THE GENUS GUNTHERANA (Acarina, Trombiculidae)

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1 Pacific Isects 2 (2) : July 31, 1960 THE GENUS GUNTHERN (caria, Trombiculidae) By Robert Domrow QUEENSLND INSTITUTE OF MEDICL RESERCH, BRISBNE BSTRCT The geus Gutheraa is elarged to iclude the chiggers from ustralia ad New Guiea previously assiged to Euschogastia s. 1. (except those of the subgeus Walchiella, which should be restored to geeric rak). Two subgeera, Eerrickiella ad Gutheraa s. s., are recogized o both ymphal ad larval characters, ad each cotais 2 species groups recogizable o larval characters aloe. The subgeeric divisio by larval characters parallels that by ymphal characters. Twetysix species have bee trasferred, ad 4 ew species described, brigig the total to 33, icludig the 3 species already i the geus (kallipygos, tidalei, trasluces'). Keys are give to the larval subgeera, species groups ad species, but the ymphs are too alike morphologically to be profitably keyed, except at a subgeeric level. The followig 26 larval ames are combied for the first time with Gutheraa: adromeda Womersley, atipodiaa Hirst, cassiope Wor., coorogesis Hirst, dasycerci Hirst, derricki Wor., dumosa Wor., echymipera Wor. & Kohls, foliata Guther, heaslipi Wor. & Heaslip, iisfailesis Wor. & Heas., mackerrasae Wor., mccullochi Wor., parva Wor., perameles Wor., peregria Wor., Petrogale Wor., pseudomys Wor., queesladica Wor., shieldsi Gu., similis Wor. & Heas., smithi Wor., trichosuri Wor., ewmai Wor., womersleyi Gu., & wogabelesis Wor. Four ew larval species are described from QueesladG. (D.) petulas from Rattus assimilis, Melomys cerviipes ad Hypsiprymodo moschatus; G. (Z>.) rex from R. assimilis; G. (G.) emphyla from Isoodo macrourus, Perameles asuta ad M. cerviipes; ad G. (G.) paosa from JR. assimilis. The followig 18 ames have also bee applied to the species above, but are rejected for various reasos bipygalis Guther, bushladi Philip, cairsesis Womersley & Heaslip, callipygea Gu., criita Wor., gateri Wor. & Heas., gutheri Wor. & Heas., hirsti Wor. & Heas., lappacea Wor., lawrecei Wor., melomys Wor. & Heas., mohri Wor., westraliesis Wor., paraa Wor., Phascogale Wor. & Heas., popei Wor., procaa Wor., Uromys Wor. & Kohls. ote is give o the itradermal feedig habits of the larvae of the geotype, G. kallipygos. INTRODUCTION glace at ay of the checklists of trombiculid mites published sice 1952 (the chief oes will be foud listed i udy, 1957), will show the amout of taxoomic idecisio that has atteded the trombiculie species with expaded sesillae. The importace of the larvae as possible vectors of mite typhus, ad the ease with which they may be col

2 196 Pacific Isects Vol. 2, o, 2 Species 1 petulas 2 wogabelesis 3 coorogesis 4 trichosuri 5 peregria 6 ewmai 7 dumosa 8 Petrogale 9 smithi 10 heaslipi ll dasycerci 12 shieldsi 13 pseudomys 14 similis 15 derricki 16 rex 17 perameles 18 mackerrasae 19 emphyla 20 iisfailesis 21 parva 22 adromeda 23 echymipera 24 queesladica 25 atipodiaa 26 cassiope 27 womersleyi 28 /o/wta 29 mccullochi 30 paosa 31 kallipygos \ Caudal Dorsal Plate/s Platelets j 1 ~ ~ Setae i First Dorsal Row !! io !! 6 6 I Body Setae Foliate ; I j ~" i ~ Cuticle : C C C C c c c ~~~ Table 1. Larval characters i i IL&PL 1 approx. : j j ;! _ ~~T~~ L>M rj Geualae ~ _ j ! 3 i! / =aulate; C=creulate. I the palpal formulae, b=weakly barbed, B=strogly barbed, =ude, P=plumose. I have bee able to check all the characters i the above table, except the lateral palpal tibial seta i parva, ad the dorsal ad lateral palpal tibial setae*

3 1960 Domrow: Gutheraa (Trombiculidae) 197 the geus Gutheraa. Tibiala III Mastisetae III Setae o Coxae Coxae III Closely Galeals Puctate Palpal Formula Palpal Subtermiala B /b B B B B B B b. b.. B4b b. b.. Bf4b b. b.. B4b b. b.. B4b B. b. b. B4b B. b. b. B4b b. b. b. B4b b. b. b. B4b B. b.. B4b b. b.. B4b P. b. bbb. B4b P. b. bbb. B4b P. b. bbb. B4b P. b. bb. B4b P. b. b. B4b P. b. bb. B4b B. b. bbb. B4b P. b.. B4b b. b. bb. B4b b. b.. B4b b. b. b. B4b b. b. b. B4b B. b. b. B4b b. b.. Bf 4b b. b. bbb. B4b B. b.. B4b b. b. b. B4b b. b. b. B4b b. b.. B4b b. b. b. B4b B. b.. B4b *i peregria, which have bee take from the origial descriptios. I coorogesis ad wogabelesis, the palpal geual seta may be so weakly barbed as to appear ude. I pseudomys, a suggestio of cuticular creulatio is to be see.

4 198 Pacific Isects Vol. 2, o. 2 lected, have resulted i a almost exclusively larval classificatio. May of the large geera of this system have bee broadly coceived, a result both of our lack of kowledge of the postlarval stages, ad of the appearace of polyphyletic characters i the larvae. It was ot util Wharto (1946) recogized a akamushi group withi Trombicula s. 1., Brea ad Wharto (1950) three groups withi Neotrombicula, ad the prelimiary paper of udy (1954), that the importace of defiig species groups withi the may large heterogeeous geera was fully realised. May of these groups have sice bee raised to subgeeric or geeric rak, while others, although clearly defied, must still be left i the large geera util their type species have bee reexamied. Recet years have also see a advace i our kowledge of the postlarval (particularly the ymphal) stages. This has bee a direct result of improved rearig techiques (Sasa & Miura, 1953, 1955; udy & Nadchatram, 1954). lthough it is usually impossible to idetify ymphs specifically, (ad occasioally eve geerically, e. g. scoschogastia Ewig ad Microtrombicuia Ewig, Schoutedeicha Jadi ad VercammeGradjea ad Gahrliepia Oudemas), without their correlated larval pelts, the ymphs are geerally of great value at a supraspecific level. I may cases they have cofirmed the validity of groups previously recogized o larval characters aloe, e. g., Schoutedeichia, Walchiella, Leptotrombidium ad Trombiculidus. Thus, while the preset classificatio of the Trombiculidae must be based o the larvae, the postlarval stages may be used as a guide, ad will be essetial i a fial atural classificatio (see Gordo, 1955). The species cocered i the preset paper are those from ustralia ad New Guiea which, havig passed through various " schogastia" geera, come at legth to Euschbgastia s. 1. udy ad Domrow (1957) were able to recogize several compact groups amog these species, ad gave them subgeeric or group rak. The arragemet was prelimiary, ad the desire to recogize groups of related species lead to a overclassificatio at the supraspecific level. Thus, some of their "groups" represeted othig more tha treds withi larger, more atural groups. The umber of species groups recogized below has bee reduced to four. Util 1955, ymphs of oly two of these groups were adequately kow, but represetatives of all four have sice bee reared or more closely studied. Dr. J. M. Brea has kidly give me some ymphs of Euschogastia s. s. from the New World, ad these cofirm Crossley's (1960) statemets that true Euschogastia ymphs have four pairs of galeal setae ad o precoxal plates. The ymphs described below show either of these two essetial characters, but coform i every way to the geotype of a mootypic ustralasia geus, Gutheraa. The remaiig 1, compact group of 26 ustralia ad New Guiea species of " Euschogastia s. 1. " is therefore removed to Gutheraa, which is divided ito two subgeera, each cotaiig two species groups. further 18 ames are held ivalid for various reasos, while four ew species are described. Some of the umerous extraustralasia species of Euschogastia s. 1. may also belog here, but my lack of familiarity with them precludes their cosideratio i the preset paper. I have tried to keep the followig keys as atural as possible, ad they accordigly 1. Walchiella Fuller, 1952 was also icluded i Euschogastia s. 1. by udy ad Domrow (1957). Their figures show that both the ymphs ad larvae are distict from those of Gutheraa ad Euschogastia, ad Walchiella is ow cosidered a full geus.

5 1960 Domrow: Gutheraa (Trombiculidae) 199 reflect much of the classificatio used. Characters which vary oly i degree have bee avoided, while those chose are readily visible uder high power, although oilimmersio is occasioally ecessary. It is ecessary to stress here that the larvae oe usually ecouters are those collected at least half egorged from their hosts, ad their moutig frequetly itroduces distortio, especially foreshorteig. There are two characters which I have avoided as far as possible. The first is the setatio of the palpal tarsus or tetaculum, i spite of its udoubted taxoomic importace. This has bee doe partly because it varies little i this compact group, but particularly because it is a character very difficult to see i the usual specime. Certaily, all the species here cosidered have five barbed setae ad a striate tarsala. The subtermiala, however, may or may ot be preset, but this is a most difficult character to observe. s it happes, however, i this particular group, the presece or absece of the subtermiala does ot seem to be of much diagostic value, for ymphs reared from larvae with a palpal tarsal formula either B 4b (e. g. perameles) or B 4b. S (e. g. smithi) are scarcely separable at specific level. The secod is the specialized leg setatio, which, as will be see from the table below, varies withi three of the four species groups. That this setatio is ot basically importat i Gutheraa is further idicated by a slide cotaiig six specimes of G. (G.) cassiope ad four of G. (G.) iisfailesis collected freelivig at Mudcod Creek, Bramsto Beach, orth Queeslad, 18 Sept. 1949, I. M. Mackerras. Five of the six cassiope larvae have two geualae I o both geua I, while the sixth has two geualae I o oe leg, but oly oe o the other. Likewise, the variatio i geualae I i the four specimes of iisfailesis is as follows3.3 oce, 3.2 twice, ad 2.2 oce. The figures give i the table below are the ormal values. s with the palpal subtermiala, differeces i this larval character, e. g. i G. (Z>.) shieldsi ad G. (Z>.) smithi, are ot reflected i the ymphal stage. bbreviatios: QIMR Queeslad Istitute of Medical Research, Brisbae QM Queeslad Museum, Brisbae SM South ustralia Museum, delaide SPHTMSchool of Public Health ad Tropical Medicie, Sydey USNM Uited States Natioal Museum, Washigto, D. C. Geus Gutheraa Womersley ad Heaslip, 1943 Gutheraa Wor. & Heas., Nome ovum for Gutheria Wor., 1939, ee Bleeker, 1862 (Pisces). Type species Neoschbgastia kallipygos Guther, by mootypy. Diagosis'. Trombiculie larvae from ustralia ad New Guiea with the followig characters. Legs 7.7.7, with 13 geualae I; tibiala III preset, except i emphyla. Palpi with femoral ad geual setae barbed; palpal tarsus with oe dorsal ad 4 vetral barbed setae, tarsala, ad ofte a subtermiala; palpal claw 3proged. Galeal setae ude or brached. Scutum trasversely rectagular, with clavate to globose sesillae, ad 5 setae, of which PL are usually the logest; shoulders ever preset as i Lauretella udy, 1956, so that L setae are always iserted at a level i frot of M seta, Body setae usually cylidrical, but foliate i 3 species. Dorsocaudal plate/s preset oly i kallipygos

6 200 Pacific Isects Vol. 2, o. 2 group. Itercoxal setae 2.2, except i particularly hairy species; coxal setae 1.1.1, except i adromeda ad Petrogale. Hosts i all cases mammals, although i oe species (iisfailesis} a bird has bee recorded as a casual host. The ymphs show rather less variatio, ad may be recogized by the followig combiatio of characters. Sesillae filametous, ciliated o distal 1/2 or 1/3; tectum ot detate; oe short tectal seta preset; all paracristal setae subequal; eyes abset. Galeal setae 2024 i umber, i cotrast with usual 8. Precoxal plates preset betwee coxae I. Body setae uiform. The above characters, particularly those of the ymphs, will separate Gutheraa from ay of the groups discussed by Crossley (1960), particularly Pseudoschdgastia Lipovsky, Euschdgastia Ewig, ad Euschbgastoides Loomis. LIST OF SPECIES INCLUDED IN GENUS GUNTHERN Subgeus Derrickiella Subgeus Gutheraa dumosa group coorogesis dumosa peregria Petrogale petulas trichosuri ewmai wogabelesis smithi group dasycerci derricki heaslipi mackerrasae perameles pseudomys rex shieldsi similis smithi cassiope group adromeda atipodiaa cassiope echymipera emphyla foliata iisfailesis mccullochi parva queesladica womersleyi kallipygos group kallipygos paosa Species Iquiredae tidalei trasluces KEY TO SUBGENER OF GUNTHERN Nymphs with arrow itersesillary space, ratio SL/SB Larvae with L setae shorter tha M seta; sesillae clavate, occasioally broadly so... Derrickiella Nymphs with wide itersesillary space, ratio SL/ SB Larvae with L setae much loger tha M seta; sesillae always distictly globose Gutheraa Subgeus Derrickiella udy ad Domrow, 1957 (Subgeotype Neoschogastia smithi Womersley, 1939) I separatig groups like Microtrombicula (see udy ad VercammeGradjea, i press) ad scoschogastia (i the sese of Domrow, 1957, ad 1960 b), eve the old primary character for dividig the larval geera ito 2 major groups seems to be breakig dow, amely whether filametous or expaded sesillae are preset. Similarly i ustralia species, the old divisio at couplet 18 (Womersley, 1952, p. 231) to give 2 larval groups withi his broad scoschogastia is ow believed to be artificial.

7 1960 Domrow: Gutheraa (Trombiculidae) 201 The kow ymphs of this subgeus are all from larvae with clavate sesillae, ad are idetical i all characters except oe of degree (the scutal ratio SL/SB) with those of the subgeus Gutheraa, whose larvae all have globose sesillae. I additio, some larval species with quite broadly clavate sesillae are ow kow to belog to both the species groups defied below, e. g., mackerrasae belogs i the smithi group, while the dumosa group is based o a species with very broadly clavate sesillae. The smithi group ad the dumosa group are here recogized o larval characters; they comprise the subgeus Derrickiella. Diagosis'. Nymphal scutal ratio SL/SB high (1.52.3) because of arrow itersesillary space. Larvae with L setae ever loger tha M. Sesillae clavate, occasioally broadly so. KEY TO SPECIES GROUPS ND SPECIES OF SUBGENUS DERRICKIELL (LRVE ONLY) 1. Body cuticle aulate ad coxae sparsely puctate; always with 3 geualae I dumosa species group 2 Body cuticle creulate ad coxae closely puctate ; if ot, with 2 geualae I smithi species group 9 2 (1). Te or more setae i first dorsal row 3 Eight or less setae i first dorsal row 4 3 (2). t least first 2 rows of 10 dorsal setae clearly defied; aterodorsal setae sleder ad ciliated, mergig ito shorter, stouter caudal setae with blut barbules; coxae III with usual sigle seta 7. dumosa Dorsal setae very umerous, irregularly spaced, short ad very thick, with strog blut barbs; coxae III with 2 setae 8. Petrogale 4 (2). With 8 setae i first dorsal row 5 With less tha 8 setae i first dorsal row 7 5 (4). Vetral palpal tibial seta ude; subtermiala abset 4. trichosuri Vetral palpal tibial seta brached; subtermiala preset 6 6 (5). Galeal seta ude 5. peregria Galeal seta brached 6. ewmai 7 (4). Posterior scutal margi shallowly bicovex; with 6 setae i first dorsal row 3. coorogesis Posterior scutal margi deeply covex; with 4 or 6 setae i first dorsal row (7). With 6 setae i first dorsal row; PL reachig beyod posterior scutal margi 2. wogabelesis With 4 setae i first dorsal row; PL ot reachig posterior scutal margi petulas 9 (1). L ad PL scutal setae ot approximated, so that SB lies i frot of PL; palpal subtermiala preset 10 L ad PL scutal setae approximated, so that SB lies behid PL; palpal subtermiala abset ll 10 (9). Dorsal setae set o distict platelets 10, heaslipi Dorsal setae ot set o platelets 9. smithi

8 202 Pacific Isects Vol. 2, o. 2 ll (9). Cuticle creulate 12 Cuticle aulate 16 12(11). First dorsal row with o more tha 8 setae; both pairs of itercoxal setae distict (except, perhaps, i rex); galeal setae barbed 13 First dorsal row with 1014 setae; posterior pair of itercoxal setae migled with vetral setae; galeal setae ude 15 13(12). With 8 setae i first dorsal row. 16. rex With less tha 8 setae i first dorsal row 14 14(13). With 6 setae i first dorsal row 15, derricki With 4 setae i first dorsal row 14, similis 15 (12). Dorsal setae slightly thickeed, with extremely strog, blut barbs; all 3 setae o palpal tibia ude 18, mackerrasae Dorsal setae sleder, with weak, sharp barbules; all 3 setae o palpal tibia brached 17, perameles 16(11). Dorsal setae foliate 13, pseudomys Dorsal setae cylidrical 17 17(16). Posterior scutal margi deep, ad evely rouded ll. dasycerci Posterior scutal margi oticeably flatteed, or eve slightly cocave medially 12, shieldsi The dumosa species group Diagosis: Larvae with aulate cuticle ad sparsely puctate coxae; always with 3 geualae I. Nymphs udistiguished Gutheraa (Derrickiella) petulas Domrow,. sp. Figs. 18 Types: Holotype larva i Queeslad Mus., Brisbae, from the aogeital regio of a allied rat, Rattus assimilis (Gould), collected i savaah coutry, Etty Bay Road, ear Iisfail, N. Queeslad, 25 July 1958, J. L. Harriso. The type series also icludes 30 paratypes as followsoe from foot, ad 23 from aroud aus of R. assimilis, i titree swamp, Etty Bay, 30 July 1958, J. L. H.; 3 from a scaletailed rat, Melomys cerviipes (Gould), i rai forest, Etty Bay, 7 ug. 1958, J. L. H.; 3 from a musk ratkagaroo, Hypsiprymodo moschatus Ramsay, i rai forest o hills (altitude less tha 300 m) immediately behid Etty Bay, 6 ug. 1958, J. L. H scoschogastia (Oculicola) scaevola Domrow, 1960& may also fit here, but is excluded for the preset because of its simple palpal setae. 3. Where possible, paratypes of the 4 ew larval species described i this paper have bee deposited i the Queeslad Istitute of Medical Research, Brisbae (QIMR); South ustralia Museum, delaide (SM); British Museum (Natural History), Lodo; Rocky Moutai Laboratory, Hamilto; Uited States Natioal Museum, Washigto (USNM) ; Istitute for Medical Research, Kuala Lumpur; Bishop Museum, Hoolulu.

9 1960 Domrow: Gutheraa (Trombiculidae) 203 Larva: Idiosoma 248x165 p. i uegorged specimes, ad 417 X 288 /JL i fully fed larvae. Cuticle aulate. Body setatio: Dorsal setae cylidrical, arraged , those i secod row beig 2225 /JL log, while termial setae are 1415 /JL log. Humeral setae 3639 /x log. Vetral setae about 51 i umber, those adjacet to aus 1617 f* log, ad taperig. Coxal setae arraged ; steral setae 2. 2; all fie ad taperig. Gathosoma: Cheliceral bases of usual shape, puctate i posterior 1/2, cheliceral blades large, 3639 /JL log, without teeth, but with usual tricuspid cap. Galeal setae ude. Palpal coxal setae strogly ciliated. I additio to the usual vetroexteral tarsala, the palpal formula is b. b.. B H 4b. S, /. e., subtermiala preset. Palpal claw 3proged. Scutum subquadrate, evely puctate except for small zoes behid M seta ad sesillary bases. terior ad lateral margis slightly cocave, except for fait covexity aroud M setal base. Posterior margi markedly covex, ad slightly irregular i outlie. Oly L setae o corers. Shoulders abset, M thus beig behid level of L. L setae slightly loger tha PL, but both very short. M seta lackig i all specimes. Sesillary bases fairly wide apart, set earer to level of PL setae, yet still earer aterior margi tha posterior. Brows weakly developed. Sesillae lackig i all specimes. The scutal stadard data are give i Table 2. Eyes double, o ocular plate /JL log; diameter of aterior les 89 /JL. Legs stout, sevesegmeted. ll tarsi with basal bar. The special sesory setatio is as follows: Tarsus I with pretarsala, subtermiala, parasubtermiala, tarsala 19 p log, ad microtarsala. Tibia I with 2 tibialae ad Figs. 18. Gutheraa (Derrickiella} petulas,. sp. Larva. 1. dorsum; 2, 3 ad 4, specialized setatio of legs I, II ad III; 5, veter; 6 ad 7, dorsal ad vetral views of gathosoma; 8, scutum.

10 204 Pacific Isects Vol. 2, o. 2 microtibiala. Geu I with 3 geualae ad microgeuala. Tarsus II with pretarsala, tarsala 16 p. log, ad microtarsala. Tibia II with two tibialae. Geu II with oe geuala. Leg III with oe tibiala ad geuala. W Meas 61 Table 2 Stadard data i micra of larval scutum of G. (D). petulas,. sp. PW SB SB PSB SD P M L PL Ses Notes: ll 31 specimes of this species ufortuately lack sesillae ad the M scutal seta. I feel, evertheless, that the species is correctly placed here. It may be recogized by the small umber of dorsal setae, the extremely deep scutum, ad the miute PL setae. I these latter respects it is closely similar to G. (ZX) wogabelesis, the followig species. 2. Gutheraa (Derrickiella) wogabelesis (Womersley), NEW COMBINTION. Schogastia (scoschogastia) wogabelesis Wor., 1952, S. ust. Mus., Rec. 10: 224 (Type larvae from a water rat, Hydromys sp., Wogabel, N. Q., i SM 4 ). part from the type series, I have also see 2 specimes of this species from SE Queeslad, from Rattus assimilis, Mt. Glorious, 6 ug. 1951, E. H. Derrick, ad 5 from various hosts i N. Queeslad. I all specimes there are 6 setae i the first dorsal row, while the scutal setae are strogly barbed. The PL scutal setae project well beyod the posterior scutal margi, which is flatly covex, rather tha slightly cocave medially as figured by Womersley (1952). 3. Gutheraa (Derrickiella) coorogesis (Hirst), NEW COMBINTION. Schogastia coorogesis Hirst, 1929a,. Mag. Nat. Hist. ser. 10, 3: 565 (Type larval series from a swamp rat, Rattus lutreolus (Gray), Robe, S.., i SM). Neoschbgastia cairsesis Womersley & Heaslip, 1943, R. Soc. S. ust., Tras. 67: 128 (Type larval series from a shortosed badicoot, Isoodo macrourus (Gould), 5 Brisbae, i SM). New Syoymy. 4. The documetatio of the species of this geus as here uderstood is readily available (Womersley, 1952; Guther, 1952; Wharto & Fuller, 1952; udy, 1957). I this paper I have therefore oly give the origial referece to each species ad syoym. 5. I my previous papers, this badicoot has bee referred to as Isoodo or Thylacis obesulus (Shaw & Nodder).

11 1960 Domrow: Gutheraa (Trombiculidae) 205 Neoschbgastia cairsesis var. gateri Wor. & Heas., 1943, R. Soc. S. ust., Tras. 67: 129 (Type larval series from rats, Cairs, N. Q., i SM). New Syoymy. Euschbgastia popei Wor., 1954, R. Soc. S. ust., Tras. 77: 76 (Type larval series from Rattus assimilis, Mt. Glorious, SE Q., i SM). New Syoymy. ll the above forms, icludig the variety of popei listed by Mykytowycz (1957), with dorsal setae i rows of 6, ad ude galeal setae, key out readily to the same place i the above key, ad I am uable to fid tagible characters to divide them further. Variability i the 2 cairsesis forms has already bee cosidered (Womersley, 1952), ad I feel it is better to accept the existece of a sigle, broadly recogizable species, tha several less tagible etities. The ymph has bee described by Domrow (1956). 4. Gutheraa (Derrickiella) trichosuri (Womersley), NEW COMBINTION. Neoschbgastia westraliese var. trichosuri Wor., 1939, R. Soc. S. ust., Tras. 63: 160 (Type larval series from a brushtailed possum, Trichosurus vulpecula (Kerr), Nambour, Q., i SM). This ad the ext 2 species are characterized by havig 8 setae i the first dorsal row, but trichosuri ad peregria have ude galeal setae, while ewmai has barbed galeals. G. (Z>.) shieldsi is distict from this species, its galeal setae beig brached. 5. Gutheraa (Derrickiella) peregria (Womersley), NEW COMBINTION. Schogastia {scoschogastia) peregria Wor., 1952, S. ust. Mus., Rec. 10: 220 (Type larval series collected freelivig, Wogabel, N. Q., i SM). 6. Gutheraa (Derrickiella) ewmai (Womersley), NEW COMBINTION. Schogastia westraliese Wor., 1934, S. ust. Mus., Rec. 5: 215 (Type larval series from domestic cat, Greebushes, W.., i SM). Schogastia {scoschogastia) ewmai Wor., 1952, S. ust. Mus., Rec. 10: 219 {ome ovum). I the paper i which these larvae were origially described, there was also a descriptio of a adult species, Microtrombidium westraliese (p. 188). However, i his 1952 moograph, Womersley placed these 2 species both i Schogastia, hece the ew ame for the larval species, over which the adult has page precedece. I have ot bee able to see the origial adult specime, but, as the descriptio reads "Eyes two o each side, almost sessile", the adult species may be excluded from Gutheraa. Nevertheless, the first available ame for the larval species is ewmai, sice rejected homoyms ca ever be used agai (rt. 36). This species is the oly oe i the dumosa group with barbed galeal setae i the larvae. 7. Gutheraa (Derrickiella) dumosa (Womersley), NEW COMBINTION. Schogastia {scoschogastia) dumosa Wor., 1952, S. ust. Mus., Rec. 10: 189 (Type larval series collected freelivig, therto, Q., i SM).

12 206 Pacific Isects Vol. 2, o. 2 Euschdgastia procaa Wor., 1954, R. Soc. S. ust., Tras. 77: 77 (Type larval series collected freelivig, Mt. Jukes, Q., i SM). G. (Z>.) dumosa was origially described from a sigle specime, which has bee badly damaged. Oly oe broke palp, a few crushed leg segmets, ad umerous characteristic body setae are ow clear o the slide. These, however, agree quite well with the type of procaa, as do the descriptios. The cuticle is ot creulate as i the smithi group. 8. Gutheraa (Derrickiella) petrogale (Womersley), NEW COMBINTION. Neoschogastia Petrogale Wor., 1934, S. ust. Mus., Rec, 5: 215 (Type larval series from wallaby, Musgrave Rage, S.., i SM). This species is easily recogized by the short, heavily spied dorsal setae, ad the presece of 2 setae o coxae III. The smithi species group Diagosis: Larvae with creulate cuticle ad closely puctate coxae; if ot, with 2 geualae I. Nymphs udistiguished. 9. Gutheraa (Derrickiella) smithi (Womersley), NEW COMBINTION. Figs Neoschogastia smithi Wor., 1939, R. Soc. S. ust., Tras. 63: 164 (Type larval series from Rattus assimilis, Imbil, Q., i SM). Neoschogastia Phascogale Wor. & Heas., 1943, R. Soc. S. ust., Tras. 61: larval series from a variety of marsupials from SEQ., i SM). 127 (Type New Syoymy. Schogastia {s co schogastia) lawrecei Wor., 1952, S. ust. Mus., Rec. 10: 223 Nome ovum for Neoschogastia gutheri Wor. & Heas., 1943, R. Soc. S. ust., Tras. 67: 126, ee Radford, 1942, Parasitology 34: 76. (Type larval series from rats, Cairs, i SM). New Syoymy. I SE Queeslad, this species has 69 setae i the first dorsal row. examiatio of 33 specimes showed 6 setae 10 times, eight 22 times, ad 9 oce. I N. Queeslad, the rage is somewhat greater. I have couted 300 specimes with the followig results6 (22), 7(3), 8(71), 9(69), 10(68), 11(41), 12(19), 13(6), 14 (oce). This uimodal distributio idicates that oly oe species is ivolved. I additio, these forms all show a decided preferece for the extreme ear edge as a attachmet site. The smaller forms with 6 ad 8 setae i the first dorsal row were origially described as Phascogale ad smithi, ad the larger forms as lawrecei. They are ow cosidered to be the same species. Nymphs of both forms have bee reared, ad are similar, but a descriptio of the lawrecei form is added to supplemet a earlier descriptio of the smithi form (Domrow, 19556). I the larval stage, G. (Z>.) smithi ad the followig species {heaslipi) are characterized by havig L ad PL ot approximated, so that SB lies i frot of PL. strog palpal subtermiala is preset. The galeal setae are ude or weakly barbed. Nymph. large creamcoloured species, idiosomal legth averagig 950 /JL i compressed specimes. Geital area oval, ^ (av. 99) log, with 2 pairs of geital discs;

13 1960 Domrow: Gutheraa (Trombiculidae) 207 Figs Gutheraa (Derrickiella) smithi (Womersley). Nymph. 9, geitalia; 10, aal plates; ll, precoxal plates; 12, scutum; 13, termial body seta; 14, vetral view of gathosoma. aterior discs 1930/* (av. 26) log, slightly larger tha posterior discs, which are 1925/* (av. 22) log. Geital plates weakly defied, with 811 brached setae each. Ier geital setae arraged 3.3, bbb or bb, the media pair beig less strogly brached tha the other 2 pairs, or ude (all 3 pairs are ever as strogly brached as the adjacet body setae). al plates 7085 ju (av. 76) log, more heavily sclerotized ateriorly, with 810 brached setae each. Gathosoma: Chelicerae typical, ju (av. 181) log; blade slightly serrate o cocave dorsal edge, 6270 ju (av. 66) log. Hypostome blut, with about 12 pairs of ude galeal setae to 22 p log. Basis capituli with about 18 pairs of brached setae vetrally. Palpi with usual 5 segmets; femur usually with 5 or 6 brached setae (but occasioally 4 or 7, ad eve 2 o oe side of oe specime) ; geu usually with 10 or ll brached setae (but occasioally 8 or 9 o oe side) ; tibia with sigle (2 o oe side of oe spe

14 208 Pacific Isects Vol. 2, o. 2 Figs Gutheraa (Derrickiella) smithi (Womersley). Nymph. 15, lateral view of tarsus ad tibia I; 16 ad 17, exteral ad iteral views of palp; 18, chelicera. cime) iteral brached seta ear isertio of tarsus, 2 ier accessory combs, a exteral ude seta ear base of claw, ad 4 (5 o oe side of oe specime) exteral brached setae; tarsus fairly sleder, with tarsala, 3 subtermialae, ad 9 (ll o oe side of oe specime) brached setae. Legth of palpal claw 4751 // (av. 49). Legs typical, 7segmeted. Coxae I with typical precoxal plates, each with 49 brached setae, but usually 7 or 8. Tarsus I ot elogate, ^ (av. 160) log ad 7385^ (av. 78) high. Legth tibia I f* (av. 103). Ratio LTI/HTI i most specimes apparetly However, most of the tarsi are ot quite flat, ad the specime illustrated has this ratio 1.84, which is probably a better figure. Ratio LTI/Ltl No preapical dorsal process o tarsus I. Scutum: rea sesilligera about as wide as log, puctate ateriorly, ad with posterior apodeme well developed. Crista loger tha distace betwee sesillary bases, termiatig i apparetly odetate tectum. Tectal seta shorter tha paracristal

15 1960 Domrow: Gutheraa (Trombiculidae) 209 setae, which are taperig ad subequal i legth. Sesillae log ad quite sleder, with barbules o basal 1/2, mergig ito 8 or 9 log braches o distal 1/2. Eyes abset. The stadard data are give i Table 3. Setatio: terodorsal setae brached ad taperig, 2837 ix (av. 34) log, posterior body setae evely brached, from 7685 /JL (av. 81) log. Leg setatio as usual; soleidia to 31 fj. log Meas 90 Table 3 Stadard data i micra of ymphal scutum of G. (Z>.) smithi (Wor.) CTL SL SB SL SB PSL PD TS SS SENS Material examied: ll morphotype ymphs i QIMR, all bred from egorged larvae removed from the extreme ear margis of Rattus assimilis, collected i farmlad, Mea Creek, ear Iisfail, 2 July 1958, J. L. Harriso. Correlated larval pelts accompay 10 of the ll specimes. 10. Gutheraa (Derrickiella) heaslipi (Womersley ad Heaslip), NEW COMBINTION Neo schogastia heaslipi Wor. & Heas., 1943, R. Soc. S. ust., Tras. 67: 120 (Type larval series from rats from Cairs, i SM). This species is characterized by the possessio of distict dorsal plates, o each of which is set a dorsal seta. Similar platelets have arise quite idepedetly i Walchiella traubi (Womersley, 1952). The isertios of the dorsal setae are strogly sclerotized i some species, e. g., the precedig ad G. (G.) paosa, but ever approach these platelets i size. 11. Gutheraa (Derrickiella) dasycerci (Hirst), NEW COMBINTION Schogastia dasycerci Hirst, 1929 c, Zool. Soc, Lod., Proc. 1929: 174 (Type larval series from a marsupial mouse, Dasycercus cristicauda (Krefft), Ooldea, S.., i SM). This species, ad the remaider of the smithi group which follow, may be distiguished from the precedig 2 species (heaslipi ad smithi) by havig L ad PL approximated so that SB lies behid PL. The palpal subtermiala is abset. This species is extremely close to the followig species, G. (D.) shieldsi, but may best be separated by the outlie of the posterior scutal margi. The palpal tarsus is ot clear i ay of the available specimes, but I believe the subtermiala to be abset. 12. Gutheraa (Derrickiella) shieldsi (Guther), NEW COMBINTION. Figs

16 210 Pacific Isects Vol. 2, o. 2 Neoschogastia shieldsi Gu., 1941 b, Li. Soc. N. S. W., Proc. 66: 158 (Type larval series from a scaletailed rat, Melomys rubex Thomas, Bulolo, N. G. i SPHTM). Neoschogastia hirsti Wor. & Heas., R. Soc. S. ust., Tras. 67: 123 (Type larval series from rats, Cairs ad Imbil, i SM). New syoymy. The very close similarity betwee these 2 forms has already bee metioed by udy ad Domrow (1957). The syoymy is here cofirmed. Nymph'. s i G. (D.) smithi except where oted. Idiosomal legth ot exceedig 550," i compressed specimes. Geital area 65/^ log, aterior discs 18/i log, slightly larger tha posterior discs, which are \ ju log. Geital plates with 68 brached setae each. Ier geital setae bbb. al plates 51 ju log, with about 6 brached setae each. Gathosoma: Chelicerae 108 ju log, blade 40 ju log. Hypostome with about 12 pairs of short ude galeal setae to 14 ju log. Basis capituli with 20 to 24 brached setae vetrally. Palpi: Femur with 69 brached setae, the exteral oes beig brached o both sides; geu with 10 or ll brached setae; tibia with sigle iteral brached seta ear isertio of tarsus, 2 ier accessory combs, a exteral ude seta ear base of claw, ad 4 exteral brached setae; tarsus fairly sleder, with tarsala, 2 subtermialae, ad 8 brached setae. Legth of palpal claw 25/*. Legs: Precoxal plates each with 4 or 5 (but apparetly 8 o oe side of oe specime) brached setae. Tarsus I quite elogate, /* log, to 59 ju high. Tibia I 81101// log. Ratio LTI/HTI 2.86, ratio LTI/Ltl Scutum: Sesillae very log ad sleder, with barbules o basal 2/3, which merge ito umerous quite short braches o distal 1/3. The stadard data are give i Table 4. Setatio: terodorsal setae to 26 /u log. Posterior body setae to 54 ju log, with braches o basal 4/5 stroger tha those o apical 1/5. Leg setatio as i G. ((?.) cassiope, but o microsetae see. Soleidia to 23 ju log Table 4 Stadard data i micra of ymphal scutum of G. (D.) shieldsi (W. & H.) CTL SL SB SL PSL PD TS SS SENS SB ll Material examied: 2 morphotype ymphs (together with correlated larval pelts) i QIMR; bred from egorged larvae collected i the ears of Melomys cerviipes (Gould), i rai forest o hills, Etty Bay, ear Iisfail, 21 July 1958, J. L. Harriso. Notes: This is the seveth Gutheraa ymph to be correlated with its larva. It differs from all other kow Derrickiella ymphs i havig umerous, quite short braches o the sesillae, rather tha a few log ciliatios, ad i the elogate frot tarsi, the ratio LTI/HTI beig 2.86 as agaist I the ymphal descriptios i this paper, I have used the term " subtermialae " for the setae o the palpal tarsus which I had previously called " ude apical setae. " This usage stems from the followig observatio, which idicates the homology of these setae. The 5 species {cassiope, kallipygos, perameles, shieldsi ad womersleyi), whose ymphs have oly 2 ude apical setae, have o subtermiala i the larval stage, while the 2 species {coorogesis ad smithi), whose ymphs have 3 ude apical setae, have a subtermiala i

17 1960 Domrow: Gutheraa (Trombiculidae) 211 Figs Gutheraa (Derrickiella) shieldsi (Guther). Nymph. 19, chelicera; 20, scutum; 21, termial body seta; 22, aal plates; 23, vetral view of gathosoma; 24 ad 25, exteral ad iteral views of palp; 26, lateral view of tarsus ad tibia I; 27 precoxal plates; 28, geitalia. the larval stage. Likewise the " basoexteral rod " of the ymph has bee homologized with the " tarsala " of the larva. I have already (1960 a) oted the costacy of the armature of the palpal tibia ad tarsus i the 2 subgeera of Leptotrombidium Nagayo et al. (Leptotrombidium s. s. ad Trombiculidus Radford). The patter i Gutheraa is similar, ad as follows. Tibia with a simple termial claw, which has a outer ude seta ad 2 ier acces

18 212 Pacific Isects Vol. 2, o. 2 sory combs ear its base; with a ier brached seta ear the isertio of the tarsus, ad a variable umber of brached setae exterally. Tarsus with exteral tarsala, 2 or 3 subtermialae, ad about 9 brached setae. The correlatio betwee the umber of subtermialae i the ymph ad larva has bee discussed above, but some itraspecific variatio does occur. I G. (D.) perameles, oly oe subtermiala was preset i oe out of 16 palpi where this character was clear, while the umber of brached setae o 22 palpi was as follows ll (oce), 9 (10 times), 8 (9), 6 (1), ad 5 (oce). The palp with 5 brached setae was also the oe with oly a sigle subtermiala. I the oe apparetly ormal G. (ZX) shieldsi ymph, 8 brached setae are preset o both tarsi. The secod specime is abormal. It lacks the tarsala o both palpi, ad o oe tarsus 8 brached setae ad oe ormal subtermiala are preset, while o the other are 7 brached setae ad oe ormal subtermiala, i additio to a secod stuted subtermiala about 1/2 the ormal size. 13. Gutheraa (Derrickiella) pseudomys (Womersley), NEW COMBINTION. Schdgastia {scoschogastia) pseudomys Wor., 1952, S. ust. Mus., Rec. 10: 171 (Type larval series from Trichosurus vulpecula (Kerr), Wogabel, Q., i SM). It has bee suggested (udy & Domrow, 1957; Womersley & udy, 1957) that this species might belog to the subgeus Trombiculidus Radford (geus Leptotrombidium Nagayo et al). I have, however, bee able to examie the etire type series, ad, eve i the absece of sesillae, would ow amed this. These last 3 species are closely related i a umber of characters to other species of this group, e. g. derricki. These characters iclude the plumose palpal femoral seta, the hyalie basoexteral process of the cheliceral base, ad all scutal characters. 14. Gutheraa (Derrickiella) similis (Womersley ad Heaslip), NEW COMBINTION. Neoschogastia similis Wor. & Heas., 1943, R. Soc. S. ust., Tras. series from rats, Cairs, i SM). 67: 124 (Type larval series of 39 specimes from N. Queeslad show 4 setae i the first dorsal row i all cases except oe, where 5 are preset. 15. Gutheraa (Derrickiella) derricki (Womersley), NEW COMBINTION. Neoschogastia derricki Wor., 1939, R. Soc. S. ust., Tras. Rattus spp., Imbil, Q., i SM). I have bee able to examie 8 specimes of this species. first dorsal row, the eighth has 7. 63: 162 (Type series from Seve have 6 setae i the 16. Gutheraa (Derrickiella) rex Domrow,. sp. Figs Types: Holotype larva i Queeslad Mus., Brisbae, from Rattus assimilis, collected i farmlad, Mea Creek, ear Iisfail, 8 July 1958, J. L. Harriso. The type series also icludes 55 paratypes as follows oe with same data as holotype; 12 from aogeital

19 1960 Domrow: Gutheraa (Trombiculidae) 213 regio of R. assimilis, savaah lad, Etty Bay Road, ear Iisfail, 25 July 1958, J. L. H.; 42 from R. assimilis, titree swamp, Etty Bay, 2930 July 1958, J. L. H. Larva: Idiosoma ju log, /* wide i uegorged specimes, but 414 X 270 ju whe fully fed. Cuticle creulate. Body setatio: Dorsal setae cylidrical ad well barbed, commecig , those i secod row beig 7186 /i log. Humeral setae 92104//, caudal setae 4551 ju log. Vetral setae irregularly arraged, about 32 i umber, those immediately i frot of aus 28 ju log, ad taperig. Coxal setae arraged 1.1.1; steral setae 2.2, with posterior pair close to vetral setae; all fie ad taperig. Gathosoma: Cheliceral bases stout, with posterolateral agles expaded ad hyalie; puctate dorsally; cheliceral blade 35 ju log, without teeth except for tricuspid cap. Galeal setae with several braches. Palpal coxal seta well brached. I additio to the usual vetroexteral tarsala, the palpal formula is P. b. b/b. B 4b. Subtermiala abset. Palpal claw with 3 weakly defied progs. Scutum trasverse, subrectagular, fairly evely puctate. terior margi almost straight, but with very slight covexity aroud M setal base. Lateral margis very short due to movemet of PL setae forward. Posterior margi 3sided, the posterolateral agles beig approx. 90. Scutal setae ot set o corers, but with well marked isertios; L ot o shoulders, ad thus i frot of level of M. PL loger tha M, which i tur is loger tha L, which are very short. Sesillary bases behid level of PL, with weakly marked brows. Sesillae clavate, with umerous fie barbs o oe side, ad fewer, stouter barbs o the other. The scutal stadard data are give i Table 5. Eyes double, o ocular plate 1920 ju log, adjacet to PL setae; diameter of aterior les 89/*. Legs: 7segmeted; all tarsi with basal bar. The special sesory setatio is as follows: Tarsus I with pretarsala, subtermial, parasubtermiala, tarsala 1516 ju log, ad microtarsala. Tibia I with 2 tibialae ad microtibiala. Geu I with 2 geualae ad microgeuala. Tarsus II with pretarsala, tarsala 1415 ju log, ad microtarsala. Tibia II with 2 tibialae. Geu II with oe geuala. Leg III with sigle tibiala ad geuala. W Meas 62 PW Table 5 Stadard data i micra of larval scutum of G. (D.) rex,. sp. SB SB PSB SD P M L PL Ses 42X15 42X15 Notes: This species is a typical member of the smithi species group, beig closely related to derricki ad similis. These 2 species have their dorsal setae i rows of 6 ad 4, respectively, while rex has them i rows of 8. I 110 specimes, 6 setae were preset i the first dorsal row oce, 7 twice, eight 106 times, ad 9 oce. The secod row varies commoly from 6 to 8 (six 33 times, seve ad eight 38 times). The more posterior setae are exceedigly variable.

20 214 Pacific Isects Vol. 2, o. 2 Figs Gutheraa {Derrickiella) rex,. sp. Larva. 29, dorsum; 30, veter; 31, 32 ad 33, specialized setatio of legs I, II ad III; 34, scutum; 35 ad 36, dorsal ad vetral views of gathosoma.

21 1960 Domrow: Gutheraa (Trombiculidae) Gutheraa (Derrickiella) perameles (Womersley), NEW COMBINTION. Neoschogastia perameles Wor., 1939, R. Soc. S. ust., Tras. 63: 160 (Type larval series from a badicoot from Kiamba, Q., i SM). Schogastia (scoschogastia) mohri Wor., 1952, S. ust. Mus., Rec. 10: 184 (Type larval series from a badicoot, Echymipera kalubu kalubu (Lesso), Sasapor, Dutch N. G. i SM). New Syoymy. Schogastia (scoschogastia) criita Wor., 1952, S. ust. Mus., Rec. 10: 185 (Type larval series from Echymipera kalubu kalubu, Sasapor, i SM). New Syoymy. These 3 very similar forms were all described from badicoots (except a sigle specime of mohri o a rat), ad were differetiated by havig 10, 12, or 14 setae i the first dorsal row. However, aroud Brisbae, the umber i perameles varies from 1115 (certaily 13, if oe grats it is hard to decide whether the extreme outer pair belog to the first row or the secod). I have examied all 7 of the origial series of mohri, ad the rage is I the oe specime of criita I have see, 15 are preset. Clearly oly oe species is ivolved. It is commo, ad apparetly restricted to badicoots, i coastal Queeslad ad New Guiea. The ymph was described by Domrow (1955 ). Correlated with the geeral hairiess of this ad the followig species (mackerrasae), the posterior pair of itercoxal setae are migled with the vetrals, while i the 3 less hairy species precedig, both pairs of itercoxals are distict. bormalities ivolvig the absece of ormal, or presece of additioal scutal setae are ucommo, ad the followig specime is therefore of iterest. It was take from Isoodo macrourus, at Idooroopilly, Brisbae, 20 Nov. 1951, E. H. Derrick coll. The ateroad posterolateral scutal setae are ormal, but i the place of the usual sigle ateromedia scutal seta there are 3 subequal setae set i the form of a flat isosceles triagle, with the apex to the rear. 18. Gutheraa (Derrickiella) mackerrasae (Womersley), NEW COMBINTION. Schogastia (scoschogastia) mackerrasae Wor., 1952, S. ust. Mus., Rec. 10: 182 (Type larval series foud freelivig, Bramsto Beach, N. Q., i SM). This species was origially described from a sigle specime, but I have sice see a further 2 larvae from Rattus assimilis, rai forest, Etty Bay, ear Iisfail, 7 ug. 1958, J. L. Harriso coll. dditioal descriptive otes were give by udy ad Domrow (1957). Subgeus Gutheraa Womersley ad Heaslip, 1943 (Subgeotype Neoschogastia kallipygos Guther, 1939) I spite of the extesive syoymy attedig it, G. kallipygos has always bee cosidered a species apart, eve to the extet of havig had a separate subfamily Gutheraiae erected for it (Guther, 1952). I additio to the larva, the ymph ad adult have bee kow sice World War II. However, apart from its characteristic caudal plate/s (which were the cause of its cotiued seclusio, ad which are here cosidered of group value oly), there is a close

22 216 Pacific Isects Vol. 2, o. 2 lesemblace btewee the larva of kallipygos ad those of the species placed i the cassiope group. I additio, the ewly reared ymphs of the cassiope group are idistiguishable at group level from that of G. kallipygos. The kallipygos group ad the cassiope group are here recogized o larval characters; they comprise the subgeus Gutheraa. Diagosis'. Nymphal scutal ratio SL/SB low (0.81.1) because of wide itersesillary space. Larvae with L scutal setae much loger tha M seta. Sesillae always distictly globose. KEY TO SPECIES GROUPS ND SPECIES OF SUBGENUS GUNTHERN (LRVE) 1. Caudal plate/s abset; 2 or 3 geualae I cassiope species group 2 Caudal plate/s preset; oe geuala I kallipygos species group (1) Posterior scutal margi clearly emargiate, the cocavity reachig beyod the lie of PL; tibiala III abset 19, emphyla Posterior scutal margi covex or bicovex, the media cocavity i the latter case ever reachig the level of PL; tibiala III preset 3 3 (2). Dorsal setae foliate 4 Dorsal setae cylidrical 5 4 (3). Oly PL scutal setae foliate 28, foliata Both PL ad L scutal setae foliate 29, mccullochi 5 (3). t least oe exceedigly log, basally barbed mastiseta o leg III 6 Setae o leg III ormal 7 6 (5). Oly telofemur III with mastiseta; dorsal setae i rows of 8 22, adromeda ll 4 termial segmets (except geu) of leg III with mastisetae; dorsal setae i rows of 6 21, parva 7 (5). Six setae i first dorsal row; palpal subtermiala abset 8 Eight or more setae i first dorsal row; palpal subtermiala preset 10 8 (7). Posterior scutal margi deeply covex 20, iisfailesis Posterior scutal margi shallowly bicovex 9 9 (8). PL ad PW subequal...26, cassiope PL about twice as log as PW 27, womersleyi 10 (7). ll 8 setae i first dorsal row set i a eve curve, the patter commecig , echymipera Some setae of first dorsal row moved forward to form a false row with the humeral setae ll 11 (10). Medial pair of first row of 8 dorsal setae moved forward, so that their bases ad those of the submedial pair are at the corers of a square, resultig i a setal patter commecig 4.6, ad ot 2.8. Scutum trapezoidal, with PW at least 15 fi loger tha W 24, queesladica Medial 4 of first row of 12 dorsal setae moved forward to form a row of 6 with the 2 humerals, resultig i a setal patter commecig 6.8, ad ot Scutum quadrate, with PW oly about \0 fi wider tha W 25, atipodiaa

23 1960 Domrow: Gutheraa (Trombiculidae) (1). Dorsal setal patter ; caudal plate/s with 3 weak setae each 31, kallipygos Dorsal setal patter ; caudal plates with 4 very strog setae each 30, paosa The cassiope species group Diagosis: Larvae without dorsocaudal plate/s. Two or 3 geualae I. Nymphs udistiguished. 19. Gutheraa (Gutheraa) emphyla Domrow,. sp. Figs Types'. Holotype larva i Queeslad Mus., Brisbae, from ear of Isoodo macrourus, Iisfail, 29 ug. 1958, J. L. Harriso coll. The type series also icludes 9 paratypes as followsoe with same data as holotype; 2 from a logosed badicoot, Perameles asuta Geoffroy, Iisfail, 15 Sep. 1958, J. L. H.; 6 from Melomys cerviipes, Iisfail, 6 Oct. 1958, J. L. H. Larva: specimes. Idiosoma 324x172/* i uegorged specimes, ad 547x270/* i fully fed Cuticle aulate. Body setatio: Dorsal setae cylidrical ad well barbed, arraged (5) , those i secod row beig 3136 ju, log, while termial setae are 2631 ju log. Humeral setae 4248 ju log. Vetral setae about 43 i umber, those adjacet to aus beig 2931 ju log, ad taperig. Coxal setae arraged ; steral setae arraged 2.2; all fie ad taperig. Gathosoma : Cheliceral bases stout, puctate dorsolaterally; cheliceral blade 2934 ju log, without teeth, but with usual tricuspid cap. Galeal setae ude. Palpal coxal seta strogly ciliated. I additio to the usual vetroexteral tarsala, the palpal formula is b. b. bb. B4b. Subtermiala abset. Palpal claw with 3 strog progs, the cetral prog beig the strogest. Scutum trasverse, subrectagular, evely puctate except for 2 small areas behid sesillary bases. terior ad lateral margis slightly cocave. Posterior margi markedly cocave. Oly PL setae set o oticeable corers. L setae ot o shoulders, ad therefore i frot of M seta. L slightly loger tha PL, ad M shorter tha both L ad PL. Sesillary bases fairly close together, ad much closer to level of PL tha to L; with weakly deed brows. Sesillae globose, with weak barbules scattered over surface. The scutal stadard data are give i Table 6. Eyes double, o ocular plate 1722 ju log; diameter of aterior les 68 ju. Legs 7segmeted. ll tarsi with basal bar. The special sesory setatio is as follows: Tarsus I with pretarsala, subtermiala, parasubtermiala, tarsala 1820 ju log, ad microtarsala. Tibia I with 2 tibialae ad microtibiala. Geu I with 3 geualae ad microgeuala. Tarsus II with pretarsala, tarsala 1719 ju log, ad microtarsala. Tibia II with 2 tibialae. Geu II with oe geuala. Leg IH with oe geuala oly. Notes: This species has 2 characters which set it apart from the other species of the cassiope groupthe completely cocave posterior scutal margi, ad the absece of tibiala III. These 2 characters also occur i the basically frica geus Schoutedeichia Jadi ad VercammeGradjea, However, that emphyla fits quite aturally i the geus Gu

24 218 Pacific Isects Vol. 2, o. 2 W Table 6 Stadard data i micra of larval scutum of G. (G.) emphyla,. sp. PW SB SB PSB SD P M L PL Ses 33X19 31X17 29X19 33X17 34X19 34X17 31X17 31X17 Meas X18 Figs Gutheraa (Gutheraa) emphyla,. sp. Larva. 37, dorsum; 38, 39 ad 40, specialized setatio of legs I, II ad III; 41, veter; 42 ad 43, vetral ad dorsal views of gathosoma; 44, scutum.

25 1960 Domrow: Gutheraa (Trombiculidae) 219 theraa as here defied, rather tha Schoutedeichia, is show by the followig combiatio of characterspalpal tarsal formula 5B, 3 geualae I preset, sesillae globose, L setae much loger tha M seta. However, withi the subgeus Gutheraa, iisfailesis is the oly other species i which more tha 2 geualae I may be preset. 20. Gutheraa (Gutheraa) iisfailesis (Womersley ad Heaslip), NEW COMBIN TION. Neoschogastia iisfailesis Wor. & Heas., 1943, R. Soc. S. ust., Tras. 67: 108 (Type larval series from a scaletailed rat, Melomys lutillus littoralis (Loberg), Iisfail, i SM). Neoschogastia melomys Wor. & Heas., 1943, R. Soc. S. ust., Tras. 67: 110, i part, holotype larva oly (From Melomys lutillus littoralis, Iisfail, i SM). scoschogastia bushladi Philip, 1947, Jour. Parasitol. 33: 387 (Type larval series from Megapodius sp., Owi Is., D. N. G., i USNM). This species is commo o a variety of rats ad badicoots i N. Queeslad, while it is also kow to occur o a groud bird i New Guiea. The type larva of bushladi is the oly oe of the approximately 40 types of this geus ot i a ustralia collectio. I was able to examie it ad 2 paratypes i Washigto recetly, ad ca cofirm the syoymy used by Womersley (1952) ad udy ad Domrow (1957), which was based o paratypes oly. Variatios i the specialized leg setatio of this species are discussed i the itroductory commets o the keys above. The type of bushladi has 3 geualae I. 21. Gutheraa (Gutheraa) parva (Womersley), NEW COMBINTION. Euschogastia parva Wor., 1954, R. Soc. S. ust., Tras. 77: 74 (Type larval series collected freelivig, Mt. Jukes, Q., i SM). This ad the followig species are characterized by possessig elogate, atteuated, ad basally fiely ciliated setae o at least oe of the 4 distal segmets of leg III. I this species, the mastisetae are arraged as follows, oe o telofemur, 2 o tibia, ad at least 2 o tarsus, while i adromeda, oly oe mastiseta is preset, that o the telofemur. I therefore believe that i Womersley (1954) figure 8G represts parva, rather tha adromeda. The two species were accorded group rak by udy ad Domrow (1957), but they are here retured to the cassiope group, pedig the rearig of their ymphs, which should cofirm the importace, if ay, of these mastisetae Gutheraa (Gutheraa) adromeda (Womersley), NEW COMBINTION. Euschogastia adromeda Wor., 1954, R. Soc. S. ust., Tras. 77: 80 (Type larval series foud freelivig, Mt. Tamborie, SE Q., i SM). 6. E. philippesis (Philip & Woodward, 1946a) also possesses similar mastisetae III. However, the scutum of this species shows distict aterolateral excavatios, ito which the sesillae may be withdraw (Philip & Woodward, 19466), ad i this respect resembles some of the merica species of Euschogastia (see Farrell, 1956). This species has bee recorded from ustralia (Womersley, 1952) but these specimes are i bad coditio, ad I would prefer to see fresh material before addig it to the ustralasia geus Gutheraa.

26 220 Pacific Isects Vol. 2, o. 2 part from G. (Z>.) Petrogale, this is the oly other species of Gutheraa with more tha oe seta o coxae III. 23. Gutheraa (Gutheraa) echymipera (Womersley ad Kohls). NEW COMBINTION. scoschogastia echymipera Wor. & Kohls, 1947, R. Soc. S. ust., Tras. 71 : ll (Type larval series from Echymipera kalubu kalubu, Dobadura, N. G., i SM). This ad the followig two species are the oly 3 of the 13 icluded i the subgeus Gutheraa which have a palpal tarsal subtermial. 24. Gutheraa (Gutheraa) queesladica (Womersley), NEW COMBINTION. Neoschbgastia queesladica Wor., 1939, R. Soc. S. ust., Tras. 63 : 162 (Type larval series from various ative rats, SE Q., i SM). I additio to havig a palpal tarsal subtermiala, this ad the followig species are further characterized by the fact that some cetral setae of the first dorsal row have moved forward to form a false row with the humerals. This chigger is commo o a variety of rats ad marsupials i N. ad NE ustralia. I have also see specimes from the eyelids of rabbits, Colo Vale, New South Wales, ug. 1959,. L. Dyce coll. There are also further specimes from rabbits i the South ustralia Museum. These are mouted with, ad labelled, as, atipodiaa, "Bargibel," ear Budarra, N. S. W., E. J. Waterhouse. 25. Gutheraa (Gutheraa) atipodiaa (Hirst), NEW COMBINTION. Schogastia atipodiaum Hirst, 1929c, Zool. Soc, Lod., Proc. 1929: 175 (Type larval series from Rattus greyii (Gray), Kagaroo Is., S.., i SM). I additio to the origial series, this species has also bee recorded from rabbits from New South Wales (Mykytowycz, 1957). The specimes referred to this species by Domrow ad Smith (1956) are o loger available, but almost certaily should have bee referred to G. (G.) queesladica. 26. Gutheraa (Gutheraa) cassiope (Womersley), NEW COMBINTION. Figs Schogastia (scoschogastia) cassiope Wor., 1952, S. ust. Mus., Rec. 10: 187 (Type larval series collected freelivig, Wodecla, N. Q., i SM). Schogastia (scoschogastia) lappacea Wor., 1952, S. ust. Mus., Rec. 10: 190 (Type larval series from rat, Cairs, N. Q., i SM). New syoymy. The holotype of melomys Wor. & Heaslip is a specime of iisfailesis Wor. & Heaslip (see above), but the paratypes are cassiope. The idetity of cassiope ad lappacea was suggested by udy ad Domrow (1957), ad a subsequet examiatio of some hudreds of larvae from a variety of ative rats ad marsupials from coastal Queeslad has cofirmed this syoymy. The cassiope form was origially characterised by possessig strog, thick dorsal setae, but the gradatio to the slightly thier (ad more usual) shape i the lappacea form is complete. I other respects the larvae are iseparable.

27 1960 Domrow: Gutheraa (Trombiculidae) 221 I additio to umerous specimes from N. Queeslad, I have also see material collected free livig from Mt. Jukes, ear Mackay, 6 Sep. 1951, E. H. Derrick; ad Brookfield, Brisbae, 21 Feb. 1951, E. H. D. further 2 specimes from New South Wales have recetly come to had. These were collected with about 30 comatacarus sp. larvae o the edges ad iside the ears of a blackstriped wallaby, Protemodo dorsalis (Gray), Gorge Creek, Boalbo, 20 Feb. 1960, J. H. Calaby. iterestig scutal abormality has bee oted i a larva of this species from Melomys cerviipes, Iisfail, 2 Oct. 1958, J. L. Harriso coll. Ulike the abormality reported above for G. (Z>.) perameles, the 5 scutal setae i this specime are ormal. However, although the left sesilla is quite ormal, there is o trace of the right oe, or of the ivolucrum i which it should have bee set. The texture of the scutum i the right half is uiformly puctate. Variatio i the specialized sesory setatio of geu I of this species has bee discussed i the itroductory commets to the keys above. Nymph: very pale yellow species, with idiosomal legth ot exceedig 612 ju i compressed specimes. Geital area oval, averagig 59 ju i legth, with 2 pairs of subequal, oval geital discs. terior discs 1117 ju log (av. 14 /JL) ; posterior discs 1115 ju, log (av. 13 /u). Geital plates ot well defied, with about 8 brached setae each. Ier geital setae arraged 3.3 i 3 specimes; b or bb. I fourth specime 2.2, b ad bb. al plates elogate, 46 / * log, with 46 setae each. Gathosoma: Chelicerae typical, 83 /JL log; blade slightly serrate dorsally, 33 ju log. Hypostome blut, with about 12 pairs of short ude galeal setae to 12^ log. Basis capituli with about 15 pairs of brached setae. Palpi rather sleder, with usual 5 segmets; femur with 5 or 6 brached setae (some of which are brached o both sides) ; geu with 7 or 8 brached setae; tibia with sigle iteral brached seta ear isertio of tarsus, 2 ier accessory combs, a exteral ude seta ear base of claw, ad 25 (average 3.6) exteral brached setae; tarsus fairly stout, with tarsala, 2 subtermialae ad 9 brached setae arraged as figured. Legth of palpal claw 26 ju. Legs typical, 7segmeted. Coxae I with ier agle cut off by trasverse strut to form typical precoxal plates; each plate with 4 or 5 brached setae. Tarsus I of usual shape, 87 ju log; tibia I 54 ju log; ratio LTI/Ltl No preapical dorsal process o tarsus I. Scutum: rea sesilligera very broad, beig wider tha legth of crista, ad ot produced far behid level of sesillary bases. Crista short, termiatig i odetate tectum. Tectal seta much shorter tha crista. Sesillae very log ad filametous, with barbules o basal half, which merge rapidly ito about 15 log ad very sleder braches distally. Eyes abset. Paracristal seta about 6 i umber, ad subequal. The stadard data are give i Table 7. Setatio: Setae o aterior part of dorsum short ad taperig, to 19 ju log. Posterior body setae very log ad siuous, form Table 7 Stadard data i micra of ymphal scutum of G. (G.) cassiope (Wor.) CTL SL SB SL SB PSL PD TS SS SENS Meas

28 222 Pacific Isects Vol. 2, o. 2 Figs Gutheraa (Gutheraa) cassiope (Womersley). Nymph. 45, chelicera; 46, aal plates; 47, geitalia; 48 ad 49, exteral ad iteral views of palp; 50, lateral view of tarsus ad tibia I; 51, precoxal plates; 52, scutum; 53, termial body seta; 54, vetral view of gathosoma. ig a frige aroud opisthosoma to 116/; log, ad evely brached alog their etire legth. Leg setatio typical, /. e., i additio to usual brached setae there are preset both figerlike soleidia of varyig legth ad thickess, ad ude taperig setae. These 2 types of sesory setae are, as usual, more umerous o the distal segmets of aterior legs. Soleidia to 17 /JL, brached setae to 22 /J. log. microseta is occasioally preset o tibia I. Material examied: Four morphotype ymphs i QIMR; 2 bred from larvae from a giat scaletailed rat, Uromys caudimaculatus (Krefft), Palmersto, N. Queeslad, 8 May 1958, M. L. Emauel coll., ad 2 obtaied i the same way from Rattus assimilis, Palmersto, 20 May 1958, M. L. E. Correlated larval pelts accompay 3 of the 4 specimes. Method: I these 2 breedig experimets, the ear cochae of the hosts were airfreighted south from Iisfail, N. Queeslad, to Brisbae (1360 km), ad were received oe or 2 days after dispatch. few egorged larvae detached durig trasit, ad other swolle specimes were removed with a eedle. These larvae were the placed o moist, sterile alumiium oxide (aludum) i a small sealed cotaier, so that codesed droplets

29 1960 Domrow: Gutheraa (Trombiculidae) 223 of moisture were always preset iside the cotaier. The temperature rage was 1828 C. (6483 F.) The larvae are small, eve whe egorged, ad very pale yellow i colour, readily visible o the dark aludum. The larvae from U. caudimaculatus were set up o 8 May Two ymphochrysalises were oted 22 days later (30 May), but these failed to moult. Fially o 19 Jue, the larval pelts were dissected away to yield 2 pale yellow, but dead ymphs. These were of small size, but the posterior circlet of log hairs was oted before they were mouted. These 2 ymphs had probably died i the first week of Jue soo after the completio of metamorphosis withi the larval pelt. few larvae were also obtaied from R. assimilis i the same way o 20 May Oe ymphochrysalis was oted 30 days later (19 Jue), ad was see i the process of moultig o 23 Jue. Moultig, however, was ot successful, ad the ymph ad pelt were mouted o a sigle slide. secod ymphochrysalis still seemed alive o 27 Jue, but had died by 3 July, ad was also mouted together with its larval pelt. These ymphochrysalises showed a distict forwardly directed aterodorsal spie as figured by Joes (1951) for Neotrombicula autumalis (Shaw), ad oted by Domrow (1956) for T. atechius (Wor.). The opisthosoma was quite poited posteriorly prior to moultig. The ymphs of Gutheraa (Derrickiella) were also reared by this method. smithi ad G. (Z>.) shieldsi described above 27. Gutheraa (Gutheraa) womersleyi (Guther), NEW COMBINTION. Figs Neoschogastia womersleyi Gu., 1940, Li. Soc. N. S. W., Proc. 65: 254 (Type larval series from a pademelo, Thylogale coxeii (Gray), Bulolo, N. G., i SPHTM). This species is very close to G. (G.) cassiope i both the larval ad ymphal stages. The similarity of the dorsal setal patter ad scutal shape i the larvae is strikig, ad the oly real differece appears to be i the relative legths of the PL scutal setae. I additio to the type series, this species has also bee collected o "rats" i New Guiea. The ymph described below was reared from this latter material. Nymph: s i G. (G.) cassiope except where oted. Colour i life ukow Idiosomal legth ot measurable i specime at had, but the scutal stadard data are almost idetical with those give for G. (G.) cassiope above. Geital area oval, 64 ju log, aterior discs 17 JJL log, posterior discs 14 ji log. Geital plates with about 9 brached setae each. Ier geital setae bbb. al plates 47 ju log, with 6 or 7 brached setae each. Gathosoma: Chelicerae 82^ log, blade 31 ju log. Hypostome with about 12 pairs of ude galeal setae to 12 JJ. log. Basis capituli with about 25 brached setae. Palpi a little stout; femur with 7 or 8 brached setae; geu with 9 or 10 brached setae; tibia with sigle iteral brached seta ear isertio of tarsus, 2 ier accessory combs, a exteral ude seta ear base of claw, ad 4 exteral brached setae; tarsus fairly stout, with tarsala, 2 subtermialae, ad 8 brached setae o oe palp ad 9 o the other. Legth of palpal claw 25 ju. Legs: Precoxal plates with 6 brached setae each. Tarsus I 96 ju log, of usual shape, but tarsal ratio uavailable because height of tarsus I umeasurable. Scutum: Sesillae lackig i available specime. Paracristal setae about 9 i umber, subequal. The stadard data are give i Table 8. Setatio: terodorsal body setae 17 p log; posterior body setae log ad sleder, 59 ju log. Leg soleidia to 20 ju log.

30 224 Pacific Isects Vol. 2, o. 2 Figs Gutheraa (Gutheraa) womersleyi (Guther). Nymph. 55, chelicera; 56, aal plates; 57, geitalia; 58 ad 59, iteral ad exteral views of palpi; 60, lateral view of tarsus ad tibia I; 6\ y precoxal plates; 62, scutum; 63, termial body seta; 64, vetral view of gathosoma. Table 8 Stadard data i micra of ymphal scutum of G. (.G) womersleyi (Gu.) CTL SL SB SL PSL PD TS SS SENS SB Material examied: Oe ymph labelled u Neoschogastia womersleyi Guth. Nymph ex larvae from rat, Dobadura, N. G., ll , G. M. K. # 294 (ee 293). This is probably Gutheraa bipygalis [ = G. (G.) kallipygos]." Notes: The ymph before me is oe of 2 idetified by Kohls as womersleyi, but later described uder Gutheraa kallipygos by Womersley (1952). s both G. (G.) womersleyi ad G. (G.) kallipygos are characteristic species i the larval stage, Kohls' idetificatio is accepted. G. (G.) cassiope ad G. (G.) womersleyi are very closely related i the larvae ; likewise i the ymphs the oly differece seems to be i the legth of the termial body setae.

31 1960 Domrow: Gutheraa (Trombiculidae) Gutheraa (Gutheraa) foliata (Guther), NEW COMBINTION. Neoschogastia foliata Gu., 1940, Li. Soc. N. S. W., Proc. 65: 255 (Type larval series from Thylogale coxeii, Bulolo, N. G., i SPHTM). This ad the followig species comprise, i essece, the foliata group of udy (1954). However, as the resemblace betwee the foliata series ad G. (G.) cassiope is quite close, ad the gradatio from cassiope Xo mccullochi complete, the 2 species are here retured to the cassiope group. ll these species have a palpal formula b. b. /b. B4b (the formulae give by Womersley ad udy, 1957, are i error). Two New World geera with globose sesillae ad foliate dorsal setae should be cosidered here, if oly to exclude them. These are Cordiseta Hoffma, 1954 ad Trombewigia da Foseca, Womersley ad udy (1957) cofirm da Foseca's statemet that Trombewigia has oly 4 setae o the palpal tarsus, while Cordiseta does ot agree with Gutheraa i scutal patter. Both are atedated by Gutheraa. 29. Gutheraa (Gutheraa) mccullochi (Womersley), NEW COMBINTION. Neoschogastia mccullochi Wor., 1944, R. Soc. S. ust., Tras. 68: 100 (Type larval series collected freelivig, dibari, N. G., i SM). scoschogastia uromys Womersley ad Kohls, 1947, R. Soc. S. ust., Tras. 71: 10 (Type larval series from Uromys caudimaculatus aruesis Gray, Dobadura, N. G., i SM). New syoymy. I have examied both the type series above, ad fid that the structure of the dorsal setae is idetical i all specimes. They are foliate, ad apparetly smooth o the vetral surface. The flatteed lateral margis are very strogly detate as i Womersley's fig. 7E (1944), while dorsally there is a secod row of several much weaker barbules alog either side of the shaft proper. The kallipygos species group Diagosis'. Larvae with dorsocaudal plate/s preset. Oe geuala I. Nymphs udistiguished. 30. Gutheraa (Gutheraa) paosa Domrow,. sp. Figs. 6572, 76. Types'. Holotype larva i Queeslad Mus., Brisbae, from Rattus assimilis, collected i farmlad, Coorumba, ear Iisfail, 15 July 1958, J. L. Harriso. Larva: Idiosomal measuremets uavailable because of fracture durig moutig procedure. Cuticle aulate. Body setatio: Dorsal setae cylidrical ad strogly barbed, arraged , those i secod row beig 56 ju log, while termial setae are as short as 28 ja. Humeral setae 43 ju log. Caudal setae set o miute basal platelets. Dorsum posteriorly with 2 elogate, opuctate, caudal plates, which are 104 p. log ad 62 p wide. These bear 4 setae each, oe o the ier margi 95 fu log, ad 3 o outer margi 99, 87 ad 78 ju log (from frot to back). Vetral setae 29 i umber, those immediately i frot of aus 29 ju log. The more aterior setae are fie ad taperig, while the posterolateral setae are coarse, ad resemble the caudal setae. Coxal setae arraged

32 226 Pacific Isects Vol. 2, o. 2 *k Figs Gutheraa {Gutheraa) paosa,. sp. Larva. 65, dorsum; 66, veter; 67, 68 ad 69, specialized setatio of legs I, II ad III; 70, scutum; 71 ad 72, dorsal ad vetral views of gathosoma.

33 1960 Domrow: Gutheraa (Trombiculidae) ; steral setae arraged 2.2; all fie ad taperig. Gathosoma: Cheliceral bases stout, puctate dorsally; cheliceral blade 29 ju log, without teeth, but with usual tricuspid cap. Galeal setae ude. Palpal coxal seta with several ciliatios. I additio to the usual vetroexteral tarsala, the palpal formula is b. b. b. B4b. Subtermiala abset. Palpal claw with 3 weak progs, of which the middle oe is strogest. Scutum trasverse, rectagular, evely puctate except for small area behid each sesillary base, ad a more extesive zoe behid M seta. terior margi cocave, but with slight covexity aroud base of M seta. Posterior margi slightly covex, but with very weak medial emargiatio. L ad PL setae set o distict corers, L ot beig o "shoulders", ad cosequetly slightly i frot of M setal base. Both PL ad L setae much loger tha M, but PL somewhat shorter tha L. Sesillary bases fairly close together, much earer posterior tha aterior margi; flaked ateriorly by well marked brows, ad posteriorly by weakly sclerotized lie. Sesillae globose, with several weak barbules o surface. The scutal stadard data are give i Table 9. Eyes double, o ocular plate IS ju log ear bases of PL setae; diameter of aterior les 5 ju; posterior les very weakly developed. Legs 7segmeted, but femur III oly divided vetrally, the vetral seta beig accordigly ormally developed ad barbed. ll tarsi with basal bar, although weak i leg III. The special sesory setatio is as follows: Tarsus I with pretarsala, subtermiala, parasubtermiala, tarsala 14 ju log, ad microtarsala. Tibia I with 2 tibialae ad microtibiala. Geu I with oe geuala ad microgeuala. Tarsus II with pretarsala, tarsala 17^ log, ad microtarsala. Tibia II with 2 tibialae. Geu II with oe geuala. Leg IH with sigle tibiala ad geuala. Table 9 Stadard data i micra of larval scutum of G. (G.) paosa,. sp. W PW SB SB PSB SD P M L PL Ses X18 Notes: That this form is a Gutheraa i the strictest sese is certai, but, as it coforms either i the characteristic dorsal setatio or i the caudal plate/s with G. kallipygos, it is described as a ew species. It may be recogized by the followig 3 characters dorsal setal patter (omittig caudal setae) , caudal plates with 4 pairs of very log setae, 2 strog setae set betwee the caudal plates. 31. Gutheraa (Gutheraa) kallipygos (Guther) Figs. 7375, Neoschogastia kallipygos Gu., 1939a, Li. Soc. N. S. W., Proc. 64: 83 (Type larval series from rats ad badicoots, N. G., i SPHTM). Neoschogastia callipygea Gu., 1938, Med. Jour. ust. 2: 204 ome udum. Neoschogastia bipygalis Gu., , Li. Soc. N. S. W., Proc. 64: 471 (ome ovum. This actio was uecessary, as the preoccupyig ame kallipygos Derrick et al was a ome udum). 68: 100 (Type larvel series col Gutheraa paraa Wor., 1944, R. Soc. S. ust., Tras. lected freelivig, dibari, N. G.). The larvae of this iterestig species occur o a variety of rats ad badicoots from New Guiea (Bulolo) to SE Queeslad. ote o their feedig habits is give below.

34 228 Pacific Isects Vol. 2, o. 2 Morphologically, this is a variable species, ad I agree with Womersley (1952) that G. paraa Wor., 1944 is a syoym. The least variable, ad most useful character i idetifyig this species, apart from the caudal plate/s, is the peculiar arragemet of dorsal setae, Two poits are of ote here, the short third row of 4 setae, which lies closely about a media lie, ad the fifth row of 2 very log setae set close together immediately i frot of the caudal plate/s. This caudal sclerotizatio takes the form of oe or 2 plates of variable structure. The 3 forms illustrated below all occur i the Bramsto BeachTisfail area of orth Queeslad. I the extreme form, the 2 plates are completely fused to form a trasverse oval, puctate shield this is the form described as G. paraa. I the itermediate coditio, the plates are divided medially, but still strogly puctate. The 3 pairs of setae o the aterolateral margis are somewhat variable i legth, ragig from 25 to l ji log. The extreme i the other directio is reached i the type of kallipygos. Here we fid 2 widely separated plates, a lack of puctae, ad 3 pairs of miute setae barely 10 ju log. The caudal sclerotizatio i all 3 forms, the, has a basic patter, ad as they also possess the same charateristic dorsal setatio, I cosider them oe species. I the same year as Guther (19396) described the larva of this species, he also Figs Gutheraa species. Caudal plates of larvae. 73, 74, 75, G. (G.) kallipygos (Guther) ; 76, G. (G.) paosa,. sp.

35 1960 Domrow: Gutheraa (Trombiculidae) 229 Figs Gutheraa (Gutheraa) kallipygos (Guther). Nymph. 77, chelicera; 78, scutum; 79, lateral view of tarsus ad tibia I; 80 ad 81, iteral ad exteral views of palpi; 82, aal plates; 83, geitalia; 84, termial body seta; 85, vetral view of gathosoma; 86, precoxal plates. published a ote o some trasparet structures foud "cemeted" to the hairs of the host. Each cotaied a dark mass or a sigle larva, ad he cocluded that they were

36 230 Pacific Isects Vol. 2, o. 2 Fig. 87. Gutheraa (Gutheraa) kallipygos (Guther). Larva eclosed i modified suckig tube. The followig structures of the mite, which is displaced to the right, are also visible the left half of the scutum with sesilla ad PL seta, the left humeral seta, the left 4 of the first row of 6 dorsal setae, at least 3 of the secod row of 4 setae, all 6 setae i the third row, the 2 elogate setae at the aterior ed of the cleft betwee the 2 caudal plates, most of the 6 legs, ad at least 4 coxae. Four hairs are embedded i the capsule. udifferetiated or late stage ova, respectively. He was uable to decide whether the adult mites lived i the est of the host, or whether, like the larvae, the adults i this

37 1960 Domrow: Gutheraa (Trombiculidae) 231 oe case were also true parasites. He later (1952) raised a ew subfamily Gutheraiae for this oe species o the basis of these fidigs, ad o a distictive larval morphological character, the presece of 2 dorsocaudal plates. Doubt was later cast (Womersley, 1952; udy, 1954) o the belief that these structures were eggs. Capsules similar to those described by Guther have sice bee take o 3 occasios. I have draw freely o ms otes made by Dr. E. H. Derrick i 1953 i recordig the first of these 3 fidigs. The belief that these structures were eggs was ot abadoed i these otes, while the typical histosiphos were called "pseudosuckig tubes." I have had accordigly to alter the wordig slightly i oe or 2 places. female badicoot {Isoodo macrourus) was captured at Mt. Nebo, 23 km NW of Brisbae, ad brought to this Istitute o 16 pr While its ectoparasites were beig removed by Dr. Derrick, some fiely graular, scaly areas were oticed o the aterior aspect of its hid limbs. This scaly material was scraped off, ad placed i polyviyl alcohol medium. It separated ito a umber of idividual oval graules rather less tha 1 mm i legth (4 specimes measured 720 x 660, 816x684, 840x600 ad 1140x600/0. Whe these were examied with the microscope, several were see to cosist of a egorged mite surrouded by a mass of tough, fiely fibrillary substace. The ecapsulatio of the mite was icomplete posteriorly, ad o applyig pressure to the coverslip, the mite teded to extrude here. Projectig forward from the chelicerae of the mite, a hyalie suckig tube was clearly visible. Like the mite itself, this tube was also surrouded by the fibrillary substace, ad its aterior ed just reached the exterior of the capsule. Cotiuous with the wall of the suckig tube, ad cosistig of the same material, was a hyalie layer that exteded backwards aroud the body of the mite. Towards the posterior ed, it thied ad disappeared. Its hyalie ature distiguished it readily from the fibrillary substace, to which it was closely applied. I most of the capsules the mite was missig, but a cavity idicated where oe had bee. suckig tube, usually shorter tha the oe illustrated, was also preset i these specimes, as well as the hyalie layer cotiuous with it that bouded the cavity. I have sice take aother series of capsules from Perameles asuta, Mt. Nebo Road, 1 Dec. 1958, E. H. Derrick. The capsules were first oted attached to the shafts of hairs well out from the ski about the rump of the host. Soo some specimes were foud attached to the hair quite close to, ad almost touchig, the ski. These were examied uder a dissectig microscope to see which ed of the capsule was earer the ski, ad it was foud that the poited, ipplelike ed was. The may miute holes, of a size to admit the poit of a small pi, were oted i the ski. The edge of oe of these holes was grasped with very fie forceps, ad at a slight pull, a complete capsule was extracted from the ski. This process was repeated may times, ad each of the holes proved to be the ope ed of a capsule. The capsules were completely embedded i the ski, their apertures beig at ski level. Very little, if ay, swellig or iflammatio surrouded the holes. O moutig, these empty capsules also showed typical histosiphos peetratig their poited aterior eds, ad occasioally traces of larval structures were see i their cavities, as well as opaque deposits. These deposits therefore appear black i specimes examied by the compoud microscope, but i umouted material ad usig reflected light,

38 232 Pacific Isects Vol. 2, o. 2 they are white, ad much resemble excretory material. These deserted capsules udoubtedly correspod to Guther's " udifferetiated ova." It ow seems certai that these capsules are ot eggs, but merely modified feedig tubes, which, i additio to formig the usual arrow chael directed forwardly from the chelicerae of the larvae ito the host tissue, also grow backwardly i a much more exaggerated fashio to form a capsule, which remais ope posteriorly, ad i which the whole body of the larva is eclosed as peetratio of the ski ad egorgemet take place. third record is 5 capsules (oe cotaiig a egorged larva) foud attached to the hair aroud the aus of a marsupial mouse, techius flavipes godmai (Thomas) 7, i raiforest at 360 r, Palmersto Natioal Park, N. Q., 18 Mar. 1959, R. D. The formatio by trombiculid mites of a suckig tube or histosipho is a pheomeo kow for over 80 years. The tube is formsd by the reactio of a secretio of the mite with tissue from the host. Its purpose is to allow the mite to obtai tissue juices for igestio from levels deeper tha its mouthparts could reach. It is progressively legtheed durig the attachmet of the mite, ad its legth may evetually much exceed that of the mite itself. Whe the mite disegages, the tube remais i the host tissue. Normally oly the chelicerae of the attached larva are iserted ito the ski, but G. kallipygos is etirely itradermal. Perhaps the sclerotized caudal plate/s serve to protect the posterior aperture of the capsule. It would seem from da Foseca's illustratios (1955), that rather similar capsules are formed by the quillihabitig chigger, poloia tigipioesis Torres ad Braga. The varied positio of the capsules alog the hairs ad i the ski may be explaied as follows. I specimes totally embedded i the ski, the wall of the capsule occasioally etraps the tips of oe or more hairs about to emerge from the ski. s these hairs grow, the capsule is slowly withdraw from the ski, util it is may times its ow legth free of the ski's surface, yet still attached to the hairs. Thus a larva may be pulled out of the ski prematurely, as i Guther's material, but ormally the larva i each capsule has time to egorge fully. Whether metamorphosis takes place i the capsules or i the soil is ukow. Larval fragmets have bee see withi some capsules, but these may be the remais of larvae that have died. Nymph: Colour i life ukow; legth of idiosoma 680 ju i mouted specimes. Geital area oval, 94 ju log, with 2 pairs of geital discs; aterior discs 24 ju log, posterior discs 22 ju log. Geital plates weakly defied, with about 10 brached setae each. Ier geital setae arraged 3.3, the posterior pair, ad occasioally at least oe of the aterior pair, beig weakly barbed. al plates 69 ju log, more heavily sclerotized ateriorly, with 13 or 14 brached setae each. Gathosoma : Chelicerae of usual facies, 111 ju log; blade slightly serrate o cocave dorsal edge, 40 ju log. Hypostome blut ateriorly, with about 12 pairs of ude galeal setae to 15 ju log. Basis capituli with about 35 brached setae vetrally. Palpi with usual 5 segmets; femur with 9 brached setae; geu with 13 or 14 brached setae; tibia with sigle iteral brached seta ear isertio of tarsus, 2 ier accessory combs, a exteral ude seta ear base of claw, ad 4 exteral brached setae; tarsus with tarsala, 2 subtermialae, ad 9 brached setae. Legth of palpal claw 33 ju. Legs typical, 7segmeted. Coxae I with usual precoxal plates, each with 8 or 9 brached 7. I listig the host ames, I have foud the followig useful, ad have tried to strike a balace betwee themlaurie ad Hill, 1954; Tate, ; Troughto, 1954.

39 1960 Domrow: Gutheraa (Trombiculidae) 233 setae. Tarsus I ot elogate, 138/* log, IS ju high. Tibial Sl /JL log. Ratio LTI/HTI 1.77; ratio LTI/Ltl No preapical dorsal process o tarsus I. Scutum: rea sesilligera wider tha log, with fairly well developed posterior apodeme. Crista shorter tha area sesilligera, termiatig i usual tectum bearig sigle tectal seta. Paracristal setae subequal. Sesillae very sleder, especially distally, with barbules o basal half, which merge ito about 16 very fie ciliatios apically. Eyes abset. The stadard data are give i Table 10. Setatio: Middorsal body setae taperig ad brached, 19 ju log; termial body setae similar, but larger, 6573 ju log. Leg soleidia to 23 ju log. Table 10 Stadard data i micra of ymphal scutum of G. (G.) kallipygos (Gu.) CTL SL SB SL PSL PD TS SS SENS SB Female: Very similar to ymph, but cosistetly larger ad more hairy. Idiosomal legth uavailable because of fracture. Geital area oval, 187/* log, with 3 pairs of geital discs; aterior discs 35 ju log, middle discs 32 ju log, posterior discs 29 ju log. Ier geital setae weakly barbed, i about 15 pairs. al plates 118 ju log, with about 35 setae each. Gathosoma: Chelicerae 165 ju log; blade 57 ju log. t least 12 pairs of ude galeal setae preset. Basis capituli with about 100 setae vetrally. Palpi: Femur ad geu with about 30 brached setae each; tibia with sigle iteral brached seta ear isertio of tarsus, 3 ier accessory combs ear claw ad a fourth additioal comb set half way alog the dorsal margi, a exteral ude seta ear base of claw, ad 57 brached exteral setae; tarsus with tarsala, a ukow umber of subtermialae, ad about 14 brached setae. Legth of tibial claw 50 ju. Legs: Precoxal plates with about 20 setae each. Tarsus I more elogate tha i the ymph, 230 ju log, 113/* high. Tibia I 144 ju log. Ratio LTI/HTI 2.04; ratio LTI/Ltl No preapical dorsal process o tarsus I. Scutum as i ymph, but paracristal setae more umerous. Eyes abset. The stadard data are give i Table ll. Setatio: Middorsal body setae 39/* log; termial body setae 86/* log. Leg soleidia to 22 ju log. Table Stadard data i micra of adult scutum of G. (G.) kallipygos (Gu.) CTL SL SB SL PSL PD TS SS Ses SB Material examied: Two ymphs o separate slides, both labelled "Gutheraa bipygalis (Guth.) reared ymph #253, N. Guiea, G. M. K., 1944." Four ymphs o oe slide labelled "Gutheraa bipygalis (Guth.) ymphs ex larvae o rat, Dobadura, N. G., G. M. K., , #385." Oe with oe larva hatched from a egg it laid, labelled "Guther aa paraa Wor., adult ad larva #83, G. M. Kohls. Sy. of bipygalis." The larva has divided caudal plates, without puctae, ad with very short setae. 32. Gutheraa tidalei (Womersley) ll Trombicula tidalei Wor., 1936, Li. Soc. Lod. (Zool.), Jour. 40: 110. (Type ymph

40 234 Pacific Isects Vol. 2, o. 2 collected freelivig, Kagaroo Islad, S.., i SM). This species ad the followig are 2 of the least kow ustralia trombiculids, yet, paradoxically eough, they are the oly 2 species (apart from the geotype) which have previously bee assiged to Gutheraa (see Womersley, 1952). The available material cosists of 2 adults of trasluces ad oe ymph of tidalei. Ufortuately, they are i a coditio that precludes rehabilitatio, ad util similar material ca be reared from larvae from suitable localities, I ca do o more tha place them as species Iquiredae. Both species have the 2 characteristic clumps of galeal setae, but the precoxal area is idistict. G. tidalei is rather larger i the ymphal stage (SL 81/i, SB 76/i, SL/SB 1.07) tha the adult of trasluces (SL 74/*, SB 51^, SL/SB 1.45). The ratio SL/ SB idicates that tidalei is a Gutheraa s. s., ad trasluces a Derrickiella. Both subgeera occur as larvae i South ustralia. 33. Gutheraa trasluces (Womersley) Trombicula trasluces Wor., 1944, R. Soc. S. ust., Tras. 68: 83 (2 Q sytypes collected freelivig at Mt. rde, S.., i SM). CKNOWLEDGEMENTS I writig this paper, I have bee able to see material from the type series of every species cocered. This would ot have bee possible without the geerous cooperatio of may people, ad I am most grateful to the followigmessrs. H. M. Hale ad H. Womersley, South ustralia Museum, delaide, for Hirst's material, as well as that described by Mr. Womersley ad Dr. Heaslip; Mr. D. J. Lee, School of Public Health ad Tropical Medicie, Sydey, for slides from the Guther collectio; Mr. M. Nadchatram, Istitute for Medical Research, Kuala Lumpur, for other relevat material. Drs. J. M. Brea, Rocky Moutai Laboratory, Hamilto, ad D.. Crossley, Jr., Oak Ridge Natioal Laboratory, Oak Ridge, have critically reviewed my mauscript, while Dr. J. R. udy, Hooper Foudatio, Sa Fracisco, with whom I oce hoped to write this paper, has also bee most geerous with his time. Drs. I. M. Mackerras ad E. H. Derrick of this Istitute have also read my mauscript. To all of these I am grateful for their advice ad iterest. Lastly, I must thak Miss P. Nicholas for her careful typig, ad Mr. G. M. Smith for the photograph. REFERENCES udy, J. R Malaysia parasites IX. Notes o the taxoomy of trombiculid mites with descriptio of a ew subgeus. Ist. Med. Res., Malaya, Stud. 26: Lauretella, a ew subgeus of trombiculid mites, with otes o biology ad medical importace. Raffles Mus., Sigapore, Bull. 28: checklist of trombiculid mites of the Orietal ad ustralasia regios. Parasitology 47: udy, J. R. & R. Domrow Malaysia parasites XXIII. revisio of Orietal ad

41 1960 Domrow: Gutheraa (Trombiculidae) 235 ustralasia species of Euschogastia Ewig, icludig Walchiella Fuller, ad descriptio of a ew subgeus, Derrickiella (caria, Trombiculidae). Ist. Med. Res., Malaya, Stud. 28: udy, J. R. & M. Nadchatram, method of rearig idividual trombiculid mites i the field. Nature 174: udy, J. R. & P. H. VercammeGradjea. I press. frica Trombiculidae. I. iterim ote o the geus Trombicula Berlese ad the subgeus Micro trombicula Ewig. Natal Mus.,. Brea, J. M. & G. W. Wharto, Studies o North merica chiggers. 3. The subgeus Neotrombicula. mer. Midi. Nat. 44: Crossley, D Comparative exteral morphology ad taxoomy of ymphs of the Trombiculidae (caria). Kasas. Sci. Bull. 40: Derrick, E. H., D. J. W. Smith, H. E. Brow & M. Freema The role of the badicoot i the epidemiology of " Q " Fever: prelimiary study. Med. Jour. ust. 1: Domrow, R. 1955a. The ymph of Euschogastia perameles (Womersley, 1939) : caria, Trombiculidae. Li. Soc. N. S. W., Proc. 80: The ymph of Euschogastia smithi (Womersley, 1939) (caria, Trombiculidae). Li. Soc. N.S.W., Proc. 80: Three ew ustralia chigger ymphs (caria, Trombiculidae). Li. Soc. N. S. W., Proc. 81: Malaysia parasites XXXI. Nymphs of scoschogastia Ewig ad Lauretella udy (caria, Trombiculidae). Ist. Med. Res., Malaya, Stud. 28: a. Malaysia parasites XL. Nymphs of Leptotrombidium ad Trombiculidus (caria, Trombiculidae). Ist. Med. Res., Malaya, Stud. 29: Malaysia Parasites XLII. Oculicola, ew subgeus of scoschogastia Ewig (caria) : Trombiculidae from eyes of mammals. Ist. Med. Res., Malaya, Stud. 29: Domrow, R. & D. J. W. Smith, caria from five hudred ative mammals from Queeslad. Li. Soc. N. S. W., Proc. 80: Farrell, C. E Chiggers of the geus Euschogastia (caria: Trombiculidae) i North merica. U. S. Nat. Mus., Proc. 106: da Foseca, F carological otes XXXVII. Schoegastia (Trombewigid) bakeri, subg,, sp.., ad otes o the parasitism of ma by chigger mites from Brazil (cari, Trombiculidae). Rev. Brasil. Et. 3: 112. Fuller, H. S The mite larvae of the family Trombiculidae i the Oudemas collectio: taxoomy ad medical importace. Zool. Verh. 18: Gordo, I Importace of larval characters i classificatio. Nature 176: Guther, C. E. M The probable vector of edemic typhus i New Guiea. Med. Jour. ust. 2: a. Trombidiid larvae i New Guiea (caria: Trombidiidae). Li. Soc. N. S. W., Proc. 64: Observatios o the life history of Neoschbgastia kallipygos Guther, Li. Soc. N. S. W., Proc. 64: Further observatios o the trombidiid larvae of New Guiea (ca

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