Evolution, 35(5), 1981, pp.

872-881

GENETIC VARIABILITY OF THE FACE FLY, MUSCA AUTUMNALIS

DE GEER, IN RELATION TO A POPULATION BOTTLENECK

EDWIN H. BRYANT
Department of Biology, University of Houston, Houston, Texas 77004
AND

HENK VAN DIJK AND WILKE VAN DELDEN

Biological Center, Department of Genetics, University of Groningen, Haren (Gn), The Netherlands

Received July 30, 1980. Revised November 11, 1980

Various theories have been promoted to rather unconvincing counter-juggling of

explain the apparent high levels of elec- mutation rate and population size, such
trophoretic variation present in most or- that the product of the two remained near-
ganisms (see reviews by Lewontin, 1974; ly constant. The traditional formulation
Nei, 1975; Nevo, 1978). Among these the of the neutral mutation model assumes
random drift of neutral or nearly neutral constant population size, however, and
mutations developed by Kimura and his Nei et al., (1975) and Chakraborty and
colleagues (e.g., Kimura, 1968; Kimura Nei (1977) have pointed out that the size
and Maruyama, 1971; Kimura and Ohta, of natural populations can be drastically
1971a, 1971b, 1971c) convincingly ac- reduced during evolution (see also Carson,
counts for observed patterns of variation 1971, 1973), resulting in the loss of alleles,
within and among populations of many particularly those already in low frequen-
organisms (e.g., Fuerst et al., 1977; Chak- cy. They show that the recovery time after
raborty et al., 1978). In focusing upon the such loss is of the order of the reciprocal
pattern of variation, however, less atten- of the mutation rate and thus historical
tion has been given to the fact that many bottlenecks can be used to explain lower
organisms with large populations, partic- variation in many species than predicted
ularly insects, have considerably less vari- by the equilibrium of neutral mutations,
ation than predicted by the neutral model even though contemporary populations of
and that the level of variation is remark- these species may be quite large.
ably consistent among widely divergent Population bottlenecks have been in-
organisms. Thus, putatively inbred or- voked in this manner to account for low
ganisms as Avena and Mus (Selander et variation in a number of organisms, in-
al., 1969; Marshall and Allard, 1970; cluding cave-dwelling fish (Avise and Se-
Clegg and Allard, 1972), organisms living lander, 1972), elephant seals (Bonnell and
in highly uniform environments (Gooch Selander, 1974), Anolis lizards (Webster
and Schlopf, 1972; Selander et al., 1974; et al., 1972), man (Haigh and Maynard
Selander, 1976), parthenogenetic organ- Smith, 1972) and the Bogota, Colombia
isms (Suomalainen and Saura, 1973; Par- population of Drosophila pseudoobscura
ker et al., 1977) and organisms as diverse (Prakash et al., 1969; Prakash, 1972,
as the horseshoe crab and man (Selander 1977; Lewontin, 1974; but see Singh et al.,
et al., 1970; Harris and Hopkinson, 1972) 1976). In these cases, however, a probable
all exhibit similar levels of variability, in bottleneck was only invoked after low
spite of varied evolutionary histories, con- levels of variation were observed in these
temporary ecologies, and population populations and we are only aware of the
structures. two studies by Taylor and Gorman (1975)
To account for this uniformity in vari- on Anolis grahami and by Schwaegerle
ation by invoking the neutral mutation and Schaal (1979) on the pitcher plant,
model (at equilibrium) would require a Sarracenia purpurea, that specifically ad-
872
 FACE FLY GENETIC VARIATION 873

dress changes in variability in relation to man blood, and fresh cattle dung to collect
known bottlenecks. eggs. The F 1 offspring and the field col-
The face fly, Musca autumnalis De lected parents were placed on dry ice and
Geer, is widespread in the Palearctic re- transported to the Genetics Institute in
gion but did not occur in the New World Haren, The Netherlands, where they were
until a presumed accidental introduction stored at -90 C until used. European flies
of this species into Nova Scotia, where it were similarly placed in population cages
was discovered in 1952 (Vokeroth, 1953). and transported to the Genetics Institute
It expanded during the next few years into for low temperature storage. During the
the United States and during the ensuing process of egg collection and transporta-
10 years spread across the country through tion, some flies died and thus the final
the northern states, appearing in Califor- sample size for the electrophoretic com-
nia about 1967 (USDA, 1967; Depner, parisons was less than the number of flies
1969). The face fly is a serious pest of cat- collected from the field and which could
tle and its presence in the New World have contributed genomes to the F 1 gen-
would not have gone unnoticed for long. eration.
Since these flies do not frequent indoor Electrophoretic procedures. -Horizon-
areas where they might be artificially tal electrophoresis was carried out in 6%
transported, and they were discovered ini- polyacrylamide gels at 0-5 C and 15 VI
tially at a single locality in the New em. Flies were homogenized in 40 pJ of
World, it has been concluded that the water containing a trace of EDTA.
species was successfully introduced only Enough homogenate could be obtained
once from Europe and that little time from a single fly to fill slots on eight dif-
elapsed before its discovery (Sabrosky, ferent gels, so that eight different enzymes
1961). There has been insufficient time for could be scored on each fly. The total
any substantial (mutational) replenish- number of enzymes examined was 14 out
ment of alleles lost during introduction of an initial number of 17: three enzymes
and hence a comparison of variation in were omitted because of difficulties in ob-
current face fly populations from the taining clear bands or in the interpretation
United States and Europe should provide of banding pattern (e.g., esterases). Indi-
an assessment of any genetic shifts that vidual females and their offpsring were
may have occurred during colonization. run to confirm inheritance patterns of
Any such shifts can then be compared electromorphs and thus we will refer to
with the theoretical predictions of bottle- these electromorphs as allozymes (or al-
neck effects given by Nei et al., (1975) and leles).
Chakraborty and Nei (1977). Four different electrophoresis buffers
were used: A) Phosphate-citrate per 1:9.3
MATERIALS AND METHODS g N~HP04 (anh.), citric acid added to
Face fiies.-Adult face flies were col- prescribed pH from 4.0 to 5.5. B) Tris-
lected in open fields near cattle at four lo- maleate: per 1:4.9 g maleic anhydride,
calities in the United States (Plattsburgh, Tris added to pH 6.0. C) Tris-citrate: per
New York; Olathe, Kansas; Ovid, Idaho; 1:9.45 g citric acid, Tris added to pH 7.0.
and Denver, Colorado) and at four local- D) Sodium-borate: per 1:18.55 g boric
ities in Europe (Heerde, The Netherlands; acid, NaOH to pH 8.5.
Ansbach, Germany; Dijon, France; and The electrophoretic buffers and staining
Ehrlenbach, Switzerland). These localities techniques used for each enzyme system
were selected to represent diverse popu- are listed in Table 1. The following stains
lations within the range of the species were used where indicated: Alcohol de-
while occurring within the same general hydrogenase: stain composed of 25 ml of
climatic regimes in both continents. 0.1 M carbonate-bicarbonate buffer pH
American face flies were placed in pop- 10.0, 10 mg NAD+, 10 mg MTT, 1 ml
ulation cages with sugar water, whole hu- ethanol and 2 mg PMS. Catalase: gel con-
874 EDWIN H. BRYANT ET AL.

TABLE 1. Enzyme abbreviation, electrophoretic buffer, and staining technique for the 14 enzymes utilized
in this study.

Staining
Enzyme Symbol Buffer technique

Alcohol dehydrogenase ADH A, pH 4.0 see text

a-Glycerophosphate dehydrogenase a-GPDH A, pH 5.0 1
Malate dehydrogenase MDH B 1
Malic enzyme ME B or C 1
Glucose-6-phosphate dehydrogenase G-6-PD C 2
Glyceraldehyde-3-phosphate dehydrogenase GA-3-PD B 1
Catalase CAT D see text
Aldehyde dehydrogenase ALDOX D see text
6-Phosphogluconate dehydrogenase 6-PGD A, pH 5.5 2
Isocitrate dehydrogenase IDH C 1
Superoxide-dismutase SOD A, pH 4.5 see text
Phosphoglucomutase PGM B 1
Acid Phosphatase ACPH D 1
Amylase AMY D see text
1 = Ayala et al. (1972).
2 = Bijlsma and van Delden (1977).

taining 0.2% starch was shaken with 50 were polymorphic over all samples. Sod
ml of a 1% solution of H 2 0 2 in buffer C. was only polymorphic in Europe and
When oxygen bubbles appeared, the per- Acph was only polymorphic in America
oxide solution was removed, the gel rinsed and thus the percent polymorphic loci per
in water, and a solution of 1% KJ and continent was 36% (5 of 14). The average
0.1 % sodium thiosulfate added. Aldehyde heterozygosities for these polymorphic loci
dehydrogenase: 25 ml of a 0.05 M Tris- in Table 3 were 0.123 ± 0.058, 0.088 ±
HCI pH 8.5 with 5 mg NAD+, 5 mg 0.049 and 0.078 ± 0.042 for the Europe-
MTT, 1 mg PMS and after 10 min, 0.1 an, American and American F 1 flies, re-
ml acetaldehyde. Superoxide-dismutase spectively. Averaged over all loci, these
(=Tetrazolium oxidase): 25 ml of 0.1 M heterozygosities were 0.053 ± 0.029,
carbonate-bicarbonate pH 10.0 with 10 0.038 ± 0.027 and 0.030 ± 0.019, and the
mg NAD+, 10 mg MTT and 2 mg PMS. average numbers of alleles per locus were
Shaken for 10 min and exposed to light. 1.55 ± 0.89,1.46 ± 0.83 and 1.33 ± 0.57,
Amylase: gel with 0.2% starch shaken respectively.
with 50 ml buffer C for 30 min and iodine- The average heterozygosity as well as
potassium iodide solution added. the average number of alleles per locus
were lower in the American flies, as might
RESULTS be expected if a founding bottleneck had
While it is sometimes useful to consider occurred. The standard errors of these es-
only those alleles above a certain frequen- timates in the two continents were broadly
cy, say 5% or 1%, low frequency alleles overlapping and thus, there was no evi-
are precisely the ones likely to be affected dence for a significant reduction in vari-
during a bottleneck. In Table 2, we thus ability during colonization of the New
give the frequencies of all alleles encoun- World.
tered in the study, for the parental flies The observed variation in the two con-
from both continents and for the F 1 flies tinents only represents a sample from the
from America. Alleles were designated by supposedly greater universe of alleles
their mobilities relative to the most com- present in these entire populations. To the
mon allele (=1.0). extent the observed number of alleles in
Of the 14 loci examined, six, or 43%, the two continents, for example, deviates
 FACE FLY GENETIC VARIATION 875

TABLE 2. Allozyme frequencies per population for the 6 polymorphic enzyme systems and for the parental
and offspring (F1J samples. H = Holland, G = Germany, F = France, S = Switzerland, N = New York,
K = Kansas, I = Idaho, C = Colorado, n = number of flies scored on enzyme and population.

European American American Ft

Enzyme-allele H G F S N K C N K c
6-PGD 0.6 .01 .05 .04 .02 .03 .04 .00
0.8 .26 .24 .25 .32 .20 .20 .13 .19 .26 .13 .11 .05
1.0 .71 .72 .66 .60 .70 .76 .83 .81 .68 .80 .86 .95
1.2 .01 .04 .04 .04 .09 .04 .02 .08 .02
(n) (44) (23) (55) (25) (33) (46) (15) (13) (62) (20) (114) (40)
IDH .87 .03 .02 .01 .04 .04
1.00 1.00 .97 .99 1.00 .98 .99 1.00 1.00 1.00 1.00 .96 .96
1.13 .01
(n) (41) (18) (53) (22) (32) (64) (13) (12) (76) (16) (100) (44)
SOD 0.5 .01 .01
1.0 .97 .99 .99 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00
1.5 .02 .01
(n) (61) (38) (78) (22) (32) (67) (13) (12) (100) (98) (161) (68)
PGM 0.83 .03 .03
1.00 .84 .97 .92 .85 .97 .98 1.00 1.00
1.17 .13 .03 .08 .12 .03 .02
(n) (38) (20) (45) (20) (31) (41) (13) (13)
ACPH 1.0 1.00 1.00 1.00 1.00 .99 1.00 1.00 1.00 .96 .97 .98 .92
1.4 .01 .04
1.8 .03 .02 ;08
(n) (28) (10) (35) (20) (43) (40) (13) (13) (61) (16) (107) (44)
AMY 0.88 .03 .06 .08 .01
0.92 .04 .04 .11 .03 .03 .04 .04 .05 .02
0.96 .01 .05 .02 .08 .03 .13 .03 .03
1.00 .94 .96 .92 .86 .83 .87 .76 .92 ' .94 1.00 .94 .97
1.04 .01 .03 .02 .01 .04 .01
(n) (43) (24) (53) (22) (30) (49) (12) (12) (60) (16) (110) (30)

from the potential number of alleles avail- locus (a) in a sample of size n would be
able in these two populations, the loss in (Ewens, 1972; Templeton, 1980):
variability during colonization may be in- () ()
accurately measured by these sample sta- E(a) = -() +(}+1
--
tistics. The availability of alleles in the ()
two continents may be explored through + ... + ,
the sampling theory of neutral mutations () + 2n - 1
developed by Ewens (1972). where () = 4N e JL and N; and JL are the
Since the zygotic frequencies of field effective population size and the mutation
collected flies did not deviate from Hardy- rate, respectively. Theoretically, as sam-
Weinberg expectations, and the average ples are combined in any order, one
genetic identity (Nei, 1973) among popu- should observe an increase in the mean
lations within continents was extremely number of alleles per locus.
high (I > 0.99), the geographic popula- The averages of six possible sampling
tions within continents may be considered sequences starting from each of the four
as replicate samples from a single pan- European populations are shown in Fig-
mictic population. Under these assump- ure 1. An estimate of () = 0.144 was ob-
tions, the expected number of alleles per tained from Table 3 of Ewens (1972), us-
 876 EDWIN R. BRYANT ET AL.

TABLE 3. Observed proportion of heterozygous flies 1.9

per enzyme system and population. ii = average
number of flies scored per enzyme. Locality abbre-
viations as in Table 2.
1.8
(f)
European flies ::::J
Enzyme U
system H G F s fl 0
...J
<, 1.7
6-PGD .34 .29 .36 .48 .368 (f)

IDR .00 .06 .02 .00 .020 w

...J
w
SOD .06 .03 .01 .00 .025 ...J
...J
PGM .26 .05 .16 .30 .192 «
ACPR .00 .00 .00 .00 .000 1.6
AMY .12 .08 .15 .18 .132 c:i
z
H .130 .085 .117 .160 .123
z
it 42 22 53 22 34.7 «
w
::;
American flies

N K C fl

6-PGD .33 .24 .27 .23 .268 40 60 80 100 120 140

IDR .03 .02 .00 .00 .012 NO. FLIES
SOD .00 .00 .00 .00 .000 FIG. 1. Increase in number of alleles per locus
PGM .06 .05 .00 .00 .028 with number of flies examined, averaged over the six
ACPR .02 .00 .00 .00 .005 possible sequences starting with each of the four geo-
AMY .27 .24 .17 .17 .212 graphic samples from Europe. Dashed line repre-
H .118 .092 .073 .067 .088 sents theoretical expectation from Ewens' formula
it 33 51 13 13 27.5 with 8 = 0.144, estimated from the maximal sample
size (n = 139) and the observed number of alleles
American F 1 flies per locus of 1. 86. Locality abbreviations as in Ta-
N K C fl ble 2.

6-PGD .31 .40 .20 .05 .240

IDR .00 .00 .03 .07 .025 be random samples from a single pan-
SOD .00 .00 .00 .00 .000 mictic population in the United States.
PGM
ACPR .08 .06 .03 .16 .082 With this in mind, we conjecture that
AMY .05 .00 .06 .07 .045 nearly all of the alleles available to Amer-
H .088 .092 .064 .070 .078 ican flies would be present in the earlier
it 72 20 118 44 63.5 established New York population and
that alleles were subsequently lost during
the expansion of the face fly into the west-
ern localities. Furthermore, the rapid
ing the mean number of alleles per locus growth of the face fly populations in the
in the entire sample (n = 139; ii = 1.86), United States would almost certainly en-
and a curve of expected number of alleles sure that the number of alleles in the entire
with increasing sample size was generated population would be far below the num-
with this estimate in Ewens' formula. ber expected in an equilibrium population
There seems to be a reasonable agreement of the same size (Nei et al., 1975). Ana-
among all four sampling sequences and lyzing an increasing number of flies from
the expected number of alleles from ran- such a non-equilibrium population would
dom sampling. When the American flies not result in the continued addition of rare
are treated in the same manner in Figure alleles. Given these considerations, we feel
2, however, there is wide disparity among that the observed 1.71 alleles per locus in
the four sampling sequences. Any se- the New York flies may be near the avail-
quence starting with New York results in able number of alleles present in the entire
no further increase in the number of al- American population. Thus, there may
leles. These populations do not appear to have been a real loss in variability during
 FACE FLY GENETIC VARIATION 877

colonization of the New World that was

undetected by our sample estimates of av-
erage variability in the two continents.
1.7 N
7--
It would be possible to estimate the
magnitude of this loss in variability if we
en 1.6
knew the effective size of the face fly pop- :::>
u
ulation in Europe. It would not be unrea- 0
...J -&=.123"
/

I
sonable to assume an effective size of at <, I
en 1.5 I
least 106_107 for this common species (Ay- w
...J
/
I
ala, 1972, e.g., estimated the population W
...J /
...J /
size for Drosophila willis toni to be near <[
I
109) . Utilizing our estimate of () = 0.144 0
1.4 /
/

for the European population, the mean z

number of alleles per locus in a population z
of size 106_107 would be 3. 1-3.4. In spite <[
w 1.3
::;:
of the moderate reduction in variability
observed in the sample of American flies
compared to the European sample, the c
I. 2 '-----,------,r-----r------,----,--------,
American flies may contain only about o 20 40 60 80 100 120
50% (1. 7/3.4) of the alleles in the ancestral NO. FLIES
European population.
FIG. 2. Increase in number of alleles per locus
DISCUSSION with number of flies examined, averaged over the six
possible sequences starting with each of the four geo-
We feel that two alternative interpre- graphic samples from America. Note that no increase
tations of our data are possible. On the in allelic variation occurred with sequences starting
one hand, the lack of a significant reduc- with New York. Dashed line represents theoretical
tion in mean variability between conti- expectation from Ewens's formula with (J = 0.123
estimated from the maximal sample size (n = 110)
nents suggests that there was little or no and observed number of alleles per locus of 1.71.
loss of allelic variation during colonization Locality abbreviations as in Table 2.
of the New World. This may be consistent
with the predictions of Nei et al. (1975),
that only severe bottlenecks « 10) will population of the face fly may thus contain
have any serious effect upon variability. only about one-half of the number of al-
Since we do not know the actual bottle- leles that would be present in an equilib-
neck size during introduction of the face rium population of the same size.
fly to the New World, it could have indeed This latter interpretation rests upon
been large enough to contain most of the finding no new alleles (in wild flies) be-
variability present in the ancestral popu- yond the New York population. We do
lation. On the other hand, the pattern of not know, however, whether additional
allelic variation among our American alleles would have been found in this pop-
samples in relation to sample size (Fig. 2) ulation had a larger sample of flies been
suggests that these populations are not in taken. We did obtain 140 offspring from
genetic equilibrium and that our sample New York females and found no new al-
may have contained nearly all of the al- leles, but we do not know how many ad-
leles present in the population, rather than ditional genomes appeared in these F 1
a sample from a larger universe of alleles. flies. The number of new genomes could
Taking an apparently conservative esti- have been as many as twice the number
mate of the effective size of these flies in of females collected and thus all of the F 1
Europe of 106-107 , it is possible that a re- flies could have carried previously unsam-
duction in allelic variation of as much as pled genomes, since more than 50 females
50% may have occurred during introduc- were collected from this locality. On the
tion to the New World. The American other hand, the egg-batch size for the face
878 EDWIN H. BRYANT ET AL.

fly is near 20 (Wang, 1964; Killough and Michigan area, we obtain an estimate of
McClellan, 1965) and assuming a larval () = 0.10 for these populations (from their
survival rate of at most 50% (Valiela, Table 1). A single individual would then
1969; Burton and Turner, 1970), it is also be expected to contain 1.09 alleles per lo-
possible that as few as 14 females contrib- cus on the average and a heterozygosity
uted to the F 1 sample. Using Table 1 of of 50% of the ancestral population. These
Ewens (1972), the New York flies should observed values were 1.1 and 55%, re-
have yielded 1.2 (n = 33 + 28) to 2.4 spectively, and are consistent with the
(n = 33 + 72) new alleles, whereas none sampling theory of neutral mutations.
were found. The lack of additional alleles Taylor and Gorman (1975) studied pop-
in these F 1 flies supports our assumption ulations of Anolis lizards introduced into
that we may have sampled nearly all of Bermuda with a sample of 71 individuals
the alleles present in the population. from Jamaica. For Jamaica we estimate
Our analysis, of course, depends upon () == 0.16, from which a sample of 71 indi-
the infinite allele model of neutral muta- viduals should contain 1.85 alleles per lo-
tions (Kimura and Crow, 1964). There is cus. The observed mean number of alleles
little concern using the infinite allele mod- per locus of 1.5 is more consistent with a
el versus the step-wise model for electro- bottleneck size of 10 rather than 71. The
phoretic variation (Ohta and Kimura, smaller variation in the derived popula-
1973) as the two give nearly identical re- tions could stem from selection against
sults for bottlenecks (Chakraborty and some alleles in the new environment,
Nei, 1977). There may be some problem which is not predicted from the neutral
if selection were operating. The small model. It is also possible that few of these
population size during a bottleneck should released individuals survived to reproduce
ensure that nearly all alleles behave as if or that the founders were related geneti-
selectively neutral, but the estimation of cally, perhaps due to subdivision in the
allelic variation in the entire population parental population (Templeton, 1980).
may not be faithful. However, much of Either of these possibilities would reduce
the available electrophoretic data is con- the effective size of the inoculum and in
sistent with the neutral mutation model fact, may be more parsimonious expla-
(e.g., Fuerst et al., 1977; Chakrabortyet nations of reduced variability in the de-
al., 1978) and our own data seem to fit rived population than selection.
neutral expectation up to the limit of our The interpretation of the effect of the
sample (Fig. 1). Since we have no way of introduction on variability in the face fly
assessing unknown selective effects in any notwithstanding, it is still of some interest
event, the neutral mutation model pro- that variability in this species was consid-
vides a useful, and perhaps, the only an- erably less than typical of other insects,
alytic approach to the problem of bottle- particularly Drosophila, where heterozy-
necks. gosity averages near 15% (Selander and
It is of interest to know whether the Kaufman, 1973; Lewontin, 1974; Powell,
predictions of the neutral model hold for 1975; Nevo, 1978). Our low estimate of
other bottleneck data, particularly when fI = 0.053 for the European flies could be
bottleneck size is known. Unfortunately, due to our omission of esterases, How-
few studies have addressed such changes ever, Prakash (1973)and Barker and Mul-
in variability during bottlenecks. Schwae- ley (1976) found low heterozygosity in
gerle and Schaal (1979) investigated the Drosophila buskii (fI = 0.044) and D.
electrophoretic variation in a population buzzatii (fI = 0.065), respectively, which
of the pitcher plant, Sarracenia purpurea, they attributed to the narrow nutritional
founded by a single individual. The spe- niches of these-species compared to other
cific origin of the single pitcher plant was Drosophila. In comparison to the conge-
unknown, but assuming it came from the neric housefly, whose larvae occupy a
 FACE FLY GENETIC VARIATION 879

number of diverse sites, the larval niche information about the distribution of the
of the face fly seems to be largely restricted face fly in Holland.
to cattle dung. In lieu of a putative rela-
tionship between the food niche and ge- LITERATURE CITED
netic variability (see also Nevo, 1978), a AVISE, J C., AND R K. SELANDER. 1972. Evolu-
comparison of enzyme variation between tionary genetics of cave-dwelling fishes of the ge-
the housefly and the face fly would be in- nus Astyanax. Evolution 26:1-19.
AYALA, F. J. 1972. Darwinian versus non-Darwin-
teresting, but we are unaware of any sur- ian evolution in natural populations of Drosoph-
vey of enzyme variation in the housefly. ila. Proc. 6th Berkeley Symp. Math. Stat. Prob,
5:211-236.
SUMMARY AYALA, F. J, J R POWELL, M. L. TRACEY, C. A.
MOURAO, AND S. PEREZ-SALAS. 1972. Enzyme
Enzyme variation of the face fly, Musca variability in the Drosophila willistoni group.
autumnalis De Geer, from Europe and the IV. Genetic variation in natural populations of
United States, was compared over 14 loci Drosophila willistoni. Genetics 70:113-139.
BARKER, J S. F., AND J C. MULLEY. 1976. Iso-
to assess any loss in variability that may zyme variation in natural populations of Dro-
have occurred when this species invaded sophila buzzatii. Evolution 30:213-233.
the New World. The average heterozy- BIJLSMA, R, AND W. VAN DELDEN. 1977. Poly-
gosity and the average number of alleles morphism at the G-6-Pd and the 6-Pgd loci in
Drosophila melanogaster. I. Evidence for selec-
per locus in the American flies were less, tion in experimental populations. Genet. Res.
but not significantly so, than in the ances- 30:221-236.
tral European flies. This suggested that BONNELL, M. L., AND R K. SELANDER. 1974.
the colonization episode had little or no Elephant seals: genetic variation and near ex-
effect upon variability in American pop- tinction. Science 184:908-910.
BURTON, R P., AND E. C. TURNER. 1970. Mor-
ulations. On the other hand, the earlier tality in field populations of face fly larvae and
established population of the face fly in pupae. J Econ. Entomol. 63:1592-1594.
New York contained nearly all of the al- CARSON, H. L. 1971. Speciation and the founder
leles present in the other populations sam- principle. Stadler Genet. Symp, 3:51-70.
- - . 1973. Reorganization of the gene pool dur-
pled. This suggested that the American ing speciation, p. 214-280. In N. W. Morton
population as a whole was not in genetic (ed.), Genetic Structure of Populations. Univ.
equilibrium. It would not therefore con- Hawaii Press, Honolulu.
tain the same number of alleles expected CHAKRABORTY, R, P. A. FUERST, AND M. NEI.
in an equilibrium population of the same 1978. Statistical studies on protein polymor-
phism in natural populations. n. Gene differen-
size. Utilizing Ewens (1972) sampling the- tiation between populations. Genetics 88:367-
ory for neutral mutations, we estimated 390.
that as many as 50% of the alleles avail- CHAKRABORTY, R, AND M. NEI. 1977. Bottleneck
able in the European population could effects on average heterozygosity and genetic dis-
tance with the step-wise mutation model. Evo-
have been lost during introduction, in lution 31:347-356.
spite of the similarity in sample estimates CLEGG, M. T., AND R W. ALLARD. 1972. Patterns
of variation between continents. of genetic differentiation in the slender wild oat
Avena bcrbata. Proc. Nat. Acad. Sci. USA
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ACKNOWLEDGMENTS DEPNER, K. R 1969. Distribution of the face fly,
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WEBSTER, T. P., R. K. SELANDER, AND S. Y. Corresponding Editor: J. R. Powell

ANNOUNCEMENT

COUNCIL FOR PHILOSOPHICAL STUDIES

The Council for Philosophical Studies is pleased to announce that it has re-
ceived funding from The National Endowment for the Humanities to conduct a
Summer Institute in the Philosophy of Biology at Cornell University, June 21 to
July 16, 1982. The Summer Institute is for professors of philosophy who are
interested in developing courses in the philosophy of biology. Teachers of biology
with suitable interests also are encouraged to apply.
The Summer Institute will combine illustrative lectures summarizing problems
and theories appropriate to this subject matter with workshops designed to ex-
amine and develop a body of curricular materials. Resident and guest lecturers
will discuss recent biological explanation, the conceptual structure of evolutionary
theory, the foundations of taxonomy, language in man and other primates,
and the neurological basis of mind.
Marjorie Grene (Davis) is the Resident Director of the Summer Institute, and
Richard Burian (Drexel) is the Assistant Resident Director. A partial list of the
visiting lecturers and consultants includes Richard Boyd (Philosophy-Cornell),
Lindley Darden (History and Philosophy of Science-Maryland), Stephen Jay
Gould (Biology, Geology and History of Science-Harvard), David Hull (Philoso-
phy-Wisconsin at Milwaukee), Emil Menzel (Primate Ethology-Stony Brook),
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Philosophy-Chicago).
Detailed inquiries may be sent to Professor Marjorie Grene, % Dept. of
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