimes From Britton and Rose a
- -
5
whorls the jointed Phyllorhipsalides the longi tudinal symmetry and periodical thickening very typical habit which give the plant Zygocactus similar Also this group angled are species which sometimes have Rhipsalis chloroptera which stems such has very slender but long joints The some winged joints pachyptera and times crispimarginata Fig the terete stem place these species 155 this group too The highest evolution this group
-
- of
to
6
4
to
2
ME Eaton
paradoxa were derived acrotonically
branch
these
3
re - to -
7
.
i
to
.
in
of
.
-w
a
to
cm
. .
angled
)
,,
is
.
-
3
cm in
up
of
.
ap
.
is
.
a
.
by
-
5
cm
12
1
of
stem
-
,
Both
diameter This inged species Rhipsalis the few joints Tonduzii which has different types parted diameter arising whorls The lower branches are angled equal diam All the joints are from
,-
nating 3 natin
growth and the joints having determinate Onted growth Thiste group acrotonical branching This branching branchineterminate parently derived from species having radial joints with determinate growth and acro clearly tonical branching This type presented Rhipsalis pentaptera This spe winged branches which are cies has long and arise parted whorls leads
along the whole length From this along winged Rhipsalis Rhipsalis trigona trichon From and alter winged
and
stems
eter
eter form
3
distinct from the first two groups the characteristics are jointed stems
very
because
the
ng
-
is
CEREEAE
PHYLOGENY
THE CACTACEAE
OF
or winged Rhipsalides remain to be investi gated , the habit of the terete Rhipsalides has clarified the different shapes of these species ( 9 ) . Although the literature states that terete species of Rhipsalis , such as R .
VI
lumbricoides , R . chrysocarpa , R . rigida , and R . Novaesis branch mesotonically , these re ports appear to be in error . The plants de may have been injured so that scribed branching occurred below the stem tip . From recent studies the specimens ex amined have definite longitudinal growth and branch acrotonically The character istics seem be very old for Erythro rhipsalis pilocarpa which one the old members the Rhipsalides shows them extremely clear development This plant undoubtedly direct descendant
:
,
of
.
.
from
to
—
of
a
or
were derived
a
is
which terete Rhip them the genus Hariota Zygocactus related Therefore this plant will now
those ancestors part salides and the genera discussed
.
be
of
is
of
an
in
est
,
,
to
.
a
all
L
OF THE HABIT
,
es
in
.
be
.
)
.
of
(
of in R .
,
the ovate often nearly orbicular crispata and others Fig 156 Although many problems the angular
tained joints
an
be
.
It
byor
one row which increases the stem length appears that the same direction the joints should terminal elongations but actually the joints arise from areole near the top and not terminally This can arise
.
,
,
to
a
.
.
.
in
Rhipsalis
crispata jointed nearly with ovate joints formed bicular determinate extreme acrotonical branching 156
Phyllorhipsalis
in
FIG
of
of
Erythrorhipsalis The tender branches pilocarpa show the primitive character many spine hairs The branch the areoles are definitely articulate and the joints
observed
because the joints are always
.
d an -
. .
.
Fig
.
of
; ;
of
b -
.
.
a
of
of
;
a -
.
.
Erythrorhipsalis pilocarpa with terminal ellongations but actually the joints arise from Fig 157 areole Longi joint showing the coalesced areoles Tip Fig 158 159 near the top and not terminally joint apical showing that none Fig 160 Top view tudinal section the the areoles are strictly terminal joint after removing the areole bristles
mmmmmmmmmmmm an is
,
.
R
, .
by
at be
,
is ) a
or (
of
II ) ,
in
to
(
of
I.
of
.
a
on
.
)
.
,
in
as
R
of
in
as
(
.
,
.
)
tip ,
.
.
.
a
of
an is
.
of
)
(
( . A A
.
.
;
.
of
a
of
of
as
-
.
R
.,
,
so
tip
,
-R
a
of
.
. .
Fig 161 hipsalis cereuscula young growing branch Tip FIG 162 mature long shoot showing the large areole which pseudowhorl forming One the young branches arrow and adjacent branch have grown past the areole wool scale has grown above the walled border the areole FIG 163 late stage de velopment the acrotonic pseudowhorl The branches are clearly unequal a b -
of
of
a
be
,
his
T
10 .
all
of
.
is
and they bear small scales and the areoles have few bristles Because they grow rapidly they are very tender After having length the last are reached the maximum oles remain clustered near the stem but
VOKAL
.
re
the
penduli
and until the last rank order single prolongation Fig 84e These jointed branches arise from the long shoots Erythrorhipsalis the same manner pilocarpa The long shoots cereuscula ig 161 are formed Erythrohipsalis a
.
of
a
is
it
Therefore all the branches are not equal and sometimes only the uppermost one will branch again typical growth This form
also
basitonically flora etc are characterized formed long shoots which attain different lengths depending upon the age and culture the specimen These long shoots will called branches Order These bear the top whorl short shoots branches Order which turn branch form poor whorl shorter joints Order III
.
.
in
,
a
in
a
true whorl
lated
of
of bya -up
of
.
or
4
.
3
,
to
tip
not
Rhipsalis cereuscula like Rhipsalis Saglionis and
shrub species
of
so
tip
By
.
is
to
)
.
.
of
(
of
.
Some specimens have not only one but more joints arising from the areoles These joints arise what appears be whorl but because the areoles are spiral downward from the stem formed
, . ,
terminally
Cactus
a
to
.tip big
stops and some areoles remain near the apex Thus new joints arise near the apex but not
terete Rhipsalides which were examined . species How the different habit each study derived may illustrated by extremely acrotonical species the Coral
in
of
a
is
a
.
)
. (F
(
),
of
tip
laterally
displaced
the branch ig spine hairs
(F
(Fig . 157 ) . The covered by tuft longitudinal 158 and Fig section 159 shows that the areoles very become close each other near the stem The last internodes are short that the areoles become clustered around the top the joint The when seen Fig 160 shows that the from above permost areoles appear merge into single central areole This caused growth the joint and the formation longitudinal growth the new areoles internodes lengthen and the areoles become Finally growth equally distant the joint slightly
.
CEREEAE
mm PHYLOGENY
OF THE HABIT OF THE CACTACEAE
at
.
,
of
of
all
of
so
).
.
(
its
is
tip
a
so
tip
grows unlike Erythrorhipsalis the stem that wall formed around the stem around the This bears scales Fig border 162 The areoles near the top also bear some spine hairs that the every long shoot and top joints the in
.
In
.
is
.
tip
branches bear tufts hairs sunk into the cylindrical top The wall around the Rhipsalis puni quite high some species ceodiscus the terminal whorl first develops inside and then breaks through of
of )
3
(
full length
the top
at
attaining
after
usually
the areoles
Fig
Immediately some
.
) in
.
as
(
.
is
in
is
,
in
II
branches Order 163 The Ery young shoots are unequal size throrhipsalis and when they reach maturity the difference size still marked The always slightly longer uppermost joint
form
branches more than the others The joints immediately Order form joints five
six
until
the
and
be
long shoot typical tuft
of
damaged
comes
a
of
.
,
so
on
Order and orders have been formed Very often the top
or
of
III ,
II
of
.
and
In
of
areoles
a
of
of
.
in
,
In
-a
4
.
of
is
of
.
in
of
the
.
ex
an
.
is
. :
cm
Average
.
5 14 . 18 6 0 5
:
.
15 12 . . 0 0
.
12 13 .5 5
.5 0
5 .2
.
نر
.
II IV
VII
2 8
.
6 10 . 16 . 5 4 2
VI
Maximum
2 5
بر
III
Minimum
1 .0
ارر
. 5.1 .
رد
Average
5 0
14
Order
.
,
VI
و
Maximum 18
II III IV
Minimum
no
,
:
had the following measurements length
length mm
Order
to
1
the other orders are more constant Rhipsalis cereuscula
Other Rhipsalides have same type branching but their habit quite different because the joints are longer As ample plant related Rhipsalis clavata
,
of on
of
in
,
of
Order the same branching
cereiform
,
a
,
.
I. ,
of
which are Order Although the long shoots length differ considerably specimen the joints and angle
These results show clearly that the length diminishes the higher orders Also the extreme acrotony these plants shown by the large number Rhipsalis orders Saglionis the first and last joints are also striking because the last ones are ngled
a
II
.
of
,
in
of
.
of
isis
top
lost
.
if
example
the long branching shoot are capable The upper most again develop joints Order which branch the usual manner but the joints formed toward the base the long shoot are longer Near the base intermediate branches are formed which branch like long shoot but are much shorter Finally just above the roots true long shoots arise
whole top
at
long
ilong the
Rhipsalis cereuscula whic Tip long shoot 164 had been injured before the branch attained full length large areole was formed Because the lower areoles hav grown into short shoots The upper right branch has been jured the top and has branched from its uppermost areole FIG
.
distin a
of
for
If , alred
).
, ,
.
whorl
all
lost ,
shoot
is
guished
can be ig 164 the top severely injured for example and
size
(F
in
equal
no
II,
of
of
.
is
hairs not developed this case the up permost areoles the shoot form branches Order but these joints are very un
اه
CEREEAE
this species the branches of Order I were about the same length as those of Order II , In
that there were no long shoots . Another species , which may have been Rhipsalis capilliformis according to the so
il
lustrations from Britton and Rose but dif fering from their descriptions , had these measurements : length
Order
Minimum
,
cm
. Average
Maximum
19. 0
II III
.
.
50
IV
.
3. 4 2. 3
VI It has scula
6 .8
8 5
.
7.5
5 7
5. 9
54
4 2
V
14 . 0
14 . 0
13 . 5
..
3 .2
2. 4
been shown that in Rhipsalis
the number of joints
in
each
cereu pseudo
whorl generally decreases in the higher orders , with the result that the higher orders have more than two parts and very only often one branch exists . It has also been shown that these joints do not arise terminally but do arise from areoles slightly displaced from the top . Also the joints of the whorls become shorter in the higher orders and this causes flowers to appear from lateral areoles rather than from the terminal tufts of the last joints . Usually in rarely
III ,
of
is
II
II all
In
.
, or
.
,
is
.
of
from
few
-
3
of
in
II,
.
.
These
more re
parted has but often only two the top the long branches become very of
”
"
of
to
,
no
is
of
.
long and may even form aerial roots branching great The angle also importance yet author has used this character With regard the plants just branching angle discussed the term refers the angle enclosed by the branches pseudowhorl This angle has been found very constant for each species and
al
,
,
at
.
shoot
even
sulcata
.
of
few
of
a
., .
Order
branches arise
, .
III
an
.
cm
II of 5 . 6
a
a
Rhipsalis
whorls
become
Rhipsalis
stricted
joints
to
.
.
.
cm
II
40
.
cm
15
to
6
of of
to
A
is
in
the apex
this specimen formed flowers but usually they arose laterally
two whorls Branching may
even
be
of
in
all
,
is
of .
,
branches again whereas the others usually remain unbranched branching especially The reduction species with thick branches common specimen similar Rhipsalis Shaferi had Only long shoots length over joints three Order were formed and long Order these were was developed and the joints had average length but only one branch complete whorl and Order had single joint had The strong branches
the flowers arise near the top these Rhipsalis teres only Order branches developed and the flowers arise from areoles even from the long shoots Some times species which only form two orders joints have very dense pseudowhorls of
no
is
In
).
.
or
(
Hariota
a
of
clarified long Fig 165 genus there are this short shoots because the joints are every about the same length except that slightly longer and whorl one branch Thus the habit
Neves- Armondii has branches of Order but sometimes only Order formed and
of
whorl
to
that the
.
so
longer than the others asymmetrical
is
species with long joints in the higher orders one of the branches of each whorl becomes
sali A branch of Hariota showing the unequal de cornioides velopment of each pseudowhorl .
FIG . 165.
PHYLOGENY
OF THE HABIT
OF THE CACTACEAE
.
of
.
.
R
of
to
of
III
angle of branching is only 20 to 30 degrees , which is very small . Although the angle of branching is not absolutely constant, it varies only slightly in each species . An indication of the con sistency and use of the angle of branching may be illustrated as follows . A Rhipsalis spp . obtained from the Botanical Institue of the University of Halle by Prof. Dr. Troll had two very different types of branches . This plant ( called Halle No. 1 ) attained a length of 1. 8 m . From the top of the long branches of Order I a woolly branching of long joints of Orders II and occurred branching The angle these shoots was capilliformis wide and very similar But from one the long shoots some lateral
of
.
,
(
,
it
of
al .
is
in
of
of
of
"
cereuscula
of
is
same
.
plant
in
of
the
habit
.
Pseudowhorl "
spp
.
Rhipsalis
of
.
R
In
,
. .
167
Fig
.
habit
the result the same develop any terete Rhipsalis Also the Rhipsalis mesembryanthemoides
in
is
,
of
.
a
this habit
ment
as
40
the
and that the latter Thus the first species has slender bunch branches and the latter has spreading branches dissimilis the grees
to
branches arose but the branching the angle definitely indicated this interesting genus The habit the joints Hariota has ready been discussed and was shown that
-
the two types importance
de - to
is
120
degrees
of
50
of
,
similar longitudinal symmetries but branching first species the angle
in
.
.
clavata and
R
.
R
cies
of
The two spe capilliformis have very
causes the dissimilar habits
It of
some branching
).
,
species are the same the angle
entirely unlike the rest the plant but the angle branching was the same Fig 166 168 was not possible discover how
of
the longitudinal symmetries
of
though
.
of
.
A
.
.
Rhipsalis spp 166 normal pseudowhorl Halle Photograph from the Botanical Institute
.
Fig
of
to
of
in
ap branches arose which were similar pearance Rhipsalis the long shoots Saglionis and also had the same type short jointed pseudowhorls These branches were
the abnormal
for
CEREEAE
may be clarified
.
of
,
of R .
at
of
of
R
.
is
. if
all
of
do
as
R
now . As shown cereuscula areoles the long shoot may top damaged form short shoots the The mesembryanthemoides short shoots have the same type tuft hairs the stem apex those cereuscula and
is
.
R
the only method
early
,
.
R
in
cereuscula flower formation from the development these as
as
not terminated
of
of
,
of
.
Aowers are also formed from the top How longitudinal growth ever because the long shoots mesembryanthemoides
of
.
is
by short shoots from the lateral branches
of
,
of
to
to
of
,
In
Ery the same manner the habit throrhipsalis has evolved the habit Epiphyllanthus and Rhipsalidopsis and from Epiphyllopsis here the habit Schlum
des
.
The complete plant
.
Rhipsalis spp
.
168 .
.
FIG
of a
an
.
etc
of
-
,
it
to in
of
.
,
bergera and Zygocactus The latter genus differs from the habit the Phyllorhipsali that has large areole and tuft simi des Rhipsalis cereuscula lar those and Hariota The large areole indicates that Zygocactus had the flat jointed habit origin other than from the Phyllorhipsali
LITERATURE ,
CITED
. E . Cactus and Succulent Journal , TTT , pp . 99. 1931-32 . 2. . Private communication . 3 . Berger , Alwin . Die Entwicklung slinien der Kakteen , Jena. 1926. 4 . Bilhuber , E . Beitr . zur Organstellung in Pflantzen reich , Bot . Archiv . 35: 305. 1938 . 5 . Brückner , E . Betr . achtungen und Beobachtungen zur Systematik der Melocacteen , 1. Anton
G
Backeberg
,
Kakteenkunde
25 ff . 1942.
, F . Der Formenkries der Strombocacti , Cactaceae , Jahrbach . d. Deutsch . Kakt. Ges. 1936 . ............... Rost und Trockenfaule der Kakteen , Kakteen -jahrbuch der Deutschen Kakt . Ges ., pp . 62. 1936. ....... ..... Areolendiagramme , in Beitr . z. Sukkulentenkunde und -pflege , pp . 52- 56. 1940.
6 . Buxbaum
.
.
bei
1942
Portulaceacceae
und
1931
119
Bonn
Biologia Generalis
353 376
.
,
Nektarien und Bienenbesuch bei Opuntia monacantha
-
.
-
:
308 327
.
Zeitschr
.
.
, . , E
11 .
Daumann
.
Osterr Bot
Cactaceae
80
............... Rhipsalidenstudien , Cactaceae , Jahrb . d. Deutsch . Kakt. Ges . 10. Chorinsky , F . Vergl . morphologische Untersuchungen der Haargebilde
VI :
7.
.
.
Bd
.
von Hanstein
1 -
herausgegeben
2
Bot Abhandl
,
Die Pflanzenstacheln
.
.
, C .
Delbrouk
,
12
.
1930
United
Boundary
States and Mexican
Survey
,
the Boundary
,
.
.
, , . A H
Washing
.
86
-
Flora
49
Kakteen
79 :
. F .
zur Kenntnis der Morphologie und Biologie
Beitrage
,
W
der
.
.
,
.
Cactaceae
1858
Ganong
15
Mutations theorie of
de
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585 602
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