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The Psocoptera (Insecta) of Wilsons Promontory National Park, Victoria PDF

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1 Memoirs ofthe Museum ofVictoria 53(2): 137-220 (1992) 31 December 1992 https://doi.org/10.24199/j.mmv.1992.53.09 THE PSOCOPTERA (INSECTA) OF WILSONS PROMONTORY NATIONAL PARK, VICTORIA, AUSTRALIA By E. R. Schmidt and I. W. B. Thornton Department ofZoology, La Trobe University, Bundoora, Victoria 3083, Australia Abstract Schmidt, E.R. and Thornton, I.W.B., 1992. The Psocoptera (Insecta) ofWilsons Prom- ontoryNational Park.Victoria.Australia.MemoirsoftheMuseumofVictoria53: 137-220. Psocoptera(psocids)werecollectedbybeatingfromawiderangeofhabitatassociationsat Wilsons Promontory National Park, Victoria. Eleven different habitats were regularly sampledovera 13-monthperiodduring 1985-1986andadditionalhabitatswereextensively sampled at other times. Sixty-eight species were collected, representing 25 genera in 1 families. Twenty-four species are newly described and further descriptions provided for ninespecies.Thefaunaisricherthanthatfoundinsurveysofotherregionsofsouth-eastern Australiabuttheadiversityindex(a= 10.74)issimilartothatfoundatMuogamarraNature- Reserve,nearSydney,NSW.ThelargenumberofplantassociationsatWilsonsPromontory appears to be the most likely explanation for the relative richness of the psocopteran fauna. Keywords: Psocoptera; Wilsons Promontory; BassStrait zoogeography, faunal diversity. Introduction highland areas was made in early 1990, and is The Psocoptera (psocids) isone ofthe smaller now being analysed. and lesser-known insect orders. About 3500 Mackerras(1970) regardedthe insect faunaof species of36 families are known and the insects south-eastern Australia as comprising predomi- occur commonly in all zoogeographical regions. nantly the southern element of Australia's Several species are cosmopolitan and others fauna, somegroups showingaffinities with New have wide tropical ranges. Many species, how- Zealand, NewCaledonia, southern cool temper- ever, have limited distributions giving rise to ate South America and (toa lesserextent) South regional faunas. Several islands and archipel- Africa. These disjunctions appear to reflect his- agos ofthe inner and outer Melanesian arcs in torical Gondwanan connections via Antarctica. the west and south-west Pacific, New Guinea Pleistoceneloweringofsea levelwassufficient and adjacent Indonesian islands contain such to expose a land bridge, the Bassian Rise, faunas, which recently have prompted infer- connectingsouthernVictoriaviaFlindersIsland encesconcerning systematicsand biogeographi- to Tasmania (Blom, 1988; Rawlinson, 1974) cal distributions (Smithers and Thornton, 1981 (Map A). Changes in world climate, associated and references therein. 1990; Thornton et al., vegetation shifts and intermittent land connec- 1988; Thornton, 1989 and references therein). tions probably occurred often, causing repeated Surveys ofthe Muogamarra Nature Reserve, isolation and fusion of habitats conducive to nearSydney(Smithers, 1977),and ofSouth Aus- speciation. Thus, south-eastern Australia, par- tralia (Smithers, 1984) indicated that the tem- ticularlyTasmania and the Bass Strait region, is perate Australian psocopteran fauna is diverse an areawell suited for investigationsofpatterns and little known. Earlier records from Victoria ofdistribution and evolution of Psocoptera. are predominantly by New (1973a, 1973b, Thirty-one described species are known from 1974a, 1974b) and Thornton and New (1977). mainland Tasmania (Edwards, 1950; Hickman, More recently a survey of two isolated inland 1934; Smithers. 1979) and 33 from the Bass regions of Victoria, The Grampians and Mt Strait Islands(Coleetal., 1989;New, 1971).The Arapiles (Endersby et al., 1990) yielded 32 relationships of the fauna of the Bass Strait species, raising the known species in Victoria to islands to those ofTasmania and southern Vic- 61. Investigations ofthe fauna ofthe Bass Strait toria were assessed by Cole et al. (1989) but islands(Coleet al., 1989)and theOtway Ranges several species remain to be described from this (Thomas, 1986) have been made and the fauna survey. Inaddition,currentstudiesofthefaunas ofTasmania iscurrentlybeingstudied.Asurvey of both mainland Tasmania and the areas of of the fauna of south-eastern Australian southern Victoria mentioned above have 137 138 E. R. SCHMIDT AND I. W. B. THORN ION revealed a numberofundescribed species in the ago(Blom, 1988). Theestablishment ofthe land south-eastern Australian fauna. As a further bridges had facilitated faunal interchange contribution to our knowledge ofPsocoptera in between the mainland and Tasmania during the Bass Strait region, a study was made ofthe periods when the climate was colder than at psocopteran fauna of Wilsons Promontory present (Blom, 1988; Cole et al., 1989; Rawlin- National Parkbasedonasystematicsurveyover son, 1974) and when these were breached the thirteen months. Tasmanian faunawasabletodevelopin relative isolation. Rises in sea level during the Pleisto- Study area cene (approximately 120 000 years ago) turned Wilsons Promontory, a hilly peninsula 235 km Wilsons Promontory into an island. As a result south-east of Melbourne, Victoria (39°S, ofthe action ofwind and tides sand has gradu- 146°20'E), is the most southerly part ofthe Aus- ally built up the gap between the island and the tralian mainland (Map A). It was permanently mainland, forming the Yanakie Isthmus (Anon, reserved as a national park in 1905 and later 1984; Edgecombe, 1985). additions increased the area of the park to its Reed (1959) has summarised the geology of prirseessefntrosmizseeoaflaelvemlostot745940m00athetchtearheisg.heTshtepleaakn,d pWoislesdonosfPtrwoomosnetpaorrayt.eTunhiets:prgormanointteorayndisreccoemn-t MtLatrobe,andthereareanumberofotherhills deposits. Granite outcrops over most of the ofabout this height. NationalParkand formsthebedrockofthearea. Mean annual rainfall at Tidal River is 1080 Therock isa uniform grey, coarse-grained, well- mm, the maximum occurring in June, the mini- jointed porphyritic granite containing felspar mum inJanuary.ThewettestmonthsareMayto crystals of Devonian age. Recent deposits are August, the driest December to March. Tem- confined to small areas of sand dunes, alluvial peraturesare moderateand frosts rare. The pre- formations and swamps. Sand dunes are better vailing wind is from the west in all months developed on the western side than on the east, except July (north-west), August (north-west whereingeneraltheyarerestrictedtooneortwo and west) and December (west and north-east). fairly low ridges running parallel to the coast Eastern and south-easternaspectsarecharacter- (e.g., Five Mile Beach. Map B). On the western ized by lower evapotranspiration rates than side, sanddunesupto 15m highoccurontheflat western and northwestern aspects. Mean and low-lyingswampyareasat theheadsofbays monthly wind speeds show no seasonal trends, and inletswherewind-blown sand has been able varyingfrom 20to 30 km h 1, similartothoseof to accumulate (e.g., at the heads of the neigh- other exposed coastal locations in southern bouring Leonard, Norman and Oberon Bays). Australia (Parsons, 1966). Nearthesouth endofDarby Beach and on parts BassStrait hasexisted sincetheearlyTertiary. ofthe Yanakiegrasslands, thickbeds ofPleisto- During the Pleistocene the granite peaks of cene dune limestone lie under recent sand Wilsons Promontory formed part ofa range of dunes. Alluvial and swamp deposits, consisting mountains, the Bassian Rise, linking Tasmania mainly ofdetritus derived from the weathering with the mainland through the Hogan, Curtis ofgranite, extend fromthefootofthehillstothe and Kent Groups of islands to Flinders and coast formingareasofflat, low-lyingand usually Cape Barren Islands (Jennings, 1959) (Map A). badly-drained country. Fivemajorworldglacialphaseshavebeen rec- As a result of extensive analysis of aerial ognisedsincethePleistocene. In the lastthreeof photographs. Smith (1978) described 27 differ- these (the Illinoian, Early Wisconsin and Late ent vegetation unitsin Wilsons Promontory. He Wisconsin) sea levels fell below minus 80 m. described the promontory as constituting the exposing the Bassian Rise (maximum depth 55 "meeting ground" and often the distributional m) and the western land bridge extending from boundary of plant species from many parts of Tasmania through King Island to the Morning- south-eastern Australia, thus having floristic ton Peninsula (maximum depth 67 m) (Blom, links with western Victoria, Tasmania and East 1988). At the peak of the last glacial phase Gippsland. (18 0m00-20 000 years ago) the sea lay 132- Thevascularflora iswellknown,andaspecies 150 belowthepresentlevel. Sincethen thesea list has been published by the National Parks level has risen and the present coastline was Service (Anon., 1972a). Lower plants are less attained about 5 000 years ago. Western Bass well known, but some information can be Strait was breached 1 1 800 years ago and the obtained from Garnet (1971), Ashton and Bassian Rise ofeastern Bass Strait 8 700 vears Frankensberg (1976) and Ashton and Webb PSOCOPTERA OF WILSONS PROMONTORY 139 Table 2. List of collecting sites at Wilsons Promontory with grid reference, vegetation type, locality and date. Sites are shown in Map B bv the site numbers allocated in this table. Site Grid Vegetation type Locality Date No. reference 1 414778 Casuarina stricta Little Oberon Bay 2 442792 Low eucalypt woodland Telegraph Saddle 3 422798 Melaleuca ericifolia scrub Waterpump shed 4 416804 Closed heath Lilly Pilly track 5 423804 Closed scrub 6 425805 Ptericiium escu/entum 7 426807 Tall open forest is *•, ., 8 427810 Closed forest 9 415806 Low open forest 10 372857 Coastal dune vegetation Darby Beach track 11 395920 Banksia serrata Millers Landing track 12 393826 Closed heath Darby Ridge 12 Apr 1982 13 442789 Banksia sp. S ofTelegraph Saddle 3 Apr 1983 14 400933 Melaleuca ericifolia scrub Millers Landing, shore 15 397926 Open heath Millers Landing track 16 502805 Closed forest Sealers Cove track 23 Apr 1984 17 414793 l.cptospermum laevigatum Botany lab.. Tidal River 18 414796 Banksia iniegrifolia Tidal River 30 Nov 1990 19 469699 Tall open forest Roaring Meg Camping area 25 Apr 1989 20 502688 Closed heath South East Point 26 Apr 1989 21 502687 Casuarina stricta South East Point 26 Apr 1989 22 502686 Coastal vegetation South East Point 26 Apr 1989 23 499689 Banksia scrub Lighthouse lookout 26 Apr 1989 24 487694 Closed heath Lighthouse track 26 Apr 1989 25 505756 Acmena smithii Little Waterloo Bay 27 Apr 1989 26 536782 Leptospermum laevigatum Refuge Cove 27 Apr 1989 27 448926 Open heath Barry Creek 13 Apr 1990 28 468918 Closed forest Chinaman Creek 29 549949 Coastal vegetation Johnny Souey Cove 14 Apr 1990 30 540005 Banksia woodland Mt Margaret track 15 Apr 1990 31 536004 Melaleuca squarrosa scrub Mt Margaret track 15 Apr 1990 32 498042 Coastal vegetation TWin Mine Cove 16 Apr 1990 33 473995 Avicennia marina of Chinaman Swamp 16 Apr 1990 34 402933 Avicennia marina Millers Landing 17 Apr 1990 35 391910 Open scrub carpark S of Millers Landing 17 Apr 1990 36 355897 Leptospermum lanigerum Cotters Lake 18 Apr 1990 37 412802 Closed heath Pillar Pt track 10 Apr 1982 38 503862 Closed forest inland of Freshwater Lake 5 Apr 1991 39 510851 Acmena smithii southern end. Five Mile Beach 6 Apr 1991 40 511851 Coastal vegetation southern end. Five Mile Beach 6 Apr 1991 41 405804 Open heath Turnoffto Squeaky Beach 7 Apr 1991 42 403808 Closed scrub Beside main road 7 Apr 1991 43 386841 Banksia serrata Beside main road 7 Apr 1991 44 346985 Coastal vegetation Stockyard Camp 7 Apr 1991 140 E. R. SCHMIDT AND I. W. B. THORNTON Map A. Bass Strait, showing position ofWilsons Promontory in relation to the bathymetry and the Bass Strait islands. Submarine contours in fathoms. PSOCOPTERA OF WILSONS PROMONTORY 141 30\Lighthouse JohnnySouey 29k Cove Refuge Cove I 1 20 21 South 22 Ecst Point Map B. Wilsons Promontory, showing collecting sites as numbered in Table 2. Contours are 200 m and 500 m. . 142 E. R. SCHMIDT AND I. W. B. THORNTON (1977). The number of species of native and Checklist of Psocoptera from Wilsons introduced vascularplantsfound in the park(up Promontory National Park, Victoria to July 1978) is shown in Table I Lepidopsocidae The structure of the vegetation is quite diverse. In general, forests are associated with Echmepteryx (Loxopholia) albigena sp. nov. themountainsofPalaeozoicgranite. On the sur- E. (L.) renoides sp. nov. rounding Quarternary sands trees are usually Trogiidae much shorter and more widely spaced or even absent, the plant communities in these areas Cerobctsisgnestfalica (Kolbe, 1880) being dominated by shrubs and herbs. Lepinotus reticulatus Enderlein, 1905 Caeciliidae Sampling localities and methods Caecilius concavistigma sp. nov. Originally eleven of the vegetation units (habi- C. ericifoliae sp. nov. tats) described by Smith (1978) were chosen for C. juneae sp. nov. samplingon groundsofaccessibility, the need to C, ptericJii Smithers, 1977 sample the widest possible range of habitats in C. quercus Edwards, 1950 the time available, and for comparison with C. semifuscatus (Tillyard, 1923) habitats of a similar survey carried out by C. wilsoni sp. nov. Smithers (1977) at Muogamarra Nature Enderleinel/a hilli Smithers, 1979 Reserve. These habitats (sites I—11, Table 2, E. setta sp. nov. Map B) were systematically sampled over 13 Amphipsocidae months (22 Jan 1985-23 Feb 1986) on 18 occasions. On each occasion in each habitat ten Taeniostigma thekettae Smithers, 1974 samples were taken, each sample obtained by Ectopsocidae beating a branch of a tree or shrub 12 times. Wherea numberofplant specieswere present in Ectopsocus acutistigma sp. nov. a habitat, care was taken to ensure that all were E. australis sp. nov. sampled. From each of the remaining habitats E. axillaris (Smithers, 1969) (Table2) 20ofthesesamplesweretaken onceon E. briggsi McLachlan, 1899 the date shown, except those ofsites 1 5, 1 7 and E. californicus (Banks, 1903) 28, which were sampled twice, and site 14 which E. edwardsi New, 1973 was sampled on three occasions. Dislodged /:'. pieridii Smithers, 1977 insects were aspirated from the beating tray E. rileyae sp. nov. (0.7 m2) and preserved in 75% alcohol. Smith Peripsocidae (1978)providesfloristic information on all habi- tats sampled; the six-digit grid reference (Table Peripsocus bifasciatus sp. nov. 2) refers to the 1:50 000 Vicmap series of Wil- P. maoricus (Tillyard, 1923) sons Promontory. Sorting and specimen prep- P. melaleucaeNew, 1971 aration involved standard techniques (New, P. millerx(Tillyard, 1923) 1977). P. Iil/yardi New, 1973 Table 1. Number of native and introduced species of vascular plants recorded from Wilsons Promontory National Park by 1978 (Smith, 1978) Group Native Introduced Total Ferns and fern allies 60 60 Flowering plants Monocotyledons 248 35 283 Dicotyledons 433 82 515 Total 741 117 858 PSOCOPTERA OF WILSONS PROMONTORY 143 Pseudocaeciliidae P. prosta sp. nov. Austropsocus antennalis Thornton and New P. umbrata New, 1974 1977 Sigmatoneuraformosa (Banks, 1918) Tanystigma inglewoodense (New. 1974) A. cornutus sp. nov. A. costalis Thornton and New. 1977 T. valvula sp. nov. A. hyalinus Thornton. Wong and Smithers. Myopsocidae 1977 Myopsocus australis (Brauer, 1865) A. sinuosus (Banks. 1939) A. tibialis Thornton and New, 1977 Systematics A. viridis (Enderlein. 1903) In the following systematic treatment draw- Cladioneura pulchripennis Enderlein. 1906 ings from permanent preparations were made HHe.teiraocchaieacniil(iEunsdebrrluenienl,lus19(0T3i)llyard, 1923) with the aid ofa camera lucida by Ms J. Brown- ing. Measurements of body parts are given in PP.seruodlosucnodtattiaelNleawp.api1l9l7o4sa sp. nov. mm, and the method used for determining the ratio of interocular distance to diameter ofeye P. tanei Smithers. 1977 (IO:Dinsystematictreatment below)wasthatof P. yenoides sp. nov. Pearman, as described by Ball (1943). The fol- Philotarsidae lowing abbreviations are used: B, body length; ct, number ofctenidia on hind tarsal segments; Aaroniella rawlingsi Smithers. 1969 Haplophallus sinus Thornton and New, 1977 F, length of hind femur; f,, length of basal fla- gellar segment; f length ofsecond flagellar seg- Latrobiellafenestrdta sp. nov. ment; FW, lengt2h, of fore wing; HW, length of L. guttata (Tillyard, 1923) hind wing;T, length ofhindtibia;t,,t2, t3, length Elipsocidae of basal, second and apical tarsal segments Drymopsocus brunneus Smithers, 1963 respectively;rt,rationoft, andt3tot2. Localities areenumerated in Table 2. Holotypes, allotypes Pentacladus eucalypti Enderlein, 1906 and paratypes are deposited in the Australian Propsocus pulchripennis (Perkins, 1899) Museum, Sydney. Remaining material is either Spilopsoeus masseyi New, 1971 deposited in the Museum ofVictoriaorretained S. serratus sp. nov. in the authors' collections. Psocidae Lepidopsocidae Pearman Blaste bistriata sp. nov. Echmepteryx Aaron B. forficula sp. nov. Echmepteryx Aaron, 1886: 17. Type species: B. lignicola (Enderlein, 1906) impkientotnum hageni Packard. B. taylori New, 1974 B. tillyardi Smithers, 1969 Echmepteryx (Loxopholia) Enderlein Clematostigma lunulata sp. nov. Lpxopholia Enderlein, 1931: 225. Type species: C. maculiceps(Enderlein, 1903) Loxopholia pirtn—ula Enderlein = Echmepteryx (Loxo- C. striata sp. nov. pholia) pinnula, Rocsler. 1944: 133. Ptycta australis sp. nov. Remarks. Since only six species ofEchmepteryx P. campbelli sp. nov. (Loxopholia) Enderlein are known from Aus- P. glossoptera New, 1974 tralia we provide a key to the Australian P. muogamarra Smithers. 1977 species. Key to Australian species ofEchmepteryx (Loxopholia) Enderlein —1. Ocelli present 2 Ocelli absent E. howensis Smithers and Thornton 2. Fore wing apically spear-shaped (in hind wing vein r arising distal to m { — Fveoirne wi,n)g shape normal (in hind wing vein m opposite orarising dista3l x to vein /,) 5 144 E. R. SCHMIDT AND I. W. B. THORNTON 3. Fore wing with brown membrane and a hyaline fascia in apical third — 4 Fore wing barely tinged pale brown, no fasciaE. hartmeyeri Enderlein 4. Genapalecream posteriorly, largebasalbrown cloud in forewinghaving — small hyaline areas E. allngena sp. nov. Gena dark brown, large basal brown cloud in fore wing lacking hyaline areas E. quadrilineata Smithers 5. Epicranial suture distinct, dark brown; coxa, femuroflegs dark brown; fore wing membrane brown except for colourless apex, venation — distinct E. brunnea Smithers Epicranial suture not distinct; coxa, femuroflegs pale cream; fore wing membrane with brown and hyaline areas, venation not distinct E. renoides sp. nov. Echmepteryx (Loxopholia) albigena sp. nov. simple, setose. Paraproct (fig. 4) with field of 6 Figures 1-9 trichobothria and 1 seta without a basal rosette; posteriorspine blunt. Hypandrium (fig. 5) semi- Material examined. Holotype <S : Refuge Cove, circular, setose. Phallosome (fig. 6). Leptospermum laevigalUm, 27 Apr 1989. Allotype 2. Dimensions. B2.5,FW2.54, HW 1.92,F0.88, 2dand I9paratypes: samedataas holotype(K73372- X sKi7t3e32786)(.ApArdd1i9t9i0o)n.al records (29, 1 nvmph): site 19. 0.015.91,5,ft2,00..045850,, tf^,/0f.2081.00,7.t30.080, rt 6.0:1:1.0. f, Dalecsochroilp)t.ioHneaodfmcarleea.mCwoiltohrabtrioownn(amfaterrkicnag1syarsiinn Dalecsochroilp)t.ioAnsomffaelmea.le. Coloration(afterca 1 yrin fig. 1. Maxillary palps pale cream, distal halfof Morphology. IO:D=2.2. General morphology apical segment grey-brown. Genae posteriorly similar to that of male. Epicranial and frons- pale cream. Scape and pedicel grey-brown, vertex sutures more clearly discernible than in antennae brown. Fore wing (fig. 2) membrane male. In hind wing vein r2+3 2.7-5.5 times, vein brown with paler markings, veins in various >'4+5 5-9 times, length ofstalkofradial fork. Epi- shades of brown. Hind wing (fig. 3) hyaline, proct (fig. 7). Paraproct (fig. 7) with 7 (and 6) veinsbrown. Prothoraxpale. Mesothoracicnota setae in basal rosettes and 1 seta without a pale brown, pale buff anterolaterally. Meta- rosette; field of setae on mesial surface with 2 thorax pale buff, brown medially. Thoracic setae at least twice as longas remaining 18; pos- pleura cream with longitudinal grey-brown nar- terior spine blunt. Subgenital plate (fig. 8). Gonapophyses row line. Legs with coxa, trochanter and femur (fig. 9).W HW cream, femur brown apically; tibia brown with T Dimensions. B2.8,F 2.48. 1.95, F0.92, narrow buff band basally. broader buff bands 1.15, t, 0.513. t,0.166.t30.158, rt 3.1:1:1.l,f, apically and at mid-length; basal tarsal segment 0.059, f, 0.047, f|/F2 1.25.' brown over basal third, otherwise creamy buff, Remarks. This species resembles Echmepteryx second segment palebrown, apical segment pale quadrilineata in details ofpattern ofbrown pig- buff. Abdomen cream, with brown terminal ment on the fore wing membrane and in head structures. pattern. E. quadrilineata, however, lacks the Morphology. IO:D = 2.2. Antennae complete, small hyaline areaswithin the large basal brown with 38 flagellarsegments. Stout brown setaeon cloud in the fore wing, and has dark brown brown stripeson vertex, on lateral areasoffrons genae. The head pattern is very similar to the and on anterioredges ofgenae. 3 ocelli, anterior typical form of Lepidopsocusfascialus Thorn- ocellus slightly smaller than lateral ocelli. Fore ton, which, however, is clearly a member ofthe wing with asymmetrical scales on membrane, genusLepidopsocusEnderleinonwingvenation. marginal scales narrow. Both fore and hind wings (figs 2, 3) spear-shaped apically, simi- Echmepteryx (Loxopholia) renoides sp. nov. lar to Echmepteryx (Loxopholia) quadrilineata Figures 10-17 Smithers. In hind wingvein r2+3 3 times, r4+5 5.3 times,lengthofstalkofradiaffork. RaspofPear- Material examined. Holotype 9: Darby Beach man's organ well developed. Basal hind tarsal Track,coastaldunevegetation, 23-26Jun 1985. Allo- sapeigcmaelnttoowtiht,hpu1l5vcitlelnuisdinaa.rrColwa.wEwpiitphroscttou(tfigs.ub4-) thIydo,pleo5t<6yf,pne2y(mnpKyh7ms3p)3h:7s7,s-itK3e9731a3n(8dM2a)Iyd."Ap1da9dr8ai5t,tvipJoeansna:l1sr9ae8c6mo)er.ddssaitt(ea461a90s. PSOCOPTERA OF WILSONS PROMONTORY 145 f#*f|fe€ mmmm Figures 1-9. Echmepteryx albigena. Male: 1, head; 2, forewing; 3, hindwing; 4, epiproct and paraproct; 5, hypandrium;6,phallosome.Female: 7,epiproctandparaproct;8,subgenitalplate;9,gonapophyses.Figure 1 not to scale. Figures 2, 3 and 4-9 to common scales. 146 E. R. SCHMIDT AND I. W. B. THORNTON Figures 10-15.Echmepteryxrenoides.Female: 10,head; I l.forewing; 12,hindwing; 13,epiproctandparaproct: 14, gonapophyses; 15, subgenital plate. Figure 10 not to scale. Figures 11,12 and 13-15 to common scales.

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