Two new species of moths (Lepidoptera, Pyralidae, Chrysauginae)
associated with the three-toed sloth (Bradypus spp.) in South America
J . D . BradleyC1 )
Abstract
Two species of chrysaugine moths, discovered as a result of an ecological study in Brazil of tree sloths and their ectoparasites, are described as new and named Cryptoses waagei sp. n. and C. mfipictus sp. n. They are differentiated from their single congener C. choloepi Dyar and near relatives Bradypophila garbei Ihering and Bradypodicola hahneii Spuler which are also found on tree sloths.
Several s p e c i e s of p y r a l i d m o t h s of t h e s u b f a m i l y Chrysauginae have long b e e n k n o w n t o live habitually in t h e f u r of t r e e s l o t h s in South A m e r i c a . The m o t h s w e r e generally believed to be p a r a s i t e s and t o i n h a b i t t h e f u r of t h e animal t h r o u g h o u t t h e i r l i f e - c y c l e . It is only c o m p a r a t i v e l y r e c e n t l y , in a r e p o r t by Waage & M o n t g o m e r y (1976) b a s e d on an ecological s t u d y of t h e m o t h Cryptoses cho Icepi Dyar, w h i c h o c c u r s in C o s t a Rica, Panama and C o l o m b i a , and t h e t h r e e - t o e d s l o t h Brady pus vaiiegatus ( ^ i n f u s c a t u s ) , t h a t f a c t u a l e v i d e n c e on t h e m o t h - s l o t h r e l a t i o n s h i p has become a v a i l a b l e . The r e l a t i o n s h i p proved to be p h o r e t i c , since only in the adult stage (imago) does t h e m o t h a c t u a l l y l i v e in t h e animal's f u r . The i m m a t u r e stages o f the m o t h ( o v u m , iarva, pupa) w e r e f o u n d not t o live on t h e animal but on its d u n g , the larva being coprcphagous (see also Waage & B e s t , 1932, for r e v i e w ) .
S u b s e q u e n t l y , w h e n e c o l o g i c a l s t u d i e s w e r e e x t e n d e d t o Brazil, it w a s o b s e r v e d t h a t t h e m o t h s l i v i n g on t h e t h r e e - t o e d s l o t h (Bra dypus tridactylus and B. variegatus) o c c u r r i n g t h e r e d i f f e r e d s i g n i f i c a n t l y f r o m C. choloepi in c e r t a i n c h a r a c t e r i s t i c s . T a x o n o m i c s t u d y of t h e Brazilian s p e c i m e n s , t o g e t h e r w i t h material in t h e c o l l e c t i o n s of t h e B r i t i s h M u s e u m
(Natural H i s t o r y ) , has s h o w n t h a t t w o d i s t i n c t s p e c i e s are involved w h i c h are c l o s e l y r e l a t e d
and apparently c o n g e n e r i c w i t h C. choloepi but h i t h e r t o u n d e s c r i b e d .
Cryptoses Dyar, 1908
Cryptoses waagei
s p . n .
(Figs. 1-4)
WINGSPAN
: o \ 2 1315 m m . Sexual d i
-m o r p h i s -m not p r o n o u n c e d , f e -m a l e u s u a l l y larger, sexes s i m i l a r in c o l o r a t i o n and f o r e w i n g p a t t s r n , w i t h antennae s h o r t l y c i l i a t e . f l a g e l l u m of male s o m e w h a t s t o u t e r . Labial palpus, head, t h o r a x , p a t a g i u m and t e g u l a ochreous-b r o w n , v a r i a ochreous-b l y s u f f u s e d w i t h g r e y - ochreous-b r o w n ; an-tennal scape and f l a g e l l u m o c h r e o u s - w h i t e , s c a l i n g of f l a g e l l u m in male c o n f i n e d m a i n l y to basal p a r t . In b o t h sexes f o r e w i n g long and n a r r o w , w i t h apex p r o d u c e d and p o i n t e d and t e r m e n very o b l i q u e ; g r o u n d c o l o u r ochreous-w h i t e , m a r k e d ochreous-w i t h s o m e ochreous-w h a t d i f f u s e longi-t u d i n a l s longi-t r e a k s or r a y s c o n s i s longi-t i n g of b l a c k and grey-black s c a l e s m i x e d w i t h dull p u r p l i s h red s c a l e s , c o m p r i s i n g b a s i c a l l y a s u b c o s t a l , a medial and a subdorsal s t r e a k , t h e s e m o r e or less c o n f l u e n t in basal area of w i n g , d i v e r g i n g d i s t a l l y and i n t e r r u p t e d before t e r m e n by a n a r r o w band of g r o u n d c o l o u r , t h e s u b c o s t a l stre?,k divided i n t o t w o p l u m e s and t h e medial s t r e a k into f o u r p l u m e s p o s t m e d i a l l y ; a s e r i e s of 5 or 6 b l a c k i s h I n t e r n e u r a l d o t s d i s p e r s e d along t e r m e n ; c i l i a g r e y , w i t h a d i f f u s e , blackish b r o w n subbasal l i n e . H l n d w i n g unif o r m ' y dark g r e y , t h e s c a l i n g r a t h e r t h i n -s p r e a d ; cilia u n i c o l o r o u -s , w i t h a b l a c k i -s h b r o w n subbasal l i n e . Legs o c h r e o u s - w h i t e , i r r o r a t e or d i f f u s e l y marked w i t h b l a c k i s h b r o w n e x t e r i o r l y . A b d o m e n o c h r e o u s - b r o w n , d a r k e r v e n t r a l l y ; male w i t h a light o c h r e o u s - b r o w n anal t u f t .
MALE GENITALIA ; Uncus conical and
s p a r s e l y s e t o s e , its apex reaching w e l l beyond
c o m p a r a t i v e l y small lateral p r o t u b e r a n c e s at b a s e . Gnathos w i t h medial process long and slender, hook-tipped. Valva c o m p a r a t i v e l y short and broad, t r u n c a t e d d i s t a l l y ; sacculus n a r r o w , reaching to about m i d d l e of v e n t r a l m a r g i n of valva, its u p p e r ( d o r s a l ) m a r g i n sharply produced at about m i d d l e . A e d e a g u s w i t h m e m b r a -nous apical part ( v e s i c a ) m i n u t e l y s p i c u l a t e . basal part w i t h o u t internal s c l e r o t i z a t i o n .
FEMALE GENITALIA ; O v i p o s i t o r e x t e n s i l e ,
pointed, w i t h Icbes (papillae anales) long ana n a r r o w . Eighth s t e r n i t e long and c o m p a r a t i v e l y slender; o s t i u m s i t u a t e d a n t e r i o r l y (on inter-segmental m e m b r a n e ) , ovate ( w i d e r than-b r o a d ) ; s t e r i g m a f o r m i n g a t h i n , w e a k l y scle-rotized r i m ; d u c t u s bursae and bursa c o p u l a t r i x v e r y w e a k l y d e v e l o p e d , m e m b r a n o u s .
MATERIAL EXAMINED
HOLOTYPE S: Brazil, Upper A m a z o n ,
Co-dajas, i v . 1 9 0 7 ( S . M . K l a g e s ) ; abdomen in situ. Paratypes: Brazil, M a n a u s , x i . 1919 (Parish) 1 9 (genitalia slide 13933); M a n a u s , 1 2 . i v . 1978, ex Brady pus s p . ( J . K. W a a g e ) , 4 $
(genitalia slide 13887), 1 5 ( g e n i t a l i a slide 13891); M a n a u s , 1 9 . i i i . 1 9 7 8 , ex Bradypus s p . ( R . C . B e s t ) , 1 9 ; same l o c a l i t y data but dated 2 7 . i i i . 1 9 7 8 , 1 <3; d i t t o 2 8 . i i i . 1 9 7 8 , 1 3 , 1 9 ; d i t t o 8. V Í. 1 9 7 9 , 1 S\ Parintins, 1 . x . 1919
( P a r i s h ) , 1 9 ( w i n g slide 13921, abdomen m i s s i n g ) ; Rio Japurá, lago A m a n a , 2.ix.1979, at light ( R . C . B e s t ) , 2 S: same l o c a l i t y data but dated 1 0 . i x . 1979, 3 τ • H o l o t y p e and 9 paratypes d e p o s i t e d in the B r i t i s h M u s e u m (Natural H i s t o r y ) ; 8 paratypes in t h e M u s e u m of Zoology of t h e U n i v e r s i t y , São Paulo: M u -seu Paraense Emílio G o e l d i , B e l é m , and Insti-t u Insti-t o Nacional de Pesquisas da A m a z ó n i a , Ma-n a u s .
Cryptoses rufipictus
s p . n .
(Figs. 5 - 9 )
w i N G S P A N : a 11-12 m m , 9 15-18 m m
Sexual d i m o r p h i s m m o d e r a t e l y p r o n o u n c e d , f e m a l e larger and w i t h general c o l o r a t i o n of f o r e w i n g darker; antennae s h o r t l y c i l i a t e in both s e x e s . Labial palpus, head, t h o r a x ,
pata-g i u m and tepata-gula pata-g r e y i s h o c h r e o u s , palpus s p r i n k l e d w i t h dark p u r p l i s h red e x t e r i o r l y , v e r t e x and f r o n s of head s u f f u s e d w i t h
b r o w n i s h r e d . In both sexes f o r e w i n g moder-a t e l y b r o moder-a d , w i t h moder-apex o b t u s e l y rounded moder-and t e r m e n m o d e r a t e l y o b l i q u e ; g r o u n d colour c r e a m , in male o v e r l a i d w i t h g r e y i s h o c h r e o u s w e a k l y t i n g e d w i t h o l i v e , in f e m a l e overlaid w i t h b r o w n i s h o c h r e o u s or r e d d i s h o c h r e o u s lacking olive t i n g e but o f t e n w i t h an a d m i x t u r e of s c a t t e r e d blackish-tipped s c a l e s , e x c e p t (in both s e x e s ) for poorly d e f i n e d , s l i g h t l y convex and sinuate a n t e m e d i a n and p o s t m e d i a n trans-verse lines, each line a r i s i n g f r o m a black s p o t on c o s t a , t h e p.pots being heaviest and m o s t c o n s p i c u o u s in the male and t h e t r a n s v e r s e lines o f t e n o b s c u r e and o b l i t e r a t e , w h i l e in the f e m a l e t h e p o s t m e d i a n line is o f t e n compara-t i v e l y clear and d i s compara-t i n c compara-t ; in bocompara-th sexes a f e w raised blackish scales in u p p e r part of cell a l i t t l e beyond a n t e m e d i a n line, s o m e t i m e s f o r m i n g a s t i g m a in m a l e ; costa n a r r o w l y edged w i t h o r a n g e - y e l l o w sparsely sprinkled w i t h black; b l a c k i s h i n t e r n e u r a l d o t s along t e r m s n ; cilia c r e a m - w h i t e , w i t h a purplish red
basal line reaching f r o m apex t o near t o r n u s , the r e d d i s h c o l o r a t i o n i n f u s i n g the area around t h e interneural dots along t e r m i n a l margin. C i l i a around t o r n u s w h o l l y s u f f u s e d w i t h grey.
H i n d w i n g u n i f o i m l y dark g r e y ; c i l i a cream-w h i t e , cream-w i t h a dark g r e y subbasal line, some-t i m e s f a i n some-t l y s u f f u s e d w i some-t h p u r p l i s h red f r o m apex to m i d d l e of t e r m e n , t h e n c e s u f f u s e d w i t h g r e y . U n d e r s i d e of h i n d w i n g g r e y , w i t h c o s t a l m a r g i n b r o a d l y s u f f u s e d w i t h purplish red and s p r i n k l e d w i t h black: a pair o f c l o s e l y a p p r o x i m a t e d , a l m o s t p a r a l l e l , b l a c k i s h subter-minal lines, t h e inner one h e a v i e s t . Legs paie o c h r e o u s , t i b i a e x t e n s i v e l y s u f f u s e d w i t h p u r p l i s h red e x t e r i o r l y e x c e p t t a r s a l s e g m e n t s , these marked w i t h b l a c k i s h a n n u l i . A b d o m e n g r e y i s h d o r s a l l y , pale o c h r e o u s m i x e d w i t h p u r p l i s h red v e n t r a l l y , w i t h a pale o c h r e o u s anal t u f t .
MALE GENITALIA; Uncus c o n i c a l and sparsely
Figs. 5 - 9 — Cryptoses rufipictus sp. n.: 5. imago, 3: 6, imago, $ ; 7, male genitalia, ventral aspect (aedeagus
quarter t o one t h i r d along v e n t r a l m a r g i n , its inner edge s m o o t h and e v e n . A e d e a g u s w i t h m e m b r a n o u s apical part ( v e s i c a ) m i n u t e l y s p i c u l a t e , basal part s t r o n g l y s c l e r o t i z e d in-t e r n a l l y , in-t e r m i n a in-t i n g apically in a long acicular process ( c o r n u t u s ) .
FEMALE GENITALIA ; O v i p o s i t o r e x t e n s i l e
slender, p o i n t e d , w i t h lobes (papillae anales) long and n a r r o w . Eighth s t e r n i t e c o m p a r a t i v e l y broad; o s t i u m s i t u a t e d a n t e r i o r l y (on inter-s e g m e n t a l m e m b r a n e ) , c i r c u l a r ; inter-s t e r i g m a
m e m b r a n o u s p o s t e r i o r l y , w e a k l y s c l e r o t i z e d a n t e r i o r l y and f o r m i n g a t h i n r i m . In some s p e c i m e n s e x a m i n e d t h e o s t i u m bursae had become i m p r e g n a t e d w i t h an opaque spherical mass of black m o u l d s p o r e s .
MATERIAL EXAMINED
HOLOTYPE $ : French Guiana, St Jean de
M a r o n i , ante 1939 (E. Le M o u l t ) ; abdomen in situ. Paratypes: same data as h o l o t y p e , 9 β ( g e n i t a l i a s l i d e s 13911, 13915, 13916), 20 5 ( g e n i t a l i a s l i d e s 13909, 13914); Brazil, R. Ma-r o n i , — 1916 (Le M . ) , 1 5 ( g e n i t a l i a slide 13910); M a n a u s , x i . 1919 ( P a r i s h ) , 1 $ \ M a -naus, 1 8 . i . 1 9 7 8 , ex Bradypus s p . ( R . C . B e s t ) , 1 ι\ same l o c a l i t y data but d a t e d 2 3 . i . 1978, 1 9 ; d i t t o 2 8 . i . 1978, 2 9 ; d i t t o 1 9 . i i i . 1 9 7 8 , 1 9 ; d i t t o 2 6 . i i i . 1 9 7 8 , 1 9 ; d i t t o 2 7 . H i . 1978, 4 $ , 2 3 ; d i t t o 28.iii.1978, 2 9 ; d i t t o 12.iv.1978. 3<3 (genitalia slide 13886), 3 9 (genitalia slide 1 3 8 9 0 ) ; Belém, 2 1 . v i . 1978, ex Bradypus s p .
( R . C . B e s t ) , 3(3 (genitalia s l i d e 13888), 4 9 ( g e n i t a l i a slides 13892, 13934); Rio Japurá. Lago A m a n a , 2 . i x . 1979, at light ( R . C . B e s t ) , 2$; s a m e locality data b u t dated 1 0 . i x . 1979:
3 ( 5 - H o l o t y p e and 46 paratypes d e p o s i t e d in t h e B r i t i s h Museum (Natural H i s t o r y ) ; 17 p a r a t y p e s in the M u s e u m of Zoology of t h e
U n i v e r s i t y , São Paulo, M u s e u Paraense Emílio G o e l d i , and Instituto Nacional de Pesquisas da A m a z ó n i a , M a n a u s .
C O M M E N T S AND DIFFRENTIAL DIAGNOSES
The imago of C . waagei is s u p e r f i c i a l l y v e r y s i m i l a r but on average smaller than that o f C . choloepi Dyar; the male of t h e latter c a n h a v e a wingspan of as little as 13 m m but
is more o f t e n nearer 15-17 m m , w h i l e the f e m a l e can reach 24 m m . Both species have a f o r e w i n g p a t t e r n c o n s i s t i n g o f radiating longitudinal s t r e a k s , but C- waagei is dis-t i n g u i s h e d by dis-the w e l l - d e f i n e d s u b dis-t e r m i n a l band w h i c h in C choloepi is p o o r l y defined and hardly d e t e r m i n a t e in t h e male and obliter-ated in t h e f e m a l e , and by t h e d i s t i n c t light-c o l o u r e d light-costal m a r g i n of the f o r e w i n g w h i light-c h in C. choloepi is s u f f u s e d w i t h b r o w n . C. rufi pictus d i f f e r s m a r k e d l y f r o m b o t h s p e c i e s in having a n t e m e d i a n and p o s t m e d i a n t r a n s v e r s e lines on the f o r e w i n g in lieu of t h e longitudinal s t r e a k s and s u b t e r m i n a l band of t h e o t h e r t w o s p e c i e s .
VENATION
(Figs. 10-15)
In both C. waagei (Figs. 10, 11) and C . rufipictus ( F i g s . 12, 13) v e i n 11 of t h e fore-w i n g is absent in t h e male but p r e s e n t in t h e f e m a l e , s i m i l a r l y as in C. choloepi (Figs. 14, 15) . In C. waagei and C. choloepi veins 3, A and 5 of the f o r e w i n g are s t a l k e d in both sexes, b u t in C . rufipicius t h e y are s t a l k e d in t h e f e m a l e only and are e i t h e r connate or c l o s e l y a p p r o x i m a t e in the m a l e . In t h e h i n d w i n g , t h e s t a l k i n g of v e i n s 7 and 8 v a r i e s in all three species but t h e s t a l k is usually longer in C-choloepi.
MALE GENITALIA
Figs. 10-17 — Wing venation: 10, Cryptoses waagei sp. n., $ ; 11, C. waagei sp. n., 2 ; 12, Cryptoses rufipictus sn.
n.. 5 : 13, C. rufipictus sp. n., 9 ; 14, Cryptoses choloeni ny a r. $; 15. C. choloepi Dyar. 9 ; 15, Bradypophila garbei
FEMALE GENITALIA
The s t r u c t u r e of the f e m a l e genitalia in ail t h r e e species is very s i m i l a r and d i f f e r e n c e s are m a i n l y c o m p a r a t i v e . In C. choloepi t h e e i g h t h s t e r n i t e is r e l a t i v e l y s h o r t and is less w e l l s c l e r o t i z e d than in the o t h e r t w o s p e c i e s ; t h e o s t i u m is m e m b r a n o u s and c i r c u l a r as in C. rufipictus but is relatively larger. C. waagei d i f f e r s f r o m both species by its broader, ovate o s t i u m .
OTHER CHRYSAUGINAE ASSOCIATED W I T H
TREE SLOTHS
Two f u r t h e r chrysaugine s p e c i e s , Bradyphi la garbei Iherir.g and Bradypodicola hahneli
Spuler, are k n o w n t o l i v e in association w i t h t r e e s l o t h s in Brazil and adjacent c o u n t r i e s in South A m e r i c a . The imagos of both s p e c i e s come w i t h i n t h e size-range of the three Cryp toses species but can be d i s t i n g u i s h e d by t h e f o l l o w i n g c h a r a c t e r s .
B. garbei is d i s t i n c t i v e by i t s a l m o s t uni-colorous dark o c h r e o u s f o r e w i n g s , w h i c h lack t h e m a r k i n g s found in Cryptoses. The w i n g venation ( F i g . 16) is reduced and s i m p l i f i e d , v e i n 2 being absent in t h e f o r e w i n g and t h e stalk of veins 7 and 8 a n a s t o m o s i n g w i t h v e i n 6 in iho h i n d w i n g . A s in t h e genus Cryptoses, vein 11 of t h e f o r e w i n g is a b s e n t in the male but present in t h e f e m a l e .
B. hahneli d i f f e r s f r o m all t h e s p e c i e s m e n t i o n e d above by i t s a l m o s t unicolorous dark fuscous f o r e w i n g s , t h e c o l o r a t i o n being d i s t i n c t l y more blackish than in B. garbei. A d i s t i n c t i v e feature of B. hahneli is t h e c o m -pressed f o r m of the head, t h e f r o n s of w h i c h is f l a t t e n e d and receding and t h e v e r t e x p r o m i -n e -n t ; i-n Cryptoses and Bradypophila t h e f r o n s is convex and the v e r t e x is not p r o m i n e n t .
In both 6 . hahneli and 3. garbei t h e pro-b o s c i s is shorter than in Cryptoses and is less e x t e n s i v e l y scaled at the base and t h e labial palpi are comparatively rough-scaled and droop m o r e or less straight d o w n w a r d s . In Cryptoses t h e p r o b o s c i s is very long and the basal p o r t i o n
is e x t e n s i v e l y s c a l e d , t h e labial palpi appear slender, the scales being a p p r e s s e d , and the palpi are held in a d e c l i v e n t , p o r r e c t or s l i g h t l y recurved p o s i t i o n .
A s a r e s u l t of e x a m i n i n g e x t e n s i v e sloth-m o t h sloth-material i t has b e c o sloth-m e apparent t h a t , in contrast to t h e v i r t u a l l y p r i s t i n e c o n d i t i o n of captured imagos of Cryptoses and B. garbei, s p e c i m e n s of B. hahneli are o f t e n w o r n and t h e e x t r e m i t i e s of t h e w i n g s s o m e t i m e s t a t t e r e d . It s e e m s that t h e o u t e r margins and f r i n g e s of the w i n g s of t h i s species may be w e a k and t h i n l y scaled and that t h e y could be damaged by m o v e m e n t t h r o u g h t h e fur of t h e s l o t h .
The male genitalia and w i n g v e n a t i o n s of B. garbei and B. hahneli are i l l u s t r a t e d by Lima ( 1 9 4 9 ) .
ACKNOWLEDGEMENTS
I am p a r t i c u l a r l y g r a t e f u l t o Dr. J . K. Waage of Imperial College Field S t a t i o n , S i l w o o d Park, A s c o t , Berkshire, England, w h o i n i t i a l l y made available f o r s t u d y l e p i d o p t e r o u s material c o l l e c t e d d u r i n g f i e l d w o r k on s l o t h s in South A m e r i c a , and also t o Dr. Nelson Bernardi of the M u s e u de Universidade de São Paulo. Brazil, w h o k i n d l y arranged t h e loan of type material of Bradypophila garbei f o r c o m p a r a t i v e s t u d y , and t o M r . Robin C. Best of t h e Instituto Nacional de Pesquisas da A m a z ô n i a , Manaus, Brazil and Dr. J . A d i s of t h e Max Planck I n s t i t u t e of L i m n o l o g y , Plõn, W e s t Germany, both of w h o m s u b s e q u e n t l y p r o v i d e d additional s l o t h - m o t h m a t e r i a l f o r s t u d y .
RESUMO
REFERENCES
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