Predation by the Introduced Phoretic Mite~

Macrocheles peregrinus (Acari: Macrochelidae)~ on the

Buffalo Fly~ Haematobia irritans exigua
(Diptera: Muscidae)~ in Australia 1

J. P. ROTH," A. MACQUEEN," AND D. E. BAY'

Environ. Entomol. 17(3): 603-607 (1988)

ABSTRACT The effectiveness of Macrocheles peregrinus (Krantz) as a predator of buffalo
fly, Haematobia irritans exigua (De Meijere), was examined. In laboratory tests, mites were
confined with eggs or larvae of buffalo fly to assess the effect of prey location and predator I
prey ratio on fly survival, to determine the main life stage attacked, and to determine the
prey preference of the mite in the presence of other fly species. In field tests, impact of the
mite on buffalo fly survival was measured in natural field pats enclosed in screen cages.
Buffalo fly mortality was highest when eggs of this species were placed on top of dung pats
rather than beneath them. Maximum mortality obtained was 83%; there was no increase in
mortality at predator Iprey ratios above one mite per two fly eggs. The mite caused highest
mortality when exposed to hatching eggs and young larvae of buffalo fly, but it showed a
strong preference for eggs of other dipteran species that have softer egg chorions. Mites
acting alone caused an average of 33% suppression of buffalo fly breeding in field pats.
Macrocheles peregrinus was judged to be a relatively ineffective predator of buffalo fly.

KEY WORDS Arachnida, phoretic predacious mite, Haematobia irritans exigua, buffalo fly

IN 1975, THE CSIHO Division of Entomology began 1983, Doube et a!. 1986). Subsequent to releases of
a study of predacious, phoretic mites associated M. peregrinus at Rockhampton, Queensland in
with native and established dung beetles in Aus- December 1980 and again in February 1981 near
tralia. The purpose of these investigations was to the Adelaide River, Northern Territory, Wallace
enhance biological control of the bush fly, Musca & Holm (1983) found that this mite was established
vetustissima (Walker), and the buffalo fly, Hae- and widely distributed in central Queensland and
matoma irritans exigua (De Meijere) (Wallace et the northeastern portion of the Northern Territory.
a!. 1979). The control of these two pests, along with The mites were primarily associated with the in-
pasture improvement, was the primary goal of the troduced dung beetle Euoniticellus intermedius
Australian dung beetle project (Waterhouse 1974). (Reiche), but they were also recovered from several
Wallace et al. (1979) found that Macrocheles gla- other introduced scarabs and one native scarab
ber (Muller), a mite abundant on native dung bee- species.
tles in southeastern Australia, is an effective pred- Our report describes studies designed to deter-
ator of the bush fly. Neither M. glaber nor any mine the effectiveness of M. peregrinus as a pred-
similar mite was found to be present within the ator of the buffalo fly in central Queensland.
range of the buffalo fly in Queensland and the
Northern Territory.
Krantz (1981) described Macrocheles peregri- Materials and Methods
nus (Krantz) from collections made in South Af- A laboratory colony of M. peregrinus was es-
rica, Namibia, and Kenya. This mite was selected tablished at Rockhampton in 1984 from local field
for introduction to Australia because of its wide collections. Laboratory colonies of the mites were
geographic distribution, its relative abundance in maintained on fresh cattle dung shaped into 200-g
Africa, and its range of phoretic hosts that included pats and placed on 25 cm of clean sifted sand in
most of the African dung beetles that have been a plastic container (30 by 20 by 12 cm). The sides
established in northern Australia (Wallace & Holm of the container were coated with fluon (ICI, Syd-
ney, Australia) to prevent mites from escaping. Col-
onies were held at 26°C and a 14:10 (L:D) photo-
I This artic!p rpports the resultsof research only. Mention of a period. Mites were supplied with bush fly eggs from
proprit'lary product dol'Snot constitute an endorsement or a rec-
onlllU'oJation for its use by USDA. a laboratory colony four times a week. Fresh dung
'Veterinary Toxicologyand EntomologyResearchLaboratory, pats were added to the colonies twice a week. The
CSDA~ARS,P.O. Drawer GE, College Station, Tex. 77841. new dung pats were placed with their edges touch-
3 Divisionof Entomology,CSIHO,Box5545, RockhamptonMail
ing the older pats to facilitate translocation by the
Centrt', Quepnsland 4702, Australia.
• Department of Entomology, Texas A&M University,College mites. Old desiccated dung pats were removed as
Station, Tex. 779·tQ. needed. All the mites used in experiments were
604 ENVIRONMENTAL
adult females; male M. peregrinus have a com-
paratively short life span, and the sex ratio heavily
favors the females. Mites to be used in tests were
removed from the colony and held without feeding
overnight on dung that had been frozen to destroy
any food source.
H. irritans exigua normally oviposits beneath
cow pats immediately after defecation. Oviposition
occurs not only on the dung but on soil and ground
litter that are in contact with the dung. Natural
oviposition was difficult to obtain for laboratory
tests; therefore, dung pats were seeded with known
numbers of Haematobia eggs using a standard
technique that simulated natural oviposition. This
was done by forming 150 g of fresh bovine dung
into cylindrical pats and supporting each pat on a
bamboo grid placed on 5 cm of dry sand in a I-liter
container. The grids were constructed from bam-
boo skewers (2 mm diameter by 8 cm long) glued
together by epoxy cement. Each grid consisted of
a lower row of four parallel bamboo sections spaced
1.5 cm apart, with a similar upper row glued at a
slight angle. Egg seeding was accomplished by im-
printing a grid on the surface of each test pat and
placing a known number of eggs on the dung at
approximately evenly distributed intervals along
the lines of contact with the grid. The dung pat
and grid were then inverted and placed egg-side
down on the sand in the test containers. The top
inner sides of each test container were coated with
fluon to prevent mites from escaping.
Buffalo fly eggs were obtained by netting adult
flies from cattle and inducing oviposition by CO2
anesthesia as described by Macqueen et al. (1986).
Prey Location. A test was designed to determine
the effect of prey location on the ability of M.
peregrinus to prey on buffalo fly eggs. In this test,
half the pats were prepared as described for the
standard technique with 30 buffalo fly eggs on the
underside, and the other half were prepared iden-
tically except that the buffalo fly eggs were placed
randomly on the top of the dung. The pats then
were infested with either 0, IS, or 45 M. peregri-
nus, respectively, per pat. After 6 d, Haematobia
pupae were collected by sifting the sand and ex-
amined the dung in each test container.
Predator/Prey Ratios. Dung pats and con-
tainers were prepared by the previously described
method for simulating natural oviposition. The
number of buffalo fly eggs per test container was
held constant at 30 eggs, and the number of mites
was varied to change predator jprey ratios. The
tests included a control with no mites and eight
mite treatments ranging from 3 to 60 mites per
container.
Buffalo Fly Stage Attacked. A test was con-
ducted to determine the period of buffalo fly sus-
ceptibility to predation by M. peregrinus. Test con-
tainers were prepared with dung as before, except
that buffalo fly eggs (30) were placed on top of the
dung pats. Treatments included a control (I) that
had buffalo fly eggs only; treatment 2, in which 20
ENTOMOLOGY Vol. 17, no. 3
mites were exposed only to the eggs; treatment 3,
in which 20 mites were exposed to hatching eggs
and first-instar larvae; and treatment 4, in which
20 mites were exposed to hatched larvae that had
been allowed 2 h to burrow into the dung. In treat-
ment 2, the mites and eggs were placed on the
dung at the same time. At approximately I h before
hatch (as indicated by the first appearance of larval
striations in the eggs), the eggs were carefully re-
moved and transferred to the surface of dung pats
in containers that had no mites. In treatment 3,
mites were not added to the containers until larval
striations appeared within the eggs; thus hatching
eggs and larvae were exposed. In treatment 4, mites
were added to the containers 2 h after hatch.
Prey Preference Tests. The first prey prefer-
ence test was conducted by exposing eggs of buffalo
fly and Orthellia lauta (Wiedemann) to adult fe-
male M. peregrinus on moistened filter paper in a
sealed Petri dish. After 24 h, the number of hatched
eggs and live larvae of each prey species was count-
ed. Treatments included a buffalo fly control with
20 eggs per Petri dish, an O. lauta control with 20
eggs per Petri dish, a treatment with 20 buffalo fly
eggs and four mites per Petri dish, and a treatment
with 20 eggs of each fly species and eight mites
per Petri dish. Each treatment in this test was rep-
licated five times.
Subsequent preference tests with buffalo fly and
bush fly were conducted using the previously de-
scribed technique for seeding buffalo fly eggs under
the dung pat. In treatments that contained bush
fly eggs, small indentations were made around the
lower edges of the dung pats near the point of
contact with the soil. Bush fly eggs from a labo-
ratory colony were placed in clusters in these in-
dentations to simulate natural oviposition sites. Two
test designs were used: the treatments in the first
test design consisted of a buffalo fly control with
50 eggs per pat, a bush fly control with 50 eggs per
pat, a combined control with 25 eggs of each species
per pat, 50 buffalo fly eggs with 20 mites per pat,
50 bush fly eggs with 20 mites per pat, and 25
buffalo fly eggs with 25 bush fly eggs and 20 mites
per pat. In this test, therefore, the total prey jpred-
ator ratio was constant. The treatments in the second
test design were similar, except that 40 eggs of each
prey species per pat were used. Thus, the combined
prey-control and the combined prey-mite treat-
ment had a total of 80 eggs per pat. Treatments
with mites had 15 mites per pat in this test.
With the exception of the preliminary test on
prey preference (which had five replications), all
the laboratory tests were replicated six times. Anal-
ysis of variance was conducted for a completely
random design, and treatment means were sepa-
rated using Duncan's multiple range test (Duncan
1955).
Survival in the Field. A field trial of the impact
of M. peregrinus on buffalo fly was conducted in
1985. This trial consisted of four tests which were
conducted on 13 February, 10 April, 1 May, and
J line 1988 ROTH ET AL.: M. peregrinus PREDATION ON BUFFALO FLY 605

TobIe 1. E£reets of egg loeotion on predotion of M. Table 2. Buffalo fly survival as a function of exposure
peregrinus on the buffalo fly to varying numbers of M. peregrinus in the laboratory

No. of % buffalo fly No. of No. of Mites per

Ep.p.location on pat % survival
M. peregrinus survivala mites prey egg
Top o 41. 7a 0 30 0 73.9 ± 7.4
Bottom o 42.2a 3 30 0.1 37.3 ± 8.7
Top 15 l.ld 6 30 0.2 27.3 ± 14.8
Bottom 15 30.ob 9 30 0.3 25.3 ± 12.6
Top 45 1.7d 12 30 0.4 24.0 ± 5.5
Bottom 45 18.3c 15 30 0.5 22.2 ± 13.9
30 30 1.0 12.2 ± 7.8
a M,'ans not followed by the same letter are significantly dif- 45 30 1.5 12.8 ± 8.9
[,'n'nt (P = 0.05 level, Duncan's [1955] multiple range test). 60 30 2.0 20.5 ± 12.4

15 May. The following procedure was used in each

test. Cattle dung pats were marked immediately
upon defecation. The pats were observed until ovi-
positing buffalo flies left the dung (ca. 3 min). After
oviposition occurred, the pats were covered with
mesh-covered metal cones to exclude all dung fau-
na (Macqueen & Beirne 1975). Ten pats were cov-
ered on each test date. Five of these pats were
seeded with 500 M. peregrinus from the laboratory
colony before being covered; these mites had been
held overnight without feeding. The remaining five
pats served as the controls. The test pats and a
shallow divot of the soil beneath the pats were
collected after 48 h in the pasture and held in an
insectary for 3 wk in cages fitted with emergence
traps to collect emergent Diptera. The data were
analyzed by ranking the number of buffalo flies
collected from each control pat in descending order
from highest to lowest and performing a paired t
test with similarly ranked mite treatments. Paired
t tests were used to separate treatment means for
each test and for combined treatment means for
the whole trial. The fauna-exclusion cones used in
the field trial were intended to keep out all dung
fauna except buffalo flies. However, several species
of Diptera (such as Orthellia [auta and Sepsidae
spp.) were able to colonize the pats during the time
when they were still exposed to buffalo fly ovipo-
sition. The Sepsidae spp. were collected so consis-
tently that we decided to analyze the data on sepsid
emergence as well as that on buffalo fly.
Results and Discussion
The results of the buffalo fly egg placement test
(Table 1) indicate that prey location on the dung
pat has a direct effect on its susceptibility to pre-
dation of M. peregrinus. Survival was less than 2%
in both treatments in which eggs on the surface of
the dung were exposed to the mites. In comparison,
survival in the mite treatments with bottom-seeded
eggs was 30% with 15 mites per pat and 18% with
45 mites per pat. Survival in the two control treat-
ments was about 42% for both top- and bottom-
seeded pats. The mites could be more adapted to
predation on species which do not oviposit under-
neath the dung pat. In fact, in our prey preference
tests, M. peregrinus showed a preference for O.
[auta and for the bush fly over buffalo fly. Neither
of these species habitually oviposit directly under
the dung.
The results of the predator /prey ratio tests (Ta-
ble 2) indicate that all the mite treatments caused
significantly more mortality than occurred in the
control, but even the highest mite/prey ratios did
not completely suppress buffalo fly development to
the degree that predation on top-seeded eggs did
in the egg placement test. This would seem to in-
dicate that when eggs were seeded underneath the
dung pat, some of the buffalo fly eggs were not
exposed to mite predation. Settling of the dung
could make some of the buffalo fly eggs and larvae
inaccessible to the mites. To what degree this might
occur with natural oviposition is not known.
Results of the stage preference test (Table 3)
indicated that the only treatment in which there
was a significant reduction in the number of buffalo
fly pupae collected was that in which mites were
added just prior to egg hatch. Exposure of the buf-
falo fly eggs to the mites during the pre hatch period
did not cause a significant reduction in buffalo fly
survival. Apparently there was some predation
when mites were added 2 h after hatch, but this
was not statistically significant. These results differ
from those of Doube et al. (1986), who reported
that predation on buffalo fly eggs did occur when
M. peregrinus was exposed to eggs seeded on top
of 50-ml dung pats. The high level of predation
that occurred on the top-seeded treatment in our
prey location test (Table 1) was similar to the level
of predation that occurred on bush fly eggs in the
prey preference test (see Table 5). It is unlikely
Table 3. Period of immature buffalo fly susceptibility
to predation by M. peregrinus
Exposure period
No. of buffalo
% survival
fly pupaea
None (control) 11.7a 39.8
16 h prehatch 1O.3a 34.4
2 h prehatch to pupae 4.8b 16.1
2 h posthatch to pupae 8.2a 27.2
a Means not followed by the same leller are significantly dif-
ferent (P = 0.05; Duncan's [1955] multiple range test).
606 ENVIRONMENTAL ENTOMOLOGY Vol. 17, no. 3

Table 4. Prey preference by M. peregrinus for eggs
and larvae of buffalo fly and O. lauta exposed to Ihe miles
on moistened filler paper
Buffalo fly
Treatments % %
egg live
hatch larvae
20 buffalo fly eggs (control) 89a 87.5a
20 O. Lauta eggs (control)
20 buffalo fly eggs + 4 mites 86a 46.7c
20 buffalo fly eggs + 20 O.
Lauta eggs + 8 mites 95a 68.5b
Means in columns not followed by the same letter are signifi-
cantly different (P = 0.05, Duncan's [1955] multiple range test).
that such high mortality could be accounted for by
predation on the buffalo fly larvae alone; therefore,
it is probable that under some circumstances the
mites will prey on prehatched buffalo fly eggs seed-
ed on the top of dung pats.
Results of the initial prey preference test are
presented in Table 4. The presence of the mites
had no significant effect on the mean percentage
of egg hatch of buffalo fly eggs. The mean per-
centage of the buffalo fly larvae alive at 24 h was
significantly lower than the control in the treatment
with buffalo fly, O. [auta, and mites. The presence
of O. [auta caused a significant reduction in the
degree to which buffalo fly larvae were preyed
upon in the combined-prey treatment. The mean
percentage of egg hatch of O. [auta eggs was sig-
nificantly reduced in the combined treatment com-
pared with the control. 0., [auta larvae did not
survive long on the moistened filter paper, and no
live larvae were found after 24 h in either the
control or the combined prey treatment.
The results of the two tests for preference of M.
peregrinus for bush fly or buffalo fly are presented
in Table 5. In the test in which the number of total
prey was constant, there was no significant reduc-
tion in buffalo fly survival by the mites, but this
may have been masked by low buffalo fly survival
in the 'control treatments. In contrast, bush fly sur-
O. Lauta vival was drastically reduced by the mites.
There was better survival in the buffalo fly con-
%
%
live
trol in the test in which the total number of prey
egg was variable. In this test the numbers of prey species
lar-
hatch
vae were high enough so that there was some compe-
tition between buffalo fly and bush fly larvae. Bush
81a 0 fly survival in the combined-fly control was signif-
icantly lower (60.4%) than in the bush fly control
(81.6%). Buffalo fly survival in the combined con-
53.6b 0
trol was not significantly different from survival in
the buffalo fly-only control. In the buffalo fly-mite
combination, mites caused a significant reduction
in buffalo fly survival (28.7%) compared with buf-
falo fly controls (64.6 and 57.9%) and the combined
prey and mite treatment (46.6%). As in the pre-
ceding test, bush fly survival in the presence of
mites (1.6 and 2.1%) was significantly lower than
bush fly or buffalo fly survival in any other treat-
ment. The significantly higher survival of buffalo
flies in the combined prey-plus-mite treatment
(46.6% compared with 28.7%) indicates that the
presence of a preferred alternative prey species can
reduce M. peregrinus predation on the buffalo fly.
The results of both tests demonstrate that M. per-
egrinus is a much more effective predator of bush
fly than buffalo fly. Although survival in the control
treatments in these laboratory tests was low, this is
a typical result when Haematobia spp. eggs are
artificially seeded in dung pats; variability is not
likely to be improved by additional replication.
Thomas & Morgan (1972) found similar levels of
mortality in control treatments with the horn fly,
Haematobia irritans (L.). In the present series of
tests, there were a total of 36 control dung pats,
and survival ranged from 31.4 to 73.9% with a
mean of 49 ± 16.4%.
The results of the field trial (Table 6) indicate
that there was considerable variation in the number
of buffalo fly eggs deposited within and between

Table 5. Prey preference by M. peregrinus for buffalo fly and bush fly in two laboratory tests that simulated natural
oviposition by these prey species

No. total prey constanta No. total prey variableb

Buffalo fly Bush fly Buffalo fly Bush fly
Treatments
% survival % survival % survival % survival
(no. eggs) (no. eggs) (no. eggs) (no. eggs)
I-Buffalo fly control 31.6a (50) 64.6b (40)
2-Bush fly control 7l.6a (50) 81.6a (40)
3-Buffalo fly + bush fly 32.0a (25) 66.0a (25) 57.9b (40) 60.4b (40)
4-Buffalo lIy + mites 25.3a (50) 28.7d (40)
5-Bush lIy + mites 4.3c (50) 1.6e (40)
6-Buffalo lIy + bush lIy + mites 25.3a (25) Oc (25) 46.6c (40) 2.1e (40)

Means are not compared between the two tests. Within the tests, means not followed by the same letter are significantly different
(P = 0.05. Duncan's [1955] multiple range test).
a No. prey (eggs). 50. This consisted of either 50 buffalo fly eggs and 50 bush lIy eggs. or 25 buffalo lIy eggs and 25 bush lIy eggs.
The mite treatments received 20 mites per pat.
b No. prey (eggs). 40 for both the buffalo lly and the bush lly, regardless of whether the treatment used one or both species. Thus,
combined species treatments had a total of 80 prey eggs, and the single species treatments had a total of 40 prey eggs. The mite
treatments received 15 mites per pat.
June 1988 ROTH ET AL.: M. peregrinus PREDATION ON BUFFALO FLY 607

Table 6. Effects of predation by M. peregrina. on H. irriran. exigua

Mean no. of adult flies"

!Jatl' H. irritans exigua Sepsidae spp.
Control Mitesb % suppression Control Mitesb % suppression

.13 Fl'b. .19H5 340.6 330.6 2.9 77.2 69.4 .10..1

10 Apr . .19H5 .188.6 77.8 58.7 22.1 48.6 78
t ~Iay .19H5 .126.2 59.4 52.9 74 .19.4 73.8
15 ~Iay .19H5 60.4 .10.4 82.8 .128.6 9.4 92.7*
TotalC 3,577 2,39.1 2,454 734
(h,'ralllO,'anc 179.8 119 33.2* 123 37 70.1*

" Vallit's arl' me'ans of fivt' dung pats.

h Eae'h dun~ pat s<,<,de'dwith 500 At. peregrinas from the laboratory colony .
•.Total and o\'<,rall m<,an are from combined data on 20 dung pats per treatment; *, significant (P = 0.05, paIred t test).

tests. The results of the 13 February test in partic-
ular were influenced by the emergence of 1,500
buffalo flies from one of the mite-treated pats.
However, the fact that the mites also performed
poorly against Sepsidae in the 13 February test,
could indicate that these mites were in poor con-
dition or that adverse climatic factors could have
been involved. Only in the 15 May test was suppres-
sion of the buffalo flies statistically significant; over-
all, mites caused a significant reduction of 33% (P =
0.05) in buffalo fly emergence. This result is prob-
ably not representative because of the lack of
suppression in the 13 February test. If the results
of just the 10 April, 1 May, and 15 May tests are
averaged, the resulting predation rate would be
64.8%. Doube et al. (1986) reported 66 and 54%
suppression, respectively, in two field tests.
Although the results of the 10 April and I May
tests indicate substantial predation, mites signifi-
cantly reduced sepsid emergence only in the IS
May test. The pooled data, including the 13 Feb-
ruary test, indicated that the mites caused a 70%
reduction in sepsid emergence. The overall number
of sepsids emerging from control pats was 30%
lower than the number of buffalo flies from these
pats. Therefore, M. peregrinus preyed more heavi-
lyon the less numerous Sepsidae than on the buffalo
flies.
The presence of alternate prey must be taken
into account in relating the results of the field trial
to the potential impact of M. peregrinus on buffalo
fly populations. In these tests, Sepsidae and other
Diptera had only a limited time to colonize the
pats before the mesh cones were applied. Under
normal field conditions, there would be larger
numbers of alternate prey available; this could re-
duce the level of predation on the buffalo fly. A
second factor that must be considered is that the
numbers of mites per pat used in these tests rep-
resent a level of colonization that may occur in-
frequently in the field, Wallace & Holm (1983)
reported that weekly mite collections in pitfall traps
exceeded 500 per trap on only 5 of 29 occasions.
We assume that similar numbers would colonize
dung pats.
The results of our study indicate that M. pere-
grinus may not be a very effective predator of the
buffalo fly. Natural prey location, prey stage at-
tacked, preference for alternative prey species, and
the need for high predator /prey ratios tend to limit
predation by these mites on this species. The es-
tablishment and dispersal of M. peregrinus is more
likely to have an effect on the bush fly (for which
the mites have demonstrated a preference), which
oviposits in more accessible parts of the dung pat.
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Received for publication 31 July 1987; accepted 11
February 1988.