Immature Stages and Biology of Fifteen Species of

Peruvian Calliphoridae (Diptera)

BERNARD GREENBERG AND MICHAEL L. SZYSKA
Department of Biological Sciences, University of Illinois at Chicago,
Chicago, Illinois 60680

Ann. Entomol. Soc. Am. 77: 488-517 (1984)

ABSTRACT Isofemale rearings from egg to adult are reported for 15 species of Peruvian
blow flies. Twelve reared for the first time are: Chrysomya chloropyga putoria (Wiede-
mann), Compsomyiops boliviano. (Mello), Compsomyiops verena (Mello), Hemilucilia flaw-
faces (Engel), Hemilucilia hermanlenti (Mello), Paralucilia fulvinota (Bigot), Calliphora
peruxrtana (Robineau-Desvoidy), Phoenicia ibis (Shannon), Sarconesia chlorogaster (Wiede-
mann), Sarconesia magellanica (Macquart), Sarconesia splendida (Townsend), and Sarco-
nesia versicolor (Bigot). Cochliomyia macellaria (¥.), Phoenicia cuprina (Wiedemann), and
Phoenicia eximia (Wiedemann) were also reared. Some of these flies are exclusively Andean
or rain forest, others span both habitats. The study includes: descriptions of first, second,
and third instars, puparia, and egg plastrons; developmental rates; successional and diel
activity of highland flies in Montaro Valley; and a key to the known third instars of Peru.
Similarities exist between species of Sarconesia, Calliphora, and Phoenicia in developmental
rates and morphology of the immature stages, suggesting that Sarconesia may be more
closely related to the Calliphorinae than previously thought.

THE ROLE OF calliphorids in disease transmission Methods

and parasitism is well documented (Greenberg
1971, 1973, James 1947, Zumpt 1965). In Peru the
primary screwworm, Cochliomyia hominivorax
(Coquerel), and the secondary myiasis producer,
Chrysomya albiceps (Wiedemann), threaten de-
veloping livestock industries in the jungle and else-
where (Baumgartner and Greenberg 1983). The
rapid spread in South America of three recently
introduced Chrysomya species—megacephala (F.),
albiceps, and chloropyga putoria (Wiedemann)—
has been documented (Aradi and Guimaraes 1982)
and the concomitant suppression of the endemic
Cochliomyia macellaria (F.) has been noted
(Baumgartner and Greenberg 1984).
Peru is divided into three major regions—the
Pacific coastal desert, Andes mountains, and the
eastern sloping rain forest—that offer a large num-
ber of habitats over a relatively short distance.
There are approximately 30 species of blow flies
in Peru.
Basic to understanding the medical and veteri-
nary importance and control of these flies is knowl-
edge of their biology. Very little bionomic work
exists on the strictly South American species and,
in view of the dwindling tropical forests, compet-
itive introductions, and other irreversible changes,
such studies are imperative.
Fifteen species of blow flies were reared, 12 for
the first time. The study includes morphological
descriptions of first, second, and third instars, pu-
paria, and egg plastrons; developmental rates,
successional and diel activity of some species; and
a key to the known third instars of Peru.
Rearings. Flies were collected along the Peru-
vian Carretera Central, which originates in Lima,
traverses the Andes, and terminates in the rain
forest. The following flies were collected at the
places indicated: Compsomyiops boliviana (Mel-
lo)12 (=Paraludlia boliviana [Mello]), Montaro
Valley; Sarconesia chlorogaster (Wiedemann),2
Montaro Valley and La Oroya (3,660 m); Sarco-
nesia magellanica (Macquart),2 Montaro Valley;
Sarconesia splendida (Townsend),2 Morococha
(4,500 m); Sarconesia versicolor (Bigot),2 Moro-
cocha. Rearings of the above flies were performed
in the Sierra during December 1979, on a small
ranch (Hacienda San Juan, San Lorenzo, alt. 3,550
m) 10 km south of Jauja, in Montaro Valley.
The following flies were collected at the places
indicated: Chrysomya chloropyga putoria
(Wiedemann),2 San Ramon; Cochliomyia macel-
laria (F.), San Ramon and Playa Hermosa (1,000
m); Compsomyiops verena (Walker)12 (=Paralu-
cilia verena [Mello]), 16 km west of San Ramon
(1,430 m); Hemilucilia hermanlenti (Mello),23 16
km west of San Ramon; Hemilucilia semidia-
phana (Rondani)12 (=Hemilucilia flavifacies [En-
1
New generic combination, Dear (1984) and personal com-
munication from Adrian Pont.
8
Species reared for the first time.
3
Baumgartner (unpublished data) has examined 1,026 speci-
mens of H. hermanlenti and concludes that it is distinct from H.
semidiaphana (Dear 1984) and H. segmentaria (Mariluis 1980),
based on a different altitudinal distribution and absence of mor-
phological intermediates.

488
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES
Fig. 1. Fly-rearing container.
gel]), 16 km west of San Ramon; Paralucilia ful-
vinota (Bigot)12 (=Myiolucilia fulvinota [Bigot]),
16 km west of San Ramon; Phaenicia cuprina
(Wiedemann), San Ramon and Puerto Bermudez
(500 m); Calliphora peruviana2 R-D, San Ramon
and Huasahuasi (2,750 m); Phaenicia eximia
(Wiedemann), Playa Hermosa and Puerto Ber-
mudez; Phaenicia ibis2 (Shannon), 16 km west of
San Ramon; S. splendida,2 15 km north of Tarma
(3,100 m); and S. versicolor,2 west of Tarma (3,800
m). The flies above were reared in the upper rain
forest at San Ramon (1,000 m) during June and
July 1980 and December 1981.
Gravid females were netted on fish and liver
baits, identified, and placed in transparent 1-gal
polyethylene containers fitted with a 15-cm di-
ameter stockinette access sleeve (Fig. 1). Cages
were provided a 180-ml specimen cup with 30 g
of fish and beef liver and two 30-ml cups for water
and sugar; food was added as needed. Fish and
liver were partially buried in sawdust, which ab-
sorbed the juices of the decomposing meat and
provided a medium for pupariation. Generally, flies
laid eggs shortly after capture, or within 4 days.
Recalcitrant females were isolated in 6-dram vials
plugged with meat-soiled cotton. This method
proved successful for obtaining eggs from C. bo-
liviana, S. splendida, S. magellanica, P. fulvinota,
and P. ibis. When discovered, an egg batch was
placed in a separate cup and covered with tissue
paper. Isofemale rearings assured more exact data
on developmental time and morphology of a single
brood.
Median time for approximately half the popu-
lation to reach the next stage was recorded for
eggs, each instar, puparia, and eclosion. Larvae
489
were examined twice daily and pupae once a day.
Pupariation usually occurred in the specimen cup,
but sawdust was placed at the bottom of the rear-
ing container for wandering prepupae. Minimal
and maximal temperatures were recorded daily.
In the Sierra, humidity was provided by adding
small amounts of water to the sawdust. All rearings
were performed under natural light. Due to time
constraints it was necessary to complete some of
the rearings in Lima.
Samples of eggs, each instar, and pupae were
preserved in a 70% ethanol/glycerine solution
(9:1), and are retained in the collections of the
Univ. of Illinois at Chicago, as are the initial fe-
male and her adult progeny. Because the cuticle
of the third instar is not easily penetrated by pre-
servatives, larvae were injected posterodorsally with
an 80% ethanol/glycerine/glacial acetic acid so-
lution (8:1:1) to prevent decomposition and shrink-
age.
The three instars were cleared in boiling 10%
KOH and stored in glycerine for detailed study of
cephalopharyngeal skeletons, spine bands and spine
types, anterior and posterior spiracles, and tuber-
cle size and arrangement. Spiracles of the first and
second instars are not described or figured, be-
cause their presence and pattern are typical—the
anterior spiracle first appears in the second instar
and the posterior spiracle has one slit in the first
instar, two in the second, and three in the third.
Eggs were rinsed in distilled water, fixed in 1%
osmium tetroxide for 6 h, dehydrated stepwise in
ethanol, critical-point dried, sputter-coated with
gold/palladium, and viewed with a Cambridge MK
2A Stereoscan electron microscope. Puparia were
treated similarly but were not fixed in osmium
tetroxide.
Concurrent with the rearings in 1979 a study
was made of diurnal activity and blow fly succes-
sion on carrion in Montaro Valley. Flies were col-
lected from 1- to 5-day-old dogs (road kills). Col-
lections consisted of netting flies for 1-min intervals
on an hourly basis from 0730 to 1500 hours. Num-
ber of calliphorids and other muscoid flies, hourly
temperature, RH, wind velocity, and cloud con-
ditions were recorded.
Description of Larvae and Puparia
Chrysomyinae
Chrysomya chloropyga putoria. Third Instar.
Anterior of segments 2-7 encircled with spines (Fig.
2). Segment 8 usually interrupted on dorsum, but
sometimes completed by a few randomly placed
spines. Segment 5 with lateral fusiform area; on
subsequent segments these areas sometimes vari-
able depending on how well spine bands devel-
oped laterad. Spine bands generally narrowed lat-
erad on segments 6-8, sometimes incomplete.
Spines on segments 9-12 restricted ventrad. Spines
multipointed, moderately pigmented (Fig. 100),
 490 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

ODT

OVT

8
Fig. 2 - 5 . C. c. putoria, third instar larva. Fig. 2. Lateral view. Fig. 3. Posterior view. IDT, Inner dorsal tubercles;
IVT, inner ventral tubercles; MDT, median dorsal tubercles; MVT, median ventral tubercles; ODT, outer dorsal
tubercles; OVT, outer ventral tubercles. Fig. 4. Anterior spiracle. Fig. 5. Right posterior spiracle. P, Peritreme.
Fig. 6. Cephalopharyngeal skeleton, first instar. Fig. 7. Cephalopharyngeal skeleton, second instar. Fig. 8. Ce-
phalopharyngeal skeleton, third instar.

noticeably larger posteriad, especially on anal pro-
tuberance. Elongate single- and double-pointed
transparent intersegmental spines surrounding
posterior segments (Fig. 101).
Anterior spiracles (Fig. 4) with 10 to 12 branch-
es (mean = 10 ± 0.8, n = 25). Posterior spiracles
heavily sclerotized with incomplete peritreme and
no button (Fig. 5). Mean spiracular width and sep-
aration 0.34 ± 0.01 mm (n = 28), and 0.18 ± 0.02
mm (n = 14), respectively.
Distance between inner tubercles on upper stig-
mal field approximately equal to distance between
inner and median tubercles, or slightly greater (Fig.
3). Median tubercles slightly closer to outer tuber-
cles than inner ones. Spine pattern on anal protu-
berance bell-shaped. Cephalopharyngeal skeleton
as in Fig. 8.
Second Instar. Spine patterns similar to third
instar, except segments 2-8 with complete bands
of spines. Segment 9 usually interrupted on dor-
sum, but may be completed by 1 or 2 rows of
spines. Lateral fusiform areas sometimes extend-
ing to segment 9. Usually with single or double
row of spines (sometimes pigmented) encircling
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 491

2.0 mm

0.2 mm

12

Fig. 9-12. C. macellaria, third instar larva. Fig. 9. Lateral view. FU, Fusiform area. Fig. 10. Posterior view.
Fig. 11. Anterior spiracle. Fig. 12. Right posterior spiracle. Fig. 13. Cephalopharyngeal skeleton, first instar. Fig.
14. Cephalopharyngeal skeleton, second instar. Fig. 15. Cephalopharyngeal skeleton, third instar.

posterior of segment 11. Intersegmental spines not
present. Cephalopharyngeal skeleton as in Fig. 7.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 6.
Puparium. See Fig. 116.
Cochliomyia macellaria. Third Instar. Ante-
rior of segments 2-9 with complete bands of spines
(Fig. 9). Segment 10 incomplete on dorsum; sub-
sequent segments restricted ventrad. Posterior of
all segments without spine bands. Fusiform areas
generally on segments 5-9, although variable.
Spines large, multipointed, heavily pigmented (Fig.
102).
Anterior spiracles (Fig. 11) with 8-12 branches
492 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

0.2 mm,

19

0.05 mm
.0.1 mm.

20
21

AS

Fig. 16-19. C. boliviano, third instar larva. Fig. 16. Lateral view. Fig. 17. Posterior view. Fig. 18. Anterior
spiracle. Fig. 19. Right posterior spiracle. Fig. 20. Cephalopharyngeal skeleton, first instar. Fig. 2 1 . Cephalophar-
yngeal skeleton, second instar. Fig. 22. Cephalopharyngeal skeleton, third instar. AS, Accessory sclerite.

(mean 10 ± 0.9, n = 26). Posterior spiracles heavi- protuberance U- or V-shaped. Cephalopharyngeal

ly pigmented, with incomplete peritreme and faint
button (Fig. 12). Mean spiracular width and sep-
aration 0.31 ± 0.02 mm (n = 14) and 0.19 ± 0.05
mm (n = 7), respectively.
Distance between inner tubercles on upper stig-
mal field approximately equal to distance between
inner and outer tubercles, or at least greater than
distance to median tubercles (Fig. 10). Lower pair
of inner tubercles on lower stigmal field not ap-
parent or barely evident. Pattern of spines on anal
skeleton as in Fig. 15.
Second Instar. Spine patterns similar to third
instar, except segment 11 sometimes incomplete
on dorsum. Cephalopharyngeal skeleton as in Fig.
14.
First Instar. Spine patterns similar to second in-
star. Cephalopharyngeal skeleton as in Fig. 13.
Puparium. See Fig. 117.
Compsomyiops boliviano. Third Instar. Ante-
rior of segments 2-10 encircled with spines (Fig.
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES
16). Segment 11 usually completed by 1 or 2 rows.
Spines on segment 12 restricted ventrad. Fusiform
areas laterad on segments 5-10. Posterior of seg-
ments usually lacking spines. Spines large, single-
and multipointed, well pigmented (Fig. 103).
Anterior spiracles (Fig. 18) with 10-13 branches
(mean = 11 ± 0.7, n = 18). Posterior spiracles
heavily pigmented with indistinct button and in-
complete peritreme (Fig. 19). Mean spiracular
width and separation 0.37 ± 0.04 mm (n = 16)
and 0.23 ± 0.03 mm (n = 8), respectively.
Distance between inner tubercles on upper stig-
mal field slightly less than distance between inner
and outer tubercles (Fig. 17). Pattern of spines on
dorsum of anal protuberance V- or U-shaped.
Cephalopharyngeal skeleton as in Fig. 22.
Second Instar. Anterior of segments 2-9 encir-
cled with spines. Segment 9 sometimes incom-
plete; spines on rest of larva, like third instar.
Cephalopharyngeal skeleton as in Fig. 21.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 20.
Puparium. See Fig. 118.
Compsomyiops verena. Immature forms of this
species are indistinguishable from C. boliviana.
Hemilucilia flavifacies. Third Instar. Anterior
of segments 2-6 with complete bands of spines
(Fig. 23). Segment 7 interrupted on dorsum, some-
times complete on dorsum by a few spines. Seg-
ments 8-11 with poorly denned rows of spines lat-
erad. Posterior of segment 11 encircled with spines,
generally 2-3 rows on dorsum. Segment 10 with
groups of transparent spines along posterior mar-
gin. Lateral fusiform areas absent. Spines single-
and multipointed, moderately pigmented (Fig.
104).
Anterior spiracles (Fig. 25) with 9-11 branches
(mean = 10 ± 0.7, n = 17). Posterior spiracles
heavily sclerotized with incomplete peritreme and
indistinct button (Fig. 26). Mean spiracular width
and distance between spiracles 0.23 ± 0.07 mm
(n = 8) and 0.15 ± 0.01 mm (n = 4), respectively.
Distance between inner tubercles on stigmal field
approximately equal to distance between inner and
outer tubercles (Fig. 24). Median tubercles closer
to outer tubercles. Spine pattern on anal protuber-
ance U-shaped. Cephalopharyngeal skeleton as in
Fig. 29.
Second Instar. Anterior of segments 2-5 encir-
cled with spines. Posterior of segment 11 with well-
developed band of spines, darker than third instar.
Cephalopharyngeal skeleton as in Fig. 28.
First Instar. Spine patterns similar to second in-
star; segment 5 sometimes interrupted on dorsum.
Spines on segments 6-12 restricted laterad. Ceph-
alopharyngeal skeleton as in Fig. 27.
Puparium. See Fig. 119.
Hemilucilia hermanlenti. Third Instar. Ante-
rior of segments 2-7 encircled with spines (Fig.
30). Segment 7 sometimes incomplete on dorsum,
and segments 8-11 with weak rows of spines lat-
erad. Posterior of segment 11 encircled with spines,
493
usually with groups of spines on dorsum. Segment
10 usually with groups of pigmentless spines en-
circling posterior margins. Lateral fusiform areas
absent. Spines, single- and multipointed, moder-
ately pigmented (Fig. 105). Spines especially large
and numerous ventrad on 12th segment (Fig. 106);
also true of H. flavifacies.
Anterior spiracles (Fig. 32) with 10-12 branches
(mean = 11 ± 0.8, n = 17). Posterior spiracles
heavily sclerotized with incomplete peritreme and
indistinct button (Fig. 33). Mean spiracular width
and separation 0.32 ± 0.07 mm (n = 8) and 0.17 ±
0.7 mm (n = 4), respectively.
Tubercle size and distribution similar to that of
H. flavifacies, except position of median tubercles
variable (Fig. 31). Cephalopharyngeal skeleton as
in Fig. 36.
Second Instar. Anterior spine bands usually
complete to segment 6; some specimens incom-
plete on dorsum and others complete to segment
7. Segments 7-12 with weak rows of spines later-
ad. Spines on posterior of segment 11 darker than
in third instar, but rest of larva similar. Cephalo-
pharyngeal skeleton as in Fig. 35.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 34.
Puparium. Similar to H. flavifacies (Fig. 119).
Paralucilia fulvinota. Third Instar. Late third
instar and prepupa blue. Anterior of segments 2-10
encircled with spines (Fig. 37). Segment 11 usually
incomplete on dorsum, sometimes completed by
weak row. Posterior segments without spines. Lat-
eral fusiform spine areas present on segments 5-
10. Spines large, single- and multipointed, heavily
pigmented (Fig. 107).
Anterior spiracles (Fig. 39) with 9-12 branches
(mean = 10 ± 0.8, n = 20). Posterior spiracles
moderately pigmented with incomplete peritreme
and faint button (Fig. 40). Mean spiracular width
and separation 0.35 ± 0.01 mm (n = 10) and
0.25 ± 0.03 mm (n = 5), respectively.
Distance between inner tubercles on upper stig-
mal field slightly less than distance between inner
and median tubercles (Fig. 38). Median tubercles
approximately equidistant to inner and outer tu-
bercles. Spine pattern on anal protuberance
U-shaped. The cephalopharyngeal skeleton as in
Fig. 43.
Second Instar. Anterior of segments 2-8 encir-
cled with spines. Segment 8 sometimes incomplete
on dorsum. Cephalopharyngeal skeleton as in Fig.
42.
First Instar. Spine patterns similar to second in-
star. Cephalopharyngeal skeleton as in Fig. 41.
Puparium. See Fig. 120.
Calliphorinae
Calliphora peruviana. Third Instar. Anterior
of segments 2-8 encircled with spines (Fig. 44).
Segment 9 incomplete on dorsum, 10 and 11 with
weak rows laterad. Posterior of segments 10 and
494 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

.0.1 mm.

27 28

Fig. 23-26. H. flavifacies, third instar larva. Fig. 23. Lateral view. Fig. 24. Posterior view. Fig. 25. Anterior
spiracle. Fig. 26. Right posterior spiracle. Fig. 27. Cephalopharyngeal skeleton, first instar. Fig. 28. Cephalopha-
ryngeal skeleton, second instar. Fig. 29. Cephalopharyngeal skeleton, third instar.

11 encircled with spines, weak laterad. Lateral fu-
siform areas absent. Spines scalelike, single- and
double-pointed, moderately pigmented (Fig. 108).
Anterior spiracles (Fig. 46) with 8-10 branches
(mean = 9 ± 0.7, n = 20). Posterior spiracles mod-
erately sclerotized, with delicate peritreme and
button (Fig. 47). Mean spiracular width and sep-
aration 0.33 ± 0.01 mm (n = 8) and 0.30 ± 0.06
mm (n = 4), respectively.
Distance between inner tubercles on upper stig-
mal field greater than distance between inner and
median tubercles (Fig. 45). Cephalopharyngeal
skeleton as in Fig. 50.
Second Instar. Spine patterns similar to third
instar. Cephalopharyngeal skeleton as in Fig. 49.
First Instar. Anterior of segments 2-7 encircled
with spines. Segment 7 sometimes incomplete on
dorsum. Segment 8 always incomplete dorsally, and
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 495

0.2 mm

33

0.05 mm 0.1 mm

34 35

Fig. 30-33. H. hermanlenti, third instar larva. Fig. 30. Lateral view. Fig. 31. Posterior view. Fig. 32. Anterior
spiracle. Fig. 33. Right posterior spiracle. Fig. 34. Cephalopharyngeal skeleton, first instar. Fig. 35. Cephalopha-
ryngeal skeleton, second instar. Fig. 36. Cephalopharyngeal skeleton, third instar.

subsequent segments with spines restricted ven-
trad. Spines not present on posterior of segment
10, but with few weak rows on posterior of seg-
ment 11. Cephalopharyngeal skeleton as in Fig.
48.
Puparium. See Fig. 121.
Phoenicia cuprina. Third Instar. Anterior of
segments 2-8 encircled with spines (Fig. 51). Seg-
ment 9 with poorly defined row of spines on dor-
sum, sometimes with 1 or 2 rows complete. Ante-
rior spine bands on segments 10-12 usually
restricted laterad. Segment 11 with band of spines
along posterior margin, with 1-3 rows on dorsum.
Lateral fusiform areas absent. Spines single-point-
ed, lightly pigmented (Fig. 109).
Anterior spiracles (Fig. 53) with 5-6 branches
(mean = 5 ± 0.5, n = 20). Posterior spiracles light-
ly sclerotized, surrounded by round, delicate, com-
plete peritreme; button present (Fig. 54). Mean
spiracular width and separation 0.37 ± 0.00 mm
496 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

t0.2 mrt\

40

g.05 mm

41

Fig. 37-40. P. fulvinota, third instar larva. Fig. 37. Lateral view. Fig. 38. Posterior view. Fig. 39. Anterior
spiracle. Fig. 40. Right posterior spiracle. Fig. 4 1 . Cephalopharyngeal skeleton, first instar. Fig. 42. Cephalopha-
ryngeal skeleton, second instar. Fig. 43. Cephalopharyngeal skeleton, third instar.

(n = 20) and 0.39 ± 0.03 mm (n = 10), respec-
tively.
Distance between inner tubercles on upper field
approximately equal to distance between inner and
outer tubercles (Fig. 52). Median tubercles some-
times slightly closer to inner tubercles (this distin-
guishes some specimens of P. cuprina from P. ex-
imia and P. ibis). Relative tubercle position variable
in all three Phaenicia species making diagnosis of
single specimens uncertain. Cephalopharyngeal
skeleton as in Fig. 57.
Second Instar. Anterior of segments 2-7 encir-
cled with spines. Segment 7 sparsely spined, band
sometimes interrupted dorsad. Segment 8 almost
always incomplete on dorsum, subsequent seg-
ments with spines restricted ventrad; otherwise like
third instar. Cephalopharyngeal skeleton as in Fig.
56.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 55.
Puparium. See Fig. 122.
Phaenicia eximia. Third Instar. Anterior spine
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 497

Fig. 4 4 - 4 7 . C. peruviana, third instar larva. Fig. 44. Lateral view. Fig. 45. Posterior view. Fig. 46. Anterior
spiracle. Fig. 47. Right posterior spiracle. ES, Ecdysial scar. Fig. 48. Cephalopharyngeal skeleton, first instar. Fig.
49. Cephalopharyngeal skeleton, second instar. Fig. 50. Cephalopharyngeal skeleton, third instar.

bands encircle segments 2-7 (Fig. 58). Segment 7
sometimes incomplete on dorsum. Segments 8-12
generally incomplete on dorsum, frequently with
weak rows laterad. Posterior of segment 11 with
spines, usually weak rows on dorsum, absent lat-
erally. Lateral fusiform areas absent. Spines lightly
pigmented, single-pointed as in other Phaenicia
(Fig. 110).
Anterior spiracles (Fig. 60) with 6-8 branches
(mean = 7 ± 0.7, n = 9). Posterior spiracles sur-
rounded by delicate, complete peritreme with but-
ton (Fig. 61). Mean spiracular width and separa-
tion 0.40 ± 0.03 mm (n = 10) and 0.46 ± 0.01
mm (n = 5), respectively.
Distance between inner tubercles on upper stig-
mal field approximately equal to distance between
inner and outer tubercles, or slightly more (Fig.
59). This separates P. eximia from P. ibis. Median
498 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

m
2.0 mm

51

0.2 mm

53

0.2 mm

57
Fig. 51-54. P. cuprina, third instar larva. Fig. 51. Lateral view. Fig. 52. Posterior view. Fig. 53. Anterior
spiracle. Fig. 54. Right posterior spiracle. Fig. 55. Cephalopharyngeal skeleton, first instar. Fig. 56. Cephalopha-
ryngeal skeleton, second instar. Fig. 57. Cephalopharyngeal skeleton, third instar.

tubercles closer to outer tubercles in 8 of 13 spec-
imens. In P. cuprina median tubercles sometimes
closer to inner ones. Cephalopharyngeal skeleton
as in Fig. 64.
Second Instar. Anterior of segments 2-6 with
complete band of spines. Segment 7 usually in-
complete on dorsum, spines on segments 8-12 re-
stricted ventrad. Posterior of segment 11 usually
with one or more weak rows of spines, otherwise
like third instar. Cephalopharyngeal skeleton as in
Fig. 63.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 62.
Puparium. Similar to P. cuprina (Fig. 122).
Phoenicia ibis. Third Instar. Anterior of seg-
ments 2-8 encircled with spines (Fig. 65). Spines
on segments 9-12 generally restricted ventrad,
segment 9 with weak row of spines laterally. Pos-
terior of segment 11 encircled with spines, usually
few groups on dorsum. Lateral fusiform spine areas
absent. Spines single-pointed, lightly pigmented
(Fig. 111).
Anterior spiracles (Fig. 67) with 6-8 branches
(mean = 7 ± 0.6, n = 28). Posterior spiracles del-
icate, lightly pigmented with complete peritreme,
well defined button (Fig. 68). Mean spiracular
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 499

.0.1 mm.

63

64
Fig. 5 8 - 6 1 . P. eximia, third instar larva. Fig. 58. Lateral view. Fig. 59. Posterior view. Fig. 60. Anterior
spiracle. Fig. 6 1 . Right posterior spiracle. Fig. 62. Cephalopharyngeal skeleton, first instar. Fig. 63. Cephalopha-
ryngeal skeleton, second instar. Fig. 64. Cephalopharyngeal skeleton, third instar.

width and separation 0.38 ± 0.03 mm (n = 28)
and 0.44 ± 0.05 mm (n = 14), respectively.
Distance between inner tubercles on upper stig-
mal field usually less than distance between inner
and outer tubercles (Fig. 66). This distinguishes it
from P. eximia. Median tubercles closer to outer
tubercles. Cephalopharyngeal skeleton as in Fig.
71.
Second Instar. Spine patterns similar to third
instar; sometimes anterior bands usually complete
to segment 7; spines more heavily pigmented.
Cephalopharyngeal skeleton as in Fig. 70.
First Instar. Spine patterns similar to second in-
star. Cephalopharyngeal skeleton as in Fig. 69.
Puparium. Similar to P. cuprina (Fig. 122).
Toxotarsinae
Sarconesia chlorogaster. Third Instar. Ante-
rior of segments 2-8 encircled with spines (Fig.
500 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

9-2 mm

68

0.1 mm

70

Fig. 65-68. P. ibis, third instar larva. Fig. 65. Lateral view. Fig. 66. Posterior view. Fig. 67. Anterior spiracle.
Fig. 68. Right posterior spiracle. Fig. 69. Cephalopharyngeal skeleton, first instar. Fig. 70. Cephalopharyngeal
skeleton, second instar. Fig. 71. Cephalopharyngeal skeleton, third instar.

72). Segment 8 sometimes interrupted on dorsum,
usually with single row dorsally. Segments 9-12
restricted ventrad. Dorsum of segment 11 with
spines, usually 2 or 3 rows, sometimes posterior
band absent. Spines small, single-pointed, lightly
pigmented to transparent (Fig. 112).
Anterior spiracles (Fig. 74) with 8 to 10 branch-
es (mean = 9 ± 0.8, n = 12). Posterior spiracles
with delicate, lightly sclerotized, oval, buttonless
peritreme (Fig. 75). Mean spiracular width and
separation 0.24 ± 0.02 mm (n = 22) and 0.26 ±
0.02 mm (n = 11), respectively.
Distance between inner tubercles on upper stig-
mal field approximately equal to distance between
inner and median tubercles occasionally slightly
larger (Fig. 73). Median tubercles closer to outer
tubercles. Cephalopharyngeal skeleton as in Fig.
78.
Second Instar. Spine patterns similar to third
instar. Cephalopharyngeal skeleton as in Fig. 77.
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 501

0.2 mm
I—I
74

Fig. 72-75. S. chlorogaster, third instar larva. Fig. 72. Lateral view. Fig. 73. Posterior view. Fig. 74. Anterior
spiracle. Fig. 75. Right posterior spiracle. Fig. 76. Cephalopharyngeal skeleton, first instar. Fig. 77. Cephalopha-
ryngeal skeleton, second instar. Fig. 78. Cephalopharyngeal skeleton, third instar.

First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 76.
Puparium. See Fig. 123.
Sarconesia magellanica. Third Instar. Ante-
rior of segments 2-7 with complete bands of spines
(Fig. 79). Segment 8 sometimes complete by 1 or
2 rows of spines, generally incomplete on dorsum.
Spines on segments 10 and 11 with weak rows
laterad. Posterior of segment 11 encircled with
spines, 2 or 3 rows on dorsum. Lateral fusiform
areas absent. All spines small, single-pointed, light-
ly pigmented (Fig. 113).
Anterior spiracles (Fig. 81) with 7 to 10 branch-
es (mean = 8 + 0.6, n = 40). Posterior spiracles
oval, surrounded by incomplete lightly sclerotized
peritreme without button (Fig. 82). Mean spirac-
ular width and separation 0.20 ±0.1 mm (n = 20)
and 0.29 ± 0.07 mm (n = 10), respectively.
Distance between inner tubercles on upper stig-
mal field approximately equal to distance between
inner and median tubercles (Fig. 80). Median tu-
bercles usually equidistant from inner and outer
tubercles. Cephalopharyngeal skeleton as in Fig.
85.
502 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Second Instar. Spine patterns similar to third
instar. Cephalopharyngeal skeleton as in Fig. 84.
First Instar. Spine patterns similar to third in-
star, except segments 2-8 usually encircled with
spines. Cephalopharyngeal skeleton as in Fig. 83.
Puparium. Similar to S. chlorogaster (Fig. 123).
Sarconesia splendida. Third Instar. Anterior
of segments 2-8 encircled with bands of spines
(Fig. 86). Segment 9 usually interrupted on dor-
sum. Segments 9-11 with weak rows of spines lat-
erad. Posterior of segment 11 encircled with weak
rows of spines. Lateral fusiform areas absent; all
spines single-pointed, lightly pigmented (Fig. 114).
Anterior spiracles (Fig. 88) with 6 to 8 branches
(mean = 7 + 0.9, n = 20). Posterior spiracles oval,
lightly pigmented with incomplete peritreme (Fig.
89), button absent. Mean spiracular width and sep-
aration 0.21 ± 0.01 mm (n = 18) and 0.23 ± 0.04
mm (n = 9), respectively.
Distance between inner tubercles on upper stig-
mal field approximately equal to distance between
inner and outer tubercles, or slightly less (Fig. 87).
Median tubercles closer to outer tubercles than in-
ner ones. Cephalopharyngeal skeleton as in Fig.
92.
Second Instar. Spine patterns similar to third
instar. Cephalopharyngeal skeleton as in Fig. 91.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 90.
Puparium. Similar to S. chlorogaster (Fig. 123).
Sarconesia versicolor. Third Instar. Anterior
of segments 2-8 encircled with spines (Fig. 93).
Segment 8 usually complete by 2 or 3 rows of
spines on dorsum. Segment 9 generally incom-
plete; segments 10-12 with spines restricted ven-
trad, except for band of spines encircling posterior
of segment 11. Lateral fusiform areas absent. Spines
single-pointed, lightly pigmented (Fig. 115).
Anterior spiracles (Fig. 95) with 6 to 8 branches
(mean = 7 ± 0.7, n = 15). Posterior spiracles oval,
lightly pigmented with incomplete peritreme, no
button (Fig. 96). Mean spiracular width and sep-
aration 0.18 ± 0.01 mm (n = 6) and 0.31 ± 0.03
mm (n = 3), respectively.
Distance between inner tubercles on upper stig-
mal field about same or slightly less than distance
between inner and median tubercles (Fig. 94). Me-
dian tubercles closer to outer tubercles. Cephalo-
pharyngeal skeleton as in Fig. 99.
Second Instar. Anterior of segments 2-7 com-
pletely encircled with spines. Segment 8 generally
interrupted on dorsum. Cephalopharyngeal skel-
eton as in Fig. 98.
First Instar. Spine patterns similar to third in-
star. Cephalopharyngeal skeleton as in Fig. 97.
Puparium. Similar to S. chlorogaster (Fig. 123).
Provisional Key to Known Third Instar
Larvae of Peru
1. Peritreme incomplete (Compsomyiops, Fig.
19; Hemilucilia, Fig. 26; Paralucilia, Fig.
40; Cochliomyia, Fig. 12; Chrysomya, Fig.
5; Sarconesia, Fig. 75) 2 apically. However, the apices anastomose more ex-
Vol. 77, no. 5
Peritreme complete (Phaenicia, Fig. 54; Cal-
liphora, Fig. 47) 7
2. Lateral fusiform areas absent. With accessory
dental sclerite (Fig. 22). (Hemilucilia, Fig.
29; Sarconesia, Fig. 78) 3
Lateral fusiform areas present (Fig. 9). With
or without accessory dental sclerite.
(Compsomyiops, Paralucilia, Cochlio-
myia, Chrysomya) 4
3. All spines single-pointed, small. (Fig. 112-
115) Sarconesia
Spines single- and multipointed, especially
large and numerous ventrad on segment
12. (Fig. 104-106 Hemilucilia
4. With accessory dental sclerite . . Compsomyiops
Without accessory dental sclerite. (Paraluci-
lia, Fig. 43; Cochliomyia, Fig. 15; Chrys-
omya, Fig. 8) 5
5. Spine pattern on anal protuberance convex
or bell-shaped (Fig. 3)
Chrysomya c. putoria
Spine pattern on anal protuberance U- or
V-shaped. (Paralucilia, Fig. 17; Cochlio-
myia, Fig. 10) 6
6. Mature third instar blue-gray
Paralucilia fulvinota
Mature third instar nonpigmented
Cochliomyia macellaria
7. With accessory dental sclerite (Fig. 50) . . . .
Calliphora peruviana
Without accessory dental sclerite 8
8. Our specimens of P. cuprina, eximia, and
ibis are difficult to separate due to similar-
ity and variability of characters.
Distance between inner tubercles on upper
stigmal field usually less than distance be-
tween inner and outer tubercles (Fig. 66)
Phaenicia ibis
Distance between inner tubercles about equal
to distance between inner and outer tuber-
cles 9
9. Median tubercles on upper stigmal field usu-
ally closer to inner than to outer tubercles
(Fig. 52) Phaenicia cuprina
Median tubercles usually closer to outer tu-
bercles (Fig. 59) Phaenicia eximia
Plastron Morphology
The anterior third of the egg plastron, between
the hatching lines and before it bifurcates around
the micropyle, was examined by scanning electron
microscopy (SEM) in the 15 species. There were
differences in the surface of the outer chorionic
network among the three subfamilies. In Chryso-
myinae, the apices of the vertical struts are peglike
and do not anastomose (Fig. 124-129). In Calli-
phorinae the struts are flared apically and appear
suckerlike; pores in the apices may provide addi-
tional air passage to the interior (Fig. 130-133).
The Sarconesia resemble the Calliphorinae we
studied in that the vertical struts are expanded
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 503

0.1 mm.

84

0.2 mm

85
Fig. 79-82. S. magellanica, third instar larva. Fig. 79. Lateral view. Fig. 80. Posterior view. Fig. 81. Anterior
spiracle. Fig. 82. Right posterior spiracle. Fig. 83. Cephalopharyngeal skeleton, first instar. Fig. 84. Cephalopha-
ryngeal skeleton, second instar. Fig. 85. Cephalopharyngeal skeleton, third instar.

tensively (Fig. 134-137). One can assume that the Bionomics

more intact outer network of the latter two groups
confers greater resistance to wetting when eggs are
submerged or deposited in decomposing carrion
(Hinton 1960). Whether this structural difference,
by itself, improves plastron respiration in a sub-
merged egg or increases the desiccation resistance
of such an egg in dry air, thus increasing the num-
ber of potential oviposition sites, are questions be-
yond the scope of this report.
Developmental rates of preadult stages, and
rearing sites, dates, and temperatures are given in
Tables 1 and 2. The terms "eusynanthropic,"
"hemisynanthropic," and "asynanthropic" de-
scribe thefly'ssynanthropy or association with man;
by extension we have applied these terms to de-
scribe the fly's habitat. Thus, eusynanthropic refers
to dense, human habitations (e.g., cities and vil-
504 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

Table 1. Andean rearings, December 1979

Median duration of immatures (h) Egg to

Pupa
Larva Larva Larva III Larval adult
Egg (days)
batch I II and prepupa duration (days)

Chrysomyinae
C. boliviano 57 23 49 144* 216 9 20.4
50 23 96 214* 333 10 26.0
Toxotarsinae
S. chlorogaster 36 24 60 203 287 12* 25.5
27 49 49 191 289 11* 24.2
24 24 — 189* 266 11 23.1
S. magellanica 26 24 38 265* 315 13 27.2
25 35 38 208* 281 12 24.8
S. splendida 35 20 — 186* 291 11 24.6
— 20 22 182* 224 9 —
S. versicolor 37 38 22 179* 239 11 22.5

Data are replicate rearings in most cases. Data preceding the number marked with an asterisk are from rearings near Jauja, in
Montaro Valley, December 1979; average maximum and minimum daily temperatures: 20.3°C (SD ± 1.5) and 13.7°C (SD ± 1.5):
developmental data subsequent to that stage were obtained in Lima: average maximum and minimum daily temperatures: 26.7°C
(SD ± 1.7) and 24.8°C (SD ± 1.6).

lages), hemisynanthropic to isolated dwellings and tigation since it is a secondary myiasis producer in
asynanthropic to areas devoid of human settle- man and animals in the Old World and a vector
ment. In addition to data on developmental rates of human pathogens (Greenberg 1971, 1973).
of 15 species of blow flies, observations on adult Cochliomyia macellaria. Three rearings were
and larval behavior are included. Developmental performed from adults collected at Playa Hermosa
rates given below are averages. (1,000 m, eusynanthropic) from fish bait. Eggs were
deposited in clusters of 75 to 150, and had an in-
Chrysomyinae cubation time of 19.2 h; Melvin (1934) reports 33
h at 17.7°C. Developmental times for first and sec-
Chrysomya chloropyga putoria. Two batches ond instars were 15.3 and 24 h, respectively. Du-
of 150 and 200 eggs were laid on fish by adults rations of the third instar/prepupa and pupal stages
collected at San Ramon (1,000 m, eusynanthropic); were 77.7 h and 5.3 days, respectively. Laake
eggs hatched in 15.5 h. Developmental times for (1936) recorded a pupal period of about 3.5 days.
the first and second instars were 16.3 and 23.5 h, Egg-to-adult interval was 10.9 days.
respectively. Durations of the third instar/prepu- C. macellaria was the most abundant blow fly
pa and pupal stages were 86 h and 4 days, respec- in San Ramon during June 1980. Eighteen months
tively. This species had the fastest developmental later, it had decreased from 89% (n = 1163) to
rate of all blow flies reared, with a minimal egg- 0.2% (n = 2,101) of all blow flies collected, appar-
to-adult interval of 9.5 days. ently having been replaced by Chrysomya albi-
C. chloropyga putoria is African and was only ceps (Wiedemann) and C. c. putoria (Baumgart-
recently discovered in Peru (Baumgartner and ner and Greenberg 1984).
Greenberg 1984). Its life cycle was previously un- Compsomyiops boliviano. Adults were collect-
known, but presumed to be similar to that of C. ed in Montaro Valley (3,550 m, hemisynanthropic)
chloropyga chloropyga (Zumpt 1965). Develop- from dead dogs in December 1979; two rearings
mental rates of putoria approximate those report- were completed. Although Palma (1973) reared
ed for chloropyga by Smit (1931), Zumpt (1965) Paralucilia fulvicrura {^Compsomyiops fulvi-
and Prins (1982), although exact developmental crura), his rearing data are included for compar-
and temperature data are lacking for chloropyga ison because of the confusion in separating species
and much confusion has surrounded the designa- of this genus and because our larval material closely
tion of these two forms. The Peruvian form shows resembles his descriptions.
little predilection for feces, whereas in east Africa Eggs were laid in clusters of 75 to 250, and
"the mature larvae will roll out of a latrine as you hatched in 53.5 h. Palma (1973) reports an incu-
open the door" (B. Laurence, personal communi- bation period of 20 to 24 h at 20 to 25°C for P.
cation). "In habits," Laurence continues, "the south fulvicrura. Two-day-old dogs were usually cov-
African taxon appears to resemble the behavior of ered with mounds of Compsomyiops eggs, filling
the last species [C. chloropyga] in southern Africa crevices and lining the underside of the body. Eggs
rather than the behavior I know of C. putoria in laid in fur were securely cemented to hairs next
east Africa." This species warrants further inves- to the skin. Developmental time for the first instar
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 505

Table 2. Upper jungle rearings, June 1980

Median duration of immatures (h) Egg to

Pupa
Egg Larva Larva Larva III Larval (days) adult
batch I II and prepupa duration (days)
Chrysomyinae
C. c. putoria 14.5 18 24 76 118 4 9.5
16.5 14.5* 23 96 137 4 10.4
C. macellaria 14 11 24 71 106 5 10.0
22.5 18 24 71 113 6 11.6
21 12.5 24 91 127.5 5 11.2
C. verena 16 27.5 24 190 241.5 9 19.7
16 25 15 136 176 8 16.0
23.5 18 22.5 133.5 174 7 15.2
H. flavifacies 22 14 23 72 109 4.5 10.0
H. hermanlenti 25 16.5 24 78 118.5 5 11.0
14 23 24 117 164 6 13.4
— 15* 12 92 119 5 —
13 22 24 122 168 6.5 14.0
13.5 18 23 72 113 6 11.3
P. fulvinota 28 18 20.5 99 137.5 5 12.0
22 26 12 183 221 5 15.1
Calliphorinae
C. peruoiana 13 37 23 323 383 12 28.5
17 26 29 246 301 12 25.3
P. cuprina 12 17 20* 96 133 8 14.0
20 23 24 120 167 7 14.8
13 11.5 14* 96 121.5 10 15.6
15.5 23 24 96 143 8.5 15.1
P. eximia 14 13 34 144 191 15 23.5
12 13 13 144 170 15 22.6
12 13 12 144 169 15 22.5
P. ibis 28 23 12 279 314 12.5 26.8
22.5 12 12.5 279 303 15 28.6
Toxotarsinae
S. splendida 25.5 13 21 231 265 9 21.1
S. versicolor 14 23 11 170 204 11 20.1
16 28 20 184 232 11 21.3

Data are from replicate rearings at San Ramon: average maximum and minimum daily temperatures, 26.0°C (SD ± 3.1) and 21.7CC
(SD ± 1.9). Data preceding the number with an asterisk are rearings at Puerto Bermudez: average maximum and minimum daily
temperatures: 29.5°C (SD ± 1.0) and 22.9°C (SD ± 1.1).

was 23 h; Palma reports 38 to 42 h. Developmental Compsomyiops verena. Adults were collected

time for the second instar was 72.5 h; Palma re-
ports 30 to 34 h. Duration of the third instar/
prepupa stage was 179 h; Palma reports the third
instar and prepupa to be 38 to 112 h and 52 to 56
h, respectively. Duration of the pupal stage was
9.5 days; Palma reports 6 to 7 days. Egg-to-adult
interval was 23.2 days.
C. boliviana was the most abundant fly around
farms in Montaro Valley, where it avidly ovipos-
ited on dead dogs, fish, and liver. Ovipositing fe-
males were observed drilling their abdomens into
dense fur with wings held out perpendicularly for
support. Some of the flies had pollen covering the
head and thorax, suggesting a probable role as pol-
linators. The habits and life cycle of C. boliviana
are apparently very similar to western North
American C. wheeleri (Hough), investigated by
Deonier and Knipling (1940). Both are montane
species.
16 km west of San Ramon (1,430 m, asynanthrop-
ic) from fish and liver bait in June 1980. Three
rearings were completed. Eggs were laid in clus-
ters of 150 to 300, and hatched in 18.5 h. Devel-
opmental times for the first and second instars were
23.5 and 20.5 h, respectively. Durations of the third
instar/prepupa and pupal stages were 153.2 h and
8 days, respectively. Egg-to-adult interval was 17
days.
C. verena was the most abundant fly in asynan-
thropic areas near San Ramon. The habits of this
fly are similar to those of C. boliviana except for
a preference for warmer temperatures (lower el-
evations). Immature stages of this species are in-
distinguishable from C. boliviana and develop-
mental data probably differ because of slightly
higher rearing temperatures in San Ramon than
in the highlands.
Hemilucilia flavifacies. Adults were collected
506 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

2.0 mm

86

0.2 mm

89

0.1 mm.

91

Fig. 86-89. S. splendida, third instar larva. Fig. 86. Lateral view. Fig. 87. Posterior view. Fig. 88. Anterior
spiracle. Fig. 89. Right posterior spiracle. Fig. 90. Cephalopharyngeal skeleton, first instar. Fig. 9 1 . Cephalopha-
ryngeal skeleton, second instar. Fig. 92. Cephalopharyngeal skeleton, third instar.

16 km west of San Ramon (1,430 m, asynanthrop-
ic). One rearing was completed of this relatively
rare fly. One hundred eggs were laid in a cluster
and hatched in 22 h. The developmental times for
first and second instars were 14 and 23 h, respec-
tively. Durations of the third instar/prepupa and
pupal stages were 72 h and 4.5 days. The egg-to-
adult interval was 10 days.
H. flavifacies is far less abundant than its sibling
H. hermanlenti at the altitude of San Ramon. It
probably has a preference for warmer tempera-
tures and higher humidities since it is more abun-
dant in the lower jungle.
Hemilucilia hermanlenti. Adults were collect-
ed 16 km west of San Ramon (1,430 m, asynan-
thropic) in substantial numbers. Five rearings were
completed. Eggs were laid in clusters of 100 to
150, and incubation time was 16.4 h. Develop-
mental times for first and second instars were 18.9
and 21.4 h, respectively. Durations of the third
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 507

0..2 mm

96

0.1 mm

98

Fig. 93-96. S. versicolor, third instar larva. Fig. 93. Lateral view. Fig.. 94. Posterior view. Fig. 95. Anterior
spiracle. Fig. 96. Right posterior spiracle. Fig. 97. Cephalopharyngeal skeleton, first instar. Fig. 98. Cephalopha-
ryngeal skeleton, second instar. Fig. 99. Cephalopharyngeal skeleton, third instar.

instar/prepupa and pupal stages were 96.2 h and
5.7 days, respectively. Egg-to-adult interval was
12.4 days. This species is second only to Comp-
somyiops verena in abundance in asynanthropic
areas near San Ramon.
Paralucilia fulvinota. Adults were collected 16
km west of San Ramon (1,430 m, asynanthropic)
from fish and liver bait. Two rearings were com-
pleted of this relatively rare fly. Eggs were laid in
clusters of 75 to 120, and hatched in 25 h. Devel-
opmental times for first and second instars were
22 and 16.3 h, respectively. Duration of the third
instar/prepupa stage was 141 h; the prepupa ac-
counted for the prolongation of this stage. Forty-
eight hours into the third instar (still feeding) the
larvae turned blue and became prepupae shortly
thereafter. Color changes in Diptera larvae are re-
ported by Tobin (1971) and Shinonaga and Kano
(1973) and are thought to be a type of camouflage
against predators. This may be true for P. fulvi-
nota, because active prepupae when disturbed or
shadowed readily contracted and lay motionless
508 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

104 105 106 107

112 113 114 115

Fig. 100-115. SEM of spines on the puparium (5th segment, ventrolateral). Fig. 100. C. c. putoria. Fig. 101.
C. c. putoria, intrasegmental area of 9th segment (dorsolateral). Fig. 102. C. macellaria. Fig. 103. C. bolimana.
Fig. 104. H. fiavifacies. Fig. 105. H. hermanlenti. Fig. 106. Hemilucilia, 12th segment (ventrad). Fig. 107. P.
julvinota. Fig. 108. C. perumana. Fig. 109. P. cuprina. Fig. 110. P. eximia. Fig. 111. P. ibis. Fig. 112. S.
chlorogaster. Fig. 113. S. magellanica. Fig. 114. S. splendida. Fig. 115. S. versicolor.
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES 509

117

118
Fig. 116-118. SEM of puparia: anterior end (left), posterior end (right). All scales 5 mm. Fig. 116. C. c.
putoria. Fig. 117. C. macellaria. Fig. 118. C. boliviano.
510 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

Fig. 119. H. flavifacies. Fig. 120. P. fulvinota. Fig. 121. C. peruviana.

September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES
122
123
Fig. 122. P. cuprina. Fig. 123. S. chlorogaster.
for short periods of time. Duration of the pupal
stage was 5 days in both rearings. Egg-to-adult
duration was 13.6 days.
Calliphorinae
Calliphora peruviana. Adults were collected
from fish and liver bait 16 km west of San Ramon
(1,430 m, asynanthropic) and at Huasahuasi (2,750
m, hemisynanthropic). Adults were uncommon and
only two rearings were performed. Eggs were laid
singly or in clusters totaling from 75 to 175. The
incubation period was 15 h. Developmental times
for first and second instars were 31.5 and 26 h,
respectively. Durations of the third instar/prepu-
pa stage were 246 and 323 h, of which 137 and
270 h were spent as a prepupa in both rearings.
511
0.5 mm
The pupal stage lasted 12 days in both rearings.
Egg-to-adult interval was 26.9 days.
C. peruviana was the largest blow fly reared and
mature larvae often attained a length of 25 mm.
It is apparently a nomadic generalist found at gar-
bage dumps and coming to carrion in most of the
higher altitude habitats we studied. Jiron (1981)
reported that this fly was attracted to cadavers in
Costa Rica.
Phoenicia cuprina. Adults were captured in San
Ramon and Pto. Bermudez (500 m, both sites eu-
synanthropic). Four rearings were completed. Eggs
were laid in clusters of 50 to 80 on fish, and hatched
in 15.1 h. Subramanian (1980) reported 8 to 10 h
at 25.6°C and Melvin (1934), 12 h at 26.1°C. De-
velopmental times for first and second instars were
18.6 and 20.5 h, respectively; Subramanian (1980)
512 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

127 128 129

130 132
Fig. 124-132. SEM of egg plastrons (anterior third). Fig. 124. C. c. putoria. Fig. 125. C. macellaria. Fig.
126. C. boliviana. Fig. 127. H. ftavifacies. Fig. 128. H. hermanlenti. Fig. 129. P. fulvinota. Fig. 130. C.
peruviana. Fig. 131. P. cuprina. Fig. 132. P. eximia.
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES
3um 133 134
136
Fig. 133. P. ibis. Fig. 134. S. chlorogaster. Fig. 135. S. magellanica. Fig. 136. S. splendida. Fig. 137. S.
versicolor.
reported 14 and 18 h, respectively. Duration of
the third instar/prepupa stage was 102 h, com-
pared with 48 h reported by Subramanian (1980).
The discrepancy is in the prepupal stage: ours last-
ed 72 h, and his only 12 h. Pupal duration was 8.4
days; Subramanian (1980) recorded 5 days. Egg-
to-adult interval was 14.9 days.
Although P. cuprina is a primary myiasis fly in
Australia, there was no evidence of parasitic activ-
ity in San Ramon or Pto. Bermudez. According to
Hall (1948), Knipling found differences in larval
characters (anterior spiracles, tubercles, spines) be-
tween Australian P. cuprina and a nonparasitic
North American strain. It is noteworthy that lar-
vae of Peruvian P. cuprina are indistinguishable
from the latter. C. Sabrosky (personal communi-
cation) agrees that the two are different and that
the Australian form was recognized as a subspe-
513
135
cies, whereas the original and typical cuprina came
from China.
Phoenicia eximia. Adults were captured in
Playa Hermosa (1,000 m, eusynanthropic) and Pto.
Bermudez (500 m, eusynanthropic) on fish. Three
rearings were completed. Eggs were laid in clus-
ters of 30 to 70, and hatched in 12.7 h. Develop-
mental times for first and second instars were 13
and 19.7 h, respectively. The duration of the third
instar/prepupa stage was 144 h in all three rear-
ings. Late prepupae developed a pinkish color, es-
pecially obvious posteriad. Duration of the pupal
stage was 15 days for all rearings. Egg-to-adult
interval was 22.9 days.
P. eximia larvae have been reared or recovered
from: rotting fruit, decaying meat, and epithelial
detritus on the feet of birds (Hall 1948). This fly
was attracted to cadavers in Costa Rica (Jiron 1981).
514 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 77, no. 5

Phoenicia ibis. Adults were captured 16 km MEAN HOURLY TEMR(°C)

west of San Ramon (1,430 m, asynanthropic) on 18 19.6 20.6 22.9 2Z5 23.6 27.5 7*jb 18 22 16
fish and flowering shrubs. Two rearings were com- 170
pleted. Eggs were deposited in small clusters of 30 160
to 40 and hatched in 25.3 h, almost twice as long
150
as the other Phaenicia species. Developmental
times for first and second instars were 17.5 and 140
12.3 h, respectively. The duration of the third in- 130
star/prepupa stage was 279 h in both rearings; this
stage was the longest of the three Phaenicia species, rf 120
with the prepupal stage lasting 255 and 254 h. Z no
Prepupae remained in a quiescent semicontracted ~ 100
state at the bottom of the rearing containers, be-
coming active when examined or touched by other ~ 90
larvae. This quiescence suggested a diapause state, 80
but all of the larvae pupariated and eclosed nor-
70
mally. P. ibis prepupae had a slightly pinkish hue
similar to that of P. eximia. Duration of the pupal 60
stage was 13.8 days. Egg-to-adult duration was 27.7 50
days.
40

Toxotarsinae
Sarconesia chlorogaster. Adult flies were col-
lected in La Oroya (3,660 m, eusynanthropic) and
Jauja (3,550 m, eusynanthropic and hemisynan-
thropic) from fish and dead dogs. Three rearings
were completed. Eggs were laid in clusters of 50
to 150, and hatched in 29 h. Developmental times
for first and second instars were 32.3 and 54.5 h,
respectively. Duration of the third instar/prepupa
stage was 194.3 h and the pupal period lasted 11.3
days. Egg-to-adult interval was 24.3 days.
Copulation was observed inside rearing cages on
three separate occasions, lasting 1, 2, and 2.5 h,
with some females mating more than once. Mat-
ings of this duration and frequency are unchar-
acteristic of blow flies, but not unlike some sarco-
phagids which this species superficially resembles
(Dear 1979). S. chlorogaster is the largest of the
four Sarconesia reared and was fairly common
around market refuse in Jauja and La Oroya.
Sarconesia magellanica. Adults were collected
in Jauja (3,550 m, hemisynanthropic) on fish and
dead dogs. Two rearings were completed. Eggs
were laid in clusters of 100 to 130, and hatched in
25.5 h. Durations of first and second instars were
29.5 and 38 h, respectively. The third instar/pre-
pupa stage lasted 236.5 h and the pupal period,
12.5 days. Egg-to-adult duration was 26 days.
In the Sierra S. magellanica was second in
abundance to C. boliiriana. Adult male S. magel-
lanica exhibited a distinctive mating behavior.
They took flight from positions close to carrion (on
rocks or vegetation) and hovered head down and
flew in circles above females attracted to the car-
rion.
Sarconesia splendida. In 1979, two rearings
were completed in Jauja (3,550 m, eusynanthrop-
ic) from adults obtained in Morococha (4,500 m,
eusynanthropic). In 1980, another rearing with
adults from Morococha was performed in San Ra-
30
20
10 II 12 13 14 15 16 17
TIME (Hr)
Fig. 138. Diel activity of C. boliviano based on a
4-day collection from dog carrion in Montaro Valley.
Mean hourly temperature in the shade is given.
mon. Fish was used for all rearings. Eggs were
deposited in clusters of 20 to 150 inside fish heads.
Incubation time was 25.5 h in San Ramon and 35
h in Montaro Valley. In one case, viable eggs were
deposited on the underwing of a dead female. De-
velopmental time for first and second instars was
13 and 21 h in San Ramon and 20 and 22 h in
Montaro Valley. Duration of the third instar/pre-
pupa stage was 231 in San Ramon and 184 h in
Montaro Valley. The pupal period lasted 9 days in
San Ramon and 10 days in Montaro Valley. Be-
cause of lower ambient temperatures, egg-to-adult
duration in the Sierra was 24.6 days, compared
with 21.1 days in San Ramon.
S. splendida is the smallest of the four Sarco-
nesia reared and is apparently the dominant blow
fly at higher altitudes. In December 1979, 16 ten-
eral adults were taken emerging from a privy in
Morococha (0900 hours at 9.5°C).
Sarconesia versicolor. Adults were collected at
Morococha (4,500 m, eusynanthropic) and near
Tarma (3,800 m, asynanthropic) on fish. Three egg
batches were obtained, and two were reared in San
Ramon and one in Montaro Valley. Eggs were de-
posited in clusters of 50 on liver. Incubation time
was 15 h in San Ramon and 37 h in Montaro Val-
ley. Developmental times for first and second in-
stars were 25.5 and 15.5 h in San Ramon and 38
September 1984 GREENBERG AND SZYSKA: BIOLOGY OF PERUVIAN BLOW FLIES
O 300
O Compsomyiops boliviano
V) no
Ui
CJ I 50
z ing from 1000 to 1500 hours, which coincided
-or
AGE OF CARRION
Fig. 139. Fly succession on dog carrion in Montaro
Valley. (A) Blowflies, >96% C. boliviano, <4% S. ma-
gellanica and S. chlorogaster. (B) M. domestica. (C)
Near Hydrotea. (D) Fannia spp. (E) Sarcophagidae;
Anthomyiidae; and Muscidae near Hylemia, Apsil,
Phaonia, and Helina.
and 22 h in Montaro Valley. Duration of the third
instar/prepupa stage was 177 h in San Ramon and
179 h in Montaro Valley. Egg-to-adult duration
was 20.7 days in San Ramon and 22.5 days in
Montaro Valley. S. versicolor is second in abun-
dance to S. splendida at higher altitudes, and both
life cycles are very similar.
Diel Activity and Fly Succession
in Montaro Valley
According to Mani (1968), differences in sun and
shade temperatures are greater at higher altitudes
than at lower ones and rarefied air promotes evap-
oration and cooling in montane species. Although
this may be true at much higher altitudes, sun/
shade differences in Montaro Valley varied only
3°C on average (silver bulb temp). Sunlight is the
major factor governing fly activity at high alti-
tudes in Montaro Valley, at 3,550 m; flight was
greatly reduced or ceased whenever the sun was
obscured. Flight depends on wing muscle temper-
ature and it is advantageous to have adequate tho-
racic heat for quick takeoff under cool conditions.
According to Digby (1955) the thorax of a fly can
be thought of as a sphere, and appreciable heat
loss from a sphere in simulated flight in a directed
wind source occurs only at the anterior and pos-
terior areas. In a fly these conductive areas are
conveniently blocked by two other spheres, the
head and abdomen, connected by constricted, rel-
atively nonconductive pivots. This construction to-
gether with the absorptive dark ground color of
the thorax should minimize heat loss in Sarconesia
515
spp. and C. peruviana, the primary blow flies of
Montaro Valley and other highland regions. Digby
further postulates that a fine pile of hairs may in
some flies, in situations with low wind velocities,
stagnate a layer of air around the thorax providing
further insulation against heat loss.
C. boliviano,, like other highland flies, is helio-
philic, and was generally active only in direct sun-
shine above about 18°C. It had a broad activity
peak in Montaro Valley, during December, rang-
roughly with maximum daily temperatures (Fig.
138). Although flies appeared extremely attuned
to rapidly changing meteorological conditions, peak
temperatures and peak flight activity did not al-
ways correspond. Activity ceased at sunset, or ap-
proximately 1800 hours, regardless of ambient
temperature.
Figure 139 shows the abundance of blow flies
on dog carrion in Montaro Valley (Dec. 1979), in
relation to other muscoids, over a period of 5 days.
C. boliviano constituted 96% of all blow flies and
98% were gravid females. The small percentage
of males visiting carrion were observed feeding
but not copulating. Adult blow fly activity, includ-
ing oviposition, remained high the first 4 days,
then fell off rapidly on day 5. Oviposition by C.
boliviano, was not observed after day 4. Musca
domestica was the second most numerous fly until
day 4, when it surpassed C. boliviano.
Although C. boliviano predominated in pasture
areas in Montaro Valley, it constituted only 36%
of blow flies (n = 91) collected from fish and liver
baits inside an open courtyard in the same area
(Hacienda San Lorenzo) and it was never observed
in marketplaces in the city of Jauja. This suggests
that C. boliviano tends to avoid human dwellings.
Sarconesia magellanica
S. magellanica was the second most abundant
blow fly in Montaro Valley during December 1979.
On dog carcasses, S. magellanica represented about
3% (Fig. 139) of the total blow fly population and
on fish and liver about 6% (n = 91). It was found
inside an hacienda (San Lorenzo) and in a hotel in
the city of Tarma, strongly suggesting endophily.
The diel activity of S. magellanica on dog carrion
parallels that of C. boliviano except for lower
numbers and a higher proportion of males (ca.
25% compared with <4%). Males of this species
are presumably attracted to carrion for mating
purposes.
Affinities of Sarconesia
Boyes and Shewell (1975) suggest that the gen-
era Sarconesiopsis {^Sarconesia) and Phoenicia
may be more closely related than previously
 516 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
thought, based on similarities in the chromosome
complements (especially sex chromosomes) of Sar-
conesiopsis chilensis (=Sarconesia magellanica)
and Phaenicia sericata. They combine other
members of Dear's (1979) Toxotarsinae with the
Chrysomyinae on the same grounds, although the
only similarity we observed between larvae of Sar-
conesia and Chrysomyinae is an incomplete peri-
treme. This character, although used to separate
subfamilies, is not always reliable; for instance,
Calliphora iceila of New Zealand has an incom-
plete peritreme (Miller 1939).
On the other hand, suggestive similarities exist
between the Sarconesia and Calliphorinae we
studied. The cephalopharyngeal skeletons of all
three instars of the four Sarconesia resemble those
of Calliphora. The oral hooks of the first instar are
L-shaped in our specimens of Sarconesia, Phae-
nicia, and Calliphora. They differ from Chryso-
myinae which are more elongate and somewhat
S-shaped. In Sarconesia the oral hooks of the sec-
ond instar are stout and broadly curved, typical of
most Calliphora. Other shared larval characteris-
tics are the accessory oral sclerite of the third in-
star and single-pointed spines.
The eggs of both groups have similarities in the
outer plastron network. The Sarconesia resemble
Hinton's (1960) description of C. vicina and P.
sericata because the apices of the vertical struts
are expanded and frequently anastomose (Fig. 134-
137). In contrast, the outer network of the six
species of Chrysomyinae we examined was vir-
tually nonexistent except for the peglike vertical
struts.
Prepupae and pupae of Sarconesia and Calli-
phora generally developed more slowly than most
of the Chrysomyinae we reared. This could be re-
lated to unsuitable temperature and humidity,
crowding, and inadequate pupariation substrate.
A more likely explanation has to do with the pu-
parium and the site for pupariation in the two
subfamilies. The puparia of the Sarconesia,
Phaenicia and Calliphora we examined are rela-
tively smooth, oval, and have a thin, flexible cu-
ticle. Zdarek and Fraenkel (1972) describe seven
events in the transformation of the larval cuticle
into the hard, immobile puparium, including the
smoothing out of the cuticle due to longitudinal
shrinkage. This event, and possibly others, are ap-
parently incomplete in the Chrysomyinae we stud-
ied. Their puparia are rugose, hard, and inflexible
with incomplete retraction of both ends. Many, if
not all the sarcosaprophagous Chrysomyinae tend
to pupariate on, in, or close to the carrion, whereas
Calliphorinae tend to migrate before burrowing
into the ground (Waterhouse 1947, Norris 1959).
These authors suggest a good reason for this be-
havior. Tests on parasitic attack rates by Hyme-
noptera bear out the fact that the tougher puparia
of Chrysomyinae are more resistant. Therefore,
the Calliphorinae may require a longer prepupal
period to migrate from carrion in order to burrow
into the ground. Once interred, the pupae can re-
Vol. 77, no. 5
main longer in this stage, relatively free from par-
asites and predators. Prepupae of several species
of Lucilia and Calliphora dispersed 12 to 15 feet
or more from the center of a sheep carcass (Cragg
1955); under unfavorable conditions of hard ground
these larvae traveled 80 to 100 feet (Green 1951).
Our rearing methods did not permit us to test this
hypothesis on Sarconesia or the other flies, but the
evidence of morphological similarities in the eggs,
larvae, and puparia of Sarconesia and Calliphor-
inae warrant extensive sampling of other species.
If the evidence holds up, it may be necessary to
reconsider the affinities of Sarconesia.
Acknowledgment
We thank Donald L. Baumgartner for his cooperation
and help and William E. Dale, Univ. Nacional Agraria
La Molina, Lima, Peru, for giving generously of his time
and facilities. In 1979, some of the life cycles were com-
pleted in his laboratory, thanks to his effort. Curtis W.
Sabrosky, Raymond J. Gagne, and Norman Woodley,
Systematic Entomology, U.S. Dept. of Agriculture, read
the manuscript and contributed helpful suggestions. Re-
search supported by NSF Grant 79-05282, Div. of In-
ternational Programs, to B. G. Publication costs sup-
ported by contract DAMD17-84-M-4012, U.S. Army
Med. Res. and Dev. Command.
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Received for publication 22 February 1984; accepted
23 April 1984.