WARNING CONCERNING COPYRIGHT RESTRICTIONS The copyright law of the United States (Title 17, United States Code) governs the making of photocopies or other reproductions of copyrighted materials. Under certain conditions specified in the law, libraries and archives are authorized to furnish a photocopy or other reproduction. One of these specified conditions is that the photocopy or reproduction is not to be “ used for any purpose other than private study, scholarship, or research.” Anyone using, a photocopy or reproduction for purposes in excess of “ fair use,” may be liable for copyright infringement.
bJl
·-,"""Cl
Borrower: FUG
Call#:
Lending String: "FDA,FTU,FXG.FHM,FQG,VRC,VA@,KUK,VPI,V WM,MFM,GUA,TMA,TKN
Location: DIRAC
Charge
Patron: Moser, Bettina
Maxcost: SO.OOIFM
...l-
.c=
Journal Title: Journal of economic entomology .
·-= "'. . = """
Volume: 84 Issue: 4 MonthNear: ; 1991Pages: 1257-1261
""""
Article Author: B A Moser, P G Koehler, R S Patterson
·->= == 0.)
~
- .. 0.)
i fJ
Shipping Address: University of Florida Smathers Library 245 Library West Gainesville, FL 32611-7001 2 1254 00152 8270 GVL
-
"' ., ·-"'.... ....~ "'" .,....z -
590.5J861
"'=
-
CD
Article Title: Effect of larval diet on cat flea (Siphonaptera; Pulicidae) developmental times and adult emergence.
"Cl
0
.!!! ....1 ....1
Imprint: [Lanham, Md., etc.] Entomological Societ
ILL Number: 116328463 1111111111111111111111111111111111111111111111111111111
'
'
Fax: 352-392-7598 Odyssey: 128.227.193.10 Ariel: 128.227.193.10 Email:
[email protected] NOTICE: THIS MATERIAL MAY BE PROTECTED BY COPYRIGHT LAW (Title 17 USC)
1256
jOURNAL OF EcONOMIC ENTOMOLOGY
Florida show that adults are virtually absent in December and January and again in June and July (Brenner 1990), suggesting that there may be two broad generations annually. Minimum generation time (female immature development time + initial preoviposition period + oothecal incubation period) under laboratory conditions is ==100 d. Total life span for females from this study, including oothecal incubation, nymphal development, and female longevity is 188.5 d, almost exactly 6 mo. Although Asian cockroaches breed continuously in colony indoors, seasonal changes in temperature and pos.sibly humidity may prevent this from occurring in the field; this would certainly be the case if the Asian cockroach spreads northwards to any appreciable degree. Our preliminary data on control of the Asian cockroach by application of pesticides to infested turf indicate that chemical control is feasible. Such treatments should also be effective in areas with low vegetation and leaf litter. Asian cockroaches are not known to burrow, and pesticide residues on foliage or in the upper layers of the litter would be most effective. Because Asian cockroaches are primarily perceived as pests when adults are flying, timing outdoor pesticide applications to seasonal peaks in adult abundance might reduce local populations to acceptable levels with a minimal number of applications. Fecundity, development, longevity, and seasonal changes in population should be studied for Asian cockroaches under conditions more similar to those encountered outdoors. Asian cockroaches of both sexes fly actively in the field and have been observed feeding on homopteran honeydew and fruits (Brenner et aL 1988, Brenner 1990). Development times of nymphs held in groups should be studied because German and field cockroach nymphs develop faster in groups than when held in isolation (Willis et aL 1958). Larger rearing containers and alternative food sources with a high sugar content might significantly increase Asian longevity and fecundity, providing more realistic estimates of biological parameters.
Acknowledgment We thank Euripedes Mena (Entomology & Nematology Department, University of Florida) for technical assistance. N.C. Hinkle (Entomology & Nematology Department, University of Florida), M.H. Ross (Department of Entomology, Virginia Polytechnic Institute & State University), and L.M. Roth (Museum of Comparative Zoology, Harvard University) kindly reviewed this
VuL 84, no. 4
manuscript. This is Florida Agricultural Experiment Station Journal Series R-00561.
T Effect of Larval Diet on Cat Flea (Siphonaptera: Pulicidae) Developmental Times and Adult Emergence
References Cited BETTINA A. MOSER, PHILIP G. KOEHLER, A:'«D RICHARDS. PATTERSON'
Atkinson, T. H., P. G. Koehler&R. S. Patterson. 1990.
Checklist of the cockroaches of Florida (Dictyoptera: Blattaria: Blattidae, Polyphagidae, Blattellidae. Blaberidae). Fla. Entomol. 73: 303-327. Brenner, R. J. 1988, Activity and distribution of cockroaches with implications to developing control strategies. pp. 33-40. In Proc. Natl. Conf. Urban Entomol. (1988), University of Maryland, College Park, Md. 1990. Asian cockroaches: implications to the food industry and complexities of management strategies. In]. R. Gorham [ed.]. Ecology and management of food-industry pests. Food Drug Admin. Tech. Bull. No. 4 (in press). Brenner, R. J., R, S. Patterson & P. G. Koehler. 1988. Ecology, behavior, and distribution of Blattella asahinai (Orthoptera: Blattellidae) in central Florida. Ann. Entomol. Soc. Am. 81: 432-436. Cochran, D. G. 1979. A genetic determination of insemination frequency and spenn precedence in the German cockroach. Entomol. Exp. Appl. 26: 259266.
Cornwell, P. B. 1968. The cockroach, a laboratory insect and industrial pest, vol. 1 Rentokil Library, Hutchinson, London. Ebeling, W. 1975. Urban entomology. Univ. California Div. Agric. Sci.. Los Angeles. Koehler, P. G, & R. S. Patterson. 1987. The Asian roach invasion. Nat. Hist. 96: 28-35. Mizukubo, T. 1981. A revision of the genus Blattella {Blattaria: Blattellidae) of Japan. I. Terminology of the male genitalia and description of a new species from Okinawa Island. Esakia 17: 149-159. Ross, M. H. 1988. Cytological studies of Blattella germanlca and Blattella asahinai. I. A possible genetic basis of interspecific divergence. Genome 30: 812-819. 1989. Asian cockroaches. What are the implications for the development of mixed populations? Pest Management 8: 20-22. Ross, M. H. & D. E. Mullins. 1988. Nymphal and oothecal comparisons of Blattella asahinai and Blattella geTTIWnica (Dictyoptera: Blattellidae). J. Econ. Entomol. 81: 1645-1647. Roth, L. M. 1985, A taxonomic revision of the genus Blattella Caudell {Dictyoptera: Blattaria: Blattellidae). Entomol. Scand. Suppl. 22: 1-221. 1986, Blatt ella asahinai introduced into Florida (Blattaria: Blattellidae). Psyche 93: 371-374. Willis, E. R., G. R. Riser & L. M. Roth, 1958. Observations on reproduction and development in cockroaches. Ann. Entomol. Soc. Am. 51: 53--69. Received for publication 15 October 1990; accepted 11 February 1991.
Department of Entomology and Nematology, Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida 32611
J. Econ. Entomol. 84(4); 1257-1261 (1991) ABSTRACT The natural diet of cat flea, Ctenocephalides felis (Bouche), larvae is primarily adult flea feces, but dried bovine blood may be substituted in the laboratory. Percentage adult emergence (79.4% on feces; 78.9% on blood) and developmental times (20.6 don feces; 17.1 don blood) did not significantly differ for the two diets. The drying temperature of blood detennined its quality; blood dried at 120"C was unsatisfactory for larval development. The dietary value of dried bovine blood was not enhanced when supplemented with brewer's yeast, rodent chow, or a combination of those constituents. Blood particle size ranging from 500u did not affect the value of blood as a diet. Rodent chow. yeast, albumen, hemoglobin, and mixtures of these constituents were unsuitable as larval diets. KEY WORDS
Insecta, Ctenocephalides jelis, larval development, blood
MoST FLEA SPECIES parasitize nest-dwelling animals, and the great majority of flea larvae live in the nest or den of their host (Marshalll981). Among the nest-inhabiting flea larvae, there is a gradation of dependence on the host and adult fleas for nutrition. The larvae of many flea species can feed solely upon organic matter that is not host-derived, although their food may often include host epidermal tissues. Larvae of other species need adult flea feces as their major dietary component. The larva of one species, Nosopsyllus fasciatus (Bose), feeds directly at the anus of the adult Ilea. Another species (Tunga monositus Barnes & Radovsky) has all the necessary nutrients for immature development in the egg (Radovsky 1972). Because adult flea feces often constitute a major dietary component in larval nutrition, adult fleas imbibe much more blood from the host than they can use. Adult flea feces are generally regarded as consisting of almost undigested or only slightly altered host blood (Sharif 1937, Akin 1984). Probably only the blood nearest the functional midgut cells i. 0.0592; blood: F = 2.11; df = 1, 5; P > 0.2062; blood-yeast: F = 2.13; df = 3, 9; P > 0.1667; blood-rodent chow: F = 0.36; df = 2, 6; P > 0.7109; laboratory diet: F = 4.65; df = 2, 6; P > 0.0604; rodent chow: F = 0.07; df = l, 6; P > 0.8058; rodent chow-yeast: F = 1.0; df = 1, 4; P > 0.3739; albumen: F = 0.82; df = 1, 4; P > 0.4169; yeast: F = 0.25; df = 1, 4; P > 0.6433). However, the mean number of adults that emerged from each diet was significantly different (F = 97.49; df = 10, 79; P > O.IX.M.H). Diets containing adult flea feces or dried bovine blood produced the most adults (Table 1). Percentage adult emergence did not differ significantly for fleas reared on adult flea feces, dried bovine blood, dried bovine blood mixed with either yeast or rodent chow, and the standard laboratory diet. Significantly fewer adults emerged from rodent chow and the mixture of rodent chow and yeast. Few or no cat flea larvae developed to the adult stage on diets of individual blood components or without blood. From these data, we conclude that no additives to dried blood are necessary in the cat flea larval diet.
1259
MOSER ET AL.. CAT FLEA LARVAL DIETS
f - - - - test tube
Fig. I.
Discussion Several studies confirm that some form of blood must be present in the flea larval diet. Strenger (1973) successfully reared cat flea larvae on dried cat and calf blood and adult flea feces; no differences among the diets was observed. Pausch & Fraenkel (1966) obtained almost 100% pupation when Xenopsylla cheopis larvae were reared on
Emergence chambers to collect adult cat fieaJS. Table 2. Effed of larval diet on emergence tiiileS of adult cat fleas
The efficiency of a diet can also be evaluated by larval development times. Time to adult emergence was shortest for diets containing adult flea feces and diets containing dried bovine blood (Table 2). The ET 50 was shortest for cat flea larvae reared on dried blood, followed closely by larvae reared on the blood-yeast diet, adult flea feces, blood-rodent chow mix, and the standard laboratory diet. The ET 50 of blood was significantly less than the ET50 of the blood-yeast, blood-rodent chow mix, and laboratory diet. Emergence times among the replicates varied for fleas reared on adult flea feces and resulted in a wide confidence interval for the ET 50 • We suspect that the quality of the flea feces was not consistent among the replicates. ET 50 S could not be calculated for the albumen, hemo-
Diet
Blood Blood-Yeast Feces Blood-Rodent chow Laboratory diet Rodent chow Rodent chow-Yeast Albumen Yeast Hemoglobin HemoglobinAlbumen
"" 57
ET~,l.
days
00
17.1 I9 7 W.6 23.4
""
Nl 28.7 31.1
105
w;
27 le
95% CL 16.6-17.6 17.9-2l.l l7.5-23.I 22.5-24.3 :l2 8-25.3 27.5-29.7 29.2-32.6
Slope .:!: SE 0.347 0.112 0.049 0.116 0.106 0.210 0.162
.:!:
± ± ± ± ± ±
0.044 0.019 0.009 0.011 0.013 0034 0.030
'0 0
"Total number of emerged adults. bET 50 is the time at which 50% of the adults have emerged (Finney 1971).
1260
Table 3. Effed of bl....:>d partiele size and drying temperature on number of adult eat fleas (X ::!: SE) emerging from ehamben Diet F~~
Blood (60"0) _,;Jgob
>ISO: ,;zso!' >250, :S355b
>355; :Ssoob >5oo'
Vol. 84, no. 4
}OCRNAL OF ECONOMIC ENTOMOLOGY
Mixed sizes ).1hed sizes (120"C")
"
' '3 3 '3 '5
No. fleas per chamber 55.6a 32_3 ± I 76a 37.3 ± 1.67a 55.6 ± 2.19a 31.6 ± 1.20a 32.3::!: 3.84a 30.6 ± 145a O.Ob
Table 4. Effeet o£ blood partiele size on emergenee times of adult eat fleas Diet F~•
••
ET51l.
days
95% CL
Slope± SE
107
17.2
16.4-17.9
0.363 ± 0.049
Blood (130"'0')
:Sisod >ISO; :S25od >5ood >250; :S35sd
:S5ood Mixed sizes >355;
"
112 99 107
95
"
16.8 17.0 \7.4 li.6 17.7 18.0
~
16.8-17.2 ~
16.8-18.5 16.6--18.8 15.2-20.6
a Total number of adults emerged. h EToo is the time at which 50% of
0.379 0.454 0.327 0.339 0.360 0.288
::!: 0.251 ± 0 Q-22 ± 0.267 ± 0.049 ± 0.075 ::!: 0.096
the adults have emerged
).-leans followed by the same letter are not significantly different (P ~ 0.05; Tukey's studentized range test [SAS Institute 1988]) "Drying temperature of blood. b Particle sizes in micrometers.
(Finney 1971). c Drying temperature of blood. d Particle size in micrometers_
dried bovine blood. However, Bruce (1948) found that cat flea larvae developed well on dried beef blood, but the addition of 5% (wt:wt) brewer's yeast to dried beef blood was superior to blood alone. The blood was oven dried at 9goc for at least 24 h. Drying blood at high temperatures can alter nutrients such as vitamins, proteins, or other growthpromoting substances. Presumably, the blood used by Bruce (1948) was deficient in nutrients that were supplied by the addition of yeast. The blood used in our study was either oven-dried at 600C or air dried at room temperature. The method of drying blood might explain why the addition of yeast did not enhance the dietary value of dried blood in our study. Hemoglobin and serum are important components of blood in the nutritional requirements of N. fasciatus larvae (Sharif 1937). Hemoglobin provides larvae with iron needed for normal growth and proper sclerotization as adults, and serum has all the essential proteins. Although blood is the most important part of nutritional requirements of flea larvae, it was not sufficient for normal and successful development of N. fasciatus (Sharif 1937). Larvae could not be reared to adults on dried blood or its constituents. When fed blood, larvae were active and strong for 30 d and then suddenly became sluggish and died in the third stage without forming cocoons. When fed on red blood corpuscles, they died in 12-21 d, mainly in the second larval stage. On serum, they lived for 19 d and reached the second larval stage. When the sole diet was fibrin, the larvae did not molt and died in 820 d. Addition of dried baker's yeast to blood gave almost 100% success in the rearing of N. fasciatus . The blood and its different constituents were dried by evaporation in a vacuum desiccator. When autoclaved blood dried at 1200C mixed with yeast was fed to N.fasctatus larvae, they did not develop into adults (Sharif 1937). Similar observations were made in our study. When blood was oven-dried at 120"C and offered to cat flea larvae as a diet, none developed to adults (Table 3), again stressing the
importance of drying blood at the appropriate temperature. Cat flea larvae died after 3-4 d when offered fresh or dried yeast (Strenger 1973). N. fasciatus larvae did not grow on yeast alone and died within 12-25 din the first stage without any appreciable increase in size (Sharif 1937). Only a small percentage of cat flea larvae reached the adult stage when reared on brewer's yeast or Swift's dog food (35 and 23%, respectively), and 68 and 89% reached maturity when fed on hemoglobin or albumen (Bruce 1948). In our study, none of the larvae emerged as adults when hemoglobin was the diet, and only 5% emerged as adults when raised on albumen. This might be attributable to the processing method for hemoglobin and albumen and their resulting water content. The water content of the larval diet is crucial for flea larval rearing (Sikes 1931). However, Sikes (1931) found albumen with the correct water content to be an unsatisfactory larval diet for Ceratophyllus wickhami. Only 25% of the larvae of X. cheopis pupated after a prolonged larval period when fed hemoglobin (Sharif 1937). Some larvae fed on inadequate diets prolonged their larval life to such an extent that it was longer than the combined larval and pupal period of others (B.A.M., personal observation, Sharif 1937). These larvae (called lagging larvae) seldom formed cocoons and usually died before developing into adults. The phenomenon of prolonged growth periods of flea larvae is probably caused by the lack of some essential dietary nutrient. Larvae receive enough nutrients from certain inadequate diets to sustain essential life functions but they are unable to metamorphose (Sharif 1937) . In summary, adult flea feces are the natural cat flea larval diet; however, dried bovine blood can be substituted to rear larvae in the laboratory. The temperature used to dry blood determines its quality as a diet. High temperatures render blood unsuitable as a diet; whereas, low temperature drying produces blood that gives optimal flea development and survival
August 1991
MOSER ET AL.: CAT FLEA LARVAL
Acknowledgment This is Florida Agricultural Experiment Station journal Series Number R-00766. References Cited Akin, D. E. 1984. Relationship between feeding and reproduction in the cat flea, Ctenocephalides fells {Bouche) (Siphonaptera: Pulicidae). M.S. Thesis. University of Florida. Bruce, W. N. 1948. Studies on the biological requirements of the cat Ilea. Ann. EntomoL Soc. Am. 41: 346----352. Dryden, M. W. 1989. Biology of the cat Ilea, Ctenocephalides fells fells. O:lmpanion Animal PracticeParasitology 19(3): 23-27. Finney, D. J, 1971. Probit analysis. Cambridge University, London. Hodson, B. W. 1958. A method for large-scale rearing of the cat flea, Ctenocephalides fells felis (Bouche). BulL w_ H. Q_ 19: 1126-1129. Joseph, S. A. 1976. Observations on the feeding habits of Ctenocephalides fe/is OTientls Jordan on human host. Cheiron 5: 73--77. 1981. Studies on the bionomics of Ctenocephalides fells orientis (Jordan). Cheiron 10: 275-280. Marshall, A. 1981. The ecology of ectoparasitic insects. Academic, London. Pa~h. R. D. & G. Fraenkel. 1966. The nutrition of the larva of the oriental rat Ilea Xenopsylla cheopis (Rothschild). PhysioL Zoo!. 39: 202-222.
DIETS
1261
RadoV!;ky, F. J, 1972. Fixed parasitism in the Siphonaptera_ J. Med. EntomoL 9: 602. Rothschild, M.B.F. & M. H115hes. 1970. Maturation of the male rabbit flea (Spilopsyllus cuniculi) and the oriental rat flea (Xenopsylla cheopls ): Some effects of mammalian hormones on development and impregnation. Trans. Zoo!. Soc. Lond. 32: 105-188. SAS Institute, 1988. SAS/STAT guide for personal computers. SAS Institute, Cary, N.C. Sharif, M. 1937. On the life history and the biology of the rat flea, Nosopsyllus fasciatus (Bose.). Parasitology 29: 225-238. Sikes, E. K. 193l. Notes to breeding fleas, with reference to humidity and feeding. Parasitology 23: 243249. Smith, C. N. & G. W. Eddy. 1954. Techniques for rearing and handling body lice, oriental rat fleas. and cat fleas_ BulL W. H. 0. 10: 127-137. Stre115er, A. 1973. Zur Emaehrungsbiologie der Larve von Ctenocephalides felis fells B. ZooL Jb. Syst. 100: 64-80. Suter, P.R. 1964. Biologie von Echidnophaga gallinacea (\Vestw.) und Vergleich mit anderen Verhaltenstypen bei Floehen. Acta Trop. 21: 193-238. Winston, P. W. & H. D. Bates. 1960. Saturated solutioru; for the control of humidity in biological research. Ecology 41: 232-237. Received for publication 23 july 1990; accepted 11 March 1991.
