The JapanSocietyJapan Society of Medical Entomology and Zoology

(Jpn. J. Sanit. Zool. Vol. 38 Ne. 3 p. 187-1951987]

Biologieal studies of (Diptera) from molluscnon-museoidcarrion in Southeast Asiabred

R. A. BEAvER

School of Pure and Applied Sciences, The University of the Seuth Pacific,

Box 1168, Suva, Fiji

(Received: September 8, 1986)

Key werds: snail carrion, Diptera, WestMalaysia,Thailand,

Abstract: Twenty-one species of non-muscoid Diptera (Psychodidae, Phori- dae, Sphaeroceridae, , Milichiidae, ) were bred from dead snails, Achatina fulica, in West Malaysia, and four species from the same habitat in Northem Thailand. Information is given on the breeding sites, pattern of occurrence, development time, hymenopteran parasitoids, and medical and

veterinary importance of the species.

didae), rather than the numbers of adults INTRoDUCTION bred, and thus include those individuals that died in the puparium, or were parasitised In the first paper of this series (Beaver, by Hymenoptera. Nomenclature of the 1986a), I dealt with 12 species of muscoid Diptera follows Delfinado and Hardy (1973- flies bred from dead snails, chiefly the giant 1977). African snai1, Achatina fulica Bowdich, in Thailand and West Malaysia. In this paper, BIOLOGy OF TIIE DIpTERA I conslder 21 species of non-muscoid flies of the families Psychodidae, Phoridae, Sphaero- Ps)Jchodidae ceridae, Ephydridae, Milichiidae and Chloro- Psychoda alternata SaylPsychoda sp.: pidae, bred from the $ame series of snails. There seem to be few records of Psychodidae Details of the Jocalities and of the experimen- from dead snails, although two species of tal procedures used are given by Beaver PhilosePedon, P. Phalaenoides (L.) and P. (1986a). humeralis (Meigen), are regular members of Five experiments were carried eut in X･Vest the dipteran community breeding in dead Malaysia, on a site in the campus of Uni- snails in Europe (Beaver, 1972; Keilin, 1919; versiti Sains Malaysia on Pulau Penang, using Schmitz, 1917). Psychoda alternata is a from 13-tl5 dead Achatina fulica. Two cosmopolitan species breeding in a wide range experiments were carried out in Thailand of habitats in whieh both moisture and a on the campus of Chiang Mai University supply of organic material are provided, such using A. fulica, and in one experiment as sewage beds, drains, etc. It may be in- ViviPara 1`aPonica Mart., and Indoplanorbis volved in the mechanical transmission of sp. also. Records are frorn dead Achatina parasitic nematodes (Tod et al. 1971), and fulica unless otherwise indicated. It should lam,ae have been involved in human enteric be noted that the numbers of individuals myiasis (Harwood and James, 1979). P. recorded for each species refer to the numbers alternata was bred in Malaysia from a single of puparia found (except for the Psycho- snail which produced 18 adults, This number 187

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cou]d not be checked against the number of scoPus have similar habits to those of pupae because the pupal skin was fragile Rsychoda species (Bohart and Gressitt, 1951). and easily broken up during extraction, unlike Adults were bred from dead Achatina fulica the harder and stronger puparial cases of in both Thailand and West Malaysia. In other dipteran families considered in this Malaysia, the species occurred more frequent- paper. Adults emerged from 10-12 days after ly in the snails than the other two psychodids the snail was exposed (Table 2). P. alternata bred (Table 1), and in larger numbers per is known to have a short development period, snail on average (Table 2). Its develop- which may be as little as 7 days when mental period is certainly longer than that temperatures are high (Ameen and Huq, of Psychoda alternata (Table 2). 1973). Psychoda sp. was bred from a single snail Phoridae in the same experiment as P. alternata, and Conicera (Tritoconicera) formosensis again 18 adults were reared. The develop- Brues: Species of Conicera breed in carrion mental period appears to be longer than that and other dead organic matter (Schmitz, is well-known of P. alternata (Table 2), but this could be 1953), and C. tibialis Schmitz "coMn an effect ef different dates of oviposition for as the fiy," able to maintain itself the snails were exposed to attack for a week. over many successive generations on human Telmatoscopus sp.: Species of Telmato- cerpses inside cofins (Oldroyd, 1964). C.

Table 1 Percentage of Achatina fulicaattacked by various species in West Malaysia.

Percentageof snails attacked Species 1* 2* 3* 4* 5*

Psychoda alternata 5515

Psychoda sp.

TelmatoscoPus sp. 4 7

Conicera formosensis 4

DehrniPnora cornuta 15153115

Gymneptera orientalis

Megaselia curtineura 86367 40 223225579272

Megaselia sp. D 29

Megaselia sp. G 4

PuticiPhora sp. F 8 5 18

PuliciPhora sp. L 16 367 co SPiniphora genitalis 10

Coproica hirtula 38 1oo 69 7 Atlotrichoma alium 5 Allotrichoma livens 15100 60 7 265

Discomyza maculipennis 95 14 69

DesmemetoPa sp. 5 9

Milichia sp. 10 2 1062 bilineatus 815 40 20

SiPhuneutina minima 75 89 7 Siphunculina sp. 5

Unidentified sp. 5 4

45 Totar No, of snails 1318.iv.752S.iv,75 2029.iv.756,v.75 28 coi6.vi.7723.vi.77 7.iL76 5.v.76 Date of exposure

Date of collection 8-12,iL76 6-12.v.76

* Experiment No.

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Tab]e 2 Number of individuals pupated per attacked snail (mean and range) and time in days (mean and range) between exposure of the snail and emergence of the adult flies for various fly species in NNTest Malaysia.

Species No.of individuals No, of days

Psychoda alternata 181833.0 11,7 (10-12) Psychoda sp. 18,2 (18-19)

TelmatoscoPus sp. (1-100) 23.4 (21-28) Conicera formosensis 27 27.9 (26-32) DohrniPhora cornuta 8.5 (1- 16> IL7 (11-13) Gymnoptera orientalis 6.0 {2- 10) 28.9 (21-30>

Megaselia curtineura 21.0 (1-228) 16.9 (14--24)

Megaselia sp. D 15.8 (1- 96) 22,9 (15-32)

Megaselia sp. G 4.8 (1- 15) 17,l (14-20) PuliciPhora sp. F 12.8 (1-61(-887)) 21.5 (17-30) PuliciPhora sp. L 51,7

Coproica hirtula 47.6 (1-366) 12.6 ( 9-20) Allotrichoma alium 10.2 (2- 21) 10.6 ( 9-12)

Allotrichotna livens 22.4 (1- 85) 9,5 ( 7-13)

Discomyza maculiPennis 14.8 (1- 82) 20.5 (14-38) DesmometoPa sp, 9.4 (1- 21) (?9-12) Milichia sp. 4.0 (1- 8) (?9-12)

Hippelates biiineatus 7.4 (1- S2) 27.3 (19-35)

SiPhunculina mininza 29,2 (1-212) 17.5 <10-35)

SiPhunculina sp. 2.2 (1- 5) --33.2 S.2.7-g7).-

formosensis is known from Taiwan and West generation within the same snail, and have Malaysia, but nothing is recorded of its been excluded from the calculation of the biology. I bred the species from only a mean value in Table 2. Bohart and Gressitt single Achatina fulica in Malaysia. The snail (1951) give a developmental period of 15+ day, and it is days. had been exposed for only 1 ' of development Gymnoptera onentalis Meijere): G. clear that the mean time -bed (de

(27.9 days) is longer than for most other orientalis was desc- from Java, and does species bred from the snails (Table 2). not seem to have been recorded before from DohrniPhora cornuta (Bigot): This is a Malaysia. Noth ing has been recorded of its cosmopolitan species, bred from a very wide biology, aithough the reJated species, G.

range of decaying and matter, molluscovora was associated with plant (Bohart) J including dead molluscs, dead in decaying molluscs in Guam (Bohart and Sarracenia pitcher plants, and sewage beds Gressitt, 1951). These authors suggest that (Bohart and Gressitt, 1951; Kloter et al. G. moZluscovora may help the spread and 1977). KIQter et al. C1977) note that the development of toxin-producing micro- species is a facultative predator of Psychoda organisms on shellfish. I bred the species in alternate in sewage beds, but the two species Malaysia from only two snails. These pro- did not occur together in these experiments. duced 2 and 10 adults respectively. Bohart I bred D. cornuta in Malaysia from only and Gressitt (1951) give a development two snails, which produced 1 and 16 adults period for G. moUuscovora of 16+ days.

respectively. Adults emerged from 11-13 and The littleevidence available for G. orientalis 27-32 days after the snails were exposed, suggests a longer period, at Ieast in the The later emergences may have been a secend conditions of the experiments, The average

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time (28.9 days) between exposure of the Achatina fulica in West Malaysia, sometimes snails and emergence of the adults was Ionger in large numbers per snail. than for almost any other species bred PulioiPhora sp. F was bred from 20 snails (Table 2). in four experiments (Table 1). Most of these Megaselia curtineura (Brues) : This species snails produced moderate numbers of indivi- has a distribution wide from Africa through duals (Table 2), but one snail produced 887 the Orienta] region to Hawaii. It has been individuals. This snail had been protected reared before from Achatina fulica, and also from oviposition by the Iarger muscoid flies, from carrion and as a laboratory cul- and from predation by ants. Adults emerged ture medium contaminant (Robinson, 1971). after 18-19 days from snails exposed for In Malaysia, I bred the species frorn 47 snails 1 day. in four experiments (Table 1). In Experi- PuliciPhora sp. L was bred from 30 snails ment 3, it was one of the two most abundant in two experiments (Table 1). This species species present, with up to 228 individuals also occurred in large numbers in certain in a single snail (Table 2). A small number snails (Table 2). The development period is of larvae were parasitised by the encyrtid, similar to that of Puliciphora sp. F. Exoristobia philippinensis Ashmead. This Spiniphora genitalis Schmitz: This species species attacks third instar larvae, but com- was described from specimens bred from pletes development within the puparium. It dead Achatina fulica in West Malaysia is normally a gregarious parasite when it (Schmitz, 1940), and is also known from attacks larger flies (R. A. Beaver, unpub- the same snail in Java (Colyer, 1955). All lished), but here a single parasitoid adult species of SPiniphora develop in dead snails, emerged from each puparium. and appear to be confined to this habitat Megaselia spp,: Records of the breeding for breeding (Schmitz, 1917, 1941). Large sites of Megaselia species are summarised land snails are preferred. The larvae pupate by Robinson (1971). They include frequent within the snail shell, usually in rows side records from dead snails and slugs. Further by side, the anterior ends pointing in the records from this habitat have been added same direction away from the axis of the by Beaver (1972) and Kneidel (1984). There shell. The posterior spiracles are elevated are occasional records of myiasis caused by on a tubular projection. Megaselia species in man and cattle (sum- I bred S. genitalis in West Malaysia from marised by Disney and Kurahashi, 1978). A. fulica, and in Thailand from this snail, Megaselia sp. D was bred from 25 snails in ViviPara iaPonica Mart. and Indoplanorbis West Malaysia, and up to 96 individuals sp. It was one of the most abundant species were found in a snail. Some larvae were in the snails (Tables 1, 2). When the larger parasitised by Exoristobia PhiliPPinensis. As muscoid flies were prevented from ovipositing with M. curtineura, a single parasitoid in the snails, S. genitalis usually dominated emerged from a puparium. the community in temis of both numbers Megaselia sp, G was bred from Achatina and biomass. Adults emerged after 16-27 fulica in both Thailand and West Malaysia. days from snails exposed for 2 days, but In the latter country, it occurred in three none of the snails exposed for a single day experiments, but in small numbers (Tables 1, were attacked. No parasitoids were bred. 2). Adults emerged after 16-20 days from snails exposed for a single day. No evidence SPhaeroceridae of parasitisation was found. Coproica hirtula (Rondani): There are PuliciPhora spp.: Most members of the a number of records of Sphaeroceridae bred genus are probably general scavengers, and from dead snails (e.g. Beaver, 1972; Deeming several species have been bred from dead and Knutson, 1966; Richards, 1930; Schmitz, snails (Beaver, 1986bi Bohart and Gressitt, 1917), but all the species recorded are Iikely 1951; Kneidel, 1984). The females are wing- te breed in other dead organic matter as less and may be carried to the breeding site well. CoProica hirtula is a cosmopelitan by the males while in coPula. Two species species found in a variety of habitats, usually of these tiny flies were bred from dead in associatlon with dung, rotting plant mate-

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rial or carrion. It was one of the species dead snails. A. alium was bred in small

most commonly bred from dead Achatina numbers in two experiments in West Malay- fulica in l'Vest Malaysia, occurring in all sia; A. Iivens occurred more frequently experiments and 87 snails (Table 1), In (Table 1) and in larger numbers (Table 2). experiment 2, up to 336 individuals were Bohart and Gressitt (1951) give a develop- obtained from a single snail; in the other ment time for Allotrichoma sp. of 5+ days. experiments up to 96. Almost all adults I found the first adults of A. Iivens emerging' emerged between 9 and 20 days after the only 7 days after the snails were exposed snails were exposed, with peak numbers (Table 2). A. aliutn may develop a little emerging on days 11-13 (Fig. 1). A few more slowiy (Table 2), but the development stragglers (or possibly a few individuals of tirne of both species is clearly very short, a second generation) emerged from day 22- A single male Kleidotonza sp. was bred from 30 (Table 2). The eucoilid parasitoid, a puparium of A. Iivens. Kleidotoma sp., was bred from some puparia Discomyia maculipennis (Wiedemann): in one experiment. Most breeding records of Discomyza species

in the Palearctic, Afrotrepical and Oriental

EPhptdridae regions are from dead snails, although the Allotrichoma atium Cresson!A. tivens Cres- fiies are clearly able te breed in other carrion son: Little is known of these small ephydrid (Beaver, 1986bs Rohart and Gressitt, 1951; fiies. A. alium is distributed in Southeast Disney, 1970, 1973; Keilin, 1919). D. mac-

Asia, Papua New Guinea and the Pacific uliPennis has a wide distribution in the Islands; A. Iivens has only been recorded Oriental region and the Pacific Islands. I before from India and Taiwan (Cogan and bred it from dead Achatina fulica in Thai- Wirth, 1977) and not from West Malaysia. land and West Malaysia, and in Thailand Bohart and Gressitt (1951) found AUotri- also from P'ivipara juponica and IndoPlanor- choma sp. breeding in pig droppings in bis sp. In NVest Malaysia, it occurred in the Guam. Cogan and Wirth (1977) indicate majority of snails (Table 1), sometimes i'n that the larvae of Allotrichoma are scaven- quite large numbers (Table 2). The larvae gers in carrion and faeces. There appear to pupate within the snail shell, often in the be no previous records ef either species from inner whorls, and often Iined up in parallel

4

eX3o9ptEoi-e2'egece.e

l8

Days after expesure

Fig, 1 Pattern of emergence of five of the more abundant species of flies

bred from mollusc carrion in Malaysia. e Coproica hirtula, O Megaselia curtineura, A Diseomyza maculipennis, V SPiniphora genitalis. - HipPelates bilineatus,

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rows like the puparia of SpiniPhora genitalis snail. (see above). As in other species of Disco- myia, the posterior spiracles of the puparium Chloropidae are raised above the bedy, keeping them HiPPelates bilineatus de Meijere: This above any fluid remaining in the snail. species is widely distributed in the Oriental Adults emerged after 14-19 days from region and on Pacific islands, but little is snails exposed for 1 or 2 days, but the known of its biology. Bohart and Gressitt development period may semetimes be (1951) found larvae in decaying molluscs, prolonged well beyond this (Fig. 1). This rotting coconuts and pig faeces. They suggest prolongation was more evident in snails that the species may be of irnportance as a containing Iarge numbers of individuals of carrier of faecal pathogens. The biology of D. maculipennis, and may be a consequence some of the North American species of of larval competition. Three species of HiPPelates has been studied in more detail, parasitoid, Spalangia endius Walker (Ptero- and this work is summarised by Greenberg malidae), Exoristobia PhiliPPinensis (Encyr- ('1973). One species, Hippelates dissidens tidae) and an unidentified pteromalid were (Tucker) has been bred from dead snails, bred from puparia. Drlymaeus dormani (Binney), in FIorida (Muma, 1954, quoted in Mead, 1961). Milichiidae Other species can be of medical importance Desmometopa sp.IMilichia sp.: Milichiid (Harwood and James, 1979). larvae are saprophagous or coprophagous, I bred H. bilineatus in four of the five and have been bred from cattle dung, swiftlet experiments in West Malaysia (Table 1), guano, dead fish, decaying plants, etc. (Bo- but in fairly small numbers per snail (Table hart and Gressitt, 1951; Sabrosky, 1977). 2). The species occurred only once in snails

The adults of certain species of Desmometopa that had been exposed for less than 5 days, are known to have a commensal re]ationship and the species seems not to be one of the with larger predatory insects, being carried early colonisers. Development seems to be around by the Iatter, and feeding on the relatively slow. Adults emerged from 19-35 body fluid of their prey. Both Desmometopa days after the snails were exposed with a sp. and Milichia sp. were bred in West mean of 27.3 days (Table 2, Fig. 1). Malaysia from a small number of snails SiPhuneulina minima (de Meijere)ISi- (Table 1) and occurred in only small num- Phunculina sp.: Like the species of HiPPe- bers within those snails (Table 2). Both lates, certain species of Siphunculina (par- species sometimes pupated inside the snail ticularly S. funicola (de Meijere)) are of shell, and sometimes in the sand. In one medical importance as possible vectors of snail, several puparia of DesmometoPa sp. bacterial conjunctivitis and yaws (Harwood were found inside empty puparia of the and James, 1979). Like S. funicola, S. phorid, SPiniPhora genitalis, which were in- minima is reported to suck blood (Becker, side the snail shell. Three of the phorid 1911), but has not been specifically impli- puparia contained two puparia of Desmome- cated as a disease carrier, nor has its biology topa sp., others only one. Since S. genitalis been studied. It may resemble S. funicola emerged after a minimum of 15 days, these in its breeding sites, which are characterised Desmometopa puparia probably represented by organic pollution (Antonipulle, 1957; a second generation, The development time Greenberg, 1973). Both S. funicola and S. of the two milichiid species in the snails is minima occurred commonly near the experi-

not certain, but appears to have been 9-12 mental site in West Malaysia, but only the days. Bohart and Gressitt (1951) give a latter species was bred from the dead snails, development period of 8+ days for Desmo- together with an undetermined species of the metoPa sp. on Guam. In one snail, some genus. puparia of DesmometoPa sp, were parasitised, S. minima occurred in four of the experi- probably by Exoristobia philiPPinensis, which ments, and in over 60% of the snails in also attacked the sarcophagid, Parasarco- three of them (Table 1). Up to 212 indi- phaga dux (Thomson) breeding in the same viduals occurred in a single snail (Table 2).

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The nermal pupation site was in the sand (up to 1 week). It probably results from for food between the flylarvae in the dishes, but puparia were occasionally competitiDn found within the snails, sometimes attached in the snail. Larvae of a particular species to puparia of Discomyza maculiPennis. which emerge from eggs laid towards the are te SiPhunculina sp. occurred in enly four snails, end of the exposure period likely have

with small numbers of individuals per snail lessfood available to them than larvaewhich access to the food. There may (Table 2). Adults of S. minima emerged have early after 10-12 days from snails exposed for also be a decline in foed quality. These 1 day, but in other snails, some stragglers factors may delay the development of the emerged up to 35 days after exposure. Adults immature stages, as is known to occur in

of Siphuncntina sp. emerged a minimum of other cyclorrhaphan dipteran larvae compet- and 1983; 27 days after the snails were exposed (Table ing for food (Baxter Morrison, 2). A single female parasitoid, Kleidotoma Peters and Barbosa, 1977). The importance of the small non-muscoid sp., was bred from a puparium of Siphun- culina minima. flies that breed in dead organic matter as passive vectors of diseases or parasites is DIscussloN probably often under-estimated. There are several reasons for this. Because they are Their taxon- The majority of the species bred from the small, they are often overlooked. dead snails are not confined to this habitat omy is generally less well-know'n than that for breeding, but breed in other carrion, and of the muscoid fiies, so that they are more often in a wide range of decaying organic dirncult to identify accurately. There is a lack ef biological information, and of vector- matter, both plant and animal. Thls might oriented studies of these flies. Of the species be expected given the patchy distribution of the species carrion of any kind in space and time bred in the present experiments, two species, of Hippelates and Siphunculina (Beaver, 1984) , However, SPini- probably have the importance. phora genitalis and Discom),xa maculiPennis, greatest potential have been incriminated are particularly associated with dead snails. Species of both genera Both occurred commonly in the snails, and as passive vectors of conjunctivitis and yaws often in relatively large numbers per snail. (Mattingly et al. 1973), and the eye- Numbers of S. genitalis were particularly frequenting and sore-frequenting habits of make high in snails which were protected from SiPhunculina in Southeast Asia trans- oviposition or larviposition by the larger mission of these diseases a relatively easy muscoid flies, and frem which ants were process, excluded. Numbers of D. maculipennis were higher where ant predation occurred, suggest- ACKNOWLEDGEMENTS ing differential predation of the two species I am most grateful to Mr. B. H. Cogan, Dr. P. S, by the ants (R, A. Beaver, unpublished). Cransten, Dr, R. H. L. Disney, Dr, K, Kanmiya The percentages of flies emerging at vari- and Mr, K. G, V. Smith for identifying the ous times after the snails were exposed are Diptera, and te Dr. Z. BouEek, Dr. J. S. Neyes shown in Fig. 1 for five of the commenest and Mr. J. Quinlan for identifying the Hymen- It isnoticeable species bred from the snails. opteran parasitoids. Faeilities for the experiments that species with shorter mean development were provided by the Scheol of Biological Sciences, times (Coproica hirtula. Megaselia curti- Universiti Sains Malaysia, and the Department of Biology, Chiang Mai University. neura) tend to have a more strongly peaked distribution of emergence times than slower- REFERENCES developing species. The curves of all species are significantly skewed to the (p

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(Biol, Sci.), (N.S,), 1: t-5. tomology in Human and Animal Health, 7th ed., Baxter, J. A. and P. E. Morrison (1983): Dynam- 548pp., Macmillan PubL ao., New York. ics of growth modified by larval population Keilin, D. (1919): On the life history and larval density in the fieshfiy Sarcophaga buUata. Can, anatomy of Melinda cognata Meigen (Diptera, J. Zool. 61: 512-517. Calliphorinae) parasitic in the snail Helicella Beaver, R. A. (1972): Ecological studies on (Heliomanes) virgata DaCosta, with an account Diptera breeding in dead snails. 1. Biology of of the other Diptera living upon molluscs. the species found in CePaea nerrtoralis (L.), Parasitology, 11: 43[V-455. Entomologist, 105: 41-52. KIoter, K. O,, L, R. Penner and W. J. Widner Beaver, R. A. (1984); Insect exploitation ef (1977): Interactions between the larvae of epherneral habitats. S, Pac. J, Nat. Sci., 6: Psychoda alternata (Dipt., Psychodidae) and 3.47. Dohrniphora cornzata (Dipt. Pheridae) in a Beaver, R. A, (1986a): Biological studies of trickling filter sewage bed. Ann. Entomol. Soc. muscoid flies (Diptera) breeding in rnelluse Am,, 70: 775-781. carrion in Southeast Asia. IPn. 1. Sanit. Zool., Kneidel, K. A. (1984): Infiuence of carcass 37: 205-211. taxon and size on species cemposition of carrion- Beaver, R. A. (1986b): Some Diptera and their breeding Diptera. Am. Midl. Nat. 111: 57-63. parasitoids bred from dead snails in Zambia. Mattingly, P. F., R. W. Crosskey and K. G. V. Entomol. Mon. Mag. 122: 195-199. Smith (1973) : Summary of Vectors. Becker, T. (1911): Chloropidae. Eine mono- In: Inseets and Other of Medical graphische Studie. III. Teil, Die indo-australi- ImPortance (ed. Smith, K. G. V.), pp. 497- sche Region. Ann. Mus. Nat. Hung. 9: 35-170. 532, British Museum (N,H.), London, Bohart, G. E. and J. L, Gressitt (1951): Filth- Mead, A. R. (1961): The Giant African Snail. inhabiting flies of Guam. Butl. B.P. BishoP Mus., A Problem in Economic Malacology, 257pp., 2ou; 1-152. Univ, Chicago Press, Chicago. Cogan, B. H. and W. W. Wirth (1977): Family Muma, H. H. (1954) : Predators and parasites of Ephydridae. In; A Catalog of the Diptera of the citrus tree snall. Citrus Mag., 16: 8-9. the Oriental Region (ed,, Delfinado, M. D. and Oldroyd, H. (1964): The Aratural History of D. E. Hardy), Vol. 3, pp, 321-339, Univ. Press Flies, 324pp., Weidenfeld and Nicolson, Lon- Hawaii, Honolulu. don.Peters, Colyer, C, N, (1955) : A new species of Spiniphora T. M. and P. Barbosa (1977): Influence (Dipt., Pheridae) from Ceylon; notes on Spini- of population density on size, fecundity and phora genitalis Schmitz. Entomel. Mon. Mag. development rate of insects in culture. Annu. 91: 48-50. Rev. Entomol. 22: 431-45D. Deeming, J. C. and L. V. Knutson ('1966): Richards, O. W. (1930): The British species of Ecological notes on some Sphaeroceridae reared Sphaeroceridae (Borboridae, Diptera). Proc. frorn snails, and a description of the puparium Zool. Soc. Lond. 1930: 261-345. of Coprompea (Apterina) pedestris Meigen. Robinsen, W. H. (1971): Old and new biologies Proc, Entomol. Soc. Wash. 68: 108-112. of Megaselia species (Dipt., Phoridae). Stud. Delfinado, M. D. and D, E, Hardy (197S-1977): Entomol. 14: 321-348. A Catalog of the Diptera of the Orientat Region, Sabrosky, C. W, (1977): Family Milichiidae. Vels. 1-3, 628 pp., 469 pp,, 864 pp. Univ. Press In: A Catalog of the Diptera ef the Oriental Hawaii, Honolulu. Region (ed. Delfinade, M. D. and D. E. Disney, R, H, L. (1970): A note on Discomyza Hardy), VoL 3, pp. 27e-274, Univ. Press Ha- similis Lamb (Dipt., Ephydridae) and other waii, Honolulu. snails reared from dead snails in Cameroon. Schmitz, H. (1917); Biologische Beziehungen Entomol, Mon, Mag. 105 (1969): 250-251. zwischen Dipteren und Schneeken. Biol. Zen- Disney, R. H. L. (197S): A note en some filth- tratbl,, 37: 24-t+S. inhabiting fiies of Cameroon. Entomot. Mon. Schmitz, H. (1940): Eine neue ostasiatische Mag. 108 (1972): 212-21S. Spiniphora. Natuurhist. Maandbl. 29: 78-79. Disney, R, H. L. and H. Kurahashi (1978): A Sclmitz, H. (1941): Phoridae. In: Die Fliegen case of urogenital myiasis caused by a species der Paldarktischen Region (ed., Lindner, E.), of Megaselia (Diptera: Phoridae). J. Med. PL 37. LieL 141, pp. 65-128, E. Schweizerbart, Entomol. 14: 717. Stuttgart, Greenberg, B. (1973): Flies and Disease. Vol. 2. Schmitz, H. {1953): Phoridae. In: Die Fliegen Biology and Disease Transmission, 447pp., der Paldiarktischen Region (ed., Lindner, E.), Princeton Univ. Press, Princeton) New Jersey. Pt. 37, LieL 171, pp. 273-320, E. Schweizerbart, Harwood, R. F. and M. T. James (1979): En- Stuttgart.

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