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Full text of "Records of the South Australian Museum"

RECORDS 

OF THE 

SOUTH AUSTRALIAN MUSEUM 



VOLUME X 

The SCRUB-TYPHUS and SCRUB-ITCH MITES 

(TROMBICULIDAE, ACARINA) of the 

ASIATIC-PACIFIC REGION 

PART 1 (TEXT) 



Published by The Museum Board, and edited by the 
Museum Director 



Price: £2 2s. 



Adelaide,{March i, 1952 
PRINTED AT THE HASSELL PRESS, 104 CURRIE STREET 

Registered in Australia for transmission by post as a periodical. 



RECORDS 



OF THE 



SOUTH AUSTRALIAN MUSEUM 



VOLUME X 

The SCRUB-TYPHUS and SCRUB-ITCH MITES 

{TROMBICULIDAE, ACARINA) of the 

ASIATIC-PACIFIC REGION 

PART 1 (TEXT) 



Published by the Museum Board, and edited by the 
Museum Director 



Adelaide, 1952 
Printed at The Hassell Press, 104. Currie Street 



CONTENTS 

Page 

PREFACE 1 

INTRODUCTION 3 

CLASSIFICATION AND SYSTEMATICS - 11 

Larvae: - 19 

Trombieulinae 19 

Gahrliepiinae 278 

Adults and Nymphs : - - - - - - 312 

Trombieulinae - - - - - - 312 

Gahrliepiinae - 392 

LIST OF SPECIES 401 

REFERENCES 408 

APPENDIX 421 

INDEX - - - 429 



ACKNOWLEDGMENT 

This investigation amd publication was gener- 
ously supported (in part) by a research grant 
from the Research Gra/nts Committee, of the 
National Institutes of Health, Public Health 
Service of the United States of America, amd 
by further grants in aid from the South Aus- 
tralian Government. 



PREFACE 

The Trombiculidae, a family of Aearina (mites), are in their larval stage 
external parasites of vertebrates and more rarely of invertebrates. 

In general the species are not host specific, as many are known to occur 
on two or more kinds of animals, and secondarily also on man. In a few eases 
the same species is to be found on both vertebrate and invertebrate bosts. 

Many speeies have been known for a very long time to be a source of 
extreme annoyance to man in that they produce an ''itch/' while in some 
countries their association with a typhus-like disease in man has been suspected 
for centuries past. During the last two or three decades, and especially during 
the course of the last war in the Pacific Region, the association of these mites 
with the disease t< tsutsngamushi ,, or " river fever* ' of Japan, China, etc., and 
*' scrub typhus" in New Guinea, Malaya and Queensland, has been very inten- 
sively studied by the personnel of the Medical Army Services of Britain, 
United States of America and Australia. As a result, two speeies, Tromhicula 
{Leptoirombidium) akamushi (Brumpt) and T. (L.) dfilie?ms (Walch) have 
been definitely incriminated as vectors of the organism of this disease. 

The possibility that others of the many species as yet only responsible for 
an "itch" in man, may be potential carriers of the disease from the primary 
host, has rendered the detailed taxonomie study of these mites a matter of the 
utmost importance to those engaged in the study of Tropical Diseases. 

The writer first became interested in this particular group of Acarids in 
1940 when, because of the association of certain species wath disease in New 
Guinea and Northern Queensland, a survey of our then knowledge of the group 
in the Austro-Malayan and Oriental Regions was published jointly with W. G. 
Heaslip in the Trans, of the Roy. Soc. of South Australia, 67, (1), 1943. 

With the development of war operation in New Guinea and the Pacific 
generally, that publication became a basis for the determination of the larvae 
met with by officers of the British, American and Australian Forces, and very 
large numbers of specimens were forwarded to the writer at the South Aus- 
tralian Museum for critical determination and study. 

As was to be expected, many new species were recognized and our know- 
ledge of the taxonomy, as well as the distribution of the Trombiculidae of the 
Asiatic-Pacific Region has been very materially increased. 

Apart from my own studies of this material, a number of new speeies 
have been described by other workers engaged on ecological and medical aspects 
of the scrub typhus problem during the later period of the war. Amongst these 
must be mentioned Lt.-Col. C. B. Philip and his colleagues Major G. M. Kohls, 



2 Records of the S.A. Museum 

Major B. Traub, of the American Army, Lt 6. W. Wharton of the U.S. Navy, 
and Squadron Leader C. D. Radford, of the British South-East Asia Command. 
To aJl these colleagues I am indebted for a large amount of material, including 
paratype specimens of their species. 

Since 1947, I have also received an immense amount of interesting material, 
including many new or rare Malayan and Borneo species from Dr. J. E. Andy, 
in charge of the Brit. East Asia Scrub-typhus Commission in Kuala Lumpur. 

Further, the close association of the above-mentioned workers, most of 
whom were able to visit Adelaide at some time during the war, and whom 1 
was again able to personally meet in 1947 in America and England, has very 
greatly stimulated my own studies and been of the greatest help. To all of 
these colleagues I desire to express my greatest appreciation. 

The realization of the paucity and scatter of published knowledge of these 
mites stimulated the writer to consider the publication of a monographic re- 
vision of the species of Trombiculidae of the Asiatic-Pacific Region. In 1947 
a plan was put forward to this effect, and a generous grant was received from 
the Research Grants Division of the United States National Institute of Health, 
to enable the writer to visit institutions in the United States, Britain, and 
Europe to study further material, and as a grant in aid of the subsequent pub- 
lications of such a revision. To the Research Grants Committee and especially 
to Dr. G. B. Philip of the Rocky Mountain Laboratory, Hamilton, Montana, 
who sponsored my application, I desire to express my deep appreciation. 
Further finance towards publication was met by the Board of the South Aus* 
tralian Museum, in whose Museum Records the work is now published. For 
this and for the facilities to carry out the studies as part of my official duties, 
my sincere appreciation is also gratefully acknowledged. 

As originally planned the publication was to include also the allied family 
of "itch mites/ ' the Leeuwenhoekiidae, as well as an atlas of micro-photographs 
of the dorsal scutum of all species, of which the material available was suit- 
able. However, the recent great increase in the cost of printing and block- 
making, and the limited finance available, has regrettably caused these portions 
to be omitted for the present. The work, therefore, is restricted to the Trorabt- 
culidae only, with line figures of the species. 

In the taxonomy of these mites, the classification now adopted has been 
based on a study of the adult and nymphal forms from the whole world, and 
consequently many of the larval genera hitherto proposed have been dropped 
or reduced at the most to subgenera based on larval characters. 

In conclusion, appreciation of the help and encouragement of many 
colleagues, mentioned from time to time in the text^ is sincerely tendered. 

H. WOMERSLET?. 



ADDENDA AND CORRIGENDA 

(See also page 427.) 

A description of the circumstances of collections during World War II 
in Assam and North Burma by members of the U.S. Typhus Commission 
(Puller, Traub, Sundermeyer, etc.) is given by Mackie et al. (Trans. R, Soc. 
trop. Med. Hyg., 1946, 40, 15), while those in Manipur, South Burma, and 
also Malaya and Sarawak, are given by Audy and Harrison (Ibid., 1951, 44, 
371). 

The collection in Manipur (Imphal) area and in South Burma was made 
by the Scrub Typhus Research Laboratory, South East Asia Command, from 
March 1945 to March 1946. Squadron Leader C. D. Radford was attached to 
this unit for a period and described fifteen new species from the Imphal area 
(Proc. Zool, Soc, Lond,, 1946, 116, 247) ; he was followed in the laboratory by 
three of his pupils, G. W. Ash, W. K. Ford, M.Sc. (Liverpool Museum), and 
later T. J. Lawrence (all at the time in the Royal Army Medical Corps). 
Lawrence prepared a manuscript account, including some new names and 
temporary designations, but he was unable to follow this up after demobiliza- 
tion. A summary of his draft manuscript (without names) formed Appendix 
7 of the mimeographed War Office report (1947) referred to in the text (vide 
abstract in Trop. Dis. Bull,, 1948, 45, 62). T. J. Lawrence's species have 
been redescribed and drawn, with the retention of his names and authorship 
(viz. T. Iwnceolata, T. macacus, T. fordi, spp. n. Lawrence in Womersley 1952). 

Radford's host-data were based on provisional identifications in the field, 
since revised by Roonwal (Trans, nat. Inst. ScL India, 1949, 3, 67). His 
R. r. rvfeseens and B. r. nitidis, and the B. r. nr. brwivneusctdvs of the War 
Office report (in the present text referred to as B. r. brwntwuscidus) are all 
correctly to be identified as B. r. buUocki Roonwal 1948 (Proc, nat. Inst. Sci. 
India, 14, 386). Hydromys hwmei is a misprint for Hadrornys humei (vide 
S. lacunosa, &. lewthwaitei spp. n.) — Hydromys is an Australian genus of 
water rats. 

Collections in South Burma ascribed to Lawrenee were made by Ash and 
Andy. The following host-data of South Burma chiggers should be amended : 
N'esokia is a synonym of Bwndieota (B. bengalensis). The hosts of S> (Al) 
oAidAji sp. n. are R t rattus subsp. (not B> iwrvegicus) in Toungoo, and the 
Pagoda Rd. locality refers to Rangoon (" R.G.N. ") This is incidentally now 
known to be the commonest ehigger in squirrels in the forest in Malaya, and 



Addenda and Corrigenda 

is the "species 'S* ,T referred to by Audy and Harrison (1951). The hosts in 
Southern Burma of 8. (A.) soakafooemiensis should read Bandicota bengalevr 
sis, R. raittus nr- khyensis (not R- rattus norvegicm). The hosts of T. fordi 
sp. n. should read R. r\ bvilocki (not B. r« bruwneusculus) and the common 
Indian shrew Buncus coeruleus (misprinted Scincus). 

The following notes and amendments apply to collections from Malaya and 
Sarawak (Borneo), which were made by the Colonial Office Scrub Typhus 
Research Unit (Institute for Medical Research, Kuala Lumpur), (this and the 
Imphal Military Laboratory are variously referred to in the text) 5 References 
made by Gater, and here reproduced, to Rattus maiaisia Kloss refer in fact 
to a subspecies, the tree rat Rattus camis maiaisia. (vide T. mwnda, T. spicea, 
S> malayensi<s t etc.). The host of 8. {A.) malayensis is Rattus canus and this 
mite appears to be one of the few host-specific chiggers — the reference to its 
occurrence on R. sabanus and R. whitehead* is erroneous. The data for 
2\ muridia sp. n. should be corrected to read: specimen No. 6,451 on R. 7,246 
R. mulleri, 25 July, 1949. Identifications of certain bats have recently been 
confirmed : R.10,299 (host of T. leveri sp. n. in Malaya) is Emballonum mwvti- 
e#la; R.11,018 (host of T. Jtfvrrisoni sp. n.) is Hipposideros sp.; R.10,566 (host 
of 8. lipoxena sp. n.) is also Hipposideros sp. (the number 13,944 refers to the 
first chigger specimen-number). 

Most collections in Malaya have come from areas within 15 miles of the 
capital, Kuala Lumpur, in Selangor State, The island Jarak and the twin 
islands Seletar (bat-cave) and Angsa are in the Malacca Straits, respectively 
at 3° 59' N., 100° G' E„ and 3 B 12' N., 101° 13' E. 

Collections made by Major S- Lai Kalra, I.M.S./I.A.M.C, in charge of the 
Field Typhus Research Detachment (formerly with the Imphal team), have 
been in the Kumaon Hills (circa 79° 30' N-, 79° 30' E.) and Kashmir (Simla 
HUls, eto., 33° 30' N.— 34° 30' N. t 72° 40' E.— 75° 30' E.). 

Page 2, in line 21 for "G. B. Philip" read "C. B. Philip". 

Page 3, in first line of paragraph 2 of Introduction for ** (146) " read " (142a) '\ 
and in Bibliography on page 417, after "142" insert "142a Sambon h. W. 
1923. The parasitic acarians of animals and the part they play in the 
causation of the eruptive fevers and other diseases of man. Preliminary 
consideration based upon an ecological study of typhus fever". — Ann. 
Trop. Med. and Parasite!., 22, 460. 

Page 6, in line 3 for "BSte range' ■ read "Bete rouge". 

Page 9, in line 16 for "221" read "222", 

Page 33, in line 7 transpose. "ne" in IcueJcenschrijveri to "en". 



Addenda and Corrigenda 

Page 46, in bottom line before Trombiculindus insert "(", and after "1951" 

insert U Y\ 
Page 55, in line 3 for "T. H. Lawrence" read "T. J. Lawrence". 
Page 81, in line 2 for "1945" read "1944", and in line 3 for " (Zephronia sp.) 

read " (Sphaeropaeus gldbus-magicus Jeekel 1951) ". 
Page 109, in the description of Trombicula parmifera sp. n. no reference is 

made to the peculiar structure of the seta on coxae III ; this is short and 

strongly swollen in the basal half and fine distally. 
Page 111, in line 19 for "C. Lundblad" read "0. Lundblad". 
Page 129, line 25 and elsewhere for "Kukit Lagond" read "Bukit Lagong". 
Page 139, following Standard Data for Trombicula southcotti sp. n. insert 

1 * Log. and Host. The type and two paratypes from the leg of a bandicoot, 

at Birdum, N.T., Aust., 2 Apr., 1942, (R.V.S.)." 
Page 149, in line 19 for "scabram" read "$cabrum'\ 
Page 188, in line 18 for "Wandecla" read "Wondecla" 

Page 205, the host of S. labuanensis sp. n. should read as "a rat" (not mar- 
supial rat — these are not found in Borneo). 
Page 281, delete "With only 1 humeral seta on each side" in line 16, and "6a" 

in line 17, and insert a fresh line "6a. With only 1 humeral seta on each 

side" between lines 17 and 18. 
Page 303, in line 16 for "bifurcate" read "trifurcate". 
Page 307, and elsewhere, for "longipilis" read "longipiH". 
Page 336, delete third line from bottom. 
Page 355, in line 8 for "Oniscomorpha sp." read "Sphaeropaeus gldbus-magicus 

Jeekel 1951". 
Page 379, re S. (A.) indica in line 12 for "reddish in life" read "whitish in 

life". 
Page 389, for "fig. Eu I" read "fig. E-I". 



THE SCRUB-TYPHUS AND SCRUB-ITCH MITES 

(TROMBICULIDAE, ACARINA) OF THE 

ASIATIC-PACIFIC REGION 

PART I (TEXT) 



ByH. Womersley, A.L.S. (Hon. causa), F.R.E.S., Entomologist, 

South Australian Museum 



Summary 

Although the modern taxonomic descriptions of these mites dates only from 1905, 
when Oudemans described two species, Thrombidium wiclimanni and Th. 
vandersandei from New Guinea - both well known causes of "itch" to travellers in 
that country - their possible association with disease appears to have been suspected 
from much earlier times. 

The classification of this family of mites, which is very extensively based on the 
larval stages, is an extremely difficult problem in taxonomy and, owing to the very 
many larval genera which have been proposed by recent workers, as well as to our 
very great lack of knowledge of the adult stages of most genera, is still in somewhat 
of a chaotic state. 

The family Trombiculidae was erected in 1944 by Ewing to include the subfamilies 
Hemitrombiculinae Ewing 1944, and Trombiculinae Ewing 1929 (External Parasites 
p. 22) for all the then known larval Trombidiidae known to parasitise vertebrates. The 
other larval Trombidiidae, although little known in that stage, and as far as then 
known parasitic on invertebrates (as in the allied family Erythraeidae) were left in the 
Trombidiidae in its restricted sense. The Trombiculidae s.l., however, are now known 
not to be entirely confined to vertebrate hosts, and several species of this family, 
sensu stricta, and of the Leeuwenhoekiidae occur in numbers on various species of 
arthropods. 

The subfamily Leeuwenhoekiinae was erected in 1944 by the present writer for the 
genus Leeuwenhoekia Ouds. s.l. on the presence in the larvae of a pair of true 
spiracles, one on each side between the gnathosoma and the first coxae, and leading 
into the tracheae ramifying through the body. 



The SCRUB-TYPHUS and SCRUB-ITCH MITES 
(TR0MBICUL1DAE, ACARINA) of the 
ASIATIC-PACIFIC REGION 

By H. WOMERSLEY, A.L.S. (Hon. causa), f.r.e.s., Entomologist, 
South Australian Museum. 



INTRODUCTION 

Although the modern taxonomic descriptions of these mites dates only from 
1905, when Ondemans described two species, Tkrombidium wichmanm and Th. 
vandersandei from New Guinea — both well known causes of "itch" to travellers 
in that country — their possible association with disease appears to have been 
suspected from much earlier times. 

Prom the writings of Dr. Louis Sambon (146) it seems that the earliest 
known recognition of the sha^shih (sand mite) as the cause of typhus-like infec- 
tion in Southern China was the account given in a Chinese work of the 16th 
Century, the Pen Te'ao Kang Mu by Li Shih-chen. In this account the mite 
was said to be red in colour, not larger than an ant, and to kill man by burrowing 
under the skin. It occurred on land and water. The fever caused is now 
regarded as clinically the same as the "river-fever" or "tsutsugamushi" of 
Japan. 

The earliest published account of the disease itself and its association with 
a mite was given in a letter from Dr. T. A. Palm (130) to the Rev. J. Lowe and 
published in the Edinburgh Medical Journal in 1873. The disease was called 
"Shima-mushi" or "island insect" disease. In 1879 Baelz (16) in a supplemen- 
tary note to an earlier article by himself and Hawakami (17) describing Japan- 
ese "flood fever" recognized it as "Shima-mushi". 

The earliest description of the disease in Japan (from Niigata Prefecture) 
was by Hakuyu Hashimoto in 1810, who called it "tsutsuga". According to 
Hatori (71) 1919-20, the identity of the disease mite of Japan with the "sand 
mite" of South China was, however, recognized a century earlier by Genkei 
Ohtomo, and his son, physicians of the Akito district of Japan. 

In 1899 Tanaka (155) considered tsutsugamushi to be almost certainly due 
to a minute mite, locally called "kedani" or "hairy mite", which was subse- 
quently named Trombidium akamushi by Brumpt in 1910. It occurred plenti- 



4 Records of the S.A. Museum 

fully on voles but would also attack man and other animals. In this paper 
Tanaka also indicated that a second species occurred which he called " Yasodani" 
or "vole mite". 

In 1906 Miyajima reared the nymphs of the M kedaiu" and some ten years 
later (113) in association with Okumura he was able to rear the adults. From 
1916 onwards the study of the Japanese Trombiculid mites was intensively taken 
up by other workers, notably Nagayo and his colleagues, Kawamura and Yaina- 
guchi, Okumura, etc. 

Thus in 1916 (120) Nagayo et al. described the nymphs and adults of the 
tsutsngamusbi mite, and in general established its life-cycle. Later in 1917 
(121) they further extended their studies and as the nymphal and adult stages 
diffei*ed from those of all other Trombiculid mites known to them, they suggested 
the generic name of Leptotronibidium for akamushi Brumpt 1910. Shortly 
afterwards the work of these students was duplicated by Kawamura (94), while 
Miyajima and Okumura (115) in a comparison of these mites from Japan, Korea 
and Formosa accepted Tanaka 's view as to the occurrence of two kinds. 

Prior to 1916, however, Oudemans had in 1912 (126) published his mono- 
graph "Die bis jetjst bekannten Larven von Thrombidiidae und Erythraeidae" 
in the Zool. Jahrb., Suppl. 14, in which he described, and mostly figured all the 
then known species of Trombidiid larvae. Of the Japanese species he had very 
little to say beyond a brief reference to Tanaka 1889, Baelz (16), and to Ashburn 
and Craig's account (11) of the disease in the Philippines. The mite itself wa« 
referred to as "Gen. ? sp. V. He suggested that these larvae, like the other 
described species which attack man, probably belonged to the genus u Micro- 
thrombidium" Haller 1882. 

In 1915 (77) Stanley Hirst redescribed and refigured the Japanese species 
and, following Oudemans, called it Microtrombiddum akamushi (Brumpt 1910). 

In 1919 Nagayo et ah, added a third species from Japan, and apparently 
realising that the adult of akamushi was very closely related to the genus IVaw- 
oicttla Berl. 1904 (type T. minor BerL 1904 from Java), they dropped their 
name Leptotrombidium in favour of Trombicula, In that paper they applied 
the name of T. akamushi (Brumpt) to Tanaka 's "kedani, T. pallida to Tanaka 's 
"yasodani", and T. palpalis to the new species. 

In 1921 Nagayo et al. (123) described two more species from Japan, 
T. intermedia and T. scivtellcms. Of the above five species they were able to cor- 
relate adults and larvae of akamushi > pallida and palpalis, both by rearing larvae 
through to the adult and by obtaining larvae from captured adult females; 
intermedia was only got to the pupal stage (nympbophane) between larva and 
nymph, and smdellaris only as far as the nymph. 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 5 

In 1930 (158) Tanaka described another Japanese species under tilH name of 
T. autumnclis japonica, relating it to the European autuvvnalis (Shaw 1790). 
lie also reared it through to the nyiaph which he described and of which he gave 
a photograph in which the eyes are seen to be placed in front of and away f om 
the sensillary area of the crista, a featnre which places the species in the genus 
Tragardhuto Berl. 1912, an# distinguishes it from autumnalis Shaw. 

In January 1950 (133) Philip and Fuller pointed out that a second species, 
closely allied had been figured by some Japanese workers (Okumura 1918; 
Kawamura 1926) which had been recognized as distinct by some, and confused by 
others, but which had remained unnamed. They dedicated the species to Tamiya 
as Trombimda tamiyai. 

More recently, Sasa 1950, (148, 149, 150) in Tokyo Ishi Shimshi, Pts, 10, 
11, and 12, described three more species from Japan, under the names of 
T. nagayoi, T. mitamtifai and T. burnsL The first two species in having a pen- 
tagonal scutum and long nude setae on the tarsi and the metatarsi of leg III 
should probably be placed in Tragardkvla. The first is closely related to, but 
appears to be distinct from, japmiica (Tanaka), while the second is almost iden- 
tical with tamiyai Philip and Kohls, only differing in the number of dorsal setae 
and the Standard Data. The third species is near to T> (£.) pallida and T* (L.) 
intermedia. 

Thus with a new species of Gahrliepia described in the present paper, eleven 
species of Trombiculidae are now known to occur in Japan. 

In Formosa, although the presence of a typhus-like disease was known as 
early as 1908 it was not until 1919 (71) that Hatori related it to the tsutsuga- 
mushi and considered T. akamushi (Brumpt) to be the vector. In that paper he 
also described three other Formosan species, T. pseudodkamusM (non Tanaka), 
T. conn, from crows, and T. gallinarum from fowls. The first of these he consi- 
dered to be the larva of an adult found in a field at Karenko, which after Miya- 
jima he regarded as the same as mediocris Berlesc 1912 from Java. In 1921 
Kawamura confirmed and extended Hatori 's work and gave further details of the 
species including the nymph of gallinarwm. According to present day nomencla- 
ture, psf.'udoa'ka?nushi Hatori (non Tanaka) is a synonym of T. wichrn/t/nm Ouds. 
1905, and gaUinarum is a species of Neoscliongastia, 

In other parts of the world these mites have long been known as causing an 
annoying '*itek" to humans. The earliest record is that of Neotrombicula, batatas 
(L. ) from Surinam. Linnaeus quotes this species as having been found by 
Itolander, and in 1905 Oudemans (126) quotes Muller 1775 as stating that, 
according to Bolander, Acancs batatas crawls on the legs of man and causes an 
itch f The earliest date given by Sig. Thor and Willmann (1947) is that or Her- 



6 Records of the S.A. Museum 

lean 1718 in Beschr. Volk. Plant. Zuriname, p. 179, where it is mentioned under 
the vernacular "Patattes luis". In other places Martinique, etc., it was known 
a*"B§te-range'\ 

In North America the common itch mite or "dagger" was first described by 
Riley in 1873 as Leptus irritans. As this name, however, had been previously- 
used for any species the synonymy became confused, until 1941-2 (44) when 
Ewing showed that it should be known as TronibictUa alfreddugem Ouds. 1910. 
Oudeman's proposal of rileyi nom. nov for the N, American species is considered 
by Ewing to be unnecessary owing to the existence of several synonyms, and he 
suggests that if alfreddugesn is invalid, then his name winabarinus 1921 based 
on the adult would be applicable. In the adult section of the present paper, how- 
ever, it is considered that the adults of alfreddugem and dnndbarinus cannot be 
separated, and that Oudenian'g name alfreddugem should be used. 

In Europe, the earliest scientifically recognizable species is Neotrombicvl<f> 
mtumnalis (Shaw 1790). This, the "Harvest Mite" of Britain, is known from 
moat European countries where it goes under various local names. It becomes 
especially numerous at harvest time when it causes considerable annoyance to 
workers in the fields. 

In New Guinea and Malaya "itch mites' • were a source of much discomfort 
to many of the earlier travellers in those parts, and Oudemans 1905 gives a very 
full account of the 'many records left by such explorers. 

The earliest of these is that of J. H. de Boudijek Bastiaanse in his "Voyages 
faits dans les Moloques a la Nouvelle Guinee et a Celebes ", Paris 1845, where he 
says that ever and anon on returning from trips ashore or to the forests he was 
plagued by an itch. 

Amongst other records of similar experiences quoted by Oudemans are those 
of C. B. H. von Rosenberg in "Reistoehten naar de Geetvinkbaai op Nieuw 
Guinea in de Jaren 1869-1870". s'Gravenbage 1872 ; L. M. D'Albertis in "New 
Guinea: What I did and what I saw", London 1880; Alfred Russell Wallace in 
"The Malay Archipelago : The land of the Orang-TJtan and the bird of Paradise' f 
1869 ; and of the two Sarasin's in "Zeitechrift der Gesellschaft fur Erdkunde za 
Berlin", Bd. 29, 1894. 

In the descriptive part of his paper Oudemans figures and describes in detail 
the two species Trombicula wichnumni, and Bchongastia vandersamdei. 

In New Guinea, while a form of endemic typhus was known from the Man- 
dated Territory, no cases had been reported from either Papua or Dutch New 
Guinea until the second World War. Dr. Carl Gunther (Med. J. Aust. 1940) 
first showed that there was sufficient evidence to indicate that the probable vector 
in New Guinea was a species of mite, Trombicida minor (non Bertese) (== hirsti 



Womersley— Asiatic-Pacific Scrub Typhus Mites 7 

Sambon 1927). Further work on this mite and on the Troinbiculidae of New 
Guinea in general by Gunther laid the foundation of our pesent knowledge of 
the mites of this island. He described a number of new species and published 
much on their ecological relationship. 

In Sumatra a kedani (tsutsugniushi)-like fever was reported by Dowden 
in 1915, and in 1921 E. W. Walch commenced an investigation of the disease at 
Deli, paying particular attention to the mites which could be associated witJi it. 
His results were published in several papers in 1923, 1924 and 1927, in which, 
besides describing a number of new species of Trombieulidae, he showed that 
Trombicufa, deUensis Walch was a probable vector in the transmission of the 
infection from rats to man. Further he was able to rear the larvae of tin* and 
other species to the nymphs, which he fully described and figured. Earlier, 
however (1915) Schuffner had already recorded an undetermined species of mite 
as attacking man. 

The first association of mites with typhus-like infections in Malaya is that 
of Fletcher, Lesslar and Lewthwaite in 1928. They recorded an outbreak of the 
disease in 1926, on an oil palm estate near Kuala Lumpur, when one case was 
attributed to an "insect bite n . In their 1928 paper they listed four species, 
namely 2\ pseiidoakamushi Walch («. T, MrsU Sambon), T> teMensw Walch, 
(«= Schongastia (Ascoschongastia) hidica Hirst), and T, (= Schongastia) 
schuffneri Walch, all of which had been described from Sumatra by Walch. 

Following on this paper, B. A. Gater 1941-2 made a very extensive survey 
of: the Trombieulidae in Kuala Lumpur and the surrounding oil and rubber 
estates, but confined it to those species found parasitic on rats. He listed three 
species T. akamushi, T. deUensis and T. hirsti Sambon as found on labourers 
and, from another district, T. aeiuscutellaris Walch. He followed Oudeman's 
classification, and in all recorded eight species of Trombicula (three new), two 
of Schongastia (one new), five of Neoschongastm (= AscoscMnga&tia) (f out- 
new), six of QahrNepia (including Gateria, all new), and three new species of 
Walchia (**= Gahrliepi<a) . 

In India the mites associated with typhus-like infections were first studied 
by Mehta (103) in 1937, although several odd species had been described earlier 
by Oudemaus 1914 (127) and by Hirst in 1915 (78). He recorded both T. deli- 
ensis and T. acuseutelhiris as attacking both rats and man, and regarded deUensis 
as the vector. 

In Australia the association of mites with the coastal and other tsutsuga- 
fuushi-like fevers was suggested by Cilento 1923 (31), Hirst 1929 (81) and 
Matthew 1938, while the studies of Heaslip 1938-9 (74, 75) indicated that Trom- 
liiada deUensis was most probably the chief vector in Northern Queensland. 



8 Records of the S.A. Museum 

The first species known from this country was T. hirst i Sambon 1927 (146) 
from Queensland. Later in 1929, Hirst recorded this species as occurring in 
South Australia, but later Womersley 1939 (191) showed that the South Austra^ 
lian species was not co-specific with Sambon 's species, but was a new species, 
T. samboni Hirst also described in 1929 T, no-vae-hoUandiac y Schongastia antu 
podimum, 8. coorongeme, and S. dasycerci, all from S. Australia. In 1934, 
Womersley (188) added 8. westraUe?ise from "Western Australia and 8. petrogale 
from the Musgrave Ranges, S, Australia. 

In 1936, (189) an adult Trombieulid, T. (Ghmtherana) Hndafai was des- 
cribed by Womersley from Kangaroo Is., S. Australia, and a larval species 
T. ma-cropus from the Northern Territory. 

In 1939, Womersley (191) reviewed and keyed the Australian and New 
Guinea species, and in addition described the T. samb&ni from S. Australia, 
recorded Guntheria kallipygos Qunther 1939 (= Ountherana bipy galas Gunther 
1939), and described Neoschongastia (== Ascoschdngastia trichosuri) N. (=4.) 
perameles, ffi (—4,) queenskmdica, N* (A.) derrick^ N. (A.) smithi from 
Qtieensland, He also erected the genus Paraschongastia (= Neoschonga&tia 
Ewing) for four of Gunther \s New Guinea species. 

In 1942, (192), an adult, Trombicida (I'fXtiro'inbimla) elegans Worn, was 
described from Lush Is., South Australia. 

In 1943 (195) a comprehensive survey of the Trombieulid^ (including 
Leeutvenhoekia) of the Austro-Malayan and Oriental Regions was made, jointly 
with W. G. Heaslip. In this work a total of 91 species (excl. Leewwenlwekia) 
with four varieties were listed, described and figured. 

Trombicula translucent n. sp. (adult.), T. sciTMoides, T. obscum (=<Uy/- 
mu^hi), T. kohlsij T. sareina, Selwngmtia pusilla (= psendo-schuffneri Walch.) 
Neoschdngdstia {AscosclibngJ) mccullachd and Guntherana parana (== bipygedh) 
n. sp. were described from Australia or New Guinea in 1944 by Womersley 
(193). 

In 1947, Womersley and Kohls (196) added Eutrombicvla gynvnodaotyhi 
n. sp., Novotrombicula owiensis n. g, et. sp. from Dutch N. Guinea, Schongastvl 
philipi n. sp., Oenoschonga&tia cana n. g. et. sp., Ascoschongastia uromys n. sp. 
and A&coschong, echymipera n. sp. from New Guinea, 

Prom Burma and Manipur, India, a number of species have been described 
during the war by Radford, 1946 (136, 137) and also by Traub and his colleagues 
and by Philip and Fuller (133). 

Owing to the many successful rearings of larval species to nymphs during 
the past decade, especially by officers of the British Scrub Typhus Research Unit 
in Malaya, it is now possible to <iarify the generic classification of these mites, 



Womersley— Asiatic-Pacific Scrub Typhus Mites 9 

and to base it on the adult or nymphal stages. With this aim in view, the adults 
and/or nymphs known from the whole world have been considered. Such a 
revision shows that many of the larval genera which have been proposed cannot 
be separated in the nymphal or adult stages, should not be used except perhaps 
for convenience, as subgenera based on larval features only. It would appear 
that in this family of mites, we have a very striking example of extreme "larval 
incongruence". 

As an adjunct to the differentiation of larval species considerable use has 
been made of the Standard Data (linear micrometric measurements) of the 
dorsal scutum, as first applied by Womersley and Heaslip 1943 (195). Where 
sufficient specimens of a species, or a population of a species, were available, the 
Standard Data have been treated statistically, and a theoretical range of the 
variation to be expected for each measurement, as expressed by M dt 3o> deter- 
mined. In the condensed data given under each species in the keys, this value 
is used. 

In the present study 221 larval species and 53 adult or nymphal forms are 
listed, of the latter of which 46 have been correlated with the larvae. From the 
rest of the world 26 species are dealt with, of which only in 10 have the adults 
and/or nymphs been correlated with the larvae. Seventy-seven new species are 
described, mainly from the larvae. 

That this list, however, is nowhere near the complete fauna of the area is 
obvious, and from the rapid increase in our knowledge of these mites during the 
past 10 years, the mite fauna, when more inaccessible parts are explored, will 
undoubtedly be much increased, possibly to double the present number of species. 
It is hoped, however, that the present study will serve as a basis for further and 
even more intensive work on these mites. 



CLASSIFICATION AND SYSTEMATICA, 

The classification of this family of mites, which is very extensively based 
on the larval stages, is an extremely difficult problem in taxonomy and, owing 
to the very many larval genera which have been proposed by recent workers, as 
well as to our very great lack of knowledge of the adult stages of most genera, 
is still in somewhat of a chaotic state. 

The family Trombiculidae was erected in 1944 by Bwing to include the 
subfamilies Hemitrombiculinae Ewing 1944, and Trombiculinae Ewing 1929 
(External Parasites, p. 22) for all the then known larval Trombidiidae known 
to parasitise vertebrates. The other larval Trombidiidae, although little known 
in that stage, and as far as then known parasitic on invertebrates (as in tie 
allied family Erythraeidae) were left in the Trombidiidae in its restricted 
sense. The Trombiculidae s,l,, however, are now known not to be entirely 
eonfined to vertebrate hosts, and several species of this family, sensu striata, 
and of the Leeuwenhoekiidae occur in numbers on various species of arthro- 
pods. 

The subfamily Leeuwenhoekiinae was erected in 1944 by the present writer 
for the genus Leewwenhoekia Ouds. s.l. on the presence in the larvae of a pair 
of true spiracles, one on each side between the gnathosoma and the first coxae, 
and leading into the tracheae ramifying through the body. 

Later Womersley (1945, Tr. Roy. Soc. S. Aust., 69, (1), 96), raised the 
Leeuwenhoekiinae to family rank, when it was shown that the adults of 
Lteuwenhoekia {Acomataearm) differed markedly in shape and details of the 
epistome from those of the Trombiculidae. Ewing 1946 (J. Parasitology, 32, 
(5), 435) apparently failing to appreciate the fundamental nature of the pre- 
sence of true stigmata and tracheae in the larvae, and the differences in the 
adults of the Leeuwenhoekiidae, did not accept this family separation, but 
divided the Trombiculidae into the four subfamilies; Trombiculinae, Leeuwen- 
hoekiinae, Hemitrombiculinae and Walchiinae. 

In 1947 Wharton (J. Parasitology, 33, (4), 380) gave a detailed diagnosis 
of the Trombiculidae, based on larval characters, after Ewing. In this paper 
he removed, quite correctly, the aberrant Hemitrombiculinae, retained the 
Leeuwenhoekiinae, and added another subfamily, the Apoloniinae, to include the 
genera Apolonuv Torres and Braga 1939 and Womersia Wharton 1947, both of 
which are closely related to Leeuwenhoelria and its allies, in possessing true 
stigmata and tracheae. 



12 Records of the S.A. Museum 

In his key to the subfamilies, Wharton stressed the segmentation of the 
legs as the primary character, the Leeuwenhoekimae having all three pairs of 
legs 6-segmented, ie. the femur is undivided; the Trombiculinae and Apolonii- 
nae having the femur of all legs subdivided into basi- and telofemur, and all legs 
therefore 7-segmented ; and the Walchiinae with only the first pair of legs 7-seg- 
mented. 

Apart from the fact that the division of the femur into basi- and telo- 
femur, is not a true articulation, the character as a subfamily one is not import- 
ant, and if strictly applied, as has been done recently by E. P. Lawrence (Ann. 
Natal Mns., XI, (3), 1949) leads to some strange relationships. Lawrence's 
new genus Sauracarella, is, in my view, because of the distinct anterior projec- 
tion of the dorsal scutum and the paired AM scutal setae, better placed in the 
Leeuwenhoekimae, despite the clavate or globose sensillae, and the 7-segmented 
legs. As has been shown elsewhere, the anterior scutal process and the paired 
AM setae are correlated with characters of the epistome in the adult. The 
occurrence of clavate sensillae in the Leeuwenhoekiidae, can just as well be 
expected to occur in this family also, as it does in the Trombiculidae. The 
absence of stigmata and tracheae is possibly also open to question. This 
feature, with the stigmata so little chitinized, is so very difficult to see at 
times, even in freshly mounted specimens, that one must be very careful before 
asserting their absence. 

Other species, difficult to assign on the leg segmentation, are Schongastia 
(Ascoschongastia) oudenumsi Walch, 8. (A.) heaslipi Worn, and Heasp. and 
#. (A.) traubi sp. n. In these species, contrary to the rest of the Trombicu- 
linae, the first pair of legs only are 7-segmented, the others 6-segmented. They 
would, therefore, have to be placed in the Walchiinae (Gahrliepiinae), a group 
from which they completely differ in the scutum and scutal setae. 

The genus Heaslipia Ewing 1944 (— Tromhiculoides Womersley and Heas- 
lip 1943) is erroneously referred by R. P. Lawrence 1949, to the Leeuwenhoe- 
kimae, on the supposed 6-segmented legs. Unfortunately, however, the sub- 
division of the femora is not shown in the original drawing from which Law- 
rence drew his conclusion- I have re-examined the specimens in the South 
Australian Museum, and the legs can now be definitely stated to be all 7-seg- 
niented. 

There has been a tendency to regard trivial characters as of generic import- 
ance, and some of these have been discussed by Lawrence (1949). 

The first that need be mentioned is the subdivision of the palpal claw. 
Ewing has used the bi- or trifurcation of this as generic. As will be seen from 



WOMERSLEY— ASIATIC-PaCIFIC SCRUB TYPHUS MlTES 13 

the following descriptions and keys, this feature is not even subgeneric, and 
both bi- and trifurcate claws occur in different species of the same genus. 

Ewing also, has raised genera on the number of dorsal setae, particularly 
in separating the genus Acariscus from Eutronibioula. This has been corrected 
by Michener and others. Similarly, the dentition of the chelicerae, cannot be 
used as has been done by Lawrence 1949 y as more than subgeneric, and even 
then must be used with considerable discretion and a certain amount of latitude. 

However, until more is known of the adults, some larval genera must still 
be diagnosed on characters, which may ultimately be shown to be of no more 
than specific value. 

The LeeuwenhoeMidae and the Trombiculidae can be separated on the fol- 
lowing larval and adult or nymphal characters • 

1. Larvae with a dorsal scutum with generally 2 AM (rarely 1 or none) and 
an anterior median scutal process (« sometimes absent), generally with 
tracheae and true stigmata (? sometimes absent). 

Adults and nymphs cordate, widest across propodosoma, without medial 
constriction. Crista anteriorly expanded into a rounded or pointed arrow- 
head like nasus furnished with two ciliated setae. Eyes present or absent; 
if present, then 2 -f 2, situated in front of and away from sensillary area 
of crista. Family LEEUWEfraoiSKnDAE Womersley 1945. 

2. Larvae with only 1 AM seta, or none, on scutum and no antero-median 
scutal process. Stigmata and tracheae absent (? Tarely present). 
Adults and nymphs figure-eight shaped, with a distinct medial constriction, 
and propodosoma not wider than hysterosoma. Crista anteriorly ending 
in a hyaline dentate epistome furnished with a single ciliated seta. Eyes 
1 + 1 ,or absent ; when present either closely adjacent to sensillary area, 
or away from and in front of this area. 

Family Trombiculidae Ewing 1944 (restricted Womersley 1945). 



Family TROMBICULIDAE Ewing 1944. 

Proc. Biol. Soc. Washington 1944, 57, 101-104; J. Parasitology, 1946, 32, (5), 
435-440. 

A fresh diagnosis of this family as defined above, based on both larvae and 
adults or nymphs is as follows : 

Larvae.. Text fig. 1, A-D. Minute, oval to round, six-legged acarids, fur- 
nished with only a single antero^dorsal scutum. The scutum of varied shape, 
furnished with a pair of filamentous, or globose, or elavate, or lanceolate sen- 
sillae, and normally 5 setae (1 antero-median, 2 anterolateral and 2 postero- 



14 



Records of the S.A. Museum 



lateral) (in the Gahrliepiinae the antero-median seta is absent) ; frequently in 
the Gahrliepiinae (rarely in the Trombiculinae) the scutum is prolonged pos- 
teriorly to include some of the setae from the dorsum; or there may be addi- 
tional setae between the antero- and postero-lateral setae. Eyes usually 2, some- 
times only 1 or none on each side, more or less close to the lateral scutal margins. 
Chelicerae two-segmented, basal segment stout, apically a chitinized curved 
blade frequently serrated or with strong teeth on the inner (dorsal) edge. Palpi 
5-segmented, basal segments each bearing a single, always branched, seta and 
fused in the median line, and with a median laminate projection extending be- 




MJU m 



♦total Mb* 



Fig. 1, Details of Larval Characters: A. right ventral and left doraal halves showing 
setal arrangement; B. left palp in ventral view; C. mandible and galeal seta; D. dorsal 
scutum showing method of obtaining Standard Data, 



Womersley— Asiatic- Pacific Scrub Typhus Mites 15 

yond basal segment* of chelieerae, and with & pair of galeae which curve dorsally 
over the chelicerae each witb a nude or branched seta ; the palpal femur is stout, 
outwardly curved or Homewhat angular, and carries dorsally a single nude or 
branched or ciliated seta ; the palpal genu carries a single dorsal similar seta j the 
palpal tibia is always furnished with 3 setae, one dorsal, one lateral and one 
ventral, which may be nude or branched; the tibia terminates in a strong claw 
which may be simple or variously furcate,- the tarsus opposes the tibial claw 
and usually bears 6 to 8 ciliated setae, a subapical nude seta and a sub-basal 
striated rod-like aensilla. No true stigmata and tracheae are present (except in 
one species of Neoschongastia) . Between -coxae I and II is always present a 
so-called pseudostigmata or "urstigma." Behind the dorsal scutum the dorsal 
setae are arranged in more or less well defined curved transverse rows; the 
first row consists of 1 or more humeral setae on each side placed laterad or 
postero-laterad of the scutum. Generally the legs are 7-seginented due to a 
secondary division of the femur into basi. and telofemur; occasionally the front 
legs only have a subdivided femur, the second and third legs being 6-aeg- 
mented. The tarsi ends in a. pair of curved claws, with a median longer and 
more slender claw-like empodium. In one unique genus the empodium is 
caruncle-like or pad like, as in the Paraaitidae. Each coxa normally carries a 
single ciliated seta but frequently some coxae may be multisetose. Tarsi I and 
II have a dorsal sensory rod, and tarsi HI and/or metatarsi III may be fur- 
nished with 1 or more long nude outstanding setae.* On the venter between 
coxae I and between coxae III is a pair of ciliated setae, and "behind coxae III 
the setae are arranged in more or l*ss transverse rows. The anal opening is 
placed subposteriorly on the venter. 

In 1943 Womersley and Heaslip devised a system of linear measurements 
of the dorsal scutum, and scutal setae, as an adjunct to the normal morpholo- 
gical characters used in the specific identification of Trombiculid larvae. In a 
more general way, however, somewhat similar scutal measurements had been 



♦ In 1951, Wharton et al (J. Parasitol, 37, (1), 13-31) published an excellent paper on 
the terminology and classification of the Trombiculid Mi tea stressing the need for a note 
exact terminology of the morphological characters of both larvae and adults. While from the 
viewpoint of comparative morphology their terminology has much to be said for it, from the 
aspect ot the practical taxonomist, many features especially the various small nude sensory 
setae of the legs, are of little importance and do not appear to offer any practical help in 
the identification and separation of species. The use of the term "spur" for a short Btriatcd 
blunt seta found on the tarsi of legs I and II is morphologically incorrect, In no sense is it 
a spur and the usual description of it as a "sensory rod" ia more descriptive. For the 
long nude setae found on the femora, tibia and tarsus of the third pair of leg*, the? use the 
tonne mastifemoraloe, masritibiaJae and mastitaraalae. ' ' Taxonomically these setae are 
important, but the above terminology has not been adopted in the present work, as beine 
somewhat unnecessary. ^ 



16 Records of the S.A. Museum 

used earlier by Methlagl 1927. "Ueber di Trombidiose in den Oesterreichischen 
Alpenlandern", Denksch. Akad. Wissen. Wien., 101, 213-250. Methlagl used 
the ratio of breadth to length of scutum and particularly the ratio of breadth 
of scutum to the depth between a line at the maximum width and the extreme 
posterior angle, which he called the "Convexity Factor/' 

The system of Womersiey and Heaslip used a number of scutal measure- 
ments, termed the "Standard Data" as follows: 

AW ? width between the centres of the anterolateral scutal setae. 

PW. width between the centres of the postero-lateral scutal setae. 

SB. distance between the centres of the sensillae bases. 

ASB. distance from a line joining seusillae bases to extreme point of 

anterior margin, 
PSB. distance from a line joining sensillae bases to extreme point of 

posterior margin. 
SD. Sum of ASB and PSB, i.e. scutal depth. 
A-P. distance between antero-lateral and postero-lateral setae in a direct 

and more or less oblique line, not vertical. 
AM. length of antero-median seta. 
AL. length of antero-lateral setae. 
PL. length of postero-lateral setae. 
Sens. Length of sensillae. 

The method of making the measurements is clearly shown in text fig. 1, D. 

The use of the Standard Data in addition to morphological features should 
be part of the routine description of larval species of Trombiculid mites, for, 
in the case of closely similar speeies it may indicate significant separation on 
one or more of the Standard Data criteria. Wherever possible, the Standard 
Data should be based on a number of specimens from the same population, and 
thus an estimate of the extreme range of variation in a species can be obtained. 
In some populations, otherwise morphologically the same but from different loca- 
lities, significant differences in Standard Data values would indicate geographi- 
cal races, differing only in the overall dimensions of the scutum, or differences 
in length of scutal setae, SB, etc. 

With these ideas in mind, where sufficient numbers are available, the 
Standard Data for each species dealt with in the present paper, have been 
treated and expressed statistically. In the keys to species, however, the Stan- 
dard Data are simplified, and the value for each character is expressed as 
"Mean zt 3 X Standard Deviation,' ' i.e. the theoretical range of variation that 
might be expected in any one population. 

In 1944 (Trans. Roy. Soc. S. Aust., 68, (1), 82-112) and later papers. 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



17 



Womersley has treated and quoted the Standard Data statistically; and further, 
where different populations of certain species, e.g. T. (L.) deliensis, etc. 
appeared to occur, given graphs which showed clearly how the populations or 
species might be separated. This method is also used in the present studies, 
where material is sufficient and suitable. 




Pig. 2. Details of Adult Characters: A, Trombicula samboni. Worn., entire doraal view 
B, diagrammatic view without legs, showing pre-coxal plates of a Leptotrombidium; C, side 
view of a Trombunda; D palp; E. crista, showing eyes, and epistnme of a TrombicvU: F. 

SSS £ C ttt 6 * m <E ft*?* "W?* 1 plates of a G^towa; H, tarsus and mota- 
tarsus of leg III ot a Trombuwla, showing method of measuring: I, tarsi of lee of a 
Cranrliepia showing process. e 



38 Records of the S.A. Museum 

In addition, another graphic method of separating closely allied species, 
eg. 8. (A.) indwa Hirst and S. (A.) soekaboemiensis Takekawa is indicated 
in Graph II showing the Regression Lines of AW.PW, for populations of both 
species. The triangles and circles indicate the frequency of specimens with 
AW.PW, at the points therein. The elipses are drawn at the range/<r using 
Tippet's Table (Biometrika, 17, 386, 1925), while the dotted diagonal are the 

axes of AW and of PW. 

Adults {and Nymphs). Text fig. 2, A-L Larger, eight-legged animals of 
pale to red colour and furnished with a thick pilosity of setae. Form rather 
sqaat, with the body divided medially by a constriction giving a characteristic 
figure of 8 shape. The cephalothorax is rather narrower than the anterior 
part of the body and triangular in shape; it carries a median longitudinal rod- 
like crista which is expanded posteriorly or subposteriorly into a wide sensillary 
area carrying a pair of filamentous or narrowly davate sensillae. Anteriorly the 
crista ends in a hyaline rounded or transverse, dentate epistome carrying a 
single ciliated seta. The eyes when present, are 1 on each side, sessile, and 
either closely adjacent to the sensillary area, or in front of and away from it. 
The ehelicerae are two segmented, the apical being a strongly dutinized curved 
blade with finely serrate inner (dorsal) margin. The palpi are Segmented, 
with the basal segments fused medially; the femur is the longest and stoutest; 
the. tibia ends in a strong simple claw at the base of which are 3 to 4 strong 
spines (2 in nymphs), and there is another, somewhat more tapering about the 
middle of the outer surface; sometimes the palpal tibia has a pair of pectines 
or combs; the palpal tarsus is conical, opposed to the tibial claw, and apically 
bears 5 or 6 short nude setae. The respiratory system is by means of tracheae 
opening at the base of the mandibles. 

Ventrally the coxae are arranged in two groups, widely separated; an 

anterior group composed of coxae I and II and a posterior group of coxae III 

aud IV. Between the coxae of the anterior group is a generally more or less 

pentagonal sternum formed by the complete fusion of the precoxae of coxae I; 

in some cases the precoxae of coxae I are clearly defined and do not quite 

conjoin to form a sternum, in others they are only partially fused to give a 

longitudinally divided sternum; no precoxae occur on legs II, III or IV. The 

legs are 7-segmented with I generally longer and stronger than the others; the 

tarsi end in a pair of curved claws, those of leg I being much shorter thau 

the rest. The legs and palpi are thickly clothed with setae usually more taper- 

ing than the body setae. The genital opening lies just behind or between coxae 

IV and in both sexes has 3 pairs (2 in nymphs) of genital discs; in the male 

often the inner genital lips are flanked by one or more pairs of specialized 

f titillating) setae. The anus is posterior of the genital opening. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 19 

LARVAE. 

Because of their importance as vectors of scrub-typhus and tsutsugamushi 
fever, our knowledge of the larval species of the Trombiculidae during 
recent years has increased tremendously. On the other hand, relatively little 
is known of the respective adults and only in a small proportion of species have 
the two stages been correlated. Consequently it is still advisable, as will be 
done in this work, to deal separately with the known larvae and adults (or 
nymphs). 

Key to the Larval Subfamilies and Genera 
of the TROMBIOULIDAE. 

1. With AM scutal seta present Femora of all legs mostly clearly 
divided into basi- and telof einur . , . . , , . . - . 2 

Subfara. Trombiculinae Ewing 1946. 

Without an AM seutai seta. Femora of leg I subdivided, of II and 
III not so, Sensillae clavate. (Tarsi I of adults and nymphs with a 
subapical dorsal process. No sternum or precoxal plates. Byes 
absent)* . . . . . . . . . . . . . . 23 

Subfam. Gahrliepiinae noni. nov. for Walchiinae Ewing 1946. 

Gen. Galirliepia Ouds, 1912. 
including 

Walchia. Ewing 1931 
Schongastiella Hirst 1915 
Gateria Ewing 1938 

2. Sensillae filamentous, more or less with distinct ciliations . , * . 3 
Sensillae globose, clavate or lanceolate, nude or setulose . . . . 13 

3. Dorsal scutum pentagonal. Palpal claw trifurcate or bifurcate. 
Tarsi and/or metatarsi of leg III frequently with 1 or more long, 
nude, outstanding setae. (Adults with 1 + 1 eyes, placed in front of 
sensillary area and away from crista). 

Gen. Tragarrdhula Berl. 1912 

= Blankaartia Berl. 1912 (nan Ouds. 1911). 
— Pentagonella Sig Thor 1936 
= Megatrombicula Micheiier 1946 

Dorsal scutum not clearly pentagonal . . . . . . . . . . 4 

4. Mandibles and palpi short and stout, apparently modified for grasp- 
ing hair. On bats. Gen . Myatrombicu-Ut Womersle^ and Heaslip 1943. 

Mandibles and palpi not so formed . . . . 5 

5. With more than the 5 normal setae, in addition to the sensillae, on 

the scutum. ' Scutum not rectangular, usually longer than wide ► . 12 
With only the 5 normal scutal setae (1 AM. 2 AL and 2 PL) in addi- 
tion to sensillae (PL may occasionally be off the scutum). Scutum 
usually more or less rectangular, wider than long . . . , . . 6 



20 Records of the S.A. Museum 

6. Palpal claw simple, Chelicerae unarmed except for the apical tricus- 
pid cap. Gen. Crotiscus Ewing 1944. 

Palpal claw furcate . - . . & . . 7 

7. Chelicerae with a row of 3 large dorsal (inner) teeth. Scutum ves- 
tigial, AM seta spiniform. Sensillae short and simple with bases 
close together and slightly anterior of line of PL. Palpal claw trifur- 
cate. Gen. Endotrombicula Ewing 1931. 

Chelicerae without teeth, only with an apical tricuspid cap . . . . 8 

8. Palpal claws tri- or quadrifurcate. Scutum small with PL setae well 
off the scutum. All palpal setae branched. Galeal setae nude. 

Gen. Tecomatlam> Hoffmann 1947. 
= Sauri-scus Lawrence 1949. 
= Trisetica Traub and Evans, 1950. 

Palpal claws bi- or trifurcate. Scutum not as above 9 

Gen. TrombiciUa Berl. 1905. 

9. AL or AM and AL scutal setae short, stout and peg-like. Palpal 
claw bi- or trifurcate. Subgen. Fonsecia Radford 1946. 

The above scutal setae normal . n 10 

10. Palpal claws bifurcate. Galeal setae nude or branched. Tarsi III 
usually with 1 long nude seta. (Adults with eyes 1 + 1, closely adja- 
cent to sensillary area). 

Subgen. Trombicnla Berl. 1905, s. str. 
'mm Eutrombicula Ewing 1938 
mm Acariscus Ewing 1943. 

Palpal claws trifurcate. (Adults without eyes) . . . . >'-« H 

11. Setae on palpal femur and genu nude ; on tibia dorsal seta branched, 
lateral and ventral nude (except ventral in palpalis, etc.). No long 
nude seta on tarsi or metatarsi of leg IIL Galeal setae branched 
(nude in myzmtha). (Adults with preeoxae on legs I, forming a 
longitudinally divided sternum). 

Subgen. Leptotrombidium Nagayo et al. r 1916. 

Without the above combination of characters. Usually without any 
long nude seta on tarsi or metatarsi III. Galeal setae nude or 
branched. Subgen. Neotrombicula Hirst 1925. 

12. Dorsal scutum produced posteriorly, tongue-like so as to take in some 
of the median setae of some of the dorsal rows. Palpal claw bifur- 
cate. Chelicerae without a prominent dorsal (inner) tooth. 

Gen. N&votrombicula Worn, and Kohls 1947. 

Dorsal scutum not produced posteriorly, but with posterior and lat- 
eral margins forming a continuous even curve ; with 2 extra setae on 
each side between AL and PL. Palpal claw trifurcate. Chelicerae 
with a large prominent dorsal (inner) tooth. 

Gen. Heaslipia Ewing 1944. 

for Tr&mbicuMdrs Worn, and Heasp. 1943 preoc. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 21 

13. Empodium expanded into a pad-like pulvillus. Sensillae globose. 

Gen. Mackiena Traub and Evans 1950. 
Empodium normal, claw-like and slender . . 14 

14. All coxae multisetose. Scutum roughly triangular with apex anterior. 
Sensillae clavate and setulose. Palpal claw trifurcate. 

Gen. Doloisia Ouds. 1910. 

Coxae I and II always unisetose, III sometimes multisetose. wScutum 
trapezoidal, pentagonal or hexagonal . . . . . . . . . . 15 

15. Empodium distally thickened, lancet or blade-like. Eyes absent. 
Scutum roughly pentagonal with posterior angle. All coxae unisetose. 
Sensillae globose. Gen. Reidlinia Ouds. 1944. 
Empodium not as above 16 

16. Chelicerae short and stout, curved and stumpy with apex divided into 
two terminal teeth and two smaller subapical teeth. Palpal claw short 
and stout, trifurcate. Sensillae globose. 

Gen. Oenoschongasti<a Worn, and Heasp. 1945. 
Chelicerae not so as above, more elongate and slender . . . . 17 

17. Palpal claw simple. Chelicerae unusually long and straight and 
slender, finely serrate dorsally. Sensillae capitate. 

Gen, Radfordiana nov. 
Palpal claw furcate . . . . . , . . * . . . , . 18 

18. Body constricted medially. Dorsally with a rounded caudal plate, 
generally longitudinally divided and bearing 3 pairs of fine setae. 
Palpal claw bifurcate. Sensillae globose, apparently nude. (Adults 
with sensillary area wider than crista is long. Precoxal plates of leg I 
distinct and not conjoined to form a sternum. Eyes absent). 

Gen. Quntheraw Worn, and Heasp. 1943. 

Body not constricted medially. No caudal plate . . . . . . 19 

19. Cuticle of dorsum overlapping posterior half of scutum with semi- 
circular striations thereon. Sensillae globose or clavate, their bases 
inserted in a transverse wall or crest. Coxae III sometimes multi- 
setose, (Adults with sensillary area kidney-shaped, with striations. 
Precoxal plates absent. Sensillae lightly clavate. or whip-like. Eyes 
absent.) Gen. Neoschditgastia E wing 1929 

= Parmclwngastia Worn. 1939. 
Cuticle of dorsum not so overlapping scutum, or if so with only trans- 
verse striations. Sensillae globose or clavate, their bases not in a 

crest. Chelicerae simple or with teeth . . . . 20 

Gen. Sclumgastia Ouds. 1910 

20. Chelicerae non-serrate, with only the apical tricuspid cap . . . ► 21 
Chelicerae with tooth-like serrations . . 22 

21. Palpal claw with 5-7 apical prongs. Sensillae clavate or capitate. 

Subgen. Euschongastui Ewing 1938. 

Palpal claw bi- or trifurcate. (Adults or nymphs with triangular 
sensillary area. Eyes and precoxal plates absent, or completely fused 
to form a sternum.) Subgen. AsicoscAongasiia Ewing 1945. 



22 Records of the S.A. Museum 

22. Chelicerae finely serrated for almost the whole length on dorsal mar- 
gin. Tarsi I with one short sensory rod. Palpal claw bi- or tri- 
furcate, (Adults and nymphs as in Ascoschongastia) , 

Subgen. Schtin-gastia Ouds. 1910 s. str. 
Chelicerae with only two, more prominent, subapical dorsal teeth. 
Tarsi I with 4 sensory rods. Palpal claw trifurcate. 

Subgen. Phrynacarus Lawrence 1949. 

23. Dorsal scutum pentagonal, not produced posteriorly to include some 
of the dorsal setae. 

Subgen. GakrUepia Ouds. 1912. 
~ Walchia Ewing 1931. 

Dorsal scutum produced posteriorly, tongue-like and generally taking 

in some of the dorsal setae. . . . . . ► . . . . . . 24 

24. With only 2 or no additional setae on scutum. 

Subgen. Schongastiella Hirst 1915. 

With more than 2 additional setae on scutum 25 

25. All additional setae on scutum marginal. 

Subgen. Gahrliepia Ouds. 1912. 

Some of the additional scutal setae not marginal. 

Subgen. Oateria Ewing 1938. 

Genus TRAGABDHUIiA Berlesa 1912. 
Tragardhula Berl. 1912, Redia, 8, (1), 4. 

Blamkaartia Berl. 1912, Redia, 8, (1), 96 (nan. Oudemans 1911. Ent. Ber., 3, 
(57), 123). 

Pentagmella S, Thor. 1936. Zool. Anz., 114, 30. 

Megiitrombictda Michener 1946. Ann. Entom. Soc. Amer., 39, 432. 

Diagnosis as defined in the key to subfamilies and genera of Trombiculidae. 

Key to the Larvae from the Asiatic-Pa ci&c Region. 

1. Tarsi III and/or metatarsi with some long nude, outstanding setae . . 2 
No such setae on tarsi or metatarsi III . . . . . . . . . . 5 

2. Only 1 long nude outstanding seta on tarsi III, none on meta- 
tarsi III -. .. .. ,.'.. .. . . .. .. 3 

Three long nude setae on tarsi III, one such on metatarsi III . « 4 

3. PW only a little longer than AW. Dorsal and lateral setae of palpal 
tibia nude. Galeal setae nude- Dorsal setae 26, arranged 
2.6.6.6.4.2, to 72fi long. 

AW 77-4=b7-8, PW 80-l±7-4, SB 30--8zt3-6, ASB 30-0±3-7, 
PSB 43-3+4-3, SD 72-3+6-1, A-P 25-9±3-5, AM 52-7±6-l r 
AL 40-25+4-7, PL 77-1+8*0, Sens. 84-0. 

Trag, acuscutellaris (Walehl922). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 23 

PW much longer than AW. Setae on palpal femur and genu 
branched; on tibia dorsal with 1 short branch; lateral nude, ventral 
branched. Galeal setae nude. Dorsal setae 34, arranged 4.6.6.6.6.4.2, 
i.e. 2 humeral setae on each side. 

AW 83-8±:10-3, PW 100-5+22-3, SB 33-0+5-7, ASB 35-5+ 
3*7, PSB 31-0+4-6, SD 66-5+7-7, A-P 32-5+3-7, AM 62-0,+ 
7-3, AL 5S-8+10-2, PL 73-2+6-4, Sens. 88*0. 

Trag. japonica (Tanaka 1930) . 

4. All setae on palpal tibia branched. Galeal setae nude. Dorsal setae 
42, arranged 2.8.8.10.8.6. 

AW 64-5+5-9, PW 82-2+8-2, SB 27-25+4*4, ASB 28-25 + 
4-4. PSB 32-6±5-0 f SD 60-9+7-3. A-P 21-55+4-2, AM 44-45 
±6-5, AL 48*0+7-4, PL 62-45+7-9, Sens. 76-0+15-0. 

Trag. tamiyan (Philip and Fuller, 1950). 

Only ventral seta of palpal tibia branched. Galeal setae nude. Dor- 
sal setae 42, arranged 2.8.8.8(10) J0(8).6.4. 

AW 60-2, PW 81-2. SB 26-6, ASB 30*8, PSB 30-8, SD 61-6, 

A-P 25-2, AM 42-0. AL 36-4, PL 46-2, Sens. 80-0. 

Trag. fvjigmo (Philip and Fuller 1950) . 

5. Femora of all legs distinctly undivided, i.e. all legs 6-segmented. 
Galeal setae stout and thickly, shortly, ciliated or pectinate. Setae on 
palpal femur and genu branched; on tibia, ventral branched, dorsal 
and lateral nude. Palpal claw bifurcate. Dorsal setae 30 in number, 
arranged 2.8.6.6.6.2, to 30^ long. 

AW 36-4, PW 53-2, SB 12-6, ASB 19-6, PSB 14-0, SD 33-6, 
A-P 16-8, AM 14-0, AL 25-2, PL 30-2, Sens. 42-0. 

Trag. geckobia sp. n. 

All legs distinctly 7-segmented, i.e. femora divided into basi- and telo- 
femur. Galeal setae nude . . 6 

6. Anterior scutal margin convex, with AM' placed well in front of line 
of AL. Lateral margins of posterior angle of scutum concave. All 
setae on palpal femur, genu and tibia branched. Dorsal setae 24, 
arranged 2.6.6.4.4.2. Palpal claw trifurcate. 

AW 51-6+618, PW 70-2+8-85, SB 20-8+3-09, ASB 25-3 + 
2-58, PSB 28-5+2-97, SD 53-5+4-32, A-P 22-3+3-78, AM 
34-6+4-29, AL 29-6+2-40, PL 35-0+3-51, Sens. 55-6+6-45. 

Trag. pentagons sp. n. 

Anterior scutal margin straight, with AM in line with AL. Lateral 
margins of posterior angle of scutum convex. Seta on palpal femur 
branched; on genu nude; on tibia, dorsal and lateral nude, ventral 
branched. Dorsal setae 24, arranged 2.8.6.6.4.2, to 30/* long. Palpal 
claw bifurcate. 

AW 45-75+4-50, PW 60-0, SB 16-5+5-2, ASB 18-0, PSB 23-25 
±4-50, SD 41-25+4-50, A-P 19-5+5-2, AM 24-0, AL 25-5 
+ 5-2, PL 24-0, Sens. 47-0+5-2. 

Trag. gj/mnodactylki (Worn, and Kohls 1947). 



24 Records of the S.A, Museum 

Tragardhitla actjscutellaris (Walch 1922). 

Irombictda acuscutellaris Walch 1922 (1923). Kitasato Archives Exper. 
Med., 5, (3), 78; Tromb. a.> Gater 1932, Parasitol., 24, 143-174; Pmta- 
gonella <*., Sig Thor 1936, Zool. Anz., 114, 30; Tromb. (Pentag.) a., Wom- 
ersley and Heaslip 1943, Tr. Roy. Soe, & Anst., 67, (1), 78; Tronib. a,, 
Metha 1937, J. Med. Res., 25, (2), 253-365; Tromb. a., Williams 1944, 
Amer. J. Trop. Med., 24, (6), 255; Tromb. a., Philip, Woodward and 
Sullivan 1946, Amer. J. Trop. Med., 26, (2) \ Tromb, a. y Radford 1946, 
Parasit, 37, (1-2) ; Pentagonella a., Sig Thor and Willmann 1947, Das 
Tierreich, Lfg., 71b., 293 and 296; Tromb. a., Jayewickrerne 1947, Nature, 
160, 578. 

Plate 1, fig. A-IX 

This speeies is widely distributed in the Asiatic-Pacific Region. Origin- 
ally described by Walch from Sumatra it has since been reported from the 
Federated Malay States (Gater), India (Mehta), Maldive Islands (Radford), 
Philippines (Philip) and has also been taken at Labuan, Borneo (R. K. Mc- 
Culloeh) ; in all cases the hosts were species of fiattus. 

The nymph was reared from larvae by Radford and briefly described. 
Jayewickrerne in Ceylon has reared them to the adult from larvae through 3 
or 4 generations, and study of that material, published in Tr. Roy. Soc. S. Anst., 
1948, 72, (1), 83-90, by the present author has shown that it is a species of 
Tmgardhula, of which the type is Trag. mlotica Trag. (adult). 

The specific characters of the larvae are given in the following revised 
description and in the key to species. In having only one long, nude, seta on 
tarsi III it is closely related to japonica (Tanaka), but differs in having 
1 humeral seta on each side and in the galeal seta being nude. 

Re-description of Larvae, Colour in life red. Shape oval. Length 
(unfed) 260/i (excluding gnathosoma), width 195/x. Dorsal scutum (fig. A) 
pentagonal, with PW only very slightly greater than AW and angle of con- 
vexity of posterior margin, i.e. PW/PSB ca. 2-0. Sensillae long and filament- 
ous with filiations on distal two-thirds. Normal scutal setae ciliated and 
tapering. Eyes 2 4-2, on ocular shields and posterior eyes the smaller. 
Palpi (fig. B) stout; femur and genu with a branched or long ciliated seta; 
trifureate. Chelicerae (fig. C) with the usual apical tricuspid cap, and on 
tibia with the dorsal and lateral setae nude, the ventral branched \ palpal claw 

Of the above Pacific-Asiatic species placed under Tragardh\da 9 only acuscutellaris (Walch) 
and japonica (Tanaka) are definitely known to truly belong there.'on the basis of the adults or 
nympha. The others are provisionally placed in the genua on the pentagonal scutum in the 
larvae. Farther Tearing studies may prove or disprove this generic placing. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 25 

the inner edge a subapical forwardly directed tooth. Galeal setae nude. 
Dorsal setae 26 in number, arranged 2.6.6.6.4.2, to 70/t long. Ventral setae 
(excluding the coxae) 24, arranged 2.2.6.4.4.4.2, to 50/a long. Legs long and 
slender, all 7-segmented ; I 350/i long (including coxae), II 392/4, III 435/1,; 
tarsi I and II with the usual dorsal sensory rod, III with one long, fine, nude, 
sub-basal seta to 80/*, long* 

The Standard Data derived from 17 specimens bred from eggs laid by a 
wild adult from Ceylon, 17th Dec. 1947 (S. H. Jayewickreme) and 1 specimen 
each from Federated Malay States and from Batavia reported upon in 1943 
(Womersley and Heaslip) and one from R. r. 7iorvegicus from the Maldive 
Islands, 30 Dec. 1944 (CD. Eadford) are as follows: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 

Bange 


Coeff . of 
Variation 


AW 


77 -4+0-58 


2-61±0-41 


69-6-85.2 


70*0-S4-0 


3-3 


PW 


80-l±0-55 


2-47rh0-39 


72-7-S7-5 


75-6-86-8 


3.1 


SB 


30-8±0-27 


l-20±0-19 


27-3-34.4 


28-0-33-6 


3*9 


ASB 


30-0±0-28 


l-24±0-19 


26- 3-33- 7 


28-0-30-8 


4-1 


P8B 


43-3±0-32 


l-43:±0-22 


39*0-47-6 


42-0-44-8 


3-3 


8D 


72»3db0-46 


2-04±0-32 


66-2-78-4 


70-0-75-6 


2-8 


A-P 


25-9±0-26 


1-16±0-18 


22-4-29-4 


25-2-28-0 


4-5 


AM 


52-7±0«49 


2-04±0-35 


46.6-58-8 


50-4-56-0 


4-0 


AL 


40-25±0-35 


l-56±0-25 


35.5^44.9 


39-2-43-4 


3.8 


PL 

Sens. 


77-l±0<61 
84-0. No varii 


2-68±0-43 
ttion recorded. 


69-1-85*1 


75-6-85*1 


3-5 



Tragardhula japonica (Tanaka 1930). 

Leptus (mtumnalis japonica Tanaka 1916, Igakai Zasshi (in jap.), 30, (22); 
1918, Ikai Jiko (in jap.), No. 1,228. 

Trombicukb atdumnaMs japonica, (Tanaka et al. 1930), Zentbt. Bakt., Abt. 1, 
Orig., 116, 361. 

Pentagonella japonica, Sig Thor 1936, Zool. Anz., 114, 30. 

Trombicula (Pentagondla) japonica^ Womersley and Heaslip 1943, Tr. Roy. 
Soc. S. Aust., 67, (1). 

Trombicula autumnalis japonica, Sig Thor and Willniann 1947. Das Tierreich, 
Lfg., 71b. 

Trombicula japomca, Philip and Fuller, 1950, ParasitoL, 40. 

Plate 1, fig. E-H. 

There are two distinct species of "akidani" or u harvest" mites figured 
by various Japanese workers, in one of which the tarsi of leg III have only a 



26 Records of the S.A. Museum 

single long, nude, outstanding seta; the other has 3 such setae on tarsi III 
and also 1 on metatarsi III. 

From Tanaka's 1930 paper, the first of these would appear to be the one 
to which he attached the name autumnatis japanica. The second form does 
not appear to have been given a specific name by Japanese workers but rather 
to have been confused with Tanaka's species. Its distinctiveness, however, 
has now been recognized by C. B. Philip and H. S. Fuller, who have described 
it as a new species tamvyai. 

That the species is now to be placed in the genus Tragardhula is shown by 
Tanaka's reference (1930) to the position of the eyes in the nymph, which he 
was able to rear, as being away from and in advance of the sensillary area, as 
in the type of Tragardhula, nUotwa BerL {nan. Ouds.), 

The position of the eyes is also clearly shown on Tanaka's figure 37 on 
Plate IV. His referring of the Japanese form to a subspecies of the European 
autumnaMs is thus shown to be untenable, for mitumnalis belongs to 
Neotrpmbicula, a subgenus of Trambicula Berl., in which eyes are entirely 
wanting. The species is closely related to acuscutellcvris but differs as siven 
in the key to species. Specimens agreeing with Tanaka's description (1930) 
and figures have been collected by Lt.-Col, C. B. Philip in both Yamagata and 
Niigata Prefectures in 1945 from Microtus montehelli. It is not yet known to 
occur outside of Japan. 

Re-description of Larvae. Shape oval. Length (engorged) to 560**, width 
to 390/*. Dorsal scutum pentagonal, with PW much longer than AW. and 
angle of convexity of posterior margin, i.e. PW/PSB — 3-24. Sensillae long 
and with about 3 cilia on mid-third, only slightly posterior of line of PL. 
Other scutal setae long, ciliated and tapering. Eyes 2 + 2, on ocular shields, 
posterior eyes the smaller. Palpi stout, (fig. F), femur and genu with 
branched setae; tibia with ventral branched, dorsal with 1 short branch, lateral 
nude. Palpal claw trifurcate, Chelicerae with only the small apical tricuspid 
cap. Galea! setae nude- Dorsal setae long, ciliated and tapering, 34 in 
number, arranged 4*6.6,6.6.4.2, to 72/w, long, i.e. there are 2 humeral setae on 
each side. Ventral setae similar but shorter, approx. 44 in number (exchiding 
those on coxae). Legs all 7-segmented, I 325/x long, II 285^, III 350^ (includ- 
ing coxae) ; tarsi I and II with usual dorsal sensory rod-, III with 1 long 
nude outstanding seta to 50/i long. All coxae unisetose. 

The Standard Data derived from 6 specimens from Microtus montehelli 
collected by Lt.-Col. C. B. Philip, from Yachi, Yamagata Prefecture, Japan, 
28th Oct. 1945 are as follows ! 





WOMERSLEY— 


-Asiatic-Pacific Scrub Typhus Mites 


2 






Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


83-8+1-40 


3*43+0.99 


73-5-94-1 


78-0-88-0 


4-1 


PW 


100 -5+3-04 


7-45+2*15 


78-2-122-8 


94-0-115-0 


7-4 


SB 


33-0+0-78 * 


l-90±0-55 


27-3-38-7 


30-0-36-0 


5-7 


ASB 


35-5±0-50 


1-22+0 '35 


31'8-39-2 


33-0-36-0 


3-4 


P8B 


31-0+0-63 


1-55+0-45 


26-35-35-65 


30-0-33-0 


5-0 


8D 


66-5+0-92 


2-56+0-65 


59-7-73-3 


63-0-69-0 


34 


A-P 


32-5+0-50 


1-22+0-35 


28-8-36-2 


30-O-33-0 


3-8 


AM 


62-0+1-00 


2-45+0-71 


54 -7-69 -3 


60-0-66-O 


3-9 


AL 


58-8+1-53 


3-42+1-08 


48-6-69-0 


54-0-63-0 


58 


PL 


73-2+0-87 


2-14+0-62 


66-8-79-6 


70-0-75-0 


29 


Sena. 


88 '0. Only one 


determination. 









Tragardhula tamiyai (Philip and Fuller 1950.) 
Trombicula tamiyai Philip and Fuller, 1950, Parasitol., Vol. 40. 

Plate 1, fig, I-M. 

Description of Larvae. Shape oval. Length 325/*. (unfed), width 200/a. 
Scutum pentagonal with the posterior angle slightly rounded laterally with 
angle of convexity of posterior margin « 2 • 52. Sensillae bases posterior of 
line of PL ; PL setae the longest ; sensillae ciliated on middle two-fifths ; anterior 
margin lightly sinuous between AL and with AM posterior of line of AL. 
Eyes 2 + 2, on ocular shields, about equal in size. Chelicerae with only the 
usual tricuspid cap. Galea! setae with 2 or 3 branches. Palpi stout, claw 
trifurcate; all setae on palpal femur, genu and tibia branched. Dorsal setae ca. 
42, arranged 2.8.8.10.6. Ventrally with 1 seta on each coxa, a pair between 
«oxae I and between coxae III and thereafter ca. 42. Legs: I 260/*, II 230/t, 
III witb 3 long, nude, outstanding setae; metatarsi III with one such seta. 

The Standard Data derived from 31 specimens from the original material 
collected by Nagayo, Tamiya and others in Yamagata, Japan, are as follows : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 

Range 


Coeff. of 
Variation 


AW 


64-5+0-35 


1-96+0-25 


58-6-70-4 


60-0-69-0 


3-0 


PW 
SB 


82-2+0-50 
27-25+0-27 


2-73+0-35 
1-48+0-19 


74-0-90-4 
22-8-31-7 


75-0-88-0 
24-0-31-7 


3-3 
5-3 


ASB 


28-25+0-37 


1-48+0-19 


23-8-32-7 


27.0-30-0 


5-2 


PSB 
SD 


32-6+0-30 
60-9±0-44 


1-66+0-21 
2-43±0-31 


27-6-37-6 
53-G-68-2 


30-0-36-0 
57-0-66-0 


5-1 
4-0 


A-P 


21-55+0-25 


1-39+0-18 


17-4-25-7 


20-0-25-0 


6-5 


AM 
AL 


44-45+0-46 
48-0+0-46 


2-15+0.32 
2-46+0-33 


38-0-50-9 
40-6-55-4 


42-0-48-0 
42-0-51-0 


4-8 
5-1 


PI/ 


62*45+0-50 


2-65+0-35 


54-5-70-4 


58.0-68-0 


4-2 


Sens. 


76-0+1-77 


5-01+1- 25 


61-0-91-0 


70-0-85-0 


6-6 



28 Records of the S.A. Museum 

Remarks. This species appears to have been confused by many Japanese 
authors with the preceding species., japomca Tanaka. It is, however, as recog- 
nized by Philip and Fuller, very different in the setae of the palpal tibia, the 
dimensions of the scutum, the number of dorsal setae and in the number of 
long nude setae on leg IIL 

TRAGABDBuiiA fujigmo (Philip and Fuller 1950.) 
Trombicula fujigmo Philip and Fuller, 1950, Parasitol., 1950, 

Trombicula n. sp- "D." Lawrence in MS. 1947. In Audy 1947. "Scrub 
Typhus Investigations in S.B. Asia, Pt. III. Appendix 7." AMD, 7. 
War Office, London. 

Plate 2, fig. A-C. 

Description of Larvae, Shape oval. Length (unfed) 260/*. width 170/4. 
Dorsal scutum pentagonal, but the posterior margin rather rounded, angle of 
convexity of posterior margin — 2-64; anterior margin sinuous with AM placed 
behind line of AL ; sensillae about in line with PL, filamentous and ciliated on 
distal two-thirds. Eyes 2 + 2, close to scutum, on ocular shields, posterior 
slightly the smaller. Chelicerae with only the apical tricuspid cap, Galeal 
setae nude. Palpi stout, with all setae on femur, genu and tibia branched, 
except dorsal and lateral on tibia. Dorsal setae 42, arranged 2.8.8.8(10).10(8).6, 
to 45/* long. Ventrally with 1 seta on each coxa, a pair between coxae I 
and between coxae III, and thereafter 6.6.6.4.4. Legs; I 260/* long, II 220p, 
III 260//.; tarsi I and II with the usual dorsal rod, III with 3 long, nude, out- 
standing setae on tarsi and 1 on metatarsi. 

The Standard Data for 2 paratypes are; AW 60*2, PW 81-2, SB 26-6, 
ASB 30 8, PSB 30-8, SD 61*6 A-P 25-2, AM 42-0, AL 36-4, PL 46-2, Sens. 
80-0. 

Remarks. The above description is from two paratype specimens from 
Croeidura vorax from Myitkyina, Burma, April 14, 1945, Its relationship to 
other species is given in the key. 

The single specimen, which Lawrence intended to describe, was from 
Ratios r. hrtcnneusctilus Hodgson from Kanglatongbi, nr, Imphal, Assam., 
Dec. 6th, 1945. This specimen was referred to by Philip and Fuller 1950 as 
being near to their species fujigmo from Burma, differing only in the some- 
what higher Standard Data and the slightly greater number of dorsal setae, 
47 as against 42. 

Through the kindness of Dr. J. R. Audy I have been able to study this 
specimen and would regard it as only a slight variant of typical fujigmo. With 



Womersley— Asiatic -Pacific Scrub Typhus Mites 29 

the discovery of more specimens it might be shown to be a local population. 
The Standard Data are: AW 75-6, PW 86-8, SB 30-8, ASB 30-8, PSB 30-8, 
SD 61-6, A-P 28-0, AM 56-0, AL 47-6, PL 70-0, Sens. — . 

Tragardhula geckobia sp. n. 
Plate 2, fig. D-E, 

Description of Larvae. Shape oval. Colour in life unknown. Length 
(unfed) 230/i, width 170/*. Dorsal scutum small, pentagonal, punctate, with 
PW much longer than AW, and angle of convexity of posterior margin, i.e. 
PW/PSB— * 3*8. Sensillae long, filamentous, ciliated on distal half and about 
in line with PL. AM scutal seta short and thin and shortly ciliated, others 
thicker and longer and more strongly ciliated. Eyes 2 + 2, large, on ocular 
shields, posterior the smaller. Chelicerae with only the apical tricuspid cap. 
Palpi stout; setae on femur and genu branched; on tibia, dorsal and lateral 
nude, ventral branched. Palpal claw bifurcate. Galeal setae stout and 
strongly, thickly ciliated or pectinate. Dorsal setae tapering and ciliated, 30 
in number, arranged 2.8.6.6.6.2, to 30/x long. Ventral setae, excluding coxal, 
2.2.8.6.4.2,2, somewhat similar but shorter than dorsal setae. Legs, all 6- 
segmented, femur undivided; I 200/* long, II 162/*, III 195/i; tarsi I and II 
with usual rod-like seta dorsally, III without any long nude seta. 

The Standard Data for the type specimen are: AW 36-4, PW 53-2, 
SB 12-6, ASB 19-6, PSB 14 0, SD 33-6, A-P 16*8, AM 14-0, AL 25-2, PL 
30-0, Sens. 42-0. 

Loc. and Host. Described from the type and 3 paratypes from a gecko, 
collected by Dr. R. V. Southcott, 60 miles south of Darwin, N.T,, Aust, 9th 
May, 1943. 

Bemarhs. As the adult is unknown, this species is placed in Tragwrdkvla 
on the bases of the pentagonal scutum. Its relation to other species is given in 
the key. 

Tragardhula pentagona sp. n. 

Plate 2, fig. F-I. 

Description of Larvae. Shape (unfed) an elongate oval. Length (unfed) 
225/*, width 150/t. Dorsal scutum pentagonal, with AM distinctly anterior of 
line of AL and the anterior margin strongly convex; surface sparsely punctate; 
with angle of convexity of posterior margin, i.e. PW/PSB = 2-44; sensillae 
placed only slightly behind line of PL, filamentous, with minute spicules on 
basal half, longer ciliations distally. Eyes 2 + 2, adjacent to posterolateral 



30 Records of the S.A, Museum 

corners of scutum, posterior eyes the smaller, Chelicerae 24/a long, with only 
the email apical tricuspid cap. Galeal setae long and nude. Palpi stout 5 
setae on palpal femur and genu long and curved with long branches; tibia with 
all 3 setae finely and long branched; palpal claw trifurcate; tarsus with sub- 
basal sensory rod and 6 or 7 ciliated setae, one of which is much stronger 
than the others and over-reaches tip of claw- Dorsal setae 24 in number, 
arranged 2.6.6.4.4.2, the scapular setae 40ft long, the others 32^ all tapering 
and strongly ciliated. Ventrally with a single seta on each coxa, otherwise 
2.2.6.4.2.2, to 32ft. long. Legs: I 240/4 long, II 21% III 240/i; tarsi I and II 
with dorsal sensory rod, III without any long outstanding nude seta. 
The Standard Data derived from the type and 10 paratypes are : 







Standard 


Theoretical 


Observed 


CocftV of 




Menu 


Deviation 


Bange 


Bange 


Variation 


AW 


49-2:£0-44 


1-79*0.33 


43*85-54-55 


46-0-61*0 


3-6 


PW 


66-9±0-63 


2-52±0-44 


59 -35-74 -45 


63-0-72-0 


3-7 


SB 


19-3±0-34 


1-37+0-24 


15-3-23-3 


18*0-21 -0 


7-0 


ABB 


27-0 


No variation recorded 






PSB 


27-2±0-42 


1.67*0-29 


22-2-32-2 


24-0-30-0 


6-1 


SD 


54-2+0-42 


1-67^:0.29 


39-2-59-2 


51-0-57.0 


3-1 


A-P 


20-5±0.45 


l-80±0-32 


15-1-25-9 


18-0-24-0 


8-8 


AM 


27-0+0.68 


2-33±0-48 


19-4.-34-6 


24-0-30-0 


9-4 


AL 


3O-45±O-70 


2-84±0-56 


21-9-39.0 


27-0-36-0 


9-6 


PL 


33-2±0-42 


l-67±0-29 


27-2-38-2 


30-0-36-0 


5-0 


Sena. 


60-l±0-38 


1.20*0-27 


56-5-63-7 


58-0-63-0 


1-9 



Loc. Eleven specimens collected on boots, Tolga-Atherton Road, Queens- 
land, 2nd Sept., 1944 (R. N. McCulloch). Two other specimens, also collected 
on boots, Wongabel, Qld., 3rd Sept., 1944 (R.N.Mc.) and a single specimen 
found attached to a female Troides priamus (Lepidoptera-Heteroeera), Ather- 
ton Tableland, Qld„ 20th Aug., 1943 (R.V.S.) are also referred to this species. 

This species is very close to gymnodactyla, Womersley and Kohls 1947, but 
differs in having a trifurcate palpal claw, the anterior margin of scutum 
strongly convex and AM in front of line of AL, as well as with the side 
margins of the posterior angle of scutum concave and not convex. 

Tragardhula gymnodactyla (Worn, and Kohls, 1947). 

Eutrombwula gymnodactyla Worn, and Kohls 1947, Tr, Roy. Soc. S. Ausfc, 71, 

(D,3. 

Plate 2, %. J-M. 

Description of Larvae. Shape oval. Length (engorged) to 450/i ? width to 
300ft. Scutum pentagonal, with its anterior margin straight or lightly sinuous 
between AL, and AM in line therewith; sides of posterior angle convex. Sen- 



Womersley— Asiatic-Pacific Scrub Typhus Mites 31 

eillae ciliated on distal half with the bases nearer to each other than to PL and 
slightly anterior of line of PL. Eyes 2 + 2, on distinct shields, posterior eyes 
the smaller. Chelicerae with only the apical tricuspid cap. Galeal setae nude. 
Palpi stout, tibial claw bifurcate ; seta on palpal femur 2-3-hranched ; on genu 
nude; on tibia only the ventral branched; tarsi short with sub-basal sensory 
rod and 5 or 6 ciliated setae, one of which is much stronger than the rest and 
over-reaches tip of tibial claw by the claw's length. Dorsal setae 28. to 3<V, 
long and arranged 2.8.6.6.4.2. Ventrally with a pair of branched setae on 
palpal segment I (gnathosoma), one on each coxa, a pair between coxae I and 
between coxae III, and thereafter uncertain, to 20/x long. Legs i I 210/i long, 
II 195/*, III 210/*.; tarsi I and II with usual dorsal sensory rod; tarsi III with- 
out any long nude seta- 

The Standard Data for the type and 3 paratypes are: 

Standard Theoretical 

Mean Deviation Bange 

AW 45-75±0-75 l-50=fc0-50 41-25-50-25 

PW 60 * No variation recorded 

8B 16-5+0-87 1-73+0-61 11-3-21-7 

ASB 18*0 No variation recorded 

PSB 23-25±0-75 l»50±0-5a 18- 75-23- 75 

SD 41-25±0-75 l-50±0«53 36-75-45*75 

A-P 19-5±0-87 1*73+0-61 14-3-24-7 

AM 24 • No variation recorded 

AL 25-5±0'87 1» 73+0-61 20-3-30-7 

PL 24-0 No variation recorded 

Seng. 47-0+1.0 1-73+0-71 41-8-52-2 

Remarks, This species was described from a gecko Gymnodactylus louisiM- 
densis from Sansapore, Dutch New Guinea. It is very close to the preceding 
new species, pentagon®, but differs in the palpal setae and bifurcate palpal 
elaw and in the shape of the dorsal scutum. 

To it are now referred 5 specimens in the South Australian Museum, col- 
lected from Phyllodactylus marmtirafots (a gecko) on Greenly Island, S. Aust., 
Dec. 1947 (F. J. Mitchell), and 4 specimens from a gecko, approximately 60 
miles south of Darwin, Northern Territory of Australia, May 9, 1943 (B.V.S.). 
The DS are actually 2.8.6.6.4.2, not 2.6,6.4.4.2 as originally given. 

Genus TROMBKTCTLA Berleee 1905. 

Berlese 1905. Acari nuovi Manip. IV, 155, taf. XV, fig-, 4; idem, 1912, "Bedia," 
8, fasc. i, p. 83, fig. 39-44. 

Genotype Trombicula minor Berl. 1905, Adult from Java- 

*= Eutrombicula Ewing 1938, J. Washington Acad. Sci., 27, 293. 

= Aoarisms Ewing 1943, Proe. Ent. Soc. Washington, 45, (3). 



Observed 
Bange 

45-0-48-0 


Coeff. of 
Vaiiatioa 

3-2 


15-0-18-0 


10-5 


21-0-24-0 
39-0-42-0 
18-0-21*0 


6-4 
3-6 

8-8 


24-0-27-0 


6-8 


45-0-48-0 


3-7 



32 Records of the S.A. Museum 

The unjustified separation of Acimscw from Eutrombicula on the number 
of dorsal setae has been pointed out by Michener (1946, Ann. Ent. Soc. 
America, 39, 432). Earlier in the same year and publication, p. 411-417, 
Michener described the nymphs of three American species of Eutrombicttio, 
all of which were found to have one eye on each side, closely adjacent to the 
sensillary area, as in the genotype of TrombicuLa (T. minor Berl.) and as is 
also the case in T. saniboni Worn, from S. Australia, and T. v)ichmanm from 
New Guinea, etc. It seems probable, therefore, that at least the abovemen- 
tioned species, in which the palpal claws of the larvae are bifurcate, will 
when the larvae of tbe genotype are known be correctly placed in the genus 
Trombiada s. str. as defined on adult and/or nymphal characters. 

Key to the Subgenera and Species of TROMBIOULA S. Str. 
from the Asiatic-Pacific Region, 

N.B.~ Where the adults and/or nymphs are known and correlated with 
the larvae the subgenus as based on adult and/or nymphal characters is given 
in parenthesis, but where this is assumed on the homology of larval characters 
the subgeneric name is preceded by a question mark; otherwise no subgenus is 
as yet to be suggested. 

1. Galeal setae branched or ciliated. Setae on palpal femur and genu 
nude j on tibia, dorsal ciliated or branched. Palpal claw bi- or tri- 
furcate. Scutum wider than long, more or less rectangular. Scutal 
and dorsal setae normal, ciliated or with strong denticles, not lanceo- 
late with folded edges or otherwise modified. (Adults and nymphs 
where known with precoxal plates on leg I, forming a longitudinally 
divided sternum. Eyes absent) . . . . . . 2 

Subgen. Leptotronibiditmi Nagayo et al., 1916, 
~= "akamushi" group of Wharton 1947 

Without the above combination of characters . . 19 

2. Postero-lateral corners of scutum well rounded, with PL placed about 
midway between AL and the posterior scutal margin . . . . . . 3 

PL x>laced at the more or less angular postero-lateral scutal corners . 8 

3. Ventral setae of palpal tibia branched. Seta of coxae III on anterior 
margin. DS 44-50, in number arranged 2.10(12).10(12).10.6(8).4.2 r 
fairly thick with strong lateral setules. 

AW 65-5dr7-0 PW 70-3+7-4, SB 29-8±:4*l, ASB 24-0, PSB 
15-0, SD 39-0, A-P 18-5:±3'8, AM 46-4zt6^7, AL 36>5±5-l, 
PL 58-l±r7-2, Sens. 67-0:±8-2. 

Trombicida (?£*) palpalis (Nagayo et al. 1919). 
Ventral setae of palpal tibia nude . . . r . . . , . . 4 

4. Dorsal setae 50 or more in number - . . , - . , . * . 5 
Dorsal setae fewer than 40 in number « . . . . . . , • . 6 



WOMERSLEY — ASIATIC- PACIFIC SCRUB TYPHUS MlTES 33 

5* Dorsal setae 52 in number, arranged ca. 2.12.4.10.4.8.8.4, to 45/a long, 
slender and tapering with filiations rather than setules. Sensillae 
proximally nude. 

AW 58*8, PW 68-6+4-6, SB 28-9+3-4, ASB 27-1^4-3, PSB 
11-2, SB 38-3+4-3, A-P 23-3+4-3, AM 51-5+9-6, AL 44-8 
+5-9, PL 54-9+4-6, Sens, 61-6+6-8, 

Trombkula (?£.) kueJcensckrijveri (WaJch 1923). 

Dorsal setae ca. 70 in number, to 43/* long, thick and strongly setulate 
or dentate, arranged 2.12.14.12.10,8.6.4.2. Sensillae proximally 
barbed. 

AW 70-5+5-1, PW 76-75+9-6, SB 35-6+5-5, ASB 26-7+ 
2-7, PSB 14-8+1-8, SD 41-5+3-0, A-P 19-0+3-8, AM 
49-8+5-6, AL 42-15+5-2, PL 53-3+7-0, Sens. 71-5+7-2. 

Trombioxda ( ? L.) pallida (Nagayo et al. 1919). 

6. Dorsal setae 38-40 in number, to 58/* long, arranged ca. 
2.10.8.8.6.4.2(0), with setules or dentations which are not as long as 
in palldda. Sensillae proximally minutely barbed. 

AW 53-8, PW 63-0, SB 25-8, ASB 24-8, PSB 12-9, SD 37-7, 
A-P 17-2, AM 51-6, AL 38*7, PL 58-1, Sens. 66-7, (Womersley 
and Heaslip 1943, after Nagayo et al., 1921.) 

AW 69-2+5-0, PW 77-8+2-7, SB 33-7+0-8, ASB — , PSB 
13-3+1-6, SD — , A-P 23<5+2-8, AM 49-8+2-3, AL 39-3+9-1, 
PL 54*8+4-0, Sens. 61-5+12-1 (after Kuwata, Berge and Philip 
1950), Trombimda ( ? L.) intermedia (Nagayo et al. 1920), 

Dorsal setae 28-30 in number , . . . , . . . . . . . 7 

7. Scutum smaller, with PW only slightly greater than AW. Setae on 
palpal genu very long and over-reaching palpal claw. DS 30 in 
number, arranged 2.8.6.6.2.6, to 50/x long, sparsely barbed. Sensillae 
ciliated on distal two-thirds, minutely barbed on basal third. 

AW 50-5, 51-5, PW 52-0, 53-0, SB 24-0, 24-5, ASB ca, 20-0, 
PSB 11-0, SD ca. 31-0, A-P 140, 150, AM 29-5, 32-5, AL 
28-5, PL 49-5, 51-0, Sens. 40-5, 42-0 (after Kuwata, Berge and 
Philip 1950, except ASB and SD, which are interpolated from 
their figure) . Trombicfda ( I L.) fuji (Kuwata, Berge and Philip 1950) . 

Scutum larger with PW distinctly greater than AW. Palpal genual 
setae shorter. DS 28, to 58/i long, arranged 2.8.6.6.4.2, thicker and 
strongly dentate. 

AW 58-5+8-9, PW 69-4+7-0, SB 31-1+4-8, ASB 28-0, PSB 
11-2, SD 39-2, A-P 20-85+4-4, AM 40*8+6-6, AL 50-4^8-4, 
PL 56-0+4-2, Sens. 68-3±:7-5. 

Trombicula ( ? L.) lanceolatasy. n. Lawrence in MS. 

8. Palpal claw trifurcate . . - ♦ . . . . 9 

Palpal claw bifurcate . . . . . * . * . - . . . . 15 

9. Ventral seta of palpal tibia branched . . . * , - . . . . 10 
Ventral seta of palpal tibia nude . . . . . . . . * . 12 



34 Records of the S.A. Museum 

10. Dorsal setae 52 in number, arranged ca. 2,13.12.12.8-4.2, to 56> long, 
tapering with strong long eiliations rather than denticles. 

AW 70-l:±6-3, PW 86-8:t9*2, SB 35-5±4*4, ASB 30-8, PSB 
14-0, SD 44-8, A-P 25-2, AM 63-8±3-7, AL 48-5±4-3, PL 63-8 
±:3-7, Sens. 85-4. Trombicida (? L.) parapalpalis sp. n. 

Dorsal setae stout, not tapering, with strong denticles 11 

11. With only 1 humeral seta on each side of first dorsal row. Dorsal 
setae ea. 42 in number, to 40/* long and arranged ca. 2.10.8.10.6.4.2. 

AW 64-4, PW 75-6, SB 308, ASB 28-0, PSB 11*2, SD 39-2, A-P 
25-2, AM 47-5, AL 40-0, PL 50-4, Sens. 61-6. 

Trcmibicula (1 L.) macacus sp. n, Lawrence in MS. 

With 2 humeral setae on each side in first dorsal row- Dorsal setae 
ca. 52 in number, arranged 4.10.8.2.8.8.6.2.2, to 40/t long. 

AW 75-6, PW 81-2, SB 36-4, ASB 28-0, PSB 15*4, SD 43-4, 

A-P 25-2, AM 42-0, AL 33*6, PL 42*0, Sens. — . 

Trombicvla (? £.) pitta sp, n. 

12. Scutum very large, with irregularly wrinkled or shagreened surface. 
Dorsal setae ca. 80-90 in number, to 50*t long, tapering and strongly 
ciliated. 

AW 99-2, PW 121-6, SB 51*2, ASB 41-6, PSB 16-0, SD 57-6, 
A-P 33-2, AM 62«4, AL 56*0, PL 70-4, Sens. 96-0. 

Trontbiada (?i.) d/ux sp. n. 

Scutum smaller, surface punctate. Dorsal setae ca, 50 or fewer in 
number , , 13 

13. Dorsal setae ca T 50 in number, arranged ca. 2.10.10 (12). 10 (8). 8.6.4, 
to 51/* long, more or less tapering and fairly thick with strong eilia- 
tions. 

AW 72*4±6-l, PW 82-2rbl0-4, SB 32-2zt5-9, ASB 29-l±5-5, 
PSB 15-lrtl-2, SD 44-0±5-7. A-P 28-3+5-8, AM 59-8+4-6, 
AL 50-6+10-8, PL 64-3+7-3, Sens. 77-75+5-75. 

Trondkula (?L.) smtettaris (Nagayo et aL 1920). 

Dorsal setae fewer in number 14 

14. Dorsal setae 28 in number, arranged 2.8.6.6.4.2, with very little 
variation. Sensillae bases in line with or only slightly in advance of 
line of PL. Posterior scutal margin more rounded and ca. 12/a behind 
PL. (For populations, see text). 

Trombi-cula (L.) deliensis (Wakh 1923) 
— = vwnderghmstei Gunther 1940. 
— waldhi Worn, and Heasp. 1943. 

Dorsal setae more variable from 34 to 40 in number, generally 38, 
and mostly arranged 2.10 (8). 8.8.6.4.2(0). Sensillae bases distinctly 
in front of line of PL. (For ranges of Standard Data in various 
populations, see text). Tr&mbicula {L.)akamuski¥>mm-pt 1910. 

= fletcheri Worn, and Heasp. 1943. 

*=* obscura Worn. 1944. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 3.5 

15. Dorsal setae very numerous, more than 100 in number, in ca. 10 
rows of 14 each, to 58/* long, slender, tapering and strongly ciliated. 

AW 88-5±7-5, PW 103-0drl0'8, SB 41-2+3-7, ASB 35*8:fc3-7 f 
PSB 13'4±3-7, SD 49-3±T-5, A-P 34-7rt4-6, AM 75-6±6-0, 
AL 62-2±3-7, PL 72-0+7*0, Bens. — . ' Trombicula (1 L.) vUlosa sp.n. 
Dorsal setae much fewer in number 16 

16. Dorsal setae ca. 42 in number, to 70/x. long anteriorly and arranged 
2.12.8.8.6.4.2. 

AW 83*2±4-2, PW 93-8+4-8, SB 36-4+6-8, ASB 35-0+4-8, 
PSB 16-1+4-2, SD 51-1+8-0, A-P 28*7+8-0, AM 51-3+12*8, 
AL 47*6, PL 58-1+8-0, Sens. 78*4. 

Trombicula (?L.) Hthwdlensis sp. n. 

Dorsal setae 28-32 in number 17 

17. Dorsal setae 32 in number, arranged 2.10.8.6.4.2, to 54/a long, tapering 
and slender, and ciliated. Ventral seta on palpal tibia branched. 
Anterior and posterior margins of scutum strongly sinuous, 

AW 70*4, PW 83-2, SB 35-2. ASB 30-0, PSB 12-6, SD 42-6, 
A-P 28-8, AM 32-0, AL 41-6, PL 48-0, Sens. — . 

Trombioula ( ? L.) bhimtalensis sp. n. 

Dorsal setae 28, arranged 2,8.6.6.4.2 » . . . . . . • , . 18 

18. PL scutal setae very long, and more than twice as long as AL or PW. 
Posterior scutal margin a shallow even curve, with SB behind line of 
PL. 

AW 59-9+5-9, PW ttdS&JMfy SB 26-75+4-7, ASB 32-0, PSB 
9-6, SD 41-6, A~P 25-6, AM 55-5+11-0, AL 55-1+6-7, PL 
108-3+24-0, Sens. 69-8+3-7. Trombicula (£.) longi&etasp.n. 

PL not much, if at all longer than PW. Posterior scutal margin 
strongly sinuous and concave medially. SB only slightly behind line 
of PL. Trombicula (L.) fuUeri (Ewing, 1945). 

19. PL and dorsal setae lanceolate or foliate with folded edges carrying a 
double row of setules. Palpal claw bi- or trifurcate. Setae on palpal 
femur and genu nude; on tibia lateral and ventral nude, dorsal 
ciliated or branched. Scutum rectangular with rounded posterior 
corners and PL midway between AL and posterior margin . . 20 

PL and dorsal setae normal, or otherwise than as above . . . . 21 

20. Scutum larger. Palpal claw bifurcate. Dorsal setae 48 in number. 
to 42ft long and arranged ea. 2.10.8.8.8.6.4.2. 

AW 75-0+9-7, PW 84-3 + 10-7, SB 37-3+6-3, ASB 30-4+4-2, 
PSB 16<0, SD 46-4rfc4-8, A-P 19-55+6-25, AM 44-8+9-6, 
AL 43-6+5-0, PL 48*9+7-2, Sens. 60-8 (57-6, 64-0). 

Trombi&ula (Neotrombk'xda) fordi sp. n. Lawrence in MS. 

Scutum smaller. Palpal claw trifurcate. Dorsal setae 52 in number, 
to 30/a long and arranged ca. 2.10.10.10.8.6.2. 

AW 64-4, PW 75-6. SB 33-6, ASB 28-0, PSB 15-4, SD 43-4, 

A-P 22-4, AM 42-0, AL 39-2, PL 36-4, Sens. 60-0. 

<, Trombimda ( t N.) tra/ubi sp. n. 



36 Records of the S.A. Museum 

21. Doraal and AM and PL scutal setae lanceolate, finely ciliated and 
without folded edges. Palpal claw trifurcate. Dorsal setae 30 in 
number, 34-42/t long and arranged ca. 2.8.6.6.4.4. 

AW 67-2, PW 72-8, SB 86-4, ASB 30-6, PSB 14-0, SD 44-8, 
A-P 14-0, AM 30-8, AL 30-8, PL 58-8, Sens. 70-0. 

Trombicula (JV.) hastata (Gater 1932). 

Dorsal and scutal setae normal, more or less tapering with fine cilia- 
tions, or phyllode-like, or broad and aciculate 22 

22. Galeal setae nude * . . . 23 

Galea! setae ciliated or branched , . . . . . . , . , . . 63 

23. Tarsi III with 2, metatarsi III with 1 long nude seta. All 3 setae on 
palpal tibia and the setae on femur and genu nude. Palpal claw 
trifurcate. Dorsal setae 26 in number, arranged 2.6.6.6.4.2, and from 
50ft to 80/a long. Scutum with posterior margin an even curve, deep 
behind PL. SB in line with PL and in midline of scutum. 

AW 70-65+7-45, PW 93-9+9-0, SB 30-3+5-6, ASB 36-15 + 
3-05, PSB 26-65+3-6, SD 62-65+5-45, A-P 33-6+6-75, AM 
55-2+9-0, AL 50-4+5-55, PL 66-85+14-05, Sens. 95-8+9-4. 

. Trombicula n&vae-hottandiae Hirst 1929. 

Tarsi III only with one long nude seta or without 24 

24. Palpal claw bifurcate . . . * . . . . . 25 

Palpal claw trifurcate . . 50 

25. Tarsi III with a long nude outstanding seta sub-basally . , . . 26 
Tarsi III without such, or if present then ciliated . . . . . . 38 

26. Scutum more or less quadrate with SB wide apart and close to AL. 
DS 22 in number, arranged 2.6.6.4.2.2, variable in length in different 
populations from 31/* to 56/a. 

AW 87-6+6*9, PW 94-8+8-0, SB 59-0+5-3, ASB 21-4+6-2, 
PSB 53-0+7-1, SD 74-4+6-3, A-P 41-7+7-2, AM 43-9+9-7, 
AL 38-0+5-6, PL 55-0+9-0, Sens. 58*65+10-45. 

Trombicula (N>) ram (Walchl923). 
Scutum more rectangular with SB not placed as above. , „ . » 27 

27. Dorsal setae 20(22) arranged 2.6.6,4.2(2) 28 

Dorsal setae 26 or more .. ... .. ». .., .. .. 31 

28. Posterior scutal margin relatively deep, its greatest distance from 

line of PL more than twice that between line of SB and PL . . . . 29 

Posterior scutal margins shallower, its greatest distance from line of 

PL equal to or less than that between SB and PL 30 

29. Ratio of PW/SD — 1-35 — 1-52. Seta on palpal femur 3-5- 
branched; on genu usually 1-, occasionally 2-branched; on tibia, 
dorsal and lateral nude, ventral 3-4-brancbecl. DS 20 in number, to 

55/i. long. Trombicula ( ? Trombicula) kirsti Sambon 1927. 



WOMERSLBY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 37 

AW 83'5nt7-3, PW 96-6±7*9, SB 42-8:±:4-6, ASB 29-2±2-6, 
PSB 38-6±5*6, SD 65-4±6-3, A-P 35-9±3'7, AM 40-6:±7<0, 
AL 45*95=t7-l, PL 54-15+4-7, Sens. 64-9±5-4. 

/. typ. hirsti Sainbon 1927. 

AW 84-5±9-4, PW 99-35+9-7, SB 44-35db4-8, ASB 26-7+4-6, 
PSB 38-45+5-2, SD 56-4+5-8. A-P 34-6+5-9, AM 47-1+8-2, 
AL 53-1+8-3, PL 57-7+8-9, Sens. 58-95+11-2. 

/. deliensis (Walchl923). 

« hidoloensis Gunther 1939, 

AW 95-0, PW 110-0, SB 43-0, ASB 33-0, PSB 48-0, SD 81 -0, 
A-P 39-0, AM 58-0, AL 55-0, PL 70-0 r Sens. 70-0. 

/. hakei Radford 1946. 

AW 109-7+8-6, PW 125-7+13-4, SB 58-7+7-1, ASB 32*0, 
PSB 43-0, SD 75-0, A-P 43-0. AM 53-9+5-6, AL 61-2+9-5, 
PL 65-9+13-6, Sens. 60 '0 + 6 -7. /. nnsanemis Dumbleton 1947. 

Ratio of PW/SD ca. 1*86. Seta on palpal tibia with 2-3 short 
branches, often appearing nude; on genu nude; on tibia dorsal and 
lateral nude, ventral branched. DS 20 in number, to 50^ long:. 

Tromhicula ( fl T.) sobrina sp. n. 

30. Setae on palpal femur, genu and tibia (except ventral) nude. 
Scutum irregularly punctate, not striate-punctate. DS 22 in number, 
to 35/x long. 

AW 73-25+4-5, PW 90-2+7-5, SB 36-0+3-4, ASB 21-0, PSB 
31*0+3-0, SD 52-0+3-4, A-P 29-0+2-0, AL 24-5+3-0, PL 
32-5+3-0, Sens. ?. Tronibimla ( f T.) atolephara ap. n. 

Setae on palpal femur 3-5 branched ■ on genu nude j on tibia dorsal 
and lateral nude, ventral 3-4-branched. Scutum striate-punctate. DS 
22 to 35/1. (For Standard Data of different populations see Table C). 

Tr&mMevla (T.) wkhmanni (Ouds. 1905). 

31. Dorsal setae fewer than 40 in number . . . . . . . . . . 32 

Dorsal setae more than 40 in number . . . . . . . , . . 36 

32. Dorsal setae 26(28) in number 34 

Dorsal setae 32 to 38 in number . . . . . . 33 

33. Dorsal scutum larger with posterior margin deeper behind PL. AM 
the shortest. DS 32 in number, arranged 2.8,8.6.4.2.2. to 75-0/jl long. 

AW 103-0, PW 118-0, SB 44-0, ASB 36-0, PSB 23-0, SD 59-0, 
A-P 32*0, AM 43*0, AL 60-0, PL 85-0, Sens. 90-0. 

Trombicula anous (Wharton 1946). 

Dorsal scutum smaller, with posterior margin shallower behind PL. 
AL the shortest. DS 38 in number, arranged 2.10.8.8.4.4.2, to 65-0/a. 
long. 

AW 86-0, PW 90-0, SB 36-0, ASB 29*0, PSB 16-0, SD 45-0, 
A-P 30-0, AM 50-0, AL 40*0, PL 72-0, Sens. 60-0. 

Trombicula pluvius (Wharton 1946). 



38 Records of the S.A. Museum 

34. Setae on palpal femur, genu and tibia all nude. DS 26-28 in number, 
to 55/* long, and arranged 2.6 (8). 6.6.4.2. Scutum irregularly punc- 
tate. 

AW 102*7±6*9, PW 123-O:±6'0, SB 47*6±4-9, ASB 29-0, PSB 
37-l±5-3, SD 65-9±5-3, A-P 33*9+4-4, AM 38*3:h6*2, AL 
47-7zt5-9 > PL 53*1±:5'9, Sens, 64-0. Trombicvla lygosomoides sp. n. 
Seta on palpal femur branched or ciliated . - . . . . * . 35 

35. Dorsal setae 26, uniformly arranged 2.6.6.6.4,2, to 39^ long. Seta of 
palpal femur 4-5-branched j of genu 1-2-branehed ; of tibia dorsal and 
lateral nude, ventral 3-4-branched. 

AW 87'8=hl-7, PW 97-7^12-5, SB 39-7±6'7, ASB 39-0+5-9, 
PSB 29-2±4*3, SD 68-2:fc8-4, A-P 31-6+4-1, AM 40-3:h3-5, 
AL 45-2±6-2, PL 52-l-±5-2, Sens. 72-2^9-9. 

Trombicvla (T.) sambom Worn. 1939. 

Dorsal setae 28, arranged 2.6.6,2 then two lateral clusters of 6 + 6. 
Seta of palpal femur 4-5-branched ; of genu 1-2-branched ; of tibia 
dorsal and lateral nude, ventral 3-4-branched. 

AW 83-3±6'5, PW 92*4+10*4, SB 40-0, ASB 27*0, PSB 34-9±_ 
3-3, SD 62*0+4*0, A-P 33*5+3-4, AM 31-5+6-4, AL 40-1:1: 
4-4, PL 46*1+7-4, Sens. 64-3+2-2. 

Trombicula (#.) sarcina (Worn. 1944). 

36. With 3 humeral setae on each side. Dorsal setae ca. 6.10.2.8.2.10.8.- 
4.2, to 42/* long. Telofemur of leg I with 2 very long outstanding 
eiliated setae, of legs II and III with 1 such. 

AW 70*0, 75-6, PW 89-6, 95-2, SB 30-8, 33*6, ASB 28-0, 28*0, 
PSB 28-0, 28-0, SD 56-0, 56-0, A-P 25-2, 25-2, AM 42-0, 42*0, 
AL 42-0, 420, PL 560, 56*0, Sens. 70-0, 70-0. 

Trambicvla, kmzdl'Wamnsis sp. n. 

With only 1 humeral seta on each side . . . . - « - . * . 37 

37. Dorsal setae 44 in number, arranged 2.6.6.10.8.6.4.2, the posterior five 
rows clustered together. Palpal femur with seta with 6 or more short 
branches,* genu with nude seta; tibia with dorsal and lateral nude, 
ventral with 3-4 branches. 

AW 83*4+6-9, PW 89-9+4-9, SB 36-0+4-2, ASB 27-0+.4-2, 
PSB 34-2+3-3, SD 61-2+6-0, A-P 31-6+2-7, AM 35-7+3*5, 
AL 40-0, PL 54-0+3-3, Sens, 72-0, 

Trombicula ( ? T.) macropus Worn. 1936. 

Dorsal setae 68 in number, arranged 2.14.12.4.6.8.10.8.4, somewhat 
clustered posteriorly. Seta on palpal femur with many short branches 
or dilations; on genu nude; on tibia dorsal and lateral nude, ventral 
3-4-branched. 

AW 100-8+6*5, PW lll-2:±:6'5, SB 55-4+5-6, ASB 29-2+3-3, 
PSB 34*4+40. SD 63-6+7-2, A-P 31-6+3-4, AM 59-8+3*3, 
AL 68-8+8-3, PL 72-0+6*3, Sens. 70*7:1:8*9. 

TromMtnda ( * T.) rim Gunther 1939. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 39 

38. AL scutal setae short and peg-like (genua Fonsecia Badford 1942). 
All setae on palpal femur, genu and tibia (except ventral) nude. 
DS 30 in number, stiff, with indistinct barbs, to 60/t long and arranged 
2.6.6.6.4.4.2. 

AW 90-0, PW 108-0, SB 32-0, ASB 21-0, PSB 30-0, SD 51-0, 
A-P 32-0, AM 60-0, AL 14-0, PL 36-0, Sens. 70-0. 

Trombwuto coluberina (Radford 1946). 

( «= Fonsecia coluberina Radford 1946) . 

AL scutal setae normal . * . . . . . . 39 

39. Scutum smaller, -with AW less than 60/x 40 

Scutum larger, AW often very much more than 60/x 42 

40. Sensillae with only a single branch, half its entire length- Seta on 
palpal femur with 2-4 short branches ■ genu with 1 branch ; tibia 
with all 3 nude. DS 38, arranged 2.8.8.6.6.4.4, to 25/* long. 

AW 39*8±12-2, PW 48-2±12-7, SB 18*6±6-6, ASB ?, PSB 
19-6:1:5*4, SD I A-P 24-7±4-9, AM 27-4±2-7, AL 23*2±4-9, 
PL 30- ±14 a, Sens, 39 -6 ±8-1, (after Philip and Traub 1950). 

Trombtcula batwi Philip and Traub 1950. 

Sensillae otherwise 41 

41. Sensillae with only 2-3 short branches or ciliations on distal half. 
DS 24 in number, arranged 2.6.6.4.4.2, to 25p long. Setae on palpal 
femur and genu nude; on tibia, dorsal with 1 or 2 short indistinct 
barbs, lateral and ventral nude. 

AW 33-0, PW 47-0, SB 12*5, ASB 16-8, PSB 17*0, SD 33-8, 
A-P 17-0, AM 21*0, AL 20*0, PL 30-0, Sens. 30-0. 

Trombicuia (N.) mutida (Gater 1932). 

Sensillae with numerous branches or ciliations distally, DS 26 in 
number, arranged 2.6.4.4.4.4.2, to 35-0/t long. Seta on palpal femur 
with a few short branches ; on genu nude ; on tibia dorsal with 1 or 
2 short branches, lateral and ventral nude. 

AW 38-0, PW 51*0, SB 13-0, ASB 18-0, PSB 22-0, SD 40*0, 
A-P 20*0, AM 23-0, AL 25-0, PL 30*0, Sens. 38*0. 

Trombicuia (N.) spieea (Gater 1932). 

Sensillae slightly thickening apically, with numerous ciliations. Dor- 
sal setae 36 in number to 46//, long and arranged ea. 2.8.6.8,4 T 2. 
Only ventral seta on tibia of palpi branched. 

AW 53-&-60-3, PW 80*4, SB 30*15, ASB 23*45, PSB 10-05- 
13-4, SD 33*5-36*85, A-P 20*1, AM 43-55, AL 33-5-36*85, PL 
60-3-63-65, Sens. 53-6. Trombicvla (N.) con$uetaap.n. 

42. Palpi with femur and genu cylindrical, tibia very small, with the 
claws represented by two short stumpy prongs which are shorter than 
tarsus. Dorsal setae 20 in number, arranged 2.6.6.4.2, from 50/i to 
4f)fi long. Scutum shield-like. 



40 Records of the S.A. Museum 

AW 63*0±4-7, PW 73-25+4-45, SB 36-7+2-5, ASB 22-4, PSB 
38-3±:5-0, SD 60-7+5-0, A-P 29-75+4-05, AM 45-6+6-5, AL 
31-85+7-45, PL 49-5+6-5, Sens. 53-7+6-7. 

Trombicula parmifera sp. n. 

Palpi and palpal claw normal . • . • 43 

43. Scutum conspicuously striate-punctate, and large 48 

Scutum smaller and not striate-punctate . . . , . . . - 44 

44. Setae on palpal femur and genu nude. All setae on palpal tibia nude. 
DS 22, arranged 2.6.6.4.2.2, to 36/*, fine with only indistinct ciliations. 
Setae on palpal femur and genu branched . . 45 

All setae on palpal tibia nude. DS 22, arranged 2.6.6.4/2.2, to 36/i, 

fine with only indistinct filiations. 
AW 78-85+7-8, PW 85-4+9-0, SB 44-1+5-2, ASB 21-0, PSB 
37-8+5-3, SD 58-8+5-3, A~P 29-4+3-6, AM 24-1+6-4, AL 
22 • 0+. 5 • 5, PL 22 • 55+7 - 8, Sens. 50 ■ 0. Trontbivula Umdbladd sp. n. 

45. Scutum rectangular, much wider than long, with AW not much 
shorter than PW. shallow behind PL, and with anterior and posterior 
margins almost rectilinear. Sensillae rather thick with ciliations on 
distal two-thirds, bases nearer to one another than to lateral margin 
and in front of line of PL. Setae on palpal femur and genu branched, 
on tibia all three nude. Dorsal setae 28 in number, to 55/* long, and 
arranged 2.8.6*6.4.2. 

AW 66-0, PW 69-0, SB 23-0, ASB 26-0, PSB 130, SD 39-0, 
A-P 29-0, AM 26-0, AL 33-0-36-0 (34*5), PL 43-0-46-0 (44-5), 
Sens. 36-0 (after Dumbleton). Trombicula rvissam Dumbleton 1947. 

Scutum not as above . .. . * .. . . . . ► . . ► . 46 

46. Scutum much wider than long, but with AW much shorter than 
PW, and the lateral margins diverging widely posteriorly; anterior 
and posterior margins almost rectilinear. Dorsal setae ca. 38 in num- 
ber, to 30/a long and arranged 2.8.6.6.6.6.4. 

AW 56-0, PW 86-8, SB 44-6, ASB 28-0, PSB 12-0, SD 40-0, 
A-P 50-4, AM 25-2, AL 22-4, PL 50-4, Sens. — . 

Tr&mfoicula pibbidporensis sp. n. 

Scutum not much wider than long, with AW only a little less than 

P W .. .. .. .. .. .. .. . . .. .. 47 

47. Scutum with posterior margin deep and rounded behind line of PL ; 
anterior margin convex with AM slightly in front of AL; sensillae 
bases nearer to lateral margins than to each other and in front 
slightly of line of PL. Setae on palpal femur and genu, and the 
ventral on tibia branched. Dorsal setae 26 in number, to 56-O/x long, 
and arranged 2.6.6.6.4.2. 

AW 72-0, PW 78-0, SB 31-0, ASB 27-0, PSB 28-0, SD 
55-.0, A-P 28-0, AM 58-0, AL 43-0, PL 74-0-78-0, Sens. 97-0 
(after Sugimoto) , Trombicula isshikii Sugimoto 1938. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 41 

Scutum with the posterior margin angular, giving a pentagonal form ; 
depth of posterior margin less than distance between SB and line of 
PL. AL setae placed rather well back from the rounded anterolateral 
shoulders and behind AM. All setae on palpal f emur, genu and tibia 
branched. Dorsal setae 44 in number, to 38/a long, and arranged 
2.4.6.6,4.6.6.4.4.2. Coxae III 1-setose. 

AW 61-6, PW 73-0, SB 22-4, ASB 28-0, PSB 28-0, SD 56*0, 
A-P 30-0, AM 36-4, AL 30-8, PL 47-6, Sens. 64-4. 

Trombieula khurdmgen$i$&j). n. 

48. Scutum fairly large with AW 97-0-122-0, and PW 112- 0-132 -0. 

AW 98*2+5-6, PW 113-5+6-5, SB 51-05+3-65, ASB 28-0+ 
3-4, PSB 26-6+4-2, SD 54-6+5*8, A-P 30-4+4-7, AM 39-9+ 
5-2, AL 39-2+3-4, PL 47-6+3-4, Sens. 57-4+6-7. 

Trombieula (N,) scincoides (Worn. 1944). 
/. typi>ca. 

AW 105-4+8-8, PW 116-7+13-0, SB 55-1+5-7, ASB 30-8+ 
4-5, PSB 28-0, SD 58-8+4-5, A-P 32-8+4-1, AM 46-95+4-15, 
AL 44-0+4-8, PL 51-3+4-9, Sens. 64-4+70-0. 

TronibicuU (N.) scincoides (Worn. 1944) 
/. ex Hollandia. 

Scutum larger or smaller than above . . . . . * « - . . 49 

49. Scutum larger than above. 

AW 110-8+8-2, PW 124-4+0-56, SB 62-4+6-3, ASB 34-8+: 
4-5, PSB 33-6, SD 68-4+4-5, A-P 36-8+3-2, AM 45-6+4-1, 
AL 47-6+4-8, PL 55-6+3-2, Sens. 63-6+4-1. 

Trombieula ( ? N.) kohlsi (Worn. 1944). 
Scutum smaller than above. 

AW 77-9+3-4, PW 89-1+8-2, SB 40-1 + 6-85, ASB 27-1+4^3, 
PSB 25-7+3-4, SD 52-7+3-4, A-P 26-1+4-3, AM 29-4+4-6, 
AL 30*8, PL 41-5+3-4, Sens. 56-0. (For other populations see 
text.) Trombieula ( ? JV.) tovelli sp. n. 

50. A long nude seta on tarsi III . * . . . . . . . . . . 51 

No long nude seta on tarsi III . . . . - * . . . . • . 53 

51. Scutum small; shield-shaped; with anterior margin deeply concave, 
and posterior margin forming a shallow angle. Sensillae ciliated their 
whole length and SB in front of PL. AL well back from shoulders 
and behind AM. All setae on palpal femur, genu and tibia nude. 
Coxae III bisetose. DS thick and strongly ciliated, to 48/* long and 
arranged 2.6.6(4). 4(6). 4.2.2. 

AW 45-3+6-6, PW 46-6+5-1, SB 18-7+2-3, ASB 28-8, PSB 
22-4, SD 51-2, A-P 25-6, AM 38-4, AL 22-4, PL 41-6, Sens, 
54 - 7 +7 • 1. Trombieula buxtoni sp. n. 

Scutum larger, not shield-shaped, more or less rectangular, or with 
deeply curved posterior margin . . . . . . . . - . 52 



42 Records of the S.A. Museum 

52. Posterior scutal margin a deep curve behind line of PL with PL 
nearer to AL than to posterior margin, and SB behind line of PL, All 
setae on palpal femur, genu and tibia fine and nude. DS 22 in num- 
ber, to 50/a long, and arranged 2.8.6.4.2. 

AW 81-0^82-0 (81*5), PW 90-0-91-0 (90-5), SB 36-0-40-0 
(38-0), ASB 26-0-30-0 (28-0), PSB 20-0, SD 46-0-50-0 (48-0), 
A-P 16-0, AM ?, AL 23-0-26-0 (24-5), PL 49-0, Sens. 69-0 
( after Dumbleton). Trombicula naultini Dumbleton 1947. 

Posterior scutal margin not so rounded except laterally, medially 
rather flattened. PL nearer to posterior than to anterior margins and 
SB in front of PL. All setae on palpal femur, genu and tibia (except 
ventral) nude. DS 32 in number, to 40/x and arranged ca. 2.6.6.6.6.4.2. 

AW 65-2-72-0 (67-5), PW 68-5-81-5 (73-9), SB 26*0^32*6 
(28-2), ASB 23-4, PSB 23-4, SD 46-8, A-P 26-0-32-6, AM 
39-1-45-6 (42-2), AL 39-1-45-6 (41-3), PL 42-5-65-2 (49-9), 
Sens. 65-0-81-5 (72-8). Trombicula cervulicola Ewing 1931. 

53. With more than 50 dorsal setae 54 

With fewer than 50 dorsal setae „ * . . 56 

54- Dorsal setae 68-70 in number 55 

Dorsal setae 58 in number, to 50^l long and arranged 2.8.8.12.12.6.4.4.2. 
Scutum rectangular between PL, and with SB very much nearer to 
line of PL than to line of AL. Seta on palpal femur branched, 
(Galeal setae stated by Walch to be 1-branched). 

AW 69-0 (72-0), PW 75-0 (80-0), SB 24-0 (22-0), ASB 27-0 
(28-2), PSB 10-0 (11-0), SD 37-0 (39-2), A-P 30-0 (37-0), 
AM 42-0 (37-0), AL 45-0 (51-0), PL 62-0 (51-0), Sens. 50-0 
(45-0). (The values in parentheses after Walch.) 

Trombicula densipildata Walch 1923. 

55. Sensillae ciliated distally, with bases fairly close together. Anterior 
and posterior scutal margins strongly sinuate. Dorsal setae ca. 70 
in number, to 30/w. long, arranged approximately 2.10.10.10.10.10.8.6.4. 

AW 64-4, PW 87-0, SB 19-6, ASB 33-6, PSB 14-0, SD 47-6, 
A-P 39-2, AM 20-0-f, AL 40-0, PL 50-4, Sens, 56-0. 

Twrnbictda taphozous ap. n. 

Sensillae only ciliated basally, wide apart and in front of line of PL. 

Setae on palpal femur and genu strongly branched, on tibia all 

3 nude. Anterior and posterior scutal margins at most lightly sinuate. 

Dorsal setae 68 in number, to 45/x. long and arranged 2.10.10.16.12.10.8. 

PW 85-0, SD 47-0, Sens. 58-0, PW/SD = 2-3 (after Kaw. and 
Yam. 1921) . Trombicula cot*vi Kawamura and Yamaguchi 1921. 

56. Scutum with posterior margin almost rectilinear between PL . . . . 57 
Posterior margin of scutum not rectilinear but deep behind line of 

-L l~i .. • - .. ., .. ., ,, ij ,. Otr 



Womersley— Asiatic-Pacific Scrub Typhus Mites 43 

57, Scutum trapezoidal with AW much less than PW. Sensillae bases 
nearer to line of PL ; at about three-quarters of scutal depth. Setae 
of palpal femur, genu and tibia all nude. DS ca. 42 in number, to 
45/* long and arranged 2,8 (9). 8 (10). 8 plus about 20. 

AW 48-0-52-0 (50-0), PW 65-0-69-0 (66-3), SB .22-0-23*0 
(22-7), ASB ea. 33-0, PSB 11-0-12-0 (11-7), SD ca. 44-0, A-P 
35-0-36-0 (35-2), AM 44-0-46-0 (45-0), AL 42-0, PL 57-0-64-0 
(60-3), Sens. 65-0-680 (66-5). (After Philip and Traub 1950). 

Trombicula imotti Philip and Traub 1950. 
Scutum almost rectangular, with AW only slightly less than PW, 
Sensillae bases about midway between lines of AL and PL. Setae on 
palpal femur and genu long branched ; on tibia, dorsal nude, lateral 
and ventral branched. DS to 60/* long, tapering and finely ciliated, 
38-40 in number, and arranged 2.8.6-6.6.6.4.2. 
AW 62-0+5-2, PW 78-3+10-5, SB 24-0, ASB 24-0. PSB 24-0, 
SD 48-0, A-P 38-3-±6-2, A,M 57-0, AL 35-0+5-2, PL 62-Ozt 
5-2, Sens. ? Trombiwlaleverisp, n. 

Scutum about as in insolU, with sensillae bases at about two-thirds 
distance from AL to PL. All setae of palpal femur, genu and tibia 
branched. DS 32 in number, to 36/a long and arranged ca 2.6.6.6.6.4.2. 
AW 47-9+2-8, PW 61-9+8-9, SB 19-6, ASB 30-2+3-5, PSB 
14-0, SD 44-2+3-5, A-P 38-1+8-1, AM 41-1+4-3, AL 42-35+ 
2-9, PL 47-6, Sens. 68-6. Tromlicuta (i\T.) tomtom sp. XL 

58. Scutum as long as wide between AL, with SB in front of line of 
PL. AL the shortest, PL the longest. All setae on palpal femur, 
genu and tibia (except ! ventral) nude. DS 38 in number, to 56/i 
long and arranged 2.8.8.8.6.4.2, 

AW 58-7, PW 81-5, SB 26-1, ASB 34-5, PSB 27-6, SD 62-1, A-P 
41-4, AM 55-2, AL 34-5, PL 76-0, Sens, — . 

Trombicula piercei Ewing 1931. 
Scutum distinctly wider than long 59 

59. Scutum pentagonal . . . . M 60 

Scutum not pentagonal 61 

60, Scutum small j its posterior angle more acute and with short internally 
radiating lines. Seta on palpal femur with 2-3 short branches; on 
genu nude; on tibia only the ventral branched. Dorsal setae 22 in 
number, and arranged 2.6.6.4.2.2. 

AW 47-6+3-8, PW 63-1+9-5. SB 17-4+3-8, ASB 20-1+5-1, 
PSB 22-4, SD 42-5+5-1, A-P 17-3+3-4, AM 31-1+5-0, AL 
25-75+6-6, PL 32-3+4-4, Sens. 50-4. 

Trombicula (T.) frittsiWh&riou 1945. 

Scutum larger, its posterior angle more obtuse, without internally 

radiating lines. Setae on palpal femur, genu and tibia as above. 

Dorsal setae 44 in number, to 47^ long, and arranged ca. 

2.13.9.8.5.5.2. 

AW 81-9+4-2, PW 98-7+4-2, SB 34-6+4-0, ASB 29-4+4-8, 
PSB 33-6, SD 63-0+4-8, A-P 30-8, AM 42-0+6-9, AL 42-7± 
4*2, P& 54-6. Sens, 84-0. Trombicula 'l*a$hniir?n$is£V~n. 



44 Records of the S.A. Museum 

61. Seutuni small, posterior margin moderately deep behind line of PL 61a 
Scutum larger, posterior margin not so deep behind line of PL and 
more or less flattened medially - . - - » 62 

61a. Posterior margin deep behind PL, broadly rounded medially and 
lightly concave laterally. Seta on palpal femur 4-5 branched; on 
genu 1-branched j on tibia all three nude. Dorsal setae stiff and short, 
to 20/* long, 32 in number, arranged 2.6.6.4.6.4.2.2. 

AW 50-4, PW 67-2, SB 19-6, ASB 25-2, PSB 19*6, SD 44-8, 
A-P 22-4, AM 19-6, AL 16-8, PL 25«2, Sens. 57-6. 

Trombiada ineurva sp. n. 

Posterior margin not so deep, medially lightly concave. Seta on pal- 
pal femur strongly branched, on genu nude, only ventral seta on 
palpal tibia branched. Dorsal seta 30 in number to 40/* long, tapering 
and ciliated, and arranged 2.8.6.6.4.2.2. 
AW 50-9dbl3-35, PW 58-95±13-5, SB 19-8±2-l, ASB 20-1, 
PSB 10-05, SD 30-15, A-P 20-1, AM 26-8, AL 26-8, PL 30*15, 
g eng m Trombicula vietzi&pn* 

62. Dorsal setae 28 in number, arranged 2.8.6,6.4.2, to 50/* long. Palpal 
femur and genu with long curved and strongly branched setae ; tibia 
with all 3 setae nude. Body ovate. 

AW 74-8±5-9, PW 850+5-5. SB 25-5zh5-0, ASB 29<0zfc4-2, 
PSB 17-6±2-6, SD 46-5+5-1, A-P 29-7+1*0, AM 31-9+4-3, 
AL 39*6+5-9, PL 53-8+9-3, Sens. 45-25+5-7. 

TrombicxiU (L.) myztmthasp.n. 

Dorsal setae 34 in number, arranged 2.12.8.6.4.2, to 50/* long. Setae 
on palpal femur and genu branched ; on tibia all 3 nude. Body cor- 
date, widest across shoulders. 
AW 72-6+4-0, PW 78-0+6-3, SB 24*0, ASB 32-4+40, PSB 
15-0, SD 46-8+4-9, A-P 30-6+4-0, AM 42-0, AL 47-4+4-0, PL 
52-8+4-9, Sens. 50-0. Trombicula (*£.) robusta (Gunther 1941). 

63. PL scutal and all dorsal setae broadly foliate or phyllode-like . . 73 
All scutal and dorsal setae normal . . . . . * - - • - 64 

64. Palpal claw bifurcate . . * . . . . • * • • • 85 
Palpal claw trif urcate - . • . . . • • • ■ • - • - 68 

65. Scutum roughly pentagonal, with the depth of posterior margin much 
greater than distanee between SB and line of PL. AL setae set well 
back from the rounded anterolateral shoulders and behind AM. All 
setae on palpal femur, genu and tibia nude. Coxae HE bi-setoee. Dor- 
sal setae stiff and spine-like, curved, ca. 50 in number, to 40/x. long and 
arranged 2.6.6.6.6.6.6.6.4.2. 

AW 53-2, PW 56-0, SB 22-0, ASB 30-8, PSB 33-6, SD 64-4, A-P 
28-0, AM 40-0, AL 25-2, PL 50-4, Sens. 56-0. 

Trombicula rajoriensis sp.n. 

Scutum not pentagonal , 66 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 45 

66. Scutum about as long as wide between AL. Dorsal setae thickened 
basally then tapering, finely ciliated, 26 in number to 87/a long and 
arranged 2.8.6.6.4. Setae on palpal femur and genu nude, on tibia 
only ventral branched. 

AW 72-8-78-4 (aver. 74-7), PW 92-4-98-0 (94-3), SB 36-4-42-0 
(38-7), ASB 44-8-47-6 (45-7)* PSB 19-6-22*4 (21-5), SD 67-2, 
A-P 44-8-47-6 (45-7), AM 72-8-75-6 (74-2), AL 50-4-56-0 
(54-1), PL 89-6, Bens.—. 

Trorribicula sylvestris Audyand Traub 1950. 

Dorsal scutum much shorter than wide between AL, almost rectilinear, 

but with posterior margin shallow . . 67 

67. Setae on palpal femur and genu branched, on tibia only dorsal cili- 
ated. Posterior shallow and evenly curved behind PL. Dorsal setae 
26 in number, to 53/a long, and arranged 2.8.6.6.4* 

AW 58-8, PW 72-8, SB 32-0, ASB 28-0, PSB 12-0, SD 40-0, A-P 
22-4, AM 53-2, AL 56-0, PL 64-0, Sens. 60-0, 

Trombicwa muridia sp. n. 

Setae on palpal femur and genu nude, on tibia only dorsal ciliated, 
Posterior margin sinuous but almost rectilinear between PL. DS 28 
in number to 66/a long, and arranged 2.8.6,6.4.2. 

AW 64-5rfc6*7, PW 71-3+6-7, SB 301±:3-5, ASB 25-0, PSB 
11-0, SD 36-0, A-P 29-8±4-0, AM 51-5dr6-9, AL 52-3±9-l, 
PL 58-25+11-5, Sens. 68-55+8-8. 

Trombkxduhodenm (6untherl940). 

68. Dorsal setae 40 or more in number 69 

Dorsal setae 28 to 34 in number 70 

69. Dorsal setae 40 in number, to 45/* long and arranged 2.10.8.8.6.4.2. 
Setae on palpal femur and genu branched ; on tibia, dorsal and lateral 
nude, ventral branched. 

AW 70-5+5-1, PW 83-5+9-0, SB 25-5+2*5, ASB 28-3+2-5, 
PSB 13-6+2-7, SD 41-9+5-4, A-P 24-4:4=5-4, AM 33-2+9-4, 
AL 38-7+10-5, PL 51-2+5-4, Sens. 63-0. 

Trorribieula gllri-coUns (Hirst 1915). 

Dorsal seta 54 in number, stiff and needle-like, arranged 
2.10.8,10.12.6.4.2. Seta on palpal femur 2-3 long branches; on genu 
1-2 long branches; on tibia, dorsal and lateral nude, ventral 
branched. 

AW 64-5+6-3, PW 79-5+10-9, SB 22-6+4-3, ASB 24-9+5-9, 
PSB 15-3+2-2, SD 40-2 + 6-2, A-P 23-6+4-1, AM 35-75+5-6, 
AL 27-8rir6-0, PL 32-0+4-9, Sens. 55-3±ll-6. 

Tronibicula plriUpi sp. n. 

70. Dorsal setae of palpal tibia ciliated or branched, lateral and ventral 
nude; setae on femur and genu ciliated or branched. Sensillae 
slightly behind line of PL, with short dilations or barbs basally, 
Irfnger distally. DS 28 in number, to 64/i, arranged 2.8.6.6.4.2. 



46 Records of the S.A. Museum 

AW 70-4, PW 78-9+9-5, SB 33-5+4-1, ASB 31*4+4-3, PSB 
14-7=t:2-l, SD 46-1+5-2, A-P 27-2+4-8, AM 59-5+8-5, AL 46-7 
+5-2, PL 71-0+4-3, Sens. 70-4. 

Trombicvla (L.) burmensis (Ewing 1945), 

All 3 setae of palpal tibia, as well as the setae on femur and genu 
ciliated or branched. Scutum rugose, with SB in front of line of 
PL and posterior margin as even shallow curve. DS 28 in number, 
to 30/x long, thick with short setulations and arranged 2.8.6*6.4.2. 

AW 56-7+6-9, PW 78-0, SB 18-0, ASB 21-0, PSB 18-0, SD 39-0, 
A-P 30-0, AM 30-0, AL 25*0+5-2, PL 36-0, Sena. 50-0. 

Trombicu<la soutkootti sp. n. 

Only the ventral of the palpal tibial setae branched 71 

71. Scutum short, with PW much greater than AW, so that the angle at 
AL is very obtuse ; posterior margin evenly crescentic ; sensillae bases 
in line with PL ; surface strongly rugose. Setae on palpal femur and 
genu ciliated. DS 28 in number, to 40/* long, and arranged 2.8.6.6.4.2. 

AW 58-8+5-3, PW 83-8+4*9, SB 20-2+6-1, ASB 21-0, PSB 
16-75+8-7, SD 38-25+8-6, A-P, 23-5+6-2, AM 30-75+4-5, AL 
27-0, PL 440+6-7, Sens. 50-0. 

Tronibicula quadriense Worn, and Heasp. 1943. 
= chiroptera Worn, and Heasp. 1943. 

Scutum more rectangular with posterior margin sinuous . . . . 72 

72. Scutum smaller, with strong sparse punctations, and sensillae behind 
line of PL. Seta of palpal genu nude. DS 34 in number, short to 28/*, 
arranged 2,8.6.6.6.4.2. 

AW 43-2, PW 53-2, SB 16-0, ASB 21-8, PSB 7-0, SD 28-8, A-P 
16-0, AM 30-0, AL 28 -8 t PL 35-6, Sens. — . 

Trambicnda pelta sp. n> 

Scutum larger, surface only lightly punctate. Sensillae bases in line 
with PL. Seta of palpal genu ciliated or branched. Dorsal setae 30 
in number, to 50/i long and arranged 2.8.6.6.4.2.2. 

AW 67-2, PW 81-2, SB 28-0, ASB 25-2, PSB 11-2, SD 36-4, 
A-P 25-2, AM 50-4, AL 44-8, PL 50-4, Sens. 56-0. 

Trombicula (N .) jayewickremei sp. n. 

73. PL and dorsal setae aciculate-f oliate with longitudinal rows of setules 74 

PL and dorsal setae broadly phyllode-like with fine punctations or with 
large-reticulations . . . , , . , . .... . . . . 75 

74. Dorsal setae 32 in number, to 84-0/* X 28-0/t, arranged 2.8.6.6.4,4.2, the 
posterior rows shorter. Ventrally without any foliate setae; the setae 
behind coxae III short, to 30/i and arranged 6.6*4.2.2.2. 

AW 84-0, PW 106-+-1120, SB 50-4, ASB 33-6, PSB 22-4, SD 
56-0, A-P 16-8, AM 75-0, AL 36-4, PL 84-0 X 28-0, Sens. — . 

Trombieula oune&ta (Traub and Evans, 1951). 
Trombicidindus cmieatus Traub and Evans, 1951. 



Womersley— Asiatic- Pacific Scrub Typhus Mites 47 

Dorsal setae 40 in number, arranged 2.6.6.8.8,6.4, not much shorter pos- 
teriorly, and the first 6 between the humerals much more slender than 
the rest. Ventrally behind coxae III with ca. 32 long, slender, and 
fine aciculate-foliate setae as on the dorsum. 

AW 95-2, 98-0, PW 117-6, SB 47-6, ASB 39-2, PSB 28-0, SD 
67-2, A-P 28-0, AM 75-6, 72-8, AL 47-6, 53-2, PL 112-0 X 42'0, 
106-8 X 42-0, Sens. — . Trombicula squamifera sp. n. 

75. PL and dorsal setae very broad and overlapping and entirely cover- 
ing dorsum, finely punctate, 28 in number, arranged 2.8.6,6.4.2. 
Ventrally behind coxae III with ca. 20 long, ciliated setae followed 
by ca. 16 setae similar to those on the dorsum. 

AW 88-0, PW 100-0, SB 400, ASB 36-0, PSB 11-0, SD 47-0, 
A-P 31-0, AM 73-0, AL 67-0, PL 75-0, Sens. 80*0 X 54-0. 

Trombicula squamosa (Radford 1947) . 

(*= Trombiculindus squamosus Radford 1947) . 

PL and dorsal setae smaller, not overlapping, with large retieulations, 
30 in number, arranged 2.8.6.6.4.2.2. Ventrally behind coxae III 
with 8.4.2.2. short fine ciliated setae only. 

AW 70-O, PW 86-8, SB 44-8, ASB 30-8, PSB 16-8, SD 47-6, 
A-P 14-0, AM 39*2, AL 36-4, PL 78-4 X 39-2, Sens. — i 

Trombicula foliacea (Traub and Evans, 1951). 
(=Trcmibiculi7idus foliaceus Traub and Evans 1951). 

Bubgonus LEPTOTROMBIDIUM Nagayo et &L 19174 

Leptotrombidwm Nagayo et al., 1917, J. Exper. Med., 25, (2), 2-3. Type 
Trombicula akwnushi Brumpt. 

The above name was used generically by Nagayo and his co-authors for 
Trombicula akamushi when, on rearing the nymphs and adults they found 
that these differed from the adults of other species of Trombidiids then known 
to them. In a later paper, and subsequently, they dropped the name Lepto- 
trombidium on realizing that the adults and nymphs conformed to Berlese's 
genus Trombicula Berl. 1904. They did not however, point out that the 
Japanese species differed from the type of Trombicula {minor Berl. 1904) in 
lacking eyes. 

In the adult section of the present work, however, it is shown that oka- 
miishi, deliensis and other allied species do not only differ from the genotype of 
Trombicula s, str. in lacking eyes, but that they are also characterized by pos- 
sessing distinct precoxal plates on leg I which meet in. the median line to form 
a longitudinally divided sternum (as was well figured originally by Walch for 
dreliewis). 

In 1946, Wharton stressed the fact that the two species, proven to be vectors 



48 Records of the S.A. Museum 

of teutsugamushi disease, akamushi and deliensis, together with other closely 
allied species, form a small group which he termed the "cukamiirShi" group. On 
the basis of the larvae, he further showed that this group might not be con- 
fined to the Asiatic-Pacific Begion as he was unable to distinguish an American 
species T~ myotis Ewing, except on the ornamentation of the surface of the 
dorsal scutum. 

Of the species here placed in Leptotrornbidi'ivni on larval characters only, 
akamushi Brumpt, pallida Nagayo et ah, deliensis Walch, hurmensis Ewing 
and myzaniha sp. n. are also known from the adult or nymphal stages. In 1921, 
Nagayo and colleagues claimed to have reared nymphs of three other Japanese 
species, palpalis, intermedia and Scutellaria, but were unable to separate them 
from pallida or akamushi No further studies of these species, however, appear 
to have been made. 

On larval characters the sub-genus may be diagnosed as follows: Palpal 
claw bi- or trif urcate. Setae on palpal femur and genu nude ; on tibia the dorsal 
ciliated, the lateral and ventral (except in palpalis, etc.) nude, Galeal setae 
strongly ciliated (nude in myzontjia)* No long nude, outstanding setae on 
tarsi or metatarsi of leg III. Dorsal scutum trapezoidal. Scutal and dorsal 
setae long, tapering, ciliated or with fairly strong denticles, 

Trombicula (?Leptotrombidium) palpalis (Nagayo et ah 1919). 

Trvmbicula palpalis Nagayo et al. 1919, Verhd. d. jap, pathoi Gesellscht., 
Tokyo. 10, 107; idem 1920, 10, 471; idem Amer. J, Hyg., 1, (5-6), 1921 \ 
Womersiey and Heaslip, 1943, Trans. Roy. Soc. S. Aust., 67, (1), 90. 

Plate 3, fig. A-F. 

A considerable number of specimens, remounted separately, from mixed 
original preparations kindly given to Lt.-Col. C. B. Philip by Dr. T. Tainiya, 
from Yamagata Prefecture, Japan, have been studied. 

This species, which is as yet unknown outside of Japan, agrees with the 
following four species in not having the PL setae placed distinctly at the 
posterolateral corners of the scutum. It differs from these, and the preceding 
three species also, in that the ventral seta on the palpal tibia is branched and 
in having the seta of coxae HI plaeed on the anterior eoxal margin. Other 
specific differences lie in the number and arrangement of the dorsal setae. 

Over thirty remounted specimens, from Okiage and Orato in Yamagata 
Prefecture, Sept. to Dec, 1920, have been studied. On the original slides this 
species was indicated by the sign ( J> ) or $ho, translated by Mr. N. B. Tindale 
as meaning " small" and presumably referring to the more slender dorsal seta 
than in pallida. No recent material has been seen. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 1 49 

Re-description of Larvae (mainly after Nagayo). Colour in life orange 
red. Shape oval. Length (unengorged) to 264/x, width to 172/*. Scutum rec- 
tangular with PL at the posterior corners; the posterior margin deep behind 
SB, straight medially and strongly curved laterally; the anterior margin sinu- 
ous ; sensillae filamentous, nude on proximal fourth, strongly ciliated on distal 
three-fourths, bases slightly behind line of PL j PL setae the longest. Eyes 2 -|- 2, 
close to lateral borders of scutum. Chelieerae with only the apical tricuspid 
cap. Galeal setae strongly ciliated. Palpi stout, with trifurcate tibial claw; 
all setae on femur, genu and tibia, except the dorsal and ventral on tibia, nude. 
Dorsal setae, more slender and tapering than in pallida or intermedia but still 
with rather strong setules, to 51/x, long, ca. 44 in number, and arranged* 
2.10.10.10.6.4.2. Ventral setae similar to dorsal, one on each coxa with that 
of coxae III placed on anterior margin, a pair of setae between coxae I and 
between coxae III and thereafter ca. 45 setae. Legs as long as and similar 
to pallidal. 

The Standard Data derived from 28 specimens are as follows : 



AW 


Mean 
65*5±0.44 


Standard 
Deviation 

2-34±0-31 


Theoretical 

Bange 

58-5-72-5 


Observed 
Bange 

63-0-70-0 


Ooeff . of 
Variation 

3-6 


PW 


70-3±O-41 


2-45^:0-30 


63-6-77. 


66-0-75*0 


3-2 


SB 


29-8±0-26 


1-37+0-18 


25-7-33-9 


27 -0-33- 


4-6 


ASB 
PSB 


24*0 
15-0 


No variation recorded 
No variation recorded 






8D 


39-0 


No variation 


recorded 






A-P 


18-5±0-24 


1-27+0-17 


14-7-22-3 


15-0-21-0 


7-0 


AM 


46-4±:Q-44 


2-24^:0.31 


3d -7-53-1 


42-0-51-0 


4-8 


AL 


36-5:±0.32 


1- 70+0 -23 


31-4-41-6 


33-0-42-0 


4-6 


PL 


58-l±0*46 


2-42r!:0-32 


50-8-65-4 


54-0-63-0 


4-2 


Sens. 


67-0±l-22 


2-74+0-87 


58-8-75-2 


65-0-70-0 


4-1 



Trombicula [1 Leptotrombidium)keukenschrijveri Walch 1923. 

Tromlicula keuhenschrijveri Walch 1923, Tr. Yth. Bien. Cong. Far East. Assoc. 
Trop. Med., 583, Singapore (1924) ; Womersley and Heaslip 1943, Tr. 
Roy. Soc. S. Aust, 67, (1), 1943. 

Trombicxila (Trombicula) keukensehrijveri, Sig Thor and Willmann 1947, Das 
Tierreich, Lfg., 71b, 268. 

Plate 3, fig. G-J. 

This species was originally described from a single specimen on man at 
Deli in Sumatra. Until now, no further specimens appear to have been 
recorded. 

Recently, however, Dr. J. R. Audy has forwarded me a slide from amongst 
the Gater material left at the I.M.R., Kuala Lumpur, and labelled by Gater as 



50 Records of the S.A. Museum 

keyJcenschrijveri, and also a number of freshly collected specimens from Kuala 
Lumpur, which agree with this species. 

A fresh description based on this material is as follows : 
Larvae. Shape oval. Length (engorged) to 495/*, width to 420^. Scutum 
as figured, with the posterior corners well rounded, and PL about ^midway 
between the anterior and posterior scutal margins; scutal setae relatively thin 
and shortly ciliated, AL the shortest, PL the longest; sensillae filamentous and 
ciliated on distal half, the bases behind line of PL. Eyes 2 + 2, posterior 
the smaller, Chelicerae simple, with only the apical tricuspid cap. Palpi 
stout, tibial claw trifurcate; setae on femur and genu nude; on tibia, dorsal 
ciliated or branched • lateral and ventral nude. Dorsal setae 2.12.10.4.10.8.4: *= 
50, to 45,*. (Walch gives 13.4.6.8.11.8.4, but figures them as 2.12.4.10.4.10.8.4.) 
Ventrally with a pair of branched setae on maxillae, one on each coxa, a pair 
between coxae I and between coxae III and thereafter ca. 46 setae to 40/a long. 
Legs 7 -segmented, I 234/a long, II 208/x, III 260/i ; tarsi I and II with usual dorsal 
sensory rod • no long nude seta on tarsi III. 

The Standard Data derived from 6 of the above specimens are i 



AW 


Mean 
58-8 


Standard Theoretical 
Deviation Range 

No variation recorded 


Observed 

Range 


Coetf . of 
Variation 


PW 


68-6±0-63 


l-53±0-44 


64-0-73-2 


67-2-70*0 


2-2 


SB 


28-9±0-47 


1- 14^0-33 


25- 5-32 >3 


28- 0-30 -8 


4-0 


ASB 


27-l±0-59 


1-44 :£0 *42 


22-8-31-4 


25-2-28-0 


5-3 


PSB 
A-P 


11-2 

38-3±0-59 

23-3±0-59 


No variation recorded 
l-44:t0-42 34-0-42.6 
l-44±0-42 19-0-27-6 


36-4-39-2 
22-4-25*2 


3-8 
6-2 


AM 


51-5ihl-43 


3*19^1-01 


41- 9*41-1 


47-6-56-0 


6-2 


AL 


44-8±0-88 


l-98±0-63 


38-9-50-7 


42*0-47-6 


4-4 


PL 


54-9±0-69 


l-53±0-48 


50*3-59-5 


53-2-56-0 


2*8 


Sens. 


61-6±1-14 


2-29±0-81 


54 '8-68-4 


58-8-64-4 


3-7 



Remarks. This species in the trifurcate palpal claw, the setation of the 
palpi, and the ciliated galeal setae belongs to the "akamushi" group of Whar- 
ton, referred here to the subgenus or genus Leptotrombidiu-m Nagayo. In the 
broadly rounded posterolateral corners of the scutum it is closely related to 
pallida Nagayo from Japan, but distinctly differs in the structure of the scutal 
and dorsal setae. 

Trombicula (Leptotrombidium) pallida (Nagayo et al. 1919). 

Trombicula pallida Nagayo et al. 1919, Verhdl. d. jap. pathol. Gesellsch., 
Tokyo, 9, 107; idem, 1921, Amer. J. Hyg., 1, (5-6), 569; Womersley and 
Heaslip 1943, Trans. Roy. Soc. SL Aust., 67, (1), 75. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 51 

Plate 3, %. K-P. 

In addition to the material reported upon in 1943 (Womersley and Heas- 
lip) over 60 specimens remounted singly from slides of mixed species given to 
Lt-Col. C. B. Philip by Dr. T. Tamiya have now been studied. The material 
was collected by Nagayo and his colleagues in 1919 and 1920 from Yamagata 
Prefecture, Japan, the localities, as translated for me by Mr. N. J*. Tindale 
being Arato, Okiage, and Nukanome. 

On these slides which contained specimens of pallida, this species was indi- 
cated by the character for *'dai" (;fc), meaning "big" or "great," and pre- 
sumably referring to the big (or thick) dorsal setae. 

In addition to the above material, 13 other specimens, 5 from a vole 
Microtus montebelli from Agamo River, Niigata Prefecture, and 8 from Apo- 
denws speciosus from Yacki, Yamagata Prel, all collected by Lt-Col. C. B. 
Philip in 1945, have been examined. No material referable to this species has 
so far been seen from anywhere outside of Japan. 

Tronibicida pallida is here placed in the subgenus Leptotrombidiwn, on 
the characters of the adult. In the larval stage it differs from its nearest allies, 
keiikenscJirijveri, intermedia, etc., as given in the key, and particularly in the 
number and character of the dorsal setae. 

A re-description of the larvae (mainly after Nagayo), together with the 
Standard Data, derived from 31 specimens, is as follows: 

Larvae. Shape oval. Colour in life orange red. Length (unengorged) to 
264/a, width to 172/i. Scutum rectangular with anterior margin sinuous and 
slightly convex medially; posterior margin slightly concave medially; PL situ- 
ated about midway between AL and extreme posterior margin and not at the 
postero-lateral corners, which are well rounded; sensillae filamentous, with 
minute barbs ou proximal third, and strong numerous ciliations on distal two- 
thirds; sensillae bases slightly behind line of PL. Scutal setae stout, blunt, 
with strong denticulations. Eyes 2 + 2, close to lateral margins of scutum, 
posterior the smaller. Chelicerae with only the apical tricuspid cap, Galeal 
setae strongly ciliated. Palpi stout somewhat angular basally; tibial elaw 
trifurcate; all setae on femur, genu and tibia nude, except the dorsal tibial. 
Dorsal -setae stout, strongly denticulate, blunt and shorter than in palpalis, 
keiihenschrijveri, etc., to 43/a long, ca. 70 in number, and arranged ca, 
2.14(12).12(14).12.10.8.6.4.2. Ventrally the setae are similar to the dorsal, 
a pair (of longer branched ones) on maxillae, one on each coxa, a pair between 
coxae I and between coxae III, and thereafter approx. 50 in number. Legs: 
I 190//, long, II 160ft, III 200/i ; tarsi I and II with dorsal sensory rod, III with- 
out any Ion? nude seta. 



52 Records of the S.A. Museum 

The Standard Data for 31 specimens are: 







Standard 


Theoretical 


Observed 


Coeff, of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


70-5±0*30 


1*70*0-21 


65-4-75-6 


66-0-75-0 


2-4 


PW 


76-75*0-57 


3-20*0-41 


67-15-86-35 


72-0-85-0 


4*2 


SB 


35-6*0-33 


1.83*0-23 


30* 1-44 »1 


33-0-39-0 


5-1 


ASB 


26-7*0-16 


0*89*0-11 


24-0-29-4 


24-0-27-0 


3-3 


PSB 


14-8*0-10 


0-59*0-07 


13*0-16»6 


13- 0-15- 


4-0 


SD 


41-5*0 -18 


1-01*003 


38-5-44*5 


39-0-42-0 


2-4 


A-P 


19-0*0-22 


1-26*0-16 


15- 0-23-0 


18-0-21-0 


6-6 


AM 


49-8*0-36 


1-86*0-26 


44-2-55-4 


48-0-54-0 


3-8 


AL 


42-15*0-31 


1*74*0-22 


36-9-47*4 


40*0-45*0 


4*1 


PL 


53.3*0*42 


2-32*0-30 


40-3-60-3 


50-0-57*0 


4-4 


Sens. 


71-5*0-76 


2-41±0*54 


64-3-78-7 


70-0-75-0 


3-4 



Trombictjla (Leptotrombidium) intermedia (Nagayo et al~ 1920). 

Trombicida intermedia Nagayo et ai. 1920. Verhdl. d. jap. pathol. Gesellscht., 
Tokyo, 10, 471; idem Amer. J. Hyg., 1, (5-6), 569, 1921; Womersley and 
Heaslip, Trans. Roy. Soc. S. Aust., 67, (1), 90 1943. 

Plate 3, fig. Q. 

Re-description of Larvae (after Nagayo et ai. 1920). Shape oval. Colour 
in life orange red. Length to 264/t, width to 172^. Scutum rectangular, with 
posterior corners well rounded, and PL about midway between anterior and 
posterior borders. SensiUae about in line or slightly behind line of PL, proximal 
third with minute barbs, distal two-thirds strongly ciliated. Normal scutal 
setae stout, with strong denticles, but not so heavy as in pallida. Eyes 2 + 2, 
the posterior the smaller. Chelicerae with only the apieal tricuspid cap. Galeal 
setae ciliated. Palpi stout, tibial claw trifurcate* setae on palpal femur, 
genu and tibia all nude, except dorsal on tibia. Dorsal setae very thick, blunt, 
and strongly denticulate, 38-40 in number, arranged 2.10.8-8.6.4.2(0), to 57j* 
long. Ventral setae, one on each coxa, a pair between coxae I and between 
coxae III and thereafter ca. 34 similar to dorsal. Legs as in pallida; tarsi I and 
II probably with the usual dorsal rod or spur, III without any long nude seta. 

The Standard Data (after Nagayo et al. 1920, and Womersley and Heas- 
lip 1939 )are: AW 53-0, PW 60-0, SB 22-5, ASB 23*5, PSB 23-0, SD 46-5, 
A-P 18-0, AM 53-0, AL 41-0, PL 57-0, Sens. 650. 

Remarks. Amongst the Japanese material which I have been able to study, 
it has not been possible to identify any specimens with certainty, as intermedia 
Nagayo et ah 

Kuwato, Berge and Philip 1950, however, record specimens from " indigen- 
ous small animal hosts'' from North slopes of Mount Fuji, Fujino Susono 



Womersley— Asiatic-Pacific Scrub Typhus Mites 53 

(Gatemba), Oct. 1948 (Major T. L. Berge), along with T. abamuski, palpalis* 
pallida, scutellaris, and a new species T. fuji, as well as a Gahrliepia sp. 

These authors give the following Standard Data for two specimens of 
intermedia, the values of which are higher than those quoted above, as inter- 
polated by Womersley and Heaslip 1943, from the published figures of Nagayo 
etal. 



AW 


PW 


SB ASB PSB 


SD 


A-P 


AM 


AL 


PL 


Sens. 


71-5 


78-5 


34-0 — 12-5 


— 


24-5 


50-5 


41-5 


55-0 


62-0 


69-0 


79-0 


33-5 — 12-5 





23-5 


49- V 


38-? 


54-5 


59- T 



These higher values are probably more indicative of the measurements to be 
expected, 

Tbocmbicula (?Leptotrombidium) fuji Kuwato Berge and Philip 1950. 

Trombiada (Leptotronibidmm) fuji Kuwato, Berge and Philip 1950, J. Para- 
sitology, 36, (1), 80). 

Plate 4, fig. A-C. 

This interesting species, which on larval characters was placed by the 
authors in the subgenus Leptotrombidmm, was described from larvae from 
Apodemus speciosus speciosus Temminck and Schlegel, Microtus montebelli 
montebelli Milne-Edwards, and Urotrickus talpoides hondonis Thomas, from 
the Pujino Maneuver Area near the base of Mt. Fuji, south-west of Tokyo, 
Japan, in Oct. 1948. 

In June, 1949, further specimens were obtained from Apodemus speciosus 
and A. geisha Thomas, in the same area. 

Larvae, Size (fully engorged) 515/x wide by 670/x long. Colour in life 
pale reddish yellow. Scutum small, roughly rectangular, sparsely punctate, 
with SB well behind line of PL, and the posterior corners well rounded with 
PL not at the corners but much nearer to AL. Sensillae filamentous, strongly 
ciliated on distal two-thirds and minutely barbed basally. PL more than twice 
as long as AL and strongly ciliated. Eyes 2 -|- 2, posterior the smaller. Palpal 
tibial claw trifurcate; setae on femur and genu nude, doi*sal ciliated. Dorsal 
setae 30-32 in number, from 50/* long anteriorly to SOp posteriorly, barbed, and 
arranged ca, 2.8.6.6.2.6.2. with slight valuation. Ventrally, all coxae 1-setose, a 
pair of branched setae on maxillae, a pair between coxae I and between 
coxae III, and thereafter ca. 24 small prcanal and 18 longer postanal setae. 
Legs all 7-segmented ; no long nude seta on tarsi III. 

The Standard Data for the type and 9 paratypes as given by the authors 
are: 



54 Records of the S.A. Museum 





AW 


PW 


SB 


A-P SB 


PSB PL-SB AM 


AL 


PL 


Sens. 


Type 


50-5 


52 


24 


14 30-5 


11 10-5 29-5 


28*5 


51 


40-5 


Extremes 


44-50 


48-55 


21-24 


13-14 30-37 


10-15 10*11 26-30 


23-27 


45-48 


33-40 


Mean 


49 


51-38 


23 


13'4 33-98 


12-2 10-2 27*5 


25-5 


46-8 


35-9 



Remarks. When the adults and/or nymphs of this species become known 
the assignment of the larvae to Leytotroyribidium will probably be confirmed. 

The figures given in this paper are re-drawn from the author's published 
figures, that of the scutum being at a magnification of 500 times for comparison 
with other species. 

Trombioula ( ? LBPTOTROMBioitJM) lancbolata sp. n. Lawrence in MS. 

Plate 4, fig. D-I. 

Description of Larvae. Shape oval. Length (engorged) to 425/x, width to 
357/a. Scutum trapezoidal, with posterior lateral Corners well rounded, and PL 
about midway between AL and posterior margin ; surface finety punctate ; sen- 
sillae filamentous, finely barbed on basal half and ciliated on distal half, 
bases fairly wide apart and well behind line of PL. Eyes 2 + 2, closely adja- 
cent to scutum and posterior the smaller, Chelicerae not serrate, with only the 
apical tricuspid cap. Galeal setae strongly branched. Palpi stout, tibial claw 
trifurcate; setae on palpal femur and genu, and ventrally and laterally on 
tibia nude, dorsally on tibia branched. Dorsal setae thick, with strong denticles, 
28 in number, arranged 2.8.6.6.4.2, to 58/i long. Ventrally, with a pair of 
ciliated or branched setae on maxillae, a slender ciliated seta on each coxa, a 
pair between coxae I and between coxae III, and thereafter ca. 13-14 fine 
ciliated setae, followed by 6.4.2. setae similar to but smaller than the dorsal 
setae. Legs all 7-segmented, I 260^ long, II 227/*, III 273/*; tarsi I and II 
with the usual dorsal sensory rod, III without any long nude seta. 

The Standard Data derived from nine specimens are as follows: 

Standard Theoretical 

Mean Deviation Range 

AW 58-5±0-98 2-95±0-69 49-7-57-3 

PW 69-4+0-78 2-33^0-55 62-4-76-4 

SB 31-l±0-53 1-61+0-38 26-3-35-8 

AftB 28 • No variation recorded 

PSB 11 • 2 No variation recorded 

SD 39-2 No variation recorded 

A-P 20-85+0-49 1-47+0-35 16-4-2(5-3 

AM 40-8±0-83 2-20+0-59 34-2-47-4 

AL 50-4+0-93 2-80±0-66 42-0-58-8 

PL 560+0-47 1-40+.0*47 51-8-60-2 

Sene. 68-3+1 -12 2-50+0-79 60-8-75-8 



Observed 


Coeff . of 


Range 


Variation 


53-2-61-6 


5-0 


67-2-72*8 


3-4 


28*0-33-6 


5-1 


19-6-22-4 


7-0 


39 -2-^4 • 8 


5-4 


47-6-56-0 


5'6 


53*2-58-8 


2 '5 


64-4-70-0 


3-7 



WOMERSLEY— ASIATIC* PACIFIC SCRUB TYPHUS MlTES 55 

Loc. and Hosts. The above description is drawn up from nine specimens 
from Nesokia bengalemis and Rattus norvegims from Insein, Paungde and 
Prome in Southern Burma, collected by T. H. Lawrence 1945. 

TheBe specimens were amongst the material deposited in the British 
Museum (N.H.) by Lawrence and marked in his list as "laneeo" but no 
description appears to have been made by him. 

Remarks. In the setation of the palpi, galeal setae, the shape of the 
acatiun and the structure of the dorsal setae, this species is very close to- 
intermedia Nagayo et al. from Japan. It also agrees with that species in the 
Standard Data, but differs in the number of dorsal and ventral setae, and 
from Nagayo 's figures, in that the ventral setae are not all alike and similar to 
dorsal. In lanceolata only the 3 posterior rows of setae conform to the dorsal 
structure. 

Tbombioitla (Leptotbombidium) parapalpalis sp. n. 

Plate 10, fig. A-E. 

Description of Larvae. Shape oval. Length (unengorged) 286^, width 
182/a. Scutum wider than long, finely punctate; anterior margin lightly 
concave ; posterior margin fairly deep behind line of PL, laterally and medially 
slightly concave; sensillae bases fairly wide apart and behind line of PL. 
Eyes 2 + 2, posterior the smaller, on ocular shields. Chelicerae with only the 
apical tricuspid cap. Galea! setae branched or ciliated. Palpi stout, tibial 
claw trif urcate ; seta on palpal femur and genu nude, on tibia both dorsal and 
ventral branched or ciliated. Dorsal setae 52 in number, arranged ca 
2.13.12.12.8.4,2, to 56/a long, and tapering with fairly long ciliations. Vent- 
rally with a pair of branched setae on maxillae, one on each coxae, a pair 
between coxae I and between coxae III and thereafter ca. 40 setae to 36/*. Legs ; 
I 285/*, II 260/1, III 312/a; tarsi I and II with dorsal sensory rod, III without 
any long nude seta. 

The Standard Data for the type and 5 paratypes are: 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


# Deviation 


Bange 


Range 


Variation 


AW 


76-1+0-86 


2.11±0-G1 


69 -8-82 -4 


73-8-78-4 


2-7 


PW 


86-8±l*25 


3-07±0-88 


77*6-96-0 


840-92-4 


3-5 


SB 


35*5±0-59 


l-45:h0-42 


SI -2-39-8 


33-6-36-4 


4-0 


ABB 


30-8 


No variation recorded 






P8B 


14-0 


No variation recorded 






BD 


44-8 


No variation recorded 






A-P 


25-2 


No variation recorded 






AM 


63-8±0-56 


l-25±0<40 


60-0-67-6 


61-6-64-4 


2-0 


AL 


48*5±0-5I 


l-44±0'42 


44-2-52-8 


47-6-50-4 


3-0 


PL 


63-8±0-56 


1*25+0-40 


60-0-67-6 


61-6-64-4 


2 


Sens. 


85-4 (Mean 


of 84-0 and 86-8). 


Only two determii 


lations. 





56 Records of the S.A. Museum 

Loc. and Hosts. The type and 5 paratopes from a rat, from Kanzalwan, 
Kashmir, India, 1949 (S- L. Kalra), and 3 other specimens from rat from 
Gurais, Kashmir, 1949 (S.L.K.). 

Remarks. Somewhat closely related to pal.pali$, but much more so to macar 
cus sp. n. Lawrence, in the palpal setation, the shape of the scutum and the 
number and arrangement of tlie dorsal setae. From mwacus, however, it can 
be separated as in the key on the larger scute, and more particularly on the 
different structure of the dorsal setae. These are tapering with long ciliations, 
whereas in macacus they are almost uniformly thick to the apex -with strong 
setules ratber than ciliations. 

Trombicula ( 1 Leptotbombidium) macacus sp. n. 

TroMbicula n. sp. "F." Lawrence in MS. In Audy 1947, "Scrub Typhus 
Investigations in S.E. Asia, Pt. III. Appendix 7.' ! AMD. 7. War Office, 
London. 

Plate 5, fig. A-F. 

Description of Larvae. Shape oval. Colour in life unknown. Length 
260/*, width 182j*. Scutum trapezoidal with the anterior margin lightly sinu- 
ous, PL at the postero-lateral angles, posterior margin not very deep behind 
line of PL and straight medially, sensillae in line with or slightly anterior of 
line of PL, sensillae filamentous and ciliated distally. Chelicerae with only 
the apical tricuspid cap. Galeal setae feathered or ciliated. Palpi stout, tibial 
claw trif ureate ; setae on palpal femur and genu nude, on tibia both dorsal and 
ventral branched, lateral nude. Eyes 2 + 2, posterior the smaller. Dorsal 
setae moderately stout and blunt with strong denticles, to 40/* long, and 
arranged 2.10.8.10.8,6.4,2. Ventrally all coxae unisetose, a pair between coxae 1 
and between coxae III, and thereafter ca. 50, which, except for the 2 or 3 
posterior rows, are much more slender and tapering than the dorsal setae. 
Legs: I 227 j* long, II 200ft, Til 247/*; tarsi I and II with the usual dorsal sen- 
sory rod {"spur" of Brennan and "Wharton) ; III without any long nude Beta 
on tarsi or metatarsi. 

Standard Data are: AW 64-4, PW 75-6, SB 30-8, ASB 280, PSB 11-2, 
SD 39-2, A-P 26-2, AM 47-6, AL 40-0, PL 50-4, Sens. 61-6. 

Loc. and Hosts. A single specimen from Macacus aMamensis, Imphal, 8 
December, 1945 (T. J. Lawrence). 

Remarks. In having the ventral as well as the dorsal seta of the palpal 
tibia branched, this species is closely related to palpaUs Nagayo. It differs in 
the structure of the dorsal setae as well as in their number and arrangement, 



Womersley— Asiatic-Pacific Scrub Typhus Mites 57 

and also in that, except for the two or three posterior rows, the ventral setae 
are very much more slender and finer than the dorsal setae, which is not the 
case in paipalis. 

Trombicula ( ? Leptotrqmbidjum) puta sp. n. 

Plate 10, fig. F-J. 

Description of Larva. Length (unengorged) 260^, width 195/*. Shape 
broadly oval. Scutum more or less rectangular, wider than long, with pos- 
terior margin fairly deep behind line of PL ; SB in line with or very slightly 
behind line of PL ; AM well back from anterior margin ; AM and AL tapering 
and normally ciliated, PL stout with strong denticles. Eyes 2 + 2, posterior 
the smaller. Chelicerae with only the apical tricuspid cap, Galeal setae 
branched or ciliated. Palpi stout, tibial claw trifurcate; seta on palpal femur 
and genu nude, on tibia only the ventral branched. Dorsal setae stout, blunt, 
with strong edges furnished with strong denticles, 50 in number to 36-40/* 
long, and arranged 4.10.8.2.8.8.6.2.2. i.e., two humeral setae on each side. 
Ventrally with a pair of branched setae on maxillae, one on each coxa, a pair 
between coxae 1 and between coxae III, and thereafter 6.6.4.6.4.2 ; all ventral 
setae fine and tapering with ciliations, except the posterior 3 rows which are 
similar to the dorsal setae, to 36/x long. Legs ■ I 227^ long, II 208/i, III 240>a; 
tarsi I and II with usual dorsal sensory rod. III without any long nude seta. 

Standard Data for the unique type are: AW 75*6, PW 81-2, SB 36*4, 
ASB 28-0, PSB 15-4, SD 43-4, A-P 25*2, AM 42-0, AL 36-4, PL 420, 
Sens. — . 

Loc. and Host. A single specimen from a rat from Kanzalwan, Kashmir, 
India, 12 Oct, 1948 (S. L. Kalra). 

Remarks. This species probably falls into the subgenus Leptotrombidium 
and come9 near to macacus sp. n. from which it differs in the two humeral 
setae on each side, in the form of the dorsal setae, and other details as in the 
key. The dorsal setae are also very similar to those found in T. fordi sp. n. 
Lawrence, and T. traubi sp. n. 

Trombicula ( ? Leptotrombtdium) nvx, sp. n. 

Plate 5, fig. G-L. 

Description of Larvae. Size large; length (engorged) 890/i, width 780/*. 
Shape almost round. Scutum large, rectangular, with surface irregularly 
shagreened; anterior margin concave, posterior margin convex but lightly con- 
cave medially; AM missing but its base well behind AL; PL at the postero- 



58 Records of the S.A. Museum 

lateral corners; sensillae filamentous, ciliated on distal two-thirds, bases only 
very slightly behind line of PL. Eyes 2 + 2, indistinct and away from 
scutal margins. Chelicerae non-serrate, with only the apical tricuspid cap. 
Galea! setae strongly ciliated or branched. Palpi stout, femur anguiate, tibial 
claw trif urcate » all setae on palpal femur, genu and tibia, except dorsal tibial, 
nude. Dorsal setae numerous, relatively short, from 40/* anteriorly to 50/* 
posteriorly, approximately 86 in number, and arranged in rows of 2.14.14. plus. 
Ventrally with a pair of ciliated maxillary setae, one on each coxa, a pair 
between coxae I and between coxae III and thereafter ca. 56; from 30/i long 
anteriorly to 58/* posteriorly, and more slender and tapering than the dorsal 
setae, Anal orifice not visible. Legs all 7 -segmented, 1 broken short in type, 
II 325/* long, III 385/*; tarsi II with usual dorsal sensory rod; III without 
any long nude seta on tarsi. 

The Standard Data of the unique type are: AW 99-2, PW 121-6, SB 
51-2, ASB 41-6, PSB 16-0, SD 57-6, A-P 35-2, AM — , AL 56-0, PL 70-4 r 
Sens. 96*0. 

Loc. and Host. A single specimen from a brown rat at Ranikhet, Kumaon 
Hills, India, 20 Oct., 1946 (S. L. Kalra), 11 other specimens from a "mouse" 
from Baltal, Kashmir, N. India, along with T. squamifera sp. n. These speci- 
mens have the following Standard Data: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 


95-7±l-01 


3-34±0«71 


85 '7-105 -7 


92-4-100-8 


3-5 , 


PW 


107-4±1-41 


4-G7±0-99 


93-4-121-4 


100-8-112-0 


4-3 


SB 


44<0±0-38 


1'25±0"27 


40*3-47-7 


42-0-44-8 


28 


ASB 


34-l±:3-2 


l-08±0-23 


30-9-37-3 


33-6-36-4 


3-2 


PSB 


19-1±:0«32- 


l-08±0-23 


15-9-22-3 


16-8-19-6 


5-6 


8D 


53-2 


No variation recorded 






A-P 


SO -8 


No variation recorded 






AM 


62-4+0-57 


l-79:fc0-40 


57-0-67-8 


58-8-64-4 


2-9 


AL 


48'2±0*52 


3-01±l-73 


43-0-53*4 


47-6-53-2 


3-6 


PL 


67 -53:0 -42 


2*45±0-58 


60-2-74-8 


64*4r-70-0 


3-6 


Sens. 


81-2:tl-61 


10-41+3-23 


71-5-90-9 


78-4-«4-0 


4-0 



Remarks. In the palpal claw and setae, and the galeal setae this species 
comes into the genus Leptatrombidium (akamtishi-deUensis group of Trombi- 
cula s.l.), but differs in size, dorsal setae etc., as in the key. 

Trombicula (f Leptotoombiditjm) sotjtellabis (Nagayo et al> 1920). 

Trombicula, scutellaris Nagayo et al. 1920. Verhdl. d. jap. pathol. Gesellsch., 
Tokyo, 10, 471 j idem Amer. J. Hyg., 1, (5-6), 569, 1921 j Womersley and 
Heaslip, Trans. Roy. Soc S. Aust., 67, (1), 88, 1943, Kuwato, Berge and 
Philip 1950, J. Parasitol., 36, (1), 82. 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 59 

Plate 6, fig. A-E. 

Again no fresh material of this species has been reported until recently, 
•when Kuwato, Berge and Philip 1950, recorded it from near the base of 
Mt. Fuji, near Tokyo, Japan. Amongst the slides of the original Nagayo 
material brought back by Lt.-Col. C. B. Philip, however, were a number of 
specimens from Okiage, Arato and Nukanome, all in Tamagata Prefecture, 
and dated Sept. 1919 and Sept, 1920. 

The above specimens are in full agreement with Nagayo 's description and 
were indicated on the slides by the ideograph ( $j), or "shin," translated by 
Mr. % B. Tindale as meaning "new" or "fresh," thus indicating a new 
species. 

Re-description of Larvae (mainly after Nagayo). Shape oval r Colour in 
life orange red. Length (unengorged) 264/1, width to 172/i. Scutum roughly 
rectangular, with PL at the postero-lateral corners ; sensillae filamentous, basal 
third with sparse minute barbs, distal two-thirds strongly ciliated, bases in line 
with PL. Scutal setae slender, tapering, with dilations. Eyes 2+ 2, close to 
the postero-lateral corners of scute. Chelicerae with only the apical tricuspid 
«ap. Galeal setae strongly feathered. Palpi stout, tibial claw trifurcate; ail 
setae on palpal femur, genu and tibia (except dorsal) nude. Dorsal setae 
slender, to 51-69/* long, with stout setules, to 50 in number and arranged 
2.10.10(12).10(8).8.6.4. Ventral setae behind coxae in, ca. 24 in number, 
shorter than but similar to dorsal setae. Tarsi of leg III without any long 
nude seta. 

The Standard Data derived from 23 specimens from Arato, Okiage and 
Nakanome, Yamagata Prefecture, Japan, Sept, 1919 and Sept. 1920, are as 
follows : 







Standard 


Theoretical 


Observed 


Ooeflr. of 




Mean 


Deviation 


Range 


BaHge 


Variation 


AW 


72- 4+0 -42 


2-04+0-30 


66-3-78-5 


70-0-75*0 


2-8 


PW 


82-2±0*72 


3*46+0-51 


71- 8-92 -6 


75-0-88-0 


4-2 


SB 


32-2±0-41 


l-96±0-29 


26-3-38*1 


30-0-36*0 


e-i 


A8B 


29-l:±0-38 


1 -82+0-27 


23-6-34-6 


24.0-32*0 


6-2 


PSB 


15.1+0-08 


0-41 ±0-06 


13-9-16-2 


15-0-17-0 


2-7 


8D 


44-0+0-40 


1-04+.O-28 


38-3-49-7 


39-0-47-0 


4-5 


A-P 


28 * 3+0- 40 


1-04+0-29 


22*5-34-1 


24-0-30-0 


6-8 


AM 


59-8+0*36 


1-55 +.0-26 


55-2-64-4 


57-0-63-0 


2-6 


AL 


50-6+0-78 


3-60+0-54 


39-8-61-4. 


42-0-57-0 


7-1 


PL 


64-3+0-52 


2-44+0-37 


57-0-71-6 


60-0-69-0 


3-8 


Sens. 


77-75+0-67 


1-91+0-48 


72-0-83-5 


75-0-80-0 


2-3 



60 



Records of the S.A. Museum 



Trombicula (? Leptotrombidium) vuxosa sp. n. 

Plate 7, fig. A-E. 

Description of Larvae. Shape oval. Colour in life orange. Length (par- 
tially engorged) to 325/i, width to 208/i. Scutum roughly rectangular, 
with AL and PL at the anterior and posterior lateral corners, anterior margin 
lightly concave, posterior margin not very deep behind PL with the median 
part straight, SB in line with or slightly behind line of PL ■ sensillae missing 
in all specimens but probably filamentous. Eyes 2 4" 2, on ocular shields, 
about equal. Chelicerae with only the apical tricuspid cap. Gale&l setae 
feathered or branched. Palpi stout,- tibial claw bifurcate; setae on femur, 
genu and tibia all nude except the dorsal on tibia. Dorsal setae to 58/a long, 
very numerous, long, slender, tapering and ciliated, more than 100 in number 
and arranged 2, then ca, 10 rows of ea. 14 setae. Ventrally a pair of ciliated 
setae on maxillae, each coxae unisetose, a pair of setae between coxae I and 
between coxae III, and thereafter ca. 100 setae. Legs all 7-segmented ; I 312/* 
long, II 260/i, III 325fi ; tarsi I and II with usual dorsal sensory rod, III with- 
out any long nude seta. 





Mean. 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Bange 


Coeff . of 
Variation 


AW 


88* 5+1-12 


2-50+0-79 


81-0-96-0 


86-8-92-4 


2-8 


PW 


103-0±l-63 


3-65±l-15 


92-1-113-9 


98-0-106-4 


3-5 


SB 


41-2+0-56 


1-25+0-40 


37-5-44-9 


39-2-42-0 


3-0 


ABB 


35-8+0-56 


1-25+0-40 


32-1-39-5 


33-6-36-4 


3-5 


P8B 


13-4+0-56 


1-25±0-40 


9-7-17-1 


11-2-14-0 


9-4 


sn 


49*3+1*12 


2-50+0-79 


41-8-56-8 


44-8-50-4 


5-0 


A-P 


34*7+0-68 


1-53+0-48 


30O-39-3 


33-6-36-4 


4-4 


AM 


75-6+0-88 


1-98+0-63 


72-7-78-5 


72-8-78-4 


2-6 


AL 


62-2+0-56 


1-25+0-40 


58-4-66-0 


61-6-64-4 


2-0 


PL 


72-0+1*05 


2-34+0-74 


63-0-81-0 


70-0-75-6 


3-2 


Sena. 


No determination 


. Sensillae lost* 









Loc. and Most. From a rat, Ranikhet, Kumaon Hills, India, 20 Oct. 1946, 
(S- L. Kalra). 



TifiOMBIOUI/A ( ? LlEPTOTROMBIDIUM ) BHtMTALENSIS $p. n. 

Plate 7, %. F-I. 

Description of Larvae. Shape oval. Size_ moderate, length (engorged) 
665/a, width 560/a. Scutum rectangular, finely punctate; anterior margin sinu- 
ously concave; AM placed well behind line of AL; posterior margin shallow 
and strongly sinuous; PL setae the longest; sensillae missing but bases about 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



Che Lioerae 



in line with PL. Eyes 2 + 2, but very indistinct, 
setae branched. Palpal claw bifid; setae on palpal feniur 
tibia dorsal and ventral branched r lateral nude. Dors*! 
long and arranged 2.10.8.6.4,2. Ventrally with paired 
on each coxa, a pair between coxae I and between coxae 
ing and only represented by the seta bases), and thereaft< 
Legs: I 240/a long, II 240/c, III 270/*; tarsi I and II 
III without any long nude seta. 

Standard Data: AW 70-4, PW 83-2, SB 35-2, 
SD 42-6, A-P 28-8, AM 32-0, AL 41-6, PL 48-0, Sens. 

Loc. and Host. The unique type from a shrew, 
India, 10 Oct., 1946 (& L. Kalra). 

BemarJcs. Can be separated as in the key to species 



missing. Galeal 

and genu nude; on 

setae ciliated, to 54/* 

setae on maxillae, one 

III (all these are miss- 

ea. 20, to 35/i long. 

wjith dorsal sensory rod, 

kSB 30-0, PSB 12-6, 

Bhimtal, Kumaon Hills, 



Trombioula (Leptotrqmbidium) longiseta sp. n 



postcro 



finely 



Trombwulan. sp. "L." Lawrence in MS. 1947, In J. E. 
Investigations in S.E. Asia, Pt. IIL Appendix V* 
London. 

Plate 8, fig. A-E. 

Description of Larvae, Shape broadly oval. Leng 
width 185/*. Scutum rectangular, with PL at the 
very long, and posterior margin rather shallow behind 
lightly concave, almost straight medially; surface 
very much behind line of AL ; sensillae filamentous, an<k 
the bases distinctly behind line of PL. Eyes 2 + 2, 
Chelieerae non-serrate, with only the apical tricuspid 
ciliated. Palpal claw bifurcate; setae on palpal femu: 
tibia, dorsal ciliated and lateral and ventral nude, 
tapering, to 70/i, and arranged 2.8.6.6.4.2. Ventrally 
ciliated maxillary setae, one on each coxa, a pair betweei 
coxae III, and thereafter ca. 28 in number, arranged ea 
Legs all 7-segmented : I 227/* long, II 227/*, III 241/£ 5 
sensory rod, III without any long nude seta. 

The Standard Data derived from 14 specimens, 1 
Burma, 1945 (G. M. Kohls, No. 740), 3 larvae from 
(T. J. Lawrence and K L. Cockings), and 10 larval 
reared nymphs by CocMngs (larvae from Imphal, 1945) 



tarki 



61 



Audy, "Scrub Typhus 
AMD. 7. War Office, 



Doi sal 



frith 



h (unengorged) 285/t, 

-lateral corners, and 

line of PL and only 

punctate; AM not 

ciliated distally with 

posterior the smaller. 

cap. Galeal setae 

and genu nude; on 

setae 28, long and 

the usual pair of 

coxae I and between 

10.8.6.4, to 50/t long* 

I and II with dorsal 



arva from Myitkyina, 
Imphal, India, 1945 
pelts recovered from 
are as follows: 



62 Records of the S.A. Museum 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


59*9±0-52 


1«96±0*37 


54-0-65-8 


57-6-64-0 


3-3 


PW 


71-5^:0*61 


2-30±0-43 


64-6-78-4 


67-2-76-8 


3-2 


SB 


26*75±0-42 


l-59±0-30 


22 -0-31. 5 


25-6-28-8 


5-9 


ASB 


32-0 


No variation recorded. 






PSB 


9-6 


No variation recorded. 






SD 


41-6 


No variation recorded. 






A-P 


25-6 


No variation recorded. 






AM 


55-5:±0'98 


3-68±0*68 


44-5-66-5 


51*2-«4-0 


6-6 


AX 


05-l:tO'6O 


2-24:±0'42 


48-4-61-8 


51-2-57-6 


4-0 


PL 


108-3±2-14 


8- 00+1* 51 


84-3-132-3 


96-0-128-0 


7-4 


Sens, 


69-8±0-41 


l-23±0-29 


66-1-73-5 


67-6-70-4 


4-7 



Remarks. This species might be regarded as only a variant of delien&is 
-with very long PL setae. It differs, however, in having the palpal claw only 
bifurcate. 

It is herewith placed in Leptotromhidium on the precoxal plates in the 
reared nymphs, as described elsewhere in this paper. 

TROMBicukA (Leptoteombidium) deliensis (Walch 1923). 

Trombictda deliensis Walch 1923, Kitasato Archiv. Exper. Med. 5, (3), 63; 

idem, 1923. Tr. Vth. Bien. Congr. Par East. Assoc. Trop. Med., Singapore 

(1924); Womersley and Heaslip 1943; Tr. Roy. Soc. S. Aust, 67, (1), 87; 

Womersley 1944, ibid,, 68, (1), 90. 
Trombicula vanderghinstei Gunther 1940, Proc, Linn. Soc, N. S. .Wales, 65, 

(3-4) 250; Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, (1), 

87. 
Trombicula Walchi Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, 

(1), 83; Womersley 1944; ibid., 68, (1), 89. 

Plate 6, %. F-L. 

This is a common and widely distributed species ranging from Kashmir 
and India, south-eastwardly to the Philippines, and from Ceylon south-east- 
wardly through Burma, Malaya, New Guinea, to the Solomon Islands and 
Northern Queensland. As a known vector in tlie transmission of the rickettsia 
of scrub typhus from rats to man it is of very great importance medically. 

From the only other species definitely known as a vector, T. (L.) akamushi, 
— a species which some recent workers would regard as only a variety of 
deliensis— it differs in the smaller number of dorsal setae, 28 arranged 
2.8.6.6.4.2 with only occasionally one more or less on the second or third dorsal 



Womersley— Asiatic-Pacific Scrub Ty 



>hus Mites 



any other, and I have 



rows. In this arrangement of dorsal setae, deliemis agrees with many 

other species, which can be separated on other morphological characters. In 

tokamu&hi the dorsal setae are more numerous, 32-40 in number, arranged ca. 

2.10.8.6.4.2 to 2.10.8.8.6.4.2, with frequent variations. 

scutal margin i& very shallow behind the line of PL anld consequently SB lies 

distinctly in front of PL; whereas in deliensis SB is not greatly in front of 

PL, sometimes appearing almost in line, and the posterior scutal margin is 

deeper and more rounded. 

This species has probably been collected more than 

been able to compare a considerable number of populations from different 

areas. 

Waleh in his original description and figures gave vlery few measurements : 

The scutum was stated to be 37/i long and 74/x wide; JStj was 4/4 behind line 

of SB ; AL 3fyi.- 7 Sens. 61/x. No measurements for the lengths of AM and PL 

were stated but his figure shows both as longer than AL, 

AL. 

Of the following twenty-four populations studied), 

may be considered as more or less typical deliensis on 
sometimes significant between one another, in relative n 
Data. 

If anything, there seems to be tendency for the 
larger as one goes from west to east, the populations from New Guinea and 
Northern Queensland having somewhat larger Standard Data* The relative 
proportions of the scutal setae, with AL the shortest 
remain fairly consistent. Apart from these normal populations, however, the 
one from Buna, New Guinea, (No. 24) is very significantly different from all 
the others in the extreme length of the scutal setae especially PL, which reach 
a mean length of 85/a. In general the dorsal setae in thiu, as in all populations 
approximate the PL setae in length. This population 
distinct, possibly local, form under the name of deliensis form iimaensis nov. 

(plate 6, fig. I-L). 

At the other extreme, another small collection of 
Abidari and seven from Buna, N.G. (No. 22) differ frbm the normal in the 
very short scutal (and dorsal) setae, with PL being equal to or only slightly 
longer than AM, with a mean of ca. 40/i. Two other specimens (No. 23) from 
Bougainville, (G. W. Wharton) are also of this form, 
mens are sufficiently distinct to warrant a form name 
specimens from Bougainville appear to have the pal 



63 



and PL as longer than 

the first twenty-one 
y showing differences, 
alues of the Standard 

dorsal scutum to be 



three specimens from 



Wh ether these speei- 
is doubtful. The two 
>al claw two-pronged 



instead of three-pronged as is normal for deliensis, al the ugh the Buna- Abidari 



64 Records of the S.A. Museum 

specimens are apparently three-pronged. This character, however, is some- 
times very difficult to be certain about even under high magnification. 

The type specimen of T. walchi Worn, and Heaslip 1943 from Batavia 
(^measured) has the following Standard Data: AW 61-6, PW 75-6, SB 
28*0, ASB 28-0, PSB 14-0, SD 42-0, A-P 28-0, AM 60-4,-AL 56-4, PL 56-0, 
Sens. — , and thus cannot be separated from typical deliensis. 

Gunther's vanderghinstei from Bulolo, New Guinea is also a typical 
deUen^is. 

Number and Docaltties or Population in the following table. 

(1) Ceylon. Seven specimens from Colombo, 1944 (C. D. Badford) and seven from Embill 
pitiya, on Battus Icaadiyanus, 1944 (8. H. Jayewickreme). 

(2) Maldi/ve Is. Pout specimens from Battus r. norvegious, 1045 (C.D.E.). 

(3) Assam. Sixteen specimens from Battus flavipectus yunna/nensis from 21 mile mark, 
Stillweli Road, 6/9/45. (B. Traub). 

(4) Mmipwr. Three specimens from Battus sp. 9/6/45 (B. Traub). 

(5) Burma. Eighteen specimens ex Traub. coll. 1945, 

(f>) Burma. Thirty-sii specimens ex T. J. Lawrence coll. 1945* 

(7) Malaya. Twenty- three specimens from Battus mullen from Kepong, Selangor, F.M.8., 
June, 1945 (J. B. Andy coll.). 

(8) Malaya. Eighteen specimens from ground squirrel, Kepong, Selangox, F.M.S., Sept., 
1950 (J. B. Audy coll.). 

(9) Malaya. Fourteen specimens from Battus bowersi, from Ulu Langat, Selangor, F.M.S., 
June, 1950 (J. B. Audy). 

{10) Malaya. Eight specimens: one from Battus annandalei from. Buhit Langan, F.B., 1949; 
five from a shrew, Ulu Langat, Dec, 1949, and two from JR. mullefi from Ulu Langat, 
Selangor F.M.8. Nov., 1949 (J. E. Audy colL). 

(11) Borneo. Seventy-five specimens from Tragulus javanious from Fort Leju, Tinjar, 
Sarawak, Aug., 1950 (ex J. B. Audy). 

(12) Borneo. Twentv-one specimens from ears of CaUoscimrus hippurus, Stapok F.B., 
Kuching, May, 1950 (ex J. B. Audy). . 

(13) PJwlippines, Five specimens ex Battus sp. Feb., 1945. (colL C. Mohr). 

(34) Philippines. Sixteen specimens ex. Battus sp. from Mercia, Negros. July, 1945 (O. B. 

Philip coll.). 
(15) Bat. Is., N.G. Eight specimens on boots Nov.. 1949 (coll. G. M. Kohls). 
(1(5) Solomon, Is. Twenty-four specimens on boots, from Island North of Bougainville, 

Aug., 1945 (Coll. G. H. McQueen), 

(17) Dutch N.G. Eighty-five specimens from Sanaapore, 1944 (colL C. Mohr). 

(18) New Guinea. Fourteen specimens from ears of rat, Milne Bay, Papua, Aug., 1943 
(coll. S. L. Allman). 

(19) New Guinea. Ten specimens from ears of rat, Kerowae, Nov., 1944 (coll, Consett. 
Davis). 

(20) Australia, Twenty -eight specimens from rats, from Cairns, N. Queensland, 1939 (G. W. 
Heaslip). 

(21) Australia* She specimens from boots, Bramston's Beach, N. Queensland, Sept., 1949 
(coll. I, M. Maekerras). 

(22) New Guinea,. Ten specimens t three from boots, Abidari, July, 1943 (coll. B. N. McCul- 
loch), and seven from boots at Buna, Aug., 1943 (coll. R. N. McCulloch), 

(23) Solomon Is. Two specimens ex Battus praetor from Jaba River, 8ept., 1944 (coll. G. W. 
Wharton) . 

(24) New Guinea. Twelve specimens: eight from rat, Buna, Dec, 1946 (coll. G. M. Kohls), 
and four from rat, Buna, Jan., 1944 (coll. Maj. Hicks). 



WOMERSLEY— ASIATIC- PACIFIC SCRUB TYPHUS MlTES 



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TH'^^OeOOOCOffCOO'^OOlOrHlDCOOO^kO-^OQCtOOCMCM 
rl rH HfONHH t- 01 »-4 OlOOr-irHOl rH rH 






W«0t>Q00)OHNC0^IO5Dt»000iOHNM^ 





WOMERSLEY — ASIATIC-PACIFIC SCRUB TYIHUS MlTES 



67 



Tromdicula (Leptotrombidium) akamushi C Brumpt 1910) 

Trombieula eoarctatum (Berl. 1888), in Kitashima and Miyajima 1918, cited 
Walch 1923, Tr. Vth. Bien, Congr. Par East. Assoc. Trop. Med. (1924). 

Kedmia tanakai Kishida 1909, cited Tanaka et al, 1)30, Zentralblt. Bakt. 
Abt. 1, 116. Orig., 353. 

Tr&mbidium akamushi Brumpt 1910, Precis de ParasitoL 2nd ed,, 506. 

Microtrombidium akamushi, Hirst 1915, J. Econ. Biol., 10, 79. 

Leptotrombidium akamushi, Nagayo et al., 1917, J. Exper. Med., 25, 255. 

Trombieula akamushi, Hirst 1917, Arachnida injurious t) Man (B. Mm N.H.). 
Econ. Ser., No. 6: Nagayo et al, 1921, Amer. J. Hygiene, 1, (5-6), 569; 
Walch 1923, Tr. Vth. Bien. Congr. Par East Assce. Trop. Med. (1924); 
Gater 1932, Parasitology, 24; Brumpt 1936, Precis de Parasitol: Womers- 
ley and Heaslip, 1943. Tr. Roy. Soc. S. Aust., 67, (1), 84; Sig. Thor and 
Willmann 1947, Das Tierreich, Lfg.. 71b, 381,. 

Microtrombidium brumpti Hirst 1915, J. Econ. Biol., 
Ann. Ent. Soc. Amer., 13, 381. 

Trombieula fletcheri Womersley and Heaslip 1943, Tr. 
(1), 86- Womersley 1944, idem 68, (1), 89. 

Trombieula obscure Womersley 1944, Tr. Roy. Soc. S. 



Plate 6, fig. M-O. 

If valid, Kishida ? s names would replace those of Leptotrombidium akamushi 
Nagayo et al, 1917. Kishida first published his paper 
which only 100 copies are understood to have been distributed. However, in 
spite of intensive search in America, Britain and elsewhere, my colleague, 
Dr. C. B. Philip, has been unable to trace anywhere it copy of the original 
paper. In 1946, Kishida republished his paper, but agiiin privately, although 
with excellent figures. It appears then that as Kishida *s publications are not 
scientific papers under the International Rules, his nanes are not valid. 



This species is very close to delienms Walch and 1 



03; syn. Ewing 1925, 
oy. Soc, S. Aust., 67, 
Aust., 68, (1), 86. 



as been considered by 



some workers to be the same, differing only in the larger number of dorsal 
setae which vary from 32 to 40 in number, as against an almost constant 28 
in deliensis. It is, however, not only distinct in this character but is distin- 
guished by the longer A-P, bringing the sensillae base* (SB) much more in 
advance of line of PL, and the posterior margin being straighter, although 
lightly sinuous, and shallower behind PL, 



68 Records of the S.A. Museum 

In the typical form from Japan, as shown in Kislrida's 1946 figure and 
in the works of other Japanese authors, as well as the Japanese material 
studied by the writer, the dorsal setae appear to be somewhat variable from 
32-40 in number. Kishida, 1946, shows them as 2.8.6.8.6.4.= 34, whilst Hirst 
1915 shows them as 2.8.6.10.8.4.2 — 40. The arrangement on the first 3 rows, 
however, appears fairly constant, the variation occurring on the posterior rows. 
Here, however, it is sometimes difficult to place some of the lateral posterior 
setae as belonging to the dorsal or ventral surface. 

In a moderately large population from Bagabag, Luzon, in the Philip- 
pines, collected by C» B. Philip, 1945, the dorsal setae vary from 32 to 39, 
mainly arranged 2.8.8.8.6.4.3 = 38, but varying from 2.8.4.8.6.4=32 to 
2.9.8.8.6.4.2 = 39. 

In 1943, Womersley and Heaslip described fletcheri from Malaya mainly 
on the. different arrangement and number of dorsal setae, viz. 2.10.8.6.4.2.2=34. 
Recently a large population, collected by Dr. J. R. Audy 1945, from various 
hosts in the neighbourhood of Sungei Buloh, P.M.S., has been studied, in which 
the dorsal setae varied from 2.10.8.6.4.2 = 32 to 2.11.8.6.6.4.2=39, but mainly 
were 2.10.8.6.6.4.2 = 38. 

Similarly populations from Owi Island, Dutch New Guinea, as well as parts 
of New Guinea, agree in having the dorsal setae ca. 38 in number and arranged 
generally 2.10.8.8.6.4. This difference from the normal Japanese form perhaps 
justifies the retention of the name fletcheri as a geographical one. 

The Standard Data expressed by the Mean dk 3 times the Standard 
Deviation, for the following populations are given in the succeeding table. 

Number and Localities of Population in the following table. 

(1) Jfrpan, 11 specimens: aome from Dr. Mitamura's original collection and some collected 
at Niigata, Yamagata Prefecture, by Dr. C. B. Philip in 1945. 

(2) Philippines. 25 specimens from Bagabag, Luzon, Aug., 1945 (coll. C. B. P.). 

(3) Philippines. 13 specimens from rats, precise locality t s 1945 (coll. C. Mohr.). 

(4) New Qu.im.ea. 14 specimens from Battua mordax, No. 467. Buna area, 1944 (coll. 
G. M. Kohls.). 

(5^) New Guinea. 11 specimens from fiattus mordax, No* 475. Buna area, 194.4 (coll, 
GJktK.). 

(6) New Guinea. 12 specimens from boots, Dumpu, 1943 (eoll. R. K. MeCulloch). 

(7) New Guinea. 21 specimens from Echytnipera cockerelli from Dobodura area (coll. 
G.M t K.). 

(8) New Guinea. 10 specimens from rats, Buna, 1944 (coll- Maj. Hicks), 

(9) Dutch New Guinea. 14 specimens, 1944 (coll. C.B.P.), 

(10) Malaya. 125 specimens from Sungei Buloh, etc., 1948 (coll. J. R. Andy). 



Womersley— Asiatic- Pacific Scrub Tvphus Mites 



Standard Data of ten populations of Tronibwula (Leptotr) akavnmliHf (Brumpt.) (allowing Means :± 3 

times Standard Deviation) 



69 



Popu- 
lation 



No. of 
Speci- 
mens 



AW 



PW 



SB> 



ASB 



PSB 



(1) 


11 


(35 


25 


(3) 


13 


« 


11 


(5) 


13 


(6) 


12 


(7) 


21 


(8) 


10 


(9) 


14 


(10) 


125 



65 -5 ±5 
57'7±;4' 

55-85:t3 
58-4±4 
63-9±9 
54-8^4-0 

66-75±:8'4 
60«5±4-6 

65*65:±4-5 
68-7±6-3 



4 
6 

65 
7 



74-5±5-l 

66-9rt4.fi 

69-85it3-65 

6'5-2±4.1 

77-S±7-8 

60- 25+4 -65 

81-75±13-2 

68-2±6-9 

75-3±8*l 

81-l±8-3 



29-8±7'2 
27-0 

26-4±4-9 
32*l±5-7 
26<0:fc3-4 
34-l±6«2 
28 -65 ±3-0 
30-9+4-1 
33-4±5-0 



27-0 
21-0 
24-0 
24-0 
27-0 
24*75±3 
26 -5±3 



27-0 
26*ltf±3*05 



13-2r±3-5 

12-0 

13-0 

12 -7+3-0 

12-5±3>4 

12-3±2-65 

13-2±3«5 

14- 15 ±3 -7 
16«65:±l-8 



Popu- 
lation 



No. of 
Speci- 
mens 



SD 



A-P 



AM 



AL 



PL 



Sens. 



(1) 


11 


(2) 


25 


(3) 


13 


(4) 


11 


(S) 


13 


(6) 


12 


(7) 

m 


21 


10 


t»5 


14 


(10) 


125 



40-2zt3-5 
33-0 
37-0 

36-7:t3-0 
39-o±:3.4 
37-1+3 
39-7+3 
38-5+8 
4M5+3 



42-8+3*9 



23-7+3-8 

24-0 

27-9+3*1 
26-45+3-65 

27-0 

24-0 

26-8±3-6 

26-7+7-9 

28-13:4-5 
28 -15+4 -3 



53-6+6-4 
50-15±5-8 
50*2+7.6 
50^7+7-9 
52-45+6-9 
43-6+5-6 
55-3+7-7 
43 -1+10 -6 
54-9+8-1 
52-4+8-1 



42-3+7 
37-8+4 
39-6+3 

39-7+3- 
38-6+4 
39-6+3 
35-0+4- 
40-0+3- 
42-6±8< 



57 


•l±6-8 


52 


■7+4.8 


52-' 


15+5-15 


49-4+7-4 


51 


7+7-5 


45 


•5+5-2 


53 


•7±6-8 


42 


4+12-2 


55 


4+7-7 


53 


8+7-4 



61-2+4-9 
59-3+3-0 

53-55+6-9 
60-0 
60-0 

53-7+2-9 
64-9+4-7 
60-3±6-l 
65-7+7«5 

64-25+11-05 



Trombieula dbscura Womersdey 1944 cannot be 
now regarded as being, the usual New Guinea form o 
dorsal setae arranged 2.8.8.8.6.4.2, and the scutal seta<£ 
normal, but not outside of the theoretical range. It 
type and 3 paratypes, from Milne Bay, N.G. The 
Maj. Hicks from Buna must also be referred to 
Standard Data are given under No. 8 in the above tablel) 



separated from, and is 

akamushi, with the 38 

somewhat longer than 

kvas described from the 

population collected by 

akamushi f. fletcheri. (The 



Trombicula (Leptotrombiditjm) ftjlleri (] Swing 1945) 



(engo 



Trombioida fxdlerl Ewing 1945, Proe. Ent. Soc, Wash 

Plate 8, fig. F-J. 

Redescription of Larvae. Shape oval. Length 
to 500/x. Scutum rectangular, with PL at the posterior 
posterior margin only moderately deep behind PL 
concave medially; PL setae the longest; sensillae filaider i i 
tally, with the bases only slightly behind line of PL, 
the smaller. Chelicerae normal, non-serrate and with d 



intgon, 47, (3), 46. 



bit 



rged) to 710/1, width 

lateral corners, and the 

strongly sinuous and 

and ciliated dis- 

Eyes 2 -f 2, posterior 

nly the apical tricuspid 



70 Records of the S.A. Museum 

cap. Galeal setae ciliated. Palpi claw trif urcate ; setae on palpal femur and 
genu nude; on tibia, dorsal ciliated, lateral and ventral nude. Dorsal setae 28 r 
to 52/a, arranged ca. 2.8(10).6(4).6.4.2. Ventrally with the usual pair of ciliated 
maxillary setae, one on each coxa, a pair between coxae I and between coxae III r 
and thereafter ca. 20 to 32/* long. Legs all T-segmented : I 225/* long, II 227^, 
III 260/*; tarsi I and II with dorsal sensory rod, III without any long nude 
seta. 

The Standard Data derived from 3 larvae and 8 larval pelts from reared 
nymphs (K. L. Cookings), all from Imphal, India, 1945, are as follows: 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Range 


Variation 


AW 


56-6±0-39 


1-29^:0.28 


52-7-60'5 


54-4-57-6 


2-3 


PW 


65-0±0-45 


l-49±0-32 


60-6-69-4 


64-0-67-2 


2-3 


SB 


25-95±0-30 


0.98^:0.21 


23.0-28-9 


25-6-28-8 


3-8 


ASB 


27-0±:0-59 


1-97 ±0-41 


21-1-32-9 


22-4-28-8 


7-3 


PBB 


10-8±:0-38 


l-26:t0-27 


7-0-14.6 


9-6-12-8 


11-6 


SD 


37-75Jz0-55 


1.82±0»39 


32-3-43. 2 


35-2-41-6 


4-8 


A-P 


22.7rt0-51 


l*70:t0-36 


17-6-27-8 


19-2-25-6 


7-5 


AM 


42-25:t0-37 


M2:t0-26 


38«9-45-6 


41-6-44-8 


2-6 


AL 
PL 


35-65:£0-38 


1-12+0-24 


32-3-39-0 


35-0-38-0 


3-1 


62 -6:1:0 -70 


2»34±0»50 


50. 6-69*6 


57*6-65-8 


3-7 


Sens. 


55-0±0-89 


2»19±:0'63 


48-4-61*6 


51-2-57-6 


4-0 



Remarks. In the dorsal setae and the setation of the palpi this species is. 
close to deMemis, but it differs in the sinuation of the posterior seutal margin. 
It is here placed in Leptotrombiddum on the presence of precoma! plates on 
coxae I in the nymphs reared by Mr. Cookings, and described elsewhere in this 
paper. 

A further collection of larvae of this speeies made by Major R, Traub from 
Burma has been examined, which show a significantly larger scutum than those 
quoted above, although they do not differ in any morphological character. 
The Standard Data for 13 of these specimens are as follows: 







Standard 


Theoretical 


Observed 


Ooefi*. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


62 -25^0-81 


2-94±0*57 


56-45-68 05 


58-8-67-2 


4-7 


PW 


72-4:1:0-60 


2-15±0-42 


65*95-78-85 


70-0-75*6 


3-0 


SB 


28-85±:0.36 


l-29ih0-25 


25*0-32-? 


2S-0-30-8 


4-5 


ASB 


28*0 


Wo variation recorded. 






PBB 


11-2 


No variation recorded. 






SD 


39-2 


No variation 


recorded. 






A-P 


23-7±0-39 


l-39±0<27 


19-5-27'9 


22-4-25 2 


5-9 


AM 


46-2±0-40 


1-40:1:0 -28 


42-0-50-4 


44-8-47-6 


3-0 


AL 


43-8zt0.52 


l-79±0-36 


36-4-47.2 


39-2-44-8 


4-3 


PL 


69-35:fc0-54 


1-95^:0.38 


63-5-75-2 


64-4-72-8 


2-8 


Stans. 


to 64-4 











Womersley— Asiatic-Pacific Scrub Typhus Mites 



In correspondence, Dr. Fuller informs me that h< 
co-types of fulleri and compared them with an authem ic 
Walch from Sumatra, and that Ewing's figure erroneously 
sillae bases as anterior of line of PL whereas they 
thereof. Dr. Fuller's specimens from Burma agreed 
co-types of fMeri Ewing and are considered to be thi 
<also examined another lot from Burma collected at the 
agreement with Fuller that they are not deUenm Which 
differing in having SB posterior of PL, there is a. marked 
curvature of the posterior scutal margin. In all the 
have examined from many localities this margin is morj* 
whereas in fulleri it is sinuous and medially concave 
E wing's figure is erroneous. 



has examined E wing's 
specimen of deliensis 
shows the sen- 
are actually posterior 
\n this respect with the 
same species. I have 
same time and am in 
However, besides 
difference in the 
material of delknsis I 
or less an even curve, 
In this respect also 



MYZANTHA. 



TrOMBICUL,A. (LePTOTROMBIDIUM) 

Trambicula sp. Gill, Moule and Riek., 1925, Aust Vet. 



MZfi 



margin 



pan- 



tarsi 



Plate 9, fig. A-C. 

Description' of Larvae. Shape ovate. Length 
scutum more or less rectangular, with the anterioi 
ally, and posterior margin sbailowly convex with the 
not sinuous; surface coarsely punctate and with a 
near anterior margin. Eyes 2 -f 2, on distinct ocular 
the smaller. Chelicerae 44/* long and with only the usiia 
Galeal setae nude. Palpi stout, femur and genu with 
strongly ciliated setae; tibia with all 3 setae nude; 
rod, and 5 or 6 ciliated setae, one being much strong* 
others and over-reaching tip of palpal claw ; palpal dav\ 
prong the longest- Dorsum with 28 rather blunt-tipped 
long and arranged 2.8.6.6.4.2. Ventrally with the usual 
maxillary setae; a single seta on each coxa; a pair 
between coxae III, and thereafter 6.4.4.4.2.2 = 22 all 
dorsal setae, to 45/x long. Legs : I 370^ long, II 33(V, 
II with the usual dorsal rod-like setae, which are 
of tarsus; no long nude outstanding seta on tarsi III. 

The Standard Data for the type, and 15 paratyped 



relatively 



71 



sp.n. 
J, 32, fig. 9 (3) 



width 27fyu Dorsal 

concave except medi- 

m dian part flattened and 

of distinct oval discs 

: ] elds, the posterior eyes 

apical tricuspid cap. 

a long curved strongly 

with the usual basal 

r and longer than the 

( trifurcate, the median 

ciliated setae, to 42/a 

i>air of curved branched 

between coxae I and 

more slender than the 

III 405ju.; tarsi I and 

short, ca. % length 



from Queensland are 



72 



Records of the S.A. Museum 







Standard 


Theoretical 


Observed 


Coeff,of 




Mean 


Deviation 


Bange 


Range 


Variation 


AW 


74.8=b0-49 


1-97+0-35 


68-9-80-7 


72-0-78-0 


2-6 


PW 


85-0±0-4G 


1-85+0-33 


79-4-90-6 


82-0-87-0 


22 


SB 


25-5±0?41 


1-66+0-29 


20 -5-30 -5 


22-0-27-0 


6-1 


ASB 


29-0±0'35 


1-39+0-25 


24'£-33-2 


27-0-30-0 


4*8 


PSB 


17-6±0-21 


0-86+0-15 


14-95-20-25 


15-0-18-0 


4-9 


SD 


46 -5:1:0 -42 


1-69+0-30 


41 '4-61-6 


43-0-48-0 


3-6 


A-P 


29-7±0-33 


1- 31+0-23 


25-8-33-6 


27-0-33-0 


4-4 


AM 


31-9±0-38 


1-44+0*27 


27-6-36-2 


30-0-42-0 


4-5 


AL 


39*6:±0-50 


1*96+0-36 


33-7-45-5 


36-0-42-0 


5-0 


PL 


53- 8+0 -81 


3-12+0-57 


44-4-63-2 


51-0-60-0 


5-8 


Bens. 


45-25+0-55 


1-92+0-31 


39-5-51-0 


42-0-48-0 


4-2 



hoc. The type from a pallid cuckoo, Clermont, Queensland, 4 Nov. 1944 
(D. A. Gill) and 15 pafatypes from a "Lousy Jack", Logan Downs, Clermont, 
28 Nov., 1944 (D.A.G.). 

Remarks. In their interesting paper, "Trombidiosis of Sheep in Queens- 
land,' ■ Aust. Vet. J., 1945, 22, the authors, Messrs. D. A. GUI, G. B. Moule and 
R. F. Riek, state that this species, therein referred tq as new but unnamed 
species of Trombicula, was common on a number of other birds, the names of 
which are given. 

In that paper an exceptionally fine microphotograph of the dorsal scutum, 
as obtained by a new technique developed by Mr. Parrish of the McMaster 
Laboratory, is published. 

As stated above, myzantha is very closely related to the preceding species, 
in having the same number of dorsal and ventral setae, although the disposition 
in the anterior rows differs. The DS are longer and the dimensions of the 
dorsal scutum greater in myzantha, tut with further material, the two species 
may possibly be shown to be synonymous. 

Also 9 specimens from a "fruit pigeon" from Owi Island, Dutch N. Guinea, 
13 Oct. 1944 (D. E. Howell) and 22 specimens from "pigeon" from a small 
island off the north of Bougainville, Aug., 1945 (G> H. McQueen). 

In both these populations the Standard Data are very significantly lower 
than those of the Queensland population, while they are also significantly differ- 
ent between themselves. 

For these island populations the form names T. (?L.) myzantha L ouriensis 
and T, i*L.) myzantha f. bougainvillensw are proposed. 

The Standard Data are as follows: 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



Owi Island Population, 



AW 

PW 

SB 

ASB 

P8B 

SD 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
63-0±0-71 
69-45±0-78 
21-2±0-22 
27-0 

17-3±0-44 
44-3±0-44 
27-7±:0-44 
26-3±0-44 
35-3±0-44 
45-0:±0-50 
45-75:fcl-43 



Standard 
Deviation 

2-12±0'50 

2-35:±0-55 

0-67±0-16 

No variation 



32±0-31 
32±0-31 
32±0-31 
32±0-31 
32±0-31 
50±0-35 
87±1-01 



Bougainville Population, 



AW 

PW 

SB 

ASB 

P8B 

SB 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
66-7±0-54 
72-6±0-58 
22-6±0-39 
27- 9*0 -43 
16-4±:0-45 
44-l±0-43 
28-l±:0-43 
29-25±0-75 
37-45 ±:0-62 
46-0±0-64 
46-l±0-55 



Standard 
Deviation 

l-79±0-38 

1-92*0 «41 

30zt0-28 

37±0-31 

49±0-32 

37±0-31 

l-44±0-31 

2-12±0-53 

2-06±:0-44 

2-13±0-45 

l-55±0-39 



Theoretical 
Range 

56-7-69-3 

62 -4-76« 5 

19-2-23-2 

recorded. 

13-4-21-2 

40-4r-48-2 

23-8-31-6 

22-4-30-2 

31-4-39-2 

40-5-49-5 

35-15-54-35 



Theoretical 
Bange 

61-3-72-1 

66-8-78-4 

18-7-26-5 

23-8-32-0 

11-9-20-9 

40-8^*8-2 

23-8-32-4 

22-9-35-6 

31-3-43-6 

39-6-524 

41-4-50-8 



Trombicula (?Leptotrombidium) robusta Gunther 1941 

Proc. Linn. Soc. N.S.W., 66, (3-4), 157, 1941, fig. 1-5; Womersley and Heaslip, 
Tr. Eoy. Soc. S. Aust., 67, (1), 85, 1943. 



Plate 9, fig. D-F. 

This species was originally described from colonie^ 
maokloti Teinminck and of Microeca sp., from Bulolo, 
now referred two specimens from the ears of a "thrusjt 
landia, Dutch N. Guinea, 10 Dec., 1944 (CJB.R)- 

The dorsal scutum is as figured, approximately rec 
terior margin very shallow behind PL and medially siniu > 
greater than AW, and SB much nearer to PL than 
PL; the surface coarsely pitted, and near to anterior 
discs; scutal setae more or less subequal. Chelicerae 
tricuspid cap. Galeal setae nude. Palpal claw trifurca 



73 



Observed 
Bange 

60-0-66-0 
66-0-72-0 
21 -0-23 «0 

15.0-18*0 
42- 0-45-0 
27-0-30-0 
24- 0-27-0 
33-0-36-0 
42-0-48-0 
42-0-48-0 



Observed 
Bange 

63-0-69-0 

69-0-75-0 

21-0-24*0 

27-0-30-0 

15-0-18-0 

42-0-45-0 

.27-0-30-0 

27-0-33-0 

33-0-40-0 

42-0-50-0 

45-0-48-0 



Coeff.of 
Variation 

3-4 

3-4 

3-1 

7-6 
3-0 
4-8 
5-0 
3-7 
3-3 
6-2 



Coeff. of 
Variation 



4-6 
3-4 



on the legs of Pitta 

N, Guinea. To it are 

-like bird" from Hol- 



&ngular, with the pos- 
PW only a little 

o AL but anterior of 
[margin a pair of oval 

with only the apical 
;e; setae on femur and 



74 Records of the S.A. Museum 

genu branched; on tibia all 3 nude (Gunther states "two nude setae near the 
base and one with very short branches half-way"). DS 34, arranged 
2.12.8.6.4.2, to 50/a long. No long nude seta on tarsi of leg III. Body cordate 
and overlapping mouth parts. 

The Standard Data derived from the type and 2 paratypes, and 2 speci- 
mens from Hollandia are: 



• 


Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff * of 
Variation 


AW 


72-6+0-60 


1-34+0-42 


68 -6-76-6 


72*0-75-0 


1-S 


PW 
SB 


78-0+0-95 
24-0 


2-12±0-68 71-7-84-3 
No variation recorded. 


75-0-81-0 


2-8 


ASB 


32-4+0-60 


1-34+0-42 


28-4-36-4 


30-0-33-0 


4-3 


PSB 


15-0 


No variation recorded. 






SD 


46-8+0-73 


1. 64+0-52 


41-9-51-7 


45-0-48-0 


3-6 


A-P 


30*6±0« 60 


1-34+0-42 


26-6-34*6 


30-O-33-0 


4-4 


AM 

AL 


42-0 
47-4+0-60 


Only one determination. 
l-34±0<42 43-4-51-4 


45-0-48-0 


4-1 


PI/ 

Sens. 


52-8+0-73 
50-0 


1-64+0-52 47-9-57-7 
No variation recorded. 


51-0-54-0 


3-1 



Along with T. (L.) myzantha sp. n. and E philipi sp. n. r this species 
forms a small group of closely allied species in having the pair of anterior 
scutal discs. 

Troimbicula (Leptotrombidium) burmensis (Ewing 1945). 
Tronibicula burmensis Ewing 1945. Proc. Ent. Soc. Washington, 47, (3), 63. 

Plate 9, fig. G-K. 

This species was somewhat inadequately described from larvae from speci- 
mens from Ting Hawk, June, 1944, and Shaduzup, July, 1944, in Burma (coll, 
H. S. Puller). I have been able to study five specimens of this species collected 
by T. J. Lawrence and K, L. Coekings, from Rattus rattus Iruneuscubus from 
Imphal, India, Sept. and Oct., 1945, from which the following redescription is 
drawn. Three of these specimens were larval pelts recovered from the reared 
nymphs by K. L. Coekings. 

Redescription of Larvae, Shape oval. Length (unengorged) 290/x, width 
227/* (well engorged, according to Ewing, loo. cit t , 830^u and 440/a). Scutum 
rectangular, finely punctate, with the posterior angles well rounded, and the 
PL setae placed not quite midway between posterior margin and AL, i.e. PL 
are not at the posterolateral corners; AM placed well back from line of AL;. 
sensillae filamentous, with short barbs basally and longer ciliations distally, their 
bases slightly behind line of PL. Eyes 2 + 2, posterior the smaller, Chelieerae 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



75 



setae strongly ciliated- 
ciliated ; on tibia dorsal 
4/z, long, and arranged 



non-serrate, with only the apical" tricuspid cap. Galeal 
Palpal claw trif urcate ; setae on palpal femur and genu 
ciliated, lateral and ventral nude. Dorsal setae 28, to 

2.8,6,6.4.2. Ventrally with the usual pair of ciliated kaxillary setae, one on 
each coxa, a pair between coxae I and between coxae III, and thereafter ca. 26 
arranged ca. 12.6.4.2.2. Legs all 7-segmented ; tarsi I jind II with dorsal sen- 
sory rod, III without any long nude seta. 

The Standard Data for the above 5 specimens are ; 



AW 

PW 

SB 

ASB 

PSB 

SB 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
70-4 

78*9+1-42 
33-5zfc0-61 
31-4±0-64 
14 -7*0-32 
4«-l±0-78 
27-2:fc0'71 
59-5*1-26 
46*7*0-78 
71*0*0-64 
70-4 



Standard 
Deviation 

No variation 

3-18*1-00 

•36*0-43 

43*0-45 

•71*0-23 

•75*0-55 

►60*0-51 

2-82*0-89 

1-75*0-55 

1-43*0-45 

No variation 



Theoretical 
Bange 

recorded. 

69-4-88-4 
29*4-37-6 
27-1-35*7 
12-6-16-8 
40-9-61-3 
22-4-32-0 
51-0-68-0 
41-5-51-9 
66-7-75-4 

recorded. 



Observed 
Bange 

76-8-84-0 
32-0-35-2 
28-8-32-0 
14-4-16-0 
43-2-48-0 
25-6-28-8 
57-6-64-0 
44-8-48-0 
70-4-73-6 



Coeff. of 
Variation 

4-0 
4-0 
4-5 
4-3 
3-8 
5-9 
4-7 
3-7 
2-0 



Remarks, This species is placed in Leptotrombidiiim on the character of 
precoxal plates on coxae I of the nymphs, reared by (Jockings and correlated 
with the larval pelts. In the shape of the dorsal scutum and the position of PL 
it is closely related to the palpalis group, but differs in having the setae on the 
palpal femur and genu strongly ciliated. 

Trombicula (Neotbombicula) fobdi sp. n. Lawrehce, T. J. in MS. 

Gen. nov. species "FD. M Lawrence in MS. In Audy 1947, "Scrub Typhus 
Investigations in S.E, Asia, Pt III. Appendix 7." AMD. 7. War Office, 
London. 

Plate 11, fig. A-P. 

Description of Larvae. Shape oval. Colour in life unknown. Length 
(partially engorged) to 325/*, width to 225/x. Dorsal scutum rectangular, pitted, 
with PL foliate and nearer to AL than to posterior margin, posterior corners 
rounded; AM and AL normally ciliated, AM well back| from anterior margin; 
seneillae fairly wide apart, filamentous and ciliated on distal half, their bases 
well behind line of PL, Eyes 2 + 2, small, a little distance from scutum and 
apparently not on ocular shields. Chelicerae not serrate, with only the usual 
apical tricuspid cap. Galeal setae strongly branched. Palpi stout, tibial claw 



76 Records of the S.A. Museum 

bifid; setae on palpal femur, genu and lateral and ventral on tibia nude, dorsal 
on tibia strongly branched. All dorsal setae, like PL, foliate, with the lateral 
margins curled in and with a double fringe of strong setules, (the appearance at 
first glance suggests the setae are bifid, but this is not so), 48 in number and 
arranged 2.10.8.8.8.6.4.2, but rows 2 and 3 variable, to 42^ long. Ventrally 
the setae are slender and ciliated, a pair on maxillae, one on each coxae, a pair 
between coxae I and between coxae III, and then ca. 10.8.8, followed by ca. 
6*4.2. which are foliate similar to the dorsal setae, to 30ft long. Legs all 7-seg- 
mented; I 225/i long, II 208/a, III 260/i; tarsi I and II with dorsal sensory rod, 
III without any long nude seta. 

The Standard Data derived from the type, 6 paratypes and 2 larval pelts 
from nymphs reared by K L. Cookings are: 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


75-0±l-08 


3-24±0-76 


65-3-84-7 


70-4-80-0 


4-3 


PW 


84»3±l-09 


3-58±0-84 


73-6-95-0 


80-0-89-6 


3-9 


SB 


37-3±0-71 


2-12±0-50 


31-0-43-6 


35-2-41-6 


5-6 


ASB 


30-4±0-53 


1-603:0 -38 


25 6-35-2 


28-8-32-0 


5-2 


P8B 


16-0 


No variation recorded. 






SD 


46-4±:0-53 


l-60±0-38 


41-6-53-2 


44-8-48-0 


3-4 


A-P 


19-55±0-69 


2-08±0-49 


13-3-25-8 


16-0-22-4 


10-6 


AM 


44-8±l*43 


3-20±l-01 


35-2-54-4 


41-6-48-0 


7-1 


AL 


43-6±0-58 


l-66±0-41 


38-6-48-6 


41-6-48-0 


3-8 


PL 


48-9±:0-91 


2'42=£0-65 


41-7-56-1 


44-8-51-2 


49 


Sens. 


60-8. Averac 


;e of two determinations of 57*6 and 64*0. 





Loc. and Hosts. Type host, Ratttis roitus brunnetiscuhis Hodgson, from 
Imphal, Manipur State, India 1945. Paratypes from same locality, hosts, and 
date. 

The type host was a "Black Rat with gray belly. Battus sp." 

In Lawrence's MS. this species is stated to be. fairly common and distri- 
buted over a wide area from Imphal down to South Burma. As many as 500 
were taken from a single host. Other hosts were the tree shrew Twpaia belan- 
geri belangeri Wagner, and the common musk shrew Savncus caertdus fulvocine- 
reus Anderson, both of these from Kanglatongbi, Dec, 1945. 

Type and paratype in British Museum, paratype in U.S. Nat. Mm, and 
King Edward VII College of Medicine, Singapore. 

TBOMBIOUnA (NbOTROMBICULA) TRAtTBI Sp. U. 

Gen. nov. species "A." Lawrence, T. J. in MS. In Audy 1947. "Scrub Typhus 
Investigations in S.E. Asia, Pt. III. Appendix 7." AMD. 7. War Office, 
London. 



Womersley— Asiatic- Pacific Scrub Typhus Mites 



77 



ife unknown. Length 



Plate 11, fig. G-K. 

Description of Larvae . Shape oval. Colour in 
(somewhat engorged) 481/a, width 350/a.. Scutum rectangular, with SB behind 
line of PL ,- PL about midway between AL and posterior margin ; AM moderately 
well behind line of AL ; both AM and AL normally eifiated, PL foliate, with 
the lateral margins incurled and bearing a double fringe of setules as in preced- 
ing species; sensillae missing but probably filamentous and distally ciliated. 
Eyes 2 -J- 2, small, and apparently not on ocular shields. Chelicerae not ser- 
rate, with only the apical tricuspid cap. Galeal setae stfongly branched. Palpi 
stout, tibial claw trifurcate ; setae on palpal femur and genu, and laterally and 
ventrally on tibia nude, dorsally on tibia branched. Dbrsal setae foliate as in 
preceding species, but shorter to 30/4. and with fewer sethiles, 52 iji number and 
arranged ca. 2.10.10.10.8.6.4.2. Ventrally with a pair of ciliated setae on maxil- 
lae, a pair of slender ciliated setae between eoyae I and! between coxae III, one 
on each cox*, and behind coxae Til with ca. 8.8.10.8, followed by 8.6.4.2. foliate 
setae similar to dorsal setae. Legs all 7-segmented j I 546/1, long, II 430/*, III 
460/a; tarsi I pnd II with dorsal sensory rod, III without! any long nude seta. 

The Standard Data of the type are: AW 64-4, PW 75-6, SB 33-6, ASB 
28-0, PSB 15-4, SD 43-4, A-P 22-4, AM 42-0, AL 39-£, PL 36-4, Sens. 60-0. 

Loo. and Hast. The type specimen from Macacus adsamensis, from Imphal, 
India, August 1945 (T. J. Lawrence). 

I have also seen three other specimens, in the collection of C. D. Radford. 

The type specimen has very unfortunately become budly damaged, by using 
oil immersion to determine details of the palpal setae. 

Remarks. This species in the structure of the dorial setae and the shape 
of the dorsal scutum, is very close to the preceding species T. (N.) fordi sp. n., 
but appears to be distinct in the smaller scutum, and the dorsal setae being more 
numerous and shorter with fewer lateral setules. 



Trombicula (Neotrombicula) hast ata (G-Ater 1932) 

Trombicula hastata Gater 1932, Parasitology, 24, 151. 

Neoschongastia hastaia, Womersley and Heaslip 1943. [Tr. Roy. Soc. S. Aust. r 
67, (1), 116. 

Plate 12, fig. A-E, 

This species was described from the type and two paratypes from Rating 
surifer surifer (Miller) from Sungei Buloh, Selangor, Federated Malay States. 



It was later, despite the absence of the sensillae, referred 



to the genus Neoschon~ 



78 Records of the S.A. Museum 

gastia, as then understood, by Womersley and Heaslip (1943), rather than to 
the genus TrombicuLa. 

Until recently no further material has been collected since the original dis- 
covery. 

Gater's description is as follows: "Body ovoid, rounded; striations mode- 
rate, slightly crenulated; pitting on scutum, epistome and legs. Fangs of eheli- 
cerae moderately stout, slightly curved, abruptly tapering to a sharp point which 
appears to set at an angle; dorao-apical and ventral teeth apparently absent; 
maxillary setae strongly pectinate. Palpi rounded; first and second palpal 
setae short, plain; on fourth segment the dorsal seta with a few short barbs, 
the other two plain; appendiculum with six pectinate setae; palpal claw long, 
trifurcate, the upper accessory prong longer than the lower. Scutum broader 
than long (mean L 44/4, W 7 V; pseudostigraatic organs 37/t apart), trapezoidal, 
anterior margin concave, posterior margin slightly concave, corners rounded; 
acutal setae five in number, median anterior small, lanceolate and placed well 
back from margin ; anterior laterals short, stout, strongly pectinate ; situated on 
corners; posterior laterals large, lanceolate and situated under two-thirds the 
length of scutum fom anterior pair. Pseudostigraata behind posetrfor lateral 
setae, as far in front of the posterior margin of the scutum as , the distance 
between anterior and posterior lateral setae, pseudostigmatie organs missing in 
these specimens. Body setae large, forty-eight in number; on dorsum thirty, 
large, translucent, lanceolate, covered with minute hairs, becoming smaller pos- 
teriorly, but two pairs of the usual type of body seta, stout, minutely pilose, on 
posterior edge of body \ on venter of the usual type, strongly serrate to pectinate, 
much longer posteriorly than anteriorly. Foreleg, mean L 157/* ; leg setae mostly 
serrate or pectinate with a few plain; cox&l setae single, serrate or strongly pec- 
tinate, the one on the midcoxa the shortest. Length (partially engorged) 0-42- 
0*46 mm.; width 0-26-0-35 mm." * " ; 

In 1943 Womersley and Heaslip interpolated the Standard Data from 
Gater's figure and data as follows: AW 69-0, PW 71-0, SB 37-0, ASB 29-0, 
PSB 15-0, SD 44*0, A-P 15-0, AM 30-0, AL 32-0, PL 42-0, Sens. — . 

The "Maxillary setae" in the above description are the galeal setae of 
present day workers. 

The dorsal setae are arranged 2.8.6.6.4.2.2 and the ventral setae posterior 
of coxae III, 2.6.4.4.4. 

Recently I have received from Dr. J. R. Audy a paratype of Gater's .mate- 
rial from the I.M.R., Kuala Lumpur, as well as three specimens collected at 



- 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MflTES 



Bukit Lagon Forest Reserve, Kuala Lumpur, 1949-50, 
Battus subamiSj same locality 1949, which possessed longj 

From these 5 specimens the Standard Data are somewhat lower than the 
values interpolated from Gater's figures by Womersley arid Heaslip 1943, and 
are as follows i 



AW 

PW 

SB 

ASB 

PSB 

SB 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
59-4±0-56 
64-4 
36-4 

28-6±0-56 
14-0 

42-6±0-56 
12-4±0'40 
30*8 
30-8 

58-2±0-56 
70-0 



Standard 
Deviation 



Theoretical 
Range 

1-25:1:0«39 55-6-63-2 

No variation recorded. 

No variation recorded. 

l-25±0-39 24-8-32-4 

No variation recorded, 

l-25:£0-39 38-8^6-4 

0-89±0-28 9-7-15-1 

No variation recorded. 

No variation recorded. 

l-25±0-39 54-4-62-0 

Onlv 1 determination. 



79 



and one collected on 
filamentous sensillae. 



TaOMBICULA NOVAE-HOLLANDIAE Hirst 1929. 



Trombicvla novae-hollandiae Hirst 1929. Proe, Zool. Soc; 



Observed 
Range 

58-8-61-6 



28-0-30-8 

42-0-44-8 
12-0-14-0 



56-0-58-8 



Ooeff . of 
Variation 

2-1 



4-3 

2-9 
7-2 



2-1 



jondon, 172 ; Womers- 



ley 1934, Rec. S. Aust. Mus., 5, (2), 213; Womerslpy and Heaslip 1943, 



Tr. Roy. Soc. S. Aust, 67, (1), 95; Sig Thor and 
Tierreich, Lfg., 71b, 270. 

Plate 12, fig. F-L 

This species was originally described from the ears 
D'Estree Bay, Kangaroo Is., S, Aust., and Womersley and 
it from Potorus ti*idoAtyVo$ from Bothwell, Tas.' (Coll. H. 
as from Cairns, Queensland (host ?). 



' ri 



Willmann 1947, Das 



of Battus greyi from 
Heaslfp 1943, recorded 
H. Pinlayson), as well 



i v. 



The scutum is comparatively large and rectangular,. Mit the posterior mar- 
gin evenly rounded, and SB about in line with PL. Cheli{jerae simple with only 
the apical tricuspid cap. Galeal setae nude. All setae <jn palpal femur, genu 
and tibia branched ; tibial claw trif urcate. Dorsal setae 2tt in number, from 80 ■ 
to 50-Oju, long, and arranged 2.6.6.6.4.2. Tarsi of leg III with 2 long, nude, 
outstanding setae; metatarsi of leg III with 1 such seta 

The Standard Data for the type and 2 paratypes ftonx Kangaroo Is., 3 
specimens from Cairns, 12 from Tasmania and 2 from Loc. ? are as follows : 



80 



Records of the S.A. Museum 





- 


Standard 


Theoretical 


Observed 


Coeff, of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


70-65±0-55 


2-48:4:0-39 


63-2-78-1 


66-0-75-0 


3-5 


PW 


93-9:fc0-67 


3-01±0-48 


84*9-102-9 


87-0-99-0 


3-2 


SB 


30-3rb0-42 


l-87±0-29 


24-7-35-9 


27-0-33-0 


6-2 


ASB 


36-15±:0-23 


1-01±0-16 


33*1-39-2 


33-0-38-0 


2-6 


PSB 


26-65±0-27 


1-19±0-19 


23-05-30-25 


24-0-29-0 


4-5 


SD 


62- 65:1:0 -41 


1-82^:0-29 


57-2-68-1 


60-0-67-0 


2-9 


A-P 


33 -6:1:0 -50 


2-24^:0.35 


26-85-40-35 


27-0-36-0 


6-6 


AM 


55*2±0-75 


2-99:t0-53 


46-2-64-2 


48-0-63-0 


5-4 


AL 


50-4±0-50 


1- 85+0 -32 


44-85-55-95 


48- 0-54« 


3*6 


PL 


66 -85:1:1 -07 


4-68:1:0*76 


52-8-80-9 


60-0-78-0 


7-0 


Sens. 


95-8±l-27 


3-12±0«90 


86-4-105-2 


90-0-99-0 


3*3 



Remarks. A very characteristic species on the shape and dimensions of 
scutum, the dorsal setae, and the long nude setae on metatarsi and tarsi III, and 
separated as in the key to species. 

Trombicuia (Nbotoombicula) rara (Walch 1923). 

Trombieula rara Waleh 1923. Tr. Vtb. Bien. Cong. Far East. Assoc. Trop. Med., 

Singapore (1924) \ Womersley and Heasiip 1943. Tr. Eoy. Soc. S. Aust, 

67, (1),90. 
Trombieula (Eutr&mbicvla) rara, Sig Thor and Willmann 1947, Das Tierreich 

Lfg. 71b, 283. 

Plate 13, fig. A-D. 

This species was originally described by Walch from 4 specimens taken from 
man in Deli, Sumatra. Womersley and Heasiip 1943 recorded a specimen sent 
to them by Dr. J. E. Dinger of the Koningen Wilhelmina Institute v. Bactereo- 
logie, Batavia as this species, but a re-examination of the specimen shows this 
determination to be in error, the species being T. hirsti Sambon. 

From Walch 's original data and figures, Womersley and Heasiip 1943 com- 
puted the Standard Data as follows: AW 85-0, PW 104-0, SB 54-0, ASB 25-0, 
PSB 49-0, SD 74-0, A-P 400, AM 40-0, AL 33-0, PL 40*0, Sens. 57-0, DS 
to 34-0. 

The following material has been studied and is referred to this characteristic 

species : 

27 specimens from a "small plain skink," Mts. near San Juan, Mindoro, 
Philippine Is., April 11, 1945 (C.B.P.). 
4 Specimens from Hollandia, Dutch New Guinea (2 on a skink, 2 collected on 
boots), Dec. 8, 1944 (CJB.P.). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



81 



3 specimens from a skink Lygosoma (Leiolopisma) rhomb oidalis Peters from 
Lake Eacham, N. Queensland, 10 Sept., 1945 (R.V.S.). 
14 specimens from a pill-millipede (Zephronia sp.) from Kepong Forest Reserve, 
Kuala Lumpur, 1948 (J. R. Audy). 



From this material the following redescription is drawn : 

Larvae. Shape a rather broad oval. Length (imengorged) 240/*, width 
180/*. . Scutum large, almost quadrate, with the sensillae bases placed very much 
nearer to AL than to PL and widely separated. Eyes B -f 2. Chelicerae with 
the usual apical tricuspid cap. Galeal setae nude. Palpi stout, with bifurcate 
tibial claw ; seta on palpal femur with a few short branches, on genu and all 3 
on tibia nude ; tarsi with the usual sub-basal sensory rod a nude subapical seta, 
and 4 or 5 ciliated setae. Dorsal setae short and spine-likje with closely adpressed 
ciliations, 22 in number and arranged 2.6.6,4.2.2, somewbjat variable in length in 
different populations from 30/* to 56/*. Ventrally with the usual pair of branched 
setae on maxillae, a single ciliated seta on each coxa, a piir between coxae I and 
between coxae III. and thereafter 2.2.4.4.2.2.2 to 25tt. I Legs ■ I 270/* long, II 
240/i, III 270/i ; tarsi I and II with the usual dorsal sensciry rod, III with a long 
sub-basal nude seta. 

The Standard Data, derived from the specimens from the Philippines, 
Hollandia and Queensland are as follows; 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


74-45±0-71 


4-08±0-50 


62-2-86-7 


64*4-78-4 


5-5 


PW 


85-4±0-70 


3-99±0-50 


73-4-97-4 


75-6-92-4 


4-7 


SB 


54 -85:1:0 -64 


3-66±0-45 


43-9-65-8 


47-6-61-6 


6-7 


ASB 


19-75±:0*12 


0-69±0-08 


17-7-21-8 


19-6-22-4 


3*5 


PSB 


51-8±0-34 


1<98±0*24 


45-9-57-7 


44«8-53-2 


3*8 


SD 


71-552:0*32 


l«85±0-23 


66-0-77-1 


64*4-72*8 


2-6 


A-P 


36-45±0-47 


2-68±0-33 


28-4-44-5 


30-8-42-0 


7-3 


AM 


27-5±0-57 


3-O2+0-40 


18-5-36-5 


22-4^33-6 


11-0 


AL 


24-8±0-58 


3-21±0-41 


15-2-34-4 


22-4-33-6 


13-0 


PL 


32-3±Q-44 


2<47:±0'3l 


24-9-39-7 


28*0-39-2 


7-6 


Sens. 


49-2±0-68 


2-81±0-48 


40-8-57-6 


44*8-53*2 


5-7 



This composite population shows a moderate degree o{ variation but there are 
no significant differences between that from the Philippines and those from other 
localities, except that the dorsal setae in the Philippine material measures to 
31/i in length, whereas in the others the length is 42^,. 

The Standard Data for eleven specimens of a popu] 
pede from Kuala Lumpur, F.M.S., are as follows ; 



ation from a pill-milli- 



82 



Records of the S.A. Museum 





Mean 


Standard 
Deviation 


Theoretical 

Ran^e 


Observed 

Bange 


Coeff. of 
Variation 


AW 


70-65±0-55 


2-48±0-39 


63-&-78-1 


66-0-75-0 


3-5 


PW 


93-9:£0-67 


3-01+0-48 


84-9-102-9 


87-0-99-0 


3-2 


SB 


30-3:±0-42 


l-87±0-30 


24-7-35*9 


27-0-33-0 


6-2 


ASB 


36-15:fc0*23 


1-01:±0-16 


33-1-39-2 


33-0-38-0 


2-8 


P8B 


2G-65±0-27 


1-19±0-19 


23-05-30-25 


24-0-29-0 


4-5 


SD 


62-65r£0-41 


l*82±0-29 


57-2-68-1 


60-0-67-0 


2-9 


A-P 


33-6±0-50 


2-24±:0-35 


26-85-40-35 


27*0-36-0 


6-6 


AM 
AL 
PL 

Sens. 


55-2±0-75 
50-4±0-50 
66-85:tl-07 
95-8:fcl-27 


2-99±0-53 
l-85±0-32 
4-68±0*70 
3-12±0-90 


46-2-64-2 
44-85-55-95 

52-8-80-9 
86 -4-105 -2 


48-0-63-0 
48-0-54-0 
60-0-78-0 
90-0-99-0 


5-4 
3-6 
7-0 
3-2 



The Standard Data of this population more closely approaches that given by 
Womersley and Heaslip for Walch's figures, except that AM, AL and PL are 
longer, as are also the dorsal setae which reach 57^u. in length. 

Remarks, The common occurrence of this species on a pill-millipede in 
Malaya is remarkable as being an exception to the assumed restricted association 
of the Trombiculid larvae to vertebrate hosts. A few other cases of Leeuwen- 
hoekiids occurring on invertebrates are, however, also known, 

A single specimen collected on boots at Noemfoor Is., Dutch New Guinea 
(D, C. Swan) has a significantly larger scutum, the Standard Data being: 
AW 103-6, PW 109-2, SB 64-4, ASB 35-4, PSB 58-8, SD 95*2, A-P 44-2, Ail 
53-2, AL 44-8, PL 58-8, Sens. — . DS to 62/*. 

This specimen may be regarded as belonging to a distinct race or population. 

The species was also collected in numbers on Owi Is,, Biak Is., Dutch New 
Guinea, by Maj. G. M. Kohls, and I have seen some specimens from there, 
taken from boots, Aug. 7, 1944, and from Lygosoma hinulia variegatum Aug. 
12, 1944. 

Trombicula hirsti Sambon 1927. 

Trombicula pseudoakamushi v> deliensis Walcb 1923, Tr. Vth. Bien. Cong. Far 
East. Assoc. Trop. Med., 601, Singapore (1924). 

Trombicida hirsti Sambon 1927, Ann. and Mag. Nat. Hist., (9), 20, 157; nee. 
Hirst 1929, Ann. and Mag. Nat, Hist., (10), 3, 564; Gater 1932, Parasito- 
logy, 24, 143; nee. Womersley 1934, Rec. S. Aust. Mus., 5, (2), 212. 

Trothbioula hirsti v, morobensis Gunther 1938 (wow. nud.), Med. J. Aust., 2, 
(6), 202. 

Trombicuki hirsti v. buloloensis Gunther 1939, Proc. Linn. Soc. N.S.W., 
64, (1-2), 78. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 83 

Trombicida minor Womersley 1939 ? Tr. Roy. Soc. S. AiJsL, 63, (2), 152 (nee. 
Berlese),- Gunther 1939 (Dec), Proc. Linn. Soc, flfew South Wales, 64, 
(5-6), 466 (nee. Berl.) \ Womersley and Heaslip, Tr.jRoy. Soc. S. Aust., 67 
(1), 92 (nee. Berl) ; Womersley 1944, Tr. Itoy. Soc. L Aust., 68 (1), 92. 

Trombicida minor v. deliensis Womersley and Heaslip J.943, Tr. Roy. Soc. S. 
Aust., 67, (1), 94. 

Trombicida toiediacris Womersley and Heaslip 1943, Tr. JJS-oy.. Soc. S. Aust., 67, 
(1), 92, (nee. Berl.). 

Trombicida (Eidrombimla) hirsti Sig Thor and Willmanjii 1947, Das Tierreich, 
Lfg. 71b, 285 (in part). 

Plate 13, fig. E-L 

Willmann (Zool. Anz., 1940) has re-examined the typ$s of Trombicvla minor 
Berl. in the Hamburg Museum, and shown that not only [were they adults from 
Tjompea, Java (not nymphs as assumed by other workerfe), but that they were 
labelled "aus Hohlenguano gesiebt". The latter fact Was not mentioned by 
Berlese, but as cave guano is usually that of bats, it seems highly probable, as 
suggested by Ewing (Naval Med. Bull., 43, (4), 837-39J Oct., 1944), that the 
larvae of T. minor Berlese, when discovered will be found to be bat parasites. 

The larvae described and recorded in recent years from Australia and New 
Guinea, are generally found on birds, or animals, such as bush fowl, rodents and 
bandicoots, etc., and it appears unlikely that they can be referred to Berlese % 
species. Whether, however, any of the known bat-infepting larvae are those 
of T. minor Berl. must await rearing experiments, and, in view of the urgent 
need for correlating the larvae with the type of Tromhicula Berlese, every 
chance should be taken of rearing the known bat larVae to nymphs and/or 
adults,- as well as of re-discovering the adults and rearing larvae therefrom. 

In view of the above remarks, the question of the correct name to apply to 
the Australian and New Guinea species becomes important, and although still 
confused for the reasons stated, the synonymy appears to be as given above. 

The form described by Walch in 1923 as T. pseudoa&aniiishiv. deliensis from 
Sumatra, is the same as Gunther ? s hirsti v. buloloensis from New Guinea, but 
the Standard Data is slightly greater than in the typical form from Queens- 
land as is shown in the key to species. Sig Thor and j Willmann 1947 place 
Wakh's form as a synonym of T. pallida Nagayo et al,, from which it is clearly 
distinct as given in the key. 

As the New Guinea and Queensland material is so littlle different in Standard 
Data (see Key) the following values are given for a combination of 50 speci- 
mens from Queensland and 23 specimens from New Guinea : 



84 



Records of the S.A. Museum 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Kange 


Range 


Variation 


AW 


S3-5±0*34 


2-43±0-24 


76-2-90-8 


77-0-90- 


2-9 


PW 


96-6+0 -37 


2-65±:0-26 


88-7-104-5 


90-0-104-0 


2-7 


SB 


42-8±0-22 


l-53±0-15 


38-2-47-4 


40-0-47-0 


3-6 


ASB 


29-2+0-12 


0-86+0 -12 


26-6-31-8 


27-0-32-0 


2-9 


PSB 


33-6±0-26 


1-86+0-18 


33-0-44-2 


36-0-40-0 


4-8 


SD 


67-8±0-29 


2-09+0-21 


61-6-74-0 


64-0-72-0 


31 


A-P 


35-9±0-17 


1-23+0-12 


32-2-39-6 


32*6-39-0 


3*4 


AM 


4G-6±0-34 


2-35+0-24 


33-6-47*6 


34-0-47-0 


5-8 


AL 


45-95±0-33 


2-37+0-24 


38-85-53-05 


40-0-50-0 


5*2 


PL 


54-15±0-23 


1-56+0-16 


49-45-58-85 


50*0-57-0 


2-8 


Sens. 


64-9±0-36 


1-80+0-24 


59-5-70-3 


60-0-68-0 


2-8 



HIRSTI f . NISSANENSJB 110V. 

Tr&mbicula hirsti Dumbleton 1947, Tr. Roy. Soc. N. Zealand, 76, 413. 

Through the kindness of Dr. L. J. Dumbleton, I have been able to study a 
series of 14 specimens, 9 from pigs, and 5 from man, of the species which he 
recorded as T. hirsti from Nissan Is. (Green 1b.), Territory of New Guinea, 
1944. 

In the Standard Data given below, the values are significantly higher and 
the population may be regarded as a distinct form: 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


109-7+0-75 


2-89+0-53 


101-0-118-4 


104-0-115-0 


2-6 


PW 


125 '7+1 -16 


4-48±0-82 


112-3-139-1 


115 -0-133-0 


3-6 


SB 


58 -7+0 '61 


2-38±0-43 


51-6-65-8 


54-0-61-0 


4-0 


ASB 


32-0 


No variation 


recorded. 






P8B 


43-0 


No variation recorded. 






SD 


75-0 


No variation recorded. 


1 




A-P 


43-0 


No variation recorded. 






AM 


53-9+0*49 


1*88+0-34 


48-2-59-6 


50-0-57-0 


3-5 


AL 


61 -2+0 -82 


3-17+0-58 


51-7-70-7 


54-0-65-0 


5-1 


PL 


65-9+.1-22 


4-55+0-86 


52-3-79-5 


57-0-72-0 


6-9 


Sens. 


60-6+0*64 


2-23+0-46 


53-9-67-3 


57-0-65-0 


3-7 



niRSTi 1 hakei Radford 1946. 

Trombieula hakei Radford 1946, Proc. Zool, Soc., London, 116, (2). 

Radford described T, hakei from a Copper-headed snake (Coluber radiaius 
Schlegel) from Imphal, Manipur State, India, 10 May, 1940. I am greatly 
indebted to him for a paratype of his species, and from an examination of this, 
and of his description and figure I am convinced that it cannot be separated 
specifically from typical hirsti Sambon, except on the size of the scutum and 
the Standard Data, which lie between that of the typical form and f . iiissanense. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



The Standard Data from the paratype available to ine, as compared with 



85 



95-0 (85-0), PW 110*0 

48-0 (34-0), SD 81*0 

(^•O), PL 70-0 (54-0), 



the values given by Radford (in parentheses) are: AW 
(102-0), SB 43-0 (45-0), ASB 33*0 (34*0), PSB 
(68-0), A-P 39*0 (34-0), AM 58-0 (45-0), AL 55-0 
Sens. 70*0 (65-0). 

It is beat considered as only a race or form of hiifsti Sambon 

Trombicula sobrina sp. n. 
Plate 13, fig. J-L. 

Description of Larvae. Shape ovate with sides somewhat flattened. Length 
(engorged) to 630/*, width to 480/1. Scutum as figured, irregularly punctate, 
nearly twice as long as wide, anterior margin lightly sijnuous; posterior margin 
more than twice as deep as distance between SB and PL and rather evenly 
curved, SB nearer to line of PL than to AL. Eyes 2 + 2, on distinct ocular 
shields, fairly close to lateral scutal margins and posterior the smaller. Cheli- 
cerae damaged in all specimens. Galeal setae f Paljpi stout, with bifurcate 
tibial claw ,- setae on femur with 2-3 short branches, oflten appearing nude ; on 
genu nude, on tibia, dorsal and lateral nude, but ventrial branched ; tarsi short 
with sub-basal sensory rod -and a subapical nude seta, ind 6 ciliated setae, five 
of w r hich have 3-4 short stout branches, the other being stouter with more 
branches and over-reaching the tip of palpal claw. Dorjsal setae rather stiff and 
blunt with short ciliations, 20 in number and arrange^ 2,6,6 4.2, to 50/i long. 
Ventrally with a pair of branched setae on maxillae, a ciliated seta on each coxa, 
a pair between coxae I and between coxae III and thereafter 6.2.2,2, to 30/i long, 
and rather more slender than dorsal setae. Legs i I 320^ Jong, II 300/*, III 320/*; 
tarsi I and II with dorsal sensory rod, Til with a long i^ude outstanding seta. 
The Standard Data for the type and 10 paratypes are| as follows : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Devi.-ition 


Range 


Range 


Variation 


AW 


97-l±l-00 


3-33±:0-71 


87* 1-107-1 


91-0-103-0 


3-4 


PW 


lU-3^3-09 


3-61+0-77 


100-5-122-1 


n07'0-119-0 


3-2 


SB 


48-l:±0-43 


l'46rt0.31 


43-8-52-4 


46-0-51-0 


3-0 


ABB 


27-55rt0»41 


l-35:£0-29 


23-5-31-6 


26-0-30-0 


4-9 


PBB 


34-G±0-53 


l-76±0-38 


28-7-39*3 


32' 0-37-0 


5-2 


SD 


61-4+0-90 


3-98±0-63 


52<5~7C4 


58-0-66-0 


4-8 


A-P 


34-2±0-52 


l-72:t0-36 


29*1-39-3 


32-0-37-0 


5-0 


AM 


39'7:£0-33 


0-82±0-23 


37-3-42 vl 


38-0^0-0 


2-0 


AL 


46-l:±0-64 


2-02:tO-45 


40 '1-52-1 


43-0-50-0 


44 


PL 


54-0±0-60 


2* 00+0 -42 


48-0-60-0 


50-0-56-0 


3-7 


Sena. 


64*0 


Only one deter 


mination. 







86 Records of the S.A* Museum 

Loc. and Sost. The type and 10 paratypes from a skink, Lygosoma Stanley- 
anum from the Wau-Edie Creek Rd., New Guinea, 11 Sept., 1944, along with 
T. lygosomoidres sp. n. collected by the late Dr. Consett Davies. 

Remarks. This species in the bifurcate palpal claw, the DS, and the punc- 
tation of the scutum, is closely allied to T. hirsti. It differs from that species, 
however, in the scutum being very much wider than long, the ratio of PW/SD 
= 1-86 (in hirsti PW/SD — 1*35 to 1-52) as in the key. 

Trombicula ablephara sp. n. 

Plate 14, fig. A-C. 

Description of Larvae. Shape oval. Length (partially engorged) to 450^ y 
width to 360/*. Dorsal scutum as figured, with strongly sinuous anterior margin 
and evenly rounded posterior margin which is less than twice as deep as the 
distance between line of SB and PL. SB nearer to PL than to AL, surface only 
sparsely and irregularly punctate, sensillae missing in all specimens ; scutal setae 
short, pointed and shortly ciliated. Eyes 2 + 2, rather small, on distinct ocular 
shields, posterior the smaller, Ghelicerae damaged, Galeal setae nude. Palpi 
stout with bifurcate tibial claw ; setae on femur, genu and tibia all nude ; tarsi 
with sub-basal sensory rod, and subapical nude seta, and 5-6 ciliated setae, one 
of which over-reaches tip of claw. Dorsal setae stiff and only slightly curved, 
shortly ciliated, 22 in number, to 35«x long and arranged 2.6,6.4.2.2. Ventrally 
with paired branched setae on maxillae, a single filiated seta on each coxa, a 
pair between coxae I and between coxae III, and thereafter ca. 4.2.2.2, to 30//. 
long. Legs: I 240/x long, II 210/*, III 240/* ; tarsi I and II with dorsal sensory 
rod, III with a long outstanding nude seta. 

The Standard Data for the type and 3 paratypes are : 

Standard Theoretical 

Mean Deviation Kange 

AW 73-25±0-75 l-50±:0-53 68-75-77-75 

PW 90*2±l-25 2-50±:0'88 82-7-97-7 

SB 36-0±0-58 1.15^0-41 32-5-39*4 

ASB 21-0 No variation recorded. 

PSfi 31-0:±0'58 1-15±0-41 27-65-34-45 

SD 52-0:tO-58 1*15±0*41 48-5-55-4 

A-P 29-0rt0-41 0-81=t0*29 26-6-31-4 

AM 24*0 No variation recorded. 

AL 24*5±0-50 l-00:fc0*35 21-5-27-5 

PL 32*5±0*50 l-OOzfcO-35 29 -5-35-0 

Sens. Missing in all specimens. 



Observed 


Coeff . of 


"Range 


Variation 


72-O-75-0 


2-0 


87-0-93-0 


2-7 


35-0-37-0 


3*2 


30-0-32-0 


5-7 


51*0-53-0 


2-2 


28-0-30-0 


2*8 


24-0-26-0 


4-1 


32.0-34-0 


3-1 



Womersley— Asiatic- Pacific Scrub Typhus Mites 



87 



Lac. and Host. The type and 3 paratypes fijom a skink Ablepharus 
bottom from Port Moresby, New Guinea, July, 19^4, collected by the late 
Dr. Consett Davies. 

Remarks. Although the sensillae are missing in ajll specimens, this species 
is undoubtedly a true Trombicula in the older sense. M differs from the closely 
allied species as given in the key. 

Another small population from a skink Lygosoma iLeiotopisma) chaUengeti 
from Mt. Laraington, National Park, Queensland, Die, 1948 (H.W,) is also 
referred to this species, although the Standard Data are significantly greater 
and suggest a different race or form. The Standard Data for 7 specimens are: 



AW 


Mean 
86-8:£l»22 


Standard 
Deviation 

3-23:±0-86 


Theoretical 
Range 

77-1-96-5 


Observed 
Range 

84.0-92-4 


Coeff . of 
Variation 

3-7 


PW 


104-8±l-48 


3-91±:l-05 


93-1-116-5 


98-0-109-2 


3-7 


SB 
A8B 


46-0±l-03 
25-2 


2-73rfc0-73 37-8-54-2 
No variation recorded. 


42-0-50-4 


5-9 


PSB 
SD 
A-P 
AM 


28-0 
53-2 

32-4±0-5G 
30-8 


No variation recorded. 
No variation recorded. 
1 -50+0 > 40 27-9-36-9 
v No variation recorded. 


30-8-33*6 


4-6 


AL 
PL 

Sens. 


33-2±0*40 
44-4±0-73 
60-7±0-93 


l-06±0-28 
l-93±:0-52 
1-62:1:0-66 


30-O-36-4 
38-6-50-2 
55-9-65-5 


30-8-33-6 
42*0-47-6 
58-8-61-6 


3-2 
4-3 
2-7 



1909 in Cr. Soc. Biol., 



Trombicula ( Trombicula) wiohmanni (Cuds. 1905). 

Trambidium wichmanni Ouds. 1905. Ent. Ber., 1, (22),|217; idem, Nova Guinea, 

1905, 5, 106, 132, pi. IV, fig. 67-68; idem, 1908, Tijds. Entom., 51, 25, 52, 38. 

AUothrombium wichmanni Ouds. 1906. Ent. Ber. 2, (p8), 58-59. 

Trombidiwm {Heteroirombidmm) wichmanni, Verdun 
Paris, 67, 246. 

Microtrombidium wichmanni Ouds. 1909, Ent. Ber, 3|, (50), 20; idem, 1912, 
Zool. Jahrb., Suppl. 14, 9. 

Trombicula mediocris Berl. 1912, Redia, 8, (1), 93; flatori 1919, Ann. Trop 
Med. Parasitol., 13, 233; Kawamura and Yamaguchi 1921, Kitasato 
Archiv. Exper. Med., 4, 169. 

Trombicula pseudoaikamushi Hatori 1920 (non Tanaka), Taiwan Igakai Zasshi, 
No. 209. Pt. II; idem, Ann. Trop. Med. Parasitol, 
Yamaguchi 1921, Kitasato Archiv. Exper. Med., 4, 

Trombicula hirsti, Gater 1932, Parasitol., 24, 147. 



±3, 233 ; Kawamura and 
169. 



88 Records of the S.A. Museum 

Trombicula urichmanni, Gunther 1940, Proc. Linn. Soe. New South Wales, 65, 

(5, 6), 480; Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust, 67, (l) r 

92. 
Trvmbicxda hatorii Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, 

(D,92. 
Trombicula (Eutrombicutia) wichmanni, Sig Thor and Willmann 1947, Das 

Tierreich, Lfg,, 71b, 277. 

Plate 14, fig. D-G. 

Oudemans originally described T. wichma/nmi from the Crowned Pigeon 
Goura ooronata from New Guinea, and from man in the North Celebes. 

In 1940, Gunther recorded it from the mouse deer, Tragulus borneanus 
from British North Borneo. Of his material four specimens were reported on by 
Womesley and Heaslip in 1943 and the Standard Data therefrom given. 

In 1919 Hatori referred to, but did not describe a larval Trombmda from 
Formosa under the name of "pseudoakamdiski (now- Tanaka)" and -claimed to 
have reared larvae from captured adults considered to be identical with 
T. medsiooris BerL Later in 1921 Kawamura and Yamaguchi working also 
on Formosan material described and figured in detail the larvae of what they 
regarded as Hatori 'a species; they also obtained adults and affirmed Hatori 's 
view that the larvae were those of mediocris BerL 

In 1943 Womersley and Heaslip after showing that the name pseudoaM- 
mushi was invalid, named the Formosan larvae as hatorii and gave the Standard 
Data based on figures and measurements of Kawamura and Yamaguchi. 

The identity of the larvae with the adult mediocris was still regarded as 
not finally proven. They were, however, shown to be closely allied to vmihmanni 
Ouds., only differing in the higher values of AM, AL, PL and Sens, and in the 
posterior seutal margin not being so sinuous medially but more evenly rounded. 

I have received from Maj. R. N. McCulloch four adult Trombicula collected 
in soil in New Guinea, from which he successfully reared a number of larvae* 
These adults, as discussed in the Adult Section of this paper, proved upon 
examination to agree with the figures and data given by Kawamura and Yama- 
guchi for their Formosan mediocris. They also agreed with Berlese's descrip- 
tion except that he gives the colour as white whereas the Formosan and New 
Guinea specimens were red in life. Berlese's material, however, may have be- 
come bleached in spirit. 

In describing his methods of rearing larvae from adults captured in the 
field, Maj. McCulloch writes: 

"The adult specimens later determined as mediocris were collected at Lae 
on April 9th and 10th, 1944. In searching for them and keeping them (and also 



Womersley— Asiatic-Pacific Scrub Typhus Mites 89 

the larvae) in the laboratory I followed the advice of Maj. G. M. Kohls. In 
jungle where larval mites were relatively numerous, litter was scraped from 
the soil and adults watched for on the surface, then in the top inch of soil and 
finally in some patches by digging and crumbling the soil down to about 6 inches. 

"In two days about 15 adult red Trombiculids ware found in or close to 
the surfaee. There were also present at various depths more numerous and 
larger mites, probably adult Erythraeidae, etc. The adult Trombiculids were 
tubed individually, each with about a teaspoonful oi moist soil, previously 
sterilized by steaming, from the locality. The test-tubis were closed by corks, 
perforated to allow aeration, covered with calico and iniverted into the tubes to 
give a tight fit so that the larvae could not force their wajf into any space between 
the edge of the cork and the glass. j 

"The specimens were kept in the laboratory wherejthe temperature varied 
between 74° F. in the early morning and 85°-88° F. ini the afternoon. A few 
drops of water were added to the soil in each tube frpra time to time and a 
little condesed moisture was always present on the inn$r wall of the tube. In 
this manner, larvae of Schongastia blestowei were kept alive for as long as 46 
days, 

"The living adult Trombiculids were examined from time to time as other 
work allowed. On May 15th it was observed that one adult had produced more 
than a dozen larvae apparently sometime earlier. No em "were found. Some of 
the other adults had died, and the remainder had then to be killed. The 
specimens of larvae and adults were then forwarded to Adelaide for identifica- 
tion. 13 

This species is certainly very closely related to kirsii Sarabon as was shown 
by Sambon 1927. The difference of the evenly curved posterior scutal margin 
in hirsti (sic minor) and the medially sinuous margin iln wichmanwi, given by 
Womersley and Heaslip in 1943 is not easy to use in practice. The two species 
differ more markedly, however, in the greater depth of the posterior scutal 
margin in hirsti. than in wichm<mni. In hirsti, the greatest depth of the pos- 
terior margin beyond the line of PL is twice or more, than the distance 
between tlie lines of SB and PL ; in wichmanni it is less than twice the distance. 

The relationship of I 7 . (T.) wichmanni (Ouds.) toi other species is shown 
in the new key to the species. Along with T. (T.) ablephata sp. n. it differs 
from T. (T.) hirsti Sambon and T. (T.) sobrina sp. n. in the much shallower 
posterior scutal margin. From ailephara it differs in having both the femoral 
and ventro-tibial setae of the palpi branched. The surface of the dorsal scutum 
is distinctly striate-punetate as is clearly indicated in Oudemans , figures, and 
in this feature urichmawni is closely related to the sctnoaifies-kohlsi^tovelli group, 
but differs in the presence of a long nude seta on tarsi III. 



90 Records of the S.A. Museum 

As in other species of whioh a fair amount of material from different 
localities is available for study, the different populations show significant varia- 
tions in the Standard Data measurements. 

Pour of these populations have been examined biometrically, as given in 
Table B. The populations are: 

I. Borneo: Six specimens from C, Gunther from the mouse deer r 
Tragulus borneanus, from Bode River, 1939; eleven specimens collected 
by R. N. McCullocli as follows: Labuan, 17 June, 1945, on boots (1); 
Limbang, 22 July, 1945, on boots (3) ; Balikpapan, 23 Aug., 1945 (1) ; 
ditto, 1 Sept., 1945, on rotten log (1) ; Brunei, 8 July, 1945, on boots 
(1), 20 July, 1945, on boots (3); (seventeen specimens in all). 
II. Celebes: Morotai, Halmahera, on boots in Sabala scrub, 13 and 14 May, 

1945, (3 spec.), (R.N.McC). 
III. New Guinea: Lae'll Mar., 1944 (1), 17 Apr., 1944 (1) on boots 
(R.N.McC); Dobodura, 6 Sept., 1943 (1 on man), (R.N.McC); Dobo- 
dura, June, 1944, on snake (5), (G. M. Kohls) ; Lae, 10 Apr., 1944, (1 
spec, reared from adult identified as T. mediocris Berl.), (R.N.McC.) r 
nine specimens in all). 
IV. Philippine Island: Bagabag, Luzon, 10 Aug., 1945, (6 spec, off Man) r 
(C. B. Philip and A. Mundan). 

From the Analysis of Variance of these populations (Table B) it is 
seen that the variation between the populations is significantly different from 
the variation within the populations at the 0-1% level of probability. 

Similarly the Difference of Means of the Standard Data of the population 
from Borneo compared with those of other populations in general, also shows, 
significant variations as in the following table. 

r 

TABLE A. 

Significance of Differences of Standard Data of 3 Populations of T. (T.) wich- 
manni compared with a fourth populations from Borneo, at 5%, 1% and 

0-1% probability. 

AW PW SB ASB PSB 8D A-P AM AL PL Sens, 



Celebes + ++ +++ +++ — ++ + — ++ +++ +++ — 

New Guinea — — 4- 4-4-4- — +++ +++ + ++ +++ 4-H-+ 

Philippines h + + + + + — +++ ++ ++ + +++ +4- 4- 



— Not significant + + Significant at 1 p.c. 

+ Significant at 5 p.c. 4-4-1- Significant at 0*1 p.c. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



91 



As the values of F in the Analysis of Variance :n the above populations, 
.although significant at the 0*1% level of probability riven in Fisher's Tables, 
are very much lower than those found in the different 
sis and akamushi, it would seem uncertain whether 



populations of T. delien- 
the populations of wick* 



-marmi are sufficiently distinct to justify racial trinomill names. For the present 



are biometric differences between the populations from 


. different 


areas. 






TABLE 


B. 






Frequency of different values of Standard Data in 


microns f c 


r four popula- 


iions of Trombicula mchmanni (Ouds.). 








N.B.- 


-The variance Ratio (F) from Fisher's tabl 


es for n/n 2 at the 0-1% 


level is given in parenthesis 


• 








AW = Anterior width of dorsal 


scutum as measured between AL 


■ 




AW 


I 


h 


Ill 


IV 


Total 


83 


1/83 




1/83 




166 


84 












85 






2/170 




170 


86 


4/344 




2/172 




516 


87 












88 


3/264 


• 






264 


89 


2/179 




1/89 




267 


90 


5/450 






4/360 


810 


91 • 


i 




2/182 


1/91 


273 


92 


J 










93 






1/93 


1/93 


186 


94 


2/188 








188 


95 










. 


96 










• 


97 




2/194 






194 


98 












99 












300 












101 




1/101 






101 




17/1507 


3/295 


9/789 


6/544 


3,135 


M±<r M 


88-65zt0-68 


98*3drl-34 


87-55:+: 


1-14 


90- 7+0 -49 


S±<r s 


2*81+0-48 


2*32+0-95 


3-43± 


0-81 


1-21+0-35 






Analysis of Variance. 






Variability. 


L 


Mean Bq. 




F. 


Between 


Groups 


3 


95-2889 


.. 12-37 (7-05) 


Within 


Groups 


31 


7-7001 



















92 



Records of the S.A. Museum 



PW = Posterior width of scutum as measured between PL. 
PW I II in 



IV 



Total 



95 






1/95 






95 


96 














97 






2/194 






194 


98 














99 














100 


1/100 










100 


101 


4/404 


• 


2/202 






606 


102 














103 














104 


9/936 




3/312 


1/104 




1,352 


105 














106 


1/306 


- 








106 


107 




• 










108 


2/216 




1/108 


1/108 




432 


109 














110 






• 


2/220 




220 


111 




1/111 




2/222 




333 


112 














113 














114 














115 




2/230 








230 




17/1,762 


3/341 


9/911 


6/654 
1 


109- 


3,668 


M±<r M 


103-65±:0-56 


113 •7±1- 34 


101-2:fcl-4 


0±:1«09 


s±* s 


2-31±0-40 


2.32^:0-95 


4-24±l-0 




2-68±:0-77 






Analysis of Variance. 








Variability. 


t. 


Mean Sq. 




F 


• 


Between 


Groups 


3 


159-8318 




17-94 


(7-05) 


Within Groups 


31 


8-9066 









SB = as measured between sensiilae bases. 
SB I II 



III 



IV 



Total 



40 
41 


1/40 
















40 


42 
43 
44 


1/42 
11/473 








1/42 
1/43 


. 






84 
516 


45 
46 


2/90 








4/180 




1/45 




315 


47 
48 
49 
50 


2/94 




1/47 
2/100 


• 


2/94 
1/48 




5/235 




470 
48 

100 




17/739 




3/147 




9/407 




6/280 




1,573 


Mrfccr M 


43-35±0' 


41 


49- 


0±l-0 


45 


•2±0 


•64 


46 


•7±0-33 


8±<r a 


l-70±0 


•29 


1*73±0-71 


1- 


92±0-45 


0- 


82±0-23 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



Variability. 



Analysis of Variance. 

f • Mean Sq. 



Between Groups 
Within Groups 



3 

31 



34-9205 
2-7459 



A8B = distance between anterior margin of dorsal scutum and a line joining centres of 
sensillae bases. 



ASB 



n 



III 



26 
27 
28 
29 
30 
31 
32 



1/27 
5/145 

11/352 



3/96 



4/104 
1/27 

3/87 



1/32 



17/542 



3/96 



9/250 



M±<r M 
S±<r 



Variability, 



30-8=0-41 
l-70±0-29 



32-0 



27-8±0'70 
2*11 ±0-50 



93 



12-7 (7-05) 



rv 



Total 



6/192 



104 
54 

232 



672 



6/192 



1,062 



32-0 



Analysis of Variance. 

f • Mean Sq. 



Between Groups 
Within Groups 



3 
31 



29-8595 
2-646 



PSB m distance between posterior margin of dorsal scutum and a line joining sensillae bases. 



P8B 



II 



III 



30 
31 
32 
33 
34 
35 
36 



2/60 
10/320 
2/68 
3/108 



3/108 



2/60 
6/192 

1/36 



17/556 



3A08 



9/288 



M±(r M 
S±<r„ 



Variability. 



32-7^:0-45 
l*86±0-32 



36-0 



32*0i:0-99 
2-193:0 -73 



Analysis of Variance. 

f . Mean Sq. 



Between Groups 
Within Groups 



3 
31 



28-8807 
2-5655 



P. 



11-28 (7-05) 



IV 



Total 



6/216 



120 

512 

68 

468 



6/216 



1,168 



36-0 



11-26 (7-05) 



94 Records of the S.A. Museum 

SD = total depth of dorsal scutum = ASB + PSR. 



SD 



56 
57 
58 
59 
60 
61 
62 
63 
64 
65 
6G 
67 
68 



1/57 
1/59 
4/244 

6/384 
2/132 
3/204 



II 



3/204 



III 



2/122 



IV 



6/408 



Total 



1/56 


56 




57 


3/174 


174 


2/118 


177 



384 
132 
816 



17/1,080 



3/204 



8/470 



6/408 



2,162 



M±<r M 

S±<r Q 



63 -5:2:0 -77 
3-18=t0-55 



68-0 



58-75^0-51 
l-67±0-42 



68-0 



Variability. 



Analysis of Variance. 

f . Mean Sq. 



Between Groups 
Within Groups 



3 
30 



120*8333 
6-0575 



19-94 (7-05) 



A~P = distance between AL and PL. 

A-P I II 



III 



IV 



Total 



32 


1/32 








8/256 






288 


33 


1/33 








1/33 






66 


34 


















35 


4/140 














140 


36 


6/216 










2/72 




288 


37 






1/37 










37 


38 


















39 


5/195 




2/78 






4/156 




429 




17/616 




3/115 




9/289 


6/228 




1,248 


M±<r M 


36-2:fc0' 


52 


38-3=b0' 


•67 


32 


•1±0-11 


38 


•0±0-63 


S±<r a 


2-14:±0 


-37 


l-16±0-47 


0- 


33±0-08 


1- 


55±0-45 



Womersley— Asiatic-Pacific Scrub Typhus Mites 

Analysis of Variance. 
Variability. f. MeanSq. ] 



95 



Between Grouj 


IS 


3 




57-7571 




20-42 


(7-05) 


Within Groups 




31 




2-8585 








AM = length of antero-median scutal seta. 


AM 


I 


II 




III 


IV 




Total 


39 








1/39 






39 


40 


1/40 


1/40 




2/80 






160 


41 








1/41 






41 


42 


1/42 












42 


43 


2/86 


2/86 




2/86 






258 


44 
















45 


2/90 












90 


46 
















47 


4/188 












188 


48 


1/48 






1/48 






96 


49 
















50 


4/200 








3/150 




350 


51 
















52 
















53 
















54 










1/54 




54 




15/694 


3/126 




7/294 


4/204 




1,318 


M±<r M 


46-3±0-82 


42-0±l- 


42-0rtl 


-15 


51- 


Ozfcl-0 


S±<r g 


3*19±0-58 


J-fi 


}±0 


•71 3-05±0-82 


2-0±0-71 






Analysis 


of Variance. 








Variability. 




f. 




Mean Sq. 




K 




Between Groups 


3 




84-0797 




9-69 


(7-45) 


Within Groups 




25 




8-6773 









AL = length of antero-lateral scutal setae. 
AL I II 



III 



IV 



Total 



43 

45 

46 
47 
48 
49 



3/141 



1/48 
5/250 



1/43 
2/90 

2/96 
1/50 



43 

90 

141 
144 

300 



96 



Records of the S.A. Museum 



AL 



II 



III 



IV 



Total 



51 




1/51 










51 


52 




1/52 






1/52 




104 


53 




- 












54 




9/486 






1/54 


1/54 


594 


55 
















56 












1/56 


56 


57 












3/171 


171 


58 
















59 






• 










60 
















61 












1/61 


61 






17/887 


3/141 




8/385 


6/342 


1,755 


M±om 




52-2±0-52 


47.0 




48-l±l-32 




57-0±0-93 


S±a s 




2*13r±0-36 


— 




3-74±0-94 




2-28±0-66 








Analysis of Variance. 






Variability 




f. 




Mean Sq. 




ft 


Between 1 


Groups 


3 




112-7279 




16-92 (7-05) 


Within Groups 


30 




6-16615 






PL = length 


of 


postero-latera] 
I 


i sental setae. 










Th 


II 




III 


IV 


Total 


50 










2/100 




100 


51 
















52 






. 










53 










1/53 




53 


54 






2/108 








108 


55 










2/110 




110 


56 










2/112 




112 


57 




1/57 


1/57 








114 


58 




1/58 










58 


59 
















60 




2/120 


• 




• 


■ 


120 


61 




7/427 






1/61 


1/61 


549 


62 




1/62 










62 


63 




2/126 








1/63 


189 


64 




2/128 










128 


65 




1/65 








1/65 


130 


66 
















67 














• 


68 












3/204 


204 






17/1,043 


3/165 




8/436 


6/393 


2,037 


M±:<r M 




61-35rb0-5 


, 55*0±1< 





54-5±:l*: 


23 


65-5r±l-23 


S±<r s 




2 -06:1:0*35 


l-73±0 


-71 


3-58:±0'i 


30 


3-01±0-87 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



97 



Variability. 



Between Groups 
Within Groups 



Analysis of Variance. 

f . Mean Sq. 



3 
30 



176*451 
6-9794 



F. 



25-28 (7-05) 



Sens. = length of sensillae. 



Sens. 




I 




II 


ni iv 


Total 


60 
61 










1/60 


60 


62 














63 














64 
65 
66 
67 








1/65 


3/192 

1/65 
1/66 


192 

130 

66 


68 
69 




4/272 




- 


2/136 


408 


70 
71 




2/140 




1/70 


• 


210 


72 
73 




5/360 








360 


74 














75 




1/75 








75 






12/847 




2/135 


5/318 3/201 


1,501 


M±<r M 




70-6:t0 


•66 


67-5±2 


•5 63-6±0-99 


67-0±l-0 


S±<r g 




2-27±:0' 


46 


3-53±l 
Analysis o 


•77 2-20±0-69 
f Variance, 


l-73±0-70 


Variability. 






f. 


Mean Sq. 


F. 


Between Groups 
Within Groups 




3 
18 


59-749 11 
5-2564 


-367 (8-49) 



A further population of nineteen specimens collected from a species of 
Lygosoma at Kerowie, N.6. at 5,000 feet by the late Dr. Consett Davis has the 
following Standard Data. It comes very close to the small population from 
the Celebes reported above, in that it does not differ significantly at the 0-1% 
level of probability. 



98 



Records of the S.A. Museum 



AW 

PW 

SB 

A8B 

PSB 

SD 

A-P 

AM 

AL 

PL 

Sena. 



Mean 
98-2±0-53 
113-25±0-44 
48-85±0-39 
27-3:t0-35 
32-4±0-15 
59-85±0-44 
36-25:±0-45 
35-95±0-46 
45-0±0-46 
53-l±0-60 
67-0:t0-80 



Standard 
Deviation 

2-32±0-38 

l-94±0-Bl 

l-71:t0-28 

l-53±0-25 

0-84±0-12 

1-92±0»31 

l-97±0*32 

2-03±033 

2-00±0«32 

2-62±0-42 

3-40±0-57 



Theoretical 
Range 

91*3-105-1 

107-45-119-05 

43-7-54-0 

22*7-31-9 

29-9-34-9 

54-1-65-6 

30-35-42-15 

29-3-42-0 

39-0-51-0 

45-3-60-9 

56-8-77-2 



Observed 
Range 

94-0-101-0 

110-0-117-0 

45-0-52-0 

25-0-29-0 

32-0-34-0 

57-0-63-0 

32-0-39-0 

32*0-39-0 

43-0-47-0 

47-0-57-0 

61* 0-72-0 



Coeff. of 
Variation 

2-4 
1-7 
3-5 
5-6 
2-6 
3-2 



From Malaya (Kuala Lumpur district) also I have been able to study a 
large number of specimens of wichmanni, and from this population of 47 speci- 
mens, including 17 of Gater's original Malayan material labelled as "Trombi- 
cida hirsti Sambon ,, J the Standard Data is as follows: The slides of Gater's 
and the other material were kindly sent to me by Dr. J. R. Audy. 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


88-5dt0-40 


2* 74;±0- 28 


80-3-96-7 


84-0-95-2 


3-1 


PW 


.J07-75±0«QO 


3-46^0-36 


97-4-118-1 


103-6-114-8 


3-2 


SB 


45-3±0-17 


1*20:±0-12 


41-7-48-9 


44-8-50-4 


2-6 


ASB 


31-75±0-21 


l-43±0-15 


27-45-36-05 


28-0-33-6 


4-5 


PSB 


34-55±0-26 


l-78±0-18 


29-2-39-9 


30-8-36-4 


52 


SD 


66-3^0-41 


2-83±0-29 


57-8-74-8 


61*6-70*0 


4-3 


A-P 


36-4 


No variation recorded. 






AM 


49-l±0-36 


2-20±0-25 


42-5-55-7 


42-0-50-4 


4-4 


Ali 


53-8±0-44 


2-80±0-31 


45-4-62-3 


47-6-58-8 


5*2 


PL 


60-2±0-39 


2-48±0-27 


52-8-67-6 


53-2-64-4 


4-1 


Sens, 


61»7±0-28 


l-33:t0<20 


57-7-65-7 


58-8-64-4 


2-1 



Trombicltla anous ("Wharton 1945). 

Acariscus anous Wharton 1945, J. Parasitol., 31, (6), 403, 

Plate 14, fig. H-J. 

This species was recently described by Dr. G. W. Wharton for specimens 
found on the wings of the Tattler, Anous stolidm and Noddy, Ueteroscelus 
incanus from Ypao Point, Guam, 27 May, 1925. I am gratefully indebted to 
him for the presentation of two paratypes from which the following details and 
the figures are taken. 

The body shape is ovoid. The length of engorged specimens to 390/*, and the 
width to 300/*. The scutum is sparsely and finely punctate, and as figured, with 
the anterior margin strongly sinuate and posterior margin moderately deep 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 99 

behind line of PL ; sensillae bases distinctly in advance of PL. Eyes 2 -f 2, on 
distinct ocular shields, posterior the smaller. Chelicerae with only the usual 
apical tricuspid cap. Galeal setae nude. Palpi stout, tibial claw rather strongly 
curved and bifurcate j seta on palpal femur with 5^6 branches, fine ; on genu with 
2-3 short branches: all setae on tibia apparently nude. Dorsal setae long and 
slender, with fine ciliations, to 70^ long, and arranged ca. 2.8.8.6.4.4.2 — 34 in 
number. Ventrally with the usual pair of branched setae on gnathosoma, a 
single seta on each coxa, a pair between coxae I and between coxae III and 
thereafter approximately 8.8.8.6,4.4.2. to 40/x long. Tarsi of leg III with a long 
nude seta. The Standard Data are: AW 103*0, PW 118-0, SB 44-0, ASB 
36-0, PSB 23-0, SD 59-0, A-P 32-0, AM 43-0, AL 60-0, PL 85-0, Sens. 90-0. 

Trombicxjla pluvius (Wharton 1945). 

Acariscua phwius Wharton 1945, J. ParasitoL, 31, (6), 401. 

Plate 14, fig. K-M. 

Dr. G. W. Wharton has very kindly furnished me with two paratypes from 
the Noddy, Anous tenmrostris, from the Jaba River, Bougainville, 9 Jan., 1945, 
from which the following details and the figures are drawn. 

Shape of body ovoid. Length (engorged) 450/x, width 360ji. Dorsal scutum 
sparsely and finely punctate, but smaller than in anous and with the posterior 
scutai margin very shallow behind line of PL j sensillae bases slightly anterior of 
PL. Eyes 2 + 2, on distinct ocular shields and posterior the smaller. Chelicerae 
simple, with only the apical tricuspid cap. Galeal setae nude. Palpi stout, with 
bifurcate, strongly curved tibial claw ; seta on palpal femur with short branches ; 
all setae on genu and tibia apparently nude. Dorsal setae long, finely ciliated, 
to 70ja long, and arranged approximately 2.10.8.8.4.4.2 = 38 in number. Vent- 
rally with the usual pair of branched setae on gnathosoma, one on each coxa, a 
pair between coxae I and between coxae ITI and thereafter approximately 
10.8.8.8.6.4.2 = 46. Tarsi of leg III with a long nude outstanding seta. The 
Standard Data are: AW 86-0, PW 90*0, SB 36-0, ASB 29-0, PSB 16-0, SD 
45-0, A-P 30-0, AM 50*0, AL 40*0, PL 72-0, Sens. 60-0. 

Loc. Cape Torokina, Empress Augusta Bay, Bougainville Island, Austra- 
lian Mandate; Ypao Point, Guam, Mariana Islands. 

Hosts. Pluvicvlk do-minica. Anous tenuirostris, Anous stolidus, *Hetero- 
scelus incmus. 

Remarks, This and the preceding species are very close together in the 
curved bifurcate palpal claw and the general form of the scutum, but differ in 
the size of the scutum and in the number and arrangement of DS. 



100 Records of the S.A, Museum 

Tbombicula. lygosomoides sp. n. 

Plate 15, fig. A-D. 

Description of Lwrvae. Shape oval. Length unfed, 345/a; width 240/i. 
Dorsal scutum lightly and irregularly punctate, much wider than long, anterior 
margin slightly concave, posterior margin flattened medially and laterally 
rather angular (see PI. 15, iig. A) : sensillae bases only slightly in advance of 
line of PL. Eyes 2 -(- 2, on distinct ocular shields, posterior eyes the smaller. 
Chelicerae with only the usual tricuspid cap, Galeal setae nude. Palpi stout, 
tibial claw bifurcate, setae on palpal femur, genu and tibia all nude j tarsi with 
basal and subapical sensory rods, and 5 or 6 ciliated setae, one of which is longer 
and stronger than the others and over-reaches the tip of the tibial claw. Dorsal 
setae 26(28) in number, arranged 2.6 (8). 6,6.4,2, to 55/* long. Ventrally with 
the usual pair of branched setae on gnathosoma, a ciliated seta on each coxa, a 
pair between coxae I and between coxae III, and thereafter 4.2.2.2, to 25/u. long. 
Legs: I 300/a long, II 280/i, III 30%; tarsi I and II with the usual dorsal rod- 
like seta, and tarsi III with a long outstanding nude seta. 

The Standard Data, in microns derived from seven specimens are : 



AW 


Mean 
l02-7±0-86 


Standard 
Deviation 

2-29±0-61 


Theoretical 

Range 

95-85-109.55 


Observed 
Bange 

100-0-106-0 


Coeff . of 
Variation 

2-25 


PW 


123-0±0-64 


l-68±0-45 


118-0-128-0 


120-0-124-0 


1-4 


SB 
ASB 


47-6±0-61 
29-0 


l-62±0<43 43*8-52-4 
No variation recorded. 


45-0-50-0 


3-4 


P8B 


37-l:£0-67 


l-77±:0.47 


31-8-42-4 


35-0-40-0 


4-8 


SB 


65-9:t0-67 


l-77±0-47 


€0-6-71*2 


64-0-69-0 


2-7 


A-P 


33.9±0-55 


l<46rt0.31 


29-5-38-3 


32-0-35-0 


4-3 


AM 


38-3±0-84 


2 -07±0- 59 


32-1-44-5 


35-0-40-0 


5-4 


AL 


47-7:fc0-80 


l-97±0.57 


41-8-53 -6 


46-0-50*0 


3-1 


PL 

Sens. 


53-l±l*18 
64-0 


3-ll±0-83 
Only one deter 


43-8-G2-4 

ruination. 


50-0-56-0 


5<8 



hoc. Seven specimens from the lizard Lygosoma Stanley anum f from the 
Wau-Edie Creek Rd., T.N.G., 11 Sept., 1944, collected by the late Dr. Consett 
Davis. 

Bematks. In the bifurcate tibial claw this species is a Eutrombicula in 
Ewing's original definition, but in the number of dorsal setae it comes into 
Acariscus as separated later by the same author, both of which genera are now 
considered synonymous with Trombicula. 

It may be separated from its allies as in the key to species and on the form 
of the dorsal scutum. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



Trombicula (Trombicula) samboni Worn. 1939. 

Trombicula sanibom Womersley 1939, Tr. Roy. Soc. S. Aust, 63 (2), 153; 

ibid, 1943, 67, (1), 95. 
Tromlicula hirsti Hirst 1929, Ann. Mag. Nat. Hist, (10), 3, 564-, nee Sambon 

1927; Womersley 1934, Rec. S. Aust. Mus., 5, (2), 212, nee Sambon 1927. 

Plate 15, fig. E-H. 

This species is distinguished from T. hirsti Sambon 1927 in the number of 
DS. T. samboni is a common species in the south-east of South Australia, 
while hirsti is a Queensland and New Guinea species. 

In the new key to species it comes close to lygosonwictes sp. n. but is easily 
distinguished by the shape of the dorsal scutum, and the setae on the palpal 
femur and genu, and ventrally on the tibia, being branched. 

From the following species, T, (N.) sarcina (Worn. 1944), which is also a 
Queensland species, it differs in having only 26, instead of 28 DS, and more 
particularly in that in sarcina the posterior 12 DS are not in regular transverse 
rows, but arranged in tw T o lateral clusters of 6 each, which have the counter- 
part on the ventral surface. 

The following Standard Data in microns is derived from 20 specimens : 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


87-8±0-87 


3.90+0.61 


76 -1-99.5 


83-0-96-0 


4.4 


PW 


97-7±0-83 


4 -18±0. 66 


85.2-110-2 


93-0-108-0 


4-2 


SB 


39-7±0-50 


2-24±0-35 


33-0-46-4 


37-0-45-0 


5-6 


ASB 


39'05±0-44 


, l-96±0-31 


33*15-44.95 


35- 0-42-0 


5-0 


P8B 


29-2±0-32 


l-44±0-23 


24.9-33.5 


27-0-33-0 


4-9 


8D 


68-2:±0-63 


2-81±0-44 


59*8-76-6 


62*0-74-0 


4-1 


A-P 


31-6:1:0 -30 


1-36:1:0 -21 


27-5-35-7 


29*0-33-0 


4-3 


AM 


40-3:t0-32 


1-18±0.22 


36-8-43-8 


38-0-43-0 


2-9 


AL 


45-2±:0-48 


2-06±r0-34 


39.0-51-4 


43-0-50-0 


4-5 


PL 


52-l±0-43 


l-92±0-30 


46.3-57-9 


50-0-56-0 


3-7 


Sena, 


72-2:1:0-95 


3. 29+0-67 


62-3-82. 1 


68-0-80*0 


4-6 



Trombicula (Neotroc&ibicula) sarcina (Worn. 1944). 

Trombicula sarcina Womersley 1944, Tr. Roy. Soc. S, Aust., 68, (1), 95; Gill, 
Monle and Riek, 1945, Aust, Vet. J., 32, fig. 

Plate 15, fig. I-L. 

This is a very striking species, in that the posterior dorsal and ventral setae 
are arranged into two lateral clusters. It is an important economic species, for 
although its natural host is the Grey Kangaroo Macropns yigwiteus (Zimmer- 



102 Records of the SA. Museum 

maim), in the Clermont district of Queensland it has been found to cause a 
serious "Trombidiosis" of sheep. (See Gill et al, A. Vet. J., 1945, p. 22). 

In the revised key to species it comes next to T, (T.) saniboni Worn, from 
South Australia, but it is at once distinguished by the posterolateral clusters 
of dorsal setae. It is also related to T. (?r.) maeropus (Worn. 1936) and 
2\ (?jP.) riot (Gunther 1939) but apart from other minor differences the pos- 
terior dorsal setae in these are not in two clusters, and the total dorsal setae 
are more in these two species. 

The following Standard Data are derived from 9 specimens of the original 
material ; 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Bange 


Variation 


AW 


83-3±0-72 


2-18±0-51 


76 -8-89 -8 


80-0-86-0 


2-6 


PW 


92-4+:l-16 


3-47:fc0-82 


82 -0-102- 2 


86-0-960 


3-7 


SB 


40-0±0-41 


l-22±0-29 


36-3-43-7 


38-0-41-0 


3-0 


ASB 


27-0 


No variation recorded. 






PSB 


34-9:t0-36 


l-09±0»26 


31-6-38-3 


32-0-37-0 


3-1 


SD 


62-0±0-43 


l»32:tO»Sl 


58-0-66-0 


59-0-64-0 


21 


A-P 


33«5±0-38 


l-13±0-27 


30-1-36-9 


32-0-35-0 


3-4 


AM 


31-B±0-71 


2-13±0-50 


25-1-37-9 


29-0-33-0 


6-8 


AL 


40-l±0-48 


l-45±0-34 


35-7-44-5 


38-0-42-0 


3-6 


PL 


46-l±0«82 


2-47±0-58 


38-7-53-5 


43-0-50-0 


5-3 


Seas. 


64-3±l-66 


4 -08:1:1 -18 


52-1-76-5 


60-0-70-0 


6-8 



Trombicula kanzalwanensis sp. n. 

Plate 21, fig. H-L. 

Description of Larvae. Shape ovoid. Length (unengorged) 260/1, width 
182/a. Scutum finely punctate, shaped as figured with deep, evenly rounded 
posterior margin; SB slightly behind line of PL, Eyes 2 + 2, on ocular shields, 
posterior the smaller. Palpi with bifurcate tibial claw; setae on femur and 
genu strongly branched, on tibia only the ventral branched. Chelicerae simple, 
with only the apical tricuspid cap, Galeal setae nude. Dorsal setae strongly 
ciliated, ca. 52 in number, arranged 3 humeral on each side to 53/t long, then 
10.2 (lateral). 8.2.10.8.4.2, to 42/a long. Ventrally with a pair of branched setae 
on maxillae, one on each coxa, a pair between coxae I and between coxae III, 
and thereafter ca. 12.8.6.6.6.4, to 45/i long. Coxae - finely punctate. Legs: 1 270^ 
long, II 240/z, III 280jx ; coxae I and II w T ith dorsal sensory rod ; tarsi III with 
a long nude outstanding seta; telofemur of leg I with a pair of very long 
straight outstanding ciliated setae ; telofemur of legs II and III with one such 
seta. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 103 

The Standard Data for the type and one paratype are: 

AW PW SB ASB PSB SD A-P AM AL PL Sens. 

Type 70-0 89-6 30-8 280 28-0 56-0 25«2 42*0 42-0 56 70 
Para- 
type 75-6 95-2 33-6 28-0 28-0 56-0 25-2 42-0 42-0 56-0 70-0 

Loc. and Host. The type and one paratype from rats, Kanzalwan, India, 
Oct. 10-llth, 1946 (S. L. Kalra). 

Remarks. A characteristic species on the humeral setae, and the long stiff 
outstanding ciliated setae on telof emora of legs. 

Tbombicula macropus Worn. 1936. 

Trombicula macropus Womersley 1936, J. Linn. Soc, London (ZooL), 40, (269), 
112; Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust, 67, (1), 97; 
Sig Thor and Willmann 1947. Das Tierreich, Lfg., 71b, 272. 

Plate 16, fig. A-D. 

As stated above, this species is closely related to T. (N») sareina (Worn.) 
and to T. (IT.) rim (Gunther), differing from both in the number of dorsal 
setae as given in the key, and from the former in that the posterior dorsal setae 
are not in two groups. It was originally described from specimens found in 
the groin of a wallaby, Mavropus sp., from Darwin, Northern Territory of 
Australia. No fresh material has come to hand and the following Standard 
Data is given from 5 specimens of the original material : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Bange 


Coeff . of 

Variation 


AW 


83-4:±1.03 


2*30±0*73 


76*5-90-3 


80-0-85-0 


2-7 


PW 


89- 8+0 -73 


1.64rfc0.52 


84-9-94-7 


88-0-91-0 


1-8 


SB 


36-0±0-63 


1-41±0'45 


31-8-40-2 


34-0-37-0 


3-9 


ASB 


27.0±0.63 


l-41:±0-45 


22-8-31-2 


25-0-29-0 


5-2 


PSB 
SD 


34.2±:0-49 
61-2±0-89 


1-09^0-35 
l-99±0-63 


30-9-37-5 
55-3-67-1 


33-0-35-0 

60-0-64-0 


3-2 
3-2 


A-P 


31-6±0-40 


0-89±0-28 


28-9-34-3 


30-0-32-0 


2-8 


AM 
AL 


35-7±0-77 
40-0 


l-15±0-54 32-2-39-2 
No variation recorded. 


35-0-37-0 


3-2 


PL 
Seni. 


54-0 ±1-09 
72-0 


2-50:fc0-77 46-7-61-3 
No variation recorded. 


50-0-56-0 


4*5 



104 



Records of the S.A. Museum 
Trqmdicula bioi Gunther 1939, 



Trombiada Hoi Gunther 1939, Proc. Linn. Soc. N.S.W., 64, (1-2), 80; Womers- 
ley and Heaslip 1943, Tr. Roy. Soc. S. Aust, 67, (1), 91. 

Plate 16, fig. E-H. 

Although closely related to the preceding species, in that the posterior 
dorsal setae are somewhat clustered together, this species differs in the total 
number of dorsal setae and the size of the dorsal scutum. 

The palpi are stout, and the bifurcate tibial claw rather short. The seta on 
the palpal femur is strongly branched, on the genu and the dorsal and lateral 
on the tibia are nude, the ventral tibial seta is branched. The chelicerae has 
only the apical tricuspid cap and the galeal setae are nude and strong. The 
posterior dorsal setae are stronger and more strongly ciliated than the antei'ior 
ones. Tarsi III has a long nude outstanding seta. 

In addition to the type and 3 paratypes reported on by Womersley and 
Heaslip 1943, an additonal 9 specimens collected from the mound of a bush 
turkey at Dobodura, N.G., 13 July, 1944, by Maj. G. M. Kohls have now been 
seen. 



The Standard Data in microns derived i 


from these 9 


specimens and 


two pars 


types are 


as follows: 














Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Hange 


Variation 


AW 


104-4±0-D3 


3-09±0-66 


95-1-113-7 


100-8-112-0 


2-9 


PW 


117-l:±l-26 


4*]7±0'89 


104 -6-129 -6 


112-0-123-2 


3*6 


SB 


55- 75:t0»59 


1-96+0*42 


49-9-61-6 


53-2-58-8 


3*5 


ASB 


33-85±0-25 


0-84+0*18 


31-3-36-4 


33-6-36*4 


2-5 


P8B 


34-l±0-34 


1*13+0-24 


30-7-37-5 


33-6-36-4 


3-3 


BD 


68 • 0+0 < 39 


1-31:1:0 -28 


64-1-71*9 


67-2-70-0 


1-9 


A-P 


33-6±0-38 


1-25+0-27 


29-8-37-4 


30-&-36-4 


3-7 


AM 


60-7:tl-16 


3-49+0-82 


50-2-71-2 


56-0-64-4 


5-7 


AL 


65-25±l-51 


4*76+1-07 


50-95-79-75 


61-6-72-8 


7-3 


PL 


67- 5+1- 48 


4-92+1-05 


52-7-82-3 


61-6-75-6 


7-3 


Sens. 


68* 4:1:1 • 03 


2-73+0-73 


60-2-76-6 


64-4-70-0 


4-0 



Trombicula coluberina (Radford 1946). 
Fonsecia coluberina Badford 1946, Proc. Zool, Soe. London, 116, (2), 249. 

Plate 16, fig. I-L. 

In 1942 (Parasitology, 34, 55-81) Radford erected the genus Fonsecia for 
the two South American species Trombicula travassosi Fonseca 1935 and 
T. ewingi Fonseca 1932, citing Eivingi as the type. The generic separation was 
based only on the fact that in both these species the AL setae and in travassosi 
also the AM seta, are short and stumpy and nude. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 105 

In the writer's opinion these features are insufficient to separate these 
species generically from Trombicula 8.1., and until the adults and nymphs are 
known even subgeneric separation seems undesirable. 

In 1946, Radford described and ascribed to his genus a third species, 
coluberina from a Copper-headed rat snake (Coluber rad/iatus Schlegel) from 
Imphal, Manipur State, India, and also recorded the same species from a cobra 
(Naia tripudians fasciatus Gray) from Imphal. 

As in emngi the AM and PL scutal setae are long and filamentous, but are 
stated to be "unfeathered." The dorsal scutum is longer than wide, with the 
posterior margin evenly rounded. The eyes are 2 + 2, but very small. No 
details of the palpi are given. The dorsal setae, similar to the scutal setae, are 
nude; they number 32, to 30/l long and are arranged 2.6.8.6.6.4. The ventral 
setae are also simple, one on each coxa, a pair between coxae I and between 
coxae III and thereafter ca. 50, posteriorly the Jongest Apparently no long 
nude seta on tarsi III. 

The Standard Data as given by Radford are: AW 68*0, PW 83-0, SB 
26-0, ASB 31-0, PSB 29-0, SD 60-0, A-P 26-0, AM 51-0, AL 20-0, PL 51*0, 
Sens. 68-0. 

A paratype kindly given to me by Dr. Radford has the following Standard 
Data: AW 67-2, PW 81-2, SB 28-0, ASB 25-2, PSB 30-8, SD 62-0, A-P 25-2, 
AM 53-2, AL 16-8, PL" 47 -6, Sens. — . Prom this specimen it can be seen that 
the scutal and dorsal, as well as the ventral setae, are not quite nude but are 
furnished with short sparse and indistinct barbs. The paired setae on the 
maxillae are shortly but distinctly branched. The chelicerae are non-serrate 
and the galeal setae fine, short and nude. The palpal claws are bifurcate; the 
seta on palpal femur finely and indistinctly barbed, the setae on palpal genu 
and tibia all nude. The dorsal and ventral setae are arranged as in the type 
description. 

All legs 7-segmented; I 300/* long, II 270/*, III 300/*; tarsi I and II with 
usual dorsal rod, III without any long outstanding nude seta. The specimen, 
only partially engorged, is 365// long and 260/* wide. 

TROMBicuiiA batui Philip and Traub 1950. 
J. Parasitol., 1950, 36, (1), 29-32, fig. 1. 

Plate 17, fig. A-C. 

This very interesting species was described by Philip and Traub from a 
number of specimens from bats Eonycteria spelaea in the Batu Caves, near 
Kuala Lumpur, Federated Malay States. 



106 Records of the S.A, Museum 

The dorsal scutum is very small, not much wider between PL than it is 
long and is coarsely punctate ; the posterior margin forms a shallow angle,- AM 
is slightly in front of AL, and the sensillae bases are slightly nearer to PL than 
to AL and anteriorly with strongly defined crests* the sensillae are very char- 
acteristic, being nude, but with a single fork at midlength. Chelicerae simple. 
Galeal setae nude. Palpal claw bifurcate- seta on femur with 3 or 4 short 
branches, on genu 1-branched, on tibia all nude. Dorsal setae short, to 26/*,. 
<?a. 36 in number and arranged 2.8.8.6.6.4.2, with short ciliations or branches. 
Ventrally, a pair of branched setae on maxillae, a ciliated seta on each coxa, a 
pair between coxae I and between coxae. Ill and thereafter ca. 40, the posterior 
setae nearly as long as the dorsal. No long nude seta on tarsi III. 

The Standard Data treated statistically from those given by the authors for 
the type and 4 paratypes are : 





Mean 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Bange 


Coeff . of 
"Variation 


AW 


39-8±l-83 


4*083:l-29 


27-6-62-0 


36-O-45-0 


10-2 


PW 


48-23:1-9 


4-25±l-34 


35-5-60-9 


45-0-54-0 


8-8 


SB 


18-6±0-98 


2*193:0-69 


12-0-25-2 


16- O-21-0 


11-8 


ASB 
PSB 


19-6±0-81 


Not given. 
1-823:0 -54 


14-2-25-0 


18.0-22.0 


9-3 


SD 
A-P 


24-7±0-73 


Not given. 
l-64i:0.52 


19-&-29-6 


23-0-26-5 


6*6 


AM 


27 -43:0 -40 


0*89±0*28 


24-7-30-1 


26-0-28-0 


3-2 


AL 


23-2±0-73 


l-64±0-52 


18-3-28-1 


22*0-26-0 


7-1 


PL 
Sens. 


30-03:1-64 
39»6±l-2l 


3 -67311 '16 

2-70±0'85 


15-9-44-1 
31-5-47-7 


27-0-34-0 
37-0-44-0 


12-2 
6-8 



The possibility of this species being the true larvae of Trombicula minor 
Berl., when the larvae and adult have been correlated, is discussed by the authors. 

Trombicxila ( ? NEOTROMBicuiiA) munda (Gater 1932). 

Trombicvla munda Gater 1932, Parasitol., 24, 143-174; Womersley and Heaslip 
1943, Tr. Roy. 8oc. S. Austr., 67, (1), 76. 

Trombicuto (Euirombicula) nmnda, Sig Thor and "Willmann 1947, Das Tier- 
reich, Lfg. 71b, 289. 

Plate 17, fig. D-F. 

This species was originally described from specimens from Rattus rathu 
diardi (Jentink) from Kuala Lumpur, Federated Malay States. Gater also 
recorded it from B. m/ulleri vaUdas (Miller) and R. mataisa Kloss, from Sungei 
Buloh, F.M,S. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 107 

It is very close to the following species, but differs in the Standard Data 
and the sensillae as given in the key to species. The sensillae in munda have 
only 2 or 3 distal branches, whereas in spicea they have numerous ciliations 
distally, appearing almost bushy. 

Two specimens of this species in the South Australian Museum and received 
from Gater were measured and the Standard Data given by Womersley and 
Heaslipl943. 

Since then I have received a few specimens from Dr. S. H. Jayewickreme, 
of Colombo, and taken from Rattus rattus kandiyanus, at Embilipitiya, Ceylon, 
Jan., 1945 (Tube 13). 

The Standard Data for 5 specimens are as follows : 







Standard 


Theoretical 


Observed 


Ooeff, of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


35-7±0-70 


l«40±0-49 


31-6-39-9 


33-6-36-4 


3-9 


PW 


51-8±1-81 


3-61^:1.28 


41-0-62-6 


47-6-56-0 


7-0 


SB 


15-l±0-69 


l-53±Q-48 


10-5-19-7 


14-0-16-8 


10-2 


ASB 


21-8±0»58 


l-25±0*39 


18-1-26-5 


19-6-22-4 


5-7 


PSB 


17-4±0-56 


1-25:^0-39 


13-7-21-1 


16-8-19-6 


7-2 


SB 


39-2 


No variation recorded. 






A-P 


20-73:0.69 


3»53±0-48 


16-1-25-3 


19*6-22-4 


7*4 


AM 


25-2 


No variation recorded. 






AL 


18-9±G-70 


l-40±0-49 


14- 7-23-] 


16-8-19-6 


7-4 


PL 


27'4±0-56 


1»25±0.39 


23-7-31-1 


25-2-28-0 


4-5 


Sens. 


28-0 


Only one determination. 







This species is here placed in the subgenus Neotrombicida on its close homo- 
logy with 3F. (N.) spicea which has been reared to the nymph. 

Trombicula (Neotrombicula) spicea (Gater 1932). 

Trombicula spicea Gater 1932, Parasitol., 24, 143-174; Womersley and Heaslip 
1943, Tr. Roy. Soc. S. Austr., 67, (1), 78. 

Trombicula (Eutrombicula) spicea, Sig Thor and Wilimann 1947, Das Tier- 
reich, Lfg. 71b, 288. , 

Eutrombicula n. sp. "A." Lawrence in MS. In Audy 1947. i{ Scrub Typhus 
Investigations in S,E. Asia, Pt III. Appedix 7." AMD. 7. War Office, 
London. 

Plate 17, fig. G-I. 

Closely related to the preceding species, and differing therefrom as given 
under that species and as in the key. 

The species was described from specimens from Rattus maJaisia Kloss, from 
Sungei Buloh, Selangor, Federated Malay States. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 109 

thereafter 6.4.4.4,4.4.2, to 43/i long. Legs- I 241,4 long, II 214/*, III 268/*; 
tarsi I and II with dorsal sensory rod, III without any long nude seta. 
The Standard Data of two larvae and one larval pelt* are s 



AW PW 


SB 


ASB 


PSB 


SD 


A-P 


AM 


AL 


PL 


Sens. 


60-3 80-4 


30-15 


23-45 


13-4 


36-85 


20-1 


43-55 


36-85 


63-65 


mmfm 


56-95 80-4 


30-15 


23-45 


10-05 


33-5 


201 


43-55 


36-85 


60-3 


_ 


*53-6 — 


30 15 


23-45 


10-05 


33-5 


20-1 


43-55 


33-5 


63-65 


53-6 



Loc. and Hosts, Described from 2 larvae and 1 larval pelt 1 * from Battus 
bower si from Ulu Langat Forest Reserve, Selangor, F.M.S., 13th June, 1950 
(Scrub Typhus Research Unit). 

Remarks. In general morphological features this species is closely related 
to T. anoxic and T. pliiviw, but differs from these in the Standard Data and 
number of dorsal setae. It also differs from all other species of TrombicuXa 
B.l. in that the sensillae gradually thicken to the apex, but can hardly be re- 
garded as lanceolate. 

Trombicula pabmifeka sp. n. 

Plate 17, fig. J-M. 

Description of Larvae. Shape broadly oval. Length (partially engorged) 
to 325/*, width to 235/i. Scutum shield-like, not much wider than long, posterior 
margin somewhat obtusely angular with the sides conxex but at extreme point 
forming a pronounced tip; surface finely punctate; setae fine and shortly 
ciliated, PL the longest, AL the shortest; sensillae filamentous with ciliations 
distally ; sensillae bases wide apart, slightly nearer to line of PL than to line of 
AL. Eyes 2 -f- 2, posterior the smaller. Chelicerae strongly hook-like but with 
only the apical cap. Galea! setae nude. Palpi elongate, both femora and genu 
cylindrical, tibia very small and provided with two or a bifurcate claw, the 
prongs of which are short and stumpy, and do not reach the tip of palpal tarsus; 
setae on femur and genu fine and finely ciliated, on tibia all 3 apparently nude; 
tarsus with 4 or 5 ciliated setae. Dorsal setae fine and tapering, shortly ciliated, 
anteriorly 50/*, shortening posteriorly to 42/t, 20 in number and arranged 
2.6.6.4.2. Ventrally with paired setae on maxillae, one on each coxa, those on 
I 42/i., on II 30/*, on III short and spine-like to 14//.; between coxae I and II a 
pair of fine setae, and thereafter 6.4.2.2.2 from 30/* anteriorly to 40/t posteriorly. 
Legs all 7-segmented, I 285/x long, II 250/t, III 273/i j tarsi I and II with dorsal 
sensory rod, III without any long nude seta. 



108 Records of the S.A. Museum 

The Standard Data was given by Womersley and Heaslip 1943 for a speci- 
men received from Gater. 

Recently I have received 4 specimens from Baitus r. kandiyanus from 
Embilipitiya, Ceylon, Jan., 1945 (S. H, Jayewickreme, No. 314 A.B.C.D.) which, 
although the sensillae are missing in all specimens, agree in the Standard Data 
with spicea. The Standard Data for these specimens are : 





Mean 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Bange 


Coeff.of 
Variation 


AW 
PW 


46-9±:l-76 
58-8±l«66 


3-52:tl-24 
3-32±l-17 


36-4-57-4 
48-9-68-7 


42-0-50-4 
56-0-61*6 


7-5 
5-6 


SB 
ASB 
PSB 
8D 


17-5±0-70 
25*2 

17-5±0-70 
48 '7:1:0 -70 


l-40±0-49 13-3-21-7 
No variation recorded. 
l-40±:0-49 13*3-21-7 
l-40±0-49 38-5-46-9 


16-8-19-6 

16-8-19-6 
42-0-44-8 


8-0 

8-0 
3-3 


A-P 

AM 


25-9±:l-34 
20 -5:1:0-93 


2-68±0-95 
l-62±0-66 


17-9-33-9 
15-7-25-3 


22-4-28-0 
19-6-22-4 


10-3 
7-9 


PL 

Sens. 


32-8±:l-34 

Missing. 


2-68±0-95 


24-8-40-8 


30-8-36*4 


8-1 



This species is here placed in the subgenus Neotrombicula, as larvae have 
been reared to the nymphal stage by Dr. S. H. Jayewickreme in Ceylon (see 
Adult Section). 

The species referred to by T. J. Lawrence as Eutr&mbicvla sp. n. "A" in 
Appendix 7 in Part III, Scrub-typhus Investigations in S.E. Asia, AMD 7, War 
Office 1947, are without doubt this species. 

Tbombicula (Neotrombicula) consubta sp. n. 

Plate 110, fig. B-H. 

Description of Larvae. Shape almost round. Length (engorged) 469^ 
width 455/*. Scutum as figured, much wider than long with SB only slightly 
behind, or practically in line with PL; posterior margin sinuous and excavate 
medially; surface finely punctate; sensillae slightly thickening from base to 
apex and with long ciliations. Eyes 2 -|- 2, posterior the smaller. Chelicerae 
non-serrate, with only the apical tricuspid cap. Galea! setae nude. Palpi as 
figured, tibial claw bifurcate ; all setae on femur, genu and tibia, except ventral 
on tibia, nude. Dorsal setae 36 in number, to 46/* long and arranged ca. 
2.8.6.8.6,4.2, shortly ciliated and tapering. Ventrally with a pair of seta on 
maxillae, one on each coxa, a pair between coxae I and between coxae III and 



110 Records of the S.A. Museum 

The Standard Data for the type and 28 paratypes are as follows : 







Standard 


Theoretical 


Observed 


Coeff . 




Mean 


Deviation 


Range 


Bange 


Variati 


AW 


63- 0+0 -29 


1-55+0 -20 


58*3-67-7 


61-6-67-2 


2-4 


PW 


73-25:±0'27 


1-48:2:0.19 


68-8-77-7 


70-0-75-6 


2-0 


SB 


36-7±:0-16 


0-85+0-11 


34-2-39-2 


364-392 


2-3 


ASB 


22-4 


No variation recorded. 






PSB 


38-3rfc0-31 


1-66+0-24 


33-3-43-3 


36-4-42-0 


4-3 


SD 


60-7±0-31 


1-66+0-24 


55-7-65*7 


58-8-^4-4 


2-7 


A-P 


29-75±0-25 


1-36+0-18 


25-7-33-8 


28-0-30-8 


4-5 


AM 


45-6:t0-43 


2- 17 ±0.31 


39-1-53*1 


42-6-47-6 


4-7 


AL 


31-85±0-46 


2-48+0-33 


24-4-39-3 


28-0-36-4 


7-7 


PL 


49-5zt0-41 


2-18+0-29 


43-0-56-0 


44-8-53-2 


4-4 


Sens, 


53* 7+0-48 


2-22+0-34 


47-0-60-4 


47-6-56-0 


4-7 



Loc. and Hosts. The type and 28 paratypes from a skink, Mabuia multi- 
fasdata, 11 Aug., 1949, from Bukit Lagong Forest Beserve, Kepong, near 
Kuala Lumpur, F.M.S. (J. K. Audy). 

Remarks. This species is remarkable in the structure of the palpi, especially 
the cylindrical femora and genu, the very small tibia, and the short stumpy 
bifid palpal claws. These features, however, do not warrant the erection of a 
genus or subgenus until the nymph or adult of the species is known* 

I am much indebted to Dr. J. R. Andy for the opportunity of studying 
the above material, as well as specimens of many other new species described 
herein. 

Tbombioula lundbladi sp. n. 

Plate 18, fig. A-D. 
Description of L&rvae* Shape broadly oval. Length (engorged) to 480/a, 
width to 420/*. Dorsal scutum as figured, with the posterior margin rather 
angular and extending well behind line of PL, surface lightly and irregularly 
pitted, SB nearer to line of PL than to AL, sensillae with 5 or 6 branches on 
distal half. Eyes 2 + 2, small, on ocular shields about 2 diams. from lateral 
scutal margins, posterior eyes about equal to anterior. Chelicerae curved, with 
the usual apical tricuspid cap. Galeal setae nude. Palpi stout, tibial claw 
bifurcate ; all setae on femur, genu and tibia nude, tarsi with curved basal and 
longer subapical sensory rods and 5 or 6 ciliated setae, one of which is longer 
than the rest and over-reaches tip of claw. Dorsal setae rather stiff and mode- 
rately thick, with short dilations, to 36/x long and arranged 2.6.6.4.2,2. Vent- 
rally with the usual pair of branched setae on gnathosoma, a single fine, 
shortly ciliated seta on each coxa, a pair between coxae I and between coxae III 
and thereafter 4.2.2.2; ventral setae between coxae 16/*, at posterior end 30/*. 
Legs: I 230/*, II 195/i, III 230pj tarsi I and II with the usual dorsal rod-like 
acta. Ill without a long nude seta. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 111 

The Standard Data in microns for 13 specimens from the type series are : 



AW 


Mean 
70-85;t0-39 


Standard 
Deviation 
l-34±0-26 


Theoretical 

Range 
66-8-74-9 


Observed 

Bange 
70-0-72-8 


Ooeff . of 
Variation 

1-9 


FW 
SB 


78-2±0*21 
41*6±:0-29 


0-78±0-15 
1-05±0*20 


75-9-80-5 
38-5-44-7 


75-6-78-4 
39-2-42-0 


1-0 

2-5 


ASB 


23-9±0-40 


l-45±0-28 


19 '6-28 -2 


22-4-25-2 


6-6 


P8B 


36-4 


No variation recorded. 






SD 


60-3±0-40 


l-45:±0»28 


56 -0-64 '6 


58-8-61-6 


2*4 


A-P 


27-6±0-29 


1*05+0*20 


24-5-30*7 


25*2-28-0 


38 


AM 


21 -5+0 -49 


l-76±0-34 


16-3-26-7 


19-6-25-2 


8-2 


AL 
PL 

Sens. 


21-l±0-40 

19-6 

42-0 


1*45±0*28 16-8-25-4 
No variation recorded. 
Only one determination. 


19*6-22-4 


6-9 



Loc. From under the scales of a dark green and striped skink from 
Hollandia, Dutch New Guinea, 8 Dec, 1944 (C, B. Philip). 

Remarks. This is a distinct and rather characteristic species in the shape 
of the dorsal scutum and in the small and fine scutal setae. It is named in 
honour of Dr. C. Lundblad, the eminent Swedish Hydrachnologist. 

Trombicula (Leptotrombidium) bodensis (Gunther 1940). 

Trombicula bodensis Gunther 1940, Proe. Linn. Soc. N.S.W., 65 (5-6) t 479; 
Womersley and Heaalip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 87. 

Plate 18, fig. E-H. 

Gunther described this species from colonies on the legs of the mouse deer 
(Tragtdus borneamis Miller) from the Bode River, near Sandakan, British 
North Borneo. 

I have recently been able to study a lot of material of this species col- 
lected by Maj. R. N. McCulloch, from the mouse deer at Kuehing, Sarawak, 
22 Sept., 1945, and also from rats at Labuan, British North Borneo, in June 
and Oct., 1945, and from Balikpapan in Dutch Borneo, 21 Aug., 1945. 

The specimens conform well with the original description and the following 
re-description is now given based on this additional material. 

Re-description of Larvae. Shape broadly oval. Length (only slightly 
engorged) 300/a, width 230/*, Dorsal scutum almost rectangular with the 
anterior and posterior margins almost straight as figured, surface lightly and 
sparsely pitted, scutal setae fairly stout and ciliated ; sensillae placed a little in 
front of a line joining PL, and with 6-7 long outstanding branches on distal 
third. Eyes 2 -f- 2, anterior the larger. Chelicerae as figured, with the usual 
tricuspid cap. Galeal setae slender and with 6-7 long branches as figured. 
Palpi stout j all setae on femur, genu and tibia except the dorsal tibial nude. 



112 Records of the S.A. Museum 

Palpal claw bifurcate. Palpal tarsus with the usual sensory rods and 5-6 ciliated 
setae. Dorsal setae, moderately thick, only slightly curved and ciliated, to 60/a 
long, and arranged 2.8.6.6,6.2(4). Ventrally, with the usual pair of gnatho- 
somal ciliated setae, a single seta on each coxa, a pair between coxae I and 
between coxae III and then approximately 10(8),4(8),4.4.4. Legs: I 180/*, 
II 125/4, III 200/*; tarsi I and II with the usual dorsal rod-like setae; no long 
nude seta on tarsi III. 

The following Standard Data in microns, is derived from 2 paratypes and 
14 recent specimens ! 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


60-2±0-33 


l-45±:0-27 


55-8-64-6 


58-8-61-6 


2-4 


PW 


68-8±0-64 


2-38±0-45 


61-7-75-9 


64-4-72-8 


3-5 


SB 


28-8:£0-35 


l-31±:0-24 


24-9-32-7 


28-0-30-8 


4-6 


ASB 


28-0 


No variation recorded* 






PSB 


14-2 


No variation recorded. 






SD 


39-2 


No variation recorded. 






A-P 


28-0 


No variation recorded. 






AM 


50-0±0-53 


1-92+0-37 


44-2-55-8 


47-6-53-2 


3-8 


AL 


43-4±0-39 


1-45.&G-27 


19*0-27-8 


42-0-44-8 


6-2 


PL 


55-6±0-82 


3-08HtO-58 


46-4-64-8 


50-4-61-6 


5-5 


Sena. 


61 -63:0-72 


l-77zt0-55 


56*3-€6-9 


58-8-64-4 


2-9 



Remarks. This species may be separated as in the revised key. In the 
bifurcate palpal claw it would fall in Euirambicula Ewing 1938, and in the 
number of dorsal setae into Acariscm Ewing 1943, but these genera are not 
now recognized as valid. 

It is here placed in the subgenus Leptotrombidium on the struetrue of the 
longitudinally divided sternum formed by the precoxal plates of leg I, in the 
nymph. 

A more recent collection from Cailoscmrus hippurus, from Kuching, Sara- 
wak, 1950, and received from Dr. J. R. Audy, has the following Standard 
Data derived from 23 specimens: 





Mean 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Bange 


Coeff. of 
Variation 


AW 
PW 

SB 


59-7^:0*39 
68 -25 ±0-39 
27-25^0-23 


1- 89+0-28 
1-88+0-28 
1-13±0-17 


54-0-65-4 
62-6-73-9 
23-85-30-65 


56-9&-63-65 

67-0-73-7 

26*8-30-15 


3-2 

2-7 
4-1 


ASB 
PSB 
SD 
A-P 


30-15 
13-4 

43-55 
27*25±0-23 


No variation recorded. 
No variation recorded* 
No variation recorded. 
1-13+0-17 23-85-30«65 


26-8-30-15 


4-1 


AM 


49-2±0-60 


2-87±0»42 


40-6-57*8 


46-9-56-95 


5-8 


AL 


38 '5:1:0 -47 


2-23±0-33 


31-8-45-2 


36-85-43-55 


5-8 


PL 

Sena. 


49-2+0-70 
67-2±0-23 


3-34+0-49 
0-86±.0*16 


39-2-59-2 
64-6-69-8 


46-9-56-95 
67*0-70-35 


6-8 
1-3 



Womersley— Asiatic- Pacific Scrub Typhus Mites 113 

Trombicula nissani Dumbleton 1947. 
Trans. Roy. Soc. N. Zealand, 76, 412. 

Plate 18, fig. I-K, 

This species was described from the type and paratype from a " discus" 
from Nissan Island (Green Is,), Territory of N. Guinea, 28 April, 1944 (coll. 
L.J.D.). The type is deposited in the South Australian Museum. 

The dorsal scutum is almost rectangular with the posterior margin only 
very slightly convex behind PL. Sensillae bases in front of PL, but very 
much nearer to line of PL than AL, Scutal setae thick and with short 
dilations or serrations, AM the shortest. Eyes 2+2, on distinct ocular shields, 
posterior eyes the smaller. Chelicerae with only the apical tricuspid cap. 
Galeal setae nude. Palpal tibial claw trif urcate (not bifid as given by Dumble- 
ton) ; setae on palpal femur and genu branched, on tibia all 3 nude. The DS 
are 28 in number, to 55/i long, and arranged 2.8.6.6.4.2. All coxae unisetose. 
Tarsi of leg III without any long nude seta. 

The Standard Data of the type and paratype are: AW 66-0, 65-0, PW 
69-0, 69-0, SB 23-0, 23*0, ASB 26-0, 26-0* PSB 13-0, 13-0, SD 39-0, 
39»0, A-P 29-0, 29*0, AM 26-0, 26-0, AL 36-0, 33-0, PL 46-0, 43-0, Sens. 

36 -0, — . (after Dumbleton). 

- 

Trombicula issHixn Sugimoto 1938. 

Trombicula isshikii Sugimoto 1938, J. Jap. Soc. Vet, Sci., 17, (1), 57-62, fig. 1-3. 

Trombicula issMlcii Womersley and Heaslip 1943, Tr. Roy. Soc S. Aust., 67 y 

(D,81. 

Plate 18, fig. L-M. 

Owing to unfamiliarity with the Japanese writing, the data given in 
Womersley and Heaslip '$ 1943 paper, and derived from Sugimoto *s paper (it 
had no summary in English or any other language) were in error. 

I have recently had the essential details translated by a Japanese student, 
and the following description is based on this translation, 

u Shape oval to circular. Colour red. Chelicerae not serrated. Galeal 
setae nude. Palpi with ciliated setae on femur, genu, and dorsally on tibia ; 
tarsi with 6 comb-like setae, tibial claw bifurcate with the upper prong longer 
than lower. Dorsal scutum trapezoidal, the front and side margins convex, 
posterior margin rounded. 

"Scutal setae fine, comb-like; the PL setae very long, AM and AL of 
equal length. SB somewhat nearer to posterior than to anterior scutal mar- 



114 



Records of the S.A. Museum 



gin. Sensillae with five branches. Eyes 2 + 2, on ocular shields, not far from 
8cutal margins, anterior eyes with a distinct cornea, posterior eyes the smaller. 
Dorsal setae small, comb-like. Total setae 42-44, with 26 on dorsum, arranged 
2.6.6.6.4.2; anterior longer than posterior. Ventral setae arranged 2.2.4.4.4.2, 
longer posteriorly than anteriorly. Legs: tarsi I and II rapidly tapering, 
coxae II and III separated, the longer leg setae serrated and comb-like. 

"Body length 0-49-0-59 mm., width 0-41-0*44 mm. Length of palpi 94/*, 
of chelae 42/*. Scutal length 55/*, width 78/*. SB 31/*, AM 58/*, AL 43/*, 
PL 74-78/*, Sens. 97/*, Tarsi I 34/*." 

Remarks. In the bifurcate palpal claw and the number of DS, isshikii 
would come into Acariscus Ewing, although the ciliated dorsal setae of the 
palpal tibia shows some relationship with the akamushi-dsliensis group. The 
convex anterior and lateral scutal margins are also unusual. No material has 
been seen. 

Type Loc. and Host. Naikosho, Shichiseiguin, Pref. Taihoku, Formosa, 
on Capella kardwicMi (Gray). Type in Taihoku Imperial University Museum. 

Tbombicula (Nbotrombicula) scinooides (Worn. 1944). 
Trombicida scincoidts Womersley 1944, Tr. Roy. Soc. S. Aust., 68, (1), 84. 

Plate 19, fig. A-E, 
This species was originally described from a series of 15 specimens from 
a skink, Lygosoma (Leiolopisma) licarinatus (MacL. 1877) from New Guinea, 
Oct. 1, 1943 (R. N. McCulloch). The specimens are now, after the lapse of 
time, somewhat more flattened, and the values of the Standard Data slightly 
greater than originally given. The newly measured values for 13 specimens 
are: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Ooeff » of 
Variation 


AW 


98*2±0-51 
113-5±0-60 


l-86±0-36 
2-17+0-43 


92-6-103*8 
107-0-120-0 


95-2-100-8 
112-0-117-6 


1-9 
1»9 


SB 


51-05:1:0-34 


1-233:0 -24 


47-4-54*7 


50-4-53-2 


2-4 


ASB 
PSB 
SD 


28-0±0-33 
26-6±0-39 
54-6-±0-54 


l-14±0-23 
1-40+0 -27 
l-93:t0-38 


24-6-31-4 

22-4-30-8 
48*8-60-4 


25-2-30-8 
25-2-28-0 
50-4-58-8 


4-1 

5-2 
3-5 


A-P 


30 -4^0-43 


1-55±:0-30 


25-7-35-3 


28-0-33-6 


5-1 


AM 
AL 


39- 9 ±0-50 
39-2±0-32 


1-742:0 '35 
1»14±0.22 


34-7-45-1 
35-8-42-6 


36-4-42*0 
36-4-42-0 


4-3 
2-9 


PL 

Seas. 


47«6:±0-32 
57- 4+0 -65 


1-14+0-22 
2-23:£0<46 


44-2-51-0 
50-7-64-1 


44-8-50-4 
56-0-61-6 


2-4 
3-9 



The chelicerae have the usual tricuspid cap only and the galea! setae are 
nude. The tibial claws of the palpi are bifurcate ; the seta on the palpal femur 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 115 

with a few short branches, often appearing nude; on the genu nude and on 
the tibia only the ventral seta is branched. There is no long nude outstanding 
seta on tarsi on leg III. 

Another population of 34 specimens from Hollandia, Dutch New Guinea, 
from various skinks, collected Noy. and Dec., 1944 by Lt-Col. C. B. Philip 
(No, 5, 6 and 7) are not different morphologically from the type series, except 
that the scutum is considerably larger, all the Standard Data being significantly 
greater at 0-1% probability, except PW, which is significantly greater at 2% 
leveL (Sens, were not compared.) 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


105«4±0-51 


2 '94:1:0. 36 


96-6-114-2 


98-0-112-0 


2-8 


PW 


116-7±0-75 


4-33±:0-53 


103 -7-129-7 


109-2-126-0 


3*7 


SB 


55 -l±Q- 32 


l-89:±:0-23 


49-4-60-8 


53-2-58-8 


3-4 


A8B 


30-8±0-26 


1-50±0*18 


26-3-35-3 


28-0-33-6 


4-8 


PSB 


28-0 


No vnmtion recorded. 






SD 


58-8+0-26 


1*50±0«18 


54-3-63-3 


56-0-61-6 


2-5 


A-P 


32-8±0-23 


1-36:1:0 -16 


28-7-36-9 


28-0-33-6 


4-1 


AM 


46 -95:2:0 -27 


l-39±0-19 


42-8-51-1 


44*8-50-4 


30 


AL 


44-0+0-28 


l-60±0<20 


39-2-48-8 


42-0-47-6 


3-6 


PL 


51-3±0*29 


1-64±0-20 


46-4^56-2 


47-6-56-0 


3-2 



Sens. 67-2 Aver, of only 2 determinations, 64-4 and 70*0. 

As discussed further on, it seems hardly justified to consider this popula- 
tion as more than a race or f onn of scincoides. 

To the typical form can be referred the following additional specimens : 
New Guinea: Nadzab, on hoots, 21 Dee., 1942 (1 spec.) ; Burma, Jan., 1944, on 

boots (2 specs.); Lae, 9 March, 1944, and 3 May, 1944 (2 specs.), 

(RJST.McC.) ; Goodenough Is., N.G., 18 Jan., 1944 on boots (D.C.S.), (4 

specs.). 
Bougainville: On a gecko, 4 July, 1944 (G. W. Wharton), Nambru, II. 44 (1 

spec). 
Philippine Is. : Mindoro, Mts. near San Juan, 11 Apr., 1945, on mottled skink 

(2 specs.) (C.B.P.) 

This species is placed in the subgenus N eotrombicida on the adult charac- 
ters. It is closely related to the two following species, which are only separated 
from it, on the very much greater differences in scutal dimensions, than occur 
between the typical populations of semcaides and its form from Hollandia, 
D.N.G. 

The Standard Data of the population of scincoides from Hollandia are 
approximately intermediate between those of typical scincoides and those of 
hohhi, and while all three populations are widely significantly different it is 
possible they may be only populations of the one species. 



116 Records of the S,A. Museum 

Trombicula ( ? Neotrgmbicula) kohlsi (Worn. 1944). 
Trombicula hohhi Womersley 1944, Tr. Roy. Soc. S. Ausi, 68, (1), 87. 

Plate 19, fig. F-G. 

Originally described from a series of 7 specimens collected on boots 
amongst Kunai grass, Buna area of New Guinea, 1943 (G. M. Kohls), this 
species differs from ft (A 7 .) scincoides (Worn.) and its form from Hollandia 
only in the very much larger dimensions of the Standard Data. 

The Standard Data of these specimens as remeasured are : 







8tandard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


110-8±:1»03 


2 • 73+0* 73 


102- 6-119*0 


106-4-114-8 


2-5 


PW 


124*4±Q-56 


1- 49+0 -40 


119 -9-128 -9 


123- 3-126- 


1-2 


SB 


62-4±0-80 


2 12+0-56 


56< 1-38.7 


58-8-64*4 


3-4 


ABB 


34-8±0-56 


1-49+0-40 


30-3-39*3 


33-6-36-4 


4-3 


P8B 


33-6 


No variation recorded. 






SD 


68-4±0-56 


1-49+0-40 


63-9-72-9 


67-2-70-0 


2-2 


A-P 


36-8±:0.40 


1-06+0-28 


33-6-40-0 


36-4-39-2 


2-8 


AH 


45-6±0-52 


1-37+0-36 


41-5-49-7 


44-8-47-6 


3-0 


AL 


47'6±0-61 


1-61+0-43 


42-8-52-4 


44-8-50-4 


34 


PL, 


55-6±0-40 


1-06+0-28 


52-4-68-8 


53-2-56-0 


1-9 


Sens. 


63.6+0 -52 


1-37+0-36 


59-5-67'7 


61-6-64-0 


21 



The relationship of this species to scincoides and tovelli is discussed below. 

Trombicula ( f Neotrombicuija) TOVELLr sp. n. 

Plate 19; fig. H-J. 

Description of Larvae. Shape an elongate oval. Length, partly engorged 
io 420/*, width 330/*. Dorsal scutum as figured with finely striate-punctate sur- 
face, anterior margin sinuous, posterior evenly curved and not very deep, SB 
nearer to line of PL than to AL. Eyes 2 -f- 2, on well developed ocular shields, 
the posterior eyes the smaller. Chelicerae w T ith the usual apical tricuspid cap. 
Palpi stout, with bifurcate tibial claw. Seta on palpal femur with a few fine 
short indistinct branches; on genu nude; on tibia dorsal and lateral nude, 
ventral branched- tarsi with basal and subapical rods and 5-6 ciliated setae, 
one of which over-reaches tip of tibial claw. Galeal setae nude. Dorsal setae 
stiff, with short ciliations, 20-22 in number and arranged 2.6.6.4.2(2) to 35/* 
long. Ventrally with the usual pair of branched setae on gnathosoma, a 
filiated seta on each coxa, a pair between coxae I and between coxae III, and 
thereafter 6.2.2.2, to 30/* long. Legs: I 240/* long, IT 225/*, III 240/*; tarsi I 
And II with dorsal sensory rod, and III without any long nude seta. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



117 



The Standard Data are as follows: 



AW 

PW 

SB 

ASB 

PSB 

SD 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
77-9+0-47 
89-l±0-47 
40-1+0-93 
27-1+0-59 
25-7+0-47 
52-7+.0-47 
26-1+0-59 
29-4+0-63 
30-8 

41-5+0-47 
56-0 



Standard 
Deviation 

1-14+0-33 

2-75+0-33 

2 -28+0 -66 

1-45+0-42 

1-14+0-33 

1-14+0-33 

1-45+0-42 

1-53+0-44 

No variation 

1-14+0-33 



Theoretical 
Range 

74-5-81-3 

85-7-92-5 

33-25-46-95 

22-8-31-4 

22-3-29-1 

49-3-56-1 

21-8-30-4 

24*8-34-0 

recorded. 

38-1-44-9 



Observed 
Range 

75-6-78-4 

84-0-92-4 

36-4-42-0 

25-2-28-0 

25-2-28-0 

50-4-53-2 

25-2-28-0 

28-O-30-8 

39-2-42-0 



Coeff. of 
Variation 

1>5 
1-3 
5-7 
5-3 
4-4 
2-2 
5-5 
5-2 

2-7 



No variation recorded. Two determinations only. 



Another population 
of Darwin, N.T., Aust, 
Standard Data i 



of eight specimens from a lizard, taken 60 miles south 
30 May, 1943 (R t V. Southeott) has the following 



• 




Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


72*8±0-53 


1-49+0-37 


68-3-77-3 


70-0-75-6 


2-0 


P# 


86-l±0-48 


1-35+0-34 


82-05-90-15 


84-0-86-8 


1-6 


SB 


37-1+0-48 


1-35+0-34 


33-05-41-15 


36-4-39-2 


3*6 


A8B 


25-2 


No variation recorded. 






PSB 


25-2 


No variation recorded. 






SD 


50-4 


No variation recorded. 






A-P 


25-2 


No variation recorded. 






AM 


26-25±0-51 


1-45+0-36 


21*9-30-6 


25-2-28*0 


5-5 


AL 


30-45+0-63 


1«79±0'45 


25-05^35-85 


28-0-33-6 


5-9 


PL 


39-2+0-53 


1-49+0-37 


34-7-43-7 


36-4-42-0 


3-8 


Sens. 


53-2+0-61 


3-61+0-43 


48-4-58-0 


50-4-56-0 


3-0 



A third population 
Eylandt, Arnhem Land 
greater than the type 
follows : 



from a skink, Sphenomorphus taeniolatus from Groote 
Expedition, 1945, has the Standard Data significantly 
series. As measured for 11 specimens, these are as 



AW 

PW 

SB 

ASB 

PSB 

SD 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
83-5+0-51 
96-5+0-69 
42-5+0-63 
26-7+0-25 
27-7+0-25 
54-5±0-44 
28-25+0 25 
28-5+0-34 
33-6 

41-2+0-55 
58-45+0-82 



Standard 
Deviation 



69+0-36 
29+0-49 
10+0*45 
46+0-31 
84+0-18 
46+0-31 
84+0-18 
13+0-24 
No variation 
1-81+0 -38 
2-34+0-58 



Theoretical 
Range 

78-4-88-6 
89-7-103-3 
36-2-48-8 
22-3-31-1 
25-2-30-3 
50-1-58-6 
25-75-30-75 
25-1-31-9 
recorded. 

35-8-46-6 
51-45-65-45 



Observed 

Range 

81-2-86-8 

92-4-100-8 

39-2-44-8 

25-2-28-0 

25-2-28-0 

53-2-56-0 

28-0-30-8 

28-0-30-8 

39-2-44-8 
56-0-61-6 



Coeff. of 
Variation 

2-0 



4-4 
4-0 



118 



Records of the S.A. Museum 



Morphologically, tovelli and its two other populations, do not differ from 
sovncoides except in the very much smaller size of the scutum, the significance 
of which is discussed below. 

Comparison of the Species T, scincoides Worn., kohlsi Worn. 

and tovelli sp. n. 

In trying to evaluate specifically the differences in the Standard Data of 
these three species and their several populations, resort has been made to 
showing the Mean, Range of Mean as expressed by M±3<r M , Theoretical Range 
M±:3 X Standard Deviation, Md=2S., and Observed Range, in Graph form; and 
also by a statistical comparison of the Difference of Means, with a value of 
"t" calculated for each item of Standard Data, all populations being referred 
to the type population of scincoides. Those of tovelli from 60 miles south of 
Darwin, and from Groote Eylandt are also compared with the type series of 
tovelU. 



Compared with type series of scincoides 




Compared with type 
series of tovelli 


1 scincoides 

ex 
1 Hollandia 

I n. j t. 


tovelli 
kohlsi j type aeries 
n. j t. | n f | t. 


tovelli 
10 nils, 8. 
of Darwin 
n.j t. 


tovelli 

ex Groote 

Eylandt 

n.| t. 


tovelli | tovelli 
60 mla. 8. | ex Groote 
of Darwin | Eylandt 

B.| t. |tt.| t. 



AW 

PW 

SB 

ASB 

PSB 

BD 

A-P 

AM 

AL 

PL 

Sene. 



144 

44 
| 45 
| 45 
145 
J45 
145 
137 
(43 
I 44 



+ 8*05 


18 


1 i 

+12-23117 


—12*29 


19 1—36*52 


1 

22 


— 20-12) 12 


+ 2'49 


18 


+11-76] 17 


—20-95 


19 1—32-12 


22 


— 18-6 | 


12 


|+ 7-06 


IS 


+15-32] 17 


—13-75 


19—24-79 


22 


-12-40 


12 


|+ 5*94)18 | 


+16-78|37 


— 1 -471 ID [— 6-86 


22 


— 2-44 


12 


|+ 5-94 


18) 


+13-06 


17 


— 1-371 19 j — 2-80 


22 


— 2-27 


12 


j + 7-78 


18 


+16-35 


171 


— 2'2l' 


19 |— 6-08 


22 


— 0-14 


12 


j+ 5*14 


18 


+ 9-69 


17 


— 5-72 


19 


— 9-37 


22 


— 4-09 


12 


I+13-36 


17 


+ 7-40 


16 


—16-79 


18 


—16-35 


21 


— 2-26 


12 


|+ 9-73 


18 


+13-5 


17 


—19-4 


19 


—13-73 


22 


—16-19 


12 


[+ 7-33 


18 


+15*3 


17 


—10*8 


19 


—14- 55 


22 —10-52 


12 


1 " '*. 


+ 6-61 


16 |- - 1-51 
i 

i 


17 


— 4-331 18 I — 1-01) 11 1 

1 1 i 1 



-6-94| 
— 2-73| 
— 3-121 
— 3-77| 
— 1-25J 
—5-77 
—3-77 
—4-09 
—0-47 1 
— 3*13j 
— 4 - 30 



+7-23 
+5-94 
+2-19 

—0-54 

+4-32 

+2-60 

+3-92 

( —1-38 

No- variation 

15 | —0-36 

il2|- +2-54 



15 
15 
15 
15 
15 
15 
15 
15 



It is evident from this table and from the graph that while all these 
populations differ very significantly from one another at the 0-1% level of 
probability, at least two groups may be separated biometrieally and on a 
specific level, the one with the largest scuta including scincoides and its popu- 
lation from Hollandia as well as kohlri; the other including tovelli and its two 
populations. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 1.19 

TrOMBICULA JtTBBULPORBNSIS Sp. 11. 

Plate 20, fig. A-B. 

Description of Larva. Shape ? Length and width ! Scutum as figured 
with AW very much shorter than PW and lateral margins strongly divergent 
posteriorly; sensillae ?, their bases very wide apart; posterior scutal margin 
lightly concave between PL, almost rectilinear; AL the shortest, PL the 
longest. Eyes 1 Chelieerae with only the apical tricuspid cap. Galeal setae 
nude. Palpi stout, tibia elaw bifurcate; seta on femur and genu ciliated, on 
tibia only the ventral ciliated. Dorsal setae to 30/t long, ca. 38 in number and 
arranged 2.8.6.6.6.6.4. Ventrally with a pair of ciliated setae on maxillae, 
one on each coxa, a pair between coxae I and between coxae III, thereafter ? 
Legs i tarsi I and II with dorsal sensory rod, III without any long nude seta. 

The Standard Data for the unique type are: AW 56-0, PW 86-8, SB 44-6, 
ASB 28-0, PSB 12-0, SD 40-0, A-P 50*4, AM 25-22, AL 22-4, PL 50-4, 
Sens. — . 

Loc. and Host. A single specimen from a rat, Jubbulpore, India, April, 
1947(S.L.Kalra). 

Remarks. As the type was inadvertently damaged under oil immersion 
only a few .details are figured. The species should, however, be easily recog- 
nized again on the characteristic scutum, etc. 

Trombictjla khurdangensis sp. n. 

Plate 20, fig. C-D. 

Description of Larva. Shape engorged, elongate oval, with slight constric- 
tion behind coxae III. Length 585ft, width 325/*.. Scutum as figured, roughly 
pentagonal but the posterior angle rather shallow, lateral margins divergent 
posteriorly; AM in front of line of AL and AL well back from the rounded 
anterolateral shoulders; sensillae filamentous and ciliated on distal two-thirds, 
their bases midway between lines of AL and PL : AL the shorter and PL the 
longer. Eyes not seen. Chelieerae with only the apical tricuspid cap. Galeal 
setae nude. Palpi stout, tibial claw bifurcate; all setae on femirr, genu and 
tibia ciliated or branched. Dorsal setae strong, curved and distinctly ciliated, 
44 in number, to 38/t long and arranged 2.4.6.6.4.6.6.4.4.2. Ventrally with a 
pair of ciliated setae on maxillae, one on each coxa, a pair between coxae I 
and between coxae III, and thereafter 8.6.8.8.8.6.6.4.2. to ca. 32/x long. Legs: 
I 2G0/I long, II 247/;., Ill 286/i ; tarsi I and II with dorsal sensory rod, III with- 
out any long nude seta. 



120 Records of the S.A. Museum 

The Standard Data for the unique type are: AW 61-6, PW 73-0, SB 
22-4, ASB 28-0, PSB 28-0, SD 56-0, A-P 30-0, AM 36*4, AL 30-8, PL 47-6, 
Sens. 64-4. 

Loc. and Host. A single specimen from a rat from Khurdang, Kashmir, 
July, 1949 (S. L, Kalra), 

Remarks. This species in the general form of the scutum is closely allied 
to T. rajoriensis sp. n., but differs in the Standard Data, the number and 
structure of the dorsal setae, and the unisetose coxa III. 

Trombicula rajoriensis sp. n. 

Plate 20, fig. G-K. 

Description of Larva, Shape (engorged) elongate oval with slight con- 
striction behind coxae III, Length 585/i, width 455/*. Scutum as figured, pen- 
tagonal with lateral margins almost parallel; sensillae filamentous, ciliated on 
distal half and with their bases in front of line of PL; margins of posterior 
angle lightly concave; AM in front of line of AL and AL rather back from 
the rounded anterolateral corners ; AL the shortest, PL the longest. Eyes not 
seen. Chelicerae with only the apical tricuspid cap. Galeal setae branched or 
ciliated. Palpi stout, tibial claw bifurcate; all setae on femur, genu and tibia 
ciliated. Dorsal setae stiff and rather spine-like, curved, with short barbs, 34 
in number, to 56/x long, and arranged 2.8.8.6.4.4.2. Ventrally with a pair of 
ciliated setae on maxillae, one on coxae I and II, and two on eoxae III, a pair 
between coxae I and between coxae III and thereafter ca. 50 setae, arranged 
ca. 2.6.6.6.6.6.6.6.4.2, to 40/a long, and shortly ciliated, not barbed. Legs: I 286/t 
long, II 260/a, III 300/*; tarsi I and II with the dorsal sensory rod (spur), III 
without any long nude seta. 

The Standard Data for the unique type are: AW 53-2, PW 56-0, SB 22-4, 
ASB 30-8, PSB 33-6, SD 64 4,. A-P 28-0, AM 40*0, AL 25-2, PL 50-4, 
Sens. 56 • 0. 

Loc. and Host. A single specimen from a rat from Rajori, Kashmir, India, 
May, 1948 (S. L. Kalra). 

Trombicula kashmirensis sp. n t 

Plate 20, fig. B-F. 

Description of Larvae. Shape (engorged) elongate oval with tendency to 
constriction behind coxae III. Length to 520/;,, width to 429^. Scutum as 
figured, with posterior margin deeply angular, giving a pentagonal shape; 



Womers ley— Asiatic-Pacific Scrub Typhus Mites 121 

PL the longest ; sensillae bases only slightly in advance of line of PL ; scutal 
surface finely punctate. Eyes 2+2, posterior the smaller. Chelicerae with 
only the apical tricuspid cap, Galeal setae nude. Palpi stout with trifurcate 
tibial claw ; setae on palpal femur and genu ciliated, on tibia only the ventral 
branched or ciliated. Dorsal setae 44 in number, to 47/* long and arranged 
ca. 2.13.9.8.5.5.2. Ventrally with paired ciliated setae on maxillae, one on 
each coxa, a pair between coxae I and between coxae III and thereafter 
6.6.4.4.4.6.4.2, to 3(M0/* long. Legs: I 325/x long, II 286,,, III 325/*; tarsi I 
and II with usual dorsal rod, III without any long nude seta.; telofemur of 
all legs with a long outstanding ciliated seta, somewhat longer than the other 
seta but not so long as in 2\ kanzalwanensis sp. n. 

The Standard Data for the type and 3 paratypes are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Pange 


Range 


Variation 


AW 


81'9±0-70 


l-40±0-49 


77.7-86-1 


81-2-84-0 


1-7 


PW 


98-7±0-70 


1- 40+0 '49 


94-5-102-9 


98-0-100-8 


1-4 


SB 


34-65±0-67 


l-34±0-47 


30.65-38-65 


33-6-36*4 


3-8 


A8B 


29-4±0-81 


l*61±0-57 


24-6-34-2 


28-0-30-8 


5*5 


P8B 


33-6 


No variation recorded . 






SD 


63-0±0-81 


1-61^:0.57 


58-2-67-8 


61-6-64-4 


2-5 


A-P 


30-8 


No variation recorded. 






AM 


42-0±l-14 


2-29;£0-81 


35-1-48-9 


39-2-44-8 


5-4 


AL 


42-7±0-70 


1- 40+0-49 


38-5-46-9 


42-0-44-8 


3-3 


PL 


54-6 (Aver. 


of two determinations 


only, viz. 53-2 


and 56-0) 




Sens. 


84-0 


One determination only. 







Loo. and Hosts. Four specimens from a rat from Khurdang, Kashmir, 
India, July, 1949 (S. L. Kalra). 

Remarks. Very similar in scutum to T. kanzalwwnenm, but easily separ- 
ated as in the key on the absence of a long nude seta on tarsi of leg III, and in 
the arrangement of dorsal setae. 

Trombioula mubidia sp. n. 

Plate 20, fig. L-M. 

De$cr'lption of Larva. Scutum transversely rectangular, as figured with 
posterior margin an even but shallow curve behind line of PL ; sensillae bases 
in line of PL; sensillae filamentous and ciliated in distal half; PL the longest. 
Eyes? Chelicerae with only the apical tricuspid cap. Galeal setae branched. 
Palpi with bifurcate tibial claw; seta on femur and genu branched; setae on 
tibia nude except the dorsal. Dorsal setae 26 in number, to 53^ long, and 



122 Records of the S.A. Museum 

arranged 2.8.6.6,4. Ventrally with paired ciliated setae on maxillae, one on 
each coxa, a pair between coxae I and between coxae III, thereafter uncertain. 
Legs, tarsi of III without any long nude seta ; of I and II with dorsal sensory 
rod. 

The Standard Data for the unique type are : AW 58-8, PW 72 -8, SB 32-0, 
ASB 28-0, PSB 12*0, SD 40-0, A-P 22-4, AM 53-2, AL 56-0, PL 64-0, 
Sens. 60 -/J. 

hoc. and Host. A single specimen from Baitus midleri. No. 7646, from 
Bukit Lagong, Kepong, P.M.S., 25 July, 1950 (J.R.A.). 

Remarks. The unique example of this species was unfortunately damaged 
in remounting, and consequently entire dorsal and ventral figures are not 
given, and the description lacks certain details. The scutum, and the palpal 
setation, however, will enable the species to be again recognized. 

In the key, it will run down to near to T. sylvestris Audy and Traub 
1950, from which it is easily separated. 

Trombicula taphozous sp. n. 

Plate 21, fig. A-B. 

Description of Larvcv. Length (engorged) 650/*, width 455/*. Scutum as 
figured; sparsely punctate, the punctations fairly large; anterior margin 
shorter than posterior, and lightly concave between AM and AL; posterior 
margin shallow behind line of PL, and concave, almost rectilinear laterally on 
each side from PL for a quarter of its leugth, then deepening to a lightly 
sinnous curve; sensillae filamentous, ciliated distally and their bases in front 
of PL ; PL the longest, AM the shortest. Eyes f 2 + 2. Chelicerae missing. 
Galeal setae nude. Palpi as figured; palpal claw trifurcate; seta on femur 
nude or shortly and indistinctly ciliated, on genu apparently nude, on tibia 
all 3 setae apparently nude. Dorsal setae difficult to ascertain owing to dis- 
placement, but ca. 2 plus 5 rows of about 10, plus 8.6.4. or ca. 70 in number,, 
to 30/* long. Ventrally with a pair of ciliated setae on maxillae, one on each 
coxa, a pair between coxae I and between coxae III and thereafter ca. 50 -f-, 
to 28{jl long. Legs: I 286>, II 260/*, III 300/*; tarsi I and II with dorsal rod, 
III without any long nude seta. 

The Standard Data for the unique type are: AW 64-4, PW 87 -0, SB 19*6, 
ASB 33-6, PSB 14*0, SD 47-6, A-P 39-2, AM 20-0+, AL 40*0, PL 50-4, 
Sens. 56-0. 

Loc. and Host. A single specimen from bat, Taphozmts mdamopogon, 
R.9626, No. 13062 from Palau Seletan Is. (P. Angsa), Malaya, 1/2/50, (J.R.A.). 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 123 

Remarks, In the key this species comes near to dewipiliata, from which it 
differs in the shape of scutum, the nude setae on palpal femur and genu, and 
the number of dorsal setae, 

Trombioula tithwalensis sp* n. 

Plate 21, fig. C-G. 

Description of Larvae. Length (engorged) 520/*, width 520/*. Shape 
rounded. Scutum large, finely punctate as figured, with SB slightly behind line 
of PL, and AM well back from line of AL almost midway between line of 
AL and PL; PL a little longer than AL; sensillae ciliated distally. Eyes 
2 + 2, rather small, and posterior eyes the smaller, Chelicerae missing. 
Oaleal setae branched. Palpi with bifurcate tibial claw; setae on femur and 
genu nude, on tibia both dorsal and ventral branched, lateral nude. Dorsal 
setae ca. 42, arranged 2.12.8.8.6.4.2, from 70//. long anteriorly to 56/x, posteriorly. 
Ventral with a pair of maxillary ciliated setae, one on each coxa, a pair 
between coxae I and between coxae III, and thereafter ca. 36, increasing in 
length posteriorly. Legs « I 260^ long, II 234/x, III 286/i ; tarsi I and II with 
dorsal sensory rod ; tarsi III without any long nude seta. 

The Standard Data for the unique type and 3 other specimens are : 



AW 
PW 


Mean 
83-2±:0-70 
93-8+0-81 


Standard 
Deviation 

l-40±0-49 

l-61±0-57 


Theoretical 
B&nge 

79-1-87-5 

89-0-89*6 


Observed 
Bange 

81-2-84-0 

92-4-95-2 


Coeff. of 
Variation 

1-7 

1-7 


SB 
ABB 


36«4:±M4 
35-0±:0-81 


2-28±0-81 
l-61±0-57 


29-6-43-2 
30*2-39*8 


33-6-39-2 
33-6-36-4 


6*3 
4-6 


P8B 


16-l±0-70 


l-40±0-49 


11-9-20.3 


14.0-16-8 


8-7 


SD 


51-l:£l-34 


2-68±0-95 


43-1-59-1 


47-6-53-2 


5-2 


A-P 


28-7±l-34 


2-68±0-95 


20-7-36«7 


25 -2-30 -a 


9-3 


AM 


51-3±2-46 


4-27±l*74 


38-5-64-1 


47-6-56-0 


8-3 


Ali 


47-6 


No variation recorded. 






PL 


58-l±l-34 


2-68±0-95 


50-1-66-1 


56-0-61-6 


4*6 


Sens. 


78-4 


Only one determination. 







Loc t amd Host. A single specimen from ear of rat, Tithwal, India, Sept. 
1948 (S.L.K.), and 3 specimens from a "mouse" from Baltal, Kashmir. 

Remarks. This species appears closest to bhimti(hUn<sis sp n. but differs in 
the dorsal setae, and Standard Data. In the illustration of the scutum the 
sensillae are erroneously shown as nude. The posterior scutal margin should 
also be shown as lightly concave laterally from PL and medially almost 
straight. 



124 Records of the S.A. Museum 

Trombicula buxtoni sp. n. 

Plate 22, fig. A-F. 

Description of Larvae, Shape an elongate oval, "widest in anterior third 
and slightly constricted behind coxae III, Length (engorged) to 525/a, width 
to 327/i. Scutum small, roughly quadrate, but the posterior margin forming 
a shallow angle ; AL placed well behind anterolateral corners ; anterior margin 
concave and AM well in front of AL. Sensillae long and filamentous, ciliated 
for their whole length, with their bases in front of PL. Byes 2 + 2, the pos- 
terior the smaller. Chelicerae non-serrate, with only the apical tricuspid cap. 
Galeal setae nude. Palpal claw trifurcate; setae on palpal femur, genu and 
tibia all strongly ciliated. Dorsal setae thick, and strongly ciliated, to 48/* 
long, and arranged 2.6.6(4).4(6).4.2.2. Ventrally with a pair of ciliated setae 
on maxillae, one on each of coxae I and II, two on each of coxae III, a pair 
between coxae I and between coxae III, and thereafter ca. 38 setae to 32/i long. 
Legs : I 285/a long, II 241/i, III 312ft ; tarsi I and II with the usual dorsal sen- 
sory rod, III with a long fine nude seta, not easy to see and not as long as 
the ordinary setae. 

The Standard Data derived from seven of eight specimens, are as follows : 

Coeff. of 
Variation 

4-8 

3-7 

42 







Standard 


Theoretical 


Observed 




Mean 


Deviation 


Range 


Range 


AW 


45 -3:1:0 -83 


2-21±:0-59 


38-7-51-9 


41-6-48-0 


PW 


46 -6:1:0 -65 


1-71^:0-46 


41-5-51-7 


44-8-48-0 


SB 


18-7:±0-29 


0-78:t0-21 


16-^21-6 


17.6-19*2 


ASB 


28-8 


No variation recorded. 




PSB 


22-4 


No variation recorded. 




SD 


51*2 


No variation recorded. 




A-P 


25*6 


No variation recorded. 




AM 


38*4 


No variation recorded. 




AX 


22-4 


No variation recorded. 




PL 


41-6 


No variation recorded. 




Sens, 


54-7:1:1-06 


2-37:1:0-75 


47-6-61-8 


51-2-57-6 



4-3 

Loo. and Host. Eight specimens (the type and 7 paratopes ) from a yellow 
coloured rat from Kanikhet, Kumaon Hills, India, Oct. 1946 (S. L. Kalra). 

Remarks. A very characteristic species in the shape and dimensions of the 
scutum, the 2-setose coxae III, the dorsal setae, and the setation of the palpi. 
It can be distinguished as in the key. 

Trombicula naultini Dumbleton 1947. 
Trans. Roy. Soc. N. Zealand, 1947, 76, 412. 

Plate 22, fig. G-J. 
The type and paratype of this species were from a gecko Naidtimts elegams 
Gray, from Invercargill, N. Zealand (coll. B. B, Given, no date), Dumbleton 



Womersley— Asiatic-Pacific Scrub Typhus Mites 125 

records also six other specimens as " Parasitic on a Gecko, Big Hill, 18.3.12, 
G. Gar row." 

In this species the scutum is considerably wider than long, with A-P rela- 
tively short, SB posterior of line of PL, and the posterior margin deep behind 
PL and considerably rounded. The eyes are only one on each side, the pos- 
terior eyes apparently being completely obsolete. The chelicerae are simple, 
with only the apical tricuspid cap, and the galea! setae are slender and nude. 
The palpi are fairly stout with trifurcate tibial claw; all the setae on femur, 
genu and tibia nude. The dorsal setae are 22 in number, arranged 2.8.6,4.2, to 
50/i long, and fairly stout with short ciliations or serrations. All coxae unise- 
tose, Tarsi III with one long nude seta. 

The Standard Data given by Dumbleton for the type and paratype are t 
AW 81-0, 82-0, PW 99-0, 91-0, SB 40-0, 36-0, ASB 26-0, 30-0, PSB 20-0, 
20-0, SD 46-0, 50-0, A-P 16-0, 16-0, AM — , — , AL 23-0, 26-0, PL 49-0, 
49*0, Sens. 69-0,—. 

For the additional specimens from Big Mill which I have seen and 
measured, the Standard Data are as follows - 







»Sltandard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


79»25:£l-16 


3-28:t0'82 


69-4-89-1 


73*0-82-0 


41 


PW 


90-75±l-74 


4-92±l-23 


76-0-105-5 


82-0-99-0 


5-4 


8B 


33-4±l-99 


5*63±1-41 


15-5-50-3 


25-0-40-0 


16-8 


ASB 


25-0±0-76 


2- 14:1:0 -53 


18-6-31-4 


24-0-30-0 


8-5 


PSB 


20-9±0-35 


0'99:±0 '25 


17-9-23-8 


20-0-23-0 


4-7 


8D 


45-9±0-66 


l-88:±0-47 


40-1-51-5 


44-0-50-0 


41 


A-P 


17-5±0-33 


0-92^:0 -23 


14-7-20-3 


16-0-18-0 


5-3 


AM 


S9-0±l-34 


3*28=fc0-95 


29-3-48-8 


33-0-42-0 


8-4 


AL 


27'6±l-02 


2-87±:0-72 


19-0-36-2 


23*0-32-0 


10-4 


PL 


47-l±l-02 


2-90rt0-72 


38-4-55-8 


42-0-51-0 


6-2 


Sens. 


74-0±l-00 


2-45r£0-71 


66-7-81-3 


69-0-75-0 


3-3 



Trombicula oervulicoiiA Ewing 1931. 
Ewing 1931, Proc. U.S.N. Mus., 80, (8), 13, pi. I, fig. 3. 

Plate 22, fig. K. 

Ewing described this species very inadequately and illustrated the scutum 
by a somewhat inaccurate drawing. It was described from specimens from a 
deer, Cervulus aureus, from Muktesar, Kumaon, India. 

Dr. G. W. Wharton some time ago re-examined the types in the U.S.N. 
Mus., and informed me in correspondence that the dorsal setae were not 26 in 
number, as stated by Ewing, but 38, arranged 2.8.8.8.6.4.2, and also that the 
lateral and ventral setae on the palpal tibia were nude. 



126 Records of the S.A. Museum 

In 1947 on a visit to Washington 1 was privileged to also examine the 
type and two paratypes of this species, and can now give the following details. 

The scutum is roughly rectangular as now figured, with the anterior margin 
lightly sinuous ; lateral margins straight or slightly coneave and not convex as 
shown by Ewing.; posterior margin fairly deep behind PL and flattened medi- 
ally, not as in Ewing 's figure; sensillae with a few indistinct branches distally, 
and bases in front of line of PL. Eyes 2 -j- 2, small. Chelicerae with only an 
apical cap. Galeal setae nude* Palpi with trifurcate tibial claw; setae on 
palpal femur and genu branched; on tibia, lateral and dorsal nude, ventral 
branched. Dorsal setae 34 to 36, arranged ca. 2.8.8.8,6.4.2, and to 50/i long. 
Leg III with a long nude outstanding seta on tarsi. 

The Standard Data of the above three specimens as measured by me in 
1947 are: AW 65-2, 65-2, 72-0, PW 71<5, 68-5, 81-5, SB 26*0, 26-0, 32*6, 
ASB 23-4, 23-4, 23-4, PSB 23-4, 23<4, 23-4, SD 46-8, 46-8, 46-8, A-P 
26-0, 26-0, 32*6, AM 39-1, 42-0, 45-6, AL 39-1, 39-1, 45-6, PL 42-5, 42-5, 
65-2, Sens. 72*0, 65-0, 81-5. 

Trombioula c/. densipiliata Walch 1923. 

Kitasato Archiv. Exper. Med., 5, (3), 79, 1923; Womersley and Heaslip, Trans. 
Roy. Soc. 8. Aust., 67, (1), 80, 1943; Sig Thor and Wilimann, 1947, Das 
Tierreich, Lfg. 71b, 267. 

Plate 23, fig. B-E, 

This species was originally described by Walch from a single specimen 
which, although he does not specify the host, was probably from a rat, from 
Deli, Sumatra. The very inadequate data given by Walch was summarized by 
Womersley and Heaslip 1943 (loc. cit.) and the Standard Data estimated from 
Walch 's figures. 

To this species are now assigned the following specimens. 

Two specimens taken in a log near the camp at Morotai, Halmahera, in the 
Celebes, 24 April, 1945, by Maj. K. N. McCulloeh. Only one of these was suit- 
able for measuring and gave the following Standard Data: AW 66-0, PW 73*0, 
SB 26-0, ASB 30'O, PSB 120, SD 42-0, A-P 30-0, AM 45-0, AL 45-0, PL 
60-0, Sens. 45-0. 

One specimen collected by Capt. L. J, Dumbleton from a rat on Nissan, 
Green Island, in the Solomons, 23 March, 1944, with the following Standard 
Data: AW 69-0, PW 750, SB 24-0, ASB 27-0, PSB 15-0, SD 42-0, A-P 
27-0, AM 42-0, AL 45*0, PL 62*0, Sens. 50*0. 

The above values generally are somewhat lower than those computed by 
Womersley and Heaslip from Walch 's drawings. The DS number ca. 54, are 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 127 

arranged 2.12.10.10.8.G.4.2, and measure up to 5fyi long, Walch gives the DS 
as numbering 58. 

The dorsal scutum is rectangular with the posterior margin almost rectili- 
near between PL (in the Nissan specimen it appears to be somewhat deeper 
behind PL, although the same distance from SB, and resembling that in nissam 
Dumbleton). The palpal claw is trifurcate, setae on palpal femur and genu 
branched; on tibia dorsal and lateral nude, ventral branched (all 3 nude 
apparently in specimen from Nissan). The ehelicerae are furnished with only 
the apical tricuspid cap, and the galeal setae are furnished with a single branch 
(not nude as given by Walch). 

The anterior eyes on each side are very large and conspicuous and both 
eyes are on distinct ocular shields. 

In the posterior scutal margin and the nude ventral seta of the palpal 
tibia the specimen from Nissan resembles the species T. 7iissani Dumbleton, but 
the number and arrangement of DS are as in densipiliata. 

Trombicula corvi Hatori 1920, 

Taiwan Igakai Zasshi, No. 209, Pt. II; Kawamura and Yamaguchi, 1921, 
Kitasato Archiv. Exper. Med. 7 4, 169; Sugimoto, 1926, J. Jap. Soc. Vet. 
Sci., 15, (1), 200-213 (in Japanese with English Summary) ; Womersley 
and Heaslip, 1943, Tr. Roy. Soc. S, Aust, 67, (1), 88. 

Plate 23, fig. A. 

This species was described from fowls from Formosa. Hatori 's original 
paper has not been seen but Kawamura and Yamaguchi give the following 
details in the form of a table. 

" Length 789 -1/x.r width 648-8^. Shape almost quadrate. Colour orange 
red. Dorsal scutum approximately trapezoidal, anterior margin shorter than 
posterior ; anterior and lateral margins rectilinear in comparison with posterior 
margin which is backwardly curved; length 37 -0/a, width 87-0^; line between 
postero-lateral seta 8/*. behind pseud ostignial line. Sensillae in distal half quite 
nude, in proximal half with 7-8 strong short branches; 58-1/a long. Eyes 1+1, 
lens 17-4/i. in diam. Total setae 108?, dorsally arranged 12.10.16.12,10.8, 
length 45/a. Palpal femur with 1 long branched seta; genu same; tibia all 
3 nude ; tarsi with 1 thick long branched, 2 rather thin long branched setae. 
Galeal setae nude. Tarsi III without any long nude seta. On birds". 

From these details "Womersley and Heaslip (1943) gave an approximate 
scale figure of the dorsal scutum. 

No material referable to this species has been seen since the original 
description, but apparently it is a form that should be easily recognized by the 
uensillae being nude distally and ciliated basally. 



128 Records of the S.A, Museum 

TEOMBicuiiA insoqlli Philip and Traub 1950. 
J. Parasitol., 1950, 36, (1), 32-33, fig. 2. 

Plate 23, fig. F^H. 

This species was one of two described by the authors from bats Eonycteris 
spelaea, from the Batu Caves, near Kuala Lumpur, Federated Malay States, 
March, 1948, 

From the description and figures given in the original publication the fol- 
lowing details will diagnose the species. 

Colour pale. Length (engorged) 536//,, width 286/*. Scutum as figured 
with AM slightly in advance of AL and the anterior margin lightly sinuous ; 
posterior margin almost rectilinear between PL; sensillae filamentous, ciliated 
on distal two-thirds and their bases much nearer to line of PL than to AL; 
surface finely punctate ; size small. Eyes 2 + 2, relatively large, posterior the 
smaller, Chelicerae with only the apical tricuspid cap. Galea! setae nude. 
Palpi stout with trifurcate palpal claw; setae on femur, genu and tibia all 
nude. Dorsal setae shortly ciliated, variable, 2. 8-9. 8-10. 8. plus ca. 20, to 
45ft long. Ventrally with paired branched setae on maxillae, one on each coxa, 
a pair between coxae I and between coxae III, and thereafter ca. 18 to 20 
setae, slightly shorter than dorsal setae on tarsi III. 

The Standard Data as given by the authors for the type and 2 paratypes 
are: AW 52, 48, 50, PW 69, 65, 65, SB 22, 23, 23, ASB ca. 48* PSB 12, 11, 
12, SD ca. 60 # , A-P 36, 35, 35, AM 44, 46, 45, AL 42, 42, 42, PL 57, 64, 60, 
Sens. 68, — , 65. 

Tbombicula (Trombicula) frittsi Wharton 1945. 

Trotribicula frittsi Wharton 1945, J. Parasitol., 31, (4), 282. 

Plate 23, fig. I-L. 

Re-description* of Larvae. A small sper3ies of ovoid shape. Length 235/x, 
width 173/i. Dorsal scutum small and pentagonal with the posterior margins 
thickened and sinuous, the apical angle with a series of internally radiating 
lines ; angle of convexity of posterior margin = 2-78. Eyes 2 + 2, close to the 
posterolateral angles, the posterior slightly the smaller. Chelicerae rather short, 
with only the apical tricuspid cap. Galeal setae long and nude. Palpi stout with 
trifurcate tibial claw; setae on femur 2-3 branched; on genu nude; on tibia, 
ventral branched, dorsal and lateral nude; tarsi small with sub-basal sensory 
rod, 4 nude setae and 2 strongly ciliated setae over-reaching tip of claw. Dorsal 

# Interpolated from Philip and Traub 's figure. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



129 



setae 22, arranged 2.6.6.4.2.2, to 30/x long, and ciliated. Ventrally with a pair 
of branched setae on palpal segment I; one on each coxa, a pair between coxae I 
and between coxae III, and thereafter 4.4.2.2. setae to 20/x, long. Legs : I 180/* 
long, II 162/i, III 180/x : tarsi I and II with the usual dorsal rod, III without 
any long nude seta. Sensillae rather thick-stemmed basally, with 5-6 branches 
distally. 

The Standard Data for five specimens are: 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


42.6:£1*75 


3-91±l-24 


30-9-64-3 


89-0-42-0 


9-2 


PW 


54-0±:l-48 


3-31±l-05 


40-05-63-95 


51-0-58-0 


6-1 


SB 


15-6:t0-60 


l-34±0-42 


11. 6-39-6 


15-0-18-0 


8-6 


ASB 


16 -2±0- 73 


l-64:£0-52 


11 -3-21-1 


15-0-18-0 


10-1 


PSB 


19-4±0-58 


1-31±0-41 


15-5-23-3 


18-0-21-0 


6-7 


8D 


35-6±l-25 


2-79-£0-88 


27-2-44-0 


33-0-39-0 


7-8 


A-P 


17-6±0-40 


0-S9±0-28 


14-9-20*3 


16-0-18-0 


5-1 


AM 


19-8:±l-20 


2-68±0-85 


11-8-27*8 


18-0-24-0 


135 


AL 


18-4±0-40 


0-89:£0-28 


16-7-21 -2 


18-0-20-0 


4-8 


PL 


29-0±:0-63 


l-41z£0-45 


24-8-33-2 


27-0-30-0 


4-8 


Sena. 


36-0 


No variation recorded. 







Loc. and Hosts, The above re-description is from five specimens collected 
by Ph, M. i/c L r A. Posekany of the U.S. Navy from somewhere in the Pacific, 
probably the Solomon Islands, 1944. Wharton recorded it from Gehyra 
oceamca, Varcmus indicus and Rattw praetor, all from Bougainville. 

Also from a scorpion, Heterometrus longimanus, from Kukit Lagong 
Forest Reserve, Kepong, near Kuala Lumpur, Sept. 1949 (J.R.A. and K.L.C.), 
in numbers, and reared through to the nymphs by K.L.O. The Standard Data 
of this material, 4 of 5 larvae from scorpions, 21 Sept, 1949, and 7 of 10 larval 
pelts, from same hosts and date, are slightly higher than as quoted above for 
the five specimens from Bougainville and are as follows: 



AW 


Mean 
47-6±3-8 


Standard 
Deviation 

l-25-JrO-27 


Theoretical 
Range 

43-8-51-4 


Observed 
Range 

44-8-50-4 


Coeff. of 
Variation 

2-6 


PW 

SB 


63-l±9-5 
17-4Hfc3-8 


3-15±0-68 
l-29±0-28 


53-6-72-6 
13-6-21-2 


58-8-70-0 
l5-4r-19-6 


5-0 
7-4 


ASB 
P8B 


20-l±5-l 
22-4 


1^69^:0^36 15-0-25-2 
No variation recorded. 


16-8-22-4 


8-4 


SD 


42-5±5-l 


l-69±0-36 


37-4-47-6 


39-2-44-8 


4-0 


A-P 


17-3±3-4 


l-14±0-25 


13-9-20-7 


16-8^19-6 


6-5 


AM 
AL 


31»l±5-0 

25-75±6-6 


1-69+0 -40 

2< 21+0. 47 


26-1-36-1 
19-15-32-35 


28-0-33-6 
22-4-30-8 


5-4 
8-5 


PL 

Sens, 


32-3±4-4 
50-4 


l-46zt0-31 27-9-36-7 
No variation recorded. 


30-8-33-6 


4-5 



130 Records of the S.A. Museum 

Remarks. In his description, Dr. Wharton states, w Tibia III (of legs) 
with a sensory seta", and figures such a rod-like seta in his fig* 1A. An 
examination of his paratype in the S.A. Museum, however, shows that such a 
unique feature is not present, and this is confirmed by the recent material from 
Malaya. Probably the ciliations of a normal seta were adpressed and so misled 
him. 

The host distribution of this species from a gecko, goanna, and rats from 
Bougainville, and from scorpions in Malaya is remarkable, and is somewhat 
paralleled by that of T. ram which is also known from a pillmillipede as well 
as from rats and man. 

Trombioula incurva sp. n. 

Plate 24, fig. A-B. 

Description of Larvae, Shape ovate. Length (engorged) 530/x, width 330/t. 
Scutum as figured with anterior margin almost straight, posterior margin 
evenly curved and deep behind PL, lateral margins concave towards PL. Sens, 
behind line of PL. Scutal setae rather short, with AL the shortest. Eyes * 
Chelicerae with only the usual apical tricuspid cap. Galeal setae nude. Palpal 
daw trifurcate j seta on femur with 4-5 long branches ; on genu with one long 
branch; on tibia apparently all 3 nude. Dorsal setae short and stiff, to 20/x 
long, arranged 2.6.6.4.6.4.2.2. Ventrally with a pair of branched setae on 
maxillae, one on each coxa, a pair between coxae I and between coxae III, 
and thereafter ca. 20-24 setae, to 15/* long. Legs: I 212^, II 186^ III 212/x; 
tarsi I and II with the usual dorsal sensory rod, III without any long nude 
seta. 

Standard Data of the type and paratype are-. AW 48-0, 54-0, PW 63-0, 
75-0, SB 19-0, 23*0, ASB 21-0, 22-0, PSB 18-0, 18-0, SD 39-0, 40-0, A-P 
19-0, 20*0, AM 20-0, 20-0, AL 16; 0, 19-0, PL 24-0, 26-0, Sens. 45-0, — . 

Locl and Hosts v The type and 1 paratype from a gecko at Mt Gillen, 
Alice Springs, N.T. of Australia, 29.10,42. (R.V.S.). 

Trombioula vietzi sp. n. 

Plate 110, fig. A-D. 
Desonptio?i of larvae. Sbape ovoid but with sides rather parallel. Length 
(engorged) 603^, width 402^. Scutum as figured, wider than long, with 
posterior margin rather shallow behind line of PL; sensillae missing, bases in 
line with PL; AM and AL equal, PL very slightly long, with strong ciliations. 
Eyes 2 4- 2. Chelicerae with only the apical tricuspid cap, Galeal setae nude. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



131 



Palpi as figured, with trifurcatu tibial claw; seta on femur strongly ciliated, 
on genu nude, on tibia, dorsal and lateral nude, -ventral branched. Dorsal 
setae shortly ciliated, short to 40/a, ca. 30 in number and arranged 2.8.6.6.4.2.2. 
Ventrally with paired setae on maxillae, one on each coxae, a pair between 
coxae I and between coxae III, and thereafter ca. 26 in number. Legs: I 
248ft, II 221/x, III 248/x; tarsi I and II with dorsal sensory rod, III without 
any long nude seta. 

Standard Data rather variable in AW and PW, for the type and 4 para- 
types are : 







Standard Theoretical 


Observed 


Coeff . of 




Menu 


Deviation Range 


Range 


Variation 


AW 


50-9±l-99 


4«45±1-41 37-6-64-2 


46-9-56*95 


8-7 


PW 


58-95±2-01 


4«49±l-42 45*45-72-45 


53-6-63*65 


7-6 


SB 


19«8±0-31 


0-69=t0-22 17-7-21-9 


18-55-20-1 


3-5 


ABB 


20-1 


No variation recorded. 






PSB 


10-05 


No variation recorded. 






SB 


30-15 


No variation recorded. 






A-P 


20-1 


No variation recorded. 






AM 


26-8 


No variation recorded. 






AL 


26*8 


No variation recorded. 






PL 


30-15 


No variation recorded. 






Sena. 


Missing. 









Loc. and Host, The type and 4 paratypes from rat's ear from Jubbulpore, 
India, Jan., 1947 (S. L. Kalra). 

Remarks. This species, despite the absence, of the sensillae is probably 
a Trombicula. It is remarkable for the strongly ciliated seta on palpal femur, 
and the long nude seta on the genu. 

In the key it will come close to T. incurva sp. n., in caption 61. It is 
dedicated to Dr. Karl Vietz, the noted Hydrachnologist. 



Trombicula leveri sp. n. 

Plate 24, fig. C-F. 

Description of Larvae. Shape ovoid. Length (engorged) to 750/*, width 
to 510/a. Dorsal scutum as figured, anterior margin only slightly convex, pos- 
terior margins almost rectilinear between PL j sensillae bases midway between 
anterior and posterior scuta] margins, sensillae missing- A.M and AL with 
longer and more outstanding ciliations than PL. Eyes 2 -\- 2, on distinct 
ocular shields, large, and separated from PL by about the length of PW; 
posterior eyes the smaller. Chelicerae missing. Galeal setae nude. Palpi 
with trifurcate tibial claw; setae on palpal femur and genu with long branches; 
on tibia, dorsal seta nude, both lateral and ventral branched. Dorsal striations 



132 Records of the S.A. Museum 

rather wide apart. Dorsal setae pointed and finely ciliated, 38 in number, and 
arranged 2.8,6.6.6.6.4.2, to 60/* long 1 . Ventrally with a pair of ciliated setae on 
maxillae, one on each coxa, a pair between coxae I and between coxae III and 
thereafter ca. 8.8.8.8.6.6.6.4.2, to 40/* long. Legs: I 330/i. long, II 300/1, III 
330/*; tarsi III without any long nude seta. 

The Standard Data for the type and two paratype specimens are; 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Eange 


Variation 


AW 


62-0±l-00 


1^73=t0-71 


56-8-67-2 


60-0-63-0 


2-8 


PW 


78-3±2-02 


3-51±l-43 


67*8-88-8 


75-O-82-0 


4-5 


SB 


24-0 


No variation recorded. 






ASB 


24-0 


No variation recorded. 






PSB 


24-0 


Only one determination. 






SD 


48-0 


Only one determination. 






A-P 


38-3+1-20 


2-08±0-85 


32-1-44-5 


36-0-40*0 


54 


AM 


57-0 


Only one determination. 






AL 


35-0±l-00 


1-73±0*71 


29-8-40-2 


33-0-36.0 


5*0 


PL 


62-0±:l-00 


l-73±0-71 


56-8-67.2 


60-0-63-0 


2-8 



Seng . Missing in all specimens. 

Lop. The type and 3 paratypes from a bat, EntbaUonura sp., Delciomo, 
Lakeba, Lau, Fiji, 7 Nov., 1945 (K. A. Lever), 

RemarJcs, Owing to the loss of the sensillae in all four specimens, the 
species is placed tentatively in Trombicula. Another specimen from a bat 
(R.10299) from Bukit Lagong, Kepong, Kuala Lumpur, Malaya, 6 Mar., 1950 
(J. R. Audy, Slide 13590) is this species. (Specimen damaged during study.) 

Trombicula piebcei Ewing 1931. 
Ewing 1931, Proc. U.S.N. Mus., 80, (8), 13, pi. I, fig. 4. 

Plate 24, fig. G. 

This species taken from a bat Ripposideros diadema griseiis from Negros, 
Philippine Islands, was very inadequately described, and only the palp figured. 
Lt.-Col. C. B. Philip (in litt.) says, "In an effort to decide on Ewing's T. piereei 
from bats on Negros, P.L, we remounted the very damaged types, but I 
couldn't even include them in my key. Best we could say was that any 
Philippine Trombiculids with sensillary pores almost on the hind margins of 
the scutum and less than twice the diameter between them might be suspected 
as this species". 

In 1947, on my visit to Washington I re-examined the type U.S.N.M., 
No. 1021, and now give the following details and figure of the scutum. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 133 

Scutum not much wider than long, with. SB in front of PL but behind 
midline of scutum; posterior margin a shallow curve rather flattened medially; 
sensillae now missing but bases about twice their diameters apart. (Ewing says 
sensillae slender, very long and barbed on distal two-thirds.) The galeal setae 
are nude. Palpal claw trifurcate; all setae on palpal femur, genu and tibia 
nude. Dorsal setae long and slender to 56/.*,, 42 in number and arranged ca. 
2.8.8.8.6.4.4.2. Tarsi of leg III without any long nude seta. 

The Standard Data of the type as measured in 1947 were: AW 58-7, PW 
81-5, SB 26-1, ASB 34-5, PSB 27-6, SD 62-1, A-P 41-5, AM 55-2, AL 34-5, 
PL 76*0, Sens. — . 

Remarks. The relationship of this species can now be given as in the 
key. The above details should enable the species to be recognized, but it has 
not as yet been again detected amongst the large amount of Philippine material 
seen by me. 

Trombicula sylvestris Audy and Traub 1950. 
Trombicula sylvestris Audy and Traub 1950, Bull. Baffles Museum, 

Plate 24, fig. H-K. 

Re-description of Larvae. Shape broadly oval. Length (fully engorged), 
780/i, width 610/x, Scutum as figured, somewhat quadrate, finely punctate, but 
with posterior margin sinuous and fairly deep behind PL; sensillae bases in 
line with PL j sensillae missing. Eyes ? Chelicerae simple, with only the api- 
cal tricuspid cap. Galeal setae branched. Palpi stout, with bifurcate tibial 
claw; setae on femur and genu nude, on tibia, dorsal branched, lateral and 
ventral nude. Dorsal setae 26 in number, arranged 2.8.6.6.4, from 87/* long 
anteriorly, to 70/x posteriorly, thick on basal half and then tapering (as are 
the scutal setae) and with adpressed ciliations. Ventrally with a pair of 
branched setae on maxillae, a long ciliated seta on each coxa, a pair between 
coxae I and between coxae III and thereafter ca. 8.6.4.4.4.2.2.2, to 42-60/1 long. 
Legs: T 390ju. long, IT 375/a, III 430/t; tarsi I and II with dorsal sensory rod, 
III without any long nude seta. 

The Standard Data for three paratypes are : 



AW 


PW 


SB 


ASB 


PSB 


SD 


A-P 


AM 


AL 


PL Sens 


72-8 


92-4 


36-4 


44<8 


22-4 


67-2 


44-8 


72-8 


50-4 


89-6 — 


72-8 


92-4 


36-4 


44-8 


22-4 


67-2 


44-8 


— 


56-0 


89-6 — 


78-4 


98-0 


42*0 


47 ■ 6 


19-6 


67-2 


47-6 


75-6 


56-0 


89-6 — 



134 Records of the S.A. Museum 

Loo. Described by Audy and Traub from specimens on Eattus sabanus 
from Ulu Langat, Kuala Lumpur, Malaya, 5 Dec, 1949. 

Remarks. The above re-description is from three paratypes sent by 
Dr. Audy and from the original description. 



Trombicula gliricolens (Hirst 1915). 

Microirambidium gliricolens Hirst 1915, Bull. Ent. Research, 6. 

Trombicula gliricolens Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr. r 
67, (1), 82. 

Trombicula (Eutrombimla) gliricolens Sig Thor and Willmann, 1947, Das 
Tierreich, Lfg., 71b, 281. 

Plate 25, fig. A-E. 

This species does not appear to have been met with or recorded since its 
original discovery on a rat at Calcutta, India. 

Through the kindness of Dr. J. R. Audy and Maj. S. L. Kalra, I have 
recently been able to study a number of specimens collected from rats, at 
Tithwal, N. Kashmir, by Si L. Kalra, 1945. 

I believe that these specimens can be referred to Hirst's species, despite 
certain disagreements with the original dscription and figure, viz. (1) the 
palpal claws are definitely trifurcate, whereas Hirst says they are bifurcate, 
and (2) the posterior scutal margin is not a more or less even curve but dis- 
tinctly concave medially. Both these characters may have been inaccurately 
observed. In all other respects, size of scutum, dorsal setae, setation of palpi, 
etc., these specimens agree with glirwolens. 

A re-description of these larvae is as follows: 

Larvae. Shape oval. Length (engorged) to 650/*, width to 42<V< Scutum 
as figured, more or less rectangular, with anterior margin lightly sinuous and 
convex medially; surface rather coarsely punctate ■ sensillae in line of PL; 
AM the shortest and well behind line of AL • PL the longest. Eyes 2 + 2, pos- 
terior the smaller. Palpi with trifurcate tibial claw; setae on palpal femur 
and genu branched or ciliated; on tibia ventral branched, dorsal and lateral 
nude. Chelicerae simple, with distinct apical tricuspid cap. Galeal setae 
branched. Dorsal setae tapering, shortly ciliated, 40 in number and arranged 
2.10.8.8.6.4.2, to 45f«. long, Ventrally with paired ciliated setae on maxillae, one 
on each coxa, a pair between coxae I and between coxae III and thereafter 



Womersley — Asiatic-Pacific Scrub Typhus Mites 135 

6.6.6.4.6.4.2, to 34^ long. Legs all 7-segniented, I 28% long, II 260/*, III 300/*; 
tarsi I and II with dorsal sensory rod, III without any long nude seta. 
The Standard Data for eleven specimens are : 





Mean 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Range 


Coeff. of 
Variation 


AW 


7C5±0-51 


l«69±0-36 


65-4-75-6 


67-2-72-8 


2-4 


PW 


83*5±0-91 


3-02±0-64 


74-5-92-5 


78-4-89-6 


3-6 


SB 


25'5±0-25 


0«84±0«18 


23-0-28-0 


25-2-28-0 


3-3 


ASB 


28-3±0-25 


0-84±0-18 


25-8-30-8 


28-0-30-8 


3-0 


PSB 


13<6:±0-27 


0^90±0-19 


10*9-16-3 


31-2-14-0 


6-6 


8D 


41.9zh0.55 


l-82±0-39 


36-4-47-4 


39*2-44.8 


4-3 


A-P 


24 -4:1:0 -55 


l-81±0-39 


19-0-29-8 


22-4-28-0 


7-4 


AH 


33-2±l-ll 


3-15+0-79 


23-8-42-7 


30-8-39*2 


9-5 


AL 


38-7±l-05 


3-50±0-75 


28-2-49.2 


33-6-42-0 


9-0 


PL 


51-2±0-55 


l-81±0-39 


45-8-56-6 


50-4-56-0 


3-5 


Sens. 


63-0 


— 


— 


61-6-64-0 


— 



Trgmbicula philipi sp. n. 

Plate 25, fig, F-J. 

Description of Larvae. Shape ovate. Size small ; length 225/*, width 170/*. 
Dorsal scutum more or less rectangular, with PW greater than AW; anterior 
margin sinuous, being convex medially and concave laterally of the middle j 
posterior margin rather shallow and more or less evenly curved; SB nearer to 
posterior than to anterior margin ; anterolateral corners produced well in front 
of AL; AM nearer to line of SB, than to line of AL; just in front of AL is a 
transverse line extending right across scutum and just behind this is a pair of 
oval discs; surface of scutum strongly rugose; scutal setae with closely 
adpressed ciliations; sensillae with only 2-3 branches in distal half. Eyes 2 -(- 2, 
on distinct ocular shields, posterior eyes the smaller. Chelicerae with only the 
apical tricuspid cap. C4aleal setae with 1 or 2 long branches. Palpi stout, 
tibial claw trifurcate; seta on femur with 2-3 long branches; on genu with 
1-2; on tibia, dorsal and lateral nude, ventral with 4-5 branches; tarsi with 
the usual basal rod and 5-6 ciliated setae, one longer than the others and 
reaching beyond tip of claw. Dorsal seta 54, stiff and needle-like with only 
•short inconspicuous adpressed ciliations, arranged ca. 2.10.8,10.12,6.4,2, to 22/* 
long. Ycntrally with the usual pair of branched setae on maxillae, one on 
each coxa, a pair between coxae I and between coxae III and thereafter about 
60 setae, somewhat similar to the dorsal setae. Legs: I 270/* long, II 225/*, 
III 270/* ; tarsi I and II with the usual dorsal rod ; III without any long nude 
seta. 



136 Records of the S.A. Museum 

The Standard Data from 25 specimens are as follows 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Bange 


Yariation 


AW 


64-5±0-42 


2-12±0-30 


58-2-70-8 


60-0-68-0 


3-3 


PW 


79-5±0-73 


3-64:t0-51 


68-6-90-4 


72-0-85-0 


4-5 


SB 


22-6±0-29 


l-44±0-20 


18-3-26-9 


21-0-24-0 


6-4 


ASB 


24-9±0-39 


l-97:£0-28 


19-0-30-8 


21* 0-30-0 


7-9 


PSB 


15-3±0-15 


0*73±0-10 


13-1-17-5 


15-0-17-0 


4-8 


SD 


40-2+0-41 


2-08±0-29 


34-0-46-4 


36-0-45-0 


5-0 


A-P 


23-6±0-28 


l-38±0-19 


19-5-27-7 


21-0-27-0 


6-0 


AM 


35-75±0-37 


1-87+0-26 


30-15-41*33 


30-0-39-0 


5-2 


AL 


27-8±0-40 


l-99±0-28 


21-9-33-7 


24-0-30-0 


7-1 


PL 


32-0±0-33 


l-64±0-23 


27-1-36-9 


30-0-36-0 


5-1 


Sens. 


55-3±l-29 


3-87+0-91 


43-7-66-9 


50-0-60-0 


7-0 



Loo. and Hast. Twenty-one specimens from Hipposideros cervinus, Hol- 
landia, D.N.G., (C.B.P. 8), 13 Nov., 1945, and four specimens from the same 
host and locality (W. D. Fitzwater), (no date). 



Trombioula quadriense Womersley and Heaslip 1943, 

Tr. Roy. Soc. S. Austr., 67, (1), 79, 1943. 

— i Trombicula chiroptera Womersley and Heaslip 1943, ibid, 80. 

Plate 25, fig. K-N. 

A re-examination and re-nieasurement of the type and paratype of 
Tr, chiroptera now shows it to be conspecific with quadriense which has page 
priority. 

This is a rather characteristic species in the form of the scutum in which 
the posterior margin is strongly crescentic and PW very mud greater than 
AW so that the anterolateral angles are strongly obtuse. The surface of the 
scutum appears strongly rugose. The eyes are 2 + 2, with the posterior the 
smaller. Chelicerae furnished only with the usual apical tricuspid cap. Galeal 
setae branched. The palpal claw is trif urcate ; the setae on the palpal femur 
and genu, and ventrally on the tibia are branched. The dorsal setae are 28, 
arranged 2.8.6.6.4.2, to 40/* long. Ventrally all coxae are unisetose and pos- 
terior of coxae ITI the setae are 2.6.2.4.2, to 27/x long. Tarsi of leg III without 
any long nude seta. The Standard Data for the type and two paratypes of 
quadriense and the type and paratype of chiroptera are : 



Womersley— Asiatic-Pacific Scrub Typhus Mites 137 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


58-8it0-80 


l-79±0-56 


53-5-64-1 


56-0-60-0 


3-0 


PW 


83-8:±0-73 


1-641:0.52 


78-9-88-7 


82-0-85-0 


1-9 


SB 


20-2±0-92 


2-05±0-65 


14*1-26-3 


18-0-22-0 


10-1 


A8B 


21*0 


No variation recorded. 






PBB 


16-75±l-46 


2-93±l-03 


8-0-25-5 


15-0-21-0 


17-5 


SD 


38-25±l*43 


2-87±l-01 


29-65-46-85 


36-0-42*0 


7-5 


A~P 


23-5:tl-04 


2-08±0-73 


17-3-29-7 


21-0-26-0 


9-0 


AM 


30-75:1:0-75 


l*5O:t0-53 


26-25-35-25 


30-0-33-0 


4-9 


AL 


27-0 


No variation recorded. 






tfi 


*4-0±l-00 


2-24±0-71 


37-3-50-7 


40*0-45-0 


5-1 


Sens. 


50-0 


Only one determination. 







Of recent material I liave seen and examined a specimen from ? Solomons, 
1944 (L. A, Posekany) and four specimens collected by R. N. McCulloeli at 
Wongabel, Queensland, Feb., 1945, two each from a bandicoot and a possum. 
Unfortunately, these specimens have been lost or misplaced. 

Trombioula pelta sp. n. 

Plate 26, fig. A-E. 

Description of Larvae. Shape elongate oval. Size large, length (en- 
gorged) 965ft, width 680/*. Scutum small, rectangular, sparsely but distinctly 
punctate; anterior margin sinuate; posterior margin convex but concave 
medially; PL seta slightly the longer, AM seta about in line with AL; sen- 
sillae missing but bases behind line of PL. Eyes not visible. Chelicerae non- 
serrate, with only the apical tricuspid cap. Galeal setae ciliated. Palpal claw 
trifurcate, seta on palpal femur branched or ciliated; on genu nude; on tibia, 
dorsal and lateral nude, ventral branched; tarsi with dorsal and apical rods 
and 4 or 5 ciliated setae. Dorsal setae ciliated, short, to 28/*., arranged ca. 
2.8.6.6.6.4.2. Ventrally with the usual pair of ciliated setae on maxillae, one 
on each coxa, a pair between coxae I and between coxae III and thereafter 
ca. 6.6.6.6.6.4.2, to 32/* long. Legs: I 241/a long, II 210/t, III 255/t; tarsi I and 
II with dorsal sensory rod, III without any long nude seta. 

Standard Data: AW 43-2, PW 53-2, SB 16-0, ASB 21-8, PSB 7-0, 
SD 28-8, A-P 16-0, AM 30-0, AL 28-8, PL 35-6, Sens. — . 

Loc. and. Host. The unique type from a rat, Nainital, Kumaon Hills, 
India, 10 Oct. 1946 (S. L. Kalra). 

Remarks, May be separated as in the key r 



138 Records of the S.A, Museum 

Tbombioula (Neotrombicula) jayewickremei sp. n. 
Plate 26, fig. F-J. 

Description of Larvae. Colour in life unknown. Shape ovoid. Length 
(engorged) to 450/l, width 350/4. Scutum rectangular; with posterior margin 
shallow behind line of PL, and eoncave medially ; surface lightly punctate, 
AM and PL about equal - sensillae filamentous, ciliated distally, and their bases 
slightly in advance of PL. Eyes 2 + 2, on ocular shields, close to scutal 
borders, posterior eyes the smaller. Chelicerae with only the small apical tri- 
cuspid cap, Galeal setae branched or feathered. Palpi stout ; claw trifureate ; 
setae on palpal femur and genu lightly branched, on tibia all setae nude r 
except ventral. Dorsal setae 30 in number, arranged 2.8.6.6.4.2.2, to 50/x long, 
tapering and ciliated. Ventrally, one seta on each coxa, a pair between coxae I 
and between coxae III, then 6.4.4.2.2, shorter than dorsal setae. Legs: I 227//. 
long, II 210/4., Ill 260/i; tarsi I and II with the usual dorsal sensory rod, III 
without any long nude seta. The Standard Data from the best preparation 
are: AW 67-2, PW 81*2, SB 28-0, ASB 25-2, PSB 11-2, SD 36-4, A-P 
25-2, AM 50-4, AL 44-8, PL 50-4, Sens. 56-0. 

Loc. and Host. Described from seven specimens, collected by Mr. S. H. 
Jayewiekreme; 3 from ears of BatUis rattm kandiyanus, from Nalanda, Cey- 
lon, Sept., 1944, and 4 from ears of the same host from Embilipitiya, Ceylon, 
Nov., 1944. 

Remarks. The relationship of this larvae is best shown in the key. It has 
been reared to the nymph by Mr. Jayewiekreme, as described in the Adult and 
Nymphal section of this paper. 

Trombicula sotjthcotti sp. n. 
Plate 26, fig. K-L. 

Description of Larvae. Shape broadly oval. Length engorged to 420^ 
width to 340/a. Dorsal scutum as figured, coarsely rugose, anterior margin 
almost straight, lateral margins slightly convex btit coming out prominently near 
PL, which are thus placed on a lateral projection, posterior margin cresoentic 
except near PL, sensillae bases nearer to posterior than to anterior margin 
and in front of line of PL, normal setae fairly thick with short setules, sensillae 
with four or five fine branches distally. Eyes 2 + 2, the anterior very convex and 
prominent, posterior inconspicuous. Chelicerae non-serrate but with the usual 
apical tricuspid -cap. Galeal setae with a few fine branches basally. Palpi 
stout but small, tibial claw trifureate; all setae on palpal segments branched,. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 139 

the dorsal and lateral on tibia indistinctly so. Dorsal setae thick with short 
setulations, to 30/a long, and arranged 2.8.6.6.4.2. Ventrally with a pair of fine 
branched setae on maxillae, one on each coxa, a pair between coxae 1 and 
between coxae III, and thereafter ea. 2.6.4.4.4.2, to 25/* long, and finer than 
dorsal setae. Legs: I 260/a long, II 225/*, III 270//,; tarsi I and II with dorsal 
rod-like seta, III with a long nude seta, ordinary leg setae only sparsely 
ciliated. 

The Standard Data from the type and two paratypes are \ 









Standard Theoretical 


Observed 


Ooeff. of 




Mean 


i 


Deviation Range 


Range 


Variation 


AW 


56-7±l- 


33 


2-31±:0-94 49-8-63*6 


54-0-58-0 


4-1 


78-0 




No variation recorded. 






SB 


18-0 




Ko variation recorded. 






A8B 


21-0 




No variation recorded. 






PSB 


18-0 




No variation recorded. 






SD 


39-0 




No variation recorded. 






A-P 


30-0 




No variation recorded. 






AM 


30-0 




No variation recorded. 






AL 


25-0±l- 


00 


l-73±0-71 19-8-30*2 


24-0-27'0 


6-9 


PL 


36*0 




No variation recorded. 






Sens. 


50-0 




Only one determination. 







Trombicula cuneata (Traub and Evans 1951). 
Trombiculindus cuneatus Traub and Evans 1951, Annals Entom. Soc. Amer. 

Plate 27, fig. A-D r 

Shape engorged broadly oval. Length engorged 416/l, unengorged 280/*; 
width 260/* and 180/* respectively. Scutum as figured, with PL aciculate- 
foliate, with 5 or 6 longitudinal finely dentate earinae, and much nearer to 
AL than to the rounded postero-lateral corners ; anterior and posterior margins 
almost straight; lateral margins slightly concave between AL and PL, then 
convex; AM and AL normal, ciliated, with AM the longer; sensillae filamen- 
tous, with a few short barbs on basal half and 3-4 short ciliations on each side 
distally ; surface punctate. Eyes 2 + 2, posterior of PL, posterior the smaller. 
Chelieerae simple, with only a prominent apical tricuspid cap. GaJeal setae 
ciliated. Palpal claw trif urcate ; setae on femur and genu nude, on tibia dorsal 
seta with a few branches ; ventral nude, lateral nude and short. Dorsal setae 
aciculate-foliate with 5-6 longitudinal finely dentate earinae, about a third as 
wide as long, ca. 30-32 in number and arranged ca. 2.8.6.6.4.4.2, those of the 
anterior rows to 75/a, long by 26a. wide and equal, posterior two rows smaller. 



140 Records of the S.A. Museum 

Ventrally, with a pair of normal ciliated setae on maxillae, one such on each, 
coxa, a pair between coxae I and between coxae III, and thereafter ca. 8.6.2.2. 
Legs all 7-segniented, I 260/4 long, II 234/*, III 286/a ; tarsi I and II with dorsal 
sensory rod (spur), III without long nude seta. 

The Standard Data for the type and 15 paratypes as given by Traub and 
Evans are : 

AW PW SB ASB PSB A-P AM AL PL Sens. 

70±4 85±:6 44±2 26±2 16:1:2 16rb2 63±:3 31+4 60±7 68+5 

Log. and Hosts, The type specimen was from Crocidura sp. from Myitk- 
yima, N. Burma, 29 Nov., 1944 (coll. U.S.A. Typhus Commission), and twenty- 
five paratypes from Crocidura or Sunicus, same locality, Nov., 1944 to Feb., 
1945, or from Crocidura from Shingbwiyang, Burma, Feb., 1945. 

Bemarks. In 1947 I was able to measure the Standard Data of the type 
and 3 paratypes in the U.S. N. Mus., tentatively labelled as Webberia cari<no- 
foliuta g. et sp. n. The clata were as follows: AW 74 -95+7 '35, PW 88-5+ 
23*7, SB 43-65+4-61, ASB 29-3, PSB 14-5+5-0, SD 43-8+5-0, A-P 
15-3+2-1, AM! 68-0, AL 31-9+11-0, PL 68-25+4-5, Sens. 75-0, which 
correspond with Traub and Evans' measurements. 

The two species recorded by T. J. Lawrence in Appendix 7, in "Scrub- 
typhus Investigations in S.E. Asia", Part III, AMD 7, War Office, March, 
1947, as from a shrew, Simcus caeruXeus fulvocimreus Anderson, from Imphal, 
1945, as "Gen. nov. species," "W ,? being decsribed in U.S.A., are the above 
species. 

The Standard Data for these two specimens are somewhat higher than for 
the type series, viz. AW 84-0, 84-0, PW 106-4, 112-0, SB 50-4, 50-4, ASB 
33-6, 33-6, PSB 22-4, 22-4, SD 56*0, 560, A-P 16*8, 16-8, AM 75-0, — , 
AL 36*4, 36-4, PL 84-0 X 28-0, 84-0 X 28-0, Sens. — , — . 

N.B. — In the figures given here, the dorsal seta on the palpal tibia are 
shown as nude, whereas actually they are lightly branched as in Traub and 
Evans' figures, and the AM and AL scutal setae are rather stouter than shown 
in the figures in this paper. 

TrombicuiiA squamifera sp. n. 

Plate 27, fig. E-I. 

Description of Larvae. Shape oval. Length (partially engorged) 520/*, 
width 350/a. Scutum large, as figured, PL nearer to AL than to posterior mar- 
gin ; PL aciculate-f oliate with longitudinal rows of setules ; sensillae filamentous, 
in specimens broken short, the bases wide apart and behind line of PL 3 AM and 



Womersley— Asiatic-Pacific Scrub Typhus Mites 141 

AL normal, strongly ciliated with AM the longer. Eyes 2 + 2, fairly large. 
Chelicerae with only the apical cap. Galeal setae strongly branched or 
ciliated. Palpi stout, tibial claw trifurcate; all setae on femur, genu and 
tibia (except dorsal) nude. Dorsal setae aciculate-foliate, the 6 of the second 
row narrower than others; 40 in number, and arranged 2.6.6.8.8.6.4, the an- 
terior ones to 112ju. long by 42/* wide, posteriorly not much smaller. Ventrally 
with a pair of long ciliated setae on maxillae, one on each coxa, a pair between 
coxae I and between coxae III, and thereafter ca. 30 setae to 70/x long followed 
by 6 and 4 setae similar to dorsal setae; on ventral surface laterally on each 
side between coxae II and coxae III is anothr aciculate-foliate seta. Legs all 
7-segmentcd; I 390jlc long, II 364/*, III 416**; tarsi I and II with dorsal rod- 
like seta, III without any long nude seta. 

The Standard Data for the type and paratype are: AW 95-2, 98 -0 r 
PW 117-6, 117-6, SB 47-6, 47*6, ASB 39-2, 39-2, PSB 28-0, 28-0, SD 67-2 r 
67-2, A-P 28-0, 28-0, AM 75-6, 72-8, AL 47-6, 53-2, PL 112-0X42-0, 
106-8 X42-0, Sens, broken short. 

hoc, and Host. The type and paratype from a u mouse" from Baltal r 
Kashmir, India, 1945, Maj. S. L. Kalra. 

RemarJcs. This species, in the form of the PL and dorsal setae is very 
close to cimeata but diifers in the Standard Data, the number and arrange- 
ment of the dorsal setae and in the ventral setae, as given in the key. 

Tromeicula squamosa (Radford 1948). 

Trombicutindw squamosus Radford 1948, Proc. Zool. Soc, London, 118, (1), 
126-128, fig. 1-3. 

Plate 28, fig, A-D. 

This very interesting species was described by Radford from 20 specimens 
from the ears of a rat taken at Ranikhet, Nainital, in the Kumaon Hills, India, 
Oct. 1946 by S. L. Kalra. The type is in the British Museum collection. 

The erection of a new genus for this species on the basis of the peculiar 
foliate or phyllode-like dorsal and posterior ventral setae is unnecessary. Such 
similar modification of the dorsal setae is of frequent occurrence in many 
genera of widely different families of Aeari. For example, in the Cheyletidae, 
Tyroglyphidae, s.l., Oribatoidea, Tetranychidae, etc., and also in certain species 
of 8chonga<stia (Ascoschongastia) (Trombiculidae) foliate setae are not un- 
common. 

I have been privileged to examine a paratype presented by Dr. Radford, 
and from his specimen I give the following re-description, as certain important 
details are omitted from the original description. 



142 Records of the S.A. Museum 

Re-description of Larvae, Shape broadly oval, fairly well chitinized. 
Colour in life probably red. Length (only partially engorged) 325/*, width 
210/i. Dorsal scutum roughly rectangular, with fine pitting; anterior margin 
sinuous, posterior margin fairly shallow and slightly concave medially; AM 
and AL slender and strongly ciliated, AM well behind line of AL, PL broadly 
phyliode-iike with strong puneturations. Sensillae filamentous, basally with 
distinct barbs, distally ciliated, bases in line with or slightly behind PL. Eyes 
2 + 2, on ocular shields, posterior the smaller. Chelicerae strong, not serrate, 
with only the usual apical tricuspid cap. Galeal setae strongly ciliated. Palpi 
stout, tibial claw trifurcate; all setae, on femur, genu and tibia (except dorsal) 
nude. Dorsal setae all broadly phyllode-like, punctate and without ciliations, 
covering entire dorsal surface posterior of scutum, 32 in number and arranged 
2.6.6.6.6.4.2, to ca, 60/x. wide by 60/* long. Ventrally with a pair of ciliated 
setae on maxillae, one on each coxa, those on coxae II shorter than on I and 
III ■ a pair of similar setae between coxae I and between coxae III, and there- 
after ca. 20 long ciliated setae arranged ca. 12*6 and 1 on each side of anus; 
the extreme lateral of first row are the longest, to 100/j. ; behind and flanking the 
anus are three rows of 6.6, and 2, phyllode-like setae similar to the dorsal, 
and laterally betewen coxae II and III also a phyllode seta. Legs all 7-seg- 
inented; I 312^ long, II 292/x, III 360/x; tarsi I and II with the usual dorsal 
sensory rod, III without any long nude seta. 

The Standard Data, for this specimen are as follows: AW 88-0, PW 100-0, 
SB 40*0, ASB 36*0, PSB 11-0, SB 47-0, A^P 31-0, AM 73-0, AL 67*0, PL 
75*0X54-0, Sens. 80-0. 

Remarks. Closely related to this species in having phyllode-like PL and 
dorsal setae are the following three species, cuneata, squamifera and foliacea, 
and all four can be readily separated as in the key. While all four form 
a group of species the erection of a genus or even subgenus on the group 
characters is, as stated above, unjustified, at least until such time as the 
nymphal or adult stages are known 

TROMBicuiiA FOiiTACEA (Traub and Evans 1951). 
Trombiculmdus f&liaceus Traub and Evans 1951. Annals Entonu Soc. Amer. 

Plate 28, fig. E~L 

Shape engorged broadly oval. Length engorged 650/*, unengorged 430/x; 
width 520/x. and 240/i respectively. Scutum subreetanguiar as figured, with 
PL phyllode-like with large reticulations and placed very much nearer to 
AL than to the rounded posterolateral corners; anterior and posterior margins 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 143 

almost straight, lateral margins lightly concave between AL and PL, then con- 
vex; AM and AL normal and ciliated with AM the longer; sensillae filamentous 
with rather short ciliations on distal half, their bases well behind line of PL ; 
surface finely punctate. Eyes 2 + 2, posterior of PL, posterior the smaller. 
Chelicerae simple, with only a rather prominent apical tricuspid cap. Galeal 
setae ciliated. Palpi with trifurcate tibial claw; setae on femur and genu nude; 
on tibia dorsal seta branched, ventral and lateral nude, the ventral short. The 
dorsal setae are broadly phyllode-like with large reticulations and an acute 
apex, about half as wide as long, ca. 28 to 30 in number and arranged 2.8.6.6. 
4.2.2., the posterior 2 rows somewhat smaller than the others. Ventrally with 
a pair of normal ciliated setae on maxillae, one such on each coxa, a pair 
between coxae I and between coxae III and thereafter ca. 16 to 20 normal 
ciliated setae. Legs all 7-segmented ; tarsi I and II with dorsal sensory rod 
(spur), III without any long nude seta; leg I to 234/a long, II 22V, HI 286/i. 

The Standard Data from the values given by Traub and Evans for the 
type and 6 paratypes treated statistically are as follows : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Bange 


Variation 


AW 


66-3+0-64 


1-70+0-45 


61 -2-71 -4 


63-0-68-0 


2-5 


PW 


82 -15+1- 01 


2-67+0-71 


74 15-90 -15 


77-0-84-0 


3-2 


SB 


38-9+0-77 


2-04+0-54 


32-8-45-0 


35-0-42-0 


5-2 


ASB 


2G-3±0-57 


1-50+0-57 


2-1&-30-8 


25-0-28-0 


4-2 


PSB 


14-6+0-37 


0-97+0-26 


11-7-17-5 


14-0-16-0 


6-7 


9D 


40.9+0-74 


1-95+0-52 


35'l-46-7 


39-0-44-0 


4»8 


A-P 


12-9+0-40 


1-07+0-29 


9-7-16-1 


12. 0-14-0 


8-3 


AM 


50-7+1-20 


2-08+0-85 


44-5-56-9 


49-0-53-0 


4-1 


AL 


30-6+1-27 


3-36+0*90 


20-5-40-7 


28-0-35-0 


10-9 


PL 


65*9+1-72 


4-56±l-22 


52-2-79« 6 


60-0-73-0 


7-0 


Sena. 


60-0 


Only one determination. 







Loc. wnd Host. The type was recorded from a shrew Crocidura sp. from 
Shingbwiyang, Burma, 13 Feb., 1945, and the paratypes from the same host 
species and locality, Feb, and March, 1945, collected by members of the U.S.A. 
Typhus Commission. 

A further specimen is that recorded by T. J. Lawrence in Appendix 7 in 
"Scrub^typhus Investigations in S.E. Asia", Part III; AMD 7, War Office, 
March, 1947, This specimen, from which the present drawings are given, was 
much more engorged than Traub and Evans' material. It was referred to as 
"Gen, nov., species Fols." "Lawrence in M.S.'' and was recorded from a 
shrew Suncus cmruleus fidvocinereus Anderson from Kanglatongbi, near Im- 
phal, 1 Nov., 1945. 



144 



Records of the S.A. Museum 



Remarks. In 1947 I was able to examine the type and 2 paratopes in the 
U. S. N. Mus., which at that time had been tentatively labelled as Webberia 
cytofoliata g. et sp. nov, by the discoverers. This name has been dropped and 
the species placed in Eadfords Trombiculindus under the specific cuneatus nov. 

As pointed out under squamosa, neither Webberia nor TromMoiiUndus are 
required, at least until the nymphs or adults are known and can be shown to 
exhibit good generic differences from Trombicula. 

N.B. — In the figures given here, the dorsal seta on the palpal tibia are 
shown as nude, whereas actually they are lightly branched as in Traub and 
Evans* figures, and the AM and AL scutal setae are rather stouter than shown 
in the figures in this paper. 

Trombicula (NEOTEOMBiouiiA) harrisoni sp 4 n. 

Plate 29, fig. A-D. 

Description of Larvae. Length (engorged) to 260/*, width to 195/*. Scutum 
roughly trapezoidal with AM and AL about in the same line; SB very close 
together and much closer to line of PL than to AL; posterior margin almost 
rectilinear between PL; PL the longest, AM and AL subequal; sensillae some- 
what thickened medially, and then tapering, with ciliations for most of their 
length. Byes 2 + 2. Chelicerae non-serrate, Galeal setae short and nude. 
Palpi as figured, femur outwardly rather angular, and seta branched; palpal 
claw trifurcate. Dorsal setae tapering, curved and shortly ciliated, to 36/x 
long, 32 in number and arranged 2.6.6,6.6,4.2. Ventrally with a pair of branched 
setae on maxillae, one on each coxa, a pair between coxae I and between coxae III 
and thereafter ca. 40 setae to 30/* long. Legs all 7-segmented, I 260/* long, 
II 234/x, III 260/x; tarsi I and II with dorsal sensory rod, III without any long 
nude seta. 

The Standard Data derived from ten specimens are as follows: 



AW 
PW 
SB 


Mean 
47-9±:0-31 
61-9±0-98 
19-6 


Standard Theoretical 
Devjation Range 

0-93:±0-22 45-1-50.7 

2-95^:0-69 53*1-70*7 

No variation recorded. 


Observed 
Range 

47-6-50-4 
58-8-67-2 


Coeft. i 
Variati 

1-9 

4-7 


ASE 
PSB 
SB 


30*2±0-37 

14-0 

44-2±0-37 


l-18:£0-26 26-7-33-7 
No variation recorded, 
l-18±0-26 40-7-47*8 


28-0-30-8 
42-0-44-8 


3. 9 
2-6 


A-P 


38-l:h0-85 


2- 70+0-60 


30-0-46-2 


33-6-42-Q 


7-1 


AM 


41-1:1:0*59 


l-44±:0-42 


36-8-45*4 


39-2-42-0 


3-5 


AL 


42- 35=fc0 -35 


0-P9:±0-25 


39-4-45-3 


42-0-44-8 


2-3 


PL, 

Sons, 


47-6 No variation recorded. 

68-6 Mean of two determinations 67 • 2 and 70-0. 







Womersley— Asiatic-Pacific Scrub Typhus Mites 145 

Loc. and Host. Described from the type and niue paratypes collected from 
bats, R.11Q18, from Bukit Lagong Forest Reserve, Kepong, Kuala Lumpur, 
F.M.S. (British Scrub-typhus Research Unit, Slide 14765 (6 spec), 14766 (4 
spec), 1950). 

Remarks. In the shape and general proportions of the scutum, this spe- 
cies is very close to T. insotti Philip and Traub 1950, also described from 
bats, from Malaya, and was at first thought to be this species. It differs from 
insolU however, slightly in the Standard Data, and more particularly in the 
form of the sensillae, in having all the setae on the femur, genu and tibia of 
the palpi branched, and in the fewer dorsal setae. 

It is named after Mr. J. L, Harrison, ecologist to the S.T.R. Team. 

The larvae were also reared through to the nymph (described elsewhere in 
this paper), by Mr. K. L. Cockings. 

Genus TECOMATLAtf A Hoffmann 1947. 

Annales Escuela Nacional de Ciencias Biologicas, 4, (4), 451-457, fig. 1-6 

(Genotype Tecamatlana^sandovalin, sp.) 

~=S<mriscus Lawrence 1949, Annals Natal Museum, 11, (3), 452, fig. 39 a-e 

(Genotype Sauriscus ewingi n. sp.) 

= Trisetica Traub and Evans 1950, J. Parasitol., 36, (4), 356-359, fig. 1-4 

(Genotype Trisetica melvini n. sp.) 

The genus Tecomatlana was based on a species from Mexican bats and 
characterized as follows; 

il Palpal claw with four prongs, the middle one being much the largest. 
Chelicerae with a row of minute reflected teeth on the dorsal and apical region 
of the chela and with a single, obtuse, broad and little tooth on the ventral 
margin. Dorsal plate small, roughly pentagonal with only three setae in addi- 
tion to the pseudostigmatic organs, which are flagelliform and filamentous. 
Eyes large in relation to other genera, the posterior ones being smaller. All 
tarsi with three more or less equal claws. 

Differs from all other genera of the subfamily Trombiculinae in its dorsal 
plate, which bears only three setae, in addition to the pseudo-stigmatic organs 
and in the particular arrangement of the palpal daw". 

Miss Hoffmann does not refer to the postero-lateral scutafl setae, which 
are present, but due to the small and reduced scutum are not placed thereon, 
as is clearly shown in her figure 2. They are, however, the first row of 1 + 1 
setae, in her series of dorsal rows. The description of "a row of minute 



146 Records of the S.A. Museum 

reflected teeth on the dorsal and apical region of the chela' ' at first suggests a 
serrated chela. The author's figure 3 shows clearly, however, that the minute 
teeth are confined to the lower edge of the usual tricuspid cap, and are not the 
normal serrations on the inner cheliceral margin seen in some other Trom- 
biculids. 

The only valid character by which Tecomatlana can be generically separ- 
ated appears to be the very much reduced scutum causing the postero-lateral 
setae to be widely situated off the scutum. Until such times as the nymphs or 
adults are known and correlated with the larvae the genus should perhaps 
stand on the above feature. It must be pointed out. however, that an indica- 
tion of such displacement, although slight, is clearly seen in the Malayan 
species Sclwngastia (Ascoschongastia) maktyensis Gater; and in Schdngustia 
(Ascoschontfastia) masta Traub and Sundermeyer 1950, the posterior lateral 
scutal setae are as far removed from the scutum as in Tecomatlana i etc. 

The four-pronged palpal claw on which Miss Hoffmann lays stress is not 
of generic value, as is evidenced by the varied furcation of this organ, occur- 
ring within other genera, and as- shown in the keys here given. 

In 1949, R. F. Lawrence, overlooking Tecomatlana, described the genus 
8a%riscits, with similar scutal characters, for a species Sawiscus eivmgi nov. 
from a gecko from S. Africa. The only character by which this might be 
generieally distinct is the presence of but a single large eye on each side 
instead of two. The posterior eyes in many Trombiculids are frequently ill- 
defined, and difficult to see, so that one may be doubtful of the generic value 
of the number of eyes. Lawrence does not state the nature of the palpal claw. 

More recently, Traub and Evans have described the genus Trisetica, for a 
species found inhabitating bat caves at Myitkyina, in Burma. They have 
apparently overlooked Lawrence's paper, and only compare their genus with 
Tecomatlana. 

The scutum is similar to that of Tecomatlana and Sawiscus, being reduced 
with the postero-lateral setae well off the scutum. The palpal claws are con- 
trasted with those of Tecomatlana in being trifurcate and somewhat long and 
straight. The apical tricuspid cap of the chelicerae lacks the minute teeth on 
its lower edge. The eyes are two on each side and large. 

It appears then that both Sauriscus Lawrence, and Trisetica Traub and 
Evans are synonymous with Tecomatl-ana Hoffmann. Incidently, the some- 
what thickened or ' ' subflagellate" sensillac of the type of Trisetica approach 
the lanceolate form found in some species of Sclwngastia, but its affinities are 
I believe nearer to Tecomatlana and Tromlicula than to Mekong astia. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 147 

Tecomatlana melvini (Traub and Evans 1950). 
Trmtica inelvini Traub and Evans 1950, J. Parasitol , 36, (4), 356-359, fig. 1-4. 

Plate 29, fig. E-H. 

Description of Larvae (after Traub and Evans). "Body subovate; dorsum 
and venter with fine parallel striations; length 0-54 mm.; width 0-38 mm. 

Dorsal plate. Anterior margin slightly sinuate, concave at the insertion of 
the anterolateral bristles, forming shoulder?!; lateral margins with a shallow 
median sinus; caudal margin very slightly concave, corners strongly rounded; 
punctate over most of its surface. Sensillae with short barbs on proximal por- 
tion, but apical two-thirds with longer cilia. Bases of sensillae inserted 
slightly posterior to midline of plate. Anterolateral setae short and plumose. 
Anteromedian bristle extending beyond apex of plate plumose. Posterolateral 
setae as far off plate as bases of sensillae are from margin % near eyes ; similar 
in shape and size to antero-median bristle. Eyes: Well-developed; anterior 
eye with diameter of 17 microns; that of posterior eye 10 microns. Cheliceroe: 
Acuminate, about four times as long as broad near base; with a single sub- 
apical notch. Palpal claw i with lateral prongs almost as long as middle prong. 
Palpal tarsus: Longer than broad at base; with four ventral bristles, three of 
which are sparsely plumed and the fourth shorter and virtually nnbarbed; 
with a proximal ventral short spur-like process ; with a dorsal long bristle which 
is apically plumed. Palpus : Trochanter and femur each with a dorsal sparsely 
plumed bristle; tibia with a dorsal naked bristle and with a ventral short 
branched seta. Galea i "With a pair of plumed proximal ventral setae. Dorsal 
setae : Similar in appearance to anterolaterals ; about 52 in number ; the rows 
irregular, but approximate arrangement of the anterior rows is: 2 j 12 : 12 : 10. . . 
Coxae-. Punctate; unisetose, the setae plumose. Sternal setae: A pair of plu- 
mose setae between bases of coxae I and II j three snch pairs between coxae III r 
the middle one of each group of three more posterior than the others. Ventral 
setae: Approximately 36 in number; short and irregularly arranged. Legs: 
I — 0-27 mm.; II— 0-23 mm.; Ill — 0-25 mm. Sensory setae as follows: I — two 
genualae, one microgenuala, two tibialae, one mierotibiala, one spur, one micro- 
spur laterad to spur, one pretarsaia. I — one genuala, two tibialae, one spur, 
one microspur slightly proximad to spur, one pretarsaia. Ill — one genuala, 
one tibiala. Tarsal daws: Elongate, acuminate, curved, resembling the blade 
of a scythe; middle claw slightly thinner than others/ } 

The Standard Data given for four specimens by the authors show consid- 
erable variation as follows: 



148 Records of the S.A. Museum 







standard 


Theoretical 


Observed 


CoefF. of 




Mean 


Deviation 


Uang^ 


Range 


Variation 


AW 


43-75±l*03 


2-06±0-73 


37-6-49-4 


42-0-46-0 


4-7 


PW 


66-75±5»22 


10-44+3-69 


35-45-98-05 


56*0-81-0 


15-7 


SB 


18-5±0*87 


l-73±0-61 


13-3-23 -7 


17-0-20-0 


9-3 


A SB 


23-75:£0-75 


l*50:fc0-53 


19-25-28-25 


23-0-26-0 


6-3 


FSB 


17-0 


Ko variation recorded. 






8D 


40-75±0-75 


l-50±0«53 


36 -25-45 -25 


40-0-43-0 


3-7 


A-P 


33-0:£2-86 


5-71^:2*02 


15-9-50*1 


26-0-40-0 


17-5 


AM 


36-7±l*76 


3'05±:l-24 


27-5-45-9 


34-0-40-0 


8-3 


AL 


29-0±l-78 


3-55:tl-26 


18-5-39-5 


26-0-33-0 


1-2 


PL 


43-75±0-75 


l-50±0-53 


39-25-48-25 


43-0-46-0 


3*4 


Sens. 


53 -75:1:1-43 


2-87 ±1-01 


45-15-62-35 


50-0-56-0 


5-3 



Genus MYOTROMBICULA Womeraley and Heaslip 194S. 
Trans. Roy. Soc. S. Austr., 1943, 67, (1), 99. 

This genus erected for a specimen found amongst the debris in a jar of 
spirit containing bats in the collections of the South Australian Museum, the 
probable locality for the bats being South Australia. It was assumed that the 
specimen came off the bats. 

The genus is peculiar in that the chelicerae are short and stumpy "with 
two large blunt teeth, apparently an adaption for grasping hair. The palpi 
are also of peculiar structure, both the femur and genu being strongly and 
outwardly angulate. The tibial claw of the palp is not furcate, but on the 
outer edge is provided with two small teeth. The dorsal scutum is rectangular, 
provided with the normal five setae and two ? filamentous sensillae. The legs 
are all 7-segmented. 

Genotype Myotrombicula vespertilionis Worn, and Heasp. 1943. 

Myotrombicula vespertilionis Worn, and Heasp. 1943. 
Trans. Roy. Soc. S. Austr., 1943, 67, (1), 99, 

Plate 30, fig. A-F. 

Shape almost round. Length 275/*, width 280fx> Dorsal scutum trans- 
versely rectangular (cf. fig. A) with anterior margin concave, posterior evenly 
convex, and SB in front of line of PL ; sensillae wanting, probably filamentous. 
Eyes apparently absent- Chelicerae short and stumpy with two blunt and 
strong teeth, as though for grasping hair (ef. fig. C). Galeal setae branched on 
distal fifth. Palpi stout, strongly incurved and both femur and genu out- 
wardly strongly angulate; tibial claw not furcate but with two small teeth on 
outer margin; setae on femur and genu strong, curved, with short indistinct 
eiliations- on the tibia, the dorsal seta is similar, the lateral apparently nude 
and the ventral with many long branches; the tarsus is short, with sub-basal 



Womersley— Asiatic-Pacific Scrub Typhus Mites 1+9 

sensory seta, subapical sensory nude seta, and 4 or 5 ciliated setae. Dorsal 
setae 2.6 (8). 8 (6). 8.4.8.6.4.2, to 23/i long, tapering with short dilations. Ventral 
with paired branched setae on maxillae, one on each coxa, a pair between 
coxae I and between coxae III and thereafter 2.4.4.6.6.4.2. Legs: all 7-seg- 
mented, I 170/a, II 145/^ III 145/i.; tarsi I and II with dorsal sensory rod, III 
without any long nude seta. 

The Standard Data for the unique type are: AW 62-5, PW 65-5, SB 
240, ASB 16-0, PSB 11-0, SD 27-0, A-P 16-0, AM 23-0, AL 19-0, PL 27*0, 
Sens. — . 

Remarks. The modifications of the chelicerae and palpi suggest their 
adaption for grasping hair, although the modifications are not as pronounced 
as in the other pilicolous families Myobiidae and Listrophoridae. 

Genus HEA8LIPIA Ewitt* 1944. 

BeasUpia Ewing 1944, Proc. Biol. Soc, Washington, 57, 101-104, nom. no-o. 
for Trombiculoid.es Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 
67, (1), 101, preoc. Jacot 1938, Psyche, 45, (2-3), 122. 

As Trombwuloides Worn, and Heasp. 1943 with Tromhimloides gateri 
Worn, and Heasp. 1943 as genotype was preoccupied by Trombicnloides Jacot 
1938 with Say's American Trombidium scabram as genotype, the name Beas- 
Upia-vta* proposed by Ewing (loc. tit.) in its place. 

The genus is closely related to Trombicula s. str. in having only a single 
and anterior dorsal scutum with long filamentous seusillae, but differs in the 
characteristic scutum which has the lateral and posterior margins forming a 
continuous even curve, and is furnished with 9 setae in addition to the sensillae, 
arranged 1 AM, 2 AL and 2 PL with two extra ones on each side between the 
AL and PL setae. The chelicerae are also characteristic with a very large and 
pronounced inner subapieal tooth in addition to the small apical tricuspid cap. 
The legs are all 7-segmented. 

Heaslipia gateru (Womersley and Heaslip 1943). 

Trombicnloides gateri Worn, and Heasp. 1943, Tr. Roy, Soc. 8, Austr., 67, 
(1),101. 

Heaslipia gateri \ Ewing 1944, Proc. Biol. Soc. Washington, 57, 101-104; Law- 
rence 1949, Annals Natal Museum, XI, (3), 467. 

Plate 30, fig. G-L. 

In addition to the details given in the original description of the larval 
species, the seta on the palpal femur is 5-branched, on the genu 2-3 branched, 



150 



Records of the S.A. Museum 



and on the tibia, the dorsal and lateral setae are nude, the ventral branched; 
the palpal tarsus is short and conical with sub-basal sensory rod and sub- 
apieal nude sensory spine, and 5 or 6 ciliated setae. AH legs are 7-segmented s 
tarsi I and II rather short with the usual dorsal sensory rod, and tarsi III is- 
furnished with a long nude outstanding seta. The palpal claw is bifurcate. 

Since the original description the dorsal scutum of the type and 2 para- 
types have become more flattened and consequently the Standard Data are 
now somewhat greater. 

The Standard Data as now re-determined for these specimens, together 
with one specimen from the Philippines are as follows : 





Mean 


Standard 
Deviation 


Theoretical 
Ranvje 


Observed 
Range 


Coeff. of 
Variation 


AW 


88* 75+1-70 


3-40+1*20 


78-55-98*95 


86-G-03-O 


3-8 


AWi 


107-0+1-00 


2-00±0-71 


101-0-113-0 


104*0-108 '0 


1-9 


AW2 


97-0+0-82 


1*63+0-58 


92-1-101-9 


95-0-99-8 


1*7 


PW 


74-25+1-44 


2-87+1- 01 


65*65-82-85 


72-0-78-0 


3-8 


MAXW. 


116-5+0-87 


1-73+0-61 


111-3-121-7 


115-0-118-0 


1-5 


SB 


33-0+1-00 


2-00+0-71 


27*0-39-0 


32*0-36-0 


6*0 


ASB 


29-0 


No variation recorded. 






PSB 


39-25+0-25 


0-50+0-18 


37-75-40-75 


39-0-40-0 


1-3 


SD 

Alr-ALl 

AL1-AL2 


68-25+0-25 
14-50±O*5O 
18-25+0-63 


0-50+0-18 
1-00+0-35 
1-26+0*44 


66.75-69-75" 

11-5-17-5 

14-5-22-0 


68-0-69-0 
14-0-16-0 
17-0-20-0 


0-7 

7-0 
6-9 


AL2-PL 


21-0 


No variation recorded. 






AM 


67-7+1*26 


1*52+0 -89 


60-2-75-2 


65-0-70-0 


3-7 


AL 


51-0±l-00 


2-00+0-71 


45*0-57-0 


50-0-54-0 


3-9 


ALi 


920+1-15 


£-31+0-82 


85-1-98-9 


90*0-94-0 


2-5 


AL8 


72-25+1-03 


2-06+0-73 


66-05-78-45 


70-0-75-0 


2*8 


Pli 


58-25+1-70 


3 -40 -£1-20 


48*05-68*45 


54*0-61-0 


5-8 


Sens. 


93-0+1-12 


2-24+0-79 


86-0-99-7 


90*0-95-0 


2*4 



The additional measurements of the Standard Data as used above are: 

AW 1 = distance between setae AL, i.e. the second pair of lateral setae. 
AW 2 = distance between the third pair of lateral setae, i.e. AL. 
Max. W. = maximum width of scutum. 
AL-AL 1 = distance between first and second lateral setae. 
AL^AL 2 — distance between second and third lateral setae. 
AL 2 -PL — distance between third lateral and postero-lateral setae. 
AL 1 and AL 2 = lengths of second and third pairs of lateral setae respec- 
tively. 

Remarks. As the nymphal or adult stages of this species are unknown, 
its generic relationship with Trombicula s. str. is uncertain, but on the peculiar 
dorsal scutum and chelieerae in the larvae, a separate genus seems at present 
advisable. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 151 

This species, the genotype, was described from specimens from Battue 
raitus argentivenier Chase, from Selinsing-Gunong, Seinanggcl, Perak, 15 
March 1932, and sent to the Adelaide Museum for study from the Institute for 
Medical Research, Kuala Lumpur, F.M.S. 

Lt-CoL Philip's specimen was from the ear of a rat, at San Jose, Mindoro. 
Philippine Island (coll. J. R. Mangrum, Jan., 1945). 

Ctenus NOVOTB.OMBICULA Womersley and Kohls 1947* 

Tr. Roy. Soc. S. Austr., 1947, 71, (1), 4. 

Larvae as in Trombicula, but with the dorsal scutum produced posteriorly, 
tongue-like, and taking in some of the median setae of some of the dorsal rows, 
as in the genus Qahrliepia, otherwise with the usual 5 setae and a pair of 
filamentous sensillae. 

Genotype Novotrmnbicula owiensis Worn, and Kohls 1947. 

NcyvoTROMBiouLA owiensis Woni. and Kohls. 1947. 
Tr. Roy. Soc. S. Austr., 1947, 71, (1), 4. 

Plate 31, fig. A-H. 

Shape, subrotund. Size small; length (unfed) 255/x, width 180/*. Dorsal 
scutum large, tongue-shaped and produced posteriorly to take in the two median 
setae of the third dorsal row, in addition to the AM, AL and PL setae; sen- 
sillae long and filamentous with ciliations on the distal half, bases wide apart 
and about midway between line of AL and PL. Eyes 2 -\- 2, on distinct ocular 
shields placed close to the lateral scutal margin; posterior eyes the smaller. 
Chelicerae with only the usual apical tricuspid cap, but on the inner margin 
below the cap with rather prominent subapical angle. Galea! setae long and 
nude. Palpi rather slender, with rather long slender and bifurcate tibial claw, 
the dorsal prong of which is small and indistinct; palpal femur and genu 
with a long slender, shortly ciliated seta; all three setae on tibia long and 
apparently nude; tarsi moderately long and conical, with a long sub-basal sen- 
sory rod and five or six long ciliated setae. Dorsal setae rather thick, slightly 
tapering, shortly ciliated and of two sizes, arranged 2.4.4 [2] .4.2.2; the second 
row with the median pair just off the scutum and to 57//, long; the inner 
members of the third row are on the scutum and are short to 24/x. Ventrally 
with a pair of branched or ciliated setae on the gnathosoma, one on each coxa, 
a pair between coxae I and between coxae III and thereafter 6.2.2.2.2. to 24^ 
long. All three pairs of coxae touching. Legs: I 255/* long, II 240/i, III 275/1 ; 
tarsi I and II with the usual sensory rod-like seta, III without any long nude 
seta. 



152 Records of the S.A. Museum 

Genus SCHOENGASTIA Oudemans 1910. 

Entom. Ber., 3, (54), 96, 1910. 

Larval Trombieulidae, with a single anterior dorsal scutum furnished with 
5 normal ciliated setae and a pair of clavate or capitate sensillae which arise 
from cupules inserted flat on the scutal surface and not in a transverse crest- 
like wall. Chelicerae not unusually slender, with the inner (dorsal) edge 
serrated or not. Palpal claw bifurcate or trifurcate. All legs 7-segmented, 
except in S. (SI) audemarm (Walch), 8. (A.) heasUpi Worn, and Heasp., and 
3. {A.) iraribi sp. n., where leg II and III are 6-segmented. 

Genotype Thromiidimn vandersamdei Ouds., 1905, Entom. Ber. r 
1, (22), 216. 

The chelicerae in the genotype are serrated and this led the American 
authority, Dr. H. E. Ewing, to erect the genus Ascoschdngastia Swing 1946 for 
those species, earlier placed in Neoschongastia Ewing 1929, which had the 
chelicerae non-serrate and at the same time were without the transverse scutal 
crest of Neoschongastia. As stressed in the section of this paper dealing with 
the adults and nymphs, species of Schongastia and Ascoschdngastia in which 
those stages are known cannot be separated generically. In this regard it 
would appear to be unwise to treat Ascoschdngastia as more than a subgenus of 
Sclwngastia on the basis of larval characters only. Similarly, although 
unknown as yet from the adult or nymphal stages, the genera Euschongastia 
Ewing 1938, Oenoschongastw Womersley and Kohls 1947 and Radfordmna 
nov. may ultimately be shown to be no more than of subgeneric status. 



CHELICERAE SERRATED ON INNER (DORSAL) MARGIN. 

Subgenus SCHOENGASTIA 8. str. Ewing 1946. 

1. Legs I 7-segmented, II and III 6-segmented. 
Schongastia (Schongastia) oudemansi (Walch 1923). 

Tronibicula oudemansi Walch 1923, Kitasato Archiv. Exper. Med., 5, (3) ; 
Fletcher, Lesslar and Lewthwaite, 1928, Tr. Roy. Soe. Trop. Med. and 
Hygiene, 22, 161. 

Schongastia oudemansi, Gater 1932, Parasitol., 24, 154; Womersley and Heas- 
lip, 1943, Tr. Roy. Soc. S. Austr., 67, (2), 102. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 153 

Neoschongastia impar Gunther 1939, Proc. Linn. Soc. N.S. Wales, 64, (1-2), 
85: Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 116; 
Radford 1942, ParasitoL, 34, (1), 74. 

ftcoschongustia lodensis Gunther 1940 ? Proc. Linn. Soc. N.S. Wales, 65, (5-6), 
482, 

Plate 32, fig. A-E. 

In 1943 Womersley and Heaslip synonymized Gunther 's bodensis from 
Borneo with his impar from New Guinea. The latter species, while agreeing with 
oudemansi of Walch in the form of the dorsal scutum, the dorsal setae and 
the lengths of the scutal setae, was considered to differ in the chelicerae not 
being serrated. On the last character it was placed in the genus Nc&schbngasiia 
as then understood, but in view of E wing's restriction of this genus, impar 
would now become a species of Aseoschongastia if the non-serration of the 
chelicerae did occur in this species. 

In correspondence, however, Dr. H. S. Fuller has drawn my attention to 
the fact that in certain specimens determined by me as impar Gunther, in the 
collections of the British Museum (Nat. Hist.) careful examination show T s that 
the chelicerae are indeed serrated. This observation, which I was able to affirm 
while hi London in 1947, led me to re-examine all the material in the South 
Australian Museum collections of both impar Gunther and bodensis Gunther 
and to compare them with specimens of oudemansi Walch, from the Federated 
Malay States, determined by B, A. Gater. 

In all this material it can now be stated that the chelicerae arc definitely 
serrated and both of Gunther *s species must be synonymized with oudemmxsi 
Walch. Amongst the species of Schongastia from the Asiatic-Pacific region, 
oudemansi is particularly characterized, as in the key, by the almost rectangular 
scutum, furnished with comparatively short setae of which PL are the shortest, 
The sensillae are clavate, finely ciliated and the sensillae bases midway 
between the anterior and posterior scutal borders with a well defined inwardly 
curved line in front of each base. The palpal claw is bifurcate, and only the 
setae on the palpal femur and genu shortly and sparsely branched, almost 
nude. The dorsal setae are short, and arranged 2,6,6v6,6*4. Tarsi III are 
without any long outstanding nude seta. (In his description of impar (loc. 
cit* p. 86) Gunther states that tarsi III bear a long nude seta, but in the four 
paratypes I have been able to examine this is not so) . 

A remarkable feature of oudemansi (also of impar and bodensis) is that, 
the femora of legs II and III are not secondarily divided as is that of leg I, and 



154 



Records of the SA. Museum 



as are all three pairs of femora in all other species of Schongastia, except 
8. (A.) heasUpi Woin. and Ileasp. and S. (A.) traubi sp. n. 

The secondary subdivision of the femora however, is not a true articula- 
tion, and it seems unwise, at present, in view of our little knowledge of generic 
characters of the adult Trombiculidae to regard the non-subdivision of the 
femora of legs II and III of larvae of the above species as of more than sub- 
generic value. 

The following Standard Data is derived from 1 specimen from Federated 
Malay States, det. as midemansi by Gater, 4 paratypes of impar Gunther from 
Bulolo, N.G., 1 paratype of bod&nsis Gunther from Borneo, as well as 19 speci- 
mens from Lae, N.G, and Sansapore, D.N.G., previously identified as impar. 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


51-4±0G5 


3-27+0-46 


41-0-61-2 


48-0-57-6 


6-3 


PW 


66-l±0-47 


2-34+0-33 


59- 1-73 »1 


64-0-73-6 


3-5 


SB 


30-65±0-37 


l-85±0-26 


25 -1-36 '2 


28-8-35-2 


6-0 


ASB 


25-6 




No variation recorded. 




PSB 


19 -45:£0 -15 


0-74+0- 10 


17-25+21-65 


19-2-22-4 


3-8 


SD 


45-2±0-19 


0-94+0- 13 


42-4-48-0 


44-8-48-0 


2-1 


A-P 


34-0+0-15 


0-73+0-10 


31-8-36-2 


32-0-36-8 


2-2 


AM 


34-7±0-33 


1-20+0-23 


31-1-38-3 


32-0-35-2 


3-5 


AL 


22-65+0-25 


1-25+.0-18 


18'9-26-4 


19-2-25-6 


5-5 


PL 


l3-55±0-22 


1-12+0-16 


10-2-16-9 


12 • 8-16 '0 


8'4 


Sens, 


38-0 with head 


•6 wide. Only 1 determination. 







SCHONGASTIA (SCHONGASTIA) LEWTHWAITEI Sp. n, 

Ascoschongastia n, sp. *C\ Lawrence, T. J., 1945. W. O. Kept., A.M.D.7, 
Appendix 1. 

Plate 111, fig. A-C, 

Description of Larvae. Shape (engorged) oval; length to 536/4,, width to 
375/t. Scutum as figured, sub-quadrate but wider than long, and with PW 
greater than AW; anterior margin lightly sinuous, posterior margin not very 
deep behind line of PL and lightly concave medially; sensillae bases large 
within strongly marked crescents, and much nearer to PL than to AL; sen- 
sillae unknown ; AM, AL and PL setae subequal with AM slightly shorter than 
AL, and PL slightly shorter than AL. Eyes 2 + 2, posterior the smaller. 
Chelicerae with apical tricuspid cap and the inner edge finely but indistinctl} 7 " 
serrate. Galeal setae nude. Palpi as figured, tibial claw bifurcate; setae on 
femur ciliated, on genu ciliated, on tibia ventral branched, dorsal and lateral 
apparenty nude. Dorsal setae ca. 42 in number to 45^ long and arranged 
2.6.6.6.6.6.4.4.2. Ventrally with paired ciliated setae on maxillae, one on each 



Womersley — Asiatic-Pacific Scrub Typhus Mites 155 

coxa, a pair between coxae I and between coxae IIT and thereafter ea. 46 to 30j/. 
long. Legs: I 268/* long, II 241/x, III 308/a; I 7-segmented, II and III 6-seg- 
mented ; tarsi I and II with dorsal sensory rod, III without any long nude seta. 
The Standard Data derived from the type and 3 paratypes are : 







Standard 


Theoretical 


Observed 


Coeffof 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


68-0±0-80 


1-60±0-50 


63-2-72'8 


67-8-70-4 


2-3 


PW 


86-0±3-09 


6 -18=1:2 -18 


67-5-104-6 


80-0-92-8 


7-2 


SB 


38-4 




No variation recorded. 




ASB 


35-2±l-31 


2-Gl±0-92 


27-4-43*0 


32.0-38*4 


7-8 


PSB 


22-4 




No variation recorded. 




SD 


57-fc±l-31 


2-61±:0-92 


49-8-65-4 


54-4-60-8 


4-5 


A-P 


44-8±l-31 


2-61±0-92 


37-0-62- 6 


41-6-48-0 


5-8 


AM 


34-4±0-80 


l-60±0-56 


29-6-39-2 


32-0-35*2 


41 


AL 


35-2 




No variation recorded. 




PL 


36-0±0-80 


l'60±0-56 


31-2-40-8 


35-2-38-4 


4-4 


Sena. 




Missing. 









Loc. and Host. Amongst the material in Lawrence's "Type Box" of 
slides, in the London School of Tropical Hygiene and Medicine, which I have 
been permitted to study by Dr. J. R. Audy, are 4 slides, Nos. 37-40, labelled 
" Ascoschongastia n. sp. 'C, Lawrence in M.S. 5 ' and referred to as such in 
Appendix 7, to War Office Report, AMD 7, March, 1947. 

Lawrence records these specimens as 3 from the type host, a tree shrew, 
Tupaia lelamgeri belangeri Wagner, and 2 specimens only from Hydromys 
Jwmei. The locality was Palel, Taimi Rd., 30 miles south of Imphal, Manipur 
State, India, Nov. 24th, 1945. 

Slides Nos. 39 and 40 are now in the collection of the South Australian 
Museum. 

Remarks. This species in the conformation of the scutum closely resembles 
S. (A.) lacunosa Gater and was at first thought to be that species. In having 
the second and third pairs of legs, however, only 6-segmented and the inner 
margin of the chelicerae finely serrate it comes much closer to 8. (S.) oude- 
ntami, from which it differs in the Standard Data, especially the length of 
the PL scutal setae. 

2. All legs 7-segmented. 

a. Palpal daw bifurcate. 

Schongastta (Schongastia) vandersandei (Ouds, 1905). 

Thrombidriwm vandersandei Ouds. 1905, Entom. Ber., 1, (22), 216 j 1909 
Tijdschr. Entom., 52, 50. 



156 Records of the S.A. Museum 

Allotkrombidium vandersandti Ouds. 1906, Entom. Berl., 2, (28), 56-59. 

Thrombidmm van der Sander Ouds. 1906 7 Nova Guinea, 5, 106, 131, tab. 4, fig. 
59-66,- 1908, Tijdsehr. Entom., 51, 25. 

Mwrothrombidium vamlersandei Ouda. 1912, ZooL Jahrb., Suppl. 14, 45-62. 

Plate 32, fig. F-J. 

This species was originally described by Oudemans from specimens found 
on man by Dr. G. A. J. van der Sande, Physician to the Dutch New Guinea 
Expedition of 1903. 

In Nova Guinea, 5, Oudemans cites a large number of references to the 
presence of these mites, in New Guinea, Malay Archipelago and the Celebes. 
It is therefore remarkable that despite these leads, this species has not 
hitherto been recognized amongst the very large amount of material collected and 
studied during the war years. Most specimens have been referred by myself 
and other students to Schongastia blestowei Gunther and other species. Gunther 
1939 (Proc. Linn. Soe. N.S.W., 64, (l-2) 7 94) compared his species blestowei in 
detail with the data given by Oudemans for vandersandei, stressing particularly 
the facts that in vandersandei the palpal claw is bifurcate, in bltestmoei trifur- 
cate, and that the dorsal setae are 50 and 64 respectively. In addition he 
states " maxillary setae plain" in vandersandei, "with 4 long branches' 7 in. 
blestoum. This difference, however, does not exist. In Oudemans' species, the 
maxillary seta is as figured by Oudemans (1906) branched and, as stated, "of 
the same kind as the other coxal hairs". The main differences between these 
two species then, as brought out in the key, are the furcation of the palpal 
claws and the number of dorsal and ventral setae. 

The original description as given very fully in Nova Guinea 1906, is as 
follows: "Larvae. Length 200-300^, measured on 40 specimens. Colour scar- 
let ; in spirits colourless. Shape of newly-hatched larvae short, scarcely longer 
than wide, with small shoulders between legs I and II. and big rounded ones 
above legs III,- abdomen posteriorly rounded. Textnre of epistome, dorsal 
shield, palps and legs smooth, finely porate, of remainder skin of body finely 
wrinkled. These wrinkles run almost parallel to the transverse rows of dorsal 
hairs. Dorsal side. Dorsal shield subhexangular, with one anterior, one pos- 
terior and four lateral sides, wider than long; anterior side the longest and 
straight; the 5 remainder sides almost equal in length; posterior side slightly 
concave. Eyes. Aside of the shield a double eye, of which the anterior one is 
distinctly semi-globular. promineut and less light-refracting. Pseudostigmata 
almost in the centre of the shield, round, cup-shaped. Pseudostigmatic organs 
(sensillae) clavate; peduncle and head almost equal in length; peduncle rod- 



Womersley— Asiatic-Pacific Scrub Typhus Mites 157 

like, thin, smooth; head swollen, distally rounded, proximally rapidly tapering 
towards the peduncle, seen from aside almost equal in width and length { seen 
on the top elliptical; yellowish brown with greenish hue; contents oily, very 
light refracting; surface of head examined under oil-immersion-system 6uely 
villous. Hairs. On the anterior margin of the shield one median hair. la 
each anterior and in each lateral corner of the shield a hair. On each shoulder 
above legs III a hair. On the abdomen 5 transverse rows of ten hairs each. 
All the hairs are slightly curved, except the above-mentioned median hair which 
i* straight. The median hair scarcely hairy; the 4 other shield hairs are bettor 
haired j all the other dorsal hairs are more or less serrate. 

Ventral side. Behind the rostrum there is a collar-shaped shield, widest 
in the centre. Coxae I triangular, with posterior external angle excavate to 
receive the "Urtrachee" which is round. Coxae II longer, internally rounded. 
Coxae III almost quadrangular » internally rounded. Anus ventral, nearer the 
posterior abdominal edge than the coxae III. Hairs. Coxal hairs thin, a little 
curved, beautifully pectinate with from three to five smaller hairs on the convex 
side. Between the proximal ends or angles of coxae I a pair of ditto. Between 
coxae III a pair of ditto. Behind this pair a single hair of stiff, slightly curved 
and externally serrate hairs resembling those of the dorsum. Further the 
whole venter is provided with hairs of the same kind, arranged in about four 
or five irregular transverse rows of about ten hairs each. 

Epistome, pentangular, provided with a fine median line. Mandibles dor- 
sally and internally slightly serrate. 

Maxillae (palpi). Coxae with coxal hair close to the trochanterofemur, of 
the same kind as the other coxal hairs. Hypostome ventrally smooth, distally 
scarcely bilobate, dorsally opened, gutter-shaped and basal ly provided with long 
smooth hair.* No trace of any epipharynx, nor of amj hypophwynx. Troch- 
anterofemur dorsally and externally swollen, with proximal pectinate hair. 
Genu short with dorsal pectinate hair. Tibiotarsus dorsally with one smooth 
hair and one external pectinate hair, terminally with strong claw which 
externally and ventrally shows a smaller accessory claw almost in the 
middle. Telotarsus short, thick, club-shaped, ventrally with three pec- 
tinate hairs and two olfactoric hairs, dorsally with one pectinate hair and one 
olfactoric hair. Legs rather thick; their joints rather short, increasing in 
lenpfth distalward. Also the number of hairs with which they are omated 
increased distalward, from one or two to about ten or twelve on each joints The 
greater part of these hairs are pectinate. The following not pectinate hairs may 
be mentioned : Legs I i genu with short smooth hair ; tibia with 9 short smooth 
hairs; tarsus with a thick, short, rod-like olfactoric hair and a long smooth tac- 

* f 'GaleaI seta" of recent authors. 



158 Records of the S.A. Museum 

tile hair, accompanied by a very short smooth olf actoric hair. Legs II : genu 
with one short smooth hair ; tibia with one short smooth hair ; tarsus with one 
short thick olfaetoric hair. Legs III: genu with one short smooth hair;. tibia 
with one short smooth hair. All the femora are distinctly divided in a basi- 
fenmr and a telpfemwr. Paratwsi minute, almost invisible. Empodium claw- 
like, thin proximally and distally nodded. Claws rather strong, curved, thicker 
than claw-like ejapt^Smni^^. 

In the above description it is to be noted that the long nude outstanding 
seta present on tarsi III of all the specimens now referred to Oudemans species, 
is neither mentioned nor figured. This is remarkable considering the meticu- 
lous work of Dr. Oudemans, but this seta is however, when lying longitudinally 
along the tarsus sometimes difficult to see, and one must presume that Oude- 
mans may have overlooked it. 1 Its closest relative amongst those species of 
Scho?igastia s. str. with bifurcate palpal claws is S. pseudo-schuffneri (Walch), 
which is distinguished as in the key by the shape and dimensions of the dorsal 
scutum, and the number and arrangement of the dorsal setae. 

The following Standard Data is derived from 63 specimens now recorded 
from various localities in New Guinea : Abidari, Mandated Territory, 28 July, 
1943 (1 spec. R. N. McCulloch) ; Dumpu, M. % Nov. 1943 (5 spec. ft. N. McC.) ; 
Lae, M.T., 8 Aug. 1944 (6 spec. B.N.McC.) ; Buna, M.T. (2 spec. No. 32-1, 3 
spec. No. 32-5, G. M. Kohls) j Milne Bay, Papua, Aug. 1943 (5 spee. S. L. All- 
man ) y Nov. 1943 (5 spec, on Rattus contains, W. D. Exton) j Hollandia, 
D.K6., 2 Mar. 1945 (2 spec, on boots, C. B. Philip), (3 spec, on man, C.B.P.), 
22 May, 1945 (6 spec, on man, C.B.P.) ; Sansapore, D.N.G., 10 Nov. 1944 
(5 spec, on Rattus contolor, C. Mohiv), 25 Nov. 1944 (3 spec, on Rattus ringens, 
1 spec, on R. concolor, CM.). 

Standard Theoretical Observed Coeff , of 

Mean Deviation Range Range Variation 

AW 66-9±0-41 3-30=£0-29 57-0-76-8 60-8-76-8 4-9 

PW 88-6±0-57 4-54±0-40 75-0-102-3 80-0-99-2 5-1 

SB 25-SrtO*10 0«3=t0-07 23-3-28-3 24-0-28-8 3-2 

ASB 35-5:t0-12 0-95:1:0*08 32-6-38-4 35*2-38-4 2-7 

PSB 28-7=t0-09 0-72±0-06 26-5-30-9 25*6-32-0 2-5 

SD 64-1^0-15 1-17±0-10 60-6-67-6 60-8-67-2 1-8 

A-P 31-7^014 l-13dfc0-10 28-3-35*1 28-8-35-2 3-6 

AM 37-3±0-22 l»63d:0-15 32-4-42-3 35-2-41-6 4-4 

AL 77-8±0-41 3-26^:0*29 68-0-87-6 70^4-83-2 4-2 

PL 58-1^0-22 2*57=fc0-23 50-4-65*8 54-4-64-0 4-4 

Sens. 32-0 with head 16/19. No variation recorded. 



l In a recent letter from ray colleague, Dr. G. L. van Kyndhoven I am indebted for the 
information that an examination of the Oudemans ■ type slide in Leiden which also contains a 
specimen of S. schuffneri (Walch) (id. H. S. Fuller) phows that on the tarsi of leg III of 
vandersandei there is a long nude outstanding seta. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 159 

The number of DS given by Oudemans is 52, but they vary somewhat, 
especially posteriorly, from about 48 to 54. They measure ca. 40/* long, while 
the ventral setae which number ca, 36, posterior of coxae III measure to» 
30/t in length. Oudemans only refers to one smooth and one branched hair 
on the palpal tibia, but as in all species of Trombiculidae there are three setae 
on this segment, a dorsal and a lateral (both nude) and a ventral which is 
the branched hair mentioned by Oudemans. 

Schongastia (Schongastia) schuffneri (Walch 1923) 

Trambicula schuffneri Walch 1923, Kitasato Archiv. Exper. Med., 5, (3) ; Tr. 
Vth, Bien. Cong. Far East. Assoc. Trop. Med., Singapore, 1923 (24). 

Neoschongastia schuffneri, Womersley and Heaslip 1943, Tr, Roy. Soc. 
S.Austr.,67, (1),117. 

Schongastia katoms Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 67, 
(1),104. 

Schongastia pusilla Womersley 1944, Tr. Roy, Soc. S. Austr., 68, (1), 96. 

Plate 33, fig. A-E. 

Through the kindness of my colleague Dr. G. W. Wharton I have now been 
able to study a slide of Walch 's original material of this species, which he 
received from Dr, H. E. Ewing who himself had received it from the Koloniaal 
Institute in Amsterdam. The slide was labelled "in virgin forest, Sumatra 
1922". In this specimen the chelicerae are serrated and the sensillae globular 
as in the genus Schongastia. The dorsal setae are arranged 2.8.2.8.8 .! and 
measure to 40/* long. The Standard Data are: AW 54-5, PW 67-0, SB 21-0, 
ASB 28-0, PSB 22-0, SD 50-0, A-P 25-0, AM 33-0, AL 56-0, PL 42-0, Sens. 
32-0 with the head globular and minutely setulose. The above Standard Data 
do not differ significantly for those given by Womersley 1944 for 28 specimens 
of S. pimtta, and this species must therefore be regarded as a synonym of 
schuffneri Walch, as it also agrees in other morphological features. 

Schongastia haAonis Worn, and Heaslip 1943, erected for the specimen 
described and figured, but not named, by Kawamura and Yamaguchi 1921 
(Kitasato Archiv. Exper. Med. TV) is probably also to be referred to Walch 's 
species. 

This species is closely related to S. vandersmdei (Ouds.), and appears to 
be almost as common and widespread in New Guinea. It differs from vander- 



160 Records of the S.A. Museum 

satndei in the smaller scutum and the number and arrangement of the dorsal 
and ventral setae. 

As in the preceding species the AL scutal setae are the longest, the sen- 
sillae bases lie in line with or slightly in front of PL, the galea! setae are nude 
and there is a long nude outstanding seta on tarsi III. The dorsal seta are 
2.8.2,8.8,6.4, and the ventral setae 2,2-4.6.6.4.4. The setae on the palpal femur 
and genu are branched; on the tibia the dorsal and lateral nude, ventral 
branched. The palpal claw is bifurcate. All legs 7-segmented. 

In additon to the 28 specimens of which the Standard Data was recorded 
in 1944 (under the name S. pusitta) , the values for another 32 specimens from 
the following localities: Batisati, N.6., Aug. 1943 (R. N. McCulloch 8 speci- 
mens) ; Dobodura, N.G., Apr. 1943 (D. C. Swan, 7 spec.) ; Bat. Is., Purdy. 
Group, June 1944 (G. M. Kohls, 3 spec); Moratai, Celebes, 1945 (R. N. Mc- 
Culloch, 2 spec.) i Holiandia, D.N.G., Feb. 1945 (R. C. Bushland, 3 spec.) ; 
Sansapore, D.N.G., Nov. 1944 (C. Mohr, 4 spec.) ■ Noemfur, D.N.G., Nov. 1944 
(D. C. Swan, 5 spec), are as follows : 





Mean 


Standard 
Deviation 


Theoretical 
Eaiige 


Observed 
Eange 


Coeff . of 
Variation 


AW 


53.1±:0-49 


2-78±0-35 


44-7^1-5 


48 • 0-61- 2" 


5-2 


PW 


70-0:±0-36 


2-04+0-25 


63-9-76-1 


67-2-73-6 


2-9 


SB 


20-5+0-24 


1-34 ±0-17 


16-5-24-5 


19-2-22-4 


6-5 


ASB 


27 -0+0 -25 


1-41+0.18 


22-8-31-2 


25*6-28-8 


5-2 


PSB 


22-55±0-ll 


0-61+0-08 


20-7-24-6 


22-4-25-6 


2-7 


SD 


49-G±0-28 


1-60+0.20 


44-8-54-4 


48-0-54-4 


3-2 


A-P 


23-7±0-24 


1-38*0-17 


19-6-27-8 


20-8-25-6 


5-8 


AM 


27-8±0'37 


2-05+0-2G 


21-7-33-9 


25-6-32-0 


7-3 


AL 


54'4rt0-49 


2-77*0-35 


46-0-62-8 


48-0-64-0 


5-1 


PL 


43-l±0-58 


3-29*0-41 


33-2-63-0 


38-4-54-4 


7-6 



Sens, 32-0 with head 12-8/19-2. No variation recorded. 

SCHONGASTIA ( SCHONQASTIA ) TAYLOBI Gllllther 1940. 

Schongastia taylori Gunther 1940, Proc, Linn. Soc N.S. Wales, 65, (3-^), 257, 
fig. 12-14; Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 
106. 

Plate 33, fig. F-J. 

Like the following species, 8. taylori differs from vandersandti and 
schuffneri in having the sensillae bases placed well behind a line joining the PL 
scutal setae. It has, however, a roughly hexagonal scutum as in those species 
but the anterior lateral sides are much shorter than the posterior-lateral sides, 
i.e, A-P is relatively short and less than half as long as the scutal depth- In 



Womersley— Asiatic-Pacific Scrub Typhus Mites 161 

addition the dorsal setae are more numerous, ea. 80 and arranged ca. 2.10.2.13. 
2.14.12.12.8.4. 

As stated by Womersley and Heaslip (1943) Gunther gives PL as longer 
than AL and figures it so. This however, is not borne out by the examination 
of the type and 4 paratypes of Gunther 'a material, AL being the longest. 
The setae on the femur and genu of the palpi are well branched, on the 
tibia the dorsal and lateral are nude and the ventral branched. The galeal 
setae are nude. Tarsi III is provided with a long nude outstanding seta. 

This species was described from Macropus (Thylogale) coxeni Gray from 
Bulolo, N.G. No fresh material has since been seen, and the following Standard 
Data is derived from the type and 4 paratypes as recorded by Womersley and 
Heaslip 1943. 



AW 
PW 


Mean 
02-4±0-40 
79-0 


Standard Theoretical 
Deviation Range 

0-89±0-28 59-7-65-1 
No variation recorded 


Observed 
Range 

61-0-63-0 


OoeffrOf 
Variation 

1-4 


BB 
ASB 


21-2±:0-20 
35-0 


0-45±<M4 19*85-22-55 
No variation recorded 


21-0-22-0 


2-1 


PSB 


23-5 


No variation recorded 






SD 


58-5 


No variation recorded 






A-P 


23-5 


No variation recorded 






AM 


41-0±0-71 


1-41:4:0-50 36-8-45-2 


40-0-43-0 


3-4 


AL 

PL 


77-6±l-29 
57-0±l-22 


2-88±:0'91 69-0-86-2 
2-43±0-97 49-7-64-3 


73- 0-81-0 
55-0-60-0 


3-7 
6-6 



Sena. 35-0 with head 15/28. Only 1 determination. 

Schongastia (Schongastia) whartoni sp r n. 
Plate 34, fig. A-E. 

Description of Larvae. Shape roughly oval. Length (unengorged) 200/c, 
width 154/a. Dorsal scutum as figured, with anterior margin lightly sinuous, 
the lateral margins short and PL near to AL, posterior margin deeply and 
evenly rounded; AL the longest; sensillae globular, nude with their bases 
behind a line joining PL, Eyes 2 + 2, on ocular shields and adjacent to the 
postero-lateral scutal angles. Chelicerae fairly long with numerous teeth on 
the inner (dorsal) margin. Palpi fairly stout; tibial claw T bifurcate, prongs 
unequal; setae on palpal femur and genu branched, on tibia dorsal and 
lateral nude, ventral branched ; tarsus with subapical and sub-basal nude rod- 
like setae and 4 or 5 ciliated Setae, Dorsal setae 36 in number, arranged 
2.8,8.6.6.4, to 35fA long. Ventrally with the usual pair of maxillary setae, one 
on each coxa, a pair between coxae I and between coxae III, and thereafter 
2.6.4.4.4.2.2. to 22/* long. Legs I 230/i long, II 210/x, in 240/.; tarsi I and II 



{Standard Theoretical 


Observed 


Coeff . of 


Deviation Range 


Range 


Variation 


l*60±0-57 45*6-55-2 


48-0-51 2 


1-6 


No variation recorded 






1-53+0-54 L3-4-22-6 


16*0-19<2 


8-5 


No variation recorded 






No variation recorded 






No variation recorded 






1-85+0-65 8-8-20-0 


12*8-16-0 


12-8 


3-06*±l-09 25-2-43-6 


32*0-38-4 


9-0 



162 Records of the S.A. Museum 

with the usual dorsal rod-like sensory seta, III with a long nude outstanding 
seta. The Standard Data for the type and 3 paratypes are : 

Mean 

AW 50 -4+0*80 

P¥ 64-0 

SB 18 ? 0:±0-77 

ASB 25-6 

PSB 22-4 

SD 48-0 

A-P 14* 4+0 -92 

AM 34-4±l-53 

AL 56-8dr0-80 1- 60+0 -57 52-0-61-6 54-4-57*6 1*4 

PL 49*6+0*92 t-MM*66 44*0-55-2 48*0-51*2 1-9 

Sens, 32-0 with head 16/22. No variation recorded. 

Loc t The type and 3 paratypes taken on boots in the scrub at Lae, 
N. Guinea, March 1944 (R. N. McCulloch). 

Remarks. In having the sensillae bases well behind line of PL, A-P much 
shorter than the scutal depth, and the palpal claw bifurcate, this species is 
closely related to & taylori Gunther. It differs in the number of dorsal setae 
and the shape of the scutum. It is a pleasure to name it after Dr. G, H. Whar- 
ton, of Duke University, Durham, N. Carolina, U.S.A. 

b. Palpal claw trifurcate. 

SCIIONGASTIA (SCHilNGASTlA) BIDENTATA Sp. n* 

Plate 34, fig. F-J, 

Description of Larva. Shape oval. Length (engorged) to 355-x, width to 
255/*. Scutum pentagonal, strongly but sparsely punctate, the posterior por- 
tion tending to be overlapped by the striated dorsal cuticle j scutal setae rod- 
like, blunt tipped, with minute adpressed cilia tious; PL setae the longest,- sen- 
siliae clavate, ciliated, their bases about in line with PL. Eyes fairly large, 
2 + 2, on ocular shields adjacent to postero-lateral angles of scutum. Cheli- 
cerae, besides the usual tricuspid cap. with only two subapical teeth on inner 
margin. Galeal setae nude. Palpi stout; tibial claw trifurcate; all setae on 
femur, genu and tibia of palpi nude. Dorsal setae stiff and rod-like, blunt, 
with short adpressed ciliations, to 22/* long, 30-31 in number and arranged 
2.8 (9). 6.6.4.4. Ventrally with the usual pair of maxillary setae, one on each 
coxa, a pair between coxae 1 and between coxae III and thereafter 2.6.4,4.2.2.2., 
to 20-22/* long. Legs I 240/* long, II 224/*., Ill 248/*; tarsi I and II with a 



Womersley — Asiatic-Pacific Scrub Typhus Mites 163 

long nude outstanding seta. The Standard Data for the type and 7 paratypes 



are 





Mean 


Standard 
Deviation 


Theoretical 
Eange 


Observed 
Range 


Coeff. of 
Variation 


AW 


53*55:±0-62 


l-75±:0-44 


48-3-58-8 


54-4-57-6 


3-3 


PW 


69*7:±:0*46 


l-30±0-32 


65-8-73-6 


67-6-70*4 


1-8 


SB 


22-0:±0-40 


l*13:±0-28 


18-6-25-4 


19*2-22-4 


5-1 


ASB 


22-8±:0-40 


l-13*±0-28 


19-4-26-2 


22-4-25*6 


4*9 


PSB 


22*4 


No variation recorded 






8D 


45-2±0-40 


l-13th0-28 


41*8-48-6 


44-8-48-0 


2*5 


A-P 


26*8±0«58 


l-66r±0*41 


21-8-31-8 


25*6-28*8 


6-2 


AM 


16*0 


No variation 


recorded 






AL 


28-8 


No variation 


recorded 






PL 


32«0 


No variation 


recorded 






Sena. 


32-8±0-80 


1-60:1=0 -32 


28-0-37-6 


32-0-35*2 


5*0 



hoc. mid Host. The type and 7 paratypes from the axilla of a slrink, 
60 miles south of Darwin, N.T., Australia, 13th- May, 1943 (R. V. Southcott). 

Remarks. Amongst the species of Schongastia this species is very charac- 
teristic in the shape of the scutum, and the dorsal setae. In having only two 
teeth on the chelicerae it is rather a connecting link between the species of 
Schongastia s. str. and Ascoschdngastia. 

Schongastia (Schongastia) vieta Gater 1932. 

Schongastia vieta Gater 1932, Parasitology, 24, 154, fig. 5; Womersley and 
Heaalip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 103. 

Plate 35, fig. A-E. 

This species is closely related to the preceding, in having a more or less 
pentagonal scutum. It differs, however, in having globose sensillae which are 
indistinctly villous, in the scutal and dorsal setae being tapering and normally 
long ciliated, and in AL being the longest of the scutal setae. The chelicerae 
have more than the two inner teeth. The palpal setae on the femur and genu 
are shortly branched, often appearing to be nude. The sensillae are indistinctly 
setulose or villous. The dorsal setae number 32 and are arranged 2.8,6.6.6.4.2, 
sometimes 2.10.8.6.4.2, and measure to 35/x in length. The ventral setae pos- 
terior of coxae III are 6.6.6.6.4.2 approximately. All coxae 1-setose. Tarsi of 
leg III with a long nude seta. 

The Standard Data from the type and 3 paratypes in the British Museum 
(Nat. Ilist.) measured while in London in 1947, and one paratype in the S.A, 
Museum collection are as follows: 



164 Records of the S.A. Museum 





Mean 


Standard 
Deviation, 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


53-8±0-96 


2-15±0-67 


47-4-60-2 


52-8-57-6 


4-0 


PW 


73-0±l-04 


2 -22:1:0 -74 


66-1-79-9 


70-4-76-8 


3-2 


SB 


19-7±0.18 


0-40^:0.13 


18*5-20-9 


19-2-19-8 


2-0 


ASB 


26-2±0-16 


0-36±0-ll 


25-1-27-3 


25-6-26-4 


1-3 


PSB 


26-9±0-48 


1-07+0-34 


23-8-30-1 


26-4-28-8 


4-0 


8D 


53- 1+0 -32 


0-72+0-23 


51-0-55-2 


52-8-54-4 


1-35 


A-P 


30-8±0-25 


0-56+0-18 


29-1-32-5 


30-0-31-6 


1-8 


AM 


28-2±0-84 


1-88+0-60 


22*6-33-8 


25-0-30-0 


6-6 


AL 


60-3±2-05 


4-60±0-45 


46- 5-74-1 


55-0-66-0 


7-6 


PL 


50-7±0-99 


2-21+0-70 


44-1-57-0 


48-4r-52-8 


4-4 


Sens. 


39-3±0-70 


0-99±0-50 


36-3-41-4 


38-4-40-0 


2-4 



Theoretical 


Observed 


Coeff. of 


Range 


Range 


Variation 


49-3-61-7 


51-2-60-8 


3-8 


71-8-84-6 


73-6-83-2 


2-7 


17-2-21-6 


19-2-22-4 


3-8 



Amongst the large collection of Trombiculids collected by T. J. Lawrence 
in Southern Burma in Nov. 1945 are 39 specimens of this species. The speci- 
mens were from Rattus rattus and Nesokia- bengalensis, and all from the neigh- 
bourhood of Toungoo, S.B. 

The Standard Data of these 39 specimens are as follows: 

Standard 
Mean Deviation 

AW 55-5±0-33 2-09+0-24 

PW 78-2+0-34 2-14+0-24 

SB 19-4+0-12 0-74±0-08 

ASB 28-8 No variation recorded 

PSB 25-6 No variation recorded 

SD 54*4 No variation recorded 

A-P 31-2±0-26 1-61±0-18 26-4-36-0 

AM 29-1+0-15 0-97+0-11 26-2-32-0 

AL 70-2zt0-40 2-48+0-28 62-8-77-6 

PL 54-8+0-23 l-41-±0-16 50*6-59-0 

Sena. 32-0 with head 16/19. No variation recorded. 

Schongastia (Schongastia) PHrLiPi Worn, and Kohls 1947. 

Bchonga&tia philipi Wonier&ley and Kohls 1947. Tr. Koy. Soc. S. Aust. 71, 
(1),6. 

Plate 35, fig. F-J. 

This species was described from the type and 9 paratypes from a skink 
Leiolopisma albertmi from Goodenough Island, N.G. (coll. Jan. 17, 1944, G. M. 
Kohls No, 486). 

The scutum is hexagonal as in the species vandersandei, schuffneri, 
blestowei and b. var. megapodiu.% but it only agrees with blestmmi and its 
variety in having a trif urcate palpal claw. 



28-8-32-0 


5-1 


28-8-32-0 


3-3 


64-0-76-4 


3-5 


51-2-60-8 


2-6 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 165 

From blestowei (and megapodius) it differs as in the key, in the smaller 
scutum, the smaller number of dorsal setae, and in the seta on the palpal genu 
being nude. 

The Standard Data for the type and 9 paratypes in the S.A. Museum 
collection as given by Worn, and Kohls 1947 are : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


CoerT. of 
Variation 


AW 


48-6±l-12 


2-51:t0-8O 


41-1-5(>*1 


45- 0-51-0 


5-1 


PW 


69-6±0-40 


-89:1:0 -28 


66-9-72-3 


68-0-70-0 


1-3 


SB 


18-4±0-40 


0-89±0-28 


15-7-21'! 


18-0-20-0 


4-8 


ASB 


24-0 


No variation recorded 






PSB 


21-0 


TNTo variation 


recorded 






SD 


45-0 


No variation 


recorded 






A-P 


23-6±0-40 


0-89 + 0- 28 


20-9-26-3 


22-0-24-0 


3-8 


AM 


22-2±0-73 


l-64±0-52 


17-3-27-1 


21-0-24-0 


7-4 


AL 


59.2±0-49 


l-09:fcO-35 


55-9-62-5 


58 -0-60-0 


1-8 


PL 


42-6±l-13 


2-51±0-80 


35-1-50-1 


39*0-45-0 


5*9 



Sens. 30-0 with head 15/18. Only 1 determination. 

Schongastia (Schongastia) blestowei Gunther 1939. 
Schongastia yeomanM Gunther 1938, Med. J. Aust., 2, (6), 202, (nom. nud,)- 

Schongastia blestowei Gunther 1939, Proe. Linn. Soc, N.S. Wales, 64, (1-2), 
92 ? Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, (1), 103; 
Womersley 1944, idem, 68, (1), 97 (in part). 

Plate 36, fig. A-B. 

Gunther originally described this species from a number of specimens from 
two men near the Suein River, Sepik District of New Guinea. At the same 
time he also recorded as the same some specimens from the ears of a bush fowl 
(Megapodius duperreyi) from Bulolo River Basin, Morobe District,. 

On the difference in size of the scutum and in the Standard Data, 
the specimens from Megapodms were distinguished from those from the Suein 
River, by Womersley and Heaslip (1943), as a variety under the name of 
nuigapflddm. In addition to those differences, the scutal, palpal and dorsal 
setae are much longer branched in megapodius than In the typical form. Also 
in the variety in front of each sensilla base is a distinct semicircle not found in 
the typical specimens. More characteristic even is the much wider distance 
between the sensillae bases in megapodius. In typical blestoivei the dorsal setae 
are rather variable ; Gunther gives the number as up to 64, whereas the figure 
given here shows 54. The latter number is approximately that found in 



166 



Records of the S.A. Museum 



vandersmdei with which Gunther contrasts blest owei (1939). The two species 
however, are very distinct on the character of the palpal claw. 

The following Standard Data for the typical form are from the figures 
given by Womersley and Heaslip 1939 for Gunther 's Suein River specimens 
with the following additional records: ? Solomons 1945 (4 spec. L. A. Pose- 
kany) [ on boots, island north of Bougainville Aug. 1945 (8 spec. G. H. Mc- 
Queen); on man. Treasury Islands, Jan. 1944 (7 spec. L. J- Dumbleton) ; a 
total of 26 specimens. 



AW 


Mean 
65 -7 ±0-51 


Standard 
Deviation 

2*58+0-36 


Theoretical 
Range 

58-0-73-4 


Observed 
Range 

60-8-73-6 


Coeff. of 
Variation 

3-8 


PW 


86*0+0*66 


3-35+0-46 


76-0-96-0 


76-8-91-0 


3-9 


SB 


25-9+0-18 


0*92+0-12 


23-1-28-7 


24-0-28-8 


3-5 


ASB 


32-8+0*32 


1-65+0-23 


27-8-37-8 


28-8-34-0 


5-0 


PSB 


27-0+0-34 


1-73+0-24 


21-8-32-2 


25-6-31-0 


6'3 


BD 


59 8+0*36 


1-85+0*26 


54-2-66-4 


57-6-64-0 


3*1 


A-P 


30-4+0*31 


1-60+0-22 


25-6-35*2 


28-8-35*2 


5-2 


AM 


34*9+0*42 


1-97+0*30 


29 -0-40 '8 


32*0-40*0 


5-6 


AL 


67-1+0-61 


3-01+:0-43 


58-l-76<l 


64-0-73-6 


4-5 


PL 

SfiUH. 


51-35+0*47 
33 -5+0 -34 


2*40+0-33 
1- 53+0-24 


44-15-58-55 
28-9-38-1 


48-0-54-4 
32*0-35-2 


4-7 
4-6 



var. megapodius Worn, and Kohls 1943. 

Tr. Roy. Soc. S. Aust., 67, (1), 104. 

Plate 36, fig. F-J. 

The following Standard Data is derived from the type slide in the S,A. 
Museum containing 3 specimens of Gunther *s 8. blestowei from Megapodms 
duperreyi, together with 4 specimens collected on boots in scrub at Lae, NiG. 
Apr. 1944 (R. N. MicCulloch) and 7 specimens from a Megapodius mound at 
Dobodura, N.G., 13 July 1944 (coll. G. M. Kohls, No. 506) ■ 14 specimens in all. 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


57-7+0-52 


l«94+0-37 


51-9-63-5 


54-4-60-8 


3-3 


PW 


80-3+0-47 


l-75±0-33 


75-1-85-5 


76-8-83-2 


2-2 


SB 


82-2+0-21 


0-80+0-15 


29-8-34-6 


32-0-35-0 


2-4 


ASB 


30-2+0-55 


2-07+0-39 


24-0-36-4 


26-0-32*0 


6>$ 


PSB 


23-0+0-29 


1-09+0-21 


19-7-26-3 


22-4-26-0 


4-7 


SD 


53-2±:0-39 


l-46±:0-27 


48-8-57-6 


51-2-54-4 


2-7 


A-P 


24-4+0-64 


2-38+0-45 


17-3-31-5 


20-0-25-6 


9-8 


AM 


38-55+0-30 


1- 04+0 -21 


35-4-41-7 


37-0-41-6 


2-7 


AL 


67-65+0-90 


3-12+0-64 


5S -3-77*0 


63-0-70-4 


4-6 


PL 


57-2+0-56 


2-01+0-39 


51- 2-63 -2 


54-4-62 >0 


3-5 


Sens. 


33-15 + 0-76 


2-77+0-54 


24-85-41-45 


30-0-38-4 


8-3 



WOMERSLEY— ASIATICPACIFIC SCRUB TVPHUS MlTES 167 

ScHONGASTCA ( SoHONQASTTA ) OCULICOLA 8p. Ii. 

Plate 37, fig. A-E. 

Description of Larva: Shape oval. Colour in life unknown. Length 
(engorged) to 420/i 7 width to 350/** Scutum relatively small, as in figure, 
with the anterior margin sinuous, lateral margins lightly concave, and the 
posterior margin laterally lightly concave, medially convex ; scutal setae slender 
and tapering with only short adpressed ciliations; PI the longest, AM the 
shortest; sensillae glohose, ciliated with their bases fairly near together and 
well in advance of line of PL. Eyes 2 + 2, indistinct, Chelicerae short, curved 
and stout, the apical tricuspid cap prominent, on the dorsal edge a strong sub- 
apical forwardly directed tooth, and more proxiinally some fine indistinct 
rounded teeth. Galea! setae nude. Palpi rather stout; tibial claw large and 
strong with 2 large subeqnal prongs, and 1 smaller shorter prong; seta on palpal 
femur and genu with indistinct short branches; on tibia, dorsal and lateral 
nude, ventral branched; tarsus with a subapical nude seta, a dorsal sub-basal 
sensory rod and 4 or 5 ciliated setae. Dorsal setae to 42^ long, fine, \y\th only 
short adpressed ciliations, 36 in number, arranged 4.6.8.8.6.4, i,e> 2 humeral 
setae on each side. Ventrally : with the usual pair of maxillary setae, one on 
each coxa, a pair between coxa I and between coxae III and thereafter 6,8.8.8.6.2 
somewhat shorter than the dorsal setae. Legs all 7-segmented; I 260/l long, 
II 227/x, III 240ju.; tarsi I and II with the usual dorsal sensory rod; III with- 
out any long nude outstanding seta. 

The Standard Data derived from two larvae and one larval pelt are: 
AW 44-8, PW 71-5 (70-4-73-6); SB 16-0, ASB 22-4, PSB 19-2, SD 41-6, 
A-P 29-9 (28-8-32-0), AM 16-0, AL 36-0, PL 48-0, Sens. 22-0 with head 
16/16. 

Host and Loc. This species is described from the type and paratype (and 
a recovered larval pelt from rearing experiments) from the conjunctival sacs 
of Leggada booduga fulvidimntris (Blyth) from Nalanda, Ceylon, July 1944. 

Mr. S. H. Jayewickreme of the Division of Medical Entomology, Colombo, 
to whom I am indebted for the above material and also the proposed name, 
has successfully reared the larvae through to the nymph as described in another 
part of this paper. 

Schongastia (Schongastia) pseudoschttffneri (Walch 1927). 

TromMcula 'jjsevdoschuffnen Walch 1927, Geneesk. Tijds. v. Ned. Indie, 67, 
(6), 922. 

Neoschongastia pseudoschuffneri, Womersley and Heaslip 1943, Tr* Roy. Soc. 
S. Aust. 67, (1), 117. 



168 Records of the S.A, Museum 

Schdvgasiia n. sp. "X." Lawrence id MS. In Audy 1947. u Scrub Typhus 
Investigations in S.E. Asia, Pt. 111. Appendix 7. M AMD. 7. War Office, 
London. 

Plate 37, fig. F-^T. 

This species does not appear to have been recognized hitherto, since the 
original description from specimens taken on rats from the Lampong District 
of Macassar. It was wrongly placed in Neosehongastia {Ascoschoixgasiia) by' 
Womersley and Heaslip through an oversight of Walch's statement that the 
chelicerae were saw-like. 

However, I have recently been able to examine 5 specimens collected from 
Battus rattus dmrcU from Batavia by Dr, R. Gispin (coll. Nov. 1948) which 
he considered could be referred to Walch's species. After a careful study of 
this material I can confirm Dr. Gispin \s identification. 

The chelicerae are serrate and tbe galeal setae nude. Palpal daw trifur- 
cate and the seta on palpal femur lightly branched, on genu rather less so, on 
tibia nude except the ventral. The scutum is not hexagonal, the posterior mar- 
gin being rounded laterally and rather flattened medially. Sensillae distinctly 
globose or capitate, and ciliated. AL the longest. Eyes 2 + 2, ocular shields 
apparently absent. Dorsal setae 34, arranged 2.8.6.6.6.4.2 or 2.8.8.6,4.4.2. 
Ventral setae posterior of coxae III are 8.4.4.4.4.2 approximately. Legs all 
7-segmented, tarsi III with a long nude seta. 

The Standard Data for these specimens are as follows: 



AW 


Mean 
57-0±:2-12 


Standard 
Deviation 

4-75±l-50 


Theoretical 
Bangc 

42-8-71-2 


Observed 
Range 

51*2-64*0 


Coeff. of 
Variation 

8-3 


PW 


78 ail* 03 


3*65±1-15 


67-2-89*0 


73-6-83-2 


5-0 


SB 


22-4 


No variation 


recorded 






A SB 


25*6 


No variation 


recorded 






PSB 


23^05^:0 -39 


0*88+0*28 


20-4-25-7 


22-4^-24-0 


3-8 


A-P 
AM 


48-65±0-39 

28-3 

26- 2+0 -64 


0-88+0-28 46-0-51-3 
No variation recorded 
l-43H=0..i5 21-9-30-5 


48-0-49*0 
25-6-28-8 


1-8 
5-4 


AL 


66 -1±1- 33 


2-97+0-94 


57-2-75-0 


64-0-70-4 


4-5 


PL 


53- 9+0 -48 


l-07±0-34 


50-7-57-1 


52-0-54-4 


2-0 



Sens, 32*0 with head 13/19. No variation recorded. 

The following specimens from Burma, mainly in material in the British 
Museum and sent to me for study and determination are also to be referred to 
8. pseudoschiiffneri (Walch) : On Battus rattus brunneusculits Hodgson, Iinphal, 
Burma, 7 Nov., 1945 (T. J. Lawrence, 4 specimens) • on Nesokia benyalensis 
24 Sept., 1945 ? 1 mile S. of stream, Paungde, Burma (T.J.L. No. 721, 5 specs., 
No. 722.. 6 specs., No. 723, 3 specs.) j on A', bengalensis, 25 Sept., 1945, 1 mile 



Womersley— Asiatic-Pacific Scrdb Typhus Mites 169 

S. of stream, Paiingde, Burma (T.J.L. No. 728, 14 specs., No. 729, 7 specs.)* 
Total of 49 specimens. 

These 49 specimens have the fallowing Standard Data ; 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coetf . of 
Variation 


AW 


53 -9+0 -24 


l-69±0-17 


4.8- 8-59-0 


51-2-57-6 


3-1 


PW 


78-4±0-35 


2-42±0«25 


71-1-85-7 


73-6-86-4 


3-1 


SB 


20-S±0-21 


1-50 ±.0-16 


16-3-25-3 


19-2-22-4 


7-2 


ASB 


29-0±0-19 


1-31±0-13 


2(5-1-32-9 


25-6-32-0 


4-5 


PSB 


25-3±0-13 


0-88±0-09 


22-65-27-95 


22-4-25-6 


3-5 


BD 


54-3±0-21 


1-47:^0-15 


49-9-58-7 


51-2-57-6 


2-7 


A-P 


31-4±0-20 


1-40+0 -14 


27-2-35-6 


28-8-35*2 


4-4 


AM 


33-0±0-34 


2-24zt0'24 


2(5 -3-39 -7 


28-8-38-4 


6-8 


AL 


74-5±0»65 


4-27±0-46 


61-7-87-2 


70-4-86-4 


5-7 


PL 


59-4±0-44 


2-9S±0-31 


50-5-68-3 


54-4-64-0 


5-0 


Sens. 


35-3 with head 13/19. No variation recorded. 







SOHONGASTIA ( ScHONGASTIA ) MALDIVDENSIS Radford 1946. 

Schongastia nmldiviensis Radford 1946, Parasitol., 37, 48, fig. 9-12. 

Plate 38, fig. A-E. 

This species was somewhat briefly described and figured from specimens 
from a lizard (Caloies versicolor) and occasionally from Battus norvcgicws 
norwgicus from the Maldive Islands. I am indebted to Dr. Radford for two 
specimens, from which I give the following description 

The length is 300/t, width 210^ (engorged or partially so). Scutum as 
figured, anterior margin almost straight, posterior evenly rounded, surface 
punctate. AL the longest, sensillae bases slightly in front of line of PL, and 
with a crescent-shaped line in front and behind; sensillae globose, ciliated. 
Eyes 2 4* 2, ocular shields apparently absent, close to the posterolateral corners 
of scutum. Chelicerae strong, curved, with strong inner serrations. Galeal 
setae nude. Palpi stout, claw trifurcate with median prong the longest; setae 
on femur and genu strongly branched ; on tibia dorsal and lateral nude, ventral 
branched. Dorsal setae ea. 34 in number and arranged ca. 2.8.6.6.6.4.2, to 32/x 
long. Ventral setae, the usual pair of maxillary, one on each coxa, a pair 
between coxae I and between coxae III and thereafter 4.6.6.6.4.4.2. Legs I 
330/x, long, II 300/t, III 330/* ; tarsi I and II with the usual dorsal sensory rod, 
III with a long outstanding nude seta* 

The Standard Data for these two specimens are; AW 51-2, 51-2, PW 67*2, 
€4-0, SB 16-0, 16-0, ASB 25-6, 25-6 PSB 25-6, 25-6, SD 51-2, 51-2, A-P 
28-8, 28-8, AM 25-6, 28-8, AL 54-4, 54-4, PL 48-0, 48-0, Sens. — , 32-0 with 
liead 15/20. 



170 Records of the S.A. Museum 

CHELICERAE NOT SERRATED. 

Subgenus AS-COSCHOENGASTIA Ewins 1946, 

Neoschbngasiia Ewing 1929, Manual of External Parasites, p. 187 (in part) ; 
Womersley and Heaslip 1943, Tr. Roy. Soe. S. Aust., 67, (1), 106; Sig 
Thor and Willmann 1947, Das Tierreich, 3, 71b, 310. 

Ascosclibngaviia Ewing 1946, Proc. Biol. Soc. Washington, 59, 69-72. 

Dr. H. E. Ewing 1946 showed that the genus Neoschbngastia with ameri- 
cana Hirst as type was synonymous with Paraschbngastia Womersley 1939, 
and that the non-serrate chelicerae, the sensillae bases being inserted in a trans- 
verse wall or erest of the scutum, and the posterior part of the scutum being 
overlapped by the dorsal cuticle with semicircular striatums, characteristic of 
Pavasclibngastia were also features of Neoschbngasiia americana. 

For the other species with non-serrate chelicerae, and a simple type of 
scutum Ewing proposed (1946) the generic name of Ascoschbngasiia, designat- 
ing a somewhat aberrant species, Neoschongastia malayensis, as type. 

As shown in the adult and nymphal section of the present work, the genera 
cannot be separated in these stages, and it seems doubtful whether the serration 
of the chelicerae in the larvae can be regarded as of more than of subgeneric 
value. Consequently here Asc-osckbngastia is given only as a subgenus of the 
older genus. 

This is supported by the fact that the number of cheliceral teeth varies 
in the different species and in some Schongastia species (e.g. bklentaia sp. n.) 
approaches the non-serrate type. 

The genus Ewchongastia was erected by Ewing, 193S (Wash. Acad. Sci., 28, (6), 293) 
with Euschoiigaxtia americoma ap. m, as type, em the basis of the palpal claw having five or 
seven prongs. In 1946 Ewing (Proe, Biol. Soc. Wash., 59, 69t-72) also erected the genus 
Axcoschongastia.) for those species with n on -per rate chelicerae and with bi- ot trifurcate palpal 
claw, but without the characters of the scutum of Neosclumgaatia 1929 (Manual External Para- 
sites, p. 137). For Asoo8ehSnga»tm he designated as type, Ncofschonga.rtm m*ila>yeiisi>s Gater 
1932, a speeies characterized by having only 3 ecutal setae, i.e. AM and AL, both PL being 
situated off the scutum. 

Wharton 1948 (Psyche, 55, (2), 90) in describing phyUoii from Peru, with 3 prongs to the 
palpal claw placed it in Euschongastia and proposed to expand Ewing 3 s usage of this- generic 
name to embrace such species, and to restrict Ascoachvnga&tia to species with PL off the 
scutum. 

As there are no generic differences to be found in the nymphs or adults which will separate 
malayenxis from the other species of Ascoselwngastiaj it seems that such a character aB 
exhibited by malayev&is larvae can hardly be valid generically. A similar shortening of the 
scutum so a 8 to exclude the PL setae, as well as a multipronged palpal claw, also occurs in the 
genus Teeomatla'na Hoffman 1947 (=- SavHsci^ Lawrence 1949). 

Further, the variation in the number of prongs in the palpal claw is also of little, if any, 
generic value and, ot most, should only "be used subgenerieaUy. 

These considerations then, show that Euschdngostm Ewing is identical with Aseochoti- 
gastia Ewing. over which it has many years' priority, and should in all probability bo used in 
place of A neoschongastia in this paper. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 171 

The subgenus Ascoschongastta Ewing may be diagnosed as follows : 
Larval Trombiculid mites in which the dorsal scutum carries 5 normal ciliated 
setae, and a pair of clavate or globose sensillae whose bases are not set in a 
transverse wall or crest. The posterior part of the scutum not overlapped by 
the dorsal cuticle and without striations. Chelicerae non-serrate on inner 
(dox*sal) margin; with only the usual apical tricuspid cap. 

1. Some or all doksal, as well as PL and sometlmes AL or AM foliate, 

SCHONGASTIA (AscOSCHONGASTU) PSEUDOMYS Sp. n. 

Plate 39, ftg. A-E, 

Description of Larvae: Shape oval. Length (partially engorged) 450/x, 
width 300/i. Dorsal scutum as figured, with PL setae long, foliate and placed 
slightly nearer to AL, than to the posterior margin; anterior seutal margin 
slightly sinuous, posterior with widely rounded corners; AL very short and 
shorter than AM, both AM and AL not foliate,* sensillae missing but bases 
behind line of PL. Eyes 2 -\- 2, on ocular shields, closely adjacent to scutum 
on a level with PL ; posterior eyes the smaller. Chelicerae with only the apical 
tricuspid cap. Galeal seta strongly branched. Palpi as figured, with trifurcate 
tibial claw, and all setae on femur, genu and tibia strongly ciliated, tarsi with 
sub-basal and subapical nude sensory setae and 4 or 5 ciliated setae. Dorsal 
setae 38 in number to 85/* long, all except the posterior 4 long and foliate, with 
the edges with short ciliations, arranged 2.8.6.6.6.6.2.2. Ventrally with all setae 
tine and ciliate, a pair between coxae I and between coxae III, and thereafter 
8.6.6,4.4.2, to 40/i long. Legs all 7-segmented, T 260/x long, II 240/*, III 270/t 
tarsi I and IT with the usual dorsal sensory rod, III without any long nude seta. 

The Standard Data for the type and two paratypes are : 





Mean 


Standard Theoretical 
Deviation Kange 


Observed 
Range 


Ooeff. of 
Variation 


AW 


70-4 


No variation recorded 






rw 


84«3±:l-07 


l-S5±0-75 78-7-89-0 


83-2-86-4 


2-2 


SB 


32'0:±1*87 


3-20±:l-31 22-4-41-6 


28-8-35-2 


10-0 


ABB 


28-8 
24-5±0-54 


No variation recorded 
0-93:hO-38 21-7-27 3 


24-0-25-6 


3-8 


SU 


52*3±1*40 


2-43±l-00 45 • 0-59 • 6 


49* £-64* 4 


4-6 


A-P 


16-0 


No variation recorded 






AM 
AL 


54-4 
25-6 


No variation recorded 
No variation recorded 






PL 


81-litl-07 


l-85-±0-75 75-5-86-7 


80-0-83-2 


2-3 


Sena. 




Missing. 







172 Records of the S.A. Museum 

hoc. and Host. The type and two paratypes in the S.A. Mus. coll. from a 
Pseudomys sp. from Johnston's Pass, Wongabel, Queensland, 13 Feb., 1945 
(R. N. McCulloch). 

Remarks. Although the sensiilae are missing from all three specimens, this 
species is probably correctly placed in the subgenus Ascoschongasim. It can 
be separated as in the key, on the palpal setae and on the foliate scutal and 
dorsal setae. 

Sohongastia ( Ascoschongastia) mccullochi (Worn. 1944). 

Neosch&ngastm mccvllochi "Womersley 1944, Tr. Roy. Soc. S. Aust., 68, (1), 
100. 

Plate 39, fig. F-K. 

This species has not been seen since the unique specimen was described. 

In addition to the details given in the original description the following 
are to be noted. 

The galeal setae are nude. The palpal claw is trifurcate, and all seta on 
the palpal femur, genu and tibia are nude. There is no long nude seta on 
tarsi III. 

The Standard Data as given in 1944 are: AW 48-0, PW 67-0, SB 19*0, 
ASB 19-0, PSB 16-0, SD 35-0, A-P 20-0, AM 16-0, AL 42*0, PL 64-0, Sens. 
32-0, with head 17 X 17 and nude. The dorsal setae are 2.6.6.6.4.2, and as 
figured. 

The type and unique specimen in the S.A. Museum collection -was collected 
on boots at Abidari, New Guinea, 28th July, 1943 (B. N. McCulloch). 

Sohongastia (Ascoschongastia) ubomys (Worn, and Kohls 1947). 

AscoscJwngastia uromys Worn, and Kohls 1947, Tr. Koy. Soc. S. Aust., 71, (1), 
10. 

Plate 40, fig. A-F. 

This species was described from the type and 7 paratypes from TJromys 
lamington from Dobodura, New Guinea. 

It belongs to a small group of species, including foliata Gunther, 
mccidlochi Worn, and pseudomys sp. n., in having some of the scutal setae 
and most of or all the dorsal setae lanceolate or narrowly foliate. The 
original description of the larvae was as follows: " Shape oval. Length (un- 
engorged) 300/x, width 200/*. Scutum roughly rectangular with fairly well 



Womersley— Asiatic-Pacific Scrub Typhus Mites 173 

produced posterior margin. AM short and normally ciliated, AL and PL 
lanceolate or foliate, to 16ju. wide with longitudinal rows of strong dentures. 
Sensillae globose, apparently nude, with the bases only slightly in advance of 
line of PL. Eyes 2 + 2, on distinct ocular shields; posterior the smaller. 
Chelicerae with only the usual apical tricuspid cap. Galeal setae nude. Palpi 
fairly stout, with bifurcate tibial claw. Setae on palpal femur, genu and 
tarsi apparently all nude. Dorsal setae all foliate or lanceolate with strong 
dentures, to 64u, long and 16/* wide, 28 in number and arranged 2.6.6.6.6.2. 
Ventrally with usual pair of branched setae on gnathosoma, a single ciliated 
seta on each coxa, a pair between coxae I and between coxae III and thereafter 
5.6.6 anterior of anus, and of which the outer two of the second six approach 
the dorsal setae in form; then posterior of anus 4.2, foliate and dentate as on 
dorsum; the anterior non-foliate ventral setae to 20/* long. Legs: I 270/* long, 
IT 220/*, III 270it; tarsi I and II with the usual sensory rod, III without any 
long nude seta. 

The Standard Data derived from the type and seven paratypes are as 
follows : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 


50-6±l-74 


4-92±l-23 


35-8-65-4 


45-0-60*4 


9*7 


PW 


69*5±0-71 


2-00±0-50 


63 -5-75 -5 


66*0-75*0 


2-9 


SB 
ASB 


19-0±0*50 
21-0 


1'41±0«35 14 -8-23 -2 
No variation recorded 


18*0-21-0 


7*4 


PSB 


1S-0 


No variation recorded 






8D 


39-0 


No variation recorded 






A-P 


21 -6^:0 -42 


l-19±0-30 


18-0-25-2 


21-0-24-0 


5*5 


AM 


16-5±0-67 


l*G4±0-47 


11 -6-21 -4 


15-0-18'0 


9-9 


AL 


49*5±0-67 


3 -<>4:t0-47 


44-0-54-4 


48-0-51-0 


3-3 


PL 
Sens. 


60-0+0-57 
28-0 with heac 


l-60±0-40 
i 17/17. Only 1 d< 


55-2-64-8 
^termination. 


57-0-63*0 


2-7 



Remarks-. No fresh material of this species has been collected. 

Schongastia (Ascoschongastta) foliata (Gunther 1940). 

Neoschdnga<stia foliata Gunther 1940, Proc. Linn. Soc, New South "Wales, 65, 
(3-4), 255; Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, (1), 
112. 

Plate 40, fig. G-L 

In addition to the details given by Womersley and Heaslip 1943, it must 
be stated that according to Gunther (1940) the palpal claw is trifurcate, the 
setae on the palpal femur and genu are branched or ciliated, and on the 



174 Records of the S.A. Museum 

tibia the dorsal and lateral setae are nude and the ventral branched. The 
ehelieerae are simple, with only the apical tricuspid cap, and the galeal setae 
are nude. The dorsal setae are 2.6,6.6.6,4,2, to 62ja long. 

The Standard Data for 5 paratypes as given by Womersley and Heaslip 
and statistically treated are I 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


"Variation 


AW 


53-2±l-52 


S-42±l-08 


43-0-63-5 


50-0-58-0 


6-4 


PW 


69'6±0-68 


l-52±0-48 


65- 1-74-1 


68-0-71-0 


2-1 


vSE 


17-l±0-37 


0-82±0-26 


14 -7-19 -5 


16-5-18-0 


4-8 


A SB 


19-3±0-44 


0-97+0-38 


16-4-22-2 


1S-O-20-0 


5-0 


PSB 


17-2±0-34 


0-74 + 0-24 


14-9-19-5 


16-5-18-0 


4-4 


SD 


36-5±0*67 


1-50+0-47 


32-0-41-0 


35 '0-38-0 


4-1 


A-P 


25-0 


No variation recorded 






AM 


20-6±0-98 


2-19+0-69 


14-0-27-2 


18-0-24-0 


10-6 


AL 


58-7±2-33 


4-04±l-f>5 


46-6-70-8 


55-0-63-0 


6-9 


PL 


86-6±l-69 


3-78+1-19 


75-2-98-0 


83-0-91-0 


4-4 


Sens. 


28-0 with head 3 


[7/17. No variation recorded. 







This species is as yet only known from the original material from, the 
ears of Macropus (Thylogale) coxeni Gray, from Bulolo, New Guinea. 

II. No Dorsal or Scutal Setae Foliate. 

(A) SENSILLAE OLOBQSE. 

Schongastia (Ascoschongastia) signata sp. n. 

Plate 41, fig. A-D. 

Description of Larvae. Shape elongate oval. Length (unfed) 220ju, width 
140/t; (engorged) to 480/* and 300/*.. Scutum as figured, rectangular, with 
anterior margin shallow behind line of PL, almost straight; sensillae globular, 
apparently nude, and with their sensillae bases placed less than their own 
diameter apart, and well in front of line of PL; A-P greater than 3/4 SD; 
AM setae the shortest, PL the longest. Eyes 2 + 2, the posterior the shorter. 
Chelicerae with only the apical tricuspid cap. Galea! setae nude. Palpal setae 
all ciliated or branched* Dorsal setae fairly long and slender, 36 in number 
arranged ca, 2.10.8.6.6.4 to 54^, long. Ventrally with the usual pair of branched 
setae on maxillae, a long ciliated seta on each coxa, a pair between 
coxae I and between coxae III, thereafter about 26 setae. Legs T 288/x, IT 224/*, 
III 27.2ft, tarsi I and II with the usual dorsal sensory rod, tarsi III without any 
long nude setae. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 175 

The Standard Data for the type and 11 paratypes are : 





Mean 


Standard- 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


41-9±0-27 


0-92±:0-19 


39-1-44-7 


41-6-44-8 


2-0 


PW 


64-0 


No variation 


recorded 






SB 


12-5±0*18 


0<62±0-13 


10-6-14-4 


11-2-12-8 


5<0 


ASB 


22-4 


No variation recorded 






PSB 


ll-6±0-29 


0* 99+0 -20 


8-6-14-6 


9-6-12-8 


8-6 


SD 


34«0±0-29 


0-99±0-20 


31-0-37-0 


32-0-35-2 


2-9 


A-P 


3l-5±<)-22 


0-79±0*16 


29-2-33-8 


30-4-32-0 


2-5 


AM 
AL 


29-4±0-58 
63-4±0*84 


l-93±0-41 
2-80;£0-59 


23-6-35-2 

55-0-71-8 


25-6-32-0 
57-G-67-2 


6-5 
2'6 


PL 


70-9±0-88 


2-77+0-62 


62-6-79-2 


67-2-73-6 


3-9 



Sens. 35 • 2 with head 19 • 6/22 • 4. No variation recorded, 

Loc, and' Hosts. The type and eleven paratypes collected on boots by Maj. 
R. N. McCulloch, June, July and October, 1945, at Brunei and Labuan, North 
Borneo. 

Remarks, In the small group of species with closely approximate sensiUae 
bases this species is, in the shape of the scutum, most closely allied to nwtaibilis 
Gater. From this it differs in having PL setae the longest, in the number of 
DS and in having all coxae unisetose. It also has the sensiUae nude, and all 
palpal setae ciliated. 

SCHONGA&TIA (AsCOSOHONGASTIA) ? GLOBULARE (Walch, 1927). 

Trombidium (Trombiculaf) globulare Walch, 1927. Geneesk. Tijdschr. v. Ned. 
Indie. 67, (6), 929, fig. 7.8.9; 

Neoschongnstia globulare, Womersley and Heaslip, 1943. Trans. Roy. Soc, 
S. Aust., 67, (1), 114. 

Plates 41, fig. E-F; 42, fig. A-B. 

This species does not appear to have been met with since Walch first 
recorded it from rats from the Macassars. According to Walch, the scutum is 
somewhat rectangular, with the anterior margin sinuous, and the posterior 
margin fairly deep and rounded behind line of PL. The sensillae bases are set 
close together, behind line of the PL, and the globular sensiUae are finely 
ciliated. PL setae are the longest. The eyes are 2 + 2, with the posterior the 
smaller. The chelicerae are not serrate. The galeal setae and all the palpal 
setae arc ciliated. Coxae I and II have one seta, III is stated to have 2 setae, 
but in his figure 8 are shown as having 3 setae. The dorsal setae are 2(0). 
8(10). 6,6.6.4.2, to 33/i and the ventral setae posterior of coxae III ca. 32 in 
number. There is no long nude seta on tarsi in. 



176 Records of the S.A. Museum 

Amongst specimens of kohlsi and lanius from B. rattus brunwu$culU9 
17 Dec. 1945 from Imphal was one specimen which on the dimensions and 
shape of the scutum, the position of the sensillae bases, and the dorsal setae 
as well as the ciliated galeal setae, bifurcate palpal claw, and all palpal setae 
being ciliated, is doubtfully referred to Walch 's species. Unfortunately the 
sensillae and scutal setae are missing, and it has 4 setae on each coxa III. This 
latter feature in other species is known to occasionally vary, but the setae not 
shown in Walch J s figure is that situated at the anterior outer corner of the 
coxa and generally is somewhat difficult to make out, being confused with the 
edges of the coxa. It may be that Walch did not see it. 

This specimen is the one figured herein, except that in fig. C the sensillae 
and scutal setae arc inserted from Walch 's data. 

Walch gives the scutum as 43/* long and 64/x wide, and the sensillae as 25/* 
long with the head 19 by 15ft. The specimen described here has the following 
Standard Data AW 57-6, PW 73-6, SB 11-2, ASB 22-4, PSB 12-8, SD 35-2, 
A-P22-4. 

8. (A.) globular e in the shape of the scutum is closely related to both 
lanius Radford and kohlsi Philip and Woodward, bat differs from both in 
having PL the longest of the scutal setae. It differs from lanius in the palpal 
claw being bifid and in the ciliated galeal setae. From kohlsi it differs in the 
multisetose coxae III and the number of dorsal setae. 

Schongastu ( Ascoschongastia) kohlsi (Philip and Woodward 1946). 

Neoschongastia JcohUi Philip and Woodward 1946, Amer. J. Tropical Medicine, 
16, (2), 159-161, pi. I, fig. B, D. 

Plate. 42, fig. C-F. 

This species was described from Rattus mindanemis mmdanensis and 
R. vigoratiiSj San Jose, Mindoro, Philippines, 1945. It also occurs commonly 
in Burma, and T have been able to study, through the kindness of Lt.-Col. Audy, 
a number of specimens from Rattus rattus brunnenseulm from the neighbour- 
hood of Imphal and collected by Sgt. T. J. Lawrence and other members of the 
British Scrub-Typhus Research Unit, 1945. 

Redescrvption of Larvae. Shape an elongate oval. Length (engorged) to 
510/t, width 405ft. Dorsal scutum as figured, anterior margin sinuous, posterior 
rather flattened medially, curved laterally and fairly deep behind line of PL ; 
sensillae bases less than their own diameter apart and placed distinctly behind 
line of PL, A-P ca. 2/3 scutal depth, AM seta the shortest, AL the longest. 



Wo\fERSLEY— Asiatic-Pacific Scrub Typhus Mites 177 

Sensillae globose with fine indistinct ciliations. Eyes 2 -f- 2, the posterior 
the smaller. Chelieerae not serrate, only furnished with the usual apical tri- 
cuspid cap. Galea! setae ciliated. Palpi stout, with bifurcate tibial claw. All 
setae on palpi ciliated, although sometimes the dorsal and lateral setae of 
tibia appear nude, palpal tarsi with the usual basal and subapical sensory 
rods, and 4 or 5 ciliated setae. Dorsal setae very numerous, ca. 86 in number 
(P. and W. give 52 to 68), and arranged ca. 2,12.14.12.14.10,10.6.4.2, to 40- 
45/x long, thick with prominent setules. Ventrally, all coxae 1 -setose, the 
usual pair of branched maxillary palpal setae, a pair of ciliated setae between 
coxae I and between coxae III, and thereafter ca. 60 setae to 25m long. Legs ; 
I 23%4 long, II 210/*, III 230/x ; tarsi I and II with the usual dorsal sensory rod, 
III without any long outstanding nude seta. 

Thq Standard Data, for eleven paratypes statistically treated from the data 
given by Philip and Woodward are : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 


52-3+0-65 


2-15+0-46 


45-9-58-7 


48-0-56-0 


4-1 


PW 


63 • 6+0-58 


1-91+0-41 


57-9-69-3 


60-0-66-0 


3-0 


SB 


9-6+0-28 


0-92+0-20 


6-8-12*4 


8-0-11-0 


9-6 


ASB 


24-3+0-38 


1-27+0-27 


20 '£-28*1 


22*0-27-0 


5-2 


PSB 


8-55+0-28 


0-93+0-20 


5-75-11-35 


7-0-10-0 


10-9 


8D 


32-8+0-38 


1-25+0-27 


29-05-36-55 


32-0-35-0 


3-8 


A-P 


19-7+0-33 


1-11+0-24 


16-4-23-0 


18-0-21-0 


5-6 


AM 


22-55+0-92 


3-04±0-65 


13-4-31-7 


20-0-28-0 


13-5 


AL 


45-55+0-46 


1-54+0-33 


40-9-50-2 


42-0-48-0 


3*4 


PL 


34-3d:0-75 


2-49±0*53 


26*8-41-8 


32'0-40-0 


7-3 


Sens. 


24-6+1-17 


3-69+0-82 


13-5-35'7 


20*0-30*0 


15-0 



That the Burma material does not differ significantly in the Standard Data 
is seen from the following data derived from 9 specimens measured : 





Mean 


Standard Theoretical 
Deviation Bange 


Observed 
Range 


Cooff . of 
Variation 


AW 


49-95+0-74 


2-23+0-53 43-25-56-65 


48*0-54 '4 


4-5 


PW 


66-65+0-38 


1-13+0- 27 63-25-70-05 


64-0-67-2 


1-7 


SB 
ASB 


9-6 
25-6 


No variation recorded 
No variation recorded 






PSB 


12-8 


No variation recorded 






8D 


38-4 


No variation recorded 






A-P 


22-4 


No variation recorded 






AM 


24-5+0-66 


1-65+0-48 19'5-29'5 


22 -4-25 '6 


6-7 


AL 


59-7+1-55 


3-88+1-12 48-1-71-5 


54-4-64*0 


6-5 


PL 


38-3+0-89 


2-35+0-63 29-3-43-3 


32-0-38-4 


6-5 



Sens. 28-8 with head 19/19. Only 1 determination. 



178 Records of the S.A, Museum 

Remarks. This species is one of a small group distinguished by the sen- 
sillae bases being less than their own diameter apart. It can be separated from 
the others of the group as in the key. 

In the general form of the scutum, it has the sensillae bases behind line of 
PL; and in having AL the longest of the scutal setae this species is closely 
related to lamus Radford 1946. From this species, however, it is distinct in 
the much more numerous dorsal setae, in the palpal claw being only bifid, and 
in having all palpal setae as well as the galeal setae ciliated or branched. 

ScnoNGA&TiA (Ascoschongastia) oomata sp. n. 

Plate 43, fig. A-E. 

Description of Larvae: Shape oval. Length (un-engorged) 364^, width 
260/a. Scutum as figured with anterior margin lightly sinuous, posterior mar- 
gin deeply rounded but rather flattened medially, A-P rather short with PL 
nearer to AL than to posterior margin, SB very close together and posterior of 
line of PL, sensillae globose and finely setulose. Eyes 2 -{- 2, on ocular shields, 
posterior the smaller and indistinct. Chelicerae simple with only the apical 
tricuspid cap. Galeal setae strongly branched. Palpi stout, tibial claw bifur- 
cate, setae on palpal femur and genu shortly and strongly ciliated; setae on 
tibia nude except ventral which has 2 or 3 branches, tarsi with subapical and 
sub-basal rods and 4 or 5 ciliated setae. Dorsal setae numerous, more than 
110, and arranged ca. 2.16.10.12.14.12.14.14.8.6.4, thick and with strong setules, 
to 2&jjL long. Ventrally with a pair of ciliated setae on maxillae, a pair between 
coxae I and between coxae III and thereafter ea. 60 setae finer but almost as 
long as the dorsal. Legs 7-segmented, I 220/* long, II 182/* long. III 220/l; 
tarsi I and II with usual dorsal sensory rods, tarsi III without any long nude 
seta. 

The Standard Data for the type and 1 paratype are; AW 59*0, PW 70-0, 
SB 10-0, ASB 28'0 3 PSB 12*0, SD 40-0, A-P 20*0 ? AM 24-0, AL 42-0, 
PL 31 '0, Sens. 30-0 with head 18/18. 

Loc. amd Host. The type and two para types (damaged) from Clarke Field, 
Philippine Islands. 1945. (G. M. Kohls). 

Remarks. In the shape of the scutum with AL setae the longest, and all 
coxae unisetose, this species is closely related to 8. (A.) kokhi Philip and Wood- 
ward. It differs from that species, however, in having much more numerous and 
shorter dorsal setae. This species also occurs in Kashmir, India, and I have 
recently been able to study four specimens in all, collected from rats from 
Kanzalwan, Kashmir, October, 1946, by Major S. L. Kalra. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 179 

Schongastia (Ascoschongastia) lanius (Radford 1946). 

Neoschongmtia. lanius Radford 1946. Proc. Zool. Soc, London, 116, (2), 262, 
fig. 23-24, 

Plates 43, fig. F-H; 44, fig. A. 

This species was originally described, but very briefly, from specimens, from 
the Black-headed shrike (Lanius naswtus Scopoli) from Imphal, Burma, Feb, 
11th, 1945. It has also been taken in fairly large numbers by Sgt. T. J. Law- 
rence and other members of the British Scrub-typhus Research Unit in Burma, 
1945 and onwards. To Lt.-Col J. A. Audy, 1 am indebted for the opportunity 
of studying this material, which -was mainly from Bathos ratUis brmMeuscwhis 
from an area about 10 to 20 miles north of Imphal. The following redescrip- 
tion and figures are from this material, 

Ee-description of Larvae* Shape elongate oval. Length (engorged) to 
510/*, width to 320/a. Scutum as figured with PL placed about midway of 
scutal depth, and sensillae well behind line of PL. AM the shortest, AL the 
longest. Sensillae globose and finely ciliated, bases less than their diameters 
apart, with strong crescents anteriorly. Eyes 2 + 2, the posterior the smaller, 
anterior with strongly reflecting cornea. Chelicerae non-serrate, with only the 
apical tricuspid cap. Galeal setae nude. All palpal setae ciliated. Palpal claw 
trifid. Dorsal setae 38 in number, arranged 10.6-6.8.4.2.2, to 38ft long. Vent- 
rally, with the usual pair of branched setae on maxillae, one seta on 
each of coxae I and II, two setae on coxae III (but variable from 2-4), a pah 
of setae between coxae I and between coxae III, thereafter ca. 28, arranged 
6.6.6.2.4.2.2. Legs I 292^ long, II 258/x, III 278/*; tarsi I and II with the 
usual dorsal rod, III without any long nude seta. The Standard Data from 
eleven specimens from Imphal are as follows: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Gooff, of 
Variation 


AW 


57-l:±:0-77 


2-66±0-54 


49-1-65-1 


51*2-60-8 


4-7 


PW 


70-9=h0-77 


2-67±0-54 


62-9-78-9 


67-2-73-6 


3-7 


SB 


9-2:t0-21 


0-72±(M5 


7 -0-1 1-4 


8-0-9*6 


7-0 


ASB 


25-6 


No variation recorded 






P8B 


12-8 


No variation recc 


►rded 






ST) 


38-4 


No variation recorded 






a-p 


17-8+0-42 


1- 44:1:0 -29 


13-5-22-1 


16-0-19'2 


8-0 


AM 


32-3±0-52 


l-73:±0-37 


27-1-37-5 


28-8-35-2 


5-3 


Ah 


56-15^0-66 


2*20^:0-47 


49-55-62-75 


54-4-60-8 


3-9 


PL 


45-6±0-42 


1-45^-0-20 


43-3-49-9 


44-8-48-0 


3-2 



Sens. 28 '8 with head 16/19. No variation recorded. 



180 Records of the S.A. Museum 

Remarks. This member of the small group of species with closely approxi- 
mated sensillae bases is closely related to koklsi Philip and Woodward, but 
differs in the trifureate palpal -claw, the nude galea! setae, the nude lateral 
.and dorsal setae on the palpal tibia, and in the smaller number of dorsal 
setae. Also A-P is not greater than 1/2 SD. In this it approaches globular e 
Walch, but differs in the trifureate palpal claw, in coxae 111 being: bisetose, not 
trisetose, and in AL not PL being the longest. It can be distinguished as in 
the key. 

Schongastia (Ascoschongastia) mutabilis (Gater 1932), 

Neoschdngastia mutabilis Gater 1932 Parasitology, 24 ; "Womersley and Ileaslip 
1943. Tr. Roy. Soc. S. Aust 67, (1), 111, pi. VII, Pig. 5. 

Plate 44, fig. B-E. 

This species was described by Gater, but without any figure, from a cluster 
of mites in the ears of Rattus sabanus vociferam (Miller) from Sungei Buloh, 
Selangor, F.M.8. It does not appear to have been met with since, and the 
following redescription and figures are from a paratype specimen from Gater 
from Tiipaia glis ferruginw in the collections of the South Australian Museum. 

Re-description of Larva. Shape an elongate oval. Length (engorged) 
430/i, width 285/*. Scutum trapezoidal, with sensillae bases (sensillae missing) 
in front of line of PL and A-P greater than 3/4 scutal depth ; AL setae the 
longest. Eyes 2 — (— 2, posterior the smaller. Chelicerae non-serrate, with only 
the small apical tricuspid cap. Galeal setae nude. Palpal claw trifureate. 
Setae on palpal femur and genu with a few short ciliations ; on tibia, dorsal and 
lateral nude or with indistinct ciliations, ventral ciliated. Dorsal setae 40, 
arranged ca. 2.8.6.6.8.2.4.2.2, to 38/* long, shortly ciliated. Ventrally with the 
usual pair of branched setae on maxillae, a pair of ciliated setae between 
coxae I and between coxae III, one such on coxae I and II, and three (vari- 
able) on coxae III, behind coxae III with ca. 32 setae. Legs I 240/a, II 185/i, 
III 220/i; tarsi I and II with the usual dorsal sensory rod, 111 without any 
long nude seta. 

Standard Data: AW 48 «0, PW 62-0, SB 9-0, ASB 26-0, PSB 13-0, 
SD 39-0, A-P 26-0, AM 29-0, AL 53-0, PL 40-0, Sens, t 

Remarks. In the shape of the scutum, with A-P greater than 1/2 scutal 
depth, and sensillae bases in front of line of PL, mutabilis is closely related to 
Mgnata sp. n„ It differs from the latter species, however, in having AL and 
not PL the longest, the sensillae being distinctly ciliated, in the number of 
dorsal setae, and in having coxae III trisetose. It is readily separated as in 
the key. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 181 

The record of Lawrence (in Audy 1947) for this species is incorrect. His 
material is a mixture of S. (A.) lanms (Radford 1946) and S. (A.) kohlsi 
(Philip and Woodward 1946). 

SCH0NGASTIA( AsCOSCHONGASTIA ) &DWARDSI (Gunther 1939), 

Neoschongastia rioi Gunthcr 1938 (nam. n%d.) Med. J. Aust., 2, (6), 202. 

Neoschdngattia edwardsi Gunther 1939, Proc. Linn. Soc, New South Wales, 64, 
(1-2), 86; Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, (1), 
112. 

Plate 44, fig. F-H. 

According to Gunther 's original description the palpal claw is bifurcate, 
the setae on the palpal femur and genu ciliated and on the tibia dorsally and 
laterally nude, ventrally branched; the galeal setae are nude; the chelieerae 
simple with only the apical tricuspid cap; and tarsi III is furnished with a 
long nude seta. 

I have now been able to examine the paratype specimen of this species in 
the Australian Museum, Sydney; it is unfortunately poorly mounted and the 
details of the palpi and mouthparts cannot be clearly seen. The Standard Data, 
however, are measurable, and are as follows : AW 76-0, PW 96-0, SB 38-0, 
ASB 32-0, PSB 29-0, SD 61-0, A-P 29-0, AM 48-0, AL 80-0, PL 72-0, Sens. 
28 «0 with head 18/29 and finely setulose. These values are somewhat higher 
than those reported by Womersley and Heaslip (1943) for Gunther 's type. 
The dorsal setae are 2.14.14.10.12.8.4 approx. 

In the bifurcate palpal claw this species is close to philippensis Philip and 
Woodward 1946 but differs from it in the presence of a long nude seta, on 
tarsi III, in the shape and dimensions of the dorsal scutum, in AL being longer 
than PL, and in having many more dorsal setae. 

It is still only known from the original material, from the bush fowl, 
Megapodhts dwperreyi from New Guinea. 

Schongastia ( Ascoschongastia ) PHiUPPENsis (Philip and Woodward 1946). 

Neoschongastia philippensis Philip and Woodward 1946, Amev. J. Tropical 
Medicine, 26, (2), 158. 

Plate 45, fig. A-C. 

This rather characteristic species was described by Philip and Woodward 
from specimens from Rattus mindmienm mindanensis and R. vigoratus from 
Mindoro, Philippines. 



182 Records of the S.A. Museum 

I am much indebted to Lt.-Col. Philip for a number of paratypes (19) and 
I have also 3 other specimens from the same locality, Feb., 1945 (C. Mohr.). 
In addition, in the South Aust. Museum collection, is a slide of 4 specimens 
from New Castle Waters, Queensland, dated 17 May, 1942, from the ears of a 
wallaby (R. V. Southcott) which can be referred to this species. In addition 
to the details given in the original description the following may be noted : the 
palpal claw is bifurcate; the setae on the palpal femur and genu branched, 
on the tibia dorsally and laterally nude and ventrally branched ; ehelicerae with 
only the apical tricuspid cap and galeal setae nude; there is no long nude 
seta on tarsi III. 

The shape of the scutum is rather characteristic, and the Standard Data 
for 22 specimens from the original material are as follows : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Bongo 


Range 


Variation 


AW 


54 -G:£0 -39 


l-82±0-27 


49-2-60-0 


50-0^57-0 


3-3 


PW 


79-2±0-46 


2-19±0-33 


72-7-S5-7 


75-0-82-0 


2-7 


SB 


23 0±0-38 


l-Sl±0-27 


17-6-28*4 


21-0-25-0 


7-9 


ASB 


21-0 


No variation 


recorded 






PSB 


12-3±0*26 


1-22;±:(M8 


8-7-15.9 


11-0-14-0 


9-9 


ST) 


33 -%±0 -26 


1-22+0-18 


29 -7-36 -9 


32-0-35-0 


3-7 


A-P 


25-0 


No variation recorded 






AM 


24-5±0-25 


1- 19+0-18 


20- 2-28-1 


21-0-27-0 


4-9 


AL 


44 *7 ±0-46 


2*16^:0*32 


38-2-S1-2 


43-0-47-0 


4-8 


PL 


53-l±0-47 


2-20±0-33 


46-5-59-4 


50-0-57-0 


4-1 



Sena. 29-0 with head 18/18. 

Schongastia (Ascoschongastia) mackerrasae sp. n. 

Plate 45, fig. D-I. 

Descripticm of Larvae. Shape an elongate oval. Length (unengorged) 
365/*,, width 220/*. Scutum as figured with well rounded postero-lateral corners, 
and PL placed very near to AL; sensillae globose and finely setulose with their 
bases behind line of PL; AL setae the shortest, PL the longest, and all scutal 
setae strongly dentate but denticles rathetr shorter and more numerous than on 
dorsal setae. Eyes 2 + 2, closely adjacent to scutum, posterior the smaller. 
Palpi stout, femur strongly angulate, with seta long and strongly ciliated; 
seta o)i genu much finer with fewer branches ; all setae on palpal tibia nude ; 
tibial claw bifurcate; tarsus with 5-6 long ciliated setae. Cheliccrae non- 
serrate, with only the usual apical tricuspid cap. Galeal setae nude- Dorsal 
setae numerous, ca. 100 to 70-56/*, strong with 5-6 strong denticles, arranged 
ca. 8.16.18. plus. Ventrally with a pair of branched setae on maxillae, a long 
fine ciliated seta on each coxa, a pair between coxae I, and 6 between coxae HI, 
the outer ones of which lie close in the angle formed by the peculiar shaped 



WOMERSLEY — ASIATIC- PACIFIC SCRUB TYPHUS MlTES 183 

coxae III with coxae II, thereafter ca. 80-100 setae to 42/x long. Legs all 
7-segmented; I 350/a long, II 312/x, III 390/*; tarsi I and II with dorsal sen- 
sory rod ; III without any long- nude setae. 

The Standard Data for the unique type are: AW 58-8, PW 75-6. SB 33-6, 
ASB 25*2, PSB 16-8, SD 42-0, A-P 14-0, AM 53-2, AL 30-8, PL 78-4, Sens. 
36*4 with head 19-6/25-2. 

Loo. amd Host. A single specimen found free living, Bramston's Beach, 
N. Queensland, Sept., 1949 (M. J. and I, M. Mackerras). 

Remarks. This species is striking in the dentate dorsal setae, and shows 
some relationship to certain members of the jolicda group. It is named in 
honour of Mrs. M. J, Mackerras, one of the discoverers. 

Schongastia ( AscosoHONGA&TU) masta (Traub and Sundermeyer 1950). 

Ascoschongastia masta Traub and Sundermeyer 1950, Proc. Helminth. Soc. 
Washington, 17, (1), 35-38, fig. 1-4. 

Plate 68, fig. A-D. 

Description of Larvav (after Traub and Sundermeyer). Shape oval, often 
with two faint constrictions on abdomen. Length 287//., width 220/i. Scutum 
small, wider than long; anterior margin concave but medially sinuate, posterior 
corners well rounded,- PL situated off the scutum, AL the shortest; sensillae 
globose with sparse strong setules, the bases in midline of scutum; surface fine 
punctate. Eyes 2 + 2, on ocular shields, posterior the smaller, Chelicerae with 
only the apical tricuspid cap. Galeal setae short and nude. Palpi stout, tibial 
claw trifurcate; seta on femur strongly branched, on genu long and nude, on 
tibia dorsal and ventral branched but lateral not specified or figured. Dorsal 
setae short, to 25/x long and ciliated, 46 in number, and arranged 2.10.8.10.8.6.2, 
Ventrally with paired branched setae on maxillae ("galea!" of T. and S.), a 
pair between coxae I and between coxae III, one on each of coxae I and II, two 
on coxae III, and thereafter about 36 setae somewhat thinner and shorter than 
dorsal. Legs: I 220/x, TT 170/x, III 190/x; tarsi I and II with dorsal sensory 
rod (spur). III without any long nude seta. 

The Standard Data as given for the type and 3 paratypes are: AW 38-42 
(aver. 40), PW 66-74(70), SB lf>-24(21), ASB 14-19(17), PSB 13-16(14), 
SD 27-35(31), A-P 32, AM 28, AL 14, PL 24, Sens. 30-32(31). 

Loc. and Host., Described from specimens from Battus rattus, from Myit- 
kyina, North Burma, 29 Nov., 1944, with additional specimens from Tupaia 
lelamgeri. (a tree shrew), 24 Dec, 1944, and a shrew Crocidura sp,. 2 Feb., 
1945, all collected by members of the U.S. Typhus Commission. 



184 Records of the S-A. Museum 

Remarks, The authors place this species in the genus Ascoschongastia 
Ewing in the restricted sense as used by Wharton 1948 and as typified by the 
genotype malwyensis Gater. The position and validity of the generic name 
Ascoschongastia is discussed earlier in this paper. 

The position of PL off the dorsal scutum is similar to that of malayensis, 
from which the present species differs in the number of DS, the trif urcate palpal 
claw, and the bisetose coxae III, as well as the globose instead of lanceolate 
sensillae. 

Sohongastia (Ascoschongastta) petrogale (Worn. 1934). 

Neoschongastia petrogale Womersley 1934, Rec. S. Aust. Mus., 5, (2), 215; 
Womersley and Heaslip 1943, Tr. Roy, Soc. S. Aust. 67, (1), 111. 

Plate 45, fig. J-M. 

No further specimens of this rather striking species, which was described 
from the scrotum of a wallaby collected by Dr. C. J. Hackett in the Musgrave 
Ranges, South Australia, in July, 1933, have been received. 

The palpal claw is trif urcate ; the setae on the palpal femur and genu are 
branched and on the tibia the dorsal and lateral nude, ventral branched. The 
chelicerae are simple, with only the apical tricuspid cap; and the galeal setae 
are nude. The globose sensillae are finely but distinctly setulose. 

The Standard Data (re-measured) for eleven specimens of the type series 
are: 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Kange 


Range 


Variation 


AW 


73-52:0-67 


1-64+0-47 


68-6-78-4 


72-0-75*0 


2-2 


PW 


98-8±l-22 


2-99±0-86 


89-8-107-6 


97- 0-104-0 


3-0 


SB 


34-7±0-84 


2-07±0-59 


28-5-40-9 


32*0-36*0 


6-0 


ASB 


32*0 


No- variation recorded 






PSB 


30*0 


No variation recorded 






SD 


62-0 


No variation recorded 






A^-P 


360 


No variation 


recorded 






AM 


41-0±0-63 


l*56=fcD-45 


36-3-45*7 


40-0-43*0 


3-8 


AL 


49-7±:l-05 


2*58:fcl-05 


42-0-57-4 


47-0-54-0 


5-2 


PL 


55-0±:0-63 


l-56±0»45 


60-3-59-7 


54*0-57-0 


2-8 


Sens. 


32-0 with head 3 


17/17. 









Sohongastia (Ascoschongastia) mohri sp. n. 
Plate 46, fig. A-C. 

Description of Larvae. Shape broadly oval. Length (engorged) to 450/a, 
width to 360/*.. Scutum rectangular, with PL setae placed on the sides, rela- 
tively near to AL and not at the broadly rounded posterolateral angles : scutum 
widest behind line of PL; posterior scutal margin lightly sinuous; sensillae 



Womerslev— Asiatic-Pacific Scrub Typhus Mites 185 

bases behind line of PL and about in mid-line of scutum. Eyes 2 -f % appar- 
ently not on ocular shields, posterior the smaller. Chelicerae simple, with only 
the apical tricuspid cap. Galeal setae nude. Palpi stout, tibial claw trif urcate ; 
all setae on femur, genu and tibia branched. Dorsal setae ca. 44 in number, 
to 65/t long and arranged 2.10.10,10.6.4.2. Dorsal cuticular striatums markedly 
crenulate (in good fresh mounts) as in following species. Ventrally with a 
pair of branched setae on maxillae, one on each coxa, a pair between coxae I 
and between coxae III and thereafter ca. 60 setae to 36^ long. Legs I 330/* 
long, II 300/*., Ill 320/4 ; tarsi I and II with the usual dorsal sensory rod; 
tarsi III without any long nude seta. 

The Standard Data for the type and 5 paratypes are ■ 





Mean 


Standard 
Deviation 


Theoretical 
Eongc 


Observed 
Range 


Coeff . of 
Variation 


AW 


74-2±l-19 


2-93±0-85 


$5* 4-83*0 


69-0-77 '0 


3-9 


PS 


87-0±]-17 


2'89±0-83 


78-3-95*7 


83-0-91-0 


3-3 


SB 


28-3±0-49 


J-21±0-35 


24- 7-31- 9 


27*0-30*0 


4-2 


ABB 


29«2±0-54 


l-33±0-38 


25-2-33-2 


27-0-30-0 


4-5 


P8B 


30-0 


No variation recorded 






SD 


59-2±0-54 


l-33±0-38 


55-2-63 -2 


57-0-60-0 


2-2 


A-P 


21-8+0-48 


1'17±0«34 


18-0-25-3 


21 -0-24-0 


5-4 


AM 


56-4±0-40 


0-89±0-28 


53-7-59-1 


56-0-58-0 


1-6 


AL 


46'0±1-11 


2-49±:0-79 


38*5-53.5 


42-0-48-0 


5 '4 


PL 


78-0:t0-95 


2-12±0-G7 


71-7-84-3 


75-0-80-0 


2*7 



Sens. 36-0 with head 16/38. Only 3 determination. 

Loc. and Hosts, The type and 5 paratypes from Eckyympera doreyaim 
from Sansapor, Dutch New Guinea, 16 Nov,, 1944 (C. Mohi\); another speci- 
men from same locality on Battus concolor 19 Nov., 1944 (CJVL). 

Remarks: In the general form of the scutum with short A-F and the 
crenulate dorsal striations this species is closely related to the following, 
S* (A.) crinita sp. n. from which it diifers as in the key. In the type, the sen- 
sillae which have become detached show an involucrum-like structure at the 
base of the stem. 

It is a pleasure to name this species after the finder, Dr. C. Mohr, from 
whom much interesting material was received. 

SCHONOASTIA (AsCOSCHONQASTTA) CRINITA sp. n. 

Plate 46, fig. D-G. 

Description of Larvae. Shape ovaL Length (unfed) 285/*, width 185/!., 
Dorsal scutum as figured, very much as in mohrt, but with PL much closer to 
AL, and with the expansion behind PL greater; finely punctate. Sensillae 
missing in all specimens; the bases well behind line of PL. Chelicerae non- 



186 Records of the S.A. Museum 

serrate, with the usual apical tricuspid cap. Galeal setae nude. Eyes 2 4- 2 T 
on ocular shields; posterior the smaller. Palpi stout, with trifurcate tibial 

claw; all setae on palpal femur, genu and tibia ciliated or branched. Dorsal 
setae more than 100, arranged ca. 2.14.12.16.12.12.12.10.8.6.4, rather stouter 
and not so tapering as in mohri, with shorter ciliations, to 60/*. long. Dorsal 
cuticle with crenulate striations. Ventrally with a pair of branched setae on 
gnathosoma, one on each coxa, a pair between coxae T and between coxae 111, 
and thereafter ca. 100 to 20/* long. Legs I 300/* long, II 270m, ni 300/*, tarsi I 
and II with the usual dorsal sensory rod, III without any long nude seta. 
The Standard Data for the type and 2 paratypes are i 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Kange 


Coeff. of 
"Variation 


AW 


83-7:±2-03 


3-51±l-43 


73-2-94-2 


80-0-87^0 


4-2 


PW 


92-3±2-33 


4-04±l-65 


80-2-104-4 


90*0-07-0 


4-3 


8B 


30-7=£0-67 


l*15±0-47 


27-2-34-2 


30*0-32-0 


3-7 


ABB 


33-3^1-33 


2-31+0-94 


26-4-40*2 


32 '0-36-0 


7-0 


P8B 


30-0 


No variation recorded 






SB 


63-3±l-33 


2-31±094 


56*4-70-2 


62-0-66-0 


3-6 


A-P 


17-0±l-00 


l-73±0-71 


11-8-22-2 


15-0-18-0 


10-2 


AM 


54-0 


One determination only. 






AL 


45'7±l-33 


2-31:1:0 -94 


38-8-52-6 


43-0-47-0 


5-0 


PL 


90-0 


No variation recorded 







Sens. Missing in all specimens. 

Lot. Described from 3 specimens from Echymipera doreycma from Sansa- 
por, Dutch New Guinea, 1945 (W. D. Fitzwater). 

Remarks. Very -close to the preceding species in the crenulate dorsal 
striations and the dorsal scutum. Differs in the much greater number of DS 
and the shorter distance between AL and PL, as well as in the scutal width 
behind PL being somewhat greater. 

SOHONGASTIA (AsCOSCHONGASTIA) RECTANGULAR^ sp. n. 

Plate 46, fig. H-L. 

Description of Larvae. Shape oval. Length (only slightly engorged) 
375/^ width 240/a. Dorsal scutum as figured with PW only slightly greater 
than AW, i.e. almost rectangular with lateral margins parallel or almost so; 
sensillae bases wide apart and about midway between line of AL, and line of 
PL, and also nearer to lateral margins than to one another; AL much longer 
than AM or PL and placed well back from antero-lateral corners; sensillae 
with large globose heads with distinct sctules; surface of scutum punctate and 



Womersley — Asiatic-Pacific Scrub Typhus Mites 187 

the posterior half overlapped by the dorsal cuticular striations. Eyes 2 + 2, 
large, on distinct ocular shields and the posterior eye the smaller. Chelicerae 
not serrate, only furnished with the usual apical tricuspid cap. Galeal setae 
nude. Palpi stout, tibial claw trifurcate; setae on femur and genu branched, 
on tibia dorsal and lateral nude, ventral branched. Dorsal setae 36, to 60ft 
long and arranged 2.8.6.8.6.4.2, slender and finely ciliated. Ventrally on maxil- 
lae with a pair of branched setae, a single seta on each coxa, a pair between 
coxae I and between coxae III, and thereafter 6.6.4.4.2, to 45/x long. Legs 
I 390>t long, II 360/i, and III 390/a; tarsi I and II with usual dorsal sensory 
rod; tarsi III with a long nude seta. 

The Standard Data from the type and 4 paratypes arc: 





Mean. 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 


76-2±0-56 


l-25±0-39 


72-4-80-0 


75-6-78-4 


1-6 


PW 


81-8±l-05 


2-34±0-74 


74-8-88-8 


78-4-84-0 


2*8 


SB 


47*0±0-56 


l'25±0-39 


43-2-50-8 


44-8-47-6 


2-7 


ASB 


28-0 


No variation 


recorded 






P8B 


31-9±0-28 


0-62i0 -20 


30-0-33-8 


30-8-32-2 


2-0 


SD 


59-9^0-28 


0-G2±0-20 


58-0-61-8 


58-8-60-2 


1-0 


A-P 


30-8 


No variation 


recorded 






AM 


54-0±2'00 


4-00+1-41 


42-0-66-0 


48-0-56-0 


7-4 


AL 


82-D±:l'42 


3-18+1-00 


73-4-92-4 


78 -4-86 -8 


3-8 


PL 


58- 2+1 -05 


2-34+0-74 


51*2-65-2 


56-0-61-6 


4-0 


Sen?. 


39-2 with head 


22-4/22-4. No 


variation, recorded. 







Loo. The type and 4 paratypes from Lake Eae-ham, N. Queensland, Dec, 
1944 (R.N. MeCulloch). 

Schongastia (Ascoschongastia) cassiope sp. n. 
Plates 47, fig. E-P; 48, fig. A-B. 

Descri'ptim of Larvae. Shape oval. Length (unfed) 240/a, width 170/x. 
Dorsal scutum as figured, with both anterior and posterior margins sinuous, AM 
fqhort, PL the longest; sensillae globose and finely setulose, with bases very 
slightly in front of line of PL. Eyes 2 + 2, fairly large, on distinct ocular 
shields, posterior eye the smaller. Chelicerae non-serrate, with only the apical 
tricuspid cap. Galeal setae nude. Palpi stout, seta on femur with 4-5 branches, 
on genu with 3-4 branches, all on tibia nude; palpal claw trifurcate. Dorsal 
setae long, thick and strongly ciliated, 32 in number, and arranged 
2.6.6.6.6.4.2, to 56/x. long. Ventrally with a pair of branched setae on max- 
illae, one on each coxa, a pair between coxae I and between coxae TIT, and 



188 Records of the S,A. Museum 

thereafter ca. 22, shorter and finer than dorsal setae. Legs all 7-segmented, 
I 240/a long, II 2]%, Ill 280/*; tarsi I and II with usual dorsal sensory rod, 
III without any long nude seta. 

The Standard Data for the type and two paratopes are | 

Coeff. of 
Variation 

2-6 

3'S 







Standard 


Theoretical 


Observed 




Mean 


Deviation 


Range 


Range 


AW 


60-7+0-93 


l-62±0-66 


55-9-65-5 


58-S-61-6 


PW 


72-8±l-62 


2*80±:1-14 


64-4-81-2 


70-0-75-6 


SB 


19-6 


No variation recorded 




ASB 


25-2 


No variation recorded 




PSB 


16-8 


No variation recorded 




SD 


42-0 


No variation recorded 




A-P 


23-3=t0-93 


l-62±0-66 


18 -5-28-1 


22-4-25-2 


AM 


40 -Oca. 


No variation 


recorded 




AL 


71-9±3-61 


6-35±2-59 


57-9-85*9 


64-4-75-6 


PL 


79-3zt4-66 


8-08:±3«30 


55-J-103-5 


70-0-84-0 



6-5 

8-8 
10-2 



Sens. 30-8 with head 19-6/19-6. No variation recorded. 



Loe. The type on boots, Wandecla, N. Queensland, 2 Oct., 1944 (R, N. Mc- 
Culloch) and two paratypes, Wongabel, N.Q., Dec, 1944 (R. N. McO.) 
Bemarks. The relationship of this species is best seen from the key. 

Sohongastia (Asoosohongastia) antipodianum (Hirst 1929). 
Schdnga$tia antipodianum Hirst 1929, Proe. ZooL Soc. London, (1), 175. 

Neoschongastia antipodianum, Womersley and Heaslip 1943, Tr. Roy. Soc. 
S. Aust., 67, (1), 114. 

Plate 47, fig. A-D. 

No fresh material of this species has been seen since the original discovery. 
It was found in the ears of Battus greyi from D'Estree Bay, Kangaroo Island^ 
South Australia and the co-type material is in the collection of the South Aus- 
tralian Museum. 

The palpal tibial claw is trifureate and all the setae on the palpal femur, 
genu and tibia are branched or ciliated. The chelicerae are non-serrate with 
only the apical tricuspid cap. The galeal setae are nude. 

The dorsal scutum is almost rectangular, with AW only slightly shorter 
than PW, and the lateral margins subparallel • AL are slightly longer than PL 
and placed in the anterolateral earners; sensillae globose, indistinctly setulose 
and their bases somewhat nearer line of PL than line of AL. Eyes large, on. 
distinct ocular shields, the posterior eyes the smaller. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 189 

The Standard Data as redetermined from 7 specimens of the co-type 
material are : 



AW 


Mean 
60-8±0-31 


Standard 
Deviation 

0-81:t0'22 


Theoretical 
Range 

58-4-63-2 


Observed 
Range 

59-8-61-6 


Ooett of 
Variation 

1-3 


PW 
SB 


69-6±0-95 
25-2 


2-52:±0-67 62-1-77-1 
No variation recorded 


67-2-72-8 


3-6 


ASB 


28-8±0-52 


l-37db0-36 


24-7-32-9 


28-0-30-8 


4-7 


PSB 


19-6 


No variation recorded 






SB 


48-4±0-52 


l-37±:0-36 


44-3-52-5 


47-6*-50«4 


2-8 


A-P 


39'2±0-61 


l>63±0-43 


34-4-44*0 


36-4-42-0 


2-7 


AM 


38-2±0-60 


l-34±0-42 


34-2-42-0 


36-4-39-2 


3-5 


Ah 


67-8±l-05 


2-34:±0-74 


60-8-74-8 


64-4-70-0 


3-4 


PL 


59-9+1-42 


3-19±l-01 


50-3-69-5 


56-0-64-4 


5-3 



Sens. 28-0 with head 22 -4/22 '4. No variation recorded, 

Schongastia (Ascoschongastia) dumosa sp. n. 
Plate 48, fig, C-F. 

Description of Larva, Shape oval. Length (unfed) 300/*, width 210/*. 
Dorsal scutum as figured, with anterior margin convex, and posterior margin 
somewhat angular; sensillae bases in front of line of PL and surrounded by 
strongly defined crests; PL the longest, AL the shortest; sensillae globose, and 
strongly setulose. Eyes fairly large, 2 + 2, on ocular shields, and the posterior 
only a little smaller than anterior. Chelicerae simple, with only the apical 
tricuspid cap. Galea! setae nude. Palpi stout, tibial claw trifurcate; setae on 
palpal femur and genu strongly branched; on tibia, dorsal and lateral nude r 
ventral branched. Dorsal setae strong and strongly ciliated or serrate, ca. 52 
in number and arranged 2.10.10.10.8.6.4.2 to 30/a long. Ventrally a pair of 
branched setae on maxillae, one on each -coxa, a pair between coxae I and 
between coxae III and thereafter ca. 3 rows of 10, plus 8.6.4, to 20/x long and 
posteriorly gradually approaching dorsal setae in form. The striations of the 
dorsal cuticle very finely crenulate. Legs I and II with dorsal sensory rod, 
III without any long nude seta. 

The Standard Data are: AW 58-8, PW 75-6, SB 22-4, ASB 22 <4 t PSB 
22-4, SD 44*8, A-P 25*2, AM 45 '0 AL 33-6, PL 56-0. Sens. 36-4 with head 
19 *6/19 -6. 

Loc. The type only, from Atherton, Queensland, on boots, April 5, 1944 
(R.N.McCulloch). 

Remarks. The unique type slide in the S,A. Mus. collection was regrettably 
badly damaged while under oil immersion. 



190 Records of the S.A. Museum 

Schongastia (Ascoschongastia) queenslandica (Womersley 1939). 

Neoschongastia queenslandica Womersley 1939, Tr. Roy. Soc, S, Aust., 63, (2), 
162; Womersley and Heaslip 1943, Tr. Hoy. Soc. S. Aust., 67, (1), 112. 

Plate 48, fig. G-K. 

The Standard Data for 10 of the 12 specimens recorded in 1943, as re- 
measured are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


57-9±0-74 


2'33±0-52 


50-9-64-9 


54-0-61-0 


4-0 


PW 


90-7±:0'84 


2-62:£0-59 


82- 8-98 -6 


87-0-97-0 


2-9 


SB 


29-8±0-53 


1» G9:±0 -38 


24-8-34-8 


27-0-32-0 


5-6 


ABB 


31-6±0-27 


0-84±0-19 


29-1-34-1 


30-0-32-0 


2-7 


PSB 


18-8±0-33 


1-03±0'23 


15'7-21-9 


18-0-20-0 


5-5 


SB 


50-4:£0-50 


l-58±0-35 


45-7-55-1 


48-0-52-0 


3-1 


A-P 


42-8:1:0-85 


2-70±0-60 


34 -7-50 »9 


39-0-47-0 


6-3 


AM 


39-4±0-43 


l-35dt0-30 


35-4-43-4 


36-0-40-0 


3-4 


AL 


70-85±l-16 


3-08±0-82 


61-6-80*1 


65-0-75-0 


4-3 


PL 


67-l±0-67 


l-88±0-47 


61*5-72-7 


65-0-70-0 


2-8 


Sens. 


40-0 with head 


20/21. 









I have recently seen 5 more specimens collected by Major R. N. McCulloch 
on boots: Cairns, Queensland, 3 spec. July, 1943; Skull Pocket, QkL, 1 spec. 
June, 1943, and Atherton, Qld., 1 spec. Sept., 1944. Also numerous specimens 
from the ears of a euro, near the Daly River, Northern Territory of Australia, 
S June, 1943 (R. V. Southeott, coll. A.C.B. 162). 

The species is characterized from its allies by having 4 setae in the first 
dorsal row, i.e. two submedial as well as the two scapular setae. The posterior 
margin of the scutum is very shallow behind PL, being almost rectilinear. The 
AL and PL setae are long, with Ah slightly the longer. Chelicerae are simple 
with only the usual apical tricuspid cap, Galeal setae nude. Palpi stout, with 
trifurcate tibial claw; setae on femur and genu branched; on tibia, dorsal and 
lateral nude, ventral branched. Dorsal setae 4.6.6.6.2.2, to 50/x long. Ventral 
setae, posterior of coxae III 8.6.6.4.4.2, to 21/x long. Sensillae globose and 
finely setulose. 

SCHOKOASTIA (AgCOSCIlONGASTIA) LAPPACEA n. UOV. 

NeoscJwiigastia melomys Womersley and Heaslip 1943, Tr. Roy. Soc, S. Aust., 
67, (1),110 (in part). 

Plate 49, fig. A-D. 

On critical re-examination of the type of N. meloinys from Melomys Uttor- 
atis from Innisfail, Queensland, it was found to be con-specific with irmisfail- 



Womersley — Asiatic-Pacific Scrub Typhus Mites 191 

ensis Worn, and Heasp. 1943 (Tr. Roy. Soc. S. Aust., 67, (1), 108) but not 
with the series of 15 specimens from rats at Cairns, Queensland, and recorded 
as melomys {loc. tit.). 

The name melomys thus becomes a homonym and invalid and the material 
from Cairns is herewith redescribed under the name of lappaeea. 

Description of Larvae. Shape oval with sides rather flattened. Length 
(engorged) 465/i, width 260/&. Dorsal scutum as figured, anterior margin lightly 
sinuous, posterior margin more or less evenly rounded but slightly flattened 
or concave medially, and fairly deep behind line of PL ; sensillae globose with 
fine indistinct dilations, the sensillae bases anterior of line of PL - PL the long- 
est and longer than PW, Eyes 2 -)- 2, rather small, posterior the smaller. 
Chelicerae simple, with only the apical tricuspid cap. Galea! setae nude. Palpi 
stout with trifurcate tibial claw; setae on femur and genu branched, setae on 
tibia nude except the ventral sometimes with a. medial branch. Dorsal setae 
32 in number to 75/* long, rather strong with strong setules, arranged 
2,6.6.6.6.4.2. Ventrally with a pair of branched setae on maxillae, one on each 
coxa, a pair between coxae I and between coxae III and thereafter 4.6.6.6,4.4, 
to 30-45/t long. Legs I 220/*, II 190/*, III 220/*; tarsi I and II with dorsal 
sensory rod, III without any long nude seta. 

The Standard Data for the type and 9 specimens from rats, Cairns, Qld. 
1939-40 (W.G.H.), 8 specimens from the ears of a euro, at Daly River, North- 
ern Territory, Anst (R. V. Southcott), and 2 specimens from boots from 
Skull Pocket, Kairi, Qld., July 1943 (E. N. McCulloch), are: 







Standard 


Theoretical 


Observed 


Ooeff . of 




Mean 


Deviation, 


Kange 


Range 


Variation 


AW 


54*0±0-45 


2-00±:0-32 


48-0-60-0 


50-4-56-0 


3-7 


PW 


70-4±0-53 


2-39±0-38 


63-3-77-5 


64-4-72-8 


3*4 


SB 


19-6±0.28 


l«25:k0*20 


15-8-23-4 


16-8-22*4 


6-4 


ASB 


25-2 


No variation recorded 






PSB 


19-6 


No variation recorded 






BD 


44-8 


No variation recorded 






A-P 


27-8±0-27 


1-19 ±0*19 


24*2-31*4 


25*2-29-4 


4-3 


AM 


81-6±0-69 


3 -08:1:0 -49 


22*4-40-8 


28*0-39-2 


9-7 


AL 


70-4±O-36 


l-60=h0<25 


65-6-75-2 


67-2-72-8 


2-3 


PL 


S0'9±:0-81 


3-64dr0-57 


70-0-91-8 


75-6-86-8 


4-5 



Sens. 33-6 with head 16 -8/19 '6. No Variation recorded. 

Sohongastia (Ascoschongastia) womersleyi (Gunther 1940). 

Neoschongastia womersleyi Gunther 1940, Proc. Linn. Soc. New South Wales, 
65, (3-4), 254 j Womersley and Heaslip 1943, Tr. Boy, Soc. S. Aust., 
67, (1), 109. 



192 Records of the S.A. Museum 

Plates 49, fig. E-G; 50, fig. A. 

Of this species I have another specimen collected from a rat (No. 9) by 
Maj. G. M. Kohls in New Guinea, 15th Nov., 1943. 

The Standard Data as remeasured for 7 of Gunther's paratypes in the 
Sonth Australian Museum collection, and the additional specimen referred to 
■above are : 





Mean 


Standard 
Deviation 


Theoretical 
Hange 


Observed 
Range 


Coeff. of 
Variation 


AW 


47-95:±0-63 


l-79±0-45 


42-55-53-25 


44- 8-50 '4 


3-7 


PW 


68-95±l-05 


2-97rt0-74 


60-05-77-85 


67-3-75.6 


4»3 


SB 
ABB 


19-25±0*35 
28-0 


0-99+0 '25 16-25-22-25 
No variation recorded 


16-8-19-6 


5-1 


PSR 


18-9±0-46 


l-29±0-32 


15-0-22-8 


16.8-19-6 


6-8 


SD 


46-9:±0-46 


l-29±0*32 


43-0-50-8 


44-8-47-6 


2-8 


A-P 

AM 

At 


31-5±0-46 
39-2:t0-61 
84-0 


l-29±0*32 27-6-35-4 
l-62±0-43 34-3-44-1 
No variation recorded 


30-8-33-6 
36-4-42-0 


4-1 
4-1 


PL 
Sens. 


134-6±1-7S 
33-6 with head 


4-72±l-26 
16-8/19-6. Onlv 


120-4-148-8 
1 determination. 


126-0-140-0 


3-5 



This is a very remarkable species in the excessively long PL setae which 
are about twice the length of PW. The scutum is as figured with the posterior 
margin relatively shallow behind PL. The sensillae are globose with fine 
setules ? and their bases nearer to one another than to the lateral scutal margins 
and only slightly in front of line PL. The chelicerae are not serrate and have 
only the usual apical tricuspid cap. The galeal setae are nude. The palpal 
tibial claw is trifurcate and the setae on the femur and genu branched, on the 
tibial only the ventral seta with branches. The dorsal setae are strongly 
ciliated, to 75jt long, and 28 in number arranged 2.6.6.6.4.2.2. 

Sohongastia (Ascoschongastia) coorongense (Hirst 1929). 

Sohongastia coorongense Hirst 1929, Ann. Mag. Nat. Hist., (10), 3, 564, 

Neosehongastia coorongense, Worn, and Heasp. 1943, Tr. Roy. Soc. S. Aust., 
67, (1), 114. 

Plate 50, fig. B-E. 

This species, the original description of which was unaccompanied by any 
figure, is still only known from the original material from the ears of a rat 
from Robe, S. Australia. The co-types are in the South Australian Museum, 

The palpal claw is trifurcate; the seta on the palpal femur has many 



Womersley— Asiatic-Pacific Scrub Typhus Mites 193 

rather short branches, on the genu somewhat fewer short branches and all 3 
setae on the tibia are nude. The chelicerae are simple and armed only with 
the usual apical tricuspid cap. The galeal setae are nude. The dorsal scutum 
is as figured with the setae relatively slender and finely ciliated and PL the 
longest, AL the shortest; the sensillae are capitate, strongly setulose, and with 
their bases surrounded with strong ehitinous lines and placed slightly in front 
of line of PL. The dorsal setae are fairly slender, shortly ciliated, 32 in num- 
ber, to 45/i long and arranged 2.6.6.6.6.4.2. The ventral setae behind coxae III 
are ca. 8.8.8. to 25/* long, then 6.6.4.2, to 40/x long. There is no long nude 
seta on tarsi III. 

The Standard Data for 15 co-types as remeasured are : 



AW 


Mean 
70-9±0-93 


Standard 
Deviation 

3 -60±:0- 66 


Theoretical 
Itange 

60 -1-81 -7 


Observed 
Range 

67*2-78-4 


Coeff . of 
Variation 

1-8 


PW 


95-95±l-50 


5 '82+1 -06 


78-5-113-4 


86-8-109-2 


6-1 


SB 


32-8±:0-67 


2-60zfc0-47 


25-0-39-8 


28-0-39*2 


7-9 


ASB 


30-8±0<37 


1*45^=0-26 


26 -5-35*1 


28-0-33-6 


4-7 


PSB 


22-4 


No variation recorded 






SD 


53-2±:0-37 


1-45:^0-26 


48-9-57-5 


50-4-56-0 


2-7 


A-P 


34-5±0-51 


1-96+0-36 


28-6-40-4 


33-6-39-2 


3-5 


AM 


49«I±0'45 


1-73+0-32 


43-9-54-3 


44-8-50-4 


3*5 


AL 


42-0±0-46 


1- 77=4:0 -32 


36-7-47-3 


39-2-44-8 


4-2 


PL 


64-4±0-95 


3-68+0-67 


53-4-75-4 


56-0-70-0 


5-7 



Sena. 33-6 with head 19-6/19-6. Ko variation recorded. 

Two specimens were recently taken along with one S. (A.) dasycerci (Hirst) 
from Battus murrayi var. on Greenly Is., South Australia, Dec, 1947 (P. J- 
Mitchell). 

Schongastta (Ascoschongastia) echymitera (Worn, and Kohls 1947). 

Ascoschongastia: eohymipera Womersley and Kohls 1947, Ti\ Roy. Soc. S. Aust. 
71, (1), 11-12. 

Plates 50, fig. F-I; 51, fig. A. 

This species was described from specimens collected from a bandicoot, 
Echymipera cocker elli from Dobodura, New Guinea. Nov. 1943 (G. M. Kohls). 
In the key it comes close to irmisfaitemis Worn, and Heasp. 1943, but can be 
distinguished therefrom by the posterior scuta! margin being very shallow, 
almost rectilinear between PL, and by the rather greater number of dorsal 
setae. 



19+ Records of the S.A. Museum 

The Standard Data as given by Woniersley and Kohls are ; 







Standard 


Theoretical 


Observed 


Coeff. of 




MTean 


Deviation 


Range 


Range 


Variation 


AW 


45*3±0-29 


l-16±0-20 


41 -8-48*8 


43-0-48-0 


2-5 


PW 


61-5;±0-53 


2-12±0-37 


55- 1-(>7 -9 


o7-0-<"6-0 


3-4 


SB 


20'4±0-39 


1-I7ir0-21 


16-9-23-9 


18 -0-21-0 


5-7 


ASB 


21-0 


No variation recorded 






PSB 


15-0 


No variation recorded 






SD 


36-0 


No variation recorded 






A-P 


31-3db0-37 


l-49±0-26 


26-8-35-8 


30-0-33-0 


4-7 


AM 


26-6±0-36 


1*45±0'26 


22-2-31-0 


24-0-30-0 


5-6 


AL 


46-5±:0'59 


2-37±0<42 


39 '4-53 »6 


42-0-51-0 


5-1 


PL 


42-6:t0-36 


l-45±0-26 


38-2^7-0 


40-0-45-0 


3*4 



Sena. 27-0 with head 17/17. No variation recorded. 

Schongastia ( Ascoschongastia) jnnisfailensis (Worn, and Heaslip 1943). 

Neoschongastia inmsfadlensis Womersley and Heaslip 1943, Tr. Roy. Soc. 
S. Aust, 67, (1),108. 

Neoschongastia melomys Womersley and Heaslip 1943, Tr. Roy. Soe. S, AuBt., 
67, (1), 110 (in part). 

Ascoschdngastia lushlandi Philip 1947, J. Parasitology, 33, (3), 387. 

Plate 51, fig. B-E. 

Re-examination of the type specimen of Neoschongastia melomys Worn, and 
Heasp., from Innisfail, Queensland, shows it to be co-specific with mnisfailemis, 
but not so with the rest of the material from Cairns, and recorded as melomys 
(see under fifr (AscoschongaMia) lappacea, n, nov.). 

Re-description of Larvae. Shape elliptical. Length (engorged) to 450/x, 
width to 320/4. Dorsal scutum as figured, with AL the longest but only slightly 
longer than PL; anterior margin sinuous and convex medially; sensillae glo- 
bose, appearing nude, but under high magnification with indistinct setules which 
are most obvious near the base of head; sensillae bases nearer to one another 
than to lateral margins and practically in line with PL. Eyes 2 + 2, on ocular 
shields, the posterior the smaller. Chelicerae non-serrate, with only the apical 
tricuspid cap. Galeal setae nude. Palpi stout with trifurcate tibial claw; 
setae on femur and genu branched, on tibia all 3 nude. Dorsal setae tapering 
and with fairly strong setules, to 45/* long and arranged 2.6.6.6.6.4.2. Ventraily 
with a pair of branched setae on maxillae, one on each coxa, a pair between 
coxae I and between coxae III and thereafter ca. 38-40, to 35/* long. Legs I 
270/x long, II 225/x, III 300/*.; tarsi I and II with the usual dorsal sensory rod. 
Ill without any long nude seta. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



195 



The Standard Data from twelve of the original specimens from Innisfail, 
Queensland, from Melomijs liti oralis are ; 







Standard 


Tkcoictical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


.Range 


Variation 


AW 


47-8±0-88 


3-03±0-62 


38-7-56-9 


44-8-56-0 


6-3 


PW 


72-l±l-15 


3-98±0-81 


60-2-84-0 


67*2-78-4 


5-5 


SB 


21-5±0-44 


l-52r±:0-31 


17-0-26*0 


19-6-23-8 


7-0 


ASB 


28-0 


No variation 


recorded 






PSB 


19-6 


No variation recorded 






SB 


47-6 


No variation recorded 






A-P 


27*9+0-27 


0.93rh0-19 


25-1-30-7 


25-2-29-4 


3-3 


AM 


2S-55±0-37 


l-18±0-26 


25-0-32-1 


28 -0-30- 8 


4-1 


AL 


54-9:±0'48 


l-44±0-32 


50-6-59-2 


53-2-56-0 


2-6 


PL 


47-6±0-49 


l-69:±0-34 


42-6-52-6 


44-2-50-4 


3-5 


Sens. 


35-0 with head 


20/21, 









Recently Philip (loc. cit.) has described from Owi Is., DJSLG,, a new- 
species under the name of Ascoschongastia btishlandi. In his discussion Philip 
points out the close relationship to iimisfailensis, the main differences being in 
the somewhat larger size of the dorsal scutum, the longer ciliations on the 
scutal setae, and the presence of an involucrum at the base of the stem of the 
detached sensillae. This peculiar structure has not been described before, and 
is probably to be found in many species but obscured by the structure of the 
sensillary cup unless the sensillae become detached. I have since seen the same 
structure in $ (A.) mohri sp. n. as mentioned under that species, and it also 
occurs in inwisfaileims* 

In other morphological details bushlandi cannot be separated from mnis- 
faUen8is f and on the slightly larger scutum, particularly PW, it should prob- 
ably be regarded as not more than a geographical race. I am greatly indebted 
to Dr. Philip for the original data of 28 specimens for which he gives the 
Standard Data in terms of observed range and mean in his paper. Treated 
more fully for comparison with that of innisf mien sis the Standard Data for 
bushlandi is as follows : 



AW 
PW 


Mean 
51-7d=0-37 
81-3+:0-72 


Standard 
Deviation 

1-07+0 -26 
3-83±0-5l 


Theoretical 

Range 
45-8-57*6 
69-8-92-8 


Observed 

Range 

47-8-55-1 

70-3-89-9 


Coeff. of 

Variation 

3-8 

4-7 


SB 


21-4±0-35 


1- 84+0 -25 


15*9-26-9 


15-2-26-7 


8-6 


ASB 


25-0+0-27 


1-43+0 -19 


20-7-29-3 


22-4-27-5 


5-7 


PSB 


21-9±:0-28 


1-49+0-20 


17-5-26-3 


18-2-24-6 


6-8 


SD 


47-0±0-45 


2-39+0*32 


39-9-54-1 


41-4r-51*6 


5-1 


A-P 


28-8+0-18 


0*94+0-12 


26*0-31-6 


27-0-31-0 


3-3 


AM 


30-4+0-46 


2-31+0-32 


23* 5-87 -3 


26-1-35-2 


7-6 


AL 


60-2+0*94 


4-42+0-67 


47-0-73-2 


46-0-68-1 


7-3 


PL 


56-3+0-48 


2-41*0-34 


49*1-63-5 


53-2-60-9 


4*3 


Bens. 


31-4+0-52 


1-88+0-37 


25-8-37-0 


27-5-34*8 


6-0 



196 



Records of the S.A. Museum 



In general bushlandi has AW slightly larger than in innisfailensis, PW 
definitely larger, and AL and PL also longer. 

In addition to the typical material from Innisfail, for which Standard 
Data are given above, there are 4 specimens in the South Australian Museum, 
which agree more with bushlandi than inmsfailensis in AW and PW, hut which 
have PSB, SD and A-P still greater. 

Of these one specimen was collected by W. D. Smith in Oct, 1938, and 
three by W. 6. Heaslip in 1940, all from the type locality and same host as 
typical inmsfarilensis* Their Standard Data are: 



AW 

PW 

SB 

ASB 

PSB 

SB 

A-P 

AM 

AL 

PL 

Sens. 



Mean 
51-l±0-70 
83-3±l-76 
24«5±0*70 
30«8 
25-2 
56-0 
36-4 

30-8^1*61 
58-8 

54-6:±0-81 
Missing. 



Standard 
Deviation 

l-40:±0-49 

3-52rirl-24 

1-40=1:0 -49 

No variation 

No variation 

No variation 

No variation 

3-23=tl*14 

No variation 

l-61±0-57 



Theoretical 
Range 

46-9-55-3 

72-8-93-8 

20-3-28-7 

recorded 

recorded 

recorded 

recorded 

21-1-40-5 

recorded 

49-8-59-4 



Observed 
Range 

50-4-53-2 

78-4r-86-8 

22-4-25-2 



28 -0-33 '6 



53-2-56-0 



Coeff . of 
Variation 

2'7 
4-2 

5-7 



10-5 



3-0 



More recently a small collection, of 14 .specimens, of ixnf ed larvae of this 
species, from the Bramston's Beach area. N.Q.. abont 60 miles south of Cairns, 
and 14 miles north of Innisfail (coll. I. M, and M. J. Mackerras, Sept., 1949) 
has been studied. 

The Standard Data of this series are : 





Mean 


Standard 
Deviation. 


Theoretical 
Bange 


Observed 
Range 


Coeff . of 
Variation 


AW 


46-l±0-52 


l-94±0-37 


40-3-51-9 


43-4-50-4 


4-2 


PW 


63-6±l-03 


3-87±0-73 


52-0-75-2 


58*8-70-0 


6-1 


SB 


19-2=£0-52 


l-94±0-37 


13 -4-25-0 


16-8-22-4 


10-0 


ASB 


23-8±0-57 


2-15+0-41 


17-35-30*25 


19-6-28-0 


9-0 


PSB 


18*0=±0-48 


1-81+0-34 


12- 6-23-4 


16-&-22-4 


10-0 


SD 


41*8:±0-46 


1-72+0-33 


36-6-47-0 


. 39-2-44-8 


4-1 


A-P 


26-8:4:0 -81 


3-05+0-58 


17-7-35-9 


22-4-30-8 


11-4 


AM 


37-2±0-62 


2 '31+0 -44 


30-3-^4-1 


33-6^*2-0 


6-2 


AL 


58.8±0-51 


l-90±0'36 


53-1-64-5 


56-0-31-6 


3-2 


PL 


50-4=£0-75 


2-U+0-53 


44-1-56-7 


47-6-53-2 


4-2 


Sens. 


31-9^=0-39 


1. 42 +0-28 


27 - 7-36 '1 


30-8-33-6 


4-4 



Womersley— Asiatic- Pacific Scrub Typhus Mites 197 

This population is significantly different from the original population, in 
all the Standard Data except AW and A-P, being lower in PW, SB, ASB, 
PSB, and SD, and Sen. and higher in AM, AL, and PL. The data for PL is 
based on 5 specimens only, as in one of these 5, only one PL was of normal 
length, the other measuring 70/x. In 4 other specimens both PL setae measured 
70/*. 

The population of f. lushlandi Philip from the Philippines, differs from 
the original population significantly in all Standard Data except SB and SD, 
being greater in AW, PW, PSB, All and AL, and smaller in ASB, PL, and 
Sens. 

This species, then, seems to be very variable in Standard Data, as seen in 
populations from different localities. 



B. SENSILLAE CLAVATE, SOMETIMES BROADLY SO, OR LANCEOLATE WITHOUT 

A BASAL STEM. 

a. Sensillae lanceolate urithout a basal stem, 

Schongastia ( Ascoschongastia) lacunosa (Gater 1932). 

Neoschongastia lacmwsa Gater 1932, Parasitology, 24; Womersley and Heaslip 
1943, Tr. Roy. Soc. S. Aust., 67, (1), 110, 

Plate 51, fig. F-I. 

Dorsal scutum as figured, with posterior margin shallow behind line of 
PL and almost rectilinear; sensillae somewhat lanceolate with fine setules, their 
bases wide apart and nearer to line of PL than to line of AL ; AM the longest, 
PL the shortest. Palpal claw bifurcate, setae on femur and genu nude or with 
a few indistinct barbs, on tibia all 3 nude or with short indistinct barbs. 
Chelicerae with only the usual apical tricuspid cap. Galeal setae nude. Dorsal 
setae to 54f», long, short setulations basally, longer distally, 36 in number and 
arranged 2.6.8.8.6.4.2. No long nude setae on tarsi III. 

The Standard Data for 4 paratypes in the British Museum (N.H.) and 
measured by me during a visit to London in 1947, together with the two in the 
South Australian Museum and reported by Womersley and Heaslip 1943 are as 
follows ,* 



198 Records of the S.A, Museum 



AW 


Mean 
64-0rtl-20 


Standard 

Deviation 

2-95±:0-85 


Theoretical 

Range 
55-2-72-8 


Observed 

Range 
59-4-66-5 


Coeff. of 
Variation 

4-6 


PW 


75-2±:0-Sl 


l-98±0-57 


69.3-81-1 


72-6-78-0 


2-6 


SB 


37-3±:0-27 


0-67±0-19 


35-3-39-3 


36-0-38-0 


1*8 


A8B 


30-7±0-43 


l-07±0-31 


27-5-33-9 


28-6-31-5 


3-5 


PSB 


24-0±0»74 


l-64±0-53 


39-l-28'9 


22 -0-26 '4 


6-8 


3D 


54-45±3-07 


2~40±0-76 


47-25-01-65 


50-6-57-2 


4-4 


A-P 


44-0+0-48 


l-17±0-34 


40-5-47-5 


43-0-46-2 


2-6 


AM 


44-95zt0-47 


l-15±0-33 


41-5-48-4 


44-0-46-5 


2-6 


AL 


35-5±:0»38 


0-93+0-27 


32-7-38-3 


35-0-37-4 


2-6 


PL 


26-5±0-48 


l-18zt0-34 


23-0-30-0 


25-0-28-6 


4-4 



Sens. 54-0 mth head 6-7 wide. 

Amongst the material in Lawrence's "Type box" of slides in the London 
School of Tropical Hygiene and Medicine, which I have been permitted to study 
by Lt.-Col. J. R. Andy, are 4 slides Nob. (37-40) labelled " Ascoschongastia 
n. sp. 'C, Lawrence in m.s. M and referred to as such in Appendix 7, to "War 
Office Report, AMD 7, March 7, 1947. 

Lawrence (he. ait.) records these specimens as 3 from the type host, a 
tree shrew Tupaia belangeri belangeri Wagner, and 2 specimens only from 
Hydromys humei. The locality was Palel, Tamu Rd., 30 miles south of Imphal, 
Manipur State, India, Nov, 24th, 1945. 

The four of the above specimens which I have been able to examine do 
not differ in any morphological features from the Malayan specimens except, 
that they are consistently and significantly larger in all details of the Standard 
Data, as given below, suggesting that the Imphal material may be regarded 
as a geographical race of the Malayan species. 







Standard 


Theoretical 


Observed 


Ooeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


68-0±0-80 


l-60±0-56 


63-2-72-8 


67-2-70-4 


2-3 


PW 


86-0±3-09 


6'18±2-18 


67-5-104-5 


80-0-92-8 


7-2 


SB 


38-4 


No variation recorded 






A8B 


35-2±l-31 


2-61±0-92 


27-4-43-0 


32-0-38-4 


7-5 


PSB 


22-4 


No variation recorded 






SD 


57-6±l-31 


2 -61 iirO-92 


49-8-65-4 


54*4-60-8 


4*5 


A-P 


44-8^=1-31 


2-61+0-92 


37*0-52-6 


41-6-48-0 


5-8 


AM 


34-4±0«80 


l*60±0-56 


29-6-39-2 


32*0-35-2 


4-7 


AL 


35-2 


No variation recorded 






PL 


36'0±0<80 


l-60±0-56 


31-2-40-8 


35-2-38-4 


4.4 


Sen3, 


Missing. 











In addition to the differences in Standard Data there is a slight, but pro- 
bably quite unimportant difference in that in the Imphal specimens PL is about 
equal in length to AL instead of being distinctly shorter. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 199 

SCHONCASTIA ( ASCOSCHONG A^TI A ) MALAYEXSIS (Gater 1932 ) . 

Neoschbngastia malayensis Gater 1932, Parasitology, 24; Womersley and Heas- 
lip 1943, Ti\ Roy. Soc. S. Aust., 67, (1), 110. 

Plate 51, fig. J-K. 

This species was originally described from specimens from Rattus malaisia 
Xloss from Sungei Buloh, Selangor, F.M.S. Recently Audy and his colleagues 
have found it at Kepong, Forest Reserve, Selangor, F.M.S. 1950, on R. sctbaniis 
and R, whiteheadi and reared the engorged larvae to nymphs. The Standard 
Data as interpolated from Gater 's figure were given by Womersley and Heaslip 
1943. In 1947 on a visit to the British Museum I was able to examine the 
type and one paratype, and I am now indebted to Dr. Audy for another para- 
type from Gater's collection in the Instit. for Medical Research at Kuala Lum- 
pur, The Standard Data for these three specimens are : 

PSB SD A-P AM AL PL Sen*. 



AW 


PW 


SB 


ASB 


Type 41-6 


66-0 


22-0 


17-6 


Paraty. 41 • 6 


66-0 


22-0 


17-6 


Paraty. 44-8 


61-6 


22-4 


16-8 



22-0 


220 


18-0 


30-0 


19-6 


22-4 


16-8 


33-6 



19-6 36-4 

The scutum is rectangular, wider than long, with AL the shortest and PL 
distinctly off the scutum; SB rather wide apart and somewhat nearer to AL 
than to PL, not the reverse as indicated by Gater 's figure. Sensillae lanceolate 
and finely ciliated to 34/a long (according to Gater). The palpal claw is bifur- 
cate and the setae on the femur, genu and tibia all nude or with one or two 
short indistinct barbs. The cheiicerae are furnished only with the apical tri- 
cuspid cap. Galeal setae nude. There is no long nude seta on tarsi of leg III. 

This species was chosen by Ewing as the type of Ascosclwngastia, but it i» 
unfortunately rather an aberrant species in having PL situated off the scutum. 

b. Sensillae with more or less a basal stem. 

SOHONGASTTA ( " ASCOSCHONGAJSTTA) LTPOXENA Sp. 11. 

Plate 52, fig. F-H. 

Descripti&n of Larvae. Shape broadly oval. Length (engorged) to 1,235ft, 
width to 975/*. Mouth parts largely hidden under the dorsum. Scutum small, 
as figured, with anterior margin sinuous, posterior margin shallow behind line 
of PL; sensillae broadly lanceolate, strongly setulose, without a distinct basal 
stem j AL setae the shortest, PL the longest. Eyes 3 absent. Cheiicerae short 



200 Records of the S.A. Museum 

and stout, as figured, "with only the apical tricuspid cap. Galeal setae nude. 
Palpi stout, tibial claw bifurcate; setae on femur and genu branched; on tibia 
only the ventral nude. Dorsal setae needle-like, apparently nude or with only 
short indistinct barbs j to 44/li long, ca. 26 in number, and arranged 2.6.6.6.4.2. 
Ventrally, with a pair of branched setae on maxillae, a ciliated seta on each 
coxa, a pair between coxae I and between coxae in, and thereafter ca. 
4.4.6.6.6.4.2, similar to but finer than dorsal. Legs: 1 220/i long, II 195/x, 
III 220/x- tarsi I and II with dorsal sensory rod; III without any long nude 
seta. 

The Standard Data for 31 specimens are ; 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
"Variation 


AW 


44-6±0-36 


l-65rt0-21 


39-7-49-5 


42-0-47-6 


3-7 


PW 


60-3±0-44 


2-43:±0-31 


53-0-67-6 


56-0-64^4 


4-0 


SB 


20-5rtO-25 


l-40±0-18 


16-3-24-7 


16-8-22-4 


7-0 


ASB 
PSB 


20-9±0-25 
14-0 


1*392:0 '18 1(5 -7-25 a 
JS T o variation recorded 


19-6-22-4 


6-7 


SD 


34-9+0*25 


l-39Hr0aS 


30 -7-39 -1 


33*&-36-4 


4-0 


A-P 


27-2±6-29 


1*61:±0*20 


22-4-32-0 


25-2-30-8 


5-9 


AM 


35-6±0-43 


2-26±;0-31 


2S-8-42-4 


30-8-39-2 


6-4 


AL 


31-9dt0-87 


l-91rh0-26 


26-2-37-6 


28-0-33-6 


6-0 


PL 

Sena. 


46*63:0-33 
61 • 6 with head 


l-82±0-23 
8 '4 wide. No vai 


41-2-52-0 
rintion recorded. 


42-0-50-4 


3-9 



Log. and Host From bats No. 13944 and R.10566, 71 from Bukit Lagong 
Forest Reserve, Kepong, Kuala Lumpur, Malaya, March, 1950 J. R. Andy). 

Remarks. In the form of the sensillae this species will come close to 
lacunosa Gater and malayensis Gater, differing from the first in the shape and 
size of the scutum, and from the second, in having PL on the scutum and the 
setae of the palpal femur and genu nude. 

As all the specimens were very fully engorged and did not make very 
satisfactory mounts it has not been possible to figure the entire dorsal and 
ventral sides, 

Sohongastia (Ascoschongastia) nadchAtrami sp. n. 
Plate 111, fig. H-J. 

Description of Larvae. Shape (engorged) ovoid with rather parallel 
sides, slightly constricted medially. Length to 938^, width to 670/x. Scutum 
as figured, only slightly wider posteriorly than long; PW greater than AW; 
anterior and posterior margins sinuous; posterior margin very shallow behind 
PL; sensillae missing, bases about in mid-line of scutum; PL very short, AL 
long and much longer than AM. Eyes 2 + 2, posterior the smaller. Chelicerae 
with only the apical tricuspid cap. Galeal setae nude. Palpi outwardly angu- 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



201 



late, tibial claw bifurcate; setae on femur, genu and dorsally and laterally on 
tibia apparently nude, ventral on tibia with short ciliations. Dorsal setae ca. 
42 in number, to 57^. long, and arranged 2.6,6.6,6.6,4.4.2. only very shortly 
ciliated or barbed, or perhaps quite nude. Ventrally with paired ciliated setae 
on maxillae, one on each coxa, a pair between coxae I and between coxae III 
and thereafter ca. 46 setae, lengthening from 33/x, to 50jk posteriorly, and more 
obviously ciliated. Legs : T 402/a. long, 11 348//, III 469ft ; tarsi I and II with 
dorsal sensory rods, III without any long nude seta. 

The Standard Data for the type and 5 paratypes are: 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation. 


AW 


54- 15+0 -56 


l-37±0-39 


50-05-58-25 


53-6-56-95 


2-5 


PW 


70-9±1-03 


2* 52+0* 73 


63-35-78-45 


67-0-73-7 


3*5 


SB 


32-4±0-71 


l-73±0-50 


27-2-37-6 


30*15-33-5 


5-3 


A8B 


35-4±0-43 


l-06±0-31 


39*0-v38-35 


33-5-36-85 


3-0 


PSB 


26-25+0-56 


l-37±0-39 


22-15-30-35 


23-45-26-8 


5-2 


8D 


61-4±1-13 


2-77=t0-80 


53-1-69-7 


56-95-63-65 


4-5 


A-P 


53 -05:1:0 -56 


l-37:£0'39 


48*95-57-15 


50-25-53-6 


2-6 


AM 


55«25±1*14 


2-80±0-81 


46-85-63-65 


50-25-56-95 


5-1 


AL 


73-l±2-91 


7-14±2-06 


51-7-94-5 


67-0-83-75 


9-8 


PL 


13-4 




No variation recorded. 




Sena. 


Missing in all 


specimens. 









Loc. and Eost. Six specimens, 3 from Raftus sab amis, Bukit Lanjar 
Forest Reserve, Selangor, F.M.S., 4 Nov., 1950, 1 from same host, Kepong F.B, 
Selangor, 23 June, 1950, and 2 from Battus rajah, from Ulu Langat FJB. 
Selangor, 21 July, 1950, 

Remarks. This species in the shape of the scutum is close to S. (A.) 
lacunosa Gater, g, (8.) lewthwaiiei sp. n. and 8. (A.) sarawaketms sp. n. It 
differs from lewthwaitei in the quite non-serrate chelicerae; from lacwwsa in 
the very short PL setae, and from sarawakensis in the Standard Data, especi- 
ally the very long AL scutal setae. 

Schongastia (Ascoschongastia) sarawakensis sp. n. 

Plate 111, fig. D-G. 

Description of Larva*. Shape (engorged) broadly oval with slight medial 
constriction. Length (engorged) to 730ft, width to 600/x. Scutum small as 
figured, almost quadrate, with AL longer than AM, and PL very short; sen- 
sillae rather short, clavate and setulose, with short but distinct basal stem, 
head as wide as long in ratio of 10 : 18 ; bases distinctly nearer to PL than to 
AL ; contours as in typical lavmiosa. Eyes 2 + 2, posterior the smaller. Cheli- 
cerae simple with only the apical tricuspid cap. Galeal setae nude. Palpi 



202 



Records of the S.A. Museum 



rather angular on femur; seta on femur nude or with indistinct barbs; on 
genu nude and long; on tibia ventral with short branches, dorsal and lateral 
apparently nude; palpal <»law bifurcate. Dorsal setae short and curved, to 
57/a long, with short sparse barbs; ca. 42 in number and arranged ca. 2.6,6.6.6.- 
6.4.4.2. Ventrally with a pair of shortly ciliated or barbed setae on maxillae, 
one on each coxa, a pair between coxae I and between coxae III and there- 
after ea. 46-48 setae. Legs: 1 321|t long, II 28V, III 375/*; tarsi I and II 
with dorsal sensory rod, III without any long nude seta. 
The Standard Data for the type and 8 paratypes are j 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Kangc 


Range 


Variation 


AW 


50-G±0-67 


2 ■ 01:4:0 -47 


44-6-56-6 


46-9-53-6 


4-0 


PW 


65 -1:1:0 -83 


2 '43:£0 -57 


57-S-72-4 


60-3-67*0 


3-7 


8B 


30-9±0-49 


l-47±0-35 


26-5-35-3 


50-15-33-5 


4-8 


ASB 


30-9±0-49 


l-47±0*35 


26-5-35-3 


30- 15-33- 5 


4-8 


PSB 


23-45 




No variation 


recorded. 




BD 


54-35z±0-49 


l-47±0-35 


49-95-58-75 


53*6-56-95 


2-7 


A-P 


46*9 




No variation 


recorded. 




AM 


50-25 




No variation recorded. 




AL 


56-95 




No variation 


recorded. 




PL 


13-4 




No variation recorded 




Sens. 


23-45 with head 10 -05/18 -75. 


One determination only. 





Loc r and Hosts. The type and 8 paratypes from moon rat EcMnosorex 
gymnurus from Tinjar, Sarawak, 15th June, 1950. 

Remarks. This species is so close to typical la&unosa Gater, that one 
might be disposed to regard it only as a variety. It differs from the typical 
form in having a significantly smaller scutum, although of the same contours, 
and in having the sensillae much shorter and more clavate with a short but 
distinct basal stem* The AL seutal setae are distinctly longer than the AM 
seta, while the dorsal setae are shorter than (compare Standard Data) in typical 
lacunosa from Malaya. 

SCHONGASTIA (ASCOSCHONGASTIA) NAUSHERAENSIS Sp. 11. 

Plate 52, fig. A-E. 

Description of Larvae. Length (engorged) 390ft, unengorged 320^, width 
227ft and 214^. respectively. Shape broadly ovaL Scutum as figured, roughly 
rectangular with posterior margin either almost rectilinear or shallow between 
PL (this is difficult to ascertain as some of the dorsal striations overlap the 
margin and obscure it) ; sensillae clavate, strongly setulose with the bases wide 
apart and only slightly nearer to PL than to AL; surface finely punctate. AM 
and AL subequal, PL longer. Eyes not discernible. Chelicerae non-serrate,. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 203 

with only the apical cap. Galeal setae nude. Palpi stout, tibial claw bifurcate ; 
setae on femur and genu strongly branched; on tibia ventral only branched. 
Dorsal setae to 30^. long, ca. 40 in number and arranged 2.10.8.8.2 lateral, 6.4.2. 
Ventrally a pair of ciliated setae on maxillae, one each on coxae I and II 
and three on coxae III, a pair between coxae I and between III, and thereafter 
ca. 26-28. Legs: I 195/x long, II 169/*, III 20 V, tarsi I and II with dorsal 
sensory rod, III without any long nude seta. 

The Standard Data for the type and eleven paratypes are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


50-5±0-44 


l-52;t0-31 


46-0-55-0 


47-6-53*2 


3*0 


P\V 


67'8:£0'58 


2*01rt0-41 


61*8-73-6 


64'4-70-0 


3-0 


SB 


37-3±0-53 


l-82±0*37 


31-8-42-8 


33-6-39-2 


5-0 


ABB 


22-3±(M2 


0-40±0'08 


21-1-23-5 


21-0-22-4 


1-8 


P8B 


22.3±0-12 


0-40±0-08 


2L1-23-5 


21-0-22-4 


1-8 


SD 


44-7±0*24 


0-82±0-17 


42-2-47-2 


42-0-44-8 


1*8 


A-P 


34-4±0.36 


1*26±0»26 


30-6-38-2 


33*6-36-4 


3-7 


AM 


20.0 


No variation recorded 






AL 


20-0 


No variation recorded 






PL 


30-25:t0-18 


0-62±0-13 


28-4-32 «1 


30-0-32-0 


2-1 


Sens. 


36-4 with head 1 


13/22. No varia 


tion recorded. 







Loo. and Host. The type and eleven paratypes from ears of a mouse, 
Naushera, India, 14 May, 1949 (S.L.K.), 

Remarks. This species is elosely related to lacunosa Gater and malayensis 
Gater, but differs in the dorsal setae, setation of palpi, Standard Data, and 
especially the trisetose coxae III. 

Schongastia (Ascoschongastia) pebtlis (Gater 1932). 

Neoschongastia debilis Gater 1932, Parasitology, 24; Womersley and Ileaslip 
1943, Tr. Roy. Soc. S. Aust., 67, (1), 118, 

Plate 53, fig. A. 

This species is still only known from the original specimen from the ear 
of Battus c.renwHventer cremoriventer (Mullcr) from Sungei Bui oh, Selangor, 
F.M.S. 

Gater 's description is somewhat brief and he only figures the dorsal 
scutum, from which the sensillae were missing. 

The scutum is almost rectangular with the posterior margin rectilinear 
between PL. The AL setae are the shortest and all are distinctly ciliated. The 
sensillae bases are about in the mid-line of the scutum or slightly in front of 
it. The ehelicerae are non-serrate, apparently with only the apical tricuspid 
cap. The palpal claws are stated by Gater to be trifurcate, the setae on the 



204 Records of the S.A, Museum 

palpal femur and genu nude, and on the tibia the dorsal and lateral slightly 
pectinate, the ventral plain. Eyes apparently absent. 

The dorsal setae number 54, and are arranged 2,12.4.8.7.10.2.5-4. The 
Standard Data from Gater's figure and data as computed by Woinersley and 
Heaslip (1943) are: AW 40-0, PW 48-0, SB 17-0, ASB 13-0, PSB 11-0, 
SD 240, A-P 21-0, AM 30-0, AL 20-0, PL 36-0, Sens. — . 

With its characteristic scutum this species should be readily recognized 
when again met with. 

SCHONGASTIA (AsCOSCHONQASTtA) MANIPUBENSIS (Radford 1946). 

Neoschongastia> manipwrenms Radford 1946. Proc. Zool. Soc, London, 116, (2). 

Plate 53, fig. B-D. 

Described by Radford from Rattus ratitm rufescens (Gray) from Imphal, 
Manipur, 13 June 1945, this species will fall into the indica group on the size 
of the dorsal scutum, AL scutal setae being the shortest, and the short rather 
stumpy front tarsi. It is strikingly different from all other members of the 
group in that PW is almost 3 times as long as AW, and in having both coxae II 
and III multisetose, II with 3 setae, III with 6 setae. The scutal setae are said to 
be nude. The dorsal setae are 30 in number, arranged 4.8.6.6.4.2, i.e. there are 
2 scapular setae on each side. Ventrally behind coxae III the setae number 
ca. 36 r No details or figures of the palpi are given by Radford, and I have not 
seen any material. \ 

The Standard Data as given by Radford are: AW 25-0, PW 74-0, SB 21*0, 
ASB 23-0, PSB 20-0, SD 43-0, A-P 32-0, AM 20-0, AL 17-0, PL 27-0, 
Sens. 38*0. The value of 32-0 for A-P, however, is not in accordance with his 
figure of the scutum and is more probably an error for 42*0. 

In the multisetose coxae II and III and the shape of the scutum this species 
might be placed in Doloiria< Ouds. 1910 as suggested by Puller 1948, if that 
genus can be validly separated from Schonga$tia when the adults become known. 
It differs from the genotype of Dotoisia, (synote Ouds. from Germany) in tftiat 
coxae I is only unisetose. 

SCHONGASTIA (AbCOSOHONGASTIA) tiABUANENSIS Sp. n. 

Plates 53, fig, E-J ; 57, fig. B. 

Description of Larvae. Shape oval. Length (partially engorged) 390/x., 
width 300/*. Dorsal scutum as figured, small, with the anterior margin lightly 
sinuous, posterior margin concave, the posterior angles produced ■ AL setae the 



Womersley— Asiatic-Pacific Scrub Typhus Mites 205 

shortest but all shorter than usual; sensillae bases in the mid-line of scutum, 
sensillae missing, but possibly elavate. Eyes not observed, possibly absent. 
Chelicerae simple, broad and stout, with only the apical tricuspid -cap. Galeal 
setae apparently nude. Palpi small, tibial claw short, bifurcate with subeqnal 
prongs; setae on palpal femur and genu with short setules, on tibia, dorsal 
and lateral ciliated, ventral apparently nude ■ tarsi with 4 or 5 ciliated setae, as 
well as subapical and sub-basal sensory rods. Dorsal setae short and setulose, 
to 22/l long, ca. 62 in number, arranged 2.10.10.10.1010.6.4. Ventrally with 
a pair of branched setae on maxillae, a single ciliated seta on each coxa, a pair 
between coxae I and between coxae III and thereafter ca. 40, similar to dorsal, 
and to 20/x long. Legs, relatively short, segments short and stout, all 7-seg- 
mented, I 180^ long, II 150/x, III 180/*; tarsi I and II with the usual dorsal 
sensory rod, III without any long nude seta, or nude setae of any kind. 
The Standard Data for the type and 4 paratypes are : 







Standard Theoretical 


Observed 


Coeff. of 




Mean 


Deviation Range 


Range 


Variation 


AW 


28*8 


No variation recorded 






PW 


41-3+0-32 


0-71+0-23 39-1-43-4 


40-0-41-6 


1-7 


SB 


13.45±0«59 


1-31+0-42 9-5-17-4 


12*8-16-0 


9-5 


ASB 


14-4 


No variation recorded 






PSB 


11-2 


No variation recorded 






3D 


25-6 


No variation recorded 






A-P 


22-4 


No variation recorded 






AM 


16-0 


No variation recorded 






AL 


10*0 


No variation recorded 






PL 


19-2 


No variation recorded 






Sens. 


Missing. 









Loc. and Host. Five specimens from a marsupial rat from Labuan, British 
North Borneo, 17th Sept., 1945 (B. N. McCulloeh). 

EemarJcs. If this species should he found to have elavate sensillae it will, 
on the structure of the scutum, tarsi and palpal claw fall into the indica group 
of species as in the key. 

SCHONGASTIA (ASCOSCHONGASTIA) AUDYI Sp. n. 

Plates 54, fig. A-E ; 57, fig. D. 

Description of Larvae. Shape oval. Length (partially engorged) to 300/*, 
width to 200/*. Scutum roughly rectangular, with the anterior margin sinuous, 
posterior margin, lightly curved behind line of PL : AL setae the shortest ; sen- 



206 Records of the S.A. Museum 

sillae clavate and setulose, their bases about the mid-line of scutum. Eyes 
2 -J- 2 on ocular shields, closely adjacent to posterolateral corners of scutum, 
posterior eyes the smaller, Chelicerae simple, with only the usual apical tri- 
cuspid cap. Galeal setae nude. Palpi stout, claw bifurcate with the prongs 
unequal, to 14^. long 1 ; setae on palpal femur and genu indistinctly barbed, on 
tibia all three nude; tarsi with subapical and sub-basal sensory rod-like setae 
and 4 or 5 ciliated setae. Dorsal setae 30 in number, to 36/i long, with short, 
setulations and arranged 2.6.6.6.6.4.2. Ventrally with a pair of ciliated setae 
on maxillae, a single seta on each coxa, a single pair between coxae I and two 
pairs between coxae III, thereafter ca. 36, to 22/a long. Legs I 208/a long, 
II 184/*, III 218/x ; tarsi I and II with the usual dorsal sensory rod, III with ft 
long nude seta* 

The Standard Data for the type and 6 paratypes are: 



AW 


Mean 
49-5±0-92 


Standard 
Deviation 

2-26±0-65 


Theoretical 
Range 

42-7-56-3 


Observed 
Range 

47-6-53-2 


Coeff . of 
Variation 

4'5 


PW 


67-8±l-41 


3-54±1.02 


56-6-77-8 


61-6-72-8 


1-4 


SB 


22-6±0-5G 


1-38±0-40 


18-5-26-7 


21-0-25-2 


6*1 


ASB 


26-6±0-63 


1-53 ±0-44 


22 -0-31 *2f 


25-2-28-0 


5-7 


PSB 


22-4 


No variation recorded 






SD 


49-0±0-63 


i*m±S>-U 


44-4-53-6 


47-6-50*4 


31 


A-P 


28-9+0-59 


l-44±0-42 


24-6-33-2 


28-0-30-8 


5-0 


AM 


31-7±0-£>9 


l-44:±0-42 


27-4-36-0 


30-8-33-6 


4-5 


AL 


25*2 


No variation recorded 






PL 


39-7±0-47 


l«14±0-33 


36-3-43-1 


39-2-42-0 


2-9 



Sena. 39-2 with head 11/22. No variation recorded. 

Loc. dnd Eost. Described from the type, from Prome, S. Burma, 1945 (no 
host and further date recorded) and from 6 paratypes from Rattus rattus nor- 
vegicus from R.G.N. Pagoda Rd., S. Burma 14-16 Sept, 1945, (coll. T. J. Law- 
rence, type Pll, paratypes 702-1, 706-4, 708-1). Also 7 specimens from 
R. r. norvegicus from Toungoo, S.B., 23 Oct., 1945 (768). 

Remarks. At first glance this species suggests tortus Gunther, but careful 
study shows that it is quite distinct, differing in the number of dorsal and 
ventral setae, the longer and less stumpy palpal claw, and the presence of two 
pairs of setae between coxae III. I am pleased to name it after Dr. J. R. Audy, 
leader of the British Medical Research Council Typhus Team in Burma, and 
to whom I am indebted for the opportunity of studying so much of their 
material. 

Another lot of five slides from the collection of Gater's material left 



Womersley — Asiatic-Pacific Scrub Typhus Mites 207 

in the IJYIR., and forwarded to me by Dr. J. E. Audy are of this species. 
These slides were labelled in grease pencil "seiurii" and one of them "not to be 
published. 7 ' The locality is not known but was most probably Sungei Buloh." 

Schongastia (Ascoschongastia) battus (Worn, and Heaslip, 1943). 

Neoschongastia rattus Womersley and Heaslip 1943, Tr. Boy. Soc. S. Ausi, 

67, (D,118. 

Plates 54, fig. F-I; 55, fig. A; 57, fig. C. 

This species was described from specimens from Rattus assimUis from 
Imbil and Brisbane, Queensland. 

It is a rather characteristic species amongst those of the indica group in 
that the scutum is as long as it is wide between PL. The setae on the femur, 
genu and tibia of the palpi are all nude, and the claws are long, to 16ft and 
bifurcate. The dorsal setae are 2.6,6.4 2, as in lorius Gunther, but the ventral 
setae are much more numerous, ca. 40-50 in number. Tarsi III with 1 long 
nude outstanding seta. 

The Standard Data, as redetermined for the type and 3 paratypes are : 









Standard Theoretical 


Observed 


Ooeff. of 




Mean 


Deviation Range 


Range 


Variation 


AW 


35-9+0* 


80 


l*60±0-57 31-1-40-7 


35-2-38-4 


4-5 


PW 


64-8±0 


80 


l-60±0-57 60-0-69-6 


64-0-67- 2 


2-5 


SB 


22-0±0' 


20 


0-80±0-28 19 -6-24 -4 


20 -8-22. 4 


3*6 


ASB 


22-4 




No variation recorded 






P8B 


35-2 




No variation recorded 






SD 


57-6 




No variation recorded 






A-P 


41-6 




No variation recorded 






AM 


25-6 




No variation recorded 






AL 


12-8 




No variation recorded 






PL 


38-4 




No variation recorded 






Sens. 


35-2 




Only 1 determination. 







Schongastia (Ascoschongastia) lorius (Gunther 1939). 

Neoschongastia lorius Gunther 1939, Proc. Linn. Soc. New South Wales, 64, 
(1-2), 86; Womersley and Heaslip 1943, Tr. Roy. Soe. S. Aust, 67, (1), 
118. 

Plates 55, fig. B-E ; 57, fig. E. 

This species was described from a parrot (Lorius r&ratus) from Bul'olo, 
N. Guinea. Womersley and Heaslip (1943), recorded the Standard Data from 
their re-examination of the type and two paratypes as follows: AW 40-0, 



208 Records of the S,A. Museum 

PW 55-0-58-0 (aver. 56-5), SB 20-0, ASB 16-0, PSB 21-5-23-0 (aver. 
22-25), SD 37-5-39-0 (aver. 38-25), A-P 21-5, AM 30-0, AL 17-0, PL 31-5- 
33-0 (aver. 32-0), Sens. 35-0 with head 10-0 wide. Gunther gave: scutal 
length 47-0, width 62-5, SB 20-0, AWL 36-0, AL 25*0, PL 37-5, Sens. 30-0. 

The dorsal setae are : 2.6.6.4.2.2, as given by Gunther. and not as quoted 
in error by Womersley and Heaslip (1943). 

In addition it should be noted that the palpal claws, which are bifurcate, 
are short and stumpy, measuring only 7fi long, as compared with those of 
G/udyi n. sp. and rattiis Worn, and Heaslip. A long nude seta on tarsi III. 

Schonqastia (Ascosciiongastia) indica (Hirst 1915). 

SMngasiia indica Hirst 1915, Bull. Ent. Res., 6, 183; Walch 1927, Geneesk. 
Tijds. v. Ned. Indie, 67, (6), 924. 

Tromhicida muris Walch 1922, Kitasato Archiv. Exper. Med., 5, (3) ; 1928, 
Fletcher, Lesslar and Lewthwaite, Tr. Roy. Soc. Trop. Med. and Hyg., 
22, 161. 

Neoschdnga&tia indica Gater 1932, Parasitology, 24, 156 ; Womersley and Heas- 
lip 1943, 67, (1), 118; Radford 1942, Parasitology, 34, (1), 76; idem 1948, 
Parasitology, 37, (1), 48; Wharton, 1946, Ecol. Mbnogr,, 16, 151-184. 

Neoschonga>$tm cocJcingsi Radford 1946 Proc. Zool. Soc. London, 110, (2), 262. 

Plates 55, fig. F-I; 56, fig. A; 57, fig. G. 

Graph 1 and 2. 

Dorsal scutum of small size, as figured, with AL the shortest; sensillae 
clavate with distinct setules and their bases in the mid-line of scutum and 
slightly nearer to PL than to AL ; posterior margin rather shallow behind PL, 
but evenly curved or lightly concave medially. Chelicerae simple, with only 
the apical tricuspid cap. Galeal setae nude. Eyes 2 + 2, on distinct ocular 
shield, posterior eyes the smaller. Palpal claw short and stumpy, bifurcate, 
the prongs equal ; setae on femur with a few short branches, on genu the same 
but often appearing nude, on tibia only the ventral branched. Dorsal setae 
rather slender, with short setules, 34 in number, arranged 4(2), 6(8), 6.6.6.4.2, 
to 25/x long; in the second row the seta on each side of the median pair is 
situated in front of the others and in line with the scapular setae, thus the 
first two rows may be read as 4.6, instead of 2.8. Tarsi III with one long nude 
setae. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 209 

The Standard Data derived from 41 specimens from Burma (T. *L Law- 
rence coll.) are as follows; 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Bange 


Range 


Variation 


AW 


38-4:±0-27 


1-71^0-19 


33-3-43-5 


38-6-42-0 


4-4 


PW 


54-9±0*30 


1-91±0-21 


49-2-60-6 


50-4-58-8 


3-5 


SB 


20 -5:1:0 -20 


1- 26:£0-14 


16-7-24-3 


18-3-22-4 


6-1 


ASB 


22-8:£0*14 


o^93ito.ao 


20-0-25-6 


21.0-25-2 


4-0 


PSB 


19'5rt0-00 


0-36=t0 ? 04 


18-4-20-6 


18-2-19«6 


1-8 


SD 


42-3ztO-15 


0-99±0-ll 


39-3-45-3 


40-6-4.4-8 


2-3 


A^P 


26-7±0-28 


1-76±0*19 


21-5-32-0 


22-^29-4 


6-6 


AM 


24-6±0-29 


l-73±0-20 


19-4-29-8 


22-4-28-0 


7-0 


AL 


J8-8±0-19 


l-22±0-14 


15-1-22-5 


16-8-19-6 


6-5 


PL 


31-l±0-22 


1-41±Q-15 


26-9-35-3 


28-0-33-6 


4-5 


Sens. 


31 -1^:0 -29 


1-61±0-21 


26-3-35-9 


28-0-36-4 


5-1 



Bemcrks. Hirst in his original figure shows quite -clearly the anterior 
position of the second seta from the middle of the second dorsal row. He also 
states: "Some of the hairs on the third leg are plain". The chaetotaxy of leg 
III however, has not been mentioned in detail by other workers, until Wharton 
(1946) described and figured it in detail, showing on the metatarsi one simple 
striated seta, and on the tarsi two long nude whip-like setae. The more distal 
of the latter two setae shown in his figure 2, is actually indistinctly branched, 
frequently appearing to be nude. There is, therefore, only a single long nude 
seta on tarsi III sub-basal in position, 

Schongcbstia (Ascoschongastia) indica, and SL (A.) soekaboemiensis Take- 
kawa are two members of a small group of closely allied species, including 
lorius Gunther, rattm Womersley and Heaslip, and audyi sp. n. in. which the 
dorsal scutum is of very similar shape, but differing in overall size, and in 
which the dorsal setae also vary in number. 

Of these, mdica and soekaboemiensis> while the scuta are different in Stan- 
dard Data, at first sight appear to be only variants of one another in that the 
extreme lower number of DS in the first few dorsal rows of sdekaboemiensis 
approaches that found in iiidica. 

As pointed out in the key and the descriptions, the DS in soekaboemiensn-s 
number 40 to 42, and are arranged 2.10(8 or 9). 8.6.6.6.4, with considerable 
variation, whereas in indica they number 34, arranged 4 (2). 6 (8). 6.6.6.4.2, with 
little variation, but the second row always has the seta on each side of the 
median pair inserted well in front of the others so that the first row may be 
read as 4, i.e. two scapular setae and two submedian. 

The differences of these two species which in the field have been pro- 
visionally identified as indica are clearly shown in the accompanying graphs. 



210 



Records of the S.A. Museum 



In Graph 1 the whole of the Standard Data are shown as follows : Mean, 
Bange of Means expressed by Mean ±3<r, Theoretical Ranges expressed by- 
Mean zt 3a- and by Mean ± 2a, and the Observed Ranges. In all the larger and 
more important characters, AW, PW, etc., it is clear that the two species are 
very significantly distinct. 



70 



€0 



50 



40 



30 



20 



10 



PW 



AW 



1 



i 



SD 



• -£ 



PL Sen* 



A~P 



AM 



SB 



ASB 






• x -£ 



4 



PSB 



X 



~E 



AL 



Graph I. Showing differences of Standard Data of population of (left) Schongastia 
(Ascoschongastm) indica (Hirst) and (right) S. (A.) soeToaboemiensis Takekawa. 

(Measurements in microns. Horizontal lines indicate Mean, Mean it 3<r M , Eange at 
M ± 3 times, and ± twice, Standard Deviation. Crosses indicate observed range.) 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



211 



In Graph 2, the distribution of variations in the two species is shown by 
the regression lines for AW/PW and PW/AW. The distribution is seen to be 
elliptical, the theoretical ranges being indicated by the ellipses drawn using 
the value for Range/n. as given in Tippet's table (Biometrika 1925), 

The circles and triangles indicate the number of specimens with value of 
AW. PW at the centre dots. 



70 



eo 



50 



PW(y> 

IN U. 




Awaa 

IN fJ. 



30 



40 



50 



Graph II, Showing [Regression Lines of AW.PW for populations of (left) Schongasttia 
(A&coschongastia) indica (Hirst) and (right) S. (A.) soehaboemiensis Takekawa, 

The circles and triangles show the frequency of specimens with AW.PW at the centre 
points. The ellipses are drawn at the range/<r using Tippet's Table (Biometrika, 17, 386, 
1925.). 



212 Records of the S.A. Museum 

Scho^gastia (Ascosohongastia) soekaboemiensis (Takekawa 1945). 

Trombicula soekaboemiensis Takekawa S. 1945, Bull, from the Nampogun 
Bookikyusi Bu. No. 126; in Report of Kiyosi Hayakawa, 1946, Singapore. 

Neoschongastia soekaboemiensis, Hayakawa 1946 in Report "A Comparative 
Study of Japanese and Tropical (Scrub Typhus) Tsutsuganiushi Disease" 
(figures only, Chart 13). 

Plates 56, fig. B-F; 57, fig. F, 

Graph 1 and 2. 

Re-description of Larvae. Shape oval. Length (engorged) to 350/z, width 
to 260/a. Scutum trapezoidal, small, anterior and posterior margins sinuous, 
AL setae the smallest and placed distinctly back from anterior angles of 
scutum ; sensillae clavate, their bases about in mid-line of scutum. Eyes 2 + 2, 
on ocular shields, posterior the smaller. Chelicerae simple, with only the apical 
tricuspid cap and a subapical rounded shoulder on dorsal (inner) margin, 
Galeal setae nude. Palpi stout; tibial claw short, bifurcate, prongs subequal, 
to 9/i long; seta on palpal femur, genu and tibia with short indistinct branches. 
Dorsal setae with distinct setules, fairly stout, to 32/* long, variable in number, 
40 to 42, and arranged 2.10(8 or 9). 8.6.6.6.4. Ventrally, a pair of ciliated 
setae on maxillae, one on each coxa, a pair between coxae I and between 
-coxae III and thereafter ca. 8.6.10.6.2.2, the anterior setae shorter and finer 
than the dorsal, the posterior setae much like the dorsal getae. Legs: I 185/i 
long, II 160/*, III 176/i, tarsi I and II with the usual dorsal sensory rod ; III 
-with one long nude outstanding seta. 

The Standard Data derived from 109 specimens are as follows : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


47*15±0-16 


l-64±:0-ll 


42-25-52-05 


44-8-50-4 


3-5 


PW 


64-l:fc0-27 


2-84;±0-19 


55-6-72-6 


58-8-70-0 


4-4 


SB 


24-4±0-14 


1-49:±0<10 


2-0-0-28 • a 


22-4-28-0 


6-1 


A8B 


24-9+0-09 


0-97±0'07 


22-0-27-8 


22-4^28-0 


3-9 


PSB 


19-6 


No variation recorded 






SD 


44-1 ±0*10 


1-01 ±0-07 


41-1-47-4 


42-0-47-6 


2-3 


A-P 


25-5=t0-10 


1-02±0-07 


22-5-28-5 


23-8-29-4 


4-0 


AM 


27-2=£0-20 


l-92±0-14 


21-4-33-0 


25-2-36-4 


7-1 


AL 


20-1±0-11 


1-09±0-07 


16-9-23-3 


16-8-22-4 


5-4 


PL 


33-8:±0-16 


l-64=£0-ll 


28-9-38-7 


30-8-42-0 


4-9 


Sens. 


34-7±:0'22 


1-75+0-15 


29 -5-39- J) 


30*8-39-2 


5-0 



Loc. <md Hosts. This species is very common in Southern Burma, occur- 
ring on Nesokia bengalemis and Battus rattus norvegicus. Some hundreds of 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 213 

specimens were collected by T, J. Lawrence of the British Medical Research 
Council Typhus Team in 1945. 

Remarks. The species is closely related to indka in the general shape of 
the dorsal scutum and has been confused in the field with that species, tinder 
the name of cocfcingsi Eadford (=indica Hirst). Prom this species, however, 
it differs in the lengths of AW and PW (see Standard Data) and in the number 
of dorsal seta. Normally the first three rows of dorsal setae are 2.10.8, as 
compared with 2.8.6 in vn&ica; the second row however, varies somewhat, fre- 
quently having 9 setae and occasionally only 8, but these are always in more or 
less of a straight line. In indica, however, the dorsal setae are more delicate 
and the second sete on each side of the median pah- in the second row is so far 
in front of the others as to he in line with the scapular setae. 

A single specimen from Bwdieota* bmgalemis, Paungde, S. Burma, Sept. 
23rd, 1945 (T. J. Lawrence) had 2 AM setae very close together , and had in 
the field erroneously been thought to be a species of Rannemama, 

The Bulletin in which this species was described was from the Japanese 
Army Institute of Preventive Medicine, Singapore during the Japanese occupa- 
tion .of Malaya. It was published in Japanese, but with well recognisable 
figures, the name in Latin and certain measurements in arable figures. I am 
indebted to Lt. Col. J, ft. Audy for a copy of the Bulletin and a tentative trans- 
lation of certain details. 

The specimens described were collected, mainly from the ears of Rattus r. 
diardi (Jentinck) and R. r. roquei Sodi (R, r. jalorensu (Bonhote)) in a 
bathing area of a camp at Soekaboemi, Java. 

At least 200 specimens are referred to, but the location of the type and 
paratypes is unknown. In the Summary it is suggested that this species may 
be a vector of "tropical" tsutsugamushi disease. 

SCHONGASTIA (ASCOSCIHONGASTIA) PERAMELES (Wom. 1939). 

Neoschongastia perameles Womersley 1939, Tr. Boy. Soc. S. Aust, 63, (2), 
160; Womersley and Eteaslip 1943, ibid,, 67, (1), 122. 

Plates 56, fig. G-I ; 57, fig. A. 

This species was described from co-type material from bandicoots. Isoodon 
toroms, from Kiamba, Queensland, in June, 1938 (coll. D. J. W. Smith) to- 
gether with specimens from the same host from Nambour, Q., May and Sept. 
1938, by the same collector. Of this material in the S.A. Museum, that from 
Kiamba has now been divided into a type and 10 paratypes. Additional speci- 
mens from the same species of bandicoot were recorded in 1943, from Cairns, 



214 Records of the S.A. Museum 

■while in the S.A, M'us. collection, not previously recorded are specimens from 
Yandina, Q, from the same hosts (colL D.J.W.S , 5 May, 1938). No more re- 
cently collected material has been seen. The dorsal scutum is rather character- 
istic with PL nearer to AL than to the posterior margin, and not at the pos- 
terolateral corners which are well rounded; the scutum is widest posterior of 
PL; AL the shortest, PL the longest. The chelicerae are simple with only 
the apical tricuspid cap. The galeal setae are nude. The palpal claw is trifur- 
cate and all setae on femur, genu and tibia are branched ; on femur with 5-6 
branches; on genu 2-3; on tibia, ventral with 3 or 4, lateral 2-3 and dorsal 1 
or 2 barbs. The dorsal setae number ca. 76, to 32/a long and arranged 2.12.12.12 
(14),10.10,8.6.2. Ventrally the setae are to 26^ long, with 10-12 lying between 
coxae IIL 

The Standard Data derived from 45 specimens, including the type and 
10 paratypes are j 





Mean 


Standard 
Deviation 


Theoretical 
Eange 


Observed 
Range 


Cocff. of 
Variation 


AW 


63-5+0-46 


3 -12+0-33 


54 -1-72 '9 


58-8-70-0 


4-9 


PW 


78-7±0-44 


2-96+0-31 


69-8-87-6 


72-6-84*0 


3*7 


SB 


25-l±0-23 


1-58+0 -17 


20-4-29*8 


22-4-28-0 


6-3 


ASB 


28-9±:0-20 


l-32±0-14 


24-9~32'9 


28-0-30-8 


4-5 


PSB 


25-8±0-16 


1-05+0-11 


22-7-28-9 


25-2-28-0 


4-0 


SD 


54-7±0-25 


1-68+0-18 


49-7-59-7 


53-2-58-8 


3-0 


A-P 
AM 


21-9=t0-23 
48-2+0-24 


1-56+0-16 
1-32+0-17 


37-2-26-6 
44-2-52-1 


19-6-25-2 
44-8-50-4 


7-0 
2-7 


AL 


38-6+0-43 


2-83±0-30 


30-1-47-1 


33-G-42'0 


7-3 


PL 

Sens. 


74-3+0-48 
42-0 with head 


3-05±0-34 
14/28. Novariati 


65-1-83-5 
ion recorded. 


70- 0-81-2 


4-1 



Remarks. The figure of the dorsal scutum given in the original publication 
is not quite correct in the shape posterior of PL. The scutum is as figured in 
1943 (loe. eit.) and herewith. 

Scuongastia (Ascoschongastta) similis (Worn, and Heasp. 1943). 

Neoschongastia mniliii Womerstey and Heaslip 1943, Tr. Roy. Soc. S> Aust, 
67, (1), 124. 

Plate 58, fig. A-E. 

This species was originally described from specimens taken from rats at 
Cairns, Queensland in 1939 (W. G. Heaslip). In the shape of the scutum 
with A-P very short and SB well behind line of PL it is closely related to 
8. (A,) derricU Worn, and Heasp. 1943 and S. (A.) dasycerci (Hirst 1929), but 
agreeing with the first only in having crenulate dorsal striations. From der- 



Womersley— Asiatic-Pacific Scrub Typhus Mites 215 

ricki, it can be distinguished by the fewer dorsal setae which in the second row 
are ea. 98/a long, in the third row 70/*, and in the posterior rows as low as 
5(V The palpal claw is trifurcate, not bifurcate, and the setae on the palpal 
femur and genu have much fewer but longer branches than in derrichi. In 
dasycerci the dorsal setae are more numerous and shorter than in either der- 
richi or similis. 

The Standard Data as redetermined for the type and 6 paratopes are : 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Bange 


Variation 


AW 


65-5±0*83 


2-20±0.59 


58-9-72*2 


64-4-70-0 


3*3 


PW 


87-2:±0*73 


l-93;t0-o2 


81 • 4-93 '0 


84-0-89-6 


2-2 


SB 


34-4±0-52 


l-37:i:0-36 


30 -3-38 -5 


33*6-36-4 


4-0 


ASB 


22-4 


No variation 


recorded 






PSB 


28-4±0-40 


1-Q6+0-28 


25-2-31*6 


28-0-30-8 


3-7 


8D 


50-8±0-40 


1-062:0 -28 


47-6-54-0 


50-4-53-2 


2-1 


A-P 


16-8 


No variation 


recorded 






AM 


53-0 


Only 1 determination 






AL 


19-6 


No variation recorded 






PL 


99-2±l-48 


3-91±l-04 


87-5-110-0 


92-4-103-6 


3-9 



Sens. 50*0 with head 11-0 wide. Original determination. 

Schongastia (Ascoschongastia) derricki (Wohi. 1939). 

Neoschdngmtia dtrricki Womersley 1939, Tr. Roy. Soc. S. Aust., 63, (2), 162; 
Womersley and Heaslip 1943, ibid, 67, (1), 121 

Plates 58, fig. P-T; 59, fig. A. 

This species was described from specimens from Rattus lutreolus and 
R. assimitis, from Imbil, Queensland in August, 1938 (D. J. W. Smith). In 
the shape and dimensions of the scutum with sensillae bases posterior of line of 
PL and the posterior scuta! margin an even curve, it is closely related to 
8. (A) daxsycerci (Hirst 1929) and 8. (A,) similis (Worn, and Ileasp. 1943). 
Together with similis it differs from dasycerci in that the dorsal cuticula stria- 
tions are markedly crenulate. From similis it can be distinguished, as given in 
the key, in the greater number of dorsal setae, the posterior of which are not 
very much shorter than the anterior. The palpal claw is bifurcate; the 
seta on palpal femur thickly but shortly ciliated, on genu rather less so, and on 
tibia the dorsal and lateral are nude and the ventral branched. The dorsal 
setae are 2.6.6.6.4.2, to 100/x anteriorly and 70> posteriorly. The ventral setae 
to 30/x long. There is no long nude seta on tarsi III. The sensillae are wider 
than in simiUs. 

The Standard Data as redetermined for the type and 9 paratopes are: 



216 Records of the S.A. Museum 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Obsorved 
Range 


Coeff . of 
Variation 


AW 


68-0±0-43 


l-35±0-30 


64-0-72-0 


67-2-70*0 


2-0 


pvt 


88-75±0-70 


2-23±0-50 


82-05-95-45 


84-0-92-4 


2-5 


SB 


34-7±0-46 


l-45±0-32 


30 -4-39-0 


33-6-36-4 


4-2 


ASB 


25-2 


No variation recorded 


. 




PSB 


25-2 


No variation 


recorded 






SD 


50-4 


No variation 


recorded 






A-P 


14*0 


No variation recorded 






AM 


52-85±0-70 


4-35±l*09 


39-8-65-9 


44-8-56-0 


8-2 


AL 


25-75±0-70 


2-21±0-49 


19-15-32*35 


22-4-28-0 


8-5 


PL 


86-25 + 1-00 


3-18±0«71 


76-7-95-8 


84-0-92-4 


3-7 



Sens. 42-0 with head 16-8/30-8. No variation recorded. 

Schongastia (Ascoschongastia) dasycerci (Hirst 1929). 

Sehdiigastia dasycerci Hirst 1929, Proc. ZooL Soc, London, (2). 

Neosc}wn>gastia, dasycerci, Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust, 
67, (1), 122. 

Plate 59, fig. B-E. 

This species was described from the ears of Dasycercus cristicauda, from 
Ooldea, South Australia. 

The dorsal scutum is as figured with A-P short, and SB behind line of 
PL. The sensillae are clavate with prominent setules especially at the base of 
the head. AL slightly the shortest, being a little less than AM. The chelicerae 
is non-serrate with only the usual apical cap. The palpal claw is trifurcate 7 - 
the setae on the palpal femur, genu and tibia are all branched or ciliated. The 
dorsal setae number 34, to 5Q/* long and are arranged 2.8,6,6.6.4,2. There are 
no long nude setae on tarsi III. 

The Standard Data as redetermined for 7 of Hirst's co-types in the South 
Australian Museum collection are i 







Standard 


Theoretical 


Observed 


Coeff- of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


55-6:£l-13 


2-99:fc0-80 


46-6-64-6 


50*4r-58-8 


5-4 


PW 


65-2±:l*33 


3-51:£0-94 


54.7-75-7 


61 -6-70-0 


5-3 


SB 


23-8±0-52 


l-40±0-37 


19-6-28-0 


22-4-25-2 


6-0 


ASB 


22-4 


No variation recorded 






PSB 


20-4±0-52 


1-37+0-36 


16-3-24. 5 


19-6-22-4 


6-7 


SD 


42-82:0-52 


l-37±0-36 


38-7-46-9 


42-0-44-8 


3-2 


A-P 


14-0 


No variation recorded 






AM 


35-6±;0-80 


2-12+0-56 


29-3-41-9 


33-6-39-2 


5-9 


AL 


21-3:h0-48 


1-07+0-34 


18-1-24-5 


19-6-22-4 


5-0 


PL 


48-0±0-40 


1-06+0-28 


44-8-51-2 


47-6-50-4 


2-2 


Sens. 


43*9±0-93 


1-62+0-66 


39-1-48-7 


42*0-44-8 


3-7 



Womersley— Asiatic-Pacific Scrub Typhus Mites 217 

A single specimen was taken along* with 2 specimens of S. (A.) coorongense 
(Hirst) from Ratius murrayi var. on Greenly Is., South Australia, Dec, 1947 
(P. J. Mitchell). 



Sohongastia (Ascoschongastia) trichosuri (Worn. 1939). 

Ne&schdngastia westralierms var. trwhosuri Womersley 1939, Tr. Roy. Soc. 
S. Aust., 63, (2), 160. 

Neoschortgastia shield^ Gunther 1941, Proc. Linn. Soe. New South Wales, 56, 
(3-4), 157; Womersley and Heaslip 1943, Tr. Eoy. Soc S. Aust,, 67, (1), 
123. 

Neoschongastia trichosuri, Womersley and Heaslip 1943, Ti\ Roy. Soe. S. Aust., 
67, (1), 122. 

Plate 59, fig. F-J. 

This species was described from a single specimen, from Nambour, Queens« 
land. In 1943, Womersley and Heaslip raised it to specific rank and gave a 
much better figure of the dorsal scutum, the figure published in 1939 being in- 
correct in the shape of the posterior scutal margin. The unique type has been 
remounted and the scutum is now seen to be much as figured in 1943 and to 
agree with that figured by Gunther 1941 for his species, shieldst, described, from 
Melomys rubex Thomas, from Bulolo, New Guinea. 

The Standard Data for trichosuri as remeftsured are now found to be : AW 
66-0, PW 95-2, SB 28-0, ASB 30-8, PSB 19*6, SD 50-4, A-P 28-0, AM 43-0, 
AL 35-0, PL 56-0. Sens. 40 «0 with head 13-0. Those for shieldsi as deduced 
afresh from Gunther 's data and figures (the types, said to be in the Australian 
Museum, Sydney have not been seen) are: AW 62 -0 t PW 75-0, SB 24-0, 
ASB 23-5, PSB 15-5, SD 39*0, A-P 15-0, AM 37*5, AL 45-0, PL 56-0, Sens. — . 

The dorsal setae in trichosuri number 34, and are arranged 2.8.6.6.6.4.2, 
and measured to 45/x, long, as shown by Gunther for shieldsi. 

In both trichosuri and shield&i the palpal claw is bifurcate, the setae on 
the palpal femur and genu branched (in Gunther's figure of shieldsi, but in 
his text he states "on &k (genu) two (branched setae) at the base and one 
towards the apex. • ' This is evidently a mistake, for in all known Trombiculidae 
there is only a simple seta on this palpal segment* On the palpal tibia the 
dorsal ventral and lateral setae are nude (Gunther only saw the ventral) 
(? branched) and one nude seta). The chelicerae are simple with only the 
apical cap, and the galeal setae are branched. 



218 Records of the S.A, Museum 

Schongastia (AscoschOngastia) hirsti (Worn, and Heasp, 1943). 

Neoschongastia hirsti Womersley and Heaslip 1943, Tr. Boy. Soc. S. Aust., 67, 
(1),123. 

Plate 60, %. A-E. 

This species was described from the type and 3 paratypes from Melomys 
cervenipes, from Imbil, Queensland, (coll. D. J. W. Smith 1938), and two 
specimens from rats, from Cairns, Queensland, 1938 (W. G, Heaslip). In 
addition I now have another specimen from rat from Wandecla, Q., 24 Jan,, 
1945 (R. N. McCulloch). 

This species, as given in the key, is closely related 8. (A.) trichosuri 
Worn, and Heasp. (= shieldsi (Gunther)) in having A-P short, SB behind the 
line of PL, and the posterior scutal margin being roughly three-sided. From 
trichosuri it differs in the rather larger scutum and in having AL setae much 
the shortest. The dorsal setae are of the same number and arrangement but 
measure to 70/a long whereas in trichosuri and according to Gunther 's figures 
of shield&i they do not exceed approximately 45/1. The palpal claw is tri- 
furcate, and all the setae on femur, genu and tibia strongly ciliated, that 
on the femur being also very long. The chelicerae have only the apical 
tricuspid cap and the galeal setae are strongly ciliated. The sensiliae are miss- 
ing from all specimens. 

The Standard Data for the above 7 specimens as remeasured are: 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


68-4±l-03 


2-73:±0-73 


60-2-76-6 


64-4-72-8 


4-0 


PW 


82-0±0-80 


2-12±0-57 


75-7-88-3 


78-4-84-0 


2-6 


SB 


31-6d=0-80 


2-12;±:0-57 


25-3-37-9 


28-0-33-6 


6-7 


ASB 


24-8:fc0-40 


1-06:4:0 -28 


21-6-28-0 


22-4-25-2 


4-3 


PSB 


28-4 


No variation recorded 






BD 


47-2:±0-40 


l-06±:0-28 


44-0-50-4 


44-8-47-6 


2-3 


A-P 


18-2:±0-48 


l-28±0'84 


14-4-22-0 


16-8-19-2 


7-0 


AM 


45-2±0-73 


l-93db0 -52 


39-4-51-0 


42-0-47-6 


4-3 


AL 


25-2±:0.61 


1-63+0-43 


20-4-30-0 


22-4r-28.0 


6-4 


PL 


70-4±0*40 


l-06±0.28 


67-2-73-6 


70-0-72-8 


1-5 


Sens. 


Missing. 











From the shape and dimensions of the scutum, with SB behind line of PL 
and PSB less than A-P, the dorsal setae and the palpal setation, there seems 
little doubt but that shieldsi and trichosuri are synonymous. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 219 

Schongastia (Ascogchongastia) phascogale (Worn, and Heasp. 1943). 

Neoschongastia phascogale Womersley and Heaslip 1943, Tr» Roy. Soc. S. Aust., 
67, (1), 127. 

Plate 60, %. F-J. 

This species was described from the type and 3 paratypes from a wallaby, 
Phascogale sp., from Brisbane, Queensland, Jan., 1938 (D. J. W. Smith), and 
fifteen other specimens from bandicoots, Isoodon torosus, from Cairns 
(W. G. Heaslip 1940), and Perwneles nasuta, Palm Woods, Queensland, Dec, 
1938 (DJ/W.S,). 

As given in the key, this species is closely related to westraliense Worn., 
trichosuri Worn, and peregrina sp. n. in having the sensillae bases almost in 
line with PL. It agrees with the first in having the palpal claw trifurcate and 
the galeal setae branched but differs in A-P being much greater than PSB and 
in the arrangement of the dorsal setae. 

The Standard Data for the nineteen specimens given by Womersley and 
Heaslip (1943) treated statistically are: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


71-6±0-37 


l-63±0-26 


66 '7-76 -5 


68-0-75-0 


2-3 


PW 


83-4±0-42 


l-81±0-29 


78-0-88-4 


79-0-86*0 


2-2 


SB 


30-0:t0-23 


1-00^:0-16 


27-0-33-0 


28*0-32*0 


3*3 


ASB 


30-0±0*27 


1-11+0O9 


26-7-33-3 


27-0-32*0 


3-7 


PSB 


23*6±0-16 


0-74±0-12 


21-4-25*8 


22-0-25-0 


3-1 


SD 


53-3±0-40 


l*65±0-28 


48-6-58-4 


49-0-56*0 


3*1 


A-P 


30-7±0-39 


l-69±0-27 


25-7-35-7 


27-0-35-0 


5-5 


AM 


53-65±0-33 


l-37±0-23 


49*55-57-75 


50-0-55-0 


2*5 


AL 


38-4±0-28 


1-14±0*19 


35*0-41-8 


36-0-49-0 


3-0 


PL 


63-l±0-70 


3-02+0.49 


54-0-72-2 


56-0-67-0 


4-8 


Sens. 


42-6±0'68 


2-73+0-48 


34- 4-50 -8 


36-0-48-0 


6-4 



with head 12*0 "wide. 

Schongastia (Ascoschongastia) westrauensis (Worn. 1934). 

Schongastia westraliense Womersley 1934, Rec. S. Aust Mus., 5, (2), 215. 

Neoschongastia westraliensis Womersley and Heaslip 1943, Tr. lioy. Soc. S* 
Aust, 67, (1),122. 

Plate 61, %. A-E. 

This species is still only known from the original material, one slide con- 
taining about 17 specimens taken from the ears of a cat at Greenbushes, West- 
ern Australia, 11 Aug,, 1931 (coll. H.W.). 



220 Records of the S.A. Museum 

The dorsal scutum is as figured with PL the longest. The posterior margin 
of the scutum, is roughly 3-sided with the median part lightly concave. The 
sensillae are clavate with distinct setules, and their bases practically in line with 
PL, The chelicerae are simple with only the apical tricuspid cap. The galeal 
setae are distinctly branched or ciliated. Eyes 2 -J- 2, posterior the smaller. 
The palpi are stout, with the tibial claw trifurcate. The setae on the palpal 
femur and genu are strongly branched; on the tibia the dorsal and lateral 
nude., ventral branched. The dorsal setae number 34, and are arranged 
2,8.6.6.6(4) ,4(6) ,2, to 56/* long. 

The Standard Data for 15 of the specimens as remeasured are: 





Mean. 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Bange 


Coeff . of 
Variation 


AW 


83-9±:0-63 


2-46±0-45 


75-5-90-3 


75-6-84-0 


3-0 


PW 


99-9±0-63 


2-43:£0'44 


92-6-107-2 


95-2-103-6 


2-4 


SB 


36-l±0-43 


l-68±0-31 


31-1-41-1 


33-6-39-2 


4-6 


ASB 


30-4±0«25 


0'95±0-17 


27-6-33-2 


28-0-30-8 


3-1 


PSB 


24-8±0-25 


0-95±0-17 


22-0-27-6 


22-4-25-2 


3-8 


SD 


55-2:t0-40 


l-54±0-28 


50-6-69-8 


50-4-56-0 


2-8 


A-P 

AM 


26-5±0-45 
53-4±:0.84 


1-73:8:0 -32 
2-90:±0-59 


21-3-31-7 
44-7-62-1 


25-2-30-8 
47-6-56-0 


6-5 
5-4 


AL 


48-l±:0'54 


2-09±0-38 


41-8-54-4 


44-8-50-4 


4-3 


PL 


79-5±0-70 


2-68±0*49 


71-5-87-5 


75-6-84-0 


3-3 



Sens. 42-0 with head 14-0 wide. No variation recorded. 

Schongastia (Ascoschongastia) pbreorina sp. n. 

Plate 61, fig. F-J. 

Description of Larvae, Shape oval. Length (unengorged) to 450/*., width 
to 375/x. Scutum large and roughly trapezoidal, with the posterior margin fairly 
deep behind line of PL and slightly sinuous or concave medially; sensillae bases, 
only slightly behind PL and A-P longer than PSB; AL the shortest, PL the- 
longest; sensillae clavate with strong setules. Chelicerae simple with only the 
usual apical tricuspid cap. Galeal Betae nude. Eyes 2+2, the posterior the 
smaller. Palpi stout, tibial claw bifurcate; seta on both femur and genu strongly 
branched ■ on tibia dorsal and lateral nude, ventral strongly branched. Dorsal 
setae 34, arranged 2.8.6.6.6.4.2 to 60/a long. Ventrally with a pair- of ciliated 
setae on maxillae, one on each coxa, a pair between coxae I and between 
coxae III, and behind coxae III with ca. 6.4.6.4.8.6.4.4 setae to 40/a long. Legs: 
I 310/£ long, II 270,*, III 330,*; tarsi I and II with dorsal sensory rod, III 
without any long nude seta. 



'27 







Records of the 


S.A. Museum 










Standard 


Theoretical 


Observed 


Coeff . ot 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


62-0±:l-73 


3»00±l-22 


53^0-71-0 


59'0-65-0 


4^8 


PW 


78-0Hbl-53 


2-G4±l-08 


70-1-85-9 


75*0-80-0 


3-4 


SB 


29-0 


No variation xeeorded 






ABB 


26-7+1-33 


2-31:t0-94 


19-8-33- 6 


24-0-28-0 


8<6 


PSB 


23-7±0-33 


0-58±0-23 


22-0-25-4 


23*0-24-0 


2-4 


SD 


50-3±l-20 


2-08±0-85 


44-1-56-5 


48-0-52-0 


4-0 


A-P 


36*0±0-82 


l-41;£0-58 


31-8-40-2 


34-0-38-0 


3-9 


AM 


46-0 


No variation recorded 






AL 


33-0±0-58 


l-00±0-41 


30-0-36-0 


32-0-34-0 


3-0 


PL 


49 


No variation recorded 






Sens. 


39-3±0-33 


0-58 ±0-23 


37-6-41-0 


39-0-40-0 


1-5 



with head 13-0 wide. 
, SOHONGASTIA (ASCOSOHONGASTIA) TRAUBI 8p. n. 

Plate 62, fig. F-L 

Description of Larvae, Shape almost round. Length (imengorged) 195p., 
width 162/l. Scutum as figured, almost rectangular ; PL setae the shortest, AM 
the longest. SB rather wide apart and about the midline of the scutum; sen- 
sillae rather narrowly clavate and ciliated. Eyes 2 + 2. Chelicerae non-serrate, 
with only the apical tricuspid cap. Galea! setae nude. Palpi with bifurcate 
tibial claw ; all setae of femur, genu and tibia apparently nude. Dorsal setae on 
distinct platelets, ca. 38 in number, to 30/a long, shortly ciliated and arranged 
2.6.2.6.8.8.4.2. Ventrally with a pair of branched maxillary setae, a ciliated 
seta on each coxa, a pair between ooxae I and between coxae III, and thereafter 
ca. 40 setae in rows of 12 to 6, followed by 8 to 10 setae on platelets as on the 
dorsum. Legs.* fairly stout, 1 7-segniented and 220/a long, II and III 6-seg- 
mented and 195/i. and 245/a respectively ; no long nude seta on tarsi III. 

The Standard Data derived from the type and 3 paratypes are: 

Standard Theoretical Observed Coeff. of 

Mean Deviation Range Range Variation 

AW 63-0±:0-81 l-62:±:0-57 58-2-67-8 61-6-64-4 2-5 

PW 75*95±0-35 0*70:£0-25 73-85-78-05 75-0-77-0 0-9 

SB 37-8±0-81 l-62ct0-57 33-0-42-6 36*4-39-2 4-3 

ASB 28-0 No variation recorded 

P8B 25-2 No variation recorded 

SD 53 "2 No variation recorded 

A-P 37-8d:0-81 l-62±0-57 33-0-42-6 36-4-39-2 4-3 

A^l 42*0 No variation recorded 

AL 34-8±0-76 l-51±0-53 30-3-39-3 33*6-36*4 4-3 

PL 22-4 No variation recorded 

Sens. 47-6 tjy 6-0 wide. One determination only. 

hoc. and Host. The type and 3 paratypes (No. 3,969, ex Suncus sp.) from 
Shiugbwiyang, Burma, 23 Oct., 1945 (coll. R. Tranb and D. D. Millspaugh for 



Womersley— Asiatic-Pacific Scrub Typhus Mites 221 

The Standard Data for the type and 5 paratopes are : 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


79-3±0-59 


l-45±0-42 


75-0-83-6 


78-4^81-2 


1-8 


PW 


99-2:±0-56 


1-38±0-40 


95 -1-103' 3 


98*0-100-8 


1-4 


8B 


35-9±0-47 


1-14^0-33 


32-5-39-3 


33-&-36-0 


3-2 


ASB 


30-3±0-47 


l-14±0-33 


26-9-33-7 


28-0-30-8 


3-8 


PSB 


20-l:fc0-47 


l-14±0-33 


16-7-23-5 


19-&-22'4 


5-7 


SB 


50*4 


No variation recorded 


. 




A-P 


25*2 


No variation recorded 






AM 


47-6 


No variation 


recorded 






AL 


40-6=t0-95 


2-34;t0-67 


33-6-47-6 


36-4-42-0 


5-8 


PL 


70-l:t0-67 


l-65±0-47 


65-2-75-0 


C7-&-72-8 


2-3 



Sens. 44-8 vrith head IX -2. No variation recorded. 

Loc. The type and five paratypes from boots, Johnston's Pass, Wongabel, 
Queensland, Feb., 1945 (R. N. MeCulloch). 

Remarks. In the shape and dimensions of the scutum and the dorsal setae 
this species is near to #, (A,) tue&traliensis (Worn.) but differs in the bifurcate 
palpal claw, and the nude galeal setae, 

ScHONGASTiA ( Ascoschongastia) heaslipi (Worn, and Heasp. 1943), 

Neoschongastia headipi Womersley and Heaslip 1943, Tr. Boy. Soc. S. Auat, 
67, (1),120. 

Plate 62, fig. A-E. 

This species which was described from the type and two paratypes from 
rats from Intake, Cairns, Queensland (coll. 1940, W.G.H.), is distinguished from 
all others in the genus Schongastia, except trauM sp. n. in that the dorsal setae 
arise from the centre of small platelets, the diameters of which are approxi- 
mately half the length of the setae. 

The dorsal scutum is as figured with the posterior margin only moderately 
deeper behind PL and only lightly concave medially; sensillae clavate, setulose 
and their bases in front of line of PL. The PL setae are the longest and AL 
the shortest but all scutal setae are subequal. The palpal claw is trif urcate j the 
seta on femur -with 4 or 5 branches, on genu with apparently only 2 branches, 
and all 3 setae on tibia nude ; on the palpal tarsus the sub-basal sensory rod is 
exceptionally long. The dorsal setae number 54, each one on a separate small 
plate, arranged 2.6.8.8.8.4.6.6.4.2, and from 27 to 52^ long. Leg 1 7-segmented, 
II and III 6-segmented, as in oudemamsi Walch. 

The Standard Data for the type and 2 paratypes as given by Woinersley 
and Heaslip 1943 and treated statistically are as follows : 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



!23 



U.S.A^ Typhus Commission). Also 2 specimens from Baitus mbanus, from 
Ulu Langat, Kuala Lumpur, Malaya, 5 Dec, 1949 (J. R. Audy). 

Remarks. This species is very closely related to £. (A.) heaslipi Worn, and 
Heasp. 1943 in having the dorsal setae arising from distinct platelets. It differs 
however, in having only a bifurcate palpal claw, nude setae on palpal femur and 
genu, and in having some of the posterior ventral setae also on platelets. To- 
gether with headipi it agrees with iS. {A.) oudemansi m having the femora of 
legs II and III undivided and it also resembles oudemansi in the shape of the 
scutum and the short PL setae, but differs in the non-serrate chelicerae and the 
dorsal platelets. 

ScndNGASTU (Ascoschongastia) lawbbncei nom. nov. 

Neostfwngastia guntheri Womersley and Heaslip 1943, Tr. Boy. Soc. S. Aust., 
67, (1), 126, (preoc. Radford 1942). 

Plate 63, fig. A-E. 

As the specific name guntheri was used by Radford 1942 (Parasitology) for 
a species of Neoschongastia (Asooschongastia) from Colorado, the name of the 
Australian species is now changed to lawrencei in honour of Dr. F. R. Law- 
rence, Director of the Natal Museum. 

The dorsal scutum is as figured, with posterior margin fairly deep behind 
line of PL and slightly concave medially ; the sensillae are clavate with promi- 
nent setules, and their bases in front of line of PL ; AL is slightly the shortest 
and PL the longest. The chelicerae are simple with only the apical tricuspid 
cap. Galea! setae long, slender and nude. Palpi stout, with trifurcate tibial 
claw; setae on femur and genu branched, on tibia all 3 setae nude. Dorsal 
setae numerous, ca. 80 in number, arranged 2,12.12.12.1210.8.6.4.2, to 75ji long. 
Tarsi III without any long nude seta. 

The Standard Data for the 17 specimens as given by Womersley and Heaslip 
1943 treated statistically are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Vaxiation 


AW 


77-6±0-50 


2-06±0-35 


71-4r-83-8 


75-0-81-0 


2-7 


PW 


97-2=h0-59 


2-44r£0-42 


89-9-104-5 


94^0-lOS-O 


2-5 


SB 


31-4:±0-23 


0-93±0-16 


28-6-34-2 


30-0-32-0 


3.0 


ASB 


30-l±0-54 


2-23:±r0-38 


23-4-36-8 


27-0-35-0 


7-4 


PSB 


27-l±0-47 


1-93+0-33 


21-3-32-9 


24-0-32.0 


7-1 


SI) 


57-2±0-78 


3-23±0-55 


48*3-36 -9 


54-0-64-0 


5-6 


A-P 


32-2+:0-48 


l-98±0-34 


26-3-38-1 


27-0-36-0 


6-0 


AM 


57-0±0-82 


3*16±0-58 


47-5-66-5 


49-0-60-0 


5-5 


AJL 


39-7±0-32 


l-32:*:0-23 


35-7-43-7 


37-0-41-0 


3-3 


PL 


67-2:fc0-40 


1-67 ±0-29 


62-2-72-2 


65*0-70-0 


2-5 


Sens. 


51-8±0-29 


1-17±0-21 


48-3-55-3 


49-0-54-0 


2-2 



with head 12*0 -wide. 



224 Records of the S.A.. Museum 

This species was described from the type and 18 paratypes from a rat from 
Cairns, Queensland (1939-40, W.G.EL) and in the South Australian Museum 
collection is another slide of 6 specimens from a rat from Wongabel, Q,, Feb., 
1945 (R.N.Mcd). 

Its relationship to other species is best seen from the key. It is close to 
wongahelen&is sp. n., smithi, and cairnseims in having SB in front of PL, but 
differs in the more numerous dorsal setae and other details. 

It is renamed in honour of Dr. R. F. Lawrence, Director of Natal Museum. 

Schongastia (Ascoschongastia) wongabelensis sp. n. 
Plate 63, fig. F-J. 

DescHption of Larvae. Shape oval. Length (engorged) 450/*, width 230jn. 
Scutum large as figured, finely and indistinctly punctate, anterior margin 
lightly sinuous ; posterior margin rounded laterally and flattened medially and 
fairly deep behind line of PL ; AL very short, PL the longest ; sensilla^ fairly 
widely clavate with prominent setules, and the sensillae bases well in front of 
PL. Eyes 2 + 2, on distinct ocular shields, posterior the smaller. Chelicerae 
simple, with only the apical tricuspid cap. Galeal setae nude. Palpi stout, 
with trifurcate tibial claw; setae on palpal femur with many long branches; 
seta on genu with 1 long branch; on tibia all three setae nude; tarsi with 4 or 
5 ciliated setae and the usual basal and subapical sensory rods. Dorsal setae 
tapering and ciliated to 45/x. long, 20-22 in number and arranged 2.6.6 (4). 4 (6). 
2,2(0). Ventrally with the usual pair of branched setae on maxillae, one on 
each coxa, a pair between coxae I and between coxae III, and thereafter ca. 40, 
arranged ;8.6'.8.8,6.2.2.2, to 20/* long. Legs: I 210/* long, II 200*4, III 250**.; 
tarsi I and II with the usual dorsal sensory rod; III without any long nude 
seta. 

The Standard Data for the type and four paratypes and six other speci- 
mens are : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 


74-6=t0-57 


l-89±0-40 


68-9-80-3 


72-8-78-4 


2-5 


PW 


102-3±0-44 


l-46±0-31 


97- 9-106*7 


100-8-103-6 


1-4 


SB 


45-6;±Q-55 


l-81±0-39 


40-2-50'8 


42-0-47-6 


4-0 


ASB 


31-3±0-34 


1-13±(>.24 


28 '9-34 -7 


30-8-33-6 


3-6 


PSB 


41-0=b0-43 


1-43 ±0-30 


36-8-45-2 


39-2-42-0 


3-4 


SD 


72- 3d:0- 66 


2-10+0.45 


66-0-78-6 


70-0-75-6 


2-9 


A-P 


45*3:1:0-74 


2-45±0-52 


38-0-52-6 


47-0-47-6 


5-4 


AM 


43-2±0-57 


l*50:t0-40 


38-7-47-7 


42-0-44-8 


3-5 


AS 


94-l±:0-46 


l-45±0*32 


39-8-28-4 


22-4-25-2 


6-0 


PL 

Sens. 


51-2:h0-55 l'8I±0-39 
44-8 with head 16-8 wide. No v. 


45-8-56-6 

ariation recorded. 


47-6-53-2 


3-5 



Womersley— Asiatic-Pacific Scrub Typhus Mites 225 

hoc. and Host. The type and 5 paratypes from a water rat from Wonga- 
bel, Q., Feb., 1945 (K. N. McCulloch), and 5 other specimens from the same 
locality, host?, Feb., 1945 (R.N.Mc). 

Remarks. A characteristic species separated as in the key. 

SCHONGASTIA ( ASCOSCHONGASTIA ) SMITHI (Wom. 1939). 

Neoschdngastia smifhi Womersley 1939, Tr. Roy. Soc. S. Aust., 63, (2), 164; 
Womersley and Heaslip 1943, ibid., 67, (1), 127, 

Plate 65, fig. A-E. 

No further specimens of this species, described from Rattua assimiUs, from 
Imbil, Queensland, have been seen. 

In the key it is closely related S. (A.) cairnsensis from which it differs in 
the number of dorsal setae and the branched galeal setae. 

The dorsal setae number 46, and are arranged 2.8.8.8.8.6.4.2. The dorsal 
scutum is deep behind PL with A-P approximately equal to PSB. 

Originally known from the type and 2 paratypes, only the type now exists, 
*but the Standard Data as given by Womersley and Heaslip in 1943 for the 
three specimens and treated statistically are : 









Standard Theoretical 


Observed 


Coeff. of 




Mean 


Deviation Range 


Range 


Variation 


AW 


70-0 




No variation recorded 






PW 


81*0 




No variation recorded 






SB 


27-0 




No variation recorded 






ASB 


30 '0 




No variation recorded 






PSB 


24-0 




No variation recorded 






8D 


54-0 




No variation recorded 






A-P 


29-7±l 


•45 


2-51:±l'03 22-1-37-3 


27-0-32-0 


8-5 


AM 


S3-0±l' 


15 


2-00±0-81 47*0-59-0 


51 -0-55-0 


3-8 


AL 


34-7±l 


•76 


3-05d:l-25 25-6-43-8 


32-0-38-0 


8-9 


PL 


63-0±l 


•58 


1-00+0-41 60-0-66-0 


62*0-64-0 


1-6 


Schp. 


43-0 




Only one determination. 


■ 





SCHONGASTIA ( AsCOSCHONQASTIA) RATJI Bp. n. 

Plate 64, fig. A-JX 

Description of Larvae, Shape, an elongate oval with slight tendency when 
unfed to a constriction posterior of coxae III. Colour reddish. Length (only 
lightly fed) 195/i, width 162ft. Scutum as figured with anterior margin sinuate, 
posterior margin shallowly rounded. Sensillae clavate on the distal half, with 
apex of club somewhat flattened, bases wide apart and in front of line of 
PL; sensillary club either nude or only indistinctly setulose; ordinary scuta! 
setae shortly and indistinctly ciliated. Eyes 2 + 2, close to posterolateral 



226 Records of the S.A. Museum 

corners of scutum. Chelicerae simple with only the apical tricuspid cap. 
Galeal setae nude. Palpi stout with bifurcate tibial claw; setae on palpal 
femur, genu and tibia apparently nude. Dorsal setae tapering shortly ciliated, 
28 in number, to 36ju long, and arranged 2.8.6.6,4.2. Ventrally with a pair of 
ciliated setae on maxillae, one on each coxa, a pair between coxae I and 
between coxae III and thereafter ca. 30 setae only slightly shorter than dorsal. 
Legs all 7-segmented, I 227/* long, II 195/a, III 227/* ; tarsi I and II with dorsal 
sensory rod, III without any long nude seta. 

The Standard Data for the type and 3 paratypes are ; 



AW 


Mean 
62-3dr0-70 


Standard 
Deviation 

l-40±0-49 


Theoretical 
Range 

58-1-66-5 


Observed 
Bange 

61-6-64-4 


Coeff. ot 
Variation 

2-2 


PW 
SB 


86-l:£0-70 
30-8 


l-40zh0-49 81-9-90-3 
No variation recorded 


84-0-86-8 


1-6 


ASB 


25-2 


No variation recorded 






PSB 


25-2 


No variation recorded 






8D 


50-4 


No variation recorded 






A-P 


23-8:t0-81 


l-61±0-57 


19-0-28-6 


22-4-25-2 


6-5 


AM 


39- 2;fcl-14 


2-28^0-81 


32-4-46-0 


36*4-42-0 


5-8 


AL 


27-J:£0-93 


l<62±:0-66 


22-3-3L9 


25-2-28*0 


5-9 


PL 

Seng. 


41-3drl'34 
47 »6 with liead 


2-68±0-95 
8-4/19-6. Nova 


33-2-49-4 
riation recorded* 


39-2-44-8 


6-5 



Loc. and Host. The type and three paratypes from beneath the belly 
scales of a skink Leiolojn&ma guichenoti Dumeril and Bibron from Middle 
River, Kangaroo Is., South Australia (coll. A. Rau, March, 1950). 

Remarks. The relationship of this species is best seen from the key. The 
form of the sensillae is somewhat unusual. It is named after the finder of the 
host. 

Schongastia (Ascoschongastia) cairnsensis (Worn, and Heasp. 1943). 

Neoschongmtia cairnsensis Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust, 
67, (1), 128. 

Plate 65, tig, F-J. 

This species is very common and widely distributed in Queensland and 
w 7 as recorded by Womersley and Heaslip 1943, from bandicoots, Isoodon torosus 
Cairns, 1939 (W.G.H.) and Brisbane, 1938 (W.G.H.) and from Cowan Cowan 
1939 (D. J. W. Smith); also from rats from Cairns 1939 (W.G.H.) , from 
Rattus assimilis, and R. lutreolus; and from R. yowngi from Cowan Cowan 1938 
(D.J.W.S.). 

Of this original material, that from bandicoots from Brisbane, 1938 
(W.G.H.) comprising 8 specimens is regarded as the type series, consisting of 
the type and 7 paratypes. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



227 



The palpal claw is trifurcate, and the seta on palpal femur 4-5 branched, 
on genu 1-2 short branches, on tibia all 3 setae nude. The chelicerae are simple 
with only the apical tricuspid cap, and the galeal setae are nude. The dorsal scu- 
tum is as figured, with the posterior margin rather shallow behind PL and lightly 
concave medially; the lateral margins are lightly concave; the scnsillac are 
somewhat broadly clavate, with pronounced setules, and their bases in front of 
line of PL, The dorsal setae number 32, to 70/a long, and arranged 2.6.6.6.6.4.2. 

The Standard Data show r s rather a large variation and for 62 specimens 
of the original material are as follows : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


56-2±0-31 


2-46+0-22 


48-8-63-6 


50-4r-61-6 


4*4 


PW 


82 '8+0-62 


4*91+0-44 


68 -1-97 -5 


72-8-98-0 


5-9 


SB 


29*5±0*27 


2*10+0*19 


23-4-35-8 


25-2-36-4 


7-1 


ASB 


29*5+0-19 


1*48±0-13 


25 '1-33 »9 


25-2-30-8 


5*0 


PSB 


20-2±0'16 


1-27+0-11 


16-4-24-0 


19-6-25-2 


6-3 


SB 


49 -7+0-26 


2*05+0-18 


43-5-55-9 


44-8-56-0 


4-2 


A-P 


33-7:±0*29 


2-27+0-20 


26-9-40-5 


28-0-39-2 


6-7 


AM 


40*9+0-24 


1-75+017 


35*6-46-2 


36-4-44-8 


4-3 


AL 


31*2+0*29 


2*22+0*21 


24 *5-37 -9 


28-0-36-4 


7-1 


PL 


51-25+.0-37 


2*76+0-26 


42-95-59-55 


44 "8-56-0 


5-4 


Sens. 


37-5dt0-91 


3*17+0*65 


28-0-47*0 


33-6-42-0 


8-4 



with, head 14*0 wide. 



var, gateri (Worn, and Heasp. 1943). 

Neoschongnstia cairnsensis v. gateri Womersley and Heaslip 1943, Tr. Roy. Soc. 
S. Aust, 67, (1), 129. 

Plate 65, fig. K. 

This was recorded as a variety of cairnsensis only differing in the larger 
dorsal scutum. 

The Standard Data of the type and 4 of the five paratopes from rats at 
Cairns, 1939 (W.G.IL) are: 





Mean 


AW 


69-4+1-05 


PW 


99*7+4-21 


SB 


35*8±0*56 


ASB 


31*9±0-69 


PSB 


25-2 


SB 


57-l±0-69 


A-P 


39-2+1-53 


AM 


42-7+1*34 


AL 


31-2+0-56 


PL 


55*3+2-65 


Sens. 


39-2 with 1 



Standard 


Theoretical 


Observed 


Coeff. of 


Deviation 


Range 


Kange 


Variation 


2-34+0-74 


62*4-76-4 


67-2-72-8 


3-4 


9-41+2-98 


71-5-127-9 


89-6-112-0 


9-4 


1-25+0*39 


32. 0-39 -6 


33-6-36-4 


3-5 


1-53+0-48 


27*3-36*5 


30-8-33-6 


4*8 


No variation recorded 






3-53±0-48 


52-5-61*7 


56*0-58-8 


2-7 


3-42+1-08 


29-0-49*4 


33-6-42-0 


8-7 


2-68+0*95 


34-7-50-7 


39-2-44-8 


6-3 


1*26+0 -40 


27-4-35-0 


30-8-33-6 


4*0 


5-30+1-87 


39-4-71-2 


47-6-58*8 


9-6 



39-2 with head 14*0 wide. Only one determination. 



228 Records of the S.A. Museum 

Key to the I*arval Species of SOHONGASTIA Ouds. 1910. 

1. Chelicerae serrate on dorsal (inner) margin 2 

Subgen. Schongastia s. str. Ewing 1946. 
Chelicerae non-serrate on dorsal margin ... . » . . . . 14 

Subgen. Asooschdnga&tia Ewing 1946. 

2. All legs 7-segmented, i.e. femora secondarily subdivided. Tarsi III 
with a long outstanding nude seta ? except in oculicola sp. n 3 

Legs II and III 6-segmented, femur not subdivided. Tarsi III with- 
out any long nude seta. Scutum trapezoidal with posterior margin 
shallow behind line of PL. 2a 

2a. Scutum smaller* Sensillae clavate. DS 30 in number, arranged 
2.6,6.6.6.4. 

AW 51-4±:9-8, PW 66-1+7-0, SB 30-65+5-55, ASB 25 6, 
PSB 19-45±2-2, SD 45-2+2-8, A-P 34-0+2-2, AM 34-7+3-6, 
AL 22-65+3-75, PL 13-55+3-35, Sens. 38-0 with head 9-6 wide, 

Schon. (Schon.) oudemansi (Walch 1923). 

— impar Gunther 1939. 

= hodensis Gunther 1940. 

Scutum larger. Sensillae ? DS 42 in number, arranged 2.6.6.6.6.- 
6.4.4.2 to 45/x long. 

AW 68-0+4-8, PW 86-0+18-5, SB 38-4, ASB 35-2+7-8, PSB 

22-4, SD 57-6+7-8, A-P 44-8+7-8, AM 34*4+4-8, AL 35-2, 

PL 36-0+4-8, Sens. ? Schon. (Schon.) lewthwaitei sp. n. 

3. Palpal claw trif urcate 7 

Palpal claw bifurcate . . 4 

4. Sensillae bases in line with or slightly in advance of PL. Scutum 
hexagonal, Sensillae globose . . fc , _ . . . . 5 

Sensillae bases placed well behind line of PL 6 

5. Sensillae nude or only indistinctly villous under oil immersion. 
Scutum larger. DS ca. 2.10.8 (10). 2.10.6.6.4.2 (4) =50-54. 

AW 66-9+_9-9, PW 88-65+13-65, SB 25-8+2-5, ASB 35-5 
±2-9, PSB 28-7+2-2. SD 64-1+3-5, A-P 31-7+3-4, AM 37-3 
+4-9, AL 77-8+9-8, PL 58-1 ±4 -4, Sens. 32-1+1-6 with head 
16/19. ScMn. (Schon.) mndersandei Ouds. 1905. 

Sensillae distinctly setulose. Scutum smaller. DS ca. 2.8.10.10.6.2 
= 38. 

AW 53-1+8-4, PW 70-0+6-1, SB 20-5+ -0, ASB 27-0+4-2, 
PSB 22-55+1-85, SD 49-6+4-8, A-P 23-7+4-1, AM 27-8+6-1, 
AL 54-4+8-4, PL 43-1+9-9, Sens 32-0 with head 12-8/19-2. 

Schon. (Schon.) schuffneri (Walch 1923). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 229 

6, Scutum hexagonal, but A-P shorter than the posterolateral sides. 
DS ca. 2.10.2.13.2.14.12.12,8.4 == ca. 80. 

AW 62-4±2<5, PW 79-0, SB 21-2:fcl-3, ASB 35-0, PSB 23-5, 
SD58-5, A-P 23*5, AM 41-0+4-2, AL 77- 6dr 10* 2, PL 57-0+7-3, 
Sens. 35-0 with head 15/19, 

Schon. (Schon.) tayhri Gunther 1940. 

Scutum not hexagonal, posterior margin deep behind line of PL and 
strongly evenly rounded. A-P approx. l-3rd the scutal depth. 
DS 2.8.8.8.6.4 = 36. 

AW 50-4+4-8, PW 64-0, SB 18-0±4-6, ASB 25-6, PSB 22-4, 
SD 48-0, A-P 14-4±5-6, AM 34-4+9-2, AL 56-8d=4-8, PL 49-6 
±5 ■ 6. Sens. 32 - with head 16/22. 

Schon. (Schon.) whartmu Bg. n. 

7. Scutum pentagonal . . * , . . . . . . . . ► • 8 

Scutum not pentagonal . . . * . . , . . - . . . . 9 

8, Sensillae clavate. Scutal and dorsal setae relatively short, rod-like, 
with short ciliations. Chelicerae with only two strong teeth on dorsal 
margin. All palpal setae on femur, genu and tibia nude. DS ca. 
2.10(8).6.6.4 = 30-32. 

AW 53-55+5-25, PW 69-7+3-9, SB 22-0+3-5, ASB 22-8+3-4, 
PSB 22-4, SD 45-2=3-4, A-P 26-8+5-0, AM 16-0, AL 28-8, 
PL 32-0, Sens. 32-8+4-8 with head 6-0 wide. 

Schon. (Schon.) bidentata sp. n. 

Sensillae globose. Scutal and dorsal setae normal with long cilia- 
tions. Chelicerae with more than two teeth. Seta on palpal femur 
branched, on genu nude. DS 2.8.6.6.4.4.2 = 32. 

AW 53-8+6-4, PW 73-0+6-9, SB 19-7+1-2, ASB 26-2+1-1, 
PSB 26-9+3-2, SD 53-1+2-1, A-P 30-8+1-7, AM 28-2+5-6, 
AL 60-3+13-8, PL 50-7+6-6, Sens. 39-3+3-0 with head 15/19. 

Schon. (Schon.) meta Gater 1932. 

9. Scutum hexagonal with the antero- and poster o-lateral sides about 
equal, posterior side lightly concave * « . ♦ . . . . • ► 10 

Scutum not hexagonal ♦■ * • ► • - 12 

10. Seta on palpal genu nude. Chelicerae rather short and curved. DS 
ca. 2.8.6.6.4 (4). 2 — 28-32. 

AW 48-6 + 7-5, PW 69-6+2-7, SB 18-4+2-7, ASB 24-0, PSB 
21-0, SD 45-0, A-P 23-6+2-7, AM 22-2+4-9, AL 59-2+3-3, 
PL 42-6+7-5, Sens, 30-0 with head 15/18. 

Schon. (Sdh/on,) philipi Worn, and Kohls, 1947 

Seta on palpal genu branched. Chelicerae longer and straighter. 

DS 50 to 64 in number „ . . U 



230 Records of the S.A. Museum 

11. Seutal and setae on palpal femur and genu only short branched. No 
•crescent in front of each sensilla base. Sensillae bases nearer to one 
another than they are distant from each PL. DS ca, 2.12.8.8.10.8.4.2 
*=54. (Guntlier gives 64.) 

AW 65-7di7-7, PW 86-0+10-0, SB 25*9+2-8, ASB 32-8±:5-0 3 
PSB 27-0+5-2, SD 59-8+5-6, A-P 30-4+4*8, AM 34-9 + 5-9, 
AL 67-1+9-0, PL 51-35+7-2, Sens. 33-5+4-6, with head 19/19. 

Schdn. (Schdn.) Mestowei Gunther 1939, 

Seutal and setae on palpal femur and genu long branched. A dis- 
tinct crescent in front of each sensilla base. Sensillae bases as far 
apart, or slightly more, than they are distant from each PL. DS ca. 
2,12 (14) .12 (10) .12.10.6.2 = i>6. 

AW 57-7+5-8, PW 80-3+5-2, SB 32-2+2-4, ASB 30-2+6-2, 
PSB 23-0+3*3, SD 53-2+4-4, A-P 24-4+7-l,AM 38-55+3-15, 
AL 67-65+9-35, PL 57-2+6-0, Sens. 33-15+8-3 with head 
19/20. 

Schdn. (Schdn-,) bUstowei var. m-egapodius Worn, and Heasp. 1943. 

12. Lateral parts of posterior seutal margin almost in line with PL and 
lightly concave, middle part strongly convex and deeper behind line of 
PL. Sensillae globose, ciliated, apparently with only short peduncles, 
and their bases placed well in advance of PL. Seta on palpal femur 
with only a few indistinct branches, on genu nude. Chelicerae short 
and stumpy. DS fine, tapering, with indistinct short cilia tions, ca. 
2,4.4.8,8,6.4.2 =38. 

AW 44-8, PW 70-4, SB 16-0, ASB 22-4, PSB 19-2, SD 41-6, 
A-P 28-8, AM 16-0, AL 36-0, PL 48-0, Sens. 22-0 with head 
16/16. Schdn. {Schdn.) oculicola&p. n. 

All the posterior seutal margin deep behind line of PL t laterally 
rounded and medially straight .. . . .. ..IS 

13. Scutum much wider than long. Setae on femur and genu of palpi 
with only a few short, indistinct branches. DS 2.8.6.6.6.4.2 =34. 

AW 57-0+14-2, PW 78-1+10-9, SB 22-4, ASB 25-6, PSB 
23-05+2-65, SD 48-65+2*65. A-P 28-8, AM 26-2+4-3, AL 
66-1+8-9, PL 53-9+3-2, Sens. 35-2 with head 13/19. 

Schdn-. (Schdn.) pseudoschuffneH (Waleh 1927). 
Scutum not much wider than long, almost quadrate, with strong 
punctations. Setae on palpal femur and genu more distinctly and 
longer branched. DS 2.8.6.6.6.4.2.2 = 36. 

AW 51-2, PW 66-0, SB 16-0, ASB 25-6, PSB 25-6, SD 51*2, 
A-P 28-8, AM 28-8, AL 54-4, PL 48-0, Sens. 32-0 with head 
15/20. Schdn. (Schdn,) rnaldivicnm (Radford 1946). 

14. Some or all dorsal setae as well as PL and sometimes AL or AM 
foliate. foliata group 15 
All seutal and dorsal setae normal and non-foliate IS 



Womeksley — Asiatic-Pacific Scrub Typhus Mites 231 

15. Galeal setae ciliated or branched. Dorsal scutum with posterior cor- 
ners -well rounded. PL nearer to AL than to posterior margin. Sen- 
sillae bases behind line of PL. Only PL and all DS except posterior 
two rows long and foliate with small serrations or denticles. Palpal 
claw trifurcate, all setae on femur, genu and tibia strongly branched. 
DS 38, arranged 2.8.6.6.6.6.2.2 to 85/t long. 

AW 70-4, PW 84-3+5-6, SB 32-0+9-6, ASB 28-8, PSB 24-5, 
+2-8, SD 52-3+7-3, A-P 16-0, AM 54-4, AL 25-6, PL 81-1+ 
5-6. Sens. — . Schon. (Asco$chdn>*)psetidomy$ Bp.n. 
Galeal setae nude 16 

16. Scutum relatively deep behind line of PL. Sens, bases slightly be- 
hind line of PL. All dorsal setae as well as AL and PL very broadly 
foliate with long and strong denticles or teeth. Palpal claw trifur- 
cate ; setae on palpal femur and genu branched j on tibia all 3 appar- 
ently nude. DS 26 in number, arranged 2,6.6.6.4.2, and to 48-60/* 
long. Sensillae capitate and apparently nude. 

AW 60-0, PW 85-0, SB 24*0, ASB 30*0, PSB 21*0, SD 51-0, 
A-P 29-0, AM 21-0, AL 42-0, PL 56-0, Sens. 32-0. 

Schon. (Ascoschon.) mccullochi (Worn. 1944). 

Scutum not so deep behind PL. Sens, bases well in front of line of 
PL. Foliate setae with smaller denticles . . . . . ► , , 17 

17. All DS foliate with the denticles longer. AL and PL foliate. Palpal 
claw bifurcate, all setae on palpal femur, genu and tibia apparently 
nude. DS 28, to 64/x long and arranged 2.6.6.6.6.2. Sens. 28/a with 
head 17/17 and apparently nude. 

AW 50-6+14 -8, PW 69-5+6-f), SB 19-0+4-2, ASB 21-0, PSB 
18-0, SD 39*0, A-P 21-6+3*6, AM 16-5+4-9, AL 49-5+4-9, 
PL 60*0+4-8, Sens. 28-0 with head 17/17, (After Worn, and 
Kohls). Schon. (Ascoschon.) uromys (Worn, and Kohls 1947). 

All DS foliate except outer members of posterior 3 rows, denticles 
shorter and more in the nature of ciliations. Only PL of scutum 
foliate. Palpal claw trifurcate; setae on femur and genu ciliated j 
on tibia dorsal and lateral nude, ventral branched. Sens, capitate, 
with fine short setules. 

AW 53-0, PW 70*0, SB 17-5, ASB 19-5, PSB 17*5, SD 37-0, 
A-P 25-0, AL 57-0, PL 86-0, Sens. 28-0, (after Gunther). 

Schon. (Ascmclwn.) fciiata (Gunther 1940). 

18. Sensillae globose . . . - . . »< . * . . . . . . 19 

Sensillae clavate, sometimes broadly so, or lanceolate . . * . - * 41 

19. Sensillae bases very close together, not more than 15/s* or 16yx. apart. 

ghbuhire group 20 
Sensillae bases widely separated . . . . . . . . . . . . 25 

20. PL the longest 21 

AL the longest . . . . . . . . . • . . . . . . 22 



232 Records of the S.A. Museum 

21. Sensillae bases in front of line of PL. Posterior scutal margin shallow 
behind PL. PL longer than PW. Coxae III 1-setose. Sens. nude. 
DS 2.10.8.6.6.4.2 = 38. 

AW 41-9+2-7, PW 64-0, SB 12-5±:l-8, ASB 22-4, PSB 11 -6±: 
3-0, SD 34-O±:3'0, A-P 31- 5*2*4, AM 29*4+5'8, AL 63-4zb 
8-4, PL 70-9±8-3, Sens. 35-2 with head 19-6/22-4. 

Sclion* (AscoscJio7u) signata sp. n. 

Sensillae bases behind line of PL. Posterior scutal margin as even 
curve and deeper behind line of PL. PL about as long as PW. 
Coxae III 3 (4) -setose. 
AW 57-6, PW 73-6, SB 11*2, ASB 22-4, PSB 12-8, SD 35-2, 
A-P 22*4, AM !, AL t, PL ?, Sens. 25-0 with head 15/19 (after 
Walch ) . Schon. (Ascoschon, ) glabvlare ( Walch 1927 ) . 

22. Coxae III 1-setose. Sens, bases posterior of PL . . . . . . 23 

Coxae III multisetose . . . . . . . . • * . • • • 24 

23. Dorsal setae ca. 86, and to 4O-50ft long. All setae on palpal tibia 
branched or ciliated. 

AW 52-3=t6-5, PW 63-6^5-7, SB 9-6±2-7, ASB 24-3±:3-8, 
PSB 8-5±2-8, SD 32-8:±3'8, A-P 19-7+3-3, AM 22-5dz9-l, 
AL 45-5±_4-6, PL 34*3+7-5, Sens. 24-6+11-1, 

Schon. (Ascoschon.) kohlsi Philip and Woodward 1946. 

Dorsal setae more than 110, and to 28/* long. All 3 setae on palpal 

tibia branched. 

AW 59-0, PW 70-0, SB 10-0, ASB 28-0, PSB 14-0, SD 42*0, 
A-P 18-0, AM 24-0, AL 42-0, PL 31-0, Sens. 30-0 with head 
18/18. • Schon. {Ascoschon.) omurta Bp. n. 

24. Coxae III 2-setose. Posterior scutal margin deep and 3-sided. Sens, 
bases behind line of PL. Palpal claw trifurcate. all setae ciliated. 
Galeal setae nude. DS 38 in number, to 38 k a long and arranged 
2.8.6.6.8.4.2.2. 

AW 57-1+8-0, PW 70-9+8-0, SB 9-2+2-1, ASB 25-6, PSB 
12-8, SD 38-4, A-P 17-8+4-3, AM 32-3+5-2, AL 56-1+6-6, 
PL 45 -6^:4 -3, Sens. 28-8 with head 16/19. 

Schon. (Ascoschon.) lanius (Radford 1946). 

Coxae III 3-5-setose. Posterior scutal margin shallower and an even 
curve. Sens, bases slightly in front of PL, Galeal setae nude. 
Palpal claw trifurcate; setae on palpal femur and genu with a few 
short ciliations ; on tibia dorsal and lateral nude or with a few 
indistinct ciliations, ventral branched. DS 40 in number, to 38/a long 
and arranged ca. 2.8.6.6.8.2.4.2.2. 

AW 48-0, PW 62-0, SB 9-0, ASB 26*0, PSB 13-0, SD 39-0, 
A-P 26-0, AM 29-0, AL 53-0, PL 40-0, Sens. ?. 

Schon. (Ascoschon.) mutaMlis (Gater 1932), 



WOMEKSLEY— ASIATJC-pAClFIC SCRUB TYPHUS MlTES 233 

25. Palpal claw bifurcate . . . . - * * . - 26 

Palpal claw trifurcate . . . - . - . . ^ . . - • . 28 

26. A long nude seta on tarsi III. Scutum larger, more or less six-sided 
with posterior margin deep behind PL and only slightly concave medi- 
ally. Galeal setae nude. Setae on palpal femur and genu branched; 
on tibia, dorsal and lateral nude, ventral branched. Sensillae capitate, 
with distinct setules and placed just behind line of PL. DS 62, to 
50/a long and arranged 2.1414.10.12.8.4. 

AW 76-0, PW 96-0, SB 38-0, ASB 32-0, PSB 29-0, SD 61-0, 
A-P 29 -0, AM 48-0, AL 80-0, PL 72 -0, Sens. 28-0. 

ScJion. (Ascosckon.) edxmrdsi (Gunther 1939). 

Ko long nude seta on tarsi III . . . . . . . . . . « • 27 

27. Dorsal setae normal and ciliated. Seutum with posterior margin 
strongly sinuate and deeply concave medially. Galeal setae nude. 
Setae on palpal femur and genu branched ; on tibia, dorsal and lateral 
nude, ventral branched. Sensillae capitate, with distinct setules, and 
in line with PL. DS 32-34, to 30^ long and arranged 2.6.6.6.6.4(2) .2. 

AW 54«6±5»o, PW 79-2+6-6, SB 23*0+5-4, ASB 21*0, PSB 
12-3±3-6, SD 33-3+3-6, A-P 25-0, AM 24-5+3-6, AL 44-7 
±6 »5, PL 53-lrh6-6, Sens. 29-0. 

Schan. (Ascoschbn.) phiHpp&nsis (Philip and Woodward 1946). 

Dorsal setae strong, with 5-6 lateral long teeth. Galeal setae nude. 
Seta on palpal femur strongly ciliated, on genu much finer with few 
branches; on tibia, all 3 nude. Sensillae capitate, with distinct fine 
setules and well behind line of PL. DS approx. 100 to 70-56/i. long 
and arranged ca. 8,16.18, plus, 

AW 58-8, PW 75-6, SB 33-6, ASB 25-2, PSB 16-8, SD 42-0, 
A-P 14-0, AM 53-2, AL 30-8, PL 78-4. Sens. 36-4 with head 
19 -6/25 -2. Bchon. (Ascmchdn.) maeJcerrasae sp. n 

28. PL setae situated off the scutum. Coxae III 2-setose. Palpal claw 
trifurcate; seta on femur branched, on genu long and nude, on tibia 
dorsal and ventral branched. DS 46 in number, arranged 2.10.8.10.- 
8.6.2 to 25/4.. 

AW 38-42(40), PW 66-74(70), SB 19-24(21), ASB 14-19(17), 
PSB 13-16(14), SD 27-35(31), A-P 32, AM 28, AL 14, PL 24, 
Sens. 30-32(31). 

Schori. (Ascoschdn.) masta Traub and Sundermeyer 1950. 

PL setae on the scutum . . . , . t . . . . . . . . 29 

29. Coxae III 2-setose. Scutum with posterior margin well rounded later- 
ally, flat medially and deep behind line of PL. Setae on palpal 
femur and genu strongly branched; on tibia, dorsal and lateral nude, 
ventral branched. Galeal setae nude. Sensillae setulose. DS 2 plus 
ca. 10 rows of 10 each, to 45/x long with long outstanding ciliations. 



234 Records of the S.A. Museum 

AW 73-0+4-9, PW 98-8+9-0, SB 34-7+6-2, ASB 32-0, PSB 
30-0, SD 62-0, A-P 36-0, AM 41-0+4-7, AL 49-7+7-7, PL 
55*0+4-7, Sens. 32-0. Schon. (AscoscMn.) petrogale (Worn. 1934). 
Coxae III 1-setose 30 

30. PL setae placed much nearer to AL 7 and not at the posterolateral 
angles of the scutum. Dorsal cuticular striations very finely crenu- 

PL setae placed at the posterolateral angles of scutum . . . . 32 

31. DS 44 in number, to 55-65,* long, and arranged 2.10.10.10,6.4.2. PL 
relatively further from AL and scutum only slightly wider posterior 
of PL than distance between PL, 

AW 74-2+8-8, PW 87-0+8-7, SB 28*3+3-6, ASB 29-2+4-0, 
PSB 30-0, SD 59-2+4-0, A-P 21-8+3*5, AM 56-4+2-7, 
AL 46-0+7-5, PL 78-0+6-3, Sens. 36-0. 

Schon. (Ascostihon.) mohri-sp. n. 
DS more than 100 in number, to 60/u. long, and arranged ca. 2.14.12. 
16.12.12.12.10.8.6.4. PL relatively nearer to AL, and scutum pos- 
terior of PL much wider than between PL. 
AW 83-7+10-5, PW 92-3+12-1, SB 30-7+3-45, ASB 33-3+ 
6-9, PSB 30-0, SD 63-3+6-9, A-P 17-0+5-2, AM 54-0, AL 
45-7+6-9, PL 90-0, Sens. ? 8chdn m (Ascoschon.) crinita sp. n. 

32. PW less than 15,* longer than AW 33 

PW more than 15^ longer than AW 35 

33. Sens, bases very wide apart and nearer to lateral scutal margins 
than to one another; and nearer to line of AL than to line of PL. 
Scutum posteriorly overlapped by dorsal cuticle. Only the setae 
on palpal femur and genu, and the ventral on tibia branched. A 
long nude seta on tarsi III. Dorsal setae 36-38, arranged 2.8.6.8.6 
(4).4.2.2, to 60^ long. 

AW 76-2+3-8, PW 81-8+7-0, SB 47-0+3-8, ASB 28-0, PSB 
31-9+1-9, SD 59-9+1-9, A-P 30-8, AM 54*0+12-0, AL 82-9 
+9-5, PL 58*2+7-0, Sens. 39-2 with head 23/25. 

Schon. (AscosoJion.) reet amgulare sp.n k 

Sens, bases as near, or nearer to one another than to lateral scutal 
margins and much nearer to line of PL than line of AL. Scutum not 
overlapped by dorsal cutiele 34 

34. Scutum deep with A-P about 2/3 PW. Sensillae bases distinctly in 
front of line of PL. Dorsal setae to 52 in number, to 60//. long, and 
arranged 2.6.8.8.8.8.6.4.2. All setae on palpal femur, genu and tibia 
branched. 

AW 60-8+2-4, PW 69-6+7-5, SB 25-2, ASB 28-8+4-1, PSB 
19-G, SD 48-4+5-2, A-P 39-2+4-3, AM 38-2+4-0, AL 67-8 
+ 7-0, PL 59-9+9-6, Sens. 28*0 with head 22-4/22-4. 

Schon. (Ascoschon.) amtipodianum. (Hirst 1929). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 235 

Scutum shallower, with A-P about 1/3 PW. Sensillae bases about in 
line with PL. Dorsal setae thicker and stronger, 32 in number to 56/j, 
long, and arranged 2,6.6.6.6.4,2. Setae on palpal femur, and genu 
branched, on tibia all 3 nude. 

AW 60-7+4-8, PW 72-8+8-4, SB 19-6, ASB 25-2 PSB 16-8, 
SD 42-0, A-P 23-3+4-8, AM 40-0, AL 71-9+14-0, PL 79-3+ 
24-2, Sens. 30*8 with head 19-6/19-6. 

Schon. (Ascoschon.) cmsiope sp. n. 

35. Dorsal cuticular striations finely and closely crenulate. Scutum with 
anterior margin convex and posterior margin somewhat angular. 
Sens, bases anterior of line of PL. Seutal setae relatively short, PL 
the longest. Setae on palpal femur and genu branched ; on tibia only 
ventral branched. DS 52, to 30/a long, arranged 2,10.10.10.8.6.4.2. 

AW 58-8, PW 75-6, SB 22-4, ASB 22-4, PSB 22-4, SD 44-8, A-P 
25-2, AM 45-0, AL 33-6, PL 56*0, Sens. 36-4 with head 19-6/ 
19 ■ 6. Schon, {Ascoschon. ) dumosa sp. n. 

Dorsal cuticular striations not crenulate , , . , . . . . 36 

36. First row of DS with 4 setae, 2 scapular and 2 submedial. Posterior 
seutal margin shallow and almost rectilinear between PL. Setae on 
palpal femur and genu, and vent-rally on tibia branched. Dorsal 
setae 26, to 50jn long and arranged 4.6.6.6.2.2. 

AW 57-9+7-0, PW 90-7+7-8, SB 29-8+5-1, ASB 31*6+2-5, 
PSB 18-8+3-1, SD 50-4+5-7, A-P 42-8+8-1, AM 39-4+4-1, 
AL 70-85+9*25, PL 67-1+5-6, Sens. 40-0. 

Schon, (Ascoschon.) queenslandica (Womersley 1939). 

First row of DS with only the two scapular setae . 37 

37. PL setae longer than PW ., .. ... .. ,. .„ .. 38 

PL setae not longer than PW 39 

38. PL not greatly longer than PW. Ventral seta of palpal tibia nude 
or with only one branch. 

AW 54-0+6-0, PW 70-4+.7-2, SB 19-6±3<8 f ASB 25-2, PSB 
19-6, SD 44-8, A-P 27-8+3-6, AJVI 31-6+9-2, AL 70-4+4-8, 
PL 80 -9 ±10 -9, Sens. 33-6 with head 16-8/19-6. 

Schon. (Ascoschon.) lappacea n. comb. 

PL twice as long as PW, and much longer than AL. Ventral seta 
of palpal tibia strongly branched. DS 28, to 75/* long, and arranged 
2.6.6.6.2(4).4(2).2. 

AW 47-95+5-4, PW 68-95+8-9, SB 19-25+3-0, ASB 28-0, 
PSB 18-9+3-9, SD 46-9+3-9, A-P 31-5+3-9, AM 39-2+4-9, 
AL 84-0, PL 134-6+14-2, Sens. 33-6 with head 16-8/19-6. 

Schon. (Ascoschon.) womersl&yi (Gunther 1939). 



236 Records of the S.A. Museum 

39. With. AL setae the shortest. Sensillae further apart, with strong 
setules. All setae on palpal tibia nude. DS 32, to 45^* long, and 
arranged 2.6.6.6.6.4.2, 

AW 70-9^10-8, PW 95 -95^17 -45, SB 32-8zfc4-3, ASB 30-8 
±4-3, PSB 22-4, SD 53-2±4-3, A-P 34-5±5-9, AM 49-l±5-2, 
AL 42-0±5-3, PL 64-4±ll-0, Sens. 33-6 with head 19-6/19-6. 

Schon. (Ascoschon.) coorongense (Hirst 1929) 

With AM the shortest 40 

40. Scutum posterior of PL very shallow, margin between PL almost rec- 
tilinear. DS 34-38, arranged 2.8,6.6,6.4.5 (2). 2(0). to 30/* long. 

AW 45-3zh3-5, PW 61-5=fc6'4, SB 20-4+3-5, ASB 21-0, PSB 
15-0, SD 36-0, A-P 31-3±4-5, AM 26-6dr4-5, AL 46-5+7-1, PL 
42-6+4-5, Sens. 27-0 with head 17/17. 

Schon. (Ascoschon.) echymipera (Worn, and Kohls 1947). 

Scutum deeper behind line of PL. DS 32, arranged 2.6.6,6.6.4.2. 

AW 47-8+9-1, PW 72-1+11-9, SB 21-5+4-5. ASB 28 0, PSB 
19-6, SD 47-6, A-P 27-9+2-8, AM 28-55+3-55, AL 54-9+4-3, 
PL 47-6+5-0, Sens. 35-0. 

Schon. (Ascosckdn.) mnisfaiUnsis (Worn, and Heasp. 1943). 

AW 51-7+5-9, PW 81-3+11-5, SB 21-4+5-5, ASB 25*0+4-3, 
PSB 21-9+4-4, SD 47-0+7-1, A-P 28-8+2-8, AM 30-4+6-9, 
AL 60-2+13-2, PL 56-3+7-2, Sens. 31-4+5-6. 

Schon. (Ascoschon.) irmisfailerms f. bushlandi (Philip 1947). 

41, Sensillae long and narrow, almost lanceolate and without a true basal 
stem . . . . . . 42 

Sensillae clavate, with more or less of a basal stem . . . . . , 43 

42, Dorsal scutum larger, subquadrate, with PL the shortest and dis- 
tinctly on scutum at posterolateral corners. Seta on palpal genu 
very long. DS ea. 2.6.8.8.6.4.2. (but variable from 44-52 according 
toGater). 

AW 64-0+8-8, PW 75-2+5-9, SB 37-3+2-0, ASB 30-7+3-2, 
PSB 24-0+4-9. SD 54-45+7-2, A-P 44-0+3-5, AM 44-95+3-5, 
AL 35-5+2*8, PL 26-5+3-5, Sens. 54-0 with head 6-7 wide. 

Schon. (Ascoschon.) lacunosa (Gater 1932). 

Dorsal scutum small, with AW not greater than 50ft, PL placed dis- 
tinctly off the scutum. DS 0,8,8.6.4 to 30/x. Posterior scutal margin 
shallow and almost rectilinear between posterior corners. Setae on 
palpal femur, genu and tibia nude or with only a few indistinct barbs. 

AW 41-8, PW 66-0, SB 22*0, ASB 17*6, PSB 8-4. SD 26-0, A-P 
22*0, AM 22-0, AL 18-0, PL 30-0, Sens. ea. 34-0. 

Schon. (Ascoschd/t.) maiayensis Gater 1932). 

Dorsal scutum also small, AW not greater than 50/i, and with AL the 
shortest. PL on the scutum, DS 2.6.6,6,4,2. to 44/x. Posterior scutal 
margin shallow behind PL and concave medially. Setae on palpal 
femur and genu ciliated,- on tibia only ventral branched. 



WOMERSLEY— ASIATIC-PACIFiG SCRUB TYPHUS MlTES 237 

AW 44-6+4-9, PW 60-3+7-3, SB 20-5+4-2, ASB 20-9+4-2, 
PSB 14-0, SD 34-9+4-2, A-P 27-2+4-8, AM 35-6+6-8, AL 
31-9±5-7, PL 46-6+5-4, Sens. 61-6 with head 8-4 wide. 

Schon. (Asemchon.) Wpoxenasp.n. 

43. Scutum small, with AW ca. 50/* or less ±- 44 

Scutum much larger with AW much more than 55/x . . . « 52 

44. Dorsal scutum with AW da, 50-55/x, with PL very much shorter than 

AM or AL | and with AL at the anterolateral angles 44a 

Dorsal scutum with AW ca. 50ft or less. AL the longest . . . * 44b 

44a. Scutum larger, with AL longest and longer than PW. 

AW 54-15+4-1, PW 70-9+7-5, SB 32-4+5-2, ASB 35-4+3-2, 
PSB 26-25+4-1, SD 61-4+8-3, A-P 53-05, AM 55-25 + 8-4, AL 
73-1+21-4, PL 13-4, Sens. ? Schon, (Ascoschon.) nadvhut rami sp.n. 

Scutum smaller, with AL the longest but not longer than PW. 

AW 50-6+6-0, PW 65-1+7-3, SB 30-9+4-4, ASB 30-9+4-4, 
PSB 23-45, SD 54-35+4-4, A-P 46-9, AM 50-25, AL 56-95, PL 
13-4, Sens. 23-45 with head 10 -05/18 -75. 

ScJwn. (Ascoschon.) Sarawak ensis sp. n. 

44b. Dorsal scutum small, AW ca. 50^, with AL at the anterolateral 
angles ; posterior margin a shallow even curve ; PL the longest. Sens, 
fairly broadly elavate, and setulose, bases wide apart. Palpal claw 
bifurcate; setae on femur and genu branched; on tibia only ventral 
branched. Coxae III 3-setose. 

AW 50-5+4-5, PW 67-8+6-0, SB 37-3+5-5, ASB 22-3+1-2, 
PSB 22-3+1-2, SD 44-7+2-4, A-P 34*4+3-8, AM 20-0, AL 20-0, 
PL 30-25+1-85, Seus. 36-4 with head 13/22. 

Schon. (Ascoschon.) 7iausheraensis sp. n. 

Scutum geenrally with AW less than 50/a, and with AL well back 
from the rounded antero-lateral shoulders. indica group 45 

45. Posterior seutal margin rectilinear between PL 46 

Posterior seutal margin curved or sinuous between PL . . . . 47 

46. Scutum rectangular, with AW not much shorter than PW. Seutal 
setae strongly ciliated. 

AW 40-0, PW 48-0, SB 17-0, ASB 13-0, PSB 11-0, SD 24-0, 
A-P 21-0, AM 30-0, AL 20-0, PL 36-0, Sens. — . (After Gater 
1932 ; Womersley and Ilcaslip 1943) . 

Schon. (Ascoschon.) d^bilis (Gater 1932). 
Scutum not rectangular, with AW ca. 1/3 length of PW, lateral 
sides strongly converging forwardly. Seutal and dorsal setae shortly 
and indistinctly ciliated. 

AW 25-0, PW 74-0, SB 21-0. ASB 23-0, PSB 20-0, SD 43-0, 
A-P 42-0, AM 20-0 AL 17-0, PL 27-0, Sens. 38 -/J, (after Rad- 
ford 1946), Schon. (Ascoschon.) manipwrensis (Radford 1946). 



238 Records of the S-A. Museum 

47. Posterior scutal margin concave. All palpal setae ciliated- DS ca. 
62 in number, arranged 2.10.10.10.10.10.6.4, to 29/a long. No long 
nude seta on tarsi III. 

AW 28-8 PW 41*3+21, SB 13*4+3-9, ASB 14-4, PSB 11-2, 
SD 25-6, A-P 22-4, AM 16-0, AL 10-0, PL 19*2, Sens. — . 

Schon. (AscoscJwn.) Idbtianensis sp. n. 

Posterior scutal margin convex or convexly sinuous. A fairly long 
nude seta on tarsi III . . . . . . 3 . . 48 

48. Palpal claws short and stumpy . . . . . . ♦ . . . . . 50 

Palpal claws longer and more pointed * . . . - . . . i$ 49 

49. Scutum much wider between PL than long. Two pairs of setae 
between coxae III. DS 30 in number and arranged 2.6.6.6.6.4. to 36/x. 

AW 49-5+6-8, PW 67-2+10-6, SB 22-6+4-1, ASB 26-6+4-6, 
PSB 22-4, SD 42-0+4-6, A-P 28-9+4-3, AM 31-7+4-3, AL 
25-2, PL 39-7+3-4, Sens. 39-2 with head 11/22, 

Schon, (Ascoschon.) audyi sp. n. 

Scutum almost as long as wide between PL. Only one pair of setae 
between eoxae III. DS 22, arranged 2.6.6,4.2.2. 

AW 35*9+4-8, PW 64-8+4-8, SB 22-0+2-4, ASB 22-4, PSB 
35-2, SD 57-6, A-P 41-6, AM 25-0, AL 12-8, PL 38*4, Sens. 
35-2, Schon. {AscoscMn.) rattus (Worn, and Heasp, 1943). 

50. DS 22, arranged 2.6.6.4.2.2. 

AW 40-0, 38-4, PW 60-8, 61-8, SB 19-2, 22-4, ASB 19-2, 19-2, 
PSB 22-4, 22-4, 22-4, SD 41-6, 41-6, A-P 22-4, 22-4, A3f 36-0, 
AL 25-0, PL 37-5, Sens. 32-0 with head 23-0/9-5. (AM, and AL 
after Gunther). Schon. (Ascoschon.) lorius (Gunther 1939). 

DS more numerous , . 51 

5L Scutum smaller, DS 34 arranged 4(2).6(8).6.6.6.4.2. i.e. 2nd from 
middle of second row situated forward in line with humeral setae. 

AW 38-4+5-1, PW 54-0+5-7, SB 20*5+3-8, ASB 22-2+2-8, 
PSB 19-5+10-8, SD 42-3+3-0, A-P 26*7+5-3, AM 24-6+5-2, 
AL 18-8+3-7, PL 31-1+4-2, Sens. 31*1+4*8. 

Schon. (Ascoschon.) indica (Hirst 1915).. 

= Trombicula nmris Walch 1922. 

= Neoschongastia cockingsi Radford 1946. 

Scutum larger. DS ca. 42 in number, arranged 2.10.8.6.6.6.4, the 
second row somewhat variable from 8 to 10, but always in more or 
less the same line. 

AW 47-15+4-9, PW 64-1+8-5, SB 24-4+4-5, ASB 24-9+2-9, 
PSB 19-6, SD 44-1+3-0, A-P 25-5+3-0, AM 27-2+5-7. AL 20-1 
+3-3, PL 33-8+4-9, Sens. 34-7+5-1. 

Schon. (Ascoschon.) soekaboemiensis (Takekawa. 1945). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 239 

52. Sensillae bases distinctly posterior of line of PL, and PL nearer to AL 
than to posterior margin - - . . 53 

Sensillae bases in line with or anterior of line of PL, or if slightly 
posterior of PL then A~P ca. equal to or greater than P8B . . . . 58 

53. Scutum widest posterior of line of PL, and PL placed nearer to AL 
than to the well rounded posterolateral corners. Palpal claw trifur- 
eate ; setae on femur, genu and tibia all branched or ciliated. Galeal 
setae nude. DS ca, 80, to 50/* long, and arranged 2.12.12.12.12. plus 
ca. 30. Ventrally with many setae between coxae III. 

AW 63-5±:9-4, PW 78-7±:8-9, SB 25-l=h4-7, ASB 28-9±:4-0 ? 
PSB 25-8±3-l, SD 54-7=fc5-0, A-P 21-9+4-7, AM 48-2+3-9, 
AL 38-6+8-5, PL 74-3+9-2, Sens. 42-0 with head 14/28. 

Schon. (Ascoschon.) perameles (Worn. 1939). 

Scutum widest in line with PL, and PL at the posterolateral -corners. 
Only 1 pair of setae ventrally between coxae III . . . . . . 54 

54. Posterior scutal margin deep and evenly rounded . . , T - - 55 
Posterior scutal margin not an even curve, but roughly three-sided - . 57 

55. Dorsal setae 16 in number, arranged 2.4.4.4.2, the anterior setae to 
95/A long, decreasing posteriorly to 40-45/*, 

AW 65-6+6-6, PW 87-2+5-8, SB 34-4+.4-1, ASB 22-4, PSB 
28-4+3-2, SD 50-8+3-2, A-P 16-8, AM 53-0, AL 19-6, PL 99-2 
+11-7, Sens. 50-0 with head 11*0 wide. 

Schon, (Ascoschon.) similis (Worn, and Heasp. 1943). 

Dorsal setae more numerous . . . . . - . . . < . « 56 

56. Dorsal setae 26, arranged 2.6.6.6.4.2. 

AW 68-0+4-0, PW 88-75+6-7, SB 34-7+4-3, ASB 25-2 t PSB 
25-2, SD 50-4, A-P 14-0, AM 52-85+13-05, AL 25-75+6-6, PL 
86 -25 ±9 -55, Sens. 42-0 with head 16-8/30-8. 

Schon. (Ascoschon.) derricki Worn. 1939. 

Dorsal setae 34, arranged 2.8.6.6.6.4.2. Palpal claw trifurcate. 

AW 55-6:1:9*0, PW 65-2+10-5, SB 23-8+4-2, ASB 22-4, PSB 
20-4+4-1, SD 42-8+4-1, A-P 14-0, AM 35-6+6-3, AL 21-3 
+3-2, PL 48-0+3-2, Sens. 43-9+4-8 with head 8-4 wide. 

Schon. {Ascoschon.) dasycerci Hirst 1929. 

57. Scutal setae long and subequal, with AM slightly shorter than AL and 
PL the longest. Dorsal setae 2.8.6.6.6.4.2, to ca, 45/*. Palpal claw 
bifurcate; setae on femur and genu branched; on tibia, ventral 
branched and dorsal and lateral nude. 

AW 61-5, PW 70-0. SB 24-0, ASB 24-0, PSB 16-0, SD 40-0, A-P 
16-0, AM 37-5, AL 45-0, PL 56-0, Sens. — . (After Worn, and 
Heasp. 1943). Schon. (Ascoschon.) trichoswi. (Worn. 1939). 

" « shisldsi ( G unther 1941 ) . 



240 Records of the S.A. Museum 

AL scutal setae very short, and much shorter than AM or PL. Palpal 
claw trifurcate ; all setae on femur, genu and tibia except dorsal tibial 
branched. Dorsal setae 34, arranged 2.8.6.6.6.4.2, to 70/x. long. 

AW 68-4+8-2. PW 82-0+6-3, SB 31-6+6-3. ASB 24-8^3-2, 
PSB 22-4, SD 47-2+3-2, A-P 18-2+3-8, AM 45-2+5-8, AL 
25-2+4-8, PL 70-4+3-2, Sens. — . 

Schdii. (Ascoschon.) hirsti (Worn, and Heasp. 1943). 

58. Sensillae bases about in line with PL . . . . . - . . . - 59 
Sensillae bases distinctly in front of line of PL . . . . . ► . . 61 

59. A-P greater than PSB. Palpal claw trifurcate. Ventral setae on 
palpal tibia nude. Galeal setae branched. DS 38 arranged 2.6.6.6.6.6 
4.2. 

AW 71-6+4-9, PW 83-4+5-4, SB 30-0+3-0, ASB 30-0+3*3, 
PSB 23-6+2-2 t SD 53-5+4-9, A-P 30-7+5-0, AM 53-65+4-1, 
AL 38-4+3-4, PL 63-1+9-1, Sens. 42-6+8-2 with head 12-0 wide. 

Schon. (Ascoschon.) phascogale (Worn, and Heasp. 1943). 

A-P ca. equal to PSB , * 60 

60. Palpal claw trifurcate. Ventral seta of palpal tibia branched. Galeal 
setae branched. DS ca. 34 arranged 2.8.6.6.6.4.2. 

AW 82-9+.7-4, PW 99-9+7-3, SB 36-1+5-0, ASB 30-4+2-8, 
PSB 24-8+2-8, SD 55-2+4-6, A-P 26-5+5-2, AM 53-4+8-7, 
AL 48-1+6*3, PL 79-5+8-0, Sens. 42-0 by 14-0 wide. 

Schon. (Ascosclion.) westraliense (Worn, 1934). 

Palpal claw bifurcate. Ventral seta of palpal tibia branched. Galeal 
setae nude. DS 34 arranged 2.8.6.6.6.4.2. 

AW 79-3+4-3, PW 99-2+.4-1, SB 35-9+3-4, ASB 30-3+3-4, 
PSB 20-1+3-4, SD 50-4, A-P 25-2, AM 47-6, AL 40-6+7-0, 
PL 70-1+4-9, Sens. 44-8 with head 11-2 wide. 

Schon. (Ascoschon.) peregrina sp. n. 

61. Dorsal setae situated on small platelets. Leg I 6-segmented . * . ► 62 
Dorsal setae not on platelets. Leg I 7-segmented . . . . . . 63 

62. Only the dorsal setae on platelets. Palpal claw trifurcate; setae on 
palpal femur and genu branched. DS 50 in number, and arranged 
2.6.2.6.8.6.4.8.6.4.2, from 27 to 52^ long. 

AW 62-0+9-0, PW 78-0+7-0, SB 29-0, ASB 26-7+6-9, PSB 
23-7+1-7, SD 50-3+6-2, A-P 36-0+4-2, AM 46-0, AL 33-0, PL 
49-0, Sens. 39-3+1-7, with head 13-0 wide. 

Schon. (Ascoschon.) heaslipi (Worn, and Heasp. 1943) 

Some of the posterior ventral setae also on platelets. Palpal claw 
bifurcate e setae on palpal femur and genu nude. DS 30 in number, 
arranged 2.6.2.6.8.8.4, to 30/* long. 

AW 63*0+4-8, PW 75-95+2-1, SB 37-8+4-8, ASB 28-0, PSB 
25-0, SD 53-2, A-P 37-8 + 4-8, AM 42-0, AL 34-8+4-5, PL 22-4, 
Sens. 47-6 with head 6-0 wide. Schon. (Ascoschon.) trauhis^.u. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 241 

63. Dorsal setae 80 in number, arranged 2.12.12.1212.10.8.6.4.2, 

AW 77-6+6-2, PW 97-2+7-3, SB 31-4+2-8, ASB 30-1+6-7, 
PSB 27-1+5-8, SD 57-2+9-7, A-P 32-2+5-9, AM 57-0+9-5, 
AL 39-7+4-0, PL 67-2+5-0, Sens. 51-8+3-5, with head 12-0 
wide. Schon. (Ascoschfriu) Vawremei n. nov. 

for guntheri Worn. & Heasp. 1943, preoc. 

Dorsal setae fewer than 50 in number * * . ♦ . . .* . , 64 

64. Sens, bases nearer to anterior than to posterior scutal margin. AL 
the shortest and very short, A-P greater than PSB. 

AW 74-6+5-7, PW 102-3+4-4, SB 45-6+5-4, ASB 31-3+3 -4, 
PSB 41*0+4-2, SD 72*3+6-3, A-P 45-3+7-3, AM 43-2+4-5, 
AL 24-1+4-3, PL 51-2+5-4, Sens. 44-8 with head 16-8 wide. 

Schon. (Ascoschon.) ntongabelen-sisB.p.n. 

Sens, bases ca. midway between anterior and posterior scutal mar- 
gins. AL the shortest but not very short . . . . - . ► . 65 

65. Dorsal setae 46 in number, arranged 2.8.8.8.8.6.4.2. Scutum deep 
behind PL. A-P ca. equal to PSB. Galeal setae branched. 

AW 70-0, PW 81-0, SB 27-0, ASB 30-0, PSB 24-0, SD 54-0, 
A-P 29-7+7-6, AM 53-0+6-0, AL 34-7 + 9-1, PL 63-0+3-0, 
Sens. 43-0. Schon. (Asooschdn.) sndthi (Worn. 1939). 

Dorsal setae 28 to 32 in number. Galeal setae nude . . . . . . 66 

66. Dorsal setae 28 in number, arranged 2.8.6.6.4.2. Scutum evenly 
curved behind PL, with A-P slightly less than PSB. Palpal claw 
bifurcate. 

AW 62*3+4-2, PW 86-1+4-2, SB 30-8, ASB 25-2, PSB 25-2, 
SD 50-4, A-P 23-8+4-8, AM 39-2+6-8, AL 27-1+4-8, PL 41-3 
+8-1, Sens. 47-6 with head 8-4 wide. 

Schon. (Ascoschon.) raid sp. n. 

Dorsal setae 32 in number, arranged 2.6.6.6.6.4.2. Scutum medially 
flattened or lightly concave behind PL, with A-P greater than PSB. 
Palpal claw trifurcate. 

AW 56-2+7-4, PW 82-8+14-7, SB 29-5=fc6-3, ASB 29-5+4-4, 
PSB 20-2+2-38, SD 49-7+6-2, A-P 33-7+6-8, AM 40-9+5-3, 
AL 31-2+6-7, PL 51-25+8-3, Sens. 37-5+9-5, with head 14-0 
wide. Schon. (Ascoschon*) cairnsensis (Worn, and Heasp. 1943). 

AW 56-2+1 '05, PW 99-7+28*2, SB 35-8+3-8, ASB 31-9+4-6, 
PSB 25-2, SD 57-1+4-6, A-P 39-2+10-2, AM 42-7+8-0, AL 
31-2+3-8, PL 55-3+15-9, Sens. 39*2 with head 14-0 wide. 

Schon, (Ascoschon.) cairnsensis v. gateri (Worn, and Heasp. 1943). 



242 Records of the S.A. Museum 

Genus BADFORDIANA no v. 
As in SchSngastia s. str. but with the chelieerae unusually long, serrated, 
and almost styliform, and the hypostome long and tongue-like. The palpi are 
long and slender with the tibial claw simple. 

Genotype Radfordiana rostrata sp* n. 

Radfordiana rostrata sp. n. 
Plate 66, fig. A-E. 

Description of Larvae. Shape broadly oval. Length (unengorged) 300/*, 
width 215/t, Scutum as figured with anterior margin only indistinctly sinuous 
between AL; PL nearer to AL than to posterior margin which is deep and 
broadly rounded behind line of PL ; sensillae globose, only indistinctly villous 
and placed behind PL; scutel setae long, slender and ciliated, with AL the 
longest Eyes 2 -f- 2, posterior the smaller. Chelieerae very long and slender, 
almost styliform, to 115/* long, with ca. 10-12 fine teeth on distal half of inner 
margin. Hypostome long and tongue-like, with the galeal setae nude. Palpi 
long and slender, tibial claw simple; setae on femur and genu shortly but 
thickly ciliated; on tibia ventral only shortly branched, dorsal and lateral 
nude; tarsi with pronounced subapical spine-like seta, sub-basal sensory rod, 
and 4—5 fine slender ciliated setae. 

Dorsal setae stout, strongly ciliated, to 4f)jx, 38 in number, and arranged 
2.8.8.8.6.4.2. Ventrally with a pair of branched setae on maxillae, one ciliated 
seta on each coxa, a pair between coxae I and between coxae III, and there- 
after 6.6.6.2.2.2, to 30fi long. Legs; I 360/j- long, II 305/x, III 335/*; tarsi I 
and II with the usual sensory rod, III with a long outstanding nude seta. 

The Standard Data for the type and one paratype from scrub, at Lae, 
New Guinea, April, 1944 (R. N. McCulloch), six specimens from mound of a 
bush turkey (Megapodhu) at Dobodura, N.G., June, 1944 (G. M. Kohls), and 
4 specimens from an M armed lizard", Hollandia t Dutch N.G., Nov., 1944 (C* B. 
Philip) are as follows: 



AW 


Mean 
61-4±0-88 


Standard 
Deviation 

3-03±0-62 


Theoretical 
Range 

52-3-70-5 


Observed 
Eange 

56-0-64-4 


Coeff . of 
Variation 

5-0 


PW 


87-7=fcl-21 


4 -19:4:0 -86 


75-1-100-3 


78-4-95-2 


4-8 


SB 


24-7±:0-47 


l-62±0-33 


19-9-29-5 


22-4-28-0 


6-5 


ASB 
PSB 


40-8±0-60 
30-8 


2-06±0-42 34-6-47-0 
No variation recorded 


36-4-42-0 


5-0 


SO 


71*6:±0*5S 


1-87+0-38 


6(5 -0-77-2 


67-2-72-8 


2-6 


A-P 


30- 1±:0 » 61 


2-11+0-43 


23-8-36-4 


25-2-33-6 


7-0 


AM 


46-S±l-02 


3-37:±0-72 


36.7-56-9 


42-0-50-4 


7-2 


AL 


69-8±0-80 


2-79:£0-57 


61-4^78-2 


64-4-72-8 


4-0 


PL 

Sens. 


56*6±:l-00 3-46:±0-70 46-2-67-0 
42-0 with head 16*8/22-4. No variation recorded. 


50*4-61-6 


6-0 



Womersley— Asiatic-Pacific Scrub Typhus Mites 243 

Remarks. It is possible that Eadforcltana may not stand as more than a 
subgenus of Schongastia s. str. but it is so distinct that until the nymphs or 
adults are known it should be kept at the generic level. 

Genus OENOSCHOENGASTIA Worn, and Kohls 1947. 

Trans. Roy. Soe. S. Aust., 1947, 71, (1), 8-9. 

Allied to Schdngastia but the ehelieerae are short, curved and stumpy with 
the apex divided into two terminal blunt and thick teeth, with a pair (some- 
times three) of rather smaller teeth subapieally. Palpal claw short, and stout, 
trifurcate. Median tarsal claw (empodium) longer than the laterals but 
equally thick. Sensillae clavate or capitate. 

Genotype Oetwschongastia cana Worn, and Kohls 1947. 

The nymphal and adult stages of this species are so far unknown. Until 
such times its relationships to the genus Schdngastia are uncertain, but in view 
of the unique characters of the larvae its retention as a distinct genus is 
necessary. 

Oenoschongastia oana Worn, and Kohls 1947. 

Oenoschongastia- cana Womersley and Kohls 1947, Tr. Roy. Soc. S« Aust., 71, 
(l),8-9. 

Plate 66, fig. F-J. 

Descriptwn of Larvae. Shape an elongate oval. Length (unfed) 252^, 
width 162^.. Dorsal scutum roughly hexagonal with indistinct and fine pitting; 
with the usual five normal dliated setae, of which AL are the longest and AM 
the shortest, AM with short branches, AL and PL with long outstanding 
branches; sensillae broadly clavate, the head indistinctly and very shortly setu- 
lose, sensillae bases about in line with PL; anterior scutal margin convex, 
posterior laterally angular. Eyes 2 + 2, large, on well-developed ocular shields 
and closely adjacent to scutum, posterior eyes the smaller. Chelicerae of 
peculiar form, short, stout and curved, apex truncate and divided into two 
strong, broad and blunt teeth, and two (sometimes three) smaller teeth placed 
subapieally on the inner edge. Galea! setae nude. Palpi stout, tibial claw 
short, stout and trifurcate ; femur with a long strong seta with long branches ; 
genu with a nude seta; tibia with the dorsal and lateral setae nude, ventral 
branched ■ tarsi short with basal and subapical sensory rods and three or four 
•ciliated or branched setae. Dorsal setae strong with strong eiliations, 36 in 
number and arranged 2.8.10.6.6,4, to 40-50/x long. Ventrally with the usual 
pair of branched setae on gnathosoma, a single seta on each coxa, a pair between 
coxae I and between coxae III, and thereafter 6.6.6.4.2, more slender and with 



244 Records of the S,A. Museum 

longer dilations than the dorsal setae, to 30/a long. Legs \ I 324/* long, II 252ft; 
III 380//.; tarsi I and II with the usual dorsal rod-like setae, tarsi III with a 
long outstanding nude seta; median claw (empodium) longer than but as thick 
as the laterals. 

The Standard Data in microns derived from 22 specimens are : 





Mean 


Standard 
Deviation 


Theoretical 

Range 


Observed 
Range 


Coeff , of 
Variation 


AW 


62-85±0-56 


2-.65±0'40 


54-9-70-8 


60-0-69-0 


4-2 


PW 


78-7:±U-71 


3-34±0-50 


68-7-88-7 


75-0-87-0 


4-2 


SB 


31-85±0-49 


2'30±0'35 


24-95-38*75 


30-0-39-0 


7-2 


A8B 


28*9:±0'33 


l-53±0-23 


24-3-33-5 


27-0-32-0 


5-3 


P8B 


23-8±0«30 


l-38±0-21 


19-G-28-0 


21*0-27-0 


5-8 


SD 


52-8±:0'45 


2*ll±:0-32 


46-5-59-1 


48-0-57-0 


4-0 


A-P 


29-2±0*32 


l-50:£0'23 


24*7-33-7 


27-0-33-0 


5-2 


AM 


32-45±0-46 


2-15±:0«32 


26-0-38-9 


27-0-36-0 


6-6 


AL 


74^3±0-50 


2-34±:0*35 


67-3-81-3 


70-0-80-0 


3-0 


PL 


59-5±0-39 


1 • 84:10 • 28 


54-0-65-0 


55 -0-65 «0 


3-1 



Sena* 36-0 with head 20/24. 

Lac. The type and 72 paratypes from the mound of a brush turkey, 
Dobodura, New Guinea, 18 May, 1944 (G.M.K., No. 325). 

The type and 22 paratypes deposited in the South Australian Museum, 35 
paratypes in the collection of the Rocky Mountain Laboratory, five paratypes 
in the U.S. National Museum and five in the British Museum. 

Genns GTJNTHERANA Worn, and Heasp. 1943. 
Oimtherana Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust, 67, (1), 132, 
nom. nov. for Gunthena Womersley 1939, ibid, 63, (2), 157 (preoc). 

Larval diagnosis. Body form (unengorged) oval, (engorged) elongate oval 
■with a distinct medial constriction. Posteriorly with an area which, when en- 
gorged or under pressure, consists of two semicircular plates, opposed in the 
medial longitudinal line, strongly punctate, and each bearing 3 fine setae; api- 
cally the junction of these plates is slightly incised. In unengorged larvae the 
two plates frequently appear as one large transversely oval plate. Anterior 
dorsal scutum roughly rectangular, with 5 normal ciliated setae and a pair of 
globose or capitate, indistinctly villous sensillae. Eyes 2 -f- 2. Chelicerae non- 
serrate with only the apical tricuspid cap. Galea! setae nude. Palpi stout, 
tibial claw trifurcate. All coxae l-setose- Legs III without any long nude 
seta on tarsi. 

Adult and Nymph (see Adult Section of Paper). 

Genotype Neoschongastia bipygalis Gunther 1939. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 245 

Guotherana bipygalis (Gunthcr 1939)* 

Neoschongasiia callipygea Gunther 1938, nam. nud., Med. J. Aust., 2, (6), 202. 

Neoschongastia hallipygos Gunther 1939, nom. nud., Proc, Linn. Soc. New South 
Wales, 64, (1-2), 83. 

Neoschongastia bipygalis Gunther 1939, Proc. Linn. Soe. New South Wales, 
64, (5-6), 471. 

Guiitheria kallipygos Womersley 1939, Tr. Roy. Soc. S, Aust., 63, (2), 187. 

Guntheria oipy gaits Gunther 1940, Proc. Linn. Soc. New South Wales, 65, 250 ; 
idem., Med. J. Aust., 2, (22), 564r-72. 

Guntkerana bipygalis Womersley and Heaslip 1943, Tr. Roy. Soc. S, Aust., 
67, (1), 100. 

Guntheram parana, Womersley 1944, Tr. Roy. Soc. S. Aust, 68, (1), 106. 

Plate 67, fig. A-H. 

This species was originally described from specimens from Rattus hraurni 
Alston, R. ri?igen$ Peters and Doris, R. mordax Thomas, Melomys mancktani 
Thomas, M. stalkeri Thomas, M. r-iihex Thomas, M. sp., Echymipera cockerelli 
Ramsay, and Peraryetes raffrayana Milne Edwards from New Guinea hy 
Gunther. Although Gunther later (1939, 64 (5-6)), figured the young larva 
within the ovum, his original description and figures were from free and 
engorged larvae. Although Gunther states (loc. tit. 471) that "specimens 
(larvae) also have been taken running free in the fur" he does not say whether 
these are engorged or not. 

In 1939 Womersley recorded the species from engorged specimens from 
Queensland from R. ciilm&rum youngi Thomas and from Isoodon torasus 
Ramsay. 

Guntherana parana- was described in 1944 by Womersley from unengorged 
larvae collected on boots at Abidari and Buna by R. N. MeCulloch. These 
larvae were considered as distinct from hipygalis Gunther, on the smaller 
Standard Data, the lack of a medial constriction, and the larger number of 
dorsal and ventral setae. In other generic features such as the posterior dorsal 
shield, (or pair of shields) and the two very long dorsal setae just anterior of 
the dorsal shield, they fitted well into the genus. 

Gunther 's original description and figure of the dorsal surface show the 
dorsal setae as 2.6 (2 or 4). 4.2A2./6./2, i.e. 28 (30 or 32). Actually there are 



246 Records of the S,A. Museum 

only 26, the last 2 strictly belonging to the ventral surface. The six posterior 
setae are the fine setae on the paired posterior dorsal plates, while the next 
pair are the very long closely adjacent setae just in front of the posterior 
plates. In parana the dorsal setae are 26, arranged 2.6.4.6 (4). 2 (4), plus 6 fine 
setae, on the posterior plates. 

Ventrally for bipygalis, Gunther gives the setae posterior of coxae III as 
4.2./4.4 or 14 in all. A careful examination of his type larva (engorged) in 
the South Australian Museum, shows however, that there are more setae than 
Gunther saw, namely 24, arranged <?a. 6.6./6A2. Probably some of the setae 
had been rubbed off and not observed by Gunther. This is the case with other 
specimens which I have examined. The setae bases can, however, be usually 
traced. This larger number of ventral setae is the same as in parcma, so that 
the only difference left between this species and bipygalis is the somewhat 
smaller scutum and the lack of the medial constriction. As the constriction 
may be a result of engorgement, (although it is just indicated in Gunther 's 
figure of the unhatched larva), it would appear best to regard parana as a 
possibly local population of bipygalis differing only in the lower Standard 
Data. 

It should be noted, however, that Gunther 's finding of the eggs of bipy- 
galis attached to the hairs of the host suggests that the young larvae before 
attachment for feeding, may be free living in the fur and not on the ground. 
Yet the specimens described earlier as parana as well as other unengorged 
larvae from New Guinea and Queensland, which on Standard Data agree more 
with typical bipygalis, were collected on boots. This is a point which needs 
further investigation, especially in regard to affording confirmation of Gun- 
ther ? s observations. 

The nymph and adults of this species are described in the Adult Section 
of this paper, and the genus in those stages defined. The nymph was reared 
from engorged larvae. The adult was correlated with larvae reared therefrom, 
and which at the time were regarded as parana. 

Of the material in the South Australian Museum (32 specimens of larvae), 
there is a high degree of variation in the Standard Data, and in the length of 
the pair of long setae just in front of the posterior dorsal plates. 

In the Standard Data the 7 specimens of unengorged larvae from Abidari 
and Buna, New Guinea, collected by H. N. McCulloch 1943, the data for AW, 
PW, SB, ASB, PSB. SD and A-P are significantly lower, and there is com- 
paratively little variation in PL. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



Z\l 



The Standard Data for these specimens are : 



AW 

PW 

SB 

ASB 

PSB 

SD 

A-P 

AM 

AL 

PL 

Sens* 



Mean 
47-2±0-73 
65*2:±0-52 
16*8 

23*2±0-52 
11-2 

34-4±0-52 
28-0 
33-6 

69-6±0-91 
95-6±l*28 



Standard 
Deviation 

1-93+0-52 

1-37+0-36 



Theoretical 
Range 

41- 4-53-0 

61-1-69-3 



No variation recorded 
1*37+0-36 19-1-27 

No variation recorded 
1*37+0*36 30-3-38 

No variation recorded 
No variation recorded 
2-52+0-67 62-1-77- 



3*40+0*91 



85*4-105-8 



Observed 
Range 

44*8-50-4 
64-5^-67-2 

22-4^25-2 
33-6-36-4 



64-4-72-8 
89-6-98-0 



Coeff. of 
Variation 

4-1 
2-1 

5-9 

4-0 



3-6 
3 6 



33*6 with head 16-8/19-6. No variation recorded. 



The Australian material from Queensland, plus the type and one paratype 
from Bulolo, New Guinea, show distinctly larger Standard Data, and the 
•values as given by "Womersley and Heaslip (1943) treated statistically are as 
follows : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 
PW 


57-5+0-95 
77-85+1-54 


4-12+0-67 
6-72+1-09 


45-1-69-9 
57-7-98-0 


47-0-65-0 
68-0-92.0 


7-2 

8-6 


SB 


19-75+0-41 


1-78+0-29 


H-4^25-1 


17-0-23-5 


9-0 


ASB 


23-5+0*28 


1*18+0-19 


20-0-27-0 


21-0-26-0 


5-0 


PSB 


12*8+0-24 


l-01it0-17 


9-8-15-8 


11-0-16-0 


8-0 


SD 


36-45+0*42 


1-80+0-30 


31-05-41-85 


33*0-42-0 


4-9 


A-P 


28-8+0-58 


2-44+0-41 


21-5-36-1 


26-0-34*0 


8-5 


AM 


32*0+0-46 


1-66+0-32 


27-0-37-0 


30-0-34-0 


5-0 


AL 


73-3+0*96 


3-84+0-68 


61-8-84-8 


64-0-77-0 


5-2 


PL 


97-2+1-66 


6*86+1-18 


76-6-117-8 


80-0-113-0 


7-0 


Sena. 


31*0 with head 15-0 wide. 









Gtenus NEOSCHOENGA8TIA Ewing 1D29. 

Manual of External Parasites 1929, 187. Genotype Schongastm americawai 
Hirst 1921, Ann. Mag-. Nat. Hist, (9), 7, 37, (larvae). 

— Paraschongastia Womersley 1939, Tr. Roy. Soc. S. Aust, 63, (2), 165 
Womersley and Heaslip 1943 ibid., 67, (1), 129, (larvae). 

Neoschdngastia Wharton and Hardcastle 1946, J. Parasit., 32, (3), 286-322, 
(larvae and nymphs) . 

Ewing erected this genus for Hirst's Schon-gastia america-m principally 
on the chelicerae not being serrate on the inner (dorsal) margin but also on 
the palpal claw being trifurcate and the dorsal scutum being "poorly chitinized 
and tending to break up into smaller platelets". Hirst in his description of 



248 Records of the S.A. Museum 

americana (loe. eit.) says, ct Scutum not so definite in shape as in known 
species of Schongastia, being weakly cnitinizecl; posteriorly it is interrupted 
in the middle, being practically divided into two portions or wings by a series 
uf (median) longitudinal parallel striatums". 

Although no figure was published with the original description, a very 
excellent one showing the above peculiar structure of the dorsal scutum is 
shown in the pamphlet f 1 Mites Injurious to Domestic Animals/' by Hirst 
(B,M. (N.H.) Econ. Ser, No. 13. 1922, p. 60). 

Although Ewing, as stated above, notes the peculiar scutal structure, he 
evidently did not regard it as of much generic importance, for in his later 
publications he included in Neoschdngcustia many species with non-serrate 
•chelieerae but with a normal scutum ; particularly did he do so in his key to 
the larval genera of the Trombiculinae of 1938 (J. Wash. Acad. ScL, 28, (6)). 

In 1939 Womersley recognized two groups within the genus- Neoschan- 
gastici Ewing, and separated off those New Guinea species of Gunther's, in 
which the scutum had a "distinct raised crest in front of the psuudostigma." 
which "forms a very distinct wall in which the bases of the sensillary hairs 
are situated", and "the posterior half of the scutum on each half has circular 
striatums but the anterior half is pitted", as a new genus Parasckonga^tia. 
Later 1943, Womersley and Heaslip designated Neo$ehdnga$ti<i yeomonsi Gun- 
ther 1939, as type of ParascMngastia. 

At the time, however, it was not recognized that these characters were also 
those of N. ameticana and it was only in 1946 that Ewing recognized that 
N. amerwana and P. yeomansi were congeneric. The present writer is in full 
agreement with, this, and accepts Paraschongastia. as A synonym of Neoschon- 
gastia. 

This position leaves all those species with a normal scutum and non-serrate 
ehelicerae hitherto placed by Ewing and by Womersley and Heaslip 1943, etc., 
in Neoschdngastia f in need of a new generic name, for which Ewing 1946 has 
proposed Ascoschomgastia with Neoschongastia malayensis Gater as the type 
species, Wharton 1946, in recently discussing the genus Neoschonflmtia has 
further stressed the striations on the dorsal scutum as being of generic import- 
ance. He points out that these striations are those of the dorsal cuticle, the 
posterior portion of the scutum being overlapped by the cuticle. This over- 
lapping, however, is not -confined to the genus Neoschongastia. As instanced 
in the present paper, it also occurs in a few species of 8chdnga<stia (AseoscJwn~ 
gastia), but here the striations are transverse, whereas in Neoschdngastia they 
are more or less circular and thus differentiated from the normal dorsal 
striations. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 249 

Neoschongastia bougainvillensis Wharton and Hardcastle J 946. 
J. ParasitoL, 1946, 32, (3), 296 fig. (larva). 

Plate 74, fig. E. 

This species was described from Hincndo tahitica from Bougainville, 
July, 1944, and from Anous stolidus and Heterocelus incmius from Guam, May 
and September respectively, 1945. It is differentiated from all other species by 
the peculiar rounded elevation on the basal segment of the chelicerae and by 
the broadly clavate sensillae fitting into a well marked pair of pits posterior 
of the sensillae bases. 

The galeal setae are nude, and on the palpi the femoral and genual setae 
are strongly branched; the tibia with the dorsal seta nude, the lateral nude or 
with an indistinct branch, the ventral distinctly branched. The scutum is as 
figured (after "Wharton and Hardcastle) with the lateral margins strongly 
concave and the posterior margin moderately shallow but concave medially. 
The DS are ca. 90 in number and from 37^ long anteriorly to 33/* posteriorly, 
in rather indefinite rows. All coxae unisetose. Ventral setae 10-30/* long, 
about 80 in number posterior of coxae III, Tarsus III without any long nude 
seta. 

The Standard Data from Wharton and Hardcastle 's data for 4 specimens 
treated statistically are ; 



AW 


Mean 
58>75±l-75 


Standard 
Deviation 

3-502:1-24 


Theoretical 
Range 

48-25-69-25 


Observed 
Range 

55-0-63-0 


Coeff. of 
Variation 

6-0 


PW 


87-75±l-93 


3-86+1-36 


76-2-99-3 


84-0-92-0 


4-4 


SB 


29«25±1-11 


2 -222:0 -78 


22 '6-35* 9 


27*0-32-0 


7-5 


ASB 


29-75±0.63 


l-26±0-44 


25-95-33-55 


28-0-31-0 


4*2 


PSB 


26-02:1-18 


3-372:1-68 


15-9-36-1 


24-0-31-0 


12-9 


SD 


55-75±l-18 


2»36±0-8S 


48-05-62-85 


54-0-59-0 


4-2 


A-P 


41-75±2»01 


4-032:1-42 


29-65-53-85 


36-0-45-0 


9-6 


AM 


60 -752:1 -65 


3-30±t>17 


50-35-70-15 


56-0-64-0 


5-5 


AL 


60«5±0-29 


0-572:0*20 


58-8-62-2 


60-0-61-0 


0-95 


PL 


60-25±0-25 


0-50±0-25 


58-75-61-75 


60-0-61-0 


0-8 


Sens. 


35-5:£0-96 


1-912:0 -68 


29 '8-41- 2 


34-0-38-0 


5-4 



Neoschongastia strongi Wharton and Hardcastle 1946. 

J. Parasitol., 1946, 32 (3), fig. (larva). 

Plate 75, fig. E. 

Described from larvae from Numenius phaeopus from Guam, July, 1945, 
and from Arenaria interpret from Peleliu, September, 1945. Distinguished 



250 Records of the 5.A. Museum 

from all other species by the nude seta on the palpal genu, the relative close- 
ness of the sensillae bases and the sensillae themselves lying in depressions an- 
terior of the sensillae bases. Galeal setae nude, as are also the dorsal and lateral 
setae of the palpal tibia. 

The dorsal setae are from 35/a to 17 p long, with a pair of humeral setae 
plus ca. 70 in irregular rows. Coxae III unisetose. Tarsus of leg III with a 
long nude seta. 

The Standard Data given by Wharton and Hardcastle for 3 specimens, 
treated statistically are: 





Mean 


Standard 
Deviation. 


Theoretical 
Eange 


Observed 
Range 


Coeff . of 
Variation 


AW 
PW 


40-0±1-0 
65-3±l*2 


l-73±0-71 
2-08±0-85 


34-8-45-2 
59 -1-71- & 


39-0-42-0 
63-0-67-0 


4-3 
3-2 


SB 


13-0±lO5 


2-00±0-82 


7-<KL9*0 


11 -0-15-0 


15-4 


ASB 


'63 -7 ±0-67 


1-15+0 -47 


30-2-37-2 


33-0-35-0 


3-4 


PSB 


14-3-b0-67 


l-15±0-47 


10-8-17-8 


13-0-15*0 


8-0 


SD 


48- 0+1-15 


2-0+0-82 


42-0-54-0 


46*0-50-0 


4-2 


A-P 


34-3±0-88 


1-53+0-62 


29-7-38-9 


33-0-36-0 


4-4 


AM 


24<3±l-45 


2*52+1-03 


16 -8 -31 -8 


22-0-27-0 


10-3 


AL 


36-7±l-20 


2-08+0-85 


30-5-39-0 


35-0-39-0 


5-7 


PL 


38-7±3-18 


5-51+2-25 


22-2-55-2 


35-0-45-0 


14-2 


Sens. 


21-5±0-5 


0-71+0-35 


19-4r-23*0 


21-0-22-0 


3-3 



Neoschongastia monticola Wharton and Hardcastle 1946. 
J. Parasitol., 1946, 32, (3), 301, fig. (larva). 

Plate 75, fig. D. 

Described from Monticola soUtarius from Okinawa, from June to August, 
1945. 

In this species the form of the sensillae is unusual for this genus, these 
being very long with a long and clavate attenuated head, furnished with long 
fine setules. The scutum is as refigured here, after Wharton and Hardcastle, 
with PL very much longer than AM or AL. The galeal setae are short and 
nude. On the palpi the femoral and genual setae are branched as is the ventral 
tibial; the dorsal and lateral tibial setae are nude. 

The DS setae number 38 to 40, are from 62/* to 39/4 long, and arranged 
ca. 2.8.6.6.6.6.4.2. All coxae unisetose, only 2 sternal setae and posterior of 
coxae III about 50 setae. A nude long seta on tarsi III. 

The Standard Data, treated statistically for the measurements of 5 speci- 
mens given by Wharton and Hardcastle are : 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



251 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


55-8±l-2 


2-68±0-85 


47-8-63-8 


54*0-60-0 


4-8 


PW 


77-8±:2-0 


4-38±l-38 


64-7-90-9 


72-0-82-0 


5'6 


SB 


32-4±0-S2 


l-83±0-82 


26-9-37-9 


30-0-34-0 


5-7 


A8B 


24-6±0-60 


1-34:2:0-42 


20*6-28-6 


24-0-27-0 


5-4 


PSB 


26-4±l-25 


2-79±0-88 


18- 0-34- S 


24-0-31-0 


10-6 


SD 


53-0±l-82 


4-06±l-28 


38*8-63-2 


48-0-58-0 


7-9 


A-P 


27-2±0-20 


0-45±0-14 


25-9-28-5 


27-0-28*0 


1-6 


AM 


42-6±2-77 


6'19drl-96 


24-1-61-1 


36-0-49-0 


14-5 


AL 


35-0±0-83 


l-87±0-59 


29*4-40-6 


33-0-37-0 


5-3 


PL 


75-2±2-80 


6-26±l-98 


56-4-94-0 


66-0-82-0 


8-3 


Sena. 


70-75:±2-80 


5-68±2-01 


53-75-87-75 


63- 0-75-0 


8-0 



Neosohongastia posekanyi Wharton and Hardcastle 1946. 

J. ParasitdL, 1946, 32, (3), 302, fig. (larva). 

Plate 75, fig. A. 

Described from Monticola* solitarius and Sireptopelia arientalis from Okin- 
awa, May and July, 1945. 

This species can be distinguished as in the key. As stated by the above 
authors it is close to goXUnarum but can be at once distinguished by the long 
nude seta on tarsi III. 

The dorsal scutum is roughly rectangular and rather long, with the mar- 
gins lightly sinuous ; AM is much stronger and with longer branches than AL 
or PL, but shorter than these. The sensillae are globular with the tip rather 
truncate, and their bases much nearer to AL than to PL. 

The galeal setae are nude, and on the palpal femur, genu and tibia only 
•the dorsal tibial is nude. All coxae unisetose. The dorsal setae number 34, 
from 52/a to 43//, long, and are arranged 2.8.6.8.4.2.2.2. Posterior of coxae III 
the ventral setae are 6.6.2.6.2.2. 

The Standard Data, treated statistically from 5 specimens, quoted by 
Wharton and Hardcastle are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


75-2±l-46 


3-27:±l-03 


65-4-85-0 


70-0-78-0 


4-3 


PW 


79-8:fcl-33 


2*97±;0.94 


70-9-88-7 


77-0-84-0 


3-7 


SB 


47-4±l-36 


3-05+0-96 


38-3-56-5 


45-0-52-0 


6-4 


ASB 


25-4±l-03 


2-30=£0-73 


18-5-32-3 


22-0-27-0 


9-0 


PSB 


33-6±l-l7 


2-61±:0-82 


25-8-41-4 


29-0-35-0 


7-7 


SD 


59 -Oil '34 


3-00±0-95 


50-0-68-0 


56-0-63.0 


5-1 


A-P 


29 -4±0- 68 


1-51^0-48 


24-9-33-9 


28-0-31-0 


5-1 


AM 


49-0±l-14 


2-55±0-Sl 


41-4-56-6 


45-0-52-0 


5-2 


AL 


74-8:±4-42 


9-88±3-l2 


45-2-104-4 


66-0-91-0 


13-2 


PL 


52-4±l-57 


3-51ztl-ll 


41-9-62-9 


49-0-56-0 


6*7 


Sens. 


29-75±l-61 


3-59±l-13 


18-95-40-55 


27-0-350 


12-0 



252 Records of the S.A. Museum 

Neoschongastia gallinabum (Hatori 192/)). 

Trombicula gaUinarum Hatori 1920. Taiwan Igakkai Zasshi, No. 209, pi. II, 
fig. 1-6 j Kawamura and Yamaguchi 1921, Kitasato Archiv, Exper. Med. 4, 
169; Fletcher, Lessler and Lewthwaite 1928, Tr. Hoy. Soc. Trop. Med. 
Hyg., 22, 16L 

Neoschdn&cwtia gaUinarum, Sugimoto, 1936, J. Jap. Soc. Veterinary Sci., 15, 
201; Wharton and Hardcastle, 1946, J. Parasitology, 32, (3), 292. 

Paraschongastia gallinanim, Womersley and Heaslip 1943, Tr. Roy. Soc. 

S. Aust., 67, (1), 130. 
Schdngastia gaUinarum, 1947, Sig Thor and Willmann, Das Tierreich, 71b, 3/J4. 

Plate 69, tig. A-E. 

This specific name was attributed by Kawamura and Yainagnehi to Hatori 
for a "red mite'' from fowls in Formosa. At the time of publishing their 
paper in 1943, Womersley and Heaslip had been unable to find any reference 
to the. use of this specific name, or of any figures or description by Hatori, and 
therefore gave Kawamura and Yamaguchi as the author. Sugimoto, 1936, in 
his paper gives the above reference to Hatori 's paper which appears to have 
contained figures and description. I have, however, not been able to obtain 
Hatori 7 s paper, but from the evidence it appears that gcMmarmn must be attri- 
buted to Hatori himself. 

Sugimoto (1936) described and figured the larvae also from Formosa. 

FJetcher et at. (1938) referred to a "red mite 7 ' of fowls and other birds 
from the Federated Malay States, but did not give a description and only 
figured the dorsum and dorsal scutum. Their figures, however, compared with 
those of Kawamura and Yamaguchi and of Sugimoto are not very convincing 
as to the identification. Kawamura and Yamaguchi gave the following details, 
of the larvae: scutal length 42/*, scutal width 65-1/*, sensillae 22-5/* long with 
head 14-5 X 14-5^ and strongly ciliated; DS 33/* long and arranged 14.10.6.8.6 
(i.e. 44 in number). 

From their figure, however, the DS are 2.11.9.8.6.2 or 38 in number. 
Sugimoto shows them as 2.10.8.8.6.6.2 or 42 in number. 

The Standard Data computed from Kawamura and Yamaguchi y s figures 
and data in 1943 by Womersley and Heaslip were: AW 52-0, PW 60-0, SB 
35-0, ASB 16-5, PSB 26-5 7 SD 43-0, A-P 31-0, AM 32-0, AL 35-0, PL 50*0 r 
Sens. 22-5 with head 14-5/14-5. 

Fletcher ei al. 193S give the DS as 2.12.10.8,6.4.2 (=44) and from their 
figures the Standard Data were computed (1943) as: AW 52-0, PW 60-0, SB 
42-0, ASB 19-0, PSB 26-0, SD 45-0, A-P 30-0, AM 34-0, AL 35-0, PL 33-0, 
Sens. 26-0. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 253 

This speeies was placed by Woniersley and Heaslip 1943 in Paraschongastia 
( <*m Neoschongastia) on the presence of the pronounced transverse crest from 
which the sensillae arise. None of the figures extant, however, shows the 
typical cuticular striations of this genus. 

The type and 19 paratypes are stated by Sugimoto (1936) to be in the 
Taihoku Imperial University Museum, Taihoku. 

Recently at my request, Dr. J. R. Audy has recovered this species from 
fowls at Kuala Lumpur, F.M.S., and from 4 specimens sent to me, the follow- 
ing redeseription and figures are given. These specimens agree very well with 
the details and figures given by Fletcher et al. 1928, Kawamurra and Yamaguchi 
1921, Sugimoto 1936, and Hatori 1920. There is no doubt but that the Kuala 
Lumpur specimens are identical with Hatori's Forraosan material, 

Be-descripti&n- of Larvae. Shape broadly oval, widest anterior of the 
middle. Length (engorged) to 390/*, width to 320/x. Scutum rather small, 
somewhat trapezoidal, with sensillae wide apart in a distinct crest and 
placed nearer to line of AL than to PL ; AM scutal seta the shortest, AL and 
PL subequal with PL slightly the shorter; sensillae globose and setulose; pos- 
terior half with semicircular striations, anterior half punctate. Eyes well 
developed, on ocular shields; posterior eyes the smaller. Chelicerae simple 
with only the apical trieuspid cap, and an inner subapieal forwardly directed 
tooth. Palpi with trif urcate tibial claw ; setae on femur, genu and tibia all 
ciliated or branched. Galeal setae ? nude. Dorsal setae 46 in number, 
arranged 2.10.8.6.8,6,4.2, to 33/i long. Ventrally with paired branched setae 
on maxillae, a pair between eosae 1 and between coxae III, a single fine and 
short ciliated seta on each coxa, and behind coxae III 40 setae, arranged 
8.8.6.4.6.6.2, to 30/* long. Legs: I 240/* long, II 200/*, III 227/*; tarsi I and 
II with usual dorsal sensory rod, HI with a long nude seta. 

The Standard Data derived from the 4 recent specimens from Kuala 
Lumpur are ; 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


55-3±0-70 


l-40:£0-49 


51 -1-59 -5 


53-&-56-0 


2-5 


PW 


67-9+0-70 


1-403:0 -49 


63 -7-72.1 


67-2-70-0 


2-0 


SB 


42-0 


No variation recorded 






A8B 


19-6 


No variation 


recorded 






PSB 


25-2 


No variation recorded 






SD 


44-8 


No variation recorded 






A-P 


28-7±0-70 


i-40HrO-49 


24-5-32-9 


28-0-30-8 


4-9 


AM 


29»9;t0-93 


l-61±0-66 


25-1-34-7 


28-0-30*8 


5-4 


AL 


45-5:±:G-70 


l-40±0-49 


41-3-49-7 


44-8-47-6 


3-1 


PL 


45-7±l-34 


2-69zfc0.95 


37-6-53*8 


42-0-47-6 


5*9 


Sens. 


33-6 with heai 


I 19-2/22-4. Ni 


o variation recordec 


[. 





254 Records of the S.A* Museum 

Neoschongastia backhousbi Gunther 1939, 
Neoschongastia fournieri Gunther 1938 (nam. mud.), Med. J. Aust., 2, 202. 
Neoschongastia lackhousei Gunther 1939, Proc. Linn. Soc. New South Wales, 

64, (1-2), 89; Wharton and Hardcastle 1946. J. Parasitol., 32, 294. 
Paraschongastia hackkousei Woinersley 1939, Tr. Koy. Soc, S. Aust., 63, (2), 

165; Gunther 1940, Proc. Linn. Soc. New South Wales, 65, (3-4), 252; 

Womersley and Heaslip 1943, Tr. Roy. Soc. S, Aust., 67, (1), 130. 

Plate 69, fig. F-J. 

No fresh material of this species has been collected since the original dis- 
covery. The palpal claw is much more slender than in yeomansi and well sur- 
passes the tip of the palpal tarsus. The setae on the palpal femur and genu 
are well branched but not so much as in yeomtmsi', on the tiiba the ventral 
seta is branched, the dorsal nude and the lateral one- or two-branched, not nude 
as stated by Gunther. The chelicerae are probably much as in yeomansi and 
other species, but Gunther fr figure is not -clear and in the type specimen they 
are broken off. The galeal setae are short and branched as figured here. On 
the dorsum the setae, are. uniform in structure without any specialization 
posteriorly. 

Coxae III is unisetose. Leg III is missing. 

The Standard Data as given in 1943 (loe. cit.) are: AW 58-0, PW 72-0, 
SB 48-0, ASB 23-5, PSB 25-0, SD 48-5, A-P 42-0, AM 37-5, AL 47-0 ? PL 
42 «0, Sens. 30-0 with head 15/20. 

Loc. Bulolo, New Guinea, ex Megapodius duperreyi 1939 coll. C. Gunther 

Neoschongastia egketta Wharton and Hardcastle 1946. 
J. Parasitol., 1946, 32, (3), 304, fig. (larva). 

Plate 74, fig. G. 

Described from Egretta intermedia from Okinawa, August, 1945, and from 
E, intermedia, Dewdgretta saorck, Anous stolidus and Pluvialis dominica from 
Ulithi Atoll, August, 1945. 

This species differs from iackh&itsei Gunther in the posterior dorsal setae 
not arising from platelets and in lacking the posterior dorsal pitting. 

The galeal setae are branched, and the setae on the palpal femur and genu 
and the ventral on the tibia are branched. The scutum is as figured from 
Wharton and Hardcastle. The sensillae are globose with rather a thick and 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 255 

relatively long pedicel and the heads lie in rather ill-defined pits posterior of 
the sensillae bases, and these pits are surrounded by the cuticular striations. 

The dorsal setae range from 45/a to 21/* in length and there are 2 humeral 
setae followed by 66-74, arranged anteriorly in rows of ea. 10. All coxae 
unisetose, and posterior of coxae about 64-70 ventral setae. No nude seta on 
tarsi III. 

Wharton and Hardcastle's data for 4 specimens treated statistically are: 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


85*32:1 -67 


2«89:fcl*18 


56-6-74-0 


62-0-67-0 


4-4 


PW 


73'25±0*72 


l-44±0-51 


67 -95-78 '55 


72-0-74-0 


2-0 


SB 


S3-0:tl-06 


2-45Hr0'87 


46-7-59-3 


50-0-55-0 


4-6 


ASB 


21-75±0-95 


l-89;±0*67 


16-0-27-4 


20-0-24-0 


8-7 


PfiB 


30'5±l-85 


3-70±l-30 


19-5-41-5 


28-0-36-0 


12-1 


SD 


52-25±2-60 


5' 19+1*84: 


36-9-68-1 


49-0-60-0 


9-9 


A-P 


31-5±l-26 


2-51±0-89 


24-0-39-0 


29-0-35-0 


8-0 


AM 


27-7±l-76 


3-05±l-S5 


18-6-36-8 


25*0-31-0 


11-0 


AL 


57-3±l<33 


2-13±0*94 


5-04-64-2 


56-0-60-0 


4-0 


PL 


37'0±l-22 


2-45±0-87 


29-7-44-3 


35-0-40-0 


6-6 


Sens. 


31-0 


Only oae measurement. 







NEOSCHONGASTIA STRUTHIDIA Sp. n. 

ParascMngastia sp., Gill, Moule and Rcik, 1945, Aust. Vet J. ? 32, fig. (5). 

Plate 70, fig. F-J. 

Description of Larvae. Shape broadly oval, almost cordate. Length, 
engorged 450/*, width 330/*. Dorsal scutum as figured with the anterior margin 
sinuous, convex medially and laterally; posterior margin rather deep, sinuous 
and concave medially; surface anterior of sensillae bases with large but sparse 
tuberculatum ; sensillae globose, apparently nude or with only minute incon- 
spicuous setulations, the bases about midway between lines of AL and PL, 
situated in the usual transverse crest-like wall; posterior with circular stria- 
tions. Eyes 2 + 2, with distinct ocular shields, the anterior eye the larger 
and more prominent. Chelicerae as figured, with the usual apical tricuspid 
cap, but the forwardly directed subapical inner tooth not as prominent as in 
other species. Galeal setae short and with a few branches. Palpi stout, tibial 
claw trifureate, median prong over-reaching tip of tarsus; setae on femur and 
genu long and strongly branched; on tibia, dorsal nude, lateral with a few 
indistinct branches, ventral slender and shortly branched; tarsus with the 
usual subapical and sub-basal sensory rods and 4 or 5 ciliated setae. Dorsal 
setae slender and very shortly ciliated, 32 in number and arranged' 2.8.6.6.4.4.2, 



256 Records of the S.A. Museum 

to 54//, long- Ventrally with the usual pair of branched setae on gnathosoma, 
one on all coxae, a pair between coxae I and between coxae III and thereafter 
6.4.6.4.2.2, to 40/i long. Legs: I 420/* long, II 360/*, III 440/1.; tarsi I and 
II with the usual dorsal rod-like setae, III without a long nude seta, the corre- 
sponding setae being shortly but distinctly ciliated; tarsi I without the con- 
spicuous dorsal subapical lobe of entoniyz®* 

The Standard Data from the type and 5 paratypes are as follows : 





Mean 


Standard 
Deviation 


Theoretical 
Bange 


Observed 
Range 


Coeff . of 
Variation 


AW 


75'5±0-92 


2-26±0*65 


68-7-82*3 


73-0-79*0 


3-0 


PW 


85-0±:0'77 


l-90±0-55 


79-3-92*7 


82-0-88-0 


2-2 


SB 


43-0±0-63 


1'55±0'45 


38-3-47-7 


42-0-45- 


3-6 


ASB 


22-5:£0-67 


l-64±:0-47 


17-6-27-4 


21-0-24-0 


7-3 


PSB 


3Q-3±0-36 


0-82:fc0-23 


27-9-32-7 


30-0-32*0 


2-7 


SD 


52 -8:1:0 -89 


2 -14:1:0 -62 


46*4-57-2 


51*0-56-0 


4-0 


A-P 


33-0 


No variation recorded 






AM 


57-6±0-60 


l-34r£0-42 


53-6-61-6 


57-0-60-0 


2-3 


AL 


85-8±:0-83 


2-04±0-59 


79-7-91-9 


85-0-90-0 


2-4 


PL 


51-75^1-43 


2-87zbl-01 


43-1-60-4 


48-»>-54*0 


5-5 



Sens. 32-0 with head 24/24, 

Loo. Described from 6 specimens from an Apostle Bird, Striithidia 
cinerea from Logan Downs, Clermont, Queensland 1944 (D. A. Gill). 

Remarks. The slide containing the type and two paratypes is in the 
McMaster Laboratory, Sydney and another slide of 3 paratypes is in the 
South Australian Museum collection. I am indebted to Mr. D. A. Gill of the 
McMaster Laboratory for the opportunity of describing this material. 

Neosohongastia retrooincta Gunther 1939. 

Neoschongastia retrocaronata Gunther 1938 (nam. nvd,), Med. J. Aust,, 2, 
(6), 202. 

Neoschongastia retracincta Gunther 1939, Proc. Linn, Soc. New South Wales, 
64, (1-2), 87; Wharton and Hardeastle 1946, J. Parasitol, 32, (3), 295. 

Parasckongastia retrotincta, Womersley 1939, Tr. Roy. Soc. S. Aust,, 63, (2), 
165; Gunther 1940, Proc. Linn. Soc. New South Wales, 65, (3-1), 247; 
Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 67, (1), 131. 

Plate 70, fig. A-E. 

Tn this species the palpal claw is trifurcate with the median prong about 
twice as long as the others and reaching well beyond the tip of the palpal 
tarsus. The setae on the palpal femur and genu are strongly branched; on 



Womhrsley— Asiatic-Pacific Scrub Typhus Mites 257 

the tibia the ventral seta is strongly branched, the dorsal nude, lateral 2-3 
branched; long dorsal seta on palpal tarsus somewhat fasciculate. Chelicerae 
as in yeomansi. Galea! setae 2-3 branched. 

Posteriorly on the dorsum and venter is a semicircular area demarcated 
by a circle of stout, finely serrated spines to 72/a long; inside the area are a 
number of large, more or less rectangular to round, finely pitted platelets from 
which arise short, 25>a long, pointed and finely serrate scetae. The dorsal setae 
are normally ciliated and ta.pering to 40/l long. 

Coxae III are furnished each with 2 setae; tarsus III has not got a long 
nude seta as stated by Gunther, the corresponding seta being finely but perhaps 
less ciliated than the others. 

The Standard Data for the type and 3 paratypes reported on in 1943 
(lo-a, cit.) and 3 specimens from scrub at Lae, New Guinea, 9th April, 1944 
(R. N. McCulloch) and 1 specimen from the mound of a bush turkey Dobodura, 
June 1944 (G M> Kohls), N.G., are as follows : 



AW 


Mean 
71-4+0-70 


Standard 
Deviation 

2-00±0-50 


Theoretical 
Bangc 

65-4-77-4 


Observed 
Bange 

70-0-75-0 


Coeff , of 
Variation 

2-8 


PW 


84-5ztl-35 


3-82±0-95 


73-1-95-9 


80-o-gi-o 


4-5 


SB 
ASB 


55-5:t0-71 
28-0 


2-00±0-50 49 -5-61 -5 
No variation recorded 


54-(MiO-0 


3-6 


PSB 


21'3±0-76 


2-01 ±0-54 


15-3-27-3 


21-0-25-0 


9-4 


8J> 


44-4=t0-53 


l-39±0-37 


40-2-48-6 


42-0-46-0 


3-1 


A-P 

AM 


35-25zt0-25 
36-0 


0-71±0-18 
Ko variation 


33-15-37-35 
recorded 


34-0-36-0 


2-0 


AL 
PL 


72-0±l-07 
51-4±l-49 


2 -83+0 -76 
3'95±l-06 


63-6-80-4 
39-7-63-1 


70-0-76-0 
44-0-56-0 


3-9 

7-7 



Sens. 36-0 with head 20/27. 

Neosciiojngastja yeomansi Gunther 1939. 

Neoschchigasiia jamesi Gunther 1939 (nom. mid.), Med. J. Anst., 2, 202. 

Neosckon-gcistia yeomanai Gunther 1939, Proc, Linn. Soc. New South Wales, 
64, (1-2), 81; nee. Wharton and Hardcastle 1946, J. Parasit., 32, (3), 293. 

Pai'aschongastia yeommisi Womersley 1939, Tr. Roy. Soc. S. Aust, 63, (2), 
166; Gunther 1940, Proc. Linn. Soc. New South Wales, 65, (3-4), 252; 
Womersley and Ileaslip 1943, Tr. Roy. Soc. S. Aust., 67, (1), 131. 

Plate 71, fig. A-K- 

In this species the palpal claw is trifurcate and short and stumpy, with 
the median and longest prong only slightly over-reaching the apex of the 
palpal tarsus. The chelicerae are more serrate, with an apical tricuspid cap 



258 



Records of the S.A. Museum 



of which the ventral tooth is rather pronounced, and on the inner (dorsal) edge 
with a prominent forwardly directed tooth. The galeal setae are short with 
4 or 5 branches. The setae on the palpal femur and genii are long, with long 
outstanding branches,; on the tibia the dorsal and lateral setae are relatively 
short and nude; the ventral tibial seta is much longer with about 4 long 
branches; the palpal tarsus carries the usual subapieal and sub-basal sensory 
rods and 5-6 ■ciliated setae, the longest of which is fasciculate, and over- 
reaches apex of claw. There is no long nude seta on tarsi III as stated by 
Gunther (a sub-basal long outstanding seta is present but distinctly, though 
shortly, ciliated). 

The dorsal scutum is as figured with AL the longest, and AM the shortest 
and situated well posterior of line of AL ; the sensillae are situated wide apart, 
about equidistant between lines of AL and PL, globose with very fine villosity, 
and their bases in a crescentie wall; the anterior half of the scutum is pitted, 
and the posterior half with circular overlapping striations. 

The scutal and anterior dorsal setae are tapering, elosely and shortly 
eiliated. The anterior dorsal setae number approximately 7fl, followed posteri- 
orly by an area, more or less pitted and carrying about 40 stout spine- or awl- 
like setae, situated on platelets, and with their margins sparsely serrated,- the 
-anterior setae are from 45 to 35/* long, decreasing posteriorly, and the pos- 
terior spines to 42^ long. 

Coxae I and II are unisetose but III are bisetose. In. 7 paratypes in 
the South Australian Museum, 5 paratypes in the collection of Wharton and 
Hardcastle, as well as in two specimens from Lae, New Guinea, there is no 
variation in the setation of coxae III. The setae on coxae I and II are very 
long. 

Ventrally, behind coxae III are about 48 tapering ciliated setae anterior 
of the anus, and then about 40 serrated spines on platelets as on the dorsum. 

The Standard Data for 7 paratypes and two specimens from Lae, N.G,, are ; 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Dcviation. 


Range 


Ru n £<?. 


Variation 


AW 


77-15+0-49 


l-47:t0-35 


72-75-81-55 


75-6-78*4 


1-9 


PW 


103 » 1:1:1 -85 


5-56±l-31 


86-4-119-8 


98*0-112-0 


5-4 


SB 


64-l±0-56 


1*68±0»39 


59-1-69-1 


61-6-67-2 


2-6 


ASB 


30 -5:±0 '31 


0-93zt0>22 


27-7-33*3 


28-0-30-8 


3-0 


PSB 


28-3:t0«31 


0-93=h0-22 


25-5-31-1 


28-0-30-8 


3-3 


8D 


58-8:fc0-47 


1-40+0-33 


54.6-83-0 


56-0-61-6 


2-4 


A-P 


36-4±0-66 


1*98+0-47 


30*5-42-3 


33-6-39-2 


5-4 


AM 


39-2:±l-22 


3-23+0-86 


32-5-45-9 


33-6-42-0 


8-2 


AL 


83-l:£0-87 


2-46+0-62 


75-7-90-5 


78-4-S6-8 


3-0 


PL 


55'6±0-40 


1*06+0-28 


52-4-58-8 


53*2-56-0 


1-9 


Sens. 


39-2 with head £ 


JO/28. No variation recorded. 







WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 259 

Loc. and Hosts. From Me-gapodius duperreyi from Bulolo, N.C4. (Gunther 
1939), and two specimens from Lae, N.G., 9/4/1946 (R. N. McCulloch). col- 
lected on boots. 

Remarks. In 1946, Wharton and Hardcastle, recorded as N. yeomansir 
Gunther, 5 specimens from Peleliu and Palau Islands in the Western Pacific, 
from Gallus gaLlus and Megapodius laperouse respectively. 

Of these specimens, which they compared with 5 paratypes of yeomansi, 
they found that one had 2 + 3 setae on coxae III, three had 3 + 3, and one 
4 + 4. 

In the present paper, two new species are recognized, which, while closely 
related to yeoman$i 9 both differ in having the setae on coxae III normally 
3 + 3, but variable from 2 to 4 on either side. Prom fourteen specimens of 
yeo-mansi known to me, there is no variation from the normal 2 + 2, The new 
species, mcqweni, from Bougainville, however, shows 3 + 3 in 5 out of 6 speci- 
mens; the other specimen has 3+4. In 31 specimens of ow-iewis sp. n, from 
Owi Island, Dutch New Guinea, 15 have 3 + 3, fourteen have 3 + 4, and two 
have 2 + 3, 

As there are, however, other differences between these three species, it 
would appear that the Peleliu and Palau specimens may be either mcqueeni or 
owiensis but almost certainly not yeomatm Gunther. Unfortunately, "Wharton 
and Hardcastle do not give figures or details sufficiently to fix their material 
at present. The Standard Data given, suggests, however, its tentative associa- 
tion with owiensis sp. n. 

Neoschongastia owiensis sp. ili 
Plate 71, fig. P-H ; 72, fig. A-B. 

Description of Larvae. Length (unengorged) to 33G>, width to 21G>. 
Shape oval, slightly wider anterior of middle* Dorsal scutum as figured, with 
scutal setae tapering with long, outstanding branches; AM the shortest and 
well behind line of AL. AL the longest; sensillae distinctly setulose, more so 
than in yeoma,mi; anterior half of scutum pitted, posterior half over-lapped 
with circular striations. Eyes 2 + 2, on distinct ocular shields. Chelicerae 
with apical tricuspid cap, and on inner (dorsal) edge with a strong, forwardly 
directed subapical tooth. Galeal setae fine with only 2 or 3 fine branches. 
Palpi stout, femur and genu with strongly branched setae; dorsal and lateral 
seta.e of tibia nude, ventral branched ; palpal tarsus with 4 or 5 ciliated setae, 
the longest slightly fasciculate, and the usual subapical and sub-basal sensory 
rods; claw short and stumpy, trifurcate. Dorsal setae with long outstanding 



260 Records of the S.A. Museum 

ciliations on anterior rows; anteriorly approximately 76, posteriorly approxi- 
mately 46 stout, awl-like spines, distinctly ciliated rather than serrated as in 
yeomansi. Vent rally a pair of branched setae on maxillae, one fairly long on 
coxae I and II, and normally 3 + 3 (frequently 3 -\- 4, rarely 2 -|- 3) on 
eoxae III; a pair of branched setae between eoxae T and between coxae III, 
and thereafter approximately 50 tapering ciliated setae anterior of anus, and 
approximately 40 spines behind anus, similar to dorsal spines. Anterior dorsal 
setae to 45/t, posterior spines to 42/a. 

Legs: I 390/a long, II 300^, III 390/x; tarsi I and II with usual dorsal 
sensory rodj III without any long nude seta. 

The Standard Data for 31 specimens are : 



AW 


Mean 
79-7:±0-39 


Standard 
Deviation 

2-16±0-28 


Theoretical 
Range 

73-2-86-2 


Observed 
Range 

75-6-84-0 


Coeft. of 
Variation 

2-7 


PW 


93-5±0-41 


2-26±0-29 


86-7-100-3 


89-&-98-0 


2-4 


SB 


62-15:t0-31 


1-70+0-22 


57-05-67-25 


58-8-64-4 


2-7 


ASB 


28-3±(M5 


0-83+0-10 


25*8-30-8 


28-0-30-8 


2-9 


PSB 


27'45±0-20 


1-11+0-14 


21 -15-30 -75 


25-2-28-0 


4-0 


SD 


55-7+0-23 


1-30+0-16 


51-8-59-6 


53-2-58-8 


2-3 


A-P 


35-3±0-27 


l-53±0-19 


30-7-39-9 


33*6-39-2 


4-3 


AM 


34-8±0-39 


2-13+0-27 


28-4-41-2 


28*0-39*2 


6-1 


AL 


80-8±0-58 


3-21±0-41 


71-2-90-4 


72-8-84-0 


4-0 


PL 


51-6+0-55 


3-04±0-39 


42-5-60-7 


47-6-56-0 


6-0 



Sens. 39-2 with head 20/28. Xo variation recorded 

Loc. Twenty-two specimens from boots, Owl Island, Dutch New Guinea, 
July and August 1944 (G. M. Kohls), and nine specimens from somewhere in 
the Western Pacific, (U.S. Navy, 3,505, A8). 

Remarks. This species is close to yeomansi. Gunther, but differs mainly in 
the setation of coxae III, the nature of the dorsal and scutal setae, and in the 
Standard Data. From mcqueeni sp, n., from Bougainville, it differs in the 
much larger dorsal scutum, and the very different Standard Data. 

Neoschonoastia mcqueeni sp. n. 

Plate 72, fig. C-G. 

Descriptwn of Larvae. Shape ovoid. Length (unengorged) to 270/x^ 
width to 18Qfi, Scutum as figured, with AL the longest and AM the shortest 
and about in line with AL; all scutal setae with long outstanding branches; 
sensillae globose and distinctly with fairly long setules, their bases wide apart 
and in a crest about midway of the scutal depth. Eyes 2 -f 2, on distinct, ocular 



Womersley— Asiatic-Pacific Scrub Typhus Mites 261 

plates. Chelieerae with apical tricuspid cap, and on inner (dorsal) edge a 
strong 1 forwardJy directed subapical tooth. Galeal setae fine with 2 or 3 fine 
branches. Palpi fairly stout,- setae on femur and genu with fairly long 
branches} on tibia dorsal and. lateral nude, ventral shortly branched; tarsi 
with the usual subapical and sub-basal sensory rods and 5 to 6 ciliated setae; 
tibial claw trifurcate but not short and stumpy as in yeomansi and omensis. 
Dorsal setae, anteriorly approximately 60 in 6 rows of fine, tapering setae with 
long outstanding setules, posteriorly with about 28 stout awl-like spines with 
distinct short eiliations; anterior setae to 42juc long, spines to 28/x. Veutrally, 
with a pair of branched setae on maxillae, one on each of coxae I and II, three 
(rarely four) on each of coxae III, a pair between coxae I and between 
coxae III and thereafter ca. 42 fine ciliated setae followed behind the anus by 
ca. 36 spines similar to those on dorsum. The setae on coxae I and II are not 
as long as in yeomatisi and owiensis. 

Legs: I 340/t long, II 300, III 330/a; tarsi 1 and II with dorsal sensory 
rod. III without any long nude seta. 

The Standard Data for the type and 5 paratypes are: 

Standard Theoretical 

Mean Deviation, Eange 

AW 65-5±:0-68 l-53:£0'48 60-9-70-1 

PW 74-5:£l-B8 3-76±l<19 63-2-85-8 

SB 47-6 No variation recorded 

A8B 26-6±0-40 0*98:1:0 '28 23*7-29-5 

PBB 22*4 No variation recorded 

SD 49-0±0-40 0-98±0'28 46«l-51-9 

A-P 32-7±0-59 W4±0-42 28'4-37-0 

AM 32.2±0-81 l»62±0-57 27-4-37-0 

AL 57-4±0-81 l-62±0-57 52-6-62-2 

PL 42-0 No variation recorded 

Peng. 28 • witn head 19/19. No variation recorded. 

Loc. The type and 5 paratypes in the collection of the South Australian 
Museum, from a small island to the North of Bougainville, August, 1946 
(GKH. McQueen). 

Remarks. This species is very closely related to both owiensis and yeo- 
mami but differs from these in the much smaller dorsal scutum and Standard 
Data, and particularly in the tibial palpal claw not being stumpy. The scutal 
and dorsal setae are long branched as in oimensis. The dorsal setae are 
also somewhat fewer in number. Coxae III is trisetose as in omensis, but 
only rarely 4-setose (not frequently as in oiuiensis) ; of the 6 specimens, only 
one showed 3 + 4 setae. 



Observed 


Coeff. of 


Range 


Variation 


64-4-67-2 


2-3 


70-0-78*4 


5-0 


25-2-28*0 


3-6 


47-6-50-4 


2-0 


30-8-33-6 


4*4 


30-8-33-6 


5-0 


56-0-58-8 


2-8 



262 



Records of the S,A. Museum 



Neoschongastia atollensis Wharton and Hardcastle 1946. 
J, Parasitol., 1946. 32, (3), 306, fig. (larva), 

Plate 74, fig. H. 

Described from Numenius phaeopus from Ulithi Atoll, August, 1945, and 
also from other birds from Okinawa, July, 1945. 

This species is closely related to namrui Wharton and Hardcastle in having 
coxae III with more than 5 setae, the number varying from 5 to 7. In namrui 
the number varies from 7 to 12. The galeal setae are nude, whereas in namrui 
they are feathered. 

The dorsal scutum is as figured, after Wharton and Hardcastle, PW much 
wider than AW. The sensillae are globose, with short pedicel and the sen- 
sillae heads lie in fairly well defined depressions surrounded by the circular 
cuticular striations. The AL setae are the longest. On the palpi the dorsal 
and lateral setae on the tibia are nude, the ventral and the femoral and genual 
are branched. 

The dorsal setae are from 21 to 49/x long, with four humeral setae and 
then ca. 120 arranged in rows of 12 or less. Coxae I and II unisetose, posterior 
of coxae III about 100 setae ventrally. No long nude seta on coxae III. 

The Standard Data as given by Wharton and Hardcastle for 6 specimens 
and treated statistically are : 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


46*5:±0-95 


2 -34:4:0 -68 


39-5-53-5 


42-0-48-0 


5-0 


PW 


69-2:±2-88 


7-07±2-04 


48-0-90-4 


56-0-77-0 


10-2 


SB 


20-5±l-58 


3'88±1-12 


8-9-32-1 


14-0-24-0 


19-0 


A8B 


19-5±l-06 


2-58=fc0-75 


11-8-27-2 


15-0-22-0 


13-2 


PSB 


31-2±l-30 


3-19±0-92 


21-7-40-7 


27-0-35-0 


10-2 


St) 


50-7:±2-03 


4*97:fcl-43 


35-8-65-6 


42-0-50-0 


9-8 


A-P 


41-7:±l-92 


4-72±l-36 


27-6-55-8 


34-0-48-0 


11-3 


AM 


26-2:tl-l4 


2-79^:0-80 


17-9-34-5 


21-0-28-0 


]0-6 


AL 


47 -7=1:2 -42 


5-92^:1 '71 


30-0-65-4 


36*0-53-0 


12-4 


PL 


33-0dbl-05 


2-68^0-77 


25-0-41-0 


28-0-35-0 


8-1 


Bens, 


28-0±4-0 


5-66±2*83 


11- 2-39 -8 


24-0-32-0 


20-2 



Remarks. The great variability shown above is largely due to the fact 
that the one specimen recorded from Okinawa had a very much smaller scutum 
than the Ulithi specimens. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 263 

Neoschongastia namuui Wharton and Hardcastle 1946. 
J. Parasitol., 1946, 32, (3), 316, fig. (larva). 

Plate 75, fig. G. 

Described from Guam from Numenius phaeopus in May, 1945, and from 
Heteroscelrus incanus and PluviaUs dominica in September, 1945; and from 
Gygis alia on Okinawa in July. 1945. 

This species is remarkable in that the scutum is slightly longer than wide, 
and with PW being shorter than AW, the lateral margins slightly converging 
posteriorly; the anterolateral shoulders are somewhat strongly produced for- 
wards. The lateral branches of AM are very long, those of PL not quite so 
long, but in both AM and PL much longer than in AL. Sensillae globose and 
setnlose, with their bases nearer to AL than to PL. The galeal setae and all 
setae on palpal femur, genu and tibia are branched. The DS setae are very 
numerous, 44^ long anteriorly to 32p posteriorly; there are about 15 setae on 
each shoulder and then about 175 setae behind these. 

Coxae III with from 7 to 12 setae and posterior of coxae III with ca. 150 
setae. Tarsus III without any long nude seta, but on the genua of all legs is 
a rather characteristic cluster of short nude setae. 

The Standard Data for 7 specimens given by Wharton and Hardcastle and 
treated statistically are; 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff, of 
Variation 


AW 


74-4±l-17 


3'10±0-83 


65- 1-78-0 


69-0-78-0 


4-2 


PW 


60-2±2-06 


5-46±3-46 


43-9-76-5 


53-0-69-0 


91 


SB 


40*0:±0-69 


l-82±:0-49 


34-5-45-5 


38*0-43-0 


4-5 


A8B 


34*4±1*92 


4-2S±l»35 


21-6-47-2 


27*0-38*0 


12-4 


PkSB 


40-0+0-51 


J-26±0-36 


36-2-43-8 


39-0-42*0 


3-2 


SD 


74-4±2*2 


4«93±l-56 


59- 7-89-1 


66-0-78-0 


6-6 


A-P 


46-2±0-51 


l*34:±0-36 


42-2-50-2 


45-0-48-0 


3*0 


AM 


61-0±1-41 


3-74±l*00 


49-8-72-2 


54-0-66-0 


6*1 


AL 


70-4±0-57 


1-51:±0*40 


65-7-74*9 


69-0-73-0 


2*1 


PI/ 


70*4±2»87 


7-59z£2-03 


47-6-93-2 


60-0-78-0 


10-8 


Sens. 


36-7±2-15 


5-70+1-52 


19-6-53-8 


240-44-0 


1-6 



Remarks. Close to N. carveri but easily distinguished by the characteristic 
scutum, sternal setae and the clusters of nude setae on the genua, of the legs. 

Neoschongastia Americana, solomonib Wharton and Hardcastle 1946. 

J. Parasitol., 1946, 32, (3), 289-292, fig. 

Plate 74, fig. D. 

Wharton and Hardcastle 1946 described a number of specimens from 
Bougainville, Two Jima. and Okinawa as a variety of the American species, dis- 
tinguished them mainly on the longer scutum. 



264 Records of the S.A, Museum 

For comparison they give tlie Standard Data of 5 specimens of N. americana 
americana from which the following statistics are now calculated : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


49-0±l-14 


2-55±0-81 


41-4-56-6 


45-0-52-0 


5-2 


PW 


66-8:hl-65 


3-70=tl-17 


55-7-77-9 


62-0-72-0 


5-5 


SB 


30-6±l-43 


3-21+1-01 


21-0-40-2 


26-0-34-0 


10-4 


ASB 


17-4±rl-29 


2-88±0-91 


9-0-26-0 


13-0-20-0 


16-5 


PSB 


26-2±:l-24 


2 -77+0 -88 


17-9-34-5 


23-0-29-0 


10-5 


8D 


43-8±l-65 


3-70±l-17 


32-7-54-9 


38-0-48-0 


8-4 


A-P 


27-6±0-75 


1<69±:0*53 


22-6-32-6 


26-0-31-0 


6-1 


AM 


32-5±l-44 


2-89±l-02 


23-9-41-1 


29-0-36-0 


8-8 


AL 


54-75+3-66 


7-32±2-58 


32-8-76-7 


49-0-65-0 


13-3 


PL 


55-5±:3-17 


6-35±2-24 


36-5-74-5 


52-0-65-0 


11-4 


Sena. 


25-3±l-46 


2-55:tl-03 


17-7-32-9 


23-0-27-0 


10-0 



The corresponding values for nine specimens from the Southern Pacific 
from Wharton and Hardcastle's data are: 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


51- 7+1 -38 


4-16+0-98 


39-2-64-2 


45-0-57-0 


8-0 


PW 


70-2+2-0 


6-01+1-42 


52-2-88-2 


57-0-77-0 


8-5 


SB 


34-0+1-43 


4-30+1-01 


21*1-46 '9 


28-0-39-0 


12-7 


ASB 


25-7+0-81 


2-14±0-57 


19-3-32-1 


24-0-28-0 


8-3 


PSB 


29-4+0-53 


l-39±0-37 


25-2-33-6 


28-0-310 


4-7 


SD 


55-2+0-83 


2-19±0-59 


48-6-61-8 


52-0-59-0 


4-0 


A-P 


35-6±0-56 


1-51+0-40 


31-1-40-1. 


34-0-38-0 


4-2 


AM 


43-6dbl»49 


4-47 + 1-05 


30-2-57-0 


36-0-50-0 


10-2 


AL 


55-6±l-90 


5-70+1-34 


38-5-72-7 


44-0-61-0 


10-3 


PL 


57-1+2-29 


6-88+1-62 


36-5-77-7 


42-0-66-0 


12-0 


Sens. 


25-8±l-0 


3-03+0-71 


18-7-34-9 


21-0-29-0 


11-8 



From these values the form from the Solomons only differs significantly 
in ASB, PSB, SD, A-P and AM, at the 1% level. In Hirst's figure (loc, ctt.) 
there is shown a long nude seta very clearly on tarsus III, which is not men- 
tioned or figured in Wharton and Hard castle. It is, however, present in a 
specimen, given to me by Dr. Wharton, from Iwo Jima. 

The DS are 32, arranged 2.8.6.4.6.2, and range in length from 68/x an- 
teriorly to 28/x posteriorly. The setae on the palpi femur and genu are 
branched, and on the tibia the dorsal and lateral setae have one or two short 
branches, and the ventral is distinctly branched. Coxae III trisetose. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 265 

Neosohongastia entomyza sp. m 
Paraschongustea, sp., Gill Moule and Reik, 1945, Aust. Vet. J., 32 (fig, 4). 

Plate 72, fig. H-K; 73, fig. A. 

Description of Larvae: Shape oval. Length, unfed, 405/x 7 width 255>a. 
/Dorsal fccutum as in fig. 66 and plate J with the usual 5 ciliated setae and a 
pair of finely setulose globose sensillae with their bases arising from a triple 
crest about midway between lines of AL and PL, Anterior margin sinuous 
with the shoulders anterior of the middle. Posteriorly the scutum with the 
usual orescenti c striations. Eyes 2 + 2, very large and on prominent ocular 
shields, closely adjacent to the lateral margins of scutum, posterior eyes the 
smaller. Chelicerae with the usual apical tricuspid cap and on inner margin a 
subapical forwardly directed point. Galea! setae .short, with a few short 
branches. Palpi stout, tibial claw rather slender and trifurcate, the median 
and longest prong surpassing apex of palpal tarsus; setae on femur and genu 
strongly branched; on tibia, dorsal apparently nude, lateral with one or two 
branches, and ventral with many branches ; tarsus with the usual subapical and 
sub-basal sensory rods and 4 or 5 ciliated setae. Dorsal setae shortly ciliated, 
to 30/^ long and 26 in number and arranged 2.8.6.4,4.2. Ventrally with the 
usual pair of branched setae on gnathosoma, one on coxae I and II, three on 
•coxae III, a pair between coxae I and between coxae III and thereafter ca. 
6.4.4.2.4.2, to 30/i long; the posterior dorsal and ventral setae are similar to 
the anterior and do not arise from platelets, Legs rather long and slender but 
I stronger and thicker than the others; tai k si I with a prominent subapical 
dorsal hump; tarsi I and II with the usual dorsal rod-like setae, III without 
any long nude seta, the corresponding setae being distinctly but shortly 
ciliated. Leg I 470/a long, II 415/*, III 496/x. 

The Standard Data from one paratype and 13 other specimens are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Eange 


Range 


Variation 


AW 


47-l±0-51 


2.]0zh0»41 


40.8-53-4 


44-8-50-4 


4-4 


PW 


68-9±0-67 


2-43±0-48 


61-6-76-2 


64-4r-72-8 


3-5 


SB 


30- 6±0-59 


2-13-£0-42 


34,2-37-0 


28-0-33-6 


7-0 


AftB 


20-8±0-38 


1 -44±0* 27 


1G-5-25-1 


19-6-22-4 


6-9 


P8B 


29-4±0-39 


l-45:fc0-27 


25-1-33 -9 


28-0-30-8 


4-9 


SB 


50-2:1:0 -46 


l-73±0-33 


4S-0-55-4 


47-&-53-2 


3-4 


A-P 


32-4+0-57 


2-12±0-40 


26*1-38-7 


28-0-36-4 


6-5 


AM 


31-85±0^74 


2-08±0-52 


25 '£-38-1 


30-8-36-4 


6-5 


AL 


37-8±0-59 


2*14:£0-42 


31 -4-44-2 


36-4-42-0 


5-6 


PL 


45-8:fcO-70 


2-53:±0-50 


38-2-53-4 


42-0-50-4 


5-5 


Sens. 


30 '8 with head 


22 -4/25 -2, No variation recorded. 







266 Records of the 5.A. Museum 

Loc. Said to be common oit birds in the Clermont district, Queensland, 
along with Tr&mbicuto (Leptotrombidium) myzantha sp. n. by Messrs. Gill, 
Moule and Reik (Aust. Vet. J., April, 1945). The birds listed a& carrying this 
species by these authors are Struthidia dnerea Gould 1837, Entomyzon cyanotic 
Latham 1801, Cractic<it3 t&rqmtus Latham 1801, Myzantha melmoctphala 
Latham 1801, Gymnorhina tibicen Latham 1801, Ocyphaps lophotes Temminck 
1822 and Corv%ts cecilae Mathews 1912. 

Remarks. The description is drawn up from the type and 1 paratype in. 
the collection of the McMaster Laboratory, Sydney, and 1 paratype in the South 
Australian Museum, all labelled from "leather head" (blue-faced honey-eater, 
Entomyzon cyanotis) from Wolfgang, Clermont, Queensland, coll. D. A. Gill, 
18 Apr. 1944) and 1 specimen from a "lousy jack" (probably an Apostle bird, 
Struthidia dnerea) from Logan Downs, Queensland, 28th Nov. 1944 (coll. 
D. A. Gill) in the McMaster collection. The 13 other specimens from which the 
Standard Data are given were from Entomyzon cyanotis from Buckleton, C.Q- 
(5 spec), Logan Downs, C.Q. and from Pomatostomus temporalis, Emerald,. 
C.Q. (2 spec), all in Nov. 1948 (H.W.). 

I am greatly indebted to Mr. D. A, Gill for the opportunity of describing 
this species. 

Neosohongastia thomasi (Radford 1946). 

Parasclwngastia thomasi Radford 1946. Proc. Zool. Soc. London, 116, (2) r 
(larva). 

Plate 73 T fig, B-D. 

This species was briefly described by Radford from specimens from a 
Black-headed Shrike (Lamus nasutus) from Ukrul Road, Imphal, Manipur, 
11 Feb., 1946. 

In the shape of the scutum it appears to be very close to entomyza sp. n. 
only differing in that A-P is much longer and the scutum consequently deeper. 
The scutal setae are longer and the globular sensillae apparently nnde. Pos- 
teriorly the scutum is overlapped by the dorsal cuticle with semicircular 
striations. The DS are slightly fewer than in entomyza and are arranged 
2.6.6.4.4.2. No details of palpi and ehelieerae are available. The eyes are 
2 + 2, large, on ocular shields with the posterior eyes the smaller. The coxae 
are strongly punctate and III trisetose. Posterior of coxae III with 8.6.4.4.2.2" 
setae ventrally. 

The Standard Data from Radford's figures are: AW 44-0, PW 63-0, 
SB 28-0, ASB 20-0, PSB 23-0, SD 43-0, A-P 34-0, AM 37-0 T AL 51-0,. 
PL 54-0, Sens. 29-0. 

This species is so very close to entomyza that it may quite possibly, wheni 
more carefully studied from fresh material be the same. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 267 

Neoschongastia pauensis Wharton and Hardcastle 1946. 
J. Parasitol., 1946, 32, (3), 311, fig. (larva). 

Plate 75, fig. B-D. 

Described from Pluvialis dominica from IJlithi Atoll in August, 1945. 

Close to JV. ewingi but differing in that there are 3-4 setae dorsally on 
each shoulder instead of one. On the palpal femur, genu and tibia the setae 
are all branched except the dorsal and lateral tibial. 

The dorsal scutum is as figured, after Wharton and Hardcastle, with the 
lateral margins strongly concave. PL the longest but only slightly longer 
than AL, and the sensillae bases slightly in front of half-way between lines 
of AL and PL. Galeal setae with 2 short branches. 

Dorsal setae from 55/a anteriorly to 30/* posteriorly ; as many as 5 humeral 
setae on each side, and then about 50 in rows of 10. Coxae III with 3-4 setae. 
Posterior of coxae III with about 60 setae. No long nude seta on tarsi III. 

The Standard Data given by Wharton and Hardcastle for 5 specimens 
treated statistically are: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 
PW 


57-5±0-87 
77-0 


l-73±0-61 52-3-62-7 
No variation recorded 


56-0-59-0 


3*0 


SB 
ASB 


37-25+1-11 

22*0 


2-18:t0-78 30-6-43-9 
No variation recorded 


35-0-40*0 


5-95 


PSB 


30- 7+2-03 


3-51:1:1 -43 


20-2-41-2 


27-0-34-0 


11-4 


SB 


52-5:t3-50 


4-d5:£2-47 


45-3-59*9 


49-0-56-0 


9-4 


A-P 


31*25^0*75 


l-50=t0-53 


26-75-35-75 


29-0-32-0 


4-8 


AM 


35*0±l-47 


2-94±l-04 


26-2-43-8 


31-0-38-0 


8-4 


AL 


52-5±3-5 


4-95±2-47 


37-7-67-3 


49-0-56-0 


9-2 


PL 


54-75±0-SO 


l-61±0-57 


49-95-59*55 


53-0-56-0 


3-0 


Sena. 


29*3±0<89 


1-53:1:0 -62 


24-8-33-8 


28-0-31-0 


5-2 



Neoschongastia salmi (Oudemans 1922). 

ScMngastia salmi f Ouds. 1922, Ent. Ber., 6, (126), 81 (brief diagnosis), idem, 
6, (128), 114; Salm 1923, Bull. Soc. Pathol. Exot., 16, 236-300,- Walch 
1927, Geneesk. Tijds. Ned. lnd., 67, 931-932; Womersley and Heaslip 
1944, Tr.Roy. Soc. S. Aust., 68, (1), 98; Sig Thor and Willmann, Das Tier- 
reieh, 71b, 304. 

Plate 73, tig. E-G. 

This species was originally briefly diagnosed by Oudemans in Ent. Ber., 
July, 1922, 6, (126), 81. Oudemans himself does not appear to have published 
any lengthy or full description, but such a description in German was drawn 



268 Records of the S.A. Museum 

up and is in the Oudemans' library and manuscripts in the possession of the 
Dutch Entomological Society under the care of Dr. G, L. van Eyndhoven. I am 
greatly indebted to Dr. Eyndhoven for a copy of this description. 

Oudemans also made very careful and detailed drawings, and these (draw- 
ings No. 1126, 1127) are in the Oudemans 1 collection in the Leiden Museum, 
but have not been published. I am indebted to Miss A. M. Buitendijk for 
copies of these. 

The species was only known from a single specimen collected by Dr. 6. 
Salm from herbage at Magelang, Java. 

In 1923, Salm (lac. cit.) gave a very full description in French, which is 
in reality a translation of Oudemans' manuscript so that it appears he must 
have received a copy of this from Oudemans himself. Only figures, very 
crude and inaccurate of an entire dorsal view and of the third leg are given, 
but not Oudemans' drawings. 

In his description Salm states that the unique specimen is in his collection. 
He gives the date of Oudemans 1 original description as Apr. 15th, 1922, but no 
description appeared as far as I have been able to ascertain until July 1st, 
1922. 

The various characteristics of the animal and the different measurements 
given by Salm agree with Oudemans' manuscript, description and the same 
measurements are also given by Walch 1927 (Joe, cit.) when he briefly refers 
to the species. 

The details as given by Oudemans* description and figures are: Length 
450/x, width 267/*. Femora divided into basi- and telofemur. Length of leg I 
225/x, II 200//., Ill 235/x. Chelicerae non-serrate, with an inner subapical for- 
wardly directed tooth. Palpi stout, tibial claw trifurcate; setae on femur and 
genu branched, on tibia dorsal and lateral nude but ventral branched. Eyes 
large, on ocular shields, posterior the smaller. Scutum as figured, with the 
sensillae globose and setulose and inserted in a strongly chitinized wall or crest ; 
anteriorly pitted, posteriorly with semicircular striations; AL scutal setae the 
longest, AM the shortest. Dorsal setae ea. 2,12.12.10.10.10.10.10.10.10.10.10, 
— ca. 116, with a few on posterior border. Ventrally with paired branched 
setae on maxillae, one between coxae I and between coxae III, one on each of 
coxae I and II, and three on coxae III. Leg III without (?) a long nude seta 
on tarsi. 

The Standard Data derived from Oudemans' figures are: AW 78-4, 
PW 100-8, SB 57-4, ASB 36-4, PSB 36-4, SD 72-8, A-P 43-4, AM 35-0, 
AL 71-4, PL 60-2, Sens. 35-0 with head 23-0/23-0. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 269 

Neoschongastia dubia Gunther 1939. 

Neoschongastia inceiia Gunther 1938 (wow. nud.) r Med. J. Aust., 2, (6), 202. 

Neasehongastia dubia Gunther 1939, Proc. Linn. Soc. New South Wales, 64, 
(1-2), 89; Wharton and Hardcastle 1946, J. Parasitol., 32, (3), 294. 

Paraschongastia dubia Womersley and Heaslip 1943, Tr. Roy. Soc. S. Aust., 
67, (1), 130. 

Plate 74, fig. A-C. 

This species is so far only known from the unique type from the New 
Guinea bush fowl, Megapodius duperreyi from Bulolo. 

It is very close to, if not merely a race of, ewingi Wharton and Hardcastle, 
from which it only differs in a somewhat larger scutum with longer scutal 
setae, and rather more dorsal setae. Acording to Wharton and Hardcastle, 
ewmgi does not possess a long nude seta on tarsi III, but a paratype in the 
South Australian Museum collection presented by Dr. Wharton, does possess 
such a seta, as given by Gunther for dubia. On page 310, Wharton and Hard- 
castle (loc. tit.) state that "three (nude setae on tarsi III) are reported for 
dubia". Reference to Gunther 's original description, however, shows that he 
only mentions one such seta. 

In their key to species of Neoschongastia Wharton and Hardcastle separ- 
ate dubia from ewingi by the lateral seta on its palpal tibia being nude instead 
of "with a few delicate barbs". In view of the lack of other more obvious 
differences, and the difficulty, even under oil immersion of detecting such fine 
barbs, it may be doubted whether this is a valid distinction. 

If emngi and dubia are different species, then the specific differences are 
in the number of dorsal setae, and the Standard Data. When more material of 
dubia becomes available it may be established that ewmgi is synonymous. 

Neoschongastia dubia is also very close to N. salmi Ouds., of which I have 
been kindly given copies of Oudemans' original figures by Miss A. M. Buiten- 
dijk of the Leiden Museum, and also a copy of Oudemans' very detailed 
description in German from Oudemans' manuscript in the possession of the 
Entomological Society of Holland, and under the care of Dr. G. L. van Eynd- 
hoven, to whom I am indebted for the copy. These drawings and description 
have not so far been published, but I understand that my colleague, Dr. E} r nd- 
hoven, will shortly do so. 

From Oudemans" drawings and description, both dubia and ewingi are 
very close to salmi. Oduemans' species appears to differ from the others only 



270 Records of the S.A. Museum 

in having considerably more dorsal setae (ca. 126), in the larger scutum with 
scutal setae almost as long as in dubia, and apparently from both in 
having no long nude seta on tarsi III- "With regard to this last feature, in 
view of the fact that it has been shown that Oudemans overlooked such a seta 
in Schongastia vanderswndsi, it may possibly be that he overlooked it also in 
salmi. 

The type of salmi from Java is said to be in the "Finders" collection, but 
its location has not yet been ascertained. 

All three species have trisetose coxae III. 

The Standard Data, of the type of dubia (after Womersley and Heaslip 
1943) are: AW 75-0, PW 98-0, SB 60-0, ASB 25-0, PSB 25-0, SD 50-0, A-P 
38*0, AM 45-0, AL 80-0, PL 71-0, Sens. — . 



Neoschongastia ewtngi Wharton and Hardcastle 1946, 
J. Parasitol., 1946, 32, 308, fig. (larva). 

Plate 75, fig. F, 

Described from Egretta intermedia, Pluviatis dommi^a and Gallus gallus, 
Ulithi Atoll, in August 1945 ; from Gallus gallus on Palau and Peleliu Islands 
in September, 1945; and from Heteroscelus incarvus from Guam, Sept., 1945. 

This species is very close to N. dubia G-unther from which it differs in not 
having a long nude seta (?) on tarsus III and the smaller scutum. Wharton 
and Hardcastle in comparing this species state "that there are no nude setae on 
tarsi III in N. emngi while three are reported from N. dubia"* Actually Gun- 
ther states that there "is only one very long slender nude seta on tarsi IIT", 
and a re-examination of the type of dubia Gunther confirms this. 

The dorsal scutum is as figured (after Wharton and Hardcastle) with the 
sensillae broadly clavate and setulose and with AL very much the longest. All 
setae on the palpal femur and genu and tibia, except the dorsal tibial, are 
branched. The galeal setae have two short branches. 

The DS are from 50 to 21/x long, with short barbs ; they number 60 to 80 
in poorly defined rows of 8 to 10, and roughly in two ill defined groups, an 
anterior of 40-50 and a posterior of about 30. Ventrally coxae III trisetose, 
and posterior of coxae III about 70 setae. 

The Standard Data as given by Wharton and Hardcastle for 9 specimens 
treated statistically are: 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



271 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Ooeff . of 
Variation 


AW 


62-7+1*26 


3-77+0-89 


51-4-74-0 


57-0-69*0 


6-0 


PW 


76-9+1-85 


5-56+1-31 


70-3-93 -5 


70*0-85-0 


7-2 


SB 


50-9+1-26 


3-79+0-89 


39-5-62-3 


45-0-55-0 


7-4 


ASB 


22-0+0-80 


2-40+0-56 


14-8-29*2 


18-0-25-0 


10-9 


PSB 


29-2+0-52 


1-56+0-37 


24*5-33*9 


27-0-32-0 


5-4 


8D 


51-4±0-97 


2-91+0-69 


42-7-60-1 


46-0-56*0 


5-6 


A-P 


33-2±0-40 


1-20+0-28 


29-6-36-8 


32-0-35*0 


3*6 


AM 


31-0+1*0 


2-64+0-77 


23-1-38-9 


28-0-35*0 


8-5 


AL 


57-25+1-43 


4*06+1-01 


45-05-69*45 


50-0-62-0 


7-0 


PL 


40-9±0-65 


1-96+0-46 


35-0-46-8 


38-0-43*0 


4*8 


Sens. 


28-3+1-33 


2-31+0-94 


21-4-35-2 


27*0-31-0 


8*2 



Neoschongastia riversi Wharton and Hardcastle 1946. 
J. ParasitoL, 1946, 32, (3), 299, fig. (larva). 

Plate 74, fig. F. 

This species was described from Eurystomns orientalis, HaUaeetus sanv- 
forcU and Falco severus from Bougainville, the first two hosts in July, and 
the last in October, 1944; and from Halycon chloris and E. orientals from 
Samar, Philippine Islands, in March, 1945. 

The species is remarkable in that there are 3 pairs of sternal setae instead 
of the usual 2 pairs, and the palpal claw is bifurcate. The galea! setae are 
small and nude. On the palpi all the setae on the femur, genu and tibia are 
branched. The dorsal scutum is small, with the anterior margin concave, and 
the posterior margin evenly rounded. The scutal and dorsal setae are finely 
barbed. The DS are 28 in number, from 29/x to 21^ long, and arranged 
2,8.6.2.4.4.2, All coxae unisetose. Posteriorly of coxae III on the venter with 
4.4.2.4.2 setae. On tarsi III there are 4 long nude setae. 

The Standard Data from 10 specimens as given by Wharton and Hard- 
castle treated statistically are: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


38*0+0*77 


2*57+0-55 


30-3-45-7 


34-0-41*0 


6*7 


PW 
SB 


57-4+0-66 
18*6+0*54 


2-20+0-47 
1*80+0-38 


50-8-64-0 
13-2-24-0 


52-0-60-0 
16-0-21-0 


3-8 
9-7 


ASB 


21*7+0-79 


2-61+0*55 


13-9-29-5 


17-0-25-0 


12-0 


PSB 


17-7+0-75 


2-49+0-53 


10-2-25*2 


10-2-25-2 


14-1 


SD 


39-5+1*07 


3-56+0*76 


28*8-50*2 


31-0-44-0 


9-0 


A-P 


24-6+.0-65 


2-16+0-46 


18-1-311 


21-0-28-0 


8-8 


AM 
AL 
PL 


37*4+1*22 
53-5+1-42 
44-0+1-09 


4-03+.0-86 
4-72+1-01 
3-34+0-71 


25-3-49-5 
39-4-67-6 
34-0-54-0 


31-0-43-0 
43-0-60-0 
40-O-49-0 


10-8 
8-1 
7-6 


Sens. 


27*0+0*82 


2*72+0*58 


18-9-35-1 


23*0-31*0 


10*1 



272 Records of the S.A. Museum 

Neoschongastia carvebi Wharton and Eardeastle 1946. 
J. Parasitol., 1946, 32, (3), 313, fig. (larva) fig. (nymph). 

Plate 75, fig. C. 

Described from larvae from Numtendm phaeopw, Demigretta sacra, Tlw- 
vialis dominion, and Arenaria interpres from Guam, the first in June and Sep- 
tember, 1945, the second and third in July, 1945 and the fourth in Septem- 
ber, 1945 ; from Okinawa from Gygis alba and Sterna dowgalUi (sic douglaii) 
in July, 1945, and from Monticola solitarius in August, 1945; from Phiiwlis 
dominica from Ulithi Atoll in August, 1945 and from Heteroscehut incanus from 
Peleliu Island in September, 1945. 

From a larva from D. sacra from Guam the nymph was reared by Wharton 
and Hard-castle (see section on Adults and Nymphs). 

The larvae of this species are distinguished from all others in that there 
are numerous setae, ca. 20-22, situated between the coxae and arranged in irregu- 
lar rows. Coxae III is multisetose, with from 6-8 setae on each; I and II 
imisetose. 

The dorsal scutum is characteristic in shape, as figured (after Wharton 
and Hardcastle) with PW being only very slightly greater than AW, and the 
lateral margins being straight and parallel except at the posterior corners; 
the anterior margin is strongly sinuous and the posterior margin straight except 
in the middle third where it is deeply incised. The PL scutal setae are very 
long and much longer than AM or AL. The sensillae are very broadly clavate 
and setulose with the bases nearer to AL than to PL. 

The galeal setae are branched. The palpal setae of the femur, genu and 
tibia are all branched except the dorsal and lateral tibial. 

The dorsal setae are long, from 98/* to 66//. and posterior of the humeral 
setae number ca. 100. On the venter the setae posterior of coxae III number 
about 100. Tarsus III without any long nude seta. 

The Standard Data from 5 specimens given by Wharton and Ilardeastle 
treated statistically are: 





Mean. 


Standard 
Deviation 


Theoretical 
Bange 


Observed 

Range 


Coeff. of 
Variation 


AW 


79 -8:2:3 -09 


6-91=t2-18 


59- 1-100 -5 


72*0-90-0 


8-6 


PW 


83-8±:2-75 


6 -14:tl -94 


65 -4-102 -2 


78-0-93-0 


7-3 


SK 


43-4:4:3 -69 


8-26^2-61 


18-6-68-2 


36-0-57-0 


19-0 


ASE 


30-0±0-55 


l-22:t0-39 


26 '3-33 -7 


28-0-33-0 


41 


PRB 


36-6±:0-60 


1 »34^=0-42 


32- 6-40-6 


36-0-39-0 


3-7 


ST) 


66-6:±l-00 


2*30:fc0*73 


59*7-73-5 


64-0-69-0 


3-4 


A-P 


45*2:2:1-35 


3«03±0-96 


36-1-54-3 


42*0-48*0 


6-7 


AM 


70-8=t3-50 


7*82:£2*47 


47*4-94-2 


60-0-81-0 


11-0 


AL 


73-8:t2-94 


6-57±2-08 


54- 1-93-5 


66-0-84-0 


9-0 


PL 


137-0±8-72 


19-49±6-16 


78-5-195-5 


108-0-162-0 


24*2 


Sens. 


40*2:±2-24 


5-02±l-58 


25-2-55-2 


36-0-48-0 


12-5 



WOMER5LEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 



273 



KEY TO THE ASIATIC-PACIFIC SPECIES OF NEOSCHONOASTIA EWING 1929, 

S. STR. 1946. 



1. Sternal setae of two pairs only. Palpal claw trifureate 
More than two pairs of sternal setae 

2. Coxae III with only 1 seta 

Coxae III with more than 1 seta . . 



3. Galeal setae nude 
Galeal setae branched 



2 

22 

3 

10 

4 
8 



basal segment of the 



4. With a distinct hemisphaerieal elevation on the 

chelicerae. DS ea. 9/J, with only 1 humeral seta on eaeh side, varying 
from 57>x long anteriorly to 33/* posteriorly. Setae on palpal femur 
and genu with branches ; on tibia with ventral branched, dorsal and 
lateral nude. 

AW 58-75=tl0-5, PW 87-75±:ll-6, SB 29*25 + 6-7, ASB 29-75± 
3-8, PSB 26-0±10-l, SD 55- 75:1:7 *1, A-P 41-75±12-1, AM 60-75 
±9-9, AL 60-5±:l-7, PL 60-25=hl-5, Sens. 35-5±5-7. 

N. bougainvilUrms Wharton and Hardcastle 1946. 

Without such a hemisphaerieal elevation on basal segment of chelicerae 5 

5. Dorsal setae more than 50, ca. 72 in number, with only 1 humeral, 

from 35/a long anteriorly to 17/x posteriorly. Seta on palpal femur 
branched; on genu nude; on tibia ventral branched, dorsal and 
lateral nude. No long nude seta on tarsi III. 

AW 40*0:t5-2, PW 65-3:fc6-2, SB 13-0±6«0, ASB 33-7±3*4, 
PSB 14-3±:3-4, SD 48-0+6-0, A-P 34-3+4-6, AM 24-3+ -76, 
AL 36-7+6-2, PL 38-7+16-5, Sens. 21-5+2-1. 

N. stron-gi.'WhBLTton and Hardcastle 1946. 

Dorsal setae fewer than 50 in number < . , . . . . . , . 6 

6. Head of sensillae spindle-like with long fine setules. DS 40 in num- 

ber, arranged 2.8.6.6.6.6.4.2, from 62/* long anteriorly to 39* posteri- 
orly. Setae on palpal femur and genu branched; on tibia ventral 
branched, dorsal and lateral nude. A long nude seta on tarsi IIL 

AW 55-8+8-0, PW 77-8+13-2, SB 32-4+5-5, ASB 24-6+4-0, 
PSB 26-4+8-4, SD 51-0+12-2, A-P 27-2+1-3, AM 42*6+18*6, 
AL 35-0+5-6, PL 75-2+18-8, Sens. 70-75+17-0. 

N. montic&la Wharton and Hardcastle 1946. 

Head of sensillae globose . . . . . . . . . . . . . . 7 

7. With a long nude seta on tarsi III. DS 34 in number, arranged 

2.8.6,8.4,2.2.2, from 52/l long anteriorly to 43/a posteriorly. Setae on 
palpal femur and genu branched; on tibia ventral and lateral 
branched, dorsal nude. 

AW 75-2+9-8, PW 79-8+8-9, SB 47-4+9-1, ASB 25*4+6-9, 
PSB 33-6+7-8, SD 59-0+9-0, A-P 29-4+4-5, AM 49-0+7-6, 
AL 74-8+29*6, PL 52-4+10-5, Sens. 29-75+10-5. 

N. posekawyi Wharton and Hardcastle 1946. 



274 Records of the S.A. Museum 

Without any long nude seta on tarsi III. DS 44 in number, arranged 
2.12.8.6.10.4,2. Setae on palpal femur and genu branched, on tibia 
!. PL the longest. 

AW 52-0, PW 60-0, SB 42-/), ASB 19-0, PSB 26*0, SD 45-0, 
A-P 30-0, AM 32-0, AL 35-0, PL 50-0, Sens. 26-0. 

N. galUnafrwm Hatori 1920* 

8. Posterior body setae arising from platelets. Striations and pitting 

weak on posterior sixth of dorsum. DS 72, arranged 2.14.14.10.8.8. 
6.6.2.2, to 30-40/1 long. Setae on palpal femur and genu branched; 
on tibia lateral and ventral branched, dorsal nude. No long nude 
seta on tarsi 111, 

AW 58-0, PW 72-0, SB 48-0, ASB 23-5, PSB 25-0, SD 48-5, 
A-P 42-0, AM 37-5, AL 47-0, PL 42-0, Sens. 30-0. 

N. backko-usei Gunther 1939. 

Posterior dorsal setae similar to anterior and not on platelets . . 9 

9. DS 68-76 in number, arranged roughly in rows of 10 except pos- 

teriorly, from 45/i long anteriorly to 21/*. posteriorly. Setae on palpal 
femur and genu branched; on tibia ventral branched, lateral and 
dorsal nude. Punctation on scutum fine and scattered. 

AW 65-3d:8*7, PW 73-25d=4-3, SB 53-0±:7-3, ASB 21-75±5-7, 
PSB 30-5d:ll-l, SD 52-25+15-6, A-P 31-5±7-5, AM 27*7+ 
10-1, AL 57-3^6-4, PL 37-0±:7-3, Sens. 31*0. 

N. egretta Wharton and Hardcastle 1946, 

DS 32, arranged 2.8.6.6.4.4.2, to 54/x long. AL almost twice as long 
as PL. Setae on palpal femur and genu branched; on tibia dorsal 
nude, lateral and ventral branched. Scutum with sparse but large 
punctations or tuberculations. 

AW 75*5+6-8, PW 85-0-b5-7, SB 43*0+4-6, ASB 22-5+4*9, 
PSB 30-3+2-5, SD 52-8 + 6-4, A-P 33-0. AM 57-6+4-0, AL 
85-8+6-1, PL 51-75+8-6, Sens. 32-0. N. st ruiMdfa sp.n . 

10. Some posterior dorsal and ventral setae specialized and spine-like, 

and arising from platelets . . . . . . . . . . . . 11 

Posterior dorsal and ventral setae similar to anterior . . . . . . 14 

11. Coxae IIT bisetose . . . . . . - . ■ . - . - • - • 12 

Coxae III tri-, rarely quadri-setose . * 13 

12. Posteriorly, both dorsaUy and ventrally with a ring of strong long 

spines arising from small platelets, and inside the ring a number of 
more or less rectangular plates each with a short nude seta. Palpal 
claw long and over-reaching tip of tarsus by tarsus length. DS 52, 
arranged" 2.8(10), 12(10), 6.8(10), 8.8(6). * Setae on palpal femur 
and genu branched; on tibia dorsal nude, lateral and ventral 
branched. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 275 

AW 71-4+6-0, PW 84-5+11-5, SB 55-5 + 6-0, ASB 28-0, PSB 
21*3+6-0, SD 44-4+4-1, A-P 35-25+2-1, AM 36-0, AL 72-0 + 
8-5, PL 51-4+11-8, Sens. 36-0, iff. reiroeincta Gunther 1939. 

Posteriorly no such circle of spines. Palpal claw shorter and 
stouter, only slightly exceeding tip of tarsus. DS ca. 100, arranged 
2, plus six rows of 8 to 16, then 10(8), 6.4.4. Setae on palpal femur 
and genu branched; on tibia dorsal and lateral short and nude, 
ventral branched. 

AW 77-15+4-4, PW 103-1+16-7, SB 64-1+5-0, ASB 30*5+ 
2-8, PSB 28-3+2-8, SD 58-8+4-2, A-P 36-4+5-9, AM 39-2+. 
9-7, AL 83-1+7-4, PL 55-6+3-2, Sens. 39-2. 

N. yeomcmsi Gunther 1939. 

13. DS ca. 120 including ca. 46 spines posteriorly. Scutum larger. 
Coxae III normally 3-setose, but frequently 4-setose, 

AW 79-7+6-5, PW 93-6+6*8, SB 62-15+5*1, ASB 28-3+2-5, 
PSB 27-45+3-3, SD 55-7+3-9, A-P 35-3+4-6, AM 34-8dr6*4, 
AL 80 - 8+9 - 6, PL 51 - 6+ 9 - 1, Sens. 39 • 2. N. owiensw sp. n. 

DS <»a. 88 including ca. 28 spines posteriorly. Scutum smaller. 
Coxae III 3-setose (very rarely 4-setose). 

AW 65-6+4-6, PW 74*5+11-3, SB 47-6, ASB 26-6+3-0, PSB 
22-4, SD 49-0+3-0, A-P 32-7+4-3, AM 32-2+4*9, AL 57-4+. 
4-9, PL 42-0, Sens. 28-0. - N. mcqueeni sp . n. 

14. With more than 5 setae on coxae III . < . . ♦ - . . . . 15 
With fewer than 5 setae on coxae III . . . , . . . . 16 

15. Galeal setae nude. PW greater than AW. Coxae III with 7 setae. 

DS ca. 120 in number in rows of 10 or fewer, and with about 4 
humeral setae on each side, 49^ long anteriorly to 21/* posteriorly. 
Setae on palpal femur and genu branched; on tibia, dorsal and 
lateral nude, ventral branched. 

AAV 46-5+7-0, PW 69-2+21*2, SB 20-5+11-6, ASB 19-5+7-7, 
PSB 31-2+9-6, SD 50-7+14-9, A-P 41*7 + 14-1, AM 26-2+8-4, 
AL 47-7+17-7, PL 33-0+8-0, Sens. 28-0+16-9. 

N. atolhiisis Wharton and Hardcastle 1946. 

Galeal setae branched. PW less than AW. Coxae III with 10 
setae. DS ca. 205, with about 15 humeral setae on each side, and 
then ca. 175, to 44/x long anteriorly, 32fc posteriorly. Setae on palpal 
femur, genu and tibia all branched. 

AW 74-4+9*3, PW 60-2+16-5, SB 40-0+5-4, ASB 34-4+12-8, 
PSB 40-0 + 3*8, SD 74-4+14-8, A-P 46-2+4-0, AM 61-0+11-2, 
AL 70*4+4-5, PL 70-4+22-8, Sens. 36-7+17-1. 

N. namrni Wharton and Hardcastle 1946. 



276 Records of the S.A. Museum 

16* Body setae fewer than 50 . . . „ . . - - . . . . 17 

Body setae more than 50 , . . . . . . . ►- .< * • 19 

17. DS 32 in number, arranged 2.8.6.4.6.4.2, from 68/x anteriorly to 28/i 
posteriorly. All setae on palpal femur, genu and tibia branched. A 
long- nude seta on tarsi III. 

AW 51-7+12-5, PW 70-2+18-0, SB 34-0+12-9, ASB 25-7+6-4, 
PSB 29-4+4-2, SD 55-2+6-6, A-P 35-6+4-5, AM 43-6+13-4, 
AL 55-6+17-1, PL 57-1+20-6, Sens, 25-8+9-1. 

N. amerkana solonionis Wharton and Hardcastle 1946. 

DS fewer than 30 . . . . • • » • • * • - • « 18 

18. DS 26 in number, arranged 2.8.6.4.4.2, to 30,i long. PL the longest. 
Setae on palpal femur and genu branched ; on tibia, ventral strongly 
branched, lateral with 2 branches, dorsal nude. No long nude seta 
on tarsi III. 

AW 47-1+6-3, PW 68-9 + 7-3, SB 30-6+6*4, ASB 20-8+4'3, 
PSB 29-4+4-3, SD 50-2+5-2, A-P 32-4+6-4, AM 31-85+6-2, 
AL 37-8+6-4, PL 45-8+7-6, Sens. 30-8. N. entomyzasp. n. 

DS 24 in number, arranged 2.6.6.4.4.2, to 50/* long. PL and AL sub- 
equal, AM the shortest. Sensillae apparently nude. 

AW 44-0, PW 63-0, SB 23-0, ASB 20-0, PSB 23-0, SD 43-0, A-P 
34-0, AM 37-0, AL 51-0, PL 54-0, Sens. 29*0, (after Radford). 

N. thomaxi (Kadford 1946) . 

19. With as many as 3 humeral setae on each side. Coxae III 3-4 setose. 
DS ea. 56 in indistinct rows of 10, anteriorly 55/x long, posteriorly 
30/i. Galeal setae barbed. Setae on palpal femur and genu branched ; 
on tibia dorsal and lateral nude, ventral branched. No long nude 
seta on tarsi IIL 

AW 57-5+5-2, PW 77-0, SB 37-25+6-5, ASB 22-0, PSB 30-7 + 
10-5, SD 52-5+14-9, A-P 31-25+4-5, AM 35-0+8-8, AL 52-5+ 
14-9, PL 54-75+4-8, Sens. 29-3+4-6. 

JV, paxiensis Wharton and Hardcastle 1946. 

With only 1 humeral seta on each s5de. Coxae III 3-setose . . . , 20 

"20. Apparently without a ltong nude seta on tarsi III. DS ca. 116-120, 
arranged ca. 2.12.12.10.10.10.10.10.10.10.10.10.+. 

AW 78-4, PW 100-8, SB 57-4, ASB 36-4, PSB 36-4. SD 72-8, 
A-P 43-4, AM 35-0, AL 71-4, PL 60-2, Sens. 35-0 (after Oude- 
mans ) . N. salmi ( Oudcmans 1922 ) . 
With a long nude seta on tarsi III 21 

21. DS 90-96 in number, arranged ca. 2.14.10.12.6.14.14.12.8.4, to 30,*. 
long. AL the longest but only slightly longer than PL. Setae on 
palpal femur and genu branched; on tibia dorsal and lateral nude; 
ventral 1 -branched. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 277 

AW 75*0, PW 98-0, SB 60*0, ASB 25-0, PSB 25-0, SD 50-0, 
A-P 38-0, AM 45*0, AL 80*0, PL 71-0, Sens. — 

& dubia Gunther 1939. 

DS 60-80 m number, in two ill-defined areas, anteriorly to 50/* long, 
posteriorly to 21/*. Setae on palpal femur and genu branched; on 
tibia dorsal nude, lateral with a few delicate barbules, ventral 
branched. 

AW 62-7+11-3, PW 78-9+16-7, SB 50-9+11-4. ASB 22-0+7-2, 
PSB 29-2+4-7, SD 51-4+8*7, A-P 33-2+3-6, AM 31-0+7-9, 
AL 57-25+12-2, PL 40-9+5-9, Sens. 28-3+6-9. 

N* ewingl Wharton and Hardcastle 1946. 

22. With three pairs of sternal setae. Palpal claw bifurcate. All coxae 
1-setose. DS 28 in number, arranged 2.8.6.2.4.4.2, from 29/* to 21/* 
long. All setae on palpal femur, genu and tibia branched. Galeal 
setae nude. 

AW 38-0+7-7, PW 57-4+6-6, SB 18*6+5-4, ASB 21-7+7-8, 
PSB 17-7+7-5, SD 39-5+10-7, A-P 24-6+6-5, AM 37-4+12-1, 
AL 53-5+14-1, PL 44-0+10-0, Sens. 27-0+8-1. 

N. riversi Wharton and Hardcastle. 1946. 

With numerous unpaired sternal setae. Palpal claw trifurcate. 
Coxae III 6-setose. DS ca. 120, with 8-10 humeral on each side, from 
93/* long anteriorly to 66/* posteriorly. Setae on palpal femur and 
genu branched; on tibia dorsal and lateral nude, ventral branched. 
Galeal setae branched. 

AW 79-2+20-7, PW 83-8+18-4, SB 43-4+24-8, ASB 30-0+ 
3-7, PSB 36-6+4-0, SD 66-6+6-9, A-P 45-2+9-1, AM' 70-8+ 
23-4, AL 73-8+19-7, PL 137-0+58-5, Sens. 40-2+15-0 (from 
Wharton and Hardcastle 's data). 

N.. wrveri Wharton and Hardcastle 1946, 

Oenus MACKIENA Traub and Evans 1950. 

J. Washington Acad. Sci, 1950, 40, (4), 126. 

This remarkable genus is characterized chiefly by the empodium of all 
legs, instead of being claw-like, being expanded to form something like a pad 
or sucker such as occurs in many other groups of Acarids, but hitherto unknown 
in the Trombidiid families. The authors relate the genus to Eiedlinia Ouds* 
1914 T but in that genus the empodium although swollen is not the same as in 
MacktCTia and may still be regarded as claw-like. 

The sensillae are globose, with only indications of a crest at their bases. 
The caudal portion of the scutum with distinct concentric striations as in* 
N eoscMngmtia, 

Genotype Mackiena empodiformis Traub and Evans 1950. 



278 Records of the S,A. Museum 

Macktkna empodiformis Traub and Evans 1950. 
J. Washington Acad. Sei., 1950, 40, (4), 126-129, fig. 1-4. 

Plate 68, fig. E-I. 

Description of Larvae (after Traub and Evans). Shape ovate-subcireular. 
Length 230/*, width 185ft; Scutum (as figured), anterior margin slightly sinuate 
with one median and two lateral lobes, lateral margins evenly convex, posterior 
margin fairly deep behind line of PL and flat medially • surface finely punctate 
annually, with concentric striations posteriorly; sensillae in line of PL, globose 
and setulose, with a Blight indistinct crest in front of bases; AL very much 
longer than AM or PL. Eyes 2 + 2, well developed, posterior the smaller. 
Chelicerae non-serrate but with well pronounced apical tricuspid cap and dorsal 
subapical tooth. Galeal setae ?. Palpi stout, tibial claw trifurcate; seta on 
femur well branched, on genu with 2 short branches, on tibia dorsal and 
lateral nude and ventral with one branch. Dorsal setae strong and shortly 
ciliated or barbed, arranged 2.6.4.6.6.4.2 = 30, to 60/x. Ventrally, a pair of 
ciliated setae on maxillae, one on each coxae, a pair between coxae I and 2 
pairs between <?oxae III, thereafter ca. 36 setae, arranged 10.8.8.-f. Legs: I 
260//., II 250/*, III 270/ji; tarsi I and II with dorsal sensory rod (spur), III 
without any long nude seta. 

The Standard Data for the type and paratype as given by the authors are : 
AW PW SB ASB PSB SD A-P AM AL PL Sens. 
Type 56*0 76*0 37-0 21-0 21*0 42-0 19-0 46-0 84-0 66.0 3.0 
Para- 
type 57-0 78-0 38-0 — — — 42-0 84-0 67-0 — 

hoc. mid Host. This species was described from only two specimens from 
a weaver-finch {Floceus many or peguenm Stuart Baker) from 20 miles north- 
west of Myitkyina, Burma, on Mogaung Rd., Feb. 23/1945 (U.S. Typhus Com.). 

Remarks. The host being a bird supports the close association of this genus 
and species with the genus Neoschongastia Ewing, as suggested by the concen- 
tric scutal striations. 

Subfamily GAHRLIEPIINAE nom. nov. 
= Walchiinae- Ewing 1946, J. Parasitology, 32, (5), 435, 

Trombiculidae in which the dorsal scutum of the larvae is without the AM 
seta, and in which tarsi I of the nymphs and adults have a short dorso-apical 
stump-like process. 

Containing only the single genus Gahrlupia Oudemans 1912. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 279 

Genus GAHRIJEPIA Ouds. 1912. 

= Typhlothrmxbmm Oudemans 1910, Ent, Ber. Ned. Ent. Ver., 3, 102. 

Gahrliepia Oudemans 1912, Ent. Ber., Ned. Ent. Ver., 6, 237; Sig Thor and 
Willmann, Das Tierreich 1947, 71b, 332. 

Schongastiella Hirst 1915, Bull. Ent. Res., 6, 183; Sig Thor and Willmann 
1947, Das Tierreich, 71b, 334. 

Walchia Ewing 1931, Proc. U.S. Nat. Mus., 80, (8), 10; Sig Thor and WilJ- 
mann 1947, Das Tierreich, 71b 7 335. 

Gateria Ewing 1938, J. Wash. Aead. Sci., 28, (6), 295; Sig Thor and Willmann, 
1947, Das Tierreich, 71b, 338. 

The genus Typhlothromhimn was founded on the single species T. nanus 
Ouds. 1912 from S. Africa. In. 1912 Oudemans proposed Gdhrliepia in ita 
place, as Typhloihrombmni had been used earlier for an adult genus of Troni- 
bidiid mites. In Gahrliepia as represented by the genotype the dorsal scutum 
in addition to the AL and PL setae, carries two pairs of additional setae* In 
1915, Hirst erected the genus Schongastiella for S. bengatensis in which the 
scutum was furnished with only one pair of setae in addition to the AL and 
PL setae. Ewing proposed the genus Walchia in 1931 with Trombicida glab- 
rwm Walch from the Dutch East Indies as the genotype for those species in 
which the scutum was somewhat pentagonal but did not carry any setae other 
than AL and PL. 

The genus Gateria was proposed by Ewing in 1938 with Gahrliepia fletck- 
eri Gater as type, for those species of Gahrliepia in which most of the addi- 
tional scutal setae are not marginal. 

In 1943, Womersley and Heaslip showed that the number of additional 
scutal setae in the genera Gahrliepia, Schongastiella and Gateria were really 
correlated with the prolongation in a posterior direction of the scutum, which 
thereby embraced some of the median setae of some of the dorsal rows. These 
three genera were therefore regarded as synonymous. 

Ewing (1938) in his key to the larval genera of the Trombiculidae gives 
for Schongastiella Hirst "eyes present;" for Walchia Ewing "eyes absent" 
and for Gahrliepia "eyes usually present/' It is frequently very difficult in 
microscopic mounts of these mites to be reasonably certain as to whether eyes 
are truly present or not, but in the species recognized by Womersley and 
Heaslip 1943 as falling into Gahrliepia (in the wide sense, i.e. including 
Schongastiella and Gateria) eyes are absent in nanus Ouds. ; in Walchia eyes 
are present in disparunguds and nwrobwsis and absent in all other species. Of 
the following species described by Badford 1946, Gateria- hirsuta-, crocidnra, 



280 Records of the S.A. Museum 

Schongwstiella brevis, lancearia, ligula and punctata, eyes apparently are absent 
except in the last where they are relatively small. 

It seems, therefore, that the presence or absence of eyes is of no more 
generic importance than the number of additional setae on the scutum, and 
that all four genera should be considered synonymous, with priority to Gahr- 
liepia Ouds., or at most, of not greater than subgeneric status. 

Further support for this is given in the section of this paper dealing 
with the adults and nymphs, where it is shown that all four genera, in the 
known reared nymphs have a characteristic short dorso-apical process on 
tarsi I but otherwise only differ in specific characters, such as the dorsal setae. 

In the following key to the known larval species from the Asiatic-Pacific 
Region, the various genera which have been proposed are only given sub- 
generic rank for convenience. 

Key to the Species, 

1. Scutum small, pentagonal, wih posterior margins forming a more or 

less acute angle. Only AL and PL setae on scutum 2 

Subgen. Walchia Ewing. 

Scutum usually larger, not pentagonal, frequently produced tongue- 
like beyond PL and taking in two or more setae from the dorsal rows 8 

2. Coxae III unisetose . . . . . . . . . . . , mm 3 

Coxae III multisetose , . . . . . . . . . . . . . 5 

3. Scutum small, AW smaller than 32-0/*. AL, Sens, bases, and PL in 
approx. the same longitudinal line. Sides of scutum between AL and 
PL convex. Sensillae clavate, apex aciculate, strongly setulose. 

AW 29-5d=6-7, PW 32-9+4-1, SB 28*1+4-5, ASB 19<2+5*9, 
PSB 44-0+4-1, SD 63-2+4-5, A-P 35-4+3-2, AL 27-4+7-3, 
PL 29*5+5-3, Sens. 25-6 with head 9 -6/19 -2. 

Gahrliepia (Walchia) morobemis Gunther 1939. 

Scutum larger, AW greater than 32-0^. AL, Sens. baBes, and PL 

not in the same longitudinal line ~ . . . . . . . . . 4 

4. Scutum with PSB not greater than PW; posterior scutal angle shal- 
low. Dorso-apical tooth of ehelieerae baekwardly directed. 

AW 36-6+3-1, PW 44-5+2-6, SB 29-7+2-4, ASB 19-7+4-5, 
PSB 44-1 + 4-2, SD 63-75+3-6, A-P 35-7+3*95, AL 30-0, PL 
30*0, Sells. — . 

GahrUepia (W<ilcJiia) mstica Gater 1932. 
Scutum with PSB greater than PW, posterior scutal angle deeper . . 4a 

4a. Only 1 humeral seta on each side Dorsal-apical tooth of chelicerai 
tricuspid cap blunt and not directed backwards. Coxae IIT 1 -setose. 
No setae between coxae IT and TIL 



Womersley— Asiatic-Pacific Scrub Typhus Mites 281 

AW 38-35±4-15, PW 48-6=b6-3, SB 32-4±4-4, ASB 22-5=fcl-5, 
PSB 56-7±7-3, SD 79-4+7-8, A-P 40-65+4-45, AL 32-9+4-9, 
PL 32-6+4-4, Sens. 32-0 with head 9-6/22-4. 

Qahrliepia (Walchia) iurmalis Gater 1932. 

With 2-3 humeral setae on each side. Ventrally between coxae II 
and III 3-4 setae. 

(Standard Data as in Gahrliepia (Walchia) brennani sp n.). 
Qahrliepia (Wafichia) brennani var. centralis nov. 

5. Scutum very small, with AW smaller than 20/i. Coxae III 6-setose. 
Dorsal setae 2.6*6.6.6.2.2. Palpal claw trifurcate. 

AW 19-5+2-1, PW 31-65+2-75, SB 17*25+3*0, ASB 19-3+ 
2-5, PSB 28-0, SD 47-3+2-5, A-P 30*4+3-0, AL 14-0, PL 14-0, 
Sens. 19-6 with head 11/14. 

Gahrliepia (Walchia) lewthwaitei Gater 1932. 

Scutum larger, AW greater than 20/a. Coxae III not 6-setose . . 6 

6. Coxae III 2 setose. With only 1 humeral seta on each side : . 6a 

6a. Coxae III more than 2-setose . . . . . . . . . . . * 7 

AW 27-45±4-8, PW 44-6+4-85, SB 23*3+3-8, ASB 20-3+3-8, 
PSB 36-1+4-55, SD 56-4+4-6, A-P 32-8+4-3, AL 22-4, PL 
28-8, Sens. 25-6 with head 9-6/19*2. 

Gahrliepia (Walchia) disparimgws (Ouds. 1929). 

With 2-3 humeral setae on each side. Scutum larger. Dorsal setae 
32-34 in number, to 30^ long and arranged 6 (4). 6.6.6.6.2.2. 

AW 37-5+12-0, PW 54-1+14-4, SB 32-8+16-8, ASB 21-8, PSB 
57-8+12-9, SD 79-6+12-9, A-£ 37-6+12-0, AL 27-3+10-8, 
PL 26-8, Sens. 33-5 with head 10-05/23-45. 

GaJirlieph (Walchia) brennani sp. n, 

7. Coxae III normally 3-setose, very rarely with 3 + 4. Palpal claw 
bifurcate. Dorsal setae 2.6.6.6.6.4.2. 

AW 28-4+3*2, PW 48-8+8-8, SB 25-0+3-6, ASB 22-4, PSB 
34-3+4-3, SD 56-7+4-3, A-P 36-7+8-1, AL 22-4, PL 28-8, 
Sens. 28-8 with head 11-2/22-4. 

Gahrliepia (Walchia) Gahrliepia (Walchia) ewingi Fuller 1949. 

= glabra- (Walch 1927, emend. Fuller 1948). 

Coxae. Ill normally 4-setosc, but very variable from 4+3 to 5 + 6. 
Palpal claw apparently trifurcate. Dorsal setae 2.8.6.6.4.4,2. 

AW 26-5+4-4, PW 37*6+6-4, SB 21-0+4-4, ASB 18-9+2-3, 
PSB 32*75+5-95, SD 52-0+6-0, A-P 30-95^4-1, AL 19-2+ 
2*3, PL 20- 7+4-1, Sens. 22-4 with head 12-8/16-0. 

Gahrliepia (Walchia) enode (iGater 1932). 

8. Dorsal scutum produced, tongue-like, behind PL, but with only the 
AL and PL setae; much longer than wide with the posterior end 
lightly concave. Eyes absent. Coxae III 2-setose. SB very much 
nearer to PL than to AL. 



282 Records of the S.A. Museum 

AW 30-0, PW 50*0, SB 25-0, ASB 18-0, PSB 58-0. SD 76-0, 
A-P 24-0, AL 28-0, PL 40 -0, Sens. — , (After Gunther 1940; 
Womersley and Heaslip 1943). 

GahrUepia (? WalcMa) rioi Gunthcr 1940. 
Dorsal scutum produced, and besides AL and PL with additional 
setae belonging to the dorsal rows . . . . . . , . . * 9 

9. Dorsal scutum with only 2 setae, in addition to AL and PL, and 

these from the second dorsal row 10 

Subgen. Schongastiella Hirst 1915. 

Scutum with 4 additonal setae besides AL and PL 15 

Subgen. GahrUepia Ouds. 1912 s. str. 

Dorsal scutum with more than 4 additional setae, and these from 

the second, third or more dorsal rows ♦ . 20 

Subgen. Qateria Ewing 1938. 

10. Dorsal scutum very sharply contracted posterior of PL, ligulate. Sen- 
sill a e more or less globose and setulose ► . . - 11 

Dorsal scutum otherwise . . . . . - , . . . * . . . 12 

11. Distance A-P of scutum at least equal to or greater than distance 
between PL and the posterior pair of scutal setae. Coxae III 1- 
setose. Dorsal setae ca. 2.8 [2]. 8.4.8.6.6.2.2. 

AW 36-8*5*6, PW 51-9*6-9, SB 35-6*4-7, ASB 26-5*4-1, 
PSB 65-5*9-6, SD 92-0*11-1, A-P 44-7*5-15, AL 42-0*3-2, 
PL 42-0, Sen's. 33-6 with head 14-0/25-2, 

GahrUepia (SchSnga&ticlta) Ugida (Radford 1946). 
Distance A-P of scutum much less than distance from PL to pos- 
terior pair of setae. Coxae III 4-setose (sometimes 5-6 setose). 
Dorsal setae 2.6 [2] .8.8.6.4.4 (2). 2(0). 

AW 31-0. PW 28-0, SB 23-0, ASB 14-0, PSB 55-0, SD 69-0, 
A-P 28-0, AL 20-0, PL 20-0, Sens, 20-0 wih head 14-0/14-0. 

GahrUepia {Schongastiella) cvylomca sp. n. 

12. Scutum about as long as broad; distance from line of PL to apex 
less than distance between PL. Sens. ? Dorsal setae 2.6 [2] .6.8.4.2. 
Coxae III 2-setose. 

AW 44*0, PW 86-0, SB 51-0. ASB 26-0, PSB 78-0, SD 104-0, 
A-P 43-0, AL 38-0, PL 41-0, Sens. — . (After Kadford). 

GahrUepia (Schongastiella) brevis (Radford 1946). 
Scutum much deeper from line of PL to apex than distance between 
fcL. Sens, clavate. Coxae III 1- or multisetose . . . . . . 13 

13. Margin of scutum posterior of PL more or less an even parabolic 
curve. Coxae III 1-setose. Dorsal setae 2.6 [2] .8.8.6.4.2.2. 

AW 36-7*4-7, PW 50-6*6-5, SB 31-7*8-0, ASB 21-1*8-1, 
PSB 581*151, SD 79*2*9-2, A-P 37-6*5-0, AL 31-45*9-7, 
PL 35-8*1-3, Sens. 33-0 with head 15/21. 

GahrUepia (Schongastiella) bengalensis (Hirst 1915) 
Margin of scutum posterior of PL more or less triangular. Coxae III 
2- or 4-setose . . . . . . . . . . . . . . . . 14r 



Womersley— Asiatic-Pacific Scrub Typhus Mites 283 

14. Coxae III 2-aetose. Dorsal setae 2.4 [2]. 6.6.6.4.2.2. Galeal and seta 
on palpal femur branched. 

AW 45-2+7-1, PW 68-5+13-2, SB 43-5+7-6, ASB 22-7+2-8. 
PSB 91-5+29-3, SD 114-1+28-8, A-P 44-9+6-6, AL 38-4, PL 
38-4, Sens. 35-2 with head 9-6/22-4. 

Qahrliepia (Schongastiella) punctata (Radford 1946). 

Coxae III 4-;setose. Dorsal setae 2.6 [2]. 6.6.6.4.2. Galeal and seta 
on palpal femur nude. 

AW 51-2, PW 76*8, SB 49*6, ASB 28-0, PSB 110-0, SD 138-0, 
A-P 51-2, AL — , PL 41-6, Sens. — . 

Gahrliepia (Schongastiella) kumaonensis sp.n . 

15. Scutum markedly ornate, with 4 additional setae 16 

Scutum at most only minutely punctate . . . . . . . , , . 18 

16. PL behind line of SB. Scutum with large, nearly contiguous square 
to hexagonal reticulations. DS 32 in number, arranged 2.6 [2]. 6 [2], 
6.6,4.2. 

AW 44-6, PW 75-6, SB 44-6, ASB 22-4, PSB 173-6, SD 196-0, 
A-P 30-8, AL 39-2, PL 53 - 2, Sens. — . 

Gahrliepia (Gakrlispui) insignesp.Ti, 

PL in front of line of SB 17 

17. Scutum uniformly with widely separated, more or less circular 
depressions. DS 34 in number, arranged 2.6.6 [2]. 6 [2] .4,4.2,2.2. 

AW 41-6+3-2, PW 66-8+5-8. SB 30-8, ASB 22-4, PSB 239-6+ 
31-7, SD 262-0+31-7, A-P 17-2+5-8, AL 26-8+4-5, PL 108-8 
+12-3, Sens. — . Gahrliepia (Gahrliepia) decora sp. n. 

Scutum medially with a longitudinal cluster of raised tubercles, and 
laterally around the margin, large squarish depressions. DS 

2.4 [2] .8 [2] .8.8.6.4.2. 

AW 40-0, PW 59-0, SB 33-6, ASB 25-2, PSB 218-4, SD 243-6, 
A-P 12-6, AL 28-0, PL 98-0, Sens. — . 

Gahrliepia (Gahrliepia) ornata sp. n. 

18. Scutum with only 8 (occasionally 6-9) setae, excluding the sensillae 19 
Scutum with more than 8 setae, excluding the sensillae . . . . 20 

19. Coxae III nnisetose. Scutum larger. Dorsal setae 2.6 [2] .6 [2] .6.6.4.4.2. 

AW 42-0, PW 90-0, SB 49-0, ASB 35-0, PSB 155-0, SD 190-0, 
A-P 65-0, AL 40-0, PL 55-0, Sens. 37-0 with head 12/30 (after 
Gater; and Womersley and Heaslip). 

Gahrliepia (Gahrliepia) cetrata (Gaterl932). 

Coxae III 4-setose. Scutum smaller. Dorsal setae 2.8 [2] .9. [2-5] .8.6. 
4.4.2. 

AW 48-9+4-7, PW 63-5+6-6, SB 41-4+1-8, ASB 22-4, PSB 
65-4+5-1, SD 87-8 + 5-1, A-P 38-4, AL 35-65+4-4, PL 35-2, 

Sens. 38-4 with head 12-8/25-6. 

Gahrliepia (Gahrliepia) saduski sp, n. 



284 Records of the S.A. Museum 

20. PL normally situated, posterior of SB and far distant from AL . . . , 21 

PL placed anterior of SB and closely adjacent to AL, longer than 
other scutal setae, Coxae III 1-setose. Additional scutal setae vari- 
able in number, normally 8 (occasionally 7-10). Dorsal setae nor- 
mally 2.4 [4]. 4 [2], 8 [2] .6.4.4.2.2. 

AW 38-4, PW 56*0, SB 43*2, ASB 22*4, PSB 107*2, SD 129-6, 
A-P 9*6, AL 41-6, PL 64-0, Sens. 33 -0 with head 9*0/23-0. 

Gahrliepia (Gateria) hirsuta (Radford 1946), 

2L Sensillae lanceolate. Coxae III 1-setose. DS 2.8 [4] .8 [2]. 8.8.6.4. 
Scutum and coxae apparently not punctate. 

AW 51-0, PW 85-0, SB 51*0, ASB 27-0, PSB 119-0, SD 146-0, 
A-P 35-0, AL 44-0, PL 68-0, Sens. 48-0 (after Radford). 

Gahrliepia (Gateria) lancearia (Radford 1946). 

Sensillae elavate 22 

22. Only 6 additional setae on scutum . . . , 23 

With 8 or more additional setae on scutum . . . . . . . . 24 

23. Dorsal scutum widest, only just behind line of PL. DS 46 in number, 
arranged 2.8 [4] .8 [2], 8.8.6.4.2. 

AW 51-0, PW 85-0, SB 51*0, ASB 27-0, PSB 119-0, SD 146-0, 
A-P 36-0, AL 44-0, PL 57-0, Sens. ? 

Gahrliepia (Gateria) crocidura (Radford 1946). 

Dorsal scutum widest well behind line of PL. DS 40 in number, 
arranged 2.8 [4] .6 [2]. 6.6.4.2.2. 

AW 50-0, PW 86-0, SB 50-0, ASB 28-0, PSB 157-0, SD 185*0, 
A-P 50-0, AL 39-0, PL 42-0, Sens. 33-6 with head 14/22. 

Gahrliepia (Gateria) nwten'sp, iu 

24. Scutum roughly pentagonal, i.e. margin behind PL forming a strong 
angle. Scutum and coxae strong! v punctate. Coxae III 1-setose, DS 
2.12 [6] .9 [3]. 8.6.4.4.2. 

AW 51-0, PW 85-0, SB 51-0, ASB 26-0, PSB 116-0, SD 142-0, 
A-P 34-0, AL 40-0, PL 57-0, Sens. — (after Radford}. 

Gahrliepia (Gateria) lontfipilis (Radford 1946). 

Scutum not pentagonal . . . . . . . . . . . « . . 25 

25. Scutum smaller, ca. half as long again as wide, widest about middle 
and posterior of PL. DS 2.6 [4]. 4 [2] .2 [2] .6.4.4.4.2. SD smaller than 
120/*. 

AW 43-0, PW 73-0, SB 41-5, ASB 17-0, PSB 99-5, SD 116-5, 
A-P 35-0, AL 18-0, PL 18-0, Sens. ? (after Gatcr; and Womers- 
ley and Heaslip. Gahrliepia (Gateria) rutila Gater 1932. 

Scutum larger, ca. twice as long as wide. SD. greater than 180/x . . 26 



Womeksley— Asiatic-Pacific Scrub Typhus Mites 285 

26. Scutum widest in posterior third, with 10 additonal setae, all but 
2 placed submarginally. 

AW 40-0, PW 70-0, SB 42-0, ASB 30-0, P8B 159-0, SD 189-0, 
A-P 53-0, AL 45-0, PL 38*0, Sens. ? (after Gator; and Womers- 
ley and Heaslip). Gahrliepia (Gateria) eittata Gater 1932. 

Scutum widest elsewhere, with more than 10 additional setae, which 

are not mainly submarginal . . . . 4 . 27 

27. Dorsal scutum widest just behind PL, with margins posterior of 
widest part lightly concave. With normally 12 additional scutal setae 
(variable from 9 to 14), arranged ca. 2.4 [4] .8 [4]. 6 [2] .6.6.4.4.2. 

AW 51-9±:4-2, PW 77-7±7-3, SB 46-7+:4-2, ASB 25-2, PSB 
163-8=fc6-7, SD 188-8±7«6, A-P 41-3+3-7, AL 37-3=fc6-5, PL 
40-8:±6-7, Sens. — . Gahrliepia {Gateria) spinidma (Radford 1946) 

Dorsal scutum widest at the middle and well behind PL, with 16 

additional setae. DS 2.4[4].10[8].4[2J.6[2].6.8.4.2. 
AW 52 '0, PW 79-0, SB 50-0, ASB 18-0, PSB 170-0, SD 188-0, 
A-P 39-0, AL 50-0, PL 37-0, Sens. I (after Gater; and Womers- 
ley and Heaslip). Gahrliepia {Gateria) fletcheri Gater 1932. 

Gahrliepia (Walohia) morobensis (Gunther 1939). 

Walchia morobensis Gunther 1939. Proc. Linn. Soc. New South Wales, 64 
(1-2), 94; Womersley and Heaslip 1943, Tr. Boy. Soc. S. Aust, 67, (1), 
135. 

Plate 76, fig. A^C. 

This species was originally described by Gunther from 4 specimens from 
Battus broumi and many from R. ring ens from Bulolo, New Guinea. 

As diagnosed the species was chiefly characterized by the scutal margins 
not being defined. A careful re-examination of Gunther's type in the collec- 
tion of the Australian Museum, Sydney, shows that at least the anterior and 
lateral margins can be seen, and possibly also the posterior margin. 

In the specimens recorded herewith and referred to this species the scutal 
margins can be plainly seen, especially in fresh or remounted specimens using 
the polyvinyl alcohol media. 

The species, on the basis of the type and the new material, can be charac- 
terized by the AL, SB and PL being almost in the same longitudinal line, 
although often the distance between the PL setae is somewhat greater than 
between SB and between AL. The posterior margin forms rather an acute 
angle, as in the figure. The palpal claw is bifurcate and all the setae on the 



286 Records of the S.A, Museum 

palpal femur, genu and tibia except the ventral on the tibia are nude. The 
sensillae are clavate, with the head strongly setulose and the apex acuminate. 
All the coxae are unisetose. Eyes absent, or only indistinctly 1 + 1. The 
dorsal setae are 2.6.6.6.4.2. There is no long nude seta on tarsi III. 

In 1943 Womersley and Heaslip (loc, cit.) gave the Standard Data for 
the type and one paratype as: AW 25-0, 25-0, PW 25-0, 25-0, SB 25-0, 27-0, 
ASB 16-5, 17-0, PSB 16-5, — f A-P 33-0, 34-0, AL 20*0, 18-0, PL 31-0, 
30-0, Sens. 25-0, 25 <0. The value of PSB 16*5 was only an estimate as at that 
time, the scutal margins could not be clearly seen. 

Through the kindness of Lt.-Col. J. R. Audy and his assistant T. J. Law- 
rence, I have been able to study a large amount of Trombiculid material from 
S. Burma, and have isolated 16 specimens which can be referred to morobensis 
Gunther. These were mainly from Nesokia tengalensis and Rattiis rattus from 
Toungoo, and neighbourhood, Oct. 1944. 

The Standard Data derived from 14 of these 16 specimens are ; 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


29«5±0-60 


2-24:£Q>42 


22-8-36-2 


25-6-32-0 


7-6 


PW 


32-9±0-90 


3-36+0-63 


22-8-43*0 


25 -6-38 '4 


10*2 


SB 


28-l±0-40 


1-50+0-28 


23-6-32-6 


24-0-28-8 


5-3 


ASB 


19-2±0-53 


l-98±0-37 


13 -3-25-1 


17-6-20-8 


10-3 


PSB 


44-0:1:0*36 


l-37±0-25 


39-9-48-1 


41-&-44-8 


3-1 


8D 


63-2:±0-40 


l-50±0*28 


58-7-67-7 


60-&-65-6 


24 


A-P 


35-4^0-28 


1« 06+0 '20 


32-2-38-6 


33-6-38-4 


3-0 


AL 


27-4±0-65 


2-42+0-46 


20-2-34-6 


22-4-28-8 


8-8 


PL 


29-5±0-48 


1-79+0 -34 


24-2-34-8 


25-6-32-0 


6-0 


Sens. 


25-6 with head 9 


►6/19*2. No variation recorded. 







Gahbltepia (Walchia) eustica Gater 1932. 

Gdhrliepia rustiea Gater 1932, Parasitology, 24, 167. 

Walchia rustiea, Womersley and Heaslip 1943. Tr. Eoy. Soc. S. Anst., 67, (1), 
136 j Sig Thor and Willmann 1947, Das Tierreich, 71b, 337. 

Plate 76, fig. D-F. 

This species was originally described from Rattus surifer surifer (Miller) 
from Selangor, Federated Malay States. 

It differs from the closely allied species, turmalis Gater in having the 
dorso-apieal tooth of the tricuspid cap on the chelieerae pointed and directed 
backward, hook-like, not blunt; in having fewer dorsal setae, 38-40 instead of 
40-44, and in the somewhat wider but shorter scutum. The palpal claw is 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



287 



trifurcate, and all the setae on the femur, genu and tibia of palpi nude. 
Gater states that the seta on the third segment (genu) sometimes has one or 
two barbs. The dorsal setae are 40 and arranged 2.6,8.8.6,4.4.2. The sensillae 
are elavate and setulose. Eyes apparently absent. 

The paratype specimen in the collection of the South Australian Museum 
has been remounted and the Standard Data re-determined as follows: AW 
41-6, PW 51-2, SB 32-0, ASB 17-6, PSB 48-0, SD 65-6, A-P 32-0, AL 28-0, 
PL 30-8, Sens. — . 

No specimens which can be referred to this species have been seen amongst 
the large amount of Burma material that I have been able to study, nor does 
the species appear to have been recorded from anywhere since the original 
discovery. 

In addition to the above paratype three other paratypes from Gater's 
collection at Kuala Lumpur, have been sent to me for study by Dr. J. E. Audy. 
In the Standard Data these specimens do not differ significantly from the 
above, the values without variation being: AW 42-0, PW 50-8, SB 31-0, ASB 
18-0, PSB 48-0, SD 66-0, A-P 30-0, AL 30-0, PL 30-0, Sens. — . 

This species has recently been found commonly on Rattus r. jaralc on 
Palau Jarak Is., in the Malacca Straits, by Dr. J. R. Audy and his team, and 
I have been privileged to study a large series collected 26th Jan. 1950. Of 
about 40 or more specimens, 16 have been measured and the following Standard 
Data determined; 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Kange 


Coeff . of 
Variation 


AW 
PW 
SB 


36-6±0-26 
44-5±:0-22 
29-7±0-20 


1^04^0-18 
0-87+0-15 
'79:t0 -14 


33-5-39-7 
41 -9-47-1 
27-3-32-1 


36*0-39-0 
42-0-45-0 
27-0-30-0 


2-8 
2-0 
2-6 


ASB 
PSB 
SD 


19-7±0-37 
44.lit0.35 
63«75:t0'30 


1.49=fc0-26 
1-39^:0.24 
l-21=fc0-21 


15-2-24-4 
39-9-48-3 
60*15-67-35 


18-0-21-0 
42-0-45-0 
63.0-66-0 


7-5 
3'1 
1-9 


A-P 


35-7±:0*33 


l-32±0*23 


31-7-39-7 


33-0-39-0 


3-7 


AL 
PL 


30-0 
30-0 


No variation recorded 
No variation recorded 






Sens. 


Lost in remounting. 









These specimens, although having slightly lower Standard Data than 
shown by Gater 'a Selangor material agree in all morphological characters. 
From the closely allied G> (W.) turmalis this species is distinct in the more 
pronounced tricuspid apical cap of the chelicerae, the fewer dorsal setae, and 
the smaller scutum with PSB equal to or less than PW. 



288 Records of the S.A. Museum 

Gahrliepia (Walchia) turmalis Gater 1932. 

Oahrliepia turmalis Gater 1932, Parasitology, 24, 168. 

Walchia turmalis, Womersley and Heaslip 1943, Tr, Roy. Soc. S. Aust», 67, (1), 
136 j Radford 1946, Parasitology, 37, 48; Sig Thor and Willmann 1947, 
Das Tierreich, 71b, 337. 

Plate 76, fig. G-I. 

This species, which is closely allied to the preceding one but differs as 
pointed out under that species, was originally described from Rattus sabanus 
vociferans (Miller) from Selangor, Federated Malay States. 

Radford 1946, recorded it from the Musk Shrew {Simons caertdeus yigan- 
teus (Geoffy.) from Colombo, Ceylon, 

It is a very common and widely distributed species in Southern Burma 
on such hosts as R&ttus rattus, R, norvegi&us concoior, and Nesokia bmgalem- 
sis, as evidenced by the very large number of specimens found amongst the 
material I have been able to examine. The localities include Ron, Insein, 
Paungola, and Toungoo. 

The palpal claw is apparently trifurcate and the setae on palpal femur, 
genu and tibia all plain or nude. The sensillae (missing in all of Gater 's 
material) are as in rustica, clavate and setulose. Gater ^s statement, "The 
setae on the hypostome are plain or serrate instead of plumose as in most other 
species" is puzzling. If he refers to what are now known as the "galeal setae" 
then these are plain or nude in most known species of Gaforliepi.a, but if he 
means the maxillary setae, I know of no species of Trombiculidae in which 
these are other than branched. The dorsal setae are from 40-44, and arranged 
2.6.6.6.6.6.6.4(2).2(0). 

The Standard Data as derived from 151 specimens from Southern Burma 
are as follows : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff. of 
Variation 


AW 


38-35±0-12 


l-38±0-08 


34* 2-42- -5 


35-2-41-6 


3-6 


PW 


48«6±0-17 


2 -102:0 -12 


42-3-54*9 


44-8-56-0 


4-3 


SB 


32 -42:0 -12 


1-482:0 -08 


28-0-36-8 


28-8-35-2 


4-6 


A8B 


22 -52:0 -04 


0-50±0-03 


21-0-24-0 


22-4-25-6 


2-2 


PSB 


56-7±0-20 


2-43:£0-14 


49-4-64-0 


48-0-64-0 


4-3 


SI) 


79-4:±0-21 


2-60:£0-15 


71-6-87-2 


70-4^86*4 


3-3 


A-P 


40-65±0-12 


1*492:0-08 


36-2-43-1 


38.4-44-8 


3-6 


Ali 


82-9±0<13 


1-622:0-09 


28'0-37-8 


28-8-38-4 


5-0 


PL 

8ens. 


32*6+0-12 
32-0 with head fl 


1-47^:0*08 
1-6/22-4. Ko va 


28-2-37-0 
riation recorded. 


28-8-35-2 


4-5 



Womersley — Asiatic-Pacific Scrub Typhus Mites 289 

Gahbliepia (Walchia) lewtjtwaitei (Gater 1932). 

Walchia lewihwadtei Gater 1932, Parasitology, 24, 170 ; Womersley and Heaslip 
1943, Tr, Roy. Soc, S. Aust, 67, (1), 135; Sig Thor and Willmanu 1947, 
Das Tierreich, 71b, 336. 

Plate 76, fig. J-L. 

This species was described from 9 specimens from Ratius r&ltus (probably 
dmrdi Jeutink) from Kuala Lumpur, Federated Malay States. 

It does not appear to have, been recorded from elsewhere, nor have I 
detected the species amongst the large amount of Gahrliepia material from 
Southern Burma. 

The species may be separated as in the key, by the very small scutum, the 
globose and not clavate sensillae and the normally 6-setose coxae III. The 
palpal claw is trifurcate and the setae on the palpal femur, genu and tibia all 
nude. Eyes apparently absent. Dorsal setae short, 34 to 40 in number and 
arranged ca. 2.6.6.6.6.4.2.2. 

The Standard Data of a specimen from the Institute of Medical Research, 
Kuala Lumpur, and collected from B. rattus from Sungei Buloh, F.M.S., 15th 
Oct., 1940, as given by Womersley and Heaslip (1943) are: AW 17-0, PW 
31-5, SB 17-0, ASB 17-0, PSB 21*5, SD 38-5, A-P 280, AL 11-5, PL 15-0, 
Sens. — , 

The Standard Data derived from 20 specimens of a large number from 
Rattus wMteheadi and R, mutteri from Selangor and Temerloh, Pahang, 
F.M.S., collected by J. R. Audy and his colleagues 1948, are as follows: 



AW 


Mean 
19*5±(M5 


Standard 
Deviation 

0-70±0-11 


Theoretical 
Itange 

17 '4.-21 -6 


Observed 
Range 

16-8-21.0 


Coeff. of 
Variation 

3-6 


PW 


31-65±0-21 


0-93±0-35 


28-9-34-4 


30-8-33-6 


2-9 


SB 


17-25^0*22 


0-99z±0-16 


14-25-20-25 


16-8-19-6 


2-7 


ASB 


19-3±0-19 


0-843:0.13 


16-8-21-8 


16-8-19-6 


4-3 


PSB 


28-0 


No variation recorded 






SD 

A-P 

AL 


47-3±0-19 
30-4±0-22 
14-0 


0-84±0-13 44-8-49-8 
1-00:4:0-16 27*4-33-4 
No variation recorded 


44-8-47-6 
28-0-30-6 


1-8 

3-2 


PL 


14-0 


No variation recorded 







Sens. 19-6 with head 1J/14. No variation recorded. 

The setation of coxae III in this population shows a slight yariation from 
the normal 6 -f- 6 setae; of the twenty specimens measured three had 5 + i> 
setae on coxae III and one had 6 -j- 5, the rest being normally 6 -j- 6> 



290 Records of the S.A. Museum 

Gahrliepia (Walohta) disparunguis (Ouds. 1929). 

SchongastieUa disparunguis Oudemaus 1929, Ent. Ber., 7, (165), 398; Sig 
Thor and Willmann 1947, Das Tierreieh, 71b, 335. 

Walchia disparunguis , "Womersley 1944, Tr. Eoy. Soc. S. Aust., 68, (1), 101. 

Plate 76, fig-. M-P. 

This species was originally described from specimens from the ears of 
Mus rattus var. from Garoet (W. Java), Aug., W, C. van Heuren. 

Oudenians' original description translated, reads as follows: "Length of 
a moderately engorged specimen 225/*, greatest breadth 145/*,. Scutum 
rounded-pentagonal with one angle directed posteriorly; in each of the other 
four angles a seta. On each shoulder is a seta and behind the scutum five 
rows of six setae in each. Pseudostigmal organ clavate, the stem about one- 
third of its length. Dorsal setae about 30/u. long, brush-like and shortly ciliated. 
Byes small, cornea half spherical Venter: all coxae (also maxillae) with a 
feathered seta; coxae III with two such. Between coxae I and between 
coxae III a pair of similar setae. Then 17 pairs of setae similar to the dorsal 
setae. Gnathosoma dorsally with 6 pairs of smooth setae; ventrally with 
more; externally on the tibiae with short smooth seta, and on the very short 
and difficult to see palpus are four setae of which one is short thick rod-like 
olfactory seta, the other three are short thick setae distally divided into four 
or five branches. Palpal claw bifid." 

Oudemans was rather uncertain about placing this species in Schongas- 
tieUa Hirst as it differed from Hirst's diagnosis in having only 4 setae besides 
the sensillae on the scutum instead of three pairs. He also noticed that the 
scutum resembled that of Tijphlothrombium Ouds. 1910 (= Gahrliepia Ouds, 
1912) although in his description it is suggested that the posterior is sharply 
defined, not tongue-like as in Gahrliepia (Womersley and Heaslip 1943) and 
similar to Wcdchia, He also was of the opinion that the shape of the scutum 
was a specific and not a generic character. Chidernans also refers to the dis- 
parity in the form and size of the 3 tarsal claws and named his species on this 
character. This feature, in which the median claw (empodium) is much 
stronger than the others and of median length, the outer longer and only 
slightly more slender, and the inner only slightly shorter than the median but 
much thinner is, however, present in all the species of Gahrliepia known to 
me, and I believe a good generic character. This species is closely related to 
glabra Walch, but can be easily distinguished as in the key, on the bi-setose 
coxae III and the shape of, and number of setae on, the scutum. 



WOMERSLEY — ASIATIC- PACIFIC SCRUB TYPHUS MlTES 291 

This species has not been recorded apparently ainee its description but that 
it is -widespread in the Asiatic-Pacific area is evident from the following speci- 
mens which I have been able to examine. 

Ceylon, 4 specimens from Millardia mellad® from Einbilipitiya, Oct., 1944, 
and 3 specimens from Bathos raitus kandiyanus from Nalanda, Nov., 1944 (S. 
U. Jayewickreme) ; Dutch New Guinea, Sansapor, 37 specimens from Aug, to 
Dec, 1944 (C. Mohr) ; Borneo, Labuan, 4 specimens from rats, Oct., 1945 (R. 
N. McC.) ; Celebes, Morotai, 2 specimens from rats, Oct., 1945 (R N. McC.) ; 
Southern Burma, On Nesokia bengalensis, Ron. 10 Oct., 1945 (747) ; on Raitus 
rattus, Toungoo, 23 Oct., 1945 (771) ; on N. bengalensis, Toungoo, 29 Oct., 
1945 (765), and on N. bengalensis, Toungoo, 26 Oct., 1945. 

The Standard Data for this species as derived from the Dutch New Guinea 
specimens are : 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Range 


Variation 


AW 


27-45+0-26 


l-60±0-18 


22 -65-32 '25 


24-0-28-8 


5-8 


PW 


44-6±:0-27 


l-62:±0-19 


39 -75-49 '45 


41-6-48-0 


3-6 


SB 


23-3±0-21 


l-26±0-15 


19-5-27<l 


22-4-25-6 


5-4 


ASB 


20 -3±lQ -21 


l-29±0-15 


16*5-24-1 


19-2-22-4 


6-3 


PSB 


36-lrt0-25 


1»53±0»18 


31-55-40-65 


32-0-38-4 


4-2 


SB 


56 -4:£0 -25 


l-53±0-18 


51-8-61-0 


54 -±-50 -2 


2-7 


A-P 


32-$±Q-23 


l-42±0-16 


28 -5-37-1 


30*4-35-2 


4-3 


AL 


22-4 


No variation recorded 






PL 


28-8 


No variation recorded 






Sens. 


25*6 with head 9- 


6/19-2. No 


variation recorded. 







A re-description of the larvae is as follows: 

Shape an elongate oval, slightly constricted behind coxae III. Size small, 
length (engorged) to 380/*, width to 250**. Scutum small, pentagonal, produced 
to a rounded point posterior of PL ; furnished only with AL and PL setae and 
paired sensillae ; sensillae fairly broadly clavate with prominent setules • anter- 
ior margin lightly concave. Eyes 2 + 2, the posterior smaller and indistinct. 
Chelicerae non-serrate, with the usual tricuspid cap. Galeal setae nude. Palpi 
fairly stout, tibial claw bifurcate; all setae on palpal femur, genu and tibia 
nude ; tarsi with the usual basal and apical sensory rods and 3-4 ciliated setae. 
Dorsal setae 36 in number, to 25/* with only short ciliations and arranged ca. 
2.6.6.2 (lateral) .6.6.4.2.2. Ventrally with a pair of ciliated setae on gnatho- 
soma, a pair between coxae I and between coxae III, one on each of coxae I 
and II, two on coxae III, and thereafter 14.12.12.6.4.2, to 20/a long. Legs 
rather short, 1 180/*, II 150/*, III 210/*; tarsi I and II with the usual sensory 
rods ; III without any long nude seta. 



292 Records of the S.A. Museum 

CtAhrliepia (Walchia) bwingi (Fuller, 1951), (=glabrum Walcli, 1927, 

preoc.) 

TromMcida glabrum Walch 1927, Geneesk. Tijdsch. y. Ned. Indie, 67, (6), 926. 

Walchia glmbrum, Ewing 1931, Proc. U.S. Nat. Mus., 80, (8), 10; Woinersley 
and Heaslip 1943, Tr. Itoy. Soc. S. Aust., 67 (1), 134 ; Womersley 1944, 
Tr. Roy. Soc. S. Aust., 68, (1), 134. 

Walchia glabra, Sig Thor and Willmann 1947, Das Tierreich, 71b, 336 ; Fuller 
1948, Bull. Brooklyn, Ent. Soc., 43, (4), 110. 

Walchia pingue Gater 1932, Parasitology, 24; Womersley and Heaslip 1943, 
Tr. Roy. Soc. S. Aust., 67, (1), 134 j Womersley 1944, Tr. Roy. Soc. S. 
Aust, 68, (1), 102. 

Walchia pingms, Sig Thor and Willmann 1947, Das Tierreich, 71b, 338. 

Plate 76, fig. Q-T. 

This species was originally described from rats from the Lampong District 
of Macassar, and in 1943 Womersley and Heaslip identified 3 unidentified 
specimens from Eattus rattus argentiventer Chasen, from Perak sent to them 
from the Institute of Medical Research, Kuala Lumpur, and one specimen 
(I, A, I.C., IV) from Batavia, as the same. 

In 1944 the species was recorded by myself from 7 specimens collected 
in the Buna area of New Guinea by G. M. Kohls and the Standard Data 
given. The Standard Data of 3 specimens from Perak, 10 from Buna, N.G. 
and 24 received more recently from Dr. R. Gispen, from Batavia, from Ratttis 
rattus cliardi are as follows: 







Standard 


Theoretical 


Observed 


Coeff . of 




Mean 


Deviation 


Bange 


Range 


Variation 


AW 


28'4:t0-18 , 


1-08±0-12 


25-2-31- 6 


25-6-30-4 


3-8 


PW 


48-8±0-48 


2-94+0-34 


40-0-57-6 


43-2-54-4 


6-0 


SB 


25-0±0-20 


l-20±:0-14 


21-4-28-6 


22»4r-25-6 


4«8 


ASB 


22-4 


No variation recorded 






PSB 


34-3±0-23 


1-42+0-16 


30-0-38-0 


32-0-35-2 


4-1 


8D 


56 • 7±0 « 23 


l-42:±0-16 


52 •4-61-0 


54-4-57-6 


2-5 


A-P 


36-7±0-45 


2-72±0-32 


28-6-44-8 


32-0-38-4 


7-4 


AL 


22-4 


No variation recorded 






PL 


28-8 


No variation recorded 






Sens. 


28 - 8 with hea 


d 11 -2/22 -4. Nq i 


rariation recorded 







This species is closely related to the following, enode Water, but differs in 
the larger scutum and particularly in the number of setae on coxae III. In 
glabra, coxae III carries normally 3 setae, and shows extremely little variation 



Womersley— Asiatic-Pacific Scrub Typhus Mites 293 

in this respect; of the 37 specimens examined only two show more than 
three setae, on one of each coxae III, these having 3 + 4 setae. In enode the 
setae on coxae III while normally 4+4, show a very great variation from 
4 + 3 to 5 + 6. "Where, however, specimens of the latter species have only 
4 + 3, the size of the scutum, the Standard Data and the dorsal setae will fairly 
easily determine the species. 

In glabra the palpal claw is bifurcate and the setae on the palpal femur, 
genu and tibia, and on the galea are nude. The dorsal setae are 2.6.6.6.6.4.2. 

From Gater's description of pmgue there seems little doubt that it i» 
identical with glabra. It has not been recognized since Gater 'a paper nor have 
any specimens that could be referred to it been seen amongst the large amount 
of material studied in the preparation of this paper. Its synonymy with glabra 
is confirmed by a study of one of Gater 's two specimens, kindly sent to me by 
Dr. J. R. Audy. 

Gahrltepta (Walchia) enode (Gater 1932). 

Walchia cnode Gater 1932, Parasitology, 24, 169, {enodis) j Womersley and 
Heaslip 1943, Tr. Roy. Soc. S. Aust, 67, (1), 135; (enodis) Radford 1946, 
Proe. Zool. Soc, London, 116, (2), 247 ; Sig Thor and Willmann 1947, 
Das Tierreich, 71b, 338. 

Plate 76, fig. U-X. 

This species which was originally described from Sungei Bui oh, Selangor, 
Malaya, from Battus mwlleri validus (Miller) does not appear to have bedn 
recorded since. 

It is, how T ever, a very common and widely distributed species in Southern 
Burma as shown by the large number of specimens amongst the Trombiculid 
material collected by Lt.-Col. J* R. Audy and his staff of the S.E.A.C. Scrub 
Typhus Research Team, and submitted to me for identification. 

As stated under G~ glabra Walch, the present species is closely related 
thereto, differing in the smaller scutum, the dorsal setae and particularly a 
different and more variable number of setae, on coxae III. 

In the 191 specimens for which the Standard Data is given, the setation 
of eoxae III varied as follows: 3 + 4, 21 specimens (11-0%) ; 3 + 5, 2 specs. 
(1-6%); 4 + 4, 119 specs. (62-3%) \ 4 + 5, 35 specs. (18-3%) ; 4 + 6, 1 spec. 
(0-7%); 5 + 5, 12 specs. (6-3%); 5 + 6, 1 spec. (0-8%). Thus the normal 
is 4 + 4. Where the setae on coxae III are 4 + 3, as in glabra, the sise of the 
scutum and its Standard Data will be determinative. 



294 Records of the S.A. Museum 

The sensillae are capitate and setulose and their bases somewhat nearer to 
PL than to AL. The palpal claw is apparently, and the setae on the palpal 
femur, genu and tibia as well as the galeal setae, nude. The chelicerae has 
only the usual apical tricuspid cap. Eyes apparently absent. The dorsal setae 
vary from 32-34 and are arranged ca. 2.8.6.6.4.4.2. 

The following Standard Data are from 189 specimens, plus a specimen from 
Sungei Buloh, Selangor, 8 Aug. 1930 from Rattiis mvileri vatidus (Miller) and 
one specimen received from Radford from Batons sp. Imphal, Manipur, 10 
June, 1945. 



AW 


Mean 
26-5+0-10 


Standard 
Deviation 

1-45+0-07 


Theoretical 
Bange 

22-1-30-9 


Observed 
Bange 

22-4r-32-0 


Coeff . of 
Variation 

5-5 


PW 


37-6+0-15 


2-13+0-11 


3J -2-44-0 


35-2-51*2 


5-7 


SB 

A8B 


21-0±0-10 

18-9±0-06 


1*44+0-07 
0*78+0-04 


16-6-25*4 
16-6-21-2 


17-6-25-4 
17*6-22*4 


6-8 
4-1 


PSB 


32-75±0-14 


1-97+0^10 


26-8-38-7 


28-8-35-2 


6-0 


SD 


52*0±0-15 


2-03+0-10 


46-0-58*0 


48-0-57-6 


3-9 


A-P 


30-95+0-10 


l-36±0-07 


26-85-35-05 


28-8-35*2 


4-4 


AL 


19-2+0-06 


0-78+0-04 


16-9-21-5 


16-0-25-6 


4-0 


PL 

Sens. 


20*7+0-10 1*39+0-07 
22-4 with head 12-8/16-0. No v 


16-6-24*8 
ariaticn recorded. 


19-2-22-4 


6-7 



In Southern Burma it occurs commonly on Nesokki bengalensis and the 
various species of Ratous. 

Gaheliepia (Walchia) bbennani sp. n. 
Plate 112, fig. A-D. 

Description of Larvae. Size small. Shape oval. Length (engorged) to 
335/a, width to 254/a. Scutum as figured and approximately of the size and 
shape of (?. (W.) turmalis Gater; with only AL and PL setae; sensillae cla- 
vate and setulose. Eyes apparently absent. Chelicerae with only the apical 
tricuspid cap. Galeal setae nude. Palpal claw bifurcate; setae on femur, genu 
and tibia apparently all nude. Dorsal setae 32-34 in number, to 30/x. long, 
shortly and sparsely ciliated, and arranged 6 (4). 6. 6.6,6.2.2, i.e., normally with 
3 + 3 humeral setae, but frequently only 2 + 2. Ventral ly with a pair of 
setae on maxillae, a pair between coxae I and between coxae III, one on each 
of coxae I and coxae II, two on coxae Til ■ posterior of coxae III ca. 36 setae 
to 26^ long. Legs: I 200/1. long, II 174^ III 200^; tarsi I and II with dorsal 
sensory rod, III without a long nude seta. 

Standard Data derived from the type and 16 paratypes are : 



Womersley— Asiatic-Pacific Scrub Typhus Mites 295 

Standard Theoretical Observed Ooeff. of 

Mean Deviation Range Range Variation 

AW 37-5±0*30 4-0±0-21 33-6-41-5 36-85-40-2 3*6 

PW 64»l±0-36 4-8=t0-25 49-3-58-9 53-25-56-95 2-9 

SB 32-8±0-42 5-6zt0-29 27-2-38'4 30-15-36-85 5-7 

ASB 21-8 No variation recorded. 

P8B 67-8±0»32 4»3±0-23 53-5-62-1 56-95-60-3 2-5 

SD 79-6±:0-32 4-3±0-23 75-3-83-9 78*75-82-1 1-9 

A-P 37-6±0-30 4<0±0-21 33*6-41-6 36-85-40 -2 3-5 

A L 27-3±0-27 3-6±9-10 23-7-30-9 36-8-30-15 4-4 

PL 26-8 No variation recorded, 

Sens. 33-5 with head 10-05/23-45. Only 1 determination. 

Loc> and Hosts. The type and sixteen paratypes from a "ground squirrel" 
Bhwwsciurus laticaudatus from Kepong Forest Reserve, Selangor, F.M.S., 11 
Sept,, 1950 (Scrub Typhus Research Unit). Also 1 specimen from Battus 
boivcrsi from Ulu Langat, Selangor, F.M.S. 13 June, 1950. 

Remarks. In the shape of the scutum and the Standard Data, this species 
is very close to G. (W.) turmalis Gater. It is strikingly different, however, in 
having coxae III normally bisetose, and in having 2-3 humeral setae on each 
side. It is named in honour of Dr. J, M. Brennan, of the Rocky Mountain 
Laboratory, Hamilton, Mont,, U.S.A., who has described many American species 
of Trombiculidae. 

var. ventralis nov. 

Three specimens with the above material differ rather remarkably in that 
coxae III are nnisetose, and ventrally between coxae II and coxae III are from 
3 to 4 setae somewhat shorter than the dorsal humeral setae. Coxae II and III 
are correspondingly wider apart than in the typical form. 

In Standard Data these 3 specimens agree with the typical specimens. For 
the moment, as in all these 3 specimens the sensillae are missing, they are re- 
garded as a distinct variety. 

The 3 specimens were from B. bower si but from the Ulu Langat Forest 
Reserve, Selangor, F.M.&, 13 June, 1950. 

Gaiirlibpia ( ? Walchia) rjoi Gunther 1940. 

Gahrliepia rioi Gunther 1940, Proe, Linn. Soc. New South Wales, 65, (5-6), 
481, 

Plate 77, fig. A. 

This species has not been recognized since its original description. Accord- 
ing to Gunther 's description and figures the dorsal scutum is distinctly produced 
beyond the PL setae; tongue-shaped, but with only the AL and PL setae; its 



296 Records of the S.A. Museum 

posterior end is slightly concave and SB are much nearer to PL than to AL. 
The sensillae were missing from Gunther's material. The chelicerae are simple, 
■with only the usual tricuspid cap., and the galeal setae are nude. The palpal 
claw is trifureate. The seta on the palpal femur is nude; on the genu 
with one long branch, and on the tibia one long branched seta (this is probably 
the ventral, the dorsal and lateral not having been seen). The dorsal setae 
number 28, and are arranged 2.6.6.6.4.2.2. Ail coxae l-setose ; except III which 
are 2-setosc; posteriorly of coxae III ca. 36 setae. 

The Standard Data as deduced by Womersley and Heaslip (1943) from 
Gunther's data are: AW 30-0, PW 50-0, SB 25-0, ASB 18-0, PSB 58-0, SD 
76-0, A-P 24-0, AL 28*0, PL 40-0, Sens. — . 

It was described from specimens from the mouse deer, Tragid%is bom- 
mnns from the Bode River, British North Borneo. Sept., 1939. 

Gahrliepja (Sciiongastiella) ligula (Radford 1946). 
Sohmgastidla ligida Radford 1946, Proc. Zool. Soc. London, 116, (2), 256. 

Plate 77, fig. B-F. 

Radford described this species from Battus rattus rufcscens (Gray) from 
Iniphal, Manipur State, India, 6th May, 1945. 

This species is very common in Southern Burma on various rats and on 
Nesokia bengalensis* I have been privileged to study some hundreds of speci- 
mens from various localities such as Ron, Prome, Toungoo, Insein, and Tbarra* 
waddy, collected by Lt.-Col. J. R. Audy and his team in 1945, 

As Radford's description is somew f hat brief a fresh and more detailed one 
is given below. 

Shape oval with a tendency to constriction just posterior of coxae III. 
Length (engorged) to 560/*, width to 320/x. Scutum elongate, ligulate behind 
PL and in addition to AL and PL carrying the two median setae of the second 
dorsal row; because of its narrow ligulate posterior part these two additional 
setae are near to the apex and close together. Eyes apparently absent. The 
sensillae are clavate (as stated by Radford and not globose as shown in his 
figure 17) and setulose. Palpi small tibial claw apparently trifureate ; all setae 
on palpal femur, genu and tibia nude. Chelicerae with only the apical tricus- 
pid cap. Galeal setae nude. Dorsal setae 48, to 35//. long and arranged 
2.812] ,8*4.8.6.6.4.2. Ventrally with the usual pair of ciliated setae on the 
maxillae, one on each coxa, a pair between coxae I and between coxae III and 
posterior of coxae III ca. 56 setae. Legs: I 210/a, II 190/x, III 240/a; tarsi I 
and II with the usual dorsal sensory rod, III without a long nude seta. 



WOMERSLEY — ASIATIOPACIFIC SCRUB TYPHUS MlTES 



297 



The Standard Data derived from 48 


specimens fi 


■om Sj Burma 


are -. 






Standard 




Theoretical 


Observed 


Ooeff. of 




Mean 


Dcvia.tion 




Range 


Bange 


Variation 


AW 


36-8±0-27 


1-88^:0.19 




31-2-42-4 


33-6-39-2 


5-1 


PW 


51-9±0-33 


2^29:±0-23 




45-0-58-8 


44-8-56.0 


4-4 


SB 


35-6±0-S2 


l-56±0-16 




30-9-40-3 


33-6-39-2 


4-4 


ASB 


26'5±0-20 


1 -39:1:0 -14 




22-4-30-6 


25-2-28 


5-2 


P8B 


65-5dt0-46 


3-21±0-33 




55- 9-75-1 


61-6-72-8 


49 


SD 


92-0±0-53 


3 -70:1:0 -38 




80-9-103-1 


86-8-100-8 


4-0 


A-P 


44-7:1:0-25 


l-72±0-17 




39-5-49-9 


42'0-47*6 


3-8 


Ali 


42*0±0-15 


l-07:£0-ll 




38-8-45-2 


39'2-44-8 


2-5 


PL 


42-0 


No variation recorded 






Sens, 


33-6 with head 14 


•0/25-2. No 


variation recorded. 







O. (£.) ligula (Radford) is also a common species in Kashmir and T have 
recently received much materia! from Maj. S. L. Kalra, collected from ears of 
rats, and from a (shrew during 1948 and 1949 from the following localities: 
Kanzalwan, Jhanghar, Gurais, Srinagar, Nau9hera and Banital. 



Gahrliepia (Schonoastihlla) ceylonica sp. n. 
Plate 77, fig. G-L. 

Description of Larvae. Shape oval, but with a marked constriction behind 
the third pair of legs. Colour in life unknown. Length to 545/j., width to 
350ft- Dorsal scutum elongate, widest in line with sensillae, slightly narrower 
between PL than between AL; posterior of PL narrowing to form a long nar- 
row lip-like projection which subapieally takes in the median pair of setae of 
the second dorsal row ; sensillae globose, finely setulose. the bases about midway 
between AL and PL. Eyes not observed. Chelicerae with only the apical 
"tricuspid cap. Palpi stout, all. setae on femur, genu and tibia apparently 
nude. Galeal setae nude. Palpal claw bifid. Dorsal setae to 28/a long, 36 in 
number and arranged 2.6 [21. 8.8.6.4.2. Ventral setae more numerous and 
shorter than on dorsum; one on coxae I and IT, four on coxae III, a pair 
between coxae I and between coxae III and thereafter ca. 8.8.8.8.8.6.4.2. Legs : 
I 156/x long, II 130/a, III 143ft; tarsi I and II with usual dorsal rod-like seta, 
III without any long nude seta. 

The Standard Data for the type larva are: AW 31-0, PW 28-0, SB 23-0, 
ASB 14*0, PSB 55-0, SD 69-0, A-P 28-0, AL 20-0, PL 20*0, Sens. 20-0 with 
head 14/14. 

Loc. a/nd Hosts. Type from Rattus r\ kandiyiWdos from Nalanda, Ceylon, 
May, 1944 (S.H.J.) and 4 paratypes from Leggad-a booduga fulvidiveritris from 
same locality May 1944 ( S.H.J. ). Two specimens also from a "mouse" from 
Nanshera, Kashmir, India, 14th May, 1949 (G, L, Kabra). 



298 Records of the S,A. Museum 

Remarks. In the peculiar iigulate form of the dorsal scutum this species 
is close to ScJiongastielki ligula Radford. It differs, however, in the scutal 
dimensions, the arrangement of the dorsal setae and especially in having 4 
setae on each coxae III, instead of the one seta in Radford's species. 

The larvae have been reared through to the nymphs which are described 
later in this paper. 

Gahbliepia (Schongastiella) brevis (Radford 1946). 
Schongastiella hrevis Radford 1946, Proc. Zool, Soc, London, 116, (2), 256. 

Plate 78, fig. A. 

This species was described from a specimen (or ? specimens) from a Tat 
Eattus r, rwfescem (Gray) from Imphal, Manipur State, India, 7 May, 1945. 
Although the scutum of this species, as figured by Radford has a roughly pen- 
tagonal shape and is prolonged posteriorly to include the 2 median setae of the 
second dorsal row, it is very wide, almost as much so as it is long. The sen- 
sillae are missing, but the bases are closer to AL than to PL. The dorsal setae 
number 28, and are arranged 2.6 [2]. 6.6.2 A2. Posterior of coxae III ventrally 
ca. 42 setae. Coxae I and II 1 -setose, III 2-setose. Eyes apparently absent. 
Mouth parts and palpi not described. 

The Standard Data (after Radford) are: AW 44-0, PW 86-0, SB 51-0, 
ASB 26-0, PSB 78*0, SD 104-0, A-P 43-0, AL 38-0, PL 41-0. Sens. — . 

This species has not been recorded, nor has any fresh material come to 
hand, since its original discovery. 

Gahrliepia (Schongastiella) bengalensis (Hirst 1915). 

Schongastietta hengalemis Hirst 1915, Bull. Ent. Res., 6, fig. 7, 8 ; Ewing 1938, 
J. Wash. Acad, Sci., 28, (6), 295; Sig Thor and Willmann 1947, Das Tier- 
reich, 71b, 334. 

Gahrliepia bengalensis, Womersley and Heaslip, 1943, Tr. Roy. Soc. S. Aust. r 
67, (1), 140. 

Plate 78, fig. B. 

In this species described from specimens in the ears of Mus. rattus, Cal- 
cutta, 20 Mar., 1915, the dorsal scutum is tongue-shaped, but rather quickly 
narrowing posterior of PL to the rounded apex, and taking in two of the- 
median setae of the second dorsal row. According to Hirst's figure there are 
two, comparatively large, eyes on each side. The chelicerae are apparently 
simple with only the apical tricuspid cap. The palpal tibial claw is shown in 
the figure as bifurcate. The setae on the palpal femur, genu and tibia are all 



Womersley— Asiatic-Pacific Scrub Typhus Mites 299 

nude. All coxae are 1-setose. The dorsal setae number 38, and are arranged 
2.6 [2] .8.8.6.4.2.2. Ventrally posterior of coxae III ca. 36 in number and 
shorter than the dorsal setae. 

To this species are now referred three specimens, two from a rat, from 
Thunia, Burma, 17 Dec., 1946 (J. R. Audy) and one from a rat from Ranik- 
het, Burma, 20 Oct., 1946 (J JR. A.). 

The Standard Data for one specimen from Thunia and the one from Ranik- 
het are: AW 38-4, 38-4, PW 51-2, 51-2, SB 35-2, 35-2, ASB 16-0, 16*0, 
PSB 70-4, 64f0, SD 86-4, 80-0, A-P 35*2, 38-4, AL 38-4, 38-4, PL 35-2, 
35-2. Sens.—. 

These values are somewhat lower in some items than those given by Wom- 
ersley and Heaslip 1943, which were derived by interpolation from Hirst's 
figure. 

It should be noticed that in Hirst's figure the sensillae bases are about 
half-way between AL and PL, whereas in the fresh material as figured they are 
definitely nearer to AL than to PL. 

More recently I have been able to examine a number of specimens of this 
species collected in Kashmir by M'aj. S. L. Kalra, as well as two specimens 
from Burma. Of these the Kashmir specimens were 5 from Kanzalwan, 10-11 
Oct., 1946, and 4 from Tithwal, Sept., 1946. The two Burma specimens were 
from Thunia, 17 Dec, 1946 (J. R. Audy) and Raniket, 20 eDc, 1946 (J.R.A.). 

The Standard Data for these eleven specimens are : 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 
PW 
SB 


36-7±0-47 
50^6+0-69 
31-7±0-80 


l-55±0-33 
2.17±0«48 

2-66:±0-57 


32*0-41-4 
44-1-57-1 
23 '7-39 -7 


34-O-39-0 
46-0-54-0 
28-0-35.0 


4-2 
4-2 
8-4 


ASB 

PSB 
SD 


21O+0-81 
58-l±l-69 
79-2±0-97 


2-70±0-57 
5-02±l-13 
3'08±0.6£. 


13-0-29-2 
43-1-73-1 
70-0-SS-4 


16-0-24-0 
54-0-70-0 
75-^-86. 


12-8 

8-0 
3-9 


A-P 


37-6±:0-52 


l-65±0-37 


32-7-42-5 


35-0-39-0 


4-4 


AL 


31-45±0-97 


3-23:±0'67 


21-75-41-15 


30-0-38-0 


10-2 


PL 


35-8±0-15 
33-0 with head J 


0-44±0-10 
.5/21. Only one 


34-5-36-1 
determination. 


35*0-36-0 


1-2 



Gaurlii^ia (Schongastiella) punotata (Radford 1946). 
Sclwngasticlla pwictaia Radford 1946, Proc. Zool. Soc. London, 116, (2). 

Plate 78, fig. C. 

This species was described from a shrew Suneus (Crocidura) caeruletis ful- 
vocinereu>s (Anderson) from Kanglatongbi, Imphal, Manipur State. 20 ApriJ, 
1945. 



300 Records of the S.A. Museum 

To it are now referred 12 specimens from Southern Burma, collected in 
1945 by T. J. Lawrence. 

The dorsal scutum is fairly large, roughly pentagonal with the posterior 
angle well produced and taking in 2 extra setae from the second dorsal row. 
The sensillae are situated nearer to AL than to PL, are clavate and setulose 
with their bases wide apart. The eyes are 2 + 2, anterior very distinct, and 
posterior the smaller. Dorsal setae 2.6 [2]. 6.6.4.2.2. Coxae III bisetose and 
venttally posterior of coxae III ca, 40 setae. Scutum and coxae strongly punc- 
tate. The galeal setae and the setae on palpal femur are lightly branched, on 
genu and tibia nude. Palpal claw apparently bifurcate. 

The Standard Data for the above 12 specimens coll. No. 62 (4 specs.), 
458 (1 spec.) and 122 (7 specs.) are as follows: 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Bangc 


Range 


Variation 


AW 


45»2±0-69 


2 -37^:0 -48 


38*1-52 3 


41-6-51-2 


5-2 


PW 


68-5±l*27 


4-41+0-90 


55-3-81*7 


60-8-73-6 


6-4 


SB 


43 -5+0 -73 


2-54rt0-52 


35-9-51-1 


38-4-44-8 


5-9 


A8B 


22<7±0-27 


0-92+0-19 


19 -9-25 «5 


22-4-25-6 


41 


PSB 


91-5±2-82 


9*78+2-00 


62- 2-120 -8 


80-0-102-4 


10-7 


SD 


114-1^2-77 


9-58+1-96 


85-3-142*9 


102-4-124-8 


8-4 


A-P 


44-9+0-64 


2-20+0-45 


38 -3-51 '5 


41-6-48-0 


4-9 


AL 


38-4 


No variation recorded 




« 


PL 


38-4 


No variation 


recorded 






8ens. 


35*2 with head 9*6/22-4. No variation recorded. 







In addition to the aboye specimens T have also one from Thunia, Burma, 
16 Dec, 1946 (J.RJV.) and one from Millardia millanlia from Embilipitiya, 
Ceylon, Jan., 1945 (S. IL Jayewickreme).. 

G-AHRLIEPIA (SCHONGASTIELLA) KTJMAONENS1S Sp. n. 

Plate 78, fig. D-GL 

This species is described from a single specimen from a shrew from Bhim- 
tal, in the Kumaon Hills, India, 10 Oct., 1946 (S. L. Kalra). 

The species is close to bengalensis Hirst and punctata Radford but differs 
as in the key in the Standard Data, shape of the scutum, dorsal setae, and the 
4-setoae eoxae III. In the last character it agrees with sadaiski sp. n, from 
Japan but differs in the other above-mentioned characters. 

Description of Larvae. Shape an elongate oval, length (engorged) 420/i, 
width 350ju,. Scutum fairly large, roughly pentagonal, widest in line of PL 
with the posterior angle produced and taking in two of the six setae in the 
second dorsal row. Sensillae missing, bases nearer AL than PL. Chelicerae 
normal, with only the usual apical tricuspid cap. Galeal setae nude. Palpi 



Womersley— Asiatic-Pacific Scrub Typhus Mites 301 

fairly stout, tibial claw bifid ; setae on palpal femur, genu and tibia all nude. 
Eyes apparently absent. Dorsal setae strongly ciliated, 32 in number and 
arranged 2,6 [2]. 6.6.6.4.2. to 36/* long. Ventrally with the usual pair of 
branched maxillary seta, one setae on each of coxae I and II, four setae on 
coxae III; behind coxae III with ea. 40 setae. Legs: I 216/a long, II 182/x, III 
238/t; tarsi I and II with the usual sensory rod, III without any long nude 
seta. 

The Standard Data of the type, in the South Australian Museum are: 
AW 51-2, PW 76-8, SB 49-6, ASB 28-0, PSB 110-0, SD 138-0, JA-P 51-2, 
AL — , PL 41-6, Sens. ~. 

Gahrliepia (Gahrliepia) saduski sp. n. 
Plate 78, fig. H-J. 

Description of Larvae, Shape broadly oval. Length (engorged) to 550/x, 
width to 450/i. Scutum as figured, comparatively small., tongue-shaped, widest 
about in line of PL, and besides the AL and PL setae embracing the median 
4(2-5) setae of the second dorsal row which itself is variable. Sensillae cla- 
yate with pronounced setules, and their bases wide apart and nearer to AL 
than to PL, Eyes apparently absent. Chelieerae simple, with only the usual 
apical tricuspid cap, Galeal setae nude. Palpi with trifurcate tibial claw with 
the prongs closely adpressed; setae on palpal femur with 4-5 strong branches, 
on genu with 2-3, on tibia the dorsal and lateral setae nude, and the ventral 
with 3 long branches; tarsi with the usual basal and apical sensory rods and 
4-5 ciliated setae. Dorsal setae strongly ciliated, ca. 32 in number and arranged 
ea. 2.8 [4] .9 [2-5]. 6.3.6.5.3, to 50^ long. Ventrally with a pair of branched 
setae on maxillae of palpi, one on each coxa I and II, and 4. setae on coxae III, 
a pair between coxae I and between coxae III, thereafter ca. 36 in number to 
40/i long. Legs: I 235/a long, II 210ft, III 230^; tarsi short, I and II with the 
usual dorsal sensory rod, III without any long nude seta. 

The Standard Data for 7 of 8 specimens are: 





Mean 


Standard 
Deviation 


Theoretical 
Range 


Observed 
Range 


Coeff . of 
Variation 


AW 


48-9±0-59 


1-56±0-42 


44-3-53 '6 


48-0-51-2 


3*2 


PW 


fi3»fi±0'83 


2-21;±0-59 


56 -9-70-1 


60-8-67-2 


3-5 


SB 


41-4-±0*23 


0*60:t0'i6 


39-6-43*2 


40-0-41-6 


1*5 


ASB 


22-4 


No variation recorded 






PSB 


65 -4 ±0-65 


l-71±0-46 


60*3-70-5 


64*0-67>2 


2*6 


8D 


87-8±0-65 


1-71:1:0 -46 


82-7-92-9 


86-4-89-6 


1-9 


A-P 

AL 

PL 

Sens. 


38-4 

35-65±0-46 

35-2 

38*4 with Ilea* 


No variation recorded 
1-21±0*32 31- 25-40*05 
No variation recorded 
d 12 *8/25 -6. No variation recorded. 


35-2-38*4 


3-4 



302 Records of the S.A. Museum 

Loc. and Hosts. The type and 2 paratypes from Mux. sp. from Misinobe, 
near Yachi, Yaniagata Prefecture, Japan, 28 Oct, 1945 (C. B. Philip) j four 
other specimens from Niigata, Japan, 18 Oct., 1945, embedded in the belly of 
Microtus montebelli, and a further specimen from the latter host and locality, 
January, 1946 (coll. T. 0. Berga). 

I am greatly indebted to CoL C. B. Philip for the opportunity of describ- 
ing this material. 

Remarks. This species can be separated from all others on the shape of 
the dorsal scutum, the additional setae thereon, and on the 4-setose coxae III. 
In the number of additional scutal setae it is somewhat variable, and of the 
eight specimens examined 3 had 4 in addition to AL and PL, 3 had 2, 1 had 
3 and 1 had 5* The number of setae on coxae III did not vary. 

Oahrlibpia (Gahruepia) insigne, sp. n f 
Plate 79, fig. A-D, 

Description, of Larvae. Shape oval. Length (partially engorged) 338/t, 
width 182/;. Scutum elongate, tongue-like, with large reticulations which are 
roughly square to hexagonal; sensillae missing but bases in front of line of 
PL ; AM shorter than PL but similarly thick and ciliated. Eyes 2 -f- 2, very 
small and close to scutal margin just anterior of PL, Palpi stout with bifur- 
cate tibial claws of which the prongs are about equal and long ; seta.e of femur 
very shortly ciliated, of genu the same, of tibia all 3 apparently nude. Cheli- 
cerae simple with only the usual apical cap. Galeal setae nude. Dorsal setae 
generally strong and thickly ciliated, 32 in number, from 42/* long anteriorly 
to 36/x posteriorly, arranged 2.6 [2] .6 [2]. 6.6.4.2, those on the scutum are thin- 
ner than the outer and the scutal setae. Ventrally, with paired fine and 
ciliated setae on maxillae, one on each coxa, a pair between eoxae I and 
between coxae ITI> thereafter ca. 56, very short and fine. Legs: 1 7-segmented, 
II and III 6-segmented; I 220/*, long, II 195/x, HI 260/x; coxae of leg III are 
broad and short, almost rectangular; tarsi I and II with the usual dorsal rod, 
HI without any long nude seta. 

The Standard Data for the unique type are: AW 44-6, PW 75-6, SB 
44-6, ASB 22-4, PSB 173-6, SD 196-0, A-P 30-8, AL 39-2, PL 53-2, Sens. — ; 
the maximum width of scutum is 98/x. 

Loc. and Host. The unique type from Twpaia glis, from Pahang Ed., 
Gombak Forest Reserve, near Kuala Lumpur, F.M.S., 18 Oct. 1948 (JJt.A.)- 

Remarks, A very striking species, but closely allied to the two following in 
baving an ornamented scutum. It differs, however, in the type of scutal orna- 



WOMERSLBY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 303 

mentation and in the size of scutum and the position of SB in front of line 
of PL. 

This, and the following two speciefs, in having four additional setae on the 
posteriorly produced scutum, agree with the type 0. nanus (Ouds.), and thus, 
if subgenera are accepted fall into Gdhrliepia s. str. Oudemans' species does 
not possess such strongly ornate scutum as in insigne, decora and ornata but is 
shown by Oudemans as very strongly punctate. 

Gahrliepia (Gahrliepia) decora sp. n. 
Plate 80, fig. A-D. 

Description of Larvae. Shape broadly oval. Length (partially engorged) 
to 410//., width 350/x. Scutum large and tongue-shaped, with many large but 
not touching, rounded, depressions in between which the surface is finely pitted 
or shagreened : sensillae missing in all specimens, but sensillae bases well behind 
line of PL,- PL comparatively near to AL; AL short and fine, PL very long, 
thick and strongly ciliated. Byes 2 + 2, small and lenticular, close to margin 
of scute. Palpi stout, tibial claw long and bifurcate, prongs subequal ; seta on 
femur and genu lightly and shortly branched, on tibia dorsal and lateral nude 
but ventral doubtful. Chelicerae simple, with only the apical tricuspid cap. 
Galea! setae nude. Dorsal setae long, thickly ciliated and tapering from 92-0/x 
long (humeral) to 36*0/* posteriorly, 34 in number, arranged ca. 2.6.6 [2]. 6 [2]. 
4,4.2.2.2, those on the scutum rather shorter than the outer. Ventrally 
with paired branched setae on maxillae, one on each coxa, a pair between 
coxae I and between coxae III and thereafter ca. 56 setae, to 30/* long. Legs: 
1 7-segmented, II and III 6-segmented ■ tarsi I and II with dorsal sensory rod, 
III without any long nude seta. Coxae finely pitted and III large and 
triangular. 

The Standard Data for the type and 6 paratypes are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


41»6±0-40 


l-06±0-98 


38-4-44.8 


39-2-42-0 


2-5 


PW 


66-8±0-73 


l-93:±0-52 


61- 0-72 -6 


64-4-70-0 


2-9 


SB 


30*8 


No variation recorded 






AflB 


82. 4 


No variation recorded 






PSB 


239-6±3-99 


10-57±2-82 


207-9-271.3 


224-0-257-6 


4-4 


SD 


262-C±3-99 


10-57±2-82 


230-3-293-7 


246-4-280-0 


4-0 


A-P 


17-2±0-73 


l-93±0-52 


11-4-23-0 


14-0-19-6 


11-2 


AL 


26-82:0-56 


1-50^:0.40 


22-3-31-3 


25-2-28-0 


5-6 


PL 


108 -8:fcl -55 


4-10+1-09 


96-5-121-1 


103-6-112-0 


3-8 


Sens. 


Missing. 











304 Records of the S.A. Museum 

Loc. and Hosts. The type and 6 paratypes from Rattus sabanus. from 
Pahang Rd., Goinbak Forest Reserve, near Kuala Lumpur, F.M.S . 15 Nov., 
1948 (J.R.A.). 

Remarks. Allied to the preceding species in the ornamental dorsal scutum 
but distinguished therefrom as in the key and as stated under that species, 

Gahrliepia (Gahrliepia) obnata sp. n. 

Plate 81, fig. A-D. 

Description of Larva. Shape broadly oval. Length (partially engorged) 
416/x, width 286/x. Scutum ornamented, as figured j with PL very much longer 
and stronger than AL and situated very close thereto, and in front of SB; 
sensillae missing. Eyes not observed. Chelicerae with only the apical tricuspid 
cap. Galeal setae nude. Palpi as figured, with bifurcate tibial claw ; seta on 
femur and genu branched, on tihia all nude except ventral. Dorsal setae thick, 
long and strongly ciliated, 2.4 [2]. 8 [2]. 8.8.8.6, to 84-60/* long; the extra seutal 
setae 56/a long and more slender. Ventrally with paired ciliated setae on 
maxillae, one on each coxa, a pair between coxae I, two rows of 4 each between 
coxa III, and thereafter 8.8.8.2.10.2.2, from 20 to 42/t long. Scutum and coxae 
finely punctate. Legs: I 286/* long, II 260/4, III 325/t; coxae III large and tri- 
angular; tarsi I and II with dorsal sensory rod, III without any long nude 
seta. 

The Standard Data for the unique type are: AW 40-0, PW 59-0, SB 33-6, 
ASB 25-2, PSB 218-4, SD 243-6, A-P 12-0, AL 28-0, PL 98-0, Sens. — . 

Loc, and Host. A single specimen from Rattus sab anus from Ulu Langal 
Forest Reserve, Kuala Lumpur, 18 Nov., 1949 (JJR.A.). 

Remarks, Close to G. (G.) decora sp. n. and G. ((?.) insigne sp. n. in the 
ornamental scutum and the peculiarly shaped coxae III. Closest to decora in 
having PL in front of SB, but distinguished by the ornamentation, the dorsal 
setae, and the Standard Data, 

Gahrliepia (Gahrliepia) cetrata Gater 1932. 

Gahrliepia cetrata Gater 1932, Parasitology, 24; Womersley and Heaslip 1943, 
Tr. Roy. Soe. S, Aust,, 67, (1), 140; Sig Thor and Willmann 1947, Das 
Tierreich, 71b, 338. 

Plate 82, fig. A. 

This species has apparently not been met with since it was originally 
described from 13 specimens from the ears and around the anus of Rattm 
edumrdsi ciliatus (Bonhote) (coll. A. K. Cosgrove, 25 Apr., 1930). The dorsal 
scutum is longer than wide and the posterior prolongation takes in the two 



Womersley — Asiatic-Pacific Scrdb Typhus Mites 305 

median setae of both the second and third dorsal rows. The sensillae are 
clavate and setulose, with their bases about midway between AL and PL. The 
PL setae are the longest. Eyes are stated by Gater to be just visible near the 
scutal margins, posterior eyes the smaller. The chelieerae are apparently 
simple with only the usual apical tricuspid cap. Galea! setae nude. The palpal 
claw is trifureate and the seta of palpal femur with a few barbs, of genu 
rarely so, of the tibia the dorsal and lateral are nude and the ventral with a 
few barbs. The dorsal setae are 36, arranged 2.6 [2] .6 [2] .6.6.4.4.2. 

The Standard Data as deduced by Womersley and Heaslip (1943) from 
Gater's data are: AW 42-0, PW 90-0, SB 49-0, ASB 35-0, PSB 155-0, SD 
190-0, Aj-F 65-0, AL 40-0, PL 55-0, Sens. 37-0 with head 12/30. 

Gahrliepia (Gateria) hirsuta (Radford 1946). 

Gateria hirsuta Badford 1946, Proe. Zool. Soc. London, 116, (2). 

Plate 82, fig. B-D. 

The description of the paratype which I have received from Radford is as 
follows r 

Shape oval. Length (partially engorged) 270/x, width 180/i. Dorsal 
scutum pentagonal, much longer than wide, lateral margins concave slightly, 
posterior margin forming a deep triangle, PL setae anterior of SB and adja- 
cent to AL at the anterolateral corners, SB wide apart, with sensillae clavate 
and setulose j posteriorly the scutum takes in the whole four setae of the 
second dorsal row, the median 2 of the 4 setae in the third row, and the median 
2 of the 6 in the fourth row. The chelieerae arc simple with only the usual 
apical tricuspid cap. Galeal setae ciliated. Eyes apparently absent. Palpal 
with tibial claw trifureate, with the prongs closely adpressed; setae on palpal 
femur and genu shortly branched; on tibia all 3 setae nude. The dorsal setae 
number 34, arranged 2.4 [4] .4 [2]. 6 [2]. 6.4.4.2.2. to 50/a long. Ventrally with 
the usual pair of branched setae on maxillae, one on each coxa, a pair between 
coxae I and between coxae III and thereafter ca. 44— 46, to 30/» long. Legs: 
I 240/* long, II 220/x, III 270/x; tarn I and II with the usual dorsal sensory 
rod; III without any long nude seta. 

The Standard Data are; AW 42-0, PW 57-0, SB 42-0, ASB 21-0, FSB 
112-0, SD 133-0, A-P 12-0, AL 45-0, PL 58-0, Sens. 32-0 with head 9/25. 

Loo. The specimen examined above is labelled as from a mole, from Kang- 
latongbi, Manipur, 22 June (C. D. Badford). 

Remarks. This species is remarkable in the manner in which PL has 
moved forward to adjoin AL at or near to the antero-lateral corners of the 
scutum. In Badford 7 s own figure the PL setae are actually shown as being 



306 Records of the S.A. Museum 

lateral of AL, in the above specimen tbey are slightly but distinctly posterior 
ofAL. 

From the above description it will be seen that I do not agree with Rad- 
ford 's interpretation that there are 2 AL setae on each side of the scutum. 
The outermost of each such pair I regard as the PL setae which have migrated 
forward in an unusual manner. This is supported by the fact that they are 
much longer than the other scutal and dorsal setae, a feature which is gener- 
ally common in the family. 

Radford gives the Standard Data measurements as: AW 57 «0, PW 68-0, 
SB 50-0, ASB 23-0, PSB 119-0, SD 142-0, A-P 45-0, AL 72-0, PL 48-0, 
Sens. 34*0, which in general, although somewhat higher agree reasonably well 
with my measurements as given in the key except that his AL corresponds to 
my PL, and his PL I consider to be one of the DS embraced by the scutum. 
My AL is his second AL which he gives as 54*0//, long and in his figure is shown 
as being slightly inside but closely adjacent, to his first AL (my PL). 

It was described from a shrew 8 uncus (Crocichira) caerule-us fulvacinereus 
(Anderson) from Kauglatongbi, Imphal, Manipur, 29 April, 1945. A second 
specimen was recorded by Radford from the same host and locality, June 21, 
1945, and 13 specimens from a mole from the same locality, 22 June, 1945. 

I have also examined another specimen, Coll. No. 61, from S. Burma 
^J. R. Audy), which has an extra pair of additional setae on the scutum, i.e. 
6 besides AL and PL. Even in Radford's paratype referred to above there are 
5 additional setae as figured here. It would appear that the setal counts of 
the second and third rows are somewhat variable in this species. 

The Standard Data of this specimen are: AW 38-4, PW 54-4, SB 41 <b, 
ASB 22-4, PSB 108-8, SD 131-2, A-P 9-6, AL 41-6, PL 64-0, Sens. — . 

The peculiar situation of the PL setae in this species, and also in other 
undeseribed material, has suggested to some of my American colleagues that 
they should be generieally separated. As, however, the nymph of this species 
is known to me, and cannot be separated, except specifically,, from other Gahr- 
liepia species, the raising of a. new genus is unwarranted. 

Gahrliepia (Gateria) lancearia (Radford 1946). 
GaUHa lancearia Radford 1946, Proc. Zool. Soc. London, 116, (2), 256. 

Plate 82, fig. E. 

Radford described this species from a specimen (or ? specimens) found on 
a mole Talpa micrura Hodgeson, at Kanglatongbi, Imphal, India, 22 June, 
1945. It does not appear to have been met with since. 

From Radford's figure and brief description the chief characteristic 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 307 

appears to be the stoutly lanceolate, not clavate, sensiliae with strong ciliations. 
The scutum Is fairly large, roughly pentagonal, with the posterior angle well 
produced and taking in 4 setae from the second dorsal row and 2 from the 
third. The sensiliae bases are at about midway between AL and PL. The 
dorsal setae number 44 and are arranged 2.8 (4). 8 (2). 8.8.6.4, Ventrally and 
posterior of coxae III ea. 56 setae. Coxae all 1-setose. No particulars of palpi 
and mouthparts are given. 

The Standard Data as given by Radford are: AW 51-0, PW 85-0, SB 
51-0, ASB 27*0, PSB 119-0, SD 146-0, A-P 35 <0, AL 44-0, PL 68-0, Sens. 
40 -0. 

Gahrliepja (Gateria) longitilis (Radford 1946). 

Gateria longipilis Radford 1946, Proc. Zool. Soc, London, 116, (2). 

Plate 82, fig. P. 

This species does not appear to hare been met with since Radford described 
it from a shrew, Simciis (Crocidwra) eaerttleus fulvocinereiis (Anderson) from 
Imphal, Manipur State, India, 8 May, 1945. No specimens were found amongst 
the large amount of material which I have been able to study from Burma. 

The dorsal scutum is strongly punctate, elongate pentagonal and taking in 
6 of the twelve setae in the second dorsal row, and 3 of the nine in the third 
row; the dorsal rows of setae are strongly curved forwards. Sensiliae missing. 
All coxae are l-setose and strongly punctate. Posterior of coxae III ca. 56 
setae. Dorsal setae 2.12 [6] .9 [3]. 8.6.4.4.2 = 47. 

The Standard Data (after Radford) are: AW 51-0, PW 85-0, SB 51-0, 
ASB 26-0, PSB 116-0, SD 142-0, A-P 34-0, AL 40-0, PL 57-0, Sens. — . 

Oakrliepia (Gaterta) ckocidura (Radford 1946). 
Gateria croddura Radford 1946, Proc. Zool. Soc. London, 116, (2). 

Plate 82, fig. G. 

No fresh records of this species have been published, nor have any speci- 
mens been detected amongst the large amount or* Burma material submitted to 
me. 

The following figure and details are derived from those of Radford. 

Scutum elongate pentagonal, punctate ; the apex produced as a long triangle 
with rounded lip, and taking in the median 4 setae of the second dorsal row, 
and the median 2 of the third row. The dorsal setae number 48, and are 
arranged 2.8 [4]. 8 [2 J. 8.8.6.4.4. Ventrally all coxae are l-setose and posterior of 
coxae III there are 58 setae. No details of the palpi and mouthparts are avail- 
able. 



308 Records of the S.A. Museum 

The Standard Data as given by Radford are: AW 51*0, PW 85-0, SB 
51*0, ASB 27 -0, PSB 119-0, SD 146*0, A-P 36-0, AL 44*0, PL 57*0, Sens. — . 

The species was described from a shrew Suncus (Crooidura) caerideus ftd- 
vocinereus (Anderson) from Imphal, Manipur State, India, 8 May, 1945. 

Gahrliepia (Gateria) romeri sp. n. 
Plate 83, fig. I. 

Description of Larva, Shape broadly oval, sides rather flattened. Length 
(engorged), 624ft, width 540/*, Scutum finely punctate as figured, strongly 
produced behind line of PL and taking in 6 extra setae of the second and third 
dorsal rows,- SB wide apart and about midway between AL and PL; widest 
part behind line of PL (and dorsal setae) shortly ciliated. Eyes 2 + 2, small. 
Chelicerae with only the apical tricuspid cap. Galeal setae nude. Palpi with 
bifurcate tibial claw; setae on femur and genu nude, on tibia ventral only 
branched. Dorsal setae 36 in number, arranged 2.8 [4]. 6 [2] .6.6.4.2.2, from 42 
to 36/a long. Ventrally with a pair of ciliated setae on maxillae, a pair between 
coxae I and between coxae III, one on each coxa, those on coxae III long to 
■56/jl, after coxae III with ca. 54 setae from 22-36//. long. Legs I and II with 
dorsal sensory rod on tarsi; no long nude seta on tarsi III; I 260/* long, II 
234/., Ill 273/*. 

The Standard Data, for the unique type are: AW 50-0, PW 86*0, SB 50*0, 
ASB 28-0, PSB 157-0, SD 185*0, A-P 50*0, AL 39-0, PL 42-0, Sens. 33*6 
with head 14/22. 

Loe. and Host. A single specimen from Rattus ratius, from Hong Kong, 
1950 (J. D. Ronier). 

Remarks, This species is very near to, and may only be a variant of 
<}. crocidura Radford. It appears to differ, however, in the Standard Data, 
particularly the larger PSB and A-P, the position of SB, and in the smaller 
number of dorsal setae. 

Gahrliepia (Gateria) rutila Gater 1932. 

Gahrliepia rutila Gater 1932, Parasitology, 24; Womersley and Ileaslip 1943, 
Tr. Roy. Soc. S. Aust., 67, (1), 138. 

Gateria rutila, Ewing, 1938, J. Wash. Acad. Sci., 28, 295; Sig Thor and Will- 
mann, 1947, Das Tierreich, 71b, 340. 

Plate 83, fig. A-C. 

Gater described this species from 7 specimens from Ratine mulleri validus 
(Miller) from Sungei Bnloh, Federated Malay States, and also recorded two 
specimens from R. edwardsi ciliatus (Bonhote) from Pahaug (coll. A. K. Cos- 
grove, 25 April. 1930). 



Womersley — Asiatic-Pacific Scrub Typhus Mites 309 

Dorsal scutum very large and tongue-like, taking in 4 of the six setae In 
the second dorsal row, 2 of the four in the third row and the 2 setae which can 
be considered as constituting the fourth row, i.e. it carries 12 setae in all ; the 
maximum width of the scutum occurs rather posterior of the line of PL. The 
sensillae are unknown, but their bases are slightly nearer AL than PL. The 
chelicerae are simple with only the usual apical tricuspid cap. Galeal setae 
nude. Eyes present but small. Palpi small, tibial claw trifurcato; all setae 
on femur, genu and tibia nude. Dorsal setae short and barbed, 34 in number, 
arranged 2.6 [4] .4 [2] . [21.6.4.4.4.2. 

The Standard Data as deduced from Gator's data by Womersley and Heas- 
lip (1943) are as follows: AW 43-0, PW 73-0, SB 41-5, ASB 17-0, PSB 
99-5, SD 116-0, A~P 35-0, AL 18-0, PL 18-0, Sens. — . Max. width 83-0. 

Gahrliepia (Gateria) ciliata Gater 1932. 

G-alirliepia ciliata Gater 1932, Parasitology, 24; Womersley and Heaslip 1943, 
Tr. Roy. Soc. S. Aiist, 67, (1), 140. 

Gateria ciliata, Ewing 1938, J. Wash. Acad. Sci„ 28, 295; Sig Thor and Will- 
maim, Das Tierreieh, 71b, 339. 

Plate 83, fig. D. 

This species was described from a single specimen only from Rattus nmlUri 
■vdlidus (Miller) from Sungei Buloh, Federated Malay States. 

Dorsal scutum very large, and broadest at about one-fourth from the pos- 
terior end, and taking in all the four setae of the second dorsal row and 2 of 
the four in the third, fourth arid fifth rows. According to Gater the palpal 
claw is trif urcate j the setae on the palpal femur, genu and tibia all nude ; the 
ehelicerae are simple with only the usual apical tricuspid cap, and the galeal 
setae are nude ; eyes present 2 + 2, the posterior the smaller \ dorsal setae 36 
in number and arranged 2. [4] .4 [2] .4 [2] ,4 [2 J. 6,4.2.4.2 

The Standard Data from Gater's figure and data as computed by Womers- 
ley and Heaslip 1943 are: AW 40-0, PW 70-0, SB 42-0, ASB 30 -0, PSB 159-0, 
SD 189-0, A-P 53-0, AL 45-0, PL 38-0, Sens. — . 

Gahrliepia (Gateria) spintjlosa (Radford 1946). 
Gateria spinulosa Radford 1946, Proc. Zool. Soc. London, 116, (2), 252. 

Plate 83, fig. E. 

This species was described from a shrew Suncus (Crocidara) caeruleus ful- 
vocwereus (Anderson) from Iruphal, Manipur State, India, 8 May, 1945. 



310 Records of the S.A. Museum 

Amongst tbe material collected by T. J. Lawrence from Southern Burma (slide 
No. 60) is a single specimen which belongs to Radford's species. It has the 
following Standard Data: AW 48-0, PW 76-8, SB 48-0, ASB 25-5. PSB 
160-0, SD 185-6, A-P 32-0, AL 32-0, PL 35 -2, Sens. — . The dorsal setae are 
2.4[4].8[4],8[4].6[1].6.6.4.2, i.e. 13 additional setae on the scutum. 

Recently received from J. R. Audy from host No. 7,074 (unspecified) from 
Bukit Lagong, Kepong, Kuala Lumpur, 13/1/50 are 12 specimens also of this 
species. 

From this material the following details are now given : 

The scutum is large, elongate, with sparse punctations. It is widest behind 
PL, but the sides are initially slightly concave behind PL, and posteriorly the 
margin becomes rounded. In the typical form the scutum carries 12 additional 
setae, but this number is variable. In Lawrence's Burma specimens there are 
13, while in the series of 12 specimens from Malaya, the number varies from 
8 to 14 with an average of 11*4. The higher number of 14 might suggest a 
variant of Gahr. {Gateria) fletckeri Gater which typically has 16 additional 
scutal setae and a scutum of about the same length and width. In fletckeri, 
■however, the arrangement of scutal setae is different, the greatest width is 
midway of the scutal length and the sides behind PL are evenly convex. 

The palpal claw is trifurcate; the seta on the femur with indistinct barbs, 
on the genu nude, on the tibia all nude except the ventral which is apparently 
branched. The eyes are 2 + 2, the larger anterior being distinct, the posterior 
more or less vestigial. The chelicerae with only the apical tricuspid cap. Galeal 
setae nude. All coxae 1-setose. 

The Standard Data for the 12 specimens from Malaya are : 







Standard 


Theoretical 


Observed 


Coeff. of 




Mean 


Deviation 


Range 


Range 


Variation 


AW 


51'9zt0-40 


l-39±0-28 


47 -7-56 ?! 


50-4-53-2 


2-7 


PW 


77'7:±Q-70 


2-42±0-59 


70- 4-85-0 


72-8-81-2 


3-1 


8B 


46-7:±0*40 


1-38+0-28 


42-5-50-9 


44*8-47-6 


2-9 


ASB 


25-2 


No variation 


recorded 






PSB 


163 -8:1:0 -64 


2-23:fc0-46 


157*1-170-5 


162-4-168-0 


1-2 


BD 


188-8;±:0-73 


2-52±0-50 


181.2-196-4 


184 -8-193 -2 


1-3 


A-P 


41-3±0-36 


1- 24+0-25 


37-<>-45-0 


39-2-42-0 


3-0 


AL 


37-3:±0-63 


2-18+0-44 


30-8-43-8 


33-6-39*2 


5-8 


PL 


40-S±0*64 


2-22+0-45 


34-1-47-5 


36-4-42-0 


5-4 


Sena. 


Aliasing in all sj 


>ecimens. 









Womersley— Asiatic-Pacific Scrub Typhus Mites 311 

Gahrliepia (Gateeia) fletcheri Gater 1932. 

GdhrUepia fletcheri Gater 1932. Parasitology, 24 ; Womersley and Heaslip 1943,. 
Tr. Roy. Soc. S. Aust., 67, (1), 140. 

Oateria fletcheri, Ewing 1938, J. Wash. Acad. Sei., 28, (6), 298; Sig Tlior 
and Willmann 1947, Das Tierreich, 71b, 339. 

Plate 83, fig. F-EL 

This species was described by Gater from numerous specimens from Rattus 
rattus diardi (Jentink), Kuala Lumpur, F.M.S. He also recorded it from 
Sungei Buloh, from Rattus sab amis vociferans Miller, Seiurus canioeps conoolor 
Blyth, Tupaia gtts ferni^pinea Raffles, and Rhinosmtrus tupaiodes laticaudatus. 

For this species, and to include also Gater 's ciliata and rutila Ewing 1938 
erected the genus Gateria, stressing that it was not the number of setae on the 
scutum, but the fact that some of the setae are not marginal, which was import- 
ant. He restricted Gahrliepia to the type species nana Ouds. from Africa, 
with 4 pairs of normal scutal setae and retained l)eiig<ilensi$ Hirst, with 3 pairs 
of scutal setae in Schongastiella, in disagreement with Gater. In 1943, Wom- 
ersley and Heaslip showed, however, that any scutal setae posterior of PL could 
be attributed to members of the dorsal rows, embraced by the prolongation of 
the scutum; and as this is the most natural and logical interpretation it seems 
better to retain them all in Gnhrliepia. 

In this species the dorsal scutum is very large, tongue-shaped, much longer 
than wide, and from the paratype in the South Australian Museum, takes in 
all the 4 setae of the second dorsal row, 8 of the ten of the third row, 2 of the 
four of the fourth row, and 2 of the six of the fifth row. (Gater states that 
the number of setae on the scutum is somewhat variable.) There are 2 eyes 
on each side fairly close to the scutum. AL setae longer than PL. Sens, cla- 
vate and setulose with bases wide apart but nearer to AL than to PL. Cheli- 
cerae simple with only the usual apical tricuspid cap. Galea! setae nude. 
Palpal claw trifurcate, the prongs -closely adpressed. The setae on the palpal 
femur and genu have a few inconspicuous barbs; on tibia all 3 setae nude. 
The dorsal setae number pa, 46, arranged 2.4[4].10[81.4[2].6,8A2. The 
Standard Data of the paratype as given by Womersley and Heaslip 1943 are : 
AW 52-0, PW 79-0, SB 50*0, ASB 18-0, PSB 170-0, SD 188-0, A-P 39-0, 
AL 50-0, PL 37-0, Sens. — . Max. width of scutum 100-0 



312 Records of the S.A, Museum 



ADULTS AND NYMPHS. 

While this paper is essentially a Monograph on the ' ' Trombicuiidae of the 
Asiatic-Pacific Region" it has become necessary, to properly evaluate, at least, 
some of the many larval genera which have been proposed, to consider all the 
adults and nymphs described from the whole world. Consequently in this 
section, the world species (relatively few compared with the described larval 
forms), are all dealt with, both descriptively and, as far as possible, with those 
of which material has not been available for study, by previously published 
figures. One species, Trombicula signata Womersley 1938, described as be- 
longing to the genus Trombicula Berl. s.l. does not belong to the Trombicuii- 
dae, but is a species of Podothrombium. 

The genera as known from adults or nymphs belong to both subfamilies, 
the Trombiculinae and the Gahrliepiinae, and may be keyed as follows t 



Key to the Adult and/or Nymphal Subfamilies and Genera of the 
TEOMBICULIDAE S. Str. 

1. Tarsi I simple without any dorsal subapical stump-like process 

Subfam. Tromlnculinae Ewing 1944. 

Tarsi I subapically with a short dorsal stump-like process. Sensillary 
area of crista wider than long, with the anterior rod shorter than 
distance between sensillae bases. Eyes absent. Epistome rounded or 
conical with fine denticulations and 1 ciliated seta. No precoxal 
plates and no definite sternum. 

Subfamily Gahrliepiinae nom. nov. 
for Walchihiae Ewing 1931. 
Gen. Oahrliepia Ouds. 1912. 

(type Typhlothrombium nanus Ouds. 1910) 
= Walehia Ewing 1931 

(type Trombicula glabrum Walch 1922) 
incl. Schongasticlla Hirst and Gateria Ew. 

2. Eyes distinctly present, one on each side 

Eyes entirely absent or vestigial and without lenses, at most repre- 
sented by pigment spots 

3. Eyes placed well away from and in front of sensillary area. Epis- 
tome rounded and subtriangular, denticulate, with 1 ciliated seta. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 313 

Sternum well defined and formed by fusion of preeoxal plates of 
leg I, entire, -without longitudinal division, wider than long. 

Gen. Tmgardhula Beri, 1912. 

(type Trombidmm nilotunm Trag. 1904 (adult)). 
— Blmkmrtia Berl. 1912 (non Ouds. 1911) (adult) 
= Pentagonella Sig Thor 1936 (larva). 

(type Trombidmm ardeae Trag. 1904). 
= Megatrombicula Michener 1947 (adult) 

(type Trambictda alleei Ewing 1926). 

Eyes placed closely adjacent to sensillary area . ~ . . . . 4 

4. Front tarsi elongate-clavate, with the greatest height distad of middle. 
Body form elongate. Crista flask-shaped, without ? an areola in sen- 
sillary area; anterior rod of crista shorter than distance between sen- 
sillae bases ; a strong ciliated seta on crista just anterior of sensillary 
area. Epistome a transverse rectangle, anterior margin dentate, one 
ciliated seta. Sternum entire, no preeoxal plates. 

Gen. Ipotrombicula nov. 

(type Tromblcula elegans Womersley 1942) (adult). 

Front tarsi tapering, with greatest height basad of middle. Body 
form squatter. Crista with anterior rod longer than distance between 
sensillac bases; subposterior sensillary area roughly diamond-shaped, 
with an internal somewhat dumb-bell shaped areola ; sensillae filamen- 
tous; no seta on crista in front of sensillary area. Epistome a trans- 
verse rectangle or rounded-conical, with dentate margin and 1 ciliated 
seta. Sternum entire, about as long as wide, no preeoxal plates. 

Gen. Troiribicxdki Berl. 1905 (adult) in part, 
(type Tronibicida minor Berl. 1905) 
(incl. Eutrambicula Ewing 1938) 

Subgen. Trombicula Berl. 1905, s, str. 

o. Legs I much longer than others, and than the body. Form more elon- 
gate. Claws strong, on legs I apically furcate. Palpal tibia in adult 
with only 1 accessory spine at base of claw. Epistome a transverse 
rectangle with dentate margin and I seta, which apparently has only 
1 short branch. Sternum entire; no preeoxal plates. 

Gen. Speotrombicula Ewing 1946 (adult), 
(type Trombicula trifurca> Ewing 1946), 
Legs all shorter than body. Form squatter. Claws normal . . . . 6 

6. Sensillary area of crista much wider than long, with anterior rod 
shorter than distance between sensillae bases. Freeoxal plates present 
on coxae I. 

Gen. Gxmtheraym Worn, and Heasp. 1943. 

(type Neoschangastia bipygalis Gunther 1939). 
Sensillary area of crista about as wide as long, with anterior rod 
much longer than distance between sensillae bases. Preeoxal plates 
on coxae 1 present or absent . . . . 7 



314 Records of the S.A. Museum 

7. Sensillary area entire, not divided into sections by longitudinal ridges, 
reniform with hilus directed forwards, with more or less circular 
striae. Sensillae spathulate, sometimes almost filiform. 

Gen. Neosclwngastia Swing" 1929 (larva), 

(type Schongastia americana Hirst 1921). 
= (Parazchdngastw} Worn. 1939) (larva). 

Sensillary area not as above . * . . . . * . ► . . . 8 

8. Sensillary area of crista triangular with apex directed posteriorly. 
Epistome rounded or conical with fine dentations and 1 ciliated seta* 

Gen. Schongastia Ouds, 1905 (larva) 

(type Thrombidium vandersaiidei Ouds. 1905) 
(inel, Ascosckongastia Ewing 1946). 

Sensillary area more or less diamond-shaped. 

Gen. Trombicula Berl. 1905 (in part) 9 

9. Precoxal plates of coxae I well developed, fused in the medial line 
to form a longitudinally divided sternum. 

Subgen. LeptotrombidiumNaLg&yo et ah 1917 (adult) 
(type Trombidium akam'mhi Brumpt. 1910). 

Precoxal plates absent. Sternum entire, undivided. 

Subgen. Neotromiicula Hirst 1925 (nymph), 
(type Acarus autumnalis Shaw 1790). 

Subfamily Trombiculinae Ewing 1944. 

This family is characterized as defined by Ewing 1944b and 1946, in 
that the anterior dorsal scutum of the larvae has a single median seta present 
on the anterior margin. In the present work it is separated in the adult and 
uymphal stages from the Gahrliepiinae (= Walchiinac) in that there is no 
dorsal subapieal clavate process on the front tarsi. It contains the following 
adult genera: Tragardhula Berl. 1912, Trombicula Berlese 1904, (subgenera 
Leptotrambidium Nagayo et al. 1917 and Neotrombicida* Hirst 1926), Speo- 
trombicula Ewing 1946, Ipotrombwula- Worn. g. nov. Schongastia Oudemans 
1905, separated as in the preceding key. 

Genus TRAGABDHtTLA Berl. 1912 

Tragardhula Berl. 1912 7 Redia, 8, (1), 4 (type Trombidium nihticum Trag. 

1904, adult), 
Blankaartia Berl. 1912, ibid. 96, non Oudemans, 1911. 
Pejhtagonella S. Thor 1936, Zool. Anz., 114. 
Megatrombicula Michener 1946, Ann. Ext. Soc, Amer,, 89, 431-445. 

The status and synonymy of this genus has recently been discussed by 
the author (Tr. Roy. Soc. S. Aust., 72, (1), 1948). 



Womersley — Asiatic-Pacific Scrub Typhus Mites 315 

Key to tlie Known Adult Species. 

1. Palpal tibia with a pectine of 8-17 spines on inner surface. Tarsi I 
ea. 3 times as long as high, 540/x X 170/x, metatarsus I 350/* long. 
Dorsal setae to 155/* long. 

Trag. velascoi (Boshell and Kerr, 1942). 

Palpal tibia without a pectine, with only the usual 3 accessory spines 
near base of claw . > . .. * « . . . .. 2 

2. Dorsal setae with apices devoid of ciliations. Trag. peruviana (Ewing 1926). 
Dorsal setae ciliated to tips . . 3 

3. Tarsi I parallel sided, more than 4 times as long as high, 370/* X 
80/*, metatarsus I 300/* long. Dorsal setae to 70b*. 

Trag. attemiaia (Mich. 1946). 

Tarsi I not parallel sided >> . . , 4 

4. Tarsi I ca. twice as long as high, 325/* X 163/*, metatarsi I 247/* long. 
Dorsal setae to 65/*. Trag. acuscutellaris (Walchl922). 

Tarsi I more than twice as long as high . . . . . . . . . . 5 

5. Dorsal setae to 160/* long. Tarsi I nearly 3 times as loug as high, 
350/* X130/A, metatarsi 1250/* long. Trag. alleei (Ewing 1926). 

Dorsal setae much shorter. 

6. Tarsi I ca. 3 times as long as high, 102 • 5/x X 32 ■ 8/*, metatarsus I 
65-6/*. Dorsal setae 25/* to 58//. (Nymph, after Tanaka). 

Trag. josponica (Tanaka 1916). 

Tarsi I ca. 3-5 times as long as high, 370/* X HO/*, metatarsus I 270/*. 
Dorsal setae 3040ft long. Trag. nilotica (Ouds. 1904) . 

Tragardhula nilotica (Trag. 19/34 adult). 

Tronibidmm niloticum Tragardh 1904, Res. Swed. Zool. Exped. Egypt and 
White Nile, 1901, No. 20, 80 (adult, mn larva). 

Tra<gardh%la nilotica Berl. 1912, Redia, 8, 4 (adult) ; Sig Thor and Willmann, 
1947, Das Tierreich, Lfg. 71 b., 353, 

Manka&rtia nilotica Berl. 1912, Redia, 8, 96 (adult) ; non Oudemans 1911, 
Entom. Ber., 3, 123 (larva) and Oudemans 1912, Zool. Jahrb. Suppl., 14; 
Cooreman 1948, Explor. van. het Nat. Park Albert, Afl, 14, 18 (adult). 

TragardhuU nilotica Womersley 1948, Tr. Roy. Soc. S. Austr., 72, (1). 

Plate 84, fig. A-F. 

Be-de^cription of Adult (from specimen from India) : Faeies as in Tram- 
l>ic-vla. Colour in life whitish (after Berlese). Length to 1,750/*, width to 
1,050/* (after Berlese). Crista elongate, 260/* with subposterior sensillary area 



316 Records of the S.A. Museum 

about as long as wide, with paired short nude sensillae to 52/x long and their 
bases 52/x apart; epistome rounded, with fine dentieulations and 1 ciliated seta. 
Eyes I on each side, very large, 72/a in diameter, and placed about midway along 
and well away from crista. Chelicerae with finely serrate inner (dorsal) 
edge. Palpi fairly stout; tibia with simple claw, and 3 accessory spines which 
are almost eiaw-like and placed about midway between base of claw and 
articulation of palpal tarsus, palpal tarsus elongate reaching beyond tip of 
claw. Legs shorter than body, I the longest, 1,650/* ; tarsi I elongate and highest 
in distal half, 375/x long X 123/x, high ; metatarsi I 292/* long. Sternum entire, 
much wider than long; precoxal plates of leg I not evident. Dorsal setae 30- 
40/«, uniform, fairly and uniformly thick to tip, with short ciliations. Genital 
discs 3 pairs. 

Loe. mid Remarks. This species which is the genotype, was originally 
described from specimens found on the leaves of an aquatic plant, Pistia, on 
the White Nile. It was associated with a larval Trombidiid which was thought 
to be its larva, but which is now regarded as not belonging to the same species, 
or even to the Trombiculidae. Womersley (1948) has recently discussed this 
question and suggested that the true larvae of Tragardhula nilottca may ulti- 
mately be shown to be Tragardhula (Pentagonelh) ardeae (Trag, 1904), which 
agrees generically in the larvae, and which is the type of Sig Thor's Pentagon- 
ella. It was found on the legs of a heron from the same habitat and locality. 

Tragardhula niloU'ca also occurs in India in the same habitat and I am 
informed by Dr. J. Cooreman of the Musee Royale Belgique, that he also has 
material from a similar habitat in the Belgian Congo, strangely enough also 
associated with larvae similar to those with which it was associated in Egypt* 
In a recent publication (1948), Dr. J. Cooreman has recorded this material,, 
which included adults, nympbochrysalides and larvae. Both adults and larvae 
agree with Tragardh's material (I was personally priviliged to see these 
while in Brussels in 1947) but no details of the nymphochrysalis are given by 
Cooreman. He considers that, the larvae and adults being found together in a 
precisely similar biotype to Tragardh's specimens, the larvae and adult* must 
be associated. With this conclusion, as pointed out above, one must disagree. 

Tragardhula acuscutellarjs (Walch 1922). 

Trombieida ami-scuteUaris Walch 1922, Kitasato Arehiv. Exper. Med., 5, (3) y 
78; Gater B. A. K., 1932, Parasitology, 24, 143-174; Mehta D. A., 1937, 
Ind- j< Med. Res., 25, (2), 353-365; Philip, Woodward and Sullivan 1946, 
Amer. J. Trop. Med., 26, (2) ■ Radford C. D. 1946, Parasitology, 37 r 
(1-2) • Jayewickreme S. H., 1947, Nature, 160, 578. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 317 

Trombicula (PentagonelUi) acuscutellarw, Sig Thor 1936, Zool. Anz., 114, 30; 
Woinersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 78. 

Tmyardhula aciiscuiellaris, Sig Thor and Willman 1947, Das Tierreich, Lfg. 
71b, 296; Womersley 1948, Tr. Roy. Soc. S. Austr., 72, (1). 

Plate 84, fig. G~N. 

Re-description of Nymph (after Womersley 1948). Colour in life bright 
red. Of typical Trcmibicula facies. Length to 800/4, width across hysteroaoma 
400/4. Crista linear with somewhat diamond-shaped sensillary area and two 
nude filamentous sensillae, 136/4 long, bases 42/* apart. Eyes 1 + 1, well 
removed from sensillary area and about half-way between extremes of crista. 
Chelicerae normal with fine inner serrations. Palpi with tibial claw strong, 
with 2 accessory spines but these on a slight boss and placed about midway 
between base of claw and articulation of palpal tarsus. Palpal tarsus slightly 
elavate, not reaching tip of claw. Legs shorter than body, I 660/4 long, 11 460/*, 
III 470/1, IV 530/* ; tarsus I less than twice as long as wide, 151/* by 91/*, meta- 
tarsus I 115/1 long. Dorsal setae numerous, 40/4 long, uniform, arising from 
closely set tubercles, uniformly thick with strong eiliations. Genital discs 2 
pairs. 

Description of Adult (after Womersley 1948). No apparent difference in 
size of males and females. Colour in life deep red. Of typical Tro-mbicula 
facies. Length 1,800ft, width across hysterosoma 1,200/4. Crista linear with 
well developed sensillary area, roughly diamond-shaped, with the sensillae 
bases at the end of transverse dumb-bell shaped areola,- crista 300/c long; sen- 
sillae bases 65/4 apart, and sensillae 195/4 long, filamentous and nude. Epistome 
well developed, rounded-conical with numerous teeth and a single seta 71/4 
long. Eyes 1 + 1, large, well away from crista and midway of crista length. 
Chelicerae with finely toothed inner edge. Palpi with strong tibial claw and 
3 accessory spines which are distinctly away from base of claw, and another 
on outer edge opposite articulation of tarsus, and on the outer surface another 
strong spine. Legs normal, I 1,275/4 long, II 910/4, III 910/4 IV 1,235/4,- 
tarsi I 325/u. long by 163/4 high, metatarsi I 247/4 long. Sternal shield between 
epimera of legs I and II, markedly shorter than wide. Genital orifice situated 
<»LoBe in to epimera of legs III and IV, with 3 pairs of discs j in the male pro- 
vided with a penis as figured, the apieal point of which is asymmetrical. Dor- 
sal setae numerous, uniformly thick, with short ciliations, to 40/4. 

Remarks. The nymphal description is from a specimen reared by C. D. 
Radford from larvae from the Maldive Islands, and that of the adult from 
captured specimens from S. H. Jayewiekreme from Ceylon (see Womersley, 
1948). The adults and nymphs can be separated from other known species aa 
in the key, 



318 Records of the S.A. Museum 

TrAgardhula japonica (Tanaka 1916). 

Lcptus autumniilis japonica Tanaka 1916, Igakkai Zasshi (in Jap.), 30, (22) ; 
1918 Ikal Jiho (in Jap.), No. 1,228. 

Tronihicula autwmnalis japonica (Tanaka et aL 1930), Zentrblt. Bakt., 116, 
(1), 361; Sig Thor and Willmann, 1947, Das Tierreieh, Lfg. 71b. 

TrwrilrieuUi japonica, Womersley and Heaslip 1943, Tr. Roy. Soe. S. Austr., 
67, (1), 79. 

Tragwrdhula japonica Womersley 1948, Tr. Roy. Soc. S. Austr., 72, (1). 

Tanaka (1930) appears to have been the only worker so far to have reared 
and described the nymph of this species. The adult is still unknown. That it 
must be placed in TragardJiula is evident from Tanaka 's description of the 
eyes, and from his photograph (PI. IV, fig. au). In the presence of eyes in 
the nymph it is not to be confused with the European autumnalis Shaw as was 
done by Tanaka, and still more recently by Sig Thor and Wilhnann 1947. The 
following description is drawn up from details given in Tanaka 's paper of 
1930. 

Nymph. Of typical Trombicula fades. Length 495//. to 790/x, width 248/a 
to 389/a. Colour in life bright red, crista, linear with vsubposterior sensillary 
area, and paired filamentous sensillae similar to those of larvae. Eyes 1 -|- 1, 
well away from crista and well in front of sensillary area. Palpi fairly stout, 
tibia ending in strong claw (accessory spines not mentioned by Tanaka, but 
doubtless two and placed as in other species of Tragardhuta.) ; tarsus oval 
three times as long as wide. Chelicerae serrated on inner (dorsal) edge. Legs 
all shorter than body; length (excluding coxae) I 664//, II 394//,, III 402//, 
IY 459/x; tarsi I slightly more than three times as long as high, 193//, by 61 '5/* 
long, metatarsi I 123/t long (those in a specimen 796//. long). Dorsal setae 
pointed and ciliated, longer posteriorly, ? to 58//.. Genitalia with 2 pairs of 
discs, the anterior pair oval and larger than the round posterior pair. 

TrAgardhula peruviana (Ewing 1926). 

Trombicula* peruviana Ewing 1926, Ent. News, 37, 112; Ewing 1931, Proc. 
U.S. Nat. Mus., 80, (8), 9; Ewing 1933, Proc. U.S. Nat. Mus., 82, (29), 2. 

Trombicula (Megatrombicula) peruviana, Michener 1946, Ann. Entom. Soe. 
Amer., 39, 434. 

Tragardh'Ula peruviana, Womersley 1948, Tr. Roy. Soc. S. Austr.. 72, (1). 

This species is as yet, only known from a single adult in the U.S. Nat. 
Mus., from El Campanicnto, Peru. The description as given by Ew r ing (1926) 
is as follows : 



Womersley — Asiatic-Pacific Scrub Typhus Mites 319 

"Colour of preserved specimens a light brownish orange. Palpi slender, 
reaching to about the tips of first femora; palpal thumb slender, not swollen, 
and slightly surpassing the palpal claw ; penultimate segment very slender and 
bearing at its free end the rather small palpal claw which is slightly over 
one-half as long as the segment bearing it. In addition to the palpal claw, the 
penultimate segment bears on its inside a comb of three long, approximate 
spines, the middle of which is longer than the other two. Eyes projecting, 
with strongly curved corneas and situated far in front of pseudostigmatic area. 
Pseudostigmatic area large, posterior lobes oval, approximate to medial line; 
pseudostigmatic organs, very slender, flagelliform, and with a few pectinations. 
Setae of body very long, especially on the shoulders and posterior margin; 
barbed conspicuously for about two-thirds their length, but the tips almost 
bare. First pair of legs much longer than, the others j tarsus I slightly swollen, 
almost straight and about one-fourth longer than tibia I j tibia I clearly longer 
than patella I; patella I and femur I subequal. Tarsus II and tarsus III about 
one and one-fourth times longer than tibia II and tibia III respectively. Tar- 
sus IV not swollen, considerably longer than tibia IV, and tibia IV »consider- 
ably longer than patella IV. Length 2-5 mm.; width 1-5 mm.** 

Tragabdhula atj»FiF.t (Ewing 1926). 

Trombicida alleei Ewing 1926, Ent News, 37, 111 ; Ewing 1931, Proc, U.S. Nat. 
Mus., 80, (8), 9; Ewingr 1933, Proc. U.S. Nat. Mus., 82, (29), 2. 

TromMculo, (Megatrombicul'a) alleei, Michener 1946, Ann. Ent. Soc. Amer., 
39, 434. 

Tragardhula alleei, Womersley 1948, Tr. Roy. Soc. S. Austr., 72, (1). 

Plate 85, fig. A-E. 

This species was described originally (Ewing 1926) from a single speci- 
men (holotype male; in collection U.S. Nat. Mus.) from Barro Colorado Island, 
Panama* Canal Zone. It was redeseribed by Michener 1946 from adults col- 
lected at Juan Mina, Canal Zone, Panama. As well as being found in the 
usual type of terrestial habitat, specimens of adults were also recorded as being 
found on the floating leaves of Pistia stratiotes, a habitat similar to that for 
TragardkuUt* ndlotica in Egypt and India, and also for Tragardhula attenua-ta 
Michener from Panama. 

Adults and nymphs were also reared by Michener from larvae. 

Description of Adult (after Michener 1946). Brilliant red in life. 
Length 1*5 to 2-5 mm. Crista rod-like, expanded gradually to the rear in to 
the triangular pseudostigmatic area, which bears the simple (and about as long 



320 Records of the S.A, Museum 

as the crista) pseudostigmata at its posterior lateral angles. Palpal tibia with 
claw, measured along upper margin, 0-30 to 0-36 times its length,- inner sur- 
face of tibia with 3 spines arising from a small projection and a fourth much 
nearer base; tarsus reaching tip of claw, slender, with 4 or 5 small bristles at 
apex and mostly simple setae. Eyes 1 + 1, large, away from crista and in 
front of pseudostigmatic area. Legs shorter than body; tarsus I swollen, 
highest near middle, less than 3 times and sometimes only a little more than 
twice as long as high, 1*30 to 1-38 times as long as preceding segment. Dorsal 
setae from 70 to 160/i long, ciliated. 

Description of Nymph (after Michener 1946). Length freshly emerged 
1-0 to 1-2 mm. Palpal tibia with only 2 accessory spines (? sometimes only 
one). Dorsal setae sparser than in adult, 50 to 70^ long. Tarsus I 1*4 to 
1*5 times as long as preceding segment. Otherwise similar to adult except for 
nymphal characteristics. 

Bionomics, Michener has made some very interesting observations on 
the habitat and life-history of this species. Adults were found at all seasons 
in wet leaf mould close to the shore of the Chagres River, under aquatic vege- 
tation left stranded, and on floating plants of Pistia strati&tes. Specimens from 
the last habitat were smaller, and more, numerous. The eggs were laid singly 
or in twos, threes or fours on leaves of Pistia, but in rearing some thirty 
larvae emerged at the same time, so it is probable that greater numbers were 
laid. This is in agreement with the fact that the female of Tragardhida nilo- 
tica from India studied by the present writer, contain some 12-15 eggs ready 
for laying. In the species of Tromhicxda s. str. the eggs appear to be laid 
singly, and rarely does one find more than 1 or 2 eggs in the gravid females. 

Remarks, An adult specimen from Juan Mina, kindly sent to me by 
Dr. Michener, has the following dimensions ; Crista as figured by Michener, 390/x 
long, with sensillae 240/1 long, nude; sensillae bases 84/t apart. The epistome 
is rounded conical with small denticles laterally, epistomal seta fine, sparsely 
ciliated, 110/x long. Tarsi of leg I 375/u. by 143p., metatarsi 273/*. Dorsal setae 
posteriorly to 84/x. Eyes smaller than in velascoi, 32/i in diameter. The 
Sternum is well defined, wider than long: no precoxal plates on leg I. 

Tragabdhula velascoi (Boshell and Kerr 1942). 

Trwnbicvla velascoi Boshell and Kerr 1942, Rev. Acad. Columbiana Cscn. 
Exact. Fisic. Nat., 5, 113. 

Tromuicula (Megatromokula) velascoi Michener 1946, Ann. Entoin, Soc. 
Amer.,39,438. 

Tragardhida velmem* Womersley 1948, Tr. Roy. Soc. S. Austr., 72, (1), 89. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 321 

Plate 85, fig. £4$ 

Originally described from Kestrepo, Intendencia de Meta, Columbia (type 
locality), and also from Villa Viceneio, and the Municipo of Bolivar, Columbia, 
this species was re-described by Michener from material from Juan Mina and 
Chiva Chiva, Canal Zone, Panama, and from Old Panama and Santa Kosa, 
Panama. 

Both adults and nymphs were collected in nearly every month of the year 
under stones and logs and in leaf mould, but frequently in relatively dry situa- 
tions in contrast to other species. 

The descriptions of the adult and nymph (after Michener) are as follow. 

"Adult. Length 1-6 to 2*5 mm. Brilliant red in life. Claw of pedipalp 
measured along upper margin 0-30 to 0-36 times length of pedipalpal tibia*, 
inner surface of tibia with eight to seventeen spines (average of twenty-five 
specimens 12*1) arranged around lateral and apical margins, those nearest apex 
consistently present, those nearer base of segment frequently absent, so that the 
row of spines is sometimes broken by large gaps; sometimes there are one or 
two spines arising near middle of segment, not in marginal row; finger of 
pedipalp with five to seven short simple bristles apicalJy, other hairs mostly 
barbed or plumose. Crista and pseudostigniatic organs similar to those of 
alleei. Body hairs about two-thirds as long on shoulders as on posterior end 
of body, plumose but the setulae smaller towards apices of hairs and sometimes 
absent from extreme apices; longest body hairs 0-15 to 0-20 mm. in length. 
Legs shorter than body, or first pair as long as body, robust; last tarsal seg- 
ment of anterior legs somewhat swollen, broadest near middle, slightly less 
than three times to over four times as long as broad, 1-49 to 1-61 times as 
long as preceding segment and ordinarily broader than that segment. 

Nymph, Length (when freshly emerged) 0*9 to 1*0 mm. Older nymphs 
reach 1*5 mm. Similar to adult except for usual nymphal characteristics. 
Claw of pedipalp sometimes nearly half as long as tibia, which bears three 
to eight spines on inner surface. Body hairs sparser than in adult, those of 
posterior end of body two or three times as long as those of shoulders ; longest 
hairs 0-09 to 0-14 mm. in length. Legs similar to adult, last segment of fore- 
leg 1-50 to 1-60 (or in one specimen, perhaps abnormal, 1-75) times length 
of preceding segment/' 

Biononms. The larvae have been reared from captured females (see Lar- 
val Section of this paper) and described by Michener 1946 

Remarks, I am indebted to Dr. Michener of the Anier. Mus. Nat. Hist., 
New York for an adult of this species, from which the crista and coxa* I and 
II with sternum are figured and the following measurements made. Crista 360/* 



322 Records of the S,A. Museum 

long, with subposterior sensillary area and sensillae long, 260/i, filamentous and 
nude with bases 78/x apart; epistome rounded-conical, 84/x long by 78/a wide at 
base, margin scarcely dentate, seta long and slender, lightly ciliated, 110/a long. 
Eyes large, to 60//, in diameter and situated away from crista and in front of 
sensillary area* Palpi with simple tibial claw; tibia with 3 strong accessory 
spines on inner tubercle. (Boshell and Kerr show 7-8 spines in the form of a 
pectine, and Michener states that in a number of specimens they vary consider- 
ably from 8 to 17 with an average of 12 ; thus the specimen examined seems to 
be an extreme form. The other spines have not been knocked off, for bases ean- 
not be seen)* The tarsus with 5-6 apical spines. Tarsi of leg I are not quite 
as long in proportion to height as shown by Mkhener, and measure 365/x long 
by 190//. high, while metatarsi I are 250/* long. The dorsal setae are somewhat 
longer anteriorly than posteriorly, to 78ft and to 65//, respectively. The setae 
on the legs are much more slender and tapering, less strongly ciliated and 
measure to 130-150/x long. The sternum is well defined, entirely enclosed, 
wider than long, and without a longitudinal division, ie, precoxal plates of 
coxae I are absent or have combined to form the sternum. 

Despite the differences between this specimen and Michener 7 s description 
I have no doubt that it is the same species. 

Tragardhula attenuata (Michener 1946). 

Trombictda (Megxrtrombicula) attenuata Michener 1946, Ann. Ent. Soc. 
Amer., 39, 440. 

Tragardhula attenuata, Womersley 1948, Trans. Roy. Soc. S. Austr., 72, (1). 

The description of the adult of this species as given by Michener 1946 is 
as follows : 

Plate 85, fig. F-H. 

"Adult. Length 2-0 mm. (varying from 1*4 to 2-1 mm. among para- 
types). Brilliant red in life. Claw of pedipalp measured along upper margin 
about O-20 (varying from 0-19 to 0-23) times length of pedipalpal tibia, 
which is extraordinarily slender ; inner surface of tibia with three blunt spines 
arising near (or basad of) middle of upper margin; finger of pedipalp with 
four or f\ye short simple bristles apically, other hairs mostly simple. Crista 
about as in other species; pseudostigmatic organs simple, longer than crista. 
Body hairs but little longer at posterior end of body than on shoulders, plu- 
mose throughout length; longest body hairs 0'13 mm. (longest hairs varying 
among paratypes from 0-05 to 0*13 mm.) long. Legs slender, anterior pair 



WOMERSLEY — ASIATIC- PACIFIC SCRUB TYPHUS MlTES 323 

longer than body, others shorter; last tarsal segment of anterior legs not 
swollen, not thicker than preceding segment, about parallel sided (sometimes 
slightly thicker beyond middle or subapically than elsewhere), four and one- 
half (varying from four to over four and one-half) times as long as broad, 
1-20 (1*19 to 1*25) times as long as preceding segment," 

Bionomics. Miehener found this species only on floating plants of Pistia 
stratioies on the Rio Hondo, along with two other species of Microtrombidiinae, 
They were found among the coarse hairs and ribs of the under-side of the 
leaves. The eggs were laid in masses of twenty-five to fifty on the under, sur- 
face of the leaves. The larvae are only known from reared specimens, and their 
hosts are unknown. 

Genua IPGTROMBIGULA nov. 

"With the general facies of Tronibmda* constricted between second and 
third pairs of legs. Crista flask-shaped, without an areola within the sub- 
posterior sensillary area; a long ciliated seta, at the base of the rod of the 
crista just anterior of the sensillary area,- epistome transverse, with straight 
front margin, denticulate, with one ciliated seta. Sensillae filamentous and 
nude. Eyes one on each side, large, closely adjacent to sensillary area. Palpi 
slender, tibia with simple claw and in adult, 3 accessory spines at base. Front 
tarsi clavate, with the greatest depth (height) distad of middle. 

Genotype Tromiicula elegans Womersley 1942. 

Ipotrombicula elegans (Womersley 1942). 
Tro?nMcula elegam Womersley 1942, Rec. S. Aust. Mus., 7, (2), 173, fig. 4 A~L 

Plate 86, fig. A-E. 

Re-description of Adult. Very elongate species but with constriction 
between second and third pairs of legs. Length 1-67 ram., width of both 
propodosoma and hysterosoma 0*58 mm,; hysterosoma about twice the length of 
propodosoma. Colour in spirit white. Crista 167/a long, linear, with a sub- 
posterior flask-shaped sensillary area without any dumb-bell shaped areola; 
with paired, nude, filamentous sensillae 96/* long, and their bases 42/x apart; 
in front of the sensillary area and on the rod of the crista is a ciliated seta 
60/t long; epistome transverse with straight front margin with fine denticula- 
tions and one ciliated seta 54/4. long. Eyes large, one on each side and closely 
adjacent to sensillary area. Chelicerae with inner (dorsal) margin finely 
serrated. Palpi slender; tibia with a strong simple, apical claw and 3 strong 
accessory spines at its base; tarsus somewhat clavate and reaching tip of tibial 



324 Records of the S.A. Museum 

claw. Legs relatively short, I 1,000/4. long, II 665,*, Ill 650/*, IV 900^; tarsus I 
clavate with the widest part distad of the middle, 245y* long by 102/x high 
(remeasured) ; metatarsus I 212// long. No distinct sternum, i.e, space between 
anterior pairs of coxae not closed posteriorly, no precoxal plates on leg I. 
Dorsal setae numerous, fine and slender, tapering with fine ciliations, 35/* 
anteriorly increasing gradually to 80a* posteriorly. Genital aperture with 3 
pairs of discs. 

Locality. This species was described from two females from Lush Island, 
South Australia, collected December, 1936 by the McCoy Expeditoin, from just 
about high water mark. No fresh material has come to hand. 

Genus SP£OTBOMBIGT7LA Ewing 1946. 

J. Parasitology, 1946, 32, (5), 437. Genotype Trombicula trifurm Ewing, 
1933, Proc. U.S. Nat. Mas., 82, (29), 1-6. 

This genus is so far only represented by the genotype. The generic diag- 
nosis as given by Ewing 1946 is as follows : 

"Adult stage. Palpi slender, slightly curved; palpal claw acute at apex, 
with a single accessory spine near base \ palpal patella not emarginate ventrally. 
Epistome a poorly sclerotized band-lite comb. One rostral seta present. Eyes 
absent. Crista broad, plate-like; pseudostigmata small, far apart; pseudostig- 
matic organs minute, setiform. Abdomen slightly constricted in front of 
middle. Dorsal abdominal setae barbed, of one kind, longest on posterior mar- 
gin. Genital opening large, with 3 pairs of genital suckers. Coxae I with a 
shelf -like expansion along its anterior margin. Coxae I and II united. Anter- 
ior legs much longer, and stronger than others and each ending in a pair of 
subequal, curved, distally trifurcate claws. Legs II, III and IV nearly equal 
and each ending in a pair of large, strongly curved, simple claws.' ' 

Speotrombicula trifurca (Ewing 1933). 

Trombicula trifurca Ewing 1933, Proc. U.S. Nat. Mus., 82, (29). 
Speotrombicula trifurca Ewing 1946, J. Parasitology, 32, (5), 437. 

Plate 86, fig. F-H. 

Ewing ? s description of this species is as follows: 

"Adult. Slender, with anterior pair of legs, mouth parts, and anterior 
part of cephalothorax darker and more sclerotized than the other parts. Mouth 
parts situated on a more or less distinct capitulum, which has parallel sides and 
is terminated below in a large trowel-shaped hypostome. Palpi slender, extend- 



Womersley— Asiatic-Pacific Scrub Typhus Mites 325 

ifcg to about the middle of first ijatcllae; palpal claw weak, slightly curved, 
about two-thirds as long as segment which bears it j palpal thumb very slightly 
swollen, reaching base of palpal daw; only one accessory spine present which 
is about one-half as broad as palpal claw. Chelicerae very long and slender, 
basal segment about four-fifths as long as capitulum; chela slender, with a 
sharp upper edge. Crista broad, plate-like with pseudostigmatic area reduced 
and near the posterior end; psendostigmata small, circular, and far apart; 
pseudostigmatic organs apparently absent. Eyes absent. Abdomen fully 
twice as long as broad and with the usual constriction at the level of third 
and fourth coxae; setae longest around posterior margin. Genital opening 
large; situated about half its diameter from fourth pair of eoxae; genital 
suckers six, first pair slightly larger than the others. Anal opening about half 
as broad and two-thirds as long as genital opening and situated about its 
greatest diameter from the latter. Coxae of first two pairs of legs united to a 
sternum and those of first pair each expanded along its anterior margin into 
a wshelf-like lamella. Anterior pair of legs much larger and longer than the 
others and each ending in a pair of trifurcate claws. Legs II and III rather 
weak, subequal; legs IV considerably stouter and longer; claws of legs II, III 
and IV simple but inflated towards the tips. Length 2*7 mm. ; width # 9 mm." 
Remarks. The species was described from four adults from Chilibrilla 
Caves, Panama, and Ewing suggests that its larvae will probably be found to 
parasitize bats. In his specific description he states that the pseudostigmatic 
organs are ct apparently absent", but in his generic description says "minute, 
setiform." The sternum according to his figure is entire, and no precoxal 
plates of leg I are evident. The epistome and epistomal (rostral) seta were 
not described until his generic revision of the species in 1946. In this paper 
he describes the epistome as * . * "a poorly sclerotized band-like comb." He 
also states .... "one rostral seta present." In his fig. 1, the outlines of the 
epistome are not delineated except the denticulate anterior margin which is 
straight. The rostral (epistomal) seta is shown as having only one short 
branch. 

Genus TROMBICUkA Berlese 1905. 

Acari nuovi, Manip. IV, 155, t. XV, fig. 4 in "Redia" 1, (2), 1905; idem, 
1912, "Redia", 8, (1), 94, €ig. 44 A-C. 

Genotype Trombkula minor Berl 1905. 
= Eutrombicula Ewing s. str. 1938, J, Washington Acad. Sci., 27, 288-295. 

With our increased knowledge oi the adult species of the Trombiculidae, 
and the correlation of adults and/or nymphs with their larvae it is possible to 



326 Records of the S.A. Museum 

define the genus Trombiwla in its non-parasitic stages with more precision as 
in the following diagnosis. 

" Adults and nymphs of a characteristic figure of 8 shape, with a pro- 
nounced median constriction, and the propodosonia usually narrower than the 
hysterosoma. Crista linear, rod-like, ending anteriorly in a hyaline, transverse 
or conical denticulate epistome with a single seta, and posteriorly with a more or 
less diamond-shaped sensillary area, with an internal areola from which arise 
a pair of filamentose sensillae. Eyes present, or absent, when present then 
one on each side and placed closely adjacent to the sensillary area. Palpal tibia 
with a single claw, and 3 accessory spines at the base of claw, with a single 
strong spine about midway between base of claw and articulation of palpal 
tarsus (in the nymphs with only 2 accessory spines at base of claw) ; rarely 
with a pectine of spines on palpal tibia. Tarsi I tapering with the greatest 
width (height) basad of middle. Sternum usually entire, no precoxal plates 
present (in aubgen. Leptotrombidmm precoxal plates present on leg I forming 
a longitudinally subdivided sternum). Genitalia with 3 pairs of discs in 
adult, 2 pairs in nymphs. " 

The specific characters in this genus are somewhat intangible, consisting 
mainly of the proportions in the length and height of the tarsi of leg I, and 
the length of the dorsal setae. 

The genus can, however, be separated into three subgenera, Trcnnbicula 
s. str. 1905, Leptotrombtdium Nagayo et at. 1916, and Neotrombictila Hirst 
1915, based on the presence or absence of eyes, and the presence or absence of 
precoxal plates on leg L as given in the following key. 



Key to the Subgenera of TBOMBICTJI*A Berl. 1905. 

1. Eyes present. No precoxal plates on leg I or these fused completely 
to form an undivided sternum. 

Subgen. Trombicuta Berl. s. str. 1905. 

Eyes absent . . . . . . . . . . . . . . . . . . 2 

2. Precoxal plates well developed on leg I and forming a longitudinally 
divided sternum. 

Subgen, Leptotrombidmm Nagayo et al, 1916 
(type Trombicida akamushi Brumpt 1910) . 

No precoxal plates on leg I or these completely fused to form an 



undivided sternum. 



Subgen. Neotrombimla Hirst 1915. 
(type Acarus auiwnnalis Shaw 1790). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 327 

Subgen. TROMBICTJI.A Berl s. str. 1906. 
Key to the Known Adult and/or Nyrnphal Species. 

1- Old World species . . , ., . . . * 2 

New World species , . . . . . . > . . . » . . 5 

2. Size small, approximately 680/t, or less 3 

Size larger, 1,000/* or smaller , ,. 4 

■3. Eyes very large, almost as wide as sensillary area. Dorsal setae uni- 
form in length, 20-25/4, tapering with moderately long setules. Tarsi 
I, 100/* long by 40/* high; metatarsi I, 90/* long, not higher than 
tarsi I. Trombicida (T.) minor Berlese 1905 (Adult) . 

Eyes smaller, not much wider than the basal cup or ring of the sen- 
sillae. Dorsal setae 14/x anteriorly to 20/* posteriorly with long strong 
setules. Tarsi 1 72-8/* long by 28-0/a high; metatarsi I 61 '6/1 long, 
and higher than tarsi I. Trombicula (T.) frittsi Wharton 1945. 

4. Dorsal setae to 40/*., tapering with moderately long setules decreasing 
towards the tip. Tarsi 12-4 times as long as high, and one and a 
half times as long as metatarsi. Trombicida (T.) samboni Womersley 1936 
Dorsal setae 25 to 55/* long, uniformly thick with setules equally long 

to tip. Tarsi I twice to two and a half times as long as high, and 
half as long again as metatarsi. 

Trombicula (T.) urichmanni (Ouds. 1905). 

5. Dorsal setae uniform in length, 36-40/1. Tarsi I 1-41 to 1-49 times 
as long as metatarsi; 1-76 times as long as high, 1-25 times as long as 
metatarsi. Trombicula (T.) batatas (Linn. 1758). 

Dorsal setae not uniform, increasing in length posteriorly . . . . 6 

6. Dorsal setae posteriorly to 110/i. Tarsi 12-8 times as long as high, 
1-25 times as long as metatarsi. Trombicula (T.) helleri Ouds. 1911. 

Dorsal setae shorter posteriorly . . . . . . . . , . . 4 7 

7. Dorsal setae posteriorly to 75/*, apically in lateral view with an indis- 
tinct spathulation showing dorsally as a small hyaline blob. Palpal 
tarsus with 9-11 nude apical setae. SB 55-70/t. 

Trombimda (T.) splendens Ewing 1913. 
= masoni Ewing 1943. 

Dorsal setae shorter posteriorly and without apical spathulation or 

blob . , 8 

8. Posterior dorsal setae to 4o/£ long. Ratio length to height of tarsi I 
«= 2-1-2-6 : 1*0 Trombicula (T.) van-ommereni Schierbeck 1937. 

Posterior dorsal setae longer . . . . . . . . . . . . 9 

S. Tarsi I only slightly more than twice as long as high, 2-05-2-11. 
SB 4S* 0-51-0/*. Palpal tarsus with 7-8 apical nude setae. Dorsal 
setae posteriorly to 56/*. Trombicida (T.) alfreddugesii (Ouds. 1911). 

Tarsi I rather more than 2| times as long as high, 2-59. SB 61-6/* 
apart. Palpal tarsus with 7-8 nude apical setae. Dorsal setae pos- 
teriorly to 56/* long. Trombicida (T,) jacoti nom. nov. 



323 Records of the S.A. Museum 

Troimbicula (Trombicula) minor Berlese 1905. 

Trombicula minor Berlese 1905, Acari nuovi, Manip. IV, 155, ft XV, fig. 4 
in "Bedia," 2, (2) ; Idem 1912, "Redia," 8, (1), 94; Ewing 1920, Ann, Ent. 
Soc. Amer., 13, 385-389 ; idem 1938, J. Washington Acad. Sci. 28, 292; 
idem 1944, US. Naval Med. Bull., 43, (4), 837-839- Uion Gunther 1939 7 
Proe. Linn, Soc. New South Wales, 64, 285-286; j non Womersley and 
Heaslip 1943, Tr. Roy. S. Austr., 67, (1), 92; Willmann 1941, Zool. 
Ana* 133, 131-136, Sig Thor and Willmann 1947, aDs Tierreich, Lfg. 
71b, 347. 

Plate 87, fig. A-C. 

Berlese described this species, the genotype, from two specimens from 
Tjompea, Java. In his original description he figures an entire dorsal view 
and a dorsal seta. Later 1912, in his monograph he added figures of the front 
tarsus and metatarsus, and of the palp, as well as an enlarged figure of the 
palpal tibia and tarsus. In the later work he stated that the types were in the 
Hamburg Museum, but that he had retained fragments of one specimen from 
which his later figures were derived. He further stated that he did not possess 
the base of the cephalothorax with the sensillary area and could not therefore 
confirm the presence or absence of eyes. Because Berlese 7 s figure of tlie palpal 
tibia shows only two accessory spines at the base of the claw, it has been gener- 
ally assumed by acarologists that his specimens were nymphs. In 1941, how- 
ever, Carl Willmann published his study of the types in the Hamburg Museum, 
and showed that the two specimens were adult females, having 3 accessory 
spines at the base of the palpal claw, and one specimen having an egg enclosed, 
and both having the 3 pairs of genital discs. 

The types were labelled "Tjompea, Java, 19, III, 1904, aus Hohlenguano 
gesiebt", this habit not having been mentioned by Berlese. 

Willmann gave a figure of the sensillary area and base of the crista. The 
sensillary area is shown as hexagonal with the sensillae bases placed on the 
lateral walls of the area. Around the sensillae bases is to be seen a clearly 
defined circular line, with a diameter of about half the width of the sensillary 
area. The inner third of this circular line is obscured by the area wall. Ewing 
1944, in discussing Willmann 's paper regards these circles as the outer rim of 
the very large sensillae bases, not as eyes. Willmann himself, however, appears 
to be uncertain as to whether they are eyes or not. To my mind they are 
definitely eyes, or at least the rims of the lobes of the sensillary area, sueh as in 
Berlese 's figure for mediocris, but with the corneas not shown, Ewing 's idea 
that they represent very large sensillary bases is not justified. The sensillae 



Womersley— Asiatic-Pacific Scrub Typhus Mites 



329 



bases in the Trombiculidae are a part of the sensillary area, and the position as 
shown by Willmann indicates displacement in the preparation. This also would 
account for the somewhat unusual position of the eyes partly beneath the wall 
of the sensillary area. The sensillae are only shown in their basal portion, but 
with distinctly short barbs. 

In general, the figure given by Willmann agrees with that of Berlese 1905, 
in which, under a lens, the large circles representing the eyes can be seen; it 
also is closely suggestive of that of mediocris and the two forms may ultimately 
be found to be the same. 

Ewing in discussing Willmann 's paper concluded as follows: 

"1. Trombicula minor Berl. is a very small species; the adults of which are 
about the same size as the nymphs of Trombicttia akamushi (Brumpt) and 
Trombicula deliensis Walch. 

2. The species is peculiar in that the pseud ostigmata are very large, the pal- 
pal claw T is but slightly tapering and the subterminal spines of the palpal 
tibia are almost setiform, long, set adjacent to each other and subparallel. 

3. Being found in cave giza.no, the inference is strong, that the larvae of 
T. minor have bats as hosts. 

4. Because of these facts it appears to the writer that Trombmda hirsti 
Sambon is a species distinct from T. minor Berlese." 

With these conclusions the present writer is generally in accord, except 
in considering that the so-called large sensillae bases of Ewing are really the 
eyes, as discussed above. 

By Willmann *s valuable re-examination of Berlese's types there seems 
reasonable ground, for the present at any rate, for regarding T> minor Berlese 
as different from T. unchmanni Ouds, (=medwcris Berl., ^hirsti Sambon). 
The most important differences are the smaller size, shorter and more uniform 
dorsal setae, size and dimensions of front tarsi, and possibly in the structure 
of the sensillary area and eyes. 

For the present it also follows that the true larvae of T. minor are un- 
known although several species of bat-infesting Troinbiculids are known from 
the Asiatic-Pacific area. However, until more material, both larvae and adults, 
are obtained from the type locality, or adults and larvae correlated by rearing 
none of these known larvae can be assumed to belong to T. minor Berlese, 

Be-descriptwn of Adult (after Willmann 1941). Colour whitish. Body 
elongate, strongly constricted medially. Crista long and straight (127/0, in the 
middle with a longitudinal furrow, widened behind to a 6-angled sensillary 



330 Records of the S.A, Museum 

area. The pseudostiginata lie directly on the outer edges of the areola. From 
them arises the long, stiff, finely ciliated sensillae. These are 107/1- long. The 
pseudostigmata are outwardly surrounded by a strongly developed smooth ring, 
almost two-thirds round, resembling a (rudimentary) eye; the sensillae bases lie 
eccentrically in the ring, and are half under the areola and half under the eye- 
like ring. Palpal tibia with strong claw and on inner edge with 3 thick, closely 
adjacent strong spines (not 2 spines as given by Berlese). Cuticle with shorty 
ciliated setae arising from small papillae. Leg I long and strong. Tarsus I 
100/<. long by 40/t high, metatarsus I 90/* long. Length of animal 680/*. 

Trombicula (Trombicula) frtttsi Wharton 1945. 
Trombicida fnttsi Wharton, 1945, J. Parasitology., 31, 4, 282-283 (larvae). 

Plate 87, fig. D-G. 

Description of Nymph. Of typical Trombicida facies, but rather elongate. 
Length (unfed) to 325/*; width across propodosoma 150/*, across hysterosoma 
162p. Crista linear with subposterior sensillary area, length 75 -6/*, SB 25 -2/*, 
sensillae 70/* long, slightly thickening soon after the base and basally finely 
barbed but in distal three-fourths with long outstanding setules. Epistome 
dentate, with one ciliated seta 22/* long. Chelieerae finely serrate on inner 
(dorsal) edge. Palpi rather elongate, with two accessory spines at base of 
claw. Eyes present but difficult to see, one on each side and closely adjacent 
to sensillary area. Dorsal and ventral setae rather sparse, dorsally anteriorly 
14/* lengthening to 20/* posteriorly, pointed with outstanding setules. Legs : I as 
long as body, thicker and stronger than rest, to 325/* long, II 195/*, III 182/* r 
IV 225/* j tarsi I tapering and slightly shorter than or equal to metatarsi, which 
is wider, 72-8/* long by 28-0 high; metatarsi I 64-4/* long by 30-8/* high. 
Genital organ with paired discs. 

Loe. Described from the type nymph and 10 paratypes reared by 
K, L. Coekings from engorged larvae from scorpions, Iletervmetrus longimwnvs 
from Kukit Lagong Forest Reserve, Kepong, Kuala Lumpur, F.M.S. The 
larvae were all collected on 21 Sept., 1949, and the nymphs emerged and were 
mounted on the 10th and 14th Oct., 1949. 

Remarks. The presence of distinct eyes, although these are not easily 
seen, places this species in the genus Trombicula s. str. Although as yet only 
known from the nymph in its small size, it is closely related to Trombicula 
minor Berl., and may even be the same. From Willmann's description and 
figures 1940, it appears to be distinct from Berlese 's species, at least in the much 
smaller sLze of the eyes, and is thus separated in the key to the known species 
of nymphs and adults given here. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 331 

Trombicula (Trombicula) batatas (Linn. 1758). 

Acarus batatas Linnaeus 1758, Gyst. Nat., 10th Ed. Genus 235, Species 22. 

Eutrombicula batatas, Michener 1945, Ann. Rept. Gorgas Labor., U.S. Gov. 
House Document, No. 896, p. 22 j 1946, idem, U.S. Gov. House Document 
No. 12, p. 13 ,• 1946, Ann. Entom. Soe. America, 39, (1), 101-118,* idem 
1946, ibid. 415. 

For further synonymy see Larval Section. 

Plate 87, fig. H-K. 
This species has been reared through to all stages, and thoroughly described 
and figured by Michener 1946. I am indebted to him for a number of speci- 
mens of nymphs. The following description is summarized from Michener 's 
studies : 

Adult. Of typical Trombicula facies with hysterosoma wider than propo- 
dosoma. Colour in life red. Size 9 770 to 990^; 3 660 to 830^t long. Crista 
slender ; sensillary area broader than long with a transverse curved ridge in 
front of sensillae ; sensillae filamentous, as long as crista, with a few short 
branches dmtally. Palpi slender; tibia with 3 accessory spines at base of 
claw and a fourth near middle of segment. Eyes one on each side, and 
adjacent to sensillary area. Legs shorter than body, I the longest; tarsi I 
1-41 to 1*49 times as long as metatarsi. Dorsal setae short, 36-40/^ long, with 
short citations. 

Nymph. As in adult, but smaller and with the usual nymphal differences 
in accessory spines at base of claw being only two and in having only 2 pairs 
instead of 3 pairs of genital discs. Length 500 to 550/a. 

Remarks. From the several specimens of nymphs of this species received 
from Dr. Michener the following data was obtained i 

Crista 162/a long; sensillae 154/1, with bases 39/a apart. EpLstome and 
epistomal setae not observed. Tarsi I 148/x long by 84/* high, metatarsi 118/a. 
Dorsal setae to 30/a long. 

Bionomics. The larvae of this species is common on domestic fowls and 
many species of native birds in ihe Panama region as listed by Michener 1946. 

Trombicula (Trombicula) helleri (Ouds. 1911). 

Microtrombidmm helleri Oudemans 1911, Ent. Ber., 30, 120; idem 1912, Zool. 
Jahrbr., Suppl., 14. 

Eutrombicula helleri, Ewing 1938, J. Wash. Acad. Sci., 28, 294 (larvae) ; 
Michener 1946, Ann. Entom. Soc. Amer., 39, 411 (adult and nymph). 
For other synonymy see Larval Section. 



332 Records of the S,A. Museum 

Plate 87, fig. L-N. 

Michener 1946 has described the adult of this species from a single female 
from which larvae identified as Oudemans' species were obtained. The speci- 
men was from leaf mould, at Santa Rosa, Colon Province, Panama, Sept. 10, 
1945. The essential points of Michener 's description are: 

Of typical Trombieula fades. Colour red. Length to 1-3 mm. Palpal 
tibia with 3 accessory spines at base of claw (4 on one side) arising from a 
small elevation on inner side. Crista slender, 190/n long with broad subposterior 
sensillary area which is 47/* wide (measured to lateral margins). Eyes one on 
each side, adjacent postero-laterally to sensillary area. Legs shorter than 
body; I 900/a long; tarsi I 1*25 times as long as metatarsi and 2-8 times as 
long as high. Dorsal setae, ciliated, twice as long posteriorly as on shoulders, 
posteriorly to 110/x slender. Genitalia with 3 pairs of discs. 

Trombicula (Trombicula) vanommebeni Schierbeck 1937. 

Tromlicula vanommerem Schierbeck 1937, Ann. Parasit. Hum. Comp f , 15, 326. 
Eutrombicula vanommereni, Michener 1946, Ann. Entom, Soc. Araer., 39, 413. 
For further synonymy see Larval Section. 

The nymphs and adults of this species are only known from specimens 
reared from larvae on lizards (Ameiva prao&ignis) from Panama, and described 
by Michener 1946. For other references to hosts of the larvae see Larval 
Section. 

The following details are from Michener 7 s paper: 

Adult. Facies as in T. (2?.) batatas. Colour red. Length 900/x to 1,100/*. 
Crista slender, 150 to 170/a long with broad subposterior sensillary area, 59 to 
75/i wide (as between lateral margins). Sensillae about as long as crista with 
a few distal branches. Eyes large, half width of sensillary area. Palpal 
tibia with 3 accessory spines at base of claw arising from a slight protuberance, 
and a fourth at middle of segment. Legs all shorter than body; 1 730/*; tarsi 1 
1-36 to 1-41 times as long as metatarsi and 2-1 to 2*6 times as long as high. 
Dorsal setae; those on shoulders about two-thirds as long as posteriorly, pos- 
teriorly 45 to 47/x long. Genitalia with 3 pairs of discs. 

Nymph. As in adult but smaller and with nymphal characteristics. 
Tarsi I 1*7 to 2-2 times as long as high, and 1-39 to 1*53 times as long as 
metatarsi. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 333 

Trombicitla (Trombicitla) alfreddugesii (Qudeinans 1910). 

Microtrambidm'm alfreddugesii Ouds. 1910, Ent. Ber. Nederl. Ver., 3, 84 
( larvae). (For further synonymy of larvae see Larval Section.) 

Trmibi-cula cinnabarinvs Ewing 1920, Ann. Ent, Soc. Amer.. 13, 387, fig. 3. 

Trombicula irrtiam, Ewing 1925, Proe. Biol. Soc. Washington, 38, 17-20; 
idem 1931, Proc. U.S.N. Mus., 80, (8), 8. 

EutrambicuJa alfreddiigesii, Ewing 1944, J. Parasitology, 30, (6), 341. 

Trombicula thalsahuate, Sig Thor and Willmann, 1947, Das Tierreich, Lfg, 
71b, 345. 

Plate 88, fig. A-F. 

This species has been fairly well described and figured by Ewing under 
the name c&nnabarin-us and is very widely distributed in the United States of 
America (Ewing 1944). Swing's description is as follows: 

"Adult, Colour in live adults vermilion, or cinnabar. Cephalothorax 
triangular and with the crista extending from the anterior end of the latter 
as a straight ridge to about the middle of the triangular pseudosigmatic area. 
Posterior lobes of pseudostigmatie area evenly rounded and near the median 
line. Pseudostigmatie organs arising from pseudostigmata and provided with 
a few minute barbs on the posterior margins near their tips. Eyes well 
developed, very near the pseudostigmata and with thick and strongly curved 
corneas- Palpi armed with three tibial spurs; the longest of which is less than 
half as long as palpal claw; thumb of palpus not swollen and not surpassing 
the palpal claw. Chelicerae with lower chela sharp and provided with about 
two dozen backwardry directed teeth; upper chela represented by a chitinous 
tubercle-like knob. Abdomen clothed with the usual setae, which are but slightly, 
if at all, longer on the posterior border than on the shoulders, and have the 
barbs smaller at the somewhat tapering tip. Legs about as usual, the first 
pair being much the largest and longest. The tarsi of this pair are one and 
forty-seven hundredths times as long as the tibia. Posterior legs reaching to 
the tip of abdomen. Length of medium sized specimens, 0-92 mm.; width 
0-52 mm." 

In his 1944 paper, Ewing also figured and described the genital aperture 
and penis of the male. 

Of this species I have been able to study the following specimens : 

Two adults reared from larvae from Neotoma micropus Baird nests from 
Laguna Madre, Texas, Sept. 21st, 1945 (D. E. Hary and V. L. Wooley) and 



334 Records of the S.A. Museum 

kindly presented to me by Dr. G. H. Wharton. Also a number of specimens 
from amongst a collection of Trombidiids and Erythraeids sent to me by 
Dr. F. Bonet from Mexico. This last lot of material comprised nymphs and 
adults of both sexes. Prom this material the following data can be added to 
the description, and the figures accompanying this paper have been drawn. 

Length of adults to 1,170/*; width across propodosoma 585/x, across hystero- 
soma 780/4. Crista (excluding epistome) to 195//. long. Epistomal seta 39/t 
long, and margin of epistome denticulate. Sensillae distally with a few branches, 
to 100/* long, and with the bases 58/i apart. Legs I to 845/x, II 520/i, III 455//., 
IV 700/* ; tarsi I 208/* long by 104/t high, metatarsi 156/a long. Dorsal setae to 
45/x long, fairly thick and not much tapering, with numerous strong setules. 
In the male sex there are 4 apically spathulate and fimbriated li titillating" 
setae (not shown or observed by Ewing) on each side of the genital aperture 
and placed on a lobe and about midway between the second and third genital 
discs. 

Trombicttla (Tsombicula) splbndens Ewing 1913. 

Tromhiwda splendem Ewing 1913, Bull. Amer. Mus., 32, 13-114, pi. 7, fig. 5 ; 
idem 1920, Ann. Ent. Soc. Amer., 13, 306, 389; idem 1926, Ent. News, 37, 
111,- idem 1932, Proc U.S. Nat. Mus., 80, 10; Sig Thor and Willmann 
1947, Das Tierreich, Lfg.71b, 349; Jenkins 1949, J. Parasitology, 35, 
201-4. 

AcaHsciis maswri Ewing 1943, Proc. Ent. Soc. "Wash., 45, 60. 

Eutrombicida, masoni (Ewing) Jenkins 1947, Ann. Ent. Soc. Amer., 40, 56-68. 

Plate 88, fig. G-K. 

Jenkins has recently discussed this species and established the synonymy 
of reared adults of masoni with splendens of Ewing. 

A re-description of the adult, from 2 males and 2 females from culture 
No. 123, Duke University, N.C., 22 Dec. 1946 and kindly given to me by 
Dr. Q. W. Wharton is as follows: 

Adult. Colour in life red. Of typical Trambicitla- faeies. Length to 
1,500ft; width across propodosoma 850/x, across hysterosoma 990/*. Crista 
elongate, to 182/* long, with diamond-shaped sensillary area, and paired 
filamentous sensillae to 180/t long and finely ciliated or nude distally 
with their bases from 58-8 to 67-6//. apart (Jenkins gives 55-70/a) ; 
epistome not clearly visible but apparently rectangular with dentate margin; 
epistomal setae to 42/*. Eyes 1 — )- 1. closely adjacent to sensillary area. Cheli- 
eerae finely serrate on inner (dorsal) edge. Palpi stout; tibia with claw and 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 335 

3 stout accessory spines at the base of claw; ventrally the tibia with a long 
nude spine-like seta; palpal tarsus with 9-11 nude apical setae. Dorsal setae 
fairly long, increasing from 30/* anteriorly to 75/* posteriorly, slender, strongly 
and long ciliated and at tip with a small and indistinct spathulatxon in lateral 
view and a hyaline blob in dorsal view. Legs I longer and stronger than the 
rest, to 99<V, II 640^, III 710/*, IV 895/* ; tarsi I from 214/* long by 117/* high, 
to 240/* by 136/* long (mean 230/* by 128/*, ratio 1-8: 1-0) - metatarsi 1 
149/* to 175/* long (mean 167/*, ratio length tarsi I to metatarsi I 1*37 :1'0). 
Coxae I without precoxal plates. Male as in female, except for the genitalia but 
precise details were not sufficiently clear. 

Trombicula (Trombicula) jacoti nom. nov. 

Trambiculoides scaber Jacot 1938, Psyche. 45, 121-132, non Trombidiu/in 
scabrum Say, 1821, J. Acad. Nat. Sci. Philadelphia. 

Plate 89, fig. A-D. 

Ewing 1946, Proc. Biol. Soc. Washington, 59, 69, has discussed the status 
of Trambi-culoides Jacot 1938, and clearly shown that the species which Jacot 
referred to Say's scabrum is not this species as recognized by other writers. 

Howard (1918) reared the larvae and figured the species regarded as 
Say's scabrum, placing it in S&ricothrombium Berlese. From Howard's figures 
and from Say's statement (1) that "the eyes are pedunculate and white" and 
(2) that "the body is ovate, broadest and very obtusely rounded before and with 
hardly perceptible hairs" it is certain that Jacot's specimens are not the same 
as Say's, a conclusion in which I am in full agreement with Ewing. 

Jacot gives "from under the bark of a large prone oak, near Savannah, 
Georgia," as the locality for his specimens, and because some of the other 
species (but not scabrum) recorded by Say were from Georgia concludes that 
this was the type locality for scabrum. As pointed out by Ewing, however, 
Say merely states "in forests, on trees, etc. not uncommon" with no definite 
locality. 

As Jacot's identification of his specimens with scabrum is therefore incor- 
rect, his generic name Trombicuboides becomes a synonym of Sericothrombwm 
Berlese. The specimens of Jacot are typical TrombicuLa as recognized in the 
present work, and a new specific name being required, that of jacoti is pro- 
posed. 

From Dr. J. Bequaert, of the Museum of Comparative Zoology, Cambridge, 
Mass., I am very much indebted for the loan of Jacot's topotype material, and 
the following re-description is drawn up from the five specimens: 



336 Records of the S.A. Museum 

Adult. Colour (in life red). Of typical Trombicula facies. Length to 
1,300/*. Width across propodosonia to 1,040/*, across hysterosoma to 780/*. 
Crista linear, 195/* long, with subposterior diamond-shaped sensillary area with 
distally ciliated filamentous sensillae to ca. 112/* long, and their bases 61 '6/* 
apart; epistome apparently rounded-eonieal with dentate margin and epistomal 
seta to 56/* long. Eyes 1 + 1, closely adjacent to sensillary area. Chelicerae 
with finely serrate inner (dorsal) edge. Palpi stout, tibia with claw and 3 
strong accessory spines at base of claw, on outer side with a long nude spine- 
like seta; tip of tarsus with ca. 7-8 nude setae. Dorsal setae tapering with 
fine ciliations, on propodosoma to 28/* long, posteriorly on hysterosoma to 56/* 
long. Legs, I longer and stronger than the rest to 975/*, II 520/*, III ?, 
IV 715/*; tarsi I 240/* by 92-6/* high (mean of 4 spec., ratio length to height 
*»2'59:1*0), metatarsi I 169/* long (ratio length tarsi I to metatarsi I ■= 
1-43:1-0). 

Male unknown. 

Remarks. In the proportions of the front tarsi and metatarsi this species 
is nearer to alfred&ugem than to splendem, but seems to be distinct therefrom 
as in the key* 

Tkombicula (Trombicula) wighmanni (Oudemansl905). 

Tronibidkim urichmanni Oudemans 1905, Ent. Ber., 7, (22), 217; idem 1906, 
Nova Guinea, 5, 106, 132. 

Trombicula mediotria Berlese 1912, Eedia, 8, (1), 95; Hatori 1919, Ann. Trop. 
Med. & Parasit, 13, 233; Kawamura and Yamaguchi 1921, Kitasato 
Archiv, Exp. Med., 2, 169; Sig Thor and Wilhnann 1947, Das Tierreich, 
Lf g. 71b, 348. 

Trombicida pseudoakamushi Hatori 1919, Ann. Trop. Med. & Parasit., 13, 233 
{non Tanaka et al 1930, Zentrbt. Bakt. Orig., 116, 335) ; Waloh 1923, 
Kitasato Archiv. Exper. Med., 5, 63; Hirst 1929, Ann. Mag. Nat. Hist., 
(10), 3, 564. 

Trombicula pseudoakamushi var. deliensis Walch 1924, Trans, Vth, Bien. 
Congr. Far East. Assoc. Trop. Med,, 691; idem 1924, Geneesk. Tijds. Ned. 
Indie, 64, (3) ; idem 1925, Kitasato Archiv, Exper. Med., 6, 235. 

Trombicula hirsti Samhon 1927, Ann. Mag. Nat. Hist,, (9), 20, 157. 

TronibicuDa liirsti var. buloloensis Gunther 1939, Proc. Linn. Soc. New South 
Wales, 64, 78; idem 1939, ibid. 466. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 337 

Trombicula minor, Gunther 1939, Proc. Linn. Soc. New South Wales, 64, (5-6), 
466 (non Berlese 1912, Bedia, 8, (1), 93) ; Womersley 1939, Tr. Boy. Soc. 
S. Austr., 63, 149 ; Womersley and Heaslip 1943, Tr. Boy. Soc. S. Austr., 
67, (1), 92 (non Berlese 1912) ; Gunther 1940, Proe. Linn. Soc. New South 
Wales, 65, (5-6), 477-478 {non Berlese 1912). 

Trombicula minor var. delieims, Womersley and Heaslip 1943, Tr. Boy. Soc. 
S. Austr., 67, (1),94. 

For further synonymy see Larval Section. 

Plate 89, fig. E-L 

Oudemans originally described the larvae of wichmanni from Goura coro- 
naia from New Guinea and from man from the North Celebes. In 1919 Hatori 
referred to but did not describe a larval Trombicula from Formosa under the 
name of " ps&udoakamushi (non Tanaka)" and claimed to have obtained larvae 
from captured adults which he considered identical with Trombicula medwcris 
Berlese from Java. 

In 1921 Kawamura and Yamaguchi, working also on Formosan material, 
described and figured in detail what they regarded as Hatori 's species and 
affirmed his opinion that the adults were mediocrk. 

In 1917, Miyajima and Okumura (Saikin Zasshi, 266; 893-908, in Japan- 
ese) described and figured the adult of a species of Trombicula which they 
doubtfully referred to mediocris. At the same time they described a "thin- 
haired" type of larvae, which they compared with the European autumnalis 
pointing out some of the differences therefrom. From the very excellent 
figures both of the larvae and adult, it is reasonably certain that the larvae 
can be referred to uriokmanni Ouds. and the adult to mediocris Berlese. 

Walch 1923 recorded larvae from Deli, Sumatra, as pscudoaka/imishi Hatori r 
and in 1924a, and 1924b, and 1925 referred it to a variety cMiensis of this 
species, and figured the crista and eyes, palpi and dorsal setae. 

In 1929, Hirst suggested the possibility that the larvae, hirsti Sambon 
1927 from Queensland, might be conspecine with Oudemans wichm&nni. Carl 
Gunther 1939 described larvae from New Guinea as T. hirsti var. btdoloensis, 
and later in the same year, as a result of having reared nymphs, considered 
hirsti as a synonym of minor Berlese. This was followed by Womersley 1939 
and Womersley and Heaslip 1943. In 1943 Womersley and Heaslip regarded 
the variety deUensis Walch as belonging to minor Berlese. For the Formosan 
pse-udoakamushi Hatori, Womersley and Heaslip after showing that the name 
was invalid proposed the name hatorn and further showed that it was closely 



338 Records of the S.A. Museum 

related to wichmanm, only differing: in the higher values of AM, AL, PL and 
Sens, and in the larger ratio of PW/SD. 

Towards the end of 1944 I received from Major R. N. MeCulloch, four 
adult Tromiicuia collected in soil, New Guinea, from which he had successfully 
reared a number of larvae which were identified at the time as T. hatorii Worn, 
and Heasp, These adults, on examination were found to agree entirely with 
the figures and data given by Kawamura and Yamaguehi for their Formosan 
mediocris as well as with Berlese 's figures and description* except that they 
were red in life, not white as given by Berlese. This had been noted by Kawa- 
mura and his colleagues, and it was suggested that Berlese \s specimens had been 
bleached by the spirit. 

The above New Guinea specimens also agree very well with the figures of 
the nymphs of pseud>oakamushi var. dcliensis given by Waleh. 

There seems little doubt therefore, that wkhmmni Ouds. is the name by 
which this species should be known, as given in the above synonymy. Whether 
it is to be further synonymized with minor Berlese 1904, as has been done by 
Gunther 1939, 1940, and by Womersley 1939, and Woraersley and Heaslip 1943, 
is somewhat doubtful, for following the re-deseription of the types of minor 
by Willmann 1941 (Zool. Anz., 133, 131-136) this species appears to be differ- 
ent (see under Trombiwda minor) . These differences, however, are rather 
small and quite possibly it may be later established that wichmanni Ouds. 1905 
is synonymous with minor Berlese 1904. Dr. H. S. Fuller has expressed to me 
in a letter his opinion that pseudoatkamnski is the same as urichma>nni, and I 
have seen a preparation in the Oudemans' collection at Leiden given to Oude- 
mans by Waleh of 3 nymphs labelled pseudoakammM* bred from larvae by 
Walch. These not only agree w r oll with Waleh ? s figures, but also in the crista 
with Willmann 5 s 1941 figure for minor Berlese. 

In describing his methods of obtaining larvae from adults captured in the 
field Major MeCulloch (m lit.) states: 

"The adult specimens, later determined as mediocris, were collected at Lae, 
on April 9th and 10th, 1944 In searching for them and keeping them (and 
also the larvae) in the laboratory, I followed the advice of Maj. G. M. Kohls. 
In jungle where larval mites were relatively numerous litter was scraped from 
the soil and adults watched for on the surface, then in the top inch of soil 
and finally in some patches by digging and crumbling the soil down to about 
six inches. In 2 days about 15 adult red Trombieulids were found on or close 
to the surface. There were also present at various depths more numerous and 
larger mites, probably adult Erythraeidae, etc. 

•'The adult Trombieulids w r ere tubed individually each with about fl tea- 



Womersley — Asiatic-Pacific Scrub Typhus Mites 339 

spoonful of moist soil, previously sterilized by steaming, from the locality. The 
teat tubes were closed by corks, perforated to allow aeration, covered with 
•calico and inverted into the tubes to give a tight fit so that the larvae could 
not force their way into any space between the edge of the cork and the glass. 
The specimens were kept in the laboratory where the temperature varied 
between 74° P. in the early morning and 85-88° F. in the afternoon. A few 
drops of water -were added to the soil in each tube from time to time, and a 
little condensed moisture was always present on the inner wall of the tubes. 
In this manner larvae of Schongastia blest oivei were kept alive for as long as 
46 days. 

''The living adult Trombiculids were examined from time to time as other 
work allowed. On May 15th it was observed that one had produced more than 
a dozen larvae, apparently sometime earlier. No eggs were found. Some of the 
other adults had died, and the remainder had then to be killed. The specimens 
of larvae were then forwarded to Adelaide for identification*" 

As previously stated the above larvae were identified as T. hatorii Worn, 
and Ileasp. 1943 (== wickmamii Onds. 1905) and the adults as mediocris BerL 
1912. 

The following description of the adult is based on the above New Guinea 
specimens; that of the nymph is from those reared from larvae- b} r Kawamura 
and Yaniaguehi, who gave little detail, and by Walch ; both from T. pseudoa- 
1mm ushi H at or i . 

Adult. Colour in life red. Of typical Trombicula facies, w'ith constriction 
between second and third pairs of legs. Length 1,100/j. (Berlese 1.550/*,; Miya- 
jima and Oknmura 1,450/x) ; width across propodosonia 525/1, across hysterosoma 
675/z, (Berlese max, 900/*; IMIyajima and Okumura max. 830/t) T Crista 197/x 
long, linear, with subposterior more or less triangular broad sensillary area, 
with paired filamentous sensillae, which are closely and shortly barbed basally, 
and with more open longer ciliations distally; (other students make no refer- 
ence to the sensillae having barbs on the basal portion) ; sensillae bases 62/x. 
apart. Epistome rounded, finely denticulate with 1 ciliated seta to 60/* long. 
(Miyajima and Okumura for their specimens from Formosa figure the epistome 
as truncate anteriorly, with no lateral denticulations. This feature, however, 
is extremely hyaline and difficult to make out). Chelieerae with inner (dorsal) 
concave margin finely denticulate. Palpi fairly stout; tibia with simple claw, 
and 3 accessory spines at base thereof, and a fourth about midway between 
base of claw and articulation of palpal tarsus ; tarsus widest medially and 
slightly over-reaching tip of claw. Eyes Fairly large with distinct corneas, one 
on each side, close to sensillary area and posterolateral of sensillae bases. 



340 Records of the S-A. Museum 

Legs all shorter than body, I the longest and strongest 945/* long, II 525/x, 
111 525/*, IV 600/x-, tarsus I 208/jl long by 100/t high, metatarsus I 135/i long 
(Berlese gives tarsus I 210/* by 80/*, metatarsus 140/*; Miyajima and Okuniura 
figure tarsus rather more than two and half times as long as wide). Sternum 
entire, about as wide as long; no precoxal plates on leg I. Dorsal setae from 25/x 
anteriorly to 55/* posteriorly (Berlese gives 25/* to 70/aj Miyajima and Okumura 
give 25-50/*), setae uniformly thick with moderately long ciliations. Genital 
aperture with 3 pairs of discs. In the male, with four spathulate. fimbriated 
"titillating" setae as described and figured for alfreddugesii Ouds. 

Nymph. As in adult but smaller and with the usual nymphal character- 
istics as only 2 accessory spines at base of palpal claw, and only 2 pairs of 
genital discs. Crista lOfyi long, SB 40/*. Sensillae ca. 90/i plus. Tarsi I 116/* 
long by 60/x high ; metatarsi I 79/*. Dorsal setae as in adult 25/i to 45/* long. 

Material seen, Three adults from Lae, New Guinea (R. N. McCulloch), 
one adult from Innisfail, Queensland, previously recorded as minor Womersley 
and Heaslip 1943 (W.G.H.) ; three adults from New Guinea, Dobodura area, 
April, 1944 (G. M. Kohls) ,- one adult from damp soil, Goodenough Island, 
August, 1944 } one adult from soil, Admiralty Island. Aug., 1944 (NAMRU II 
132,G,W.W.). 

Two nymphs reared by Dr. C. Gunther, in New Guinea, Bulolo area, 
1939, and one reared by Maj. Kohls, Dobodura area, New Guinea, 1944 (No. 19, 
G.M.K.). 

Remarks. With regard to the habitat in which the adults were found 
Maj. McCulloeb writes me as follows: "That they were all mediocris is most 
interesting in view of the fact that they came from ground where the over- 
whelmingly abundant larvae were Schongastia blestowei. Because of its rela- 
tively high mite infection this ground was used for weeks as a test area, and 
at least 50 larvae were collected on boots and identified daily. About 99% were 
S. Westowci, the rest either 8. parva Worn. 1944 (= sclmffneri Walch) or 
T. haiori (previously identified by me as wickmanni) but referred by Womers- 
ley to hatorii." 

Trombicula (Trombicula) samboni Womersley 1939. 

TrombicuU samb&ni Womersley 1939, Tr. Roy. Soc. S, Austr., 63, (2), 153. 
TromHruU Mrsti, Ilirst 1929, Ann. Mag. Nat. Hist., (10), 3, 564, wonSambon, 

1927, ibid. (9), 20, 157; Womersley 1934, Rec. S. Aust. Mus., 5, (2) 212, 

nee Sambon, 1927. All as larvae. 
Trombicula samboni, Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 

67, (1),95, (adult). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 341 

Plate 89, fig. J-O. 

Adult. Colour in life red. Of typical Trombicula facies with constriction 
between second and third pair of legs. Length to 1,200/*, width across propo- 
dosoma 690/*, across hysterosoma 860/*. Crista well developed, 170/* long, rod- 
like with wide, roughly diamond-shaped subposterior sensillary area, with 
paired sensillae 118/* long and shortly ciliated on distal half, their bases 47/a 
apart. Eyes one on each side, large and closely adjacent to sensillaiy area, 
deeply pigmented in life, but rather difficult to see in preparations. Epistorae 
rounded-conical, finely denticulate, with 1 setae. Chelicerae with the inner 
(dorsal) edge finely serrate. Palpi moderately slender; tibia with 3 accessory 
spines at base of claw. Legs all shorter than body ; 1 the longest and strongest, 
775/* long. II 520/x, III 600/1, IV 680/*; tarsus I 206/a long by 86/x wide, metatarsus 
I 137/* long. Sternum entire, about as wide as long; no precoxal plates on 
leg I. Clothing of dense, ciliated, pointed setae, to 40/t long posteriorly, the 
ciliations long and outstanding. Genital opening with 3 pairs of discs. 

Nymph. Similar to adult in all details except size, with only 2 pairs of 
genital discs and 2 accessory spines at base of palpal tibial claw. Length to 
1,000ft; width across propodosoraa 600/i, across hysterosorna 720/*. 

Remarks. This species is only known from South Australia, where since 
the extermination of the native fauna it has established itself as larvae on the 
introduced rabbit. 

Subgenus I*EPTOTROMBnHtTM Nagayo et al. 1917. 

J. Exper. Med., 25, (2), 255, 1917. 

Genotype Trombidiunt akanm&M Brumpt 1910. 

Nagayo, Miyajima, Mitamura and Iinaroura, 1917 proposed this generic 
name for the Japanese, species akamuslii Brumpt, on various differences from 
"other trombidia". In 1921, however (Amer. J. Hygiene, 1, (5, 6), 588), 
they adopted Berlese's name of Tromlmida. 

In 1946 (Proc. Entom. Soc. Washington, 48, 171-178) Wharton pointed 
out that the proved vectors of tsutsugamushi disease belonged to a small group 
of larval species, which he defined as the "akamushi" group of the genus 
Trambwula. In the present studies it is further shown that, where the larvae 
of this group of species have been reared to and correlated w T ith adults or 
nymphs, they are also distinct generically from other Trombicula species. Thus 
a new generic name is required, and that proposed by Nagayo et al. 1917, for 
akamushi is available. The diagnosis of the genus is as follows: 

Adult and Nymph. Facies as in Trombicula with marked constriction 
between propodosoma and hysterosorna. Crista elongate with roughly diamond- 



342 Records of the S.A. Museum 

shaped sensillary area, with internal areola and a pah- of filamentous sensiilae. 
Epistoine rectangular with the anterior margin slightly concave or sinuous, 
with fine denticulations on the shoulders and with 1 ciliated seta. Eyes absent. 
Tarsi of leg I tapering, widest basad of middle. Coxae of leg I with precoxal 
plates, which form a longitudinally divided sternum. 

Key to the Known Adults and Nymphs. 

1. Dorsal setae ending in a hyaline knob, or small spathulation . . . . 2 
Dorsal setae not so . . . . i r , 6 

2. Dorsal setae posteriorly terminating in a hyaline knob . . . . 3 
Dorsal setae posteriorly terminating in a small spathulation . . . > 4 

3. Dorsal setae shorter, more uniform from 30yx to 45/a long. Tarsi 
hardly twice as long as high; half as long again as metatarsi. 

Trombicula (L.) delienm (Walch 1922). 
Dorsal setae longer, from 30/x anteriorly to 90^ posteriorly. Tarsi I 
more than twice as long as high ; half as long again as metatarsi. 

Trombicula (L.) palUda (Nagayo et al. 1917). 

4. Posterior dorsal setae to 60-80/a long 4 . 5 

Posterior dorsal setae to 56 fi long. Metatarsi I about as long as tarsi 

are high. Trombicula (L.) burmemis Ewing 1945. 

5. Metatarsi I longer than tarsi are high. 

Trombicula (L.)/w7imEwing 1945. 

Metatarsi I about as long as tarsi are high, 

Trombicula (L.) lon-giseta sp. n. 

6. Dorsal setae posteriorly to llOju; ending in a strong sigma-like 
spinule. Tarsi T more than twice as long as high ; half as long again 

as metatarsi. Trombicula (L.) myzantha sp. n. 

Dorsal setae shorter, posteriorly to 70/x, uniformly thick, and not end- 
ing in such a spinule. Tarsi 1 two and a half times as long as high; 
half as long again as metatarsi. 

Trombicula (i.) akamushi (Brumpt, 1910). 
(==* fletcheri Worn, and Heasp. 1943). 

TftoaraiciTiiA (Leptotrombidium) deliensis (Walch 1922). 

Trombicula deliensis Walch 1922, Kitasato Arehiv. Exper. Med., 5., (3) 4 
(larva) ; for other larval synonymy see Larval Section. 

Tr&mbicida deliensis, Walch 1922, Geneesk. Tijds. Neder. Indie, 64, (3), 514; 
idem 1924, Tr* Vth. Bien. Congr. Far East. Assoc. Trop. Med., Singapore, 
.583; Radford 1946, Parasitol., 37, (102), 42-45; idem 1946, ibid. 46-54; 
Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 98; Sig Thor 
and Willmann 1947, Das Tierreich, Lfg. 71b. 



WOMKRSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 343 

Plate 90, fig. A-G. 

The nymph of this species was first reared from larvae and described by 
Waleh from Deli, Sumatra, when the entire animal, crista, ehelieera, palp and 
dorsal seta as well as the sternum were figured. The crista was shown as having 
a somewhat dumb-hell shaped sensillary area with the posterolateral corners 
angular : the anterior rod was only indicated by a fine line, the onter borders 
not being: shown; no epistome was indicated, but a single ciliated epistomal seta 
was figured. The sensillae were shown as shortly barbed on the basal part, 
with longer filiations distally The dorsal setae were figured with almost even 
thickness and ciliations, with at the apes a hyaline knob or blob. The sternum 
was shown as being divided longitudinally. 

Eadford (1946) also reared nymphs from larvae in Ceylon, and later was 
able to correlate them with adults found in the field* In his first paper Ead- 
ford gives fairly good figures of this species, and of certain details. The crista 
is shown more clearly and the posterior triangular projection of the sensillary 
area is indicated. 

Dr. Radford has very kindly presented me with a number of adults as 
well as two nymphs of this species from which the following re-description and 
figures are drawn .- 

Adwtt. Length 800/* to 1,030/* excluding capitulum (Radford gives 1,200/* 
including capitulum and 900/* without) ; width across propodosoma 300/* to 470/* 
(Radford 600/*), across hysterosoma 400/* to 660** (Radford 675/*). Facies as 
in Trombicidu with pronounced median constriction. Colour in life red. Crista 
linear, 151-160/* long (Radford 144/*), with diamond-like subposterior sensillary 
area, and paired filamentous sensillae,, distally ciliated, the sensillae bases 61/* to 
66/* apart (Radford 68/*) , Epistome transverse rectangular, with denticulations 
anteriorly and 1 ciliated seta. Palpal tibia with 3 accessory spines at base of 
claw. Chelieerae finely serrate on inner edge. Legs: I 660/* to 820/* long 
(Radford 675/0, II 440/* to 460/* (Radford 450/x), TTI 400** to 500/* (Radford 
450/*), IV 530/* to 600/* (Radford 630/*) ; tarsi I 182/* to 191/* long by 97/* to 
106/* high; metatarsi I 115/* to 121/* long. Precoxal plates of leg I well 
developed, and conjoined to form a longitudinally divided sternal plate. 
Dorsal setae posteriorly to 45/* long (Radford 74**), uniformly thick and 
•ciliated, and apically with a conspicuous hyaline knob or blob. Genital discs 
3 pairs j in the male without any specialized setae. 

Nymph. Generally as in adult, but with the characteristic genital and 
palpal differences. Length 600/*; width across propodosoma 320/*., across hystero- 
soma 380/* (Walch gives 456/*, 264/* and 276*;. respectively. Radford gives no 



344 Records of the S.A. Museum 

values). Crista 106/a long (Walch 74ft; Kadford 102/;.) j width between sensillae 
bases 45/i, (Walch 33/a; Radford 51/*). Tarsi I 106/x long by 63/* high; meta- 
tarsi I 66/* long (Walch gives ratio of length to height of tarsi I as 2*2, and 
of length of tarsi I to metatarsi I 1*60; Radford gives no values). Posterior 
dorsal setae as in adult, to 45/* long (Walch gives 56/u,, Radford no value). 
Palpal tibia with only 2 accessory spines at base of claw. Genital discs 2 pairs. 
Remarks. The peculiar tip to the dorsal setae is also characteristic of 
pallida Nagayo et at.; and of A 7 , clavicata Andre. The first of these speeies, 
however, differs in the dorsal setae being thinner and longer, while N. clavi- 
cata Andre is a French species. Leptotrombidiwm akamushi which in the larval 
stage is closely allied to delie?isis is, however, very distinct in the adult, in the 
form of the dorsal setae. 

Tromjbicula (Leptotrombidium) pallida (Nagayo et ail. 1919). 

Tronibicida pallida Nagayo et al. 1919, Verdhl. d. jap. pathol. Gesellsch., 
Tokyo, 9, 107: idem 1921, Amer. J. Hyg., 1, (5-6), 569; Womersley and 
Ileaslip 1943, Tr. Roy. Soc. S. Austr., 67, (1), 98; Sig Thor and Willmann, 
1947, Das Tierreich. Lfg. 71b, 349. 

Plate 90, fig. N-S. 

Nagayo et al. (1921) claimed to have reared this speeies through from 
larvae to adult and to have recognized the adults captured in the field. They 
were unable, however, to find specific differences between the adults and nymphs 
of this species, and those of akamushi y pal.palis, and scutellaris, except that in 
pailida the minute setules on the basal part of the sensillae were much longer 
and more prominent than in the other species, and that the nymphs and adults 
of akamushi were much lighter in colour, 

I have before me a preparation from Prof. Dr, Takenouehi containing an 
adult caught in the field in Yamagata Prefecture, Japan, in July, 1920, and 
stated in his letter to be "probably T. pallida". In addition I have two slides 
of specimens, collected also in Yamagata Prefecture, by Col. C. B. Philip, which 
agree with that from Takenouehi. All three specimens have the posterior 
dorsal setae much longer than in deliensis, and these setae have the similar 
hyaline terminal knob, a feature not observed by Nagayo and his colleagues. 
A re-description drawn up from these specimens is as follows: 

Fades generally as in Trombicida, with the prominent constriction between 
coxae II and III. Colour reddish. Length 1,000-1,200/*; width across propodo- 
soina 490-610/i, across hysterosoraa 613-800/*. Crista linear 151/x long, with 
sensillae shortly barbed basally, rather longer barbs or cilia distally, 121/j. long, 
and their bases 51yj. apart. Palpi stout; tibia with 3 accessory spines at base of 



Womersley— Asiatic-Pacific Scrub Typhus Mites 345 

claw. Chelicerae finely serrate on inner margin. Epistome transverse rec- 
tangular, anterior margin straight or slightly concave, denticulate, with 1 
ciliated seta. Legs: I 700^ long, II 420-500/x, III 460-510/*, IV 530-600/x; 
tarsi I 196//. long by 94/u. high, metatarsi I 118/a long. Sternum about as wide 
as long, longitudinally divided, by the junction of preeoxal plates of leg I. 
Dorsal setae gradually increasing posteriorly to 90/* long, slender and parallel 
sided with strong setules and an apical hyaline knob or swelling. Genital 
discs 3 pairs. Male unknown. 

Trombicula (Leptotkombidium) akamushi (Brumpt 1910). 

TromMdmm aJcamuahi Brumpt 1910, Precis de Parasit., 2nd. ed., 506 (larva). 

For further larval synonymy see Larval Section. 

Leptotronibidmm akammhi, Nagayo et at, 1917, J. Exper. Med., 25, (2), 255. 

Leptus akamushi, Miyajima and Okumura 1918, Kitasato Archiv. Exper. Med., 
1,(1), 1-14. 

Trombicula akamushi Nagayo et al. 1921, Amer. J. Hyg. ? 1, (5, 6), 588; Hatori 
1919, Ann. Trop. Med. Parasit,, 13, (3), 233; Kawamura and Yamaguehi 
1921, Kitasato Archiv. Exper. Med., 4, 169; Wakh 1923 (1924), Tr, Yth. 
Bien. Congr. Far East. Assoc. Trop. Med., Singapore; Womersley and 
Heaslip 1943, Tr. Roy, Soc. S. Austr., 67, (1), 84; Sig Thor and Willmann 
1947, Das Tierreich, Lfg. 71b. 

Trombicula cmrctata, Kitashima and Miyajima 1918, Kitasato Archiv. Exper. 
Med., 2, 190-191, pi, 8, fig. 1 {non Berlese 1912); Ewing 1920, Ann. 
Entom. Soc. Amer., 13, 111 (non Berlese 1912) ; idem 1933, Proc. U.S. 
Nat. Mus., 82, 2 (non Berl. 1912). 

Plate 90, fig. H-M. 

This species, the larvae of which are the proved vectors of the tsutsuga- 
inushi fever of Japan and the Pacific area, is probably the best known and best 
figured of the Trombieulidae. 

Although there lias been much confusion amongst authors as to what is the 
true adult of akamushi, the studies of Nagayo and his colleagues, of "Walch, and 
particularly of Miyajima and Okumura have given very excellent descriptions 
and figures. 

From the above literature the life cycle of this species is generally fairly 
well established, and is discussed elsewhere in this paper. 

Nagayo et al, however, in 1921, reported the rearing of nymphs and 
adults of three other closely related species of larvae, viz r pallida, palpaUs, and 



346 Records of the S.A, Museum 

scwtellarhj but they were unable to distinguish the nymphs and adults on satis- 
factory morphological characters, except that in pallida the pigment coloura- 
tion was deeper and the minute barbs on the basal part of the sensillae were 
distinctly longer than in the other species, They did, however, separate them 
on the seasonal occurrence of the larvae. 

In the present paper it has already been shown that pallida can be 
separated from the other above mentioned species on the structure of the dorsal 
setae* Of palpalis and scutellaris I have not seen any adults or nymphs, and 
cannot separate these for the present. 

Of authentic material of akamiushi I have a preparation of 4 adults from 
Kawamura and collected in Niigata Prefecture, Japan. In addition I have 
been able to study 6 nymphs reared in New Guinea by Maj. G. M, Kohls from 
larvae identified as T. fietcheri Worn, and Heasp. 1943, a species now regarded 
as synonymous with akamushi Brumpt. Three of these nymphs -were from 
larvae on a quail, Dobodura, N.G., June, 1944 (No. 376, G.M.K.) and three 
from larvae on a rat, New Guinea 1944 (No. 156, G.M.K.) . 

Apart from the usual differences found between nymphs and adults these 
nymphs do not differ from the Japanese adults, thus supporting the synonymy 
of fietcheri with akamiiM. The following description and figures are from 
these specimens: 

Re-de&cription of Adult Of typical Trombicula faeies, with usual median 
constriction. Colour in life light to pale yellowish. Length to 1,000/x; width 
across propodosoma to 490/i, across hysterosoma to 535/i T Crista linear. 165/t 
long, with subposterior sensillary area with paired filamentous sensillae, 150/*. 
long, with minute barbs bus&lly and longer ciliations distally; sensillae bases 
to 65fu apart. Epistome transverse rectangular with anterior edge slightly 
concave medially and with fine denticles. Eyes absent. Palpi stout ; tibia with 
3 accessory spines at base of claw and a fourth midway between base of claw 
and articulation of palpal tarsus. Chelieerae with inner edge finely serrated. 
Legs all shorter than body; tarsi I 205ft by 90/* high (Hatori 180/* X 70^) J 
metatarsi I 136/* long. Sternum distinct, about as wide as long, divided longi- 
tudinally by the junction of the precoxal plates of leg I. Dorsal setae uniformly 
thick, 35/i to 45/x long posteriorly, ciliated to tip. Genital discs 3 pairs. Male 
unknown. 

Be-description of Nymph. As in the adult except for nymphal character- 
istics. Crista 104//, long ; sensillae 120/* long, their bases 44/* apart. Tarsi 1 100/* 
long by 58/* high, metatarsi I 63/* long. Dorsal setae as in adult, 20** to 45/x. 
long. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 347 

Trombicxtla (Leptotrombidium) myzantha sp. n. 
Plate 91, fig. A~G. 

This species is known from both larvae and nymphs. These nymphs, 
reared from larvae from various birds (see hosts in Larval Section) by Mr. 
D, A. Gill from Yerongpilly, Queensland, 18th Dec,, 1944, and two others from 
the same locality, 5th April, 1945, 

The species is separated from all other known adults or nymphs on the 
peculiar anterior and posterior dorsal setae, as given in the key. 

Description of Nymph. Of typical Tromhimla facies, with median con- 
striction between second and third pairs of legs. Colour in life red. Length 
1,066/* to 1,330/* (aver, of 5 specimens, 1,225/*) ; width across propodosoma 600/* 
to 800/* (aver. 718/*), across hysterosoma, 640/* to 860/* (aver. 796*/.), Crista 
typical of the genus, 190/* to 212/* long (aver* 199/0, with subposterior sensil- 
lary area, with paired filamentous sensiilae, 120/* long, with minute barbs 
hasally and short ciliations distally; sensiilae bases 45/* apart. Eyes absent. 
Chelicerae finely serrated on inner edge. Palpi rather slender; tibia with 
2 accessory spines at base of claw. Epistome transverse rectangular, anterior 
margin denticulate, with 1 ciliated seta. Legs all shorter than body, I 750/* to 
860/* long (aver. 796/*), II 470/* to 560/* (aver. 502/*), III 490/* to 570/* (aver. 
523/*), IV 530/* to 660/* (aver. 593/*) ; tarsi I fairly elongate, widest basad of 
middle, 197/* to 220/* (aver. 212/*) long by 82/* to 90/* (aver. 88/*) high, meta- 
tarsi I 121/* to 136/* (aver, 128/*) long. Sternum about as wide as long, with 
median longitudinal bar at the junction of the precoxal plates of leg I. Dorsal 
setae of two forms, anteriorly to 30/* long, somewhat clavate, becoming more 
slender and longer posteriorly to 110/*, and terminating in a distinct sigma- 
shaped spinule, not clavate; all setae with short ciliations, 

Trombicxjla (Leptotrombidium) fulleri (Ewing, 1945). 
Trombicula fulleri Ewing 1945, Proc. Ent. Soe. Washington, 47, (3), 46. 

Plate 92, fig. F-J. 

Description of Nymph*. Of typical Trombicula facies. Length 710/*; 
width across propodosoma 383/*, across hysterosoma 454/u Crista with triangu- 
lar sensillary area and dumb-bell shaped areola, 128/* long, sensiilae filamentous, 
barbed basally, and with short ciliations distally, to 120/* long, and bases 57/* 
apart. Chelicerae finely serrated on inner margin. Palpi stout, tibia with 
-simple claw and 2 stout spines at base of claw. Eyes absent. Legs.- I longer 
and stronger than the rest 497/*, II 312**, III 312/*, IV 383/* ; tarsi I 130/* long 



348 Records of the S.A. Museum 

by 80/i. high, metatarsi I 105/* long. Body setae fairly long and strongly 
ciliated, posteriorly to 60-80/* long and apically with a small spathulation. 

Remarks. This species is described from eight nymphs reared from larvae 
from Imphal 1945, by K. L. Cockings, the larval pelts being correlated with the 
larval description. 

TKOMBICULA (LEPTOrTROMBlDIUM) LONGISETA sp. n. 

Plate 92, fig. A-E. 

Description of Nymph. Of typical Trombicula facies. Length to 715/l; 
width across propodosoma 390/*, across hysterosoma 455/*. Crista elongate, to 
122/*, with subposterior sensillary area, furnished with paired sensillae to 98/* 
with short barbs on basal half and ciliations distally ; sensillae bases 53/* apart. 
Epistome not discernible, epistomal seta to 28/* long. Chelicerae finely serrate 
on dorsal edge. Eyes absent. Palpi stout, with two accessory spines at base 
of claw. Legs i I stronger than rest and to 585/*, II 390/*, III 390/*, IV 455/* ; 
tarsi I slightly less than twice as long as high, 136/* by 77/*, metatarsi I 80/*. 
long* Leg I with distinct precoxal plates forming a longitudinally divided 
sternum. Dorsal setae anteriorly ciliated and tapering, from 25/* long, length- 
ening posteriorly to 60-80/* long ; the posterior setae apically with a thickened 
tip as in fig. E. Genitalia with paired discs. 

Remarks. The nymph of this species is described from two specimens 
reared from larvae from Imphal, Burma, No. 169, 16 Aug., 1945, and No. 289, 
28 >Ang., 1945, by Mr. K. L. Cockings, and the identification checked with 
the larval pelts. 

In the larva, the species is closely related to deliensis, differing in the 
excessively long PL but in the nymph, it comes nearer to futteri, 

Trombicula (Leptotrombidium) burmensis (Ewing 1945). 

Troinbicula iurmensis Ewing 1945 (as larvae), Proe. Entom. Soc. Washington, 
47, (3), 64. 

Plate 91, fig. H-L. 

Description of Nymph, Of typical Trombicwla facies. Length 710*t ; width 
across propodosoma 420/*, across hysterosoma 560/*. Crista with diamond- 
shaped sensillary area and dumb-bell shaped areola ; to 112/* long ; sensillae fila- 
mentous, barbed basally, shortly ciliated distally, to 110** long, and their base* 
56/* apart. Eyes absent. Palpal tibia with simple claw and two strong blunt 



Womersley— Asiatic-Pacific Scrub Typhus Mites 349 

spines at base of claw. Legs: I longer and stronger than rest, to 500/* long, 
II 312/i, III 350/*, IV 448/*; coxae I with distinct precoxal plates forming a 
longitudinally divided sternum ; tarsi I 128/* long by 80/* wide : metatarsi I 78/* 
long. Dorsal setae fairly uniform, strongly ciliated, apically with spathulate 
tip, posteriorly the setae reach to 56/* in length. 

Remarks. This nymphal description is drawn up from 3 nymphs reared 
from larvae from Imphal, India, by K. L. Cookings, the recovered pelts being 
correlated, and their identification as larvae checked. As with all the species 
of Trombicula,, specific separation of the nymphs or adults is extremely difficult 
and unsatisfactory. The key given above is only an attempt to separate the 
species. 

Subgenus NEOTBOMBICITLA Hirst 1926. 

Ann. Applied Biol., 13, 140-143. Genotype Acarus auUmwlis Shaw 1790. 

Hirst raised this subgenus in 1926 for the nymph of Trambicula autumn- 
alia (Shaw 1790), on apparently slight but rather poorly specified differences 
from typical Trombmila as in akamushi Brumpt. These differences were 
mainly the more elongate form, and the absence of eyes. 

It is now possible to more precisely define the genus, as below, and to in- 
clude a number of other species. 

Diagnosis of Qenus. Rather more elongate but typical of Trombicula with 
the median constriction between legs II and III. Crista linear, with roughly 
diamond-shaped or rounded sensillary area furnished with two filamentous sen- 
sillae. Epistome rounded or subtriangular with fine teeth or denticles and 1 
ciliated seta. Eyes absent. Palpi rather slender ; tibia with 3 accessory spines 
at base of claw (2 in nymph) and a fourth between base of claw and articula- 
tion of palpal tarsus; occasionally palpal tibia also with pectine, Chelicerae 
with inner edge serrated. Legs shorter than body. Sternum entire, about as 
wide as long, undivided; no precoxal plates evident. Dorsal setae various. 
Genital aperture with 3 pairs of discs in adult (2 in nymph). 

Key to Known Adult and/or Nymplial Species. 

1. Palpal tibia with pectine, in addition to the 3 accessory spines at base 
of claw. Posterior dorsal setae spathulate apically, to 40/x, long, cili- 
ated. Trambicula (2V.) scincoides (Worn. 1944). 

No such pectine on palpal tibia 2 

2. Sensillae nude r 3 

Sensillae ciliated, at least distally . . r . 5 



350 Records of the S.A. Museum 

3. Posterior dorsal setae to ca. 40/* long . . . . . * - - . . 4 
Posterior dorsal setae to 110/* long, with few long outstanding setules. 
Front tarsi twice as long as high. 

Trombicula (AT.) inopinatum (Ouds. 1909). 

4. Posterior dorsal setae clavate or spathulate apically in dorsal view, 
•ciliated to tip, to 30/* long. Tarsi of leg I nearly 3 times as long as 
high. Trombicula (N.) rara (Walch 1924). 

Posterior dorsal setae normal, to 36/* long. Tarsi of leg I not quite 
twice as long as high. Trombicnda (AT.) hastata (Gater 1932). 

5. Dorsal setae with a prominent apieal trilobed knob, stem with strong 
setules. Trombimla (N.) jayeurickremei sp. n. 

Dorsal setae otherwise . . • * • . • • 6 

6. Posterior dorsal setae long and slender, to 350/*, ciliated, apical fifth 
spine-like, but with a few subterminal fibrillae. 

Trombicula (iV.) canestrinii (Buffa 1899). 

Posterior dorsal setae otherwise . . . . . - . . . - . * 7 

7. Dorsal setae parallel sided, terminating in a hyaline knob, from 22/* 
long anteriorly to 70/* long posteriorly. Sensillae fine and slender 
with short ciliations. Trombicula (JV,) clavicata Andre 1932. 

Dorsal setae not so . . . . . . , . . . . . . . 8 

8. Front tarsi at least 5 times as long as high 9 

Front tarsi not more than 3£ times as long as high 10 

9. Front tarsi ca. 5 times as long as high. Dorsal setae 40-50/* long 
anteriorly, 110/* posteriorly. Trombicula (A 7 .) formicarum (Beii. 1910). 

Front tarsi ca. 8 times as long as high. Epistome with occasionally 2 
setae. Dorsal setae long and slender, to 250/*. with ciliations. Palpal 
tibia with 4 ( ?) accessory spines at base of claw. 

Trombicula (N.) Camilla (Wharton 1938) 

10. Sensillae rather thick for whole length, with short ciliations on entire 
length. A pair of long ciliated specialized setae in front of sensillary 
area. Dorsal setae of two sizes, anteriorly mainly 14/* long, pos- 
teriorly 40/*. Trombicula (N.) spicea (Gater 1932). 

Sensillae much more filamentous. No such specialized setae in front 

of sensillary area .... . ► - . - - . t . * . . 11 

11. Dorsal setae of about equal length throughout, 62-65/*. Colour in life 
light yellow to white (after Vitzthum). 

Trombicula (N.) russica (Ouds. 1932). 

Dorsal setae much longer posteriorly than anteriorly . . . . . . 12 

12. Posterior dorsal setae to 40-45/* long , .. .. 13 

Posterior dorsal setae to ca. 100/* long . . . . . . . . . . 15 

Posterior dorsal setae 30-35/x. long . . . . . . . . 16 



Womersley — Asiatic-Pacific Scrub Typhus Mites 351 

13. Posterior dorsal setae to 45/n long, with parallel sides and at the 
tip expanded into an oval hyaline knob, furnished with setules. The 
longer setae also interspersed as well as posterior. Tarsi I ea. twice 
as long as high, metatarsi I 3-5ths length of tarsi. 

Trombicula (N.) fordi sp. n. 

Posterior dorsal setae to 40/^ apieally pointed with outstanding 

BCllUCtS • ■ < . . I .. . t . . .. ,. .. Art 

Posterior dorsal setae to 35/* long 16 

14. Tarsi T half as long again as metatarsi, and twice as long as high. 

Trombicula (N.) sarcina ("Worn. 1944). 

Tarsi I only slightly longer than metatarsi. 

Trombicula (N>) sp. n. (Cooreman in lift.) 

15. Front tarsi ca, 3-3 times as long as high, Sensillae with only about 
3 distal cilia, nude basally. Dorsal setae 45^., anteriorly increasing 
gradually to 97ju. posteriorly. Trombicula (N.) moesrica (Andre 1932). 

Front tarsi 2*5 times as long as high. Sensillae minutely barbed 
basally, and with 10-12 branches distally. Dorsal setae 30^ anteriorly, 
increasing to 110/t posteriorly. 

Trombicula (2V.) auMmnalis (Shaw 1790). 

16. Posterior dorsal setae to 30/*, long, tapering with outstanding cilia- 
tions. Tarsi I ovoid, half as long again as high, and almost twice as 
long as metatarsi I. Trombicula (N.) consueta sp. n. 

Posterior dorsal setae to 34/z. long, parallel sided almost to tip. Tarsi 
T twice as long as high and as long as metatarsi I. 

Trombicula (N.) harrisoni sp. n. 

KB. — The following four species, although doubtless to be placed in the 
subgenus Neotrombicula cannot be satisfactorily keyed on the data available 
in the original descriptions, and 1 have not been able to examine material; 
they are comrciata (Berl.), caveniarum (Ewing), manriquei (Ewing) and 
goldii (Ouds.). 

Trombicula (Neotrombicula) scincoides (Womersley 1944). 

Trombimda soincoides Womersley 1944, Tr. Roy. Soc. S. Austr., 68, (1), 84 
(larvae). 

Plate 93, fig. A-F. 

Originally only known from the larvae. I have since received from Maj. 
G. M. Kohls four nymphs which he was successful in rearing from fully 
engorged larvae, identified as this species, from a lizard from Dobodura, New 
Guinea, March 1st, 1944. 



352 Records of the S.A. Museum 

In addition from amongst a lot of miscellaneous acarina collected from soil 
near the 2/8 A.G.H., New Guinea, in April, 1944, were 3 adults and 1 nymph 
which can be referred to the same species as the reared nymphs. The descrip- 
tions of the nymphs and adults are as follows ; 

Adult. Of typical Trombicula fades. Colour in life red. Length to 
1*125 mm.; width across propodosoma 0-825 mm.; across hysterosoma 0*9 nun. 
Palpi fairly stout, with a long and strong tibial claw, at the base of which are 3 
strong blunt accessory spines; on the palpal tibia on the inner surface also 
is a pectine of from 7 to 9 strong spines ; palpal tarsus elongate and reaching 
beyond tip of tibial claw. Chelicerae stout with about a dozen backwardly 
directed inner teeth* Crista elongate, linear, 227/* long, bifurcate apically 
and with a subposterior diamond-shaped sensillary area; sensillary area with 
a more or less dumb-bell shaped areola on which the sensillae bases are 
63/i apart; ratio of SB (distance between sensillae bases) and entire length 
(including sensillary area) of crista — 3-6. Sensillae 180/* long, filamentous, 
ciliated distally. Bpistome rounded, conical, with fine denticulations and 1 
ciliated seta. Byes absent. Legs all shorter than body; I much thicker and 
stouter than others, 975/* long, II 600/*, III 675/t, IV 870ft; tarsi I 213/x to 239/* 
long (aver, of 3 a- 227/*) by 109/* to 121/* (aver. 115/*) high, giving ratio of 
3*6: 1-0; metatarsi I 167/* to 174/a long (aver. 115/*), giving a ratio of length 
of tarsi to metatarsi of 1-35 : 1-0. Sternum entire; no precoxal plates on leg I. 
Doraal setae moderately thick-stemmed, increasing in length posteriorly from 
24/i to 40/*. rather strongly curved and apically broadened or spathulate, with 
strong eiliations. Genital discs 3 pairs. In male sex, on each side of genital 
aperture with 3 nude strong curved spines, and a pair of large expanded and 
fringed titillating setae. 

Nymph, Similar to adult except for nymphal characteristics and smaller 
8ize» Length to 0-83 mm., width across propodosoma 0-525 mm., across 
liysterosoma 0-62 mm. Eyes absent. Chelicerae and palpi as in adult but only 
two accessory spines at base of palpal tibial claw and 3 spines in the pectine of 
palpal tibia. Crista 150/* long, sensillae 140/* long with bases 45/* apart; ratio 
of length of crista to SB = 3*3:1*0. Legs as in adult, I 675/j., IT 450/*, 
III 450/*, IV 530/* long; tarsi I 147/* long by 83/* high (aver, of 4 specimens), 
giving a ratio of 1-77: 1*0; metatarsi I 117/t long, ratio of length of tar«i to 
metatarsi — 1-26: 1-0. Sternum as in adult. Dorsal setae as in adult, vary- 
ing from 20/* anteriorly to 35/* long posteriorly. 

Remarks. This is rather a characteristic species and can be fairly readily 
recognized by the pectine on the palpal tibia and the structure of the dorsal 
setae. 



Womersley — Asiatic-Pacific Scrub Typhus Mites 353 

Trombicula (Neotrombicula) inopinatum (Ouds. 1909). 



As larvae. 



1893. Trombidium gymnopterorum Berlese, Order Prostigmata, 93, tab. 13, 

fig. 2, 3; tab. 15 (non Linne). 
1903. May 14th. Thrombidium gymnopterorum, Ouds., Tijdsch. v. Entom., 

45, 143. 

1903. July 17. Thrombidium gymnopterorum Ouds., ibid., 46, 5. 

1904. March 14. Thrombidium gymnopterorum, Ouds. and Heim, in CB. 
Acad. Sci. Paris, 1. 

1909. May 28. Thrombidium inopinatum Ouds., Tijds. v, Entom., 52, 43, 

tab. 37-^tO. 
1909. May 29. Thrombidium meriddonale Ouds., Ibid., 52, 45. 
1909. July 31. (Seance 17 July). Trom. gymnopterorum (fuMginosum) et 

Allotrombidium gymnopterorum (fuMginosum) Bruyant, CR. Soc. Biol. 

Paris, 67, 207. 
1909. July 31 I (Seance 17 July). Trombidium inopinatum, Bruyant, ibid., 

67, 208. 
1909. Aug. 7 f (Seance 24 July). Trombidium (Relerotrombidium) inopina- 
tum Verdun, ibid,, 67, 246. 
1909. Aug. 7 ? (Seance 24 July). Trombidium (Heter ot rombidium) nieri- 

dionale Verdun, ibid, } 67, 246. 
1909. Nov. 1. Microthrombidmm inopinatum Ouds. Entom. Ber., 3, (50), 

17, 20. 

1909. Nov. 1. Microthrombidmm meridionale, Ouds. ibid., 3, (50), 21. 

1910. Jan. 18. Microtrombidium pusillum Bruyant, Zool. Anz., 35, (11), 351. 
1912. Microtrombidium pimttum, Ouds. Zool. Jahrb. Suppl., 14, 31. 

1912. Acarus autumnalis, Ouds., ibid., Suppl. 14, 31, 

As nymph. 

1913. Microtrombidium autumnalis, Ouds. Arch. Naturg., 79A, 127 (Publ. 
20/3/1914). 

Plate 98, fig. H-M. 

The status of this species has by most acarologists been accepted as synony- 
mous with autumnalis Shaw, chiefly on the opinions expressed on several occa- 
sions by Oudemans himself, that the larvae, described by him as inopmatum 
and as meridionale, the gymnopterorum Berlese (non Linne), w T ere the same 
species as Shaw's. 



354 Records of the SA. Museum 

In addition to the original description and figures of the larvae of this 
species (Tijds. v. Entom., 52, 43, taf. 7, fig. 37-40) I have a photostat copy of 
Oudemans 1 original drawings, No. l t 104, in the Leiden Museum, kindly given 
to me by Miss A. M. Buitendijk. 

In the figures as published the galea! setae are not shown, but in the origi- 
nal they appear to be shown as ciliated. Oudemans states that the palpal claw 
is trifurcate, and that the setae on the palpal femur and genu are branched, 
but on the tibia only the ventral is branched. The posterior margin is angular 
with the side portions initially from PL slightly concave, thus the scutum is 
pentagonal. AM scutal seta reaches to just beyond the apex of the posterior 
scutal margin, and the sensillae bases are in the posterior half of the scutum 
and slightly behind line of PL. Oudemans quotes the dorsal setae as 8.8.6.6.4.4, 
but the modern reading of these, and as shown in his figures are 2.8.6.6*6.4.4 = 
36. The ventral setae are, posterior of coxae III, 10.8.8.8.6.2 = 42. Tarsi of 
leg III bear a long nude seta. 

These details agree with those British specimens which Richards (ParasitoL 
1950, 40. (192)- 105, 118) recently placed as his group I of autiumnaUs with 
the same characters (see under autwmnalis). It seems probable there- 
fore that Oudemans 5 inopinatum is a good species, and that Richards' speci- 
mens of his "first group" are the same. This is further supported 
by the fact that the nymph reared by Bruyant from the larvae diagnosed 
by him as mopinatum, but which Oudemans later regarded as awtumnalis 
{pusiUum)-, differs in a number of important points from the nymphs and 
adults of mihimnulis as described very fully by recent workers, Andre, Rich- 
ards, etc., and as observed and figured in this paper by myself. 

From Oudemans ' description and figures the essential features by which 
the nymph of inopinatiim differs from auUimmlis are ; 

The sensillary setae are nude and not ciliated. The front tarsi are only 
slightly longer than the metatarsi, 137/* and 123/a (in auiumnaUs. 180+l and 
145/*) ; the front tarsi are ca. twice as long a* high (2£ times in wutumnalis). 
The dorsal setae are gradually increasing in size to 110/a posteriorly in both 
species, but in inopinatum they have fewer and longer, more outstanding, 
setules. 

This species therefore appears to be valid, and occurs in Europe asso- 
ciated with the common "harvest mite" iV*. autumnalis Shaw. In the absence of 
eyes it will fall into Hirst's Neotrombiada. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 355 

TrombicttIiA (Neotrombicula) raba (Waleh 1924). 

TromMcida rara Walch 1924, Tr. Vth. Bieii. Cong, Far East. Assoc. Trop. 
Med., (Singapore 1923) • Womersley and Heaslip 1943, Tr. Roy. Soc. 
a Austr., 67, (1), 90; Sig Thor and Willmann 1947, Das Tierreich, Lfg. 
71b 5 283 (as larvae). 

Plate 93, fig. G-L. 

The nymphs of the species have been reared in some numbers by K. L. 
Gockings at Kuala Lumpur, from larvae from a pill-millipede, Qmscom&rpha 
sp. 

The following description is from a number of these nymphs kindly for- 
warded to me for study and description by Dr. J. R, Audy. 

Nymph. Of typical Tromhicula faeies. Colour in life pale and yellowish 
(Audy states in litt. that some, reared from larvae from a skink and one on 
two from rats were dark red). Length to 975/x; width across propodosoma 520/x, 
across hysterosoma 650//.. Legs I 710/l long and stronger than the rest, II 410/*, 
III 455/t, IV 585/*. Crista elongate, to 202/x long (including epistome 31/0 
with subposterior sensillary area furnished with a pair of very long, 154/4, fine 
filamentous, and apparently nude sensillae } epistome triangular or conical with 
dentate margin and setae to 42/* long. Eyes absent. Chelicerae 103/* long, and 
slender, indistinctly finely serrate on inner margin. Palpi normal, tibia with 
2 accessory strong spines at base of claw. Tarsi of leg I 175// long by 65//, 
high (ratio — 1-7: 1*0), metatarsi I 117/* long (ratio length tarsi I to meta- 
tarsi l = l'5:l-0). Dorsal setae on propodosoma, short 20/*, tapering with 
pectinations, on hysterosoma rather longer, and posteriorly curved apically and 
spathulate or clavate in dorsal view (in side view showing a thickening only), 
to 30/* long. Genitalia with 2 pairs of discs. No precoxal plates on tarsi I. 

Remarks. As given in the key this species can be distinguished by the 
nude sensillae and spathulate posterior dorsal setae. 

This species has also been reared by officers of the U.S.S.T.C. and a 
specimen or specimens of nymphs are in the collection of the Army Medical 
School at Washington. 

Trombicula (Neotrombicula) hastata (Gater 1932). 

Trombicula hastceta Gater 1932, Parasitology, 24. 

Neoschongastia hastata, Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 
67, (1), 110, (as larvae ). 



356 Records of the S.A. Museum 

Plate 94, fig. A-D. 

Descnptian of Nymph. Of typical Trombicida facies. Colour in life ?. 
Length to 520/*, width across propodosoma 312/*, across hyaterosoma 364/*. 
Crista linear, 98/* long, with subposterior diamond-shaped areola; sensillae 
long, to 103/*, apparently nude to tip, bases 36/* apart ; epistome not seen. Eyes 
absent. Chelicerae with fine serrations on dorsal edge. Palpi fairly elongate, 
tibia with single daw and two strong accessory spines near base. Legs : I 450//, 
long and stronger than the others, II 300/*, III 312/*, IV 364/* ; precoxal plates 
absent; tarsus I 112/* long by 64-4/* high, metatarsus I 75-6/* long; claws nor- 
inal. Dorsal setae to 36/* long, fairly thick with strong outstanding setules, not 
greatly tapering. Genital discs 2 + 2. 

Loc. Beared by Dr. Audy and his colleagues from larvae on Raitus 
sdbanvs and R. whiteheadi from Kepong Forest Eeserve, Kuala Lumpur, March 
1950. 

Remarks. The nymphs are only separated from those of other species on 
rather intangible characters as given in the key. 



Trombicula (Nbotrombicula) jayewickremei sp. n. 

Plate 94, fig. E-K. 

Description of Nymph. Of typical Tr&nibicula facies. Colour in spirit 
white. Length (newly emerged) to 780/*; width across propodosoma to 390/i, 
across hysterosoixta 450/*. Crista linea, 140/* long, with diamond or oval-shaped 
sensillary area with dumb-bell shaped areola; sensillae filamentous, ca. 100/* 
long, with minute barbs on basal half, and a few long ciliations distally ; sen- 
sillae bases 39/* apart Eyes absent. Chelicerae with inner edge finely serrated 
Palpi slender ; tibia with 2 strong spines at base of simple claw. Epistome rect- 
angular with denticulate anterior margin concave medially, and 1 ciliated seta. 
Legs I 590/* long and stronger than rest, II 375/*, III 325/t, IV 422/* ; tarsi I 
tapering, 118/* long by 42/* high, metatarsi I 62/* long. Sternum entire, precoxal 
plates not evident. Dorsal setae uniform, on hysterosoma to 30/* long on elon- 
gate peduncles situated in well defined pits; the setae terminate in a threc-lobed 
club as figured, which is in side view about 3 times as wide as the stem ; the 
stem carries strong setules which are longest around base of club; on the 
propodosoma the setae are slightly shorter, and merely thickened at the apex, 
not clubbed. 



358 Records of the S.A. Museum 

Trombicula (Nbotrombicula) clavicata (Andre 1937). 

ThrombicuU clavicata Andre 1937, Bull. Mus, Nat. Ilist., Nat., 2nd. Ser., 9. 
(5), 316. 

Plate 95, fig. A-E. 

This species was described from a single adult collected in the rotten 
stump of a vine by Dr. F. Grandjean at Banyuls-sur-Mer, France, May, 1935. 

The following description is abridged from that of AndrS: 

Adult. Colour white, with the general facies of Tromhicula. Length 
1,020//, width 640// on propodosoma, 600/* on hysterosoma. Crista 227// long, 
linear, with rounded subposterior sensillary area, with paired filamentous sen- 
sillae which have minute barbs proximally and somewhat longer ones distally. 
Epistome rectangular with fine denticles and I short seta. Eyes absent. Palpi 
slender ; tibia with 3 accessory spines at base of claw and a fourth more basad. 
Legs all shorter than body; I 850// long, tarsi almost three times (2-8) as long 
as high, 210/a by 75ft ; metatarsi I 140/t long. Body furnished with dense setae 
which are ciliated and apically terminated in a hyaline knob ; length of dorsal 
setae varying from 22// anteriorly to 70// posteriorly. Genitalia with 3 pairs 
of discs. 

Bemarks. Through the great kindness of Dr, Marc Andre, I have been 
privileged to examine the type of clavicata, and can add the following data. 
The crista to the base of the epistome measures 195// in length. The sensillae 
are 130// long, ciliated for their whole length, with the eiliations basally very 
short but lengthening distally. The sensillae bases are 35// apart. The sternum 
is entire, and there are no precoxal plates evident. 

The figures here given are after AndrS, but redrawn from the type. 

Trombicula (Neotrombioula) fobmicahum (Bcrlese 1910). 

Trombicula formicarum Berlese 1910, Bcdia, 6, (2), 369; idem 1912, Redia, 8 r 
(1), 90; Andre 1931, Archiv. Zool. Ital., 16, 1,358; idem 1932, Assoc. 
France Adv. ScL, Brussels, 273 ; Sig Thor and Willmann 1947, Das Tier- 
reich, Lfg., 71b. 

Plate 95, fig. H-J. 

This little known species has been recorded from Holland (Wasmann), 
Bavaria (Kneissl) and France (Andre). It was originally described from 
Holland from the nest of an ant La&i'm mixtus. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 357 

Remarks, The above description is from 10 nymphs reared from larvae 
from Rattus rattus handiycmus from Nalanda, Ceylon, (Sept., 1944), and a 
nymph from larvae from the same host, from Embilipitiya, Ceylon (Nov., 
1944). The nymphs were reared by Mr. S. H. Jayewickreme, after whom the 
species is named. 

The nymph is remarkable for the form of the dorsal setae, 

Trombicula (Neotrombicula) canestrixii (Buffa 1899). 

Trombidium cwnestrinii Buffa 1899, Atti Soc. Veneta Trent di Sci. Nat. 

Tromibicula carmtrinii, Berl. 1912, Redia, 8, (1), 88; Sig Thor and Willmann, 
Das Tierreich, Lfg. 71b. 

Thrombicula cwnestrinii Andre 1931, Arch. ZooL Ital., 1,358; 1932, Assoc. 
Franc. Adv. Sc, Brussels, 273. 

Plate 94, fig. L-R. 

This species was redescribed by Berlese (1912) from four specimens from 
the type locality, Cansiglio, Italy. This locality was alpine at an altitude of 
1,000 metres. I am indebted to Dr. Marc Andre for a certain amount of 
undetermined material amongst which were three specimens which can be 
with certainty assigned to Buffa *s species. One specimen was collected by 
Dr. F, Grand jean from moss at Vallombrosa, near Florence, Italy, May, 1934, 
and the other two by Dr. Andre from moss and vegetable debris at Mangaillard, 
France, 16th June, 1939. The following re-description is drawn up from 
Berlese 's and my own observations on the above fresh material. 

Adult. Colour white to light rose. Of typical Trombicula fades. Length 
to l,350/j. t width to 700/u Crista 292/* long, linear with roughly diamond-shaped 
sensillary area and paired sensillae, 150/x long, with distal dilations and their 
bases 97/x. apart. Epistome conical but rather flattened and with fine denticula- 
tions and 1 ciliated seta. Eyes absent. Chelicerae with finely serrate inner 
edge. Palpi relatively slender ; tibia with 3 accessory spines at base of claw 
and a fourth midway towards articulation of palpal tarsus. Sternum entire, 
as long as broad; no precoxal plates evident. Legs all shorter than body; 
tarsi I 3 times as long as high (3-5 Berlese), 312/x by llO/i., metatarsi I 260//. 
long. Dorsal setae long and slender with long ciliatious, but not extending to 
tip ; posterior setae to 350/*, with apical fifth or sixth spine-like, generally nude, 
but frequently with apical and subapical fibrils. Genital organ with 3 pairs of 
discs. 



Womerslev — Asiatic-Pacific Scrub Typhus Mites 359 

No re-descriptions, since Berlese 1912, have apparently been published and 
the following details are from Berlese J s work: 

Adult. Colour white. Length 1,900/*, width 1,000/*. Eyes absent or 
vestigial. Palpi slender, tibia with 3 accessory spines at base of claw. Tarsi 
of leg I nearly 5 times longer than wide, 470/* by 100/*, metatarsi 360/* long. 
Dorsal setae with moderately short setae to the tip ; from 40 to 50/* long anter- 
iorly, increasing to 110/* posteriorly, 

Tbombicula (Neothombicula) Camilla (Wharton 1938). 

Trombiada Camilla Wharton, 1938. Fauna of the Caves of Yucatan, X, Aea- 
rina. Carnegie Inst of Washington, No. 491, p, 141, fig. 1-3. 

Plate 96, fig. A-F. 

This species is remarkable for the very long front legs with their elongate 
tarsi. In the figures given here, those of Wharton 1938 have been reproduced 
with the addition, of figures of the epistome, front tarsi and metatarsi and a 
dorsal seta, which have been recently supplied by Dr. Wharton. In the figure 
of the epistome from the specimens in Wharton's collection, two epistomal 
setae are shown. In a recent letter, however, Dr v Wharton informs me that 
in the type in the U.S.N.M., only one epistomal seta is present. Occasionally 
such an aberration occurs in other adult species of Trombiculidae but here the 
epistome itself is hyaline, and denticulate as in the Trombiculidae, and not as 
in the Leeuwenhoelciidae, where two epistomal setae are characteristic. 

The description as given by Wharton, with additional data is as follows : 
"Adult. Large size, length 3-5 mm., width 2*2 mm. Palpi extend to the 
third segment of the first pair of legs ; palpal claw strong, half as long as the 
penultimate segment which bears it; accessory spines four, curved, subequal, 
forming a comb; palpal thumb not swollen. Chelicerae slender; chelae short 
and slender, extending beyond second joints of palpi. Crista rod-like expanded 
at anterior end and fringed as a rounded conical denticulate membrane, and 
with one seta (occasionally two) ; expanded at posterior end into a diamond- 
shaped psendostigniatic area, behind which it ends in two strongly diverging 
lateral arms. Pseudostigniata small, situated at lateral angles of pseudostig- 
matic area ; pseudostigmatic organs long and setif onn and nude. Eyes absent. 
Abdomen 2-5 mm. long; 2*2 mm. wide; setae moderate, barbed, posteriorly to 
260/*. Genital opening 0-5 mm. long, anterior end between hind coxae. Genital 
armature consists of a spathulate plate smaller than the genital opening with a 



360 Records of the S.A. Museum 

small shovel-shaped plate at base. Anal opening 0*2 nun. long, 0-2 mm. behind 
the posterior edge of the genital plate. Anterior legs slightly longer than 
posterior, about 2-5 mm. long*; front tarsi elongate, ca. 8£ times as long as 
wide, ca. 950/a by 110/*, metatarsi 700/*. Tarsal claws simple, those on first leg 
small and subequal, those of the other legs larger arid unequal/ * 

Type ioc. Oxolodt Cave, Yucatan. 

Type specimen. U.S.N.M'. No. 1,263. 



Trombicula (Neotrombicula) spicea (Gater 1932). 

Trombicula spicea Gater 1932, Parasitol., 24, 143-174; Womersley and Heaslip 
1943, Tr. Roy. Soc. S. Austi\, 67, (1), 76 (as larvae). 

Plate 96, fig. G-L. 

The nymphs of this species have been reared from larvae from the ears of 
Rattus r. kandiyamui from Embilipitiya, Ceylon, by Mr. 8. H. Jayewickreine, 
to whom I am indebted for material and the opportunity of describing the 
nymph. The identity of the larva which has not been recorded from anywhere 
since Gater's original description and record of the species from Selangor, 
F.M.S., has been checked against two of Gater 's paratypes, and the association 
of the larvae with the nymphs has been checked by examination of the larval 
pelts. 

In Ceylon the larvae of this species were associated with Schongastia (A&co- 
schongmtia) indica Ilirst, as was also recorded by Gater for his Malayan 
material. 

Description of Nymph. Of typical Trombicula facies, but somewhat slen- 
der. Colour in life? Length (freshly emerged) 650/*, width across propodo- 
soma 273/*, across hysterosoma 273/*. Crista linear, 112/* Jong, with broad 
somewhat diamond -shaped sensillary area, furnished with a pair of long, 98/x, 
uniformly thick sensillae which are barbed in basal part, a.nd shortly ciliated 
distally; sensillae bases 36/* apart; just anterior of sensillary area and on each 
sider of anterior rod is a long strongly ciliated seta, much longer (31/*) than the 
surrounding stae. Epistome rounded-conical with strong denticles and 1 
ciliated seta 22/* long. Chelicerae finely serrated on inner margin Palpi 
slender, tibial claw with 2 accessory spines. Eyes absent. Legs I longer and 
stronger than rest, 495/* long, II and III 285/*, IV 365/*; tarsi I 97/* long by 
45/* high, metatarsi I 84/* long. Sternum entire, undivided. No preeoxal 
plates. Dorsal setae of two distinct sizes, anteriorly 14/* long, posteriorly 40/* 
long, tapering, with outstanding setules. Genital discs 2 pairs. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 361 

Trombicula (NbotrombicuijA) (* russica Ouds. 1902). 

Larvae. 

Trombidiwm (Otonyssus) aurantiacum Ouds. 1897, {now. Kolenati), Tijds. 
Entoin., 40, 118. 

Thrombidium russicum Ouds. 1902, Entom., Ber., 1, (7), 43; 1903, Tijds., 
Entoni., 45, 142-143, tab. 12, % 39-42; idem 1903, ibid. 46, 5; idem 

1909, ibid. 52, 31, tab. 7, fig. 31-36. 

Allothrombidium russicwm Ouds. 1906, Entom. Ber., 2, (29), 87. 

Allothrombidium muscae Ouds. 1906, Entom. Ber., 2, 43. 

Thrombidium muscae Ouds. 1909, Tijds. Entom., 52, 35, tab. 6, fig. 26-30. 

Trombidium {Heterothrombidium) muscae Verdun, 1909, C.R. Soc. Biol. 
Paris, 67, 246 ; ibid., 67, 246. 

Microthrombidium muscae Ouds. 1909, Entom. Ber., 3, (50), 20. 

Microthrombidium russicum Ouds. 1909, Entom. Ber,, 3, (50), 20; idem 

1910, ibid., 3, (52), 47; idem 1912, Zool. Jahrb., Suppl. 14, 5. 

Adult. 

Trombicula russica Vitzthum 1932, in ZooL Jahrb., Suppl. 63, 687-690; Sig 
Thor and Willmann 1947, Das Tierreich, Lfg. 71b, 345. 

The adult of a species under the name of russica was briefly described by 
Vitzthum. The description as translated from Sig Thor and Willraann, 1947, 
is as follows : 

"Colour pale yellowish, almost white. Eyes absent completely. Body hairs 
as in autumnalis but not much longer posteriorly as in that species, from 62- 
65,*. Length of 2 1,200/a. 

From the Lutherothgrotte of St. Canzian Caves near Trieste. Coprophilous 
in bat guano." 

Remarks. As Oudemans has synonyinized his muscae and russica, k fcwo 
species which from his description and figures as published in 1909 (Tijds. 
Entom. 52) are obviously quite distinct, it is somewhat uncertain to which 
larvae Vitzthum 's adult must be related. 

Trombicula (Neotrombtcula) fordt sp. n. Lawrence, T. J. 

See Larval Section of this paper. 

Plate 96, fig. M-Q. 

Through the kindness of Lt. Col. Audy I have been privileged to study 
four nymphs reared from larvae of this species in Burma by Mr. K. L. Cock- 



362 Records of the S.A. Museum 

ings. The larvae were from Baft us rattus brunneusculus from ImphaU 
20/12/45. 

Description of Nymph. Size fairly large. Facies as in typical Trombkula. 
Length (newly emerged) 640/jl; width across propodosoina 355/*, across hystero- 
soma 385/x. Crista elongate with subposterior sensillary area, 102/* long 
excluding epistome, with epistome 115/t; epistoine rounded-conical with dentate 
margin, and 1 ciliated seta. Sensillae fine and filamentous, with minute barbs 
basally and short eiliations distally, to 60/* long and bases 29/x. apart. Chelicerae 
with inner margin indistinctly serrate. Eyes absent. Palpal claw with 2 
accessory spines at base and a nude spine on outer side; tarsi with 4 nude 
apical spines. Legs I longer and stronger than the others 420/*, II 280/*„ 
III 2S0fL. IV 325/i 3 tarsi I 108/* long by 51/* high, metatarsi 64/i. long. Dorsal 
setae, anteriorly 25/* pectinate and normal, posteriorly and generally inter- 
spersed to 45/4, with parallel sides which at the extreme tip expand into a 
small roughly oval tip, furnished with strong setules (see figure). Genital 
discs two pairs. 

Remarks. Characterized mainly on the peculiar dorsal setae and as in the 
key. 

Trombicula (Neotrombicula) sarcina (Womersley 1944). 

Trombicula sartim Womersley 1944, Tr. Roy. Soc. S. Austr., 68, (1), 95, 
(larvae), 

Plate 97, fig. A-F. 

This species has been reared to the nymph by Mr. D. A. Gill from larvae 
collected from sheep at Clermont, Queensland in Dec. 1944. The following- 
description is from three of these nymphs kindly sent to me for study by 
Mr. Gill, 

Nymph. Length to 660/*; width across propodosoma 430/x, across hystero- 
soma 470/*. Colour in life reddish. Crista as in the genus, 105/* long, sensillae 
with a few (3 to 4) eiliations distally, filamentous, ca. 100/* long, with bases 
33/* apart; ratio crista length to SB =3-2: 1-0. Palpal tibia with only two- 
accessory spines at base of claw. Epistome rounded, or subtriangular with 
fine denticulations and 1 ciliated seta. Chelicerae finely serrated on inner 
edge. Eyes absent. Legs all much shorter than body, with I the longest and 
strongest 5 I 460/i long, II 290/*, III 320/*, IV 370/*; tarsi I 114/* long by 57/*- 
high, ratio 2*0: 1-0,* metatarsi I 83/* long, ratio length tarsi to metatarsi = 
1-37: 10. Sternum entire, no precoxal plates on leg T. Dorsal setae shorty 
with long outstanding setules varying from 20/* anteriorly to 40/* posteriorly. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 363 

Trombicula (Neotrombicula) moesica (Andre 1932). 

Thrambicula eanestrimi v. moesica Andre 1932, Assoc. France Adv. Sci., 

Brussels, 273; 1937, Bull. Mus. Nat, Hist. Nat,, 2nd ser., 9, (5), 313. 
Trombicula moesica, Sig Thor and Willmann 1947, Das Tierreich, Lfg. 71b. 

Plate 97, fig. G-J. 

This species was described from a single adult female collected by % M. 
Bemy in the Gradje Cave, Yugoslavia, August 1930. 

Through the great kindness of my colleague Dr. M. Andre I have been 
able to examine this unique specimen and the following re-description is from 
that of Andre and my own observations. 

Adult 2 . Colour white. Of typical Trombicula femes. Length 1,420/x, 
width 780^. Crista 227/t long, with a more or less diamond-shaped sensillary 
area, with paired filamentous sensillae, 260/a long and with 3-4 long fine 
branches distally. Sensillae bases 78//. apart. Epistome 58/* deep, rounded- 
conical with strong dentieulations and one seta to 65/x long and ciliated. Eyes 
absent. Chelicerae on inner (dorsal) edge finely serrated. Palpi slender ; 
tibia with 3 accessory spines at base of claw (only 2 shown in Andre's figure) 
and a fourth midway between base of claw and articulation of palpal tarsus. 
Tarsi of leg I 30<fy long by 91//, high; metatarsi 227^ long. Dorsal setae uni- 
formly thick, ciliated to tip, anteriorly 45^ long, posteriorly 97/x. Genitalia 
with 3 pairs of discs. Sternum entire; no precoxal plates evident. 

Remarks. Andre compared his species with the three adult European 
species of Tr&mibityida then known, viz. canestrinii Buffa 1899, jormicarum 
Berl. 1910 and autumnalia (Shaw 1790), particularly in regard to the relative 
length and height of the first- tarsi, and the lengths of the dorsal setae. On 
the close agreement of the proportionate length to height of the front tarsi, 
3-5 in earnest rimi, he regards his species as a variet3 r of this species. Apart from 
the very slight difference in the front tarsi there is a much more obvious and 
important difference in the structure of the dorsal setae. In canestrimi as 
figured by Berlese, the dorsal setae and particularly the posterior setae are 
very much longer and the apical fifth or sixth is shown as nude and spine-like. 
In the specimens of eanestrinii which I have been able to study, this is the 
general appearance under medium magnification; under higher magnification 
however, many of them show that the extreme part of the spine-like tip is itself 
fibrillated. In moesica besides being shorter, the dorsal setae have fairly long 
setules extending right to the tip. 



364 Records of the S.A. Museum 

Trombicula (Neotrombicula) autumnalis (Shaw 1790). 

Acarus awtumndtis Shaw 1790, Nat. Misc., 2, 42. 

Leptus mtiwmnalisy Brandis 1897, Festschr. d. Provincial Irrenasstalt, Nietle- 
ben; Bruyant 1910, Zooi. Anz., 35, 347-352; Picard 1927, Bull. Soc. Zool. 
France, 52. 

Micro trombidium autumnalis, Oudemans 1912, Zool. Jahrb. Suppl., 14; idem 
1913, Acarol. Maulwerfnestern, Arehiv. Naturgesch., (9), 127 (these 
probably refer to inopinatum Oudfl. 1909). 

Trombiculu autumnalis, Hirst 1926 (Neotrombicula) , Ann. App. Biol., 13, 140- 
143; Andre 1929, (Thrombimtla) , Bull. Soc. Zool. France, 54; idem 1929, 
Bull. Mus. Nat. Hist. Nat., 2nd ser., 1; idem 1930, Mem. Soc. ZooL France; 
idem 1931, Arehiv. Zool. Ital., 16; idem 1937, Bull. Mus. Nat. Hist. Nat., 
2nd ser., 9, 313; Warburton 1928, Parasitology, 20, 228-236; Sig Thor and 
Willmann 1947, Das Tierreich, Lfg., 71b. 

Plate 98, fig. A~G, 

The first attempt to rear the nymph of this species from the larvae appears 
to be that of Brandis 1897. The nymph obtained was, on account of its thick 
coating of hairs, considered as that of Trombidium holoserid%tmi\. Bruyant 
1910 also succeeded in obtaining a single nymph, which was regarded as 
Microtrombid/ium pusillum Herm. by Oudemans 1912. In 1913, however, 
Oudemans on re-examination of the specimen showed that it differed from 
Microtrombidium pusillum in the structure of the front tarsi, palpi and dorsal 
setae. He regarded it as distinct and referred it to Microtrombidium autumn- 
alis (Shaw). In 1916, Zool. Anz., 46, 253, Kneissl obtained nymphs from 
larvae identified as T. inopinatum Ouds. 1909 (a species the larvae of which 
are generally regarded as conspecific with auturrmalis (Shaw). He recog- 
nized these nymphs as belonging to the genus Trombicula BerL 1904 and 
unhesitatingly referred the nymph reared by Bruyant and figured by 
Oudemans 1913 to autumnalis. Stanley Hirst 1926 also bred a nymph from 
larvae identified as Leptus autumnalis which he showed belonged to Trom- 
bicula BerL, and designated this species as the type of a new subgenus 
Neo trombicula.. 

Our most complete study of the nymph and adult of this species is that of 
Marc Andre (1930), and at the present time autumnalis is probably the best 
known and described species, from all stages. The following re-descriptions 
of the adult and nymph are mainly from Andre's studies, supplemented or 



Womersley— Asiatic-Pacific Scrub Typhus Mites 365 

checked by lny own observations on British material, obtained while in London 
in 1947 from Dr. W. 8. Richards. 

Re-description of Nymph. Facies of TrombictUa but rather elongate with 
the hysterosoma narrower than the propodosoma. Length 930/*, width of 
hysterosoma to 310/* (after Andre). Colour in life, pale to light yellowish. 
Crista linear, with a subposterior diamond-shaped sensillary area; with paired 
filamentous sensillae each with 5 or 6 distal setules. Epistorae somewhat tri- 
angular, with denticulations, and 1 long ciliated seta. Eyes absent. Chelicerae 
with inner edge finely serrated. Palpi moderately stout; tibia with 2 acces- 
sory spines at base of -claw. Legs all shorter than body, I the longest and 
strongest, with, tarsi I 180/x long by 70 p high (Andre), metatarsi 145/i. long* 
Dorsal setae ciliated to tip, 30/t long anteriorly, increasing to 110^ posteriorly 
(Andre). Sternum entire, roughly pentagonal and as long as wide; no pre- 
coxal plates evident. Genital orifice with 2 pairs of discs. 

Re-description of Adult. As in nymph in shape. Length to 2,050/a, width 
across hysterosoma to 850/n. Colour white to light yellow. Crista, sensiUary 
area, sensillae and epistome as in nymph. Eyes absent. Palpi as in nymph 
but with 3 accessory spines at base of tibial claw. Legs as in nymph. Dorsal 
setae as in nymph. Genital organ with 3 pairs of discs. Sternum as in nymph; 
no precoxal plates present. 

Remarks. W. S. Richards has recently (Parasitology, 1950, 40, (12), 105, 
and 118) published two very important papers on the variation in the larvae r 
and on the distribution and biology of the European harvest-mite in Britain. 
In the first he shows that very considerable variation occurs in morphological 
characters of the larvae regarded as this species, such as; (1) the ciliated or 
nude galeal setae and the dorsal and lateral setae on the palpal tibia, (2) the 
duplication or otherwise of some of the scutal setae, (3) the lengths of the 
scutal setae, (4) the number and arrangement of the humeral and dorsal setae, 
and (5) the Standard Data of the scutum. Despite all these variations, he is 
mclined to the view that all the larvae he studied are to be referred to the 
one species Trwtibieula autumnalis (Shaw). 

In my experience, some of the variations suggest that at least two species 
occur in Britain. Dealing with the variations, the occasional (apparently more 
frequent in Britain) duplication of one or more of the scutal setae is to be 
found in many species of Trombiculidae, and is a teratological occurrence of 
no taxonomic value, provided that it is recognized for what it really is. For 
example, the occasional occurrence of 2 AL in a Trombieulid specimen is not to 
bo confused with the 2 AM in a Leeuwenhoekiid. Similarly the occurrence of 



366 Records of the S.A. Museum 

2 humeral setae on each side may only be a freak variation and of no specific 
importance. 

The specimens with ciliated or branched galeal and dorsal and lateral 
palpal setae together with dorsal setae 2.8.6.+, with a total of 36, seem to sug- 
gest the species T. wopinatum rather than true autamtwlis. The larvae of 
T. inopinatum was described in 1909 (Tijds. v. Entom., 52, 43, tab. 7, fig. 37- 
40, Onds.)- In Oudemans' original drawing No. 1,105, in the Leiden Museum, 
for a photostat of which I am indebted to Miss A. M. Buitendijk, are given 
figures of the. nymph reared from Bruyant'a larva, and labelled as Mkrotrom- 
hul/ium inopinatum Ouds. 

The nymph as thus figured by Gudemaus (these figures were published 
1913 Acarologisches aus Mauiwurfnestern, Arch. Naturg., LXXIX — A, taf. 
XII, fig. 1-6) under the name of MicrotronMdium awtumndlis (Shaw) does 
not coincide with the details of tovtiumnalix as delineated by recent students, 
Andre, Richards, and myself herein. In particular, the sensillae are shown 
and stated to be nude, whereas in mtumnalis they are ciliated. The dorsal 
setae although of one kind, and gradually lengthening posteriorly are more 
sparsely beset with longer setules; they reach posteriorly, however, to approxi- 
mately the same length of 100-110/t. (In W, S. Richards' first paper, loc> ci>t T 
in a footnote, he gives the mean length of the posterior setae in two lota of 
adults as 10-5+:0-86 ? and 9«7dil-5V This is probably an error for in some 
of his English material which I have examined, these setae, are up to lOO/i as 
given by Andre ei ah.). It would seem then that some of Richards' larval 
material might possibly be referred to Oudemans 1 inopinatum which is probably 
a good species, distinguishable in both larvae and adults from autumnalis. 
Richards was not very successful in rearing nymphs and those which he obtained 
appear to conform to typical autumnalis. The metatarsi of leg I in the nymph 
of vnopinatwn as figured by Oudemans are very little shorter than the tarsi, 
123^ to 137/1, whereas in autumnalis, the lengths are 195^ and 260/i respectively. 

Richards unquestionably had adults of both sexes of autuvwuiitis but in his 
figure of the male (fig. 22) he has omitted and probably failed to observe the 
pair of specialized titillating setae on each side of the penis opening. 

Whether the specimens with mtumnalis type of galeal and palpal tibial 
setae but with 2.8.6.+, and more than 32 dorsal setae are only variations of 
this species or should be referred to something else must, as also with ino'pina- 
turn, await further and more detailed work on the lines of rearing larvae from 
known adult females. Richards' second paper on the " Distribution and Bio- 
logy of the Harvest Mite" also suggests that the two major forms (with and 
without galeal setae etc. ciliated) occur at different seasons and in different 



WOMERSLEY— ASIATIC-PACIFIC SCRUB TYPHUS MlTES 367 

climatic areas although some overlapping occurs, e.g. the inopinatum form was 
not met with north of Oxford, whereas the typical, and its variations occurred 
throughout England and in Scotland. 

Trombiotjla (Nbotrombicuiia) coarctata (Berlese 1888). 

Tronibidimn cmrctatum Berlese 1888, Acari Austro-amerieani Bull. Stic entom. 
Ital., 20, 179, pi. 5, fig. 5; idem 1893, A.M.S., Prost, 98; Berlese and 
Leonardi 1902, Zool. Anz., 25, 17. 

Trombicula coarctata, Berlese 1912, Redia, 8, (1), 91-92, fig. 42; non Kitashima 
and Miyajima 1918, Kitasato Archiv. Exper. Med., 2, 190-191, pi. 8, fig. 1; 
non Ewing 1920, Ann. Entom. Soc. Anier., 13, 382-389 • non Ewing 1926, 
Entom. News, 37, 111; Ewing 1931, Proc. U.S.N. Mus., 80, 7, (in part) ,- 
non Ewing 1933, Proc. U.S.N. Mus., 82, 2; Sig Thor and Willmann 1947 y 
Das Tierreich, Lfg. 71b, nan fig. 416. 

Plate 95, fig. K-L. 

This species was originally described by Berlese from two specimens from 
under stones, one from Buenos Aires, and one from Rio Apa (Paraguay). It 
was later (1902) recorded by Berlese and Leonardi from Chili. Since then 
however, no authentic material appears to have been seen and studied. 

The following description is a compilation from Berlese's original paper 
and his later notes of 1912. 

Nymph. Colour white. As in typical Tronib&iUa with a constriction 
between second and third pairs of legs ; shoulders well rounded. Eyes absent. 
Crista linear with subposterior sensillary area about as wide as long. Sen- 
sillae ? (probably filamentous). Palpi slender, tibial claw with 2 accessory 
spines at base (hence Berlese \s specimens would be nymphs) ; tarsus elongate 
elavate. Legs slender and short; posterior hardly reaching beyond abdomen; 
tarsi I widest basally, apex subacute, subtriangular; tarsi I 180ft long by 75//. 
high, metatarsi I 130/a long. Body densely covered with setae, which are only 
slightly tapering, to 70ft long posteriorly and with outstanding ciliations. 
Length to 1,150/a, width to 850/*. 

Rema/rks, In 1918 Kitashima and Miyajima considered that the adults of 
the Japanese **kedani M mite were conapecific with the S. American species. 
Ewing (1920) reported having, along with Miyajima, studied Japanese speci- 
mens and compared them with the American species, splendens Ewing and 
cimuibannus Ewing (= alfreddugesii Ouds.). They also concluded that coarc- 
tata and the Japanese material were the same species, and published a re- 
description and figures of coarctata based on Japanese specimens. 



363 Records of the: 5.A. Museum 

Apart from the very widely different geographical localities, there are how- 
ever, certain subtle differences to be found between the descriptions and figures 
of Berlese. and those of Kitashiuia and ftTiyajima, and of Ewing. 

Berlese gives the colour as "white" whereas the Japanese species accord- 
ing to Kitashima and Miyajinia is "light reddish colour." Another important 
specific difference is seen when we compare the figures of the dorsal setae. 
Berlese shows them as slightly tapering with relatively few outstanding setules. 
In Ewing's figure, the setae are heavily ciliated and somewhat bushy towards 
the tip. In coarctata Berlese the posterior dorsal setae reach to 70/i in length. 
Ewing gives no dimensions for the Japanese specimens. Again, although the 
front tarsi are closely similar, those of the Japanese form are somewhat longer, 
2-5, in proportion to height, than the ratio of a 2-4 given by Berlese. The 
structure of the epistome was not studied or delineated by earlier workers, 
but Ewing figures it for Miyajima 's material as rectangular with anterior edge 
straight. Although this form of epistome is not unknown in the Trombiculidae, 
it is unusual; the genral form is rounded-conical. Berlese did not stress or 
describe the sensillae of the S. American species. 

Unquestionably then the Japanese material was not con-specific with coa*c- 
tata Berl. but was probably identical with akamushi Brumpt 

Tbombicula (Neotrombicula) cavernabum (Ewing 1933). 
Trombicvia caverTiarum Ewing 1933, Proc. U.S. Nat. Mm, 82, (29), 4. 

Plate 95, fig. F-G. 

This species was described from two males and one. female from Chilibrillo 
Caves, Panama. 

The original description is as follows . 

"Adult. Moderate in size and of the shape typical of the genus. No capi- 
tulum present. Palpi reaching beyond the middle of the first patellae j palpal 
claw weak, but little over half as long as the arched segment which bears it; 
accessory spines four, subequal, forming a comb; palpal thumb slightly swollen 
and extending to middle of palpal claw. Chelicerae very slender; chelae 
slender, sharp, tips extending slightly beyond the tips of second palpal seg- 
ments. Crista rod-like, expanded near its posterior end into a diamond-shaped 
pseudostigmatic area slightly beyond which the crista ends in two .strongly 
diverging lateral arms. Pseudostlgmata moderate, situated near lateral angles 
of pseudostigmatic area ; pseudostigmatic organs very long, setif orm, with two 
barbs. Eyes absent. Abdomen about one and a half times as long as broad; 



Womersley— Asiatic-Pacific Scrub Typhus Mites 369 

setae short for the genus. Genital opening about one-half of its diameter from 
posterior eoxae. Genital armature of male composed of a basal plate, a pro- 
truding penis, and an inwardly projecting manubrium j basal plate horse-shoe- 
shaped; penis a cone-shaped, sclerotized structure, situated between the two 
arms of the basal plate; manubrium somewhat longer than penis and arising 
from base of same. Anus somewhat smaller than genital opening and situated 
about one and a half times its greatest diameter behind the same. Anterior 
legs only slightly larger than posterior. Tarsal claws all simple, those of the 
first legs quite small and subequal, those of the other legs unequal, the pos- 
terior claw being larger than the anterior. Length, 1-12 mm.; width 0*7 mm. n 
Remarks* No dimensions other than total length and width are given, and 
it is very difficult to compare this species with others from what data is avail- 
able. In the absence of eyes and the structure of the sternum it belongs to 
the subgenus Neotronibicula, and on. the proportions of the front tarsi and 
metatarsi is close to formicarum Berl. and Camilla Wharton. 

TrOMBICULA (NEOTROMBICUIiA) MANRIQTTEI (Ewing 1937), 

Tromb&ula mimriquei Ewing 1937, Proc. Biol. Soe. Washington, 50, 169 ; Sig 
Thor and Willmann 1947, Das Tierreieh, Lfg. 71b, 353. 

This species is as yet, only known from a single specimen, described by 
Ewing, and collected from humus in a tree hole at Villa vicencio (Quenane) 
Meta Department, Colombia. Type in U.S.N.M. 

Swing's description is as follows: 

* 4 Palpus slender, extending about one-half its length beyond chelicerae. 
Palpal thumb slender, well clothed with setae and slightly surpassing tip of 
palpal claw; palpal claw rather slender without tooth, moderately curved; 
accessory spines three, subequal, arranged in a comb. Crista extending for- 
ward as a pointed rod between bases of chelicerae and expanded posteriorly into 
a laterally rounded pseudostigmatic area, the latter without posterior lobes. 
Pseudostigmatie organs simple, flagelliform, longer than crista. Eyes absent. 
Abdomen densely beset with tapering barbed setae which increase progressively 
in length from anterior to posterior border. Anterior legs longest; posterior 
next in length; legs of second and third pairs subequal. Claws on tarsi I 
unequal, also those on tarsus IV unequal - those on tarsus II subequal, and on 
tarsus III subequal. 

"Length of body 0-96 mm.; greatest width 0*53 mm." 

SemarJcs. Ewing states> M Although this specimen is small for an adult of 
the genus Trcmibicula, the presence of three accessory spines near the base of 



370 Records of the S.A. Museum 

the palpal claw and of three pairs of genital suckers surrounding the genital 
opening indicates it is an adult. This species is nearest T. coarctata Berlese 
and T. spleitd-ens Ewing, from both of which it differs in having no posterior 
lobes to the pseudostigmatie area." 

Trombicula (NBOTOOMBtcxjijA) goldh (Oudenians 1910). 

Microtrombidium goldm Oudenians 1910, Entom. Ber. Nederl. Ver., 3, 84; 
idem 1912, Zool. Jahrbs. Suppl. 14, 18-15 (larva), 

Trombicula goldii Ewing 1931, Proc. TT.S.N. Mus., 80, 7 (larva). 

Trombicula (Eutronibicula) goldii, Sig Thor and Willinann 1947, Das Tier- 
reich, Lfg. 71b, 279 (larva). 

Trombicula goldii, Boshell and Kerr 1942, Riv, Acad. Columbiano Cien. Exact. 
Fisic, Nat., 5, 112; Michener 1946, Ann. Ent. Soc. Ameiv, 39, (1), 101-118 
(adult). 

Plate 95, fig. M. 

This species, in the adult stage is only known from Colombia, S. America, 
where it is apparently fairly common. 

A translation of Boshell and Kerr's description is as follows: 
Adult. Size small with pronounced constriction between abdomen and 
cephalothorax; colour carmine; legs much shorter than body; anterior legs 
much thicker than others. Palpi long and stout, reaching to base of patella of 
leg I ; segment II widened ; claw of palpi strong and simple, two-thirds length 
of fourth segment ; a row of 3 unequal spines on small protuberance at base of 
claw. Thumb of palpi wide, lightly clavate and reaching tip (of claw) ; fur- 
nished with pectinate setae. Mandibula apophysis (chela) strong, sickle-shaped, 
with finely serrate concave edge. Eyes absent. Crista well developed, rod- 
like anteriorly with a wide triangular subposterior area with two pseudostig- 
mata in the posterior lateral angles. Pseudostigmatie organs flagelliform, long, 
with some small dilations on distal third. Dorsal setae of one type, dense, but 
longer posteriorly than on shoulders; situated on elevated discs, ending in a 
point and ciliated. On the dorsal surface, postero-lateral of the anus, one 
finds a pair of areas, refractive, somewhat in relief and resembling eyes. 
Genital aperture dimorphic and easy to see in cleared or partially cleared 
specimens. The female has a pair of peculiar cup-like structures, approxi- 
mately half the diameter of the genital discs, and surpassing the posterior pair 
of discs. The males have an ovoid structure without precise details, approxi- 
mately twice the size of the genital discs and placed in the middle of these. 
Leg I approximately twice as thick as IV; I and IV about equally long, II 
and III distinctly shorter than IV. Tarsi I not wider (higher) than patella I, 



Womersley— Asiatic-Pacific Scrub Typhus Mites 371 

symmetrical, conical, with the distal diameter a little leas than half the basal 
diameter. Tarsi I approximately 1-4 times as long as the patella 5 other seg- 
ments about equal. The other tarsi relatively longer and more slender with 
extremities truncate. Each tarsus with a pair of equal, slender and curved 
claws ; those on tarsi I about half the size of the others. 

Locabtiie^ Kestrepo, Villvicencio (Meta) at 400-900 metres; Municipality 
of Bolivar (Santanda) at 500-1,000 metres; Municipality of Muzo (Boyaca) 
at 700-1,000 metres. Found in humus, tree holes and under bark of fallen 
logs, at all seasons of the year. 

The captured adults were correlated with a nymph and adult reared in 
the laboratory. 

Retnarks. There is no doubt that this is a true Trombiculid despite the 
author's statement "with pronounced constriction between cophalothoras and 
abdomen"; this is surely meant to refer to the usual constriction between the 
second and third pairs of legs. 

The author's description of the difference in the genitalia of the two sexes 
is interesting and important but would be better understood if figures had 
been given. 

Trombicula (Neotbombicula) sp. n. (Cooreman in lift.). 
Plate 98, fig. N-CK 

In 1947, on a visit to the Musee Royal D'Histoire Nat. de Belgique, Brux- 
elles, my friend Dr. J. Cooreman drew my attention to a single specimen of 
an adult Trombicula which he had collected from a "microcaverne endoxyle" 
in lllmus campesiris at Boitsfort, near Bruxelles, March 25, 1947. At the time 
it was thought that it might be Trombicula -inopinatum Ouds., but our examina- 
tions of the specimen shows that it is probably a new species, which Dr. Coore- 
man will be duly describing. 

In the meantime I am greatly indebted to Dr. Cooreman for the following 
details and figures which will enable the species to be separated from others. 

Length 720/a, width 380//,. Of typical Trombicula faeies. Eyes absent. 
Crista 150/x. long, with triangular dentate epistome, with smooth epistomal 
seta; sensillae long and fine, with sparse barbs distally. Legs: I 420/* long, 
II 250/*, III 240/x, IV 325/ t ; tarsi 1 115//. long, ? wide, metatarsi I 100/a long. 
Dorsal setae figured, anteriorly 20-30/*, posteriorly 40/x, 

Remarks. The species is close to sarcind Worn, from Australia, but differs 
in the smooth epistomal seta and the relative lengths of front tarsi and meta- 
tarsi. In the smooth epistomal seta and shorter posterior dorsal setae it differs 
from both mituninalis and inopinatum, while the distally barbed sensillae sep- 
arates it from inopinatum. 



372 Records of the S.A. Museum 

TrOMBICULA (NEOTliOMBICUIiA) HARRISONI Sp. Q. 

Plate 113, fig. A-E. 

Description of Nymph, Of typical Trombicula facies. Length 355/*; width 
across propodosonia 195/*, across hysterosoma 225/*. Crista elongate, length 
including epistome 112/*; subposterior sensillary area as figured, with SB 31/*,; 
sensillae filamentous, but rather thickened medially, to 90/*, with short ciliations 
on distal four-fifths; epistome rounded-conical, dentate, with ciliated seta 20/* 
long. Eyes absent. Chelicerae strongly serrate on inner edge. Palpi rather 
slender; tibia with two accessory spines at base of claw. Dorsal setae from 
10-14/* long anteriorly, lengthening to 34/* posteriorly; anterior setae tapering 
with outstanding setules, posteriorly more parallel sided almost to tip. Leg 1 
slightly longer than body, to 375/*, II and III 195/*,. IV 225/*; tarsi of leg I 84/* 
long by 42/* high, metatarsi I 84/* long. 

hoc. The type nymph is one reared by the British Scrub-typhus Kesearch 
Team, in Malaya from larvae of the new species liwrrisoni, from bats No. 11018, 
from Bukit Lagong Forest Reserve, Kepong, Malaya, 1950, and correlated 
with the larval pelt. 

Remarks. As with most nymphs and adults of the Trombiculidae, specific 
separation from other species is based on rather intangible characters. An 
attempt to separate the above species has been made in the key. The larvae, 
however, is abundantly distinct. 

Genus GUNTHEEANA Womersley and Heaslip 1343. 

Ountherana Womersley and Heaslip 1943, Trans. Roy. Soc. S. Austr., 67, 
(1), nom. nov. for Guntheria Womersley 1939, Rec. S. Aust. Mus., 6. (1), 
157, preoc. 

Genotype Neoschongastia kallipygos Gunther 1939 (larva). 

The nymphs and adults of this genus have not hitherto been described. In 
the present work the adult and nymphal stages of the genotype Guntherana 
hipygalis (Gunther 1939) are described and the genus defined for these stages. 
A second species from Kangaroo Island, South Australia, G. tiyidahi (Worn. 
1946) known from a captured adult is now referred to the genus. 

Diagnosis of Genus (from Adult or Nymph). Form as in the genus Trom- 
bmila. Crista linear with very widely transverse sensillary area, with a dumb- 
belKshaped areola, and shallow posterior angle, filamentous sensillae, and the 
anterior arm of the crista shorter than the distance between sensillae bases. 
Epistome anteriorly concave with fine denticles and 1 seta. Eyes absent. Pre- 



Womersley— Asiatic-Pacific Scrub Typhus Mites 373 

coxal plates of leg I well defined, and barely touching in middle line so that 
there is no true sternum. Tarsi of leg I without any dorsal subapieal process. 
Dorsal setae tapering, slender and ciliated. 

Genotype (?. bipygalis (Gunther 1939). 

Key to the Adult and Nymphal Species. 

1. Colour in life red. Dorsal setae fairly thick, to 7(V l° n £ posteriorly. 

Tarsi I less than twice as long as high. G. bipygalis (Gunther 1939). 

Colour in life white . . . , . . . . . - ► - . . . - 2 

2. Tarsi I ca, 2-J times as long as high. DorsaJ setae rather fine, to 100/i 

long posteriorly. G. tindolei ( Womersley 1946) . 

Tarsi I twice as long as high. Dorsal setae much thicker, anteriorly 
on propodosoma to t&ja long, posteriorly on hysterosoma forming a 
conspicuous fringe with setae 65-70/* long. 

G> trawslueens (Woroersley 1944). 

Guntherana bipygalis (Gunther 1939). 
Neoschongastia caUipygea Gunther 1938, nam. nud. Med, J. Aust., 2, (6), 202, 

Neoschdngmtia kaUipygos, 1939, nom. mid. Derrick, Smith, Brown and Free- 
man, Med. J. Aust., Jan., 150. 

Neoschongastia kallipygos, 1939, Gunther, Proc. Linn. Soc. New South Wales, 
64, (1-2), 83. 

Neoschongastia bipygalis 1939, Gunther, Proc r Linn. Soc t New South Wales, 
64, (5-6), 471. 

Guntheria kallipygos Womersley 1939, Tr. Roy. Soc. S. Austr., 63, (2), 157. 

Guntherama bipygalis, Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 
67, (1), 132. 

Gunthemna param Womersley 1944, Tr. Roy. Soc. S. Austr., 68, (1), 

Plate 99, fig. A-F. 

Description of Nymph. Of typical Trombicula facies. Colour in life red- 
dish. Length 6O0/x, width across propodosoma 360/*, across hysterosoma 420/x.. 
Crista linear with very wide, 101/*, and transverse sensillary area which has 
only a slight posterior angulation ; sensillae bases 90/* apart, and the filamentous 
distally finely ciliated sensillae to ca. 150/* long. Apex of crista forked, epistome 
somewhat rectangular with the anterior margin concave and finely denticulate, 
with 1 seta. Eyes absent. Chelicerae finely serrate on inner margin. Palpi 
fairly stout, tibia with a pair of accessory spines at base of claw. Legs rather 
short, T the longest and strongest to 540/i long, II 390/*, III 390/x, TV 450/*; 



374- Records of the S.A. Museum 

tarsi I thick and slightly tapering, 144/4 long hy 90/i high, metatarsi I without 
any dorsal subapical process ; both tarsi and metatarsi I with numerous slender 
sensory rods amidst the ciliated setae. Tarsal claws stout and unequal. Dorsal 
setae slightly tapering, from 36-75/4 long, with outstanding ciliations. Genital 
discs 2 + 2. 

Description of Adult 9 . Shape and colouration as in nymph. Length 
1,200/*, width across propodosoma 630/4, across hysterosoma 750/*. Crista as in 
nymph but 180ft long, with sensillary area 158/*. wide, and sensillae bases 136/t 
apart, sensillae ca. 200/* long, ciliated distally. Eyes absent. Chelieerae ser- 
rate on inner margin, Epistome as in nymph. Palpi with 3 accessory spines 
at base of claw, and another midway between base of claw and articulation of 
palpal tarsus. Precoxal plates of leg I as in nymph but more chitinized. 
Legs I 900/*, II 660/4, III 660/*, IV 930/*; tarsi I 240/2 long by 130/4 high; meta- 
tarsi I 130/4 long. Dorsal setae as in nymph, 36-72/a. Genital discs 3 + 3. 

Loc. The nymph is described from two specimens reared from larvae by 
Major G. M. Kohls, in New Guinea, 1944 (No. 253). A third and fourth nymph 
reared from larvae \oi what were thought to be Ascoschongastia wamersleyi 
Gunther, by Major Kohls (Dobodura, New Guinea, 11th May, 1944, No. 293) 
is really this species, and the identification of the larvae must have been in 
error. 

The adult description is from a female from New Guinea, 1944 (G. M. 
Kohls No. 83) from which the larvae were obtained which were described by 
Woniersley, 1944 as Guntkermia parana n. sp. G. parana is now regarded as 
con-specific with ftipygalis. 

Remark*. The eggs of the larvae described as parana were not observed 
and no further information is therefore available as to their peculiar structure 
and location on the hairs of the host as described by Gunther 1939 (Proc, Linn. 
Soe. New South Wales, 64, (5-6), 471) for bipygatis. 

Guntherana tindalei (Womersley 1936). 

Trombicula tindalei Womersley 1936, J. Linn. Soc. London (Zool.), 40, (269), 
110. 

Plate 99, fig. G-L. 

Although the larva of this species is unknown the unique nymph is 
unhesitatingly placed in Gwrriherana on the characters of the very widely 
transverse sensillary area, the absence of any subapical process on the first 
tarsi, and the precoxal plates of leg L It may be distinguished as given in the 
key, although it is certainly not very different from the genotype. A re- 
description of the nymph is as follows: 



Womersley— Asiatic-Pacific Scrub Typhus Mites 375 

Re-description of Nymph. Shape as in typical Trombicida. Colour in 
life whitish. Length 1,180/4; width across propodosoma 400ft, across hystero- 
soma 700/4. Crista linear, 120/4, sensillary area broad 84/4 with only a slight 
posterior angle, and paired filamentous sensillae 170/*- long and distally with 
sparse outstanding ciliations; sensillae bases 75/t apart. Eyes absent, Cheli- 
eerae with finely serrate inner margin. Epistome somewhat rectangular, with 
the anterior margin lightly concave and denticulate; with 1 seta. Palpi 
slender, tibia with 2 stout accessory spines at base of the slender claw. Legs 
shorter than body, I thicker than II-IV ; tarsi I tapering anteriorly, 140/4 long 
by 58/4 high, metatarsi I 85/4 long; all tarsi with paired unequal claws; tarsi 
and metatarsi I with numerous slender rod-like setae in addition to the ciliated 
setae. Precoxal plates of leg I well chitinized, and barely conjoined medially, 
without a distinct sternum. Dorsal setae long, fine and slender, with out- 
standing ciliations, length from 30-100/*. Genital discs 2 pairs, 

Loc. A single specimen from under a log at the Ravine, Flinders Chase, 
Kangaroo Island, South Australia, Dec, 1934 (N. B. Tindale). 

Remarks. Upon remounting it is found that the transverse bar with 7 
or 8 divisions described originally as lying at the posterior end of the crista, is 
not part of the crista, but probably some food-body lying within the gut. Re- 
mounting shows it not to be attached to the crista in any way, and it has now 
been moved in position. 

Guntherana TRANSLUCENT (Womersley 1944). 
Tromhimda tramlucens Womersley 1944, Tr, Roy. Soc. S. Austr., 68, (1), 83. 

Plate 100, fig. A-F. 

This species was described from two specimens from moss, from Mt. Arden, 
South Australia, collected by H. M. Cooper, 1943. 

On the structure of the sensillary area it is now placed in Gimtlierana. 

It is a rather characteristic species in that the longest dorsal setae form a 
very distinct posterior fringe. The colour in life is a translucent white. 

The details of the species are as follows : 

Adult. Length 850/4; width across propodosoma 425/*., across hysterosoma 
510/x. Crista 109/4 long, with triangular sensillary area with posterior apex; 
sensillae filamentous, to 75/4 long, ciliated distally, their bases 50/4 apart. Palpi 
with 3 accessory spines at base of tibia claw. Epistome transversely rectangu- 
lar, with fine denticulation and 2 setae, Chelicerae finely serrate on inner mar- 
gin. Eyes absent. Legs all shorter than body with I longer than others and tarsi I 
elongate and slightly tapering towards apex; 136/4 long by 68/4 high, ratio 
length to height — 2-0:1-0; metatarsus I 78/4 long, ratio length tarsus I to 



376 Records of the S.A, Museum 

metatarsus I =-1-7: 1-0- Dorsal setae strongly ciliated, tapering, on propodo- 
soma 13/a long, on hysterosoma and anteriorly 13/x, posteriorly 65-70/1 and form- 
ing a fringe. Precoxal plates of eoxae I present. 

Genus SCHONGASTIA Oudemans 1910. 

Entom. Bei\, 3, (54) , 86-S7; 1912, ZooL Jahrb., Suppl. 14. 

Genotype ThrambitMam vandersandei Ouds. 1905, Nova Guinea, 
5, (larvae). 

(Including Ascoschongastia Ewing 1946, type NeosehdngasUa* malayensis Gater 
1932). 

Key to the Known Adult and/or Nymphal Species, 

1. Sensillae distinctly but narrowly clavate or lanceolate . . . . 2 
Sensillae filamentous, but sometimes fairly but uniformly thick . . 5 

2. Sensillae distinctly but narrowly clavate with short setules. Dor- 
sally the posterior setae to 30/a with stout setules. Front tarsi 2£ 
times as long as high, and ca. half as long again as metatarsi. 
(Adult) i Schongastia cttgerica (Andre 1932 ) , 

Sensillae narrowly clavate, but without distinct basal stem, gradually 
widening from base, and widest at ca, -} from apex. Front tarsi ca, 
twice as long as high. Dorsal setae to 25/x posteriorly. 

Schongastia (Ascoschongmtia) auttyisp.Q- 

Sensillae lanceolate with a short thin basal stem, then thicker until 
near tip, where they taper 3 

3. Front metatarsi small; slightly more than half the height of front 
tarsi, Sensillae fairly wide with moderately long ciliations. Dorsal 
setae uniform, pectinate, to 20/a. (Nymph). 

Schongastia (Ascoschongastia) kohlsi (Philip and Woodward 1946). 

Front metatarsi as high or almost as high as tarsi . . , . , . 4 

4. In each angle in front of sensillary area a ciliated seta twice the 
length of neighbouring setae. Sensillae with fairly long setules. 
Tarsi I ca. twice as long as high and one-fourth again longer than 
metatarsi. Dorsal setae fairly uniform, to 30/a. (Adult). 

Schongastia {Ascoschongastia) indica Hirst 1915. 
(= Trombicula muris Waleh 1932). 

No such seta in the angles in front of sensillary area. Stem of sen- 
sillae very fine and body not so wide as in indiea. Tarsi I ca. two-thirds 
as long again as metatarsi, and rather more than twice as long as high. 
Dorsal setae uniform to 20/x. (Nymph). 

Schongastia (Ascoschongastia) lamas (Radford 1946). 

5. Sensillae fine and nude 6 

Sensillae with barbs for entire length, or only basally, and then with 
ciliations distally, or with outstanding ciliations for "the whole length 7 



Womersley— Asiatic-Pacific Scrub Typhus Mites 377 

6. Posterioi* dorsal setae with a subapical setule longer than the other 
setules and as long as or longer than the setae themselves. Dorsal 
setae from 20 to 30ft long, excluding the subapical setule. Tarsi I 
1-8 times as long as high and only slightly longer than metatarsi I. 
(Nymph)* Schongastia. (ScMngastia) Oudemansi (Walch 1922). 

(= impar Gunther 1939 — bodcmis Gunther 1940. 
=muris Walch 1927 nee oudemansi Walch 1927). 

Dorsal setae pectinate without above subapical setule, with outstanding 
setules, to 30/i. long. Tarsi I twice as long as high, metatarsi slightly 
longer than tarsi are high. (Nymph). 

Schongastia (SchbngaMa) maldivienm'R&dioT&19ifr 

Dorsal setae pectinate without above subapical setules, with outstand- 
ing setules of moderate length to 56^ long. Tarsi I 1*64 times as long 
as high and metatarsi I not as long as tarsi I are high. (Nymph). 

Schongastia (Schongastia) irieia Gater 1938. 

7. Sensillae with only barbs for their entire length. Dorsal setae 15 to 
25/x long. Tarsi 11-7 times as long as high, and 1-3 times as long as 
metatarsi. (Nymph) (after Walch). 

Schongasiici {Schongastia) schuffmri (Walch 1922). 

Sensillae with outstanding ciliations for the whole length. DS poster- 
iorly to 16/*. Front tarsi 1-7 times as long as high and one-sixth as 
long again as metatarsi. 

Schongastia {Ascosclidngastia) malayensis Gater 1932. 

Sensillae ciliated distally, basally with or without barbs . . - . 8 

8. Sensillae very fine , . . . 9 

Sensillae fairly and more or less uniformly thick with short ciliations 

or barbs basally and long ciliations distally 10 

9. With a long nude seta in each angle in front of sensillary area. Sen- 
sillae minutely barbed basally and with 8-10 long ciliations distally. 
Dorsally setae from 20-30^ posteriorly. Tarsi I twice as long as high 
and twice as long as metatarsi. 

Schongastia ( Schongastia) , ocuAicola sp. n. 

Without such seta in angles of crista. Seusillae distally with many 
short ciliations. Dorsal setae from 25/x to 90/t posteriorly. Tarsi 12-2 
times as long as high and 1-33 times as long as metatarsi, 

Schongastia westraliensis (Worn. 1934) . 

10. Posterior dorsal setae to 96//, long and ending in a terminal setule 
ea. one-fifth entire length. Tarsi I 1-75 times as long as high and 
1-6 times as long as metatarsi. (NjTnph.) 

Schongastia (Ascoschongai'tia) nmtabilis (Gater 1932). 

Posterior dorsal setae to 50ft long, tapering with normal outstand- 
ing setules, without especially long terminal setules. Ta^si I nearly 
twice as long as high, and 1-17 times as long as metatarsi. (Nymph.) 

Schongastia (Ascoschdngastia) nadchatrami sp. n. 



378 Records of the S.A. Museum 

Schongastia (Ascoschongastia) indica Hirst 1915. 

Schongastia indica Hirst 1915 > Bull. Entom. Research, 6, 183. 

Trombicula mum Walch 1922, Kitasato Archiv. Exper. Med., 5, (3) ; nan 
muris Walch 1923 (24), Trans. Vth. Bien. Cong. Far East. Assoc. Trop. 
Med., (Singapore), 621, fig. 31-33. 

Tromhicula oudemansi Walch 1923 (24), Trans. Vth. Bien. Cong. Far East. 
Assoc. Trop. Med., 623, fig. 34r-37; non oudemami Walch 1922, Kitasato 
Archiv. Exper. Med., 5 ? (3) j non oudemmsi, Fletcher, Lesslar and Lewth- 
waite, 1928, Trans. Roy. Soc. Trop. Med. & Hyg., 22, 161. 

Schongastia indica, Walch 1927 Geneesk. Tijds. v. Ned. Indie, 67, (6), 924. 

Neoschongastia indica, Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 
67, (1), 118; Radford 1946, Parasitol., 37, (1-2), 51, fig. 26 a-e ; Wharton 
1946, Science, 104, (2,691), 76-77. 

Ascoschrdngastia indim, Wharton 1946, Ecol. Monog. 16, (3), 153-184. 

Plate 100, fig. G-K. 

The adults and nymphs of this species are well known. Walch was the 
first worker to succeed in rearing nymphs from larvae, but unfortunately, the 
difficulty of ensuring that the larvae are only of the one species before rearing, 
and thus establishing true correlation of nymphs with larvae led him to be 
mistaken in his resulting conclusions. More careful rearing of larvae to 
nymphs and adults, and of larvae from eggs laid by known adults by other 
workers, notably C. D. Radford, G. M. Kohls, and in particular carefully con- 
ducted research by G. W. Wharton 1946, has definitely established all the 
different stages, egg f larva, nymphochrysalis, nymph, imagochrysalis, and adult 
of both sexes, in this species. 

Walch in his experiments, in all probability had larvae of two species in 
the tubes, viz. indium Hirgt (— muris Walch) and oudenum&i Walch. In 1946 T 
the nymph of indica was described by Radford from larvae from Addu Attol, 
Maldive Islands, from which locality oudemansi is unknown. Thus, it was seen, 
that the form of the sensillae as figured by Walch for oudenumsi are as iu 
indica and his figures for muris (indica) are really applicable to oudemam*si+ 
More recently Wharton (1946) working on this species at Guam, Mariana Is.,- 
has not only established all the stages as stated above but has described the 
external morphology in a very detailed manner. Further he has by experi- 
ment and careful observation contributed very greatly, not only to our know- 
ledge of the bionomics of this particular species, but inf erentially also to that of 
the Trombiculidae as a whole. 



WOMERSLEY — ASIATIC-PACIFIC SCRUB TYPHUS MlTES 379 

The descriptions of the adult and nymph from the specimens examined, one 
adult and one nymph from the Maldives, and 5 nymphs reared by Major G. M. 
Xohls from larvae found on Dactytiopsila sp., from Dobodura, New Guinea, 
1944 (No. 261) are as follows: 

Adult. SB 42/*, Crista 90/* long, ratio crista to SB •= 2-14. Sensillae 
96/*. Tarsi I 109/* by 54/*, ratio length, height — 2«0; metatarsus I 88/* long, 
ratio tarsus I, metatarsus I = 1*24, Dorsal setae fairly uniform at 18/* to 25/* 
long. 

Nymph. SB 30/*. Crista 85/* long (Maldives 75/*), (Radford 93/*), ratio 
crista to SB — 2-8. Sensillae 75/* (Maldives 55/*), (Eadford 68^). Tarsi I 
4i0/* by 30/*. 

This species is rather small, reddish in life, and of the usual figure of eight 
form. Crista rather short with the subposterior sensillary area triangular with 
posterior apex, and a pair of lightly clavate or spathulate sensillae with 
setules; the epistome is rounded with fine teeth and 1 long ciliated Beta. Palpi 
somewhat slender ; tibia with the usual 3 accessory spines at base of claw, and 
1 other about midway between base of claw and articulation of the palpal tarsus, 
all on the inner side of palpi. Cheli^erae finely serrate on inner margin. Eyes 
«quite absent. The sternum is entire, roughly pentagonal, without any evidence 
of individual preeoxal plates. The front tarsi are from 2 to 2*5 times as long as 
high, widest proximally, then slightly tapering. Dorsal setae short, fairly uni- 
form, with long outstanding setules the whole length. In the male sex (accord- 
ing to G. W. Wharton 1946) on the genitalia on each side and arising between 
the second and third genital discs is a long curved ciliated and specialized seta. 

BchSngasiia (Ascoschongastia) Mica* is apparently a widely distributed 
species, latvae having been recorded from India (Hirst), Ceylon (Radford), 
Maldives (Radford). Burma (Audy), Sumatra (Walch), Malaya (Gater), New 
-Guinea (Womersley and Heaslip, Gunther and others), Australia (Womersley 
and Heaslip). and it occurs also in the Philippines, The nymphs and adults 
have been reared by Walch, Radford, Kohls, Wharton and Carver, and Jaye- 
wickreme. In the field, however, adults have only rarely been found, possibly 
due to their association with tree-climbing rats, for adults have been taken in 
the debris of rats 1 nests in the tops of coconut palms (Radford). 

Sohongastia algerica (Andre 1932). 

Thrombkula algerim AndrS 1932, Bull. Soc. Zool. Prance, 59, 284-88. 

Plate 100, tig. L-O. 

In 1932 Dr. Mare Andre described and figured the adult of ThromMctda 
■algerica from date palms in Algeria. Through his very great kindness 1 have 



380 Records of the S.A. Museum 

been able to study the type specimen and to compare it with adults of indica 
from Ceylon. 

Tliis species is very close to, and may possibly be synonymous with indica, 
in the short rather thickened to clavate sensiilae, the form of the crista, of the 
epistonie, and in leg dimensions and the length and form of the dorsal setae. 
The main points of difference are that the sensillae are rather more definitely 
clavate and with shorter eiliations than in indica, that the specialized seta on 
each side of the crista in front of sensillary area is absent, and that the ratio 
of length to height of the front tarsi is somewhat greater. 

The data as given by Andre are : 

Length 1,030/*, width 405-470/*. Dorsal setae 20-30/1 long. Leg I 540^ r 
tarsi 150/a long by 60/t high, metatarsi UO/a long. Tibia with 3 accessory spines. 
at base of claw and articulation of palpal tarsus, Epistome as in indwa. In 
Sig Thor and Willnianns 7 recent work (aDs Tierreich Lfg. 71b, 1947), the 
palpal tibiae are stated as "median hinter der Kralle 4 Dornen. M This includes 
the 3 spines at base of claw and one further away as stated above. 

SCHONGASTIA. (SCHONGASTIA) OUDEMANSI (Walch 1922). 

Trombicida oudemansi Walch 1922, Kitasato Archiv. Exper. Med., 5, (3), 
(larva) ; non Waleh 1924, Trans. Vth. Bien. Congr. Par East. Assoc. Trop. 
Med. (Singapore), 623 (nymph). 

Trombicula nniris Walch 1924, Trans. Vth. Bien. Congr. Far East. Assoc. Trop. 
Med. (Singapore), 621 (nymph) \ non Walch 1922, Kitasato Archiv. Exper. 
Med., 5, (3), (larva). 

Schdngastia indica Hirst 1915, Bull. Entom. Res., 6, 183 (larva) • Walch 
1927, Geneesk. Tijds. v. Med. Indie, 67, (6), 924, (larva). 

Neosclionga&tia impar Gunther 1939, Proc. Linn. Soc. New South Wales, 64, 
(1-2), 85, (larva) ; Womersley and Heaslip 1943, Tr, Roy. Soc. S. Austr. T 
67, (1), 116 (larva). 

Neoschongastia bod&ms Gunther 1940, Proc. Linn. Soc. New South Wales, 65, 
(5-6), 482, (larva). 

Plate 101, fig. F-L. 

This species was first reared to the nymph by Walch, but unfortunately 
he was apparent^ misled in his association of the larvae with the correspond- 
ing nymph (see remarks under Ascoschongastia indica) and his description and 
figures of oudemmsi nymph refer and correspond to indica (muris) and vice 
versa. The species has more recently been reared to the nymph by Major G. M. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 381 

Kohls from larvae oii rats, at Dobodura, New Guinea (25 April, 1944, No, 253), 
which larvae were determined as impar Gunther. As shown under the discus- 
sion of the larvae, however, the chelicerae of both impar and hodensis are ser- 
rate, and these two species are both synonymous with oudemansi Walch. 

The description of the nymph from 4 specimens received from Msgor 
Kohls is as follows : 

Description of Nymph. Length 470/*, width across propodosoma 240>*, 
across hysterosoma 270/*. Crista typical of Schongastia, to 90/* long, with tri- 
angular sensillary area, and paired nude filamentous sensillae to 120/* long and 
bases 40/i apart. Eyes quite absent. Chelicerae finely serrated on inner mar- 
gin. Palpal tibia with 2 accessory spines at base of claw. Legs all shorter than 
body, with I the longest and strongest; I 400^ long, II 260/*, III 280**; IV 320/x; 
tarsus I 90/i long by 50/* high, ratio length to height 1-8 ,- metatarsus I 80/*, ratio 
length tarsus I to metatarsus I = 1-12. Dorsal setae fairly uniform, to 30/* 
long, with strong outstanding setules, one of which at the apex is exceptionally 
long, and as long as the posterior setae themselves; in Walch '& fig. 36 (1924) 
which appears may be correctly given as oudcmtmsi, a subapical seta is shown 
as slightly longer than the others, but otherwise the above characteristic 
appears to have been overlooked by Walch. 

Loa This species is known from Deli, Sumatra (Walch), New Guinea 
(Gunther, Kohls), and Borneo (Gunther). 

Schongastia ( Ascoschongastta) maldivtensis (Radford 1946). 

Schongastia maldivmnsis 1946, Parasitology, 37, (1-2), 48 (larva), 51, (nymph). 

Plate 102, fig. A-L. 

Radford's description of the larvae and more particularly of the nymph 
of this species are unfortunately rather inadequate although the figures are 
more satisfactory. The species was described from a lizard and also from 
rats from Addu Attol, Maldive Islands, The following remarks and data are 
from a nymph reared from larvae and kindly presented to me by Dr. Radford- 
The crista as seen, and as figured by Radford has the typical triangular form of 
the genus Schongastia. The sensillae are about as long as the crista (given as 
93/*), and are nude with their bases 30;* apart. The epistome is rounded, with 
fine denticulations, and one seta. The chelicerae are finely serrate on the inner 
margin. Eyes absent. Palpi rather slender; tibia with 2 accessory spines at 
base of claw. The dorsal setae are short, to 25/* thickest medially, and with 
setules. In the specimen examined, the crista is 59/* long, SB 26/* and the 
posterior dorsal setae to 30/*, slender, not thickened medially as in Radford's 



382 Records of the S.A. Museum 

figure, with outstanding setules, Tarsi I 90/t long by 45/* high, metatarsi I 52/* 
long. 

Loc. So far, only known from Addu Attol, Maldive Islands. A solitary 
adult collected by S. L. Kalra, at Gudalur, India, 23 Aug. 1947 has been kindly 
sent to me for study by Dr. C. D. Radford. This specimen appears to be the 
adult of Radford's rmldiviensis, but further material of both adults and larvae 
from the same locality are required to put this beyond question. The specimen, 
however, is described and figured herewith, and the figures may be compared 
with those given of a nymph from the Maldives. 

Description of the Adult $ , Length 850/* ; width across propodosoma 
450/*, across hysterosoma 520/*. Colour in life unknown. Crista elongate, 182/*, 
with a triangular posterior sensillary area, with fine, nude, filamentous sen- 
sillae, ca. 100/* log, and their bases 58/* apart. Epistome rounded-conical with 
strong denticulations, and 1 ciliated seta. Eyes quite absent. Palpi stout, with 
stout claw with 3 accessory spines at base of claw and another midway on 
outer margin of tibia; tarsus widest medially, and not reaching tip of claw* 
Legs I stronger and longer than rest, 715/*, II and III 455/*, IV 590/*; tarsi I 
182/* long by 104/* wide, metatarsus I 130/i long. No precoxal plates on coxae I 
and with enclosed sternum. Dorsal setae fairly uniform, with strong ciliations 
.and posteriorly reaching 40/* in length. Genital aperture with 3 pairs of discs. 

ScHONGASTIA (SchSNGABTIA) SCH0FFNERI (Walch 1923). 

Trombicula schwffneri Walch 1923, Kitasato Archiv. Exper. Med., 5, (3), 
(larva); 1924, Trans.. Vth. Bien. Congr. Far East. Assoc. Trop. Med., 
(Singapore), (nymph); 1927, Geneesk. Tijds. v. Neder. Indie, 64, (3), 
(nymph). 

Neo&chonffa&tia sehuffneri, Womersley and Heaslip 1943, Tr, Roy. Soc. S. Austr., 
67, (1), 117 (larva). 

Plate 101, fig. M-O. 

The nymph is still only known from those reared by Walch, and the 
following data and the figures are from his studies. 

Nymph. Length 426/t, width across propodosoma 202/*, across hysterosoma 
212/x. Colour in life orange red. Dorsal setae colourless, more or less tapering, 
eiliated, from 15 to 25ft long. Crista 52/* long, with sensillae 111/* long and 
with minute barbs basally, slightly longer distally, sensillae bases 25/* apart. 
Palpal tibia with 2 accessory spines at base of claw. Eyes absent. Epistome 
rounded with fine denticulations and 1 seta. Chelicerae serrate on inner mar- 
gin. Legs all shorter than body, I 273/* long and stronger than others, II 147**, 



Womersley — Asiatic-Pacific Scrub Typhus Mites 383 

in 132/*, IV 177ft; tarsus I 1*7 times as long: as high, and 1-3 times as long 
as metatarsus I. Sternum entire, without trace of precoxal plates. 

In the form of the sensillary area this species belongs to Schongastia. It 
can be recognized from other known adults or nymphs by the characteristically 
barbed sensillae as in the key. 

Schongastia WESTRAUEN3IS (Womersley 1934). 

Microtrombidium westraliense Womersley 1934, Rec. S. Aust. Mus., 5, (2), 188. 

Plate 104, fig. A-F, 

As this species has the characteristic facies of the Trombiculidae and a 
triangular sensillary area of the -crista it is now placed in the genus Schongastia 
Ouds. It is still only known from the unique specimen from under a stone, 
Mundaring Weir, Western Australia, August 9, 1931 (H.W.). 

The species is separated as in the key and a re-description is as follows : 
Adult. Of typical Trombiculicl facies. Size moderate, length to 1-2 mm.; 
width across propodosoma 800/*, across hysterosoma 930/*. Crista 165/* long, 
with subposterior triangular sensillary area with posterior apex, and carrying 
a pair of filamentous sensillae 120/* long, their bases 48/* apart, and ciliated 
distally. Epistome rounded with fine denticulations and 1 seta. Eyes absent. 
Chelicerae finely serrate on inner edge. Palpi moderately stout; tibia with 3 
accessory spines near base of claw, and one other more basad. Sternum entire, 
without trace of preeoxal plates. Legs all shorter than body, I the longest and 
strongest, 840/* long, II 530/*, III 530/*, IV 670/* ; tarsi I elongate, 200/* long by 
90/* high, ratio = 2*22; metatarsus I 150/* long, ratio length tarsus L metatarsus 
I « l ■ 33, Dorsal setae tapering, 25/* anteriorly gradually increasing to 90/*. 
Colour in life reddish, 

Schongastia (Schongastia) oculicola sp- n, 

Plate 103, fig. G-L, 

Description of Nymph, More elongate but otherwise of typical Trombicu- 
lid facies. Length of newly-emerged specimens to 580/*; width across propo- 
dosoma 342/*, greater than across hysteromosa 260/*. Crista linear 70/* long, 
with triangular sensillary area enclosing a dumb-bell shaped areola, and fur- 
nished with filamentous sensillae, 75/* long, basally with minute indistinct 
barbs and distally with 8-10 long branches j sensillae bases 31/*. apart ; on each 
side of the stem of the crista is a single long seta, nude or apparently nude. 
Epistome not visible in preparations but with 1 ciliated seta (in one specimen 
2 setae). Eyes entirely absent. Palpi stout; tibia with strong claw flanked at 



384 Records of the S.A. Museum 

its base by two accessory spines which are almost as large, long and strong as 
the claw ; a third strong spine midway between base of claw and articulation of 
tibia; chelicerae with fine inner serrations. Legs I the longest and strongest, 350/* 
long, II 220,*, III 220,*, IV 260/*; tarsi I 91/* long by 45-5/* high, metatarsi I 
45*5/* long. Sternum entire; no precoxal plates. Dorsal setae uniform, 20-22/* 
long, tapering with few long outstanding setules. Genital discs 2 pairs. 

Remarks. The above description is from 4 specimens reared by Mr, S. H. 
Jayewickreme from larvae from the conjunctival sacs of Leggada hooduga fiil- 
mdiventris from Nalanda, Ceylon, July 1944. The correlation was cheeked 
against recovered larval pelts. 

Schongastia (Ascoschongastia) MALAYENsrs (Gater 1932). 

NeoscJwngastia malayensis Gater 1932, Parasitology, 24; Womersley and Heas- 
lip 1943, Tr. Eoy. Soc. S. Austr., 67, (1), (as larvae). 

Plate 104, fig. G-J. 

Description of Nymph. Of typical Trombicula fades. Colour in life ? 
Length to 416/*, width across propodosoma 260/a, across hysteromosa 286/*. 
Crista linear, 84/* long, with subposterior triangular sensillary area with the 
apex posteriad ; sensillae to 56/* long with outstanding ciliations for the whole 
length, and the bases 42/* apart; epistome rounded conical, dentate with seta 
16/* long. Byes absent. Chelicerae with finely serrate dorsal edge. Palpi 
fairly stout; tibia with single claw and two strong accessory spines at base. Legs : 
I stronger than rest to 325/* long, II 234/a, III 234/*, IV 260/*; claws normal; 
precoxal plates absent; tarsus I 70/* long by 42/* high, metatarsus I 61-6/* 
long. Dorsal setae uniform, 16/* long, tapering, with outstanding setules. Geni- 
tal discs 2 + 2. 

Loc. Reared by Dr. Audy and his colleagues from larvae on Rattus canus 
from Kepong Forest Reserve, and Bk. Langan, Selangor, F.M.S., March, 1950, 

Remarks. As with other species of the genus, &. (A.) malayensis in the 
nymph can only be separated on somewhat intangible features as in the key. 

Sohongastia (Ascoschongabtia) lanius (Radford 1946). 

Neoschongastia lanius Radford 1946, Proc. ZooL Soc. London, 116, (2), 262 ? 
fig. 23-24 (as larvae). 

Plate 102, fig. M-P. 

Description of Nymph. Unfed, length to 780/*, width across propodosoma 
383/*, across hysterosoma 412*t. Colour in life ? Crista elongate, 118/* long 
with subposterior, laterally rounded, sensillary area; with SB 32*/, and sen- 



Womersley— Asiatic-Pacific Scrub Typhus Mites 385 

sillae very thin basally then thickening until near the tip where they taper to a 
point, to 100/t long and furnished with short sparse filiations; epistome more 
or less conical, very indistinct and dentate, epistomal seta 22/* long. Eyes 
absent. Chelicerae finely serrate on inner margin. Palpi as figured, with 2 
accessory strong spines at base of claw and a strong seta on outer side; the 
tip of the palpal tarsus with a nude S-shaped seta, and the sensory rod on the 
outer side unusually long. Legs I 486/* long, II 284/*, III 312/i IV 355/*; tarsi I 
102/*. long by 48/* high, metatarsi I 64/*. Dorsal setae pectinate, short, poster- 
iorly to 22/t, with sparse strong ciliations. 

Remarks. A number of nymphs have been reared by Mr. K. L. Cookings, 
from larvae identified as lanius Radford and the identification checked against 
the recovered larval pelt. The larvae were from Rattus raMus bnmnemcuhis 
from Imphal, Burma, 1945. 

Schongastia (Ascosciiongastia) kohlsi (Philip and Woodward 1946). 

Neoschongastia hohlsi Philip and Woodward 1946, Amer. J. Trop. Med., 2($> 
(2), 159, fig. B. and D. (as larvae). 

Plate 101, fig. A-E. 

As reported in the Larval Section of this paper, the larvae of this species 
ia common in the Imphal district of Burma, and I am indebted to Lt.-Col. J. R. 
Audy and his colleague, K. L. Cockings, for the opportunity of describing here 
the nymph, a number of specimens of which Mr. Cockings was successful in 
rearing. As with other groups of Trombieulidae the specific characters of the 
nymphs (and adults) arc anything but easy to determine. It is believed, how- 
ever,, that this and other species may be separated as in the key. 

Description of Nymph. Facies as in typical Tromliml-a. Colour in life > 
Length (newly emerged) 780/4 ; width across propodosoma 385/t,, across hystero- 
soma 397/x. Crista elongate, length excluding epistome 96/*., with epistome 
118/*; with typical shape of sensillary area, with SB 22u, and sensiUae 70/* 
long, broadened and lanceolate from ca- one-quarter from base, and tapering to a 
point apically and with short barbs basally and longer, but short ciliations on 
body. Epistome rounded conical, dentate, with one seta 18/* long. Eyes 
absent. Chelicerae with finely serrate inner margin. Palpi normal, tibia with 
two strong accessory stout spines at base of claw, and another strong spine on 
outer surface; tarsi with only 1 nude apical spine, and the. sensory rod on 
outer surface longer than usual. Legs I longer and stronger than rest, 397/*, 
II 280/l, III 280^, IV 355/*; tarsi I 112/* lon£ by 58/t high, metatarsi I 64/* long, 
and not more than half the height of tarsi wide. Dorsal setae uniform, pectin- 
ate as figured with strong setules, to 20a long posteriorly. 



386 Records of the S.A. Museum 

Schongastia (Ascoschongastia) mutabilis (Gater 1932). 

Neoschongastia mutabilis Gater 1932, Parasitology, 24; Womersley and Heaslip 
1943, Tr. Itoy. Soc. S. Austr., 67, (1), 111, pi. VII, fig. 5 (as larvae). 

Plate 103, fig. A-F, 

This species is so far only known from a single nymph reared by Mr. K. h. 
Cockings from what was thought to be a larva of tanius Radford, and from a 
rat from Imphal, 22/12/1945. The pelt, as far as the scutum and palpal setae 
are seen, appears to be that of 8, (A.) nwtabiMs (Gater), and the nymph as 
in the following description is very different from a number of other reared 
nymphs from lani-us larvae. 

Description of Nymph. Of typical Trombiculu faciei Length 640/* ; width 
across propodosonia 326/*, across hysterosoma 355/*. Crista elongate with sub- 
posterior sensillary area, length excluding epistome 86/*, with epistome 112**; 
epistome rounded -conical with 1 ciliated seta 25/* long, and dentate margin; 
sensillae thick with parallel sides, short barbs basally and long setules distally, 
to 80/* long and bases 32/* apart. Eyes absent. Chelicerae with finely serrate 
inner margin. Palpi with 2 strong accessory spines or claws at base of claw, 
and a nude spine on outer surface ; sensory rod on outer surface of tarsi rather 
long, apical spine single. Dorsal setae of two forms, shorter anterior setae 
25/* long, with strong setules and ending in a long fine setule, about one-fifth 
of total length. Genital discs two pairs. Legs I longer and stronger than 
rest, 360**, II 255/*, III 255**, IV 310/*; tarsi I 99** long by 57/* high, metatarsi I 
61/i long. 

Schongastia (Ascoschongastia) audyi sp. n. 

Plate 105, fig. F-I. 

Description of Nymph. Of typical Trombicula facies. Length to 405**; 
width across propodosoma to 195**,, across hysterosoma to 260/1. Crista elon- 
gate, 70/* long, with subposterior, roughly triangular sensillary area, with paired 
narrowly clavate or lanceolate sensillae furnished with short setules; sensillae 
bases 28/* apart. Epifitome probably rounded-conical and dentate; epistomal 
seta 20/i long. Eyes absent. Chelicerae serrate on inner edge. Palpi small, 
tibia with simple claw and two accessory spines at base of claw. Dorsal setae 
short and rather sparse, mainly 15/* long, but on extreme posterior to 25/*; 
with rather prominent outstanding setules. Leg I longer and stronger than 
rest, 305/* long, II and HI 175/*, IV 227/i; tarsi I as figured, 70/* long by 33/* 
high ; metatarsi I 42/* long and as high as tarsi I. No precoxal plates on leg I, 
hut sternum enclosed posteriorly. Genital discs 2 pairs. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 387 

Loq. Described from 6 nymphs reared by Mr. K. L. Cookings in Malaya, 
from larvae from Callosciwus mgrovittatus and C. notatus, from Bukit Lagong 
Forest Reserve, Kepong, Malaya 1950. The nymphs were correlated with the 
larval pelts. 

Remarks. Can be distinguished from other n3 r mphal species as in the key. 

Schongastia (Schongastia) vieta Gater 1932* 

SoJwngastia vieta Gater 1932, Parasitology, 24, 154; Womersley and Heaalip 
1943, Tr. Roy. Soc. S. Austr., 67, (1). 103; Sig Thor and Willmann 1947, 
Das Tierreich, 71b, 308. (As larvae). 

Plate 105, fig. A-E. 

DescriptiQii of Nymph. Of typical Trombwnla facies. Length 360/i; width 
across propodosoma 208/*., across hysterosoma 266/a. Crista elongate, to 98/i 
(excL epistome) with subposterior, roughly triangular sensillary area; with 
paired long and apparently entirely nude sensillae to 112ft long, their bases 
36/* apart. Epistome rounded-conical, dentate, with epistomal seta 56/*, long. 
Eyes absent. Chelieerae with finely serrate inner edge. Palpi with simple 
tibial claw and two accessory spines at its base. Dorsal setae gradually increas- 
ing in length posteriorly to 56//. long, not very tapering, with long outstanding 
setules. Legs I longer and stronger than the rest, to 312/t, II and III 195/x, 
IV 260/*; tarsi I ovoid, 78-4/x long by 47-6//. high; metatarsi I 44-8/x, lon)g. 
Sternum entire, with no precoxal plates on leg T. 

Loc. Described from two nymphs reared by Mr. K. L, Coekings of the Bri- 
tish Scrub typhus Research Team in Malaya from larvae from Battus r. diardt 
from Kuching, Sarawak, 1950. The nymphs were correlated with the larval 
pelts. 

Remarks. As with most species, the nymphs can only be separated from 
others on very intangible characters as in the tentative key given here. 

Schongastia (Ascoschongastia) nadchatrami sp, n. 
Plate 114, fig. D-F. 

Description of Nymph. Of typical Tronibicula facies. Length to 804/t, 
width across propodosoma 402/jl, across hysterosoma 469/*. Crista elongate 196//. 
including epistome, with subposterior sensillary area, sensillae bases 60/t apart ; 
sensillae 100//,, ciliated distally; epistome dentate and rounded-conical with 
one ciliated seta. Eyes absent* Chelieerae finely serrate on inner margin. 
Palpi normal, with single tibial claw and a pair of accessory spines at base of 
claw. Dorsal setae thickly disposed, gradually lengthening from 23//, anteriorly 



388 Records of the S.A. Museum 

• 
to 50/x posteriorly, with strong eiliations. Legs • I longer and stronger than the 

rest, to 777//,, II 536/i, III 536/a, IV 602/*; tarsi I elongate oval, 161/x long by 
87 fi high, metatarsi I 137/* long ; no precoxal plates. 

Loc. Two nymphs reared by Mr. Nadchatram of the Scrub Typhus Re- 
search Unit, I.M.R. Kuala Lumpur from larvae from Rattus ssabanus from Ulu 
Langat, F.R, Selangor, 1950. The nymphs were checked against the larval 
pelts. 

Genus NEOSOHONQASTIA Ewing 1929. 

NeoscMngastia Ewing 1929. A Manual of External Parasites, 187; "Wharton 
and Hardcastle 1946, J. Parasitol., 32, (3), 286-322. 

Parasckongastia Womersley 1939, Tr. Roy. Soc. S. Austr., 63, 165, 

Genotype Schongastia americana Hirst 1921, 

Ewing erected this genus for those larval species of Schongastia in which 
the chelicerae did not possess teetli on the dorsal (inner) margin. In 1939 the 
genus Para$chbngastia Womersley was separated off for certain species in which 
there was a distinct ridge on the dorsal scutum anterior of the sensillae. The 
type of this genus was Neoselwngastia yeomansi Gunther 1939. It has since 
been shown, however, that A T . yeomansi and N. americana are congeneric, and 
therefore Pamschdngastia is synonymous with Neoschowgastia. For the 
remainder of the species in Ewing *s Neoschongastia the generic name of Asco- 
schongastia has been assigned (Ewing 1946). 

The adult or nymphal stages remained unknown until Wharton and Hard- 
castle 1946, described the nymphs of two species from Guam and Okinawa in 
the Western Pacific. One of these, Neoscbongastia carveri Wharton and Hard- 
castle, was reared from larvae definitely determined as the same species. The 
second one, however, is as yet specifically uncertain as the authors had larvae of 
both If. americana solomonis Wharton and Hardcastle and N. monticola Whar- 
ton and Hardcastle, in the rearing tubes. 

A third species of which the nymphs are known is Neoschongastia- galling 
arum (Kawamura and Yamaguchi 1921) from Formosa (Hatori) and Feder- 
ated Malay States (Fletcher, Lesslar and Lewthwaite). 

A diagnosis of the genus based on the nymphal stages is as follows, after 
Wharton and Hardcastle 1946 : 

Diagnosis of Genus. Nymph. With the facies of Trambicula. Sensillary 
area entire, not divided into sections by longitudinal ridges, and reniform with 
the hilus directed anteriorly. Striae extending over sensillary area. No eyes. 
Six setae on palpal segment IV. Two types of opisthosomal body setae, a long 
and a short. Pseudostigmatie organs or sensillae spathulate, although at times 
almost filiform. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 389 

Key to the Known Nymphal Species. 

1. Sensillae whip-like with thicker stem and fine filamentous lash, nude 
except for indistinct and sparse barbs on stem. Apex of senaillary 
area truncate. Epistome with 5 strong broad teeth and 2 setae. 
Palpal tarsus short, broader and truncate apically. Dorsal setae of 
two lengths, 27 and 50/*. 

Neoschdngastia curveri Wharton and Hardcastle 1946. 

Sensillae narrowly or more widely spathulate with long or short 
eiliations . . . . * . . , 2 

2. Sensillae very narrowly spathulate, with long ciliations. Posterior 
apex of sensillary area pointed. Epijstome with 7 teeth and 1 seta- 
Palpal tarsus of normal shape. Dorsal setae of two lengths 20-23/* 
and 50-70/*. 

Neoschdngastia sp. (either americana sol-omonis Wh. and Hie. 1946, or 

moniiwto Wh. and Hie. 1946. 

Sensillae broadly spathulate or clavate, with short ciliations. Pos- 
terior apex of sensillary area pointed. Epistome rounded and den- 
tate. Dorsal setae, of two kinds, mainly tapering to 16-20/a but inter- 
spersed posteriorly with uniformly thick setae to 45/4,. 

Neo$oKonga$ti(b gallvnurum (Hatori 1920). 

Neoschongastia carveri Wharton and Hardcastle 1946. 

J. Parasitology, 1946, 32, (3), 313. 

Plate 106, fig. E/ftl. 

Description of Nymph (after Wharton and Hardcastle). Shape not quite 
typical of Tronibicula in the unique specimen as the shoulders were indistinct 
Length 850/x; width across propodosoma 370/*, across hysterosonia 350ft. Body 
thickly clothed with ciliated setae, and integument between setal bases striated. 
Crista with somewhat reniform sensillary area with hilus directed anteriorly; 
epistome with 7 strong teeth and two ciliated setae; sensillae 37/4 apart, and 
ea. 100/x long, filamentous, thicker on basal third and with a few minute barbs, 
distal two-thirds lash-like ; cuticle over sensillary area folded and striate. Eyes 
absent. Palpi short and stout ; genu wider than long ; tibia with two accessory 
spines at base of claw.; tarsus short, truncate and widest apically. Chelieerae 
serrated on inner (dorsal) edge. Legs I the longest; tarsus I 147/x long by 50/4. 
wide; metatarsus I 67/* long. Dorsal setae of three types; the anterior setae 
finer than posterior setae which are either long, 50/4, or short, 27/*, arising from 
papillae, and tapering with short ciliations. Genitalia with 2 pairs of discs, 
the anterior pair being oyal and longer than the posterior pair. 



390 Records of the S.A. Museum 

Log. Only known from a single nymph reared from larvae from a reef 
heron Demigretta sacra from Port Ayayan, Guam, July 17, 1945, by B. K. 
Carver. The larval skin was identified and so correlated with the nymph. 
(Wharton and Hardeastle 1946). 

Neqsohongastia sp. Wharton and Hardeastle 1946. 
J. Parasitology, 1946, 32, (3), 320. 

Plate 106, fig; J-M. 

Description of Nymph (after Wharton and Hardeastle 1946). Shape as in 
typical Trombfcula. Length 670-710/x; width across propodosoma 300-390/1, 
across hysterosoma 340-390/*. Body thickly clothed with ciliated setae. Integu- 
ment posteriorly with discs from which the setae arise, with striations between. 
Crista with reniform, striated sensillary area, the posterior angle of which is 
pointed; sensillae short, 60ft approximately, lightly spathulate and with long 
fine ciliations ; their bases 37-43/* apart. Eyes absent. Epistome with 5 strong 
teeth and 1 seta. Chelicerae with the inner (dorsal) edge finely serrated. 
Palpi stout, genu about as long as wide, tibia with two strong accessory spines 
at base of claw; tarsus of normal shape and length. Legs all shorter than 
body, I the longest; tarsi I 120-133/1 long by 50^-60/1. wide, metatarsus I 77/* 
to 96/* long. Dorsal setae arising from discs, of two types, from 50-70/* long- 
and from 20-23/* long, stouter and blunter than in carveri, thickly furnished 
with short ciliations. Genitalia with two pairs of subcircular discs. 

Lot. From larvae on a rock thrush, Montieola soliiarius from Okinawa, 
16 June, 1945. As the authors had two species of larvae, Neoschongastm ameri- 
cana solomowis Wharton and Hardeastle, and Neoschongastn montieola -Wharton 
and Hardeastle in the rearing tube from which nymphs were obtained, they were 
unfortunately unable to assign the nymphs to either one of these species. 

Neoschongastia gallinarum (Hatori 1920). 

Tromhi-cula galMfnwrwm Hatori 1920, Taiwan Igakkai, No. 209, pi. II, fig. 1-6 
(larvae) ; Kawamura and Yamaguchi 1921, Kitasato Archiv. Exper. Med., 
4, 169; Fletcher, Lesslar and Lewthwaite 1928, Trans. Roy. Soc. Trop. 
Med. and Hyg., 22, 161. 

Neoschongastia gallinarwm* Sugimoto 1936, J. Jap. Soc. Vet. Sei., 15. 201; 
Wharton and Hardeastle 1946, J. Parasitol., 32, (3), 292. 



Womeksley— Asiatic-Pacific Scrub Typhus Mites 391 

Paraschonrjastiu gallinamm, Woinersley and Heaslxp 1943, Tr, Roy. Soc. S. 
Austr., 67, (1), 130. 

Plate 106, fig. A-D. 

The nyniph of this species was originally reared by Kawamura and Yama- 
guchi from Formosan material. They did not very adequately describe it, 
although they gave certain data in tabular form, and fairly good figures of the 
crista, palpi, etc. The unusual clavate sensillae were particularly drawn atten- 
tion to. 

Apart from the reference the nymph does not appear to have been observed, 
or at least, referred to by later workers. 

The larvae have been reported from Malaya, from fowls, as were Hatori's 
Formosan material. 

Recently Dr. J. R. Audy at my request, kindly undertook to relocate this 
species in Malaya, and was successful in finding it on domestic fowls, in the 
LM.R. compound at Kuala Lumpur. 

From these larvae Audy has successfully reared a nymph, which he has 
sent on to me for study. 

In the few features mentioned by Kawamura and Yamaguchi, this speci- 
men fully agrees with their description and figures. It is to be noted that it 
is only a few setae posteriorly on the dorsum which reach 42-45/1. in length, 
most of the dorsal setae are from 16-20/* long and more tapering. 

A re-description from the nymph examined is as follows : 

Description of Nymph. Of typical Trombivula facies, with a marked con- 
striction between propodosoma and hysterosoma. Length 650/*, width across 
propodosoma 325/*, across hysterosoma 350/*. Crista short and linear with a 
posterior triangular sensillary area with paired, clavate, setulose sensillae 45/* 
long by 8/* wide, with their bases 34/*, apart; length of crista, excluding epis- 
tome, 78/*; epistome rounded-conical with dentate margin and one seta to 16/i 
long. Eyes absent. Palpi moderately slender, tibia with curved claw and 
two stout spines at base of claw. Chelicerae with finely serrate inner (dorsal) 
edge. Dorsal setae rather sparse, of two kinds, mainly tapering to 16-20/* 
long, with outstanding setules, but posteriorly interspersed with uniformly 
thick setae, to 42-45/1 long, and with outstanding setules. Legs all shorter 
than body, I 338/*, II 235/*, III 208/*, IV 235/*; tarsi I 84,* long by 39/* high, 
metatarsi I 52/* long. Genitalia with two pairs of ventral discs. 



392 Records of the S,A. Museum 



• 



Subfamily GAHRLIEP1INAE nom. nov, 1 

for Walchimae Ewing 1946, J. Parasitol., 32, (5), 435-440. 

The subfamily Walchiinae was raised by Ewing for the genera Walchia 
©wing 1931, Gahrliejria Ouds., 1912, Schongastiella Hirst 1915 and Gateria 
Ewing 1938 on larval characteristics only, the adult or nymphal stages being 
then unknown. His diagnosis was as follows ; 

" Piercing part of chelicerae blade-like, upcurved, without a row of dorsal 
teeth and extending beyond apex of basal eheliceral segment. Tracheae and 
spiracles absent- Dorsal plate without median or subinedian seta, frequently 
extending backward over part of dorsum of abdomen and bearing one or more 
pairs of dorsal abdominal setae. Pseudostigmatie organs not simple setae but 
modified into true sensillae. Abdomen frequently constricted near middle, but 
never with a pair of postero-dorsal plates. Legs rather short; tarsi three- 
clawed, but one or two claws frequently much reduced. '* 

In 1943, Womersley and Heaslip did not recognize the genera Schongas- 
tiella and Gateria as distinct from Gahrliepia. In the present paper, the 
nymphs of species belonging to both Walchia and Schongastiella are described 
for the first time. In addition, although the larvae have not been specifically 
determined, I have seen the nymphs of at least two species of Gateria from 
Burma collected by the American team during the war. 

All the above nymphs are very closely allied, only differing specifically in 
the details of the dorsal setae and the sensillae. They all agree, however, 
in possessing on the front tarsi, a peculiar subapical dorsal stump-like process, 
not found in any other group of the Trombiculidae. This feature is to be 
regarded as distinguishing in the nymphal (and probably also the adult) stage 
the subfamily Gahrliepiinae from the Trombiculinae, while as no other charac- 
ters -can be found to separate the nymphs of Walchia, Gahrliepia, Schongastiella 
and Gateria, these should be considered as synonymous, or at most, but sub- 
genera on the characters of the larval scutum. Thus the subfamily Gahrlie- 
piinae will contain only the genus Gahrliepia Ouds. 1912 with Typhlothrom- 
bium manvs Ouds. 1910 as genotype. 



1 As the type genus of Ewing 'a subfamily is here considered as being congeneric and 
synonymous with Gahrliepia Ouds- 1912, it becomes necessary under Art. 5 of the International 
Bules of Zoological Nomenclature to change the subfamily name. 



Womersley — Asiatic-Pacific Scrub Typhus MlTES 393 

Ofinus GAH&LIEPIA Ouds, 1912. 
Ent. Ber., 1912, 5, 237; Zool. Jahrb., SuppL 14, 1912, 

Genotype Typhlothrombidium nanus Ouds. 1910 (Larva). 

(Including Gahrliepia Ouds. 1912. Entom. Ber. Ned. Ent. Ver. 5, noni. nov. 

for Typhlothrombium Ouds. 1910, Ent. Ber. Ned. Ent, Ver. 3, 102, preoc. ; 

Schongastiella Hirst 1915, Bull. Ent. Res., 6, 183; Gateria Ewing 1938, 

J. Wash. Acad. Sci., 28, (6), 295; Gahrlieph Womersley and Heaslip 

1943, Tr. Roy. Soc. S. Austr., 67, (1), 136, Wahhia Ewing 1931, Proc. 

U.S. Nat. Mus., 80, (8), 10; Womersley and Heaslip 1943, Tr. Roy. Soc. 

S, Austr., 67, (1),184). 

Diagnosis of Genus from Nymph. Shape typical of Tr&tribicvia\. Crista 
linear, with widely transverse dumb-bell like sensillary area and paired fila- 
mentous sensillae. Eyes absent. Palpal tibia with 2 accessory spines at base of 
claw. No precoxal plates. Sternum entire. Epistome rounded, denticulate, 
with 1 seta. Tarsi of leg I with a characteristic slump-like process dorsally and 

subapically. , 

Nymphal Genotype SehbngastieUa disparungnis Ouds. 1929- 

Key to the Known Species of Nymphs. 

1. Sensillae with fairly long to long ciliations for the whole or part of 
their length . . ♦ . . ♦ . . . ► • . * . • • • • 2 
Sensillae with only short ciliations . . . . . . * . • - 3 

2. Sensillae with only short barbs on the basal fifth, and then long out- 
standing ciliations. Dorsal setae graduating from lfy long anter- 
iorly to 32/* posteriorly. Tarsi of leg I half as long again as high, and 
half as long again as metatarsi. 

Gahrliepia (Walchia) disparungnis Ouds. 1929. 

Sensillae with only fairly long ciliations for their whole length. 
Dorsal setae to 28/x long posteriorly. Tarsi and metatarsi of leg I of 
about the same proportions as above. 

Gahrliepia (WaSchia) lewthwwitei Gater 1932. 

Sensillae with short barbs on basal half, and then long stiff setules. 
Dorsal setae to 30p long posteriorly. Tarsi of leg I two-fifths as long 
again as high, and half as long again as metatarsi. 

Gahrliepia {Walchia) mstica Gater 1932. 

3. Dorsal setae of two distinct sizes, 14/x long anteriorly and to 35/a pos- 
teriorly, stout, with strong setules. Tarsi half as long again as high, 
and half as long again as metatarsi. 

Gahrliepia (Schongastiella) acylomcasy.n. 

Dorsal setae gradually increasing in length posteriorly . . . . 4 



394 Records of the S.A. Museum 

4. Dorsal setae posteriorly to 64/a long, thick with strong setules. Sen- 
sillae barbed on basal third then with short ciliations. Tarsi 11-6 
times as long as high and slightly more than this longer than meta- 
tarsi. Gahrliepia {Schongastiella) Uguttk (Radford 1946). 

Dorsal setae more slender , . 5 

5. Sensillae slightly lanceolate, being wider medially than basally, with 
short ciliations. Dorsal setae to 30/u Tarsi I 14 times as long as 
highj and 1 • 5 times as long as metatarsi. 

Gahrliepia (WaMiia) enode (Gater 1932). 

Sensillae not so, filamentous . . . . ► . 6 

6. Sensillae rather thick, with short ciliations on basal half lengthening 
distally. Dorsal setae from 20 to 25/i long, tapering, with outstanding 
setules. Tarsi 11*6 times as long as high and as long as metatarsi. 

Gahrliepia {Schongastiella) punctata (Radford 1946 )* 
Sensillae rather thick, basal third with barbs, distally with longer 
ciliations. Dorsal setae posteriorly to 30/* long. Tarsi 11-8 times as 
long as high, and 1-8 times as long as metatarsi. 

Gahrliepia {Gahrliepia) ciliata Gater 1932, 

Sensillae rather thick, but tapering, barbed on basal third, then with 
long ciliations. Dorsal setae posteriorly to 24/* long, curved, with 
strong setules Tarsi I 1-4 times as long as high and 1*68 times as 
long as metatarsi. Gahrliepia {Gakrliepia) ornatasp.n. 

Sensillae fine, with short ciliations. Dorsal setae to 32^ long, with 
outstanding setules. Tarsi 1 1-5 times as long as high, and 1-65 times 
as long as metatarsi. Gahrliepia (Gateria) hirsuta (Radford 1946). 

Gahuliepia (Walchta) dispAbunguis (Oudemans 1929). 

Schongastiella disparungms Ouds. 1929, Ent. Ber., 17, (165), 398. 

Walchia disparungttis, Womersley 1944, Tr. Roy. Soc. S. Austr., 68, (1), 101. 

Plate 107, fig. I-M. 

Description of Nymph. Shape as in typical Trombiwda, Colour in life* 
whitish. Length to 450/a; width across propodosoma 270/x, across hysterosoma 
300/i* Cri&ta linear, 56/t long, rather indistinct apically ; epistome rounded "with 
denticulations and 1 seta; sensillary area widely transverse, 65^, with a short 
posterior apex, and paired filamentous sensillae with short ciliations basally r 
longer distally, to 90//, long and their bases 54/a apart. Eyes absent. Chelicerae 
with fine serrations on inner margin. Palpi rather slender, tibia with 2 strong 
spines at base of claw. Legs rather short. I the longest and strongest and 300/t 
long, II 210/<. ; ITT 210/j, IV 270/*; tarsi I ovoid, 75/i. long by 54/i high, meta- 
tarsus I 54/x long; tarsi I provided dorso-apically with a characteristic short 



398 Records of the S.A. Museum 

Description of Nymph. Of typical Tro7nbiculn faeies. Length 420/&; 
"width across propodosoma 227/a, across hysterosoina 256/*,. Crista short, length 
51/*, epistome not discernible, sensillary area wider than anterior arm of crista, 
is long, and with sensillae bases 41/* apart; sensillae short, 64/*, strongly but 
shortly ciliated and lightly thickened in middle three fourths, Chelicerae with 
inner edge finely serrate. Eyes absent. Palpal claw with 2 accessory spines at 
base, apparently no nude seta or spine on outer surface; tarsi with I apical 
nude seta. Legs I longer and stronger than rest, 326/*, II 213/*, III 255/*, 
IV 280/i; tarsi I with the dorsal and subapical process characteristic of the 
subfamily, 73/* long by 51/* high, metatarsi I 48/* long. Dorsal setae to 30/* 
long, with long setules as figured. Genital discs two pairs. 

Remarks. Larvae from B. r. brunneusculus from Imphal, Manipur State, 
India, Nov. 15, 1945. 

Gahkliepia (Gateria) hirsuta (Radford 1946). 

Gateria hirsuta Radford 1946, Proc. Zool. Soc. London, 116, (2), 249 (as 
larvae). 

Plate 109, fig. D-H. 

A single nymph of this species has been sent to me for study by Lt.-Col. 
Audy, which was reared by Mr. K. L. Cockings and the recovered pelt deter- 
mined as a typical hirsuta Radford. 

As with other species of Walchia, Oateria, and &chdngastiella>, the nymph 
cannot be separated generically from Oakrliepia and all are characterized by 
the peculiar process on tarsus I. 

Description of Nymph. Faeies of typical Trombicula. Length (newly 
emerged) 497ft; width across propodosoma 241/*, across hysterosoma 256/*. 
Crista short, 48/* long, epistome indiscernible, sensillary area wider than arm 
of crista, with SB 40/* apart ; sensillae filamentous, 80/* long, with short barbs 
basally and short ciliations distally. Eyes absent. Palpal claw with two 
accessory stout spines at base, and no outer spine on tibia; tarsus with only 1 
nude apical spine. Legs I longer and stronger than rest, 340ft II 170/*, III 180/*, 
IV 212/*; tarsi I short and stout, 80/* long by 54/* high, with a short dorso- 
apical blunt process; metatarsi I 48/* long. Dorsal setae stout with strong 
setules, to 30/*. long, setules not as numerous as in enode. Genital discs two 
pairs. 

Loc. and Host. The larvae from which the above nymphs was reared was 
taken from a rat, at Imphal, Manipur State, India , 17 Dec, 1945 (K. L. Cook- 
ings). 



Womersley— Asiatic-Pacific Scrub Typhus Mites 395 

clavif orni process ; tarsi I and II with many long slightly curved , sensory rods 
as well as the filiated setae. Tarsal claws unequal. Dorsal setae uniform, 
with long outstanding ciliations, from 16/* anteriorly to 30/x posteriorly and 
arising from closely adjacent platelets. 

Loc. Described from type and two paratype nymphs reared from engorged 
larvae from rats from Dobodura, New Guinea, 6 June, 1944 (No. 385), by 
Major G. M. Kohls. 

Gahrliepia (Schongastiella) punctata (Radford 1946). 

Schongastiell® punctata Radford 1946, Proe. Zool. Soc, London, 116, (2), as 
larvae. 

For further larval synonymy see Larval Section. 

Plate 108, fig. K-O. 

This speeies from Manipur State, India, has hitherto only been known from 
the larvae. 

I have now been priviliged to study a number of nymphs reared from 
larvae in Ceylon by Mr, S. H. Jayewickrenie. The nymph is here described 
and figured. 

In 1943 Womersley and Heaslip suggested that the larval genera Schon- 
gastiella Hirst and Gateria Ewing could not be generieally separated from Gahr- 
liepia Oudemans, on the basis of the number and arrangement of accessory setae 
on the scutum, as had been done by Ewing. 

In the present paper has been described the nymph of a species of Gahr- 
liepia (W. dwparunguis (Ouds.)), and the genus defined for the nymphal 
and probably the adult stage by the structure of the crista, absence of eyes, 
and particularly by the presence of a peculiar stump-like process dorsal and 
subapical on the front tarsus. In the nymphs of Schonffastwlla punctata 
described below, as well as in the nymphs of other species of Gahrliepia, 
Gateria and Walchia described later, they are also to be found. It appeals then, 
that not only are the characters used to separate these genera in the larvae un- 
sound, but that they cannot be separated generieally at all in the later stages. 
These genera must therefore be regarded as synonymous with, or at most only as 
subgenera of, Gahrliepia, on larval features. 

Description of Nymph. Of typical Trombicula faeies. Colour (in spirit) 
whitish. Length (of new emerged unfed nymph) 580/i; width across propodo- 
soma 325/i, across hysterosoma 350/4, Crista short, linear, 70/*, with broad sen- 



396 Records of the S.A. Museum 

sillary area j sensillae rather thicker medially, 84/* long, basal third with minute 
barbs, then with strong, moderately long ciliations. Sensillae bases 45/* apart. 
Byes absent. Epistome eonical-trianglar, denticulate, with 1 seta, Chelicerae 
normal, inner edge finely serrated. Palpi rather slender; tibia with 2 acces- 
sory strong spines at base of claw. Legs I longer and stronger than the rest, 
325/*, II 240/t, III 227/a, IV 292/*; tarsi I 78/* long by 50/* high with subapical 
dorsal stump, metatarsi 50/x long. No distinctly defined sternum and no pre- 
coxal plates. Dorsal setae uniform, short, 20-25/*, tapering with outstanding 
setules, and on conspicuous papillae. Genital discs 2 pairs. 

Remarks. Described from 4 nymphs bred from larvae from around the 
vibrisae of Millardia meltada meltada, Bmbilipitiya, Ceylon, Jan., 1945 (2 
specimens), and from same habitat on Bandicota malabarica, Nalanda, Ceylon, 
July, 1944 (2 specimens). 

Gahrleepia (Walchia) rustica Gater 1932, 

Gahrliepia rustica Gater 1932, Parasitology, 24, 167. 

Walchia rustica, Womersley and Heaslip 1943, Tr. Roy. Soc. S. Austr., 67, 
(1), 136; Sig Thor and Willmann 1947, as Tierreich, 71b, 337 (as 
larvae). 

Plate 107, fig. A~D. 

Description of Nymph. Of typical Tronibicuia facies. Colour in life ? 
Length 338/*, width across propodosoma 160/*, across hysterosoma 182/*,. Crista 
linear as figured, 44-8/* long, sensory area dumb-bell shaped, as broad as entire 
length of crista and with only shallow posterior angle ; sensillae very long, 88/*, 
with short ciliations on basal half and long outstanding ciliations distally; 
epistome and epistomal setae not seen. Eyes absent. Chelicerae with finely 
serrate inner edge. Palpi normal, tibia with single claw and two stout spines 
at base of claw. Legs, I stronger than others, to 312/* long, II 195**, IH 182/*, 
IV 221/*; tarsi I 70/* long by 50-4/* high and furnished with the subapical dorsal 
stump-like process characteristic of the genus; metatarsi I 47*6/* long. Dorsal 
setae uniform, fairly long to 30/*, tapering with strong outstanding setules. 
Genital discs 2 -\- 2. 

Lac, Described from some half-dozen nymphs sent to me by Dr. J. R. 
Audy and reared from larvae from Rattus r. jar ah from Patau Jarak Is., in 
Malacca Straits, 26 Jan.. 1950. 

Remarks. Distinguished as in the key. 



Womerslev— Asiatic-Pacific Scrub Typhus Mites 397 

Gahkliepia (Walchia) lewthwaitei (Gater 1932). 
Walohia lewthwaitei Gater 1932, Parasitology, 24 (larvae). 

Plate 107, fig. E-H. 

Description of Nymph. Of typical Trombimda facies. Length 450/*; width 
across propodosoina 227/*, across hysterosoma 292/*. Crista linear and short 
with wide subposterior sensillary area with a short angular posterior apex; 
total length of crista 56/*, SB 48/*; sensillae 84/* long with short ciliations in 
basal fourth, longer in remainder. Eyes absent. Chelicerae with finely ser- 
rated inner (dorsal) edge. Palpi fairly slender and normal, tibia with slender 
claw, and two accessory spines at its base. Legs, I the longest and thicker 
than the rest, to 292/* long, II 208/*, III 208/*, IV 260/*; tarsi I short and 
broad, 64*4/* long by 42*0/* high, metatarsi I 44*8/* long, and not as high as 
tarsi; tarsi I furnished with the dorso-apical stumpy process of the genus. 
Dorsal setae moderately sparse, tapering, to 28/* long posteriorly, and with 
outstanding setules. 

Loo. Described from the type nymph and 6 paratypes reared from speci- 
mens on R. whiteheacU and R. mulleri from edge of Menyala Forest Reserve, 
near Port Dickson, F.M.S., 21 Oct. 1949. The nymphs emerged 31 Oct. and 
1st Nov., 1949. 

Remarks. It does not seem possible to satisfactorily separate the nymph of 
this species from those of disparungtm in a key. It is very similar in dimen- 
sions of tarsi and metatarsi I but the dorsal setae are shorter and more uniform 
in length. 

The nymphs were reared by Mr. K. L. Coekings and checked against the 
recovered pelts. I am indebted to Dr. J. R. Audy and Mr. Coekings for the 
material from which the nymphai stage is described. 

Gahruepia (Walchia) enode (Gater 1932). 

Walohia ewde Gater 1932, Parasitology, 24; enodis Womersley and Heaslip 
1943, Tr. Roy. Soe. S. Austr., 67, (1), 135 (as larvae). 

Plate 108, fig. A-E. 

This species has been reared to the nymph by Mr. "K. L. Coekings from 
Imphal material and I am indebted to Lt.-Col. Audy for the opportunity of 
studying this material and describing the nymphai stage. 

The nymphs were correlated with the cast larval pelts. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 399 

GahrJjTftpia (Schongasttella) LiGUiiA (Radford 1946), 

Schdngastutia. ligula Radford 1946, Proc. Zool. Soc. London, 116, (2), 256 (as 
larvae). 

Plate 108, fig. F-J. 

This species is common in the region of Imphal, India, and Mr. K. Ii. 
Cookings has successfully reared a number of specimens through to the nymph r 
and checked the identification with the recovered larval pelts. I am indebted 
to Lt.-Col. Audy for the opportunity of examining and describing these 
nymphs. 

Description of Nymph. Facies of typical Trombicula form. Length 400/*; 
width across propodosoma 214/*; across hysterosoma 242/*. Crista short, 57**; 
epistome and seta indiscernible; sensillary area wide, with SB 57/* apart, and 
sensillae 120/* long, filamentous with short dilations. Eyes absent. Chelicerae 
with finely serrate inner edge. Palpal tibia with two accessory stout spines at 
base of claw; no nude seta on outer surface; tarsi with a single nude spine at 
tip. Dorsal setae pectinate with strong setules, from 30/* long anteriorly to 
60/i posteriorly. Legs I longer and stronger than rest, 312/*, II 185/*, III 171/*, 
IV 213/*; tarsi I 80/* long by 51/* high, and furnished with the characteristic 
dorso-apical stump-like process; metatarsi I 48/t long. Genital discs two pairs. 

GAHRIJEPIA (SCHONQASTIELLA) CEYIiONICA sp. DL. 

Plate 109, fig, A-C. 

Descripti&n of Nymph. Of typical Trombicula faciei Colour in life 
unknown. Length to 520/* (newly emerged) ; width across propodosoma 260/*, 
across hysterosoma 220/*. Crista linear, 70/* long, anterior arm short,, sub- 
posterior sensillary area wide, with sensillae bases 54/* apart; sensillae long, 
filamentous but moderately thick, with barbs on basal half and short ciliations 
diatally; no specialized setae in front of sensillary area. Epistome rounded- 
conical with denticulations and 1 ciliated seta. Eyes absent. Chelicerae with 
finely serrate dorsal margin. Palpi fairly stout, tibia with 2 accessory spines 
at base of claw. Legs I longer and stronger than rest, 325/* long, II 195/*, 
III 195/*, IV 220/*; tarsi I 78/* long by 52/* wide and provided with a subapicaj 
dorsal stump-like process; metatarsi I 52/* long. Dorsal setae stout, not much 
tapering, and with numerous strong setules; of two sizes, anteriorly 14/* long, 
posteriorly to 35/* long. Genital discs 2 pairs. 

Remarks. The above description is from one of four nymphs reared by 
Mr. S. H. Jayewickreme from larvae from the ears of Eattus r. Uan^diyanus 
from Nalanda, Ceylon, May, 1944, 



400 Records of the S.A, Museum 

Gahrliepia (Gahrliepia) ciliata Gater 1932. 

1932 Parasitology, 21 

Plate 115, fig, A-C. 

Description of Nymph. Of typical Trombictdu facies. Colour in life ? 
Length 536/*; width across propodosoma 268/*, across hyteterosoma 335/*. Crista 
rather short but elongate, 60/* long, with subposterior sensillary area; sensillae 
bases 43/* apart; sensillae rather thick, to 82/* long, basal third with short barbs, 
distal part with longer ciliations; epistome difficult to see but apparently coni- 
cal with one seta. Byes absent. Inner edge of chelicerae finely serrate. Palpal 
tibia with single claw and two accessory spinas at base. Dorsal setae from 18/* 
long anteriorly to 30/* posteriorly, rather straight with outstanding setules. 
Legs: I 402/i, longer and stronger than the others, II and III 228//,, IV 260/*.; 
tarsi of leg I 90/* long by 50/* high, metatarsi I 50/* long ; tarsi I furnished with 
the usual dorteal apical stump-like process. Genital discs two pairs. 

hoc. A single nymph; provisionally identified by Mr, Nadehatram as 
cetrata Gater, but which is actually ciliata Gater, was reared by him from 
larva from Rattus sabwtus from Ulu Langat Forest Eeserve, Selangor, F.M.S., 
12 Aug., 1950. The nymph was checked with the larval pelt. 

Gahrliepia (Gahrliepia) ornata sp. n. 
Plate 115, fig. D-F. 

Description of Nymph. Of typical Trombictda facies. Colour in life? 
Length 496/* ; width across propodosoma 268/* across hysterosoma 320/*, Crista 
rather short, 67*t, elongate, with subposterior sensillary area as figured: sen- 
sillae bases 40/* apart; sensillae rather thick but tapering to 84/* long, with 
short barbs on basal third then with long ciliations; eptstome rather indistinct 
but apparently conical and dentate with single ciliated seta. Eyes absent. 
Chelicera with finely serrate inner edge. Palpi slender, tibia with simple 
claw and two accessory spines at its base. Dor!sal setae rather short, from 16/* 
long anteriorly to 24** posteriorly, tapering, curved and with strong setules. 
Legs: I longer and stronger than rest, to 335/*, II 228/*, III 228/*, IV 281/*; 
tarsi of leg I rather oval, 84/* long by 60/* wide, furnished with the usual sub- 
apical subdorsal stump-like process of the genus; metatarsi I 50/* long. Geni- 
talia with two pairs of discs. 

Loc. This very unusual larval species has been reared to the nymph by 
Mr. Nadehatram of the Scrub-typhus Research Unit from larvae on Rattus sp. 
Ulu Langat Forest Reserve, Selangor, F.M.S., Sept., 1950. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 401 

LIST OF SPECIES 
(Including Appendix) * 

L. indicates known from larvae, N. from nymph, and A. from adult. 

* Species known as adult or nymph from other than the Asiatic-Pacific 
Region. 

Tragardhula acuscutellaris (Walch 1922), L.N.A. 

Tragardhula japonica (Tanaka 1930). L.N. 

Tragardhula tamiyai (Philip and Fuller 1950). L. 

Tragardhula fujigmo (Philip and Fuller 1950). L. 

Tragardhula migayoi (Sasa 1950). L, 

Tragardhula mitamurai (Sasa 1950). L. 

Tragardhula geckobia sp. n. L. 

Tragardhtda pentagons sp. n. L. 

Tragardhula gymnodactyla (Worn, and Kohls 1947). L. 

Tragardhula velascoi (Boshell and Kerr 1942). * L.N.A. 

Tragardhula> attenuata (Michener 1946). # L.A. 

Tragardhula alleei (Swing 1926). * L.N.A. 

Tragardhula nilotica (Ouds. 1904), * A. 

Tragardhula perumama (Ewing 1926). * A. 

Ipotrombicula elegans (Worn. 1942). A. 

Speotrombicula trifurca (Ewing 1933). * A. 

Trombicula (Trombicula) minor BerL 1905. A. 

Trombicula (Leptotrombidium) palpalis (Nagayo et al 1919). L. 

Trombicula (? Leptotrombidium) keukenschrijveri (Walch 1922). L. 

Trombicula (Leptotrombidium) pallida (Nagayo et al. 1919). L.A. 

Trombicula {Leptotrombidium) bumsi (Sasa 1950). L. 

Trombicula (1 Leptotrombidium) intermedia (Nagayo et al. 1919). L. 

Trombicula (? Leptotrombidium) fuji Kuwato et al. 1950). L. 

Trombicula ( ? Leptotrombidium) lanceolata sp. n. r Lawrence in MS. L. 

Trombicula (1 Leptotrombidium) parapalpalis sp. n. L. 

Trombicula (? Leptotrombidmm) macacus sp. n., Lawrence in MS. L. 

Trombicula (? Leptotrombidium) puta sp. n. L. 

Trombicula ( f Leptotrombidium) dux sp. n. L. 

Trombicula (f Leptotrombidium) scutellaris (Nagayo et al. 1920). L, 

Trombicula (Leptotrombidium) deliensis (Walch 1922). L.N.A. 

Trombicula (Leptotrombidium) ahamushi (Brumpt 1910). L.N.A. 

Trombictda (? Leptotrombidium) villosa sp. n. L. 



402 Records of the S,A. Museum 

Trombicula ( ? Leptotrombidium) tithwalensis sp. n. L, 

Trombicula (1 Leptotrombidium) bhimtalensis sp. n. L. 

Trombicula ( ? Leptotrombidium) longiseta sp. n, L.N. 

Trombicula (Leptotrombidium) fulleri (Ewiug 1945). L.N, 

Trombicula (Neotrombicula) fordi sp. n. s Lawrence in MS, L.N. 

Trombicula ( ? Neotrombicula) traubi sp, n. L. 

Trombicula {Neotrombicula) hastata (Qater 1932). L.N. 

Trombicula novae-hollandiae Hirst 1929, L. 

Trombicula (Neotrombicula) rara (Wakh 1922). L.N. 

Trombicula (Trombicula) hirsti Sanibon 1927. L. 

Trombicula (Trombicula) hirsti f. deliensis (Walch 1923). L. 

Trombicula (Trombicula) hirsti f. hakei Radford 1946. L. 

Trombieida (Trombicula) hirsti f. nissanensis Dumbleton 1947. L. 

Trombicula ( ? Trombicula) sobrina sp, n. L. 

Trombicula ( t Trombicula) ablephara sp, n. L. 

Trombicula (Trombicula) wichmanni (Ouds. 1905). L.N.A. 

Trombicula anous (Wharton 1946). L. 

Trombicula pluvius (Wharton 1946). L, 

Trombicula lygosomoides sp. n. L. 

Trombicula (Trombicula) samboni (Worn. 1939). L.N. A. 

Trombicula (Neotrombicula) sarcina (Worn. 1944). L.N. 

Trombicula kmzalwanemis sp. n. L. 

Trombicula ( i Trombicula) macropus Worn. 1936. L. 

Trombicula rioi Gunther 1939. L. 

Trombicula (Fonsema) coluberina (Radford 1946). L. 

Trombicula batui Philip and Traub 1950. L. 

Trombicula (Neotrombiada) munda (Gater 1932). L. 

Trombicula (Neotrombicula) spicea (Gater 1932). L.N. 

Trombicula (Neotrombicula) consueta sp. n. L.N. 

Trombicula parmifera sp. n. L. 

Trombicula lutidbladi sp. n. L. 

Trotnbicula nissani Dumbleton 1947. L. 

Trombicula jubbulporensis sp. n. L. 

Tro?>ibicula isshikii Sugimoto 1938. L. 

Trombicula khurdangensi-s sp. n. L. 

Trombicula (Neotrombicula) scincoides (Worn. 1944). L.N. A. 

Trombicula (? Neotrombicula) kolilsi (Worn. 1944). L. 

Trcmibicula ( \ Neotrombicula) tovelli sp. n, L. 

Trombicula buxtom sp. n. L. 

Trombicnda naultini Dumbleton 1947. L. 



Womersley— Asiatic- Pacific Scrub Typhus Mites 403 

Trombicufa cervulicola Ewing 1931. L. 

Trombicula densipiliata Walch 1923. L. 

Trombicula taphozous sp. n. L. 

Trombicula corvi Kaw. and Yam. 1921. L- 

Trombicula insolli (Philip and Traub 1950). L. 

Tronibicula leveri sp. n. L. 

Trombicula (Neotrombicida) harrisoni sp. n. L.N. 

Trombicula piercei Ewing 1931. L. 

Trombicula {Trombicula) frittsi Wharton 1945. L.N. 

Tr&mbicula kashmiren&is sp. n. L. 

Trombicula incurva sp. n. L. 

Trombicula vietzi sp. n. L. 

Trombicula schmitzi (Ouds. 1914). L. 

Trombicula (Leptotrombidium) myzantha sp. n. L.N. 

Trombicula (? Leptotrombidium) robusta (Gunther 1941). L. 

Trombicula rajoriensis sp. n. L, 

Trombicula sylvestris Andy and Traub 1950. L. 

Trombicula muridia> sp. n. L. 

Trombicula bodensts (Gunther 1940). L. 

Trombicula gliricolens (Hirst 1915). L. 

Trombicula philipi sp. n. L. 

Trombicula (Leptotrcrmbidium) burmensis (Ewing 1946). L.N. 

Trombicula southcotti sp. n. L. 

Trombicid^a quadriejise Worn, and Heasp. 1943. L. 

Trombicula pelta sp. n. L. 

Trombicuh' (Neotrombicula) jayewiclcremei sp. n. L.N. 

Trombicula (Trombicidindus) cuneata (Traub and Evans 1951). L. 

Trombicula squamifera sp- n. L. 

Trombicula (Trombiculindus) squamosa (Radford 1947). L. 

Trombicula (Trombicidindus) foliacea (Traub and Evans 1951). L, 

Trombicula thori sp. n. L. 

Trombicula (Trombicula) batatns (Linn. 1758). * L.N.A. 

Trombicula (Trombicula) helleri (Ouds. 1911). * L.A. 

Trombicula (Trombicula) splendent Ewing 1913. * A. 

Tro-mbicula (TromMcuHa) vanommereni Schierbeck 1937. * L.N.A. 

Trombicula (Trombicula) alfreddugesii (Ouds. 1911). * L.N.A. 

Trombicula (Trombicula) jacoti. nom. nov. * A. 

Trombicula (Neotrombicula) inopinatum (Ouds. 1909). * L.N. 

Trombicula (Neotrombicula) auiumnalis (Shaw 1790). * L.N.A. 

Trombictda (Neotrombicula) camestrinii (Buffa 1899). * A. 



404 Records of the S*A. Museum 

Trombicula ( Neotrombicula) clamcata (Andre 1937). * A. 

Trombicula (Neotrmnbicula) formicarum (Berl. 1910). * A. 

Trombicida (Neotrombicula) Camilla Wharton 1938). * A. 

Trombimla (Neatrombicida) russica (Ouds. 1902). * A. 

Trombicida {Neotrombicida) moesim (Ajidre 1932). * A. 

Trombicula {Neotrombicula) coarctata (Berl. 1888), * N. 

Trombicula (Neotrombicula) goldii (Ouds. 1910). # L.A, 

Trombicula (Neotrombicida) camrnarum (Ewing 1933). * A. 

Trombicida sp. Cooreman in litt. * A. 

TromMcula (Neotrombicula) manriquei (Ewing' 1937). * A. 

Tecomatlana melvini (Traub and Evans 1950). L. 

Myotrombicula* vespertilianis Worn, and Heasp. 1943. L. 

Eeaslipia gateri (Worn, and Heasp. 1943). L.N, 

Novotrombicida owiensis Worn, and Kohls 1947. L. 

Schdngastia (Schdngastia) oudemansi (Waleh 1923). L.N. 

Schdngastia (Schongastia) lewthivaitei sp. n. L. 

Schongastia (Schongastia) vandersandei (Onds. 1905). L. 

Schimgastia (Schongastia) schuffneri (Waleh 1923). L.N. 

Schongastia (Schongastia) taylori Gunther 1940. L. 

Bchongastia (Schongastia) ivhartoni sp. n. L. 

Schongastia (Schongastia) bid-entata sp.n , L. 

Schongastia (Schongastia) vieta Gater 1932. L.N. 

Schongastia (Schongastia) philipi Worn, and Kohls 1947. L. 

Schongastia (Schdngastia) blesiowei Gunther 1939. L. 

Schdngastia (Schongastia) blesiowei v. megapodius Worn, and Heasp. 1943. L. 

Schongastia (Schdngastia) ocvMcola sp. n. L.N. 

Schongastia (Schongastm) pseudoschufineri (Waleh 1927). L. 

Schdngastia (Schdngastia) maldiviensis (Radford 1946). L.N.A. 

Schdngastia (Ascoschdngastia) pseudomys sp. n, L. 

Schdngastia. (Ascoschdngastia) mccidlocM (Worn. 1944) L. 

Schdngastia (Ascoschdngastia) urom.ys (Worn, and Kohls 1947). L. 

Schdngastia (Ascoschdngastia) foliata (Gunther 1940). L. 

Schdnga<stia (Ascoscliongastia) signata tp.n, L. 

Schongastia (Ascoschdngastia) globular c (Waleh 1927). L. 

Schdngastia f Ascoschdngastia) Icohlsi (Philip and Woodward 194G, L.N. 

Schdngastia (Ascoschdngastia) comata sp.n. L. 

Schdngastia (Ascoschdngastia ) laniits (Radford 1946). L.N. 

Schdngastia (Ascoschdngastia) mutabilis (Gater 1932). L,N. 

Schdngastia (Ascoschdngastia) edwardsi (Gunther 1939). L, 

Schdngastia (Ascoschdngastia) philippensis (Philip and Woodward 194G), L. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 405 

Schdngastia (Ascoschdngastia) mackerrasae sp. n. L. 

Schdngastia^ (Ascoschdngastia) masta (Traub and Sundermeyer 1950). h. 

Schdngastia (Ascoschongastia) petrogale (Worn. 1934). L. 

Schdngastia (Ascoschdngastia) mohri sp. n. L. 

Schdngastia (Ascoschongastia) crinita sp. n. L. 

Schdngastia (Ascoschongastia) rcctangulare sp. n. L. 

Schdngastia (Ascoschongastia) cassiope sp. n. L. 

Schdngastia (Ascoschongastia) antipodiamm (Hirst 1929). L. 

Schdngastia (Ascoschongastia) dumosa sp, ri. L. 

Schdngastia (Ascoschdngastia) qucenslandica (Worn. 1939). L. 

Schdngastia (Ascoschdngastia) lappacea nom. uov. L. 

Schdngastia' (Ascoschdngastia) womersleyi (G wither 1940). L. 

Schdngastia^ (Ascoschdngastia) coorongense (Hirst 1929). L. 

Schdngastia (Ascoschdngastia) echymipera (Worn, and Kohls 1947). L. 

Schdngastia (Ascoschdngastia) inmsfailensis (Worn, and Ileasp. 1943). L, 

Schdngastia (Ascoschdngastia) lacunosa (Gater 1932). L. 

Schdngastia (Ascoschdngastia) malayensis (Gater 1932). L.N. 

Schdngastia (Ascoschdngastia) lipoxena sp. n. L, 

Schdngastia (Ascoschdrigastia) nadchatrami sp. n. L. 

Schdngastia (Ascoschdngastia) sarawakensis sp. n. L. 

Schdngastia (Ascoschdngastia) nausheraens'is sp. n. L. 

Schdngastia (Ascoschdngastia) debilis Gater 1932. L. 

Schdngastia (Ascoschdngastia) mampurensi^ (Radford 1946). L. 

Schdngastia (Ascoschdngastia) labuanensis sp. n. L. 

Schdngastia (Ascoschdngastia) audyi sp. n. L.N. 

Schdngastia (Ascoschdngastia) rattus (Worn, and Heasp, 1943). L. 

Schdngastia (Ascoschdngasiia) lorius (Gtmther 1939), L. 

Schdngastia (Ascoschdngastia) indica (Hirst 1915). L.N.A. 

Schd7igasiia (Ascoschdngastia) soekaboemiensis (Takekawa 1945), L. 

Schdngastia (Ascoschdngasiia) perameles ("Woin. 1939). L. 

Schdngastia (Ascoschdngastia) similis (Worn, and Heasp. 1943). L. 

Schdngastia (Ascoschongastia) derricki (Worn. 1939). L. 

Schdngastia (Ascoschdngastia) dasijcerci (Hirst 1929). L. 

Schdngastia (Ascoschdngastia) trichosnri (Worn. 1939). L. 

Schdngastia (Ascoschdngastia) hirst i (Worn, and Ileasp. 1943). L. 

Schdngastia (Ascoschongastia) phascogalc (Worn, and neasp. 1943). L. 

Schdngastia (Ascoschdngastia) peregrina sp. n. L. 

Schdngastia (Ascoschdngastia) heasHpi (Worn, and Heasp, 1943). L. 

Schdngastia (Ascoschongastia) tranbi sp. n. L. 

Schdngastia ( Ascoschongastia) lawrencci nom, nov. L. 



406 Records of the S.A. Museum 

Schongastia (Ascoschdngastia) iwngabelensis sp. n. L. 

Schongastia (Ascoschdngastia) smithi (Worn. 1939). L. 

Schongastia (Ascoschdngastia) rani sp. n. L. 

Schongastia (Ascoschdngastia) omrnsensis (Worn, and Heasp. 1943). L. 

Schongastia (Ascoschdngastia) cairnsensis v. gateri (Worn. & Heasp. 1943). L. 

Schongastia algerica (Andre 1932). # A. 

Schongastia- umtraliensis (Worn. 1934). A. 

Schongastia samoansis sp. n. A. 

Radfordiwna rostrata g, et. sp. n. L. 

Oenoschdngastia cana Worn, and KohLs 1947, L. 

Oimtherana bipygalis (Gunther 1939). L.N. A. 

Ghmtheram tindalei (Worn. 1936) N. 

Guntherana translttcens (Worn. 1944). A. 

Neoschongastm bougainmllensis Wharton and Hardeastle 1946. L. 

Neoschdngastia stron.gi Wharton and Hardeastle 1946. L. 

Neoschdngastia monticola Wharton and Hardeastle 1946. L. 

Neoschdngastia posekanyi Wharton and Hardeastle 1946. L. 

Neoschdngastia gallinarum (Hatori 1920). L.N. 

Neoschdngastia backhoxisei Gunther 1939. L. 

Neoschdngastia egret t a Wharton and Hardeastle. 1946. L. 

Neoschdngastia struthidia sp. n. L. 

Neoschdngastia retrocincta Gunther 1939. L. 

Neoschdngastia yeomansi Gunther 1939. L„ 

Neoschdngastia owi-ensis sp. n. L. 

Neoschdngastia mcqueeni sp. n. L. 

Neoschongastm atoUensis Wharton and Hardeastle 1946. L. 

Neoschdngastia namrvi Wharton and Hardeastle 1946. L. 

Neoschdngastia amencana solomonis Wharton and Hardeastle 1946. L. 

Neoschdngastia entomyza sp. n. L. 

Neoschdngastia thomasi (Radford 1946). L. 

Neoschongastm pauensis Wharton and Hardeastle 1946. L. 

Neoschdngastia salmi (Ouds. 1922). L. 

Neoschdngastia dubia Gunther 1939. L. 

Neoschdngastia ewingi Wharton and Hardeastle 1946. L. 

Neosclidngastia riverti Wharton and Hardeastle 1946. L. 

Neoschdngastia cwrveri Wharton and Hardeastle 1946. L.N. 

Neoschdngastia sp. Wharton and Hardeastle 1946. N- 

MacMena empodiformis Traub and Evans 1950. L. 

Gahrliepia (Walchia) morobensis (Gunther 1939). L. 



Womersley— Asiatic-Pacific Scrub Typhus Mites 407 

GahHiepia (Walchia) rustica Gater 1932. L.N. 
Gahrliepia {Walchia) turmaUs Gater 1932. L. 
Gahrliepia (Walchia) lewthwaitei sp. n. L.N. 
Gahrliepia (Walchia) disparunguis (Ouds. 1929). L.N. 
Gahrliepia (Walchia) emngi (Fuller 1951, nom. hoy.). L. 
Gahrliepia (Walchia) enode (Gater 1932). L.N. 
Gahrliepia (Walchia) brennam sp. n. L. 
Gahrliepia (Walchia) brennani v. ventralis nov. L. 
Gahrliepia (Walchia) rim Gimther 1940. L. 
Gahrliepia (SchongastieUa) ligula (Radford 1946). L.N. 
Gahrliepia (SchongastieUa) ceylonica sp. n, L.N. 
Gahrliepia (SchongastieUa) brevis (Radford 1946). L. 
Gahrliepia (SchongastieUa) bengalensis (Hirst 1915). L. 
Gahrliepia (SchongastieUa) punctata (Radford 1946). L.N. 
Gahrliepia (SchongastieUa) kurnaonensis sp. n. L. 
Gahrliepia (Gahrliepia) saduskis^.n. L. 
Gahrliepia (Gahrliepia) ciliata Gater 1932. L.N. 
Gahrliepia (Gahrliepia) insigne sp, n, L. 
Gahrliepia (Gahrliepia) decora sp. n. L. 
Gahrliepia (Gahrliepia) ornata sp. n. L.N. 
GahHiepia (Gahrliepia) cetrata Gater 1932. L. 
Gahrliepia (Gateria) hirsuta (Radford 1946). L.N. 
Gahrliepia (Gateria) lancearia (Radford 1946). L. 
GaJirUepia (Gateria) longipilis (Radford 1946). L. 
Gahrliepia (Gateria) crocidura (Radford 1946). L. 
Gahrliepia (Gateria) ronieri sp.n. L. 
Gahrliepia (Gateria) rutila (Gater 1932). L. 
Gahrliepia (Gateria) spimdosa (Radford 1946). L. 
Gahrliepia (Gateria) ftetcheri (Gater 1932). L. 

SUMMARY. 

Total species mentioned . . - . . - . - . . • - . . 254 

Exotic species (other than Asiatic-Pacific) . . . , . . . . . . 26 

New species . • • • » • - • • » • • ; • • • • • ?? 

Asiatic-Pacific species as Larvae . . . . . . . . • - . . 221 

Asiatic-Pacific species as Adults and/or Nymphs 53 

Species in which Larvae have been correlated with Adults and/or Nymphs : 

1. Asiatic-Pacific . . . . . . • • • • - • • • - * 46 

2. Other than » . . • . . . » . * * » • • - • 10 



408 Records of the S.A. Museum 



REFEKENCES. 



1. Andre, Marc, 1926. Notes coniplementaires et synonymiques concernant 
divers Trombidiidae, Bull. Mus. Nat. d'Hist (No. 6). 

2. Andre, Marc, 1926a. Trombidiidae de la Faune frangaise. Bull. Soc. 
Zool. France. 

3. Andre, Marc, 1930. Contribution a le 'etude d'un Acarien; Le Thrombi- 
culu autummlis (Shaw). Mem. Soc, Zool. de France, 29, (2), 39-138, 
fig. 1-25. 

4. Andre, Marc 1931. De rimmunisation chez l'espece huinaine contre les 
atta