Morphology of Cacti [I]

Citation preview

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

.

.

,

.

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15

Mutations theorie of

de

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207

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bei der Succulenten

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der Keimpflanze

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Deutsche Kakt

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Monatsschrift

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der Dornenales Wasserkondersatoren

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105 109

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53

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embryologie vegetale

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Reiche Naturwiss Wochenschrift Rudolph Beitr zur Kenntnis

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innovation

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150 163

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1931

gique

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Opuntien

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recapitulation

Panaschieate

. d .

Notizen

,

Histologische

I.

Linsbauer

et

1937

-

1

Afd

Zur Bedeutung

Knoche

Comparative Morphology

The

Cactus and Succulent Journal

lungsstadien

Entwick

ersten

. . .

32

585 602

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Verbanderung

der

Engler Prantl Pfanzen familien Zur Entwicklungsgeschichte

Kaufmann

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. 12 :

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60 . , K, . 12 . . , B . 33 , , : . . . K . N in 38 La . . . W . . . : .

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24 . 23

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25

.

26

.

27

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Karwinsky

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Beitr

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Schilderungen

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Contributions the embryos and seedlings Georgesku Beitrage

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1894

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Kakt Ges

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Schweiz

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,

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first

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Pfanzenfamilien

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Die Nat

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Cactaceae

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156 205

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6a

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In

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. W

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Nachtrag

Cactus and Succulent a

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Tiegel

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eine seltsame Wuchsform

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103 and 127

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Wiss Bot 343 391 1904 Morphologische studienan Samlingen

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... .. ... ... . .... .

über

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... ... .. ... ... . , .. Arten

Monatsschr

Untersuchungen

die

... .. ... .. , .... . f. A . .

Kakteenstacheln

Weisse Jahrb

128

Sukkulentenkunde

. .f

Zeitschr

1

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1907

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von

,

Funktion

und

PA

Bau

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Weingart

44 .

43 . . 42

Vol

1925 1926

Morphologio

d

Vergl

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Kakteen

Die

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horto botanico herbarioque regio Physikalischen classe der Könige

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