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Relationships of the genus Heterocheila (Diptera: Sciomyzoidea) with description of a new family PDF

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Preview Relationships of the genus Heterocheila (Diptera: Sciomyzoidea) with description of a new family

DAVID K. MCALPINE AustralianMuseum, Sydney RELATIONSHIPS OF THE GENUS HETEROCHEILA (DIPTERA: SCIOMYZOIDEA) WITH DESCRIPTION OF A NEW FAMILY McAlpine,D.K., 1991-RelationshipsofthegenusHeterocheila(Diptera:Sciomyzoidea) withdescriptionofanewfamily.-TijdschriftvoorEntomologie134:193-199-[ISSN0040- 7496]. Published 18December 1991. A morphological comparison is made of the maritime kelp-living genus Heterocheila RondaniwiththefamiliesofSciomyzoidea,particularlytheHelcomyzidae.Itisconcluded that ithas noparticularlycloserelationshipwithanyoneofthesefamilies,andthenew familyHeterocheilidae (halfbridgeflies) isestablishedforit.Heteromyzaorientalis Macquart, 1843 is a newsynonymofHeterocheilabuccata (Fallen, 1820). D. K. McAlpine,AustralianMuseum,BoxA285,SydneySouth2000,Australia. Keywords. -Diptera; Heterocheilidae, new fam.ily;Holarctic. InthecourseofmyrecentworkontheCoelopi- Useful descriptive and illustrative material of dae (kelp flies) (D. McAlpine 1991) it became the adult morphology of Heterocheila has been apparent that (1) there is disagreement in recent given by Czerny (1930), Hennig (1958), Steyskal literatureastothefamilypositionofHeterocheila, (1958, 1962, 1987),andGriffiths (1972).Backlund and (2) it is difficult on morphological grounds to (1945) and Egglishaw (1960b) have described the justifytheinclusionofHeterocheilainthevarious egg, larva and puparium of Heterocheila, and the families (Coelopidae, Dryomyzidae, Helcomyzi- latter alsogives importantbiological information. dae) whereithasbeenrecentlyassigned. Forthese I haveexamineda series ofadultsofbothsexesof reasons the present investigation was made. H.buccata,andcollectedthespeciesinthefield.W. TheholarcticgenusnowknownasHeterocheila N. Mathis has provided materialofH. hannai. Rondani, 1857, has often been referred to by the The family Helcomyzidae is here considered to synonymous name Oedoparea Loew, 1862. See includeonly thegenera HelcomyzaCurtis,Maori- both Steyskal (1965), and Gorodkov (1984) for myia Tonnoir & Malloch, and Paractora Bigot. I complete synonymy. Included species are: (1) H. have given reasons for excluding the group from buccata (Fallen, 1820) (syn. Heteromyzaorientalis the Dryomyzidae (D. McAlpine 1991) and sug- Macquart, 1843), from Europe; (2) H. hannai gestedacloserrelationshiptotheCoelopidae.Use- (Cole, 1921) (syn. H. nudiselaCurran, 1933),from fuldescriptiveandillustrativematerialoftheadult Pacific North America. Both species live in morphology of Helcomyzidae has been given in strandedkelp (seeespecially Egglishaw 1960b). most of the papers mentioned above for Hetero- Heterocheila (or Oedoparea) has often been cheila morphology, also byMalloch (1933). Eggli- placed in the family Helcomyzidae (e. g. Malloch shaw(1960a)hasdescribedthelarvaandpuparium 1933; Hennig 1937, 1973; Steyskal, 1958, 1965; ofHelcomyza, andgiven biological information. I Gorodkov 1984), or in the Dryomyzidae, broadly have examined adult material of all genera and defined to include Helcomyza Curtis and allied mostdescribed species ofHelcomyzidae. genera (e.g. Czerny 1930; Steyskal 1987;J. McAl- In the following text I use the superfamily Sci- pine 1989). Egglishaw (1960b), Dobson (1976), omyzoidea to include the families Sciomyzidae, andGriffiths (1972) havepreferredtoplaceHete- Huttoninidae, Helosciomyzidae, Dryomyzidae, rocheila in the Coelopidae (with some reserva- Helcomyzidae,Coelopidae,Ropalomeridae,Sepsi- tions), but I have given definite reasons for its dae, Chamaemyiidae (including Cremifaniidae), exclusion from the Coelopidae (D. McAlpine Eurychoromyiidae (probably excluding the Gay- 1991), and these need not be repeated here. omyia complex, seeJ. McAlpine 1989), and Laux- 193 TijdschriftvoorEntomologie,volume 134, 1991 aniidae(includingCelyphidae).J.McAlpine(1989) tinct from the investigated dryomyzids and the includesthelastthreefamiliesinaseparatesuper- apparentgroundplan ofCoelopidae. On the other familyLauxanioidea. hand the number of spermathecae is unrecorded forthe helcomyzidgenus Paractora. Increasingknowledgeofspermathecalnumbers Similarities of Heterocheilato Helcomyzidae in Schizophorahas demonstratedthat it is a fairly unstable character. The change from three to two Heterocheila shares with Helcomyzidae s. str. spermathecae must havehappenedmany times in thebeachhabitatanddependenceonstrandedkelp, the evolution of the Schizophora, if, in fact, the utilising it for larval nutrition as well as shelter. changehasalwaysbeenintheonedirection.Inthe These habits are shared with a number of other Sciomyzoidea, variation in spermathecal number Diptera, notably the Coelopidae, the sepsidgenus occurs within the families Huttoninidae, Coelopi- Orygma Meigen, and the sphaerocerid genus (or dae, and Sciomyzidae. subgenus) ThoracochaetaDuda. Similarityofhab- Other traits shared between Heterocheila and its andhabitatseems tohavecausedsomeconver- the Helcomyzidae are, so far as I am aware, found gent similarities among these shoreline flies, so inawidespectrumofsciomyzoidfliesandhavenot that Orygma and Heterocheila havebeen referred been postulated as evidence ofcloser relationship to the Coelopidae in thepast. between these two taxa. Such traits include the InelevatingHelcomyzidaetofamilystatus,Mal- developmentofmollisetae(seeD.McAlpine1991) loch (1933) emphasised the significance of the onvariouspartsofthemales,thegeneralbrownish sclerotised precoxal bridge, which connects the greypruinescentcoveringofthecuticle,character- prosternum to the propleuron on each side. Hen- isticofmany shore-dwelling flies, and the shining nig (1958) considered this condition to represent parafacial ridges, also found in many flies of mar- asynapomorphypeculiartotheHelcomyzidaeplus itime or sandy habitats. Ropalomeridae among theSchizophora, which he thereforeconsidered toconstitute a monophyletic Differences between Heterocheila and group. It is now known that prothoracic precoxal Helcomyzidae bridgesoccurinatleastsomerepresentativesofat least 19 schizophoran families (Speight 1969, and The principal morphological differences be- my observations), and many arisals of the condi- tweenadultsofthesetaxaaregivenintable1.They tionareacknowledged.WithinthesuperfamilySci- are discussed below in numerical sequence, with omyzoidea,thetraitoccursalsoinsomesciomyzids theadditionofsomecommentsonlarvalmorphol- as an independent apomorphy. The presence of ogy(18). precoxal bridges is not now usually considered as 1. Hackman & Väisänen (1985) have investi- proof for monophyly ofHelcomyzidae and Ropa- gatedthecostalchaetotaxyoftheDiptera (includ- lomeridae (Griffiths 1972;J. McAlpine 1989),and ing Heterocheila, op. cit.: fig. 16) and assigned it additional evidence is needed if monophyly of some taxonomievalueat the family andsubfamily Heterocheila with Helcomyzidae is to be estab- levels. The additional dorsal and ventral rows of lished.TheprecoxalbridgeinHeterocheiladiffers costalsetulaepresent inHeterocheilaareinagree- somewhat from that of Helcomyzidae in being mentwithDryomyzidaeandHelosciomyzidae(but markedly narrower. not Coelopafrigida (Fabricius) with which Hack- In both Helcomyza and Heterocheila the basal man&Väisänencompareit).Theabsenceofthese crossvein (crossveinbm-cuorbaseofM3+4accord- series in all generaofHelcomyzidae is incontrast ing to divergent applications of the Comstock- to the above taxa. The presence of outstanding Needham notation) lies more obliquely, with ap- anteroventralspinesatintervalsonthemidregion proach to a longitudinal orientation, than in Dry- of the costa in all Helcomyzidae separates them omyza. This condition is unlikely to be a ground- sharply from the Dryomyzidae and Heterocheila, plan condition for Helcomyzidae s. str. in view of and aligns them with Helosciomyzidae and some thefactthattheundoubtedhelcomyzidMaorimyia Coelopidae (e. g. Lopa convexa McAlpine, Gluma hasthebasalcrossveinmorenearlytransversethan keyzeriMcAlpine,andRhiswhitleyiMcAlpine,see infouravailableDryomyzaspecies.Ialsofindsome D. McAlpine 1991). variation in this character in both Coelopidae and 2. The discontinuity of the parafacial suture on Sciomyzidae. It appears to be a rather unsatisfac- its lowerpartandpeculiarangularshiningparafa- tory indicatorofphylogenetic affinities in the Sci- cial ridge are apparently autapomorphies of the omyzoidea. Helcomyzidae. The condition in Heterocheila is Heterocheila, Maorimyia, and Helcomyza have probably partly plesiomorphic, though the exten- two spermathecae in the female abdomen, as dis- sive shining ridge may be an autapomorphy. 194 McAlpine: Heterocheila 3. The incised, vertically orientated postgenal lopidae, andperhaps other families,butexcluding fold in Heterocheila, is an unusual feature in the Heterocheila, Ropalomeridae, Sepsidae, Chamae- Sciomyzoideaandpresumablyanautapomorphy.A myiidae, Lauxaniidae and Eurychoromyiidae. Con- similarconditionoccursintheheleomyzidNephel- vergence in this character cannot beexcluded, but lum dendrophilum (Malloch) (D. McAlpine 1985: seemslesslikelyforthecomplexDryomyzidaeplus Fig. 30). Helcomyzidae plus Coelopidae, plus perhaps He- 4. The Helcomyzidae share with the Heloscio- losciomyzidae,asthesefamiliesshowseveralother myzidae the narrow median emargination of the somewhat inconsistent similarities. Thus, the size face adapted to receive the prelabrum. The emar- of tergite 6 in Heterocheila makes difficulties for gination is absent in Heterocheila and Dryomyzi- thehypothesis thatitisclosetotheHelcomyzidae dae,butasomewhatsimilaremarginationoccursin orCoelopidae. some,Coelopidae,thoughitisdoubtfulifitisinthe 9.TheaedeagusofHelcomyzidaeismoresimilar groundplan of the latter. Perhaps the median to that of Dryomyzidae than to that of Hetero- emargination represents a separate apomorphy in cheila(Griffiths 1972).Mystudyoftheaedeagusin each group in which itoccurs. Coelopidae(D.McAlpine1991)andHeleomyzidae 5. The shape of the hypopleural channel in the (e. g. D. McAlpine 1967) shows that its structure Helcomyzidae represents a distinctive autapo- may be quite unstable above the species level. morphy, absent in Heterocheila (D. McAlpine There are apparently consistent differences in 1991: Fig. 9). both hypandrium and aedeagus between Hetero- 6. The significance of this character has been cheila and Helcomyzidae. The hypandrium of discussed above under "Similarities'. Heterocheilahastwopairsofprocessesposteriorly 7. The male-restricted apical ventral process of (gonites orparameres) which are absent in helco- the fore basitarsus is a trait difficult to evaluate myzids.TheaedeagusofHeterocheilahasapairof phylogenetically,possiblybecauseofirregular loss spreading lateral lobes at the junctionofthe basi- inanumberoflineages(D.McAlpine 1991).Inthe phallus and distiphallus, which is absent in the Sciomyzoidea it is apparently restricted to Helco- helcomyzids examined, but it lacks the patch of myzidae, Coelopidae, and Dryomyzidae, though pubescence near the middle of the length of the notuniformlypresentinthelastfamily.Itdoesnot distiphallus, which is present in helcomyzids. appear tobeasynapomorphyas italsooccurs in a Griffiths (1972) hasemphasised thesubstantial number of Heleomyzoidea and in Heloclusia im- differences in external male genitalia between perfecta Malloch,a somewhatprimitive represen- Heterocheilaandcertainhelcomyzids.Itispossible tative of the Nerioidea. This condition seems un- that these structures should be assigned relatively likely to have originated more than once. I low reliability as indicators of relationship. To thereforeregarditasaveryancienttraitanditmay judge from my studies of the Heleomyzidae (D. provideevidenceforrelationshipbetweenSciomy- McAlpine1967,1985)andtheCoelopidae,thecop- zoidea,Heleomyzoidea,andNerioidea.Inthiscase, ulatory structures are so unstable above species its absence in Heterocheila would be a derived level that it is difficult to homologise the various state, but not necessarily a synapomorphy with processes and lobes across the family. Therefore other sciomyzoids in which it is also absent. there is little logic in inferring autapomorphies 8.Thephylogeneticsignificanceofthereduction betweenspecialconditionsofcertaincoelopidsand of tergite 6 ofthe male abdomen in Sciomyzoidea thoseoccurringinHeterocheila,which iscertainly does not appear to have been satisfactorily ex- not so close as a sister group to the Coelopidae. plained.J. McAlpine (1989) regards reduction of However, I findgreater consistency in hypandrial tergite 6 as a groundplan apomorphy of his Scio- structurewithinthesmallertaxonomiediversityof myzoidea (as distinct from Lauxanioidea) but in- the Helcomyzidae. cludes in Sciomyzoidea such taxa as Heterocheila, 10.Thepresenceofafemale-restrictedenlarged, Orygma Meigen (Sepsidae), and Rhytidops isolated,posteriorlydirectedbristleoneachlateral Lindner (Ropalomeridae) whichhavealarge (and marginoftergites 2 to4 andsometimes 5 is char- in Orygma, according to McAlpine, primarily acteristic for most helcomyzids, including species J. unreduced) tergite 6. For this andother reasons, I inall3genera.ItisabsentintheDryomyzidae,but include the lauxanioid families Chamaemyiidae, thepresenceofsimilarfemale-restrictedbristlesin Lauxaniidae,andEurychoromyiidaeintheSciomy- two rather plesiomorphic but not closely related zoidea. coelopidspecies (D. McAlpine 1991) may indicate If evolutionary change in the size of tergite 6 that it was present in the common ancestor of takes place only in the direction of reduction, this Helcomyzidae andCoelopidae. might seem to support the ideaofa monophyletic 11-17. I do not rate these differences in chaeto- groupincludingDryomyzidae,Helcomyzidae,Coe- taxy very highly, taken individually. Collectively, 195 . TijdschriftvoorEntomologie,volume 134, 1991 Table 1. DifferencesbetweenHelcomyzidaeandHeterocheila (adults). Helcomyzidae Heterocheila 1 Midregionofcostawithspacedanteroventralspines, Midregionofcostawithoutanteroventralspines,with withoutcontinuousseriesofdorsalandventralsetu- regular series of dorsal and ventral (as distinct from lae. anterodorsalandanteroventral) setulae. 2. Shining parafacial ridge not continued posteriorly Shining parafacial ridge and associated suture con- below cheek; parafacial suture obsolete or inter- tinuedbelowcheektopostgenalregion. ruptedbehindangularsectionofridge. 3. Postgenal foldabsent. Postgenalfolddistinctlyincised. 4. Epistomal marginof face with narrow sinuation to Epistomalmarginoffacewithoutnarrowsinuation. receiveprelabrum. 5. Hypopleural channel rather narrow and parallel- Hypopleuralchannelexpandingrapidlyanteriorly. sided. 6. Prothoracicprecoxalbridgebroad,wellsclerotised. Prothoracic precoxal bridge narrow, not uniformly sclerotised. 7. $ fore basitarsus with terminal ventral thumbnail- $ (and 9) fore basitarsus without terminal ventral likeprocess. process. 8. $abdominaltergite6muchreduced. (5 abdominaltergite6large. 9. Aedeagus with densely pubescent zone or zones, Aedeagus without dense pubescence, with pair of lat- withoutlaterallobes. erallobes. 10. 9 withonelargebristle neareach lateralmarginon 9 (and$) withoutlateralmarginalbristlesonabdom- tergites2-4. inaltergites. 11. Generallyonly2 welldeveloped fronto-orbitalbris- Fronto-orbitalbristles 3. tlespresent. 12. Prostigmatalbristlepresent (sometimesreplacedby Prostigmatalbristleabsent. severallonghairs). 13. Median series of acrostichal bristles or setulae not Singleregularmedianseriesofshortacrostichalbristles differentiated. present (inadditiontoprescutellarpair). 14. Prosternumsetulose. Prosternumbare. 15. Metasternumsetulose. Metasternumbare. 16. Abdominalsternite 1 setulose. Abdominalsternite 1 bare. 17. 9 hindtibiawithsubapicalanteriorbristle. 9 (and$) hindtibiawithoutsubapicalanteriorbristle the seven points indicate a notable degree of div- segments 3 or 4 to 12 (no such spinose creeping ergencebetween the taxa. weltspresentinHelcomyzaustulata,butthecuticle 18. Larval morphology ofHelcomyzidae is only extensively covered with backwardly directed recorded for Helcomyza ustulata Curtis, and it is plates), no spine above posterior spiracle (a large uncertain if this is representative of the family as spine in this position in Helcomyza), each poste- awhole. However, inviewoftheapparentlyclose rior spiracle with two or three groups of short relationshipbetween the three helcomyzidgenera hydrofuge hairs (these absent in Helcomyza). and the very distinctive features of the known There is also difference in the arrangement and larva,thesemayhavesomesignificanceforhigher spinose armature of the ridges or processes sur- classification. According to Egglishaw (1960a, rounding the anus, and the position of the anus. 1960b) the third instar larva of Heterocheila buc- EgglishawconsidersthelarvaeofHeterocheilaand catahascreepingweltsofsmallspinesventrallyon Helcomyza to be so different that they cannot be- 196 McAlpine: Heterocheila long in the samefamily. I am unable to find in his larger male tergite 6 (see J. McAlpine 1968, for descriptions and figures any shared distinctive morphologicaldetails). Heterocheila lacks the fol- traits, which might provide evidence of relatively lowingdistinctivecharactersofEurychoromyiidae: close relationship between them, though they body form remarkably stout; chaetotaxy greatly share many points of resemblance to numerous reduced; head structure highly modified; antennal other schizophoran larvae. segment 1 (scape)elongate;scutellumabbreviated; mesopleural bristles present; tibiae lacking termi- Discussion and Conclusion nalspurs;distalsectionofvein7indistinguishable. There is also considerable difference in the male The few distinctive points of similarity shared postabdomens and in other characters. Whereas between Heterocheila and the family Helcomyzi- Heterocheilaisrestrictedtocooltemperateshores, dae are unconvincing as indicators of close phy- eurychoromyiids are only known from Bolivia, a logenetic relationship. Each of these traits occurs land-locked tropical country. I conclude that the elsewhere in the Sciomyzoidea, and the combina- few points ofresemblance between these taxa are tion or maritime habitat, a shining section of the not indicativeofclose relationship. parafacial ridge, sclerotised precoxal bridges, and Heterocheila is excluded from the Sciomyzidae twospermathecaeoccursalsointhegenus Tethina (perhaps including Phaeomyiinae) because of the Haliday (Chloropoidea: Tethinidae) though this prelabrum,andits larvae feedon neithermolluscs must beonly remotely relatedtoHelcomyzidae. It nor diplopods. Otherwise there are no special must be acknowledged that convergence between points of resemblance and no recent author has HeterocheilaandHelcomyzidaeinthesecharacters suggested aclose relationship. is not an improbableevent. HeterocheilaisexcludedfromtheHuttoninidae Ihaveindicatedapossiblesister-grouprelation- (bestregardedasaseparatefamily,notasubfamily ship between Coelopidae and Helcomyzidae (D. of Sciomyzidae or Helosciomyzidae; further dis- McAlpine 1991) and this relationshipseems tobe cussed in the forthcoming 2nd edition of'The In- further supported by costal chaetotaxy. The nu- sects of Australia') because it lacks the following merous character differences separating Hetero- distinctivefeaturesofHuttoninidae: antennalseg- cheila and Helcomyzidae indicate a markedevolu- ment 1 without setulae on medial surface; vein 6 tionarydivergence, andcharacters 1,6,7,9and 17 abbreviated;distalsectionofvein7scarcelydistin- inTable 1 may indicate that Heterocheila is prob- guishable;abdominalsternite1vestigialorabsent; ably not tobe included in the monophyleticgroup abdominal tergites 1 and2 notpartlyseparatedby Helcomyzidae plus Coelopidae, oreven the some- a membranous dorsal line. Huttoninids appear to what less surely foundedgroup Dryomyzidae plus be mainly forest-living and are endemic to New Helcomyzidaeplus Coelopidae. Zealand. Heterocheilaresemblesatleastsometaxaofthe Heterocheila is excluded from the Helosciomy- Ropalomeridae in its sclerotisedprothoracicprec- zidaebecauseitlacksthefollowingapparentlycon- oxal bridge, large male tergite 6, and two sper- sistent features of that family: costa with promi- mathecae. These traits, however,do notprovide a nent spaced anteroventral spines; fronto-orbital strongercase for relationship than those formerly plate with at most two bristles; prothoracic pre- seen to support the now discarded hypothesis of coxalbridgeabsent;abdominalsternite 1vestigial; close relationshipbetween Helcomyzidae and Ro- abdominal tergite 6 of male much reduced; each palomeridae. Heterocheila differs from the Ropa- posterior spiracleoflarva with 4 branched hydro- lomeridae inter alia in its less markedly divergent fuge hairs (Helosciomyza Hendel and Polytocus postvertical bristles, differently shaped head cap- Lamb). Helosciomyzids are apparently restricted sule, unmodified scutellum, absence of setulae on to the south-temperate zone. In my experience margin of metathoracic spiracle, unmodified theyare found in forests,grasslands, swamp-mar- femora,anddistinctbutdesclerotiseddistalsection gins,androckyshores,butarenotknowntoinhabit ofvein7.AlsoHeterocheilalivesonnorth-temper- kelp beds. ateshore-lines,whiletheropalomeridsare mainly Heterocheila differs from the families Chamae- inhabitants of tropical forests of the Americas. myiidae (including Cremifaniidae) and Lauxanii- Most recent workers (e.g.J. McAlpine 1989) con- dae (including Celyphidae) in its complete vein 6 sidertheRopalomeridaetobecloselyrelatedtothe andnumerousothertraits.Thesefamiliesaremor- Sepsidae.Thereisnoadequatereasonforincluding phologically moreremotefrom Heterocheilathan Heterocheila in this alliance. are most other sciomyzoids McAlpine 1989), (J. Heterocheila also resembles the little known theyarenotassociatedwithkelp,andfurthercom- neotropical family Eurychoromyiidae in the pro- parison is deemed unnecessary. thoracicprecoxalbridge,andthelatterhasaneven The exclusion of Heterocheila from all recog- 197 TijdschriftvoorEntomologie,volume 134, 1991 nisedfamiliesofSciomyzoideanecessitatestheset- rite generally understood to be the inverted ster- tingupofanewmonogenericfamilyofthissuper- nite8); surstylusarticulatedbasally,withbasalan- family. teriorexpansionorsecondary lobe; cereiseparate; basiphallus with broadly expanded membranous Heterocheilidaefam. n. posteriorlobe; distiphallus with complex scleroti- sation and no pubescence. Female postabdomen Type genus: Heterocheila Rondani, 1857: 104 (present onlymoderatelyextensile,withallsegmentsshort; designation). tergites andsternites6-8,epiproct,andhypoproct Diagnosticdescription all well sclerotisedand setulose; cerei separate,el- ongate. Actively flying insects of maritime environ- ments; habitus and many structural details typical Nomenciatural notes of Sciomyzoidea; cuticle largely pruinescent and non-shining; parts of thorax, legs, and abdomen I have examined the holotype of Heteromyza with mollisetae,particularlydeveloped in male. orientalisMacquart,1843,intheNationalMuseum Head.- roundedanteriorly; facesomewhatcon- ofNaturalHistory,Paris,andfindittobeidentical vex centrally, with shallow fovea below each an- withHeterocheilabuccata (Fallen, 1820) (syn.n.). tenna; parafacial ridge shining, extending, to- ThegiventypelocalityofMacquart'sspecies,Java, gether with associated suture, below cheek to is erroneous; the species is probably restricted to postgenal region; cheek deep, setulose; postgenal Europe, where it is apparently absent from foldlong,distinctlyincised,locatedtowardsposte- warmer,southern areas. rior surface of head capsule; postvertical bristles I propose the common names half-bridge flies, oftensubparellel (slightlyvariableinorientation); for representatives of the family Heterocheilidae, fronto-orbital bristles 3, directed outwards. An- and bridge flies for representatives of the family tenna, at rest, subporrect; segment 1 with setulae Helcomyzidae. These names refer to thedevelop- extending on to medial surface; segment 3 ment ofthe prothoracicprecoxalbridge. rounded-oval; aristawith minutepubescence, seg- ment5 short, notmuchexserted;prelabrum mod- Acknowledgements erately developed, narrowly separated from face, but not fitting into sinuation on lower margin of I am indebted to M. C. D. Speight for help in latter. finding Heterocheila in the field, to L. Tsacas and Thorax. - Sternopleural suture not ascending M. Baylac for the opportunity to examine type posteriorly; hypopleural channel rapidly expand- material in the National Museum ofNatural His- inganteriorly;marginofmetathoracicspiracleand tory,Paris,andtoW.N.Mathisand R.Vockeroth J. sternopleuron bare; prosternum broadly triangu- forstudy material. lar, with narrowly sclerotised precoxal bridges; othersignificantcharactersofchaetotaxy andves- titure as in Table 1. Each tibia with one preapical References dorsalbristle;allbasitarsiwithoutterminalventral process; last 2 tarsal segments depressed and dis- Backlund, H. O, 1945. Larvae andpupaeofHeterochila 4st,aelrlibyeeseyxoofpnacdnldoesenedld.yCpolofasctseaudbucsnopbsitrnaouklweeisnt,ahnedxatnaernaedngitunelgraortdosoevrresiiaenls blHKiuuencnlgcgcealrotiam1gy5aFz(a6alF)ly:,susi5ctoougplmrpaa.ptfaairseCkuadrtSwiäisltlhs(kDiatpphtee.t,ssuDiprLpyuoonsmdeydzFiödlararheva)a.ndo--f eachofdorsal,anteroventral,andventralsetulaeor Czerny,L.,1930.Dryomyzidae.-In:Lindner,E. (editor). hairs, without spaced anterior or anteroventral Die Fliegen der Paiaearktischen Region 38a: 1-8. E. spines; vein6 visible approximately to wing mar- Schweizerbart'scheVerlagbuchhandlung,Stuttgart. gin; vein 7 beyond alula represented by long, Dobson, T, 1976. Chapter 16. Seaweed flies (Diptera: curvedcrease in membrane. Cinoseelcotpsi:da4e4,7-e4t6c.3).. N-orInt:h-CHhoelnlga,ndL.P(uebdliitsorh)i.ngMaCroimn-e Abdomen. - Tergites 1 and 2 incompletely se- pany,Amsterdam. parated by a dorsal transverse membranous line; Egglishaw, H. 1960a. The life-history ofHelcomyza J., sternite 1 short,butratherwellsclerotised,bare;7 ustulataCurt.(Dipt.,Dryomyzidae).-Entomologist's pairs of abdominal spiracles situated in pleural monthlyMagazine96: 39-42. membrane. Male postabdomen: tergite 6 shorter Egglishaw,H.J.,1960b.StudiesonthefamilyCoelopidae than preceding tergites but not greatly reduced, (Diptera).-TransactionsoftheRoyalEntomological Society 112: 109-140. almost symmetrical, setulose; sternites 6 and 7 Gorodkov,K.B., 1984.FamilyHelcomyzidae.-In:Soos, placed on left side; tergites 7 and 8 absent or not A. &L. Papp (editors). CatalogueofPalaearcticDip- definitely identifiable ('8T' ofSteyskal is the scle- tera9: 149-150. Elsevier,Amsterdametc. 198 McAlpine: Heterocheila Griffiths,G. C D., 1972. Thephylogeneticclassification McAlpine,J.F, 1989.Phylogenyandclassificationofthe ofDipteraCyclorrhaphawithspecialreferencetothe Muscomorpha. - In: McAlpine,J.F (editor). Manual structure of the male postabdomen. - W.Junk, The ofNearticDiptera3: 1397-1518.-CanadianGovern- Hague. mentPublishingCentre,Hull,Quebec. Hackman, W. & R. Väisänen, 1985. The evolution and Rondani,C,1857.DipterologiaeItalicaeprodromus.Vol. phylogenetic significance of the costal chaetotaxy in 3; Species Italicaeordinis dipterorum ingeneracha- theDiptera. -AnnalesZoologicaeFennicae22: 169- racteribusdefinita,ordinatimcollectae,methodoana- 203. liticadistinctae,etnovisvelminuscognitisdescriptis, Hennig,W., 1937.Coelopidae.-In: Lindner,E. (editor). Pars prima: Oestridae, Syrpfhidae [sic], Conopidae, Die Fliegen der Palaearktischen Region 52: 39pp. E. 264pp., 1 fig.-Parmae[=Parma]. Schweizerbart'scheVerlagbuchhandlund,Stuttgart. Speight,M.C. D., 1969.Theprothoracicmorphologyof Hennig,W., 1958.DieFamilienderDipteraSchizophora acalyptrates (Diptera) and its use in systematics. - undihrephylogenetischenVerwandtschaftsbeziehun- TransactionsoftheRoyalEntomologicalSociety 121: gen. -BeiträgezurEntomologie8: 505-688. 325-421. Hennig,W.,1973.31.Diptera(Zweiflügler).-Handbuch Steyskal,G.C,1958.NotesonnearcticHeleomyzidaeand derZoologie4(2) 2: 337 +4unnumberedpp. Dryomyzidae(DipteraAcalyptratae).-Papersofthe Malloch,J. R., 1933.Acalyptrata (Helomyzidae,Trypeti- Michigan Academy ofScience, Arts, and Letters 43: dae, Sciomyzidae, Sapromyzidae, etc.). - Diptera of 133-143. PatagoniaandSouthChile6: 177-391,pi. 2-6. Steyskal,G.C, 1962.NotesonpalaearcticDryomyzidae, McAlpine,D.K., 1967.TheAustralianspeciesofDiplo- HeleomyzidaeandSciomyzidae (Diptera). - Notulae geomyza and alliedgenera (Diptera,Heleomyzidae). Entomologicae42: 71-72. - Proceedings of the Linnean Society ofNew South Steyskal,G.C, 1965.Heleomyzidae.-In:Stone,A.etal. Wales92: 74-106. (editors). ACatalogoftheDipteraofAmericanorth McAlpine,D. K., 1985.TheAustraliangeneraofHeleo- of Mexico: 678-679. - United States Department of myzidae(Diptera:Schizophora)andareclassification Agriculture,WashingtonDC. oMfutsheeufmam3i6l:y2i0n3t-o25tr1i.bes. - RecordsoftheAustralian Stey(sekdailt,orG).. CM,an1u9a87l.oDfryNoemayrzctiidcaeD.ip-tIenr:aM2c:Al9p2i3n-e9,26J.. F-. McAlpine, D. K., 1991. Review of the Australian kelp Canadian Government Publishing Centre, Hull, Qu- flies (Diptera:Coelopidae).-SystematicEntomology ebec. 16: 29-84. McAlpine,J. F., 1968. Taxonomie notes on Eurychoro- myiamallea(Diptera:Eurychoromyiidae).-TheCan- Received: 4June 1991 adianEntomologist 100: 819-823. Accepted: 25July 1991 199

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