This dissertation has been microfihned exactly as received 6 8-2971
DEAL, Andrew Stuart, 1918- THE EFFECT OF TEMPERATURE AND MOISTURE ON THE DEVELOPMENT OF FANNIA CANICULARIS (L.) AND FANNIA FEMORALIS (STEIN) (DIPTERA: MUSCI- DAE. The Ohio State University, P h . D „ 1967 Entomology
University Microfilms, Inc.. Ann Arbor, Michigan Copyright by
Andrew Stuart Deal
1967 THE EFFECT OF TEMPERATURE AND MOISTURE
ON THE DEVELOPMENT OF
FANNIA CANICULARIS (L.) AND FANNIA FEMORALIS (STEIN)
(DIPTERA: MUSCIDAE)
DISSERTATION
Presented in Partial Fulfillment of the Requirements for Degree Doctor of Philosophy in the Graduate School of the Ohio State University
By
Andrew Stuart Dealg BoSo, MoSo
******
The Ohio State University 1967
Approved by
Adviser Department of Zoology and Entomology ACKNOWLEDGMENTS
The author gratefully acknowledges the cooperation and assistance of many people in making this study possible. Significant cbntributions were made by individuals and departments of the University of California at Riverside where the laboratory phases of the work were carried out.
Thanks go to Dr« G, E, Carman of the Department of Entomology and to
Dr. I, Mo Hall of the Department of Biological Control for use of space, materials and equipment. Dr, J, M, Rible of the Agricultural Extension
Laboratory gave valuable assistance in making moisture determinations of the fly rearing medium. Others of the Riverside staff who-rendered valuable assistance were Drs, L, D, Anderson, E, C, Bay, W, H, Ewart,
Go P, Georghiou, Daniel Gonzalez, R, N, Jefferson, E, L, Reeves,
L, A, Riehl, and Mr, E, L, Atkins. Special thanks go to Dr, George B,
Alcorn and Mr, H, W, SchwaIm of the Agricultural Extension Adminis trative Staff for granting the author time away from his regular duties to carry out the work, and to Miss Lucy M, Allen, Program Leader,
Extension Education, and Mr. J. E. Tippett, Agriculturist Emeritus, for guidance in working out sabbatical leave details. Assistance from Dr,
T, M, Little in analyzing the data is sincerely appreciated.
I wish to express my gratitude to Dr, Ralph H. Davidson, Professor of Entomology at The Ohio State University for his guidance and
encouragement throughout the duration of this project, I also wish to
express my appreciation to Drs, Davidson, Donald J. Borror and Frank W,
il Fisk for their suggestions during preparation of this manuscript.
I am very grateful to my secretary, Mrs. Lucille M. Sanchez, and to my laboratory technician, Mr. W. R. Bowen, for patiently and effectively carrying out many of my regular duties during my leave of absence, and to Mrs. Freida M. Bailey for special help.
I wish also to thank my wife, Audrey, for her assistance with some of the laboratory work, the typing, and for her enduring patience with author during the course of this work.
Funds used for purchase of much of the materials and equipment for use in this project were provided by Julius Goldman's Egg City,
Moorpark, California.
iii VITA
July 3, 1918 ...... B o m - Birch Tree, Missouri
1950 o « ...... B.S., University of California, Berkeley, California
1951 00000.00000 MoSo, University of California, Berkeley, California
1951-1952 0 0.00000 Entomologist, Bio Research Laboratory, California Spray Chemical Corporation, Richmond, California
1952-1956 00 0 0 .0 0 0 Farm Advisor (Entomology), University of California Agricultural Extension Service, El Centro, Imperial County, California
1956-present ...... Extension Entomologist, University of California, Riverside, California
PUBLICATIONS
Yellow clover aphid in state. Calif. Agric. 8(9): 5, 1954.
Ground pearls on grape roots. Calif. Agric. 8(12): 5, 1954.
The Egyptian alfalfa weevil. Calif. Agric. 9(6): 8, 1955.
The omnivorous leaf roller, Platvnota stultana Wlshm.. on cotton in southern California: Damage and control. Jour. Econ. Ent. 50(1): 59-64, 1957.
The "omnivorous leaf roller," Platvnota stultana Wlshm.. on cotton in California: Nomenclature, life history and bionomics (Lepidoptera, Tortricidae)o Ann. Ent. Soc. Amer. 50(3): 251-259, 1957.
A survey of beet leafhopper populations on sugar beets in the Imperial Valley, California, 1953-1958. Jour. Econ. Ent. 52(3): 470-473, 1959.
Insecticidal control of lygus bugs and effect on yield and grade of lima beans. Jour. Econ. Ent. 59(1): 124-126, 1965.
IV Trials of Ruelene for cattle grub control in southern California. Jour, Econ. Ent, 58(2): 361-362, 1965.
Timing lygus bug control increases lima bean yield and quality, Calif. Agric. 19(7): 2-3, 1965.
Fly control in cattle feedlots with residual sprays, Calif, Agric, 19(9): 6-7, 1965.
The Egyptian alfalfa weevil and its control in southern California, Jour, Econ, Ent, 48(3): 297-300, 1955, TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS ...... ii
VITA ...... iv
PUBLICATIONS ...... iv
LIST OF TABLES ...... viil
LIST OF ILLUSTRATIONS ...... ix
Chapter
I. INTRODUCTION ...... 1
II. REVIEW OF THE LITERATURE ...... 4
Distribution ...... 5
Importance to M a n ...... 7
Biology ...... 13
Rearing Media ...... 19
III. RELEVANT MORPHOLOGICAL ASPECTS ...... 25
The E g g s ...... 25
The L a r v a e ...... 29
The P u p a e ...... 36
IV. MATERIALS AND M E T H O D S ...... 42
Colony Establishment ...... 42
Rearing ...... 42
Adult Maintenance ...... 43
Egg Collection ...... 45
Vi Chapter Page
Obtaining Larvae ...... 47
Incubation at Various Temperatures ...... 49
Larval and Pupal Development at Various Temperatures 51
Temperature Cabinets ...... 53
Effect of Moisture on Development ...... 53
V. THE EFFECT OF TEMPERATURE - RESULTS AND DISCUSSION . . 61
Effect of Temperature on Incubation ...... 61
Effect of Temperature on Larval and Pupal Development ...... 69
VI. THE EFFECT OF MOISTURE - RESULTS AND DISCUSSION . . . 77
VII. CONCLUSIONS ...... 81
VIII. LITERATURE CITED ...... 87
vii LIST OF TABLES
Table Page
1. Computation of moisture content of C.S.M.A. medium at end of ten days in moisture control apparatus .... 60
2. Effect of constant temperature on incubation period of Fannia canicularis eggs ...... 62
3. Effect of temperature on hatch of eggs of Fannia canicularis ...... 65
4. Effect of incubation at 93 Fahrenheit for various periods of time on hatching of Fannia canicularis eggs . . . 65
5. Effect of constant temperature on incubation period of Fannia femoral is e g g s ...... 67
6. Effect of temperature on hatch of eggs of Fannia femoralis ...... 68
7. Effect of incubation at 100 F for various periods of time on hatching of Fannia femoral is e g g s ...... 68
8. Effect of temperature on duration of instars and the total life cycle of Fannia canicularis ...... 71
9. Effect of temperature on the number of flies developing to the adult stage in Fannia canicularis ...... 72
10. Effect of temperature on duration of instars and the total life cycle of Fannia femoral i s ...... 74
11. Effect of temperature on the number of flies developing to the adult stage in Fannia femoral is ...... 75
12. Effect of moisture content of the rearing medium on larval and pupal development of Fannia canicularis . 79
13. Effect of moisture content of the rearing medium on larval and pupal development of Fannia femoralis . . 79
viii LIST OF ILLUSTRATIONS
Figure Page
1. Geographical distribution of Fannia canicularis ...... 6
2. Geographical distribution of Fannia femoralis ...... 8
3. Eggs of Fannia femoral i s ...... 26
4. Drawing of end view of Fannia canicularis egg showing position of wings or flanges after egg is laid . . 26
5. Drawing of dorsal aspect of Fannia canicularis egg showing reticulated pattern on the surface ...... 28
6. Drawing of ventral aspect of Fannia canicularis egg showing longitudinal ribbing on surface ...... 28
7. Drawing of dorsal aspect of Fannia canicularis larva showing long rat-tail-like processes which are characteristic of this s p e c i e s ...... 30
8. Drawing of the dorsal aspect of Fannia femoralis larva showing characteristic dorsal and lateral processes. . . . 30
9. Photomicrograph of the posterior spiracular stalks of the first instar larva of Fannia femoral is showing the two spiracular openings characteristic of this stage . 31
10. Drawing of the posterior spiracular stalks of the first instar larva of Fannia femoral is shown in Figure 9 ... . 31
11. Photomicrograph of the left posterior spiracular stalk of the second instar larva of Fannia femoralis ...... 32
12. Photomicrograph of the left posterior spiracular stalk of the third instar larva of Fannia femoralis ...... 32
13. Drawing of the left posterior spiracular stalk of the third instar larva of Fannia femoralis shown in Figure 12 showing location of the four spiracular openings 33
14. Photomicrograph of the cephalic region of the first instar larva of Fannia femoral is showing the horseshoe-shaped pharyngeal skeleton ...... 33
IX Figure Page
15. Photomicrograph of the cephalic region of the second larval instar of Fannia femoral is showing the narrowly U-shaped pharyngeal skeleton and its connection with the mouth hooks ...... 35
16. Photomicrograph of the cephalic region of the third larval instar of Fannia femoralis showing the narrowing of the U-shape and extension of the U-shape to the r e a r ...... 35
17. Photomicrograph of the shed skin of the second larval instar of Fannia femoral is showing the pharyngeal skeleton and mouth hooks attached ...... 37
18. Puparium of Fannia femoral is on the left, and that of F. canicularis on the r i g h t ...... 37
19. Drawing of the ventral aspect of Fannia canicularis larva 38
20. Drawing of the ventral aspect of Fannia femoralis larva 38
21. Adult male F anni a canicularis ...... 40
22. Adult female F anni a canicularis ...... 40
23. Adult male Fannia femoralis ...... 41
24. Adult female Fannia femoralis...... 41
25. Thirty-two ounce polystyrene container with ventilated lid used for rearing stock f l i e s ...... 44
26. C.S.M.A, larval medium with the mold, Rhizopus nigricans growing over the surface ...... 44
27. Sleeve cage of the type used for holding stock colonies of adult flies ...... 46
28. Four ounce polystyrene cup containing fermented C.S.M.A. medium used as oviposition cup for collection of large numbers of e g g s ...... 48
29. Cup used for obtaining first instar larvae by mass incubation of eggs ...... 48
30. Small incubation cup disassembled to show lid, circle of blotting paper and bottom section (cut-off vial). . 50
31. Wooden rack holding cups used in incubation experiments at lethal temperatures ...... 50
X Figure Page
32. Four ounce polystyrene cup filled with C.S.M.A. medium and used for rearing larvae at various temperatures . . 52
33. Pupae placed on inverted cover of rearing cup inside 32 ounce cup ...... 52
34. View of one section of moisture control apparatus .... 56
35. Drawing of a portion of the moisture control apparatus to show detail of construction ...... 58
36. Diagram of the layout of one section of the moisture control apparatus to show how cups were connected to pump ...... 58
XI CHAPTER I
INTRODUCTION
The word "fly" to the average citizen means the common house
fly, Musca domestica L. In fact, until recently it is doubtful that
the majority of urban dwellers knew, or even thought about the exis
tence of other species of flies. The people of California, however,
and particularly those of southern California, have gained experience with flies during the past few years. Since World War II, the human
population of this state has increased at a rapid rate. This increase
has not only been due to births, but to the movement of people in from
other states at an estimated rate of seven thousand per day. This
influx has resulted in a so-called "urban sprawl". Thousands of new
homes have been built on the outskirts of cities, and even new cities
formed. As large agricultural areas have quickly become urban
residential districts, poultry ranches, dairies, beef feed lots, hog
ranches, canneries, vegetable and fruit packing sheds, and frequently
harvested fields containing crop residues are brought within the city
limits. Since flies of several species may breed in wastes from such
enterprises, a fly nuisance and public health problem soon arises. In
some cases the agricultural enterprises move of their own choice or
are sometimes forced out by pressures of complaints from adjacent
residents and the resultant action of local public health authorities. 2
The overall agricultural community, however, is not moving far. The economics of shipping and marketing make it more profitable to main tain large concentrations of poultry and meat animals near the consumer markets. The very economics of animal production plus high land values dictate that the largest number of animals that can be maintained at the peak of health and production be kept on the smallest possible area. It would be difficult indeed to move the vast acreages of California fruits and vegetables completely away from people. These factors all complicate the fly control situation.
One fly that has been second only to the common house fly in its nuisance rating and one which is well known to many in the area is the little or "lesser” house fly, Fannia canicularis (L.). This fly breeds in a variety of organic waste materials and does particularly well in poultry droppings. Literally millions of this species some times breed on improperly managed poultry ranches and then move into surrounding residential areas where their persistent "dancing” and hovering flight in breeze-ways, porches, and inside houses aggravates the human occupants.
Another fly which is less well-known than Fannia canicularis, perhaps because it makes less of a nuisance of itself as an adult, is
Fannia femoralis (Stein). The adult of this species is smaller than that of F. canicularis, and does not move as readily from its breeding area to surrounding areas, but it is still to be reckoned with by the
poultryman and others. Like F. canicularis, it does exceedingly well
in poultry manure, and a "count” of F . femoralis larvae or pupae in the manure on a poultry ranch can cause the ranch to be condemned by 3
health authorities just as readily as a "count" of F. canicularis or
Musca domestical
Like Musca domestica, the two Fannia species have exhibited the
ability to become resistant to the common chemical insecticides used
against them and this factor has further complicated the fly control
problem. Entomologists have long known that so-called "sanitation" or
"manure management" practices which either remove the breeding medium
or render it unfit for fly development by drying or other physical
changes are the most effective control methods. Unfortunately, some
of these methods are not always practical or possible. Certain of the
older types of animal housing construction do not lend themselves well
to the use of modern mechanical manure removal equipment, and hand
labor for this purpose is difficult to obtain, expensive, and slow.
At certain times the demand for manure for agricultural use is low and
it must be stockpiled where it often continues to be a problem as far
as flies are concerned. In periods of wet weather manure cannot be
moved and it will not dry out, thus making the fly problem greater.
Since chemical insecticides have begun to fail as a means of
controlling Fannia species, and since the accepted management prac
tices used in fly control are not always practical, other possible
methods of control are needed. It is well known that the biology of
insects is greatly affected by factors in the environment such as
temperature and moisture. The purpose of this study is to discover the
favorable as well as the unfavorable effects of these two factors on the
development of Fannia canicularis and F anni a femoralis. CHAPTER II
REVIEW OF THE LITERATURE
pgRgral Classification
The species canicularis was erected by Linne^ in 1761 and placed in the genus Muscao The genus Fannia was erected by Robineau-Desvoidy in 1830 with a single species saltratrix (-scalaris). In 1834 Bouch/ erecteti the genus Homalotnyia with canicularis as the type, and with two other species also included, Macquart, in 1835, placed canicularis in the genus Anthotnyia, but left scalaris under Fannia. Haliday (1840) treated Homalotnyia as distinct from Anthomyia and included Fannia within this genus. Until the late 1 8 9 0 most authors continued to use the genus Homalomyia and treated Fannia as a synonym or subgenus, Finally
Stein (1907), in the anthomyiid section of his catalogue, listed
Homalomyia as a synonym of Fannia and the latter classification remains to this day,
Stein described femoralis in 1897 and placed it in the genus
Homalomyia, It became Fannia femoralis in 1907 when Stein synonomized
Homalomyia with Fannia,
Chillcott (1960) divides the genus Fannia into eleven species groupings based largely on male genitalic structure, but also on the basis of structural characters of the ovipositor and spermathecae, and, to a limited extent, on larval structure. His canicularis group is
further divided into the canicularis, glaucescens and pusio subgroups,
4 5
Logically he places the species canicularis in the canicularis subgroup, but he places the species femoralis in the pusio subgroup on the basis
that the latter lack the upper orbital bristle and have a trimaculate
abdominal pattern. While the two species are obviously closely related
on the basis of minute characters, they are quite different in
appearance.
Distribution
Fannia canicularis
This species, like the common house fly, Musca domestica, is
almost world-wide in distribution (Hennig, 1955; Chillcott, 1960). It
is known as the little house fly and in some areas as the "lesser"
house fly. Its range has extended further north than Musca domestica
and it is the only common house fly in Iceland (Chillcott, I960).
In North America the species ranges from Laborador across south
ern Canada, up through western Canada to Alaska, and over most of the
original forty-eight states. In general the heaviest distribution is
across southern Canada and the northern tier of the United States
(Fig. l), but it also occurs in Mexico (Chillcott, 1960).
In the eastern hemisphere F. canicularis is reported from as far
north as Greenland (Henriksen, 1917), Norway (SSinme, 1958), and the
Arctic islands (Meijere, 1909), to New Zealand and Australia (Malloch,
1923). It is known from many other places, for example: England
(Graham-Smith, 1916; Shillito, 1947); France (Lesne, 1921; Roubaud,
1927); Austria (Falcoz, 1927, 1930); Germany (Wilhelmi, 1919, 1920;
Nieschulz, 1935); Yugoslavia (Baranov, 1939); Hungary (Kodocsa, 1934); Fig, 1,--Geographical distribution of Fannia canicularis (after Chillcott, I960). 7
Korea (Kobayashi, 1919, 1930); Japan (Illingworth, 1926); Morocco
(Regnier, 1931; Seguy, 1941); Tripoli (Onorato, 1922); and Rhodesia
(Jack, 1935) to name a few.
Fannia femoralis
-The distribution of F. femoralis is much more restricted than
that of F. canicularis according to Chillcott (1960). He lists
records from Montana, Utah, Kansas, Illinois, North Carolina, Georgia,
Alabama, Louisiana, Texas, New Mexico, Arizona, California and Mexico
City. It is also reported from Maryland by McAtee (1929), and
Tennessee (Reed, 1958). In South America it is reported from Bolivia
and northern Argentina by Engel (1931).
In general the major distribution of this species seems to be
along the southern tier of the United States and into Mexico and
South America (Fig. 2).
Importance to Man
Fannia canicularis
This species is probably capable of breeding in all habitats
recorded for the subfamily, Fanniinae, to which it belongs, although
it apparently establishes itself in areas of human habitation where
decaying organic matter is easily available (Chillcott, I960). As a
result of its activity in some of these habitats, it becomes merely a
nuisance. In others it constitutes a serious hazard to the health of
man and animals, but in some cases it may be considered beneficial.
It has been recorded from several kinds of animal manures:
chicken (Smith, 1954a, 1954b; Tilden and St. Germaine, 1956; Anderson Fig. 2,--Geographical distribution of Fannia femoralis (after Chillcott, 1960). 9
and Poorbaugh, 1964); pig, calf, horse (Thomsen and Hammer, 1936); cow
(Baranov, 1939; Scîmme, 1958; Ogden and Kilpatrick, 1958); rabbit, (St.
Germaine, 1956). It has also been found in maceration tanks for animal and human feces in Japan (Illingworth, 1926); latrines (Howard
1900; Kobayshi, 1919); and in cesspools in Paris, France (Lesne, 1921).
It is known to infest the nests of birds (Falcoz, 1923; Keilin, 1924;
McAtee, 1929; Norberg, 1936); and the nests of deer mice (Barnes,
1958). Chillcott (i960) reported that he had specimens which had been reared from squirrel, meadow mouse, and kittiwake nests. It is re ported from a wide variety of decaying plant materials such as oozing sap from diseased elm trees (Shillito, 1947); rape stalks and sugar beets (Hewitt, 1912); decaying grass or piled grass clippings (Hewitt,
1912; St. Germaine, 1956); iris buds (Kodocsa, 1934); tomatoes (Nelson,
1938); decaying nasturtiums (Laurence, 1958); and high protein cow feed (Smith, 1958). Chillcott (i960) reports that he has seen specimens that had been reared from decaying plums, peas, corn and onions. Falcoz (1927, 1930) reared it from fungi. It has also been found in decaying animal matter such as dead grasshoppers (Nocedo,
1-918; Regnier, 1913; Jack, 1935); dog carcasses (Reed, 1958); cat
carcass (Illingworth, 1926); snails (Hewitt, 1912; Keilin, 1919;
Mokrzecki, 1923); and wool of sheep infested with golden-haired blow
fly, Pollenia stygia (Fallen) (Miller, 1922). Tiensuu (1936) found
adults feeding on aphid honeydew.
One of the most important aspects of F. canicularis is its role
in causing myiasis in man and animals. Several types have been reported
such as aural, intestinal, urinary, and vesicular in man, and cutaneous 10 in monkeys (Hewitt, 1912; Onorato, 1922; S^guy, 1925; Green, 1956;
James, 1947). Burnett, et £l (1957) reported F. canicularis as a vector of the nematode, Thelazia califomiensis Price, an eye worm in mammals, including man, in California.
Surprisingly, F. canicularis, in addition to being coprophagus or saprophagus in its habits, is also listed as being parasitic on several species of insects. Seguy (1932) discusses several genera of flies, including Fannia, which have similar habits, and places them into several groups according to these habits. He classifies Fannia along with other genera as follows:
1. Larvae saprophagus or coprophagus, occasionally parasites.
2. Larvae ubiquitous, attacking other insects normally or occasionally,
3. Larvae normally parasites of grasshoppers.
a) Occasional parasites of the eggs, young and adults.
It is not clear why he places Fannia in more than one category, but
presumably its activity depends upon the presence of susceptible host
species and suitable environmental conditions. F . canicularis is
listed as a parasite of the eggs of Dociostaurus marocanus (S^uy,
1932); of the adults of Schistocerca gregaria (Rdgnier, 1931); of adults of Schistocerca peregina (Nocedo, 1918); and of the larvae of the black walnut curculio, Conotrachelus iuglandis Lee. (Brooks,
1922).
Since poultry manure is a favorite breeding place for Fannia, a discussion of its role in the fly problem is in order. Tilden and St.
Germaine (1956), speaking of poultry production in Santa Clara County, 11
California, say that, "Chicken raising for profit has evolved into a highly specialized and competitive industry. Few poultrymen keep their
laying hens on the ground. For economic reasons, single and multiple cage operations have become the rule. Feed manufacturers have developed
feeds which get top production from laying hens and which put weight on meat birds rapidly. The age of specialization in chicken ranching has
resulted in more eggs, more meat, and unfortunately, more flies.The
same authors go on to explain that few flies are found breeding in
scattered poultry droppings of a ground or litter operation, but that manure in dropping pits beneath modern cage operations can, under
certain conditions, produce tremendous numbers of flies.
Smith (1954a) stated that a health department survey showed up
to 17,560 F. canicularis larvae per square foot of surface under cages
and in aisles of a poultry ranch. Anderson (1966) quotes some
estimated figures that about 60 million tons of manure were produced
by poultry in the United States in 1963. With that kind of production
of fly breeding medium the potential production of Fannia canicularis
and other species is tremendous!
Many others have reported on F. canicularis populations, Howard
(1900) rated Homalomyia (Fannia) canicularis as moderately abundant on
human excrement in samples from army camps all across the country. In
his survey he found Musca domestica to comprise 98.87» of all flies
collected, but the remainder, consisting of 1.27. of the whole,
^Tilden, J. W., and J. St. Germaine, (1956), Flies of Public Health Interest in Santa Clara County. California Vector Views, II No. 3, 12. 12 comprised various species, the most significant one being Homalomyia
(Fannia) canicularis. Somme (1958) found that Fannia spp. (mostly canicularis) made up 10.2% of flies in stables in Norway. Illingworth
(1922) foundF. canicularis the most abundant species, "literally swarming" in the hotel where he stopped in Waimea, Hawaii. Ogden and
Kilpatrick (1958) reported that F. canicularis populations in untreated dairies in Cache Valley, Utah reached 1500 per 500 square feet of floor space. St. Germaine (1956), in sampling piled lawn clippings, found as many as 3 , 0 0 0 fly larvae and pupae in a single pint of decomposing grass. Many of the flies in lawn clippings were F. canicularis. Smith (1954b) reported that F. canicularis bred in moist
poultry manure and that 10 to 60% of the complaints about flies in
California were a result of flies from chicken ranches. Smith (1958)
in reporting on a case of F. canicularis breeding in spilled high
protein dairy feed estimated populations of 4 5 0 , 0 0 0 larvae and pupae
per 36 square feet of area.
Fannia femoralis
References in the literature to the habits and the economic
importance of F . femoralis are almost completely lacking. It would appear that this fly, except for the problem of its presence on
poultry ranches as cited in the introduction of this paper, and a few
other cases, has caused little trouble to man. Tilden (1957) listed
F. femoralis from cow manure, but rated it as scarce. Reed (1958)
found this species in dog carcasses in Tennessee. Illingworth (1926b) 13 found it breeding in a cat carcass in California. McAtee (1929) collected F. femoralis. along with F. canicularis, from the nest of the starling, Stumus vulgaris in Maryland.
Biology
Life History of Fannia canicularis
A number of workers have reported on the life history of F, canicularis. The majority of these reports appear to be based on observations. Only two or three seem to have actually studied the life cycle of this insect under controlled conditions.
Wilhelmi (1919) says that oviposition continues from the beginning of warm weather until mid-October. He gives the size of the eggs as 0.8 mm long and says that they hatch in two days giving rise to dirty white bespined larvae. He says that the eggs are laid in moldy, decaying plant material and that the adults enter freely into houses and stables like M. domestica.
Lodge (1918) goes into more detail on the life history of F, canicularis. but fails to give the temperature under which he made his
studies. He says that the incubation period is 1 to 3 days and that
if eggs are allowed to get too dry that they will not hatch. His
observation was that if the eggs were moistened that they would hatch
sooner, a longitudinal split often beginning as soon as the water was
applied. He says nothing of different larval instars, but states that the "larval stage" lasts from 11 to 21 days with an average of 14 to
15 days. He gives the pupal stage as lasting from 13 to 34 days with 14 an average of 21 to 22 days. He says that the largest number of adult flies emerged after 19 to 26 days.
Thomsen and Hammer (1936) give little detail of their work, but state that in experiments the shortest duration of development was 18 days at a temperature of 21 to 26 C (approximately 70 to 80 F) and 21 days at 18 to 23 C (mainly 22 C) (approximately 66 to 73 F).
Lewallen (1954) was the first to give a detailed description of the life cycle of F. canicularis under controlled temperature and relative humidity conditions. He worked at a temperature of 80 F and a relative humidity of 6 5 % . He describes the various stages briefly and gives their duration as follows:
The eggs hatch in 36 to 48 hours.
The larval stage lasts 8 to 10 days.
First Instar: Translucent, white, with only tip of mouth hooks black; 1.5 mm long; duration 24 hours.
Second Instar; Translucent, white; middle portion of cephalo- pharyngeal skeleton black, as are the mouth hooks; 3 mm long; duration 24 hours.
Third Instar: Light-brownish color; cephalo-pharyngeal skeleton entirely black; mature larva 6 to 7 mm long; duration 5 to 7 days.
Pupal Stage: Characterized as merely quiescent larvg that has ceased feeding, drawn in the cephalic region, and become somewhat shortened, with integument hardened and dark. Stage lasts 8 to 12 days.
Adults: 5 to 6 mm in length.
Steve (1960) also studied the life history of F. canicularis under controlled conditions. He, like Lewallen, reared the flies at a temperature of 80 F and a relative humidity of 6 5 % . He gives the duration of the egg stage as from 1% to 2 days. He fails to give the 15 duration of each of the larval instars, but gives the duration of the entire larval stage as 8 to 10 days and says that the total develop mental period from egg to adult was 18% to 22 days. Steve also measured the various stages. He gives the size of the eggs as ranging from 0.8 mm to 0.95 mm. He goes on to say that his measurements agree with those of Lewallen except for the egg which the latter gave as
2.0 mm.
Fay, e^ al^ (1963) reared F. canicularis at a temperature of 75 F and 70% relative humidity. They give the average time required for the various developmental periods as follows: egg, 24 to 36 hours; first and second larval instars, 1 day each; and third instar, 4 to 6 days. They do not state the duration of the pupal stage, but give the median developmental period from egg to adult as 24 days with a range
of 19 to 29 days.
Life History of Fannia femoralis
No references on this subject could be found in the literature.
Overwintering of Fannia canicularis
Several workers have published papers on the overwintering methods of Muscat domestica and other species of flies, including Fannia
canicularis. There is a great deal of controversy among these writers
as to whether overwintering in most species occurs as the adult, or as
larvae or pupae. Hewitt (1915) ran repeated tests with M. domestica
in England and Canada, but failed to get pupae to live through the
winter outdoors. He failed, also, to find flies breeding under natural
conditions in stables in the winter and concluded that flies overwinter 16 as adults in warm rooms and other protected places where food was available.
Lyon (1915), working at Harvard University, ran experiments with
M. domestica where he placed pupae in various kinds of media in glass jars and exposed them to winter conditions both indoors and outdoors.
He found that no flies emerged from the jars outdoors, though some were in a sheltered location, and the weather was mild. He concluded that the common house fly could not easily overwinter as a pupa outdoors.
Howard (1917) working in Minnesota placed M. domestica pupae in moist soil in small, 2X2/4 inch wooden boxes and exposed them to low temperatures. He found that the pupae were killed in 1 week at 12 F; at 30 F, 237o survived, but all died at longer exposure than one week.
At 40 F 21% survived and produced adults, but none survived more than
1 week at 40 F . The author concluded that Minnesota winters are not favorable to the overwintering of the house fly in any except the adult stage, and that stage, only in places where there is sufficient high temperature and where food conditions are favorable.
Wilhelmi (1920) took the viewpoint that M. domestica. F. canicularis, and other species of flies overwinter by more or less living with man in his houses. He was referring to adults living actively in warm rooms where food was available. To this extent, he agrees with Hewitt (1915) and Howard (1917), but he goes a step further and says that overwintering as an immature or preadult
(embryonal, larval or pupal) is also probable in F. canicularis. He does not explain further.
Several workers disagree sharply with the authors cited above in 17 saying that M. domestica, F. canicularis and other species definitely overwinter as larvae or pupae. Skinner (1913) made some observations in the entomological rooms of the Academy of Natural Sciences of
Philadelphia. He noted that no flies were observed during the winter in the rooms, but as soon as the first warm days of early spring arrived, the windows were opened and flies came in. Some were caught and found to be all fresh specimens, even teneral in character, except in color. The ptilinum was not completely retracted and it was evident that they had just emerged. He observed that the colors were bright and the wings perfect and not frayed in any way, and there was no sign of dust, dirt or decay on them. He concluded that until disproved, "house flies pass the winter in the pupal stage and no other way."^
Bishopp et £l (1915), who studied the overwintering of flies in
Texas, concluded that no adults overwinter outdoors, but that pupae do.
Dove (1916), working alone, concurred with Bishopp.
Graham-Smith (1916) discusses overwintering of flies at length.
He says definitely that F. canicularis lived through the winter as a larva or pupa in experiments which he conducted.
Kisliuk (1917), in work done at College Park, Maryland, and at
Columbus, Ohio, with Musca and Fannia found that the adults were not able to survive the winter even inside houses, but the pupae success fully lived through the winter outdoors in manure piles.
^Skinner, Henry, (1913), How Does the House Fly Pass the Winter? Entomological News, XXIV, No. 7, 304. 18
Hutchison (1918), working at Bethesda, Maryland, concluded that
adult house flies could not overwinter outdoors, nor could they live more than 40 days indoors. He cites two experiments with larvae placed
in manure piles beneath emergence cages. In one of these, larvae
placed under a cage on January 10 resulted in the emergence of several
adult F. canicularis between the dates of March 22 and June 4. In a
similar test, the last larvae were introduced on October 24. Between
April 2 and June 4 many flies emerged among which were 33 F. canicu
laris .
Mellor (1919) discusses the overwintering of flies at length.
He lists several species as overwintering as larvae or pupae. F.
canicularis is among these.
Roubaud (1927) made a study of the life cycles of several
species of flies and distinguished among them two types. The first
type he calls "homodynamic" and gives such species as M. domestica and
Stomoxys calcitrans as examples. He says that these are endowed with
a continuous developmental activity from generation to generation and
that it is only suspended by cold. The second type he calls "hetero
dynamic" and says that F. canicularis belong in this group. In this
type several active generations are followed by an "affected"
generation or one which he describes in French as being affected by
"asthenobiosis" (sick life). He speaks of this condition as diapause
and says that it does not depend upon cold, but coincides solely with
the appearance of winter. He speaks of F. canicularis more specifi
cally as having four successive generations per season. The first
three generations are active. The first develops in about 20 to 23 19 days at 20 C; the second, descended from the previous one, develops in
20 to 25 days, but with a pupal delay slightly more accentuated (10 days compared to 8^). The third generation presents the same char acteristics as the second. As for the fourth generation, he says that it is affected at the third larval stage by a "pseudo-winter” diapause which "does not yield to the prolonged influence of cold."^ He goes on to say that, "we therefore have an example of one species with several active generations in the spring and fall followed abruptly by 2 a generation of torpid larvae called to obligatory hibernation."
Rearing Media
Workers rearing muscoid flies have used a variety of media.
These have ranged from very common single materials to complex mix tures. The choice of media has often been governed by the avail ability of materials. Hutchison (1916) used fresh horse manure, or he boiled it for one-half hour to kill any fly eggs that may have been laid in it before it could be collected. Glaser (1924) tried fer menting bran or hay or horse manure. He found horse manure to be the best, but noted that flies did well in it only from April to mid-Dec ember when they died. Glaser (1927) later concluded that horse manure was deficient in certain nutrients during the winter and early spring.
He did not know whether the deficiency was due to the lack of bac terial flora or to the horses’ winter diet. He found that the
^Roubaud, E., (1927), Sur 1’hibernation des quelques mouches communes. Bull. Ent. Soc. France, p. 25.
^Ibid., p. 25. 20
addition of a few cubic centimeters of a heavy suspension of yeast
cells to the horse manure every few days corrected the deficiency, and
the flies bred continuously through the year. Grady (1928) had the
same experience with horse manure and followed Glaser’s method of
adding yeast with similar success. Hockenyos (1931) first followed
Glaser’s and Grady’s method of using horse manure and yeast. He later noticed that during the summer, fly maggots were more abundant in heaps
of hog manure than in piles of horse manure and attempted to adapt the
hog manure for laboratory use. He found that hog manure alone tended
to pack and exclude the necessary air, but that a mixture of one part
fresh straw-free hog manure with three or four parts fresh straw-free
horse manure gave the best results. Grady (1937), trying to improve
on his horse manure-yeast mixture of 1928, added malt extract.
Kobayashi (1930) working in Korea took an entirely different
approach than previous workers mentioned above by rearing flies on a medium of biscuits of soybean origin moistened with peptone water.
Richardson (1932) introduced a slightly more complex mixture which he
felt was better than horse manure because it was free of mites and
other predators that were often found in horse manure and it was clean
to handle. It consisted of wheat bran, alfalfa meal, bakers’ yeast,
Diamalt and water. These became the basic ingredients for a series of
media known as the Peet-Grady formula, the N.A.I.D.M. (National
Association of Insecticide and Disinfectant Manufacturers) medium, and
presently, as the C.S.M.A. (Chemical Specialties Manufacturers
Association) medium. A series of workers including Doty (1937),
Woodbury (1943), Harrison (1949), Pimentel et ^ (1951), Smith and 21
Harrison (1951), Babers ^ ^ (1953), Lewallen (1954), McKenzie and
Hoskins (1954), Moreland and McLeod (1957), Steve (i960), Keiding and
Arevad (1964), and Schoof (1964) all used variations of this formula.
In fact, the formula is presently used widely by those working with
flies around the world.
In spite of the satisfactory results with the C.S.M.A. medium,
as it is now best known, a number of workers have modified it, or
compared it with other materials. Basden (1947) compared Richardson's
medium, the Peet-Grady medium, and one of his own which had middlings
because they allowed the puparia to be obtained in a cleaner condition.
Grass meal was used because alfalfa meal was not readily available.
He found dry malt extract more convenient to use and store than
the liquid extract. Incho (1954) and Goodhue and Cantrel (1958)
found that the addition of vermiculite improved the Peet-Grady
or C.S.M.A. medium. The former added two inches of vermiculite
to the surface after seeding the medium with eggs. The latter
mixed it directly with the medium. B o m (1954) and Champlain £t a^
(1954) in rearing stable flies (Stomoxys) had a problem with mold
growing over the surface of the C.S.M.A. medium. They found that
the addition of sand to the surface controlled the mold and provided
a place for the larvae to pupate. McGregor and Dreiss (1955) rearing
stable flies found that the addition of five parts wood shavings to
one part C.S.M.A. made a most suitable medium. B^ggild and Keiding
(1958) used the Peet-Grady formula, but found that the addition of
milk improved it. Louw (1964) also favored the use of milk and
specified full cream powdered milk. Levinson (1960), reasoning that 22 bacteria are numerous in food upon which flies feed in nature, tried
C.S.M.A. medium both with and without the bacterium, Escherichia coli.
He found that the addition of bacterium to autoclaved C.S.M.A. medium greatly increased the growth of house fly larvae. Fay ot £l (1963) were apparently satisfied with C.S.M.A. medium, but added coiled strips of corrugated cardboard to the top of the medium as a pupation site. The majority of larvae pupated in the grooves of the cardboard.
The cardboard was later stripped from its backing and the pupae fell free into a collecting container. Monroe (1960), Robbins and Shortino
(1962), Spiller (1964), and Ashrafi (1964) all found that the addition of cholesterol to the C.S.M.A. medium improved it. The first three noted improved ovarian development and egg hatch, while Ashrafi found growth improved.
Sawicki (1964) compared the attributes of C.S.M.A. medium with a
YMA (yeast, milk, agar) medium. He felt that the latter was better in that it should contain no insecticides as C.S.M.A. may. He also noted that there was no rise in temperature and that separation of pupae was easier. He did concede that the YMA was attacked by fungi and bacteria and often smelled bad, and the high price of powdered agar probably made it more expensive than C.S.M.A. medium.
Many very simple media and some rather strange ones have also been used. Eagleson (1943) was aware of the successful use of horse manure and of Richardson's medium, but he preferred a simpler one consisting only of crimped oats and water. He found that the oat hulls contributed an essential flocculence to the medium and prevented the formation of tight, gummy masses. Glaser (1938) used a mixture of 23 minced, fresh swine liver for the sterile culture of flies. He heated the liver until the proteins coagulated then mixed it with fine pine sawdust and Harris Brewer's yeast. Nagel (1962) and Pimentel and
Al-Hafidh (1965) used an agar-skimmed milk-brewer's yeast medium which was supplemented with fresh beef liver. Frings (1948) used dog biscuit to which yeast and water had been added and was fermenting.
He added sawdust or shavings to make the medium more suitable for rearing blow flies. Phormia regina. Hafez (1949) followed one of the simplest methods. He collected house fly eggs on a cotton wool pad which had been soaked in diluted milk. After the eggs were laid, the pad was transferred to a two-pound jam jar containing a fresh milk pad. Full grown, healthy larvae were obtained after four or five days.
Nagasawa and Hashizume (1955) and Nagasawa and Kishino (1959) reared
Musca domestica vicina Macq. using the "okara" culture medium, "okara" being a residual product of "tofu" making. They used 50 grams of
"okara", 5 grams of rice bran and 0.5 grams of powdered beer yeast.
Fisher and Jursic (1958) reared house-SIies on a mixture of ground absorbent cellucotton, evaporated milk, dried brewer's yeast and water. At the end of the fifth day, sawdust was placed on top of the medium as a pupation site. Kuenen (1958) used a similar formula of powdered skim milk, dried yeast, cellulose tissue and water, but added some 0.1 normal KOH. Spiller (1963) used equal parts of ground
(flocked) tissue paper and dried whole cream milk powder and yeast. 24
One of the most interesting of all the media is one used by
Tharumarajah and Thevasagayam (1961) in Ceylon, Coconut poonac, the cake left over after the extraction of oil from coconut pulp was placed in battery jars and used very successfully in rearing Musca domestica vicina. The flies laid their eggs in the poonac and the larvae lived and pupated in the same medium. CHAPTER III
RELEVANT MORPHOLOGICAL ASPECTS
During the course of the study of the effect of temperature and moisture on the development of Fannia, several interesting morpholog ical aspects of F. canicularis and F. femoral is have been brought out.
Some of these aspects have been very basic to the study itself. For example, a study of the effect of temperature on larval development requires easy and accurate recognition of the various larval instars.
A discussion of some of these aspects is, therefore, felt to be worthwhile.
The Eggs
The eggs of Fannia spp. are quite different from those of Musca domestica in that they have a pair of thin laterodorsal wings or flanges of chorion which extend almost the entire length of the egg
(Fig. 3). These flanges are folded up over the dorsal surface of the egg at the time it is laid but they almost immediately unfold into an almost horizontal position (Fig. 4). Chillcott (1960) speculates that they serve as a flotation mechanism when eggs are laid in a liquid medium. The flanges also serve to divide the dorsal from the ventral
surfaces of the egg which are quite different in pattern. The dorsal
surface is divided down the center line by a longitudinal ridge. On either side of the ridge the surface is characterized by a raised
25 26
Fig. 3.--Eggs of Fannia fetnoralis. The laterodorsal wings or Flanges and the reticulated pattern on the dorsal surface are also typical of F. canicularis.
Fig. 4.--Drawing of end view of Fannia canicularis egg showing position of wings or flanges after egg is laid. 27 reticulated or somewhat diamond pattern (Fig. 5). The ventral surface
is characterized by a longitudinal ribbing running the entire length of the egg (Fig. 6). The eggs are elongate oval with the anterior end
slightly more pointed than the posterior end.
Eggs of F. canicularis are approximately 0.8 mm long while those
of F, femoral is are approximately 0.75 mm. F. canicularis eggs are
also larger in diameter than F. femoralis eggs and the flanges or wings of the egg are narrower in proportion to the diameter than in
F. femoralis.
Fannia eggs are characterized by having a mucilage or adhesive
on the surface which causes them to adhere very tightly to surfaces
once the adhesive has dried. This adhesive is water soluble and it is
almost impossible to collect eggs from surfaces unless the eggs are
first moistened. The adhesive of F. canicularis eggs is apparently
more readily water soluble than that of F . femoralis as the amount of
water and the period of time required to loosen them is less than for
the latter.
Fannia eggs may be laid singly or in masses depending upon
conditions. They are laid directly on diamalt or milk-moistened
cotton pads, or on fermented C.S.M.A. medium in the laboratory. The
diamalt moistened pads or C.S.M.A. are preferred over the milk
moistened pads. The flies will also lay eggs in a row along the rim
of the polystyrene cup as described under "Egg Collection" in the
chapter on materials and methods. 28
Fig. 5.— Drawing of dorsal aspect of Fannia canicularis egg showing reticulated pattern on the surface.
Fig. 6.-- Drawing of ventral aspect of Fannia canicularis egg showing longitudinal ribbing on surface. 29
The Larvae
The larvae of Fannia are unlike the larvae of other members of the Muscidae. Rather than being the typical smooth, almost cylindrical, tapering, white muscoid type, Fannia larvae are somewhat depressed dorsoventrally, have a spiny appearance, and are brown when fully developed. They are actually not spiny, but have rows of soft pointed processes located dorsomedially, dorsolaterally and ventrolaterally.
These processes are somewhat plumose at their bases, especially those on the caudal, segment. Toward the apex the processes have shorter spinules or projections making them less plumose and more rat-tail like in appearance. This characteristic is more marked in F. canicularis (Fig. 7) which has comparatively longer processes than
F. femoralis (Fig. 8).
A pair of posterior spiracles are located dorsomedially on the caudal segment and are on short stalks. These are useful in dis tinguishing the various larval instars. Chillcott (1960) says that the number of openings are equal to the number of instars. On close examination, it was found that the first instar actually has two openings, and the second and third instars four openings. In the first instar the spiracular stalk is short and almost pear-shaped with the two openings close together at the apex (Figs. 9 and 10). In the second instar the stalks become somewhat longer and more cylindrical with the four openings close together at the apex (Fig. 11). In the third instar the stalk divides at the apex into three relatively long and one short finger-like process with an opening at the apex of each process (Fig. 12 and 13). 30
Fig. 7.--Drawing of dorsal aspect of Fannia canicularis larva showing long rat-tail-like processes which are characteristic of this species.
Fig. 8.--Drawing of the dorsal aspect of Fannia femoral is larva showing characteristic dorsal and lateral processes. 31
Fig. 9.--Photomicrograph of the posterior spiracular stalks of the first instar larva of Fannia femoralis showing the two spiracular openings characteristic of this stage.
Spiracular openings Spiracular stalk
Trachea
Fig. 10.--Drawing of the posterior spiracular stalks of the first instar larva of Fannia femoralis shown in Figure 9. Note position of the two spiracular openings close together at the apex of each stalk. 32
Fig. il.--Photomicrograph of the left posterior spiracular stalk of the second instar larva of F anni a femoralis. Note the four spiracular openings close together at the apex of the stalk which is characteristic of this stage.
Fig. 12.— Photomicrograph of the left posterior spiracular stalk of the third instar larva of Fannia femoralis. Note that the stalk is divided at the apex into three long and one relatively short finger-like process, each with a spiracular opening at its apex. 33
-Spiracular stalk
Trachea
Fig. 13.--Drawing of the left posterior spiracular stalk of the third instar larva of Fannia femoralis shown in Figure 12 showing location of the four spiracular openings.
Fig. 14.--Photomicrograph of the cephalic region of the first instar larva of Fannia femoral is showing the horseshoe-shaped pharyngeal skeleton. 34
Another characteristic of Fannia larvae which is useful in distinguishing the instars is the shape of the pharyngeal skeleton.
In fact, this is almost essential in distinguishing the first and second instars without the aid of a compound microscope. First, second, and early third instar larvae are quite transparent and the pharyngeal skeleton can be easily seen at relatively low power mag nification with an ordinary dissecting microscope or a hand lens.
The pharyngeal skeleton in the first instar larva appears in the cephalic region as a black horseshoe-shaped structure, the open end caudad. Any connecting structure between the pharyngeal skeleton and mouthhooks is obscure (Fig, 14). In the second instar the pharyngeal skeleton has become sclerotized further back, appearing narrowly U- shaped and black with the open end of the U to the rear. There is obvious connection with the mouthhooks (Fig. 15). Further sclero- tization of the pharyngeal skeleton toward the rear in the third instar reveals a slight narrowing of the U and an extension of the two legs of the U toward the rear (Fig. 16).
When a larva is preparing to molt it becomes sluggish, rather opaque in appearance, and stops feeding. Soon rhythmic muscular contractions move along the body from anterior to posterior. The process may continue for more than an hour until finally the cuticle splits along the median dorsal line. With what appears like one last great contraction the larva frees itself of the old mouth hooks and pharyngeal skeleton and rather rapidly crawls out of the old skin
(Fig. 17). 35
Fig. 15.--Photomicrograph of the cephalic region of the second larval instar of Fannia femoral is showing the narrowly U-shaped pharyngeal skeleton and its connection with the mouth hooks.
Fig. 16.--Photomicrograph of the cephalic region of the third larval instar of Fannia femoralis showing the narrowing of the U-shape and extension of the legs of the U to the rear. 36
The Pupae
Pupation takes place inside of the old skin of the third instar
larva. As the larva prepares to pupate, it crawls from the more moist medium into drier material, stops feeding and becomes quiet. It
shortens in length slightly and the dorsum tends to become somewhat
arched. Gradually the mouth hooks and remainder of the cephalic
region are withdrawn leaving the prothoracic area thin and flattened
with a slight depression or groove down the dorsomedial line. While
the foregoing is taking place the old cuticle is hardening and dark
ening to become the puparium (Fig. 18).
A useful feature of both the larvae and pupae is the character
of the venter which makes it very easy to distinguish the two species.
F. canicularis larvae and pupae have a median transverse ridge or
Carina on each of the abdominal sternites, while F . femoral is lacks
this ridge. It will also be noted that F. canicularis has an anal
opening forming a narrow "V" with the apex of the "V" pointed anter
iorly. F, femoralis, on the other hand, has a more oval or wide "V"
shaped anal opening with the apex pointed anteriorly (Figs. 19 and 20).
Well-fed adult male Fannia canicularis specimens are almost as
large as the common house fly, Musca domestica, but with a much more
slender thorax and abdomen. The thorax is brownish-grey in color with
medium brown vittae dorsocentrally located. There is also medium
brown on the lateral aspects of the thorax. The abdomen is brownish
grey-pollinose, the median vitta distinct and triangulated. The
second to the fourth segments are translucent yellow basally. F .
canicularis females are similar to the males in general appearance 37
■■ ■4 ‘
'î’> 1^"v
- rf.
Fig. 17,--Photomicrograph of the shed skin of the second larval instar of Fannia femoralis showing the pharyngeal skeleton and mouth hooks attached.
Fig. 18.--Puparium of Fannia femoralis on the left, and that of F. canicularis on the right. 38
Fig. 19.— Drawing of the ventral aspect of Fannia canicularis larva. Note median transverse ridge on each abdominal stemite and the anal opening forming a narrow *'V" with the apex pointing anteriorly.
Fig. 20.--Drawing of the ventral aspect of Fannia femoral is larva. Note absence of median transverse ridge on abdominal sternites and oval or wide "V” shaped anal opening with apex pointed anteriorly. 39 except that the vittae are usually paler and only the second and parts of the third abdominal segments are yellowish (Pigs. 21 and 22).
F. femoral is adults are smaller than those of F. canicularis.
The thorax and abdomen are thicker in proportion to the length than
in F. canicularis, giving the fly a more "stocky" appearance (Figs. 23
ahd 24). F. femoral is adults range from an almost shiny brownish-
black to a bluish-grey in color in both sexes.
F. canicularis is easily disturbed and flies readily, while F.
femoral is is almost sluggish in movement and sometimes will not fly
unless touched or its resting place vigorously shaken. Both species move in a hovering, "dancing" flight. F. canicularis. particularly,
is frequently seen in breezeways, open buildings, rooms in dwellings,
or under the shade of trees flying about in groups or swarms and never
seeming to alight on surfaces. Most of these flying groups are males,
and if food or breeding material is nearby, the females can usually be
found resting there, feeding and ovipositing. 40
Fig. 21.--Adult male Fannia canicularis.
Fig. 22.— Adult female F anni a canicularis 41
Fig. 23.--Adult male Fannia femoral is■
Figt 24.--Adult female Fannia femoralis. CHAPTER IV
MATERIALS AND METHODS
Colony Establishment
Stock colonies of flies were started from pupae collected from poultry droppings in a chicken house at Moorpark, California. The pupae were easily obtainable from the surface of a concrete floor along the edge of an aisle after carefully raking the dry droppings
aside. The pupae were sorted according to species and placed in small
plastic cups inside small holding cages where they were allowed to remain until the adults emerged. When the adults were sexually mature they were allowed to oviposit and the eggs carefully collected to avoid contamination by various mites and other undesirable arthropods.
Colonies were established by placing these eggs in a modified C.S.M.A. medium obtainable from the Ralston Purina Company, St. Louis,
Missouri.
Rearing
The containers used for rearing of stock flies were 32 ounce 3 polystyrene cups 4 /4 inches tall and 4% inches in diameter at the
lip. These were filled about one-half full of the larval medium and
covered with a polyethylene snap-on lid fitted with a 3-inch diameter
circle of 100 mesh brass sieve cloth in the center for ventilation
(Fig. 25). Each cup contained enough food for the development of 200
42 43 to 300 flies. The brass sieve cloth was easily fastened over a 2 \ inch hole cut in the lid by merely pressing the edge of the sieve cloth into the plastic with a hot soldering iron.
The C.S.M.A, mix was modified and mixed in the following proportions;
300 grams dry C.S.M.A, 26 grams dry powdered brewer's yeast 600 milliliters water
The dry ingredients were first mixed thoroughly in a dishpan, then water was added and thoroughly mixed. The quantity indicated made enough for three of the rearing cups described above. It was found that eggs placed on fresh medium usually produced few, if any, flies.
A heavy growth of bread mold, Rhizopus nigricans Ehr., develops over the surface of the medium by the second day after mixing, and the temperature of the interior of the medium is quite high because of the fermentation reaction (Fig. 26). If the mold is allowed to develop and then is stirred in with a small spatula, before the fly eggs are introduced, the larvae and pupae develop very successfully.
Adult Maintenance
As soon as the adults emerged in the rearing cups they were transferred to holding cages. The holding cages were constructed of
3/ 3/4 inch plywood and were 13% Inches wide by 12 4 inches deep. They 3 were 12/4 inches tall at the front and the glass top sloped back to a height of 16 inches at the rear. The back was covered with unbleached muslin. The front was fitted with a hinged door 9^/4 inches tall by
10% inches wide for ease of cleaning. In the center of the door was a 44
Fig. 25.— Thirty-two ounce polystyrene container with ventilated lid used for rearing stock flies.
Fig. 26.--C.S.M.A. larval medium with the mold, Rhizopus nigricans growing over the surface. 45 muslin sleeve 6 inches in diameter by 18 inches long for use in in serting and removing food, flies or oviposition cups (Fig. 27).
Adult flies were provided with two types of food. One was made from Fleischmann's Regular Diamalt, a pure diastatic malt syrup obtained from Standard Brands Incorporated, Stamford, Connecticut.
The diamalt was diluted by adding 5 parts water to 1 part diamalt.
The other type food was 1 part canned evaporated milk diluted with 1 part water. Each of these mixtures were provided fresh every two days on a cotton pad in a 4 ounce plastic cup. It was found that the flies could live very well on the diamalt diet alone, but very few eggs were laid until the milk was also provided.
The larval rearing cups and adult stock cages were kept in a small rearing room in an insectary building where the temperature and relative humidity were automatically controlled at 75 F and 50 to 60% respectively. The lights were controlled by a clock set for a 13 hour period of illumination.
Larval rearing medium could not be placed in the stock cages with adults and allowed to remain while the larvae and pupae developed, as it was found that eggs continued to be laid in it day after day. This resulted in too many larvae for the quantity of food and many flies emerged undernourished and small.
Egg Collection
Eggs for inoculating the 32 ounce larval rearing cups or for use in experiments were collected in 4 ounce oviposition cups usually within one to a few hours during each day. These cups were of clear 46
Fig. 27.— Sleeve cage of the type used for holding stock colonies of adult flies. 47 polystyrene plastic and were filled about 2/3 full of the two-day old
C.S.M.A. mix used for rearing larvae. The medium at this stage had begun to be broken down by microorganisms and fungi, had a definite ammonia odor, and was extremely attractive to adult flies. Frequently^ so many eggs were laid on the medium and around the lip of the cup that after moistening with water,large masses of eggs could be scraped up with a spatula or camel’s hair brush (Fig. 28).
Obtaining Larvae
When newly hatched first instar larvae were needed for exper iments, eggs were collected from the oviposition cups and placed in specially prepared cups for mass incubation. These were 4 ounce poly styrene cups Identical to those used for oviposition except that a cotton pad saturated with water was placed in the bottom and covered with a circle of blotting paper. The blotting paper was cut to fit the cup and before placing on the saturated cotton was wet with a 0*2% formalin solution to inhibit the growth of mold. The fly eggs were collected for a few hours of one day and then distributed thinly and evenly over the surface of the blotting paper (Fig. 29). A poly ethylene snap-on lid was then wet with many small droplets of water on the underside and snapped onto the cup. When the larvae hatched, many crawled up onto the underside of the lid and became entrapped in the small droplets of water. The larvae could then be easily picked off the lid with a small camel's hair brush when needed. 48
Fig. 28.--Four ounce polystyrene cup containing fermented C.S.M.A. medium used as oviposition cup for collection of large numbers of eggs.
Fig. 29.— Cup used for obtaining first instar larvae by mass incubation of eggs. Note water saturated cotton pad in bottom covered with circle of blotting paper. 49
Incubation at Various Temperatures
One phase of this study involved the determination of the effect
of various temperature levels on length of incubation period and on number of eggs hatched. The lowest temperature used was 40 F, the
highest 90 F . Also studied were each of the 10 degree increments be
tween the low and the high. Small cups made from 3% dram plastic vials
were used to hold the eggs for these experiments. The vials were sawed
off ^ inch from the bottom with a hacksaw and the rough edge sanded
smooth with a power sander. Circles of blotting paper cut with a power
cork borer fitted with a number 12 cutter were inserted in the bottom of
each cup. The small snap-on lids which came with the vials would no
longer snap on, because the beaded lip had been removed in the cutting
down process, but they were laid loosely on the cups as a cover (Fi^3().
When preparing an incubation experiment, the small blotting pads
in the cups were first wet with 4 drops of a 0.2% formalin solution
from an eye dropper bottle. This provided a moist surface to receive
the eggs and prevented mold from growing without interfering with
hatch of the fly eggs. The cups were held in a rack constructed of
1 inch X 6 inch X 6 inch pine lumber in which several rows of k inch
deep holes had been bored with a 15/16 inch wood bit (Fig. 31). Eggs
were collected as described under "Egg Collection" above. Five rows
of 5 eggs each for a total of 25 were then laid out on the blotting pad
in each cup using a camel's hair brush. The blotting pads were
moistened with the 0.2% formalin solution daily, or more often at
higher temperatures, to prevent desiccation of the eggs. The eggs
were examined once or twice daily, depending upon the conditions of 50
Fig. 30.--Small incubation cup disassembled to show lid, circle of blotting paper and bottom section (cut-off vial).
Fig. 31.--Wooden rack holding cups used in incubation experiments at lethal temperatures. 51 the experiment, and the hatched eggs counted. Experiments at each temperature level were replicated six times.
Larval and Pupal Development at Various Temperatures
Another phase of the study involved the effect of a range of ten degree increments of temperature from 40 F to 90 F on the development of the larval and pupal stages of the flies. Four-ounce polystyrene cups like those described for use in collecting eggs were used in this series of experiments. The cups were filled to within ^ inch of the top with the two-day-old C.S.M.A. medium. Twenty-five newly hatched first instar larvae were placed on top of the medium in each cup. Each temperature treatment was replicated six times. The cups were covered with their polyethylene snap-on lids which had been fitted with 1% inch diameter circles of 100 mesh brass sieve cloth for ventilation (Fig. 32). The contents of the cups were examined daily under a dissecting microscope in order to determine the stage of development of the flies. Moisture was added when necessary to keep the C.S.M.A. medium suitable for fly development (approximately 60 to
707. moisture). When the flies began to pupate the covers were removed from the cups and inverted. The pupae were counted daily and deposited on the covers. The inverted covers with the pupae were then placed back on top of the cup and the entire unit set inside a 32 ounce poly styrene cup, with seive cloth-fitted cover as described undbr "Rearing^ above. This procedure prevented any adults which might emerge from escaping, yet allowed the pupae from a particular cup to be kept safely adjacent to the remainder of the larvae in the cup (Fig. 33). 52
Fig. 32.--Four ounce polystyrene cup filled with C.S.M.A. medium and used for rearing larvae at various temperatures.
Fig. 33.--Pupae placed on inverted cover of rearing cup inside 32 ounce cup. 53
When adult flies had emerged in numbers too large to count easily as they crawled about in the containers, they were anesthetized with car bon dioxide. Anesthetization seemed to have no ill effect, as the flies all revived within three to four minutes after the gas was turned off.
Temperature Cabinets
Temperature cabinets used for most of the work were of the large walk-in type. Temperatures were thermostatically controlled to the required temperatures - one degree F. Relative humidities in all cabinets were also automatically controlled to a level of approximately
50% - 5%. The temperature cabinet used for the experiments with incuba tion at lethal temperatures was a Percival Dual Purpose Environmental
Growth Lab, Model E-57; manufactured by Percival Refrigeration and
Manufacturing Company, Boone, Iowa.
Effect of Moisture on Development
Many general statements are made by various authorities regarding flies breeding in "wet" manure, "moist" feed, or "damp" lawn and garden trimmings. Hutchison (1914) showed that moisture content of manure influenced bahavior of fly larvae about to pupate. He found that 98 to
99% of the third stage house fly larvae could be made to leave manure if the latter was kept moist, but only 70% migrated when the manure was kept more or less dry. Copeman (1916) found 115 F lethal to all fly larvae regardless of moisture content, but the time required to accomplish death was much less with wet heat than with dry. Bruce
(1939), maintaining a constant temperature of 80 F, placed puparia
in jars of fine sand in which the water content ranged from 0 54 to 17%, Adults emerged from sand with a moisture content up to 14%, but rarely from anything higher. Fay (1939) found 120 F to be lethal to house fly maggots, but showed that they could develop at 110 to
116 F if the medium remained moist. Feldman-Muhsam (1944a) found that in winter conditions in manure piles were such that flies died because of excessive moisture. Feldman-Muhsam (1944b) found that cow manure dried very quickly forming a crust. Both larvae and pupae could be found beneath the surface. He observed that this drying of the surface of the manure pile obviously inhibited the prepupal migration and he believed it to delay and possibly prevent development in many cases.
The foregoing workers have all attempted to show how moisture affects flies. None, with the exception of Bruce (1939) gave specific moisture levels, but merely used the terms "moist” or "moisture" or
"dry". None gave a specific moisture level which is optimum for fly breeding or upper and lower limits of the moisture range in which flies breed. Actually such figures are very difficult to determine. Some of the water added to a fly rearing medium at the beginning of an experiment under ordinary room conditions evaporates during the course of the experiment, so that the per cent moisture content is less at the end of the test. Also as the flies, fungi, bacteria, and perhaps other microorganisms break the medium down and assimilate it, some water is used and some metabolic water is produced. The water content of the medium is thus in a constant state of change.
In this study an attempt was made to control the moisture levels
in a fly rearing medium within as narrow a range as possible by 55 preventing water loss through evaporation. This was done by confining the medium in closed containers which were supplied from above with air which had been bubbled through a reservoir of water. This saturated air supply, in addition to reducing moisture loss, provided oxygen for the flies in the medium and forced carbon dioxide, ammonia and other undesirable gases to the outside through an outlet tube. An apparatus was constructed to accomplish the above which allowed four moisture levels, each replicated six times, to be run simultaneously (Fig. 34).
The apparatus consisted of a series of 8 ounce polystyrene cups which held the fly rearing medium, and each of these were placed inside a 32 ounce polystyrene cup and capped. Each 32 ounce cup was connected to the water reservoir by means of plastic aquarium tubing and the flow of air regulated by an aquarium valve. Air was supplied by four electric vibrator-type aquarium pumps. Each 32 ounce cup had
\ inch of moist sand in the bottom to absorb excess moisture and prevent mature larvae, seeking a place to pupate, from drowning in condensed water. The covers of the 32 ounce cups were specially con structed to minimize the dropping of condensed water from the under sides of the lids or the ends of the aquarium tubing into the fly rearing medium. Eight-ounce, round-bottomed, polyethylene bowls were inverted and "welded” with a hot soldering iron to the center under sides of the regular snap-on lids of the 32 ounce cups. These
Inverted bowls formed an "umbrella" over each cup of fly medium, yet did not restrict the flow of air and the various gases to and from the medium. The outlet tube from each cup ended in a 9 dram polystyrene vial half filled with water and with a \ inch hole in the snap-on 56
Fig. 34.--View of one section of moisture control apparatus. The entire apparatus included another section which was a duplicate of the one above. 57 lid (Fig. 35). The ends of the outlet tubes were submerged slightly in the water so that the escaping air and gases caused bubbles. These bubbles served as indicators for regulating the flow so that all cups received approximately the same amount of air. Figure 36 shows a diagram of the layout of one section of the apparatus.
The procedure followed in carrying out a moisture experiment was to combine the required amount of dry C.S.M.A. mix with water by weight to make a medium of the desired moisture content. Mixing was done thoroughly by hand in a plastic dishpan and then each of the desired number of 8 ounce cups were filled about 2/3 full of the medium and capped immediately with the snap-on lids to prevent moisture loss.
When the mixing procedure was completed the cups were uncapped one at a time, placed in the moisture control apparatus, and the pumps turned on. They were placed in such a manner as to make a randomized block design and thus prevent the possibility of bias due to the chance that cups nearer the pumps might receive more air than others.
On the second day the cups were removed from the apparatus one at a time, the mold stirred in with a small spatula, 25 newly hatched first instar fly larvae placed in the medium, and the cups returned to the apparatus. Since the apparatus was kept in a rearing room at a constant temperature of 75 F, the flies pupated in about 8 to 10 days.
As soon as pupation was completed, the 32 ounce cups with 8 ounce cups inside were removed from the apparatus and capped with ventilated snap on lids (lids with hole covered with 100 mesh sieve cloth). The cups were then set aside until the adults emerged. When emergence was completed, the flies were counted and the numbers recorded as an 58
Fig. 35.— Drawing of a portion of the moisture control apparatus to show detail of construction.
Fig. 36.--Diagram of the layout of one section of the moisture control apparatus to show how cups were connected to pump. The valve and indicator vial at the end of the line served as a relief valve for excess air. 59
indication of the effect of the various moisture levels on development.
Prior to the time that the experiments were carried out with fly
larvae in the cups, the procedure was carried out several times in
order to determine the limits within which the moisture was controlled.
This was done by running a dry matter analysis on samples of the "dry”
C.S.M.A, medium to be used in the experiment, weighing each cup of medium before and after a ten-day run, and finally running a dry matter analysis on the medium at the end. The difference in per cent
dry matter at the beginning and at the end of the experiment showed
the amount of breakdown and loss as gases due to action of micro
organisms. From these figures a\.td the other known weights at the
beginning and end of the experiment, the range of moisture contents
could be computed. Table 1 shows some typical figures.
As shown in Table 1, the four moisture levels varied in the
amount of dry matter loss due to the action of microorganisms. The
approximate 827, level was obviously less favorable for microbial
activity and there was a minimum of breakdown of solid materials.
There was also less change in moisture content than at the other
levels. This was followed by a relatively greater amount of solid
material breakdown and moisture change at the approximate 28, 46, and
647. levels. Apparently microorganisms do very well at around the 64
to 74% moisture level, as the dry matter loss and moisture range were
greatest here. Although the moisture contents at the low, high and
intermediate levels varied (by a little over 97. at around the 64%
level of moisture), known ranges of moisture content were established
under which fly rearing could be attempted and effects evaluated. TABLE 1
CCMPUTATION OF MOISTURE CONTENT OF C.S.M.A, MEDIUM AT END OF TEN DAYS IN MOISTURE CONTROL APPARATUS
Selected Wet Wt. Wt. % Water Total Wt. Actual 5^ Wet Wt. Wt. Dry Wt. Actual io Moisture Replica Mix At Water in Dry Water at Moisture Mix at Matter Water Moisture Levels tions Start Added C.S.M.AÎ Start At Start End at End* at End at End
. 1 4 3 . 1 1 8.62 9 . 3 8 11.86 2 7 . 5 1 3 9 . 8 5 2 6 . 5 9 1 3 . 2 6 3 3 . 2 7 tr 1 3 . 1 2 20^ 2 4 2 . 9 4 8 . 5 9 1 1 . 8 1 2 7 . 5 0 39.64 2 6 . 5 2 3 3 . 1 0 ir 3 42.88 8 . 5 8 1 1 . 8 0 2 7 . 5 2 3 9 . 3 3 2 6 . 6 2 1 2 . 7 1 3 2 . 3 2 II 4 4 2 . 9 1 8 . 5 8 1 1 . 8 0 2 7 . 5 0 39.48 26.40 1 3 . 0 8 3 3 . 1 3
I! 1 5 0 . 3 3 2 0 . 1 3 2 2 . 9 6 4 5 . 6 2 40.53 1 8 . 7 0 2 1 . 8 3 5 3 . 8 6 II 4 0 9 6 2 48.34 1 9 . 3 4 2 2 . 0 6 45.64 3 8 . 9 5 1 7 . 9 8 2 0 . 9 7 53.84 II 3 4 5 . 9 9 18.40 2 0 . 9 9 4 5 . 6 4 3 7 . 3 9 1 7 . 1 5 20.24 5 4 . 1 3 II 4 9 . 6 2 1 8 . 5 2 4 1 9 . 8 5 22.64 4 5 . 6 3 4 0 . 3 9 2 1 . 8 7 5 4 . 1 5
It 1 77.88 46.73 4 9 . 6 5 6 3 . 7 5 6 5 . 0 3 1 7 . 3 3 4 7 . 7 0 7 3 . 3 5 II 4 5 . 8 2 6 0 5 6 2 7 6 . 3 7 48.69 6 3 . 7 6 6 3 . 3 5 16.64 46.71 7 3 . 7 3 II 3 7 4 . 5 1 4 4 . 7 1 4 7 . 5 1 6 3 .7 6 - 61.46 1 6 . 2 3 4 5 . 2 3 7 3 . 5 9 II 4 7 4 . 7 0 44.82 4 7 . 6 2 6 3 . 7 5 6 1 . 8 3 1 6 . 8 1 4 5 . 0 2 7 2 . 8 1
II 1 1 8 1 . 7 3 1 4 5 . 3 8 148.79 81.87 177.23 3 0 . 0 3 1 4 7 . 2 0 8 3 . 0 6 n 8 1 . 8 8 1 7 2 . 2 1 8 2 . 9 8 8 0 5É 2 1 7 7 . 0 7 141.66 1 4 4 . 9 8 2 9 . 3 1 1 4 2 . 9 0 II 3 1 7 4 . 0 2 1 3 9 . 2 2 142.48 8 1 . 8 8 170.14 2 8 . 6 9 141.45 8 3 . 1 4 II 1 7 5 . 2 4 1 4 3 . 2 6 4 i 4 o .19 143.48 81.88 1 7 1 . 5 3 2 8 . 2 7 8 3 . 5 2
^Determined by standard laboratory dry matter analysis technique. CHAPTER V
THE EFFECT OF TEMPERATURE
Results and Discussion
Temperature is a very important physical factor influencing the development of Fannia canicularis and F. femoralis. The two species respond similarly to certain temperature levels, while at others, particularly the extremes, they respond quite differently. Temperature can be a definite limiting factor in the development of the two species both at the lower and higher levels. Responses to temperature in the laboratory correlate very well with the behavior of populations in the field and may help explain the more northern distribution of
F. canicularis, while F. femoralis ranges farther to the south.
Effect of Temperature on Incubation
Fannia canicularis
Eggs of this species were incubated at constant temperatures of
40, 50, 60, 70, 80 and 90 F . All eggs failed to hatch at 90 F. At
80 F the eggs hatched in 1% to 2^ days with an average incubation period of 2 days (Table 2). The incubation period appeared to be slightly longer at 70 F ranging from 1% to 3 days with an average of
2.1 days, but this difference was not statistically significant. The incubation period was significantly longer at 60 F with a range of 3 to 4% days and an average of 3.3 days. At 50 F the incubation period
61 TABLE 2
EFFECT OF CONSTANT TEMPERATURE ON INCUBATION PERIOD OF FANNIA CANICULARIS EGGS
Temperatures Degrees Fahrenheit
1+0 50 6 0 70 8 0 90
Range Day Range Day Range Day Range Day Range Day Range Day Replicate in Major in Major in Major in Major in Major in Major Number Days Hatch Days Hatch Days Hatch Days Hatch Days Hatch Days Hatch
1 17 - 2 1 1 6 . 0 1+ - l6 1 0 . 0 3 - H 4.25 1&- 3 2 . 0 2 - 2 | 2 . 0 Do Not Hatch
2 24.0 1 0 2 . 2 5 2 — — — —• 7-27 1 0 . 0 3 3.0 H 2 . 0 — 3 17 - 2 0 2 0 . 0 8 - 9 8 . 0 3 - 4 3.0 2 - 2 i 2 . 0 2 2 . 0 —
k 6 . 0 1 0 1 2 1 0 . 0 2 . 2 5 2 — — — — 6 - 3 - 4 3.25 H 2 . 0 — 5 Did Not Hatch 1 0 - 1 3 1 1 . 0 3 - 4 3.0 2 2 . 0 2 2 . 0 —
2 . 2 5 2 — — — 6 1+ l+.O 9è- 14 1 0 . 0 3 - 4 3 . 2 5 li- 2 . 0 —
Total Ave. Total Ave. Total Ave. Total Ave. Total Ave. Total- Ave.
1 1 +.1+ 4 - l6 2 . 1 2 . 0 — 4 - 27 9.7 3 - H 3 . 3 li- 3 1 &- 2 i — — * 63 was increased over the 60 F level by almost three times, the range being from 4 to 14 days, with the major hatch occurring at 9.7 days.
Amazingly, some eggs of F. canicularis hatched at a constant temper ature of 40 F, however, the hatch was very erratic. The incubation period at this temperature ranged from 4 to 27 days with the majority of the eggs hatching in an average of 14.4 days. This could not be proven to be significantly different from the hatch at 50 F however, because of the erratic differences between replications, particularly at 40 F.
It should be brought out that a few eggs of this species normally hatch almost immediately after they are laid, indicating that the females hold some eggs in the oviduct for considerable periods of time during which the embryos are developing. The fact that an occasional egg does hatch "prematurely" no doubt alters somewhat the true picture of the effect of temperature on incubation.
The effect of the various temperature levels on the numbers of eggs hatched is shown in Table 3. Since each of the six replicates at a given temperature level contained 25 eggs, the total possible hatch was 150. As indicated in the table, 124 eggs or 82.7% hatched at
80 F. There was no significant difference at the 70 F level where 123 or 82.0% of the eggs hatched. Although a slight reduction in hatch is
indicated at 60 F where 121 or 80.7% of the eggs hatched, and at 50 F where only 111 or 74.0% of the total eggs hatched, these figures were not shown to be significantly different from those at the higher temperatures. At 40 F where the hatch was very erratic, the number hatched was reduced to 31 eggs or only 20.7% of the possible total of 64
150, a definitely significant difference from the four higher temperatures.
Since no eggs of _F. canicularis hatched at a constant temperature of 90 F it was concluded that this temperature is lethal to the embryo.
In order to determine the point during incubation at this temperature at which death of the embryos occurred, several experiments were con ducted. Groups of 25 eggs were exposed for 3, 6, 9, 12, 15, 18 and 21 hours at 90 F and at the end of each of the various periods returned to
80 F for final incubation. The foregoing regime produced no definite pattern of hatching behavior, so the temperature was increased to
95 F. It was found that this temperature was highly lethal to F. canicularis eggs. The temperature was then lowered to 93 F and groups of 25 eggs exposed for one hour increments from one to ten hours and then returned to 80 F. Each treatment was replicated six times. This series of treatments produced a rather consistently increasing pattern of embryo mortality as shown in Table 4.
Statistical analysis of the figures for the control and the one hour exposure show no significant difference, indicating that exposure to 93 F for one hour has little or no effect. From this point on, how ever, each additional hour of exposure results in a reduction in hatch which approaches almost zero at seven hours.
Fannia femoral is
Eggs of F. femoralis were incubated at temperature increments of
10 degrees through 90 F just as were the F. canicularis eggs (Table 5).
Eggs of F, femoralis show a significantly different response to the higher and also to the lower temperatures than do F. canicularis. 65
TABLE 3
EFFECT OF TEMPERATURE ΠHATCH OF EGGS OF FANNIA CANICULARIS
Temperatures Degrees Fahrenheit Replicate Number 40 50 60 70 80 90
1 11 22 22 22 22 Do Not Hatch 2 11 15 18 15 18 — — — 3 2 21 22 22 25 — 4 1 20 23 21 20 " " — 5 0 16 18 24 20 » “ ™ 6 6 17 18 19 19 ...
Total Eggs 31 111 121 123 124 - - -
TABLE 4
EFFECT OF INCUBATION AT 93 FAHRENHEIT FOR VARIOUS PERIODS OF TIME ON HATCHING OF FANNIA CANICULARIS EGGS
Hours of Exposure Replicate Number 1 2 3 4 5 6 7 8 9 10 Control
1 22 8 5 2 2 1 0 0 1 0 23 2 21 15 0 2 2 I 0 0 1 0 22 3 19 15 8 6 6 2 1 0 1 0 23 4 22 16 4 2 2 1 0 0 1 0 17 5 19 12 6 1 4 0 0 0 1 0 22 6 17 12 3 0 0 0 1 0 0 0 23
Average 20.0 13.0 4.3 2 .1 3.0 0 .8 0.3 0.0 0.8 0.0 21.7 66
F. femoralis eggs hatch quite well at 90 F in from 3/4 to 3 days with
the majority of eggs hatching at about 1.1 days. There is no
significant increase in incubation period when the temperature is
dropped to 80 F. At this temperature F. femoralis eggs hatched in
from 1 to 3 days with the majority hatching at about 1.2 days. At
70 F the incubation period almost doubles with a range of 2 to 5 days
and the major hatch occurring at 2.0 days. It is interesting to note
that at 60 F the incubation periods of the two species of Fannia
studied are almost identical. The range of time over which the F.
femoralis eggs hatched was slightly longer than that of F. canicularis,
but the majority hatched in 3.3 days as did the eggs of the latter
species. A most interesting phenomenon is the fact that F. femoralis
eggs do not hatch at 50 F or below as did those of F. canicularis.
The effect of the various temperature levels on the numbers of
F . femoralis eggs hatched is shown in Table 6. Statistical analysis of
the figures for 70, 80 and 90 F shows no significant difference in egg
hatch. At 60 F there was a significant reduction in hatch, thus
indicating a deleterious effect of low temperatures on this species.
While F. femoralis eggs hatch quite well at 90 F, it was found
that a constant temperature of 95 F reduced the hatch to only 50 to
6 0%, and 100 F reduced it to zero. A series of six experiments
were then conducted where eggs were exposed to 100 F for various periods
of time from one to ten hours and then returned to 80 F for completion
of the incubation period. The results are shown in Table 7. It can
be seen that even a two hour exposure at 100 F reduces the hatch TABLE 5
EFFECT OF CONSTAlilT TEMPERATURE ON INCUBATION PERIOD OF FANNIA FEMORALIS EGGS
Temperatures Degrees Fahrenheit
40 50 6 0 TO 8 0 90
Range Day Range Day Range Day Range Day Range Day Range Day Replicate in Major in Major in Major in Major in Major in Mbjor Number Days Hatch Days Hatch Days Hatch Days Hatch Days Hatch Days Hatch
1 h - T 4.0 2 - 2 + 2 . 0 l%-2 ^ 1 . 2 5 1 - 3 1 . 2 5
2 . 0 1 . 0 1 1 . 1 2 3 - ^ 3 . 0 2 - 5 1 - Iv 3 Do Not Hatch 3 - 4 U.O 2 2 , 0 1 - 2 1 . 0 1 - 2 1 . 0 k 3 - 5 3-0 2 - 3 2 . 0 3 1 . 2 5 1 1 . 0
1 - 2 5 3 3 . 0 2 2 . 0 1 . 2 5 1 - Iv 1 . 0
2 2 . 0 ii-i?4 1 . 2 5 3 / 4-2 1 . 2 5 6 2 i - 3 i 3 . 0
Total Ave. Total Ave. Total Ave. Total Ave.
3 4 - 3 1.1 T 3 . 3 2 - 5 2.0 1 - 3 1.2 /
o\ - 4 68
TABLE 6
EFFECT OF TEMPERATURE ON HATCH OF EGGS OF FANNIA FEMORALIS
Temperatures Degrees Fahrenheit Replicate Number 40 50 60 70 80 90
1 Do Not Do l)ot 20 23 21 21 2 Hatch Hatch 20 16 21 19 3 — — « " — — 14 24 19 20 4 — — — ™ — — 23 25 23 25 5 — — — — — 20 23 22 25 6 — — - - “ - 18 22 19 23
Total Eggs 115 133 125 133
TABLE 7
EFFECT OF INCUBATION AT 100 FAHRENHEITFOR VARIOUS PERIODS OF TIME m HATCHING OF FANNIA FEMORALIS EGGS
Hours of Exposure Replicate Number 1 2 3 4 5 6 7 8 9 10 Control
1 14 8 8 5 1 0 0 0 0 0 23 2 24 7 1 0 1 0 0 0 0 0 25 3 21 13 5 0 0 0 0 0 0 0 24 4 22 9 4 0 2 0 0 0 0 0 22 5 14 13 1 1 0 0 0 0 0 0 22 6 18 9 0 0 0 0 0 0 0 0 24
Average 18 .8 9.8 3.1 l.CI 0 .66 0 0 0 0 0 23.3 69
considerably and that it drops off rapidly up to six hours where no
eggs hatch at all.
Effect of Temperature on Larval and Pupal Development
As is true with the eggs, the larvae and pupae of Fannia also
respond very differently to differences in temperature level. At
temperatures of 60 to 70 F the two species respond very similarly.
Above and below this range they respond quite differently. Certain
temperatures are lethal to the larvae of both species. Evidence of a
survival mechanism is brought into play in F. canicularis at 70 F and
below.
Fannia canicularis
Since the eggs of F, canicularis do not hatch at 90 F, some were
incubated at 80 F and first instar larvae obtained. These larvae were
placed in C.S.M.A. medium at 90 F to see if they would develop.
Within four days it was obvious that this temperature affected these
larvae adversely. They failed to develop normally, taking even longer
to reach the third instar than larvae held at 70 F. By the fourteenth
day they were still rather imnature third instar larvae and by the
fifteenth day some had died. A few larvae had pupated by the twenty-
first day, but none of them emerged as adults. This experiment was
replicated six times. From these results it may be concluded that
90 F is not only lethal to the eggs, but also to the larvae and/or
pupae of F, canicularis.
Fannia canicularis larvae and pupae developed very normally at 70
80 F making a total life cycle from egg to adult an average of 18.4 days as shown in Table 8.
At 70 F the total life cycle in the majority of the flies was increased by about five days to an average of 23.9 days. At this temperature level a very interesting phenomenon occurred, however. Up to one-fifth of the flies were delayed in their development at the late third instar. These larvae had moved into the dryer material at the top of the rearing cups and had become quiescent as if preparing to pupate. Instead of completing pupation on schedule, they remained in the quiescent larval state for fourteen to as long as twenty-eight days. At the end of this period pupation took place and finally emergence as adults.
This phenomenon is, no doubt, a survival mechanism of some sort and might be described as a diapause. It also occurs at temperatures below 70 F and is probably a means of helping this species to survive periods of winter cold. At any rate this mechanism is not at all like the one described by Roubaud (1927) for F. canicularis, where three normal generations were followed by a fourth which went into an obligate diapause (see Chapter II).
At 60 F the development of F. canicularis was slowed down to the point that the total life cycle was almost doubled, requiring an average of 40.8 days. When placed at 50 F, the cold hardiness of this species was again demonstrated when only slightly less than 507. of the flies came through in an average of 101 days.
As shown in Table 9 the total number of flies emerging was
influenced by temperature. As already stated 90 F is lethal to TABLE 8
EFFECT OF TEMEEEATÜEE ON DURATION OF INSTARS AND TEE TOTAL LIFE CYCLE OF FANNIA CANICULARIS
Days
Uo 50 6 0 TO 8 0 90
2 7 1 6 2.1 lè-2è Egg* U - Ih.k U - 9.7 3 - H 3 . 3 li- 3 2.0 Do Not Hatch First Does Not
Instar Develop 5 - 8 6 . 5 2 - 3 2 . 7 1 1.0 1 1.0 —-— Second
Instar 2 - 4 - — 1 ? 8.0 3 . 0 2 2.0 1 1.0 Third
Instar — — 2 5 ~ 2 9 . 3 12 - 1 3 1 2 . 7 6 - 8 6.8 5 - 6 5 . 7
Pupa 4 5 1 8 2 0 — - 4 7 . 5 - 1 9 . 1 12 12.0 8 - 1 0 8 . 7 -— Average Total
1 8 . 4 Days — - 101.0 40.8 2 3 - 9 ---
♦From Table 2. 72
TABLE 9
EFFECT OF TEMPERATURE ON THE NUMBER OF FLIES DEVELOPING TO THE ADULT STAGE IN FANNIA CANICULARIS
No. Flies Emerged Replicate Number 40 50 60 70 80 90
1 Do Not 17 20 22 17 Do Not 2 _ Develop 13 16 22 16 Develop 3 10 21 22 22 — — — 4 8 21 23 19 " — 5 13 20 21 19 — — 6 11 20 23 17 W M
Totals 72 118 133 110 73
F. canicularis. This species develops somewhat normally at 80 F,
However, 70 F produced the highest number of flies of all the tempera
ture levels tested.
As the temperature level was dropped to 60 F, the total number
of flies emerging was also decreased significantly. A drop in
temperature to 50 F produced only 72 flies or slightly less than 50%
of the total possible 150 flies.
As shown earlier, a few eggs of F . canicularis hatched at 40 F,
but no larvae developed at this temperature.
Fannia femoralis
The total life cycle of F. femoralis at 90 F is an average of
13.3 days as shown in Table 10. When the temperature is decreased to
80 F the life cycle increases by about one day to 14.7 days. This
species appears to be more markedly affected by low temperatures than
F. canicularis as a reduction in temperature to 70 F increases the
total life cycle by about one week to 21 days. At 60 F the life cycle
is greatly lengthened to 37.6 days or about the same as F. canicularis
at this temperature. No diapause was exhibited at the lower tempera
tures in this species as occurred in F . canicularis.
The effect of temperature on the total flies emerging is shown
in Table 11. While F. femoralis will develop at 90 F, only about one
fifth of the 150 flies survived to emerge. This would indicate that
90 F is on the borderline of being lethal for F. femoralig;
The largest number of flies of this species emerged at 80 F,
indicating that this temperature may be close to optimum for F. TABLE 10
EFFECT OF TEMEERATÜEE ON DURATION OF INSTARS AND THE TOTAL LIFE CYCLE OF FANNIA FEMORALIS
Days
li-O 5 0 6 0 TO 8 0 9 0
1 1 . 1 1 . 1 Egg* Do Not Hatch Do Not Hatch 2?- 7 3 . 3 2 - 5 2 . 0 2^A ^ 3 First
Instar --- 1 - 3 2 . 3 1 1.0 1 1.0 1 1.0 Second
Instar --- 3 “ ^ 3 . 3 1 - 2 1 . 3 1 2 1.1 1 1.0 Third Instar ——— 11—lA 12.2_ 5-6 5.7 h k.o k 4.0
Pupa --- 1 6 - 1 8 1 6 . 5 10-12 11.0 7 8 7.5 6-7 6 . 1 Average Total
Days ——- 3 7 . 6 21.0 1 U . 7 1 3 - 3
*Prom Table 5 75
TABLE 11
EFFECT OF TEMPERATURE ON THE NUMBER OF FLIES DEVELOPING TO THE ADULT STAGE IN FANNIA FEMORALIS
No. Flies Emerged Replicate Number 40 50 60 70 80 90
1 Do Not Do Not 18 19 19 0 2 Develop Develop 17 16 22 1 3 m w i m - — — 23 25 25 14 4 W M — “ — 20 19 22 2 5 — — — — — — 19 22 24 6
6 ------18 22 15 7
Totals 115 123 127 30 76 femoralis. Experience in rearing stock colonies of this species would also indicate that 80 F is more favorable than lower temperatures as the flies were more active and reproduced more prolifically. CHAPTER VI
THE EFFECT OF MOISTURE
Results and Discussion
Moisture can be demonstrated to be an important limiting factor in the development of both Fannia canicularis and F. femoralis. As with temperature, the two species respond similarly to certain moisture levels, while at others, particularly the extremes, they respond quite differently. Development is limited by both high and low moisture levels in both species of flies.
Effect of Moisture on Incubation
While no attempt was made to specifically study the effect of moisture on the eggs of Fannia. it was found that moisture was necessary to keep the eggs in good condition during the incubation period. As indicated in Chapter IV, it was necessary to moisten the eggs daily to prevent desiccation, especially at the higher tempera tures. Failure to add water to the blotting pad beneath eggs held at
80 F resulted in the eggs turning brown and failing to hatch. Failure to_add moisture when incubating eggs at 90 F resulted in an obvious drying and shrivelling.
Effect of Moisture on Development of Larvae and Pupae
By means of the moisture control apparatus described in Chapter
IV attempts were made to rear the larvae of Fannia canicularis and
77 78
Fannia femoralis in C.S.M.A. medium maintained within the following approximate moisture ranges:
28% to 33% 37% to 43% 46% to 54% 64% to 74% 82% to 83%
The results are shown in Tables 12 and 13. Neither species developed at the low moisture range of 28 to 33%. Thirty-nine out of a possible
150 F. canicularis reached the adult stage at the moisture range of
37 to 43%, while a significantly lower number, only 7, of F. femoralis reached the adult stage at this moisture range. At the 46 to 54% moisture level 80 F. canicularis emerged while only 64 F. femoralis emerged, but the difference could not be shown to be statistically significant. A significant difference did occur at the 82 to 83% moisture level where 91 F. femoralis developed to the adult stage, compared to only 13 F. canicularis.
These data show then that F. canicularis is capable of developing in media with moisture contents in the range of 37 to 43%, although not well. F. femoralis does very poorly at this moisture level. At the high end of the scale where the moisture content runs 82 to 83%,
F. canicularis does very poorly while F. femoral is develops fairly well. The optimum moisture level for both species appears to fall between 64 and 74%.
It would appear from these results that F. femoralis is capable of living under wetter conditions than _F_. canicularis. This capability may be enhanced by the habit of the larvae of the former species of 79
TABLE 12
EFFECT OF MOISTURE CONTENT OF THE REARING MEDIUM ON LARVAL AND PUPAL DEVELOPMENT OF FANNIA CANICULARIS
Approximate Moisture Levels Replicate Numbers 287.-337. 377.-437. 467.-547. 647.-747. 827.-837.
1 0 5 7 17 4 2 0 5 16 21 8 3 0 7 13 23 0 4 0 6 16 22 0 5 0 6 13 17 1 6 0 10 15 20 0
Total Flies Emerged 0 39 80 120 13
TABLE 13
EFFECT OF MOISTURE CONTENT OF THE: REARING MEDIUM m LARVAL AND PUPAL DEVELOPMENT OF FANNIA FEMORALIS
Approximate Moisture Levels Replicate Numbers 287.-337. 377.-437. 467.-547. 647.-747. 827.-837.
1 0 3 9 19 11 2 0 I 15 23 12 3 0 0 11 23 13 4 0 1 10 25 20 5 0 2 10 19 21 6 0 0 9 22 14
Total Flies Emerged 0 7 64 131 91 80
crawling up and pupating on the sides of the container in which the
rearing medium is held. This habit is not exhibited by mature larvae
of F. canicularis and may result in the drowning of pupae when the moisture content of the medium is high. CHAPTER VII
CONCLUSIONS
Fannia canicularis is a cosmopolitan species. It is found wherever the common house fly, Musea domestica, occurs as well as many areas not frequented by the latter. Its range extends into the cold regions of the far north such as Alaska, Greenland, Norway and the Arctic islands. This species is known as the little or ’’lesser” house fly. It breeds in a variety of organic waste materials including poultry droppings. It readily moves into residential areas where it has a nuisance rating second only to that of the common house fly.
Fannia femoralis breeds in organic waste materials similar to those frequented by Fannia canicularis. It is particularly attracted to poultry droppings and often reproduces in great numbers there. The range of F. femoralis is much more restricted than that of F. canicularis, being confined in general to the warmer southern regions.
The adults of this species do not move out from the breeding sites to the extent that F. canicularis does and therefore do not constitute
such a serious nuisance problem.
Although the genus Fannia has been placed in the family Muscidae
by taxonomists, some of its morphological aspects as well as its habits
are quite different from the common house fly. Musea domestica, and
some other members of the family. The eggs of Fannia are very
distinctive in that they have a pair of thin laterodorsal wings or
81 82 flanges of chorion which extend almost the entire length of the egg.
The dorsal surface is divided by a longitudinal ridge and is further characterized by having a raised reticulated pattern. The ventral surface is characterized by a longitudinal ribbing running the entire length of the egg.
Fannia eggs adhere very tightly to the material upon which they are laid. This characteristic is due to a water soluble adhesive on the surface of the egg. The adhesive of Fannia canicularis is more readily water soluble than that of F. femoralis as less time is required to loosen the eggs from the substrate after water is applied.
The larvae of Fannia are unlike the larvae of other members of the Muscidae. Rather than being the typical smooth, almost cylindrical, tapering, white rauscoid type, Fannia larvae are somewhat depressed dorsoventrally, have a spiny appearance and are brown when fully developed.
The posterior spiracles of Fannia larvae are located dorso- medially on the caudal segment and are on short stalks. The number and arrangement of spiracular openings are useful in distinguishing the various larval instars. Contrary to the statements of Chillcott
(1960) and other authors, the number of spiracular openings is not equal to the number of instars. The first instar actually has two openings, the second and third instars four each.
The shape of the pharyngeal skeleton of Fannia larvae is very characteristic in each instar. This character is almost essential in distinguishing the first and second larval instars without the aid of
a compound microscope. 83
Puparia of Fannia are quite different from those of other
Muscidae in that they are not of the characteristic smooth, "water melon” shape. Pupation in Fannia takes place inside the old skin of the third instar larva. As the larva prepares to pupate, it stops feeding and becomes quiet. It shortens slightly in length and the dorsum becomes somewhat arched. Gradually the mouth hooks and remainder of the cephalic region are withdrawn leaving the pro- thoracic area thin and flattened with a slight depression or groove down the dorsomedial line. While the foregoing is taking place the old cuticle is hardening and darkening to become the puparium. The dorsal and lateral processes of the larva remain giving the puparium its characteristic spiny appearance.
Larvae and pupae of F. canicularis and F. femoralis can usually be distinguished by the difference in length of the dorsal and lateral processes» those of F. canicularis being longer. Frequently, however, the processes are obscured by the clinging of foreign material to them. In such cases two features which may be used in distinguishing the species are found on the ventral surface. F, canicularis larvae and pupae have a median transverse ridge or carina on each of the abdominal stemites while F. femoralis lacks this ridge. Also in F. canicularis the anal opening forms a narrow "V" with the apex of the
"V" pointed anteriorly. F. femoralis. on the other hand, has a more oval or wide "V"-shaped anal opening with the apex pointed anteriorly,
F annia canicularis and F. femoralis adults are both characterized by moving in a hovering, "dancing" flight. F. canicularis. particu larly, is frequently seen in breezeways, open buildings, rooms in 84 dwellings or under the shade of trees, flying about in groups or swarms
and never seeming to light on surfaces. Unlike the common house fly,
Musea domestica. the Fannia species rarely alight on persons or on
human food.
Fannia can be very readily reared on a modified C.S.M.A, medium
containing the ingredients in the following proportions:
300 grams dry C.S.M.A. medium 26 grams dry powdered brewers yeast 600 milliliters water
Temperature is a very important physical factor influencing the
development of Fannia canicularis and F. femoralis. The two species
respond similarly to certain temperature levels, while at others,
particularly the extremes, they respond quite differently. Temperature
can be a definite limiting factor in the development of the two species
both at the lower and higher levels.
At the higher temperatures F . femoralis has a significantly
shorter life cycle than F . canicularis. The time from egg to adult
in the former being approximately 14 days at 80 F, while F. canicularis
takes approximately 18 days at the same temperature.
At 90 F F. femoralis develops from egg to adult in about 13 days
while F. canicularis will not complete its life cycle at this
temperature. Eggs of the latter species fail to hatch at 90 F. One
hundred degrees F, on the other hand, is lethal to the eggs of F.
femoralis which do not hatch at constant exposure to this temperature.
As the temperature is lowered from the optimum, F. femoralis
exhibits a much more marked effect than does F . canicularis. If the
temperature is held at 70 F the life cycle of F . femoralis increases 85 in length by about 7 days to a total of 21, while that of F. canicularis increases by only about 5 days to a total of 23. At
60 F the life cycles of the two species are about the same, at 38 to
40 days.
At a constant temperature of 50 F the life cycle of F. canicularis increases to about 3 months while F. femoralis fails to develop at all. While a few eggs of F. canicularis will hatch at 40 F, the larvae of this species fail to develop at this temperature.
At 70 F and below, F. canicularis exhibits a survival mechanism which might be described as diapause. Up to one-fifth of the larvae become quiescent in the late third larval instar and pupation is delayed from 14 to 28 days. Following this quiescent period pupation takes place and adults emerge. This mechanism apparently provides a means of survival of this species during periods of adversity.
It can be concluded from these results that Fannia canicularis is adapted for living under much cooler conditions than is F. femoralis. The geographical distribution of the two species as shown in Figs. 1 and 2 bears this out.
Moisture is also shown to be an important factor in influencing the development of Fannia canicularis and F . femoralis. As with temperature, the two species respond similarly at certain levels, while at others they respond quite differently.
Neither species of Fannia completed its life cycle when the moisture content of the rearing medium was maintained between 28 and
33%, indicating that this level is too dry for both. At a range of
37 to 43%, 7 F . femoralis out of a possible 150 flies came through 86 indicating that this level is on the borderline of being too dry for this species. On the other hand 39 F. canicularis came through at this moisture level indicating that this species is a little better adapted for dryness than F, femoralis.
The optimum moisture level for both species lies between 64 and
747» where 131 F. femoralis and 120 F . canicularis completed develop ment .
When the moisture content of the rearing medium was maintained between 82 and 837» a significant difference in response of the two species again occurred. At this level the emergence of 91 F. femoralis compared to only 13 F. canicularis shows that the former is more capable of developing at high moisture levels than the latter. CHAPTER VIII
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