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A new genus and three new speciesof Phoridae (Diptera) parasitizing ants(Hymenoptera) in SulawesiR.H.L. Disney aa Field Studies Council Research Fellow, Department of Zoology,Cambridge University, Downing Street, Cambridge, CB2 3EJ, UKPublished online: 17 Feb 2007.

To cite this article: R.H.L. Disney (1986) A new genus and three new species of Phoridae (Diptera)parasitizing ants (Hymenoptera) in Sulawesi, Journal of Natural History, 20:4, 777-787, DOI:10.1080/00222938600770551

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JOURNAL OF NATURAL HISTORY, 1986, 20, 777--787

A new genus and three new species of Phoridae (Diptera) parasitizing ants (Hymenoptera) in Sulawesi

R. H. L. DISNEY

Field Studies Council Research Fellow, Department of Zoology, Cambridge University, Downing Street, Cambridge CB2 3E J, UK

(Accepted November 1985)

Iridophora clarki gen. nov., sp. nov., Megaselia sembeli sp. nov. and M. kodongi sp. nov. are described. The females ofL clarki attack the workers of the ant lridomyrmex cordatus (Smith). The same ant species is also attacked by M. sembeli, and the morphology of this species indicates that it is close to P. beirne Brues, the type- species of the genus Plastophora. The loss of the type-material of the latter species prevented synonomy of the genus when it was re-evaluated (Disney, 1978). The detailed examination of M. sembeli now indicates that the genus Plastophora is a synonym of Megaselia (syn. nov.). A consequence is that P. dubitata Brues, 1935 becomes a homonym of M. dubitata (Malloch, 1912) and the replacement name M. bruesi nom. nov. is proposed. The females of M. kodongi attack workers of a species of Pheidologeton which forages on the forest floor. The habit of Iridomyrmex cordatus in using covered ways up trees appears to be related to its extreme vulnerability to attack by these two phorides.

Introduction From 6 January to 27 March I studied Phoridae (scuttle flies) in the Toraut Forest,

Dumoga-Bone National Park, Sulawesi-Utara as a participant in the Royal Entom- ological Society of London's Project Wallace expedition. As Phoridae are well known to parasitize ants (e.g. Borgmeier, 1963, Disney, 1982, 1983), a particular effort was made to collect scuttle flies observed attacking ants. These previous studies have shown that the parasitoid species either oviposit into the ant's head or its gaster. The purpose of the present paper is to describe three such parasitoid species.

In recording the deposition of type-material 'in collection of author ' below I would point out that this collection will ultimately be deposited in a museum. While it is in continual use for research purposes, it remains in my care (currently in the Cambridge University Zoology Museum).

Iridophora gen. nov.

Type-species. Iridophora clarki sp. nov. Etymology. The name is derived from that of the ant host of the type-species. Head. Frons with three ocelli and median furrow. 4-4-4 bristles present, but

somewhat weak. Antials and pre-ocellars widely separated. No supra-antennals and

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hairs confined to lateral margins of frons. Third antennal segment elongated, tapering, and with a short, two-segmented arista in the male. No arista in female.

Thorax. Scutum sparsely haired. Mesopleuron divided and bare. Abdomen. Tergites 1-6 normally developed in both sexes. Female with elongated

ovipositor with fine pointed tip. Articulation between the basal segment and the rest allows the latter to fold back dorsally.

Wings. Veins 1 and 3 closely approximated. Vein 2 (inner branch of fork of vein 3) missing. Sc weak. Costa with a single row of hairs.

Legs. All tibiae lack hair palisades and bristles. All tarsi with fifth segment elongated and strongly tapered.

Affinities. There is considerable taxonomic confusion among the phorid genera which lack bristles on the tibiae and have a divided mesopleuron. However, the elongated third antennal segment and tapered last tarsal segment will immediately distinguish Iridophora from more than 150 of these genera. The reduced terminal arista or no arista, the absence of vein 2 and the presence of 4-4-4 bristles on the frons place this genus close to Tubicera. However, the latter has hair palisades on the hind tibiae and normal last tarsal segments. Cataclinusa also has normal last tarsal segments and lacks a median furrow on the frons. Aristocerina has a median furrow but has normal last tarsal segments, veins 1 and 3 well-separated, and well-developed three-jointed arista. The genera Microselia, Lepta and Dacnophora all possess last tarsal segments similar to those of Iridophora. However, these all possess hair palisades on their hind tibiae and a pre-apical arista on a third antennal segment that is not elongated. Furthermore, in most species vein 2 is clearly developed.

Iridophora elarki sp. nov.

(Figs 1, 2)

Deposition of types. HOLOTYPE (9)- Sulawesi-Utara, Dumoga-Bone National Park, Toraut Forest, 17.i.1985, coll. R. H. L. Disney, deposited in collection of author. PARATYPES (219, 10c~) same data and depository as HOLOTYPE except dates various i- iii.1985 and 2d', 29 deposited in Museum Zoologicum Bogoriense, Bogor, Java.

Etymology. The species is named after Paratrooper Andy ('Nobby') Clark, who was an indefatigable member of the Services support team on Project Wallace.

Female Head. Frons brown with 7-12 marginal hairs each side. Antenna (Fig. l(c))

brownish. Palps pale basaUy and brownish apically, with a single apical bristle and a weaker pre-apical lateral bristle; labella and labrum pale.

Thorax. Brownish but lower parts of pleura pale. Propleuron with a single hair posteroventrally. A single hair and a bristle on humerus. Two bristles on notopleuron and a pre-alar bristle. A single pair of pre-scuteUar dorsocentrals. Scutellum with a posterior pair of fine bristles and an anterior pair of hairs.

Abdomen. Tergites 1-6 brown with sparse minute hairs only. Venter brownish-grey. Ovipositor as Fig. 1 (a).

Mounts in both Berlese Fluid and in glycerol were examined by ultraviolet light (about 420 mp) and produced a blue fluorescence in the region of the articulation between the basal and distal sections of the ovipositor. A small patch of blue

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New genus and three new species of Phor idae 779

°%°°

; ~ - - ~ ~ ~ ' ! i . . . ~ ' . - ~ L - " " ' ' " - -

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....... / ( a ) ' ' .... ( )

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(c) , o.,

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FIG. 1. lridophora clarki gen. nov., sp nov. 9. (a) Ovipositor in side view; (b) terminal segments of front tarsus; (c) antenna; (d) terminal segments of middle leg; (e) tarsus of hind leg; (f) right wing. Scale bars = 0-1 mm.

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780 R. H. L. Disney

2(a) ,

3(a)

I !

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3(b) 2(c) . . . . . . ~ . ' i I

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FIG. 2. Iridophora clarki gen. nov., sp. nov. ~. (a) Base of hind femur; (b) antenna; (c) hypopygium in side view.

FIG. 3. Megaselia sembeli sp. nov. (a) g Palp; (b) 9 palp; (c) g hypopygium in side view.

FIG. 4. Megaselia kodongi sp. nov. o~: hypopygium in side view. Scale bars=0-1 mm.

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New genus and three new species of Phoridae 781

fluorescence was also observed just beyond the tapered (internal) dark structure before the tip of the distal section (see Fig. 1 (a)). This fluorescence indicates the presence of resilin in these positions (Anderson and Weis-Fogh, 1964). The presence of resilin is normally associated with the performance of very rapid movements. The rubber-like properties of resilin contribute a spring-like mechanism to the structure in question. I initially described this ovipositor as looking 'like a spring-loaded mechanism for shooting it rearwards' (Disney, 1985). This would, therefore, appear to be nearer the truth than I realized at the time!

Wings. (Fig. 1 (f)). Length 0.754)-95 mm. Costal Index 0"34).4mm. Costal ratios 2.66-2.76:1. Costal cilia 0"03~.04mm. Veins pale brown a n d vein 7 very pale. Membrane lightly tinged brownish grey, a little darker along front margin. Halteres with brownish grey, somewhat large, knob and paler stem.

Legs. Pale brownish and paler still on distal part of tibiae and tarsi. All femora somewhat slender. Tarsal segments 5 and 6 as Fig. 1 (b), (d), (e).

Male Among more than 10000 specimens of Phoridae collected in traps in the type-

locality of Iridiphora clarki, the males of only one species resemble the females described below. While there are interesting differences, the resemblances in detail, as well as overall jizz (Gestalt), indicate that these males are indeed the same species as the females. The most obvious difference is in the presence of a reduced arista in the males along with a row of spines below the base of the hind femur. Sexual dimorphism in antennae and the development of spines at the base of the hind femur are both widespread in the Phoridae.

Description. Closely resembles female but with following differences. Antenna (Fig. 2 (b)) with short two-jointed arista. Last tarsal segments all resemble those of middle legs of female, that is those of front and hind legs are less attenuated. Hind femur with ventral comb of three curved spines at base (Fig. 2 (a)). Wing length 0.84-0.93 mm. Costal Index 0.34).4. Costal ratios 2.00-2.30: 1. Costal cilia 0.024).03 mm. Hypo- pygium (Fig. 2 (c)) with brownish epandrium clearly paler than abdominal tergites 1-6. Anal tube almost colourless. Hypandrium dark with pale areas away from margins.

Natural history. The females of this species attack workers of the ant Iridomyrmex cordatus (Smith). This ant utilizes covered ways constructed by termites on the trunks of trees and will take over the outer parts of the carton nests of termites. If part of the covered way is ruptured some of the ants will collect fragments of the covered way but show no obvious signs of repairing the damage, whereas when occupied by termites such damage is repaired within a few hours. After several days I have observed somewhat inexpert repairs of a covered way occupied by L cordatus, which suggest that they may eventually endeavour to repair a damaged covered way.

On exposure of the ants, Iridophora clarki and Megaselia sembeli (see below) were observed attacking them within half an hour. Iridophora clarki hovers near the ants and appears to dart at an ant's head. I presume this results in the insertion of an egg, but the action is so fast, the fly so small and the light intensity in the forest so reduced that I was unable to be sure of this. All 22 females collected were procured as they hovered at workers of Iridomyrmex cordatus. They were collected on the following dates 8, 9 (two colonies), 13, 15, 17 (2 colonies) and 27.i.1985, 9 and 22.ii.1985 and 6.iii.1985. No females were collected in any other circumstances either in traps or at a wide range of other ant species.

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Megaselia kodongi sp. nov.

(Figs 4, 6)

Deposition of types. HOLOTYPE (~) Sulawesi-Utara, Dumoga-Bone National Park, Toraut Forest, 12.i.1985, coll. R. H. L. Disney, deposited in collection of author. PARATWES (2~) same data and depository as nOLOa'¥PE.

Etymolooy. The species is named after Fenje Kodong (University of Sam Ratulangi, Manado) who was a great help to Project Wallace.

Male Head. Frons broader than high and brown. Lower supra-antennal bristles reduced

but still a little stronger and longer than hairs of frons. Antials a little higher on frons than upper supra-antennals and a little nearer the latter than to anter-laterals, which are a little higher on the frons. Median row of bristles almost straight and nearly equidistant. Fifty-sixty hairs on frons. Third antennal segment and palps pale brownish. Palp bristles a little more robust than lower supra-antennals. Labrum and labella simple and pale.

Thorax. Brown on top but paler on sides of scutum and scutellum. Pleura pale. Notopleuron with three bristles. Mesopleuron with half-a-dozen hairs and a bristle by hind margin. Scutellum with an anterior pair of hairs and a posterior pair of bristles.

Abdomen. Tergites 1--6 dark. Venter dusky with hairs on segments 4-6. Hairs of tergites largely restricted to posterior margins and sides, those of tergite 6 hind margin being stronger. Hypopygium (Fig. 4) with dark epandrium and dusky anal tube which at its base is ventrally embraced by a half-collar-like development from the epandrium.

Win#s. Length 1-06mm. Costal Index 0.494).50. Costal ratios 4.3:3-2:1. Costal cilia 0-03-0.04 mm. Sc runs to R1. Vein 4 originates just beyond fork of vein 3. No hair at base of vein 3. Axillary ridge with two bristles, the inner being shorter. Veins brown, with vein 7 being very obscure. Membrane lightly tinged brownish grey. Halteres brownish.

Legs. All very pale yellowish apart from darker tip of hind femur. Middle tibia spur almost as long as metatarsus. Hind tibia with about eight posterodorsals, the lower four being somewhat spine-like. Three to four hairs below basal half of hind femur clearly longer and stronger than anteroventrals of distal third.

Female As male but wing length 1.06-1.30mm. Costal Index 0.54-0.57. Costal ratios

4-4-4.9: 3.5-4.4: 1. Mesopleuron with one or two bristles. Abdomen. (Fig. 6). Short tergites 1-5 and 2-4 with a dark band just before hind

margin. Tergite 6 Y-shaped. Ovipositor well developed. Venter with short hairs on segments 5 and 6 and a reduced pair on 4.

Affinities. In the keys of Borgmeier (1967 a) M. kodongi runs to either couplet 49 or 53 of Group II, depending on the costal index, and thus would be identified as M. simplicior (Brues). However it differs from M. simplicior in the very different costal ratios, paler legs, much stronger end hairs of the anal tube and many other features. Megaselia corkerae Disney (1981) will also run to couplet 53 in the same key but it has a much shorter and fatter anal tube in the male and unmodified ovipositor segments with normal cerci in the female. The strongly developed ovipositor of M. kodongi would have placed the species in the genus Plastophora in the past. However, all species with a forked vein 3 have been transferred to Megaselia (Disney, 1978) and the validity of

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FIG. 5. Megaselia sembeli sp. nov.: ovipositor in dorsal view. Scale bar=0.1 mm.

FIG. 6. Megaselia kodongi sp. nov.: ovipositor in dorsal view. Scale bar = 0"1 mm.

Plastophora is further considered below. If the species is taken through Colyer and Elberg's (1969) key to the word Plastophora it runs to M. cultrata Brues, a species known from a single female from the Philippines. The two species are clearly somewhat similar but M. cultrata has longer abdominal tergites and the sixth is not Y-shaped (see fig. 39 in Borgmeier, 1967 a). Megaselia kodongi will not key to any known species in the rest of the literature for the world fauna of the large genus Megaselia.

Natural history. All three specimens were caught at anastomosing columns of the ant Pheidologeton sp. foraging on the forest floor. The females were landing and racing up to worker ants. The male was endeavouring to distract one of the females. The females appeared to be trying to oviposit in the gasters of the ants.

Megaselia sembeli sp. nov.

(Figs 3, 5)

Deposition of types. HOLOTYVE (~) Sulawesi-Utara, Dumoga-Bone National Park, Toraut Forest, 8.i.1985, coll. R. H. L. Disney, deposited in collection of author.

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PARATYPES (69~, 193`) same data and depository as HOLOTYPE except dates are various i-iii.1985 and 2~, 23' deposited in Museum Zoologicum Bogoriense, Bogor, Java.

Etymology. The species is named after Dr D. T. Sembel (University of Sam Ratulangi, Manado) who was an outstanding help to Project Wallace in many ways.

Male Head. Frons broader than high, although mid-line height is only just less than

breadth. Upper supra-antennals clearly shorter than antial bristles, but a little longer than longest bristles on palps. Lower supra-antennals clearly shorter and finer than upper pair. Antials slightly higher and a little closer to upper supra-antennals than to anterolaterals, which are somewhat higher on frons. Median row of bristles with pre- ocellars distinctly further apart than either is from mediolateral, and the latter clearly lower on the frons. Third antennal segment and arista pale brownish. Palps pale yellowish and with complex sensory pit on external face (Fig. 3 (a)). Labella simple and, along with labrum, pale in colour.

Thorax. Scutum and scutellum brown on top and pale at sides. The pleura are brown above and sharply demarcated from the pale lower half (or more). This feature, along with the pale margin of the scutum, gives a striped look to the side of the thorax. Notopleuron with three bristles. Mesopleuron with around a dozen hairs plus a long strong bristle near hind margin. Scutellum with an anterior pair of hairs and posterior pair of bristles.

Abdomen. Tergite 1 yellowish in middle and darker at sides. Tergite 2 yellow in an anterior median patch, otherwise dark. Tergites 3 and 4 entirely dark. Tergites 5 and 6 yellow in anterior half and dark posteriorly. Hairs short and sparse on first four tergites, a little more numerous on posterior halves of 5 and 6, the latter bearing stronger hairs near posterior margin. Venter yellowish with a few minute hairs only on segments 4-6. Hypopygium (Fig. 3 (c)) with dark epandrium, paler hypandrium and yellowish anal tube.

Wings. Length 1-0-1.3 mm. Costal Index 0.424).44. Costal ratios 1.3-1-4: 1. Costal cilia 0.034)-05 mm. Vein 3 unforked. Sc obscure (only discernible with critical lighting). No hair at base of vein 3. Two bristles on axillary ridge, the inner one being shorter. Veins greyish brown, but vein 7 very faint. Membrane lightly grey tinged. Haltere with darkish stem and yellow knob.

Legs. All pale yellow apart from tip of hind femora and tibiae, which are somewhat darkened. Spur of mid tibia about as long as metatarsus. Spur of hind tibia at most as long as breadth of tibia at point of insertion. Hind tibia with 8-10 posterodorsals, the lower ones being a little more robust. The four-seven hairs below base of hind femur are deafly longer than and stronger than anteroventrals of distal third, although the first is somewhat shorter.

Female As male but with bristles of median row on frons more nearly equidistant. Palps

brownish, somewhat more swollen in distal half and with sensory organ internal (Fig. 3 (b)). Costal Index 0.424).45. Costal ratios 1-2-1.4 : 1.

Abdomen. (Fig. 5). Tergites part yellow and part brown pigmented. Venter yellowish but darkened on segment 6 and below edge of tergites on segment 4 and 5. A pair of conspicuous bristles developed on posterior margin of tergite 6 and a similar

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pair either side is situated on side below edge of tergite on dark patch. Tergites 7 (illustrated in Fig. 5) and 8 long, narrow and heavily sclerotized along median band at least.

Affinities. Megaselia sembeli is distinguished from nearly all Indo-Australian Megaselia species which possess hairs, and a bristle on the mesopleuron by having an unforked vein 3. The only two species covered by Borgmeier's keys (1967 a, b) with this feature are M. sernota Beyer and M. nigribasis Beyer. Meoaselia semota has entirely dark abdominal tergites, is generally a darker species, and differs in numerous other details. Megaselia nigribasis has some yellow abdominal tergites but the mid-femur is brown and the hind femur is almost black. It also differs in numerous other details (such as the position of the antial bristles).

The modification of the ovipositor places the females in the genus Plastophora. I have previously (Disney, 1978) transferred all members of this genus with a forked vein 3 to Megasetia, leaving three species, P. beirne Brues (the type-species of the genus), P. dubitata (Brues) and P. cornigera Beyer, in Plastophora. The reason for not synonymizing the genus at the time was that the type-material of P. beirne from New Guinea had been lost. Even though this material was re-described by Schmitz (1929), it was felt wise to await new material of it, or a closely related species, before finally synonymizing the genus. Megaselia sembeli in fact runs to P. beirne in the key of Colyer and Elberg (1969). The latter species has a more robust proboscis, a longer costa (Costal Index > 0.5), and more extensive yellow coloration. Also there is no indication of bristles on abdominal segment 6 in Schmitz's fig. 41. However, while clearly a different species it is equally evident that these two species are very closely related. In terms of assessing the validity or otherwise of the genus Plastophora, M. sembeli is sufficiently close to P. beirne to be regarded as congeneric. The characters peculiar to the latter species are dearly not of generic significance.

A point-by-point comparison of M. sembeli with a range of species regarded as Megaselia by all previous authorities reveals no features which suggest that Plastophora is a distinct genus. The latter genus could only be maintained knowing it to be polyphyletic and consequently rendering Megaselia paraphyletic. I accordingly synonymize the genus Plastophora Megaselia (syn. nov.) and transfer the remaining species to Meoaselia as follows:

MegaseIia beirne (Brues, 1905) comb. nov. Megaselia cornigera (Beyer, 1966) comb. nov. Megaselia bruesi nom. nov.

Plastophora dubitata Brues, 1936 Meoaselia dubitata (Brues, 1936) nec Malloch, 1912

I have had to provide a new name for the last species in order to avoid a consequent secondary homonym.

Natural history. All specimens were caught at exposures of the ant Iridomyrmex cordatus, usually at the same time as attacks by Iridophora clarki (see above). The females landed near the ants, ran up to them from the side and appeared to oviposit into the gasters of the ants. The males also landed and attempted to distract females manoevring for an attack on an ant. The species was collected on eight occasions as follows. 8.i.1985 (35~, 33'), 9.i.1985 (5~, 1~), 13.i.1985 (6~, 7~),15.i.1985 (1~), 27.i.1985 (11~), 22.ii.1985 (97, 53'), 6.iii1985 (2~, 1~). 8.iii.1985 (1~, 2~). This gives an overall ratio of females to males of 3-5 : 1. The females were deafly gravid and up to four mature eggs have been recorded in the abdomen. These eggs measure 0.50-0.55 mm in length.

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Discussion The vulnerability of Iridomyrmex cordatus to attack by Iridophora clarki and

Megaselia sembeli is striking and led to the suggestion (Disney, 1985) that this ant's habit of utilizing covered ways constructed by termites has evolved as a direct response to the pressure of attack by phorids.

The Pheidologeton species attacked by Megaselia kodongi has the habit of altering its area of search each day. Thus whilst they were observed foraging on the same patch of the forest floor at different times on the same day, the next day no sign of this species could be found in this area. This contrasts with several ant species, belonging to a variety of genera, whose columns were observed utilizing the same routes from one day to the next over several weeks. I observed no phorids attacking these columns during many hours of daylight observation. Pheidologeton sp. does have permanent routes near its nest; these are converted into covered ways by means of soil particles. A further curious habit of this ant species is the high frequency of snails collected and transported back to the nest.

I would suggest that the relationship between an ant species' foraging habits and its vulnerability to attack by parasitoid phorids merits further observation and experiment.

Acknowledgements This paper is based on material collected whilst the author was a participant on

Project Wallace, sponsored by the Royal Entomological Society of London and the Indonesian Institute of Sciences (Results of Project Wallace No. 5).

My participation on Project Wallace was made possible by grants from the Field Studies Council Study Leave Fund (financed by an anonymous trust), The British Ecological Society, the Royal Society and the Percy Sladen Memorial Fund (administered by the Linnaean Society). The ants were identified by Barry Bolton (British Museum (Natural History)).

References ANDERSON, S. O. and WEIS-FOGH, T., 1964. A rubber like protein in arthropod cuticle. Advances

in Insect Physiology 2, 1-65. BEYER, E. M., 1966. Neue und wenig bekannte Phoriden, zumeist aus dem Bishop Museum

Honolulu. Pacific Insects g, 165-217. BORGMEIER, T., 1963. Revision of the North American Phorid Flies. Part I. Studia Entomologica

Petr6polis 6, 1-256. BORGMEIER, T., 1967 a. Studies on Indo-Australian Phorid flies, based mainly on material of the

Museum of Comparative Zoology and the United States National Museum (Diptera, Phoridae). Studia Entomologica Petrrpolis 9, 129-328.

BORGMEIER, T., 1976 b. Studies on Indo-Australian Phorid flies, based mainly on material of the Museum of Comparative Zoology and the United States National Museum (Diptera, Phoridae). Part II. Studia Entomologica Petrdpolis 10, 81-276.

BRUES, C. T., 1905. Phoridae from the Indo-Australian Region. Annales Musei Nationalis Hungarici 3, 541-555.

BRUES, C. T., 1936. Philippine Phoridae from the Mount Apo region in Mindano. Proceedin~ls of the American Academy of Arts and Sciences 70, 365-466.

COLYER, C. N. and ELBERG, K., 1969. New data on Phoridae (Diptera). Eesti NSU Teaduste Akadeemia Toimetised XVIII Kfide Biologica 1969, 154-169.

DISNEY, R. H. L., 1978. A new species of Afrotropical Megaselia (Diptera, Phoridae), with a re- evaluation of the genus Plastophora. Zeitschriftffir Angewandte Zoologie 65, 313-319.

DISNEY, R. H. L., 1981. A new species of Megaselia from Nepenthes in Hong Kong, with re- evaluation of genus Endonepenthia (Diptera: Phoridae). Oriental Insects 15, 201-206.

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New genus and three new species of Phor idae 787

DISNEY, R. H. L., •982. Three new species of scuttle-fly (Diptera, Phoridae) that parasitize ants (Hymenoptera: Formicidae) in North America. Journal of Zoology, London 197, 473-481.

DISNEY, R. H. L., 1983. Scuttle-flies--Dipteral Phoridae (except Megaselia). Handbooks for the Identification of British Insects 10 (6), 1 81.

DISNEY, R. H. L., 1985. Watching scuttle flies on Project Wallace. Antenna 9, 139-141. MALLOC~I, J. R., 1912. The insects of the dipterous family Phoridae in the United States National

Museum. Proceedings of the United States National Museum 43, 411-529. ScI-t~vz, H., 1929, Die Phoriden. Maastricht: C1. Goffin (Also published as Revision der Phoriden.

Berlin: Ferd. Dfimmlers, 211 pp.)

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