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Six new species of gall midges (Diptera: Cecidomyiidae) from Melaleuca (Myrtaceae) in Australia PDF

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Preview Six new species of gall midges (Diptera: Cecidomyiidae) from Melaleuca (Myrtaceae) in Australia

PROC. ENTOMOL. SOC. WASH. 99(2), 1997, pp. 312-334 SIX NEW SPECIES OF GALL MIDGES (DIPTERA: CECIDOMYIIDAE) FROM MELALEUCA (MYRTACEAE) IN AUSTRALIA Raymond J. Gagne, Joseph K. Balciunas, and DAMffiN W. Burrows (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, NHB 168, Wash- ington, D. C. 20560, USA.; (JKB) USDA-ARS Australian Biological Control Laboratory, James Cook University, Townsville, 4811, Queensland, Australia; present address: West- em Weeds Quarantine Laboratory, Agricultural Research Service, USDA, 800 Buchanan St., Albany, CA 94710, U.S.A.; (DWB) Australian Centre for Tropical Freshwater Re- search, Australian Biological Control Laboratory, James Cook University, Townsville, 4811, Queensland, Australia. — Abstract. A new genus, Lophodiplosis Gagne, is described for five new species of gall midges from Australia associated with Melaleuca spp. At least two of these are candidates for biological control of the introduced pest, Melaleuca quinquenervia, or paperbark, in Florida. In addition, a new species of Lasioptera is described. The new species are: Lophodiplosis indentata and Lophodiplosis denticulata that form blistergalls on leaves of Melaleuca spp.; Lophodiplosis bidentata, responsible for rosette bud galls on Melaleuca spp.; Lophodiplosis cornuata, forming trumpet-shaped leaf galls on Me- laleuca viridiflora; Lophodiplosis trifida, an inquiline in galls of L. indentata, L. denti- culata, and L. bidentata; and Lasioptera uncinata, an inquiline in galls of L. indentata and L. cornuata. Keys are given to adults of Australian genera of the supertribe Ceci- domyiidi and to adults and pupae of the six species found on Melaleuca spp. during this study. Key words: gall midges, Cecidomyiidae, Melaleuca, Australia Melaleuca quinquenervia is one of more (Florida Conservation Foundation (1993) than 200 species of a large genus of Myr- where it was introduced in 1906 (Schmitz taceae that is mostly endemic to Australia et al. 1991). It is a lesser pest on some Ha- (Barlow 1988, Holliday 1989). While most waiian Islands (Balciunas, unpublished). In Melaleuca species are small shrubs, M. southern Florida this plant now occupies at quinquenervia grows into a robust tree up least 500,000 acres (Bodle et al. 1994), m to 25 high in Australia (Bodkin 1991) causing extensive environmental and eco- and up to 29 m high in Florida (Rockwood nomic damage (Balciunas and Center and Geary 1991). It is one of many Austra- 1991). Even small trees produce a great lian trees that are popular in plantings in number of seeds that in Florida, unlike in tropical and subtropical regions of the Australia, result in a thick carpet of seed- world. Unfortunately, M. quinquenervia has lings that in a few years form dense, mon- become a pest in some locations in the ospecies forests (Balciunas, personal obser- United States. It is considered the most vation). In Australia, native insects suppress troublesome terrestrial weed in Florida the growth of saplings (Balciunas and Bur- I VOLUME 99, NUMBER 2 313 rows 1993). In Florida, conventional con- Townsville (19°28'S) and Cairns (16°54'S). trol measures, such as cutting, burning, and Some collections were also made in the vi- herbicides, have proven to be ineffective, cinity of Brisbane (27°30'S) in southern costly, or environmentally inappropriate in Queensland and of Cooktown (15°28'S) in managing this pest. Since 1986, a consor- northern Queensland. Plant species sampled tium of Florida and U.S. agencies has sup- were: Melaleuca arcana S.T Blake, M. ported a U.S. Department of Agriculture dealbata S.T. Blake, M. ''fluviatilis'' (Bar- project in Australia to locate, test, and ex- low 1988), M. leucadendra (L.) L., M. port potential biological control agents to quinquenerx'ia (Cav.) S.T. Blake, M. saligna help control this weed. The faunal surveys Schauer, and M. viridiflora Sol. ex Gaertner. of M. qiiinquenenna, the largest plant ever Galls from the field were either preserved targeted for a classical biological control ef- in alcohol or kept in small plastic containers fort (Balciunas et al. 1994), have found that were checked daily for emergence. Lar- more than 450 herbivorous insects associ- vae and pupae were excised from sample ated with this tree in Australia (Balciunas galls, adults were reared and their pupal ex- et al. 1995). Included among these insects uviae saved, and all specimens were pre- are gall midges that form or are associates served in 70% ethanol. Some larvae and of various leaf and bud galls. This paper adults were mounted for microscopic study treats six species of gall midges found as- in Canada balsam, using the method out- sociated with three kinds of galls (Figs. 1- lined in Gagne (1989a, 1994), some were 6) on M. quinquenen'ia and some of its prepared for SEM viewing. In the following close allies in the Melaleuca leucadendra descriptions, anatomical terminology ofthe group (Blake 1968). adult stage follows McAlpine (1981) and All six species ofCecidomyiidae are new that of the larval stage follows Gagne to science and described here. Four species (1989a). Specimens used in this study are are gall makers and the remaining two are deposited in the Australian National Insect presumed to be inquilines or successors. Collection, Canberra (ANIC) or the U.S. The gall midges causing leafblisterand bud National Museum of Natural History, rosette galls show potential as biological Smithsonian Institution, Washington, D.C. control agents and will be investigated fur- (USNM). Other abbreviations used in the ther. The leaf blister galls were the most text are for collectors in the lists of speci- commonly found in our survey and abun- mens studied: ADM = A.D. Moore; DWB dant from August to November with the = D.W Burrows; JKB = J.K. Balciunas; emergence of new leaves. Heavily infested RJG = R.J. Gagne. leaves may be rolled or otherwise distorted, Lophodiplosis Gagne, new genus but even small numbers of galls are a tax — on the host's energy. The rosette bud galls Adult. Head: Eyes large, connate, eye appear to have a more deleterious effect on bridge about 12 facets long; facets hexag- Melaleuca quinquenen'ia by killing branch onal, closely adjacent throughout. Vertex terminals. with short to long dorsal protuberance bear- ing 2-6 setae. Frons with 6-12 setae. La- Methods and Materials bella elongate-hemispherical in frontal Surveying, collecting, and rearing for view, the apices incurved and acute, each this study in Australia were done by the with 5-12 setae. Palpus 4 segmented. An- staff of the Australian Biological Control tenna with 12 flagellomeres, first and sec- Laboratory under the direction of JKB and ond flagellomeres connate, apex of twelfth DWB. Galls were collected on members of flagellomere with narrow, elongate exten- the Melaleuca leucadendra complex mainly sion. Male flagellomeres binodal, some- in coastal areas of Queensland between times weakly so, with 1 circumfilum on ba- ——— 314 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON sal node, 2 on distal node, circumfila sep- with either weak, scattered setae or strong arate or running into one another, the loops setae mixed with scales posteriorly and an- short to long. Female flagellomeres becom- terior pair of trichoid sensilla; ovipositor ing successively slightly shorter from base short-protrusible, less than P/i length of to apex of antenna, nodes parallel sided or seventh tergite, with ventral setae on inter- constricted near middle, necks prominent segmental membrane, evenly distributed se- and without setulae; circumfila made up of tae on ninth segment, with or without setae two wavy horizontal bands connected by 2 on tenth tergum; cerci short to long, more vertical bands, closely embracing node for or less bilaterally flattened, with 2-4 ven- all or most of extent. troapical, thick, setiform sensoria on each, Thorax: Scutum with 2 lateral and 2 dor- and otherwise covered with setulae; hypo- socentral rows of setae interspersed with proct va—riably shaped. sparse scales. Mesanepisternum covered Pupa. Vertex with conspicuous projec- with scales on dorsal halfto two thirds. Me- tion. Two pairs of vertexal papillae present, sepimeron with vertical row ofsetae. Wing: one of each pair with elongate seta and en- Rj curved apically to join C posterior to larged base. Antennal bases evenly rounded wing apex; C broken beyond juncture with or angular. Face with or without horns, with R5; Rs apparent as spur of R5, variable in or without pair of setae mesally. Prothorac- position; Cu forked; M3+4 present as fold. ic spiracle several times as long as wide. Tarsal claws curved beyond midlength, Abdominal segments uniformly spiculose, one- or two-toothed or simple; empodia al- without s—pines. most reaching bend in claws; pulvilli about Larva. Third instar: Body flattened-cy- one fourth length of empodia. lindrical, rounded at both ends. Integument Male abdomen: First through sixth ter- mostly rugose. Antenna less than twice as gites entire, rectangular, with posterior row long as wide. Spatula with two acutely tri- of setae, usually with lateral setae, anterior angular anterior projections, intervening pair of trichoid sensilla, and otherwise concavity smooth to minutely dentate. Pap- mostly covered with setiform scales. Sev- illar pattern generally as for Cecidomyiidi I enth and eighth tergites more weakly scler- (Gagne 1989a): lateral papillae on each side otized; seventh tergite usually with 1-sev- of spatula reduced in some species to the eral posterolateral setae, a few scales, with two setose pairs; dorsal and pleural papillae anterior pair of trichoid sensilla; eighth ter- with setae little longer than wide; terminal gite bare except for anterior pair oftrichoid segment with 2 or 4 pairs of papillae. sensilla. Cerci variable, triangular or ovoid. Second instar: Spatula present. Other- Hypoproct bilobed, variable in shape and wise as for third instar. vestiture. Aedeagus short to moderately Type species. Lophodiplosis indentata long. Gonocoxite with small, obtuse me- Gagne. sobasal lobe. Gonostylus tapered gradually Etymology. Lophodiplosis is of femi- from wide base to narrower, toothed apex, nine gender and combines the Greek "lo- with scattered setae, setulose basally, re- phos," meaning "crest" with "diplosis." mainderofsurface marked with minute lon- The prefix refers to the protruding vertex of gitudinal ridges. the pupa. The suffix has been commonly Female abdomen: First through seventh used for genera of the supertribe Cecido- tergites entire, rectangular, with posterior myiidi and means "double," with reference row or rows of setae, usually with lateral to the binodal male flagellomeres. group ofsetae, anterior pair oftrichoid sen- Relationships. Lophodiplosis belongs silla, and otherwise covered with setiform to the supertribe Cecidomyiidi. Uniquely scales; eighth tergite shorter, narrower, and derived characters (sometimes secondarily more weakly sclerotized than preceding. lost) of the supertribe are the dorsal protu- VOLUME NUMBER 99, 2 315 berance of the adult head and the binodal, ite;femaleseventhstemitenotappreciablylon- tricircumfilar male antennal flagellomeres. ger than sixth 4 This genus differs from all other known 2. Empodia much shorter than tarsal ciaws; male flagellomeres with necks; female without dor- genera ofCecidomyiidi exceptAcacidiplos- sal pair of lobes at base of intersegmental is in the development of the pupal vertex membrane of ovipositor into a protuberance or crest instead of or in Skusemyia (see Kolesik 1995a) addition to the antennal horns. This devel- - Empodia as long as tarsal claws; male flagel- lomeres without necks; female withdorsalpair opment appears to be another strategy to of lobes at base of intersegmental membrane allow pupae to cut their way out of the ofovipositor 3 galls. Although the five species placed here 3. Surface of flagellomeres covered with closely are otherwise diverse, we regard the pupal anastomozing circumfila; ovipositor short, pli- able Eocincticornia (see Kolesik 1995b) head character as a shared, derived char- - Surface offlagellomeres with only weakly an- acter until we have better knowledge of the astomozing circumfila in male but not in fe- Australian cecidomyiid fauna. The presence male; ovipositor long, needle-like, sclerotized of a similar modification in Acacidiplosis, Asphondylia (see Gagne 1994) an African genus restricted to Acacia spp., 4. Tarsal claws curved near basal third 5 - Tarsal claws curved beyond midlength 6 presumably arose separately in that genus 5. Head without vertexal protuberance .... because not all Acacidiplosis species have Diadiplosis (see Gagne 1994) it (Gagne and Marohasy 1994) and the two - Head with vertexal protuberance genera share no other obvious derived char- Feltiella (see Gagne 1995) acters. Acacidiplosis differs from Lophodi- 6. Head with vertexal protuberance; male flagel- lomeres with 3 circumfila occasionally inter- plosis in having no vertexal protuberance in connected Lophodiplosis the adult, less than four palpal segments, - Head without vertexal protuberance; male fla- and small, asetulose, dorsoventrally com- gellomeres with 2 circumfila 7 pressed and mesally joined female cerci. 7. Palpus 3 segmented, third segment inserted subapically on second; female cerci bilaterally compressed .... Zeuxidiplosis (see Harris 1966) Key to Adults of Australian Genera - Palpus 3 or 4 segmented, if with only 3, the OF THE SUPERTRIBE CECIDOMYIIDI third segment inserted apically on second; fe- male cerci dorsoventrally compressed 8 Lophodiplosis is the first genus of Ceci- 8. Abdominal second through seventh tergites domyiidi outside of the tribe Asphondyliini without lateral setae at midlength to be based exclusively on Australian spe- Stenodiplosis (see Gagne 1994) cies. Except for the Skuse species, which - Abdominal tergites with lateral setae at mid- remain essentially unknown (Gagne length Contarinia (see Harris 1979) 1989b), all the non-asphondyliine Cecido- The Species of Lophodiplosis myiidi identified from Australia to date are satisfactorily placed in cosmopolitan genera Besides the five new species keyed and (Gagne 1989b). The nine genera keyed be- described below, two Australian species re- low are doubtless only a small sample of ported earlier from Eucalyptus, a genus what will be found in Australia, but the key close to Melaleuca, may also belong to Lo- serves to place Lophodiplosis for future phodiplosis. These are ''Cecidomyia'' par- identification. Following each generic name ilis Skuse (1888: 87) and ''Cecidomyia'' is the most up-to-date and comprehensive eucalypti Skuse (1890: 381), both described reference. For a list ofAustralian species in on the basis of superficial characters. "C<?- each of the genera, see Gagne (1989b). cidomyia"" parilis was originally described from a female collected by Skuse from \. Male gonostylus situated on dorsum of gono- Middle Harbour, New South Wales. This coxite; female seventh stemite at least IVi times length of sixth stemite 2 pinned specimen, with abdomen and ap- - Male gonostylus situated on apex ofgonocox- pendages intact, is in the ANIC and labeled. 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ^ Figs. 1-6. Galls ofLophodiplosis spp. on Melaleuca spp. Figs. 1-2, Blister leafgalls ofL. indentata on M. dealbata: 1, Ix; 2, 6x, leafincross sectionshowing larval feedingchamberandexittubeafteradulthasemerged. Figs. 3-4, Rosette bud gall ofL. bidentata on M. quinquever\'ia: 3, Ix; 4, 6x, groupoflarval cells withadjacent modified leaves from center ofbud, one cell entire, three with apex dehisced following successful departure of adult (one in x-section), and one with hole made by parasitoid. Figs. 5-6, Trumpet leafgalls ofL. comuata on M. viridiflora; 5, Ix; 6, 6x, two galls, one in cross section. "Dipl. parilis, 9 type, F.A.A. Skuse, Mid- type are slide mounted, we cannot know dle Harb., S." Skuse (1890) later identified whether Froggatt's specimens really belong as this species females bred from leaf blis- to "C" parilis or even if "C." parilis fits ter galls on Eucalyptus corymbosa collected in Lophodiplosis. "Cecidomyia" eucalypti by Froggatt at Waverley, near Sydney. The known from evidently a single female leaf blisters suggest those of L. indentata ("Description drawn from fresh speci- on Melaleuca. Four female specimens im- men") from Botany, New South Wales, was C mediately follow the type of parilis in reared from woody swellings on stems of the ANIC, each bearing a single label read- Eucalyptus haemastoma. There is no spec- ing, "bred Nov. 20." These may be Frog- imen labelled eucalypti in the Skuse collec- gatt's specimens, but until these and the tion ANIC. VOLUME NUMBER 99, 2 317 Another Australian species, '"Cecido- phodiplosis denticulata, and Lophodiplosis bi- myia' frauenfeldi Schiner (1868: 7) was dentata Lophodiplosis trifida reared from rosette-like malformations cov- Key to Pupae of Lophodiplosis spp. and ering the swollen branch buds of a species Lasioptera on Melaleuca of Melaleuca found at "Naraby Lagoon" [prob. for Narabeen Lagoon per note by 1. Vertex convex, without protuberance (Fig. 64) Skuse 1888: 62], Port Jackson [Sydney, Lasioptera uncinata New South Wales]. The adults were de- - Vertex with protuberance (Figs. 44-53) .... 2 scribed as having more than 12 flagellom- 2. Frons without pair of setae anterior to labrum eres, with those of the male stemmed, m(Fuicgsh. s4h4o,rte4r6);venvterratlexprowtiutbher1ancceenstr(aFligas.nd452, which indicates they belong to neither Lo- 47) 3 phodiplosis nor Lasioptera, the two genera - Frons withpairofsetaeanteriortolabrum;ver- we have found on Melaleuca. tex withasingle medianprotuberancethatmay be divided apically (Figs. 48, 50, 52) 4 Key to Adults of Lophidiplosis spp. and 3. Vertex with central protuberance tapering to point (Fig. 44) Lophodiplosis indentata Lasioptera on Melaleuca - Vertex with central protuberance cylindrical, concave apically (Fig. 46) \. R5 about half wing length (Fig. 58); inquiline Lophodiplosis denticulata or successor in blister galls of Lophodiplosis 4. Vertex withprotuberanceshorterthanheightof indentata and trumpet galls of Lophodiplosis antennal bases (Fig. 48, 49) cornuata Lasioptera imcinata - R5 reaching costa beyond wing apex (Fig. 11) - Vertex with protuberanceLmouphcohdiplloonsgiers btihdaenntata 2 height ofantennal bases (Figs. 50, 52) 5 2. Tarsal claws simple (Fig. 10); male flagello- 5. Vertex with protuberance conical from base to meres with short, indistinct internodes and cir- slightly notched apex (Fig. 50) cumfilar loops barely longerthan wide (Fig. 8) Lophodiplosis cornuata 3 - Tarsal claws toothed (Fig. 24); male flagello- - Vertex with protuberance strongly angled for most of length, with three apical points (Fig. meres with distinct internodes and circumfilar loops much longer than wide (Fig. 21) 4 52) Lophodiplosis trifida 3. Tarsal claws amber colored; vertexal protuber- Lophodiplosis indentata Gagne, anceelongatewithapicalandlateralsetae(Fig. 7); male circumfila interconnected (Fig. 8); new species - Tfarrosmallecalfawbslisdtaerrkgbalrloswn;. .veLrotepxhaoldipprlootsuibserianndceentata —Figs. 1, 2, 7-17, 44, 45 Adult. Head (Fig. 7): Vertex with elon- shortwith only apical setae (Fig. 23); malecir- cumfilanot interconnected(as inFig. 21); from gate dorsal protuberance bearing 4-5 setae, trumpet leafgalls Lophodiplosis cornuata 2-3 at or near apex, the remainder near 4. Tarsal clawsdarkbrown, with2 teeth(Fig. 24); midlength. Male flagellomeres (Fig. 8) bin- female eighth tergite with distinct row ofsetae odal, internodes very short, necks moder- posteriorly; male cerci triangular (Fig. 25); - fTarrosmalbucdlarwosseattmebegrallcsol.or.edL,opwhiotdhipslinogsliestboiodtehntata catuemlfyillaonign,tenrocdoennsecwtietdh, sbpaasressensoettuolnae;sacmire- (Fig. 18); female eighth tergite with only scat- horizontal plane, loops short. Female fla- tered setae posteriorly; male cerci quadrate or gellomeres (Fig. 9) cylindrical with mod- secondarily lobed (Figs. 20, 41) 5 erately long necks; circumfila appressedex- 5. Vertexal protuberance elongate with lateral se- cept for short loops at apex. tae (as in Fig. 7); aedeagus shorter than hy- ploonpgreorctth(aFnigc.er2c0i);(assetianeFaitg.ba1s3e);offrfoemmalleaefcbelricsi- mmThoinramxa:leWsin(gn =(Fi1g0.),112).,5-l3e.n1gthm,m1.i9n-2f.e3- ter galls Lophodiplosis denticulata males (n = 10); Rs apparent only as spur - Vertexal protuberance short with only apical ofR5, situated closer to arculus than to apex setae (as in Fig. 23); aedeagus longerthan hy- ofRj. Tarsal claws (Fig. 10) amber colored, poproct (Fig. 41); setae at base offemale cerci much shorter than cerci (Fig. 43); possible in- untoothed. quiline ingallsofLophodiplosis indentata, Lo- Male abdomen: First through sixth ter- 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-15. Lophodiplosis indentata. 7, Male head. 8, Male third flagellomere. 9, Female third flagellomere. 10, Tarsal claw and empodium. 11, Wing. 12, Female postabdomen, seventh segmentto end, lateral. 13, Female cerci, detail. 14, Aedeagus and hypoproct, ventral. 15, Male genitalia, dorsal. — VOLUME 99, NUMBER 2 319 gites with mostly single, posterior row of dentate, the shaft narrowest at midlength, setae, partly double on sixth tergite, with 0- broadening but less sclerotized posteriorly 5 lateral setae, and otherwise mostly cov- and ending abruptly at broad base. Lateral ered with scattered setiform scales. Seventh papillae on each side of spatula reduced in tergite with 0-5 dorsolateral setae, 0-sev- most specimens to the two setose pairs, oc- eral setiform scales mesally, and an anterior casionally one orboth asetose papillae pres- pair of trichoid sensilla. Eighth tergite bare ent; dorsal and pleural papillae with setae except for anterior pair of trichoid sensilla. little longer than wide; terminal papillae re- Genitalia (Figs. 14-15): cerci more or less duced to two pairs, with setae slightly lon- acute-triangular, with several setae posteri- ger than wide; anal papillae absent. orly; hypoproct bilobed, laterally curving Second instar (based on specimen with toward venter and nestling aedeagus, with fully developed third instar inside): Body 1-2 pairs of weak setae on posterolateral shape as for third instar but shorter. Other- margins; aedeagus short, cylindrical, taper- wise as for third instar including spatula of ing slightly to rounded apex, reaching ap- similar shape. proximately to base of concavity of hypo- Holotype. 6, from blister leaf galls of proct; gonostylus with setulae present only M. quinquenervia, Australia, Queensland, at base. Tully Heads Road #3, 15.3 km ESE Tully, Female abdomen (Figs. 12-13): First 17-Vn-1995, JKB, FSNQMqn95137, de- through fifth tergites with mesally single to posited in AN—IC. laterally double, uninterrupted, posterior Paratypes. All specimens from blister row of setae, sixth and seventh tergites with leaf galls, Queensland, Australia, most de- mesally double to laterally triple row of se- posited in ANIC, with representatives de- tae; third through seventh tergites with in- posited in the USNM. From M. quinque- creasing number of lateral setae, 3-4 on nervia: 29, Forrest Beach, 16 km SE In- third to 15-20 on seventh; first through sev- gham, 6-Vn-1987, JKB, NQMqn87073; 1 enth tergites covered with setiform scales; pupal exuviae, Forrest Beach Swamp, 15.6 eighth tergite with weak, scattered setae km ESE Ingham, 4-X-1994, DWB, posteriorly and anterior pair oftrichoid sen- FSNQMqn94136; 4 larvae, Forrest Beach silla; ovipositor short-protrusible, distal half Swamp, 15.6 km ESE Ingham, lO-V-1994, about IVk times as long as seventh tergite, J.R. Makinson, FSNQMqn94058; 6, Card- with short setae ventrally on intersegmental well Swamp Site, 2 km SE Cardwell, membrane, ninth segment with evenly dis- 31-VIII-1992, em. 8-IX-1992, ADM, tributed setae; tenth tergum without setae; FSNQMqn92056; 3 larvae, pupa. Edge WNW cerci short, bilaterally flattened, closely ap- Hill, Woodward Park, 4.2 km Cairns proximated mesally with several setae, one GPO, 14-Vin-1994, R Geyson, NQMqn940 pair of setiform sensoria, and otherwise 80; 39, Centenary Park, Cairns, 3-Vin- covered with setulae; hypoproct entire, lon- 1987, JKB, NQMqn87085; larva, Tully ger than wide. Heads Rd #3, 15.3 km ESE Tully PO, — Pupa (Figs. 44-45). Vertex with one 27-VI-1994, JKB, FSNQMqn94094; 5 lar- dorsoventrally flattened, pyramidal projec- vae, Tully Heads Rd #3, 15.3 km ESE Tully tion and two shorter, ventral projections. PO, 2-Vin-1994, DWB, FSNQMqn941 14; Antennal bases with sharp, transverse, ven- 4(?, 39, 2 pupal exuviae, Tully Heads Road tral crest. Face without setae orconspicuous #3, 15.3 km ESE Tully, 17-VII-1995, JKB, protubera—nce. FSNQMqn95137 (19 more 9 and 7 p. exuv. Larva. Third instar (Figs. 16-17): In- in ale; from same lot as L. denticulata); 1 tegument mostly rugose. Spatula with two pupal exuviae, Tully Heads Road #3, 15.3 acutely triangular anterior projections, in- km ESE Tully, 14-VIII-1995, JKB, tervening concavity smooth to minutely FBNQMqn95163; 4 larvae, Murrigal Cul- 320 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON ^> 17 16 18 ^^A4. 19 21 Figs. 16-2L 16-17, Lophodiplosis indentata. 16, Larval spatula and associated papillae. 17, Larval eighth and terminal segments. 18-21, Lophodiplosis denticulata. 18, Tarsal claw and empodium. 19, Male genitalia, dorsal. 20, Aedeagus and hypoproct, ventral. 21, Male third flagellomere. vert, 14.8 km S Tully, 4-X-1994, DWB, DWB; 3 larvae, Doolandella, 16 km SSW FSNQMqn94137; 6 larvae, Murrigal Gra- Brisbane GPO, 30-V-1994, M. Purcell. vel Pit, 19.3 km S Tully, 2-VIII-1994, From M. dealbata: 14c?, Cardwell North, 3 DWB, FSNQMqn941 12; larva, Bruce Hwy, kms N Cardwell, 31-VIII-1992, em. 7-25- S of Murrigal, 16 km S Tully, 10-III-1992, VII-1992, ADM, DWB, FSNQMdl92057; 2 ADM, NQMqn92017; 5 larvae, Feluga Site larvae, Murrigal Culvert, 14.8km S. Tully, 1, 9.4 km E Tully, 27-VI-1994, JKB, 18-IX-1995, DWB, FSNQMdl95188; 2(5, FSNQMqn94092; larva, Bilyana, Double 9 2 pupal exuviae, Tully Heads Road #4, , Barrel Creek Swamp, 22 km NE Tully, 16.9 km ESE Tully, 17-VII-1995, JKB, 10-III-1992, ADM, NQMqn92018; 3 lar- FSNQMdl95135; 2 larvae, Tully Heads vae, 3 pupae, Townsville, 21-11-1995, RJG; Road #4, 16.9 km ESE Tully, 27-III-1995, 3 larvae, Feluga Site 1, 9.4 km E Tully, JKB, FSNQMd195076; 26, 9,4 pupal ex- 26-X-1994, JKB, FSNQMqn94142; 4 lar- uviae, Murray River Swamp, 12 km S Tul- vae, 4 pupae. South Queensland, XI-1995, ly, 14-VIII-1995, JKB, FSQNMdl95159; I — VOLUME 99, NUMBER 2 321 S 39, 4 pupal exuviae, Forrest Beach lata (q.v.). The gall surface is rounded ex- , School, 16 km ESE Ingham, 14-VIII-1995 cept on M. dealbata where it is usually JKB, FSQNMdl95154. From M. viridiflora: pointed in the center (Figs. 1-2). 3 larvae, Murrigal section of Bruce Hwy, This species shares with L. denticulata an 19.3 km S Tully, 2-VIII-1994, DWB, exceptionally long adult vertexal projection, FSNQMvr94115; 39,1 pupal exuviae, Fe- the male hypoproct partially enfolding the luga Site 1, 9.4 km E Tully, 18-VII-1995, aedeagus, and the lack ofpupal facial setae. JKB, FSNQMvr95140; c?, 29, 2 pupal ex- It differs from L. denticulata in having un- uviae, Feluga Site 1, 9.4 km E. Tully, toothed tarsal claws, very short flagello- 15-VIII-1995, DWB, FSNQMvr95165 (in mere internodes and shorter, interconnected same lot as specimens of L. denticulata). circumfilar loops, longer male gonopods, From M. arcana: 2 larvae, 1 pupal exuviae. and a tapering, pyramidal protuberance on Cape Flattery Swamp, 41.1 km N Cook- the pupal vertex. town, 25-VII-1994, JKB, NQMac94074. Lophodiplosis denticulata Gagne, From M.fluviatilis: 3 larvae, pupa. Charters new species Towers, 24-11-1995, RJG. From M. saligna: larvae. Flying Fox Swamp, 57.2 km WNW — Figs. 18-21, 46, 47 Cooktown, 27-VII-1994, JKB, FSNQMsl9 Adult. Head (as for Fig. 7): Vertex with 4109. elongate dorsal protuberance bearing 4-5 — Etymology. The name indentata is Lat- setae, 2-3 at or near apex, the remainder in for "untoothed," with reference to the near midlength. Male flagellomeres (Fig. simple, untoothed tarsal claws of this spe- 21) binodal, internodes and necks moder- cies. — ately long, nodes setulose; circumfila dis- Gall (Figs. 1-2). This species forms a crete, bases of each on same horizontal leafblister gall, a convex, occasionally low- plane, loops moderately and uniformly conical swelling that shows equally on both long. Female flagellomeres (as in Fig. 9) surfaces of the leaf. Leaves infested with cylindrical with moderately long necks; cir- these galls may be curled and otherwise cumfila appressed except for short loops at distorted, especially when the leaf surfaced apex. is completely covered with the galls. One Thorax: Wing (as in Fig. II), length, or more larvae may be found in a gall but 1.8-2.3 mm in males (n = 3), 2.2-2.3 mm each larva is in a discrete cell. As the larva in females (n = 2); Rs apparent only as a matures, the tissue above one end of the spur off R5, situated closer to arculus than gall changes in nature to form a cylindrical to apex of R,. Tarsal claws (Fig. 18) amber passage extending fromjust outside the lar- colored, with fine basal tooth. val cell to the abaxial surface epidermis. Male abdomen: First through sixth ter- Pupation occurs in the larval cell. The full- gites with mostly single, posterior row of grown pupa pushes its way through a thin, setae, sometimes mostly double on sixth circular cap that develops on the larval cell tergite, with 0-5 lateral setae, and other- wall, then proceeds through the tunnel and wise mostly covered with scattered setiform breaks through the thin leaf epidermis scales. Seventh tergite with dorsolateral forming the cover of the tunnel and where setae and O-several setiform scales mesally. the pupa lodges itself. The adult then breaks Eighth tergite bare except for anterior pair out of the pupa, usually leaving the pupal oftrichoid sensilla. Genitalia (Figs. 19-20): exuviae caught part way out ofthe gall exit. cerci more or less quadrate, longest at pos- Remarks. Lophodiplosis indentata was terolateral angle, with several setae poste- commonly reared from leaf blister galls on riorly; hypoproct bilobed, laterally curving several species of Melaleuca in Queens- toward venter and nestling the aedeagus, land, twice in association with L. denticu- with 1-2 pairs of weak setae on posterolat-

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