V
O L U M
E
6 6 , N
U M
B E R
2
8 1
Journal of the Lepidopterists’ Society
6 6 (2 ), 2 0 1 2 , 8 1 – 1 1 0
N E W
B R O M E L IA D -F E E D IN G
A N D
STRYMON S P E C I E S F R O M B I G B E N D N A T I O N A L P A R K , T E X A S , U S A
I T S V I C I N I T × ( L ×C A E N I D A E : T H E C L I N A E )
N
IC K
V . G
R IS H IN
H o w a r d H u g h e s M e d ic a l I n s titu te a n d D e p a r tm e n t o f B io c h e m is tr y , U n iv e r s ity o f T e x a s S o u th w e s te r n M e d ic a l C e n te r,
5 3 2 3 H a r r y H in e s B lv d , D a lla s , T X , U S A 7 5 3 9 0 - 9 0 5 0 , g r is h in @ c h o p .s w m e d .e d u
A N D
C
H R IS T O P H E R
J. D
U R D E N
C u r a to r E m e r itu s o f E n to m o lo g y a n d G e o lo g y , U n iv e r s ity o f T e x a s a t A u s tin , T e x a s N a tu r a l S c ie n c e C e n te r, T e x a s M e m o r ia l M u s e u m ,
2 4 0 0 T r in ity S t., A u s tin , T X , U S A 7 8 7 0 5 . d r d n @ m a il.u te x a s .e d u
ABSTRACT. Strymon solitario G r i s h i n & D u r d e n , new species i s d e s c r i b e d f r o m s o u t h w e s t T e x a s , U S A . T h i s serapio g r o u p
s p e c i e s o f Strymon i s d i s t i n g u i s h e d f r o m i t s c l o s e s t r e l a t i v e s b y a c o m b i n a t i o n o f : ( 1 ) r e s t r i c t e d a r e a s o f l i g h t s c a l e s b a s a l l y f r o m
s u b m a r g in a l d a r k e r lu n u le s o n th e v e n tr a l h in d w in g — w ith th e s e lig h t m a c u le s n o t r e a c h in g 1 /3 o f th e d is ta n c e b e tw e e n th e lu n u le s
a n d p o s tm e d ia n o r a n g e b a n d ; (2 ) s m a ll o v a l w e ll-s e p a r a te d o r a n g e m a c u le s in th e p o s tm e d ia n v e n tr a l h in d w in g b a n d ; (3 ) th e p r e s e n c e o f p o s tb a s a l o r a n g e m a c u le s o n th e v e n tr a l h in d w in g in s o m e s p e c im e n s ; (4 ) lim ite d to a lm o s t a b s e n t v io le t b lu e d o r s a l w in g
o v e r s c a lin g ; (5 ) o c c a s io n a l o r a n g e o v e r s c a lin g o n th e d o r s a l fo r e w in g in fe m a le s ; (6 ) r o u n d e r w in g s , p a r tic u la r ly in m a le s ; (7 ) n o t
p r o m in e n tly d e v e lo p e d a r e a s o f d a r k s c a le s o n le g tib ia e in m o s t s p e c im e n s , a n d (8 ) a r id , d e s e r t a n d r o c k y s lo p e s h a b ita t. R e la t i o n s h i p s w i t h o t h e r Strymon s p e c i e s a r e d i s c u s s e d a n d l i f e h i s t o r y d a t a a r e p r e s e n t e d a n d i l l u s t r a t e d . Hechtia texensis ( B r o m e l i a c e a e ) is e s ta b lis h e d a s th e la r v a l fo o d p la n t in th e B ig B e n d N a tio n a l P a r k a r e a . O v e r 4 0 s p e c im e n s a n d liv e in d iv id u a ls o f th e n e w
s p e c ie s a r e illu s tr a te d to s h o w th e e x te n t o f w in g p a tte r n v a r ia tio n .
Additional key words: jacqueline, megarus, s c r u b - h a i r s t r e a k , T a m a u l i p a s , B o q u i l l a s , T h e S o l i t a r i o , s p e c i e s c o n c e p t
T h e w o r d “ h a ir s tr e a k ” fo r m a n y p e o p le is a lm o s t
s y n o n y m o u s w i t h Strymon H ü b n e r , 1 8 1 8 ( L y c a e n i d a e :
T h e c l i n a e ) , a s Strymon melinus H ü b n e r , 1 8 1 8 i s
a b u n d a n t, w id e s p r e a d fr o m C a n a d a to A r g e n tin a , a n d
fl i e s d u r i n g m o s t o f t h e w a r m p a r t o f t h e y e a r . I t w i l l
n o t b e a n e x a g g e r a t i o n t o s a y t h a t Strymon i s t h e b e s t k n o w n g e n u s o f A m e r ic a n h a ir s tr e a k s . I n te r e s tin g ly ,
w h i l e o n l y v e r y f e w b u t t e r fl i e s a r e c o n s i d e r e d t o b e
c r o p p e s t s o f a n y s i g n i fi c a n c e , a t l e a s t t w o Strymon
s p e c i e s a r e s e r i o u s c o m m e r c i a l p i n e a p p l e p e s t s , i . e . S.
megarus ( G o d a r t , [ 1 8 2 4 ] ) a n d S. ziba ( H e w i t s o n , 1 8 6 8 )
(H a r r is 1 9 2 7 , C a r te r 1 9 3 4 , S ilv a e t a l. 1 9 6 8 ,
B e u te ls p a c h e r 1 9 7 2 , O te r o & M a r ig o 1 9 9 0 , R o b b in s
2 0 1 0 ). U s a g e o f p in e a p p le fa m ily (B r o m e lia c e a e ) p la n ts
b y b u t t e r fl y c a t e r p i l l a r s i s n o t c o m m o n . T h e o n l y
E u m a e in i (L y c a e n id a e : T h e c lin a e ) r e c o r d e d to fe e d o n
t h e s e p l a n t s b e l o n g t o Strymon ( serapio a n d ziba
s p e c ie s g r o u p s , R o b b in s & N ic o la y 2 0 0 2 ), a n d it is
l i k e l y t h a t t h e serapio g r o u p u s e s B r o m e l i a c e a e a s t h e
o n ly fo o d p la n t (R o b b in s 2 0 1 0 ).
F o llo w in g C le n c h (1 9 6 1 ), R o b b in s a n d N ic o la y
( 2 0 0 2 ) c h a r a c t e r i z e Strymon b y t h e a n t e r i o r l y d i r e c t e d
te e th o n th e p o s te r io r d o r s a l s u r fa c e o f m a le g e n ita lic
v a lv a e (F ig . 7 c ). T h is c h a r a c te r m ig h t r e p r e s e n t a n
e v i d e n c e f o r m o n o p h y l y o f t h e g e n u s a n d n i c e l y u n i fi e s
m o s t s p e c ie s th a t h a v e b e e n h is to r ic a lly p la c e d in
Strymon. O n l y S. ziba s p e c i e s g r o u p l a c k s w e l l -
d e v e lo p e d te e th (F ig . 7 # 6 c ). H o w e v e r, it w a s k e p t in
Strymon b e c a u s e i t i s l i k e l y t h a t S. ziba h a s l o s t t h e
v a l v a l t e e t h a n d t h u s t r u l y b e l o n g s t o Strymon, a s i t
e x h ib its v e r y s tr o n g w in g p a tte r n s im ila r itie s w ith
Strymon megarus f r o m t h e S. serapio g r o u p . S o m e
s p e c i m e n s o f S. ziba a n d S. megarus m i g h t n o t b e
r e lia b ly s e p a r a b le b y w in g p a tte r n s (R ic k a r d & G r is h in
2 0 1 0 ).
R o b b i n s a n d N i c o l a y ( 2 0 0 2 ) d e fi n e S. serapio a n d S.
ziba g r o u p s b y t w o c o r n u t i i n t h e p e n i s t i p ( F i g . 7 d ) ,
p a i r e d i n serapio g r o u p ( F i g s . 7 # 1 d , # 3 d , # 5 d ) a n d
u n p a i r e d i n ziba g r o u p ( F i g . 7 # 6 d ) . L a r v a e o f b o t h
g ro u p s fe e d
o n
B r o m e l i a c e a e , serapio g r o u p
e x c l u s i v e l y . M o s t S. serapio g r o u p s p e c i e s p o s s e s s
d o w n -tu r n e d p e n is tip (F ig s . 7 # 3 – # 5 ) a n d a s im p le
s c le r o tiz e d lo o p o f th e d u c tu s b u r s a e (F ig . 8 # 5 ). I n
c o n t r a s t , g e n i t a l i a o f S. ziba a r e v e r y d i s t i n c t i v e . M a l e s
h a v e a n u p -tu r n e d p e n is tip w ith 2 v e r y la r g e c o r n u ti
(F ig . 7 # 6 ), a n d fe m a le s la c k th e s c le r o tiz e d lo o p o n th e
d u c tu s b u r s a e , b u t th e p o s te r io r e n d o f th e c o r p u s
b u r s a e is e x p a n d e d a n d p r o m in e n tly s c le r o tiz e d ,
fo r m in g a v e r y c h a r a c te r is tic s tr u c tu r e te r m e d "h o o d "
b y J o h n s o n a n d c o lle a g u e s (J o h n s o n e t a l. 1 9 9 0 , A u s tin
& J o h n s o n 1 9 9 7 ) (F ig . 8 # 7 ).
U n til r e c e n tly (R ic k a r d & G r is h in 2 0 1 0 ), B r o m e lia d f e e d i n g Strymon s p e c i e s w e r e n o t k n o w n f r o m t h e U S ,
m a y b e s o m e w h a t s u r p r is in g ly , b e c a u s e S p a n is h m o s s
8 2 8 2
J
( Tillandsia usneoides ( L . ) L . ) i s a c o m m o n p l a n t o v e r
th e e a s te r n p a r t o f th e c o u n tr y a n d o th e r s p e c ie s o f
Tillandsia o c c u r i n A r i z o n a , T e x a s a n d F l o r i d a . I n t h e
s o u t h w e s t e r n d e s e r t o f T e x a s , Hechtia texensis S .
W a t s o n , a n d H. glomerata Z u c c . o c c u r ( a l l p l a n t s
m e n tio n e d a b o v e a r e in th e fa m ily B r o m e lia c e a e ). T h e
d i s c o v e r y o f a r e s i d e n t Hechtia- f e e d i n g Strymon
s p e c ie s in T e x a s w a s m a d e in d e p e n d e n tly b y s e v e r a l
c o lle c to r s , a n d th e d a ta w e r e p u t to g e th e r o n ly r e c e n tly .
T h is s y n th e s is r e s u lte d in n e w r e s e a r c h a n d c o lle c tin g
e ffo r ts th a t c u lm in a te d in th is p u b lic a tio n .
M
A T E R IA L S A N D
M
E T H O D S
T h is s tu d y is b a s e d o n c o m p a r is o n o f o v e r 1 4 5 0
s p e c im e n s fr o m
t h e serapio a n d ziba g r o u p s o f
Strymon i n t h e N a t i o n a l M u s e u m o f N a t u r a l H i s t o r y
(U S N M ), S m ith s o n ia n I n s titu tio n , W a s h in g to n , D C ;
A m e r ic a n M u s e u m o f N a tu r a l H is to r y (A M N H ), N e w
×o r k , N ×; N a t i o n a l H i s t o r y M u s e u m , L o n d o n
(B M N H ), G r e a t B r ita in ; B u r k e M u s e u m (U W B M ),
S e a ttle , W A ; T e x a s M e m o r ia l M u s e u m
(T M M C ),
A u s tin , T X ; T e x a s A & M U n iv e r s ity I n s e c t C o lle c tio n
(T A M U ), C o lle g e
S ta tio n , T X ; s e v e r a l p r iv a te
c o l l e c t i o n s ; s p e c i m e n s fi e l d - c a u g h t b y t h e a u t h o r s
u n d e r p e r m its (B ig B e n d N a tio n a l P a r k a n d T h e
S o lita r io r e g io n ); p h o to g r a p h s o f liv e in d iv id u a ls m a d e
b y s e v e r a l o b s e r v e r s ; a n d im a g e s in J a n z e n &
H a llw a c h s o n -lin e d a ta b a s e (2 0 1 1 ). I n a d d itio n , th e
p r i m a r y t y p e s p e c i m e n s o f S. serapio a n d S. megarus
w e r e i n s p e c t e d u s i n g t h e p h o t o g r a p h s ( a n d S. serapio
s p e c im e n b y N V G ) o b ta in e d fr o m
B M N H
a n d
M H N M , r e s p e c tiv e ly , a n d illu s tr a te d h e r e in . T h e la te
G e o r g e T A u s t i n k i n d l y s h a r e d s o m e Strymon
d is tr ib u tio n r e c o r d s in M e x ic o fo r s p e c im e n s fr o m
F lo r id a M u s e u m o f N a tu r a l H is to r y M c G u ir e C e n te r
fo r L e p id o p te r a a n d B io d iv e r s ity (M G C L ), G a in e s v ille ,
F L . U n p u b lis h e d n o te s , le tte r s a n d la b n o te b o o k s b y
R o y O . K e n d a ll a n d C o n n ie A . K e n d a ll r e le v a n t to th e
S. serapio g r o u p w e r e s t u d i e d a t t h e T A M U K e n d a l l &
K e n d a ll c o lle c tio n a n d d is c u s s e d h e r e . S ta n d a r d
e n to m o lo g ic a l te c h n iq u e s w e r e u s e d fo r s p e c im e n
p r e p a r a tio n a n d d is s e c tio n (R o b b in s 1 9 9 1 ). G e n ita lia
a n d w in g v e n a tio n te r m in o lo g y fo llo w K lo ts (1 9 7 0 ) a n d
C o m s to c k (1 9 1 8 ), r e s p e c tiv e ly . P la n t n a m e s fo llo w th e
P L A N S o n -lin e d a ta b a s e (U S D A , N R C S 2 0 1 1 ). L e n g th
m e a s u r e m e n ts a r e in m e tr ic u n its a n d w e r e m a d e fr o m
p h o t o g r a p h s o f s p e c i m e n s m a g n i fi e d o n a c o m p u t e r
s c r e e n . M o s t p h o to g r a p h s w e r e m a d e u s in g N ik o n
D 2 0 0 c a m e r a , fo r s p e c im e n s th r o u g h a 1 0 5 m m f/2 .8 G
A F -S V R M ic r o -N ik k o r le n s ; fo r “ d r y ” g e n ita lia o f th e
h o lo ty p e w ith a n a d d itio n a l 2 ×te le c o n v e r te r T C -2 0 E ;
a n d
“ w e t” g e n ita lia th r o u g h
a “ lig h t-th r o u g h ”
m ic r o s c o p e .
O U R N A L
O F
R
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
E S U L T S
H is to r ic a l R e c o r d s
b r ie f h is to r y o f r e c o r d s fo r th is a p p a r e n tly
u n d e s c r ib e d s p e c ie s fo llo w s . T h e e a r lie s t r e c o r d k n o w n
t o u s i s f r o m S e p t e m b e r 2 5 , 1 9 4 4 , “A m e r i e s B o q u i l l a s " ,
R io G r a n d e R iv e r, B ig B e n d N a tio n a l P a r k (T e x a s :
B r e w s te r C o .) a n d is a s e r ie s o f 1 6 s p e c im e n s a t th e
B u r k e M u s e u m ( W A :S e a ttle ) c o lle c te d b y G e o r g e
S c h e n k . T h e B o q u illa s a r e a is th e r e g io n o f
T e x a s /M e x ic o w e s t o f B o q u illa s C a n y o n to H o t S p r in g s
(o r T o r n illo C r e e k ), a d is ta n c e o f a p p r o x im a te ly 2 0
m ile s a r o u n d th e M e x ic a n v illa g e o f B o q u illa s , a n d
n o r th o f th e R io G r a n d e fo r 2 – 3 m ile s . T h e a r e a is
m o s t l y l i m e s t o n e w i t h Hechtia texensis p l a n t s a b u n d a n t
o n r o c k y s l o p e s . Hechtia texensis ( F a l s e A g a v e ,
A g u a p ié ) is r e s tr ic te d to th e lim e s to n e s o ils in th e
v ic in ity o f th e R io G r a n d e a n d S o lita r io in T e x a s , b u t
g r o w s o n lim e s to n e s o ils s o m e d is ta n c e to th e s o u th
a lo n g th e b a s e o f th e S ie r r a d e l C a r m e n (C J D & R o
W a u e r, p e r s . c o m m .) a n d is r e p o r te d fr o m
b o th
C h ih ia h u a a n d C o a h u ila in M e x ic o .
T h is 1 9 4 4 r e c o r d s u r fa c e d o n ly r e c e n tly . L ik e ly
in d e p e n d e n tly o f th is r e c o r d , R .O . K e n d a ll c o lle c te d a
fe m a le s p e c im e n o n M a r 2 7 , 1 9 6 8 a lo n g th e R io
G r a n d e V illa g e N a tu r e T r a il (B ig B e n d N a tio n a l P a r k ),
w h ic h is in th e B o q u illa s a r e a . C o n tin u e d e ffo r ts b y th e
K e n d a lls r e s u lte d in a n o th e r s p e c im e n , m a le , c o lle c te d
b y C o n n ie A . K e n d a ll o n S e p te m b e r 1 4 ? (p e r la b e l, b u t
S e p 2 0 p e r K e n d a ll n o te s fr o m T A M U a r c h iv e ) 1 9 7 1 ,
O ld S a n V ic e n te [c r o s s in g o n R io G r a n d e ] (B ig B e n d
N a tio n a l P a r k , a ls o in B o q u illa s a r e a ). T h e s e s p e c im e n s
b e a r th e fo llo w in g d e te r m in a tio n la b e ls b y C le n c h a n d
b y M ille r, r e s p e c tiv e ly : [“ T h e c la ”// s p . f// m a y b e n e w
/ / d e t . H . C l e n c h 1 9 6 9 ] a n d [ Strymon s p . / / p o s s i b l y
n e w m// d e t. L e e D . M ille r // 1 9 8 0 ] .
C JD c o n d u c e d fa u n a l su rv e y s fo r th e T e x a s G e n e ra l
L a n d O f fi c e ( 1 9 7 3 ) a n d f o r T h e N a t u r a l A r e a s S u r v e y
(1 9 7 5 ) o n T h e B ig B e n d R a n c h , in P r e s id io a n d
B r e w s te r C o u n tie s (s o m e 5 0 a ir m ile s w e s t fr o m th e
B o q u illa s ), w h ic h s u p p o r te d th e s e ttin g a s id e o f T h e
B ig B e n d R a n c h S ta te N a tu r a l A r e a . H e p h o to g r a p h e d
(M a y 1 7 , 1 9 7 3 ) a n d c o lle c te d (M a y 2 0 , 1 9 7 3 ; J u n e 8 ,
1 9 7 5 ) a s e r ie s o f s p e c im e n s fr o m T h e S o lita r io in
P r e s id io & B r e w s te r C o u n tie s a n d r e p o r te d (D u r d e n
1 9 7 6 ) t h e s e a s a n e w s p e c i e s o f Strymon w h i c h l o o k e d
s u p e r fi c i a l l y l i k e Tmolus azia, w h i c h h e h a d t a k e n
p r e v io u s ly D e c e m b e r 2 6 , 1 9 7 0 a t G o m e z F a r ia s ,
T a m a u lip a s , M e x ic o . T h e S o lita r io is th e r in g -s h a p e d
m o u n ta in r a n g e o f w h ic h F r e s n o P e a k is th e h ig h p o in t.
T h e S o lita r io , a c ir c u la r fe a tu r e v is ib le fr o m s p a c e is a n
u n -r o o fe d lo p o lith — a b u lg e o v e r a n ig n e o u s in tr u s io n
n o w la r g e ly r e m o v e d b y e r o s io n . C J D d id n o t a s s o c ia te
A
V
O L U M
E
6 6 , N
U M
B E R
2
t h e s p e c i m e n s w i t h B r o m e l i a d s , a l t h o u g h Hechtia i s a
fr e q u e n t c o m p o n e n t o f th e s lo p e v e g e ta tio n in T h e
S o lita r io a n d a lo n g F r e s n o C r e e k . J a m e s A . S c o tt
(1 9 8 6 ) c ite s th e s e r e c o r d s b u t, w ith o u t e x a m in in g
s p e c im e n s o r p ic tu r e s r e p o r ts th e m
a s Strymon
bebrycia ( H e w i t s o n , 1 8 6 8 ) o c c u r r i n g i n t h e B i g B e n d
A re a .
A ll fu r th e r in fo r m a tio n a b o u t th e in s e c t (u n til 2 0 0 5 )
c o m e s fr o m th e n o te s in th e K e n d a ll & K e n d a ll
c o lle c tio n a r c h iv e , T e x a s A & M U n iv e r s ity (T A M U ),
C o lle g e S ta tio n , T e x a s . N o ta b ly , in a le tte r to R ic h a r d S .
P e ig le r, D e p a r tm e n t o f E n to m o lo g y , T A M U , d a te d 1 7
O c to b e r 1 9 7 8 H a r r y K . C le n c h c a lle d K e n d a ll
s p e c im e n s fr o m
1 9 6 8 a n d 1 9 7 1 Strymon serapio
(G o d m a n & S a lv in , 1 8 8 7 ). T h is d e te r m in a tio n b r o u g h t
B r o m e l i a c e a e ( i . e . Hechtia) a s p o s s i b l e l a r v a l h o s t
p la n ts to a tte n tio n .
E d w a r d C . K n u d s o n c o lle c te d a s in g le m a le o n 2 9 N o v -1 9 8 5 n e a r H o t S p r in g s (B ig B e n d N a tio n a l P a r k ,
B o q u i l l a s a r e a ) , t h a t h e r e f e r r e d t o a s “u n f a m i l i a r
L y c a e n i d r e s e m b l i n g a s o m e w h a t l a r g e Tmolus azia,
b u t i t w a s n o t Tmolus echion, b u t t h e s i z e t h e r e o f . ” T h i s
s p e c im e n w a s lo a n e d to R o b e r t K . R o b b in s , w h o in a
l e t t e r d a t e d J a n u a r y 3 1 , 1 9 8 6 c a l l e d i t Strymon serapio.
A n n B . S w e n g e l o b s e r v e d a b o u t a d o z e n s p e c im e n s
a lo n g th e R io G r a n d e V illa g e N a tu r e T r a il (B ig B e n d
N a tio n a l P a r k , B o q u illa s a r e a ) o n A p r il 1 6 1 9 8 9 ,
h i l l t o p p i n g a n d p e r c h i n g o n l e c h u g u i l l a ( Agave
lechuguilla T o r r . , ( A g a v a c e a e ) ) , c r e o s o t e b u s h ( Larrea
tridentata ( D C . ) C o v i l l e , ( Z y g o p h y l l a c e a e ) ) a n d
Jatropha [ dioica C e r v . ] ( E u p h o r b i a c e a e ) . S o m e
s p e c im e n s w e r e p h o to g r a p h e d a n d th is r e c o r d w a s
r e p o r te d in 1 9 8 9 L e p id o p te r is ts ' S o c ie ty S e a s o n
S u m m a r y ( N e w s . L e p i d . S o c . 2 : 2 2 1 9 9 0 ) a s Strymon
bebrycia, w h i c h i s q u i t e s i m i l a r b e l o w .
R o y O . a n d C o n n ie A . K e n d a ll d id m o r e w o r k in th e
B o q u illa s a r e a in 1 9 8 7 , a n d c o lle c t e d 1 f e m a le o n
M a r c h 2 6 , 2 fe m a le s o n M a r c h 2 7 (b e tw e e n th e B a r k e r
H o u s e th a t is u s e d a s lo d g in g fa c ility fo r r e s e a r c h e r s in
th e p a r k , a n d B o q u illa s C a n y o n ) a n d 1 fe m a le o n
M a rc h 2 8 (b y th e B a rk e r H o u se ). O n M a rc h 3 1 1 9 8 7
a n o th e r fe m a le w a s s e e n , b u t e s c a p e d c a p tu r e . A ll
t h e s e f e m a l e s w e r e k e p t a l i v e w i t h fl o w e r b u d s o f
Hechtia texensis, b u t n o e g g s w e r e l a i d .
O n A p r il 5 , 1 9 9 2 R .O . K e n d a ll fo u n d o n e la s t in s ta r
l a r v a f e e d i n g o n Hechtia texensis fl o w e r b u d s , r o a d t o
B o q u illa s C a n y o n o v e r lo o k a c r o s s r o a d fr o m th e B a r k e r
H o u s e ( B B N P ) . A b o u t 1 0 0 0 fl o w e r s t a l k s w e r e
e x a m in e d d u r in g tw o d a y s , b u t n o a d d itio n a l la r v a e
w e r e fo u n d . T h e la r v a s p u n -u p o n A p r il 1 0 a n d p u p a te d
o n A p r i l 1 3 , 1 9 9 2 . P a r a s i t o i d Metadontia amoena ( S a y ,
1 8 3 6 ) (H y m e n o p te r a : C h a lc id id a e ) h a tc h e d fr o m th a t
p u p a o n M a y 9 1 9 9 2 . T h is p a r a s ito id is k n o w n to u s e a
8 3
w i d e r a n g e o f T h e c l i n a e a s h o s t s , i n c l u d i n g Strymon
melinus ( p e r K e n d a l l c o l l e c t i o n n o t e s ) .
S te p h e n M . S p o m e r c o lle c te d a m a le o n M a r c h 2 5 ,
1 9 9 4 a lo n g T X R a n c h R d 2 6 2 7 , 2 5 m i S E o f U S H w y
3 8 5 (c a . 3 m i N W o f R io G r a n d e ). T h is lo c a tio n is 2 0 m i
n o r th fr o m th e B o q u illa s a r e a .
T h e n , a fte r a 1 0 -y e a r h ia tu s , th e B ig B e n d H a ir s tr e a k
p r o je c t c a m e to life a g a in w ith th e e ffo r ts o f s e v e r a l
p h o to g r a p h e r s (R ic h K o s te c k e , A n d r e w
S p e n c e r,
M a r tin R e id , R o W a u e r ) e x p lo r in g B ig B e n d w h o
o b ta in e d g o o d q u a lity d ig ita l p h o to g r a p h s o f th is
Strymon. U n d e r a B i g B e n d N a t i o n a l P a r k r e s e a r c h
p e r m it, N V G c o lle c te d a s e r ie s o f s p e c im e n s in 2 0 0 5 ,
2 0 0 7 a n d 2 0 0 9 a n d r e a r e d t h e i n s e c t o n Hechtia
texensis fl o w e r b u d s f r o m o v a o b t a i n e d f r o m c a p t i v e
fe m a le s .
A d d itio n a lly , a n a ly s is o f p h o to g r a p h s (b y K im D a v is
& M ik e S ta n g e la n d a n d D a n H a r d y ) fr o m n e a r C iu d a d
V ic to r ia (M e x ic o : T a m a u lip a s , c a . 5 0 0 m ile s s o u th e a s t
fr o m B o q u illa s ) r e v e a le d p h e n o ty p e q u ite s im ila r to
t h a t f r o m t h e B i g B e n d A r e a , a n d Hechtia s p . p l a n t s
w e r e a ls o p h o to g r a p h e d in th e im m e d ia te v ic in ity o f
th e h a ir s tr e a k s ig h tin g (W a r r e n e t a l. 2 0 1 1 ).
I t b e c a m e a p p a r e n t d u e t o d i f fi c u l t i e s w i t h p l a c i n g
t h e s e s p e c i m e n s a m o n g k n o w n t a x a ( bebrycia, serapio,
p o s sib le n e w s p e c ie s ), th a t a c a r e fu l a n a ly s is o f th e
s itu a tio n is w a r r a n te d . T h e a n a ly s is r e s u lte d in th e
c o n c lu s io n
th a t th e se
s p e c im e n s r e p r e s e n t a
b io lo g ic a lly d is tin c t s p e c ie s , w h ic h is d e s c r ib e d h e r e a s
Strymon solitario G r i s h i n &
D u r d e n , new species
Diagnosis:
T h e p r e s e n c e o f a n te r io r ly d ir e c te d te e th o n th e
p o s te r io r d o r s a l s u r fa c e o f m a le g e n ita lic v a lv a e (F ig .
7 # 1 c ) p l a c e s t h i s s p e c i e s i n Strymon. D o w n - t u r n e d
p e n is tip w ith p a ir e d c o r n u ti in m a le s (F ig . 7 # 1 d ),
lo o p e d d u c tu s b u r s a e in fe m a le s (F ig s . 8 # 1 – # 4 ) a n d
l a r v a l f o o d p l a n t f r o m t h e B r o m e l i a c e a e ( Hechtia s p p . )
p o s i t i o n i t i n t h e S. serapio s p e c i e s g r o u p .
D o r s a l w i n g s u r f a c e i s s i m i l a r t o S. serapio ( F i g s .
1 0 # 1 d – # 6 d , # 2 0 d ), e x c e p t th a t in m a le s v io le t-b lu e
o v e r s c a l i n g i s l e s s d e v e l o p e d t h a n i n t h e m a j o r i t y o f S.
serapio m a l e s , e s p e c i a l l y j u s t b a s a l l y f r o m t h e o r a n g e
s u b m a r g in a l s p o t in C u 1– C u 2 c e ll (F ig s . 3 # 1 d ,
4 # 1 d – # 1 2 d , 9 # 1 5 , 1 0 # 2 2 – # 2 3 , # 2 6 ). T h is s h o r ta g e o f
b l u e s c a l e s r e s e m b l e s S. ziba ( F i g . 1 0 # 7 d ) , n o t a c l o s e l y
r e la te d (a s r e v e a le d b y its g e n ita lia s tr u c tu r e s , F ig s .
7 # 6 , 8 # 7 ) , b u t a s u p e r fi c i a l l y s i m i l a r s p e c i e s . S o m e
fe m a le s p o s s e s s o r a n g e s c a le s in th e p o s tm e d ia n a r e a o f
fo r e w in g s (F ig s . 3 # 2 d , 5 # 4 d , # 8 d , # 1 1 d , 9 # 1 1 , 1 0 # 2 7 d ),
a b s e n t i n a l l e x a m i n e d S. serapio s p e c i m e n s , a n d m o s t
fe m a le s h a v e m o r e p r o n o u n c e d s la te o v e r s c a lin g in th e
8 4 8 4
p o s tb a s a l a r e a ( e .g . F ig s . 1 0 # 2 4 d – # 2 5 d ) th a n a ty p ic a l
S. serapio f e m a l e ( F i g . 1 0 # 2 0 d ) , i n w h i c h m a c u l e s
f o r m e d b y s l a t e s c a l e s a r e c o n fi n e d t o t h e m a r g i n a l a r e a
o f a h in d w in g .
V e n t r a l w i n g s u r f a c e a n d g e n i t a l i a a r e s i m i l a r t o S.
jacqueline N i c o l a y & R o b b i n s , 2 0 0 5 ( F i g s . 1 0 # 1 8 v ,
# 1 9 v , # 2 1 v , 7 # 5 ) a n d S. megarus ( F i g s . 1 0 # 9 v – # 1 2 v ,
# 1 4 v – # 1 7 v ). T h e m o s t n o ta b le v e n tr a l p a tte r n
d iffe r e n c e fr o m th e la tte r tw o s p e c ie s is th e s m a ll
a m o u n t o f lig h t s c a lin g b e tw e e n th e p o s tm e d ia n o r a n g e
b a n d o f s p o ts o n v e n tr a l h in d w in g a n d m a r g in a l d a r k e r
lu n u le s (F ig s . 3 # 1 v – # 2 v , 4 # 1 v – # 1 2 v , 5 # 1 v – # 1 2 v ,
9 # 1 – # 1 0 , # 1 2 – # 1 4 , # 1 6 – # 1 9 , 1 0 # 8 , # 2 2 v – # 2 7 v ). L ig h t
s c a l e s a r e c o n fi n e d t o t h e v i c i n i t y o f d a r k e r m a r g i n a l
lu n u le s fr a m in g th e m , a n d d o n o t r e a c h 1 /3 o f th e
d is ta n c e fr o m th e lu n u le s to o r a n g e s u b m a r g in a l b a n d .
T h is p a tte r n g iv e s th e v e n tr a l s u r fa c e a g r a y e r, m o r e
u n i f o r m l y fl a t a n d l e s s c o n t r a s t i n g a p p e a r a n c e .
G e n ita lia a r e n o t o b v io u s ly d is tin c tiv e (F ig s . 4 # 1 – # 2 ),
a lth o u g h th e te r m in a lly n a r r o w e r, p o in te d a n d m o r e
a s y m m e t r i c s a c c u s ( a s i n S. megarus, F i g . 4 # 3 ) , s e e m s
t o s e p a r a t e t h e n e w s p e c i e s f r o m S. jacqueline ( F i g .
4 # 5 ). N o t e n o u g h m a te r ia l e x is ts to e x a m in e th e e x te n t
o f S. jacqueline g e n i t a l i a v a r i a t i o n .
D o r s a l w i n g s u r f a c e i s n o t s i m i l a r t o S. jacqueline
(F ig s . 1 0 # 1 8 d – # 1 9 d , # 2 1 d ), d u e to th e la c k o f e x te n s iv e
b lu e a r e a s . O c c a s io n a l p r e s e n c e o f o r a n g e s c a le s in th e
p o s t b a s a l a r e a o f t h e S. solitario f e m a l e s i s a c h a r a c t e r
s h a r e d o n l y w i t h S. jacqueline ( F i g s . 1 0 # 1 9 d , # 2 1 d ) .
D o r s a l w i n g s u r f a c e i s l e s s s i m i l a r t o S. megarus t h a n t o
S. serapio, b e c a u s e i n S. megarus m a l e s , e x t e n s i v e
v io le t-b lu e a r e a s u s u a lly c o v e r th e p o s te r io r h a lf o f th e
h in d w in g
a n d
p ro n o u n c e d
b a s a lly
e lo n g a te d
s u b m a r g in a l b r o w n m a c u le s a r e p r e s e n t w ith in th e s e
a r e a s (F ig s . 1 0 # 9 d – # 1 2 d ). I n th e n e w s p e c ie s , v io le tb lu e s c a lin g is le s s d e v e lo p e d , a n d r a th e r v io le t-b lu e
m a c u le s a r e a p p a r e n t o n th e b r o w n b a c k g r o u n d o f th e
h in d w in g s u b m a r g in a l a r e a (F ig s . 3 # 1 d , 4 # 1 d – # 1 2 d ,
9 # 1 5 , 1 0 # 2 2 , # 2 3 , # 2 6 ).
V e n tr a l w in g s u r fa c e a n d m a le g e n ita lia a r e n o t
s i m i l a r t o S. serapio ( F i g s . 1 0 # 1 v – # 6 v , # 2 0 v , 7 # 4 ) , a s i n
th e la tte r (fr o m
T e x a s so u th to P a n a m a ) o ra n g e
p o s tm e d ia n s p o ts fo r m a m o r e c o n tin u o u s b a n d o n th e
h in d w in g , i.e . th e th r e e s p o ts in c e lls R s – M 1, M 1– M 2
a n d M 2– M 3, a r e e lo n g a te d a n d a r e a lm o s t in lin e (F ig s .
1 0 # 1 v – # 6 v , # 2 0 v ) , w h ile in th e n e w s p e c ie s s p o ts
a p p e a r to b e m o r e s e p a r a te a n d r o u n d e r, w ith th e
R s – M 1 a n d M 1– M 2 s p o ts b e in g o ffs e t b a s a d , a n d d is ta d ,
r e s p e c t i v e l y ( e . g . F i g . 3 # 1 v – # 2 v ) . A d d i t i o n a l l y , S.
serapio d o e s n o t h a v e t h e p o s t b a s a l o r a n g e s p o t s o n t h e
h in d w in g s th a t a r e fr e q u e n tly d e v e lo p e d in th e n e w
s p e c ie s (F ig s 3 # 1 v , 4 # 1 v – # 6 v , 5 # 1 v – # 6 v , 9 # 6 , # 9 , # 1 2 ,
# 1 3 , # 1 4 , # 1 6 , # 1 7 , 1 0 # 2 2 v , # 2 7 v ) . M a l e g e n i t a l i a o f S.
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
serapio ( F i g . 7 # 4 b ) a r e c h a r a c t e r i z e d b y a s h o r t
r o u n d e d a n d m o s tly s y m m e tr ic s a c c u s (lo n g e r,
a s y m m e tr ic a n d te r m in a lly p o in te d in th e n e w s p e c ie s :
F ig . 7 # 1 b – # 2 b ) a n d a la r g e te g u m e n o c c u p y in g th e
d o r s a l h a l f o f t h e g e n i t a l i c c a p s u l e ( s m a l l e r , s i m i l a r t o S.
megarus ( F i g . 7 # 3 a ) t e g u m e n i n t h e n e w s p e c i e s : F i g .
7 # 1 a – # 2 a ).
W ild -c a u g h t s p e c im e n s o f th e n e w s p e c ie s w e
e x a m in e d fr o m th e B ig B e n d a r e a e x h ib it u n iq u e le g
p a tte r n in w h ic h b la c k s c a lin g o n th e tib ia is le s s
d e v e lo p e d (F ig s . 6 a – c , l) th a n in a ll s im ila r s p e c ie s
(F ig s . 6 g – k ) a n d if p r e s e n t, fo r m s s p o ts o f a fe w g r a y is h
s c a le s r a th e r th a n c le a r p a tc h e s o f b la c k s c a le s p r e s e n t
in o th e r c lo s e ly r e la te d s p e c ie s .
D e s e r t h a b ita t o f th e n e w s p e c ie s (F ig s . 2 a – c ) is
s i m i l a r t o t h a t o f S. jacqueline a n d n o t s i m i l a r t o b r u s h
a n d f o r e s t h a b i t a t s o f S. serapio a n d S. megarus.
I n s u m m a r y , th is s p e c ie s c o n tin u e s th e tr e n d
r e c e n tly e s ta b lis h e d fo r o th e r n e w ly d e s c r ib e d
Strymon: i t s c h a r a c t e r s r e v e a l a n e w c o m b i n a t i o n o f
f e a t u r e s k n o w n i n o t h e r s p e c i e s . E . g . S. jacqueline i s a
“c h i m e r a ” o f S. dindus ( F a b r i c i u s , 1 7 9 3 ) d o r s o a n d S.
megarus v e n t e r , a n d S. giffordi N i c o l a y & R o b b i n s ,
2 0 0 5 c o m b i n e s S. veterator ( H . H . D r u c e , 1 9 0 7 ) d o r s o
w i t h S. megarus v e n t e r . L i k e w i s e , i n t h e n e w s p e c i e s ,
o c c a s io n a l p r e s e n c e o f o r a n g e s c a le s o n th e d o r s a l
fo r e w in g , v e n tr a l w in g p a tte r n , a n d d e s e r t h a b ita t
p r e f e r e n c e a r e s h a r e d w i t h S. jacqueline, d o r s a l w i n g
p a t t e r n i s s h a r e d w i t h S. serapio, a n d v e n t r a l w i n g
p a t t e r n i s s i m i l a r t o S. megarus. M a l e g e n i t a l i a c o m b i n e
f e a t u r e s o f S. jacqueline a n d S. megarus. T h e m o s t
p ro n o u n c e d
u n iq u e
tr a its
a re
c o n s p ic u o u s ly
u n d e r d e v e lo p e d lig h te r s c a le s in th e h in d w in g
p o s tm e d ia n a r e a , m u c h w e a k e r d a r k b a n d s o n th e le g
tib ia e (in B ig B e n d p o p u la tio n ), a n d le s s p r o n o u n c e d
v io le t b lu e s c a lin g , e s p e c ia lly in th e p o s te r io r
p o s tm e d ia l a r e a o f th e h in d w in g . T h e s e c h a r a c te r s g iv e
th e n e w s p e c ie s a m o r e n e u tr a l, lo w e r c o n tr a s t
a p p e a r a n c e w ith le s s w h ite , le s s d a r k a n d le s s b lu e th a n
t h e d e s c r i b e d serapio g r o u p s p e c i e s .
Description. Male (N=30): F W l e n g t h ( b a s e R t o a p e x ) = 1 3 . 8
m m ( h o lo ty p e ) , m e a n 1 2 .6 m m , s ta n d a r d d e v ia tio n 0 .8 m m , r a n g e
1 1 .0 – 1 3 .9 m m ( n = 1 2 ) .
Dorsal wing pattern ( F i g s . 3 # 1 d , 4 ) : T h e ground color
b r o w n i s h g r a y , t h e s c a l e s w i t h a c o p p e r r e fl e c t a n c e . G r o u n d c o l o r
fa d e d in s o m e p a r a ty p e s , b u t th e a n d r o c o n ia l p a tc h s till d a r k .
Forewing: a n androconial patch 2 . 7 m m l o n g b y 2 . 0 m m w i d e i n t h e
h o lo ty p e , w ith e n la r g e d d a r k a p ic a lly 3 - to 5 -d e n tic u la te o v e r s c a le s .
E n la r g e d lig h t y e llo w is h -g r a y g la n d u la r u n d e r s c a le s w ith a p u c k e r e d
r o u n d e d a p e x . B a s a l fi f t h o f a n d r o c o n i a l p a t c h a l m o s t a s l i g h t i n t o n e
a s th e d is c a l g r o u n d c o lo r. T h e b lu e o v e r s c a lin g o f s c a tte r e d lig h t
v io le t-b lu e s c a le s . I n th e h o lo ty p e th is o v e r s c a lin g in fo r e w in g c e ll
C u 2 – 2 A a r o u n d a n d b a s a d fr o m th e o r ig in o f C u 2 v e in o c c u p y in g h a lf
o f th e w in g le n g th , e v e n m o r e e x te n s iv e a r o u n d 2 A , r e a c h in g th e
p o s tm e d ia n a r e a d is ta d a n d a lm o s t h a lf o f th e C u 2 – 2 A c e ll c e p h a la d .
I n c e ll 2 A o v e r s c a lin g in th e b a s a l h a lf. I n m a n y p a r a ty p e s b lu e
V
O L U M
E
6 6 , N
U M
B E R
2
o v e r s c a lin g r e d u c e d to tw o s tr e a k s a lo n g v e in s C u 2 a n d 2 A , o r a lm o s t
a b s e n t. P r o m in e n t o r a n g e o v e r s c a lin g a lo n g th e fo r e w in g c o s ta , w id e r
d is ta d b u t n o t e x p r e s s e d o r lo s t in s o m e p a r a ty p e s . I n th e h o lo ty p e ,
hindwing o v e r s c a l i n g o f t h e s a m e s c a t t e r e d l i g h t v i o l e t - b l u e s c a l e s i n
th e p o s te r io r p a r t o f th e c e ll, b a s e o f c e ll M 3– C u 1, b a s a l 3 /5 o f c e ll
C u 1– C u 2 , a n d a lm o s t e n tir e c e ll C u 2 – 2 A w ith a fe w s c a le s in th e b a s a l
h a lf im m e d ia te ly p o s te r io r to 2 A . M a c u le s o f v io le t-b lu e s c a le s in th e
s u b m a r g in a l a r e a o f h in d w in g in c e lls M 1– M 2 , M 2 – M 3 a n d M 3– C u 1. I n
s o m e p a r a ty p e s th e b lu e o v e r s c a lin g r e d u c e d , a n d in C u 2– 2 A it m a y
b e r e d u c e d to th e b a s a l a r e a a n d a s m a ll s u b m a r g in a l b lu e m a c u le .
B a n d s o f d a r k e r s m a ll s c a le s im m e d ia te ly o v e r th e v e in s th r o u g h th e
a r e a s w ith o v e r s c a lin g . T h e r e d d is h -o r a n g e s u b m a r g in a l c u b ita l
m a c u le o n th e h in d w in g d is ta lly p u p ille d w ith d a r k s h a d e . S iz e o f th e
o r a n g e m a c u le v a r ia b le . A n a l lo b e w ith p r o m in e n t r e d d is h -o r a n g e
s c a le s . T h e s u b m a r g in a l lin e o f b lu e s c a le s o n h in d w in g p o s te r io r to
M 2 in h o lo ty p e , p o s te r io r to M 1, o r M 3 in s o m e p a r a ty p e s . D a r k o u te r
m a r g in lin e d a r k e r th a n th e g r o u n d c o lo r p o s te r io r to M 3 v e in . A
w h ite -tip p e d a n d w h ite -m a r g in e d g r o u n d c o lo r lo n g ta il a t C u 2 a n d a
s h o r t o n e a t C u 1. A b d o m in a l fo ld p a le b r o w n is h -g r a y , p o s te r io r ly
lig h te r in c o lo r. F r in g e w h ite fr o m a p e x to to r n u s in fo r e w in g , fr o m
o u te r c o s ta l a n g le to to r n u s in h in d w in g a n d d a r k , a lm o s t b la c k o n th e
a n a l lo b e o f th e h in d w in g . I n n e r m a r g in fr in g e o f g r o u n d to n e , lig h te r
o n th e h in d w in g . C o s ta l e d g e s c a lin g v e r y n a r r o w in fo r e w in g a n d o f a
c o n tr a s tin g o r a n g e c o lo r. C o s ta l e d g e in h in d w in g is a fr in g e o f th e
v e n tra l g ro u n d to n e .
Ventral wing pattern ( F i g s . 3 # 1 v , 4 ) : T h e ground color l i g h t
p i n k i s h g r a y , w i t h fringe a s o n u p p e r s i d e . Forewing w i t h o u t b a s a l
m a c u le s a n d e n d o f c e ll m a c u le . I n m a n y s p e c im e n s , C u 1 – C u 2
(e x c e p t s u b m a r g in a l a r e a ) a n d 2 A c e lls lig h te r g r o u n d c o lo r th a n th e
r e s t o f th e w in g . F o r e w in g p o s tm e d ia n b a n d o f 5 – 6 m a c u le s in c e lls
fr o m R 2– R 3 (m is s in g in s o m e p a r a ty p e s ) to C u 1– C u 2. N o m a c u le in
C u 2– 2 A . E a c h m a c u le fr a m e d d is ta lly w ith b la c k a n d th e n w ith w h ite
s c a le s a n d d is ta lly c o n v e x . T h e p o s tm e d ia n b a n d c u t a t v e in s b y s c a le s
o f g r o u n d c o lo r, r e m o v a l o f th e s e s c a le s d u e to w e a r in s o m e p a r a ty p e s
r e v e a ls p r o m in e n t d a r k e r v e in s . T h e p o s tm e d ia n b a n d s m o o th ly b e n t,
d is ta lly c o n v e x , th e m a c u le s m o s tly a lig n e d w ith th e ir n e ig h b o r s . I n
s o m e p a r a ty p e s M 1– M 2 a n d /o r M 3– C u 1 m a c u le s a r e o ffs e t d is ta d a n d
p r o x im a d , r e s p e c tiv e ly . S u b m a r g in a l b a n d o f d a r k m a c u le s (lu n u le s ) in
R 3– M 1 to C u 2– 2 A c e lls , fa d e s a n te r io r ly in s o m e s p e c im e n s . T h e m o s t
p r o m in e n t g r a y m a c u le in C u 1– C u 2. S o m e g r a y m a c u le s , m o s tly in
c e lls M 3– C u 1 to C u 2– 2 A d is ta lly fr a m e d b y lig h te r s c a le s . A d a r k
m a r g in a l b a n d o f d iffu s e la r g e g r a y m a c u le s p r e s e n t in h o lo ty p e a n d
s o m e p a r a ty p e s fr o m R 3– M 1 to C u 2– 2 A c e lls . T h e s e m a c u le s lig h te r
th a n s u b m a r g in a l g r a y m a c u le s a n d a lm o s t b le n d in to a b a n d s h o w in g
a s a d a r k e r m a r g in a l a r e a o f fo r e w in g . T h in lig h t s u b m a r g in a l lin e a n d
d a r k o u te r m a r g in a l lin e fr o m a p e x to to r n u s in fo r e w in g . O r a n g e
s c a lin g a lo n g th e c o s ta a n d o u te r m a r g in . T h e b a s a l b a n d o f o r a n g e
m a c u l e s o n t h e hindwing m u c h r e d u c e d o r o b s o l e t e . I t s s t r o n g e s t
e x p r e s s io n is th e p r e s e n c e o f 3 m a c u le s , th e la r g e s t o n e in S c + R 1– R s
c e ll, th e s e c o n d la r g e s t in th e p o s te r io r o f th e d is c a l c e ll, a b o u t 3 /5
fr o m th e b a s e o f th e c e ll a n d a s m a ll m a c u le o f a fe w s c a le s in th e
a n te r io r p a r t o f th e d is c a l c e ll, p r o x im a lly fr o m th e p o s te r io r s id e o f
c e ll m a c u le , a t 1 /2 fr o m th e b a s e . T h e la r g e s t m a c u le s w ith d a r k s c a le s
a r o u n d th e e d g e s , m o s tly a lo n g th e b a s a l s id e . J u s t 2 , 1 (in S c + R 1– R s )
o r n o s u c h m a c u le s m a y b e p r e s e n t. E n d -o f-c e ll h in d w in g m a c u le in
m o s t s p e c im e n s , c o m p o s e d o f a n a r r o w a s s e m b la g e o f d a r k e r s c a le s ,
s o m e tim e s in te r m ix e d w ith o r a n g e s c a le s . T h e h in d w in g p o s tm e d ia n
b a n d o f r e d -o r a n g e m a c u le s s tr o n g ly d e v e lo p e d in c e lls fr o m R s – M 1
to 2 A a n d m u c h m o r e ir r e g u la r th a n o n th e fo r e w in g . T h e m a c u le s a r e
w e ll s e p a r a te d fr o m e a c h o th e r, r o u n d e d n e a r v e in s . M a c u le s o f
v a r y in g e x p r e s s io n a n d w id th d is ta lly fr a m e d w ith d a r k a n d th e n w ith
lig h t s c a le s . S c + R 1– R s m a c u le , a n d to a le s s e r e x te n d s o m e o th e r
m a c u le s w ith d a r k s c a le s o n th e p r o x im a l e d g e in s e v e r a l s p e c im e n s .
E x a c t lo c a tio n a n d r e la tiv e p o s itio n in g o f th e m a c u le s v a r ia b le , b u t a s
a tr e n d , th e R s – M 1, M 3– C u 1, a n d a n te r io r s e g m e n t o f C u 2– 2 A m a c u le s
a r e s h ifte d p r o x im a lly ; M 2– M 3, C u 1– C u 2, p o s te r io r s e g m e n t o f C u 2– 2 A
a n d a n te r io r s e g m e n t o f 2 A a r e o ffs e t d is ta lly . T h e b a n d h a s a n
a p p e a r a n c e o f d o u b le ts o f c lo s e r a s s o c ia te d m a c u le s : S c + R 1– R s w ith
R s – M 1, M 1– M 2 w ith M 2– M 3, a n d M 3– C u 1 w ith C u 1– C u 2. M a c u le s in
C u 2– 2 A a n d 2 A c e lls in a b r o a d e x te r n a l a p e x V -s h a p e d a r r a n g e m e n t
8 5
w ith s lig h tly c o n v e x s id e s . S u b m a r g in a l b a n d o f d a r k m a c u le s
(lu n u le s ) in S c + R 1– R s to 2 A c e lls , fa d in g in M 1– M 2 a n d M 2– M 3 in
s o m e s p e c im e n s . T h e la r g e s t m a c u le in R s – M 1 c e ll. M a c u le s a r e
fr a m e d w ith w h ite s c a le s . W h ite s c a le s b e tw e e n th e d a r k m a c u le s a n d
th e p o s tm e d ia n b a n d o f o r a n g e m a c u le s n o t e x te n s iv e , d o n o t r e a c h
1 /3 o f th e d is ta n c e p r o x im a d o f g r a y m a c u le s , a n d la r g e ly p r e s e n t a s
fr a m in g o f th e s e g r a y m a c u le s . O r a n g e -r e d c u b ita l s u b m a r g in a l
m a c u le d is ta lly p u p ille d w ith b la c k a n d p r o x im a lly fr a m e d w ith d a r k
g r a y a n d th e n lig h te r s c a le s . B la c k a n a l lo b e e d g e d w ith o r a n g e -r e d
s e g m e n t a n d w h ite s c a lin g . T h e m a r g in a l o r a n g e -r e d s e g m e n t b y th e
to r n u s L -s h a p e d o n th e le ft w in g a n d c o n s is ts o f a m a c u le in th e d is ta l
1 /3 o f C u 2– 2 A c e ll a n d a lo n g tr ia n g u la r m a c u le in 2 A a lo n g th e in n e r
m a r g in o f h in d w in g w ith th e s m o o th ly c u r v e d p r o x im a l m a r g in fa llin g
s e v e r a l s c a le r o w s d is ta l to p o s tm e d ia n o r a n g e b a n d . T h is 2 A m a c u le
a b r u p tly c u t b y a m a r g in a l tr ia n g le o f w h ite a t th e in n e r c o r n e r o f th e
a n a l lo b e w ith its a c u te a p e x o n th e s u b m a r g in a l b a n d o f d a r k
m a c u le s . D a r k m a r g in a l b a n d o f d iffu s e la r g e g r a y m a c u le s lig h te r in
c o lo r th a n th e s u b m a r g in a l b a n d . T h e s e m a c u le s n o t w e ll s e p a r a te d
fr o m
e a c h o th e r a lo n g v e in s a n d c r e a te a n im p r e s s io n o f a
s u b m a r g in a l a r e a b e in g d a r k e r th a n th e g r o u n d c o lo r o f h in d w in g .
T h is fe a tu r e v e r y p r o m in e n t in m o s t s p e c im e n s , h o w e v e r, th e m a le
r e a r e d in th e la b s h o w s a n u n u s u a lly d a r k g r o u n d c o lo r n o t c o n tr a s tin g
w ith th e m a r g in a l a r e a . T h e w h ite s u b m a r g in a l lin e c u t b y d a r k e r
s c a lin g o n a ll v e in s a n d w id e n s fr o m v e in S c + R to to r n u s , m o s t
p r o m in e n t in c e lls C u 1– C u 2 a n d C u 2– 2 A , le s s in M 3– C u 1. D a r k o u te r
m a r g in a l lin e w id e n s fr o m th e a p e x to w a r d s to r n u s .
Head w i t h t h e c o l l a r o f l o n g g r a y s c a l e s , f r i n g e d a n t e r i o r l y w i t h a
fe w o r a n g e s c a le s . T h is o r a n g e fr in g e a b s e n t in s o m e s p e c im e n s . A
tu ft o f lo n g w h ite s c a le s s u r r o u n d e d b y g r a y p r o m in e n t b e tw e e n th e
a n te n n a e , b u t e ls e w h e r e o n th e c r o w n th e s c a le s g r a y , v e r y s h o r t a n d
a p p r e s s e d . T h e fr o n s c o p io u s ly c lo th e d w ith w h ite s c a le s in tr u d e d b y
s p a r s e b la c k h a ir s . T h e b a s a l jo in t o f th e p a lp h e a v ily s c a le d w ith
w h ite . A p e x o f p a lp b a s a lly w h ite w ith s h o r t s c a le s g r a d in g d is ta lly in to
g r a y s c a le s b o th a b o v e a n d b e lo w . T h e e y e s b r o w n -h a ir y . F a c e b e h in d
e y e c o p io u s ly w h ite s c a le d . A n te n n a l s e g m e n ts a p ic a lly b la c k , b a s a lly
w h ite , w h ite a r e a s la r g e r v e n tr a lly , th e b o u n d a r y ir r e g u la r in m id s e g m e n t, s h a r p b e tw e e n s e g m e n ts . C lu b v e n tr a lly s c a le d fr o m b a s e
b e y o n d its w id e s t p a r t to w ith in 3 s e g m e n ts o f a p e x . C lu b la te r o d o r s a lly s c a le d to w ith in 6 s e g m e n ts o f a p e x , th e m e s io d o r s a lly n u d e
a n d m ic r o h ir s u te a r e a e x te n d s fo r a t le a s t 1 0 s e g m e n ts fr o m th e a p e x .
T h e 5 a p ic a l s e g m e n ts o f th e c lu b o r a n g e . T h e n e x t p r o x im a l 7
s e g m e n t s b l a c k w i t h t h e fi r s t w h i t e b a n d a p p e a r i n g i n t h e n e x t
p r o x im a l s e g m e n t a t th e b a s e o f th e c lu b .
Body v e s t i t u r e v e n t r a l l y w h i t e , d o r s a l l y g r a y . A b d o m e n d o r s a l l y
g r a y , v e n tr a lly w h ite . L e g s (F ig s . 6 a – c , l, m ) b a s a lly th ic k ly c lo th e d
w ith w h ite s c a le s a n d h a ir s . T ib ia l a n d ta r s a l s p in e s a r e a c o n tr a s tin g
s h in y b la c k . E a c h ta r s o m e r e w h ite , d o r s a lly b la c k a t th e b a s e , T ib ia e
m o s tly w h ite , w ith fe w d a r k s c a le s . D a r k s c a le s o n tib ia e in 3 d o r s a l
m a c u le s : a t th e b a s e , th e d a r k e s t s m a ll m a c u le , in th e m id d le , m o r e
d iffu s e , fr e q u e n tly o b s o le te m a c u le a n d th e d is ta l m a c u le a t a b o u t 1 /4
fr o m th e ta r s u s . T h e m id d le a n d d is ta l m a c u le s u s u a lly r e d u c e d to a
fe w d a r k s c a le s , b u t in a m a le r e a r e d in th e la b m o r e d e v e lo p e d a n d
m o r e s im ila r to th o s e v is ib le o n th e p h o to g r a p h s o f s p e c im e n s fr o m
n e a r C iu d a d V ic to r ia (M E X I C O : T a m a u lip a s , F ig s . 6 d – f).
Genitalia: ( F i g . 7 ) T y p i c a l f o r Strymon. V a l v a e w i t h a n t e r i o r l y
d ir e c te d te e th o n th e p o s te r io r d o r s a l s u r fa c e , ta p e r r e g u la r ly to th e
d is ta lly to r te d a c u te a p e x . S a c c u s a s y m m e tr ic to th e r ig h t, te r m in a lly
p o in te d , s lig h tly lo n g e r th a n w id e . T e g u m e n le s s th a n h a lf o f th e
g e n ita l c a p s u le h e ig h t, b r u s h o r g a n s d e v e lo p e d . V in c u lu m a t th e le v e l
o f v a l v a e b r o a d e r t h a n i n S. serapio, b u t n a r r o w e r t h a n i n S. megarus.
G n a th o s a r m s s le n d e r, th e te r m in a l ta p e r e d p o r tio n a fte r th e b u lb o u s
s u b te r m in a l w id e n in g s h o r t, 1 /3 o r le s s o f th e a r m le n g th . P e n is
s le n d e r, its tip d o w n -tu r n e d w ith 2 p a ir e d c o r n u ti in th e s h a ft.
Female (N=36): F W
le n g th ( b a s e R to a p e x ) = 1 2 .9 m m
( a llo ty p e ) , m e a n 1 3 .1 m m , s ta n d a r d d e v ia tio n 0 .8 m m , r a n g e
1 1 .6 – 1 4 .3 m m ( n = 1 2 ) .
Dorsal wing pattern ( F i g s . 3 # 2 d , 5 ) : Ground color s i m i l a r t o
m a le , m o r e b r o w n , w a r m e r a n d s lig h tly lig h te r o n a v e r a g e . B lu e
o v e r s c a l i n g v e r y l i g h t v i o l e t - b l u e , b a s i c a l l y s l a t e i n c o l o r , o n forewing
l a r g e l y c o n fi n e d t o t h e a r e a a r o u n d 2 A v e i n , i n t h e p o s t e r i o r b a s a l
8 6 8 6
p o s itio n o f th e C u 2– 2 A c e ll, r a r e ly in th e a n te r io r p a r t b a s a lly fr o m
a n d a r o u n d C u 1 v e in o r ig in , a n d m a x im a lly 2 /3 b a s a l p a r t o f th e 2 A
c e ll. I n o n e p a r a ty p e th e e n tir e 3 /4 o f C u 2– 2 A c e ll c o v e r e d in s la te
o v e r s c a lin g . T h e g r o u n d c o lo r s till b r o w n -g r a y , n o t s la te . M o s t
s i g n i fi c a n t l y s o m e f e m a l e s w i t h o r a n g e - r e d o v e r s c a l i n g i n t h e
p o s tm e d ia n a n d m o s tly in th e s u b m a r g in a l fo r e w in g a r e a . T h e s e s c a le s
p r e s e n t m a x im a lly fr o m M 1– M 2 to C u 2– 2 A c e lls , m o s t d e v e lo p e d in
C u 1 – C u 2 , o r m i n i m a l l y c o m p l e t e l y l a c k i n g . O n t h e hindwing, t h e
s la te o v e r s c a lin g u s u a lly m in im a l, r e d u c e d to b a s a l a r e a o f th e d is c a l
c e ll a n d C u 2– 2 A . S u b m a r g in a l m a c u le s o f s la te s c a le s m a y b e c le a r ly
d e v e lo p e d in s o m e s p e c im e n s (c e lls M 1– M 2, M 2– M 3 a n d M 3– C u 1),
s o m e tim e s p u p ille d w ith g r o u n d c o lo r s c a le s . I n o n e p a r a ty p e ,
h in d w in g s la te o v e r s c a lin g m o r e e x te n s iv e w ith tr a c e s o f s la te s c a le s
p r e s e n t in m o s t c e lls . T h e s u b m a r g in a l lin e o f s la te s c a le s o n h in d w in g
a n d p o s te r io r o f M 1 in a llo ty p e , p o s te r io r o f M 2, o r M 3 in s o m e
p a r a t y p e s , u s u a l l y w i d e r t h a n i n m a l e s . V a g u e l y d e fi n e d o r a n g e - r e d
s u b m a r g in a l m a c u le m a y b e p r e s e n t ( e .g . in a llo ty p e ) in C u 1– C u 2.
O r a n g e -r e d s c a lin g p r o x im a lly o f a n a l lo b e w e ll-d e v e lo p e d , u s u a lly
s t r o n g e r e x p r e s s e d t h a n i n m a l e s , s o m e t i m e s o v e r fl o w i n g t o t h e d i s t a l
p a r t o f C u 2– 2 A , m ir r o r in g th e h in d w in g p a tte r n . O th e r fe a tu r e s
s im ila r to m a le .
Ventral wing pattern ( F i g s . 3 # 2 d , 5 ) : S i m i l a r t o m a l e , e x c e p t t h a t
o n forewing C u 2 – 2 A c e l l m a y e x p r e s s a n o r a n g e - r e d p o s t m e d i a n
m a c u le in its a n te r io r p a r t, th is m a c u le s tr o n g ly o ffs e t p r o x im a lly
c o m p a r e d to th e C u 1– C u 2 m a c u le p o s itio n , a n d fr e q u e n tly v is ib le a s a
t r a c e o f d a r k e r s c a l e s o n l y . O n t h e hindwing, p o s t b a s a l o r a n g e
m a c u le s d e v e lo p e d e v e n le s s th a n in m a le s a n d m o s tly o b s o le te .
Head: C o l l a r w i t h f e w t o m a n y o r a n g e s c a l e s i n t h e a n t e r i o r f r i n g e
a n d th e s e e x te n d fo r w a r d in to th e e d g e o f th e in te r a n te n n a l tu ft. T h is
tu ft c e n tr a lly w h ite w ith p r o m in e n t b la c k h a ir s c a le s . E ls e w h e r e o n th e
s h o r t-s c a le d g r a y c r o w n p r o m in e n t b la c k h a ir s c a le s , n o t p r e s e n t in th e
m a le . A n te n n a e c o lo r e d a s in th e m a le . M e s ia l n a k e d m ic r o h ir s u te
a r e a m o r e e x te n s iv e , r e a c h in g 1 2 s e g m e n ts b e h in d th e a p e x , a lm o s t to
th e b a s e o f th e c lu b . C lu b e x te r n a lly s c a le d to w ith in 6 s e g m e n ts o f
th e a p e x .
Body v e s t i t u r e a s o n t h e m a l e e x c e p t b e i n g l i g h t e r a b o v e a n d t h e
la s t 2 /3 o f th e a b d o m e n m o s tly b r o w n is h g r a y . P a tte r n s o n le g s th e
s a m e a s in m a le s .
Genitalia: ( F i g . 8 ) T y p i c a l f o r t h e serapio g r o u p Strymon, b u r s a
c o p u l a t r i x l a r g e c o n t a i n i n g t w o p r o n o u n c e d “b o a t ” - s h a p e d s i g n a w i t h
a n te r io r p o in tin g s p in e s , n o t s c le r o tiz e d a r o u n d d u c tu s s e m in a lis ,
w h ic h a r is e s fr o m th e p o s te r io r tip o f b u r s a c o p u la tr ix . D u c tu s b u r s a e
a b o u t th e le n g th o f b u r s a c o p u la tr ix , r e la tiv e ly w id e , b a s a lly w ith a
n a r r o w ly tw is te d s in g le -tu r n lo o p , w ith tw o s m a ll te e th in s id e th e
a n te r io r p o r tio n . D u c tu s b u r s a e g r a d u a lly b r o a d e r to w a r d s la m e lla .
D u c tu s b u r s a e m o s tly s c le r o tiz e d , b u t s o m e p o r tio n s in th e m e d ia n
p a r t o f d u c tu s b u r s a e p o o r ly s c le r o tiz e d a n d a p p e a r a s tr a n s p a r e n t
a re a s.
Dimorphism of wing shape: t h e m a l e m o r e f e m a l e - s h a p e d i n
w in g o u tlin e th a n o th e r s im ila r s p e c ie s . S h a p e d iffe r s p r in c ip a lly in
th e s tr a ig h t to s lig h tly c o n v e x r a th e r th a n s lig h tly c o n c a v e a t C u 2
p o s te r io r 2 /3 o f th e o u te r m a r g in o f th e fo r e w in g (F ig s . 4 , 1 0 ), a n d
m o r e c o n v e x r a t h e r t h a n s t r a i g h t ( e . g . S. jacqueline) m e d i a n ( f r o m
v e in M 1 to C u 2) s e c tio n , e x c e p t o n e p o s s ib ly w in g -s h a p e a b e r r a n t d u e
to p u p a tio n p r o b le m s m a le w ith th e s tr a ig h t m a r g in , F ig . 4 # 5 . M o s t
o t h e r s p e c i e s o f Strymon m u c h m o r e d i m o r p h i c i n w i n g s h a p e .
Type specimens:
Holotype m a l e b e a r i n g t h e f o l l o w i n g l a b e l s : 3 p r i n t e d w h i t e
la b e ls : [ N 2 9 ° 1 1 .6 4 4 ' W 1 0 2 ° 5 7 .0 4 5 ' // n r B a r k e r, R d to B o q u illa s // B ig
B e n d N a t. P a r k // B r e w s te r C o . T X // 2 6 -M a r-2 0 0 5 U S A // le g . G r is h in
N .V .] , [ L e g fo r D N A // # 4 0 6 ] , [ N P S r e s e a r c h p e r m it // B I B E - 2 0 0 5 S C I - 0 0 0 6 ] , p r i n t e d r e d l a b e l [ H O L O T ×P E / / Strymon solitario m / /
G r i s h i n & D u r d e n , 2 0 1 1 ] . Allotype f e m a l e i s f r o m t h e s a m e l o c a l i t y
a n d c o lle c to r a s th e h o lo ty p e m a le , 2 3 -A p r-2 0 0 5 . A d d itio n a lly , th e r e
a r e 29 male and 35 female paratypes: 3 mm, 1 0 ff t h e s a m e l o c a l i t y ,
d a te a n d c o lle c to r a s h o lo ty p e , th e s e fe m a le s a r e in v e r y p o o r
c o n d i t i o n a s t h e y w e r e u s e d f o r o v i p o s i t i o n i n c a p t i v i t y ; 6 mm, 3 ff I b i d . ,
2 3 - A p r - 2 0 0 5 , 1 m I b i d . , 2 4 - A p r - 2 0 0 5 , 2 mm, 6 ff I b i d , 1 5 - A p r - 2 0 0 7 , 1 m,
e x o v u m , e x f, I b i d , h a t c h e d 2 0 - M a y - 2 0 0 5 , 1 f I b i d . , 1 5 - S e p - 2 0 0 7 , 1 m,
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
1 f I b i d . , 8 - M a r - 2 0 0 9 ; 1 m 1 f, T E X A S : B r e w s t e r C o . : “A m e r i e s
B o q u i l l a s ”, R i o G r a n d e R i v e r , B i g B e n d N a t i o n a l P a r k , 2 5 - S e p - 1 9 4 4 ,
l e g . G e o r g e S c h e n k ; 2 ff I b i d . 2 6 - S e p - 1 9 4 4 ; 2 mm, 2 ff I b i d . , 2 7 - S e p 1 9 4 4 ; 1 m, 2 ff I b i d . , 2 8 - S e p - 1 9 4 4 ; 1 f I b i d . , 2 9 - S e p - 1 9 4 4 ; 1 m, 1 f
I b i d . , 8 - O c t - 1 9 4 4 ; 1 m, 1 f I b i d . , 9 - O c t - 1 9 4 4 ; 1 f, T E X A S : B r e w s t e r
C o . B ig B e n d N a t. P a r k , R io G r a n d e V illa g e N a tu r e T r a il, 2 7 -M a r1 9 6 8 , l e g . R o y O . K e n d a l l & C o n n i e A . K e n d a l l ; 1 m, T E X A S :
B r e w s te r C o . B ig B e n d N a t. P a r k , O ld S a n V ic e n te c r o s s in g o n R io
G r a n d e , 1 4 - S e p - 1 9 7 1 , l e g . R o y O . K e n d a l l & C o n n i e A . K e n d a l l ; 1 m,
T E X A S : B r e w s te r C o . B ig B e n d N a t. P a r k , b e tw e e n B a r k e r H o u s e
a n d B o q u illa s C a n y o n 2 6 -M a r-1 9 8 7 , le g . R o y O . K e n d a ll & C o n n ie A .
K e n d a l l ; 2 ff I b i d . , 2 7 - M a r - 1 9 8 7 ; 1 f I b i d . , 2 8 - M a r - 1 9 8 7 ; 2 mm,
T E X A S : P r e s id io C o ., T h e S o lit a r io , G r a y 's R id g e 2 9 .4 3 ° N 1 0 3 .8 0 ° W ,
2 0 - M a y - 1 9 7 3 , l e g . C h r i s t o p h e r J D u r d e n ; 5 mm, T E X A S : B r e w s t e r C o .
T h e S o lita r io , r id g e s o u th o f T r e s P a p a lo te s 2 9 .4 4 ° N 1 0 3 .7 7 ° W , 8 - J u n 1 9 7 5 , le g . C h r is to p h e r J D u r d e n .
I n a d d itio n to th e s e s p e c im e n s , p h o to g r a p h s o f a d o z e n liv e
in d iv id u a ls fr o m th e B ig B e n d A r e a o f T e x a s a n d n e a r C iu d a d
V ic to r ia , M e x ic o : T a m a u lip a s , w e r e e x a m in e d (s e e F ig . 9 fo r im a g e s
a n d lo c a tio n s ). H o w e v e r, s in c e th e s e w e r e n o t c o lle c te d a n d th e ir
t a x o n o m i c p l a c e m e n t r e m a i n s u n c e r t a i n , t h e y a r e excluded f r o m t h e
ty p e s e r ie s .
T h e holotype a n d allotype a r e d e p o s i t e d i n t h e USNM
c o lle c tio n . P a r a ty p e s a r e d e p o s ite d in th e T A M U , M G C L , B M N H ,
U W B M , T M M C , a n d o th e r c o lle c tio n s (s e e M a te r ia ls a n d M e th o d s
fo r a b b r e v ia tio n s ).
Variation:
W h e n w in g p a tte r n s a r e s im ila r a m o n g m a n y
f r e q u e n t l y n o t v e r y c l o s e s p e c i e s ( e . g . S. ziba a n d S.
megarus) a n d g e n i t a l i a d o n o t o f f e r c l e a r - c u t
c h a r a c t e r s , a s i n m a n y Strymon t a x a , i t b e c o m e s
e s s e n tia l to s tu d y th e e x te n t o f v a r ia tio n in o r d e r to
d e fi n e t h e h i a t u s b e t w e e n s p e c i e s . W e a s s e m b l e d a
la r g e s e r ie s (c a . 7 0 s p e c im e n s ) o f th is r a r e ly
e n c o u n te r e d in s e c t th a t o ffe r s g o o d m a te r ia l fo r
v a r ia tio n s tu d ie s .
Wing pattern variation ( F i g s . 4 , 5 ) : T h e m o s t
n o ta b le v a r ia tio n is th e e x te n t o f p o s tb a s a l o r a n g e s p o ts
o n v e n tr a l h in d w in g . T h e s e s p o ts c a r r y ta x o n o m ic
i m p o r t a n c e , a s s o m e serapio g r o u p s p e c i e s ( e . g . S.
serapio) c o n s i s t e n t l y l a c k t h e s p o t s , w h i l e o t h e r s ( e . g . S.
megarus) c o n s i s t e n t l y h a v e t h e m p r e s e n t . T h e q u e s t i o n
m ig h t a r is e w h e th e r in d iv id u a ls la c k in g th e s p o ts m ig h t
b e a s p e c ie s d iffe r e n t fr o m th a t w ith th e s p o ts . I n
s e v e r a l s p e c im e n s ( e .g . h o lo ty p e ) th e s p o ts a r e
d e v e lo p e d d iffe r e n tly o n le ft a n d r ig h t w in g s , a n d s o m e
h a v e s p o ts p r e s e n t o n o n e w in g , b u t la c k th e m o n th e
o t h e r . I t i s m o r e l i k e l y t h a t i n t h i s s p e c i e s ( a s i n S.
jacqueline) , d e v e l o p m e n t o f s p o t s ( t o c o m p l e t e l a c k o f
th e m ) is v a r ia b le . O n s o m e w in g s 2 s p o ts a r e p r e s e n t,
o n o th e r s o n ly o n e to n o n e . S p o t d e v e lo p m e n t is m o r e
p r o n o u n c e d in m a le s , a n d m o s t fe m a le s c o m p le te ly
la c k a n y tr a c e o f o r a n g e s c a lin g in th e h in d w in g
p o stb a sa l a re a .
C o lo r o f o r a n g e s p o ts , m a c u le s a n d b a n d s w a s fo u n d
to b e v a r ia b le a s w e ll, w ith h u e r a n g in g fr o m o r a n g e r e d to o r a n g e -y e llo w . I t is s u s p e c te d th a t th is c o lo r
m ig h t b e a t le a s t in p a r t e n v ir o n m e n ta lly in d u c e d , a s
th e m a le s p e c im e n r e a r e d in th e la b (F ig . 4 # 1 2 v )
V
O L U M
E
6 6 , N
U M
B E R
2
p o s s e s s e s m u c h y e llo w e r, u n u s u a l b a n d s th a n w ild c a u g h t s p e c im e n s . D e v e lo p m e n t o f o r a n g e m a c u le s in
p o s tm e d ia n b a n d is a ls o v a r ia b le , a n d s o m e s p e c im e n s
h a v e th o s e b a n d s m u c h r e d u c e d ( e .g . F ig s . 4 # 1 1 v ,
5 # 1 1 v ). S o m e s p e c im e n s e x h ib it o r a n g e s c a lin g a t th e
d is ta l e d g e o f th e d a r k e r c e ll b a r o n v e n tr a l h in d w in g ,
w h ile th e m a jo r ity o f s p e c im e n s p o s s e s s o n ly d a r k a n d
lig h t e n d -o f-c e ll s c a le s . P o s tm e d ia n o r a n g e m a c u le in
C u 2– 2 A c e ll o n v e n tr a l fo r e w in g is la c k in g in m a le s , b u t
m a y b e p r e s e n t in fe m a le s . I f p r e s e n t, th is m a c u le is
b a s a lly o ffs e t c o m p a r e d to th e r o w o f m a c u le s .
I n m a le s , e x te n t o f v io le t b lu e d o r s a l o v e r s c a lin g a n d
its h u e v a r ie s fr o m b e in g v e r y u n d e r d e v e lo p e d a n d
p r e s e n t a lo n g C u 2 a n d 2 A v e in s a n d th e b a s a l a r e a o f
th e w in g ( e .g . F ig . 4 # 8 d ) , to m o r e e x te n s iv e , w ith a
s i g n i fi c a n t p o r t i o n o f f o r e w i n g c e l l C u 2 – 2 A c o v e r e d i n
b lu e s c a le s ( e .g . F ig . 4 # 7 d ) . T h e h u e v a r ie s fr o m
p u r p l i s h t o v i o l e t - b l u e , t y p i c a l l y r e d d e r t h a n t h a t i n S.
serapio, b u t a b o u t t h e s a m e c o l o r a s i n S. megarus.
V a r ia tio n in h u e o f b lu e -v io le t s tr u c tu r a l c o lo r a n d
c o p p e r t o n e s m a y b e i n fl u e n c e d b y h u m i d i t y b u t i s
r a th e r s ta b le . V a r ia tio n in p ig m e n te d g r o u n d c o lo r
fa d e s n a tu r a lly , d a y b y d a y w ith e x p o s u r e to th e s u n
d u r in g th e life o f th e in d iv id u a l a n d la te r w h e n
illu m in a te d in th e c o lle c tio n .
I n fe m a le s , in te r e s tin g v a r ia tio n is th e e x te n t o f
o r a n g e o v e r s c a lin g in th e p o s tm e d ia n a r e a o f d o r s a l
f o r e w i n g s . T h e s e o r a n g e s c a l e s , a b s e n t i n S. megarus,
b u t u s u a l l y p r e s e n t i n S. jacqueline, a r e r a t h e r w e a k l y
e x p r e s s e d in a b o u t a q u a r te r o f fe m a le s , b u t a b s e n t in
th e m a jo r ity o f th e m . S la te -b lu e d o r s a l o v e r s c a lin g is
a ls o v a r ia b le , b u t is n o t e x te n s iv e in a ll fe m a le s s e e n .
I t is n o te w o r th y , th a t p h o to g r a p h s o f M e x ic a n
s p e c im e n s (F ig . 9 # 1 2 – # 1 6 ) s h o w q u ite s im ila r w in g
p a tte r n s , b u t d iffe r in le g p a tte r n s in th a t tib ia e p o s s e s s
d a r k e r a n d m o r e c o n tr a s ty b a n d s o f s c a le s (F ig s . 6 a – f),
m o r e s i m i l a r t o t h o s e o f S. megarus, S. jacqueline a n d
S. serapio ( F i g s . 6 g – k ) t h a n s p e c i m e n s f r o m t h e B i g
B e n d a r e a (F ig s . 6 a – c , l, m ).
Male genitalic variation ( 5 d i s s e c t i o n s , F i g . 7 ) :
M o s t v a r i a t i o n i s c o n fi n e d t o t h e s a c c u s , w h i c h d i f f e r s
in s h a p e a n d th e d e g r e e to w h ic h its tip is p o in te d
(F ig s . 7 # 1 , # 2 ). H o w e v e r, th e s a c c u s is a lw a y s
a s y m m e tr ic a n d is n e v e r th a t s h o r t a n d r o u n d e d a t th e
t i p a s i n S. serapio ( F i g . 7 # 4 ) .
Female genitalic variation ( 6 d i s s e c t i o n s , F i g . 8 ) :
M in o r v a r ia tio n w a s o b s e r v e d in th e s h a p e o f th e
s c le r o tiz e d lo o p o f th e d u c tu s b u r s a e a n d in th e b e n d in
th e d u c tu s b u r s a e , a s illu s tr a te d in F ig s . 8 # 1 – # 4 .
Habitat, distribution and behavior:
Type Locality ( F i g s . 2 a , b ) and habitat: T h e t y p e
lo c a lity is U S A : T E X A S : B r e w s te r C o ., B ig B e n d
8 7
F I G . 1 . R e c o r d s o f Strymon solitario ( c i r c l e s ) a n d Strymon
megarus ( s t a r s ) f r o m t h e U n i t e d S t a t e s ( T e x a s ) a n d M e x i c o .
S ta te s w ith r e c o r d s a r e la b e le d .
N a tio n a l P a r k , c a . 2 m ile s w e s t o f B o q u illa s C a n y o n . I t
is a lim e s to n e h ill im m e d ia te ly to th e n o r th o f th e
p a v e d p a r k r o a d le a d in g to B o q u illa s C a n y o n , c a . 1 m ile
e a st o f S H 1 1 8 , n e a r B a rk e r H o u se , G P S a ro u n d
2 9 ° 1 1 '5 0 " N 1 0 2 ° 5 7 '0 6 " W , e le v a t io n 6 2 0 m . A r o a d s id e
o f f e r s a s m a ll p a r k in g s it e w it h G P S N 2 9 ° 1 1 .6 4 4 '
W 1 0 2 ° 5 7 .0 4 5 '. T h e s e G P S c o o r d in a t e s a r e lis t e d o n t h e
h o lo ty p e la b e l. G P S d a ta fo r th e B a r k e r H o u s e , w h ic h
is r e fe r e n c e d s e v e r a l tim e s th r o u g h o u t th is p a p e r is
2 9 ° 1 1 '3 1 .2 6 " N 1 0 2 ° 5 6 '3 4 .1 1 " W . T h is h is t o r ic h o u s e is
c u r r e n tly b e in g u s e d a s lo d g in g s ta tio n fo r r e s e a r c h e r s
i n t h e p a r k , b u t i s c l o s e d t o t h e p u b l i c . Hechtia texensis
p la n ts c o v e r m o s t o f th e h ill, b u t a r e p a r tic u la r ly d e n s e
a lo n g s m a ll c a n y o n s a n d g u lc h e s . I t is in th e s e s m a ll
g u lc h e s o n th e e a s te r n s id e o f th e h ill th a t th e m a jo r ity
o f Strymon s p e c i m e n s w e r e c a p t u r e d . M a l e s w e r e a l s o
fo u n d h illto p p in g a t th e h ig h e s t p o in t o f th is h ill. A lo n g
w i t h Hechtia, d r y l i m e s t o n e s l o p e s a n d b e d r o c k h a r b o r
l e c h u g u i l l a , P r i c k l y p e a r c a c t u s , O c o t i l l o ( Fouquieria
splendens E n g e l m . , F o u q u i e r i a c e a e ) , C r e o s o t e B u s h ,
Jatropha dioica a n d
a l s o Leucophyllum minus
(S c r o p h u la r ia c e a e ) a lo n g th e to p . T h is lo c a tio n c a n b e
8 8 8 8
F I G . 2 . T y p i c a l h a b i t a t a n d l a r v a l h o s t p l a n t o f Strymon solitario. T e
r o a d to B o q u illa s C a n y o n , c a . 1 m i E o f S H 1 1 8 , n e a r B a r k e r H o u s e , G
o f t h e h i l l s l o p e w i t h Hechtia texensis p l a n t s , v i e w t o w a r d s n o r t h - w e s
h a ir s tr e a k im a g e ta k e n a t th is lo c a tio n is p a s te d in to s ig n ify th e e v e
l e c h u g u i l l a a n d O p u n t i a ( b e h i n d ) . Hechtia texensis: c) a c l u s t e r o f p l a
a fl o w e r s t a l k w i t h fl o w e r b u d s , g) m a l e a n d h) f e m a l e fl o w e r s . T h e i m
M a r c h -2 0 0 5 ) a n d d ), e ) a n d h ) p h o to g r a p h e d o n 2 3 -A p r il-2 0 0 5 .
J
O U R N A L
x a s : B r e w s te r C o .,
P S c a . 2 9 ° 1 1 '5 0 " N
t . Hechtia fl o w e r s
n t . b) a c l o s e - u p o
n t s , d) a l e a f r o s e t
a g e s a r e ta k e n o n
O F
T H E
B ig B e n d N a
1 0 2 ° 5 7 '0 6 " W
ta lk s a r e v is ib
f th e h a b ita t
t e , e) p a r t o f
2 6 -M a r c h -2 0
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tio
, e
le
s h
E P I D O P T E R I S T S
’ S
O C I E T ×
n a l P a r k , n o r th o f th e p a r k
l e v a t i o n 6 2 0 m . a) o v e r v i e w
a s g r a y -b r o w n b lo tc h e s . A
o w i n g Hechtia m i x e d w i t h
a m a l e fl o w e r s t a l k , f) a t i p o f
0 5 , e x c e p t th e h a ir s tr e a k (2 7 -
V
O L U M
F ig .
q u illa s
lo w e a
2. A l l o
E
6 6 , N
3 . T h e
C a n y o
c h s p e c
t y p e f,
ty
n ,
im
2 3
U M
B E R
p e
c a .
e n
-A
s p e c im
1 m i E
1. H o
p r il-2 0
2
8 9
e n
o f
lo ty
0 5 .
s o f Strymon solitario. T e
S H 1 1 8 , n r. B a r k e r H o u s e .
p e m, 2 6 - M a r c h - 2 0 0 5 , e n l a
L i v e i m a g e o n F i g . 9#3. L
x a s :
da n
r g e d
e g p
B r e w
d vd
la te
a tte r
r i c h i n M a l v a c e a e p l a n t s d u r i n g w e t s e a s o n , a n d Celotes
limpia B u r n s , 1 9 7 4 w i t h Systasea zampa ( W . H .
E d w a r d s , 1 8 7 6 ) c a n b e q u ite a b u n d a n t. A s fo r o th e r
Strymon s p e c i e s , Strymon melinus o c c a s i o n a l l y o c c u r s
h e re .
I n a d d itio n to th is h ill, w h ic h s im p ly o ffe r e d a g o o d
p a r k i n g s i t e a n d c o n v e n i e n t c o l l e c t i n g a c c e s s , Strymon
s p e c im e n s w e r e fo u n d a t s e v e r a l lo c a tio n s in th e g e n e r a l
B o q u illa s a r e a — fr o m B o q u illa s C a n y o n in th e e a s t to
H o t S p r i n g s a n d O l d S a n V i c e n t e , e i t h e r o n fl o w e r s , e . g .
a lo n g th e r o a d s id e , o r in g u lc h e s a m o n g th e la r v a l
fo o d p la n ts .
s te
e n
r a l
n F
r C o
o te d
v ie w
ig s .
., B
o r s
fr o
6c,
ig B e n d N a tio n a l P a r k , n o r th o f th e p a r k r o a d to B o a l a n d v e n tr a l v ie w s , r e s p e c tiv e ly ; la b e ls a r e s h o w n b e m th e le ft o n th e in s itu g e n ita lia , s a c c u s is n o t v is ib le .
l
I n th e S o lita r io r a n g e (o n th e lin e b e tw e e n P r e s id io
a n d B r e w s te r C o u n tie s , c a . 5 0 m ile s w e s t fr o m th e ty p e
lo c a lity ), o p e n d e s e r t r o c k y s lo p e s w ith le c h e g u illa a n d
Hechtia texensis a r e b r o k e n b y g u l c h e s h a r b o r i n g r e l i c t
w o o d l a n d s c r u b o f Quercus pungens L i e b m . , Q.
vaseyana B u c k l e y ( F a g a c e a e ) a n d h i l l t o p s c r u b w i t h
Prunus havardii ( W . W i g h t ) S . C . M a s o n ( R o s a c e a e ) .
H i l l t o p p i n g m a l e s w e r e c o l l e c t e d a t fl o w e r s o f Acacia
greggii A . G r a y ( F a b a c e a e ) .
Distribution ( F i g . 1 ) : W h i l e i t i s l i k e l y t h a t m a n y
l i m e s t o n e s l o p e s a n d b e d r o c k w i t h Hechtia a c r o s s w e s t
T e x a s a n d n o r th -c e n tr a l M e x ic o s u p p o r t p o p u la tio n o f
9 0 9 0
t h i s b u t t e r fl y , t h e i n s e c t i s c u r r e n t l y k n o w n f r o m o n l y a
h a n d fu l o f s ite s . I n a d d itio n to th e B o q u illa s a r e a in B ig
B e n d N a tio n a l P a r k , B r e w s te r C o . T E X A S , it h a s b e e n
r e c o r d e d in s im ila r h a b ita t 1 ) c a . 2 0 a ir m ile s n o r th ,
a b o u t 3 m ile s fr o m th e R io G r a n d e r iv e r ; 2 ) c a . 1 7 a ir
m ile s w e s t, fr o m th e g r a s s y b r u s h la n d s , n e a r th e m o u th
o f P in e C a n y o n , C h is o s M o u n ta in s (p h o to g r a p h , F ig .
9 # 1 , Hechtia i s r e c o r d e d f r o m t h e n e a r b y m o u n t a i n s ,
e .g . N u g e n t M o u n ta in ) ; 3 ) c a . 5 0 m ile s w e s t in th e
S o lita r io r a n g e , a t th e b o u n d a r y o f P r e s id io a n d
B r e w s te r C o u n tie s (r e c o r d e d fr o m b o th c o u n tie s ).
M o s t s u r p r is in g ly , p h o to g r a p h s o f s e v e r a l h a ir s tr e a k
s p e c im e n s fr o m M E X I C O : T a m a u lip a s , n o r th w e s t o f
C iu d a d V ic t o r ia , a lo n g L o s T r o n c o n e s C a n y o n , t a k e n
1 8 - a n d 2 7 -N o v -2 0 0 4 , b y K im
D a v is a n d M ik e
S ta n g e la n d (F ig s . 9 # 1 2 – # 1 4 , 1 0 # 8 ); a n d 8 m ile s s o u th s o u th w e s t o f C iu d a d V ic to r ia , a lo n g B a lc ó n d e
M o c t e z u m a R o a d ( 2 3 ° 3 5 '5 2 .8 4 " N
9 9 ° 1 2 '3 1 .3 2 " W ) ,
ta k e n 6 -N o v -2 0 0 7 b y D a n H a r d y r e v e a le d a v e r y c lo s e
m a t c h t o Strymon s p e c i m e n s f r o m t h e B i g B e n d a r e a .
H a r d y a l s o p h o t o g r a p h e d Hechtia sp. i n t h e i m m e d i a t e
p r o x im ity o f h is h a ir s tr e a k s ig h tin g .
W e te n ta tiv e ly p la c e th e s e p h o to g r a p h e d in d iv id u a ls
w ith th is n e w s p e c ie s , h o w e v e r, s in c e n o s p e c im e n s
w e r e c o lle c te d fo r m o r e d e ta ile d a n a ly s is , a n d le g
p a tte r n o f p h o to g r a p h e d in s e c ts d iffe r s s o m e w h a t fr o m
t h e u n i q u e p a t t e r n o f t h e B i g B e n d Strymon ( F i g s . 6 a – c
v s . 6 d – f ) , t h i s p l a c e m e n t a w a i t s c o n fi r m a t i o n .
N e v e r t h e l e s s , c o m p a r e d t o S. megarus ( F i g s . 1 0 # 9 – # 1 2 ,
# 1 4 – # 1 7 ) , C i u d a d V i c t o r i a Strymon a p p e a r m u c h c l o s e r
to th e B ig B e n d p o p u la tio n s b y th e w in g p a tte r n .
Phenology: O v e r t h e y e a r s , s p e c i m e n s i n t h e B i g
B e n d a re a h a v e b e e n re c o rd e d fro m F e b ru a ry to Ju n e
a n d fr o m S e p te m b e r to N o v e m b e r w ith m a jo r ity o f
r e c o r d s b e in g in la te M a r c h – A p r il a n d th e n S e p te m b e r,
w ith e a r ly a n d la te d a te s b e in g 1 8 -F e b -2 0 0 5 a n d 9 -O c t1 9 4 4 . N o r e c o r d s a r e a v a ila b le fo r J u ly , A u g u s t,
D e c e m b e r a n d
J a n u a r y . T a m a u lip a s (M E X I C O )
p h o to g r a p h ic r e c o r d s a r e o n ly fr o m N o v e m b e r, b u t th is
i s l a r g e l y b e c a u s e m o s t b u t t e r fl i e r s v i s i t t h e s e l o c a t i o n s
la te in th e fa ll. S in c e in th e la b d e v e lo p m e n t o f
im m a tu r e s w a s d ir e c t, it is p o s s ib le th a t in w a r m e r a r e a s
o f B ig B e n d a d u lts c o u ld b e e n c o u n te r e d th r o u g h o u t
th e y e a r. H o w e v e r, d u r in g v e r y d r y y e a r s , a s in 2 0 0 6 ,
e x te n s iv e s e a r c h e s fo r a d u lts in s p r in g w e r e n o t
s u c c e s s fu l. T h e r e fo r e it is lik e ly th a t th e in s e c t s ta y s
q u ie s c e n t (d ia p a u s e , a e s tiv a tio n ) a t o n e o r m o r e its life
s ta g e s to w ith s ta n d h a r d s h ip o f u n s u ita b le w e a th e r a n d
t h e l a c k o f b l o o m i n g Hechtia t o b e u s e d a s l a r v a l h o s t
p la n ts .
Adult behavior: A d u l t s w e r e o b s e r v e d f e e d i n g o n
fl o w e r s o f t h e f o l l o w i n g p l a n t s : Hechtia texensis ( 1 5 - A p r 2 0 0 5 , B o q u i l l a s a r e a , N V G ) , Chromolaena odorata ( L . )
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
K in g &
H . R o b . ( A s te r a c e a e ) ( 1 8 ,2 7 - N o v - 2 0 0 4 ,
M X : T A M , K i m D a v i s & M i k e S t a n g e l a n d ) , Acacia
greggii ( 2 0 - M a y - 1 9 7 3 , 8 - J u n - 1 9 7 5 , t h e S o l i t a r i o , C J D ) ,
Larrea tridentata ( F i g s . 9 # 7 – # 1 0 , 7 - M a r - 2 0 0 9 ,
B o q u i l l a s a r e a , M a r t i n R e i d ) , Giliastrum rigidulum
( B e n th .)
R y d b .
(P o le m o n ia c e a e )
(2 6 -M a r-1 9 8 7 ,
B o q u i l l a s a r e a , K e n d a l l ) , Sphaeralcea angustifolia ( C a v . )
G . D o n (M a lv a c e a e ) (2 8 -M a r-1 9 8 7 , B o q u illa s a r e a ,
K e n d a l l ) . I n a d d i t i o n t o fl o w e r s , a d u l t s c o l l e c t m o i s t u r e
fr o m s to n e s (F ig . 9 # 3 ), le a v e s , a n d c a c ti (F ig . 9 # 1 ,
p r o b a b l y e x t r a fl o r a l n e c t a r ) , i n a m a n n e r s i m i l a r t o t h a t
d e s c r i b e d ( V i l a & E a s t w o o d 2 0 0 6 ) f o r S. jacqueline.
M a le s h illto p a n d e x h ib it te r r ito r ia l b e h a v io r, i.e . e n g a g e
in a tta c k s o n e a c h o th e r, r e tu r n in g to th e p r io r p e r c h
a fte r w a r d s . F e m a le s , u n le s s th e y m o v e a r o u n d to fe e d
o n fl o w e r s , s t a y c l o s e r t o t h e g r o u n d i n t h e p r o x i m i t y o f
la r v a l fo o d p la n ts.
Larval food plants: A s e s t a b l i s h e d b e l o w ( l i f e
h i s t o r y s e c t i o n ) , fl o w e r b u d s , fl o w e r s a n d f r u i t s o f
Hechtia texensis ( B r o m e l i a c e a e ) s e r v e a v i a b l e l a r v a l
f o o d s o u r c e ( F i g s . 2 b – h ) . T w o Hechtia s p e c i e s a r e
r e c o g n i z e d i n T e x a s ( U S D A , N R C S 2 0 1 1 ) : H. texensis –
F a l s e A g a v e ( i n c l u d i n g H. scariosa L . B . S m i t h – R o u g h
F a ls e -A g a v e ) in P r e s id io , B r e w s te r C o s (a n d a ls o in
M e x ic o : C h ih u a h u a , C o a h u ila ), h a s s e p a ls a c u te , w h ite
o r s c a r i o u s , t o 6 m m ; fl o w e r s 8 – 1 0 m m l o n g , l e a f m a r g i n s
s c a r c e l y r e p a n d , a l l t h e i r s p i n e s r e m o t e . H. glomerata G u a p illa , in S ta r r a n d Z a p a ta C o s . (a n d a ls o in M e x ic o :
T a m a u lip a s , H id a lg o , Q u e r e ta r o ), h a s s e p a ls o b tu s e o r
a p ic u la te , b r o w n , 4 m m ; le a v e s r e p a n d -s e r r a te to w a r d
b a s e w ith s p in e s r e la tiv e ly c lo s e . A n a d d itio n a l m o r e
r o b u s t s p e c i e s Hechtia stenopetala K l o t z c h – P a p a l o m é
o r G u a p ila is fr e q u e n t o n d r y lim e s to n e c liffs a n d k a r s t
in th e S ie r r a M a d r e O r ie n ta l fr o m
T a m a u lip a s
s o u t h w a r d . Hechtia p l a n t s l o o k s o m e w h a t l i k e s m a l l
a g a v e p la n ts , fo r w h ic h it g o t its E n g lis h n a m e fa ls e
a g a v e . Hechtia m o s t l y r e p r o d u c e s b y o f f s h o o t s w i t h i n
th e c lu m p o f p la n ts . T h e c lu m p s ta r ts fr o m a s e e d a n d
s in g le le a f r o s e tte s w ith o u t c lu m p s a r e r e a d ily o b s e r v e d .
F l o w e r i n g i s m o s t l y c o n fi n e d t o A p r i l , b u t f o l l o w i n g f a l l
r a i n s s o m e fl o w e r s t a l k s a p p e a r a s w e l l . A c c o r d i n g t o
G r e e n ( 1 9 7 3 ) , i n B i g B e n d N a t i o n a l P a r k Hechtia
texensis g r o w s p r i m a r i l y o n t h e D e a d H o r s e M o u n t a i n s ,
M a r is c a l M o u n ta in a n d in M a r is c a l, B o q u illa s a n d S a n ta
E le n a C a n y o n s . S e c o n d a r ily , th e p la n ts e x is t in th e
H a n n o ld H ill a r e a , e a s t a n d s o u th N u g e n t M o u n ta in ,
s o u th C h ilic o ta l M o u n ta in , u p th e J u n ip e r C a n y o n r o a d
a n d n e a r G le n n S p r in g s . O n th e w e s t s id e o f th e C h is o s
m o u n ta in s , p la n ts w e r e fo u n d o n G o a t M o u n ta in , T r a p
M o u n ta in , M u le E a r s P e a k a n d B o r th C a s to lo n P e a k .
Hechtia a l s o g r o w s i n t h e L i m e s t o n e H i l l s a l o n g t h e R i o
G r a n d e fr o m M a r is c a l M o u n ta in to B o q u illa s a n d
u p s tr e a m to F r e s n o C r e e k a n d in to T h e S o lita r io .
V
O L U M
E
6 6 , N
U M
B E R
2
Rearing studies and life history:
T h e h i n t t h a t Hechtia m i g h t b e a l a r v a l h o s t p l a n t
w a s g iv e n in 1 9 7 8 , w h e n H a r r y K . C le n c h c a lle d
s p e c im e n s c o lle c te d b y K e n d a ll in th e B o q u illa s a r e a o f
B i g B e n d N a t i o n a l P a r k Strymon serapio. S i n c e serapio
u s e s B r o m e l i a c e a e , a n d Hechtia texensis i s a n a b u n d a n t
m e m b e r o f th is fa m ily a r o u n d B o q u illa s , th e fo o d p la n t
h y p o t h e s is w a s f o r m e d . K e n d a ll's 1 9 8 7 e x p e r im e n t s
w i t h 3 f e m a l e s c o n fi n e d w i t h fl o w e r b u d s o f Hechtia
fa ile d to in d u c e o v ip o s itio n (K e n d a ll &
K e n d a ll
c o lle c tio n a r c h iv e , T A M U ). H o w e v e r, in 1 9 9 2 K e n d a ll
s u c c e e d e d i n fi n d i n g a s i n g l e c a t e r p i l l a r f e e d i n g o n
Hechtia texensis fl o w e r b u d s , r o a d t o B o q u i l l a s C a n y o n
o v e r lo o k a c r o s s r o a d fr o m th e B a r k e r H o u s e (B B N P ).
T h a t la r v a p u p a te d o n A p r il 1 3 , 1 9 9 2 a n d a p a r a s ito id
Metadontia amoena ( S a y , 1 8 3 6 ) ( H y m e n o p t e r a :
C h a lc id id a e ) h a tc h e d fr o m th e p u p a o n M a y 9 1 9 9 2 .
T h is p a r a s ito id is k n o w n to u s e a w id e r a n g e o f
T h e c l i n a e a s h o s t s , i n c l u d i n g Strymon melinus ( p e r
K e n d a ll c o lle c tio n a r c h iv e ).
T o f o l l o w u p t h e Hechtia h y p o t h e s i s , a n d b e i n g
p r e p a r e d f o r t h e d i f fi c u l t i e s e x p e r i e n c e d b y K e n d a l l , 1 0
fe m a le s c a u g h t o n M a r c h 2 8 , 2 0 0 5 n e a r th e B a r k e r
H o u s e (T E X A S : B r e w s te r C o . B ig B e n d N a t. P a r k ),
w e r e c o n fi n e d w i t h f r e s h fl o w e r s t a l k s , fl o w e r b u d s a n d
fl o w e r s o f Hechtia. 5 0 0 m l t i g h t l y c l o s e d g l a s s j a r s w e r e
u s e d . T h e fi r s t j a r c o n t a i n e d 5 f e m a l e s , t h e s e c o n d - 2
fe m a le s a n d th r e e ja r s o n e fe m a le e a c h . J a r s w e r e
e x p o s e d to n a tu r a l lig h t a t a ll tim e s a n d w e r e p la c e d
u n d e r 6 0 W
r e g u la r lig h t-b u lb fo r 5 h o u r s e v e r y
e v e n in g . O n e o f th e fe m a le s in its o w n ja r e x p ir e d th e
n e x t d a y (p o s s ib ly o v e r h e a te d ), a t le a s t 2 fe m a le s in a 5 fe m a le ja r a n d 2 fe m a le s in th e ir o w n ja r s la id s e v e r a l
e g g s e a c h o n M a r c h 2 9 th u p o n b e in g e x p o s e d to th e
lig h t a n d h e a t fr o m th e lig h t b u lb , a n d a fte r b e in g fe d
w ith d ilu te d h o n e y s o lu tio n th e n ig h t b e fo r e (F ig . 1 1 a ).
O v a w e r e p l a c e d o n s t a l k s , b e l o w fl o w e r b u d s , a t
b r a n c h i n g p o i n t s , o n c l u s t e r s o f j u s t f o r m i n g fl o w e r
b u d s , a n d 3 o v a o n th e g la s s o f th e ja r. A s in m a n y
h a i r s t r e a k s ( e . g . Satyrium S c u d d e r , 1 8 7 6 ) , p o s i t i o n f o r
e a c h o v u m
w a s c a r e fu lly s e a r c h e d fo r w ith th e
o v ip o s ito r, th is s e a r c h w a s ta k in g a t tim e s u p to 1 – 2
m in u te s b e fo r e th e p r o p e r s ite w a s s e le c te d a n d a n
o v u m
e je c te d a n d g lu e d to th e s ite . W h ile th is
o v ip o s itio n b e h a v io r is ty p ic a l fo r h a ir s tr e a k s w ith
o v e r w in te r in g o v a , w h ic h m a k e s s e n s e a s th e e g g n e e d s
to s ta y p u t fo r s e v e r a l m o n th s u n d e r a ll w e a th e r
c o n d i t i o n , Strymon o v a d e v e l o p e d w i t h o u t d e l a y .
O v ip o s itio n c o n tin u e d fo r th r e e d a y s (M a r c h 2 9 , 3 0 ,
3 1 , 2 0 0 5 ) a n d th e n s to p p e d (1 5 o v a to ta l), w h ile
fe m a le s s u r v iv e d fo r a b o u t 2 w e e k s . I t is p o s s ib le th a t
fo r n o r m a l m a tu r a tio n o f o v a in s id e fe m a le s d iffe r e n t
c o n d itio n s /fo o d
so u rc e
a re
n e e d e d , o r m a y b e
9 1
l a b o r a t o r y c o n d i t i o n s w e r e n o t s u f fi c i e n t t o i n d u c e
o v ip o s itio n , a n d o n ly e x c e s s e g g s w e r e la id d u r in g th e
fi r s t f e w d a y s . F e m a l e s w e r e f e d e v e r y o t h e r d a y a n d
ja r s v e n t ila t e d a n d c le a n e d d u r in g t h is t im e . N e w p la n t
s e g m e n ts w e r e u s e d to r e p la c e d e te r io r a te d o n e s a n d
o v a w e r e ta k e n o u t a n d s to r e d in 1 0 0 m l g la s s ja r s
(G e r b e r b a b y -fo o d ja r ).
A ll o v a h a tc h e d o n A p r il 5 , 6 , a n d 7 . 1 s t in s ta r la r v a e
w e r e v e r y a c tiv e a n d , p o s s ib ly b e c a u s e th e e g g s w e r e
la r g e , c o n ta in in g e n o u g h n u tr ie n ts , c o u ld s p e n d u p to
1 2 h o u r s c r a w lin g a r o u n d b e fo r e s ta r tin g to fe e d . S o m e
l a r v a e h a d t o b e p u t i n s i d e o p e n i n g fl o w e r s a n d t h e i r
w a y o u t b lo c k e d b y th e p e ta ls b e fo r e th e y s h o w e d a n y
a tte m p t a t fe e d in g . N e v e r th e le s s a ll h a tc h e d la r v a e
s ta r te d fe e d in g a n d n o n e p e r is h e d b e fo r e A p r il 1 0 .
T h r e e la r v a e e x p ir e d d u e to m o ld a c c u m u la tin g o n th e
fl o w e r s o n A p r i l 1 0 t h .
C a t e r p i l l a r s f e d o n b u d s a n d fl o w e r s , b o t h w h i t e a n d
g r e e n p a r t s o f t h e fl o w e r . S m a l l c a t e r p i l l a r s c a n b u r y
th e m s e lv e s c o m p le te ly in s id e th e g r e e n s te m o f th e
fl o w e r , w h e n t h e y a r e f e e d i n g o n t h e b o t t o m p a r t o f t h e
fl o w e r . T h i s b e h a v i o r i s i n t e r e s t i n g , a s a c c o r d i n g t o
J a n z e n & H a l l w a c h s d a t a b a s e ( 2 0 1 1 ) , Strymon megarus
la r v a e a r e le a f m in e r s in C o s ta R ic a , e v e n in th e la s t
in s ta r (s e e F ig . 1 0 # 1 5 ).
C a te r p illa r s p r o d u c e d y e llo w -b r o w n is h s e m i-liq u id
fr a s s w h e n fe d o n p e ta ls , a n d g r e e n g r a in y fr a s s w h e n
fe d o n g r e e n p a r ts , G r a in y fr a s s p a r tic le s d o n o t
s e p a r a te fr o m e a c h o th e r a n d fo r m in te r e s tin g c h a in s o f
f r a s s g r a i n s ( F i g . 1 1 f ) , s o m e w h a t l i k e Polygonia
H ü b n e r, [1 8 1 9 ] (N y m p h a lid a e ) e g g s d e p o s ite d o n to p
o f e a c h o th e r. S o m e la r v a e c a n b e s u r r o u n d e d b y p la n t
ju ic e s a n d a p p e a r to b e c o m p le te ly im m e r s e d in liq u id .
T h is , la s tin g fo r a b o u t a d a y b e fo r e th e liq u id d r ie s o u t
o r is a b s o r b e d , d o e s n o t s e e m to h a r m th e la r v a e . T h is
is a ls o c o n s is te n t w ith th e o b se r v a tio n th a t la r v a e o f a
c l o s e r e l a t i v e S. megarus a r e l e a f m i n e r s .
T h e l a r v a e w e r e t r a n s f e r r e d o n f r e s h fl o w e r b r a n c h e s
a t le a s t e v e r y o th e r d a y to a v o id m o ld o n th e ir fo o d
s o u r c e . S o m e fl o w e r b u d s w e r e n i c k e d b y s c i s s o r s t o
s tim u la te la r v a l fe e d in g , a n d la r v a e m o s t fr e q u e n tly
s ta r te d fe e d in g a t th e p la c e s o f th e n ic k . H o w e v e r,
s o m e l a r v a e s t a r t e d t o b o r e n e a r t h e b a s e o f a fl o w e r ,
fo r m in g c a v itie s s im ila r to th e o n e s h o w n o n F ig . 1 1 e .
T h is is lik e ly to b e th e ir n a tu r a l b e h a v io r.
D u e t o i n s u f fi c i e n t n u m b e r s o f l a r v a e a l l a t t e m p t s
h a v e b e e n m a d e to p re se rv e th e m ra th e r th a n to
e x p e r im e n t w ith fo o d c o n d itio n s a n d fe e d in g b e h a v io r.
A ll o f th is m a y h a v e a ffe c te d th e la r v a l b e h a v io r m a k in g
it u n n a tu r a l. D e s p ite th e s e a tte m p ts , th e m a jo r ity o f
h a tc h e d la r v a e d ie d e a r ly o n , n o t r e a c h in g th e 3 r d
in s ta r. H ig h e r th a n n a tiv e le v e ls o f h u m id ity a r e
s u s p e c te d a s th e c a u s e , a s 1 ) r e la tiv e h u m id ity is
9 2 9 2
F IG . 4 . W in
n o r th o f th e p
w in g s u r fa c e s
6. 1 5 - A p r i l - 2
h o lo ty p e , a ls o
J
g p a
a r k
a r e
0 0 7
s h o
tte r n v a r ia
r o a d to B o
s h o w n fo r
; 3, 4, 10,
w n o n F ig
tio n in
q u illa s
e a c h s
11. 2 3
. 3#1.
Strymon solitario m
C a n y o n , c a . 1 m i E
p e c im e n . 1 – 1 1 a r e
- A p r i l - 2 0 0 5 ; 5. 2 4 -
a le s . A ll s p e c im
o f S H 1 1 8 , n e a r
fi e l d - c a u g h t s p e
A p r i l - 2 0 0 5 ; 12.
e n s a r e
B a r k e r
c im e n s ,
R e a r e d
O U R N A L
fr o m T e x a s : B
H o u s e , le g . N
d a te s a r e a s f
e x o v u m in th
O F
T H E
r e w s t
.V . G r
o llo w s
e la b ,
L
e r C
is h in
: 1,
h a tc
E P I D O P T E R I S T S
o .,
. D
7,
h e
B ig B
o r s a
8, 9.
d 2 0 -
e
l (
2
M
n d N a t
d) a n d
6 -M a r c
a y -2 0 0
’ S
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io n a l P a r k ,
v e n t r a l ( v)
h - 2 0 0 5 ; 2,
5 . 1. i s t h e
V
O L U M
F
I G
E
6 6 , N
U M
. 4 . ( c o n t.) W
B E R
2
i n g p a t t e r n v a r i a t i o n i n Strymon solitario m a l e s
9 3
9 4 9 4
F IG . 5 .
P a r k , n o r
t r a l ( v) w
9. 2 3 - A p
J
W in g p a tte
th o f th e p a
in g s u r fa c e s
r i l - 2 0 0 5 ; 5.
r n
r k
a
1
v a r ia
r o a d
r e s h o
5 -S e p
tio n
to B
w n
te m
in
o q
fo r
b e
Strymon solitario
u illa s C a n y o n , c a . 1
e a c h s p e c im e n . T h
r - 2 0 0 7 ; 12. 8 - M a r c
fe m a le s . A ll s p e
m i E o f S H 1 1 8 ,
e d a te s o f c a p tu r
h - 2 0 0 9 . 8. i s t h e
c im
n e a
e a r
a llo
e n s
r B a
e a s
ty p e
O U R N A L
a r e fr
r k e r H
fo llo w
, a ls o
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
o m T e x a s : B r e w s te r C o ., B ig B e n d N a tio n a l
o u s e , l e g . N . V . G r i s h i n . D o r s a l ( d) a n d v e n s : 1, 2, 4, 6, 10, 11. 1 5 - A p r i l - 2 0 0 7 ; 3, 7, 8,
s h o w n o n F i g s . 3#2, 6c, l a n d 9#3.
V
O L U M
F
I G
E
6 6 , N
U M
. 5 . ( c o n t.) W
B E R
2
i n g p a t t e r n v a r i a t i o n i n Strymon solitario f e m a l e s .
9 5
9 6 9 6
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
F I G . 6 . L e g p a t t e r n s o f B r o m e l i a d - f e e d i n g Strymon. M i d l e g i n l a t e r a l v i e w i s s h o w n o n a l l i m a g e s . a) , b) , a n d c) S. solitario,
T e x a s : B r e w s te r C o ., B ig B e n d N a tio n a l P a r k , n o r th o f th e p a r k r o a d to B o q u illa s C a n y o n , c a . 1 m i E o f S H 1 1 8 , n e a r B a r k e r H o u s e ,
m 2 7 - M a r c h - 2 0 0 5 , f 1 5 - A p r i l - 2 0 0 7 a n d f 2 3 - A p r i l - 2 0 0 5 , c l o s e - u p o f l i v e s p e c i m e n s s h o w n o n F i g s . 9#2, #11 a n d #3, r e s p e c t i v e l y .
d) , e) S. solitario f, m, M e x i c o : T a m a u l i p a s , N W C i u d a d V i c t o r i a , L o s T r o n c o n e s C a n y o n , 1 8 - N o v e m b e r - 2 0 0 4 , p h o t o g r a p h s b y K i m
D a v i s a n d M i k e S t a n g e l a n d , c l o s e - u p o f l i v e s p e c i m e n s s h o w n o n F i g s . 9#17 a n d #16 r e s p e c t i v e l y . f) S. solitario f, M e x i c o :
T a m a u lip a s , 8 m i S S W o f C iu d a d V ic t o r ia , B a lc ó n d e M o c t e z u m a R o a d , 2 3 ° 3 5 '5 2 .8 4 " N 9 9 ° 1 2 '3 1 .3 2 " W , 6 - N o v e m b e r - 2 0 0 7 , p h o t o
b y D a n H a r d y , c l o s e - u p o f a l i v e s p e c i m e n s h o w n o n F i g . 9#19. g) S. jacqueline m p a r a t y p e , P e r u : C a j a m a r c a , P u e n t e C h e t i l l a
0 7 ° 1 2 'S 7 8 ° 4 5 'W e le v a t io n 1 0 5 0 m , 1 7 - S e p t e m b e r - 1 9 9 9 , le g . R o b b in s , L a m a s & A h r e n h o lz . I n U S N M c o lle c t io n . C lo s e - u p o f s p e c i m e n s s h o w n o n F i g . 1 0 # 1 8 . h) S. megarus m, G u a t e m a l a : E s c u i n t l a , O c t o b e r . S c h a u s & B a r n e s c o l l e c t i o n . I n U S N M c o l l e c t i o n .
i) S. serapio m, T e x a s , H i d a l g o C o . E s t e r o L l a n o G r a n d e S t a t e P a r k , 1 9 - D e c e m b e r - 2 0 0 9 , p h o t o b y M i c h a e l A . R i c k a r d , c l o s e - u p o f
t h e l i v e s p e c i m e n s h o w n o n F i g . 9#1. j) S. serapio m, C o l o m b i a : V a l l e d e l C a u c a , R i o A n c h i c a y a , e l e v a t i o n 1 0 0 m , 1 3 - F e b r u a r y 1 9 8 2 , l e g . C . J . C a l l a g h a n . I n U S N M c o l l e c t i o n . k) S. ziba m, H o n d u r a s : A t l á n t i d a , L a C e i b a , 2 8 - D e c e m b e r - 1 9 8 2 , r e a r e d o n p i n e a p p l e , l e g J . M i r a n d a . I n U S N M c o l l e c t i o n . l) l a t e r a l v i e w o f S. solitario f a l l o t y p e , s e e F i g . 3 # 2 f o r d a t a . m) l a t e r a l v i e w o f a n e x
o v u m S. solitario m r e a r e d s p e c i m e n , s h o w n o n F i g . 4 # 1 2 , s e e F i g . 4 l e g e n d f o r d a t a .
V
O L U M
E
6 6 , N
U M
B E R
2
9 7
F I G . 7 . M a l e g e n i t a l i a o f t h e serapio g r o u p Strymon s p e c i e s . a a n d b d e n o t e l e f t l a t e r a l a n d v e n t r a l v i e w s , c i s a m a g n i fi e d l e f t
l a t e r a l v i e w o f t h e l e f t v a l v a t i p ( i n o r i e n t a t i o n s i m i l a r t o a) s h o w i n g t e e t h o n t h e v e n t r a l s u r f a c e p o i n t i n g d o w n ( t e e t h r e d u c e d i n
S. ziba) , d i s a m a g n i fi e d v e n t r a l v i e w o f t h e p e n i s t i p . 1, 2. S. solitario, T e x a s : B r e w s t e r C o . , B i g B e n d N a t i o n a l P a r k , n o r t h o f t h e
p a r k r o a d t o B o q u i l l a s C a n y o n , ~ 1 m i E o f S H 1 1 8 , n e a r B a r k e r H o u s e , l e g . N . V . G r i s h i n 1. 2 6 - M a r c h - 2 0 0 5 , G e n i t a l i a N o . N V G
# 5 1 6 . T h e s p e c i m e n i s s h o w n o n F i g . 4 # 9 ; 2. 8 - M a r c h - 2 0 0 9 , G e n i t a l i a N o . N V G # 5 2 9 . 3. S. megarus, M e x i c o : T a m a u l i p a s , R o a d
to O c a m p o ( H w y A 7 0 ) c a 1 6 k m W o f H w y 8 5 , 5 - J a n u a r y - 1 9 7 4 , le g . R .O . K e n d a ll & C .A . K e n d a ll. I n T A M U c o lle c tio n . G e n ita lia
N o . N V G # 5 1 8 . T h e s p e c i m e n i s s h o w n o n F i g . 10#11. 4. S. serapio, M e x i c o : [ T a m a u l i p a s ] , e x l a r v a 1 9 7 7 , S u e G a r d n e r , l a r v a l f o o d
p l a n t Tillandsia utriculata, f r o m R . O . K e n d a l l & C . A . K e n d a l l c o l l e c t i o n . I n T A M U c o l l e c t i o n . G e n i t a l i a N o . N V G # 5 1 7 . 5. S.
jacqueline, p a r a t y p e , P e r u : C a j a m a r c a , L a C a p i l l a , 1 0 k m W C h i l e t e 0 7 ° 1 2 ' S 7 8 ° 5 7 ' W e l e v a t i o n 7 0 0 m , 1 7 - S e p t e m b e r - 1 9 9 9 , l e g .
R o b b i n s , L a m a s & A h r e n h o l z . I n U S N M c o l l e c t i o n . G e n i t a l i a N o . 2 0 0 2 : 1 3 m R . K . R o b b i n s . 6. S. ziba, M e x i c o : T a m a u l i p a s , T a y l o r
R a n c h c a . 3 2 k m N N W o f C iu d a d M a n te , 5 - J a n u a r y - 1 9 7 4 , le g . R .O . K e n d a ll & C .A . K e n d a ll. I n T A M U c o lle c tio n . G e n ita lia N o .
N V G # 5 2 1 .
9 8 9 8
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
F I G . 8 . F e m a l e g e n i t a l i a o f t h e serapio g r o u p Strymon s p e c i e s . a a n d b d e n o t e l e f t l a t e r a l a n d v e n t r a l v i e w s , c i s a l e f t l a t e r a l v i e w
o f p a p i l l a e a n d a p o p h y s i s , d i s a m a g n i fi e d v i e w o f d u c t u s b u r s a e b a s e . O n l y l a m e l l a a n d d u c t u s b u r s a e a r e s h o w n i n 3, 4, a n d 6.
1-4. Strymon solitario, T e x a s : B r e w s t e r C o . , B i g B e n d N a t i o n a l P a r k , a l o n g t h e r o a d t o B o q u i l l a s C a n y o n , n e a r B a r k e r H o u s e , 1.
2 6 - M a r c h - 2 0 0 5 , l e g . N . V . G r i s h i n , N V G # 5 1 5 s p e c i m e n s h o w n o n F i g . 5#11; 2. 2 7 - M a r c h - 1 9 8 7 , l e g . R . O . K e n d a l l & C . A . K e n d a l l .
I n T A M U c o l l e c t i o n . G e n i t a l i a N o . N V G # 5 2 3 . T h e s p e c i m e n i s s h o w n o n F i g . 1 0 # 2 5 ; 3. 2 6 - M a r c h - 2 0 0 5 l e g . N . V . G r i s h i n . G e n i t a l i a N o . N V G # 5 1 4 ; 4. 2 8 - M a r c h - 1 9 8 7 , l e g . R . O . K e n d a l l & C . A . K e n d a l l . I n T A M U c o l l e c t i o n . G e n i t a l i a N o . N V G # 5 2 4 . 5. S.
megarus, M e x i c o : T a m a u l i p a s , P a s o d e l A b r a n e a r E l A b r a , 1 8 - D e c e m b e r - 1 9 7 3 , l e g . R . O . K e n d a l l & C . A . K e n d a l l . I n T A M U c o l l e c t i o n . N V G # 5 2 2 s p e c i m e n s h o w n o n F i g . 10#17. 6. S. megarus, M e x i c o : T a m a u l i p a s , G o m e z F a r i a s , 2 5 - A u g u s t - 2 0 0 3 . G e n i t a l i a
N o . N V G # 5 2 7 . 7. S. ziba, M e x i c o : T a m a u l i p a s , R a n c h o P i c o d e O r o v i c . o f L o s K i k o s , 9 - J a n u a r y - 1 9 7 4 , l e g . R . O . K e n d a l l & C . A .
K e n d a ll. I n T A M U c o lle c tio n . G e n ita lia N o . N V G # 5 2 1 .
V
O L U M
E
6 6 , N
U M
B E R
2
F I G . 9 . L i v e a d u l t s o f Strymon solitario. L o
T e x a s : B r e w s te r C o ., B ig B e n d N a tio n a l P a r k ( B
D e p a r tm e n t) a r e s h o w n a b o v e th e r e d lin e (im a
t o r i a ) a r e b e l o w t h e r e d l i n e ( i m a g e s 13–20) . S
t o g r a p h s . 3, 4, 10–11. T h e s e t h r e e s p e c i m e n s
t r a t e d o n F i g s . 3#2 a n d 6c, l.
9 9
c a tio n , d a te a n d p h o to g r a p h e r a r e in d ic a te d b e
B N P ) a n d B la c k G a p W ild life M a n a g e m e n t a r e
g e s 1–12) , s p e c i m e n s f r o m M e x i c o ( f r o m T a m a u
o m e s p e c i m e n s a r e i l l u s t r a t e d b y t w o ( 10–11 a
a r e s h o w n o n F i g s . 5#8, 4#5, a n d 5#11, r e s p e c
lo w e a c h im a g e . S p e c im e n s fr o m
a (W M A , T e x a s P a r k s a n d W ild life
lip a s , in th e v ic in ity o f C iu d a d V ic n d 15–16) o r t h r e e ( 18–20) p h o t i v e l y . 3. i s t h e a l l o t y p e , a l s o i l l u s -
1 0 0 1 0 0
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
F I G . 1 0 . S o m e B r o m e l i a d - f e e d i n g Strymon s p e c i e s . D o r s a l ( d) a n d v e n t r a l ( v) w i n g s u r f a c e s a r e s h o w n f o r e a c h s p e c i m e n . P r i m a r y ty p e s a r e d is p la y e d in fu ll e x p a n s e , w ith a ll th e la b e ls s h o w n . O n ly a h a lf fo r o th e r s p e c im e n s is illu s tr a te d . S p e c ie s n a m e s a n d
g e n e r a l lo c a tio n a r e in d ic a te d o n th e p la te a n d d e ta ile d h e r e . A ll p in n e d s p e c im e n s a r e to s c a le . L iv e in d iv id u a ls a r e s c a le d a p p r o x i m a t e l y t o m a t c h c o r r e s p o n d i n g s p e c i e s . 1. S. serapio m, T e x a s : H i d a l g o C o . E s t e r o L l a n o G r a n d e S t a t e P a r k , 1 9 - D e c e m b e r 2 0 0 8 , fi r s t U S r e c o r d , p h o t o g r a p h e d i n n a t u r a l c o n d i t i o n s , p h o t o g r a p h s b y M i c h a e l A . R i c k a r d . 2. S. serapio m, s y n t y p e . P a n a m a :
C h ir iq u í, D a v id , le g . C h a m p io n , G o d m a n & S a lv in c o lle c tio n 1 9 1 2 – 1 9 2 3 . I n B M N H c o lle c tio n , p h o to g r a p h c o u r te s y o f B M N H .
3. S. serapio m, M e x i c o : V e r a c r u z , O r i z a b a . P o s s i b l y W m . S c h a u s c o l l e c t i o n . I n U S N M c o l l e c t i o n . 4. S. serapio m, M e x i c o :
T a m a u l i p a s , V i l l a G o m e z F a r i a s , e l e v a t i o n 5 0 0 m , 2 8 - D e c e m b e r - 1 9 7 2 , l e g . W . W . M c G u i r e . I n U S N M c o l l e c t i o n . 5. S. serapio m,
C o s t a R i c a : C a r t a g o , J u a n V i n a s , N o v e m b e r , P o s s i b l y W m . S c h a u s c o l l e c t i o n . I n U S N M c o l l e c t i o n . 6. S. serapio m, P a n a m a :
C h ir iq u í, B u g a b a , J u ly , W m . S c h a u s c o lle c tio n . I n U S N M c o lle c tio n .
V
O L U M
E
6 6 , N
U M
B E R
2
1 0 1
F I G . 1 0 . ( c o n t . ) S o m e B r o m e l i a d - f e e d i n g Strymon s p e c i e s . D o r s a l ( d) a n d v e n t r a l ( v) w i n g s u r f a c e s a r e s h o w n f o r e a c h s p e c i m e n .
P r im a r y ty p e s a r e d is p la y e d in fu ll e x p a n s e , w ith a ll th e la b e ls s h o w n . O n ly a h a lf fo r o th e r s p e c im e n s is illu s tr a te d . S p e c ie s n a m e s a n d
g e n e r a l lo c a tio n a r e in d ic a te d o n th e p la te a n d d e ta ile d h e r e . A ll p in n e d s p e c im e n s a r e to s c a le . L iv e in d iv id u a ls a r e s c a le d a p p r o x im a t e l y t o m a t c h c o r r e s p o n d i n g s p e c i e s . 7. S. ziba m, M e x i c o : T a m a u l i p a s , V i l l a G o m e z F a r i a s , 1 4 - J u l y - 1 9 7 3 , l e g . W . W . M c G u i r e . I n
U S N M c o l l e c t i o n . 8. S. solitario m, l i v e i n d i v i d u a l , M e x i c o : T a m a u l i p a s , N W C i u d a d V i c t o r i a , L o s T r o n c o n e s C a n y o n , 2 7 - N o v e m b e r 2 0 0 4 , p h o t o g r a p h b y K i m D a v i s a n d M i k e S t a n g e l a n d , t h e s a m e i n d i v i d u a l s h o w n o n F i g . 9#12, 9. S. megarus m, p o s s i b l e h o l o t y p e .
P o s s i b l y B r a z i l . L a b e l “P . m e g a r u s , G o d ” m a t c h e s G o d a r t ' s h a n d w r i t i n g . I n M N H N c o l l e c t i o n , p h o t o g r a p h b y R e n e L a h o u s s e . 10.
S. megarus m, M e x i c o : J a l i s c o , C h a m e l a , 2 9 - M a r c h - 1 9 3 9 . F r e d . H . R i n d g e c o l l e c t i o n . I n U S N M c o l l e c t i o n . 11. S. megarus m, M e x ic o : T a m a u lip a s , R o a d to O c a m p o ( H w y A 7 0 ) c a 1 6 k m W o f H w y 8 5 , 5 - J a n u a r y - 1 9 7 4 , le g . R .O . K e n d a ll & C .A . K e n d a ll. I n T A M U
c o l l e c t i o n . 12. S. megarus m, P a n a m a : C a n a l Z o n e , P a r a í s o , 5 - A u g u s t - 1 9 7 7 , l e g . G . B . S m a l l . I n U S N M c o l l e c t i o n .
1 0 2 1 0 2
F IG . 1 0 . (
S p e c ie s n a m
O a x a c a , C a
megarus f,
n a c a s te P r o
L o n g itu d e :
c o lle c tio n .
megarus f,
le c tio n .
J
c o n t . ) S o m e B r o m e l i a d - f e e d i n g Strymon s p e c i e s . D o r s a
e s a n d g e n e r a l lo c a tio n a r e in d ic a te d o n th e p la te a n d d
n d e la r ia L o x ic h a 1 5 ° 5 4 'N 9 6 ° 3 1 'W e le v a t io n 5 0 0 m , 3
M e x ic o : V e r a c r u z , T la c o ta lp a n le g . O .W . B a r r e tt. I n U S
v in c e , G u a n a c a s te N a tio n a l P a r k , J a n z e n & H a llw a c h s V
- 8 5 .6 1 8 7 1 , e x la r v a , le a f m in e r fo u n d a s la s t in s ta r fe e d in
16. S. megarus f, P a n a m a : C a n a l Z o n e , F a r f a n , 1 7 - F
M e x ic o : T a m a u lip a s , P a s o d e l A b r a n e a r E l A b r a , 1 8 -D e
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
l ( d) a n d v e n t r a l ( v) w i n g s u r f a c e s a r e s h o w n f o r e a c h s p e c i m e n .
e t a i l e d h e r e . A l l s p e c i m e n s a r e t o s c a l e . 13. S. ziba f, M e x i c o :
0 - J u n e - 1 9 7 2 , l e g . E . W e l l i n g . I n U S N M c o l l e c t i o n . 14. S.
N M c o l l e c t i o n . 15. S. megarus f, C o s t a R i c a : n o r t h e r n G u a o u c h e r S p e c im e n D a ta b a s e 9 2 - S R N P - 2 5 4 , L a titu d e : 1 0 .8 3 7 6 4
g o n Bromelia pinguin. H a t c h e d 1 3 - F e b r u a r y - 1 9 9 2 . I n U S N M
e b r u a r y - 1 9 6 3 l e g . S.S . N i c o l a y . I n U S N M c o l l e c t i o n . 17. S.
c e m b e r- 1 9 7 3 , le g . R .O . K e n d a ll & C .A . K e n d a ll. I n T A M U c o l-
V
O L U M
E
6 6 , N
U M
B E R
2
1 0 3
F I G . 1 0 . ( c o n t . ) S o m e B r o m e l i a d - f e e d i n g Strymon s p e c i e s . D o r s a l ( d) a n d v e n t r a l ( v) w i n g s u r f a c e s a r e s h o w n f o r e a c h s p e c i m e n . S p e c i e s
n a m e s a n d g e n e r a l l o c a t i o n a r e i n d i c a t e d o n t h e p l a t e a n d d e t a i l e d h e r e . A l l s p e c i m e n s a r e t o s c a l e . 18, 19, 21. S. jacqueline m, f a n d f,
p a r a t y p e s , P e r u : C a ja m a r c a , P u e n t e C h e t illa 0 7 ° 1 2 'S , 7 8 ° 4 5 'W e le v a t io n 1 0 5 0 m , 1 7 - S e p t e m b e r - 1 9 9 9 , le g . R o b b in s , L a m a s & A h r e n h o lz . I n
U S N M c o l l e c t i o n . 20. S. serapio f, M e x i c o : T a m a u l i p a s , E l A b r a , 2 2 - D e c e m b e r - 1 9 7 3 , l e g . W . W . M c G u i r e . I n U S N M c o l l e c t i o n . 22. S. solitario m, T e x a s : P r e s i d i o C o . T h e S o l i t a r i o , G r a y ' s R i d g e 2 9 . 4 3 ° N 1 0 3 . 8 0 ° W , a t fl o w e r s Acacia greggii, 2 0 - M a y - 1 9 7 3 , l e g . C . J . D u r d e n , # 7 3 1 4 0 D 3 .
23. S. solitario m, T e x a s : B r e w s t e r C o . T h e S o l i t a r i o , r i d g e s o u t h o f T r e s P a p a l o t e s 2 9 . 4 4 ° N 1 0 3 . 7 7 ° W , a t fl o w e r s Acacia greggii, 8 - J u n e - 1 9 7 5 ,
l e g . C . J . D u r d e n , # 7 5 1 5 9 A 1 5 . 24. S. solitario f, T e x a s : B r e w s t e r C o . B i g B e n d N a t i o n a l P a r k , b e t w e e n B a r k e r H o u s e a n d B o q u i l l a s C a n y o n ,
2 7 - M a r c h - 1 9 8 7 , l e g . R . O . K e n d a l l & C . A . K e n d a l l . I n T A M U c o l l e c t i o n . 25. S. solitario f, T e x a s : B r e w s t e r C o . B i g B e n d N a t i o n a l P a r k , b e t w e e n B a r k e r H o u s e a n d B o q u i l l a s C a n y o n , 2 7 - M a r c h - 1 9 8 7 , l e g . R . O . & C . A . K e n d a l l . I n T A M U c o l l e c t i o n . G e n i t a l i a s h o w n o n F i g . 8#2.
1 0 4 1 0 4
F IG . 1
S p e c ie s
C o . B ig
I n U W B
J
0 . (c o
n a m e
B e n d
M c o
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
n t . ) S o m e B r o m e l i a d - f e e d i n g Strymon s p e c i e s . D o r s a l ( d) a n d v e n t r a l ( v) w i n g s u r f a c e s a r e s h o w n f o r e a c h s p e c i m e n .
s a n d g e n e r a l l o c a t i o n a r e i n d i c a t e d o n t h e p l a t e a n d d e t a i l e d h e r e . 26. a n d 27. S. solitario m a n d f, T e x a s : B r e w s t e r
N a t io n a l P a r k , “ A m e r ie s B o q u illa s , R io G r a n d e R iv e r ” , 2 9 ° 1 2 'N 1 0 2 ° 5 5 'W , 2 5 - S e p t e m b e r - 1 9 4 4 , le g . G e o r g e S c h e n k .
lle c tio n .
n a tu r a lly h ig h e r in th e D a lla s a r e a w h e r e th e
e x p e r im e n ts w e r e c o n d u c te d th a n in d r y B o q u illa s
d e s e r t; 2 ) r e a r in g w a s d o n e in c lo s e d g la s s ja r s ; 3 )
Hechtia fl o w e r b u d s d e v e l o p m o l d e a s i l y a n d l a r v a e d i d
n o t d is tin g u is h b e tw e e n fr e s h a n d s lig h tly m o ld y fo o d
s o u r c e . I t is p o s s ib le th a t o n ly fr e s h p la n t m a tte r is
d i g e s t e d p r o p e r l y . W h i l e c a t e r p i l l a r s f e d o n fl o w e r b u d s
a n d s ta lk c u ttin g s r e a d ily , th e y c o n tin u e d to d ie o ff w ith
e v e r y in s ta r, e v e n w h e n th e fo o d s o u r c e w a s r e p la c e d
d a ily . T h e fo o d p la n t m a y r e le a s e a lk a lo id s w h e n in ju r e d
a n d th e s e m a y in h ib it la r v a l h e a lth .
A l l c a t e r p i l l a r s w e r e u n u s u a l l y a c t i v e f o r fl o w e r fe e d in g h a ir s tr e a k s . T h e y fr e q u e n tly a b a n d o n e d th e
fe e d in g p o s itio n a n d c r a w le d a b o u t th e ja r, e ith e r
r e tu r n in g to th e s a m e fe e d in g s ite , o r r e s u m e d fe e d in g
a t a d iffe r e n t lo c a tio n . T h is e x tr a m o b ility m ig h t b e
c a u s e d b y i n s u f fi c i e n t f o o d q u a l i t y , o r b e a d i s p l a y o f a
n a tu r a l b e h a v io r, fo r in s ta n c e , c a te r p illa r s r e s tin g a t th e
b a s e o f a p l a n t a n d c r a w l i n g u p t h e fl o w e r s t a l k t o f e e d .
T h e l a t t e r m i g h t e x p l a i n t h e d i f fi c u l t y o f fi n d i n g
c a te r p illa r s in th e w ild . I n th e w ild m o s t fe e d in g m a y b e
n o c tu r n a l. W e c o n d u c te d s e v e r a l h o u r s e a r c h e s fo r
c a te r p illa r s o n A p r il 1 5 2 0 0 5 a n d M a y 2 1 2 0 0 5 (w ith
J a m e s P . B r o c k ) , e x p l o r i n g s e v e r a l t h o u s a n d fl o w e r
s t a l k s w i t h o u t s u c c e s s . H o w e v e r , fl o w e r d a m a g e
c o n s is te n t w ith c a te r p illa r fe e d in g in th e la b w a s
o b s e r v e d o n m a n y o c c a s io n s , in c lu d in g th e c a v itie s lik e
th e o n e s h o w n o n F ig . 1 1 e . W e w e r e n o t a b le to d e te c t
m i n e s o n Hechtia l e a v e s , s o i t i s u n c l e a r w h e t h e r t h i s
s p e c i e s c a n e n g a g e i n m i n i n g b e h a v i o r a s d o e s S.
megarus ( C a r t e r 1 9 4 9 , S a n c h e s e t a l . 1 9 8 5 , J a n z e n &
H a llw a c h s 2 0 1 1 ).
L ife c y c le fr o m o v ip o s itio n to a d u lt to o k a b o u t 5 0
d a y s a t 7 0 F . L a r v a w e n t th r o u g h 5 in s ta r s . T h is m ig h t b e
a
r e s u lt o f u n n a tu r a l r e a r in g c o n d itio n s , a s th e
o v e r w h e lm in g m a jo r ity o f L y c a e n id a e a r e k n o w n to
h a v e 4 in s ta r s w ith ju s t a h a n d fu l o f e x c e p tio n s (B a llm e r
& P r a tt 1 9 8 8 , D u a r te e t a l. 2 0 0 5 , D u a r te & R o b b in s
2 0 0 9 ) . T h e r e f o r e t h e n u m b e r o f i n s t a r s i n S. solitario
n e e d s t o b e i n v e s t i g a t e d f u r t h e r . B r i e f d e s c r i p t i o n o f S.
solitario i m m a t u r e s f o l l o w s .
Ovum ( N = 1 5 , F i g . 1 1 b ) i s 0 . 9 – 1 . 2 m m d i a m e t e r ,
d e v e lo p s 7 d a y s . U n u s u a lly la r g e fo r th e fa m ily , it is
s c u lp tu r e d
in c o n s p ic u o u s ly , w ith
a
p r o m in e n tly
d e p r e s s e d m ic r o p y le a r e a . C o lo r is lig h t-iv o r y .
1st instar ( N = 1 5 , F i g s . 1 1 c , d ) i s c a . 3 m m l o n g , t a k e s
5 d a y s , w ith th e la s t 2 4 h o u r s p r io r to m o lt, q u ie s c e n t,
n o t fe e d in g . G lo s s y , c o v e r e d w ith p r o m in e n t s e ta e ,
e s p e c ia lly lo n g c a u d a d , it is u p to 0 .8 m m le n g th . C o lo r is
d a r k k h a k i, p in k is h c a u d a d . P in k is h tin t d e v e lo p s o n a ll
s e g m e n t s c l o s e r t o t h e fi r s t m o l t , e s p e c i a l l y l a t e r a l l y i n
p r o x im ity o f s p ir a c le s .
2nd instar ( N = 1 0 , F i g . 1 1 e ) i s c a . 6 m m l o n g , 6 d a y s ,
w ith th e la s t 2 4 h o u r s p r io r to m o lt, q u ie s c e n t, n o t
fe e d in g . S im ila r to p r e v io u s in s ta r, b u t w ith s e ta e
c o m p a r a tiv e ly s h o r te r, it is o f le s s p r o n o u n c e d p in k is h
c o lo r a tio n . G r o u n d c o lo r is s till d a r k k h a k i.
3rd instar ( N = 7 , F i g s . 1 1 f , g ) i s c a . 9 m m l o n g , 6
d a y s , w ith th e la s t 2 4 h o u r s p r io r to m o lt, q u ie s c e n t, n o t
fe e d in g . U p o n fe e d in g , c o lo r lig h te n s to k h a k i a n d e v e n
lig h te r p a tte r n b e c o m e s m o r e p r o m in e n t o n e a c h
s e g m e n t. T h e p a tte r n c o n s is ts o f a m e d ia n s tr ip e (o v e r
h e a r t) a n d tw o lo n g itu d in a l s tr ip e s o n e a c h s id e p la c e d
a t a s lig h t a n g le to th e m e d ia n s tr ip e . C a te r p illa r
a p p e a r s s e m i-tr a n s p a r e n t, n o t v e r y s tr o n g ly p ig m e n te d .
4th instar ( N = 5 , F i g . 1 1 h ) i s c a . 1 2 m m l o n g , 6 d a y s ,
w ith th e la s t 2 4 h o u r s p r io r to m o lt, q u ie s c e n t, n o t
fe e d in g . T r a c h e a e b e c o m e m o r e v is ib le th r o u g h s e m i-
V
O L U M
r o
s o
th
o n
E
6 6 , N
U M
F IG . 1 1 . L ife h is
a d to B o q u illa s C
l u t i o n . b) O v u m
e b a s e o f t h e fl o w
a n o a k le a f, d o r
B E R
to r
a n
o n
e r
s o -
2
y o f Strymon solitario. A l l s p
y o n , ~ 1 m i E o f S H 1 1 8 , n r. B
a Hechtia fl o w e r s t a l k . C a t e r p
b u d t o t h e l e f t o f t h e l a r v a ; f)
la te r a l a n d d o r s a l v ie w s , r e s p e
1 0 5
e c im e n s fr o m
a r k e r H o u s e .
i l l a r s : c) , d) 1
, g) 3 r d i n s t a r
c tiv e ly .
T e x a s : B r e
a) C a p t i v e
s t in s ta r, o n
; h) 4 t h i n s t
w s t
fe m
d)
a r ;
e r C o . B
a le s fe e
r e a d y to
i) 5 t h i n
ig B e n d N
d in g o n a
m o l t ; e) 2
s t a r . j) , k)
a tio n a l
p a p e r to
n d in s ta
p u p a m(
P a r k ,
w e l w
r, fe e d
a d u lt
n o
ith
in
im
r th o
d ilu
g c a v
a g e F
f th e p a r k
te d h o n e y
ity s e e n a t
i g . 4#12)
1 0 6 1 0 6
tr a n s p a r e n t c u tic u le a s a w e b -lik e lig h te r la te r a l p a tte r n
in th e v ic in ity o f s p ir a c le s , o th e r w is e th e s a m e a s
p r e v io u s in s ta r.
5th instar ( N = 2 , F i g . 1 1 i ) i s c a . 1 5 m m l o n g , 9 d a y s
to p u p a tio n , 6 d a y s fe e d in g , 1 .5 d a y s s e a r c h in g fo r
p u p a tio n s ite , 1 .5 d a y s a s p r e p u p a . I t is th e s a m e c o lo r
a s p r e v io u s in s ta r, ju s t la r g e r. T r a c h e a e a r e c le a r ly
v is ib le . S e ta e c o v e r in g th e b o d y a r e s h o r t a n d th in
c o m p a r e d t o t h o s e i n Strymon melinus.
Pupa ( N = 1 , F i g s . 1 1 j , k ) i s 8 . 0 m m l o n g , 3 . 7 m m w i d e
a t th e a b d o m e n , d e v e lo p s 1 2 d a y s . P r o b a b ly o n th e
s m a lle r s id e , a s th e a d u lt h a tc h e d o n 2 0 -M a y -2 0 0 5 is
th e s m a lle s t s p e c im e n in th e ty p e s e r ie s . P u p a tio n
o c c u r r e d o n a n o a k le a f, c h o s e n a m o n g o th e r o b je c ts
(d r y a n d g r e e n le a v e s , tw ig s , b a r k p ie c e s ) th a t w e r e
p la c e d in a ja r a fte r th e c a te r p illa r s to p p e d fe e d in g a n d
s ta r te d to m o v e a b o u t in s e a r c h fo r p u p a tio n s ite . I t is
m o s t lik e ly th a t in n a tu r a l c o n d itio n s p u p a tio n d o e s n o t
o c c u r o n fl o w e r s e i t h e r , a n d c a t e r p i l l a r fi n d s a p u p a t i o n
s ite c lo s e to th e g r o u n d . P u p a s to u t, o f a ty p ic a l s h a p e
fo r th e fa m ily , is c o v e r e d in s m a ll s e ta e , p a r tic u la r ly
c o n s p ic u o u s o n th e a b d o m e n . I t is a tta c h e d to th e le a f
b y a s in g le g ir d le a n d h o o k e d b y a c r e m a s te r to a lo o s e
s ilk p a d . C o lo r is w h e a t (iv o r y -b r o w n ), a b d o m e n lig h te r,
y e llo w is h e s p e c ia lly o n th e s id e s . E a c h a b d o m in a l
s e g m e n t d o r s a lly h a s tw o d a r k s m a ll m a c u le s o n e a c h
s i d e a n d a d a r k e r m i d d l e s t r i p e ( s e e - t h r o u g h “h e a r t ”
lin e ). M e d ia n d a r k e r m a c u le s a r e p r e s e n t d o r s a lly a t
th e c o n n e c tio n b e tw e e n th e th o r a x a n d a b d o m e n a n d
th e h e a d a n d th o r a x . T h o r a x h a s in d is tin c t g r a y is h
m o ttle d p a tte r n o n b o th s id e s , b a s a lly fr o m th e w in g
c a se s.
S in c e im m a tu r e s d e v e lo p e d w ith o u t d e la y in th e la b ,
it r e m a in s u n c le a r w h a t s ta g e m a y b e q u ie s c e n t
(d ia p a u s e o r a e s tiv a tio n ). I t is m o s t lik e ly th a t a t le a s t
o n e s ta g e e n te r s d ia p a u s e , a s w in te r c o n d itio n s , a t le a s t
in T h e S o lita r io a n d a r o u n d th e P in e c a n y o n a r e to o
h a r s h to s u p p o r t c o n tin u o u s d e v e lo p m e n t. S in c e n o
a d u lts w e r e o b s e r v e d in 2 0 0 6 , w h ic h w a s a v e r y d r y
y e a r , a n d n o Hechtia w a s i n b l o o m , t h e r e s h o u l d a l s o b e
s o m e m e c h a n i s m s f o r t h e s e Strymon t o s t a y i n
d ia p a u s e fo r lo n g e r th a n o n e y e a r. I t s e e m s u n lik e ly
th a t fr a g ile a d u lts a n d s u c c u le n t c a te r p illa r s a r e
c a p a b le o f th is , s o e ith e r o v a o r p u p a e c a n e n te r
d ia p a u s e . O v a w e r e u n e x p e c te d ly la r g e , a b o u t 1 .5
t i m e s t h e s i z e o f Strymon melinus o v a , a n d o v i p o s i t i o n
b e h a v io r w ith le n g th y s e a r c h fo r o v ip o s itio n s ite (a s in
t h e s p e c i e s w i t h o v e r w i n t e r i n g o v a , e . g . Satyrium)
a r g u e fo r th e p o s s ib ility o f o v a e n te r in g d ia p a u s e th a t
m ig h t b e in d u c e d b y d r y a n d c o ld c o n d itio n s . H o w e v e r,
w e c o u ld n o t r u le o u t th e p o s s ib ility th a t p u p a e , a s in
m a n y
s p r i n g - fl y i n g
e p h e m e r a l h a ir s tr e a k s ( e .g .
Callophrys B i l l b e r g , 1 8 2 0 ) m i g h t e n t e r d i a p a u s e t h a t
J
O U R N A L
O F
T H E
L
E P I D O P T E R I S T S
’ S
O C I E T ×
c a n l a s t f o r s e v e r a l y e a r s , a s i n Anthocharis B o i s d u v a l ,
R a m b u r & G r a s lin , [1 8 3 3 ] (P ie r id a e ), fo r in s ta n c e .
E a r ly e m e r g e n c e o f a d u lts (1 8 -F e b a s th e e a r lie s t) d a te
is m o r e c o n s is te n t w ith th e p u p a l h y p o th e s is , a s it
w o u ld m e a n th a t th e o v u m w o u ld h a v e h a tc h e d n o
la te r th a n e a r ly J a n u a r y , a n d p r o b a b ly a s e a r ly a s
D e c e m b e r, fo r th e a d u lt to a p p e a r in m id -F e b r u a r y . I n
a d d i t i o n , n o fl o w e r i n g p l a n t s a r e a v a i l a b l e i n J a n u a r y . I t
is a ls o p o s s ib le th a t b o th m e c h a n is m s (o v a , p u p a e ) c a n
b e u s e d d e p e n d in g o n c o n d itio n s . F u tu r e r e s e a r c h w ill
a d d r e s s th e s e q u e s tio n s .
Etymology:
T h e s p e c ie s is n a m e d a fte r th e g e o lo g ic fe a tu r e in
th e B ig B e n d R a n c h S ta te N a tu r a l A r e a , T e x a s (T h e
S o lita r io ), w h e r e s p e c im e n s w e r e c o lle c te d b y C J D .
A lth o u g h h o lo ty p e w a s u ltim a te ly s e le c te d fr o m th e
B i g B e n d N a t i o n a l P a r k , “ solit ario” r h y m e s w i t h
serapio, w h i c h i s t h e s p e c i e s g r o u p S. solitario b e l o n g s
to . T h e n a m e is a n o n -L a tin iz e d n o u n in a p p o s itio n a n d
is in d e c lin a b le .
D
IS C U S S IO N
Relationships to and comparisons with other taxa:
I t i s f r e q u e n t l y d i f fi c u l t t o k n o w w h e t h e r a n i n s e c t
r e p r e s e n ts a n e w b io lo g ic a l s p e c ie s , o r is a n e x tr e m e
g e o g r a p h ic v a r ia n t o f a k n o w n s p e c ie s . D u e to th e la c k
o f f o r m a l , v e r i fi a b l e a n d q u a n t i t a t i v e c r i t e r i a t h e
a n s w e r to th is q u e s tio n o fte n r e m a in s h y p o th e tic a l.
P e r h a p s , th e e a s ie s t a p p r o a c h is a n a tte m p t to p la c e th e
u n u s u a l p h e n o ty p e w ith in d e s c r ib e d ta x a . I f th is
a tte m p t is n o t p a r tic u la r ly s u c c e s s fu l, it is lik e ly th a t a
n e w s p e c ie s is d is c o v e r e d .
S i g n i fi c a n t d i f fi c u l t i e s w e r e e n c o u n t e r e d w i t h t h e
B i g B e n d H a i r s t r e a k i d e n t i fi c a t i o n . W h i l e i t i s a p p a r e n t
t h a t i t b e l o n g s t o Strymon ( t e e t h o n t h e m a l e v a l v a e ) ,
a n d t o t h e S. serapio g r o u p ( d o w n - t u r n e d p e n i s t i p
w ith p a ir e d c o r n u ti, B r o m e lia c e a e a s la r v a l fo o d p la n ts ),
it is n o t e a s y to id e n tify it a s a n y s in g le d e s c r ib e d
s p e c ie s , b e c a u s e
it
a p p a r e n tly
p o sse sse s
a n
a m a l g a m a t i o n o f c h a r a c t e r s k n o w n f r o m s e v e r a l S.
serapio g r o u p t a x a . E v e n h i s t o r i c a l l y , t h e r e h a v e b e e n
p r o b le m s w ith th e id e n tity o f th e s p e c im e n s fr o m th e
B ig B e n d N a tio n a l P a r k c o lle c te d b y K e n d a ll.
I n d e p e n d e n tly , r e s e a r c h e s v e r y e x p e r ie n c e d w ith
“ Theclinae” , s u c h a s C l e n c h a n d M i l l e r w r o t e t h e I D
l a b e l s o n t h e t w o o f t h e K e n d a l l s p e c i m e n s : [ “T h e c l a ”
// s p . f // m a y b e n e w // d e t. H . C le n c h 1 9 6 9 ] a n d
[S tr y m o n s p . // p o s s ib ly n e w m// d e t. L e e D . M ille r //
1 9 8 0 ].
L a t e r , t h i s s p e c i e s w a s t e n t a t i v e l y i d e n t i fi e d a s S.
serapio. A l t h o u g h i t d o e s e x h i b i t c e r t a i n t r a i t s o f S.
serapio, f o r i n s t a n c e d o r s a l w i n g p a t t e r n , a s w e t r i e d t o
d e m o n s tr a te h e r e , a ll o th e r c h a r a c te r s a r e in c o n s is te n t
V
O L U M
E
6 6 , N
U M
B E R
2
w i t h S. serapio a n d m a n y r e c e n t l y d e s c r i b e d s p e c i e s
p l a c e d u n d e r “serapio” b y R o b b i n s & N i c o l a y ( 2 0 0 2 ) .
V e n tr a l w in g p a tte r n , in p a r tic u la r d e v e lo p e d p o s tb a s a l
o r a n g e s p o ts o n h in d w in g in m a n y s p e c im e n s , p r e s e n c e
o f o r a n g e s c a le s o n th e d o r s a l fo r e w in g , tib ia l le g
p a tte r n , s h a p e o f s a c c u s , s iz e o f te g u m e n a n d d e s e r t
h a b i t a t d o n o t a g r e e w i t h S. serapio, b u t a g r e e w i t h
s e v e r a l o t h e r d e s c r i b e d Strymon s p e c i e s . W e t h i n k t h a t
t h e e v i d e n c e p r e s e n t e d h e r e i s s u f fi c i e n t t o s u p p o r t t h e
d i s t i n c t n e s s o f t h e B i g B e n d H a i r s t r e a k f r o m S. serapio
a n d ta x a c lo s e ly r e la te d to it (s y n o n y m s a n d s u b s p e c ie s
a c c o r d in g
to
R o b b in s &
N ic o la y
2 0 0 2 ) a n d
c h a r a c te r iz e d b y th e a b s e n c e o f b a s a l o r a n g e s p o ts o n
th e h in d w in g , v e r y s h o r t s a c c u s a n d a b u lk y te g u m e n .
U p o n c a r e fu l c o m p a r is o n o f B ig B e n d s p e c im e n s
w ith a ll 4 6 d e s c r ib e d ta x a p la c e d b y R o b b in s & N ic o la y
( 2 0 0 2 ) a n d R o b b i n s ( 2 0 0 4 ) i n t h e “serapio” g r o u p a n d
a r r a n g e d i n t o 1 5 s p e c i e s , o n l y S. megarus w i t h i t s 5
s y n o n y m s a n d s u b s p e c i e s , a n d S. jacqueline w a r r a n t
f u r t h e r c o n s i d e r a t i o n . B r i e fl y , t h e f o l l o w i n g f e a t u r e s
r u l e o u t o t h e r t a x a : S. serapio ( 1 1 n a m e s , d i s c u s s e d
a b o v e ) , S. glorissima K . J o h n s o n & S a l a z a r , 1 9 9 3 ( 2
n a m e s ) , S. gabatha ( H e w i t s o n , 1 8 7 0 ) ( 4 n a m e s ) , S.
monopeteinus S c h w a r t z & J . ×. M i l l e r , 1 9 8 5 , S. azuba
( H e w i t s o n , 1 8 7 4 ) ( 3 n a m e s ) , S. eremica ( H a y w a r d ,
1 9 4 9 ) ( 4 n a m e s ) , a n d S. veterator ( H . H . D r u c e , 1 9 0 7 )
(3 n a m e s ) la c k th e p o s tb a s a l o r a n g e s p o ts o n th e
h i n d w i n g v e n t e r ; S. oreala ( H e w i t s o n , 1 8 6 8 ) , S. dindus,
S. lucena ( H e w i t s o n , 1 8 6 8 ) ( 6 n a m e s ) , S. cardus
( H e w i t s o n , 1 8 7 4 ) a n d S. ahrenholzi N i c o l a y &
R o b b in s , 2 0 0 5 h a v e v e r y p r o m in e n t p o s tb a s a l s p o ts o r
b a n d s o n th e h in d w in g v e n te r, to g e th e r w ith d e v e lo p e d
e n d o f c e l l b a r ; a n d S. giffordi N i c o l a y & R o b b i n s , 2 0 0 5
p o s s e s s e s q u ite d iffe r e n t d o r s a l w in g p a tte r n w ith m o s t
o f h in d w in g b e in g s o lid ly v io le t-b lu e (m a le s ) o r s la te
(fe m a le s ). I n a d d itio n to th o s e fe a tu r e s , m a n y o th e r
c h a r a c te r s , to o n u m e r o u s to b e d is c u s s e d h e r e a n d
fr e q u e n tly in d iv id u a l to e a c h ta x o n , h e lp u s to r u le o u t
th e a b o v e -m e n tio n e d ta x a a s th e d e te r m in a tio n o f th e
B ig B e n d h a ir s tr e a k , a n d th e r e a d e r s a r e r e fe r r e d to
th e
o r ig in a l d e s c r ip tio n s a n d
e x a m in a tio n
o f
s p e c im e n s .
M a n y B ig B e n d H a ir s tr e a k s p e c im e n s p o s s e s s
o r a n g e p o s tb a s a l s p o ts o n th e h in d w in g v e n te r, w h ile
th o s e s p o ts a r e n e v e r v e r y la r g e , b u t fr e q u e n tly d o t-lik e
a n d m a n y s p e c im e n s la c k th e s p o ts . S o m e s p e c im e n s
h a v e s p o ts o n o n e h in d w in g a n d la c k th e m o n th e o th e r
h in d w in g . T h is fe a tu r e (p r e s e n c e a n d a b s e n c e o f s m a ll
s p o t s ) i s c h a r a c t e r i s t i c o f S. jacqueline. W h i l e d o r s a l
h in d w in g s m ig h t h a v e v io le t-b lu e (m a le s ) a n d s la te
(fe m a le s ) o v e r s c a lin g , it is n e v e r v e r y p r o m in e n t, in
f a c t i s e v e n l e s s p r o m i n e n t t h a n i n m o s t S. serapio
s p e c i m e n s , a n d serapio i s b e i n g r e f e r r e d t o a s t h e
1 0 7
s p e c ie s m o s t c lo s e ly r e s e m b lin g th e B ig B e n d
H a ir s te r a k o n th e d o r s a l w in g s u r fa c e .
T h u s , o u r a n a l y s i s o f 4 6 d e s c r i b e d serapio g r o u p t a x a
le a v e s u s w ith 2 p o s s ib le n a m e s fo r th e B ig B e n d
H a i r s t r e a k : S. megarus a n d S. jacqueline. T h e f o l l o w i n g
e v id e n c e s u g g e s ts th a t th e B ig B e n d H a ir s tr e a k m ig h t
b e d i s t i n c t f r o m S. megarus. F i r s t , s o m e f e m a l e s
d i s p l a y o r a n g e s c a l e s o n d o r s a l f o r e w i n g . S. megarus
(a ll 5 ta x a , R o b b in s & N ic o la y 2 0 0 2 ) fe m a le s a lw a y s
la c k o r a n g e s c a lin g . S e c o n d , p o s tb a s a l o r a n g e s p o ts o n
t h e h i n d w i n g v e n t e r a r e s m a l l e r t h a n t h o s e i n m o s t S.
megarus, o r l a c k i n g a l t o g e t h e r . T h i r d , v i o l e t - b l u e
o v e r s c a lin g is le s s d e v e lo p e d in th e n e w ta x o n
c o m p a r e d t o m o s t S. megarus. F o r t h , b l o t c h e s o f l i g h t
s c a le s o n th e h in d w in g b e tw e e n th e o r a n g e p o s tm e d ia n
b a n d a n d s u b m a r g in a l d a r k lu n u le s a r e n o t p r o m in e n t,
w h i l e b e i n g w e l l - d e v e l o p e d a n d c o n s p i c u o u s i n S.
megarus.
P e r h a p s th e s tr o n g e s t e v id e n c e fo r b io lo g ic a l
d i s t i n c t n e s s o f t h e n e w t a x o n p o p u l a t i o n s f r o m S.
megarus i s t h a t i n T a m a u l i p a s ( M e x i c o ) S. megarus fl i e s
in th e p r o x im ity o f th e B ig B e n d h a ir s tr e a k p h e n o ty p e
w ith a b o u t 3 5 m ile s s e p a r a tin g th e tw o d is tin c t
p h e n o ty p e s (s o u th o f C iu d a d V ic to r ia a n d n e a r E l A b r a
a n d G o m e z F a r ia s ). I t is p o s s ib le th a t th e tw o ta x a a r e
e v e n s y m p a t r i c , b u t S. megarus o c c u r s i n b r u s h a n d
f o r e s t h a b i t a t , n o t d e s e r t s a l t h o u g h Hechtia i s p r e s e n t
o n d r y k a r s t l i m e s t o n e a t t h e s e l o c a l i t i e s . T h e s e S.
megarus s p e c i m e n s e x h i b i t l a r g e r , b l o t c h y o r a n g e s p o t s
o n th e h in d w in g , e s p e c ia lly in th e p o s tb a s a l a r e a , m o r e
e x te n s iv e v io le t-b lu e d o r s a l o v e r s c a lin g , w ith th e
d iffe r e n c e b e in g e s p e c ia lly p r o m in e n t in fe m a le s in th e
h in d w in g s u b m a r g in a l a r e a (F ig . 1 0 # 1 7 ), w e llp r o n o u n c e d lig h t a r e a s b e tw e e n th e s u b m a r g in a l d a r k
lu n u le s a n d p o s tm e d ia n o r a n g e b a n d o n th e h in d w in g
v e n te r (F ig s 1 0 # 1 1 , # 1 7 ). B ig B e n d H a ir s tr e a k
p h e n o ty p e la c k s e x te n s iv e d o r s a l m a r g in a l b lu e a r e a s
a n d lig h t s c a le s o n th e h in d w in g v e n te r a r e r e s tr ic t e d
to a r o u n d th e d a r k e r s u b m a r g in a l lu n u le s a n d a lo n g
th e m a r g in o f th e o r a n g e p o s tm e d ia n b a n d .
I t is e s s e n tia l to c o n s id e r v a r ia tio n in e a c h ta x o n to
d e fi n e t h e b o u n d a r i e s b e t w e e n t h e m . T o a d d r e s s t h i s
q u e s tio n , w e a s s e m b le d a ty p e s e r ie s o f c lo s e to 7 0
s p e c im e n s fr o m
th e B ig B e n d a n d e x a m in e d
p h o to g r a p h s o f 4 in d iv id u a ls fr o m n e a r C iu d a d V ic to r ia
(M e x ic o : T a m a u lip a s ) th a t in m a n y c h a r a c te r s a g r e e
w ith th e B ig B e n d s p e c im e n s v e r y w e ll. T h is p o o l o f
s p e c i m e n s f r o m a n a r e a o f 5 0 0 m i l e s d e fi n e s i n d i v i d u a l
a n d g e o g r a p h ic v a r ia tio n in th is s p e c ie s . F ig s 4 , 5 , a n d
9 s h o w th e e x te n t o f th is v a r ia tio n . W h ile it is a p p a r e n t
th a t th e s iz e , s h a p e a n d c o lo r o f v e n tr a l o r a n g e m a c u le s
a n d th e e x te n t o f th e d o r s a l v io le t-b lu e o v e r s c a lin g
v a r i e s q u i t e s i g n i fi c a n t l y , i t i s e q u a l l y c l e a r t h a t t h e
1 0 8 1 0 8
b e tw e e n
th e
n e w
s p e c ie s
a n d
S.
h ia tu s
jacqueline/ megarus r e m a i n s
w e l l - d e fi n e d . T h o s e
d ia g n o s tic fe a tu r e s lis te d in th e p r e v io u s p a r a g r a p h
h o ld tr u e fo r a ll e x a m in e d s p e c im e n s , in c lu d in g liv e
in d iv id u a ls fr o m c e n tr a l T a m a u lip a s .
A l t h o u g h t h e m a i n d i a g n o s t i c f e a t u r e s a r e s u f fi c i e n t
t o d i f f e r e n t i a t e b e t w e e n t h e n e w s p e c i e s a n d S.
jacqueline/ megarus, s e v e r a l o t h e r m o r e
m in u te
d iffe r e n c e s e x is t. F o r in s ta n c e , th e n e w s p e c ie s is
a d d i t i o n a l l y d i f f e r e n t i a t e d f r o m S. jacqueline b y : 1 ) t h e
w in g s h a p e b e in g m o r e r o u n d e d w ith th e fo r e w in g
m a r g in m o r e c u r v e d th a n a r e la tiv e ly s tr a ig h t m a r g in o f
a l m o s t t r i a n g u l a r - s h a p e d w i n g s i n m a l e s S. jacqueline;
2 ) in m o s t s p e c im e n s , b a s a l m a r g in o f th e o r a n g e
m a c u le in v e n tr a l h in d w in g c e ll M 3– C u 1 is b a s a lly o ffs e t
c o m p a r e d to th e b a s a l m a r g in o f th e C u 1– C u 2 m a c u le ,
a n d th e m a r g in s o f th e tw o m a c u le s a r e m o r e in -lin e in
S. jacqueline; 3 ) i n m o s t s p e c i m e n s , o r a n g e m a c u l e i n
v e n tr a l h in d w in g c e ll R s is n o t p r o m in e n tly o ffs e t
b a s a lly c o m p a r e d to th e m a c u le in c e ll S c + R 1– R s , a s
c h a r a c t e r i s t i c o f S. jacqueline.
Biogeographic significance:
O c c u r r e n c e o f a B r o m e l i a d - f e e d i n g Strymon i n t h e
U n ite d S ta te s p e r h a p s s h o u ld n o t c o m e a s a s u r p r is e ,
b e c a u s e B r o m e lia d fa m ily p la n ts o c c u r h e r e . H o w e v e r,
d e s p i t e a w i d e d i s t r i b u t i o n o f S p a n i s h m o s s ( Tillandsia
usneoides) o v e r t h e e n t i r e e a s t e r n U S ( T e x a s t o
M a r y l a n d ) a n d o t h e r Tillandsia s p e c i e s i n A r i z o n a ,
T e x a s a n d F l o r i d a , S. serapio h a s b e e n r e c o r d e d t o u s e
Tillandsia a s l a r v a l h o s t ( s e e F i g . 7 # 4 l e g e n d ) i s n o t a
r e s id e n t o v e r m o s t p a r t o f its h o s t p la n t r a n g e .
A p p a r e n t l y , t h e serapio g r o u p Strymon a r e r e s t r i c t e d t o
s u b tr o p ic a l b io ta . T h u s , th e d is c o v e r y o f th e n e w
s p e c ie s in th e B ig B e n d a r e a b y G e o r g e S c h e n k in 1 9 4 4 ,
R o y O . K e n d a ll (a p p a r e n tly in d e p e n d e n tly ) in 1 9 6 8 ,
C J D i n 1 9 7 3 a n d o t h e r s s i n c e t h e n i s a s i g n i fi c a n t o n e .
I t i s c l e a r t h a t t h i s Strymon i s a w e l l - e s t a b l i s h e d
r e s id e n t in th e r e g io n p r o v id e d th e v a s t a r e a s d e n s e ly
c o v e r e d w i t h i t s f o o d p l a n t Hechtia texensis a n d r e c o r d s
fr o m th e fo llo w in g y e a r s : 1 9 4 4 [S c h e n k , 2 5 -S e p – 9 -O c t,
1 6 ] , 1 9 6 8 [ K e n d a l l 2 7 - M a r 1 f] , 1 9 7 1 [ K e n d a l l 2 0 - S e p
1 m] , 1 9 7 3 [ D u r d e n 1 7 – 2 0 - M a y , p h o t o , 2 m] , 1 9 7 5
[ D u r d e n 8 - J u n , 5 m] , 1 9 8 5 [ K n u d s o n 2 9 - N o v 1 m] , 1 9 8 7
[ K e n d a l l , 2 6 – 2 8 - M a r , 4 f] , 1 9 8 9 [ S w e n g e l , 2 6 - A p r
p h o to ], 1 9 9 2 [K e n d a ll, 5 -A p r, la s t in s ta r la r v a ] 1 9 9 4
[ S p o m e r 2 5 - M a r , 1 m] , 2 0 0 5 [ K o s t e c k e , G r i s h i n , S p e n c e r
p h o to s , 1 8 -F e b – 2 4 -A p r, 2 5 ], 2 0 0 7 [G r is h in , 1 5 -A p r – 1 5 S e p , 9 ; H e n s l e y , 1 9 - N o v p h o t o 1 m] , 2 0 0 9 [ R e i d 7 - M a r ,
G r i s h i n 8 - M a r , 1 m, 1 f, W a u e r 1 4 - M a r , p h o t o s ] .
R e c e n tly , th e H a ir s tr e a k h a s b e e n fo u n d in a ll s e a r c h e s
f o r i t d u r i n g t h e b l o o m o f Hechtia. H o w e v e r , Hechtia
d o e s n o t b lo o m in s o m e d r y y e a r s . N o s p e c im e n s w e r e
J
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f o u n d i n t h e a b s e n c e o f f r e s h Hechtia fl o w e r s t a l k s . I t i s
p a r tic u la r ly in te r e s tin g th a t th e a p p a r e n tly m o s t s im ila r
r e l a t i v e , S. jacqueline a l s o i n h a b i t i n g a r i d a r e a s , h a s
b e e n d e s c r ib e d fr o m a d iffe r e n t c o n tin e n t (P e r u ), a n d
o n ly r e c e n tly . I t s e e m s lik e in a c c e s s ib ility a n d d r y n e s s
o f t h e h a b i t a t t h a t i s r e l a t i v e l y d e v o i d o f o t h e r b u t t e r fl y
s p e c i e s a n d t h e r a r i t y o f t h e s e Strymon s p e c i e s e x c e p t
in th e m o s t fa v o r a b le a n d r a in y y e a r s h in d e r s th e ir
d is c o v e r y a n d s u b s e q u e n t r e s e a r c h . I n a d d itio n to th e
B ig B e n d a r e a (T e x a s : B r e w s te r a n d P r e s id io c o u n tie s ),
i t m i g h t b e f r u i t f u l t o c o n t i n u e s e a r c h e s f o r t h e serapio
g r o u p Strymon i n e x t r e m e s o u t h T e x a s ( a n d i n C o a h u i l a
a n d C h i h u a h u a , M e x i c o ) , s i n c e Hechtia s p e c i e s a r e
k n o w n to o c c u r in Z a p a ta a n d S ta r r c o u n tie s .
C
O N C L U S IO N S
E v id e n c e is p r e s e n te d th a t th e B r o m e lia d -fe e d in g
Strymon f o u n d i n a r i d , d e s e r t a n d r o c k y s l o p e s a r e a s o f
B ig B e n d N a tio n a l P a r k a n d its v ic in ity is a n
u n d e s c r ib e d b io lo g ic a l s p e c ie s , w h ic h w a s n a m e d
h e r e in . T h is n e w ta x o n c o m b in e s c h a r a c te r s o f s e v e r a l
d e s c r i b e d Strymon s p e c i e s f r o m t h e serapio g r o u p . S.
jacqueline a p p e a r s t o b e m o s t s i m i l a r i n f a c i e s a n d
h a b i t a t . S. jacqueline s h a r e s v e n t r a l w i n g p a t t e r n s ,
p r e s e n c e o f o r a n g e s c a lin g o n fe m a le d o r s a l fo r e w in g ,
g e n ita lia s tr u c tu r e s a n d h a b ita t p r e fe r e n c e s . H o w e v e r,
d o r s a l p a tte r n o f th e n e w s p e c ie s d iffe r s d r a s tic a lly
f r o m S. jacqueline a n d h a s v e r y r e s t r i c t e d v i o l e t b l u e
a n d s la te a r e a s p r e s e n t o n ly a s o v e r s c a lin g . T h is d o r s a l
p a t t e r n i s m o s t s i m i l a r t o S. serapio. A n o t h e r r e l a t e d
s p e c i e s , S. megarus, s h a r e s v e n t r a l a n d p a r t l y d o r s a l
w in g p a tte rn s , b u t o c c u r s in b r u s h a n d fo r e s t h a b ita t,
n o t d e se rts. T h e n e w
s p e c ie s is m o s t e a s ily
d i s t i n g u i s h e d f r o m b o t h S. jacqueline a n d S. megarus
b y th e r e s tr ic te d lig h t s c a le a r e a s ju s t b a s a lly fr o m th e
s u b m a r g in a l d a r k lu n u le s o n th e h in d w in g v e n te r,
p r e s e n t o n l y a s f r a m i n g o f t h e l u n u l e s . I n S. jacqueline
a n d S. megarus t h e s e a r e a s a r e m o r e e x t e n s i v e a n d
u s u a lly r e a c h a t le a s t 1 /3 o f th e d is ta n c e fr o m d a r k
m a r g in a l lu n u le s to p o s tm e d ia n b a n d o f o r a n g e s p o ts .
I n g e n e r a l, th e n e w s p e c ie s is le s s c o n tr a s te d , a n d is
m o r e fl a t a n d u n i f o r m l y c o l o r e d i n a p p e a r a n c e : l e s s
d e v e lo p e d b lu e s c a lin g , s m a lle r lig h t a r e a s , fe w e r d a r k
s c a le s o n th e le g tib ia e . W h ile g e n ita lia d o n o t o ffe r
p ro fo u n d
d iffe r e n c e s
fro m
m a n y
c o n g e n e rs,
c o m b in a tio n o f s h o r te r te r m in a l ta p e r e d a r e a s o f
g n a th o s a r m s w ith c le a r ly a s y m m e tr ic te r m in a lly
p o in t e d s a c c u s i s c h a r a c t e r i s t i c o f t h e m a l e s o f t h i s
ta x o n .
A
C K N O W L E D G E M E N T S
W e a r e in d e b te d to th e U n ite d S ta te s N a tio n a l P a r k S e r v ic e
a n d th e p e r s o n n e l o f B ig B e n d N a tio n a l P a r k fo r a s s is ta n c e (in
p a r tic u la r, to W ild life B io lo g is t R a y m o n d S k ile s ), a n d c o n tin u -
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in g s u p p o r t w ith r e s e a r c h p e r m its (B I B E -2 0 0 4 -S C I -0 0 1 1 ,
B I B E -2 0 0 5 -S C I -0 0 0 6 , B I B E -2 0 0 7 -S C I -0 0 0 7 , B I B E -2 0 0 8 -S C I 0 0 4 4 ) th a t w e r e a b s o lu te ly e s s e n tia l fo r th is w o r k . W e a r e g r a te fu l to E d w a r d G . R ile y (A s s o c ia te C u r a to r, T e x a s A & M U n iv e r s ity in s e c t c o lle c tio n ) fo r th e lo a n o f s p e c im e n s a n d
p e r m is s io n to p h o to g r a p h s p e c im e n s u n d e r h is c a r e . W e a c k n o w le d g e T e x a s P a r k s a n d W ild life D e p a r tm e n t (N a tu r a l R e s o u r c e s P r o g r a m D ir e c to r D a v id H . R is k in d ) fo r th e p e r m it
# 0 8 -0 2 R e v m a k in g th e w o r k in T e x a s S ta te P a r k s p o s s ib le . F o r
w o r k d o n e b e fo r e c r e a tio n o f th e B ig B e n d R a n c h S ta te N a tu r a l
A r e a (C J D ) th a n k s a r e d u e to R e d O liv e r, S te v e K e n n e d y , M ik e
M c K a n n a n d R o s e a n n e R o w le tt o f T h e T e x a s G e n e r a l L a n d O ffi c e ( 1 9 7 3 ) a n d D o n K e n n a r d , M a r y B u t t e r w i c k a n d S t u a r t
S tr o n g o f T h e T e x a s N a tu r a l A r e a s S u r v e y , L y n d o n B . J o h n s o n
S c h o o l o f P u b lic A ffa ir s , T h e U n iv e r s ity o f T e x a s a t A u s tin
(1 9 7 5 ). T h a n k s a r e d u e a ls o to th e la te R o y O . a n d C o n n ie A .
K e n d a l l a n d t o W . W . M c G u i r e f o r fi e l d d i s c u s s i o n a n d d i s c o v e r ie s m a d e a t E l A b r a , G o m e z F a r ia s a n d R io S a b in a s (T a m a u lip a s , M e x i c o , 1 9 7 2 t o 1 9 7 4 ) a s t h e y w e r e i n t r o d u c e d t o C J D ’s f a v o r ite lo c a litie s , w h ic h w e r e in tu r n r e c o m m e n d e d to h im b y
th e la te H a r r y C le n c h . W e a r e in d e b te d to J o n a th a n P . P e lh a m
(C u r a to r ia l A s s o c ia te o f L e p id o p te r a , B u r k e M u s e u m o f N a tu r a l H is to r y a n d C u ltu r e , U n iv e r s ity o f W a s h in g to n S e a ttle ,
W A ) f o r d i s c u s s i o n s o f t h e o l d e s t k n o w n S. solitario s p e c i m e n s
fr o m th e B ig B e n d a n d c r itic a l r e a d in g o f th e m a n u s c r ip t. S p e c ia l th a n k s to K im D a v is , M ik e S ta n g e la n d , M ic h a e l A . R ic k a r d ,
D a n H a r d y , R ic h K o s te c k e , M a r tin R e id , R . C r a ig H e n s le y , a n d
A n d r e w S p e n c e r fo r s h a r in g im a g e s o f liv e a d u lts a n d p e r m is s io n to u s e th e ir p h o to g r a p h s a s illu s tr a tio n s . N V G th a n k s A n d r e w D . W a r r e n f o r b r i n g i n g t h e s i g n i fi c a n c e o f t h i s i n s e c t t o
o u r a tte n tio n , n u m e r o u s d is c u s s io n s , in s ig h tfu l c o m m e n ts a n d
c o n t i n u o u s h e l p s h a p i n g o u r o p i n i o n s a b o u t Strymon, a n d f o r
th e p h o to g r a p h s o f th e S c h e n k s p e c im e n s , th e la te G e o r g e
A u s t i n f o r d i s c u s s i o n s o f Strymon a n d r e c o r d s o f S. megarus
fr o m M e x ic o p e r M G C L c o lle c tio n ; R o b e r t K . R o b b in s fo r
s tim u la tin g d is c u s s io n s , e n c o u r a g e m e n t, a n d m a n y h e lp fu l
c o m m e n ts o n th e m a n u s c r ip t; C h a r le s B o r d e lo n , J r. fo r c r itic a l
r e a d in g o f th e m a n u s c r ip t; S te v e S p o m e r fo r d is c u s s io n o f th e
s p e c i m e n h e c o l l e c t e d ; R e n e L a h o u s s e f o r t h e S. megarus t y p e
i m a g e s ; B l a n c a H u e r t a s f o r h e l p a n d S. serapio t y p e i m a g e s ; R o
W a u e r fo r im a g e s a n d d is c u s s io n s .
L
A
, G . T .
Strymon H
L y c a e n id a
L M E R , G . R
o f th e L y c
A
B
E
B
L
C
C
A
A
C
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D
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D
R
D
D
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U
C
D
F
F
G
G
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IT E D
K . J O H N S O N . 1 9 9 7 . T h e c lin a e o f R o n d ô n ia , B r a z il:
ü b n e r, w ith d e s c r ip tio n s o f n e w s p e c ie s (L e p id o p te r a :
e ). I n s e c ta M u n d i 1 1 (3 /4 ): 2 0 1 – 2 3 5 .
L
. & G . F . P R A T T . 1 9 8 8 . A s u r v e y o f th e la s t in s ta r la r v a e
a e n id a e o f C a lifo r n ia . J . R e s . L e p . 2 7 : 1 – 8 1
U T E L S P A C H E R , C . R . 1 9 7 2 . S o m e o b s e r v a t io n s o n t h e L e p id o p t e r a o f
b r o m e lia d s . J . L e p id . S o c . 2 6 (3 ): 1 3 3 – 1 3 7 .
A N D , K . P . 1 9 9 4 . Strymon oreala ( H e w i t s o n , 1 8 6 8 ) ( L e p i d o p t e r a : L y c a e n i d a e ) r e a r e d f r o m fl o w e r o f t h e b r o m e l i a d Quesnelia lateralis
in s o u th e r n B r a z il. N e w s L e p id . S o c . 1 9 9 4 (1 ): 5 .
R T E R , W . 1 9 3 4 . N o t e s o n t w o p e s t s o f p in e a p p le n o t k n o w n in
H a w a ii. P r o c . H a w . E n to m o l. S o c . 8 (3 ): 3 9 5 – 3 9 7 .
R T E R , W . 1 9 4 9 . I n s e c t N o t e s f r o m S o u t h A m e r ic a w it h s p e c ia l r e f e r e n c e t o Pseudococcus brevipes a n d m e a l y b u g w i l t . J . E c o n o m i c
E n t. 4 2 (5 ): 7 6 1 – 7 6 6 .
M S T O C K , J . H . 1 9 1 8 T h e w in g s o f in s e c t s . T h e C o m s t o c k P u b lis h in g
C o m p a n y, Ith a c a . 4 3 0 p p .
U C E , H . H . 1 8 9 0 . D e s c r i p t i o n s o f s e v e n n e w s p e c i e s o f Thecla. E n to m o l. M o n . M a g . 2 6 (3 1 3 ): 1 5 1 – 1 5 2 .
U C E , H . H . 1 9 0 7 . O n N e o t r o p ic a l L y c a e n id a e , w it h d e s c r ip t io n s o f
n e w s p e c ie s . P r o c . Z o o l. S o c . L o n d o n 1 9 0 7 (3 ): 5 6 6 – 6 3 2 .
A R T E , M ., R . K . R O B B IN S &
O . H . H . M IE L K E . 2 0 0 5 . I m m a tu re
s t a g e s o f Calycopis caulonia ( H e w i t s o n , 1 8 7 7 ) ( L e p i d o p t e r a , L y c a e n id a e , T h e c lin a e , E u m a e in i), w ith n o te s o n r e a r in g d e tr itiv o r o u s h a i r s t r e a k s o n a r t i fi c i a l d i e t . Z o o t a x a 1 0 6 3 : 1 – 3 1
A R T E , M . & R . K . R O B B I N S . 2 0 0 9 . I m m a t u r e S t a g e s o f Calycopis
bellera ( H e w i t s o n ) a n d C. janeirica ( F e l d e r ) ( L e p i d o p t e r a , L y -
U S T IN
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c a e n i d a e , T h e c l i n a e , E u m a e i n i ) : T a x o n o m i c S i g n i fi c a n c e a n d N e w
E v id e n c e fo r D e tr itiv o r y . Z o o ta x a 2 3 2 5 : 3 9 – 6 1
U R D E N , C . J . 1 9 7 6 . B u t t e r fl i e s o f t h e Solitario – F r e s n o C r e e k – B o f e c illo s M o u n ta in s r e g io n , w e s te r n B ig B e n d (P r e s id io a n d B r e w s te r
c o u n t i e s ) T e x a s . P p . 1 2 5 – 1 3 1 . In S m i t h , G . ( e d . ) , T h e S o l i t a r i o . A
n a tu r a l a r e a s u r v e y . N o . 9 . L y n d o n B . J o h n s o n S c h o o l o f P u b lic A ffa ir s . T h e U n iv e r s ity o f T e x a s , A u s tin .
×A R , H . G . 1 9 1 4 . R e p o r t o n t h e L e p i d o p t e r a o f t h e S m i t h s o n i a n B i o lo g ic a l S u r v e y o f th e P a n a m a C a n a l Z o n e . P r o c . U . S . N a t. M u s .
4 7 (2 0 5 0 ): 1 3 9 – 3 5 0 .
A B R IC IU S , J . C . 1 7 9 3 . E n t o m o lo g ia s y s t e m a t ic a e m e n d a t a e t a u c t a . S e c u n d u m c la s s e s , o r d in e s , g e n e r a , s p e c ie s a d je c tis s y n o n im is , lo c is ,
o b s e r v a tio n ib u s , d e s c r ip tio n ib u s . 3 (1 ): 2 6 9 . H a fn ia e : C h r is tia n
G o ttlie b P r o ft, F il. e t S o c ., 3 ( 1 ) :[ v i] + 4 8 8 p p .
A ×N E L , C . & K . J O H N S O N . 2 0 0 0 . A n e w s p e c i e s o f Strymon H ü b n e r
fr o m F r e n c h G u ia n a (L e p id o p te r a , L y c a e n id a e ). B u lle tin d e la S o c ié té e n to m o lo g iq u e d e F r a n c e 1 0 5 (4 ): 3 7 5 – 3 7 9 .
E ×E R , C . 1 8 3 7 . 5 : 4 2 . I n H ü b n e r J . ( e d . ) , Z u t r ä g e z u r S a m m l u n g e x o tis c h e r S c h m e tte r lin g e 5 : 1 – 5 2 . J a c o b H ü b n e r : A u g s b u r g .
O D A R T , J . B . [ 1 8 2 4 ] . 9 : 6 3 8 . I n L a t r e ille P . A . a n d J .B . G o d a r t ( e d s .) ,
E n c y c lo p é d ie M é th o d iq u e . H is to ir e n a tu r e lle . E n to m o lo g ie , o u
h is to ir e n a tu r e lle d e s c r u s ta c é s , d e s a r a c h n id e s e t d e s in s e c te s
9 (2 ): 3 2 9 – 8 2 8 . v e u v e A g a s s e , P a r is .
O D M A N , F . D . & O . S A L V IN . 1 8 8 7 . 2 : 9 3 . I n B io lo g ia C e n t r a li- A m e r ic a n a . I n s e c ta . L e p id o p te r a - R h o p a lo c e r a . D u la u & C o ., B e r n a r d
Q u a r itc h , L o n d o n .
R E E N A . W . 1 9 7 3 . T h e e c o lo g y , m o r p h o lo g y , a n d d is t r ib u t io n o f
Hechtia scariosa i n t h e B i g B e n d N a t i o n a l P a r k , i n c l u d i n g a b r i e f
h is to r y o f th e a r e a . M S c T h e s is . S u l R o s s U n iv e r s ity , A lp in e , T X .
1 1 7 p p .
A R R I S , W . V . 1 9 2 7 . O n a l y c a e n i d b u t t e r fl y a t t a c k i n g p i n e a p p l e s i n
T r in id a d , B .W .I . B u ll. E n to m o l. R e s . 1 8 ( 2 ) : 1 8 3 – 1 8 8 .
A ×W A R D , K . J . 1 9 4 9 . N u e v a s e s p e c i e s [ s i c ] d e “ L y c a e n i d a e ” d e l a A r g e n tin a ( L e p . R h o p .) . A c ta z o o lo g ic a L illo a n a 8 : 5 6 7 – 5 8 1 .
E W IT S O N , W . C . 1 8 6 8 . D e s c r ip t io n s o f s o m e n e w
s p e c ie s o f L y c a e n id a e . J o h n V a n V o o r s t, L o n d o n . 3 6 p p .
_ _ _ _ _ _ . 1 8 7 0 . R e m a r k s o n a n d d e s c r ip tio n s o f n e w s p e c ie s o f b u tte r fl i e s c o l l e c t e d b y M r . B u c k l e y i n E c u a d o r 4 : 4 9 – 7 9 . J o h n V a n
V o o r s t: L o n d o n ..
_ _ _ _ _ _ . 1 8 7 4 . I llu s tr a tio n s o f D iu r n a l L e p id o p te r a . L y c a e n id a e . 6 :
1 5 1 – 1 8 5 [s ic ]. J o h n v a n V o o r s t, L o n d o n .
_ _ _ _ _ _ . 1 8 7 7 . I llu s tr a tio n s o f D iu r n a l L e p id o p te r a . L y c a e n id a e 7 :
1 8 5 – 2 0 9 [s ic ]. J o h n v a n V o o r s t, L o n d o n .
A N Z E N , D . H . & W . H A L L W A C H S . 2 0 1 1 . D y n a m ic d a t a b a s e f o r a n in v e n to r y o f th e m a c r o c a te r p illa r fa u n a , a n d its fo o d p la n ts a n d p a r a s ito id s , o f A r e a d e C o n s e r v a c io n G u a n a c a s te (A C G ), n o r th w e s te r n C o s ta R ic a < h ttp ://ja n z e n .s a s .u p e n n .e d u > .
O H N S O N , K . , R . C . E I S E L E , & B . N . M A C P H E R S O N 1 9 9 0 . T h e “h a i r s t r e a k b u t t e r fl i e s ” ( L y c a e n i d a e , T h e c l i n a e ) o f n o r t h w e s t e r n A r g e n t i n a . I I . Strymon, s e n s u s t r i c t o . B u l l . A l l y n M u s . 1 3 0 : 1 – 7 7 .
O H N S O N , K . & K . R . K R O E N L E IN . 1 9 9 3 . T h r e e r e m a r k a b le n e w s p e c ie s
o f Strymon H ü b n e r f r o m B r a z i l a n d E c u a d o r ( L e p i d o p t e r a , L y c a e n id a e ,T h e c lin a e ) . R e p . M u s . N a t. H is t., U n iv . W is c . ( S te v e n s
P o in t) 3 5 : [i– ii]+ 1 – 8 .
O H N S O N , K . & J . A . S A L A Z A R . 1 9 9 3 . N e w s p e c ie s , s t a t u s e s a n d c o m b in a t i o n s i n n o r t h e r n S o u t h A m e r i c a n Strymon ( L e p i d o p t e r a , L y c a e n id a e , T h e c lin a e ) . R e p . M u s . N a t. H is t., U n iv . W is c . ( S te v e n s
P o in t) 2 6 : 1 – 1 3 .
I R B ×, W . F . 1 8 7 1 . A S y n o n y m i c C a t a l o g u e o f D i u r n a l L e p i d o p t e r a .
Jo h n V a n V o o rst, L o n d o n 6 9 0 p p .
L O T S , A . B . 1 9 7 0 . L e p i d o p t e r a , p p . 1 1 5 – 1 3 0 . In T u x e n S . L . ( e d . ) , T a x o n o m i s t ’s g l o s s a r y o f g e n i t a l i a i n i n s e c t s . M u n k s g a a r d , C o p e n h a g e n .
A T H ×, P . I . 1 9 3 0 . N o t e s o n S o u t h A m e r i c a n L y c a e n i d a e , w i t h d e s c r i p tio n s o f n e w s p e c ie s . T r a n s . E n to m o l. S o c . L o n d . 7 8 (1 ): 1 3 3 – 1 3 7 .
E C R O M , J . F . & K . J O H N S O N . 1 9 9 7 . A d d i t i o n s t o t h e Strymon f a u n a o f
C o l o m b i a ( E u m a e i n i ; Strymoni n a ) . R e v i s t a d e T h e c l i n a e c o l o m b ia n o s 2 (1 6 ): [ii] + 1 – 4 7 .
_ _ _ _ _ _ . 1 9 9 7 . A n e w s p e c i e s g r o u p o f S o u t h A m e r i c a n Strymon a n d
i t s p o s s i b l e a f fi n i t y t o e l e m e n t s o f t h e A n t i l l e a n Strymon f a u n a
1 1 0 1 1 0
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(E u m a e in i, S tr y m o n in a ). R e v is ta d e T h e c lin a e c o lo m b ia n o s 2 (1 8 ):
[ii] + 1 – 1 7 .
I C O L A ×, S . S . & R . K . R O B B I N S . 2 0 0 5 . F i v e n e w d r y - a r e a S o u t h A m e r i c a n Strymon s p e c i e s ( L y c a e n i d a e : T h e c l i n a e ) a n d t h e i r b i o g e o g r a p h i c s i g n i fi c a n c e . J . R e s . L e p . 3 8 : 3 5 – 4 9 .
T E R O , L . S . & L . C . M A R I G O . 1 9 9 0 . B u t t e r fl i e s : b e a u t y a n d b e h a v i o r
o f B r a z ilia n s p e c ie s . M a r ig o C o m u n ic a c a o V is u a l, R io d e J a n e ir o .
1 2 8 p p .
I C K A R D , M . A . & N . V . G R I S H I N . 2 0 1 0 . Strymon serapio n e w f o r t h e
U .S . ( L y c a e n id a e : T h e c lin a e ) . N e w s L e p . S o c . 5 2 ( 3 ) : 7 9 – 8 4
O B B IN S , R . K . 1 9 9 1 . E v o lu t io n , c o m p a r a t iv e m o r p h o lo g y , a n d id e n t ifi c a t i o n o f t h e e u m a e i n e b u t t e r fl y g e n u s Rekoa K a y e ( L y c a e n i d a e :
T h e c lin a e ). S m ith . C o n tr. Z o o l. # 4 9 8 . 6 4 p p .
_ _ _ _ _ _ . 2 0 0 4 . L y c a e n id a e . T h e c lin a e . T r ib e E u m a e in i. P p . 1 1 8 – 1 3 7 .
In L a m a s G . ( e d . ) , C h e c k l i s t : P a r t 4 A . H e s p e r i o i d e a – P a p i l i o n o i d e a . In J . B . H e p p n e r ( e d . ) , A t l a s o f N e o t r o p i c a l L e p i d o p te r a . V o lu m e 5 A . A s s o c ia tio n fo r T r o p ic a l L e p id o p te r a . S c ie n t i fi c P u b l i s h e r s , G a i n e s v i l l e .
_ _ _ _ _ _ . 2 0 1 0 . T h e “ u p s i d e d o w n ” s y s t e m a t i c s o f h a i r s t r e a k b u t t e r fl i e s
(L y c a e n id a e ) th a t e a t p in e a p p le a n d o th e r B r o m e lia c e a e . S tu d ie s
o n N e o tr o p ic a l F a u n a a n d E n v ir o n m e n t 4 5 (1 ): 6 5 – 7 5 .
O B B IN S , R . K . & A . A IE L L O . 1 9 8 2 . F o o d p la n t a n d o v ip o s it io n r e c o r d s
fo r P a n a m a n ia n L y c a e n id a e a n d R io d in id a e . J . L e p . S o c . 3 6 (2 ):
6 5 – 7 5 .
O B B I N S , R . K . & S . S . N I C O L A ×. 2 0 0 2 . A n o v e r v i e w o f Strymon H ü b n e r ( L y c a e n id a e :T h e c lin a e : E u m a e in i) . J . L e p . S o c . 5 5 ( 3 ) : 8 5 – 1 0 0 .
A N C H E S , N . F . , S . A . C H O A I R ×, & A . V I L A R D E B O . 1 9 8 5 . A t a q u e d e The-
J
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cla basalides ( G e y e r , 1 8 3 7 ) ( L e p i d o p t e r a : L y c a e n i d a e ) e m f o l h a s
d e a b a c a x i n a P a r a ib a , B r a s il. A n a is d a S o c ie d a d e E n to m o ló g ic a d o
B r a s il 1 4 (1 ): 1 6 7 – 1 6 9 .
C H A U S , W . 1 9 0 2 . D e s c r i p t i o n s o f n e w A m e r i c a n b u t t e r fl i e s . P r o c . U .
S . N a t. M u s. 2 4 (1 2 6 2 ): 3 8 3 – 4 6 0 .
C H W A R T Z , A . & J . ×. M I L L E R . 1 9 8 5 . A n e w s p e c i e s o f h a i r s t r e a k ( L y c a e n id a e ) fr o m H is p a n io la . B u ll. A lly n M u s . 9 9 : 1 – 6 .
C O T T , J . A . 1 9 8 6 . T h e b u t t e r fl i e s o f N o r t h A m e r i c a : a n a t u r a l h i s t o r y
a n d fi e l d g u i d e . S t a n f o r d U n i v e r s i t y P r e s s , S t a n f o r d , C a l i f o r n i a .
5 8 3 p p .
I L V A , A . G . D 'A ., C . R . G O N Ç A L V E S , D . M . G A L V Ã O , A . J . L .
G O N Ç A L V E S , J. G O M E S , M . N . S IL V A , & L . D E S IM O N I. 1 9 6 8 . Q u a rto
C a tá lo g o d o s in s e to s q u e v iv e m n a s p la n ta s d o B r a s il, s e u s p a r a s ito s e p r e d a d o r e s : in s e to s , h o s p e d e ir o s e in im ig o s n a tu r a is . M in is te r io d a A g r ic u ltu r a , R io d e J a n e ir o .V o l. 1 , 6 2 2 p p . V o l. 2 , 2 6 5 p p .
S D A , N R C S . 2 0 1 1 . T h e P L A N T S D a ta b a s e ( h ttp ://p la n ts .u s d a .g o v ,
2 0 J a n u a r y 2 0 1 1 ). N a tio n a l P la n t D a ta C e n te r, B a to n R o u g e , L A
7 0 8 7 4 – 4 4 9 0 U S A .
A R R E N , A . D ., K . J . D A V IS , N . V . G R IS H IN , J . P . P E L H A M &
E . M .
S T A N G E L A N D . 2 0 1 1 . I n t e r a c t i v e l i s t i n g o f A m e r i c a n b u t t e r fl i e s . [ 9 V I I I - 1 1 ] < h t t p : / / w w w . b u t t e r fl i e s o f a m e r i c a . c o m > .
I L A , R . & R . E A S T W O O D . 2 0 0 6 . E x t r a fl o r a l n e c t a r f e e d i n g b y Strymon
jacqueline N i c o l a y & R o b b i n s , 2 0 0 5 ( L e p i d o p t e r a : L y c a e n i d a e :
E u m a e in i). R e v is ta p e r u a n a d e B io lo g ía 1 3 (1 ): 1 2 5 – 1 2 8 .
Received for publication 2 February 2011; revised and accepted 21 September 2011