You are on page 1of 8

Ontogeny and Comparative Anatomy of Thorns of Hawaiian Lobeliaceae

Author(s): Sherwin Carlquist


Source: American Journal of Botany, Vol. 49, No. 4 (Apr., 1962), pp. 413-419
Published by: Botanical Society of America
Stable URL: http://www.jstor.org/stable/2439083 .
Accessed: 13/07/2013 16:56
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.

Botanical Society of America is collaborating with JSTOR to digitize, preserve and extend access to American
Journal of Botany.

http://www.jstor.org

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

ONTOGENY AND COMPARATIVE ANATOMY OF THORNS OF


HAWAIIAN LOBELIACEAE1 ,2
SHERWIN CARLQUIST
ClaremontGraduateSchool,Claremont,California
A B S T R A C T
CARLQUIST, SHERWIN.

(ClaremontGraduateSchool,Claremont,Calif.) Ontogenyand com-

parative anatomy of thorns of Hawaiian Lobeliaceae. Amer. Jour. Bot. 49(4): 413-419. Illus.
1962.-Species of Rollandia and Cyanea (sections Uenuinae and Palmaeformes), endemic Hawaiian genera of Lobeliaceae, are unique in the familyin possessing thorns and thorn-likestructures on leaves, and in some cases, on stems and flowers.These thorns always originate in con-

junctionwith a unicellular,non-glandular
trichomewhichterminatesthe thorn.Ontogenetic
studies show that divisions leading to the formationof the thorn occur in the ground meristem
as soon as the trichomeis differentiated.Periclinal divisions predominate at first,but anticlinal

and diagonalonesare also presentat all stages.Thicksecondarywallsareformed


on thetrichome
and other epidermal cells near the thorn tip. Periderm formson old thorns of stems. Vascular
tissue and laticifersare absent in thorns.Thorns in Cyanea and Rollandia seem best interpretedas
snecializations within these -aenera.

IN LOBELIACEAE,
thorns on leaves, stems or largethornand a simpleunicellulartrichomemay
flowersare presentin only2 genera,the Hawaiian be found.Delbrouck (1875) admits.alltheseinterendemicsCyanea and R?ollandia(Wimmer,1956). grades as "Stacheln." For purposesof simplicity,
The writerstudiedthornsof these species as one "thorn," as used in Lobeliaceae, refersto a triphase of an anatomicalinvestigationof Hawaiian chome raised upon a conical base formedfrom
Lobeliaceae, the anatomy of whichhas not been groundmeristem.
studied previously. In addition, a survey of
Thorns of Lobeliaceae are limited to some
literatureon spines and thorns(e.g., Delbrouck, species of Rollandia and species of 2 sectionsof
1875; Schnee, 1939) reveals a remarkablelack of the genus Cyanea. Several species of Rollandia
anatomical studieson thornsof various kinds. A bave no thorns,althoughsome (R. calycina)have
numberofpapersdeal withthornsor spineswhich small ones. Thorns are especially prominentin
are modifiedleaves, leaf-portions,
or branch-tips. Rollandia St. Johnii,but they are comparablein
The thorns described in the present paper are size onlyto the smallerones in the thornyspecies
distinctive in being, basically, unicellular tri- of Cyanea. Species of Clyaneawhich bear thorns
chomes raised on a conical mound of tissue de- belong to the sectionsPalmaeformesand Genurived fromthe groundmeristemof stems,leaves, inae, as recognizedby Hillebrand (1888), Rock
or flowers.Delbrouck (1875) described thorns (1919) and Wimmer(1956). The most prominent
whichresemblethose of Lobeliaceae in a number thorns occur in species of Genuinae such as
of Boraginaceae,such as Echium vulgare.He also C. ferox,C. Marksii, and C. nolimetangere,
alshows that leaf teeth of Pandanus gramineus, though in some of the Palmaeformes,such as
Agave americana, and Lomatophyllummacrum C. aculeatiflora
and C. hamatiflora,
theyare almost
possess a terminaltrichomeearly in their on- as conspicuous. In C. Marksii and C. nolitogeny.The term"thorn" does not have a precise metangere
thornsmay be a centimeteror morein
botanical usage. The term "spine" seems to be length.For a detailedsurveyofthorndistribution
applied mostly to portionsof leaves or to spe- in Cyanea,see Rock (1919) or Wimmer(1956).
cialized leaves. "Thorn" as used in this paper has
MATERIALS
AND METHODS-Materials of Loa topographical,not a morphologicalsignificance. beliaceae were collectedin the fieldand preserved
The problem of definingthorns and thorn-like in formalin-acetic-alcohol.Voucher specimens
structuresin Lobeliaceae is complicatedby the were also prepared. Specimensdocumentingthe
fact that every intergradebetween a relatively presentstudyare as follows(herbariumabbreviations accordingto Lanjouw and Stafleu, 1959):
Cyanea aculeatiflora
Rock, Carlquist 571 (RSA);
1 ReceivedforpublicationOctober14, 1961.
This investigation
was aidedby a grantfromClaremont C. atra Hillebr.,Carlquist569 (RSA); C. Marksii
Rock, Carlquist 593 (RSA); C. tritomantha
A.
University
College.
2
Withouttheassistanceof a numberofindividuals,
the Gray, Carlquist 601 (RSA); Rollandia calycina
materialsfor this study could not have been collected. G. Don, Carlquist 575 (RSA); R. St. Johnii
Sincereappreciationis expressedto Mr. Edwin Bonsey, Hosaka, Carlquist H2 (UC). Additional species
of Makawao, Maui; Mr. Norman Carlson, of Kona, could have been studied,but sectionsof thornsin
Hawaii; Dr. Harold St. John,of Honolulu,Oahu; and
Mrs.Ella StephensofVolcano,Hawaii. The writerwishes the above reveal the same basic structureand
observationson thornsin otherspecies
also to thankDr. JosephF. Rock,authority
on Hawaiian superficial
Lobeliaceae,forhis interestin thisstudy.
suggestedno differences
fromthis pattern.

413

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

----___
..
,....

.:

__- .1..
.

.
i..
I..
j.
i.
.

i1:
II

. -.
:X
.:...

...:

....

.!

. ..

: .:

-_...

..
. :. . ... .
% ..
. . . . ..;

..

. '.
..

'"

.' '.N!,
.
:

N7

.-..:.:
:!
..: -...."..

.....
...
A.
. . ..c ...
.:.....

.... ..
.
.. .

. ..

..

I..

...

::

ill

!!,:t

.
.
11

m
-- .

..
I

__--

------- ___-_____-__._-__-__-____

- ----- ---

---

_'--- -

---

..

.::..:.:..:.:
N.
. :..::
...
:.F::..,:.. ":..
.. ... .

:
..:
AI
" :.1.

...

----

-----

----

iV.
IN.i

--_

-__
-

_-

-_

----

i..

--

...

:
.U..:.
.:

..

..

.i

..

..

....
.

..

...

:L
..

..

i
i

..:

:.
..
.

..

1;

. ...

...

. ...
...

..!

:::
.. ...
.

.I
.:

..

.:i!`-

. .. - :!
.%'.4::ii

..
.

..
.

. .-.g
.
.
..

"
-::,..v:
..

..
.:i

:
:;

:.:

!.
::.
:..:;
.
!-

.!..::

. :1
ii

`i 1:
.i
..:

i.
'Vi
I
II
...I

],
.1:

...

fll-...
-

-i

..__

;
II
:

11
I,
.

..X.'I
.

:..

I...;
.

- - -_

.:

_- ----

.......:.........:....
.. ...

....::_:...::%:::::-:::.::..._

%.:: ':.. .. .

.'%.

..::..N.
.. ..I........
..:.N.
. :
.: .:.....
.. . ..:.:
..-:::..

.1

"

IL...

..

: ..
.. .

..:

..

... .. . ..I..
...
: :
:
:
:::::: IN.
. .....
.. .. ..... .... . ...
.... .. . . ...
:::::.-

. .. . ..
:.:::::.:
.: :::'.-:::--:-:::'::::
:::l::X
..:: ..ii-::::..::%.::!%.::..:::..:::!.::.
:: :. ::.; ':.:.:: -.:.-l:X....'V4 ... X4... :::...:4..::....:.:..::4..:::.:%::4.1":::: .::.-.::...'::.: :..::

..
...
.. ... .:::. .' N....Z'::.. .. ':'..:.....:.:..
...:.:
- . .. :::.......
. .......
.....::::,::.
: -...:
..N
:::..NN.::: ..... ...
. :.:...
.........
.. ... ...
... ':: ..
... .....
... ... . :' ... .
.. .. .. .... ..:..... . ... .%
- ': ..
..::.":: '::

:
. . . ... .......... ..
... ... .... ... ...... .. .. . ..... ....... :::jjj
L
.:. ::: .:
.....I
. . . ... .. ... ... :X-s-:::
... .... ...... :: .....m.. .. .. .. .. ..... ..... I..,
,.. :: :' .. .
f.
:."::."
..
... ':: N. ::- ... :-::%:--::x
:'.: .. ., "..' - '.. ,
.::.:.:: "': '::-::X::!!::...::: s
I ::
.
..:s .....
.......%X....N. .:...::: :::--%:::--:::......
...
-.'::
1.::,::;::?"
:.%- .:. .: -:::--':'--:X: ..........
.. .:.. .: '... -:: -.....
:::...:: -::...::: ..:: :%
::
. . . .::::
.,:,..,::. :::..::: :..-::::
:. .. . - . . : .
... W%::::4..:
-...:...::
:.--%:-.::%-...-:: ..::...:: -::- :4m.%:4:mm'::
X:
.. : . "
.
.. .
. .. :' : N.... ::-..... :-N:.-.:::
:..-.-N:.--::.--:: ......... N .::....:: ........ W..'44
. . . ..: .:'..::.':: ...-...
:::...:::.:.....:..-:.:%%-..--r.%..::
.: .:.I..:. 4". .'--':X--::X-':X-`::
.
.!
.. . . .:. ...
:
.. ... ... .......
':' .::. :::.::. .. ... ....:X:
.. ... ".." ..'
:.:..:.:..
::' ..
..
.. N.:..:-::.-::.
... - .. .. ,...:::: :..'N ... :N'.: ':' :4-1.:..
.. ..
N. .::: N.
- .
:..::::.::::
.
N .......... :'
: .: :.:.,.::::. -N.---:::--N'
.;: : ..:..... N....... . . ..
. .. -... i..N.- ....... :....:. .....:...
.
.:
....
:..:.: :....
.:::%: ..-:N:-.
.........
..
" .. ': . . ': . ..... :.:4.:: . .. ..
... ..:
.
.
X.-NN
.:i:::!
.
.:::.:::
.
... . .
.:... --.:..:V
::: .:.: N... :...:.
; .
... N ': -:::.::.::: :::%.::.
.
.. . .:: X%':
% '... .... ...
...
.
..: ..:. - - .
.%%
.. - %- ... .
.:%...
..
% v...%.......... ....:;::---""'...
-.:, -,:.%:.!.
.
.
-::-.:
.. .
.. ::%
:'
...1%.
1: .. :::."..........
.....:....%
%-...%-:.....
.
.
.
'
::.
::::.
..:.'N
. ... ........ ...% . ...
. ... . .. :.:.: ...... . .. . ..
. .. . .
.
....
..
.1 ::: N .. - ::: :
. ..
.: . . . .,
. .. .. ......
i.
::: :: .::::4:
.I
. . . :
..
. .. .. . .'.
I
. - . .::..
, .
.. ...' .: .. N .. . .-- %-V
: .. - .I . ...
......
.: ... V .: - : -:- .:.. . ..
.
. N.
:'. - -:::`.
.....
..
. . . ......... % N..
.
..
-- ' . :.". . .
.. .... -..%..:..%.. ..... :. ..
......-::.
--...!
;....

;. .. .. . .. . .
:: . ............ ..
. .. . ..:.:
.:.. . . . ..: .. .
. .:
.:,...--...
.

.
` : : :'. :,.: ,:":. .:. . : . .
. . ... .. .....
.
:' - .. - .
"' N N
.
e..F
% ..: :::..: ".:..:-::%.::..4
.:'-.:1:
..
.
.. .
. .. . .. - ': :
. . .
!: ..
% .....
:.: ::':':.:::'l::::'::.':::::::.
- .:.% - .. - .... .
.:.. ..:.
.
i. ".. ,::.:: ......:
.. ...: ..:: ..:...:. .:.- .. ... - ....... .
.. .
.... .: .. ..
...
... ':'. ":' .:.:...
:: ...N..::.
.:: ':: N %..
- : .. ...
.::.
:.:
..
.i:
. .::. : " .:. .:. N.
I .. .. : ::- , -:. ':.: :: ,
.
; .
:... :' ..: "
.. . .: .... .: . ... ....
.
. . ' ..
:
... .
:.
. ..
.. ....
.::. - .:- ..
.. .. . ....:'
.. ..
- N
.!F
% .
. . . .. ..
.
..: . 1. ... .. ....
.
: .... .... ...
. .
.:
....:
t.:::.
. .. ...
:'
. ...

..:..::...... ..". -

:.

-':: '

....

..

:'
.. .- . ...... .. .....:.
.....:
.. ...
.. ... . .....
... ....
.. .
.. ... % .. N
.. ..... . .. ... .:. .
..
.
.
.......
" ,:: ..: :...
.
.:':::!:::,..,"'---!i
!4 .....-.:::
:: ... .:.."..: .... .

. .-::..::::
...... ....
":...:--::...::--.
. ..:........1...
. ....
:::..::
-::.
..:...
%.
:::.
.%%....
..N....
...:...:.....
N...
.......:..
:....
....
.. ..........
......
.1:'"..
.:.N
..:..
-:::--::
.::.
..
.. N...
":.....,
:::
:::
:::..:::.
:::%.::.
-.:....::.."::
:...........
%
....: .-.: ! : ::.....-::.:.::
....
.. .:..
....... . .. :.": .::. .-N '::
IN:
.. : . : :.. ...'
.1.4
.. . -::...-:: ::: N .:: :::- ::: .::. N.. .... .:

.;
.
.. .. ..::. ...: .... %. -N:%. -:.:.. .. . ......
:.:: ..
::::
...:.--:.:
.... : ......
.
....: .. ....
.... :.. :."... .%..
.... ....
... ..M.::.' . .. .. . . -. .:
. . ..
. .. .. .. .... ....
%. ..:
... ....
.... ....
... ..:%
...... .......- ..
. :. :'
. .:: .:.:,::: ,:.:: :4 ... ... ... ....
::-:-':. ... : :... ".......%
...:..
%. ::": ... ................. ..I..,.:::..:::
.... ... ........
.:
::: 4 N : ::.::'-X::
..:.. ..::. N ...: :: ..........
; : :::.. , :, :
..
-.. :l.:.:::::l-.':'::' .. :; :::
. ::::.
%:::.,:::..:::
.. . . :!.1
.:. .::: .::: :::: :::%:::.'::: :::, -.::::..:: ::
::i....'...l:
.. :... " ...:4.:1:4.:...:..%:4:.%::-.%-::..:::..:
- N. .:::. :::.
- :.. :' V..... :--'X..'::'-'::
.:: ::::7:: :::.::: -X::.:::'.::: -::: :.:.::i!!!!!I!f:!:i
N.
... .... ... . ....I... ....
..:
....:. ...- N.........
.... :%.-::!--.::-%--:i
-:::--N:!-:-:-.:
-........ :..::::i
.: .. ..... .... ::':-:::'-':X-'::: :::::::- X'% .. i:
: .: ' N ..:::--:: :::-.:::N:l:-:: - .:.. :%
. !! !!;ii
... :... ... -:::.%:::.::: .::. -::: N.
.-.'::
::: ':: -:::.:::,::.-.::.N. ...
N.......
......... ..
.::: ::::
" :::: .: ... ..- ...
: .:::,:::.
':: :::%::. ..:.. :. :u
.., .xN..:::%-.:
.. .:: ...
. . .. .... .. . ...
N. :: ....... ',.:.... . %.,.. ::: :::7i::.":..::'...'::--::
-X'--X' N.:
. . ... ..: :::.-:::::::- :::...:. :::. % % i!i
ll.
. . .:.. : .::- :..-.-.:- : ... :..-.:.. - ... - ..
.
.: ..... :-'XX%:::-.:".:".::
" ' '.
. .
::::::: ..... i:: ...
.:::: ..
.. ..: .: .:..- .:::.-:::7::::;.:.':::.
..... '.
.M .::. %%-... .... ..%.
..
...
.-....
::: ...:7.:
:...::::.:.:.
: ..: .'...:.:
. - ...' - :
.
.. .. -X.. .... ::: - :-::--::...
! ...... ::-":- ...':.":.
.. ... ....... .............
... ....N.:.:

...h,......:.....;;;i!
:.:. :......N--::'...::..N:
!i.It;i

N.,
..:::

..

:!fll:!-Hi
: . ...
.
........
:
: ..:: ..:::..
: :::.......N.-!!F;::
. ....,;.:: ;."T :

. .::
L:

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


.... .

::z :
:N

..

..... . .:4.
.
..

.:,:.

.m

...

.. :. N'l:X: I
. .:,:.
"I..
.;

. .. . .... ... ...I....


' %.';.. ,:::
.::i:i.'..i:.i:..Iii.'.ii:.z.

.
..%.::..:

. .'.::,
.m
:i!::;::l::i:!!i:ip:

X
'
f.

..

4:

....

....
.;g.

'_-.
I1..N
,::.
-.
. .1
.::.
..

...1 %.

- :'
i::it-

.
X. ':
..'
.:.

k
.
.:- '"-:- .. ".

.......
....."
...
......
. .:.

..

:..v :'.

::

.1

m
.

..

-.,.

m....::i!:!;!;:;:A, ."

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

.
:-

:: '

4%

:.
I -1N. ..
m4
- .%
:.
.
.:,

.1..4.%.....N
.."I
. x.

k'

:4
m
:.
:

....';"

: ...

....

..'.
.
IK:.l.
.
:I
.. -U.

'.

:Z-:

':

..'
.1
.

:'

!:

.' ..
..

.'.

.
: ,:. .:,
.:.
.4. .% .:,...
. .

b .... .

.4

-" :V _:i
: : -". :, .

.,....... ,,!-Y

:..'A
Y. . -. .
--,;6
.
k. ... .
.
.
1 .. :4
":
.
.' '.

.1 .*
,'

:1 ' .0k'M %
.. .....
, ".
.-... .. .1
....
N . . .... ..'
:'... ... ..
.:K.-A: .--_
"'*
, ....
.: %,
..':..
..
.%.-'. ' :-,

,..

.i

:
.

.
" .):
..
::..
'..
.
:' ':
.
,
%" `
. ..,
. :...:
..

.:,::

...
..

..

' : %
%Olk
..
.
I
..::
'
. .
"
4
" i
..
.
: .: .

:'I

m
.t.

:.

. ... 'r .-'.


: ..
-

-.*
..

.%.
,

I..

:, ,
"'
-::
.. . lkl 1.. :..l.
. .::,
::.
..'
n
- .' ..'
", .'

':.1
il...
.1:

'i-:'

:.:.: . :: 1'm
:
.:. . .. .....
.m
..
:: : ::%
.:
: N "'%
:.:. .:.:..
. 1.
. . : ..
. %."
...
-,xt
. .,
...
.X.A I .
S .
-.
.7 4
:
':
I'll
I
'L, : . ... .m

.4

, %s.:
'

.'...x
1.
...
:

..'

..: . %..,.
:-.'i I
.M.
'
.::,:);:.
" ': ::
I
A
% t:,
.... ".
.. .

.i...

t,

'"' .. '
..
1... '
*m_I j

`4

.li
.

::
:'

4k,x.6
:: 44;::
i.
....
:'

.::.

..4.W. Z

N-.4
..

. Im.

..-..

: .. '- :'I........... N."i

.
.::'.1

. '. x

:.,::"r..

..

:.-_
:..........
"'::

.%
:
.

'. .

. :: :
:-': A :x

..... ....:::....

1:
:'N'M.
.

"
zi.1

":
.: .
.
f

. : !
:,'A.
:
. -%.
m'-Iil.!

.:

.
-iI

..

II

.1

'
-.: ,.

..
.....:::
.2"..' ...... ...
:. .: :: :: : ,..
N ::
:: ":.:%
-- -.
:..:: ..
...%.4::
V
. !:. i?::;::;5 :.::::::
....I....%--..:..
... :::
...: ..
.... ..::, ,:..';
N "::--::.-:::-:':': :: ...
:::!: Hi':-H :Hi:
::4
:ii KH. .
........ :X--':: .:.
...
i% :::. ::.,:::;. ' ...
..:.:: :..
.:: .:.....:
..::. :: :..:.:..::. ... . ..... %:X
.:
.......: 4 -:-:: :.::.:. .:: :
.:: :.:::...
': -!. :' .. :::. ...- ':' ...
%::X-'::'
:'m
...
.. : ....:... . ':'.
.11 ;!!?
.. -:-- ..
: : .. .., ....
:::.%::-.:
.
..
..
.
..
....
..
.:
::: .m.
:
, .': ...... ':: :" .. ::: ':: ... N. .::; .:;-"';.
::
I
....
.
. . :::: :; .. - .... .. .
L ,'ji-i
.: "::%:: ... :..:.::%!!::: :.".:. :-.%:'::-::%:H::
,--::
'. .,:,:::.,:.. :'... ::::%:::
. .. ... .::;i:f
.'
:,::: :,::.- .:::'
. .. ... .. .. . .:
' " .":
.. . .. :: . :.::::
..W
..
::.4: '?-;
:
.: "::
:1 . - ... :' ":...' %,
' % ::.:,: .. .::::.:.': ..::s
:." . - .
::
.. ..:
. .: . _.. .
,
...:
':: N '..:. 1 p...
.. . .. :. :::X:::
:
.
I.
M. ...
,.
::
. -...V
::s.
.. : . .. .. ... . - '...... : : ':
"
..
...... ......%
.,::. . . .::7
: . .. 1 .. :
'..... .. ..'..........::. . . . '.:7:-.7::::.;.:
..
11.!:V.l
..
...
...
4...:
.
.......
..
..
. . .. .. ... ..
:
. .
X
:.:.%:7 N. ':X
. . ..... . ... . ......4
.........
... ...
....
. ...... .. .. : : 1;
...

..

. .:.:.....:
............
: - ..i

.
:X._;'S
,
:

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

..
...:

6.

- - .,---

..

. . - - ..';
.. .. . -. .:. ......
:":'
... .. .......
......;
: :'::"N"
N.
...
-.%.:.
-...:..
....
...":-I..
......
......
...:...
..:.....
..::t
...
N....
.,. ..
...
..I....
' . ..
...
--N.!-::::
-........
....
':.:...
..":::
.
....
.-..-..
. .%...
.. .:..:.
.......N.
....
:..
..:::-%::
....:1
N
-.:..::N.
.....
. .:
... ::--::.
.... .::i
-::: = .::- ...:.- .. .
.
:
...
- ... ... ...I .1.1

..
.. ... .. ..N
1::
.."

F--2
_"-'.'-',"'M :.
. ......
..i!j:!
....
..........
V...
%..

...

-_

&
'IM 4

.
..' .
.
:'.. ... :' .......".
.::. ':: .,:::
..
" -- ': !:::!:i!:
.. .. ............
.. .....
... ... mm
.. mm
.. m.. .. .
%, . " . . ..
:. :.:
-.: ..... :.. .:%:..
..
..%:::..... .:::...:..:....
.: :... - -..
:
-..:...:::
.:..::: ::: N.
..
. :.:. : i:
. ::.....:::::::;::
:::- ::...::: :::':::.-N:: .: .. .....
... ... ....:..
:
..
.:
. ....:..... .. N :N..:..::::::: :::-::-N-':.. .: .. ... ...
. . :..... ..:.::
... . .:M .N '.
.
.... . .......... :::
.
N .::: ' :::-.::'.:::-..:
:' .
:'
' " " "' ..:.
"'
"
... .:. .::. ..: -... .. .. .. .
. '.I..
.. ... .:. ..N.. :::.
....
.
..... . . .. :--::---:: ':' '' .... - . .
,.:. ., :.:
N -.:. :.: ':' -::. N
.. ....-:.
... .. ...' !:!!! :j
.:. .N...... ::%....
. .. . ..
.. %. :::: ... .. ..
. ...
.
.
%
...:.. ..:.
... ... . ..%........
..
.
.. ...
.. . ...-..
- ..
.: --V..
.. . ..
': - " :" -.
:'. ,:::.. :':.-.::
. . .. ::
:..
....
:
:':
..:
.
..
.
..
': ..
...:
- %sis...
: ::.
.
. .. .
.
. .
.. .. . .
....
....
..' ...I
7 ..
.' S'...
X4u1 .'.-':" "--::;i4....
... :.

. ..
.':'
:
:..
. :
: ..:::,..:..::.
:
:..

.----

:
.. ....
.:-.

,I

:'.

m-.

,:.
.IF

....:
...
.....
.:::
%.::.,...:
. .......':'.':::.:::'.::::.:::'."::.
......
.......
...
....
...
.....
:--::--:-:-:::::::.::....
.....
...:.:...
.......
......
:.:%"
: ;:;:;:. .......
: : ;::os::'m
....:.
: -:."
..:.-:-.."'..:...... .-:--.%::-.._
......
.."
::'.::
::..:-.::.-.::
"..:.
...
::...::...::...:::,.:.:::;
,:::;:::::ii:ii::i.::
.::
.....
. .....
.:..:..
:..:.::::.
:..
-:---:..:::
::%.:::.
.....
......
...
.....'
....::....:....%
.-:..:.
-::.
: ':.:::i:!,::!:;!::::!;.lei.
.

lk,
,..-,.,-...

.N
M
: :::::;:
: ":
....
.::::-::.
::,::::.::
:'-1::..:
-,.::.
.-:..--:.....
:Du:%.
'--....
. ."-:-'.:.:'i:':!
...-....
: ......
.:..;.%::.
-:.::
-::-::..:.:
:::...:.
".... N.::.
....
.....:.
:i;!i!;
is;.''
"."
.:ND
.-I.
"--':::..':'
.%:::
-N
IX:'
.:::..":
.:.
.::
N.
N.
%::..-:...%
....
..: ::::.:.:: ...-:. ...
....''
.:::
.:::
.:::
..:::..::::
.::..:::
... ......
.......
...
-:N..
: .:N
--'::--.':'
.:...
...
..:!
::.:':::.:..:..... ...."-:..-':'-.......::
....:.:
::.:.
'...;
..-::.:::::..::::.,;.::,:;.:.:::..
:..
:s;m::.
.::-.::.:-l:-::ll
r:I #I!,
.. . ... ..'.........

. . .... ......:"::.
%.- --::..
':: ":
N %.
.: - ::: ... .. .. .
....::.

m
.%;
i

. ....%

. - ....

.! . ...
. .....
.. .....
.....
.iiii_'-.-.
-.....:............
. .... ". .. :."

...
..

10

::i'i:n::i::,:::;:fi;::

-.. . .-.... . . . .-

::-...:i;
,N
:%...
:.

- . ..- ::::::.::.::..::

..

::..'

...".N.

-..- . ....:::::..:::,.
..lll.ll...:!77:_::
.1....I.. .... .......'N
.
............1.
....
. :-%-' ... %-':..----. -:.--':--:-'::
... m:1.11.1111...
.:R!..1 ... :::. :%,. :..:.: ...."
::%:::.:t::%'.
.:---'---'M'--!::.-l
..::;-..::.--:!..m,
...-.-..
.
. . .;:
.... :::"::::.:::":::.,::..:::..:::.:::.::.:.:-::::-::::-..:%-:%--.:.-::
... :.....:..: ...... ... ....
-.. '.:.: ......
...s.: . .. ....
. .....:!::::::.
........
............!%.
.:4::: -"::%":::'::% ...
....:...:.. :::'..:: ..........5:::.:-:'41-":::
%:::.:.:::.:... ... :'
... .....
I.., ......
.......:::::! .. ...
.%%
... 4- ... :.-..'::..
::-:X-::::-:::
..::.,:::.:::
.::: ...X....
.!::::.::-%--..:'-:.--..:--.::
.. .. .. ... .. N..
..
.: -.X--..:-:.-.':..:
.... : -':--::--::-:.-.:::-....-..:.:.....:..-:
::.::: ....:---:-:--::.;..::::::,:::..::..:::..:::.:::,::::.,:::!:::....:..:::.....
.. ... .. .. ...,-.....
... ... .. ::...::..::.
..
...1 !.
: :::...
:::
......... ........
... ... ..
.
.. ... .......
...."
.. ::%"
:,..: ...%.:::% "-%":..
i.
-.-.,---.
..:..
.... .. : :- .........!:
.: .. -:- ..
..:''::.,:,ii:::iifi:,
::%
. . . .. ... ...
.. .....
:. ...
.... ... ...
. .. ....... ::.;:::.::.::. . . .. .......
:::. .:
:::.X -... .... ..
.;
:::::-:.:-::::X!!
..
... ::'...:::..:.
... ...
.. ...'I.,
-:::.:::..:
..........
..... .-.::::;:.".:P:':
:. ,:::
..... ....I....
..
:
..... .. .... ....::.::..:.:.::::!!::::..:,:..,::!
:": - .
.I... ...... ... .!
:--::::-f:::..::::.....!::i:,:ii,:iii.'7'iii::fii',;i!:
-x--::::-:
:.
-:::!-::::x-:x:'.'
..'.............."..:::..::.:::..:::--::::.:::c:
..
:l'::P0:...:::'.:......
:::.::::::::::::!:.:,:.::.::.:.::::..:::.:::..:.. ... ... ....
.. .....
: :.!:: .: .. . . .. ...........
... . ...
... ...
....
.,:::.f-1. -.:::::-s:::::.--.
:X '-%.
...
.
..... .. .:::.
:
. ..
..
.... ... N....
::
..... ..
,:...:
-:..::--:.:-l..U'.'l
..
.....
. ....
.:...:..::--:::::.
-.
.
............N
-"
.
..
.... .X....N: ... IN ...
...
... ....
......".......::..,::!::..:%....:....::::
.....Ni!s:..::
..
.. '..-.-.:-4.:-:.:
..
.:::.:::
........ ... .......
... . .. ...'%v4h.:
.. :.--.:.---.:...:.....- :::.:.: ......
-:::::i:::I:ii:i:iT
.:.::
-:::::.:.:::
..:.:.
: .:..
::!s:4....:...'::::'
-.
...
..:
..%.. .. .... . .. ...
:.._'-':'-::":::'-:.... .......
.%.......
:.
........ .::.. ..:..%.: _.::::::-:'::.:::'::%::::.::::.:!
"::: .... .N......:
......
.......:':'..:':..:::.:::
-..!..::::.
.N...M..:..:: .. .:.....:
... ..
.............
... ........
..... :
..
. .....
.. :...
%.'-:-'::::.::::':-."'-."'-'-'-:
...
.:
::..-::--.
..
.."...
.....
....
...".
- - ':'. .:..:: .::: .1..
.:
... :....
.:.::.::.:"::::::
....
"
M.N %.. ::.::::..
...:
....:..
-.:..--::%-:%.--.
.......:
.
_..::::::::
. .::- ..:..,..
...
-:- .: ... :::.::%':::%":::"::':
.:...,.:--;i -:;:!6;:
..
-::.:...::
...... N.
.. :.....:.:.
.. . - ... ':..-:....::-....
....:..
... ... ...':.:'::.".'::.:...
.... ..... ::: ": ...
.... ... ...... .:-...... :W. ! :: -'.
- -::': %
....
".!-.:'.
-:.....
..:
...
'4::.:::::
...
IN....,
...
....
%----%._:.:.::
..
...
..
:..
-:::::.::::
....
.--..%%::
N
...
..4.. ':.:::.:':::!.::::.':::
...:.
..::%:: . ....'
.... ... ... ...
.. ... .. ... ...
....-. .: .::% ..:";
.'-.-..::-.::..::
.... .:
'::::::-:.':::-:.:::-..
:. - ..... ...... :.:...-:
...w...:
.. ::::.:.%..::..
.... N... - "'
': X: ...::...::.
..
'::-:..:X:::.':::--.....
... ...
:.% ...... ..,:.::
.. ...
..'.
.. ... ::::-.
...
....
.: ...%......
-.
. ...::::-:::...
F.....".:...-"'::::."':':.
..
-,-- -:::::-.
':...::::.:':
.:::.::
-:::::
-:::::::-'::.."::.:'::::
.. I..
.... ::.::...:..
.: .:.
...:.:::...:
. :
: ::::::.::::.::.::.
. .... .... %X:--.:.%:..--..--s..::.': .. .......l.: .
--:::-::-::
...
...:...
': . .....
....:,:.
-.:
.
.. '::..:..:..:':::.':"::..:
.....'....
.:.
..
.:.. -::. -::I.:.
N..'
':%-':
:
.-:...-::.:.:..
:' %-:-.....
..... .....
:.%-..- .. .. ..:
.:.Z::...::
....
...: :::.-..:::::
-....
..: . . ......Y..:
..
. -...:. .:.
.V.
------:-:-:
: :-- ,--..N
. .... :-:::-:-::."':':
.
: ... ::.'-:x..-x.'': ..
.
. ...:....
.. ::.::-.
:...:::;::.:
.......
,:..:::
.
I..
.::':v:".";:::.
.....
. ...............:
....
. ...........
.. .. ....... .: ....
.....
.---....:.:...:
.......
....':
.. ...... N -.44.
- -'..
%...... ...
. . ..
.......,....
:%,.i
.. ....-::
. .... .... ..c. :...
.
.. .....
.. ..
... .... .:
. ....... %....-...,
....:..
.-M-N..
.-':"::::'7':
-::.:.::.::.
.....
.....:.......:...:.......
:..'.
......
: %.:..:::: .........---%..4.........
..-:..-.:
. ......
..
--... ...:.. . ...
.......I
....
.........
:'.:.;:7:-". ........
%%..:.
..:.:...:
.....
..
.
..
.
::..:::.
:::::::::7.-.-:-.....:X
..
.'f
4-...
.. N -..
......
:4-N-l,
.
.. . .. .......... .. ... .%..... ...:
....
N -::..:::..::-::
'.:-:.::7::
.
. .:. .. .. . ............
-I......
. . .. ..: ...... .%.-:.--. .-:
.: ..: :..::-::
:::::..::::.....
" .......
"
:'
:"
:
-.--::--::-:::.:::: .
-::. 'N .'.:........:.':'.:":::":::'::'::...::.:..:'..:::::.......
.. .. ...... V.- ............:..
:
:::i
..
.:
....
.:...:_
- ::.:.- ..-.:.
. -...
.. .:
: :': ... ::- ::'.:.- ..: ......
N.",- .. .. .
..
. . ..... . ... :.
..... ..w ...: . ...... %...
': ...........x ..... ...
. .:: ::. :.!..:..
:- ::--.:.::..::.
.
.
. :"' .: ,... :': . .....:' ... "-..%..:..
... .....'::.... N.. .
.......::..:: ..
.. ...'.... ......:...%:::
..
- "%:'s.:::.,..
.
.::::: ...:.".::...
N:.:::.:.
. ... .. .. ...
.... :..:..
...
.......::::..:::::.
. ..:..i.....
V
...":.::..-..%...%-.i..:..%.:...........I:.."..::
1:.
...
....
%::?
....:..::..::
%.
.-........:..............
....... .... ....:.: .....
::.::":::":
.. .. :::':::'.:':::::
.. ... : .:.. -.%-.:': ::.::::.:.::
. .. . .....
.%:.:....:...:..%---.%:.....:::::.::.:
.. ... .::.:::.
... ... :.:::-:
".....
.
s %...:......:..:.:.
::
.1.
..."
:'.
:--....
... ..:. .:. .:::
.: ::: . ..... .... ....::..:.
.: -:.":::,::
..
..
...
....
.....I..
........
..........
..:.
;:
.:
.::.::.. ::-:::::
X.: ...%......%...
.. . ... .... .... :j;:;jj
... .....--.:.:z:::.
- .::. ':: - .:..:. ':::::x:::
"::":::
'::
.::.
: .:-':::':::.':::..::-:.:::::::::%::...:: .... .. %. -:..
....
.::: .:::.:.:: ..:: ,% ....: ......:..
4.
.:...
"..::: ...::::::N.::-::: -::::::.::::
.... .... :s:.:..:.. .:....:.-::..'::.
':: N. .:::.:.:::..:
.. .. .....::
:: ::.. :---.--::
.. . ...--:::-.'::-'::::::::::.::
N. N.
...: ..N.: ... ...
%...
':.-:':-:'.:%:...:::-::'-:::-:.
......
....
. :" N.
:::..:::
...
:.:....::...
.
.....
..
....
...
....
..
....N..
'::::-...::'-:
....:
'
... .. ... .:::.::N......
... .. ... ..
" " '
N. N.
-..:: --::...: ... ... ....
........
. -': -...
..... ............ :-, ..... :.-::.-::..:::.l:-.
.".::
.. ..
::.: .:
::: -:::-::
;.:..
::.:%::.::.:.:.%:
":: ."!
'..: .. :.::: ..
"-::. ':: -::...:. .::...::...::
... ...-,.....
':',:......:
. "."'. "'::'-'.':::si':.'.::j-:::
,.:: ..
:::::::::::..::
:..::
:::,::::.:::
I..
:.:::"::::::.:.:::.::.::::.::.;
.. . .......
.- M :X..'% .:: . . :.: ..: : .. ..--..::-....--::.N.
.... .... :::.::'.-.::.-:
.:.:::!.:::!.,:::%...:..::..::...,.:...::
....- :...%:::...::....:.:..:::...:4.-.
.
N .... ... . ...X--':- . . ... .: . .
. .: ........
:... . . .::...:
:
N
.
:!.::: ::::..::....::..::....:...:..
....:::--.::.. ... .. ....
4.
.
. .... .. ....-..% ::%.:
m ::::-::::.:::w
..... .. .. ::--::.-.:.------:
:
%. % .
.
-- .--....
- -.. .:" .. ":-:::
...
-.::-:::.- :: .:.--::.---: ..: . . . . ... :.:.:::..:.
::::: :::. - .. ..::...:: --:::--':::--X:-%X
il!i ... .
:..:...::....:...::.....:....:,....
.:: ..-.-:..-::.-:4
..
:.:::.I
:
.......:..:.:"
N.-...
....:.:.:%..::
':::%::
:.::......:...:
....
...
.::?:i:j.:.
:%..:
X:.-.:-::: .. ......::.
.......
':'. ':". 'X--':::-':
.. ....
::-:-.:::-':'-..:.
.: -::. .:. ..:. .. ...:.
..........:-::-:!-::.--:::--::...--":-':'..': ........ ::::--.:..--:::
.. ... .... --:..%--:..
. .. .: ..
......
%..::.:.. . ::-!:':::!!:::-... :' .........
... ... - .. ...
.... . .......
::.s: .
.
... .: .. ...... :.:....:.::.
.: "-!.:.:::"::-:
N .:: ':: N :::..::
::...:.
..... . !.:::..::.:..:::!:
:--. ... ......
.. ...
...
.. ..... ::: :: -:::::::..::.::
... ........
. ... ...
:. .:: -::...:. .: . .. .. .... 4::---.:::::
: :....::-..:: .::...::.....::.::...
::. .:: .:::. .:: .:...:.
. ..: ...:' .
...
s.
.. ..
..::.....:
... N ---':N-':'---:
. ..' :::--:::-X.:::... .... ........ ...:,::.... .:..:...:..::
... .. ..
...:
-:- ::-:::
:::.::
!....
.: ':: :::::-.::.::-..::..
:. . ..::,..:
.'..
N
: . ......:...:
:...
... . ....... .:.:.::4:
.....
..-::. -:.- :
...... :..::: -::.-:-..:::..:--..::
..
-l::--..::'--:-'::
..:. ---.:: .'-::'
..:. ..
. .:N--...':: ...-.......
... .. N. :':':..:....: :::-.:::'.::.:
.... ::: N N %.:
. . ..
- ':
..::...:.
X':::.::'::::::
i... . ..::...:: -,::. .::..:::.::
....:: ... . ... .. ... .-::
:::.::
::--:::.:::.::
.::...:..:.
.....%.
...N ........
::: .::- -::. ::: -::--::. .::!
;':
N. "'
N N. .::. -":
N. ..:. .... -.:.-.:..--.:.--.- . ..--:::-s:...::::.j:::
..::
.. ....4..4
.:N%
% :::l-::-.:.---.:
........ ::--'!:,:.:.:i...
..:...
N. N. :::.::::.::
.... .. .:...:" - .. - .:: - "N" - %::
.
: .. .......... 4:....' .... .. :.:...:..
N .::: -:: .:. ::' ..,:
;.;..!Z:
.
: :.: . . .
.
......
....
'. ...--N:-:"':: .:....::
': :
. ":::.":--:
. .:: ':: -::: ':: N. .:: ::: %
. .
. .
....
.:'
..
::-- ':' ':: ':: ::: ::: :ii
....
.. " .:- '::--::-::: N ::: -.-- N. -. ..
.....
:::.-::: ::::-.::. .::. N. :-.. .:::-:.:
:'i...
. ... .. ...- ... ..:
.
.
. .N. .. ..... .
. -'N .
" ..
...
:
.
.....
..
....
..
.
.:
..
..
'-:..
.
-.:.
.
:.
:'
,:::1
....... ..":..
. ..::: " . ': - .
.. ..
... N ..: . ':: .:. .:"' ..:....
" -::
'::':'
. --;:'
.....
."::.%.:..
...::N........:
:'. .............
.:: % ..:
... ..
.
. .:
... .. N N - : ':: :::.. .. .1. ." -.....".
::: .:. .
..
..
...
........:
.
.....
.
.. N ..
.
. . . .
. .:.. N ...:.. I .. .: - . . .
. ...
.. :' :' :.. .. ...:
..:..
.. .. .......
....
:
-'
-... ,::i!!::!;..'..,.::..'..!"..L
..
.... ...%
-.
. . .. .. . .......:.:......
- .. ....'-:.:
.: ... .:. N : ..:: ...... ... .
.: ..
..
::..."
: .
. ..
.:':..
.: I...,
-:::::
.: . . . .. ... .. ..':X
. .... N ;N.:' %.- : ...1.1
.. I..
.....:.......:-.
...-':'.
: :'
..
. ..
..
. .:" ::
....
.. : . .. ...
..
.
.'.....
:
. .. ..
7:::7::::
...
..- .... :. .....
..
.
.... . .i.`.
.-...
.. ... .. .
V
...
..
.:N- :::.
.
.
... .
:::
....
. :%.
.. .. . - . % ..
..: ..
... .. .
.:-V:':"::'.
.
...
.
.m
.
.
.-. ... .... .
.
. . . .
. .. . ...: .. .. . ..
...
V.
.
..
.% ...
...
:77:::7;::::::::. ..'.
. . ..:
..
.
-.
...
.. ..: ...,.:.
...
.. ... .:.::: :::.:...:%......
.. M %-NsQ:.--:..
.
.. . :
..
..::......
.
:
.:.::7..
.
:::
-4.
.:
..
..
.
..
.:
.
....
...
..
...
.
..
...
.
: .:. % -:..
.: ....:..
...
..
::.%%
.. ---::-x= 4'-%:X:.
--:::
.
.
:N N ".
.
..: .... ..
... ...
... :..... ... ... .... ...:
.. ....
... . -....:........ - . .- ..%--.....:.
:::-%:::-:'::::
:NMN ':..:
. .. . .. . ..
.......
.
... . ..
. .... :...: ' ---ii::::i
.. .
.-.. N ..
:- ::-'-::: ::
. :..
.:. .: .. .:.. % -- . ... . - :,::,
.... .
:
.. ... .....
.:.. .:. -::..:::--::- .:: .:: ...-.:.
-.......
......
%.: .... ...... : ... -..%
.. .. '.
.
..:. ......: .... ........
........... ..... . . ..
:'. ......
.: .:: ::: .:. .: ..:. .:.. ... .. .-'_'N
iN

7
.

i
i.N!

te.

..

...:

.I:i
.

.
.

.i

__!

..

-'.

...

1i

1..

i4:4

1-1

- i.

:0 I.
.
.1.

.
"I....

!"'.

. iI
. ii

iii
.;---i
Ii
I
.I'.i

i'.

_'___

.
.

.. .
. . .
: . .:.

...

..
M-.
.
'.. '
__::

.: .: . .
: - ..
;v
:
N .?:-::4.:

I.. . .11

:19^

-'

-.
-".',:
.
I

.4:

:."

:.
-

.-:

':

.. .

.:"
-I's
,"i' i -. iz.
11 :1 ..' :.
... V
1,

April, 1962]

CARLQUIST-THORNS

fIllJU?L%L~~~~~~12

IN

LOBELIACEAE

415

1314

Fig. 9-14. Thorndevelopment


in CyaneaMarksii:interpretive
drawingsofthesectionsshownin Fig. 1-6. Brokenlines
indicateprobablerecentdivisions.Note thatthornsformoverstrandsofprocambium.
in Fig.
Dome-shapedformations
9, 11,and 12 are mostlyeminences
associatedwithveins.whereas'lheprojection
in Fig. 14 is an actualyoungthornbase.
All, X370.

Material was sectionedby means of the usual suitable stages forillustration.Cyanea Marksii is
paraffintechniquesand stained with a safranin- more densely covered with spines than other
fastgreencombinationcorresponding
to Northen's species; Fig. 8 shows the appearance of foliar
modification
ofFoster'stannicacid-ferricchloride thornsin this species.
technique(Johansen,1940).
As seen in Fig. 1, 9, formationof a foliarthorn
ANATOMICAL
DESCRIPTIONS-Ontogeny-Cyaneabegins with differentiation
of a protodermalcell
Marksii (Fig. 1-8; 9-14) was selected for illus- as a unicellulartrichome.In species with large
trating ontogenetic studies because: (1) the thorns,such as C. Marksii,divisionsinitiatingthe
numerousthorns,on both surfacesof leaves in spinebase occursimultaneously
withthe originof
this species,are moreeasily studiedfromsections the trichome.Althoughanticlinal divisionsmay
of vegetativeshoot apices than thornson stems be seen at any stage, periclinalones predominate
could be; and (2) this species has many stems, duringearlystages. Numerouspericlinaldivisions
providingmore shoot apices than the palmiform may be seen in Fig. 1-6, as interpretedin Fig.
species. Sectionsof shootapices of the palmiform 9-14. As the mound of tissue underlyingthe trispecies, C. aculeatiflora and C. tritomantha,chome expands, cell divisions occur in various
showed the same features but provided fewer planes, increasingthe volume of the thornbase.
Fig. 1-8. Thorndevelopment
in CyaneaMarksii.-Fig. 1-6. Stagesin expansionoftrichomeand underlying
tissues.
Fig. 1 showsearlydevelopmentof trichome.Shape of trichomes
in Fig. 2-6 is probablyalteredby fixation,
because
maturestages (Fig. 7) are invariablyconical,and someconicalyoungtrichomes
may be found.Activecell divisionin
celllayersis occurring
subepidermal
in thesectionsshownin Fig. 1-6 (see interpretive
drawingsand legend,Fig. 9-14).
-Fig. 7. Trichome-tipped
thornon abaxial surfaceofyoungleaf,suchas thatshownin Fig. 8. Laminabelow.-Fig. 8.
Youngleaf,seenfrombelow.Note largerthornson petiole;thornsare mostlyrestricted
to petioleand zonesalongveins.
Fig. 1-6, X600. Fig. 7, X66. Fig. 8, X4.

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

416

AMERICAN

JOURNAL

OF BOTANY

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

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

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

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

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

.... . ....

. . ... ...

..

...
.... ...

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

... .... ...

.....

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

..

.. ... ... ..

...

.... ... ...

...

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

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

........

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

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

...

..

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

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

..........

. . ...

..

...

..

... ...

... :17

.... .. . ....

..

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

.
.

.. ... ... ...

..
.. ..

..

..

. ...

..

....

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

..

..

..

.
. ..

. ..

.....

...

..

....... ...

. ...

...

.. .

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

.. . ...

....

. . ..... ..

..

... ...

....... ..
..

..

...

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

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

........ ..

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


.

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


. ...

..

...

..

. . .. .. .. ... .. ...
::.:
...........
.. ... ... .. ...

...........

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

April,19621

CARLQUIST-THORNS

Early stages in thorn formationshow divisions


only underneaththe trichome,but divisionsextend to peripheralareas later. A wide thornbase
is not built up at first.As seen in Fig. 7, a long,
spindle-shapedthorn base forms. Widening of
the thorn base takes place only duringthe last
stages of thornmaturation,and is caused by cell
enlargement.Divisions leading to thorn formation continuewell past the stage at which procambium elementsin the lamina have differentiated into protoxylemand protophloem (Fig.
6, 14).
The trichomewhich terminatesa thorn elongates greatlyduringmaturation,and develops a
thick secondarywall, the outer surfaceof which
is striate parallel to the long axis of the cell.
Epidermal cells adjacent to the trichomebase
also develop secondarywalls. This characteristic
wall thickeningaccounts for the stoutness of
thornson the living plant. On herbariumspecimens, epidermalcells with thick walls appear as
a light-colored
cap on the thorn.
Althoughthornson leaves originateabove and
below futurevascular bundles,theirdevelopment
is whollythe 'resultof groundmeristemdivisions,
and does not involvevascular tissue in any way.
Study of largenumbersof sectionspermittedthis
conclusion.Laticifers,whichare abundant in the
vegetativebody of these Lobeliaceae, are absent
in thorns.Significantin this regard is the fact
that in Cyanea and Rollandia, laticifers are
closely associated with phloem, so absence of
vascular tissue in thornsmightbe expectedto be
accompaniedby absence of laticifers.Breakingof
the thorn on a living plant does not result in
latex flowunless the entirethorn is broken off,
thus damaging laticifer-bearing
tissues beneath.
Thus, thornsin Lobeliaceae are unlike the latex
hairs of Cichorieae described by Zander (1896)
and one may answernegativelyWimmer's(1956)
query as to whetherthornsofLobeliaceae contain
a "special sap."
Thornson stemsare longer-livedthan those on
leaves, and undergofurtherdevelopments.Remnantsofthornsmay be seen on stemsofold plants
of palmiformCyaneas such as C. macrostegia
(Fig. 15-17). The stemof a youngplant (Fig. 15)
bears thornswhichare anatomicallythe same as
foliarthornsof C. Marksii. During aging of the
stem, the terminaltrichomeof these thorns is
lost (Fig. 17, right). Thick epidermal cell walls
characterizeold spines. Only a small increase in
size, comparedwith spines on a young stem,can
be noticed. Eventually, the epidermisis broken
in places, eitherby expansionof the spine or by

IN LOBELIACEAE

417

injury by decay-causingmicroorganismsor by
epiphytes.With damage of the epidermis,formation of a peridermoccurs (Fig. 17). This periderm
developmentis only moderatelyactive, and although several layers of phellem may be developed, peridermaction never producesnotable
corkyexerescences.In very old thorns(Fig. 16),
the peridermis partlylost,presumablyby decay.
At no time do thornsdevelop internalsclerenchyma. Firmness of the old thorns can be attributedonly to succulenceof the groundtissue
parenchyma.In thisrespect,thornsofLobeliaceae
are quite unlike the thornsand spines of thornscrub and desertplants. Such thornsand spines
are composed mostly of sclerenchyma(Schnee,
1939).
Comparativedata-Studies were undertakento
determinewhat, if any, is the range in thorn
anatomyamongHawaiian Lobeliaceae. The most
obvious variation is one of size, for thornsmay
grade from relatively large structuressuch as
those describedabove to trichomesonly slightly
raisedupon an eminence.In all species examined,
unicellular trichomesnot accompanied by any
evidence of thorn formationare present (viz,
Fig. 21).
Unicellular,non-glandulartrichomeshave been
reportedin a number of Lobeliaceae (Metcalfe
and Chalk, 1950; Wimmer,1956). Such trichomes
were the only sort observed in Hawaiian Lobeliaceae of the presentstudy.Close groupingsof
unicellular trichomes,which might be termed
tufted hairs, were occasionally observed in
Rollandia calycina.
Thorn-likestructuresmay be presenton the
exteriorsurfaceof calyxand corolla,as in Cyanea
aculeatiflora(Fig. 18, 19). These "thorns" are
oftenvery softand delicate, and are characteristically long and narrow. Moreover, the unicellulartrichomewhichterminatessuch thornsis
thin-walled,as are the epidermal cells which
surroundit.
Structureswhich are not thorn-like,but are
mound-likeor knobby in shape occur on leaves
and stemsof Cyanea atra. Sectionsrevealed that
these structures(Fig. 20) appear to result from
relatively late periderm action beneath unicellular trichomes.Whether this peridermis a
normal phenomenonor resultsfrominjury (e.g.,
fractureof the trichome) could not be determined.However,emergencesof the natureof the
one figuredin Fig. 20 mightbe termedcorkwarts
rather than thorns. True cork warts (Fig. 21)
were observed in Hawaiian Lobeliaceae on the
midribof a leaf of Rollandia calycina.Periderm

Fig. 15-19.ThornsofCyaneaspp.-Fig. 15-17.Cyaneamacrostegia.-Fig.15. Stemofyoungplant,showingprominent


spines.-Fig. 16. Bark of old plant,showingdegeneratethorns(light-colored
spots).-Fig. 17. Median sectionof a
thorn,fromlongisection
ofbark ofan old stem.Terminaltrichome
base is stillpresent,at right;periderm
actionevident
belowsurface.-Fig. 18-19. Cyaneaaculeatiflora.-Fig.18. Flower,showingspinynatureofcalyxand corolla.-Fig. 19.
Mediansectionofsmallerthornfromcorollalobe; fromlongisection
ofcorolla.Fig. 15, X3; Fig. 16, X2; Fig. 17, X40;
Fig. 18, X1; Fig. 19, X81.

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

418

AMERICAN

JUORNAL

0t_|(iC

.~~~~~~~~~~~.
......'.!

Z^i.s,

',~~~~~~~~~~~~~~~~
"'

-"

L';

....
_~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
1

=34

.. .......

[o.4

OF BOTANY

.o : ..... ...

~~~~~~~~.

'.S.._...~~~~~~~~~~~~~~~~~
,'

..

.. . ..:.. ..

....,.

ie

.... .. .:... .. ....


.. .......

...:

..;...

w~~~~~~~~~~~~~~~~~~..
. :...::.
..f

i.....::..: ::........ .

. . .... ...

..

. ... ... ......i


..

.. .... ..

...

:.:

S::.

::;:.:
..........
......
...
w
:.
:..:.
..
.. ..

..

-: ;

. w

. .

..

:~~~~~~~~~~~~

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

..:|.:... ..:::.:

Fig. 20-23.-Fig. 20. Cyanea atra. Transection of leaf, showing cork wart formationat base of trichome remnant
of lea.f miArib. Uniserinat trichomes
Fi.
-1 Conrkwart. frnmtfransaection
(below). Fig.- 91-99. Rdlln."i. caycna

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

April,1962]

CARLQUIST-THORNS

formedas a responseto injury by decay or attack by fungior microorganisms


was observedin
various of the Lobeliaceae studied. Cork-wart
formationas a resultof injuryto a trichomebase
would not be unexpected.
In the genus Rollandia, thorns are relatively
small (Fig. 22, 23). Cell divisionswhichresultin
the formationof the eminencesupon which trichomesare borne appear to occur relativelylate
in leaf ontogeny.As in thornsof Cyanea, epidermal cells surroundingthe trichomedevelop thick
walls. In Rollandia calycina, grouping of trichomes upon an emergence was occasionally
observed. Only single trichomeswere seen on
thornsof Rollandia St. Johnii. Thorns are confinedto the abaxial surfacesof leaves in Rollandia
St. Johnii,and are presentmostlyon the abaxial
surface,only sparinglyon the adaxial surface,of
leaves of Rollandia calycina.
Significanceofthorns-The functionwhichmay
be served by thornsof Cyanea and Rollandia is
not readily apparent. Althoughinsect predators
are presumablypresent(especiallyin the formof
larvae) in the Hawaiian rain forest,arguments
could be advanced that these thornswould be
relativelyineffective
againstdepredationby these
animals. Land shells (land snails) are indigenous
to the Hawaiian Islands, and could conceivably
be deterred by these thorns. However, the
Hawaiian Islands are lackingin otherherbivorous
animals whichwould attack leaves and stems of
Lobeliaceae. The fact that thornsof Lobeliaceae
are succulentand lacking in latex suggeststhat
thornswould be a betterfoodsourceforpredators
than leaves or stems themselves,which contain
large numbersof laticifers.Darwinian reasoning
would suggest,however,that lobeliaceousthorns
are not withoutfunction,for they have become
highlydevelopedin severalspecies,and would be
expected to have some selective advantage.
Interestingly,seedlings of some species (e.g.,
are providedwith more and
Cyanea tritomantha)
largerthornsthanthe adult plants.

419

IN LOBELIACEAE

From a phyleticstandpoint,the distributionof


spinesin certainspecies of Rollandia and in some
species of each of 2 sections of Cyanea is of
interest. Thorns are absent in the remaining
Hawaiian Lobeliaceae. Absence of thorns elsewherein Lobeliaceae, and the specializedcharacter of the species in which they occur suggest
that presence of thorns is most logically interpreted as a specialization.From the anatomical
standpoint,such an interpretationis supported
by the fact that thornsrepresentderivatives,or
elaborations,of trichomeformations.The fact
that thorns occur independentlyin Rollandia
(whichis probablycloselyrelatedto Cyanea) and
Cyanea suggests that this group of Hawaiian
endemicshas a tendencytowardthornformation
whichhas been exploitedin a remarkableway by
some species groups,less prominentlyby others.
This assumptionis considerablyeasierto envision
than a propositionthat thornswere primitively
present and have been lost in the majority of
Hawaiian Lobeliaceae.
LITERATURE

CITED

C. 1875. Die Pflanzen-Stacheln.Bot.


Abhandl.2: 1-119.
HILLEBRAND, W. 1888. Flora of the Hawaiian Islands.
Privatelypublished.London.
JOHANSEN, D. A. 1940. Plant microtechnique.
McGraw
Hill. New York.
LANJOUW, J., AND F. STAFLEU.
1959. Index herbariorum.Part I. The herbariaofthe world.Ed. 4. InternationalBureauforPlant Taxonomy,Utrecht.
ofthe
METCALFE, C. R., AND L. CHALK. 1950. Anatomy
dicotyledons.
ClarendonPress,Oxford.
ROCK,J.F. 1919. A monographic
studyoftheHawaiian
species of the tribe Lobelioideae,familyCampanulaceae. Mem.BerniceP. BishopMus. 7(2): vii + 394.
SCHNEE, L. 1939. Ranken und Dornen. In K. Linsbauer,[ed.],HandbuchderPflanzenanatomie
9: 1-24.
WIMMER, F. E. 1956. Campanulaceae-Lobelioideae.
In
DELBROUCK,

A. Engler, L. Diels, H. Stubbe & K. Noack, [eds.],


Das Pflanzenreich 106: 1-260.
ZANDER, R. 1896. Die Milchsafthaareder Cichoriaceen.

Bibl. Bot. 37: 1-44.

This content downloaded from 86.143.155.98 on Sat, 13 Jul 2013 16:56:44 PM


All use subject to JSTOR Terms and Conditions

You might also like