Some contributions to the life cycle of <italic>Papilio zelicaon</italic> Lucas with special reference to the hardening of the wings

PDF

Download

Open document

Flip pages

Download a page range

Download transcript

Copy asset link

Request this asset

Transcript (if available)

Content SOME CONTRIBUTIONS TO THE LIFE CYCLE OF FAPILIO
ZELICAQN LUCAS WITH SPECIAL REFERENCE
TO THE HARDENING OF THE WINGS
A T h e s is
P r e s e n t e d t o th e
F a c u l t y of th e D epartm ent of Zoology
U n i v e r s i t y of S o u th e rn C a l i f o r n i a
I n P a r t i a l F u l f i l l m e n t
o f th e R eq u irem en ts f o r th e D egree
M a ste r o f A rts
t)y
B ern ard E. Brown
August 1950
UMI Number: EP67188
All rights reserved
INFORMATION TO ALL USERS
The quality of this reproduction is dependent upon the quality of the copy submitted.
In the unlikely event that the author did not send a complete manuscript
and there are missing pages, these will be noted. Also, if material had to be removed,
a note will indicate the deletion.
Dlssgftaiton PyfclisWng
UMI EP67188
Published by ProQuest LLC (2014). Copyright in the Dissertation held by the Author.
Microform Edition © ProQuest LLC.
All rights reserved. This work is protected against
unauthorized copying under Title 17, United States Code
ProQuest LLC.
789 East Eisenhower Parkway
P.Q. Box 1346
Ann Arbor, Ml 48106 - 1346
% /f/ fi?7 7
This thesisj written by
Mr. B ernard Eldon Brown
under the guidance of h..%3... Faculty Committee,
and approved by all its members, has been
presented to and accepted by the Council on
Graduate Study and Research in partial fulfill
ment of the requirements for the degree of
M aster o f A rts
Date.{
Faculty Committee
Chairman
TABLE OF CONTENTS
PAGE
INTRODUCTION 1
FEEDING 2
COLOR OF CHRYSALIS 5
WING HARDENING
H i s t o r y 9
E x p e rim e n ta l
M a t e r i a l 11
Method 13
E x p erim e n ts on th e d r y in g of th e w ings 14
E xperim ent on o x i d a t i v e r e a c t i o n 15
E x perim ent on s e v e re d wing 15
E x p erim ent on k i l l i n g th e b u t t e r f l y 16
E xperim ent w i t h th e a n e s t h e s i a 16
D is c u s s io n 17
C o n c lu sio n 18
SUMMARY 19
TABLES 21
LITERATURE CITED 25
INTRODUCTION
T his i s a s tu d y t o expand th e knowledge c o n c e rn in g
F a p i l i o z e l i c a o n L u ca s, and w i t h s p e c i a l r e f e r e n c e t o th e
h a rd e n in g o f the w ings upon emergence from th e c h r y s a l i s .
The s tu d y i s d i v i d e d i n t o t h r e e p a r t s . The f i r s t p a r t
c o n c e rn in g th e fe e d in g h a b i t s o f F a p i l i o z e l i c a o n . and th e
second p a r t r e l a t i n g t o th e c o l o r o f th e c h r y s a l i s a re s e c
t i o n s w hich d e a l s p e c i f i c a l l y w i t h t h i s one s p e c i e s . T his
i n f o r m a ti o n i s o f i n t e r e s t and u se to f u r t h e r work on t h i s
s p e c i e s , b u t p ro b a b ly h a s no g e n e r a l v a l i d i t y i n th e o r d e r
L e p id o p te r a as a w h o le .
The t h i r d s e c t i o n d e a l s w i t h th e h a rd e n in g of th e
w ings upon em ergence. The f i n d i n g s p r e s e n t e d h e re p ro b a b ly
a r e t r u e f o r a l l L e p id o p te r a n s in g e n e r a l .
FEEDING
F a p i l i o z e l i c a o n Lucas i s c a l l e d th e f e n n e l sw allo w
t a i l b e c a u s e th e c a t e r p i l l a r i s found t y p i c a l l y on f e n n e l .
T h is i s an exam ple of food a d a p t a t i o n s i n c e f e n n e l i s a
r e c e n t i n t r o d u c t i o n from E u ro p e . P a r i s h (1920) in h i s p a p e r
on "im m ig rant P l a n t s i n S o u th e rn C a l i f o r n i a " co u ld n o t say
j u s t when f e n n e l was i n t r o d u c e d , b u t i t was s t i l l " r a r e " i n
San B e rn a rd in o County in 1890, and o n ly " c a s u a l" i n Los
A ngeles C ounty i n 1898. I t seems q u i t e l i k e l y t h a t th e p l a n t
was i n tr o d u c e d as a g a rd e n e sc a p e e i n th e p e r io d betw een
1880 t o 1900.
Comstock (1921) i n h i s work on b u t t e r f l i e s o f C a l i f o r n i a
s t a t e s t h a t th e A nise S w a llo w ta il (P . z e l i c a o n ) i s common
th r o u g h o u t th e s t a t e and t h a t i t s f a v o r i t e fo o d i s w ild a n i s e
( Carura k e l l o g g i G ra y ). T h is p l a n t i s n o t found s o u th o f S a n ta
C la r a C ounty. The p o s s i b i l i t y a r i s e s t h a t b e f o r e th e i n t r o
d u c t i o n o f f e n n e l th e b u t t e r f l y was n o t in t h i s a r e a . There
i s no e a s y way to ch eck on t h i s #
I n t h i s s tu d y th e c a t e r p i l l a r i n th e f i e l d was found
o n ly on f e n n e l (P oeniculum v u lg a r e H i l l ) . However i t was
fo u n d t h a t th e y w i l l e a t c a r r o t , c e l e r y , p o iso n hem lock and
two o t h e r u m b e l l s : Qenanthe and L e p t o t a e n a , and C i tr u s
( o r a n g e ).
E s s i g (1926: 634) n o t e s t h a t th e y w i l l e a t w ild
c a r r o t , w ild p a r s l e y , w a te r hem lock and dog f e n n e l . He a l s o
3
s t a t e s th e y a r e found on C i t r u s ( e s p e c i a l l y o r a n g e , b u t a re
n o t p e s t s ) . I n r a i s i n g th e c a t e r p i l l a r s some d i f f i c u l t y was
found i n t r y i n g t o g e t them t o l i v e on o ra n g e . They u s u a l l y
d i e , however one went i n t o th e c h r y s a l i s a f t e r an orange d i e t ,
and a s m a ll second s t a g e l a r v a a t e a l l th e r i n d o f f of an
o ran g e s t i l l g r e e n . A p p a re n tly some i n d i v i d u a l s can and w i l l
l i v e on C i t r u s .
T e s ts t r y i n g t o g e t th e c a t e r p i l l a r s t o f e e d on p o p l a r
and l i l a c were f a i l u r e s . O ther s p e c i e s in th e genus F a p i l i o
w i l l fe e d on th e p l a n t s a t l e a s t i n e m e r g e n c ie s . C a t e r p i l l a r s
i n t h i s s tu d y s t a r v e d on t h e s e p l a n t s w ith no s i g n of e v e r
h a v in g e a t e n any of th e p l a n t s .
T h is i n f o r m a ti o n s u p p o r t s a c o n t e n t i o n t h a t t h i s
s p e c i e s w i l l ( i n c a p t i v i t y a t l e a s t ) e a t a lm o st any p l a n t i n
th e U m b e llif e r a e f a m i l y , and some i n d i v i d u a l s w i l l e a t from
a t l e a s t one s p e c i e s i n th e C i t r u s g e n u s , (T able I ) . Most
o f th e c a t e r p i l l a r s (b u t n o t a l l ) w hich were f e d upon C i tr u s
and p o is o n hem lock d i e d . One m ight i n f e r from t h i s t h a t t h e s e
two p l a n t s a re n o t alw ays s u i t a b l e as a foo d p l a n t f o r th e s e
c a t e r p i l l a r s .
I n t h e w ild s t a t e th e U m b e llife r a e p l a n t s ch o sen by
th e fem ale p ro b a b ly depend upon o t h e r f a c t o r s i n a d d i t i o n
to th e s u i t a b i l i t y o f th e p l a n t as fo o d . Such f a c t o r s a s
l o c a t i o n and e c o l o g i c a l n ic h e u n d o u b te d ly i n f l u e n c e th e
fem ale w hich l a y s h e r eggs on a p l a n t w hich f o r h e r i s n o t a
4
food p l a n t . The f a c t o r s in d u c in g th e fem ale t o o v i p o s i t on
s p e c i a l p l a n t s would be i n t e r e s t i n g i n th e m s e lv e s .
I n th e e a r l y s p r i n g and l a t e f a l l most o f th e c a t e r
p i l l a r s a re found on young p l a n t s o r f r e s h g ro w th . D u rin g
th e se a s o n when the p l a n t s were d r y e r and more m ature most
o f th e c a t e r p i l l a r s were found on th e stem o f th e flo w e r
c l u s t e r s . A lth o u g h even th e n i f young g ro w th was a v a i l a b l e
c a t e r p i l l a r s were found upon i t .
A marked d e c r e a s e i n th e c a t e r p i l l a r s found i n th e
f i e l d was o b se rv e d d u r in g J u l y . T h is c o u ld be c o r r e l a t e d w ith
o t h e r f a c t o r s , b u t i t i s most p ro b a b ly due t o a sm a ll g n a t
s i z e d w asp. T h is wasp l a y s i t s eggs upon v e ry young c a t e r
p i l l a r s . D u ring th e second l a r v a l s ta g e th e wasp l a r v a
em erges from th e c a t e r p i l l a r , k i l l i n g i t . The wasp would form
a cocoon and i n a b o u t two weeks emerge as an a d u lt #
COLOR OF CHRYSALIS
W hatever th e f o o d , and i n t h i s s tu d y th e c a t e r p i l l a r s
w ere found o n ly on f e n n e l , th e egg i s l a i d on th e food p l a n t
o f th e c a t e r p i l l a r , w hich i s n o t a food p l a n t f o r th e a d u lt #
In th e c ase o f f e n n e l , p l a n t s ch osen a r e u s u a l l y younger o n e s ,
i s o l a t e d , b u t n e a r a l a r g e p a tc h o f fe n n e l* Plowed f i e l d s
t h a t have b ro k e n s ta n d s o f f e n n e l , r o a d s i d e s and r a i l r o a d ways
p r e s e n t good p r o s p e c t i v e l o c a t i o n s f o r th e c a t e r p i l l a r s . A
d e n se s ta n d o f f e n n e l i s u s u a l l y c o m p le te ly b a r r e n of l a r v a e .
The r e a s o n b e h in d t h i s would be i n t e r e s t i n g t o know, s in c e
i f s c e n t i s in v o lv e d one would e x p e c t t o f i n d a d en se s ta n d
as a t t r a c t i v e as an i s o l a t e d p l a n t , e s p e c i a l l y s in c e i n most
d e n se s ta n d s th e i n d i v i d u a l p l a n t s a re u s u a l l y q u i t e a c c e s
s i b l e t o t h e b u t t e r f l y .
The e g g , l a i d on t h e s e p l a n t s , h a tc h e s i n f o u r d a y s .
The c a t e r p i l l a r e m e rg es, e a t s th e egg c a s e , th e n s t a r t s
f e e d in g on th e p l a n t . T here f o llo w f o u r c a t e r p i l l a r s t a g e s .
The f i r s t s ta g e i s a s m a ll ( 0 .5 cm.) b l a c k c a t e r p i l l a r w ith
a w h ite g i r d l e . T h is i s fo llo w e d by a l a r g e r s t a g e (3 /4 cm.)
a l s o b l a c k and w h i t e . The t h i r d s ta g e s t i l l h as th e w h ite
g i r d l e b u t adds r e d d i s h d o ts and w h ite s p o ts o v e r th e f e e t .
The f i n a l c a t e r p i l l a r s t a g e i s a l a r g e (4 cm.) handsome
g re e n an im al w ith b l a c k and r e d o r y e llo w b an d s on i t . T his
l a r g e c a t e r p i l l a r f a t t e n s up and th e n i n n a tu r e l e a v e s th e
fo o d p l a n t to se e k a s p o t t o become th e pupa. F or th e c h r y s -
6
a l l s a web mat i s made on some p l a n t o r s u r f a c e t o hook th e
c r e m a s te r I n t o when i t i s d e v e lo p e d . A l i f e - l i n e i s th e n
form ed around th e th o r a x and th e c a t e r p i l l a r s e t t l e s down and
becomes th e c h r y s a l i s , t h i s form ing u n d e r th e s k i n . The o ld
s k i n i s th e n s h e d , and th e c h r y s a l i s h a r d e n s . The p u p a l
changes th e n t a k e p l a c e .
I t i s i n t e r e s t i n g t o n o te t h a t t h e r e a r e two d i f f e r e n t
c o lo r e d c h r y s a l i d s : g r e e n and brow n. These two sh a d es may
v a r y somewhat b u t a r e q u i t e d i s t i n c t and r a t h e r good camou
f l a g e . I f brow n, th e c h r y s a l i s re s e m b le s th e dead p e t i o l e s
of f e n n e l , and i f g r e e n , f i t in w e l l w i t h th e g re e n f o l i a g e .
I t f i r s t a p p e a re d t h a t t h e r e i s a te n d e n c y to m atch
th e b a c k g ro u n d , t h a t i s , brown c h r y s a l i s on brow n, g r e e n on
g r e e n . Of th e c h r y s a l i d s used i n t h i s s tu d y m ost were o f
n e c e s s i t y on g r e e n f e n n e l ( a l t h o u g h i n each c ase brown p a p e r
back g ro u n d s w ere a l s o a v a i l a b l e ) . A r e c o r d k e p t o f th e number
o f c h r y s a l i d s on v a r i o u s b a ck g ro u n d s showed no c o n s i s t e n t
c o r r e l a t i o n , namely o f 79 c h r y s a l i d s on a g r e e n backgrou nd
54 were g r e e n and 25 were brow n. On a brown backgroun d 5
were g r e e n and 4 were brow n. The te n d e n c y to m atch is. a p p a r
e n t l y a m a t t e r of c h a n c e , a lth o u g h m a tc h in g th e b ackg ro und
co u ld p o s s i b l y have s u r v i v a l v a l u e , (T able I I ) .
I n th e c h r y s a l i d c o l o r a t i o n how ever, t h e r e was n o t i c e d
a te n d e n c y f o r th e p e rc e n ta g e o f g r e e n ones t o become more
p r e v a l e n t as summer came o n , (T ab le I I ) . T his m ig h t be b e -
7
c a u se o f some s e a s o n a l f a c t o r , such as t e m p e r a t u r e , in d u c e s
th e c o l o r o f th e c h r y s a l i d s in c e th e g re e n -b ro w n r a t i o i n
c re a s e d from 2 :1 t o a b o u t 4 :1 and f i n a l l y a l l g r e e n .
Of th e v a r i o u s s e a s o n a l f a c t o r s te m p e r a tu re would
seem most o b v io u s . C h i l l i n g a n im a ls a b o u t t o e n t e r th e c h r y s
a l i s a p p a r e n t l y had no e f f e c t . A l l of th e J u l y f i r s t l o t
w ere c h i l l e d and a l l t u r n e d o u t g r e e n . P ou r o t h e r c h r y s a l i d s
o f v a r i o u s age were a l s o c h i l l e d w i t h no c o l o r change e l i c i t e d *
The n e x t e x p l a n a t i o n t h a t came t o mind s u g g e s te d t h a t
th e p a t t e r n m ight be s e t by c h i l l i n g th e l a r v a e a t v a r io u s
tim e s d u r in g t h e i r g ro w th . Two l o t s of f i v e l a r v a e were
p la c e d i n s i m i l a r j a r s and g i v e n s i m i l a r t r e a tm e n t e x c e p t
t h a t one j a r was c h i l l e d o v e r n ig h t i n a r e f r i g e r a t o r on f i v e
d i f f e r e n t o c c a s i o n s . The r e s u l t was t h a t in b o th j a r s th e
e n t i r e l o t was g r e e n i n c o l o r .
I t would be d e s i r a b l e t o be a b le to c o n t r o l th e r a t i o
o f brown to g r e e n c o lo r e d c h r y s a l i d s , so as t o i n s u r e t h a t
th e b u l k of them be g r e e n . I t would t h e r e f o r e be u s e f u l t o
know what f a c t o r o r f a c t o r s c o n t r o l c h r y s a l i d s t o be brow n,
s i n c e t h e s e emerge a t u n p r e d i c t a b l e t i m e s .
The c o l o r o f th e c h r y s a l i s h as a d e f i n i t e r e l a t i o n to
th e tim e of e m e rg en ce, (T ab le I I I ) . I n g e n e r a l th e g re e n ones
emerge i n a b o u t two w e e k s • The brown ones rem ain dorm ant f o r
an u n p r e d i c t a b l e l e n g t h o f t im e .
Of 34 c h r y s a l i d s k e p t o v e r w i n t e r 31 were brown.
8
V a rio u s o t h e r l o t s r e c o r d e d showed t h a t th e g r e e n ones w ere
u s u a l l y b u t n o t alw ays emerged w i t h i n two w eek s, w hereas th e
brown ones were u s u a l l y s t i l l d o rm a n t, (T able I I I ) .
WING HARDENING
H i s t o r y
The l i t e r a t u r e has n o th in g t o o f f e r t h a t i s d i r e c t l y
r e l e v a n t t o t h i s s u b j e c t ; however t h e r e a re s e v e r a l p a p e rs
on L e p id o p te r a w in g s, th e f i n d i n g s o f w hich w i l l be re v ie w e d
t o p r e s e n t th e s e t t i n g f o r th e problem o f wing h a r d e n i n g .
I n th e developm ent o f th e L e p id o p te r a n wing M ercer
(1896) s t a t e s t h a t i t i s formed i n th e v e ry f i r s t l a r v a l
s ta g e as an i n v a g i n a t i o n of th e h y p o d e rm is. U n t i l th e f i f t h
l a r v a l s t a g e i t m e re ly grows l a r g e r w i t h i n th e body c a v i ty *
By th e f i f t h l a r v a l s ta g e th e wing becomes d e f i n e d . The
hypoderm is becomes c o v e re d w i t h c u t i c l e (d e v e lo p e d from th e
o u t e r hypoderm al l a y e r ) . U n d e rn e a th th e hypoderm is p r o l i f
e r a t e s a l a y e r o f hypoderm al c e l l s c a l l e d th e basem ent
m em brane. At t h i s tim e th e b a s a l membrane cem ents th e wing
as a s i n g l e l a y e r . T h is i s done by means o f c e r t a i n c e l l s ,
th e hypoderm al p i l l a r s . When th e wing i s exp an ded, th e
basem ent membrane l a y e r s s l i d e o v e r one a n o th e r and a llo w
th e b lo o d to e n t e r , b u t t h e s e fu s e d p i l l a r s p re v e n t th e
wing from b lo w in g up l i k e a b a l l o o n .
At em erg en ce, H ering (1 9 2 6 :1 1 ) s t a t e s t h a t t h e r e i s
a pumping o f b lo o d and a i r i n t o th e w in g . The pumping o f
b lo o d i s e a s i l y o b se rv e d b u t i t i s d o u b t f u l as to how a i r
c o u ld be pumped i n t o th e w ing a lt h o u g h as th e t r a c h e a e
ex p an d , a i r would n a t u r a l l y flo w i n t o them .
10
S n o d g rass (1935:215) i n h i s " P r i n c i p l e s o f I n s e c t
M orphology" s t a t e s m e re ly t h a t th e w ings a r e expanded by
b lo o d and t h a t a f t e r h a rd e n in g b lo o d c i r c u l a t i o n i s r e s t r i c t e d
t o th e b a s a l p o r t i o n o f th e w ings o r a t l e a s t w ings once
h a rd e n e d no l o n g e r b l e e d i f th e t i p s a re c u t o f f .
C on cernin g th e a c t u a l cause o f h a rd e n in g t h e r e i s
some c o n tr o v e r s y as t o j u s t what h a p p e n s. Cuignon (1936)
s t a t e s t h a t th e p o st-n y rap h al developm ent o f th e wing i s
n e i t h e r an u n f o l d in g n o r an u n p l a i t i n g . He s t a t e s t h a t th e
p o st-n y m p h al dev elo pm ent c a n n o t ta k e p la c e i f ;
1. The w ings c a n n o t be a g i t a t e d .
2 . P u l s a t i o n o rg an s o f th e th o r a x a r e n o t i n t a c t .
3. T here i s no a c c u m u la tio n o f b lo o d i n th e
p e n d an t w in g s .
4 . T here i s no p o s s i b i l i t y f o r a i r t o e n t e r by th e
• m e so th o ra x s t i g m a t a .
A ll of t h e s e p o i n ts a re d o u b t l e s s l y sound; however
c o n c e rn in g p o in t t h r e e , M offat (1900) s t a t e s t h a t one can
c u t o f f th e t i p s o f th e w ings so t h a t p r e s s u r e f o r c e s f l u i d
t o e sc a p e and th e w ings s t i l l r e a c h norm al s i z e .
C o n tin u in g w i t h M. G u ignon’ s (p. 720) o b s e r v a t i o n s ,
he s t a t e s t h a t as th e wing expands th e c h i t i n o f th e wing
u n d e rg o e s a p r o g r e s s i v e d e s s i c a t i o n and i t s e l a s t i c i t y
d i m i n i s h e s . ( . . . "m ais, a u f u r e t a m esure q u i e l l e se
d i s t e n d , l a c h i t i n e de l * a i l e s u b i t une d e s s i c a t i o n p r o -
11
g r e s s i v e e t son é l a s t i c i t é d i m in u e " .)
I t must be c o n fe s s e d t h a t i t a p p e a rs t h a t th e b u t t e r f l y
i s d r y i n g i t s wings d u r in g th e i n a c t i v e h o u r a f t e r em ergence.
■ A lso f o r th e dead specim en w hich r e q u i r e s s t r a i g h t e n i n g o f
a b e n t wing one has o n ly to p u t th e specim en in a h u m id ity
cham ber, a llo w th e ,w in g s t o go lim p , s t r a i g h t e n ou t th e d e
f e c t , and d ry i n th e new p o s i t i o n . The e x p e rim e n ts d e s c r i b e d
below would te n d t o c o n t r a d i c t t h i s i d e a o f d r y i n g , and
s u g g e s t t h a t h u m id ity i s n o t a f a c t o r in th e h a rd e n in g o f
th e w in g s.
W igglew orth (1 9 4 7 :2 9 ) i n h i s " P r i n c i p l e s o f I n s e c t
P h y s io lo g y " n o te s t h a t . . . " L i t t l e i s known o f th e a c t u a l
c h e m is tr y o f h a rd e n in g ; p ro b a b ly ( l i k e b la c k e n in g ) i t i s an
o x i d a t i v e change i n th e e x i s t i n g s e c r e t i o n s , c a t a l y z e d by
o x i d a t i v e enzym es, r a t h e r th a n an im p re g n a tio n w ith o t h e r
s u b s ta n c e s ; f o r n e i t h e r h a rd e n in g n o r d a rk e n in g ta k e s p la c e
n o rm a lly i n an atm o sp h ere o f n i t r o g e n . "
E x p e rim e n ta l
M a t e r i a l . The o n ly s p e c i e s u sed i n t h i s s tu d y was
F a p i l i o z e l i c a o n L u c a s.
The l o c a t i o n o f t h i s s p e c i e s i n th e f i e l d h a s a l r e a d y
b e e n d i s c u s s e d u n d e r f e e d i n g . The specim ens were c o l l e c t e d
a b o u t once a week from S eptem b er 1949 u n t i l August 1950.
T here w as, how ever, a p e r io d when t h e r e were none t o be fo u n d ,
from a b o u t th e m id d le o f December u n t i l th e m id d le o f M arch.
12
No s p e c i a l c o l l e c t i n g s c h e d u le was f o llo w e d . E a r l y i n th e
s p r i n g one s ta n d o f f e n n e l was th o r o u g h ly c l e a r e d o f a l l
a v a i l a b l e c a t e r p i l l a r s . W hether due t o t h i s or o t h e r f a c t o r s ,
no f u r t h e r f i n d s were made u n t i l l a t e J u l y . I n o t h e r s ta n d s
a p r a c t i c e of t a k i n g o n e - h a l f o f a l l c a t e r p i l l a r s was made.
These s ta n d s were n e v e r th e n c o m p le te ly d e v o id of m a tu rin g
a n im a ls . I t i s n o t known t o th e w r i t e r how f a r t h i s b u t t e r f l y
w anders from th e o r i g i n a l s ta n d t o l a y h e r e g g s . The tim e
in v o lv e d i n r a i s i n g a specim en from c o l l e c t e d egg to em er
gence i s o n ly f i v e weeks i f th e c h r y s a l i s i s g r e e n . The
brown c h r y s a l i d s can l a s t a t l e a s t f o r s e v e r a l m onths.
The c a t e r p i l l a r s were k e p t in g a l l o n j a r s . A p ie c e
of new spaper was dampened and put i n t o th e bo tto m o f th e j a r ,
t o m a in ta in h u m id ity , and f o r e a s e i n c l e a n i n g .
The h ig h h u m id ity p r e v e n te d to o r a p i d d r y in g of th e
f e n n e l . For young c a t e r p i l l a r s up t o tw e n ty c o u ld be pu t
i n t o one j a r and have a d e q u a te food f o r one^ week.
F o r th e c a t e r p i l l a r s in th e l a s t s t a g e th e food was
e a t e n i n a b o u t f o u r day s by f i v e c a t e r p i l l a r s , and th u s had
t o be r e p l e n i s h e d a b o u t tw ic e a w eek.
Young c a t e r p i l l a r s may be l e f t u n c o v e re d , b u t th e
o l d e r ones w ander when th e y p r e p a r e to p u p a te , and m ust be
s e a l e d i n th e j a r #
M ethod. In g e n e r a l d u r in g th e summer se a s o n th e g r e e n
c o lo r e d c h r y s a l i d s l a s t a b o u t two w e ek s, and th e n th e
13
b u t t e r f l y em e rg es. The wing c o lo r i n g s t a r t s show ing th r o u g h
th e c h r y s a l i s c a s e a b o u t a day b e f o r e em erg en ce. Emergence
i s u s u a l l y in th e m o rn in g , th u s by c h i l l i n g th e c h r y s a l i s a
day o r so one can r e t a r d d e v e lo p m e n t. T h is a llo w s one t o
remove th e c h r y s a l i s from th e r e f r i g e r a t o r in th e m o rn in g ,
c o n f i d e n t t h a t emergence w i l l o c cu r i n a b o u t one h o u r .
The c h r y s a l i s c a s e i s b ro k e n b a c k o f th e head c ase
and a lo n g a seam betv/een th e a n te n n a e and wing c o v e r s . T his
whole s e c t i o n lo w e rs u n d e r p r e s s u r e from th e f e e t , and th e
b u t t e r f l y emerges and may c ra w l as h ig h as p o s s i b l e . F i n a l l y
th e an im al comes t o r e s t , hangs u p sid e down, th e w ings h an g in g
as f l a b b y s a c k s •
The f i r s t e x p e rim e n t d e s c r i b e d below t e s t e d th e th e o r y
t h a t th e wings h a rd e n as a r e s u l t of d r y i n g . To t e s t t h i s
i d e a th e b u t t e r f l i e s upon em ergence, o r b e t t e r s l i g h t l y
b e f o r e , were p la c e d i n a h u m id ity j a r . T his was an o r d i n a r y
l a r g e mouthed j a r . T h is j a r was f i x e d w ith a s c r e e n a c r o s s
th e b o tto m a b o u t two in c h e s from th e b o tto m . About one and
o n e - h a l f in c h e s l e v e l of. h o t w a te r was th e n p la c e d i n th e j a r ,
w h ich was th e n s e a l e d w i t h l a r d . As th e w a te r c o o le d th e
r e l a t i v e h u m id ity would become 100^ and c o n d e n s a tio n would
ta k e p la c e on th e w a l l s o f th e j a r . A crum pled p a p e r was in
th e j a r t o a llo w th e emerged b u t t e r f l y t o hang u p sid e down,
f r e e from o b s t r u c t i o n s . T here was an o b s e r v a t i o n window made
by ru b b in g a s e c t i o n i n s i d e th e j a r w ith " C h r y s ta l M i s t , "
14
a s a lv e used to p r e v e n t fo g g in g o f s p e c t a c l e s .
In th e open a b u t t e r f l y would n o rm a lly a tte m p t f l i g h t
i n a b o u t one and o n e - h a l f h o u r s . One t e s t o f th e e f f e c t of
th e h u m id ity chamber on th e h a rd e n in g of th e w ings was to
r e l e a s e th e b u t t e r f l y a f t e r a s i m i l a r tim e and see i f i t
c o u ld f l y a t o n c e .
A more d i r e c t e f f e c t of th e h u m id ity of h a rd e n in g
was o b se rv e d b y p l a c i n g th e c h r y s a l i s i n s i d e a p e r f o r a t e d
box ( 4 i by 2 i by i t i n c h e s ) . I n t h i s box th e b u t t e r f l y cou ld
hang u p s id e down, b u t th e w ings to u c h e d th e b o tto m and c u rv e d ,
th u s when h a rd e n in g to o k p la c e th e c u rv e d t i p s would be a
perm anent f e a t u r e o f th e w ing .
O ther e x p e r im e n ta l m ethods u sed were s im p le , and need
no s p e c i a l e x p l a n a t i o n . These m ethods w i l l be d e s c r i b e d o r
a p p a r e n t from th e e x p e r im e n ts .
E x p e rim e n ts on th e d r y in g o f th e w i n g s . On t e n
d i f f e r e n t o c c a s io n s ( l i s t e d by d a te in T able IV) a b u t t e r f l y
was p u t i n t o th e h u m id ity chamber b e f o r e em ergence. Upon
em ergence th e b u t t e r f l y clim b ed up th e p a p e r , pumped out
i t s w in g s-a n d a llo w e d them t o h a r d e n . On an a v e r a g e , th e
b u t t e r f l i e s w ere removed i n a b o u t one and o n e - h a l f h o u r s . I t
was c o n s id e r e d t h a t th e wings were n o rm a lly h a rd i f th e
b u t t e r f l y c o u ld f l y when to s s e d i n t o th e a i r . T h is tim e of
one and o n e - h a l f h o u rs was d e r i v e d from a s e r i e s o f o b s e r
v a t i o n s i n w hich b u t t e r f l i e s w ere a llo w e d t o h a rd e n t h e i r
15
w ings n o r m a lly , and f l y when s t a r t l e d by waving a hand a t
them .
A second method was t o p la c e th e b u t t e r f l y i n a sm a ll
box so t h a t a crum pled wing would r e s u l t i f perm anent
h a rd e n in g to o k p l a c e . T h i r t e e n specim ens were used f o r t h i s
p u r p o s e . I t was removed a t th e end of from one and o n e - h a l f
h o u rs to f i v e h o u r s . The l o n g e r tim e was som etim es n e c e s s a r y
s in c e one c o u ld n o t alw ays assume th e c h r y s a l i s j u s t put
i n t o th e chamber would emerge i n one h o u r , and i n th e box
t h e r e was no way t o se e a t e x a c t l y what tim e th e b u t t e r f l y
d i d em erge. I f t h e r e was a perm anent bend i n th e wing i t was
c o n s id e r e d t h a t h a rd e n in g had ta k e n p la c e d e s p i t e 100^
r e l a t i v e h u m id ity .
E x p erim en t on o x i d a t i v e r e a c t i o n . Only one e x p e r i
ment was ru n t o t e s t W ig g le s w o rth ’ s s u g g e s tio n t h a t
h a rd e n in g was due to o x i d a t i o n . I n t h i s case a b u t t e r f l y ,
new ly em erged, was drowned i n b o i l e d w a te r (c o o le d and
p re su m a b ly f r e e from o x y g e n ). The w ings on t h i s b u t t e r f l y
f a i l e d t o h a r d e n . A lso s e v e re d w ings put i n t o th e b o i l e d
w a t e r f a i l e d t o h a r d e n . T h is e x p e rim e n t was n o t r e p e a t e d
b e c a u s e i t k i l l e d th e b u t t e r f l y , and as th e n e x t e x p e r i
ment shov/s, dead b u t t e r f l i e s f a i l t o h a rd e n t h e i r w in g s .
E xperim ent on th e s e v e re d w in g s . Three e x p e rim e n ts
were ru n by s e v e r i n g th e wing o f a newly emerged b u t t e r
f l y and a wing p a r t i a l l y ex p an d e d . I n a l l t h r e e c a s e s t h e s e
16
s e v e re d w ings rem ain ed s o f t and p l i a b l e , ev en a f t e r norm al
e x p o su re t o a i r (o x y g e n ).
E x p erim en t on k i l l i n g th e b u t t e r f l y . Two e x p e rim e n ts
were ru n i n w hich th e b u t t e r f l y was k i l l e d by d e c a p i t a t i o n
im m e d ia te ly a f t e r emergence w i t h th e o b se rv e d r e s u l t t h a t
th e w ings d i d n o t h a rd e n .
E x p erim en t w ith th e a n e s t h e s i a . These e x p e rim e n ts
s t a r t e d as an a c c i d e n t . I n an e f f o r t t o k i l l th e newly
emerged b u t t e r f l y w i t h e t h e r and th u s a v o id a b a d ly b le d
dead b u t t e r f l y , a newly emerged b u t t e r f l y was p u t f o r one
h o u r i n t o a j a r w ith some e t h e r . At th e end of th e h o u r
th e b u t t e r f l y was rem oved. I t r e v i v e d , b u t th e w in g s,
p r e v i o u s l y pumped up f a i l e d t o h a r d e n , a lth o u g h t h i s b u t t e r
f l y l i v e d f o r f i v e d a y s . T h is e x p e rim e n t was r e p e a t e d on
t h r e e o c c a s io n s w i t h th e same r e s u l t s , a lt h o u g h th e b u t t e r
f l y d i d n o t alw ays l i v e f o r f i v e days — one d ie d i n two
d a y s . I n a l l t h e s e c a se s th e w ings w ere in c a p a b le o f
s u p p o r t i n g f l i g h t when th e b u t t e r f l y was throw n i n t o th e
a i r .
17
DISCUSSION
T his i n v e s t i g a t i o n , w hich began as an e f f o r t to
d e te r m in e w hich e c o l o g i c a l f a c t o r s in d u ce th e w ings to h a r d e n ,
has d e v e lo p e d i n t o a problem t h a t w i l l p ro b a b ly r e q u i r e the
e f f o r t s o f a b io c h e m is t t o c o m p le te ly u n r a v e l .
F i r s t o f a l l th e e c o l o g i c a l p ro c e s s o f d r y i n g , a s
s u g g e s te d by G uignon, does n o t a p p ea r to be th e b e s t a n s w e r.
The r e s u l t s o f th e e x p e rim e n ts i n t h i s s t u d y makes i t a p p e a r
p ro b a b le t h a t th e w ings h a rd e n in d e p e n d e n t of r e l a t i v e
h u m id ity .
I f i t i s n o t a d r y in g p ro c e s s th e n e x t s u g g e s t i o n in
th e l i t e r a t u r e i s t h a t some s o r t of an o x i d a t i v e p ro c e s s
c a u s e s th e w ings t o h a rd e n . W ig g lesw o rth n o te s t h a t th e
w ings do n o t h a rd e n i n n i t r o g e n . I f as seems l i k e l y , th e
n i t r o g e n k i l l s th e b u t t e r f l y t h i s i s n o t to o s u r p r i s i n g ,
s i n c e th e w ings o f a dead b u t t e r f l y w i l l n o t h a rd e n ev en in
a i r . T h is would n o t e l i m i n a t e W igg lesw o rth * s o b s e r v a t i o n
t h a t h a rd e n in g i s q u i t e l i k e l y "an o x i d a t i v e change in
e x i s t i n g s e c r e t i o n s , c a t a l y s e d by o x i d a t i v e enzym es. The
e x p e rim e n ts w i t h th e a n e s t h e t i z e d b u t t e r f l i e s would te n d t o
c o n firm t h i s i d e a . Enzymes a r e g e n e r a l l y p rod uced f o r a
l i m i t e d tim e in re s p o n s e t o some p h y s i c a l o r c h em ic al body
c h a n g e . S in c e wing h a rd e n in g ta k e s p la c e d u r in g th e f i r s t
h o u r a f t e r em erg ence, i t i s p l a u s i b l e t o assume t h a t th e
enzyme i s a l s o pro duced o n ly d u rin g t h i s tim e . I f t h a t be
18
t r u e i t would fo llo w t h a t i f th e b u t t e r f l y were knocked
o u t ( b u t n o t k i l l e d ) so t h a t b lo o d and a i r c i r c u l a t i o n
sto p p e d d u r in g t h i s c r i t i c a l h o u r th e enzyme would n e v e r be
c a r r i e d t o th e p a r t o f th e wing w here i t was n e ed e d . Even
i f the- b u t t e r f l y r e v iv e d l a t e r t h e r e would n o t be th e
n e c e s s a r y c a t a l y s t needed t o cause th e w ings t o h a r d e n . A
s e v e re d wing would n o t h a r d e n , a p p a r e n t l y f o r l a c k o f th e
same c a t a l y s t .
T here i s one o t h e r p o s s i b i l i t y , t h a t of a d e p o s i t i o n
of c a lc iu m p r e c i p i t a t e s o r some o t h e r p r e c i p i t a t i o n , and an
im p re g n a tio n of th e c u t i c l e and s u p p o r t i n g t r a c h e a e w i t h t h i s
s u b s t a n c e . W ig g le sw o rth (1947) d i s c o u n t s t h i s p o s s i b i l i t y ,
b u t gave no s p e c i f i c r e a s o n s . He, o f c o u r s e , b e l i e v e d i n th e
o x i d a t i v e c a t a l y s t .
W orkers such as M e rc e r, who s t u d i e d th e wing s t r u c t u r e ,
b u t w i t h no e s p e c i a l r e g a r d f o r th e problem of h a r d e n i n g ,
seem t o t h i n k th e wing i s c o v ered w i t h c h i t i n b e f o r e th e p u p a l
s t a g e , and n o te no change i n t h i s s t r u c t u r e l a t e r ; t h a t i s
th e y make no m en tio n o f any s o r t of change due t o im p re g
n a t i o n s .
CONCLUSION
The h a rd e n in g of th e wing o f F a p i l i o z e l i c a o n Lucas
i s n o t due t o d r y i n g . I t i s p o s s i b l e t h a t s t i f f e n i n g i s due
t o im p re g n a tio n s o f c a lc iu m o r some o t h e r s u b s t a n c e , b ro u g h t
i n a t t h e tim e th e wing i s expanded by th e b lo o d . I t i s more
19
p ro b a b le t h a t th e wing c h i t i n h a rd e n s by some o x i d a t i v e p r o c e s s
u n d e r c o n t r o l o f an enzyme produced d u rin g th e hou r w h ile
h a rd e n in g t a k e s p la c e and i s c a r r i e d i n t o th e wing by th e
blo o d which expands the wing sack*
SUMMARY
The f e e d i n g of F a p i l i o z e l ic a o n Lucas i s n o t l i m i t e d
e n t i r e l y to t h e f a m ily U m b e ll i f e r a e , s i n c e th e l a r v a e w i l l
e a t a t l e a s t one s p e c i e s o f C i t r u s . In th e U m b e llif e r a e the
l a r v a e a re found t o e a t any member o f the f a m ily so f a r t e s t e d .
Only th e p o is o n hem lock ( Gonium maculaturn L .) was found o f t e n
to prove u n s u i t a b l e a s f o o d , b u t even on t h i s p l a n t a t
l e a s t h a l f o f the l a r v a e l i v e d t o com plete p u p a tio n .
In t h i s a r e a th e c a t e r p i l l a r i s found t y p i c a l l y on
f e n n e l (p o en icu lum v u lg a r e H i l l ) which i s a p l a n t n a t i v e t o
Europe* The n a t i v e p l a n t s t a t e d to be th e u s u a l food o f
P* z e l i c a o n i s n o t found i n th e a r e a so u th o f S a n ta C la ra
County.
I t was found t h a t t h e r e a r e two colors, o f c h r y s a l i s .
The g ree n c o lo r e d c h r y s a l i s o r d i n a r i l y co m p lete s th e p u p a l
s t a g e i n a b o u t two w eeks, th e brown c h r y s a l i s nay rem a in in
th e p u p a l s t a g e f o r m onths. I t was o b se rv e d t h a t t h e r e i s an
i n c r e a s e i n th e r a t i o o f g re e n t o brown c o lo r e d c h r y s a l i d s
from a b o u t E:1 i n th e e a r l y s p r i n g t o a lm o s t e n t i r e l y g re e n
by the l a s t o f J u l y . E f f o r t s to c o r r e l a t e t h i s change w ith
2 0
c o o lin g (o r c o o l w e a th e r) were u n s u c c e s s f u l*
The h a rd e n in g of th e w ings o f th e b u t t e r f l y was found
t o be in d e p e n d e n t o f d ry in g * O ther p o s s i b i l i t i e s f o r th e
cau se o f h a rd e n in g seemed t o be b e s t e x p la in e d by an enzyme
w hich d u r in g the f i r s t h o u r a f t e r em ergence cause c h a n g e s ,
p ro b a b ly o f an o x i d a t i v e n a t u r e , i n th e c h i t i n c o v e rin g and
t r a c h e a e of th e wing t h a t makes th e n h a rd and r i g i d .
21
TABLE I
FEEDING HABITS OP FAPILIO ZELICAON LUCAS
PLANTS TESTED CATERPILLARS USED RESULT
P o p l a r (Populua S p .)
L i l a c (S y rin g a v u l g a r i s )
P o is o n Hemlock
(Gonium maculatura L .)
3 l a r g e
3 medium
3 sm a ll
1 l a r g e
3 medium
3 sm a ll
5 l a r g e -
4 s m a l l -
C e le ry (Apium g r a v e l o l e n s L .)
6 mixed
C a r r o t (Daucus c a r o t a L .) 10 mixed
Oenanthe sa rra e n to sa P r e s l . 6 mixed
L e p to ta e n a d i s s e c t a N u t t . 6 mixed
Orange ( C i tr u s s p . ) 8 mixed
de ad i n 8 days
dead i n 6 days
d e ad in 4 d ays
dead i n 6 days
dead i n 2 -5 days
dead i n 3 -5 days
A l l would f e e d
upon th e p l a n t
b u t u s u a l l y d ie d
w i t h i n 3 d a y s ,
one r e a c h e d th e
p u p a l s t a g e •
Would f e e d and
grow t o p u p a te .
Would fe e d and
grow to p u p a te .
Would fe e d and
grow t o p u p a te .
Would f e e d and
grow t o p u p a te .
S ix d ie d ,^ o n e
sm a ll one a t e
th e r i n d o f f of
a g re e n o r a n g e ,
one p u p a te d .
22
TABLE I I
PUPA AM D COLOR OF BÂCKGROUMD
DATE C H S CIREEM O fd liR C M V IT
No. g r e e n brown - c o lo r e d g re e n
c h r y s a l i s
BACKGROUND
brown
12 May 6 8 -
31 May 6 2 1
3 June 5 5 1
8 June 3 3 -
11 June 7 0 -
12 June 8 1 1 -
16 June 5 0 1 -
20 June 6 3 1 1
26 June 10 3 2 1
1 J u l y 7 0 ( a l l c h i l l e d ) -
26 J u l y 15
0 —
A lso c h i l l e d were g re e n c h r y s a l i s from :
3 June 1
20 June 1
26 June 2
23
TABLE I I I
NUMBER OP CHRYSALIDS REMAINING IN A LOT
j & I P T I B r R T T W f C ) ort ] N I C H R I S I M T E I S I C f l
Rem aining on 12 June Green Brown
c o lo r e d c h r y s a l i d s
O v e rw in te r l o t (34)
12 May l o t (6 g r e e n .
8 brow n;
31
6
-3 June l o t (5
6
g r e e n .
brow n) 1 6
L ot On 29 June
g re e n brown
On 5 J u l y
g r e e n brown
On 26 J u l y
g re e n brown
12 May (6 g r e e n .
8 brow n; 1 6 1 6 1 6
3 June (5-6 ) 0 5 . 0 5 0 5
8 June (3 -2 ) 0 2 0 2 0 2
11 June (7 -0 ) 0 0 - - - -
12 June (9 -1 ) 0 1 0 1 0 1
20 June (6-3)
5 2 0 2 0 2
26 June (12 -4 ) 12 4 10 1
(3 d ie d )
2 1
Over w i n t e r (3-
.31) 3 31
24
TABLE IV
TIM iG R J ü X a iT I r L B Ü D I P C H R B L A jR ID E N IN G C U T T H IS 'JriI?G 5
Date R e le a s e d from h u m id ity chamber a f t e r :
1% NoVf 1949 1 hour 30 m in u te s
26 Nov. 1 hour 25 m in u te s
3 Deo. 1 hour
11 Dec* 1 hour 45 m in u te s
17 May 1950 1 ho ur
30 m in u te s
< 3:5 jW Ü & T f 1 hour
20
m in u te s
24 May 2 h o u rs 40 m in u te s
25 May 2 h o u rs 5 m in u te s
25 May 1 hour
30
m in u te s
26 May 1 hour 30 m in u te s
(T able c o n tin u e d on n e x t page)
25
Date R e le a s e d from p e r f o r a t e d box in th e
h u m id ity chamber a f t e r :
28 May 1 hour
30 m in u te s - n o t y e t
em erged, r e u s e d
n e x t day.
. 31 May 1 hour
30
m in u te s two sp e c im e n s ,
one n o t y e t
em erged.
11 June 6 h o u rs two s p e c im e n s .
lô June 5 h o u rs two sp e cim en s.
19 June 1 hour
30 m in u te s - n o t y e t
em erged, used n e x t
day.
20
June 4 h o u rs two sp e c im e n s.
29 June 3 h o u rs two sp e c im e n s.
30
June 2 h o u rs
30
June 1 hour 30 m in u te s
LITERATURE CITED
Comstock, J . A . , B u t t e r f l i e s of C a l i f o r n i a . B u l l . So., C a l i f .
Acad. S c i . , 2 0 ( 1 ) : 5 -6 , 1921.
E s s i g , G .E ., I n s e c t s of W e ste rn N o rth A m erica. N.Y.
M a c m illia n C o ., 1035 p p . , 1926.
G uignon, G a b r i l , Sur l e developm ent P o st-n y m p h a l d e s a i l e s d e s
L e p i d o p t e r e s . C.R. Acad. S c i . P a r i s . 2 0 4 :7 2 0 -7 2 1 , 1936.
B e r i n g , M a r t in , B io lo g ie d e r S c h m e t t e r l i n g e . B e r l i n , 480 p p . ,
19 26..
M e rce r, W illiam F a i r f i e l d , The Developm ent o f th e Wings i n th e
L e p i d o p t e r a . J o u r . New York Sntora. S o c . , 8 : 1 - 2 0 . , 1896.
M o f f a t, J . A . , Remarks on th e S t r u c t u r e o f th e U ndeveloped
Wings o f th e S a t u r n i i d a e . Entom. S o c. o f O n ta rio ,
2 5 : 6 3 - 6 5 . , 1894.
P a r i s h , S . B . , The Im m igrant P l a n t s o f S o u th e rn C a l i f o r n i a .
B u l l . So. C a l i f . Acad. S c i . , 2 1 ( 4 ) : 3 - 3 0 . , 1920.
S n o d g ra s s , R .E ., P r i n c i p l e s o f I n s e c t M orphology. McGraw-
H i l l N.Y. 667 p p . , 1935.
W ig g le s w o rth , V .B ., P r i n c i p l e s of I n s e c t P h y s io lo g y . Methuen
and Co. L td . London 434 p p . , 1947.

Asset Metadata

Creator Brown, Bernard Eldon (author)

Core Title Some contributions to the life cycle of Papilio zelicaon Lucas with special reference to the hardening of the wings

Contributor Digitized by ProQuest (provenance)

Degree Master of Arts

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag Biological Sciences,OAI-PMH Harvest

Format application/pdf (imt)

Language English

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c39-276676

Unique identifier UC11316825

Identifier EP67188.pdf (filename),usctheses-c39-276676 (legacy record id)

Legacy Identifier EP67188.pdf

Dmrecord 276676

Document Type Thesis

Format application/pdf (imt)

Rights Brown, Bernard Eldon

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the au...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus, Los Angeles, California 90089, USA

Linked assets

University of Southern California Dissertations and Theses