1976
Gemetchu : Biology of Phlebotomus longipes
671
J. M e d . Ent.
20 February 1976
Vol. 12, n o . 6: 671-678
SALT MARSH TABANIDAE (DIPTERA): COMPARISON OF ABUNDANCE AND DISTRIBUTION IN SPARTINA AND JUNCUS HABITATS 1
By W . E. D a l e a n d R. C. A x t e l P Abstract: Greater diversity in species and higher numbers of adult female Tabanidae were collected (sticky traps and box traps) in Spartina than in Juncus marshes in the central coast of North Carolina during a 2-yr study. The 3 most abundant species for both types of marshes were Tabanus nigrovittatus Macquart, Chrysops atlanticus Pechuman, and C. fuliginosus 'This research was supported in part by an Agency for International Development, U. S. Department of State fellowship and by NOAA Office of Sea Grant, Department of Commerce, under Grant No. 04-3-158-40. Paper No. 4539 of the Journal Series of the North Carolina Agricultural Experiment Station. 'Predoctoral Fellow and Professor, respectively. Present address of W.E.D.: Department of Entomology, Universidad Nacional Agraria, La Molina, Lima, Peru.
Wiedemann. T. nigrovittatus was collected from 10 May to 25 September and had 1 peak of abundance from 5 to 14 June; C. atlanticus was collected from 23 April to 26 September, with 1 peak of abundance from 31 May to 8 June; C. fuliginosus was present from 8 April to 8 June, with 1 peak of abundance in the 1st week of May. Horizontal distribution in Spartina marsh habitats was determined by sticky traps. The highest numbers of T. nigrovittatus were trapped in the marsh; the lowest were obtained from the upland behind and inside nearby woods. Highest numbers of C. atlanticus were collected in the upland behind the woods and slightly less in the inner marsh; the lowest numbers were obtained from the upland inside the woods, the ecotone and the outer marsh. Highest numbers of C. fuliginosus were trapped in the inner marsh and the lowest from upland behind the woods.
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490-501. and peritrophic membrane of the adult female, Phlebotomus longipes Parrot and Martin (Diptera: Psychodidae). Ann. Macdonald, G. 1957. Epidemiology and control of malaria. Trop. Med. Parasitol. 68: 111-24. Oxford University Press, London. 201 p. McConnell, E. 1964. Leishmaniasis in the Sudan Republic. Gillies, M. T. 1956. A new character for the recognition of 17. Infections in Phlebotomus orientalis Parrot (Diptera: nulliparous females of Anopheles gambiae. Bull. Wld Hlth Psychodidae) after feeding on human kala azar patient. Organ. 15: 451-59. J. Trop. Med. Hyg. 67: 88-89. 1964. The study of longevity in biting insects. Int. Reo. Gen. Mellanby, K. 1946. Man's reaction to mosquito bites. Ext. Zool. 1: 47-77. Nature, Lond. 158: 554-55. Hertig, M. & P. T. Johnson. 1961. The rearing of PhleMutinga, M. 1971. Phlebotomus longipes, a vector of cutabotomus sandflies (Diptera: Psychodidae) I. Technique. neous leishmaniasis in Kenya. Trans. Roy. Soc. Trop. Med. Ann. Ent. Soc. Amer. 54: 753-64. Hyg. 65: 106-07. Hertig, M. & E. McConnell. 1963. Experimental infection of Panamanian Phlebotomus sandflies with Leishmania. Exp. Newstead, R. 1911. The papataci flies (Phlebotomus) of the Maltese Islands. Bull. Ent. Res. 2: 47-78. Parasitol. 14: 92-106. Roth, L. M. 1948. A study of mosquito behaviour. An Heyneman, D. 1963. The comparison of experimental experimental laboratory study of the sexual behaviour of Leishmania donovani infections in Phlebotomus papatasi (DipAedes aegypti (L.). Amer. Midi. Natur. 40: 265-352. tera: Psychodidae) with natural infections found in manShortt, H. E., A. C. Craighead & C. S. Swaminath. 1928. baited P. orientalis captured in kala azar endemic region A brief resume of recent kala azar research with special of the Sudan. Amer. J. Trop. Med. Hyg. 12: 725-40. reference to India. Indian J. Med. Res. 16: 221-39. Hindle, E. 1931. The development of various strains of Solomon, M. E. 1951. Control of humidity with potassium Leishmania in Chinese sandflies. Proc. Roy. Soc. B 108: hydroxide, sulphuric acid, or other solutions. Bull. Ent. 366-83. Res. 42: 543-54. Johnson, P. T. & M. Hertig. 1961. The rearing of Phlebotomus sandflies (Diptera: Psychodidae). II. Development Theodor, O. 1934. Observations on the hibernation of Phlebotomus papatasii (Dipt.). Bull. Ent. Res. 25: 459-72. and behaviour of Panamanian sandflies in laboratory 1936. On the relation of Phlebotomus papatasii to the temculture. Ann. Ent. Soc. Amer. 56: 764-76. perature and humidity of the environment. Bull. Ent. Res. Kirk, R. & D. J. Lewis. 1951. The Phlebotominae of the 27: 653-71. Ethiopian Region. Trans. Roy. Ent. Soc. Land. 102: 383-510. Wanson, M. 1942. Sur la biologie des Phlebotomes ConL e m m a , A., W. A. Foster, T. Gemetchu, P. M. Preston, golais. Rec. Trav. Sci. Med. Congo Belg. 1: 23-44. A. Bryceson & D. M. Minter. 1969. Studies on Waterson, J. 1922. A contribution to the knowledge of the leishmaniasis in Ethiopia. I : Preliminary investigations into bionomics of sandflies. Ann. Trop. Med. Parasitol. 16: 69-92. the epidemiology of cutaneous leishmaniasis in the highWilcocks, C. & P. E. C. Manson-Bahr. 1972. Manson's lands. Ann. Trop. Med. Parasitol. 63: 455-73. tropical diseases. Bailliere Tindall, London. 1164 p. Lewis, D. J., M. J. Mutinga & R. W. Ashford. 1972. Phlebotomus longipes Parrot & Martin (Diptera: Phleboto- Williams, P. 1966. Experimental transmission of Leishmania mexicana by Lutzomyia cruciata. Ann. Trop. Med. Parasitol. 60: midae) and a new related species. / . Ent. (B) 41: 119-24. 365-73. Martin, R. 1939. Observations sur les Phlebotomus D'Ethiopie. (Deuxième memoire). Arch. Inst. Pasteur Alg. 17:
J. Med. Ent.
672
R e c o r d s on seasonal a b u n d a n c e a n d h a b i t a t p r e f e r e n c e s for a d u l t s of t h e s e species a r e l i m i t e d in N o r t h C a r o l i n a (Brimley 1938), a l t h o u g h r e c e n t s t u d i e s h a v e r e v e a l e d t h e d i s t r i b u t i o n of t a b a n i d l a r v a e in various m a r s h h a b i t a t s (Dukes et al. 1974a, b ) . T h i s study was c o n d u c t e d to c o m p a r e the a b u n d a n c e of t h e a d u l t t a b a n i d s i n Spartina a n d Juncus salt m a r s h e s i n c o a s t a l N o r t h C a r o l i n a a n d t o d e t e r m i n e the seasonal a b u n d a n c e a n d horizontal d i s t r i b u t i o n of T. nigrovittatus, C. atlanticus a n d C. fuliginosus, w h i c h w e r e t h e m o s t c o m m o n species collected in p r e l i m i n a r y studies.
Spartina salt marshes. T h e 2 a r e a s ( A a n d B) w e r e 3 0 0 m a p a r t a l o n g N e w p o r t R i v e r , 3 k m N W of M o r e h e a d City. A r e a A was studied in 1971. T h e marsh was 125-140 m wide. A m a n - m a d e drainage d i t c h , 3 - 4 m w i d e , b u i l t for m o s q u i t o c o n t r o l , r a n in a S W to N E direction in the u p l a n d perimeter a n d was connected to N e w p o r t R i v e r b y a n a t u r a l d i t c h , 2 m w i d e . T h e e c o t o n e w a s 1-10 m w i d e a n d c o n t a i n e d spoil piles a t r e g u l a r i n t e r v a l s . The woods was 53-62 m wide. Area B was studied in 1972. T h e m a r s h w a s 7 6 - 1 0 0 m w i d e . T h e s a m e m a n - m a d e d i t c h as f o u n d i n a r e a A o c c u r r e d a l o n g t h e u p l a n d l i m i t s . T h e e c o t o n e w a s 1-10 m w i d e a n d h a d r e g u l a r l y p l a c e d spoil piles. T h e w o o d s was 126-148 m wide. I n both areas, the m a r s h plant cover was 9 8 % S. alterniflora (x = 2 7 0 p l a n t s / m ; m e a n h e i g h t = 1 m ) a n d 2 % J. roemerianus ( m o s t l y i n p a t c h e s n e a r the upland). T h e plants in the ecotone were m o s t l y J. roemerianus, Spartina patens ( A i t o n ) M u h l e n b e r g a n d Distichlis spicata ( L i n n a e u s ) . Pinus taeda L i n n a e u s w a s t h e d o m i n a n t species o n t h e spoil piles. T h e w o o d s c o n s i s t e d of m o s t l y P. taeda, Cornus florida L i n n a e u s a n d Wisteria sinensis (Sims) Sweet. 2
Juncus salt marshes.
A r e a C, studied in 1971, was
l o c a t e d a t t h e m i d d l e of a p e n i n s u l a (3.2 k m w i d e b y 5.6 k m
long), flanked
Sound, 24 k m N E from Davis. marshland
by Jarrett
of M o r e h e a d
Bay
and
Core
City a n d 800 m
T h e s t u d y a r e a w a s p a r t of a n e x t e n s i v e which
covered m o r e
than
1/2
of
the
peninsula. Within the marsh, there were 2 parallel M A T E R I A L S A N D METHODS
m a n - m a d e ditches 5 m wide, r u n n i n g in a S W to
Studies were conducted during the spring, s u m m e r a n d fall of 1971 a n d 1972 i n t h e v i c i n i t y of M o r e h e a d City, Carteret County, N o r t h Carolina.
N E direction; p e r p e n d i c u l a r to these m a i n ditches
Description
of study sites
F o u r salt m a r s h e s a n d a d j a c e n t a r e a s w e r e s t u d i e d .
a n d a t 2 0 0 - m i n t e r v a l s w a s a series of m a n - m a d e ditches (2-3 m wide) forming a drainage
network
for m o s q u i t o c o n t r o l .
11-84 m
wide.
T h e ecotone was
T h e woods was ca 800 m
wide. 2
A p u r e s t a n d of J. roemerianus (X = 3 1 3 6 s t e m s / m ;
E a c h study area could be divided into 3 habitat types: the
(1) t h e l o w e s t p o r t i o n s of t h e m a r s h n e a r e s t
water
sources,
flooded
twice
a
day,
with
m e a n height =
1.3 m ) c o v e r e d 6 1 % of t h e s u r f a c e .
N e a r t h e e c o t o n e t h e r e w a s a m i x t u r e of J.
roemeri virginica
anus, Fimbristilisspadicea
( L i n n a e u s ) , Salicornea
d o m i n a n t v e g e t a t i o n ; (2) p o r t i o n s a t slightly h i g h e r
Linnaeus
glauca
elevations,
T h e e c o t o n e p l a n t c o v e r w a s c o m p o s e d of J.
Spartina
Juncus
alterniflora L o i s e l e u r ( s m o o t h c o r d g r a s s ) t h e flooded
roemerianus
at
irregular
intervals,
with
Scheele (black needle rush)
the
d o m i n a n t v e g e t a t i o n ; a n d (3) p o r t i o n s a t t h e h i g h e s t elevations, n o t flooded, w i t h pine-deciduous woods the dominant vegetation.
Between the marsh a n d
and
Setaria
anus a n d D. spicata. P.
contained
the marsh proper.
wide. areas
were
alterniflora a n d 2 w e r e m o s t l y Juncus
mostly
Spartina
roemerianus.
Hoop
H o l e c r e e k , i n f r o n t of B o g u e S o u n d a n d 2 k m
m e d i a t e e l e v a t i o n s ; t h i s w a s less often f l o o d e d t h a n study
roemeri
T h e w o o d s c o n s i s t e d of m o s t l y
taeda.
of A t l a n t i c B e a c h .
of t h e
Beauvois.
A r e a D , studied in 1972, w a s l o c a t e d a t
woods was a transition zone or ecotone, at inter
Two
(Linnaeus)
no
W
It, a n d the neighboring marsh,
ditches.
The
ectone
was
T h e woods was 50-150 m wide. 2
0.5-4 m
A mixture
of J. roemerianus (x = 2 1 1 0 s t e m s / m ; m e a n h e i g h t 0.3 m ) a n d D. spicata
=
c o v e r e d 7 3 % of t h e m a r s h .
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Tabanus nigrovittatus M a c q u a r t , Chrysops atlanticus Pechuman a n d Chrysops fuliginosus Wiedemann c a u s e c o n s i d e r a b l e a n n o y a n c e b e c a u s e of t h e i r b i t e s a n d persistent attacks on h u m a n s a n d animals in salt m a r s h e s a n d a d j a c e n t a r e a s a l o n g t h e e a s t e r n c o a s t of t h e U n i t e d S t a t e s ( S t o n e 1938, P h i l i p 1947, 1 9 6 5 ) . I n a s u r v e y of p u b l i c o p i n i o n c o n d u c t e d o n t h e c o a s t of N o r t h C a r o l i n a , G e r h a r d t e t a l . (1973) f o u n d t h a t , a m o n g t h e b i t i n g D i p t e r a , tabanids were considered 2 n d to mosquitoes in i m p o r t a n c e as h u m a n p e s t s . T h e a b o v e species h a v e n o t b e e n s h o w n t o b e i m p o r t a n t v e c t o r s of p a t h o g e n s , a l t h o u g h G o l d f i e l d et a l . (1968) i s o l a t e d t h e v i r u s of w e s t e r n e q u i n e e n c e p h a l i t i s f r o m T. nigrovittatus i n N e w J e r s e y a n d s u g g e s t e d t h a t this species m a y b e a b l e to transmit eastern e q u i n e encephalitis virus a m o n g horses. R e l a t e d coastal species c a n t r a n s m i t h o g c h o l e r a ( T i d w e l l e t a l . 1972).
Vol. 12, no. 6
Dale & Axtell:
1976
673
Tabanidae abundance and distribution
A m i x t u r e of S. alterniflora, D. spicata a n d S. virginica c o v e r e d t h e r e m a i n d e r of t h e m a r s h . T h e p l a n t s i n t h e e c o t o n e w e r e m o s t l y D. spicata a n d Borrichia frutescens (Linnaeus) de Candolle. T h e woods c o n s i s t e d of m o s d y / . vomitoria, Quercus virginiana M i l l e r a n d Juniperus virginiana L i n n a e u s .
collecting container
Trapping methods and trap arrangement T h e h o r i z o n t a l d i s t r i b u t i o n of t h e t a b a n i d s , as w e l l as t h e i r a b u n d a n c e , w a s m e a s u r e d b y s t i c k y traps (Joyce & H a n s e n s 1968). E a c h t r a p consisted of 4 b l a c k m a s o n i t e 0.3 b y 0.3 m p a n e l s (1 f t ) , m o u n t e d o n a w o o d e n r e d p o l e (FIG. l ) . T h e b a s e of t h e p a n e l s w a s 1.2 m a b o v e t h e g r o u n d . T h e 8 surfaces e x p o s e d b y e a c h t r a p w e r e r e c o a t e d w i t h Stickem Special® , at weekly intervals. Additional d a t a on the relative a b u n d a n c e were obtained by b o x t r a p s (FIG. 2), m o d i f i e d f r o m G r a n g e r ( 1 9 7 0 ) . T h e t r a p s w e r e p a i n t e d b l a c k . T h e b a s e of t h e b o x w a s 1 m a b o v e t h e g r o u n d a n d t h e flies e n t e r e d t h r o u g h t h e o p e n b o t t o m . A p i e c e of N o Pest® strip containing dichlorvos insecticide was placed i n s i d e t h e c o l l e c t i n g c o n t a i n e r t o kill t h e flies. 2
3
0
FIG. 2.
Spartina
0.15
(UOm
Diagram of a box trap.
salt marshes.
I n a r e a A , 18 sticky t r a p s
w e r e a r r a n g e d i n 3 p a r a l l e l r o w s (30 m a p a r t ) f r o m the u p l a n d out into the marsh.
There were 3 traps
behind the woods, 3 in the woods, 3 in the ecotone, a n d 9 in the marsh.
T h e distance between adjacent
traps within a row was 30 m .
A box trap
located in the woods a n d another in the
was
marsh;
b o t h w e r e 3 0 m f r o m t h e n e a r e s t sticky t r a p .
In
a r e a B , 15 sticky t r a p s w e r e a r r a n g e d i n 3 p a r a l l e l
0.15
0.30 m
r o w s (30 m marsh.
apart) from
the upland out into
the
T h e r e were 3 traps behind the woods, 3
in the woods, 3 in the ecotone, a n d 6 in the m a r s h . T h e distance between adjacent traps within a row FIG. 1.
Diagram of a sticky trap.
'Michel & Pelton Co., Oakland, Calif. 94608, U.S.A. Shell Chemical Co., Agricultural Division, San Ramon, Calif. 94583, U.S.A. 4
was 30 m in the m a r s h - e c o t o n e a n d 6 0 - 9 0 m in a n d behind the woods.
A box trap was located in the
ecotone a n d a n o t h e r in the m a r s h ; b o t h w e r e 30 m f r o m t h e n e a r e s t sticky t r a p .
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4
674
J. Med. Ent.
Vol. 12, no. 6
TABLE 1. Total numbers and relative frequency (%) of adult Tabanidae collected by sticky and box traps in 2 types of salt marshes in 1971 and 1972. TYPE OF SALT MARSH
Spartina (AREAS A & B)
Sticky trap Chrysops fuliginosus Wiedemann Tabanas nigrovittatus Macquart Chrysops atlanticus Pechuman Leucotabanus annulatus (Say) Chrysops brimleyi Hine Tabanus lineóla Fabricius Hybomitra lasiophthalma (Macquart) Diachlorus ferrugatus (Fabricius) ChrysopsflavidusWiedemann Hybomitra daeckei (Hine) Chrysops obsoletas Wiedemann Tabanus pumilus Macquart Other species Totals
salt marshes.
(49.7) (42.5) ( 5.5) ( 0.6) ( 0.4) ( 0.3) ( 0.3) ( 0.1) ( 0.1) ( 0.1) 5 2 ( 0.1) 51 ( 0.1) 78 56,348
I n a r e a C , 2 4 sticky
traps
w e r e a r r a n g e d i n 3 p a r a l l e l r o w s (30 m a p a r t a n d 30 m b e t w e e n adjacent traps within a row).
There
were 6 traps in the woods, 9 in the ecotone a n d 9 in the marsh.
A b o x t r a p (50 m f r o m t h e sticky
traps) was located in the woods, ecotone a n d marsh. A r e a D h a d 9 sticky t r a p s a r r a n g e d i n 3 p a r a l l e l r o w s (30 m a p a r t , 6 0 m b e t w e e n t r a p s w i t h i n a r o w ) a n d all w e r e l o c a t e d
in
the marsh.
Two
box
traps
(65 m a p a r t ) w e r e in t h e m a r s h a n d 50 m from t h e n e a r e s t sticky t r a p s . Data
collection and
14 6369 36 1 0 65 3 1 0 0 6 4 48 6547
Sticky trap
( 0.2) (97.3) ( 0.6)
2519 7610 4631 23 0 61 3 48 6 28 0 1 28 14,959
( 1.0)
( 0.1) ( 0.1)
( 0.4) ( 0.3) ( 0.2)
i n v o l v e d d a t a f r o m 31 M a y t o 2 0 J u l y i n a r e a A a n d f r o m 4 J u n e t o 10 A u g u s t i n a r e a B for T. nigrovittatus, f r o m 31 M a y t o 11 J u n e i n a r e a A for C. atlanticus a n d f r o m 6—18 M a y i n a r e a A for C. fuliginosus. T h e n u m b e r of flies p e r species t a k e n o n e a c h of t h e c o l l e c t i o n d a t e s w a s i n d e p e n d e n t l y a n a l y z e d as a r a n d o m i z e d complete block, with 6 treatments ( t r a p p o s i t i o n s ) a n d 3 r e p l i c a t e s (rows) for a r e a A , a n d 5 t r e a t m e n t s a n d 3 r e p l i c a t e s for a r e a B . Honestly significant difference (h.s.d.) values ( T u k e y ' s test) a t t h e 0 . 0 5 l e v e l of s i g n i f i c a n c e w e r e u s e d for p a r t i a l 2 - t r e a t m e n t c o m p a r i s o n s .
analysis
D u r i n g p e a k fly p o p u l a t i o n s , b o t h t y p e s of t r a p s were e x a m i n e d twice a week; at other times, counts were at weekly intervals.
T h e flies i n m o s t cases
w e r e identified
i n t h e field.
counted;
few
very
Box trap 6 ( 0.3) 1897 (95.4) 39 ( 2.0) 0 0 18 ( 0.9) 0 0 0 0 0 0 29 1988
(16.8) (50.9) (31.0) ( 0.2)
males
O n l y females
were
trapped.
RESULTS A N D DISCUSSION
Comparison of Spartina
and Juncus
marshes
D u r i n g this 2-year study, 79,842 a d u l t
tabanids
were
b e l o n g i n g t o 32 species a n d 6 g e n e r a w e r e c o l l e c t e d .
After
T h e most a b u n d a n t
( m o r e t h a n 50) a r e listed i n
c o u n t i n g , all flies w e r e r e m o v e d a n d t h e S t i c k e m ®
T A B L E 1.
renewed.
w e r e c o l l e c t e d a n d i n t h e Juncus s a l t m a r s h e s o n l y
Relative
the
2 0 species.
combing
the
species i n t h e g e n u s Tabanus
c o l l e c t i o n d a t a for t h e e n t i r e s e a s o n for a r e a s A
and
of m a r s h
abundance
was
according
salt m a r s h e s 31 species
to
type
tabanid
I n t h e Spartina
determined
by
B (Spartina s a l t m a r s h e s ) a n d a r e a s C a n d D salt m a r s h e s ) .
(Juncus
T h e r e l a t i v e a b u n d a n c e of i n d i v i d u a l
species w a s d e t e r m i n e d b y s u m m i n g t h e d a t a c o l
I n t h e Juncus m a r s h e s t h e n u m b e r o f w a s 4 4 % less a n d i n
Chrysops 3 8 % less t h a n i n t h e Spartina m a r s h e s . Spartina
salt
marshes
produced
t a b a n i d s t h a n t h e Juncus m a r s h e s .
3.7
The
times
more
Davis &
(1966) f o u n d a b o u t 4 0 t i m e s m o r e insects i n
Gray
Spartina
l e c t e d b y b o t h t y p e s of t r a p s for all t h e a r e a s a n d
t h a n i n Juncus m a r s h e s b u t t h e i r d a t a i n c l u d e d v e r y
e x p r e s s i n g t h e r e s u l t s a s p e r c e n t a g e of t h e
few
total.
O n l y t h e sticky t r a p c o l l e c t i o n d a t a w e r e u s e d t o express
the
seasonal
abundance
for
the
3
most
c o m m o n species. The
horizontal
tabanids
(due
to
the
inadequate
sampling
methods). T h e 3 m o s t a b u n d a n t species i n o u r
collections
( a c c o u n t i n g for 9 8 % of t h e t o t a l ) for b o t h t y p e s of distributions
C. atlanticus a n d C. fuliginosus
of
T.
nigrovittatus,
i n Spartina salt m a r s h e s
m a r s h e s w e r e T. nigrovittatus, atlanticus.
These
species,
C. fuliginosus,
along
with
T.
a n d C. lineola
( a r e a s A a n d B) w e r e a n a l y z e d b y u s i n g d a t a f r o m
Fabricius, have been reported to predominate
t h e sticky t r a p s for c o n s e c u t i v e c o l l e c t i o n d a t e s i n
o t h e r s a l t m a r s h e s of t h e A t i a n t i c c o a s t of t h e U n i t e d
which
States ( J a m n b a c k
of a
given
species p e r t r a p p e r d a y w e r e 2 o r m o r e .
the
average
numbers
of flies
This
1960).
& Wall
1959, W a l l
in
&
Doane
I n o u r a r e a s , t h e r e w e r e v e r y few T.
lineola.
Downloaded from http://jme.oxfordjournals.org/ by guest on June 5, 2016
Juncus
28,030 23,942 3081 398 219 161 158 69 55 54
Juncus (AREAS C & D)
Box trap
Dale & Axtell:
1976
Seasonal
675
Tabanidae abundance and distribution
abundance
C. Juliginosus (FIG. 5) w a s c o l l e c t e d f r o m 8 A p r i l t o 8 J u n e . D a t a w e r e o b t a i n e d o n l y for a r e a s B a n d D . T h e e m e r g e n c e of t h e first a d u l t s c o i n c i d e d w i t h i n c r e a s i n g t e m p e r a t u r e s w i t h t h e a d v e n t of s u m m e r , as observed b y J a m n b a c k & W a l l (1959).
T h e r e w a s 1 p o p u l a t i o n p e a k , i n t h e 1st w e e k of M a y , followed b y a s h a r p decrease in the adults completely disappeared Hansens
&
Robinson
(1973)
numbers;
1 month
observed
in
later. New
J e r s e y t h a t after t h e p e a k w a s r e a c h e d , a c t i v i t y for this species d e c l i n e d
abruptly.
T h e s e a s o n a l a b u n d a n c e of t h e s e flies is i n f l u e n c e d by the changing weather, particularly the tempera t u r e . I t is a p p a r e n t f r o m c o m p a r i n g o u r d a t a t o previously published d a t a (Anderson 1971, Anderson & K n e e n 1969, B a i l e y 1947, Blickle 1954, B r e n n a n 1 9 3 5 , H a n s e n s 1952, H a n s e n s & R o b i n s o n 1 9 7 3 , J a m n b a c k 1969, J a m n b a c k & W a l l 1959, J o n e s & A n t h o n y 1964, M a c C r e a r y 1940, P e c h u m a n 1 9 5 7 , 1972, S t o n e 1938) t h a t T. nigrovittatus, C. atlanticus a n d C. fuliginosus e x h i b i t a d e l a y i n t h e t i m e of p e a k a b u n d a n c e from southern to n o r t h e r n latitudes along t h e e a s t e r n c o a s t of t h e U n i t e d S t a t e s . Horizontal
distribution
T. nigrovittatus c o l l e c t i o n s f r o m s t i c k y t r a p s s h o w e d significant differences a m o n g t h e t r a p p o s i t i o n s i n 8 0 % of t h e 15 c o l l e c t i o n p e r i o d s i n a r e a A i n 1971 a n d i n all of t h e 15 c o l l e c t i o n p e r i o d s i n a r e a B i n 1972 (TABLE 2 ) .
I n b o t h years t h e results
were
essentially t h e s a m e a n d t h e r e w a s n o c h a n g e in t h e d i s t r i b u t i o n of t h e flies d u r i n g t h e s e a s o n . flies
were
trapped
in
the
outer
marsh,
More nearest
FIG. 4. Average number of flies/trap/day for Chrysops atlanticus Pechuman, in Spartina (areas A & B) and Juncus (areas C & D) marshes during 1971 and 1972.
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T h e a b u n d a n c e of T. nigrovittatus i n Spartina a n d Juncus m a r s h e s is i l l u s t r a t e d i n FIG. 3. F o r b o t h Spartina m a r s h e s t h e r e w a s 1 p e a k of a b u n d a n c e f r o m 5 - 1 4 J u n e , a p p r o x i m a t e l y 1 m o n t h after t h e e m e r g e n c e of t h e first a d u l t s , f o l l o w e d b y a s u d d e n d e c r e a s e of t h e p o p u l a t i o n to l o w levels w h i c h w e r e m a i n t a i n e d u n t i l t h e e n d of t h e s u m m e r . T h i s is essentially t h e s a m e p a t t e r n d e s c r i b e d for M a s s a c h u setts b y W a l l & D o a n e ( 1 9 6 0 ) . A s i m i l a r p e a k o c c u r r e d i n t h e Juncus m a r s h a r e a C , a l t h o u g h n u m b e r s w e r e v e r y l o w . I n t h e o t h e r Juncus m a r s h (area D ) , there were fluctuations in the low n u m b e r s t h r o u g h o u t the season. C. atlanticus (FIG. 4) w a s c o l l e c t e d from 2 3 April t o 2 6 S e p t e m b e r . T h e r e w a s 1 p e a k of a b u n d a n c e , f r o m 31 M a y t o 18 J u n e . F i e l d o b s e r v a t i o n s o n b i t i n g a c t i v i t y for 2 c o n s e c u t i v e y e a r s r e v e a l e d a r e t u r n of h i g h n u m b e r s of flies d u r i n g t h e l a s t 1 / 2 of A u g u s t b u t t h i s w a s n o t s h o w n b y t h e t r a p d a t a . B y c o m p a r i s o n t o h i g h e r n e t t i n g c o u n t s , i t is a p p a r e n t t h a t t h e traps w e r e relatively p o o r collectors o f C. atlanticus.
J. Med. Ent.
676
Vol. 12, no. 6
CO © CO © IN CM CO CO O O 120 • O O CO CO CT) *-* O CO CM CO CM — — IN CO Q CO « O TO * CO CO TO TJ- CO 100' IN © IN CO © ©
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Gemetchu : Biology of Phlebotomus longipes
671
J. M e d . Ent.
20 February 1976
Vol. 12, n o . 6: 671-678
SALT MARSH TABANIDAE (DIPTERA): COMPARISON OF ABUNDANCE AND DISTRIBUTION IN SPARTINA AND JUNCUS HABITATS 1
By W . E. D a l e a n d R. C. A x t e l P Abstract: Greater diversity in species and higher numbers of adult female Tabanidae were collected (sticky traps and box traps) in Spartina than in Juncus marshes in the central coast of North Carolina during a 2-yr study. The 3 most abundant species for both types of marshes were Tabanus nigrovittatus Macquart, Chrysops atlanticus Pechuman, and C. fuliginosus 'This research was supported in part by an Agency for International Development, U. S. Department of State fellowship and by NOAA Office of Sea Grant, Department of Commerce, under Grant No. 04-3-158-40. Paper No. 4539 of the Journal Series of the North Carolina Agricultural Experiment Station. 'Predoctoral Fellow and Professor, respectively. Present address of W.E.D.: Department of Entomology, Universidad Nacional Agraria, La Molina, Lima, Peru.
Wiedemann. T. nigrovittatus was collected from 10 May to 25 September and had 1 peak of abundance from 5 to 14 June; C. atlanticus was collected from 23 April to 26 September, with 1 peak of abundance from 31 May to 8 June; C. fuliginosus was present from 8 April to 8 June, with 1 peak of abundance in the 1st week of May. Horizontal distribution in Spartina marsh habitats was determined by sticky traps. The highest numbers of T. nigrovittatus were trapped in the marsh; the lowest were obtained from the upland behind and inside nearby woods. Highest numbers of C. atlanticus were collected in the upland behind the woods and slightly less in the inner marsh; the lowest numbers were obtained from the upland inside the woods, the ecotone and the outer marsh. Highest numbers of C. fuliginosus were trapped in the inner marsh and the lowest from upland behind the woods.
Downloaded from http://jme.oxfordjournals.org/ by guest on June 5, 2016
490-501. and peritrophic membrane of the adult female, Phlebotomus longipes Parrot and Martin (Diptera: Psychodidae). Ann. Macdonald, G. 1957. Epidemiology and control of malaria. Trop. Med. Parasitol. 68: 111-24. Oxford University Press, London. 201 p. McConnell, E. 1964. Leishmaniasis in the Sudan Republic. Gillies, M. T. 1956. A new character for the recognition of 17. Infections in Phlebotomus orientalis Parrot (Diptera: nulliparous females of Anopheles gambiae. Bull. Wld Hlth Psychodidae) after feeding on human kala azar patient. Organ. 15: 451-59. J. Trop. Med. Hyg. 67: 88-89. 1964. The study of longevity in biting insects. Int. Reo. Gen. Mellanby, K. 1946. Man's reaction to mosquito bites. Ext. Zool. 1: 47-77. Nature, Lond. 158: 554-55. Hertig, M. & P. T. Johnson. 1961. The rearing of PhleMutinga, M. 1971. Phlebotomus longipes, a vector of cutabotomus sandflies (Diptera: Psychodidae) I. Technique. neous leishmaniasis in Kenya. Trans. Roy. Soc. Trop. Med. Ann. Ent. Soc. Amer. 54: 753-64. Hyg. 65: 106-07. Hertig, M. & E. McConnell. 1963. Experimental infection of Panamanian Phlebotomus sandflies with Leishmania. Exp. Newstead, R. 1911. The papataci flies (Phlebotomus) of the Maltese Islands. Bull. Ent. Res. 2: 47-78. Parasitol. 14: 92-106. Roth, L. M. 1948. A study of mosquito behaviour. An Heyneman, D. 1963. The comparison of experimental experimental laboratory study of the sexual behaviour of Leishmania donovani infections in Phlebotomus papatasi (DipAedes aegypti (L.). Amer. Midi. Natur. 40: 265-352. tera: Psychodidae) with natural infections found in manShortt, H. E., A. C. Craighead & C. S. Swaminath. 1928. baited P. orientalis captured in kala azar endemic region A brief resume of recent kala azar research with special of the Sudan. Amer. J. Trop. Med. Hyg. 12: 725-40. reference to India. Indian J. Med. Res. 16: 221-39. Hindle, E. 1931. The development of various strains of Solomon, M. E. 1951. Control of humidity with potassium Leishmania in Chinese sandflies. Proc. Roy. Soc. B 108: hydroxide, sulphuric acid, or other solutions. Bull. Ent. 366-83. Res. 42: 543-54. Johnson, P. T. & M. Hertig. 1961. The rearing of Phlebotomus sandflies (Diptera: Psychodidae). II. Development Theodor, O. 1934. Observations on the hibernation of Phlebotomus papatasii (Dipt.). Bull. Ent. Res. 25: 459-72. and behaviour of Panamanian sandflies in laboratory 1936. On the relation of Phlebotomus papatasii to the temculture. Ann. Ent. Soc. Amer. 56: 764-76. perature and humidity of the environment. Bull. Ent. Res. Kirk, R. & D. J. Lewis. 1951. The Phlebotominae of the 27: 653-71. Ethiopian Region. Trans. Roy. Ent. Soc. Land. 102: 383-510. Wanson, M. 1942. Sur la biologie des Phlebotomes ConL e m m a , A., W. A. Foster, T. Gemetchu, P. M. Preston, golais. Rec. Trav. Sci. Med. Congo Belg. 1: 23-44. A. Bryceson & D. M. Minter. 1969. Studies on Waterson, J. 1922. A contribution to the knowledge of the leishmaniasis in Ethiopia. I : Preliminary investigations into bionomics of sandflies. Ann. Trop. Med. Parasitol. 16: 69-92. the epidemiology of cutaneous leishmaniasis in the highWilcocks, C. & P. E. C. Manson-Bahr. 1972. Manson's lands. Ann. Trop. Med. Parasitol. 63: 455-73. tropical diseases. Bailliere Tindall, London. 1164 p. Lewis, D. J., M. J. Mutinga & R. W. Ashford. 1972. Phlebotomus longipes Parrot & Martin (Diptera: Phleboto- Williams, P. 1966. Experimental transmission of Leishmania mexicana by Lutzomyia cruciata. Ann. Trop. Med. Parasitol. 60: midae) and a new related species. / . Ent. (B) 41: 119-24. 365-73. Martin, R. 1939. Observations sur les Phlebotomus D'Ethiopie. (Deuxième memoire). Arch. Inst. Pasteur Alg. 17:
J. Med. Ent.
672
R e c o r d s on seasonal a b u n d a n c e a n d h a b i t a t p r e f e r e n c e s for a d u l t s of t h e s e species a r e l i m i t e d in N o r t h C a r o l i n a (Brimley 1938), a l t h o u g h r e c e n t s t u d i e s h a v e r e v e a l e d t h e d i s t r i b u t i o n of t a b a n i d l a r v a e in various m a r s h h a b i t a t s (Dukes et al. 1974a, b ) . T h i s study was c o n d u c t e d to c o m p a r e the a b u n d a n c e of t h e a d u l t t a b a n i d s i n Spartina a n d Juncus salt m a r s h e s i n c o a s t a l N o r t h C a r o l i n a a n d t o d e t e r m i n e the seasonal a b u n d a n c e a n d horizontal d i s t r i b u t i o n of T. nigrovittatus, C. atlanticus a n d C. fuliginosus, w h i c h w e r e t h e m o s t c o m m o n species collected in p r e l i m i n a r y studies.
Spartina salt marshes. T h e 2 a r e a s ( A a n d B) w e r e 3 0 0 m a p a r t a l o n g N e w p o r t R i v e r , 3 k m N W of M o r e h e a d City. A r e a A was studied in 1971. T h e marsh was 125-140 m wide. A m a n - m a d e drainage d i t c h , 3 - 4 m w i d e , b u i l t for m o s q u i t o c o n t r o l , r a n in a S W to N E direction in the u p l a n d perimeter a n d was connected to N e w p o r t R i v e r b y a n a t u r a l d i t c h , 2 m w i d e . T h e e c o t o n e w a s 1-10 m w i d e a n d c o n t a i n e d spoil piles a t r e g u l a r i n t e r v a l s . The woods was 53-62 m wide. Area B was studied in 1972. T h e m a r s h w a s 7 6 - 1 0 0 m w i d e . T h e s a m e m a n - m a d e d i t c h as f o u n d i n a r e a A o c c u r r e d a l o n g t h e u p l a n d l i m i t s . T h e e c o t o n e w a s 1-10 m w i d e a n d h a d r e g u l a r l y p l a c e d spoil piles. T h e w o o d s was 126-148 m wide. I n both areas, the m a r s h plant cover was 9 8 % S. alterniflora (x = 2 7 0 p l a n t s / m ; m e a n h e i g h t = 1 m ) a n d 2 % J. roemerianus ( m o s t l y i n p a t c h e s n e a r the upland). T h e plants in the ecotone were m o s t l y J. roemerianus, Spartina patens ( A i t o n ) M u h l e n b e r g a n d Distichlis spicata ( L i n n a e u s ) . Pinus taeda L i n n a e u s w a s t h e d o m i n a n t species o n t h e spoil piles. T h e w o o d s c o n s i s t e d of m o s t l y P. taeda, Cornus florida L i n n a e u s a n d Wisteria sinensis (Sims) Sweet. 2
Juncus salt marshes.
A r e a C, studied in 1971, was
l o c a t e d a t t h e m i d d l e of a p e n i n s u l a (3.2 k m w i d e b y 5.6 k m
long), flanked
Sound, 24 k m N E from Davis. marshland
by Jarrett
of M o r e h e a d
Bay
and
Core
City a n d 800 m
T h e s t u d y a r e a w a s p a r t of a n e x t e n s i v e which
covered m o r e
than
1/2
of
the
peninsula. Within the marsh, there were 2 parallel M A T E R I A L S A N D METHODS
m a n - m a d e ditches 5 m wide, r u n n i n g in a S W to
Studies were conducted during the spring, s u m m e r a n d fall of 1971 a n d 1972 i n t h e v i c i n i t y of M o r e h e a d City, Carteret County, N o r t h Carolina.
N E direction; p e r p e n d i c u l a r to these m a i n ditches
Description
of study sites
F o u r salt m a r s h e s a n d a d j a c e n t a r e a s w e r e s t u d i e d .
a n d a t 2 0 0 - m i n t e r v a l s w a s a series of m a n - m a d e ditches (2-3 m wide) forming a drainage
network
for m o s q u i t o c o n t r o l .
11-84 m
wide.
T h e ecotone was
T h e woods was ca 800 m
wide. 2
A p u r e s t a n d of J. roemerianus (X = 3 1 3 6 s t e m s / m ;
E a c h study area could be divided into 3 habitat types: the
(1) t h e l o w e s t p o r t i o n s of t h e m a r s h n e a r e s t
water
sources,
flooded
twice
a
day,
with
m e a n height =
1.3 m ) c o v e r e d 6 1 % of t h e s u r f a c e .
N e a r t h e e c o t o n e t h e r e w a s a m i x t u r e of J.
roemeri virginica
anus, Fimbristilisspadicea
( L i n n a e u s ) , Salicornea
d o m i n a n t v e g e t a t i o n ; (2) p o r t i o n s a t slightly h i g h e r
Linnaeus
glauca
elevations,
T h e e c o t o n e p l a n t c o v e r w a s c o m p o s e d of J.
Spartina
Juncus
alterniflora L o i s e l e u r ( s m o o t h c o r d g r a s s ) t h e flooded
roemerianus
at
irregular
intervals,
with
Scheele (black needle rush)
the
d o m i n a n t v e g e t a t i o n ; a n d (3) p o r t i o n s a t t h e h i g h e s t elevations, n o t flooded, w i t h pine-deciduous woods the dominant vegetation.
Between the marsh a n d
and
Setaria
anus a n d D. spicata. P.
contained
the marsh proper.
wide. areas
were
alterniflora a n d 2 w e r e m o s t l y Juncus
mostly
Spartina
roemerianus.
Hoop
H o l e c r e e k , i n f r o n t of B o g u e S o u n d a n d 2 k m
m e d i a t e e l e v a t i o n s ; t h i s w a s less often f l o o d e d t h a n study
roemeri
T h e w o o d s c o n s i s t e d of m o s t l y
taeda.
of A t l a n t i c B e a c h .
of t h e
Beauvois.
A r e a D , studied in 1972, w a s l o c a t e d a t
woods was a transition zone or ecotone, at inter
Two
(Linnaeus)
no
W
It, a n d the neighboring marsh,
ditches.
The
ectone
was
T h e woods was 50-150 m wide. 2
0.5-4 m
A mixture
of J. roemerianus (x = 2 1 1 0 s t e m s / m ; m e a n h e i g h t 0.3 m ) a n d D. spicata
=
c o v e r e d 7 3 % of t h e m a r s h .
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Tabanus nigrovittatus M a c q u a r t , Chrysops atlanticus Pechuman a n d Chrysops fuliginosus Wiedemann c a u s e c o n s i d e r a b l e a n n o y a n c e b e c a u s e of t h e i r b i t e s a n d persistent attacks on h u m a n s a n d animals in salt m a r s h e s a n d a d j a c e n t a r e a s a l o n g t h e e a s t e r n c o a s t of t h e U n i t e d S t a t e s ( S t o n e 1938, P h i l i p 1947, 1 9 6 5 ) . I n a s u r v e y of p u b l i c o p i n i o n c o n d u c t e d o n t h e c o a s t of N o r t h C a r o l i n a , G e r h a r d t e t a l . (1973) f o u n d t h a t , a m o n g t h e b i t i n g D i p t e r a , tabanids were considered 2 n d to mosquitoes in i m p o r t a n c e as h u m a n p e s t s . T h e a b o v e species h a v e n o t b e e n s h o w n t o b e i m p o r t a n t v e c t o r s of p a t h o g e n s , a l t h o u g h G o l d f i e l d et a l . (1968) i s o l a t e d t h e v i r u s of w e s t e r n e q u i n e e n c e p h a l i t i s f r o m T. nigrovittatus i n N e w J e r s e y a n d s u g g e s t e d t h a t this species m a y b e a b l e to transmit eastern e q u i n e encephalitis virus a m o n g horses. R e l a t e d coastal species c a n t r a n s m i t h o g c h o l e r a ( T i d w e l l e t a l . 1972).
Vol. 12, no. 6
Dale & Axtell:
1976
673
Tabanidae abundance and distribution
A m i x t u r e of S. alterniflora, D. spicata a n d S. virginica c o v e r e d t h e r e m a i n d e r of t h e m a r s h . T h e p l a n t s i n t h e e c o t o n e w e r e m o s t l y D. spicata a n d Borrichia frutescens (Linnaeus) de Candolle. T h e woods c o n s i s t e d of m o s d y / . vomitoria, Quercus virginiana M i l l e r a n d Juniperus virginiana L i n n a e u s .
collecting container
Trapping methods and trap arrangement T h e h o r i z o n t a l d i s t r i b u t i o n of t h e t a b a n i d s , as w e l l as t h e i r a b u n d a n c e , w a s m e a s u r e d b y s t i c k y traps (Joyce & H a n s e n s 1968). E a c h t r a p consisted of 4 b l a c k m a s o n i t e 0.3 b y 0.3 m p a n e l s (1 f t ) , m o u n t e d o n a w o o d e n r e d p o l e (FIG. l ) . T h e b a s e of t h e p a n e l s w a s 1.2 m a b o v e t h e g r o u n d . T h e 8 surfaces e x p o s e d b y e a c h t r a p w e r e r e c o a t e d w i t h Stickem Special® , at weekly intervals. Additional d a t a on the relative a b u n d a n c e were obtained by b o x t r a p s (FIG. 2), m o d i f i e d f r o m G r a n g e r ( 1 9 7 0 ) . T h e t r a p s w e r e p a i n t e d b l a c k . T h e b a s e of t h e b o x w a s 1 m a b o v e t h e g r o u n d a n d t h e flies e n t e r e d t h r o u g h t h e o p e n b o t t o m . A p i e c e of N o Pest® strip containing dichlorvos insecticide was placed i n s i d e t h e c o l l e c t i n g c o n t a i n e r t o kill t h e flies. 2
3
0
FIG. 2.
Spartina
0.15
(UOm
Diagram of a box trap.
salt marshes.
I n a r e a A , 18 sticky t r a p s
w e r e a r r a n g e d i n 3 p a r a l l e l r o w s (30 m a p a r t ) f r o m the u p l a n d out into the marsh.
There were 3 traps
behind the woods, 3 in the woods, 3 in the ecotone, a n d 9 in the marsh.
T h e distance between adjacent
traps within a row was 30 m .
A box trap
located in the woods a n d another in the
was
marsh;
b o t h w e r e 3 0 m f r o m t h e n e a r e s t sticky t r a p .
In
a r e a B , 15 sticky t r a p s w e r e a r r a n g e d i n 3 p a r a l l e l
0.15
0.30 m
r o w s (30 m marsh.
apart) from
the upland out into
the
T h e r e were 3 traps behind the woods, 3
in the woods, 3 in the ecotone, a n d 6 in the m a r s h . T h e distance between adjacent traps within a row FIG. 1.
Diagram of a sticky trap.
'Michel & Pelton Co., Oakland, Calif. 94608, U.S.A. Shell Chemical Co., Agricultural Division, San Ramon, Calif. 94583, U.S.A. 4
was 30 m in the m a r s h - e c o t o n e a n d 6 0 - 9 0 m in a n d behind the woods.
A box trap was located in the
ecotone a n d a n o t h e r in the m a r s h ; b o t h w e r e 30 m f r o m t h e n e a r e s t sticky t r a p .
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4
674
J. Med. Ent.
Vol. 12, no. 6
TABLE 1. Total numbers and relative frequency (%) of adult Tabanidae collected by sticky and box traps in 2 types of salt marshes in 1971 and 1972. TYPE OF SALT MARSH
Spartina (AREAS A & B)
Sticky trap Chrysops fuliginosus Wiedemann Tabanas nigrovittatus Macquart Chrysops atlanticus Pechuman Leucotabanus annulatus (Say) Chrysops brimleyi Hine Tabanus lineóla Fabricius Hybomitra lasiophthalma (Macquart) Diachlorus ferrugatus (Fabricius) ChrysopsflavidusWiedemann Hybomitra daeckei (Hine) Chrysops obsoletas Wiedemann Tabanus pumilus Macquart Other species Totals
salt marshes.
(49.7) (42.5) ( 5.5) ( 0.6) ( 0.4) ( 0.3) ( 0.3) ( 0.1) ( 0.1) ( 0.1) 5 2 ( 0.1) 51 ( 0.1) 78 56,348
I n a r e a C , 2 4 sticky
traps
w e r e a r r a n g e d i n 3 p a r a l l e l r o w s (30 m a p a r t a n d 30 m b e t w e e n adjacent traps within a row).
There
were 6 traps in the woods, 9 in the ecotone a n d 9 in the marsh.
A b o x t r a p (50 m f r o m t h e sticky
traps) was located in the woods, ecotone a n d marsh. A r e a D h a d 9 sticky t r a p s a r r a n g e d i n 3 p a r a l l e l r o w s (30 m a p a r t , 6 0 m b e t w e e n t r a p s w i t h i n a r o w ) a n d all w e r e l o c a t e d
in
the marsh.
Two
box
traps
(65 m a p a r t ) w e r e in t h e m a r s h a n d 50 m from t h e n e a r e s t sticky t r a p s . Data
collection and
14 6369 36 1 0 65 3 1 0 0 6 4 48 6547
Sticky trap
( 0.2) (97.3) ( 0.6)
2519 7610 4631 23 0 61 3 48 6 28 0 1 28 14,959
( 1.0)
( 0.1) ( 0.1)
( 0.4) ( 0.3) ( 0.2)
i n v o l v e d d a t a f r o m 31 M a y t o 2 0 J u l y i n a r e a A a n d f r o m 4 J u n e t o 10 A u g u s t i n a r e a B for T. nigrovittatus, f r o m 31 M a y t o 11 J u n e i n a r e a A for C. atlanticus a n d f r o m 6—18 M a y i n a r e a A for C. fuliginosus. T h e n u m b e r of flies p e r species t a k e n o n e a c h of t h e c o l l e c t i o n d a t e s w a s i n d e p e n d e n t l y a n a l y z e d as a r a n d o m i z e d complete block, with 6 treatments ( t r a p p o s i t i o n s ) a n d 3 r e p l i c a t e s (rows) for a r e a A , a n d 5 t r e a t m e n t s a n d 3 r e p l i c a t e s for a r e a B . Honestly significant difference (h.s.d.) values ( T u k e y ' s test) a t t h e 0 . 0 5 l e v e l of s i g n i f i c a n c e w e r e u s e d for p a r t i a l 2 - t r e a t m e n t c o m p a r i s o n s .
analysis
D u r i n g p e a k fly p o p u l a t i o n s , b o t h t y p e s of t r a p s were e x a m i n e d twice a week; at other times, counts were at weekly intervals.
T h e flies i n m o s t cases
w e r e identified
i n t h e field.
counted;
few
very
Box trap 6 ( 0.3) 1897 (95.4) 39 ( 2.0) 0 0 18 ( 0.9) 0 0 0 0 0 0 29 1988
(16.8) (50.9) (31.0) ( 0.2)
males
O n l y females
were
trapped.
RESULTS A N D DISCUSSION
Comparison of Spartina
and Juncus
marshes
D u r i n g this 2-year study, 79,842 a d u l t
tabanids
were
b e l o n g i n g t o 32 species a n d 6 g e n e r a w e r e c o l l e c t e d .
After
T h e most a b u n d a n t
( m o r e t h a n 50) a r e listed i n
c o u n t i n g , all flies w e r e r e m o v e d a n d t h e S t i c k e m ®
T A B L E 1.
renewed.
w e r e c o l l e c t e d a n d i n t h e Juncus s a l t m a r s h e s o n l y
Relative
the
2 0 species.
combing
the
species i n t h e g e n u s Tabanus
c o l l e c t i o n d a t a for t h e e n t i r e s e a s o n for a r e a s A
and
of m a r s h
abundance
was
according
salt m a r s h e s 31 species
to
type
tabanid
I n t h e Spartina
determined
by
B (Spartina s a l t m a r s h e s ) a n d a r e a s C a n d D salt m a r s h e s ) .
(Juncus
T h e r e l a t i v e a b u n d a n c e of i n d i v i d u a l
species w a s d e t e r m i n e d b y s u m m i n g t h e d a t a c o l
I n t h e Juncus m a r s h e s t h e n u m b e r o f w a s 4 4 % less a n d i n
Chrysops 3 8 % less t h a n i n t h e Spartina m a r s h e s . Spartina
salt
marshes
produced
t a b a n i d s t h a n t h e Juncus m a r s h e s .
3.7
The
times
more
Davis &
(1966) f o u n d a b o u t 4 0 t i m e s m o r e insects i n
Gray
Spartina
l e c t e d b y b o t h t y p e s of t r a p s for all t h e a r e a s a n d
t h a n i n Juncus m a r s h e s b u t t h e i r d a t a i n c l u d e d v e r y
e x p r e s s i n g t h e r e s u l t s a s p e r c e n t a g e of t h e
few
total.
O n l y t h e sticky t r a p c o l l e c t i o n d a t a w e r e u s e d t o express
the
seasonal
abundance
for
the
3
most
c o m m o n species. The
horizontal
tabanids
(due
to
the
inadequate
sampling
methods). T h e 3 m o s t a b u n d a n t species i n o u r
collections
( a c c o u n t i n g for 9 8 % of t h e t o t a l ) for b o t h t y p e s of distributions
C. atlanticus a n d C. fuliginosus
of
T.
nigrovittatus,
i n Spartina salt m a r s h e s
m a r s h e s w e r e T. nigrovittatus, atlanticus.
These
species,
C. fuliginosus,
along
with
T.
a n d C. lineola
( a r e a s A a n d B) w e r e a n a l y z e d b y u s i n g d a t a f r o m
Fabricius, have been reported to predominate
t h e sticky t r a p s for c o n s e c u t i v e c o l l e c t i o n d a t e s i n
o t h e r s a l t m a r s h e s of t h e A t i a n t i c c o a s t of t h e U n i t e d
which
States ( J a m n b a c k
of a
given
species p e r t r a p p e r d a y w e r e 2 o r m o r e .
the
average
numbers
of flies
This
1960).
& Wall
1959, W a l l
in
&
Doane
I n o u r a r e a s , t h e r e w e r e v e r y few T.
lineola.
Downloaded from http://jme.oxfordjournals.org/ by guest on June 5, 2016
Juncus
28,030 23,942 3081 398 219 161 158 69 55 54
Juncus (AREAS C & D)
Box trap
Dale & Axtell:
1976
Seasonal
675
Tabanidae abundance and distribution
abundance
C. Juliginosus (FIG. 5) w a s c o l l e c t e d f r o m 8 A p r i l t o 8 J u n e . D a t a w e r e o b t a i n e d o n l y for a r e a s B a n d D . T h e e m e r g e n c e of t h e first a d u l t s c o i n c i d e d w i t h i n c r e a s i n g t e m p e r a t u r e s w i t h t h e a d v e n t of s u m m e r , as observed b y J a m n b a c k & W a l l (1959).
T h e r e w a s 1 p o p u l a t i o n p e a k , i n t h e 1st w e e k of M a y , followed b y a s h a r p decrease in the adults completely disappeared Hansens
&
Robinson
(1973)
numbers;
1 month
observed
in
later. New
J e r s e y t h a t after t h e p e a k w a s r e a c h e d , a c t i v i t y for this species d e c l i n e d
abruptly.
T h e s e a s o n a l a b u n d a n c e of t h e s e flies is i n f l u e n c e d by the changing weather, particularly the tempera t u r e . I t is a p p a r e n t f r o m c o m p a r i n g o u r d a t a t o previously published d a t a (Anderson 1971, Anderson & K n e e n 1969, B a i l e y 1947, Blickle 1954, B r e n n a n 1 9 3 5 , H a n s e n s 1952, H a n s e n s & R o b i n s o n 1 9 7 3 , J a m n b a c k 1969, J a m n b a c k & W a l l 1959, J o n e s & A n t h o n y 1964, M a c C r e a r y 1940, P e c h u m a n 1 9 5 7 , 1972, S t o n e 1938) t h a t T. nigrovittatus, C. atlanticus a n d C. fuliginosus e x h i b i t a d e l a y i n t h e t i m e of p e a k a b u n d a n c e from southern to n o r t h e r n latitudes along t h e e a s t e r n c o a s t of t h e U n i t e d S t a t e s . Horizontal
distribution
T. nigrovittatus c o l l e c t i o n s f r o m s t i c k y t r a p s s h o w e d significant differences a m o n g t h e t r a p p o s i t i o n s i n 8 0 % of t h e 15 c o l l e c t i o n p e r i o d s i n a r e a A i n 1971 a n d i n all of t h e 15 c o l l e c t i o n p e r i o d s i n a r e a B i n 1972 (TABLE 2 ) .
I n b o t h years t h e results
were
essentially t h e s a m e a n d t h e r e w a s n o c h a n g e in t h e d i s t r i b u t i o n of t h e flies d u r i n g t h e s e a s o n . flies
were
trapped
in
the
outer
marsh,
More nearest
FIG. 4. Average number of flies/trap/day for Chrysops atlanticus Pechuman, in Spartina (areas A & B) and Juncus (areas C & D) marshes during 1971 and 1972.
Downloaded from http://jme.oxfordjournals.org/ by guest on June 5, 2016
T h e a b u n d a n c e of T. nigrovittatus i n Spartina a n d Juncus m a r s h e s is i l l u s t r a t e d i n FIG. 3. F o r b o t h Spartina m a r s h e s t h e r e w a s 1 p e a k of a b u n d a n c e f r o m 5 - 1 4 J u n e , a p p r o x i m a t e l y 1 m o n t h after t h e e m e r g e n c e of t h e first a d u l t s , f o l l o w e d b y a s u d d e n d e c r e a s e of t h e p o p u l a t i o n to l o w levels w h i c h w e r e m a i n t a i n e d u n t i l t h e e n d of t h e s u m m e r . T h i s is essentially t h e s a m e p a t t e r n d e s c r i b e d for M a s s a c h u setts b y W a l l & D o a n e ( 1 9 6 0 ) . A s i m i l a r p e a k o c c u r r e d i n t h e Juncus m a r s h a r e a C , a l t h o u g h n u m b e r s w e r e v e r y l o w . I n t h e o t h e r Juncus m a r s h (area D ) , there were fluctuations in the low n u m b e r s t h r o u g h o u t the season. C. atlanticus (FIG. 4) w a s c o l l e c t e d from 2 3 April t o 2 6 S e p t e m b e r . T h e r e w a s 1 p e a k of a b u n d a n c e , f r o m 31 M a y t o 18 J u n e . F i e l d o b s e r v a t i o n s o n b i t i n g a c t i v i t y for 2 c o n s e c u t i v e y e a r s r e v e a l e d a r e t u r n of h i g h n u m b e r s of flies d u r i n g t h e l a s t 1 / 2 of A u g u s t b u t t h i s w a s n o t s h o w n b y t h e t r a p d a t a . B y c o m p a r i s o n t o h i g h e r n e t t i n g c o u n t s , i t is a p p a r e n t t h a t t h e traps w e r e relatively p o o r collectors o f C. atlanticus.
J. Med. Ent.
676
Vol. 12, no. 6
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