No.

CONTENTS

. A synopsis of the frogs of the genus Ayla of north-western

Australia, with the description of a new species. Michael J.

Tyler. pp. 1-11.

(Issued 5th December 1969)

. An annotated catalogue of the Otitidae (sens. lat.) (Diptera)

recorded from Australia, Indonesia and the Pacific Islands.

Harry F. Lower. pp. 1-93.

(Issued 6th March 1970)

. The Rhodacaridae (Acari: Mesostigmata); classification, external

morphology and distribution of genera. David C. Lee. pp.

1-219.

(Issued 22nd July 1970)

. A revision of the Australian pentatomid bugs of the genus Cephalo-

platus White. (Hemiptera—Pentatomidae—Pentatominea).

Gordon F. Gross. pp. 1-58.

(Issued 23rd October 1970)

. Cyatholaimidae (Nematoda) from the Coast of Western Australia.

W. Grant Inglis. pp. 1-13.

(Issued Ist December 1970)

. The genus Ctenotus (Lacertilia, Scincidae) in South Australia.

G. M. Storr. pp. 1-15.

(Issued 18th February 1971)

. Acarine and other microfossils from the Maslin Eocene, South

Australia. R. V. Southcott and R. T. Lange. pp. 1-21.

(Issued Ist June 1971)

. Nematode parasites of Oceanica. XII. A review of Heterakis

species, particularly from birds of Taiwan and Palawan.

W. Grant Inglis, Gerald D. Schmidt and Robert E. Kuntz.

pp. 1-14.

(Issued 7th October 1971)

. Superficial mandibular musculature, vocal sacs and the phylogeny

of Australo-Papuan leptodactylid frogs. Michael J. Tyler.

pp. 1-20.

(Issued 12th May 1972)

No.

. 10.

13,

. 14.

Planigale gilesi (Marsupialia, Dasyuridae); a new species from the

interior of south-eastern Australia. Peter F. Aitken. pp. 1-14.

(Issued 12th June 1972)

. An annotated checklist of the native land mammals of the Northern

Territory. Shane A. Parker. pp. 1-57.

(Issued 8th January 1973)

. Morethia (Lacertilia, Scincidae) South Australia. Michael Smyth.

pp. 1-14.

(Issued 31st August 1972)

A new species of Troposodon Bartholomai; from the Early

Pleistocene Kanunka fauna, South Australia. (Macropodinae;

Marsupialia.) Colin R. Campbell. pp. 1-18.

(Issued Ist February 1973)

Rhodacaridae (Acari: Mesostigmata) from near Adelaide

Australia. I. Systematics. David C. Lee. pp. 1-36.

(Issued Ist March 1973)

INDEX

Acari, Rhodacaridae, genera, world . RP? ee Se

eriophyid, microfossil, South POEM 2. yh ocarecoe | SL

Rhodacaridae, near Adelaide .. .. .. 1. 1. we we ee ee ee

Aitken, Peter F.—

Planigale gilesi, Marsupialia, new species .. .. .. .. .. .. --

Anura, Hyla, north-western Australia .. .. nS RE ke IRS OTe

leptodactylid frogs, Australo- -Papuan a

Bugs

Secvblemiptorates: 271 6 te ORS 4 br re ee

Campbell, Colin R.—

Troposodon, Marsupialia, new species, fossil .. .. .. .. .. ..

Cephaloplatus, Hemiptera, generic revision .. .. .. .. .. ....

Ctenotus, Lacertilia, South Australia .. .. .. . Ley ese eS

Cyatholaimidae, Nematoda, coast, Western Aapeeralin* aes te oh ad 5 le

Diptera, Otitidae, Australia, Indonesia, Pacific Islands .. .. .. .. .. ..

Flies

See Diptera ...... . eee

Fossils, mite, rotifer, von braie dob eden Souk yaaa aes

Troposodon kenti, abs cease new ia Pleistocene, South

AMStralia: 6: (oe fee Pee ee AE ee to ore

Frogs

SEeZARUTASS eh eran, ree renee eee

Gross, Gordon F.—

Cephaloplatus, Hemiptera, generic revision... .. .. .....

Hemiptera, Cephaloplatus, generic revision .. .. . meee he

Heterakis, all species, particularly from birds, Teat i ales ge AL es

Hyla, Anura, north-western Australia . 2. 6. 6. 6. ck ee ce ee ee ee

Inglis, W. Grant—

Cyatholaimidae, Nematoda, coast, Western Australia ..

Inglis, W. Grant, Schmidt, Gerald D. and Kuntz, Robert E.—

Heterakis, all species, particularly from birds, Taiwan, Palawan

Kuntz, Robert E.—

See Inglis, W. Grant .

Number

—=

Aan BW

Lacertilia, Ctenotus, South Australia .. 2. 0. 6. ok ee ee ee eee

Morethia, South Australia .. 2. 6. 6. oe ee ee ee ee ee ee ee

Lange, R. T.—

SGC SSOUP COT ORs Vem cB ance SR et alee eee: | gat i

Lee, David C.—

Rhodacaridae, genera, world .. .. ..

Rhodacaridae, near Adelaide .. .. .. .

Leptodactylidae, Anura, Australo-Papuan, musculature, liieaens pea

Lists, Cephaloplatus, species (all Australian) .. .. .. .. .. 0... .. ee

Ctenotus, species, South Australia .. .. .. .. -. ee ee ee ee

Heterakis, species, world . ze niet Me Ale ee eet

Hyla, species, north- western Australia... .. ...... .

Mammals, native, land, Northern Territory .

Otitidae, species, Australia, Indonesia, Pacific Rie ars

Rhodacaridae, genera, world .

Lizards

Seevleacentiliat... ot Sia: Vee ee ee lew

Lower, Harry F—

Otitidae, Australia, Indonesia, Pacific Islands .. .. .. .. .. ..

Mammals, Planigale gilesi, Marsupialia, new species .. .. ..

native land, checklist for Northern Territory .

Troposodon kenti, Marsupialia, new species, fossil .

Mites

See Acari... ...

Morethia, Lacertilia, South Acustealia ee Re rhe had ee

Musculature, Australo-Papuan, leptodactylid tres acd ne Si ee

Nematoda, Cyatholaimidae, coast, Western Australia .. ..

Heterakis, all species, particularly from birds, Taiwan, Palawan

New taxa, family-group, Acari .. .. .. .. .. 2...

genus-group, Acari .. la tisk ets Ae colle ae

IRGMIGe a, See Tey eee A en A Err

species-sroupeAca r=) ny ee en oy

PATUG A ade Ae eine

Hemiptera... .. .. .... V4 bot oe 4 cs. ».

IeCencitiates eee ero e te ee wh beets t es

Mammalia eo, JF SS UP,

WNematodan. a: ase he as aie

Otitidae, Catalogue, Australia, Indonesia, Pacific Islands .. .. .. .. ..

Parker, Shane A—

native land mammals, checklist, Northern Territory .. .. .. ..

Phylogeny, Australo-Papuan, leptodactylid frogs .. .. .. .. .. 0... o.

Planigale gilesi, Marsupialia, new species, South Australia .. .. .. 2...

Number

WNHKH KH ODAA CO PhW

—_

Boh

joa

iS)

Rhodacaridae, genera, world .. .. ..

near Adelaide .. .. .. .

Rotifer, ? Keratella sp., fossil ..

Roundworms

Gal CUD OC pert ike te Me te eS ia ee ns oa cm ee

Schmidt, Gerald D.

See Leis, NN (Gir tee ae cei Pe en

Smyth, Michael—

Morethia, Lacertilia, South Australia .. .. 2. 0... 2. 0.

Southcott, R. V. and Lange, R. T.—

microfossils, Eocene, South Australia .. .. .. 1. .. 0. 0...

Storr, G. M.—

Ctenotus, Lacertilia, South Australia... .. 2... 0... .... 4.

Troposodon, Marsupialia, fossil, revised generic diagnosis .. .. .. .. ..

Tyler, Michael J.—

Hyla, north-western Australia .. 2. 6. 6. ek ee ce ee ee ee ee

leptodactylid frogs, Australo-Papuan ..

Vocal Sacs, leptodactylid frogs, Australo-Papuan .. ..

A. B. JAMES, GOVERNMENT PRINTER, AQELAIDE

_—

RECORDS oF THE

SOUTH AUSTRALIAN / em

MUSEUM ey,

A SYNOPSIS OF THE FROGS OF THE

GENUS HYLA OF NORTH-WESTERN

AUSTRALIA, WITH THE DESCRIPTION

OF A NEW SPECIES

By MICHAEL J. TYLER

Honorary Associate in Herpetology

South Australian Museum

SOUTH AUSTRALIAN MUSEUM |

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER |

5 December 1969

A SYNOPSIS OF THE FROGS OF THE GENUS HYLA OF NORTH-

WESTERN AUSTRALIA, WITH THE DESCRIPTION OF A NEW SPECIES

BY MICHAEL.J. TYLER,

HONORARY ASSOCIATE IN HERPETOLOGY, SOUTH AUSTRALIAN MUSEUM

Summary

In comparison with studies undertaken in other parts of Australia, the hylid frog fauna of the north-

west has received little attention. The most recent checklists of the species of Hyla are those

compiled by Moore (1961), recognizing six species in northern Western Australia and nine in the

Northern Territory.The majority of species listed are widely distributed in Australia, but the

recorded presence of Hy/a aurea and H. adelaidensis in the Northern Territory evoked comment

from Moore because the disjunct distribution conflicted with existing knowledge.

In a recent study Tyler (1968a) investigated the taxonomic status of the members of the H. lesueuri

complex occurring in north-western Australia and described a new species apparently confined to

the area. The possible existence of an endemic element within the hylid frog fauna has been

reinforced by the subsequent collection of a further undescribed species.

This study has endeavoured to establish the number of species of Hy/a occurring in the north-west

and to compare their distribution patterns.

A SYNOPSIS OF THE FROGS OF THE GENUS HYLA OF

NORTH-WESTERN AUSTRALIA, WITH THE DESCRIPTION

OF A NEW SPECIES

By MICHAEL J. TYLER, Honorary ASSOCIATE IN HERPETOLOGY,

SouTH AUSTRALIAN MUSEUM

INTRODUCTION

In comparison with studies undertaken in other parts of Australia, the

hylid frog fauna of the north-west has received little attention. The most

recent checklists of the species of Hyla are those compiled by Moore (1961),

recognizing six species in northern Western Australia and nine in the

Northern Territory. The majority of the species listed are widely distributed

in Australia, but the recorded presence of Hyla aurea and H. adelaidensis in

the Northern Territory evoked comment from Moore because the disjunct

distribution conflicted with existing knowledge,

In a recent study Tyler (1968a) investigated the taxonomic status of

the members of the H. lesueuri complex occurring in north-western Australia

and described a new species apparently confined to the area, The possible

existence of an endemic element within the hylid frog fauna has been

reinforced by the subsequent collection of a further undescribed species.

This study has endeavoured to establish the number of species of Hyla

occurring in the north-west and to compare their distribution patterns.

MATERIALS AND METHODS

The specimens reported are lodged in museum collections which are

abbreviated in the text as follows:—British Museum (Natural History):

B.M.: Museum of Comparative Zoology: M.C.Z.; National Museum of

Victoria: N.M.V.; South Australian Museum: S.A.M.; United States

National Museum: U.S.N.M.; Western Australian Museum: W.A.M.

Letters preceding registration numbers are departmental catalogue references.

Methods of measurement and descriptive techniques conform to those

used by Tyler (1968b). The following abbreviations appear in the text:—

E-N/IN (ratio of the eye to naris distance to the internarial span); HL/HW

(head length to head width ratio); HL/S-V (head length to snout to vent

length ratio); TL/S-V (tibia length to snout to vent length ratio).

Issued 5 December, 1969

2 REC, S. AUST. MUS., 16 (1): I-11 December, 1969

Hyla meiriana new species

Ayla adelaidensis, Mitchell (1955) p. 405, (1964) p. 339.

HOLOTYPE: S.A.M. R. 9082. An adult female collected at a rock

pool 98 miles north of Mainoru, Northern Territory, by A. Fleming,

R. Edwards and H. Bowshall on August 19, 1967,

FIG. 1.

Hand and foot of Hyla meiriana sp, nov.

DEFINITION: An extremely small species with a maximum snout to

vent length of 22.5 mm, characterised by an extremely high E-N/IN ratio

(1.286-1.600), short and unwebbed fingers with prominent, transversely

oval discs, and extensively webbed toes (the webbing reaching the base of

the discs of all toes except the fourth),

DESCRIPTION OF HOLOTYPE: The head is longer than broad

(HL/HW 1.097); its length equivalent to considerably more than one-third

of the snout to vent length (HL/S-V 0.383). The snout is not prominent;

it is rounded when viewed from above and rounded and projecting slightly

in profile. The nares are high and oblique, their distance from the end of

TYLER—NORTH AUSTRALIAN HYLA 3

the snout less than that from the eye. The distance between the eye and

the naris is considerably greater than the internarial span (E-N/IN 1,600).

The canthus rostralis is poorly defined and slightly curved. The loreal

region is concave. The eye is not prominent, its diameter slightly greater

than the eye to naris distance, The tympanum is prominent, its diameter

equivalent to two-thirds of the eye diameter and separated from the eye by

a distance equivalent to approximately one-third of its own diameter, The

vomerine teeth are in two raised and slightly oblique series. A line on a

level with the posterior margins of the choanae would bisect them. The

tongue is roughly circular and lacks a posterior indentation.

The fingers are rather short and are equipped with extremely narrow

lateral fringes; in decreusing order of length 3> 4> 2> 1. There is no

inter-digital webbing. The terminal discs are prominent and transversely

oval in shape (Pig. 1).

The hindlimbs are long and slender with a TL/S-V ratio of 0.539.

Toes in decreasing order of length 4> S> 3> 2> 1, The interdigital

webbing reaches the base of the terminal discs of all toes except the fourth

where it extends as far as the subarticular tubercle at the base of the

penultimate phalanx, and is united to the disc by a narrow lateral fringe

(Fig. 1).

There are numerous broad, flattened, very poorly developed tubercles

over the entire dorsal surface of the head and body. Each tubercle is

composed of numerous small granules, particularly conspicuous in the sacral

region, There is a very weak supratympanic fold obscuring the superior

margin of the tympanic annulus. There is a prominent oval inner and a

small but prominent rounded outer metatarsal tubercle. The throat, chest

and lower surfaces of the limbs are smooth and the abdomen is granular.

DIMENSIONS: Snout to vent length 20,6 mm; tibia length 11.1 min:

head length 7,9 mm; head width 7.2 mm, eye to naris distance 2.4 mm;

internarial span 1.5 mm; eye diameter 2.5 mm; tympanum diameter 1,6 mm.

In preservative the dorsal surface is dark brown with indistinct black

markings surrounding the individual tubercles. When the skin is moist the

granular areas within the tubercles possess a distinct metallic irridescence,

The canthus rostralis bears a short blackish stripe, and heavy stippling on

the mandibular margins produces a pattern of light and dark patches. The

anterior and posterior surfaces of the thighs and the posterior surface of the

tibia are strikingly variegated with black on a cream background, The

throat and chest are uniformly stippled with black on a pale cream back-

ground and there is irregular stippling on the ventral surface of the thighs.

VARIATION: There are 32 paratypes collected at the type locality

with the holotype: —S.A.M. R. 9014-34, 9074-81, 9083-85.

REC. S. AUST. MUS., 16 (1): 1-11 December, 1969

FIG. 2.

H. meiriana sp. nov.

TYLER—NORTH AUSTRALIAN HYLA 5

Adult male paratypes have snout to vent lengths ranging from 16,2 mm

to 18.2 mm, whilst the range for females is 17.6-20.7 mm. None of the

females are gravid, In their proportions they exhibit only slight variation.

The E-N/IN ratio is consistently high with a range of 1.286-1.563 and the

mean 1.407. The head is longer than broad in all specimens with an

HL/HW range of 1.057-1.222 and the mean 1.132. The TL/S-V range

is 0.541-0.640 and the mean 0,584.

The colouration and pattern of markings of the paratypes closely

rescmbles those of the holotype, Divergences worthy of note are the presence

of a dark transocular bar in some specimens, and the fact that variegations

on the lateral surfaces of the thighs frequently extend on to the dorsal surface.

An additional 34 specimens also represent this species: S.A.M. R, 3235,

9734; U.S.N.M, 12870-25, Oenpelli Creek, 5 miles $,S,E. of Oenpelli, N.T,

S.A.M. R, 9097-9100, Kununurra, W.A. N,M.Y, D, 10773-74, 10811-16,

10818-26, Jaspers Gorge, N.T. W.A.M, R. 13758, 13758 G-J,

Kalumburu, W.A.

The four specimens from Kununurra have E-N/IN and HL/HW

ranges within those of the paratypes, but the hind legs are shorter (the

TL/S-V range being 0,476-0.556 with a mean of 0.515). One of these

specimens (S.A.M, R. 9100) is the largest representative of the species

being a gravid female with a snout to vent length of 22.5 mm, Another

member of this series is depicted in Fig. 2.

The specimens from Jaspers Gorge differ from the type series in having

more extensive webbing of the feet (reaching mid-way up the penultimate

phalanx of the fourth toe), and in their colouration. The dorsum in this

series is a much darker brown, with the dorsal surface of the thigh similar

to the colour of the head and back (the light markings on the posterior

face do not extend upon il). The ventral surfaces are much more heavily

and extensively marked; the throat is usually a uniform dark brown and only

infrequently stippled with brown, and the ventral surface of the thighs is

suffused with brown in most specimens,

COMPARISON WITH OTHER SPECIES

Of the Australian species with completely unwebbed fingers the only

one Whose adults are within the size range of H. meiriana is H. microbelos

of Queensland. A single specimen has been available for comparison

(M.C.Z. 70013), an adult male collected at Cooktown which is approxi-

mately 100 miles north of the type locality (Cairns),

6 REC. §. AUST. MUS., 16 (1): 1-11 December, 1969

Hyla meiriana may be distinguished by the presence of vomerine teeth

and outer metatarsal tubercles (absent in H. microbelos) and by its more

extensively webbed toes. In preservative H. meiriana has a dark brown

dorsal ground colouration and striking post-femoral markings, whereas

H. microbelos is a very pale grey and lacks these markings.

Of the Papuan species H. dorsalis attains a similar adult size but may

also be distinguished by having less extensive webbing between the toes,

as revealed by comparison of Fig. | with the illustration of H. dorsalis

provided by Tyler (1968b, Fig. 25). The shape of the snout differs in being

evenly rounded and not particularly prominent, whereas in H. dorsalis it is

pointed and projecting. None of the 62 specimens of H. meiriara examined

bear the median, longitudinal, pale brown band visible on the dorsum of

H. dorsalis.

HABITAT

The type locality is an aboriginal ceremonial wind-dreaming site 98

miles north-east of Mainoru in an area where the annual rainfall is

approximately 50-60 inches. The pool is located on a sandstone plateau

on which there are occasional outcrops of exposed quartzite. The vegetation

surrounding the pool consists predominantly of sparse eucalypts and clumps

of coarse grasses near the water, with occasional Pandanus and paper bark

trees on the periphery. The bed of the pool is completely free of silt and

the water is described as crystal clear and extremely soft.

Messrs. J. Coventry and C. Tanner, who obtained the series at Jasper’s

Gorge, noted that there the species was living in red silt in rock pools.

FIELD NOTES

The type series was collected at night at the edge of the water. The

collectors noted that whereas other species occurring at the same site

(Hyla latopalmata and H, wotjulumensis) jumped into the water when

disturbed, the H. meiriana moyed away from the water on to the dry slopes

where they sought refuge amongst the vegetation.

DISTRIBUTION

Hyla meiriana is currently known from five localities in the Northern

Territory and Western Australia. The nature of the terrain is such that this

species probably occurs in numerous disjunct populations completely isolated

from one another.

TYLER—NORTH AUSTRALIAN HYLA 7

NORTHERN TERRITORY RECORDS OF HYLA AUREA AND

H. ADELAIDENSIS

Moore (1961) reported the presence in the British Museum collection

of seven specimens of H. aurea raniformis from Port Essington that had

previously been examined by Gunther (1858), Boulenger (1882) and

Parker (1938). Moore (1961, p. 319) stated, “I would not believe the

locality to be correct, were it not for the fact that Copland (1957) has

seen specimens from Darwin, and Loveridge (1949) had specimens from

Knuckey’s Lagoon, which is 9 miles from Darwin”,

The Darwin specimens which Copland (1957, p. 58) includes in his

list of the H. aurea raniformis which he examined are N.M.V. D5529-30.

The former bears a tag labelled “Phractops sp..” but lacks an identification

in the museum register. The latter is labelled, “Hyla sp. young,” and is

registered as, “Hyla sp.", For reasons which are not apparent both were

despatched to Copland in 1956 as examples of Hyla aurea. D5529 is a

representative of Limnodynastes ornatus, and D5530 a Crinia sp. As there

are no other frogs in the N.M.Y. collection from the vicinity of Darwin

labelled aurea, the inclusion of these registration numbers in Copland’s

list can be attributed to a clerical error, and this particular record discounted,

Loveridge (1949) provided a brief description of the specimens from

Knuckey's Lagoon (M.C.Z. 25994-5) commenting that they were too

shrivelled to merit measuring. Examination has shown them to be examples

of Cyclorana dahli,

The presence of Hyla aurea in the Northern Territory therefore rests

solely on the British Museum series (B.M. 1936. 12.13.135-141), This

is not the only species whose presence in the Northern Territory has been

queried (Glauert, 1947), and it is pertinent to note that all of the specimens

involved were reported to have come from the same locality (Port

Essington) and the same source (Dr. Fleming). In view of the identity of

the specimens on which the subsequent reports were based it would seem

justifiable to now remove H. aurea from the Northern Territory checklist.

With the exception of the reports of aurea from the Northern Territory

considered above, the only additional record for any of the species first cited

by Gunther (1858) is probably that of Hyla adelaidensis reported by Mitchell

(1955, 1964). ‘The specimens involved have been examined and are

considered to represent the new species Hyla meiriana,

December, 1969

REC. 8S. AUST. MUS., 16 (1)

‘NAAT WsAISAM-YIIOU Jo susayjed uONNgiIsic]

“E “Old

g@ dnoi5

TYLER—NORTH AUSTRALIAN HYLA 9

REVISED HYLID CHECKLIST

(All species occur in northern Western Australia and the Northern Territory )

Hyla bicolor (Gray). Hyla nasuta (Gray).

Hyla caerulea (White). Hyla peroni (Tschudi).

Hyla coplandi Tyler, Hyla rubella Gray.

Hyla laitopalmata (Gunther). Hyla wotjulumensis Copland

Hyla meiriana new species.

PATTERNS OF DISTRIBUTION

The hylid frogs occurring in north-western Australia can be divided into

three groups according to their respective patterns of distribution. The

groups and their member species are as follows:

GROUP A: Species which are confined to areas with an annual

rainfall exceeding 30 inches (H. bicolor, H. meiriana,

A. nasuta).

GROUP B: Species which extend from the coast to approximately

the level of the twenty-inch isohyett (H. coplandi, H.

latopalmata, H. peroni, H. wotjulumensis) .

GROUP C: Species which are widely distributed throughout the

entire area and extend into the arid parts of Central Australia

with an annual rainfall of less than ten inches (H. caerulea,

H, rubella).

The first two groups form quite distinctive units but the third consists

of a pair of species which are so widely distributed and so morphologically

variable that each may ultimately merit sub-division.

The paucity of specimens from north-western Australia permits only

the broadest generalisations concerning distribution. At localities such as

Wotjulum, north of Kings Sound in Western Australia where several fairly

extensive collections have been made, eight of the nine species listed above

have been found, and the general pattern appears to be one of a gradual

reduction in the number of species away from the high rainfall coastal

localities. Thus all species occur in the area to which GROUP A are

confined, and members of GROUP C share the area occupied by GROUP B

(Fig. 3). The rather ubiquitous distribution of the species in GROUP C

affects the interpretation of these patterns. It is therefore relevant to briefly

summarize the problems involved.

10 REC, §&, AUST. MUS., 16 (1): [11 December. 1969

The status of the Central Australian population of Hy/a caerulea has

been the subject of controversy. Spencer (1896) considered it a distinct

species which he described as H. gilleni. Copland (1957) relegated it as

a sub-species of caerulea, and Moore (1961) failed to find grounds for even

the recognition of sub-species. The most recent contributor (Mertens, 1964)

has resurrected caertilea gilleni.

A comparable situation exists in the case of Leptodactylid with a similar

distribution pattern (Limnodynastes ornatus)., Parker (1940) described

L. spenceri from Central Australia, distinguishing it from the coastal ornatus

by its more extensive toe webbing, Moore (1961) suppressed spenceri,

but subsequent contributors (with the exception of Warburg, 1967) have

not adopted this proposil.

Another species which has a similar distribution is H. rubella, Speci-

mens from low rainfall areas tend to be larger, have broader heads and more

highly developed lateral digital fringes than those from peripheral high-

rainfall areas, However, there has not been any proposal that they should

merit taxonomic recognition at the specific or sub-specific level.

Examination of north-western specimens of caerulea and rubella indicate

that if distinct central and peripheral populations are recognized, the southern

boundary of the latter is similar to that of the species in GROUP B, but

for the purpose of the present discussion no subspecies ure recognized,

The hylid fauna of the north-western Australia has hitherto ( by

impheation if not by specific statement) been regarded as simply an extension

of the fauna of the north-east, with a gradual westward reduction in the

number of species. Utilizing the basic zoogeographical patterns of distribu-

tion of frogs adopted by Moore (1961), the north-western Hyla would be

divided into the Centralian species (corresponding to GROUP C), and those

confined to the “north-east crescent” (all remaining species), Of those in

the latter category, only Jatepalmata and peroni exhibit a continuous range

ucross the north of the continent, and then southwards along the eastern

seaboard. Ayla bicoler and nasuta range over the same area, but (as

suggested by Moore, 1961) they are probably separated into two disjunct

populations, haying yet to be reported from the southern margin of the

Gulf of Carpentaria, This area may not be a barrier to wotjulumensis, and

it is possible that this species occurs in north-western Queensland,

There is now an indication that the north-west possesses a distinctive

endemic clement in ils bylid frog fauna. Intensive collecting is needed to

establish the geographical distribution of the endeniic species more precisely,

but at present potential support for the recognition of north-western Australia

as a separate ynit within the Torresian zone is indicated.

TYLER—NORTH AUSTRALIAN HYLA a

SUMMARY

Hyla meiriana new species is described and reported from five localities

in the Northern Territory and Western Australia. Recent records of

H. adelaidensis and H. aicea in the Northern Territory are demonstrated to

be based on misidentified specimens. A checklist of north-western Hyla is

presented and distribution patterns of the component species are discussed.

ACKNOWLEDGMENTS

For the loan of the specimens cited in this paper | am indebted to Miss J. M. Dixon

(National Museum of Victoria), Dr. J. A. Peters (United States National Museum),

De. G, Storr (Western Australian Museum), and Dr. E. E, Williams (Museum of Comparative

Zoology). 4 am further indebted to Miss Dixon for abstracting data from her departmental

registration catalogue.

Mr. W. Harrington generously undertook the preparation of Fig. 3 and J am also

indebted to Mr, R, Ruehle for Fig. 2,

| would like to express my thanks to Mr. K. Cole and Messrs, R. Edwards, H. Bowshall

and A, Fleming for their collections obtained at Kununurra and Arnhem Land respectively,

und for detailed information on the type locality of the new species,

Finally my thanks are due to the Director and Board of the South Australian Museum

for the provision of laboratory and secretarial facilities, and to Mr, F. J, Mitchell for

advice and constructive criticism of the manuscript.

REFERENCES

Boulenger, G. A., 1882: Catalogue of the Batruchia Salientia s, Ecatdata in the collection

of the British Museum, 2nd ed., British Museum, London, 503 pp,

Copland, §, J., 1987: Australian tree-frogs of the genus Hyla. Proce, Linn, See, N.S, 82:

4-108,

Glauert, L., 1947; Some unfortunate errors in collecting localilies. HY, ust, Nat, 1: 48,

Gunther, A., 1858: Catalogue of the Batrachia Salientia in the collection of the British

Museum. London, xvi + 160 pp.

Loveridge, A., 1949: On some reptiles and amphibians from the Northern Territory, Trans.

Roy. Sac, S. Aust, 72; 208-215,

Mertens, R., 1964: Die systematische Stellung des zentralaustralischen Laubfrosehes //yla

gilleni. Senek, biol, 45: 15-21.

Mitchell, F. J,, 1955: Preliminary account of the Reptilia and Amphibia collected by the

National Geographic-Commonwealth Government-Smithsonian Institulion Expedi-

tion to Arnhem Land (April to November, 1948). Rec, S. Ausf. Mus. Is

373-408.

1964: Reptiles ane Amphibians of Arnhem Land. IN: Records of the American-

Australian Scientific Expedition to Arnhem Land. 4, Zoology, Ed, R. L.. Specht,

Melbourne Uniy. Press. pp, 309-343,

Moore, J, A. 1961: The frogs of eastern New South Wales, Bull, Amer, Mus. Nat. Hist.

121 (3): 149-386,

Parker, H, W,, 1938: The races of the Australian frog Hyla aurea Lesson, Ann. Mag, nat.

Hist, (10) 17: 66-93,

Spencer, B., 1896: Reporr on the work of the Horn Scientific Expedition to Central Australia,

2: Amphibia 2: 152-175, London, Dulau.

Tyler, M.J., 1968a: A taxonomic study of hylid frogs of the Myla lesueuvi complex occurring

in north-western Australia. Ree, S, Aust. Mus, 15: 711-727.

1968b; Papuan hylid frogs of the genus Hyla, Zool Verh, 96: 1-203.

Warbure, M,, 1967: On thermal and water balance of three Central Australian frogs.

Comp, Biochem. Physial, 2: 2745

RECORDS OF THE

SOUTH AUSTRALIAN

MUSEUM

AN ANNOTATED CATALOGUE OF THE

OTITIDAE (sens. lat.) (DIPTERA) RECORDED

FROM AUSTRALIA, INDONESIA AND THE

PACIFIC ISLANDS

By HARRY F. LOWER

Formerly of the Waite Agricultural Research Institute,

University of Adelaide, South Australia

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 2

6 March 1970

AN ANNOTATED CATALOGUE OF THE OTITIDAE (sens. lat.) (DIPTERA)

RECORDED FROM AUSTRALIA, INDONESIA AND THE PACIFIC

ISLANDS

BY HARRY F. LOWER,

FORMERLY OF THE WAITE AGRICULTURAL RESEARCH INSTITUTE,

UNIVERSITY OF ADELAIDE, SOUTH AUSTRALIA

Summary

The aim of this catalogue is to provide a list of all the described species of Otitidae recorded from

the area detailed below, together with all published taxonomic references to them. The catalogue

inclues all species described up to the end of 1964, and every reference mentioned has been read by

the author.

AN ANNOTATED CATALOGUE OF THE OTITIDAE (sens, lat.)

(DIPTERA) RECORDED FROM AUSTRALIA, INDONESIA AND

THE PACIFIC ISLANDS

By HARRY F. LOWER

FORMERLY OF THE WAITE AGRICULTURAL RESEARCH INSTITUTE,

UNIVERSITY OF ADELAIDE, SOUTH AUSTRALIA*

1. INTRODUCTION

The aim of this catalogue is to provide a list of all the described

species of Otitidae recorded from the area detailed below, together with all

published taxonomic references to them. The catalogue includes all species

described up to the end of 1964, and every reference mentioned has been

read by the author.

Although it is well over 100 years since the first species were described

from the area, confusion of genera, species, and their respective distribu-

tions still exists. The vast taxonomic literature on the family is scattered

both in time, and in a wide range of publications, and is in a variety of

European languages; few collections of otitid material in institutions have

received modern taxonomic study; and some types of genera and species

were described, at the time, in a few lines sufficient for their validation, but

Wholly inadequate for recognition purposes. Where such genera and species

occur in this catalogue, a note at the end of each points out the difficulties

encountered and directs attention to the most fruitful lines to be followed in

order to clear up doubts. As would be expected with small insects, prac-

tically every large collection of the family contains undescribed species.

The selected area corresponds with no accepted geographic region,

It was chosen primarily because of the author’s interest in the sub-family

Platystomimae which appears to have evolyed in the area, and to which,

most of its members are naturally confined. The area is extensive: on the

west und east, it is bounded by the 90°E and 135°W meridians, respectively;

its north and south limits are latitudes 30°N and 60°S. No part of

mainland Asia is included; where such is referred to, it is because the

distribution of certain species extends from adjacent land masses to the

mainland. The land masses of the area are thus Australia and New Zealand,

Indonesia, the Australian mandated territories of Papua and New Guinea,

the Philippine Islands, and the numerous scattered islands of the western

Pacific Ocean.

* Now Department of Agriculture, Adelaide

6 March 1970

REC, S&S AUST. MUS., lo (2): 1-93 March, 1970

io]

2, SOURCES

This catalogue is based on the work of Hendel who, in several fascicles

of Genera Insectorum (notably No. 157, Hendel, 1914a) and in his Die

Arten der Platystominen (Hendel, 1914b), reviewed all the species of which

he was aware up to that time, To these, I have added species overlooked

by him, corrected names where these have later been shown to be in error,

and have added species described since then chiefly by American and

European workers to whom material was submitted for identification either

by private persons or scientific expeditions. The results of several European

scientific surveys have also been incorporated.

Comparison of Hendel’s two texts referred to above shows that he

intended publication of Die Arten der Platystominen to precede that of

fascicle 157 of Genera Insectorum, since, in the former he described his

new genera and species, and referred to them in the latter as already

published. As it so happened, however, fascicle 157 of Genera Insectorum

was published on 15th April, 1914 while Die Arten der Platystominen was

not published until 15th June of the same year, This resulted in all of

Hendel’s new names in Genera Insectorum being nomina nuda which were

not validated until the later publication of Die Arten der Platystominen

(Abh, zool-bot Ges. Wien 8). The possibility of prior publication by others

therefore exists. While this possibility is a remote one, it still should be

borne in mind by anyone working on the family.

Hendel’s work laid the foundation on which later taxonomists have

built, His ability to find morphological characters for delimiting genera,

und to make clear the relation of one genus to another, were gifts of a high

order, He did, however, have occasional lapses, as, for example, in

re-ullocating some of the earlier workers’ species to the genera he accepted,

He seems, not infrequently, to have seen neither the types, nor specimens

known to be conspecific with them, and the original descriptions, which he

often quotes verbatim, were his only guide, These descriptions, written

before modern concepts of generic and specific limits had been elaborated,

were, all too frequently, insufficient for identification purposes. Such

blemishes may possibly be attributed to the disturbed conditions prevailing

in Europe at the time he was preparing his works for publication, While

these doubtful species are here included as Hendel placed them, they should

preferably be treated as “genus unknown", until their placement is either

confirmed, or they can be placed in their correct genera by future workers.

To avoid repetition, citation of Hendel’s two major works, in the body

of the text, indicates that references to a genus or species published before

1914 will be found theretn. Die Arten der Platystominen is abbreviated

to Hendel 1914a and fascicle 157 of Genera Insectorum to Hendel 191 4b,

LOWER—AUSTRALIAN REGION OTITIDAE 3

The number following each corresponds with the pagination. This has

enabled me to reduce references to works published just before 1914, too

late to be included by him, and to those which have been published since.

Students of the Otitidae of Australia and New Guinea will find the

series of papers by J. R. Malloch in Proc. Linn. Soc. N.S.W. essential,

In addition to the genera and species from this area listed by Hendel,

Malloch includes a considerable number of genera and species described by

himself and others after 1914.

3. NOMENCLATURE

At the present time the nomenclature of the Otitidae is receiving new

study, Hendel treated the group as a single family consisting of several

subfamilies; the modern trend, as exemplified by Steyskal in the U,S,A., is

to raise some, at least, of these to family status. To what extent this lead

will be followed, the future alone can decide. In these circumstances, the

author has retained the Hendelian system; in practice, this will cause no

difficulties for workers in the group.

The sub-family, Platystominae (family Platystomatidae of Steyskal),

with 423 described species, is the dominant one in the area, to which nearly

all of these species are endemic. In addition, 19 species of other sub-familics

(Otitinae, 1; Ulidiinae, 18), have been recorded.

Of these 19 species, six are known to be introductions, and some

degree of doubt attaches to the remainder. The position is further confused

by misidentifications through which species not occurring in the area have

been recorded as so doing. The probability is high that some records may

still be based on misidentifications.

In this catalogue, the arrangement and numbering of genera are those

used by Hendel in fascicles 157 (1914, Platystominae) and 106 (1910,

Ulidiinae) of Genera Insectorum. Missing numbers represent genera none

of whose species have been recorded from the area. This applies, for

example, in the Platystominae, to the first eight genera: the list thus begins

with Hendel’s genus No. 9,

A letter following a number indicates that the genus has been erected

since Hendel's work was published, or is one of which he was unaware when

he wrote. An asterisk preceding a specific name indicates an “Australian”

species, that is, one recorded from Australia, or New Guinea and its

adjacent islands.

Within genera, species are arranged in alphabetical order.

Family: OTITIDAE

Type genus: Qtites Latreille, 1804 Nouv. Dict. Hist, nar. 24: 196.

4 REC, S. AUST. MUS., 16 (2): 1-93 March, 1970

I. Sub-family: PLATYSTOMINAE

Type genus: Platystoma Meigen, 1803 Mag. Insektenkunde (Illiger) 2: 277.

Note: Both Curran (1934) and Malloch (1939) claimed that

Platystoma was not available for use in the Diptera since it “is pre-occupied

in Mollusca.” Neither gave any reasons for making this assertion, nor

referred to any authority. In Neave (Nomenclator Zoologicus 3: 802-803),

Platystoma Meigan, 1803 pre-dates, by many years, the earliest of the six

later applications of Platystoma to other animal groups. Steyskal in all his

publications uses Platystema without any query as to the correctness of such

usage, and the name was accepted by workers who preceded Curran

(e.g., Williston, 1888 and later editions; Hendel A, B). In these circum-

stances, Curran’s statement appears to be incorrect; Malloch apparently

followed him without checking the correctness of the statement since he used

the exact wording of Curran.

GENERA

9, Genus: POECILOTRAPHERA Hendel, 1914

Hendel, 1914a Gen. Ins. 157: 33.

Type species: Urophora taeniata Macquart, 1843.

1. P. taeniata (Macq. 1843)

Urophora taeniata Macq., 1843 Dipt. exot, 2: 222 Pl. 30, Fig. 6. Bezzi,

1913 Mem. Indian Mus. 3: 80 (No. 259).

Poecilotraphera taeniata (Macq. 1843), Hendel, 1914b :21; 1914a :33.

Enderlein, 1924 Mitt. Zool, Mus. Berlin 11: 100.

Distribution: Borneo, Java, China.

The only other described species is P. comperei (Coq, 1904) from

India,

Il. Genus: XIRIA Walker, 1857

Xiria Walker, 1857 J. Proc. Linn. Soe. Lond. 1: 36.

Type species: NXiria antica Walk., 1857.

Hendel, 1914b :24; 1914a :35,

1, X, antica Walker, 1857

Xiria antica Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 36 Pl. 2, Fig. 2.

Bezzi, 1913 Mem. Indian Mus. 3: 67 (No. 20). Hendel, 1914b :24;

1914a :37.

Distribution: Sumatra.

LOWER—AUSTRALIAN REGION OTITIDAE 5

2. X. clarissa Frey, 1930

Xiria clarissa Frey, 1930 Notul. ent. Helsingf. 10; 62 Pl. 1, Fig. 7.

Distribution: Philippines.

3. X. lavinia van der Wulp, 1898

Xiria lavinia v. d. Wulp, 1898 Tijd. Ent. 51: 210 Pl. 10, Figs. 8, 9.

Enderlein, 1912 Zool. Jahrb. 33: 361. Hendel, 1914b :25; 1914a :37.

Distribution: Java, Sumatra.

4. X. limbata de Meijere, 1924

Xiria limbata de Meijere, 1924 Tijd. Ent. 67 (Suppl.): 40.

Distribution: Sumatra.

5. X. obliqua Osten-Sacken, 188]

Xiria obliqua Osten-Sacken, 1881 Ann. Stor. nat. Mus. Genova 16: 463.

Hendel, 1914b :25; 1914a :37,

Distribution: Sumatra.

*6, X,. papuana Hennig, 1940

Xiria papuana Hennig, 1940 Arb. morph. Taxon. Ent. Berl. 71: 316.

Distribution: New Guinea.

*7. X. strigata Hennig, 1940

Xiria strigata Hennig, 1940 Arb. morph. Taxon. Ent. Berl. 71: 317.

Distribution: New Guinea.

8. X. violacea (Wied., 1830)

Trypeta violacea Wied., 1830 Ausser. Zweifl. Ins. 2: 476. Bezzi, 1913

Mem. Indian Mus. 3: 81 (No. 281).

Niria violacea (Wied., 1830) Hendel, 1914b :26; 1914a :37.

Distribution: Java, Sumatra.

12. Genus: LASIOXIRIA Hendel, 1914 (Monotypic)

Hendel, 1914a Gen. Ins. 157: 37.

Type species: L. hirsuta Hend. 1914.

Hendel, 1914b :28.

*1. L. hirsuta Hendel, 1914

Lasioxiria hirsuta Hendel, 1914b :28; 1914a :37. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 101.

Distribution: New Guinea.

Malloch, 1939 suggested that Lasioxiria might be a synonym of

Dasyortalis Hendel, 1913.

6 REC. S. AUST, MUS., 16 (2): 1-93 March, 1970

13. Genus: CONICIPITHEA Hendel, 1914 (Monotypic)

Hendel, 1914a :40.

Type species: Dacus addens Walker, 1860.

J. C. addens (Walker, 1860)

Dacus addens Walker, 1860 J. Proc. Linn. Soc. Lond. 4: 149. Bezzi, 1913

Mem. Indian Mus. 3: 66 (No. 5).

Conicipithea addens (Walk., 1860) Hendel, 1914b :29. Malloch, 1939

Proc, Linn, Soc, N.S.W. 64: 103. Hardy, 1959 Bull, Brit. Mus. (nat.

Hist.) Ent. 8 (5): 163.

Distribution: Amboina (Moluccas), Macassar (Celebes).

14. Genus: PHILOCOMPUS Osten-Sacken, 1881

Osten-Sacken, 1881 Ann. Soc. ent. France (6th Ser.) 1: 134,

Type species: Philocompus cupidus Osten-Sacken, 1881. Hendel,

1914b :30; 1914a :42. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 104.

1. P. aeneus de Meijere, 1906

Philocompus aeneus de Meijere, 1906 Ann. Mus. nat. Hung, 4: 187 Pl. 2,

Fig. 16. Hendel, 1914b :30; 1914a :42.

Distribution: Bali, Java.

2. P. cupidus Osten-Sacken, 1882

Philocompus cupidus Osten-Sacken, 1882 Berl. ent. Zeit. 26: 217. Hendel,

1914b :30; 1914a :42. Frey, 1930 Notul. ent. Helsingf. 10: 46.

Distribution: Philippines.

3, P. divergens (Walk., 1860)

Dacus divergens Walker, 1860 J. Proc. Linn. Soc. Lond. 4: 149. Bezzi,

1913 Mem. Indian Mus. 3: 70 (No. 84).

Philocompus divergens (Walk., 1860) Hendel, 1914b :42; 1914a :42,

Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 170,

Distribution: Macassar (Celebes).

15. Genus: ANTINEURA Osten-Sacken, 1881

Osten-Sacken, 1881 Ann. Soc. ent. France (6th Ser.) 1: 134.

Type species: Antineura stolata Osten-Sacken, 1882. Hendel, 1914b

732: 1914a :400. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 104.

LOWER—AUSTRALIAN REGION OTITIDAE T

+1. A. biroi de Meijere, 1906

Antineura biroi de Meijere, 1906 Ann. Mus. nat. Hung. 4: 188 Pl. 2, Fig.

17. Hendel, 1914b :33; 1914a :41. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 104.

Distribution: New Guinea,

*2. A, devia (Walk., 1861)

Dacus devius Walker, 1861 J, Proc. Linn. Soc. Lond. 5: 250. Bezzi, 1913

Mem. Indian Mus. 3: 70 (No. 78).

Antineura devia (Walk., 1861) Hendel, 1914b :36; 1914a :41. Hardy,

1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 169.

Distribution: New Guinea.

3. A. grandis (Dol., 1858)

Herina grandis Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 126.

Antineura grandis (Dol, 1858) Hendel, 1914b :35; 1914a :42. Malloch,

1939 Proc. Linn. Soc, N.S.W, 64; 104.

Distribution; Moluccas.

*4. A. kerteszi de Meij., 1906

Antineura kerteszi de Meijere, 1906 Ann. Mus. nat. Hung. 4: 189 Pl. 2,

Fig. 18. Hendel, 1914b :34; 1914a :42. Enderlein, 1924 Mitt. zool.

Mus. Berlin 11: 110. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 104

Pl. 4, Fig. 4.

Distribution: New Guinea.

5. A. pubiseta (Walk., 1861)

Dacus pubiseta Walker, 1861 J. Proc. Linn. Soc, Lond. 5: 294. Bezzi,

1913 Mem. Indian Mus. 3: 78 (No. 214).

Antineura pubiseta (Walk., 1861) Hendel, 1914b :36; 1914a :42. Hardy,

1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 181.

Distribution; Moluccas.

6. A. sericata Osten-Sacken, 1882

Antineura sericata Osten-Sacken, 1882 Berl. ent. Zeit, 26: 216. Hendel,

1914b :33; 1914a :42. Bezzi, 1917 Philippine J. Sc. (D) 12: 133.

Distribution: Philippines.

7. A. stolata Osten-Sacken, 1882

Antineura stolata Osten-Sacken, 1882 Berl. ent. Zeit. 26: 215. Hendel,

1914b :32; 1914a :42. Frey, 1930 Notul. ent. Helsingf., 10: 49.

Distribution: Philippines.

8 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

8. A. strigifer (Walk., 1862)

Dacus strigifer Walker, 1862 J. Proc. Linn. Soc. Lond. 6: 13. Bezzi, 1913

Mem. Indian Mus. 3: 80 (No, 252).

Antineura strigifer (Walk, 1862) Hendel, 1914b :37; 1914a :42, Hardy,

1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 183.

Distribution: Moluccas.

I have rejected Hendel’s sub-division of this genus into the two sub-

genera, Antineura (sens. str.) and Adantineura. At least two species were

described from single specimens and neither species has since been recorded;

the genus is not well known, and still awaits revision.

ISA. Genus: PSEUDOCLEITAMIA Malloch, 1939 (Monotypic)

Malloch, 1939 Proc. Linn. Soc, N.S.W. 64: 104.

Type species: Pseudocleitamia setigera Malloch, 1939.

*1, P. setigera Malloch, 1939

Pseudocleitamia setigera Malloch, 1939 ibid.

Distribution: New Guinea.

16, Genus: KENASPIS Osten-Sacken, 1881

Osten-Sacken, 1881 Ann. Soc. ent. France (6th Ser.) 1: 134.

Type species: Xenaspis polistes Osten-Sacken, 1882.

1. X. extranea Bezzi, 1917

Xenaspis extranea Bezzi, 1917 Philippine J. Sci. (D) 12: 134. Frey, 1930

Notul. ent. Helsingf. 10: 49.

Distribution: Philippines.

2. X. homichlodes Hend., 1914

Xenaspis homichlodes Hendel, 1914b :41; 1914a :44,

Distribution: Borneo,

3. X. pictipennis (Walker, 1849)

Oxycephala (?) pictipennis Walker, 1849 List. Dipt. Brit. Mus. 4: 1162.

Xenaspis vespoides de Meijere, 1904 Bijd. Dierk. 17: 107, Figs. 19, 20.

Polistomima gigantea Enderlein, 1912 Zool. Jahrb, 33: 353, Fig. C.

Hendel, 1914b :39; 1914a :44,

Distribution: India, Indonesia.

LOWER—AUSTRALIAN REGION OTITIDAE 9

4. X. polistes Osten-Sacken, 1882

Xenaspis polistes Osten-Sacken, 1882 Berl. ent. Zeit. 26: 220. Hendel,

1914b :38; 1914a :44. Bezzi, 1917 Philippine J. Sci. (D) 12: 134.

Frey, 1930 Notul. ent. Helsingf. 10: 49.

Distribution: Philippines.

5. X. stigma (Enderlein, 1912)

Polistomima stigma Enderlein, 1912 Zool. Jahrb, Syst. 33: 352 Fig. B.

Xenaspis stigma (End., 1912) Hendel, 1914b :42; 1914a :44,

Distribution: Sumatra.

6. X. walkeri (End., 1912)

Polistomima walkeri Enderlein, 1912 Zool. Jahrb. Syst. 33: 350 Fig. A.

Xenaspis walkeri (End., 1912) Hendel, 1914b :42; 1914a :45.

Distribution: Sumatra.

This is a widely-dispersed genus: of its fifteen described species, six

have been recorded from India as far north as the Himalaya, one from

Burma, three from the Indonesian islands, two from the Philippines, two

from Taiwan, and one occurs both in India and Indonesia. It is likely that

new species will yet be found in New Guinea.

16A. Genus: XENASPOIDES Frey, 1930

Frey, 1930 Notul. ent. Helsingf. 10: 49.

Type species: Xenaspoides ichneumonea Frey, 1930

1. X. cyanea Frey, 1930

Xenaspoides cyanea Frey, 1930 Notul. ent. Helsingf. 10: 50 Pl. 1, Fig. 2.

Distribution: Philippines.

2. X. ichneumonea Frey, 1930

Xenaspoides ichneumonea Frey, 1930 Notul. ent. Helsingf. 10: 50 Pl, 1,

Fig. 1.

Distribution: Philippines.

17. Genus: LAMPROPHTHALMA Portschinsky, 1892

Portschinsky, 1892 Horae Soc. ent. Rossicae 26: 225.

Type species: L. metallica Ports., 1892.

10 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

1. L. cavenda Bezzi, 1917

Lamprophthalma cavenda Bezzi, 1917 (? publication) Frey, 1930 Notul.

ent. Helsingf. 10: 46.

Distribution: Philippines.

I have been unable to find Bezzi’s original description. My only

reference to the species is the specific name, author and date given in a list

by Frey (loc. cit.).

2. L. doleschalli (End., 1912)

Senopterina doleschalli Enderlein, 1912 Zool. Jahrb. Syst. 33: 357.

Lamprophthalma doleschalli (End., 1912) Hendel, 1914b :49; 1914a :45.

Distribution: Sumatra.

3. L. meijereana (End., 1912)

Senopterina meijereana Enderlein, 1912 Zool. Jahrb. Syst. 33: 356.

Lamprophthalma meijereana (End., 1912) Hendel, 1914b :49; 1914a :45.

Distribution: Sumatra.

4. L. sepedonoides (Walk., 1864)

Dacus sepedonoides Walker, 1864 J. Proc. Linn. Soc. Lond. 7: 228.

Lamprophthalma sepedonoides (Walk., 1864) Hardy, 1959 Bull. Brit. Mus.

(nat. Hist.) Ent. 8 (5): 181.

Distribution: Moluccas.

5. L. tuberculifrons de Meijere, 1933

Lamprophthalma tuberculifrons de Meijere, 1933 Tijd. Ent. 76: 111.

Distribution: Java.

17A. Genus: APACTONEURA Malloch, 1930 (Monotypic)

Malloch, 1930 Ins. Samoa 6 (5): 223.

Type species: Apactoneura flavicornis Malloch, 1930.

1. A. flavicornis Malloch, 1930

Apactoneura flavicornis Malloch, 1930 Ins. Samoa 6 (5): 223 Fig. 3.

Distribution: Samoa.

18. Genus: PLAGIOSTENOPTERINA Hendel, 1914

Hendel, 1914a Gen. Ins. 157: 46. Malloch, 1931 Proc. U.S. nat. Mus.

78: 12.

Type species: Dacus aeneus Wiedemann, 1819.

LOWER—AUSTRALIAN REGION OTITIDAE in

*1. P. aenea (Wied., 1819)

Dacus aeneus Wiedemann, 1819 Zool. Mag. 3: 29; 1830 Ausz. zweifl. 2:

512.

Plagiostenopterina aenea (Wied., 1819) Hendel, 1914b :54; 1914a 248.

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 107. Malloch, 1928 Proc.

Linn. Soc. N.S.W. 53: 353; 1939 ibid 64: 114.

Distribution: Java; Krakatau; Sumatra; Borneo; Philippines; New

Guinea; Queensland (Aust.); Bismarck Archipelago; Ceylon; India; Taiwan.

A wide-spread common species. See Meringomeria Enderlein, 1924

(Genus No. 18a).

2. P. armata Malloch, 1931

Plagiostenopterina armata Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 20.

Distribution: Philippines.

*3. P. basalis (Walk., 1849)

Dacus basalis Walker, 1849 List Dipt. Brit. Mus. 4: 1072. Bezzi, 1913

Mem. Indian Mus. 3: (No. 29).

Plagiostenopterina basalis (Walker, 1849) Hendel, 1914b :64; 1914a :49.

Hardy, 1959 Bull. Brit, Mus. (nat. Hist.) Ent. 8 (5): 164.

Distribution: Northern Territory (Australia).

4. P. calearata (Macq., 1843)

Herina calcarata Macquart, 1843 Dipt. exot. 2: 207 Pl. 28, Fig. 3.

Plagiostenopterina calcarata (Macq., 1843) Hendel, 1914b :63; 1914a :49.

Bezzi, 1913 Philippine J. Sci. (D) 8: 321. Malloch, 1931 Proc. U.S.

nat. Mist. 78 (15): 20.

Distribution: Indonesia; Philippines.

5. P. diptera Malloch, 1931

Plagiostenopterina diptera Malloch, 1931 Proc. U.S, nat. Mus. 78 (15): 18.

Distribution: Philippines.

6. P. discolor Malloch, 1931

Plagiostenopterina discolor Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 21.

Distribution: Philippines.

7. P. dubiosa Malloch, 1931

Plagiostenopterina dubiosa Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 15.

Distribution; Philippines; Ceylon; Singapore.

12 REC. S, AUST, MUS., 16 (2): 1-93 March, 1970

8. P. egregia de Meijere, 1924

Plagiostenopterina egregia de Meijere, 1924 Tijd. Ent. 67 (Suppl.): 41.

Distribution: Sumatra.

*Q. P. enderleini Hendel, 1914

Plagiostenopterina enderleini Hendel, 1914b :56; 1914a :49. Enderlein,

1924 Mitt. zool. Mus. Berlin 11: 108. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 114 PI. 4, Fig. 14.

Distribution: New Guinea; Sumatra; Ceylon.

10. P. farinosa Hendel, 1914

Plagiostenopterina farinosa Hendel, 1914b :68; 1914a 749.

Distribution: Moluccas.

11. P. hebes Hendel, 1914

Plagiostenopterina hebes Hendel, 1914b :70; 1914a :49.

Distribution: Singapore.

12. P. imitans (Walk., 1860)

Dacus imitans Walker, 1860 J. Proc. Linn. Soc. Lond. 4: 150. Bezzi, 1913

Mem. Indian Mus. 3: 73 (No. 132).

Plagiostenopterina imitans (Walk., 1860) Hendel, 1914a :49. Hardy, 1959

Bull. Brit. Mus (nat. Hist.) Ent. 8 (15): 175.

Distribution: Celebes.

13. P. inapta (Walk., 1860)

Dacus inaptus Walker, 1860 J. Proc. Linn. Soc, Lond. 4: 151, Bezzi, 1913

Mem. Indian Mus. 3: 73 (No. 135).

Plagiostenopterina inapta (Walk., 1860) Hardy, 1959 Bull. Brit. Mus, (nat.

Hist.) Ent. 8 (5): 175.

Distribution: Celebes.

14. P. inermis Malloch, 1931

Plagiostenopterina inermis Malloch, 1931 Proc. U.S. nat. Mus.'78 (15): 21.

Distribution: Philippines.

*15. P. lativentris (Walk., 1859)

Dacus lativentris Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 115. Bezzi,

1913 Mem. Indian Mus. 3: 74 (No. 144).

LOWER—AUSTRALIAN REGION OTITIDAE 13

Plagiostenopterina orbitalis Malloch, 1939 Proc. Linn. Soc. N.S.W. 64; 114

PL. 4, Fig. 15. Plagiostenopterina lativentris (Walk., 1859) Hardy, 1959

Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 177.

Distribution: New Guinea.

16. P. longivitta (Walk., 1859)

Dacus longivitta Walker, 1859 J. Proc, Linn. Soc. Lond. 3: 115. Bezzi,

1913 Mem. Indian Mus. 3: 74 (No. 158).

Plagiostenopterina longivitta (Walk., 1859) Hendel, 1914a :49. Enderlein,

1924 Mitt. zool. Mus. Berlin 11: 108. Hardy, 1959 Bull. Brit. Mus.

(nat. Hist.) Ent. 8 (5): 178.

Distribution: Aru Islands; India.

17, P. marginata (v. d. Wulp, 1880)

Senopterina marginata v. d. Wulp, 1880 Tijd. Ent. 23: 179 Pl. 10, Fig. 13.

Plagiostenopterina marginata (v. d. Wulp, 1880) Hendel, 1914b :66;

1914a :49,

Distribution; Java; China.

See Meringomeria Enderlein, 1924 (Genus No. 18a).

18. P. medionotata de Meijere, 1924

Plagiostenopterina medionotata, 1924 Tijd. Ent, 67 (Suppl.): 40.

Distribution: Sumatra.

19. P. neurostigma Bezzi, 1928

Plagiostenopterina neurostigma Bezzi, 1928 Diptera . . . Fiji Islands :89.

Distribution: Fiji.

20. P. nigricostata (Doleschall, 1858)

Herina nigricostata Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 126.

Plagiostenopterina nigricostata (Dol., 1858) Hendel, 1914b :70; 1914a :49.

Distribution: Moluccas.

21. P. pallidipes Frey, 1930

Plagiostenopterina pallidipes Frey, 1930 Notul. ent. Helsingf. 10: 51.

Distribution: Philippines.

*22. P. parva Mall., 1931

Plagiostenopterina parva Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 15:

1939 Proc. Linn, Soc, N.S.W. 64: 114.

Distribution: New Guinea.

14 REC. 8. AUST. MUS., 16 (2): 1-93 March, 1970

23. P. plagiata (Bezzi, 1917)

Elassogaster plagiata Bezzi, 1917 Philippine J. Sci. (D) 12: 135. Frey,

1930 Notul. ent. Helsingf. 10: 46.

Plagiostenopterina plagiata (Bezzi, 1917) Malloch, 1931 Proc. U.S. nat.

Mus. 78 (15): 19.

Distribution: Philippines.

24. P. planidorsum (Walker, 1860)

Charax planidorsum Walker, 1860 Trans. ent. Soc. Lond, 5: 325, Hendel,

1914b :53 (footnote).

Plagiostenopterina planidorsum (Walker, 1860) Frey, 1930 Notul. ent.

Helsingf. 10: 51.

Distribution: Philippines; Burma,

25. P. rutila Hendel, 1914

Plagiostenopterina rutila Hendel, 1914b :61; 1914a :49.

Distribution: Lombok.

26. P. samoaensis Malloch, 1930

Plagiostenopterina samoaensis Malloch, 1930 Ins. Samoa 6 (5): 230; 1931

Proc. U.S. nat. Mus. 78 (15): 15.

Distribution: Samoa.

27. P. trivittata (Walk., 1849)

Dacus trivittatus Walker, 1849 List Dipt. Brit. Mus, 4: 1072. Bezzi, 1913

Mem. Indian Mus. 3: 80 (No. 266).

Plagiostenopterina trivittata (Walk., 1849) Hendel, 1914b :65; 1914a :49.,

Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 183.

Distribution: Singapore; Malacca; Hongkong; Philippines.

See Meringomeria Enderlein, 1924 (Genus No. 18A).

28. P. trivittigera Malloch, 1931

Plagiostenopterina trivittigera Malloch, 1931 Proc. U.S. nat. Mus. 78

(15): 17,

Distribution: Singapore.

I8A, Genus: 7?MERINGOMERIA Enderlein, 1924

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 108.

Type species: Dacus trivittatus Walker, 1849 List Dipt. Brit. Mus.

4: 1072.

LOWER—AUSTRALIAN REGION OTITIDAE 15

The status of this genus is doubtful. It was erected by Enderlein in

1924 for four species. Three of these he transferred from Plagiostenopterina,

namely P. trivittata (Walker, 1849); P. marginata (v. d. Wulp, 1880);

and P. aenea Hendel (sic), 1914, The latter species is unknown to me;

it is possibly a /apsus calami for P. aenea (Wied., 1819). The fourth

species was Meringomeria interrupta Enderlein, 1924, which he described

from Sikkim in India—Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 109.

This genus is inserted here to draw attention to the need for a thorough

investigation of the four species. It is probable that all four will eventually

be placed in Plagiostenopterina.

18B. Genus: 7PERONOTROCHUS Enderlein, 1924

This genus was erected by Enderlein, 1924 Mitt. zool, Mus. Berlin

11: 109 for the single species Plagiostenopterina calcarata (Macq., 1843).

The status of the genus is in doubt.

Frey, 1930 (Notul. ent. Helsingf. 10: 52) accepted the genus as valid

and described the following species:

1. P. inermis Frey, 1930

Peronotrochus inermis Frey, 1930 Notul. ent. Helsingf. 10: 52.

Distribution: Philippines.

18C. Genus; IMUGANA Enderlein, 1937

Enderlein, 1937 S.B. Ges. naturf. Fr. Berl. (year 1936): 435.

Type species: Imugana pompiliformis Enderlein, 1937,

1. I. metallica Enderlein, 1937

Imugana metallica Enderlein, 1937 $.B, Ges. natur}. Fr. Berl. ( year 1936):

436,

Distribution: Philippines.

2, I. pompiliformis Enderlein, 1937

Imugana pompiliformis Enderlein, 1937 S.B. Ges. naturf. Fr. Berl. (year

1936): 435.

Distribution; Philippines.

19. Genus: ELASSOGASTER Bigot, 1859

Bigot, 1859 Ann. Soc. ent. Fr. (Ser. 3) 8: 546.

Type species: Elassogaster metallicus Bigot, 1859.

16 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*1. E. albopilosus de Meijere, 1915

Elassogaster albopilosus de Meijere, 1915 Tijd. Ent. 58: 133.

Distribution: New Guinea.

*2, E. didymoides Hendel, 1914

Elassogaster didymoides Hendel, 1914b :76; 1914a :52. Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 117.

Distribution: New Guinea.

*3. E. didymus (Osten-Sacken, 1881)

Senopterina didyma Osten-Sacken, 1881 Ann, Mus. Stor. nat. Genova 16:

465,

Elassogaster didymus (Osten-Sacken, 1881) Hendel, 1914b :75; 1914a :51.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 116.

Distribution: New Guinea.

*A_ E. evitta Malloch, 1939

Elassogaster evitta Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 116 Pl. 4,

Fig. 16.

Distribution: New Britain.

5. E. flavipes (Schiner, 1868)

Senopterina flavipes Schiner, 1868 Novara Dipt. :288.

Elassogaster flavipes (Schiner, 1868) Hendel, 1914b :78; 1914a :52.

Distribution: Singapore.

6. KE. hyalipennis Malloch, 1931

Elassogaster hyalipennis Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 23.

Distribution: Philippines.

*7. E, lineatus de Meijere, 1915

Elassogaster lineatus de Meijere, 1915 Tijd. Ent. 58: 132.

Distribution; New Guinea.

*8. E. marginalis Malloch, 1940

Elassogaster marginalis Malloch, 1940 Ann. Mag. nat. Hist 6: 68.

Distribution: Solomon Islands.

LOWER—AUSTRALIAN REGION OTITIDAE 17

*9, E. nigripes Malloch, 1940

Elassogaster nigripes Malloch, 1940 Ann. Mag. nat. Hist. 6: 70.

Distribution: Solomon Islands.

10. E. potens Frey, 1930

Elassogaster potens Frey, 1930 Notul. ent. Helsingf. 10: 52.

Distribution: Philippines.

*11. E. sepsoides (Walker, 1861)

Dacus sepsoides Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 163. Bezzi,

1913 Mem, Indian Mus, 3: 79 (No. 236).

Elassogaster sepsoides (Walker, 1861) Hendel, 1914b :82; 1914a :52,

Bezzi, 1913 Philippine J. Sci. (D) 8: 321. Malloch, 1928 Proc. Linn.

Soc. N.S.W. 53: 351 Fig. 4. Frey, 1930 Notul. ent. Helsingf. 10: 52.

Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 22; 1939 Proc. Linn. Soc.

N.S.W. 64: 115. Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5):

181.

Distribution: New Guinea; Taiwan; Amboina; Ceylon; Queensland

(Australia).

12, E. signatipes (Walker, 1861)

Dacus signatipes Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 163.

Elassogaster signatipes (Walker, 1861) Hendel, 1914a :52. Bezzi, 1913

Mem. Indian Mus, 3: 79 (No. 239). Hardy, 1959 Bull. Brit. Mus (nat.

Hist.) Ent. 8 (5): 181.

Distribution: Amboina.

13, E. simplex Frey, 1930

Elassogaster simplex Frey, 1930 Notul. ent. Helsingf. 10: 53,

Distribution: Philippines.

*14. E. sordidus (Walker, 1861)

Dacus sordidus Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 251. Bezzi,

1913 Mem. Indian Mus. 3: 79 (No. 244).

Dacus varialis Walker, 1865 Bezzi, 1913 ibid. 3: 79 (No. 275).

Elassogaster sordidus (Walker, 1861) Hardy, 1959 Bull. Brit. Mus. (nat.

Hist.) Ent. 8 (5): 182.

Distribution: New Guinea.

Dacus varialis Walker, 1865 is a synonym of Dacus sordidus Walker,

1861. Hendel, 1914a Gen. Ins. 157: 52 refers to it as Elassogaster varialis

(Walker, 1865). Hardy, 1959 showed the synonymy.

18 REC. 8. AUST. MUS., 16 (2): 1-93 March, 1970

*15. E. terrae-reginae Malloch, 1928

Elassogaster terrae-reginae Malloch, 1928 Proc. Linn. Soc. N.S.W. 53; 352;

1931 Proc. U.S. nat. Mus. 78 (15): 22; 1939 Proc. Linn. Soc. N.S.W.

64: 116.

Distribution: Queensland; New South Wales (Australia).

19A. Genus: PICROMETOPUS Frey, 1930 (Monotypic)

Frey, 1930 Notul. ent. Helsingf. 10: 53.

Type species: Picrometopus bicolor Frey, 1930.

1, P. bicolor Frey, 1930

Picrometopus bicolor Frey, 1930 (loc. cit.).

Distribution: Philippines.

20. Genus: MICROEPICAUSTA Hendel, 1914 (Monotypic)

Hendel, 1914a Gen. Ins. 157: 52.

Type species: Microepicausta gracilis Hendel, 1914.

*1. M. gracilis Hendel, 1914

Microepicausta gracilis Hendel, 1914b :85; 1914a :52.

Distribution: New South Wales (Australia).

21. Genus: SCELOSTENOPTERINA Hendel, 1914 (Monotypic)

Hendel, 1914a Gen. Ins. 157: 54.

Type species: Scelostenopterina femorata Hendel, 1914.

1. S. femorata Hendel, 1914

Scelostenopterina femorata Hendel, 1914b :86; 1914a :55. Bezzi, 1917

Philippine J. Sci. (D) 12: 136. Frey, 1930 Notul. ent. Helsingf. 10: 51.

Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 24.

Distribution: Sula Islands (Indonesia); Philippines,

21A. Genus: MINDANATIA Malloch, 1931 (Monotypic)

Malloch, 1931 Proc, U.S. nat. Mus. 78 (15): 25.

Type species: Mindanaia latifasciata Malloch, 1931.

|. M, latifasciata Malloch, 1931

Mindanaia latifasciata Malloch, 1931 ibid 78 (15): 25.

Distribution: Philippines.

LOWER—AUSTRALIAN REGION OTITIDAE 19

22, Genus: ICTERACANTHA Hendel, 1912

Hendel, 1912 Supp. Ent. Berlin 1: 14.

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 112.

Type species: Trypeta chalybeiventris Wiedemann, 1830.

1. I. chalybeiventris (Wied., 1830)

Trypeta chalybeiventris Wiedemann, 1830 Auss. zweifl. Ins. 2: 479.

Dacus bicolor Walk., 1849 List Dipt. Brit. Mus. 4: 1071.

Dacus bicolor Walk., 1849 Bezzi, 1913 Mem. Indian Mus, 3: 68 (No. 52).

Trypeta chalybeiventris Wied. 1830 Bezzi, 1913 ibid 3: 69 (No. 52).

Icteracantha chalybeiventris (Wied., 1830) Hendel, 1914b :87; 1914a :55.

Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 165.

Distribution: Type locality unknown.

2. I. eyaneiventris (v. d. Wulp, 1881)

Herina cyaneiventris v. d. Wulp, i881 Dipt. Sumatra Exp. Leiden: 51 PI. 3,

Fig, 6.

Scelacanthina cyaneiventris (y. d, Wulp, 1881) Enderlein, 1912 Zool. Jahrb.

33: 349.

Icteracantha cyaneiventris (vy. d. Wulp, 1881) Hendel, 1914b :88; 1914a

755.

Distribution: Sumatra,

23. Genus: DUOMYIA Walker, 1849

Walker, 1849 List Dipt. Brit. Mus. 4: 800. Hendel, 1914a, Gen. Ins.

157: 56, Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 350; 54: 506.

Type species: Duomyia obscura Walker, 1849.

*1, D. annulipes Hendel, 1914

Duomyia annulipes Hendel, 1914b :98; 1914a :57.

Distribution: Western Australia.

*2. D. azurea Hendel, 1914

Duomyia azurea Hendel, 1914b :97; 1914a :57.

Distribution: Western Australia.

20 REC. S$. AUST. MUS., 16 (2): 1-93 March, 1970

*3. D. decora (Macq., 1846)

Senopterina decora Macquart, 1846 Dipt. exot. Supp. 1: 208 Pl. 18, Fig. 10.

Ortalis punctifrons Macquart, 1847 ibid Supp. 3: 61 Pl. 7, Fig. 4.

Duomyia decora (Macq., 1846) Hendel, 1914b :95; 1914a :57. Enderlein,

1924 Mitt. zool. Mus. Berlin 11: 112. Malloch, 1929 Proc, Linn. Soc.

N.S.W. 54: 510.

Distribution: New South Wales (Australia).

4. D. fidschiensis Enderlein, 1924

Duomyia fidschiensis Enderlein, 1924 Mitt. zool. Mus. Berlin 11; 112.

Bezzi, 1928 Dipt. Fiji: 91.

Distribution: Fiji.

5. D. grandis (Schiner), 1868

Senopterina grandis Schiner, 1868 Novara Dipt.: 289.

Duomyia grandis (Schiner), 1868 Hendel, 1914b :95; 1914a :57.

Distribution: Chile (sic.).

*6, D. irregularis Malloch, 1929

Duomyia irregularis Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 509 Fig. 16.

Distribution: Northern Australia.

*7, D. laeta (Walk., 1849)

Lamprogaster laeta Walker, 1849 List Dipt. Brit. Mus, 4: 805.

Duomyia laeta (Walk., 1849) Hendel, 1914b :96; 1914a :58.

Distribution: Australia (sic.).

See Lamprogaster laeta (Guerin, 1830).

*8. D. maculipennis Hendel, 1914

Duomyia maculipennis Hendel, 1914b :93; 1914a :58. Malloch, 1929

Proc. Linn. Soc. N.S.W. 54: 511 Fig. 2C.

Distribution: Queensland; New South Wales (Australia).

*9. D. mithrax Hendel, 1914

Duomyia mithrax Hendel, 1914b :92; 1914a :58.

Distribution: Queensland (Australia).

*10, D. nigricosta Malloch, 1929

Duomyia nigricosta Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 511.

Distribution: Queensland (Australia),

LOWER—AUSTRALIAN REGION OTITIDAE 21

*11. D. nigripes (Macq., 1850)

Senopterina nigripes Macquart, 1850 Dipt. exot. Supp. 4: 283 Pl. 26, Fig. 6.

Urophora nigripes (Macquart, 1851) Mem. Soc. Sci. Lille: 260 Pl, 26,

Fig. 13.

Urophora nigripes (Macquart, 1851) Bezzi, 1913 Mem. Indian Mus. 3: 76

(No. 184).

Duomyia nigripes (Macq., 1851) Hendel, 1914a :102.

Distribution: Tasmania (?).

*12. D. obscura Walker, 1849

Duomyia obscura Walker, 1849 List Dipt. Brit. Mus. 4: 800.

Senopterina gigas Macquart, 1850 Dipt. exot. Supp. 4: 282 Pl. 26, Fig. 4.

Duomyia obscura Walk., 1849 Hendel, 1914b :90; 1914a :58. Malloch,

1928 Proc. Linn. Soc. N.S.W. 53: 351; 1929 ibid 54: 510.

Distribution: Queensland; New South Wales (Australia).

*13. D. punctifrons Malloch, 1929

Duomyia punctifrons Malloch, 1929 Proc. Linn. Soc. N.S.W. 54; 510.

Distribution: New South Wales (Australia).

*14. D. scutellaris (Macq., 1850)

Senopterina scutellaris Macquart, 1850 Dipt. exot. supp, 4: 282 Pl, 26,

Fig. 5.

Duomyia scutellaris (Macq., 1850) Hendel, 1914b ;102.

Distribution: New South Wales (Australia).

*15. D. sericea Hendel, 1914

Duomyia sericea Hendel, 1914b :99; 1914a :58.

Distribution: Queensland (Australia).

*16. D. spinifemorata Malloch, 1929

Duomyia spinifemorata Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 508

Figs. la, 1b.

Distribution: Northern Territory (Australia),

*17. D. thalassina Walker, 1849

Duomyia thalassina Walker, 1849 List Dipt. Brit. Mus. 4: 801. Hendel,

1914b :92; 1914a :58. Enderlein, 1924 Mitt. zool. Mus. Berlin 11:

112. Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 511.

Distribution: Queensland (Australia).

22 REC. 8. AUST. MUS., 16 (2): 1-93 March, 1970

*18. D. tomentosa Hendel, 1914

Duomyia tomentosa Hendel, 1914b :100; 1914a :58. Malloch, 1929 Proc.

Linn. Soc. N.S.W. 54: 507 Fig. 2b.

Distribution: Queensland (Australia).

Note: Evidence favours the opinion that this genus is confined to the

Australian mainland. No species have been recorded from New Guinea or

New Zealand, and there is no authentic record of any from Tasmania. The

four species, D. grandis Schiner 1868, D. fidschiensis Enderlein 1924,

D. nigripes (Macq. 1850) and D. scutellaris (Macq. 1850), which have

been recorded from other places, are all suspect in varying degrees.

The status of D. grandis is extremely doubtful. It was described by

Schiner from Novara material labelled “Chile”. It is the only Duomyia ever

to have been recorded from South America, and the species has never been

recorded since. Hendel (A) queried the type locality; Malloch (1928)

suggested that the species did not belong to Duomyia. Malloch’s suggestion

appears to be much the more probable. Were D. grandis an Australian

species wrongly labelled as having been collected in Chile, it is not unreason-

able to expect that it would since have been recorded from Australia, which

it has not. If, on the contrary, it is a member of a South American genus

superficially resembling Duomyia then all difficulties disappear.

D. fidschiensis may be an introduction from Australia into Fiji, and

the name is possibly a synonym; alternatively, the species may not belong

to Duomyia. Some of Enderlein’s work is marred by superficiality and

lack of a critical approach, and decisions arrived at by him have often been

shown to be wrong by later workers. A detailed examination of the type

of D. fidschiensis, and a careful comparison with all known species of

Duomyia, will have to be undertaken before finality can be arrived at.

D. nigripes and D. scutellaris were collected by the Verreaux brothers,

between 1842 and 1846, for the Musée d’Histoire Naturelle de Paris. They

formed part of the 140 species of Diptera described by Macquart in the

fourth supplement of his Diptéres Exotiques. The type locality of all the

material was given as “Tasmania”. It is significant that these are the only

species of Duomyia ever to be recorded from Tasmania. Hardy (1929,

Proc. Linn. Soc. N.S.W, 54: 61-64), assembled evidence which strongly

suggests that all 140 species were collected in a coastal area round Sydney,

New South Wales. The types are no longer in existence. Hendel (B)

merely quotes Macquart’s descriptions verbatim with the comment “Die

Arten D. nigripes Macquart und scutellaris Macquart konnten nicht in die

Tabelle aufgenommen werden.” Malloch (1928, 1929) mentions neither

in his revisions of the genus.

There can be little doubt that species yet await description, particularly

in Western Australia.

LOWER—AUSTRALIAN REGION OTITIDAE 23

25. Genus: PSEUDEPICAUSTA Hendel, 1914

Hendel, 1914a Gen. Ins. 157: 62; 1914b :112. Enderlein, 1924 Mitt.

zool. Mus. Berlin 11: 115. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 118.

Type species: Herina chalybea Doleschall, 1858.

*1, P. angulata Hendel, 1914

Pseudepicausta angulata Hendel, 1914b :118. Enderlein, 1924 Mitt. zool.

Mus. Berlin 11: 115. Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 26;

1939 Prac. Linn. Soc. N.S.W. 64: 119.

Distribution: New Guinea; Celebes.

*2. P. apicalis Malloch, 1939

Pseudepicausta apicalis Malloch, 1939 Proc. Linn, Soc. N.S.W, 64: 119

Pl. 4, Fig. 17.

Distribution: New Guinea.

3. P. bataviensis (Schiner, 1868)

Senopterina bataviensis Schiner, 1868 Novara Dipt. 7288.

Pseudepicausta bataviensis (Schiner, 1868) Hendel, 1914b :115; 1914a 264.

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 115. Malloch, 1931 Proc.

U.S. nat. Mus. 78 (15): 26.

Distribution; Java,

*4. P. chalybea (Doleschall, 1858)

Herina chalybea Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 125.

Dacus obtrudens Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 116. Bezzi,

1913 Mem. Indian Mus. 3: 76 (No. 191).

Pseudepicausta chalybea (Dol., 1858) Hendel, 1913 Gen. Platy. Figs, 119,

120; 1914b +113: 1914a :64 Pl. 6, Figs. 119, 120. Bezzi, 1917

Philippine J. Sci, (D) 12: 136. Enderlein, 1924 Mitt. zool. Mus. Berlin

11: 115. Frey, 1930 Notul. ent. Helsingf. 10: 54, Malloch, 1931 Proc.

U.S. nat. Mus, 78 (15): 26; 1939 Proc. Linn. Soc. N.S.W. 64; 118.

Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. & (5): 179.

Distribution: Amboina; Ternate; Deslac and Nusa Islands; Sunda

Islands; New Guinea.

5. P. contrahens (Walker, 1860)

Dacus contrahens Walker, 1860 J. Proc. Linn. Soc. Lond. 4: 151. Bezzi,

1913 Mem. Indian Mus. 3: 69 (No. 62).

Pseudepicausta contrahens (Walk., 1860) Hendel, 1914a :64. Hardy, 1959

Bull. Brit. Mus. (nat, Hist.) Ent. 8 (5): 168.

Distribution: Macassar.

24 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*6. P. detrudens (Walker, 1865)

Dacus detrudens Walker, 1865 J. Proc. Linn. Soc. Lond. 8: 135. Bezzi,

1913 Mem. Indian Mus. 3: 70 (No. 77).

Pseudepicausta detrudens (Walk., 1865) Hendel, 1914a :64; 1914b :113.

Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 169.

Distribution: Indonesia; Salawatty Is. (New Guinea).

Both Hendel and Hardy found differences between the assumed type

and Walker’s description. There is some uncertainty as to whether Walker's

specimen labelled “? type” was the specimen on which he based his

description,

7. P. exigens (Walker, 1860)

Dacus exigens Walker, 1860 J. Proc. Linn, Soc. Lond. 4; 151. Bezzi, 1913

Mem. Indian Mus. 3: 71 (No. 95).

Pseudepicausta exigens (Walk., 1860) Hendel, 1914a :64. Hardy, 1959

Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 171.

Distribution: Macassar.

8. P. experta (Walker, 1862)

Dacus expertus Walker, 1862 J. Proc. Linn. Soc. Lond. 6: 12, Bezzi, 1913

Mem. Indian Mus. 3: 71 (No. 97).

Pseudepicausta experta (Walk., 1862) Hendel, 1914a :64. Hardy, 1959

Bull. Brit. Mus. (nat. Hist.) Ent, 8 (5): 171.

Distribution: Halmahera Is. (Moluccas).

9. P. geniculata (v. d. Wulp, 1898)

Senopterina geniculata y. d. Wulp, 1898 Tijd. Ent. 41: 206. de Meijere,

1911 Tijd. Ent. 54: 369.

Pseudepicausta geniculata (v. d. Wulp, 1898) Hendel, 1914a :64; 1914b

1114.

Distribution: Java.

10. P. limpidipennis (Doleschall, 1858)

Herina limpidipennis Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 126.

Pseudepicausta limpidipennis (Dol., 1858) Hendel, 1914a :64; 1914b :117.

Distribution: Amboina.

*11. P. mutilloides (Walker, 1859)

Dacus mutilloides Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 115. Bezzi,

1913 Mem. Indian Mus. 3: 76 (No, 176).

LOWER—AUSTRALIAN REGION OTITIDAE 25

Pseudepicausta mutilloides (Walk., 1859) Hendel, 1914a :64; 1914b :116.

Frey, 1930 Notul. ent. Helsingf. 10: 54. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 119 (“multilloides”) (sic.). Hardy, 1959 Bull. Brit. Mus.

(nat. Hist.) Ent. 8 (5): 178.

Distribution: New Guinea.

*12. P. pompiloides (Walker, 1859)

Dacus pompiloides Walker, 1859 J. Proc. Linn, Soc. Lond. 3: 116. Bezzi,

1913 Mem. Indian Mus. 3: 77 (No, 210).

Pseudepicausta lagarosia Hendel, 1914a :64; 1914b :118. Malloch, 1939

Proc, Linn. Soc. N.S.W. 64: 118.

Pseudepicausta pompiloides (Walk., 1859) Hardy, 1959 Bull. Brit. Mus.

(nat. Hist.) Ent. 8 (5): 180.

Distribution: New Guinea.

P. pompiloides was the species described by Hendel as new under the

name of P. lagarosia. Hardy showed the synonymy.

13. P. quadrisetosa (de Meijere, 1911)

Senopterina quadrisetosa de Meijere, 1911 Tijd. Ent. 54: 368,

Pseudepicausta quadrisetosa (de Meij., 1911) Hendel, 1914a :64; 1914b

:115.

Distribution: Jaya.

*14. P. solocifemur Enderlein, 1924

Pseudepicausta solocifemur Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 115.

Distribution: New South Wales (Australia).

*15. P. wallacei Hendel, 1914

Pseudepicausta wallacei Hendel, 1914b :117; 1914a :64. Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 119.

Distribution: New Guinea.

26. Genus: SCOTINOSOMA Loew, 1873

Loew, 1873 Mon. N. Amer. Dipt. 3: 45.

Type species: S, bistrigata Hendel, 1914.

In 1873, Loew summarized the characters of his new genus, Scotinosoma

but named no species nor designated a type species. S. bistrigata hence

became the type species by subsequent designation by Hendel in 1914.

Hendel, 1914a :65; 1914b :120. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 117.

26 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*1. §. attenuata (Malloch, 1931)

Pseudepicausta attenuata Malloch, 1931 Proc. U.S. nat. Mus. 78: 27.

Scotinosoma attenuata (Malloch, 1931) Malloch, 1939 Proc. Linn. Soc,

N.S.W. 64: 117.

Distribution; Queensland (Australia).

Malloch first placed this species and No. 3 in Pseudepicausta but later

transferred them to Scotinosoma.

*2. §. bistrigata Hendel, 1914

Scotinosoma bistrigata Hendel, 1914b :120; 1914a :65. Malloch, 1939

Proc. Linn. Soc. N.S.W, 64: 117.

Distribution: Queensland (Australia).

*3. §. completa (Malloch, 1931)

Pseudepicausta completa Malloch, 1931 Proc. U.S. nat. Mus. 78: 27,

Scotinosoma completa (Malloch, 1931) Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 117.

Distribution: Queensland (Australia).

*4. §. erasa Malloch, 1939

Scotinosoma erasa Malloch, 1939 Proc. Linn. Soc. N.S.W. 64; 117.

Distribution: Queensland (Australia).

5. S. typicum Bezzi, 1917

Scotinosoma typicum Bezzi, 1917 Philippine J. Sci. (D) 12: 136. Frey,

1930 Notul. ent. Helsingf. 10: 46.

Distribution: Philippines.

26A. Genus: GUAMOMYIA Malloch, 1942 (Monotypic)

Type species: Guamomyia fascipennis Malloch, 1942.

Malloch, 1942 Insects of Guam B. P. Bishop Mus. Bull. 172: 206,

Fig. 3.

1. G. fascipennis Malloch, 1942 ibid

Distribution: Guam.

27. Genus: RHYTIDORTALIS Hendel, 1914

Hendel, 1914b :121; 1914a :66. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 106.

Type species: Rhytidortalis cribrata Hendel, 1914.

LOWER—AUSTRALIAN REGION OTITIDAE 27

*1. R. rugifrons (Thomson, 1868)

Senopterina rugifrons Thomson, 1868 Dipt. Eugen, Resa: 577.

Rhytidortalis rugifrons (Thomson, 1868) Hendel, 1914b :122; 1914a :68.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 106,

Distribution: Australia (sic, ).

Two species only have so far been recorded. The type species is from

Taiwan,

27A. Genus: CLEITAMOIDES Malloch, 1939

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 106.

Type species: Cleitamoides kerteszi (Hendel, 1914).

*1. C. kerteszi (Hendel, 1914)

Cleitamia kerteszi Hendel, 1914b :130. Enderlein, 1924 Mitt. zool. Mus.

Berlin 11: 117.

Cleitamoides kerteszi (Hendel, 1914) Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 107.

Distribution: New Guinea.

“2. C. latifascia (Walker, 1859)

Dacus latifascia Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 114. Bezzi,

1913 Mem. Indian Mus. 3: 74 (No. 146).

Cleitamia latifascia (Walk., 1859) Edwards, 1915 Trans. Zool. Soc. Lond,

20: 415.

Cleitamoides latifascia (Walk., 1859) Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 107. Hardy, 1959 Bull. Brit. Mus (nat. Hist.) Ent. 8 (5):

177.

Distribution: New Guinea.

*3. C. liturata (Walker, 1861)

Dacus lituratus Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 251.

Cleitamia liturata (Walk., 1861) Osten-Sacken, 1881 Ann. Mus. Stor, nat.

Genova 16: 468. de Meijere, 1913 Nov, Guin, 9: 375. Hendel, 1914a

:73; 1914b :129.

Cleitamoides liturata (Walk., 1861) Malloch, 1939 Proc. Linn. Soc. N.S.W.

64: 107. Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5): 178.

Distribution: New Guinea.

28 REC. S. AUST, MUS., 16 (2): 1-93 March, 1970

30. Genus: CLEITAMIA Macquart, 1835

Macquart, 1835 Suites d Buffon 2: 440, Hendel, 1914a Gen. Ins. 157:

71. Malloch, 1939 Proc. Linn, Soc. N.S.W. 64: 107.

Type species: Ortalis astrolabei Boisduval, 1833.

*1. C. amabilis Osten-Sacken, 1881

Cleitamia amabilis Osten-Sacken, 1881 Ann. Mus, Stor. nat. Genova 16:

468. Hendel, 1914b :127; 1914a :72, Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 109.

Distribution: New Guinea.

*2. C. astrolabei (Boisduval, 1833)

Ortalis astrolabei Boisduval, 1833 Voy. Astrolabe :688 Pl. 12, Fig. 17.

Cleitamia astrolabei (Boisd., 1833) Macquart, 1833 Suites a Buffon :440

Pl, 19, Fig. 4. de Meijere, 1913 Nov. Guin. 9: 375. Hendel, 1914a

:72: 1914b :125. Malloch, 1939 Prac. Linn. Soc. N.S.W. 64: 109

Pl. 4, Fig. 8,

Distribution: New Guinea.

*3. C. biarcuata (Walker, 1865)

Poticara biarcuata Walker, 1865 J. Proc. Linn. Soc. Lond. 8: 133.

Cleitamia biarcuata (Walk., 1865) Hendel, 1914b :133; 1914a :73.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 110.

Distribution: New Guinea.

*4, C. catharinae de Meijere, 1913

Cleitamia catharinae de Meijere, 1913 Bijdr. Dierk. 19: 63; 1915 Tijd. Ent.

58: 129. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 107 (footnote).

Distribution: New Guinea.

In his 1915 paper, de Meijere synonymized his species with Poticara

(—Cleitamia) tricurvata Walker, 1864. Malloch (1939) pointed out that

there were considerable differences in the descriptions of the two species.

In view of these, he accepted both catharinae and tricurvata as distinct until

both types had been compared. So far as I know this has not yet been done,

but the two descriptions differ sufficiently for catharinae to be regarded as

a valid species until the contrary is shown.

*5. C. cheesmanae Malloch, 1939

Cleitamia cheesmanae Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 110 PI.

4, Fig. 10.

Distribution: West Irian (formerly Dutch New Guinea).

LOWER—AUSTRALIAN REGION OTITIDAE 29

*6, C. cyclops Malloch, 1939

Cleitamia cyclops Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 110 Pl. 4,

Fig. 9.

Distribution: West Irian (formerly Dutch New Guinea).

*7, C, delandi Malloch, 1939

Cleitamia delandi Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 111 PL. 4,

Fig. 12.

Distribution: New Guinea.

*8. C. excepta Malloch, 1939

Cleitamia excepta Malloch, 1939 Proc, Linn. Soc. N.S.W. 64: 111 Pl. 4,

Fig. 11.

Distribution: West Irian (formerly Dutch New Guinea).

*9. C. gestroi Kertesz, 1899

Cleitamia gestroi Kertesz, 1899 Ann. Mus. Stor. nat. Genova 19: 566 Pl. 7,

Fig. 8. Hendel, 1914a :73; 1914b :133. Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 109.

Distribution: New Guinea.

“10. C, insignis de Meijere, 1915

Cleitamia insignis de Meijere, 1915 Tijd. Ent. 58: 128 Pl. 1, Fig. 14,

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 110.

Distribution: New Guinea.

*I1. C. orthocephala Hendel, 1914

Cleitamia orthocephala Hendel, 1914b :126; 1914a :73. Malloch, 1939

Proc. Linn. Soc. N.S.W, 64: 109.

Distribution: New Guinea.

*12. C. ostensackeni Kertész, 1898

Cleitamia ostensackeni Kertész, 1898 Term. Fiiz. 21: 494; 1899 Ann. Mus.

Stor, nat. Genova 19: 563 Pl. 7, Fig. 5. Hendel, 1914b :131; 1914a :73.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 109.

Distribution: New Guinea.

*13. C. rivellioides Osten-Sacken, 1881

Cleitamia rivellioides Osten-Sacken, 1881 Ann. Mus. Stor. nat. Genova 16:

469. Kertész, 1899 Ibid 19: 564 Pl. 7, Fig. 6. Hendel, 1914b :130;

1914a :73. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 110.

Distribution: New Guinea,

30 REC. S, AUST. MUS., 16 (2): 1-93 March, 1970

*14. C. roderi Kertész, 1899

Cleitamia roderi Kertész, 1899 Ann. Mus. Stor. nat. Genova 19: 565 Pl. 7,

Fig. 7. Hendel, 1914a :73; 1914b :132. Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 110.

Distribution: New Guinea.

*15. C. similis Kertész, 1899

Cleitamia similis Kertész, 1899 Ann. Mus. Stor. nat. Genova 19: 652 Pls

Fig. 4. Hendel, 1914a :73; 1914b :128, Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 109.

Distribution: New Guinea.

*16. C. tricurvata (Walker, 1864)

(See C. catharinae de Meijere, 1913).

Poticara tricurvata Walker, 1864 J. Proc, Linn. Soc, Lond. 7: 227.

Cleitamia tricurvata (Walk., 1864) Hendel, 1914a :73; 1914b :33.

de Meijere, 1915 Tijd. Ent. 58: 129, Enderlein, 1924 Mitt. zool. Mus.

Berlin 11: 116. Malloch, 1939 Prac. Linn. Soc. N.S.W. 64: 109.

Distribution: New Guinea.

*17. C, trigonalis de Meijere, 1913

Cleitamia trigonalis de Meijere, 1913 Nov. Guin. 9: 375; 1915 Tijd. Ent.

58: 129. Hendel, 1914a :73; 1914b :130. Malloch, 1939 Proc. Linn,

Soc. N.S.W. 64: 112.

Distribution: New Guinea.

30A. Genus: KENOGNATHUS Malloch, 1930

Malloch, 1930 Ins. Samoa 6 (5): 225.

Type species: Xenognathus bryani Malloch, 1930.

1. X. bryani Malloch, 1930

Xenognanthus bryani Malloch, 1930 Ins. Samoa 6 (5): 226 Figs. 4, 5.

Distribution: Samoa.

2. X. inermis Malloch, 1930

Xenognanthus inermis Malloch, 1930 Ins. Samoa 6 (5): 228 Fig. 6.

Distribution: Samoa.

31. Genus: LAGLAISIA Bigot, 1880

Bigot, 1880 Ann. Soc. ent. France (Sth Ser.) 10: 92.

Type species; L. caloptera Bigot, 1880.

LOWER—AUSTRALIAN REGION OTITIDAE 31

*|, L. biroi Hendel, 1914

Laglaisia biroi Hendel, 1914b :136; 1914a :75. Malloch, 1939 Proc, Linn.

Soc. N.S.W. 64: 112.

Distribution: New Guinea.

*2. L. caloptera Bigot, 1880

Laglaisia caloptera Bigot, 1880 Ann. Soc. ent. France (Sth Ser.), 10: 92.

Hendel, 1914a :75; 1914b :134. Malloch, 1939 Proc. Linn. Soc. N.S.W,

64: 112.

Distribution: New Guinea.

*3. L. fascipennis de Meijere, 1915

Laglaisia fascipennis de Meijere, 1915 Tijd. Ent, 58: 134 Pl. 1, Fig. 15.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 112.

Distribution: New Guinea.

*4. L. kochi de Meijere, 1907

Laglaisia kochi de Meijere, 1907 Tijd. Ent. 50; 120 Pl. 4, Fig. 1; 1913 Nov.

Guin. 9: 374, Hendel, 1914a :75; 1914b :135. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 112.

Distribution: New Guinea.

*5, L. stylops Enderlein, 1924

Laglaisia stylops Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 116. Malloch,

1939 Proc. Linn. Soc. N.S.W. 64: 113.

Distribution: New Guinea.

*6, L. telescopica Enderlein, 1924

Laglaisia telescopica Enderlein, 1924 Mitt. Zool. Mus. Berlin 11: 116.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 113.

Distribution: New Guinea.

32. Genus: LORIOMYTA Kertész, 1899 (Monotypic)

Kertész, 1899 Ann. Mus. Stor. nat. Genova 39: 567.

Type species: L. guttipennis Kertész, 1899.

*|. L. guttipennis Kertész, 1899

Loriomyia guttipennis Kertész, 1899 Ann. Mus. Stor. nat. Genova 39: 567.

Hendel, 1914a :75; 1914b :137. Malloch, 1939 Proc. Linn. Soc, N.S.W.

64: 113.

Distribution: New Guinea.

32 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

35. Genus: LOXONEUROIDES Hendel, 1914 (Monotypic)

Hendel, 1914b :141.

Type species: L. varipennis Hendel, 1914.

*1. L. varipennis Hendel, 1914

Loxoneuroides varipennis Hendel, 1914b :141; 1914a :80.

Distribution: Australia (sic.).

38. Genus: POGONORTALIS de Meijere, 1911

de Meijere, 1911 Tijd. Ent. 54: 370.

Type species: Trypeta doclea Walker, 1849.

*1. P. commoni Paramonov, 1957

Pogonortalis commoni Paramonov, 1957 Ann. Mag. nat. Hist. (Ser. 12)

10: 780.

Distribution: Western Australia.

This species was described from a single specimen (3) caught at

Albany, Western Australia. No other specimens have been recorded.

*2. P. doclea (Walker, 1849)

Trypeta doclea Walker, 1849 List Dipt. Brit, Mus. 4: 1035.

(Syn.) P. barbata, P. barbifera Hendel, 1914a :84; 1914b :144.

Pogonortalis barbifera Hendel, 1914 Bezzi, 1913 Mem. Indian Mus. 3: 71

(No. 87). Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 612.

Pogonortalis doclea (Walk, 1849) Malloch, 1930 ibid 55: 429; 1939 ibid

55: 429: 1939 ibid 64: 120. Paramonov, 1957 Ann. Mag. nat. Hist.

(Ser. 12) 10: 779. Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent.

8 (5): 213.

Distribution: All Australian States except Tasmania.

3. P. fulvofemoralis Malloch, 1942

Pogonortalis fulvofemoralis Malloch, 1942 Bernice P. Bishop Mus. Bull.

172: 205.

Distribution: Guam.

*4, P. howei Paramonoy, 1957

Pogonortalis howei Paramonoy, 1957 Ann. Mag. nat. Hist, (Ser. 12) 10:

780.

Distribution: Lord Howe Island (Australia).

LOWER—AUSTRALIAN REGION OTITIDAE 33

*5. P. similis Hendel, 1914

Pogonortalis similis Hendel, 1914b :143; 1914a :85. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 120.

Distribution; New Guinea.

*6. P. uncinata de Meijere, 1911

Pogonortalis uncinata de Meijere, 1911 Tijd. Ent. 54: 370. Hendel, 1914a

-85: 1914b :145. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 119

(included in key but no further details).

Plagiostenopterina aberrans (Frey, 1930) Hennig, 1941 Arb. morph, taxon.

Ent. Berl. 7: 315.

Distribution: Java.

39. Genus: RIVELLIA Robineau-Desvoidy, 1830

Robineau-Desvoidy, 1830 Essai Myodaires :729.

Type species: Musca syngenesiae Fabricius, 1781 (The type species is

of European origin).

Curran, 1934 N. Amer. Dipt. :281. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 120,

1. R. abana Curran, 1929

Rivellia abana Curran, 1929 Amer. Mus. Nov. No. 339: 11.

Distribution: New Caledonia.

2. R. aequifera (Walker, 1862)

Ortalis aequiferus Walker, 1862 J. Proc. Linn. Soc. Lond. 6: 15.

Rivellia aequifera (Walk., 1862) Hendel, 1914a :89; 1914b :182.

Hendel regarded this species as doubtful.

Distribution: Moluccas.

*3. R. affinis Hendel, 1914

Rivellia affinis Hendel, 1914b :161; 1914a :87. Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 121.

Distribution: New Guinea.

*4. R,. basilaris (Wiedemann, 1830)

Trypeta basilaris Wiedeman, 1830 Auss. zweifl. Ins. 2: 510.

Rivellia basilaris (Wied., 1830) Bezzi, 1913 Mem. Indian Mus. 3: 68

(No. 32); 1913 Philippine J. Sci. (D) 8: 321. Hendel, 1914a :87;

1914b :155. Bezzi, 1928 Dipt. Fiji :91. Frey, 1930 Notul, ent.

34 REC. 8: AUST. MUS., 16 (2): 1-93 March, 1970

Helsingf. 10: 54. Malloch, 1930 Ins. Samoa 6 (5): 221. Curran, 1936

Proc. Calif. Acad. Sci. (4th Ser.) 22 (1): 23.

Distribution; Sumatra; Java; Singapore; Taiwan; Japan, Fiji; Solomon

Islands.

5. R. concisivitta (Walker, 1862)

Ortalis concisivitta Walker, 1862 J. Proc. Linn. Soc. Lond, 6: 16.

Rivellia concisivitta (Walk., 1862) Hendel, 1914a :88; 1914b :182,

Distribution; Moluccas.

Hendel regarded this species as doubtful.

*6, R. connata (Thomson, 1868)

Herina connata Thomson, 1868 Dipt. Eug. Resa :575.

Rivellia connata (Thomson, 1868) Hendel, 1914a :88; 1914b :152. Bezzi,

1928 Dipt. . . . Fiji :91. Malloch, 1928 Proc, Linn. Soc. N.S.W. 53:

351; 1930 ibid 55: 491 Fig. 1; 1930 Ins. Samoa 6 (5): 221; 1939 Proc.

Linn. Soc. N.S.W. 64: 120.

Distribution; Australia; Samoa; Fiji; a common, widely-spread species

in all parts of Australia.

*7, R. connexa Hendel, 1914

Rivellia connexa Hendel, 1914b :163; 1914a :88. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 121.

Distribution: New Guinea.

8. R. decatomoides (Walker, 1862)

Ortalis decatomoides Walker, 1862 J. Proc. Linn. Soc. Lond. 6: 16.

Rivellia decatomoides (Walk., 1862) Hendel, 1914b :182. Hendel, (1914a,

1914b) regarded this species as doubtful.

Distribution: Moluccas.

*9. R. dimidiata de Meijere, 1908

Rivellia dimidiata de Meijere, 1908 Tijd. Ent. 51: 122 Pl. 4, Fig. 2.

Hendel, 1914a :88; 1914b :164. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 121 Pl. 4, Fig. 19,

Distribution: New Guinea.

*10. R. distobasalis Hardy, 1959

Trypeta basalis Walker, 1859 (nec. Trypeta basalis Walker, 1852) J. Proc.

Linn. Soc. Lond, 3: 120. Bezzi, 1913 Mem. Indian Mus. 3: 67 (No. 30),

Rivellia distobasalis Hardy, 1959 Bull. Brit, Mus. (nat. Hist.) Ent. 8 (5S):

211.

Distribution: Aru Islands (New Guinea).

LOWER—AUSTRALIAN REGION OTITIDAE 35

*11. R. ferruginea Hendel, 1914

Rivellia ferruginea Hendel, 1914b :159; 1914a :88. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 121.

Distribution: New Guinea.

*12. R. fulvescens Malloch, 1940

Rivellia fulvescens Malloch, 1940 Proc. R. ent. Soc, Lond, (B) 9: 72,

Distribution: Solomon Islands.

*13. R. fusca (Thomson, 1868)

Herina fusca Thomson, 1868 Dipt. Eug. Resa :575.

Rivellia fusca (Thomson, 1868) Osten-Sacken, 1882 Berl. ent. Zeit. 26: 211.

Bezzi, 1913 Philippine J. Sci. (D) 8: 321. Hendel, 1914a :89; 1914b

:156. Frey, 1930 Notul. ent. Helsingf. 10: 55. Curran, 1936 Proc.

Calif. Acad. Sci. (4th Series) 22: 23. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 121.

Distribution: Java; Philippines; New Guinea; Solomon Islands;

Taiwan,

14. R. hendeliana Bezzi, 1917

Rivellia hendeliana Bezzi, 1917 Philippine J. Sci. (D) 12: 138. Frey, 1930

Notul. ent. Helsingf. 10: 55.

Distribution: Philippines.

15. R. imitans Malloch, 1930

Rivellia imitans Malloch, 1930 Ins. Samoa 6 (5): 220.

Distribution: Samoa.

*16. R. isolata Malloch, 1930

Rivellia isolata Malloch, 1930 Proc. Linn. Soc. N.S.W. 55; 492 Fig. 2.

Distribution: New South Wales (Australia).

17. R. lavata Hendel, 1914

Rivellia lavata Hendel, 1914b :165; 1914a :89. Malloch, 1930 Ins. Samoa

6 (5): 221.

Distribution: Samoa.

18. R. marina Malloch, 1940

Rivellia marina Malloch, 1940 Proc. R. ent. Soc. Lond. (B) 9: 19,

Distribution: Admiralty Islands.

36 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

19. R. obliqua (Walker, 1861)

Ortalis obliqua Walker, 1861 J. Proc. Linn, Soc. Lond, 5: 297.

Rivellia obliqua (Walk., 1861) Hendel, 1914b :172; 1914a 789.

Distribution: Moluccas.

Hendel was doubtful of this species.

20. R. pipartita Hendel, 1933

Rivellia pipartita Hendel, 1933 Dtsch. ent, Z. (year 1933) :42.

Distribution: Jaya.

*21. R. polita Hendel, 1932

Rivellia polita Hendel, 1932 Mem. Mus. Hist. nat. Belg, 4: 30,

Distribution: New Guinea.

*22. R. radiata Hendel, 1914

Rivellia radiata Hendel, 1914b :161; 1914a :90. Malloch, 1939 Prac.

Linn, Soc. N.S.W. 64: 121.

Distribution: New Guinea.

*23. R. rufibasis Malloch, 1939

Rivellia rufibasis Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 121 Pl. 4,

Fig. 18.

Distribution: New Guinea.

24, R. sauteri Hendel, 1914

Rivellia sauteri Hendel, 1914b :158; 1914a :90. Frey, 1930 Notul. ent.

Helsingf. 10: 55.

Distribution: Taiwan; Philippines.

*25. R. similis Hendel, 1914

Rivellia similis Hendel, 1914b 7157; 1914a :90.

Distribution: New Britain,

26. R. sumbawana Hennig, 1941

Rivellia sumbawana Hennig, 1941 Arb. morph. taxon. Ent. Berl. 8: 23.

Distribution: Sumbawa Island.

27. R. vacillans (Walker, 1860)

Ortalis vacillans Walker, 1860 J. Proc. Linn. Soc. Lond. 4: 157.

Rivellia vacillans (Walk., 1860) Hendel, 1914a :90; 1914b :182.

Distribution: Celebes.

Hendel was doubtful of this species.

LOWER—AUSTRALIAN REGION O'TITIDAE 37

*28, R. virgo Hendel, 1914

Rivellia virgo Hendel, 1914b :167; 1914a :90. Malloch, 1930 Proc. Linn.

Soc. N.S.W. 55: 492 Fig. 3.

Distribution: New South Wales (Australia).

#99. R. viridis Hendel, 1914

Rivellia viridis Hendel, 1914b :152; 1914a :90.

Distribution: New South Wales (Australia).

Hendel suggested that this species was possibly synonymous with Ortalis

mentissa Walker, 1849 List Dipt. Brit. Mus. 4: 991.

39A. Genus: ZEALANDORTALIS Malloch, 1930

Malloch, 1930 Rec. Canterbury (N.Z.) Mus. 3: 243, Harrison, 1959

N.Z. Dept. Sci. ind. Res. Bull. 128: 176.

|. Z. interrupta Malloch, 1930

Zealandortalis interrupta Malloch, 1930 Rec. Canterbury (N.Z.) Mus. 3:

243. Harrison, 1959 N.Z. Dept. Sci. ind. Res. Bull. 128: 176 Fig. 209,

Distribution: New Zealand.

2. Z. philpotti Harrison, 1959

Zealandortalis philpotti Harrison, 1959 N.Z. Dept. Sci. ind. Res. Bull. 128:

178 Fig. 210.

Distribution: New Zealand.

These are the only species of Otitidae recorded from New Zealand.

42. Genus: EUXESTOMOEA de Meijere, 1913

de Meijere, 1913 Nov. Guin. 9: 377, Hendel, 1914a 391; 1914b :187.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 105.

Type species: Ortalis promptus Walker, 1859.

*1. E. bipunctata Hendel, 1914

Euxestomoea bipunctata Hendel, 1914b :188; 1914a :93. Malloch, 1939

Proc. Linn. Soc, N.S.W., 64: 106 Pl. 4, Fig. 6.

Distribution: New Guinea.

*2. K. discifera de Meijere, 1913

Euxestomoea discifera de Meijere, 1913 Nov. Guin. 9: 377. Hendel, 1914a

793; 1914b :189. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 106,

Distribution: New Guinea.

38 REC. S. AUST, MUS., 16 (2): 1-93 March, 1970

*3. E. prompta (Walker, 1859)

Ortalis promptus Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 118.

Euxestomoea prompta (Walk., 1859) de Meijere, 1913 Nov, Guin. 9: 377;

1915 Tijd. Ent. 58: 135. Hendel, 1914a :93; 1914b :187. Malloch,

1939 Proc. Linn. Soc. N.S.W. 64; 106.

Distribution: New Guinea.

Although the section on Euxestomoea in de Meijere, 1913 Neva

Guinea was the work of Hendel, and de Meijere used the wording

“Euxestomoea Hendel, Euxestomoea remained a M.S, name until de Meijere

validated it by his publication of it in Nova Guinea, As Neave (Nomen-

clator Zoologicus 2: 383) points out, the correct citation is therefore

Euxestomoea de Meijere, 1913.

44, Genus: LOXONEVRA Macquart, 1835

Macquart, 1835 Suites Buffon 2: 446 Pl. 19, Fig. 10.

Loxoneura Hendel, 1914a :96; 1914b :190.

Type species: Dictya decora Fabricius, 1805,

Because the letters v and u in Macquart’s Suites a4 Buffan are somewhat

alike in print, authors have mistakenly adopted the spelling Loxoneura.

Inspection of Macquart’s work shows beyond all doubt that he named the

genus Loxonevra. Loxonevra Macquart, 1835 is the only spelling recog-

nized by Neave (Nomenclator Zoologicus 2: 1003).

1. L. decora (Fab., 1805)

Dictya decora Fabricius, 1805 Syst. Antl. :327.

Loxonevra decora (Fab., 1805) Macquart, 1835 Suites Buffon 2: 446

Pl. 19, Fig. 10. Bezzi, 1913 Mem. Indian Mus, 3: 81 (No, 280),

Hendel, 1914a :98: 1914b :191. Bezzi, 1917 Philippine J. Sci. (D)

12: 138. Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 118. Frey, 1930

Notul. ent. Helsingf. 10: 55.

Distribution: Java; Sumatra; Nepal (India); Philippines.

2. L. fascipennis Hendel, 1914

Loxonevra fascipennis Hendel, 1914b :195; 1914a :98.

Distribution: Borneo,

3. L. perilampoides Walker, 1858

Loxonevra perilampoides Walker, 1858 Trans. ent. Soc. Lond, (new ser.)

4: 226. Hendel, 1914a :98; 1914b :198.

Distribution: Sumatra; Sikkim (India).

LOWER—AUSTRALIAN REGION OTITIDAE 39

46. Genus: ACHIOSOMA Hendel, 1914

Hendel, 1914b :200; 1914a :100. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 130.

Type species: Achias dacoides Walker, 1865.

*1. A. aspiciens (Walker, 1864)

Dacus aspiciens Walker, 1864 J. Proc. Linn. Soc. Lond, 7: 229.

Achiosoma aspiciens (Walk., 1864) Hendel, 1914a :101; 1914b :201.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 130.

Distribution; New Guinea.

*2. A. costalis Malloch, 1939

Achiosoma costalis Malloch, 1939 Proc. Linn, Soc, N.S.W. 64: 130.

Distribution: New Guinea.

*3. A. dacoides (Walker, 1865)

Achias dacoides Walker, 1865 J. Proc. Linn. Soc. Lond. 8: 133.

Achiosama dacoides (Walker, 1865) Hendel, 1914a :101; 1914b :200.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 130.

Distribution: New Guinea.

*4. A. nigrifacies Malloch, 1939

Achiosoma_ nigrifacies Malloch, 1939 Proc. Linn. Soc, N.S.W. 64; 131

Pl. 5, Fig. 26.

Distribution: New Guinea.

47. Genus: ACHIAS Fabricius, 1805

Fabricius, 1805 Syst. Antl. :247. Hendel, 1914a :101; 1914b :201.

Malloch, 1928 Proc, Linn. Sac. N.S.W. 53: 351; 1939 ibid 64: 132.

Type species: Achias oculatus Fabricius, 1805.

*1. A, albertisi Osten-Sacken, 1881

Achias albertisi Osten-Sacken, 1881 Ann. Mus. Stor. nat. Genova 16: 473,

Hendel, 1914a :103; 1914b :210. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 137.

Distribution: New Guinea.

*2. A. amplividens Walker, 1859

Achias amplividens Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 122. Hendel,

1914a :103; 1914b :272. Malloch, 1939 Prac. Linn. Soc. N.S.W.

64: 135.

Distribution: Aru Island (New Guinea).

40 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*3. A. apictipennis Hennig, 1940

Achias apictipennis Hennig, 1940 Arb. morph. taxon. Ent, Berl. 7: 316,

Distribution: Queensland (Australia).

*4 A. australis Malloch, 1939

Achias australis Malloch, 1939 Proc, Linn. Soc. N.S.W. 64: 137 Pl. 5,

Fig. 28.

Distribution: Queensland (Australia).

*5. A. brachyophthalmus Walker, 1865

Achias brachyophthalmus Walker, 1865 J. Proc. Linn, Soc. Lond. 8: 119.

Hendel, 1914a :103; 1914b :218. Malloch, 1939 Proc. Linn, Soc.

N.S.W. 64; 135 PI. 5, Fig. 27.

Distribution: New Guinea.

*6. A. diversifrons de Meijere, 1913

Achias diversifrons de Meijere, 1913 Nov. Guin. 9: 371, Hendel, 1914a

103: 1914b :214, Malloch, 1939 Proc. Linn. Soc, N.S.W. 64: 136.

Distribution; New Guinea.

*7. A. fulviceps de Meijere, 1913

Achias fulviceps de Meijere, 1913 Nov. Guin. 9: 373. Hendel, 1914a :103;

1914b :205. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 134.

Distribution: New Guinea.

*8. A, furcatus Hendel, 1914

Achias furcatus Hendel, 1914b :216; 1914a :103. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 136.

Distribution: New Guinea.

*9. A. gjellerupi de Meijere, 1915

Achias gjellerupi de Meijere, 1915 Tijd. Ent. 58: 130.

Distribution: New Guinea.

*10. A, kurandana Hennig, 1940

Achias kurandana Hennig, 1940 Arb. morph. taxon. Ent. Berl. 7. 315.

Distribution: Queensland (Australia).

*11. A. latividens Walker, 1859

Achias latividens Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 121. Hendel,

1914a :103; 1914b :208. Enderlein, 1924 Mitt. zool. Mus. Berlin 11:

126. Malloch, 1939 Proc. Linn. Soc. Lond. 64: 137.

Distribution: New Guinea.

LOWER—AUSTRALIAN REGION OTITIDAE 41

*12. A. longividens Walker, 1859

Achias longividens Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 121. Hendel,

1914a :103; 1914b :210. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64:

137.

Distribution: Aru Island (New Guinea).

*13, A. macrocephalus Hendel, 1914

Achias macrocephalus Hendel, 1914b :215; 1914a :104. Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 137 (as microcephalus (sic.)).

Distribution: New Guinea.

14. A. oculatus Fabricius, 1805

Achias oculatus Fabricius, 1805 Syst. Antl. :247. Hendel, 1914a :104;

1914b :203.

Distribution: Java (sic.),

This and A. australis Malloch are the only species not recorded from

New Guinea. All the other described species are endemic to New Guinea.

But see *15. A, platychirus Hendel (below).

*15. A. platychirus Hendel, 1914

Achias platychirus Hendel, 1914b :204; 1914a :104. Malloch, 1939 Proc.

Linn. Soc. N.S.W, 64: 134.

Distribution: New Guinea,

Hendel (1914b :202 (footnote)) suggested that his A. platychirus

might be a synonym of A. oculatus Fab., 1805. Should this be demon-

strated, A. oculatus would be a New Guinea as well as a Java species.

*16. A. punctulatus de Meijere, 1913

Achias punctulatus de Meijere, 1913 Nov. Guin. 9: 372. Hendel, 1914a

7104; 1914b :215. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 137.

Distribution: New Guinea.

*17. A. rothschildi Austen, 1910

Achias rothschildi Austen, 1910 Novit. Zool. 17: 459 Pl. 15, Figs. 5-9.

Hendel, 1914a :104; 1914b :206. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 137.

Distribution: New Guinea.

42 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*18. A. strigatus de Meijere, 1913

Achias strigatus de Meijere, 1913 Nov. Guin. 9: 372. Hendel, 1914a :104;

1914b :205. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 134.

Distribution: New Guinea.

*19. A. subnudus Malloch, 1939

Achias subnudus Malloch, 1939 Proc. Linn, Soc. N.S.W. 64: 134.

Distribution: New Guinea,

*20. A. thoracalis Hendel, 1914

Achias thoracalis Hendel, 1914b :213; 1914a :104. Malloch, 1939 Proc.

Linn. Soc. N.S.W, 64: 136.

Distribution: New Guinea.

*91. A. venustulus Walker, 1865

Achias venustulus Walker, 1865 J. Proc. Linn. Soc. Lond. 8: 119, Hendel,

1914a :104; 1914b :219. Malloch, 1939 Proc, Linn. Soc. N.S.W. 64:

134.

Distribution: New Guinea.

48. Genus: LAMPROGASTER Macquart, 1843

Macquart, 1843a Mem. Soc. R. Sci. Lille (1842) :368; 1843b Dipt.

exot. 2: 211. Hendel, 1914a :104; 1914b :220, Malloch, 1928 Proc.

Linn. Soc. N.S.W. 53: 348; 1929 ibid 54: 513; 1930 ibid 55: 432; 1939

ibid 64: 138.

Type species: Lamprogaster flavipennis Macquart, 1843.

1. L. amitina Frey, 1930

Lamprogaster amitina Frey, 1930 Notul. ent. Helsingf. 10: 55.

Distribution: Philippines.

2. L. angusta Enderlein, 1924

Lamprogaster angusta Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 128.

Distribution: Moluccas.

*3. L. apicalis (Walker, 1849)

Chromatomyia apicalis Walker, 1849 List. Dipt. Brit. Mus. 4: 804.

Lamprogaster apicalis (Walk., 1849) Hendel, 1914a :107; 1914b :227.

LOWER—AUSTRALIAN REGION OTITIDAE 43

Helocnemia apicalis (Walk., 1849) Enderlein, 1924 Mitt. zool. Mus. Berlin

11: 128. Lamprogaster apicalis (Walk., 1849) Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 138,

Distribution; Western Australia.

*4. L. austeni Sharp, 1900

Lamprogaster austeni Sharp, 1900 Willey Results 4: 391 Pl. 35, Figs. 11-1 1c.

L. xanthoptera Hendel, 1914a :107; 1914b :220; (footnote) ibid :225.

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 127. Malloch, 1928 Proc.

Linn. Soc. N.S.W. 53: 349; 1929 ibid 54: 515.

L. austeni Sharp, 1900, 1939 ibid 64: 143.

Distribution: Bismarck Archipelago.

*5. L. basalis Walker, 1861

Lamprogaster basalis Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 248.

Hendel, 1914a :107; 1914b :229,. Enderlein, 1924 Mitt. zool. Mus.

Berlin 11: 127. Malloch, 1939 Prac. Linn. Soc. N.S.W. 64: 144.

Distribution: New Guinea.

*6. L. bicolor Macquart, 1847

Lamprogaster bicolor Macquart, 1847 Dipt. exot. Supp. 2: 89. Hendel,

1914a :107; 1914b :237. Enderlein, 1924 Mitt. zool. Mus. Berlin 11:

127. Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 349,

Distribution: New South Wales (Australia).

7. L. celebensis Enderlein, 1924

Lamprogaster celebensis Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 127.

Distribution: Celebes.

*8. L. costalis Walker, 1861

Lamprogaster costalis Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 247.

Hendel, 1914a :108; 1914b :231. Malloch, 1939 Prac. Linn. Soe.

N.S.W. 64: 141.

Distribution: New Guinea.

*9, L. decolor Malloch, 1939

Lamprogaster decolor Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 144

Pl. 5, Fig. 30.

Distribution: New Guinea.

44 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*10, L. elongata v. d. Wulp, 1885

Lamprogaster elongata v. d. Wulp, 1885 Tijd. Ent. 28: 228. Hendel 1914a

:108; 1914b :223. Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 126.

Non Malloch, 1929 Proc. Linn. Soc, N.S.W. 54: 515 (L. pseudelongata

Malloch, 1930) see note. 1930 ibid 55: 432 Fig. 1, 1939 ibid 64: 144.

Distribution: New Guinea and surrounding islands.

In the 1929 reference above, Malloch misidentified as L. elongata a

species sent to him from Queensland. He corrected this mistake in his

1930 reference where he described the Queensland species under the name

of L. pseudelongata. It is to this species, therefore, that the 1929 reference

actually applies.

#1]. L. flavipennis Macquart, 1843

Lamprogaster flavipennis Macquart, 1843 Dipt. exot. 2: 211 PI. 28, Fig. 7.

Hendel, 1914a :108; 1914b :234. Enderlein, 1924 Mirt. zool, Mus.

Berlin 11; 127. Bezzi, 1928 Dipt. . . . Fiji :91. Malloch, 1928 Prac.

Linn. Soc. N.S.W. 53: 349.

Distribution; Victoria and New South Wales (Australia); Fiji.

Bezzi’s identification of L. flavipennis from Fiji requires confirmation.

12. L. fulvescens Malloch, 1931

Lamprogaster fulvescens Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 11.

Distribution: Philippines.

*13, L. fulyipes Malloch, 1939

Lamprogaster fulvipes Malloch, 1939 Proc. Linn. Soc, N.S.W. 64; 145

Pl. 5, Fig, 31.

Distribution: New Guinea,

*14. L. fuscibasis Malloch, 1930

Lamprogaster fuscibasis Malloch, 1930 Proc. Linn, Soc. N.S.W. 55: 433.

Distribution: Queensland (Australia).

*15. L. grossa Malloch, 1939

Lamprogaster grossa Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 142.

Distribution: New Guinea.

*16. L. hilaris (Walker, 1849)

Chromatomyia hilaris Walker, 1849 List Dipt. Brit. Mus. 4: 804.

Lamprogaster hilaris (Walk., 1849) Hendel, 1914a :108; 1914b :244.

Distribution: ? Australia (sic.).

LOWER—AUSTRALIAN REGION OTITIDAE 45

This is a very doubtful species. It is represented by the remains of

Walker's type, and has apparently not been collected since. Hendel (1914b)

quotes Walker's description verbatim without further comment; Malloch

does not refer to the species at all. Even the type locality ts uncertain.

*17, L. indistincta Malloch, 1928

Lamprogaster indistincta Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 349.

Distribution: Northern Australia.

*18. L. instabilis (Walker, 1861)

Dacus instabilis Walker, 1861 J. Proc. Linn, Soc. Lond, 5: 250. Bezzi,

1913 Mem. Indian Mus. 3: 74 (No. 140).

L. gracilis Hendel, 1914b :225. Malloch, 1939 Proc. Linn. Soc. N.S.W.

64: 141. L. instabilis (Walk., 1861) Hardy, 1959 Bull. Brit. Mus, (nat,

Hist.) Ent. 8 (5): 177.

Distribution: New Guinea.

*19. L. jucunda (Walker, 1849)

Chromatomyia jucunda Walker, 1849 List Dipt. Brit. Mus. 4: 802.

Lamprogaster jucunda (Walk., 1849) Hendel, 1914a :108; 1914b :238,

Enderlein, 1924 Mitt. zool. Mus, Berlin 11: 127,

Distribution; New South Wales (Australia).

Although not referred to by Malloch, Enderlein had specimens sent to

him from Sydney, N.S.W.

*90, L. laeta (Guérin, 1830)

Platystoma laeta Guérin, 1830 Vey. Coquille :299.

Lamprogaster laeta (Guérin, 1830) Hendel, 1914a :108; 1914b :244.

Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 516.

Distribution: New South Wales (Australia); Queensland (Australia).

This species is not Chromatomyia laeta Walker, 1849 (List Dipt. Brit.

Mus. 4: 805), a confusion for which Hendel was responsible. In his 1914b

publication, although he had seen neither the type nor specimens con-specific

with it, he transferred Walker's species to Lamprogaster but stated that

he believed it to be Duomyia sp.! In the same publication, he also

transferred Platystoma laeta Guerin, 1830 to Lamprogaster. By so doing,

he produced L. laeta (Guerin, 1830) and L. laeta (Walker, 1849),

In 1929, Malloch (Proc. Linn. Soc. N.S.W. 54: 516) established the

validity of L. laeta (Guérin, 1830). He received specimens of it from

46 REC. 8. AUST. MUS., 16 (2): 1-93 March, 1970

Queensland (Australia), In his paper, he provided a modern description

and commented that this was “the first known record since the original

description”.

Walker's species has no definite status until critical examination of the

type enables it to be correctly placed generically.

*21. L. lepida Walker, 1857

Lamprogaster lepida Walker, 1857 Trans. ent. Soc. Lond. (new series) 4:

226. Hendel, 1914a :108; 1914b :235. Malloch, 1928 Proc. Linn.

Soc. N.S.W. 53: 349; 1929 ibid 54: 516.

Distribution: Queensland (Australia); Celebes,

*22. L. macrocephala Hendel, 1914

Lamprogaster macrocephala Hendel, 1914b :230; 1914a :108. Malloch,

1939 Prac. Linn. Soc. N.S.W. 64: 145,

Distribution: New Guinea.

*23. L. maculipennis Macquart, 1847

Lamprogaster maculipennis Macquart, 1847 Dipt. exot. Supp. 2: 89 Pl. 6,

Fig. 5. Hendel, 1914a :108; 1914b :231. Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 143.

Distribution: Eastern Australia.

24. L. obliqua Frey, 1930

Lamprogaster obliqua Frey, 1930 Notul. ent. Helsingf. 10: 56.

Distribution: Philippines.

*25, L. patula Walker, 1861

Lamprogaster patula Walker, 1861 J. Proc. Linn. Soc. Lond, 5: 247.

Hendel, 1914a :107; 1914b :222. Enderlein, 1924 Mitt. zool. Mus.

Berlin 11: 128 (as Ceratopelta patula (Walk., 1861). Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 140.

Distribution: New Guinea.

26. L. placida (Walker, 1849)

Chromatomyia placida Walker, 1849 List Dipt. Brit. Mus. 4: 802.

Lamprogaster placida (Walker, 1849) Hendel, 1914b :243; 1914a :109.

Bezzi, 1917 Philippine J. Sci. (D) 12: 139. Frey, 1930 Notul. ent.

Helsingf. 10: 46. Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 11,

Distribution: Philippines.

LOWER—AUSTRALIAN REGION OTITIDAE 47

*27. L. poecila Hendel, 1914

Lamprogaster poecila Hendel, 1914b :239; 1914a :109,

Distribution: Northern Australia.

*28. L. pseudelongata Malloch, 1930

Lamprogaster pseudelongata Malloch, 1930 Proc. Linn. Soc. N.S.W. 55:

432 Fig. 2; 1929 ibid 54: 515.

Distribution: Queensland (Australia).

Malloch, 1929 referred this species to L. elongata y. d. Wulp, 1885. He

corrected the mistake in his 1930 reference where he described L. pseude-

longata as sp. nov. His 1929 reference is hence to L. pseudelongata.

29. L. pumicata v, d. Wulp, 1885

Lamprogaster pumicata vy. d. Wulp, 1885 Tijd. Ent. 28: 230 Pl. 7, Fig. 12.

Hendel, 1914a :109; 1914b :238. Malloch, 1929 Proc. Linn. Soc.

N.S.W, 54: 516; 1939 ibid 64: 143.

Distribution: New Caledonia.

*30. L. quadrilinea Walker, 1859

Lamprogaster quadrilinea Walker, 1859, Walker, 1859 J. Proc. Linn. Soc.

Lond. 3: 111. de Meijere, 1913 Nov. Guin. 9: 370. 1915 Tijd. Ent.

58: 132. Hendel, 1914a :109; 1914b :242, Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 142.

Distribution: New Guinea.

“31. L. rufipes Hendel, 1914

Lamprogaster rufipes Hendel, 1914b :233; 1914a :109. Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 143.

Distribution: New Guinea.

*32, L. severa Hendel, 1914

Lamprogaster severa Hendel, 1914b :240: 1914a :109, Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 143.

Distribution: New Guinea.

*33. L. stenoparia Hendel, 1914

Lamprogaster stenoparia Hendel, 1914b :227; 1914a :109. Malloch, 1928

Proc. Linn. Soc. N.S.W. 53; 350; 1929 ibid 54: 515; 1939 ibid 64: 144,

Distribution: Queensland (Australia).

48 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

34, L. superna Walker, 1862

Lamprogaster superna Walker, 1862 J. Proc. Linn. Soc. Lond. 6; 12.

Hendel, 1914a :109; 1914b :240.

Distribution: Moluccas.

35. L. taeniata v. d. Wulp, 1885

Lamprogaster taeniata vy. d. Wulp, 1885 Tijd. Ent. 28: 229 Pl. 7, Figs. 10

and 11. Hendel, 1914a :109; 1914b :241. Malloch, 1939 Proc, Linn.

Soc. N.S.W. 64: 145.

Distribution: Moluccas.

*36. L. trisignata v. d. Wulp, 1885

Lamprogaster trisignata v. d. Wulp, 1885 Tijd. Ent. 28: 231 Pl. 7, Fig. 13.

de Meijere, 1913 Nov. Guin. 9: 370, Hendel, 1914a 109; 1914b :232.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 143.

Distribution: New Guinea.

*37,. L. unimacula Hendel, 1914

Lamprogaster unimacula Hendel, 1914b :239,

Distribution: Queensland (Australia).

*38. L. vella (Walker, 1849)

Chromatomyia vella Walker, 1849 List Dipt. Brit. Mus. 4: 803.

Lamprogaster vella (Walker, 1849) Hendel, 1914a :109; 1914b :233,

Distribution: Queensland (Australia).

*39, L. viola Malloch, 1929

Lamprogaster viola Malloch, 1929. Malloch, 1929 Proc. Linn. Soc. N.S.W.

54: 515.

Distribution: Queensland (Australia).

*40. L. yiolacea (Macquart, 1843)

Cryphocera violacea Macquart, 1843 Dipt. exot. 2: 212 Pl. 29, Fig. 4.

Lamprogaster violacea (Macq., 1843) Hendel, 1914a :109; 1914b :244.

Distribution: New Guinea.

*41. L. zelotypa Hendel, 1914

Lamprogaster zelotypa Hendel, 1914b :226; 1914a :110. Malloch, 1928

Proc, Linn, Soc. N.S.W. 53: 350; 1929 ibid 54: 515; 1939 ibid 64: 141.

Distribution: Quensland (Australia); New Guinea.

Lamprogaster is essentially a New Guinea-Australian genus. Inspection

of otitid collections in Australian museums shows that many species await

description.

LOWER—AUSTRALIAN REGION OTITIDAE 49

49, Genus: MONTROUZIERA Bigot, 1860 (Monotypic)

Bigot, 1860 Ann. Soc. ent. France (3rd series) 8: 224.

Type species: M., lifua Bigot, 1860.

1. M, litua Bigot, 1860

Montrouziera lifua Bigot, 1860 Ann. Sec. ent. France (3rd series) 8: 224.

Hendel, 1914a :110; 1914b :245,

Distribution: New Caledonia.

49A. Genus: RHEGMATOSAGA Frey, 1930 (Monotypic)

Frey, 1930 Notul. ent. Helsingf. 10: 63.

Type species: Rhegmatosaga latiuscula (Walker, 1857),

1. R. latiuscula (Walker, 1857)

Noeeta latiuscula Walker, 1857 J. Proc. Linn. Soc. Lond, 1: 133.

Syn.: Rhegmatosaga insignis Frey, 1930 Notul. ent. Helsingf. 10; 63 Pl. 1,

Fig. 8; 1932 Ann. Mag, nat. Hist. 10: 256,

Rhegmatosaga latiuscula (Walker, 1857) Hardy, 1959 Bull. Brit. Mus. (nat.

Hist.) Ent. 10: 256.

Distribution: Philippines.

Frey designated his insignis as the type species of his gen. nov.,

Rhegmatosaga. Since insignis is a synonym of Noeeta latiuscula Walker,

latiuscula is the type species of Rhegmatosaga.

49B. Genus: TYLOPTERNA Bezzi, 1917

Bezzi, 1917 Philippine J, Sci. (D) 12: 131.

Type species: Tylopterna monstruosum Bezzi, 1917,

|. T. monstruosum Bezzi, 1917

Tylopterna monstruosum Bezzi, 1917 Philippine J. Sci. (D) 12: 133. Frey,

1930 Notul. ent. Helsingf. 10: 64.

Distribution: Philippines.

51. Genus: LENOPHILA Guérin-Ménéville, 1843

Guérin-Ménéville, 1843 Rev. Zool. 6: 200.

Celetor Loew, 1873 Mon. N. Amer. Dipt. 3: 41. Hendel, 1914a :1 13;

1914b :246, Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 505,

Lenophila Guerin, 1843 Bezzi, 1913 Mem. Indian Mus. 3270:

Type species: Ortalis dentipes Macquart, 1843.

Sa REC. 8S. AUST. MUS,, 16 (2): 1-93 March, 1970

*1, L. caerulea (Macquart, 1846) (conj. nov.)

Tephritis caerulea Macquart, 1846 Dipt. exot. Supp. 1: 212 Pl. 18, Fig. 15,

Trypeta cluana Walker, 1849 List Dipt. Brit. Mus. 4: 1019.

Lamprogaster caerulea (Macq., 1846) Schiner, 1868 Dipt. Novara Reis,

285,

Celetor caerulea (Macqg., 1846) Loew, 1873 Mon. N. Amer. Dipt. 3: 41,

Hendel, 1914a :115; 1914b :247. Malloch, 1929 Proc, Linn. Soe.

N.S,W. 54: 506. Hardy, 1959 Bull. Brit. Mus. (nat. Hist.) Ent. 8 (5):

211.

Distribution: Most Australian States.

This species has the peculiar habit of resting on the long, sharp-edged,

siliceous leaves of “grass-trees” (Xanthorrhoea spp.) with the outspread

wings pressed flat against the leaf surface. This habit, aided by the markings

on the wings, gives the insects the appearance of spiders. When the plant

is approached, the insects make no attempt to fly; they merely close their

wings rapidly, and drop down in the spiny centre of the plant where they

are more than adequately protected. This behaviour has been noted by

several entomologists, but the association between insect and plant is

unknown.

*2. L. dentipes (Macquart, 1843)

Ortalis dentipes Macquart, 1843 Dipt. exot. 2: 210 Pl. 28. Fig. 5.

Ceratitis? dentipes (Macq., 1843) Guérin, 1843 Rev. zool. 6: 200,

Lenophila dentipes (Macq., 1843) Guérin, 1843 ibid 6: 200.

Tephritis strigipennis Macquart, 1850 Dipr. exot. Supp. 4: 290 Pl. 27, Fig. 2.

a dentipes (Macq., 1843) Schiner, 1868 Dipt., Novara Reis.

Celetor dentipes (Macq., 1843) Loew, 1873 Mon. N. Amer. Dipt. 3: 41.

Lenophila dentipes (Macq., 1843) Bezzi, 1913 Mem. Indian Mus. 3: 70

(No. 75).

Celetor dentipes (Macq., 1843) Hendel, 1914a :115: 1914b :246.

Enderlein, 1924 Mitr, zool. Mus. Berlin 11: 120. Malloch, 1929 Proc.

Linn. Soc. N.S.W. 54: 506.

Distribution: New South Wales (Australia).

This genus has long been wrongly known as Celetor. When Loew

erected Celetor, he designated Celetor caerulea (Macq., 1846) as the type

species, being unaware that Guérin (1843) had already proposed Lenophila,

and by giving a full re-description of Lenophila dentipes (Macq., 1843) had

made it the type species of the genus. Bezzi (1913) was aware of Guérin's

LOWER—AUSTRALIAN REGION OTITIDAE 51

work but in his paper in Mem. Indian Mus. he merely cited the correct name

without comment, so that the incorrect Celetor has hitherto been used by

almost all earlier workers in the Otitidae. The above facts necessitate the

transfer of both species to Lenophila with Ortalis dentipes Macq., 1843 as

the type species of the genus.

52. Genus: SCHOLASTES Loew, 1873

Loew, 1873 Mon. N. Amer. Dipt. 3: 38. Hendel, 1914a :117; 1914b

:248. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 127.

Type species: Platystoma cinctum Guérin, 1830.

*]. §S. aitapensis Malloch, 1939

Scholastes aitapensis Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 128, Pl. 5,

Fig. 24; 1940 Ann. Mag. nat. Hist. 6: 20; 1942 Bernice P. Bishop Mus.

Bull, 172: 207; 1946 ibid 189: 199.

Distribution: New Guinea; Guam; Solomon Islands; Shortland Island;

Admiralty Islands.

*2_ S. bimaculatus Hendel, 1914

Scholastes bimaculatus Hendel, 1914b :252: 1914a :117. Enderlein, 1924

Mitt. zool. Mus, Berlin 11: 121. Bezzi, 1928 Dipt. . . . Fiji :92.

Malloch, 1930 Ins. Samoa 6 (5): 223; 1939 Proc. Linn. Soc. N.S.W.

64: 129.

S. solomonensis Curran, 1936. Malloch, 1940 Ann. Mag. nat. Hist. 6: 73.

Distribution: New Britain; Fiji; Samoa; Hawaii.

3. S. carolinensis Enderlein, 1924

Scholastes carolinensis Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 12).

Distribution: Caroline Islands.

*4. §. cinetus (Guérin, 1830)

Platystoma cinctum Guérin, 1830 Voy. Coquille Zool. :299 PI. 21, Fig. 9.

Scholastes cinctus (Guérin, 1830) Hendel, 1914b :249; 1914a :117. Bezzi,

1913 Mem. Indian Mus. 3: 71, 72 (Nos. 99 and 102); 1917 Philippine

J. Sci. (D) 12: 139; 1928 Dipr. . . . Fiji :91. de Meijere, 1915 Tijd.

Ent. 58: 132. Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 120.

Malloch, 1930 Ins. Samoa 6 (5): 22; 1939 Proc. Linn. Soc. N.S.W. 64:

128 Pl. 5, Fig. 23.

S. whitneyi Curran, 1936, Malloch, 1940 Ann, Mag. nat. Hist. 6: 73.

Distribution; New South Wales (Australia); New Guinea: New Britain;

Java; Philippines; Fiji.

REC. S. AUST, MUS., 16 (2): 1-93 March, 1970

5. S. frauenfeldi (Schiner, 1868)

Dacus frauenfeldi Schiner, 1868 Dipt. Novara Reis. :285. Bezzi, 1913

Mem. Indian Mus. 3: 72 (No. 114).

Scholastes frauenfeldi (Schiner, 1868) Hendel, 1914a :117; 1914b :251.

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 121.

Distribution: Java; Caroline Islands.

6. S. furcatifascia Enderlein, 1924

Scholastes furcatifascia Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 121.

Distribution: Borneo.

7. §. hirtiventris Malloch, 1942

Scholastes hirtiventris Malloch, 1942 Bernice P. Bishop Mus, Bull. 172: 208.

Swezey, 1946 ibid 189: 199.

Distribution: Guam.

8. §. lonchifera Hendel, 1914

Scholastes lonchifera Hendel, 1914b :253; 1914a :117. Bezzi, 1928 Dipt.

_ . Fiji :92. Malloch, 1932 Bernice P. Bishop Mus. Bull. 98: 205;

1939 Proc. Linn. Soc. N.S.W. 64: 129.

Distribution: Cook Islands; Society Islands; Fiji.

*9, §. palmyra Curran, 1936

Scholastes palmyra Curran, 1936 Proc. Calif. Acad. Sci. (4th Ser.) 22 (1):

24 Pl. 1, Fig. 5.

Distribution: Solomon Islands.

10. S. sexvittatus (Walker, 1861)

Lamprogaster sexvittata Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 261.

Scholastes distigma Hendel, 1914b :252. Frey, 1930 Notul. ent. Helsingf.

10: 56 (as S. distigma, Hendel).

Scholastes sexvittatus (Walker, 1861) Malloch, 1940 Ann. Mag. nat. Hist.

6: 74.

Distribution: Java; Admiralty Islands.

*11, S. taylori Malloch, 1939

Scholastes taylori Malloch, 1939 Proc, Linn. Soc. N.S,W. 64: 129 Pl. 5,

Fig. 25.

Distribution: New Guinea,

LOWER—AUSTRALIAN REGION OTITIDAE 33

12. S. trifasciatus Enderlein, 1924

Scholastes trifasciatus Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 122,

Distribution: Mariana Islands.

58, Genus: PSEUDORICHARDIA Hendel, 1911

Hendel, 1911 :21 (footnote); 1914b :267: 1914a :123.

Although Hendel gave no complete description of Pseudorichardia until

1914b :267 his footnote in Gens. Ins. (113: 21) complies with the mini-

mum requirements of Article 25 (a) of the Rules.

Type species: Richardia flavitarsis Macquart, 1850.

|. P. aristalis Bezzi, 1928

Pseudorichardia aristalis Bezzi, 1928 Dipt. . . . Fiji :93. Fig. 27.

Steyskal, 1952 Occas. Pap. Bernice P. Bishop Mus. 21: 64.

Distribution: Fiji.

2. P. bezziana Steyskal, 1952

Pseudorichardia bezziana Steyskal, 1952 Occas. Pap. Bernice P. Bishop

Mus. 21: 64.

Distribution: New Hebrides.

3. P. flavitarsis (Macquart, 1850)

Richardia flavitarsis Macquart, 1850 Dipt. exot. Supp. 5: 121 Pl. 7, Fig. 3.

Pseudorichardia flavitarsis (Macquart, 1850). Hendel, 1911 Gens. Ins.

113: 21 (footnote); 1914a jbid 157: 124; 1914b :267. Bezzi, 1928

Dipt. . . . Fiji :92 Fig. 26. Malloch, 1929 Ann. Mag. nat. Hist. (ser.

10) 4: 100: 1930 Ins. Samoa 6 (5): 222; 1932 Bernice P. Bishop Mus.

Bull.: 206. Steyskal, 1952 Occas. Pap. Bernice P. Bishop Mus. 21 (5):

66.

Distribution: Malay Archipelago; Samoa; Tahiti; Marquesa Islands;

Society Islands.

4, P. interrupta (Bezzi, 1928)

Pseudorichardia flavitarsis (Macq., 1850) var. interrupta Bezzi, 1928.

Dipt. . . . Fiji, :92 Fig. 26.

Pseudorichardia interrupta (Bezzi, 1928) Steyskal, 1952 Occas. Pap.

Bernice P. Bishop Mus. 21: 66.

Distribution: Fiji.

54 REC. S. AUST. MUS,, 16 (2): 1-93 March, 1970

59. Genus: BREA Walker, 1859

Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 117. Hendel, 1914a :125;

1914b :260. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 123.

Type species: Brea contraria Walker, 1859.

+], B. angustilimbata de Meijere, 1915

Brea angustilimbata de Meijere, 1915 Tijd. Ent. 58: 129.

Distribution: New Guinea.

*2. B. basalis Enderlein, 1924

Brea basalis Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 129, Malloch,

1939 Prac. Linn. Soc. N.S.W. 64: 125.

Distribution: New Guinea.

*3. B. contraria Walker, 1859

Brea contraria Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 117. Hendel,

1914a :126; 1914b :269. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64:

124 Pl. 4, Fig. 21.

Distribution: Aru Islands (New Guinea).

*4, B. discalis Walker, 1859

Brea discalis Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 117. Hendel,

1914a :126; 1914b :272. Malloch, 1939 Proc. Linn. Soc. N.S.W,

64: 124.

Distribution: Aru Islands (New Guinea).

*5, B. discifera Hendel, 1914

Brea discifera Hendel, 1914b :270; 1914a :126. Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 124.

Distribution: New Guinea.

*6, B. flavipes de Meijere, 1913

Brea flavipes de Meijere, 1913 Nov. Guin. 9: 371; 1915 Tijd. Ent. 58: 130.

Hendel, 1914a :126; 1914b :272. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 124.

Distribution: New Guinea.

*7, B. magnifica Hendel, 1914

Brea magnifica Hendel, 1914b :271; 1914a :127. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 125 Pl. 5, Fig. 22.

Distribution: New Guinea.

LOWER—AUSTRALIAN REGION OTITIDAE 35

*8. B. nouhuysi de Meijere, 1913

Brea nouhuysi de Meijere, 1913 Nov. Guin. 9: 370 Pl. 10, Fig. 42; 1915

Tijd. Ent. 58: 130. Hendel, 1914a :127; 1914b :270.

Distribution: New Guinea.

*Q, B. ralumensis Enderlein, 1924

Brea ralumensis Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 129. Malloch,

1939 Proc. Linn. Soc. N.S.W. 64: 125.

Distribution: New Britain.

61. Genus: CHAETORIVELLIA de Meijere, 1913 (Monotypic)

de Meijere, 1913 Nov. Guin. 9: 376. Hendel, 1914a :128; 1914b

:276. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 127.

Type species: Ortalis trifasciata Doleschall, 1859.

*1. C. trifasciata (Doleschall, 1859)

Ortalis trifasciata Doleschall, 1859 Nat. Tijd. Ned. Ind. 17: 121.

Chaetorivellia trifasciata (Doleschall, 1859) de Meijere, 1913 Nov. Guin.

9: 376; 1915 Tijd. Ent. 58: 132. Hendel, 1914a :129; 1914b :276.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 127.

Distribution: New Guinea.

62. Genus: DASYORTALIS Hendel, 1914

Hendel, 1914b :277; 1914a :129. Malloch, 1929 Prac. Linn. Soc.

N.S.W. 54: 506; 1939 ibid 64: 102.

Type species: Ortalis complens, Walker, 1859.

*|. D. angustifrons Hendel, 1914

Dasyortalis angustifrons Hendel, 1914b :279; 1914a :131. Malloch, 1939

Proc, Linn. Soc. N.S.W. 64: 103.

Distribution: New Guinea.

*2. D. barbata Hendel, 1914

Dasyortalis barbata Hendel, 1914b :279; 1914a :131. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 103.

Distribution: New Guinea.

56 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*3, D. complens (Walker, 1859)

Ortalis complens Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 118,

Dasyortalis complens (Walker, 1859) (Nom. nud.) de Meijere, Nov. Guin.

9: 378. Hendel, 1914b :277; 1914a :131. Malloch, 1929 Prec. Linn.

Soc. N.S.W. 54: 506; 1939 ibid 64: 102 Pl. 4, Figs. 1, 2.

Distribution: New Guinea; Solomon Islands; Moluccas.

Curran, 1936 (Proc. Cal. Acad. Sci. 22: 54) and Malloch, 1939

(Proc. Linn. Soc. N.S.W. 64: 103) have described two doubtful varieties of

this species—var. fasciata Curran, and var. separata Malloch. Both require

further study before their status can be determined.

*4. D. goniceps Hendel, 1914

Dasyortalis goniceps Hendel, 1914b :281; 1914a :131. Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 103.

Distribution: New Guinea.

5. D. ? leucomera (Walker, 1863)

Ortalis leacomera Walker, 1863 J. Proc. Linn. Soc. Lond. 7: 219.

Dasyortalis leacomera (Walker, 1863) Hendel, 1914a :131; 1914b :277

(footnote).

Distribution: Moluccas.

Examination of the type will be necessary to determine the genus to

which this species belongs.

6. D. signifacies (Walker, 1861)

Trypeta signifacies Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 165, Bezzi,

1913 Mem. Indian Mus. 3: 79 (No. 240).

Dasyortalis signifacies (Walker, 1861) Hardy, 1959 Bull, Brit. Mus. (nat.

Hist.) Ent. 8 (5): 223.

Distribution: Moluccas.

7. D. ? tarsalis (Walker, 1861)

Ortalis tarsalis Walker, 1861 J. Proc. Linn, Soc. Lond. 5: 296.

Dasyortalis tarsalis (Walker, 1861) Hendel, 1914a :131; 1914b :277

(footnote).

Distribution: Moluccas.

Examination of the type will be necessary to determine the genus to

which this species belongs.

LOWER—AUSTRALIAN REGION OTITIDAE ST

65. Genus: TROPIDOGASTRELLA Hendel, 1914

Hendel, 1914b :285; 1914a :134.

Type species: Tropidogastrella tropida Hendel, 1914.

1. T. cristiventris (Gerstacker, 1860)

Gorgopsis cristiventris Gerstacker, 1860 Stett. ent. Zeit. 21: 135.

Tropidogastrella cristiventris (Gerstacker, 1860) Hendel, 1914b :287; 1914a

:136. Enderlein, 1924 Mitt. cool. Mus. Berlin 11: 130. Bezzi, 1913

Philippine J. Sci. (D) 8: 321. Frey, 1930 Notul, ent. Helsingf. 10: 56.

Distribution: Moluccas,

2. T. decora (de Meijere, 1911)

Zygaenula decora de Meijere, 1911 Tijd. Ent. 54: 371.

Tropidogastrella decora (de Meijere, 1911) Hendel, 1914a :136; 1914b

:289, Frey, 1930 Notul. ent. Helsingf. 10: 56.

Distribution: Jaya,

The type species was recorded from Taiwan.

66. Genus: ASYNTONA Osten-Sacken, 1881

Osten-Sacken, 1881 Ann, Soc. ent. France (series 6) 1: 135. Hendel,

1914a :136: 1914b :290. Malloch, 1939 Prac. Linn. Soc. N.S.W. 64: 121.

Type species: Lamprogaster tetyroides Walker, 1859.

*1. A. flaviceps Hendel, 1914

Asyntona flaviceps Hendel, 1914b :291; 1914a :138. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 122.

Distribution: New Guinea.

*2. A. tetyroides (Walker, 1859)

Lamprogaster tetyroides Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 112.

Asyntona tetyroides (Walker, 1859) Hendel, 1914b :290; 1914a :138.

Enderlein, 1924 Mitt, zool. Mus. Berlin 11: 130. Malloch, 1939 Proc.

Linn, Soc. N.S.W. 64: 122 PIL. 4, Fig. 20.

Distribution: New Guinea,

67. Genus: ZYGAENULA Doleschall, 1858

Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 117. Hendel, 1914a :138;

1914b :292. Malloch, 1939 Proc. Linn, Soc. N.S.W. 64: 123.

Type species: Zygaenula paradoxa Doleschall, 1858.

See note at end of Mesoctenia, No. 67A.

58 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*1. Z. coalescens Hendel, 1914

Zygaenula coalescens Hendel, 1914b :293.

Mesoctenia coalescens (Hendel, 1914) Malloch, 1939 Proc, Linn. Soc.

N.S.W. 64: 123.

Distribution: New Britain.

*2. Z. hylaris Hendel, 1914

Zygaenula hylaris Hendel, 1914b Abh. zool. -bot. Ges. Wien 8: 294.

Mesoctenia hylaris (Hendel), 1914 Malloch, 1939 Proc. Linn. Soc, N.S.W.

64: 123.

Distribution: New Guinea.

3, Z. paradoxa Doleschall, 1858

Zygaenula paradoxa Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 118, Hendel,

1914a :139; 1914b :292, Enderlein, 1924 Mitt. zool. Mus, Berlin 11:

130. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 123.

Distribution: Moluccas.

67A. Genus: ? MESOCTENIA Enderlein, 1924 (Monotypic)

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 130. Mallech, 1939

Proc. Linn, Soc. N.S,W. 64: 123.

Type species: Mesoctenia ralumensis Enderlein, 1924.

*1. M. ralumensis Enderlein, 1924

Mesoctenia ralumensis Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 131.

Malloch, 1939 Proc. Linn. Soc, N.S.W. 64: 123. (A possible synonym

of Zygaenula coalescens.)

Distribution: New Guinea.

Note: Although Zygaenula and Mesoctenia are shown in this catalogue

as being distinct genera, the status of neither can at present be defined, nor

species allotted to either with certainty. All that can be done is to detail

the events that have led to the confusion, leaving clarification of the situation

to the future.

Doleschall (1858 Nat. Tijd, Ned. Ind. 17: 117) erected Zygaenula for

the reception of his species, paradoxa. By monotypy Z. paradoxa is the

type species of the genus, Zygaenula,

Hendel (1914a) revised the genus. He began by synonomizing

Gorgopsis bucephala Gersticker, 1860 (misprinted in A as G. hucephala),

with Z. paradoxa, quoting verbatim, Gersticker’s description of G. bucephala

as that of paradoxa (Hendel 1914a: 292). To Zygaenula, he transferred

LOWER—AUSTRALIAN REGION OTITIDAE 54

Lamprogaster celyphoides Walker, 1859, He also suggested that “Vielleicht

gehért hieher auch Ortalis dispila Thomson, 1868” (ibid: 292 footnote),

though he omitted this species from his key to, and discussion on, the genus.

He described two species Z. coalescens and Z. hilaris. Thus, as he saw it,

Zygaenula comprised the species, paradoxa (type species of the genus),

celyphoides, coalescens and hilaris, with Q, dispila in abeyance. Hendel

must have considered that the species he added to Zygaenula had the generic

characters of paradoxa.

In 1924, Enderlein (Mitt. zool. Mus, Berlin 11; 130) erected the

monotypic genus, Mesoctenia tor his new species, ralumensis (thid :131)

which he designated as the type species of Mesoctenia. He distinguished

his genus from Zygaenula (of which he claimed to have three males and

one female of paradoxa from Amboina) “durch die Anwesenheil einer

Lingsrethe kurzer kriftiger dornartiger Borsten auf der Unterseite (aufsen)

der etwas verdickten Mittelschenkel”.

In 1939, Malloch (Proc. Linn, Soc. N.S.W. 64: 123) discussed both

genera. From Enderlein’s description, he synonomyzed M, ralumensis with

Z. coalescens, But he accepted Mesoctenia as a valid genus with Z.

coalescens (==M. ralumensis) as its type species. To Mesoctenia he trans-

ferred Z, celyphoides and Z. hilaris. He left paradoxa in Zygaenula as “it

is unknown to me”, and he ignored O. dispila,

As revised by him, Zygaenula regained its monotypic status with

Z. paradoxa as its type species, while his enlarged Mesoctenia contained

coalescens (type of the genus), celyphoides, and hilaris, Since he admitted

knowing nothing of Z. paradoxa, it is difficult to understand how Malloch

could distinguish between Zygaenula and Mesactenia, His reliance on

Enderlein appears to be his only reason, His ignoring of O. dispila would

seem to indicate that he regarded it as too doubtful for placement in a genus.

Because of the impossibility of making correct decisions based on the

above facts alone, the author has been forced to adopt a makeshift arrange-

ment: Zygaenula, as revised by Hendel, is, for the time being, accepted;

Mesoctenia is ot doubtful validity,

68. Genus: NAUPODA Osten-Sacken, 1881

Osten-Sucken, 1881 Ann, Soc. ent. France (Ser. 6) 1: 135, Hendel.

19}4a :140; 1914b 2295. Malloch, 1929 Proc. Linn. Soc. N.S.W. 34: 313;

1939 jbid 64: 122.

Type species; Naupoda platessa Osten-Sacken 1882,

*[. N. insularis Paramonov, 1957

Naupoda insularis Paramonoy, 1957 Ann. Mag. nat. Hist. (Ser. 12) 10: 781.

Distribution: Lord Howe Island (Austraha).

60 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

2. N. platessa Osten-Sacken, 1882

Naupoda platessa Osten-Sacken, 1882 Berl. ent. Zeit. 26: 223 Fig. 6. Bezzi,

1913 Philippine J, Sci. (D) 8: 321. Hendel, 1914a :141; 1914b :296.

Frey, 1930 Notul. ent. Helsingf. 10: 57.

Distribution: Philippines.

*3. N. regina Hendel, 1914

Naupoda regina Hendel, 1914b :298; 1914a :141. Malloch, 1929 Proc.

Linn. Soc. N.S.W. 54: 513. 1939 ibid 64: 122.

Distribution: New Guinea; Queensland (Australia).

4, N. simmondsi Bezzi, 1928

Naupoda simmondsi Bezzi, 1928 Dipt, . . . Fiji :95.

Distribution: Fiji.

5. N. strigifera de Meijere, 1919

Naupoda strigifera de Meijere, 1919 Bijdr. Dierk. 21: 33 Pl. 2, Fig. 13.

Bezzi. 1928 Dipt. . . . Fiji :95.

Distribution: Sumatra.

6. N. unifasciata Bezzi, 1917

Naupoda unifasciata Bezzi,. 1917 Philippine J. Sci. (D) 12: 141. Frey,

1930 Notul. ent. Helsingf. 10: 57.

Distribution: Philippines.

*7. N. ventralis Curran, 1936

Naupoda ventralis Curran, 1936 Proc, Calif. Acad. Sci. (Ser. 4) 22: 26.

Distribution: Solomon Islands.

8. N. ypsilon v. d. Wulp, 1898

Naupoda ypsilon v. d. Wulp, 1898 Tijd. Ent. 41: 206 Pl. 10, Fig. 2.

Hendel, 1914a :141; 1914b :297.

Distribution: Java.

9. N. ypsilonoides de Meijere, 1924

Naupoda ypsilonoides de Meijere, 1924 Tijd. Ent. 67 (supp.) :42.

Distribution: Java.

LOWER—AUSTRALIAN REGION OTITIDAE 61

69. Genus: PTEROGENIA Bigot, 1859

Bigot, 1859 Rev. Mag. Zool. (Ser. 2) 11: 315 Pl. 11, Figs. 2a-2c.

Hendel, 1914a :141; 1914b :304. Malloch, 1929 Proc. Linn. Soc. N.S.W.

54: 513; 1939 ibid 64: 125.

Type species: Pterogenia singularis Bigot, 1859.

|. P. basilutea (Walker, 1857)

Lamprogaster basilutea Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 131.

Pterogenia basilutea (Walker, 1857) Hendel, 1914a :145: 1914b :329.

Distribution: Borneo.

2. P. bifasciata Enderlein, 1924

Pterogenia bifasciata Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 133.

Distribution: Sumatra.

3. P. boettcheri Frey, 1930

Pterogenia boettcheri Frey, 1930 Notul. ent. Helsingf. 10: 58.

Distribution: Philippines.

*4. P. brevis (Walker, 1865)

Platystoma breve Walker, 1865 J. Proc. Linn. Soc. Lond, 8: 120.

Pterogenia brevis (Walker, 1865) Hendel, 1914a 7145; 1914b :325,.

Distribution: New Guinea.

5. P. centralis Bezzi, 1917

Pterogenia centralis Bezzi, 1917 Philippine J. Sci. (D) 12: 148. Frey,

1930 Notul. ent. Helsingf, 10: 60.

Distribution: Philippines.

6. P. dayak Bigot, 1859

Pterogenia dayak Bigot, 1859 Rev. Mag. Zool. (Ser. 2) 11: 316 Pl. 16,

Fig. 2. Hendel, 1914a :145: 1914b :314.

Distribution: Borneo.

7. P. divisa (Walker, 1857)

Lamprogaster divisa Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 131.

Pterogenia divisa (Walker, 1857) Hendel, 1914a :146; 1914b :324,

Distribution: Borneo.

8. P. flavicornis Frey, 1930

Pterogenia flavicornis Frey, 1930 Notul. ent. Helsingf. 10: 59.

Distribution: Philippines.

62 REC. S. AUST. MUS,, 16 (2): 1-93 March, 1970

*Q. P, fuliginosa Hendel, 1914

Pterogenia fuliginosa Hendel, 1914b :309; 1914a :146. Malloch, 1939

Proc, Linn. Soc. N.S.W. 64: 126.

Distribution: New Guinea.

10. P. glabra (Walker, 1857)

Lamprogaster glabra Walker, 1857 J. Prac. Linn. Soc. Lond. 1: 30.

Pterogenia glabra (Walker, 1857) Hendel, 1914a :146; 1914b :324.

Distribution: Singapore.

11. P. glabrella Hendel, 1914

Lamprogaster glabra Walker, 1857 pp. J. Proc. Linn. Soc. Lond. 1: 131.

Pterogenia glabrella Hendel, 1914b :310; 1914a :146.

Distribution: Sarawak,

12. P. glabrina Hendel, 1914

Lamprogaster glabra Walker, 1857 pp. J. Proc. Linn. Soc. Lond. 1: 131.

Prerogenia glabrina Hendel, 1914b :318; 1914a :146.

Distribution; Sarawak.

Walker, 1857, under the name of P. glabra included three closely-

related species—P. glabra, P. glabrella and P. glabrina. Hendel, 1914b

distinguished the three species, naming the two which were distinct from

glabra.

13. P. guttata (Walker, 1857) (No. 1)

Lamprogaster guttata Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 31,

Pterogenia guttata (Walker, 1857) Hendel 1914b :325; 1914a :146.

Distribution: Singapore.

13a. P. guttata (Walker, 1857) (No. 2)

Lamprogaster guttata Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 132.

Pterogenia guttata (Walker, 1857) Hendel, 1914b :326; 1914a :146,

Distribution: Borneo,

In 1857, Walker (Joc. cit.) applied the name Lamprogaster guttata to

two of his sp. nov. Whether these are one and the same species, or whether

they are distinct, can be determined only after the types have been examined,

It will then be possible to place them generically. I know of no evidence

to justify Hendel’s transference of them to Pterogenia, but this is accepted

here for convenience.

LOWER—AUSTRALIAN REGION OTITIDAE 63

14. P. hamifera Frey, 1930

Pterogenia hamifera Frey, 1930 Notul. ent. Helsingf. 10: 58 Pl. 1, Fig. 3.

Distribution: Philippines.

*15. P. latericia Hendel, 1914

Pterogenia latericia Hendel, 1914b :312: 1914a :146. Malloch, 1939 Proc.

Linn. Soc. N.S.W. 64: 126.

Distribution: Queensland; New South Wales (Australia).

16. P. laticeps Bezzi, 1917

Pterogenia laticeps Bezzi, 1917 Philippine J. Sci. (D) 12: 145; 1928 Dipt,

Fiji :97. Frey, 1930 Notul. ent. Helsingf. 10: 59,

Distribution: Philippines.

17. P. luteipennis Bezzi, 1917

Pterogenia luteipennis Bezzi, 1917 Philippine J. Sci. (D) 12: 147. Frey,

1930 Notul. ent. Helsingf. 10: 59.

Distribution: Philippines.

18. P. niveitarsis (Bigot, 1859)

Agastrodes niveitarsis Bigot, 1859 Rev. Mag. Zool. 11: 312.

Pterogenia niveitarsis (Bigot, 1859) Hendel, 1914b 1307; 1914a :146.

Distribution: Ceylon; Borneo.

*19. P. nubecula Hendel, 1914

Pterogenia nubecula Hendel, 1914b 134; 1914a :146, Malloch, 1939 Prac.

Linn. Soc. N.S.W. 64: 126.

Distribution: Queensland; New South Wales (Australia),

*20. P. nudiseta Bezzi, 1928

Pterogenia nudiseta Bezzi, 1928 Dipt. F. yi. 297.

Distribution: Lord Howe Island (Australia).

21. P. parva Bezzi, 1917

Pterogenia parva Bezzi, 1917 Philippine J. Sci. (D) 12: 144. Frey, 1930

Notul. ent. Helsingf. 10: 59,

Distribution: Philippines.

*22. P. pectoralis Hendel, 1914

Pterogenia pectoralis Hendel, 1914b :316; 1914a :146. Malloch, 1939

Proce. Linn, Soc. N.S.W. 64: 126.

Distribution: New Guinea.

64 REC. 8S. AUST. MUS., 16 (2): 1-93 March, 1970

23. P. punctata (Walker, 1857)

Lamprogaster punctata Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 132.

Pterogenia punctata (Walker, 1857) Hendel, 1914b :324; 1914a :146.

Distribution: Borneo.

24. P. ruficrus Hendel, 1914

Prerogenia ruficrus Hendel, 1914b :321; 1914a 7146.

Distribution: Java.

*25, P. scutellaris (Walker, 1859)

Lamprogaster scutellaris Walker, 1859 J. Proc. Linn, Soc. Lond, 3: 112.

Pterogenia scutellaris (Walker, 1859) Hendel, 1914b :327; 1914a :146.

Distribution: New Guinea.

*26. P. similis Malloch, 1939

Pterogenia similis Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 126.

Distribution: Queensland (Australia).

27. P. singularis Bigot, 1859

Pterogenia singularis Bigot, 1859 Rev. Mag. Zool. (ser. 2) 11: 315 Pl. ll,

Figs. 2a-c. Hendel, 1914a 7146; 1914b :305.

Distribution: Amboina, Celebes.

28. P. suberuciata Frey, 1930

Pterogenia subcruciata Frey, 1930 Notul. ent. Helsingf. 10: 60 Pl. 1, Fig. 4.

Distribution: Philippines.

29. P. tristis Bezzi, 1917

Pterogenia tristis Bezzi, 1917 Philippine J. Sci. (D) 12: 143. Frey, 1930

Notul. ent. Helsingf. 10: 59.

Distribution: Philippines.

30. P. truncatula (Walker, 1857)

Lamprogaster truncatula Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 31.

Pterogenia truncatula (Walker, 1857) Hendel, 1914b :325; 1914a :146.

Distribution: Singapore.

31. P. valida Bezzi, 1917

Pterogenia valida Bezzi, 1917 Philippine J. Sci. (D) 12: 142, Frey, 1930

Notul. ent. Helsingf. 10: 58.

Distribution: Philippines.

LOWER—AUSTRALIAN REGION OTITIDAE 65

32. P. variipennis Walker, 1861

Pterogenia variipennis Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 292.

Hendel, 1914a :146; 1914b :326.

Distribution: Moluccas.

33. P. vittata (Walker, 1857)

Lamprogaster vittata Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 31.

Pterogenia vittata (Walker, 1857) Hendel, 1914b :328: 1914a :146.

Distribution: Sumatra; Singapore.

34. P. vittifinis Walker, 1861

Pterogenia vittifinis Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 292. Hendel,

1914a :146; 1914b :327.

Distribution: Moluccas.

35. P. zonata (Walker, 1857)

Lamprogaster zonata Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 30,

Pterogenia zonata (Walker, 1857) Hendel, 1914a :146; 1914b :328,

Distribution: Singapore.

69A. Genus: NEOHEMIGASTER Malloch, 1939

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 126.

Type species: Hemigaster albovittatus Rondani, 1875.

1. N. albovittata (Rondani, 1875)

Hemigaster albovittatus Rondani, 1875 Ann. Mus. Stor. nat. Genova 7: 431.

Pterogenia albovittata (Rondani, 1875) Hendel, 1914b :322.

Neohemigaster albovittata (Rondani, 1875) Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64: 127.

Distribution: Borneo.

69B. Genus: CHAETOSTICHIA Enderlein, 1924

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 133.

Type species: Chaetostichia aduncivena Enderlein, 1924,

1. C. aduncivena Enderlein, 1924

Chaetostichia aduncivena Enderlein, 1924 (ibid) 11: 134.

Distribution: Java.

66 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

70. Genus: EUPROSOPIA Macquart, 1847

Macquart, 1847 Dipt. exot. supp. 2: 89. Hendel, 1914a :147; 1914b

:329. Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 136. Malloch, 1928

Proc. Linn. Soc. N.S.W. 53: 343; 1928 ibid 53: 612; 1929 ibid 54: 512;

1939 ibid 64: 145; 1931 Proc. U.S. nat. Mus. 78 (15): 1.

Type species: Euprosopia tenuicornis Macquart, 1847.

1. E. albifacies (Doleschall, 1858)

Pachycephala albifacies Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 117.

Euprosopia albifacies (Doleschall, 1858) Hendel, 1914a :149; 1914b :360.

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 136.

Distribution: Amboina.

*2. E. albolineata de Meijere, 1913

Euprosopia albolineata de Meijere, 1913 Nov. Guin, 9: 367 Pl. 10, Fig.

39. Hendel, 1914a :149; 1914b :345.

Distribution: New Guinea.

*3. E. alticeps Malloch, 1940

Euprosopia alticeps Malloch, 1940 Ann. Mag. nat. Hist. 6: 81.

Distribution: Solomon Islands.

4, E. atomaria (Walker, 1861)

Platystoma atomaria Walker, 1861 J. Proc. Linn. Soc. Lond. 5: 293.

Euprosopia atomaria (Walker, 1861) Hendel, 1914a :149.

Distribution: Moluccas,

*5. E. aureovitta Malloch, 1939

Euprosopia aureovitta Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 151 Pl. 5,

Fig. 42.

Distribution: New Guinea.

*6. E. australis (Walker, 1849)

Platystoma australe Walker, 1849 List Dipt. Brit. Mus. 4: 1061.

Euprosopia australis (Walker, 1849) Hendel, 1914a :149: 1914b :363.

Distribution: Australia (sic.).

7. E. basalis (Walker, 1860)

Platystoma basale Walker, 1860 J. Proc. Linn. Soc. Lond. 4: 148.

Euprosopia basalis (Walker, 1860) Hendel, 1914a :149; 1914b :361.

Distribution: Celebes.

LOWER—AUSTRALIAN REGION OTITIDAE 67

*8. E. biarmata Malloch, 1929

Euprosopia biarmata Malloch, 1929 Proc. Linn. Soc. N.S.W. 54: 512; 1930

ibid 55: 431.

Distribution: Queensland (Australia).

*9. E. bilineata de Meijere, 1906

Euprosopia bilineata de Meijere, 1906 Noy. Guin. 5 zool. Dipt. :92 Pl. 1,

Fig. 14. Hendel, 1914a :149; 1914b :349. Malloch, 1939 Proc. Linn.

Soc. N.S.W. 64: 149 Pl. 5, Fig. 35.

Distribution: New Guinea.

10. E. brevicornis Hendel, 1914

Euprosopia brevicornis Hendel, 1914b :335; 1914a :149.

Distribution: Borneo.

11. E. calypterata Enderlein, 1912

Euprosopia calypterata Enderlein, 1912 Zool. Jahrb. 33: 359 Fig. D.

Hendel, 1914a :149; 1914b :350.

Distribution: Sumatra.

12. E. chalybea Frey, 1930

Euprosopia chalybea Frey, 1930 Notul. ent. Helsingf. 10: 60 Pl. 1, Fig. 6.

Distribution: Philippines.

*13. E. conjuncta Hendel, 1914

Euprosopia conjuncta Hendel, 1914b :339; 1914a :149. Malloch, 1928

Proc. Linn. Soc. N.S.W. 53: 345; 1929 ibid 54: 512; 1930 ibid 55: 429,

Distribution: Queensland; Northern Territory (Australia).

*14. E. connexa Malloch, 1940

Euprosopia connexa Malloch, 1940 Ann. Mag. nat. Hist. 6: 79.

Distribution: Solomon Islands,

15. E. curta (Osten-Sacken, 1882)

Notopsila curta Osten-Sacken, 1882 Berl. ent. Z. 26: 210.

Euprosopia curta (Osten-Sacken, 1882) Hendel, 1914a :149: 1914b :332.

Bezzi, 1917 Philippine J. Sci. (D) 12: 149. Frey, 1930 Notul. ent.

Helsingf. 10: 62.

Distribution: Philippines; Taiwan.

68 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*16, E. depressifrons Malloch, 1940

Euprosopia depressifrons Malloch, 1940 Ann. Mag. nat. Hist. 6: 86.

Distribution: Solomon Islands.

17. E. dorsata Hendel, 1914

Euprosopia dorsata Hendel, 1914b :340; 1914a :149. Malloch, 1931 Proc.

U.S, nat. Mus. 78 (15): 9.

Distribution: Ceylon, Philippines.

*18. E. dubitalis Malloch, 1939

Euprosopia dubitalis Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 149 PI. 5,

Fig. 33.

Distribution: New Guinea.

19. E. frontalis (Walker, 1861)

Platystoma frontale Walker, 1861 J. Proc. Linn, Soc. Lond. 5: 268.

Eupresopia frontalis (Walker, 1861) Hendel, 1914a :149; 1914b :362.

Distribution: Celebes.

*20. E. fusifacies (Walker, 1859)

Platystoma fusifacies Walker, 1859 J. Proc. Linn. Soc. Lond, 3: 113.

Euprosopia fustfacies (Walker, 1859) Osten-Sacken, 1881 Ann, Mus. Stor,

nat. Genova 16: 473. Hendel, 1914a :149; 1914b :347, Malloch, 1939

Proc. Linn, Soc. N.S.W. 64; 151 Pl. 5, Fig. 41.

Euprosopia squamifera de Meijere, 1913 Nov. Guin. 9: 368 Pl. 10, Fig. 40.

Distribution: New Guinea.

21. E. gigas Bezzi, 1917

Euprosopia gigas Bezzi, 1917 Philippine J. Sci. (D) 12: 151. Frey, 1930

Notul, ent, Helsingf. 10: 62.

Distribution: Philippines.

*22. E. impingens (Walker, 1865)

Platystoma impingens Walker, 1865 J. Proc. Linn. Soc. Lond. 8: 134.

Euprosopia impingens (Walker, 1865) Hendel, 1914b :345; 1914a :149.

Edwards, 1915 Trans. zool, Soc. Lond. 20: 416. Malloch, 1939 Prac,

Linn. Soc, N.S.W. 64: 151 Pl. 5, Fig. 40.

Lepidocompsia impingens (Walker, 1865) Enderlein, 1924 Mitt. zool. Mus,

Berlin 11: 137.

Distribution: New Guinea.

The status of Enderlein’s monotypic genus, Lepidocompsia, cannot at

present be determined.

LOWER—AUSTRALIAN REGION OTITIDAE 69

*23. E. innocua Malloch, 1939

Euprosopia innocua Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 152 Pl. 5,

Fig. 43.

Distribution: New Guinea.

Malloch (ibid) suggests that this species is possibly Tetrachaetina

burgersiana Enderlein, 1924 q.v.

*24. E. insulicola Malloch, 1940

Euprosopia insulicola Malloch, 1940 Ann. Mag. nat. Hist. 6; 84.

Distribution: Solomon Islands.

*25. E. lepida Curran, 1936

Euprosopia lepida Curran, 1936 Proc. Calif. Acad. Sci. (4th Ser.) 22 (1):

25 Pl. 1, Fig. 2.

Distribution: Guadalcanal Islands (Solomon Islands).

26. E. lepidophora Bezzi, 1917

Euprosopia lepidophora Bezzi, 1917 Philippine J. Sci. (D) 12: 153. Frey,

1930 Notul, ent. Helsingf. 10: 47.

Distribution: Philippines.

27. E. longicornis Bezzi, 1917

Euprosopia longicornis Bezzi, 1917 Philippine J. Sci. (D) 12: 154. Frey,

1930 Notul. ent. Helsingf. 10: 62. Malloch. 1931 Proc. U.S. nat. Mus.

78 (15): 9.

Distribution: Philippines.

*28. E, macrotegularia Malloch, 1928

Euprosopia macrotegularia Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 345

Fig. 1; 1929 ibid 54: 512: 1930 ibid 55: 430.

Distribution: Queensland (Australia).

*29. E, maculipennis (Guérin), 1831

Platystoma maculipennis Guérin, 1831 Voy. Coquille Liv. 24, Pl. 21, Fig.

8, p. 299.

Euprosopia maculipennis (Guérin, 1831) Hendel, 1914b :358; 1914a :149,

Enderlein, 1924 Mitr. zoal. Mus. Berlin 11: 136. Malloch, 1928 Proc.

Linn. Soc. N.S.W. 53: 346; 1928 ibid 53: 612; 1930 ibid 55: 430.

Distribution: Queensland (Australia).

For details of publication date of E. maculipennis see Musgrave, 1932

Bibliography of Australian Entomology 1775-1930 p. 133.

70 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

30. E. metallica Malloch, 1931

Euprosopia metallica Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 6.

Distribution: Philippines.

*31, E. miliaria? Hendel, 1914

Platystoma pectorale Walker, 1862 J. Proc. Linn. Soc. Lond. 6: 13,

Euprosopia miliaria Hendel, 1914b :353, 1914a 3149, Malloch, 1928

Proc. Linn. Soc. N.S.W. 53: 346; 1929 ibid 54: 512; 1939 ibid 64: 148.

Distribution: New Guinea; Queensland (Australia).

Some doubt attaches to the specific name. Walker (1862) described

this species as Platystoma pectorale. Although Hendel (b) was aware of

this, when he transferred the species to Euprosopia, he substituted the specific

name miliaria for pectorale, but gave no reasons for doing so. Neither

Malloch (1928) could, nor Steyskal (in correspondence) can, give any

explanation for his action.

Since attempted restoration of pectorale would only further cloud the

issue, miliaria is here temporarily retained.

32. E. millepuncta Bezzi, 1917

Euprosopia millepuncta Bezzi, 1917 Philippine J. Sci. (D) 12: 152. Frey,

1930 Notul. ent. Helsingf. 10: 62. Malloch, 1931 Proc. U.S. nat. Mus.

78 (15): 10.

Distribution: Philippines.

*33. E. minor Malloch, 1940

Euprosopia minor Malloch, 1940 Ann. Mag. nat. Hist. 6: 82.

Distribution: Solomon Islands,

*34. EK, minuta Malloch, 1939

Euprosopia minuta Malloch, 1939 Proc. Linn, Soc. N.S.W. 64: 148 Pl. 5,

Fig, 32,

Distribution: New Guinea.

35. E. mohnikei (Doleschall, 1858)

Pachycephala mohnikei Doleschall, 1858 Nat. Tijd. Ned. Ind. 17: 116.

Notopsila mohnikei (Dol. 1858) Osten-Sacken, 1882 Berl. ent. Z, 26: 209.

Oncoscelia mohnikei (Dol. 1858) Enderlein, 1924 Mirt. zool. Mus. Berlin

11: 136,

Euprosopia mohnikei (Dol. 1858) Hendel, 1914a :150; 1914b :359,

Distribution: Amboina.

See note under Notopsila.

LOWER—AUSTRALIAN REGION OTITIDAE 71

*36. E. multivitta (Walker, 1859)

Platystoma multivitta Walker, 1859 J. Proc. Linn, Soc. Lond. 3: 113.

Euprosopia multivitta (Walker, 1859) Hendel, 1914b :362; 1914a :150.

Distribution: New Guinea.

37. E. nobilis Frey, 1930

Euprosopia nobilis Frey, 1930 Notul. ent. Helsingf. 10: 61 Pl. 1, Fig. 5.

Distribution: Philippines.

*38. E. penicillata Hendel, 1914

Euprosopia penicillata Hendel, 1914b :343; 1914a :150. Edwards, 1915

Trans, zool. Soc, Lond. 20: 416. Malloch, 1939 Proc. Linn. Soc, N.S.W.

64: 151.

Distribution: New Guinea.

*39. E. potens (Walker, 1862)

Platystoma potens Walker, 1862 J, Proc. Linn. Soc. Lond. 6: 12.

Euprosopia potens (Walker, 1862) Osten-Sacken, 1881 Ann. Mus. Stor.

nat. Genova 16: 472, de Meijere, 1913 Nov. Guin. 9: 369. Hendel,

1914a :150; 1914b :341. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64:

150 Pl. 5, Fig. 37.

Distribution: New Guinea; Moluccas.

40. E. producta (Walker, 1861)

Platystoma productum Walker, 1861 J. Proc. Linn. Soc. Lond. 5; 293.

Euprosopia producta (Walker, 1861) Hendel, 1914a :150; 1914b :361.

Distribution: Moluccas.

*41. E, protensa (Walker, 1864)

Platystoma protensa Walker, 1864 J. Proc. Linn. Soc. Lond. 7: 228.

Euprosopia protensa (Walker, 1864) Hendel, 1914a :150; 1914b :344,

Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 136. Malloch, 1939 Proc.

Linn. Soc. N.S.W, 64: 150 PI. 5, Fig. 36.

Distribution: New Guinea.

*42. E. punctifacies Malloch, 1928

Euprosopia punctifacies Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 346

Fig. 2; 1929 ihid 54: 512; 1930 ibid 55: 430.

Distribution: Queensland (Australia).

72 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*43. E. rufiventris Hendel, 1914

Euprosopia rufiventris Hendel, 1914b :334; 1914a :150. Malloch, 1939

Proc. Linn. Soc. N.S.W. 64: 148.

Distribution: New Guinea.

*44. E. scatophaga Malloch, 1930

Euprosopia scatophaga Malloch, 1930 Proc, Linn. Soc. N.S.W. 55: 431;

1931 Proc. U.S. nat. Mus. 78 (15): 7.

Distribution: Queensland (Australia).

45. E. semiarmata Malloch, 1931

Euprosopia semiarmata Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 3.

Distribution: Philippines.

*46, E. separata Hendel, 1914

Euprosopia separata Hendel, 1914b :338; 1914a :150. Malloch, 1928

Proc, Linn. Soc. N.S.W. 53: 344. 1929 ibid 54: 512.

Distribution: Queensland (Australia).

*47. E. setinervis Malloch, 1939

Euprosopia setinervis Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 149 PI. 5,

Fig. 34.

Distribution: New Guinea.

48. E. sexpunctata (Osten-Sacken, 1882)

Notopsila sexpunctata Osten-Sacken, 1882 Berl. ent. Z. 26: 210.

Euprosopia sexpunctata (Osten-Sacken, 1882) Hendel, 1914b :359; 1914a

:150. Bezzi, 1917 Philippine J. Sci. (D) 12: 149. Frey, 1930 Notul.

ent. Helsingf. 10: 61. Malloch, 1931 Proc. U.S. nat. Mus. 78 (15): 8.

Distribution: Moluccas; Philippines.

49. E. tarsalis (Walker, 1864)

Platystoma tarsale Walker. 1864 J, Proc. Linn, Soc. Lond. 7: 237.

Euprosopia tarsalis (Walker, 1864) Hendel, 1914b :360; 1914a :150,

Distribution: Moluccas.

*50. E. tegularia Malloch, 1928

Euprosopia tegularia Malloch, 1928 Proc. Linn. Soc. N.S.W. 53: 346, Fig. 3.

Distribution: Solomon Islands.

LOWER—AUSTRALIAN REGION OTITIDAE 73

*51. E. tenuicornis Macquart, 1847

Euprosopia tenuicornis Macquart, 1847 Dipt. exot. supp. 2: 90 Pl. 6, Fig. 4.

Hendel, 1914a :150; 1914b :336. Enderlein, 1924 Mitt. zool. Mus.

Berlin 11: 136. Malloch, 1928 Proc. Linn. Soc, N.S.W. 53: 612; 1930

ibid 55: 430; 1931 Proc. U.S. nat. Mus, 78 (15): 7.

Distribution: Queensland; New South Wales (Australia).

*52. E. tigrina Osten-Sacken, 1881

Euprosopia tigrina Osten-Sacken, 1881 Ann. Mus. Stor. nat, Genova 16:

473, Hendel, 1914a :150; 1914b :331. Malloch, 1929 Prac. Linn. Sac.

N.S.W. 54: 513; 1939 ibid 64: 147.

Distribution: New Guinea,

53. E. trivittata Bezzi, 1917

Euprosopia trivittata Bezzi, 1917 Philippine J. Sci. (D) 12: 150. Frey,

1930 Notul. ent. Helsingf. 10: 61. Malloch, 1931 Prac. U.S. nat. Mus.

78 (15): 10.

Distribution: Philippines.

54. E. ? truncata (Enderlein, 1924) (conj. nev.)

Oncoscelia truncata Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 137.

Distribution: Celebes,

See note at end of ? Notopsia.

*55. E. ventralis (Walker, 1859)

Lamprogaster ventralis Walker, 1859 J. Proc. Linn. Soc. Lond. 3: 131,

Euprosopia ventralis (Walker, 1859) Hendel, 1914b :343; 1914a :150.

Malloch, 1939 Proc. Linn. Soc. N.S.W. 64: 150 PIL. 5, Figs. 38, 39.

Distribution: New Guinea.

7OA. Genus: ? NOTOPSILA Osten-Sacken, 1882

Osten-Sacken, 1882 Berl. ent. Z, 26: 209,

The validity or otherwise of this genus has yet to be determined. It

was erected by Osten-Sacken (loc. cit.) as a nom. nov. for Pachycephala

Doleschall, 1858 (Nat. Tijd. Ned, Ind. 17: 116—Pachycephala mohnikei)

which was pre-occupied in Aves by Pachycephala Vigors, 1825.

Having overlooked Osten-Sacken’s change, Enderlein in 1924 (Mitt.

zool. Mus. Berlin 11: 136) erected his genus Oncoscelia to replace

Pachycephala Doleschall, designating P. mohnikei Dol., 1828 as the type

species. At the same time, he transferred the Ceylonese species, Euprosopia

74 REC. $8. AUST, MUS., 16 (2): 1-93 March, 1970

nigropunctata Hendel, 1914 (Abh. zool.-bot. Ges. Wien 8: 356) to Oncos-

celia and described as sp. nov. O. truncata from the Celebes (ibid 11: 137).

As N. nigropunctata does not occur in the area covered by this

catalogue, no further consideration of it is necessary. Hendel’s placement

of mohnikei is accepted, and a new combination, Euprosopia truncata 1s

proposed for Oncoscelia truncata Enderlein, until the matter has been

cleared up.

70B. Genus: ? TETRACHAETINA Enderlein, 1924 (Monotypic)

Enderlein, 1924 Mirt. zool. Mus. Berlin 11: 138.

Type species: 7. burgersiana Enderlein, 1924.

*1. T. burgersiana Enderlein, 1924

Tetrachaetina burgersiana Enderlein, 1924 Mitt. zool. Mus. Berlin 11: 138.

Distribution: New Guinea.

This genus is doubtful. Malloch, 1939 (Proc, Linn. Soc, N.S.W. 64:

153) claimed that it “is not tenable”; he also suggested that his own

Euprosopia innocua (Malloch, 1939 ibid) might be a synonym of T.

burgersiana. Tetrachaetina will probably later be rejected as a synonym of

Euprosopia.

76. Genus: EUTHYPLATYSTOMA Hendel, 1914

Hendel, 1914b :398.

Type species: Platystoma rigidum Walker, 1857.

1. E. plumatum Hendel, 1914

Euthyplatystoma plumatum Hendel, 1914b :399; 1914a :160.

Distribution: Singapore.

2. E. rigidum (Walker, 1857)

Platystoma rigidum Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 32.

Euthyplatystoma rigidum (Walker, 1857) Hendel, 1914b :398; 1914a ;160

Pl. 15, Fig. 284. Malloch, 1939 Proc. Linn. Soc. N.S.W. 64; 153.

Distribution: India; Singapore; Celebes,

3. E. superbum (v. d. Wulp, 1881)

Platystoma superbum y. d. Wulp, 1881 Dipt. Sumatra Exped. :50 PI. 3,

Fig. 5.

Euthyplatystoma superbum (v. d. Wulp, 1881) Hendel, 1914b :401; 1914a

:160.

Distribution: Sumatra.

LOWER—AUSTRALIAN REGION OTITIDAE 75

77. Genus: VALONIA Walker, 1857 (Monotypic)

Walker, 1857 J. Proc. Linn. Soc. Lond. 1: 34.

Type species: Valonia complicata Walker, 1857,

1. V. complicata Walker, 1857

Valonia complicata Walker, 1857 J, Proc. Linn. Soc. Lond. 1: 34 Pl. 1,

Fig. 6. Hendel, 1914a :161; 1914b :402. Enderlein, 1924 Mitt. zeol.

Mus. Berlin 11: 150.

Distribution: Singapore.

II. Sub-family: OTITINAE

Type genus: Otites Latreille, 1804 Nouv. Dict. Hist. nat. 24: 196.

1. Genus: CERATOXYS Rondani, 1861

Rondani, 1861 Dipt. Ital. Prodr. 4: 10. Curran, 1934 Fam, Gen. N.

Amer. Dipt. :281.

Ceratoxys was proposed by Rondani to replace Meckelia R.-D., 1830

(Mém. Prés. Acad. Sci. Paris 2: 714) which was pre-occupied by Meckelia

Leucart, 1828 (Vermes). Anacampta Loew, 1868 (Z. Naturw. Berlin 32:

7) is a synonym.

1. C. latiuscula (Loew, 1873)

Anacampta latiuscula Loew, 1873 Smithson. misc. Call. 11 (3): 130.

Distribution: North America (introduced into Hawaii).

Il. Sub-family: ULIptNaAg

Type genus: Ulidia Meigen, 1826 Syst. Beschr. Eur. zweifl. Ins. 5: 385.

3. Genus: PHYSIPHORA, Fallén, 1810

Fallén, 1810 Sp. Ent. nov. Dipt. Dispon. Methodus :11.

Syn.: Chrysomyza Fallén, 1817 Dipt. Suec. Scenopinii :3.

Type species: Musca demandata Fabricius, 1798.

Until recently this genus has been wrongly known as Chrysomyza

Fallén, 1817. After erecting Physiphora in 1810, Fallén thought that it was

pre-occupied (which it was not), and suggested Chrysomyza in 1817 as a

substitute. Since Physiphora is valid, Chrysomyza is to be rejected as a

synonym.

76 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

*1. P. aenea (Fabricius, 1794)

Musca aenea Fab., 1794 Ent. Syst. 4: 335.

Chrysomyza aenea (Fab., 1794) Hendel, 1910 Gen. Ins. 106: 21 (for all

references up to 1909).

Chrysomyza sp. Grimshaw, 1902 Fauna Hawaiiensis 3 (2): 85 (probably

C. aenea).

Chrysomyza aenea (Fab., 1794) Bezzi, 1913 Philippine J. Sci. (D) 8: 321.

Knab, 1916 Brooklyn ent. Soc. Bull, 11: 43 Fig. 1. Hardy, 1920 Proc.

Linn. Soc. N.S.W. 45: 472 (? Identification). Malloch, 1930 J/ns.

Samoa 6 (5): 215. Curran, 1934 N, Amer. Dipt. :277 Figs. 27, 46.

Hennig, 1941 Ent. Beihefte 8: 117. Malloch, 1942 B.P. Bishop Mus.

Bull. 172: 205.

Physiphora aenea (Fab., 1794) Steyskal, 1952 Occ. Pap. B.P. Bishop Mus,

20 (15): 285.

Distribution: India; Java; Sumatra; Borneo; New Guinea; New South

Wales; Queensland; Northern Territory (Australia); New Caledonia; New

Hebrides; Solomon Islands; Philippines; Taiwan; Samoa; Marshall Islands;

Marianas Islands; Guam; Hawaii.

All records of this species need careful checking since it is easily

confused with P. demandata (Fab., 1798).

*2, P. aperta Steyskal, 1952

Physiphora aperta Steyskal, 1952 Occ. Pap. B.P. Bishop Mus, 20 (15); 285

Figs. la, b.

Distribution: Solomon Islands.

*3. P. demandata (Fab., 1798)

Musca demandata Fabricius, 1798 Ent. Syst. (Suppl.) :564.

Chrysomyza demandata (Fab., 1798) Hendel, 1910 Gen. Jns, 106: 21.

Knab, 1916 Brooklyn ent. Soc. Bull. 11: 41 Fig. 2.

Physiphora demandata (Fab., 1798) Steyskal, 1952 Occ. Pap. B.P, Bishop

Mus, 20 (15): 284.

Distribution: Cosmopolitan: Europe; Africa: North America;

Australia,

The one Australian record for this species consists of specimens caught

and identified by me in various parts of South Australia. I have also reared

adults from larvae infesting wet, badly-made ensilage. Despite wide collecting

over more than forty years, | have not seen, in South Australia, a single

specimen of aenea. These facts suggest to me the possibility that some at

LOWER—AUSTRALIAN REGION OTITIDAE 77

least of the specimens recorded from Australia as aenea may well be mis-

identifications. The two species may be readily distinguished by the wing

venation. Knab, 1916 (Brooklyn ent. Soc. Bull. 11: 43 Figs. 1, 2) gives

good illustrations of the wings of both.

4. Genus: EUXESTA Loew, 1867

Loew, 1867 Berl. ent, Zeit. 11: 297 Pl. 2, Figs. 7-20. Hendel 1910

Gen, Ins. 106: 22 (see for all references prior to 1909). Malloch, 1930

Ins. Samoa 6 (5): 215; 1932 B.P. Bishop Mus. Bull. 98: 208. Curran,

1934 N. Amer. Dipt. :277 Figs. 64; 85.

Type species: Urophoera quadrivittata Macquart, 1835.

|. E. hyalipennis Malloch, 1932

Euxesta hyalipennis Malloch, 1932 B.P. Bishop Mus. Bull. 98: 209.

Steyskal, 1952 Occ. Pap. B.P. Bishop Mus. 20 (15): 280.

Distribution: Marquesas Islands.

2. E. lafooni Steyskal, 1952

Euxesta lafooni Steyskal, 1952 Occ. Pap. B.P. Bishop Mus. 20 (15): 280.

Distribution: New Hebrides.

3. E. pruinosa Malloch, 1932

Euxesta pruinosa Malloch, 1932 B,P. Bishop Mus, Bull, 98: 210. Steyskal,

1952 Occ, Pap. B.P. Bishop Mus. 20 (15): 282.

Distribution: Marquesas Islands.

4. E. quadrivittata (Macquart, 1835)

Urophora quadrivittata Macquart, 1835 Hist. nat, Dipt. 2: 456.

Euxesta quadrivittata (Macquart, 1835) Knab, 1916 Brooklyn ent. Soc.

Bull. 11: 44. Steyskal, 1952 Occ. Pap. B.P. Bishop Mus. 20 (15): 282.

E. exilis Knab, 1916 (loc. cit.).

Distribution: Florida; Cuba; Jamaica (introduced into Hawaii and

Philippines).

5, E. semifasciata Malloch, 1930

Euxesta semifasciata Malloch, 1930 Ins. Samoa 6 (5): 216 Fig. 1.

Distribution: Ellice Islands.

The possibility that one or more of the species, Ayalipennis, lafooni,

pruinosa, and semifasciata may be introductions from Central America or

the Caribbean area should not be overlooked.

78 REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

7. Genus: PSEUDEUXESTA Hendel, 1910

Hendel, 1910 Gen. Ins. 106: 30.

Type species: Pseudeuxesta prima ( Osten-Sacken, 1881).

*1. P. prima (Osten-Sacken, 1881)

Pseudeuxesta prima Osten-Sacken, 1881 Ann. Mus. Stor. nat. Genova 16;

470,

Euxesta semifasciata Malloch, 1930 Ins, Samoa 6 (5): 216.

Pseudeuxesta prima (Osten-Sacken, 1881) Hendel, 1910 Gen. Ins. 106: 32;

1931 Verh, zool.-bot. Ges. Wien 81: 4. Malloch, 1939 Proc. Linn. Soc.

N.S.W. 64; 98. Steyskal, 1952 Occ. Pap. B.P. Bishop Mus. 20 (15):

286.

Distribution: Celebes; Seychelles; Ceram; New Guinea; Solomon

Islands; Palau Islands; Truk Islands; Marshall Islands; Marianas Islands;

Hawail.

17. Genus: ACROSTICTA Loew, 1867

Loew, 1867 Berl. ent, Z. 11: 293 Pl. 2, Fig. 5; 1873 Smithson. misc.

Coll. 11 (3): 151. Hendel, 1910 Gen. Ins. 106: 50.

Type species: Acrosticta scrobiculata Loew, 1867.

1. A. apicalis (Williston, 1896)

Euxesta apicalis Williston, 1896 Trans. ent. Soc. Lond. 12: 375 Pl. 12,

Fig. 128.

Acrosticta pallipes Grimshaw, 1901 Fauna Hawaiiensis 3 (1): 44; 1902

ibid 3 (2): 85. Hendel, 1910 Gen. Ins. 106: 52. Bezzi, 1928

Dipt. Fiji :89. Acrosticta apicalis (Williston, 1896) Malloch, 1930

Ins. Samoa 6 (5): 217; 1932 B.P. Bishop Mus. Bull. 98: 206. Bryan,

1934 Proc. Haw. ent. Soc. & (3): 430. Steyskal, 1952 Occ. Pap. B.P.

Bishop Mus. 20 (15): 279.

See Steyskal, 1952 for complete bibliography.

Distribution: West Indies; (introduced into Hawaii).

21. Genus: NOTOGRAMMA Loew, 1867

Loew, 1867 Berl. ent, Z. 11: 289; 1873 Smithson. misc. Coll. 3: 148.

Hendel, 1910 Gen. Ins. 106: 58.

Type species: Notogramma cimiforme Loew, 1867,

LOWER—AUSTRALIAN REGION OTITIDAE 79

1. N. cimiforme Loew, 1867

Notogramma cimiformis Loew, 1867 Berl. ent. Z. 11: 289, Loew, 1867

described N. cimiformis; in 1873 (Smithson. Misc. Coll. 11 (3): (148)

he mistakenly synonymized it with N. stigma (Fab., 1798). Since then,

both species have been confused. Steyskal, 1963 (Proc. ent. Soc. Wash.

65: 196) has distinguished the two species and emended cimiformis to

cimiforme in conformity with the gender of Notogramma. N. stigma has

never been correctly recorded from any Pacific Island; all such records

should be referred to N. cimiforme. (See Steyskal, loc. cit. for complete

bibliography. )

Distribution: West Indies; Central and South America. (Introduced

into the following places: Hawaii; Marianas Islands; Guam; Palau Islands

and Wake Island. )

80 REC. 8. AUST. MUS., 16 (2): 1-93 March, 1970

APPENDIX

GENERA OF ULIDIINAE ERECTED AFTER 1910

Hendel’s work on the sub-family, Ulidiinae, appeared in 1910 (Gen.

Ins. fascicle 106). Since that year, three new genera were erected by

Malloch who, however, did not refer them to Hendel’s arrangement of

genera. In these circumstances, I am unable to do so, and therefore present

them in this appendix.

1. Genus: HETERODOXA Malloch, 1932

Malloch, 1932 B.P. Bishop Mus. Bull. 98: 211.

Type species: Heterodoxa uapouae Malloch, 1932.

1. H. fatuhivae Malloch, 1932

Heterodoxa fatuhivae Malloch, 1932 ibid 98: 214 Fig. 49d.

Distribution: Marquesas Islands.

2. H. hivaoae Malloch, 1932

Heterodoxa hivaoae Malloch, 1932 ibid 98: 213 Fig. 49b.,

Distribution: Marquesas Islands.

3. H. uahukae Malloch, 1932

Heterodoxa uahukae Malloch, 1932 ibid 98: 214 Fig. 49c.

Distribution: Marquesas Islands,

4, H. uapouae Malloch, 1932

Heterodoxa uapouae Malloch, 1932 ibid 98: 212 Fig. 49a.

Distribution: Marquesas Islands.

2. Genus: NEOEUXESTA Malloch, 1930

Malloch, 1930 Ins. Samoa 6 (5): 218.

Type species: Neoeuxesta fumicosta Malloch, 1930.

1. N. fumicosta Malloch, 1930

Neoeuxesta fumicosta Malloch, 1930 ibid 6 (5): 218. Steyskal, 1952 Occ.

Pap. B.P. Bishop Mus, 20: 283.

Distribution: Samoa.

LOWER—AUSTRALIAN REGION OTITIDAE

2. N. guamana Steyskal, 1952

Neoeuxesta guamana Steyskal, 1952 ibid 20: 283 Fig. 1d.

Distribution: Guam.

3. Genus: PERISSONEURA Malloch, 1932

Malloch, 1932 B.P. Bishop Mus. Bull. 98: 207.

Type species: Perissoneura diversipennis Malloch, 1932.

1. P. diversipennis Malloch, 1932

Perissoneura diversipennis Malloch, 1932 ibid 98: 207 Fig. 28.

1952 Occ. Pap. B.P. Bishop Mus. 20: 284.

Distribution: Marquesas Islands.

Steyskal,

Achias

Achiosoma

Antineura

Acrosticta

A pactoneura

Asyntona

Brea

Ceratoxis

Chaetorivellia

Chaetostichia

Cleitamia

Cleitamoides

Conicipithea

Dasyortalis

Duomyia

Elassogaster

Euprosopia

Euthyplatystoma

Euxesta

Euxestomoea

Guamomyia

Heterodoxa

Icteracantha

Imugana

Laglaisia

Lamprogaster

Lamprophthalma

Lasioxiria

Lenophila

Lepidocompsia

Leriomyia

Loxoneuroides

Loxonevra

Meringomeria

Mesoctenia

Microepicausta

Mindanaia

Montrouziera

Naupoda

Neoeuxesta

Neohemigaster

Notoegramma

Notopsila

Perissoneura

Peronotrochus

Philocompus

Physiphora

Picrometopus

REC. S. AUST. MUS., 16 (2): 1-93

INDEX TO GENERA

Fabricius, 1805 .. .. .. 4. e

Hendel, 1914. .. .. «2 se.

Osten-Sacken, 1881 .. .. ..

Loew, 1867 ..

Malloch, 1930... .. ss as

Osten-Sacken, 1881 .. .. .. -.

Walker, 1859. .. 4. -e

Rondani, 186] .. .. .. ..

de Meijere, 1913 . ..

Enderlein, 1924 ,.

Macquart, 1835 ..

Malloch, 4935 4.4 64. sna

Hendel, 1914... ..

Hendel, 1914 . .. .. ..

Walker, 1859 . 1. 1. wa ae

Bigot, 1859-00, 22 ate aie ee ye om

Macquart, 1847 ..

Hendel, 1914 .

Loew, 1867 .. ..

de Meijere, 1913 .

Malloch: 194%. .4 ues es ae

Malloch, 1932... .. .. 4.

Hendel, 1912 . .. ..

Enderlein, 1937 ..

Bigot, 1880... -. .. an ee ee

Macquart, 1843 .. ke 4

Portschinsky, 1892 .. ..

Hendel, 1914...

Guérin, 1843 .

Enderlein, 1924 .. .. .. ..

Kertész, 1899 . 2. 2. 2.

Hendel, 1914. 2. 6. ee.

Macquart, 1835 .. .. .. 2. s

Enderlein, 1924 ..

Enderlein, 1924 .. 2. 1. 1. en ea

Hendel, 19145. 24 5 26 e6 we ae we

Malloch, [931.400 23 9 pe es he

Bigot, 1860 .. .. .. 2...

Osten-Sacken, 1881 .. 2. 6. 2. ee ae

Malloch, 1930 . .. 2. 2. eee ee

Malloch, 1939 . .. 2. 1. we we ae be

Loew, 1867 .. .. .. ..

Osten-Sacken, 1882 .. aide

Malloch, 1932 . .. .. 1. 2. we ee

Enderlein, 1924 .. .. .. ... ‘

Osten-Sacken, 1881 .. 1... 4.

Fallen, 1810 .. ..

Frey, 1930. .. .. ,. . ce a

March, 1970

LOWER—AUSTRALIAN REGION OTITIDAE

Plagiostenopterina

Poecilotraphera

Pogonortalis

Pseudepicausta

Pseudeuxesta

Pseudocleitamia

Pseudorichardia

Pterogenia

Rhegmatosaga

Rhytidortalis

Rivellia

Scelostenopterina

Scholastes

Scotinosoma

Tetrachaetina

Tropidogastrella

Tylopterna

Valonia

Xenaspis

Xenaspoides

X enognathus

Xiria

Zealandortalis

Zygaenula

Hendel, 1914 .

Hendel, 1914 . .

de Meijere, 1911

Hendel, 1914 ,

Hendel, 1910 . 84 tut wee

Malloch; 1939. 2. ie wae ee

Hendel, 1911 .

Bigot, 1859 ..

Frey, 1930.

Hendel, 1914 . es

Robineau-Desvoidy, 1830 .

Hendel, 1914 . $3

Loew, 1873 ..

Loew, 1873 .. ..

Enderlein, 1924 ..

Hendel, 1914 .

Bezzi, 1917 ..

Walker, 1857 . ;

Osten-Sacken, 1881 ..

Frey, 1930.

Malbose, 1030: ae py, eae fae SS

Walker, 1857 .

Malloch, 1930. .. .. s. se ue ee ce

Doleschall, 1858 .. .. 2... ..

REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

INDEX TO SPECIES

Page

dhana Curran, 1929 (Rivellia) .. 2. ee ee we ee ee eee 33

addens (Walker, 1860) (Conicipithea) .. 6. 2. ee ee ee ee 6

aduncivena Enderlein, 1924 (Chaetastichia) .. 2. 11 «6 we 65

aenea (Wiedemann, 1819) (Plagiostenopterina) . .. .-. +. +. 11

aenea (Fabricius, 1794) (Physiphora) . .. 6. 2. 22 e+ 2 oe 76

aeneus de Meijere, 1906 (Philocompus) . bof. opereiss 6

aequifera (Walker, 1862) (Rivellia) . 6. 1. 6. we we ee es 33

affinis Hendel, 1914 (Rivellia) © 6. 66 ce we we ne ee eee 33

aitapensis Malloch, 1939 (Scholastes) 2. 6. ee se ee ee ee 51

albertisi Osten-Sacken, 1881 (Achias) . 1. 66 we we ee ee ae 39

albifacies (Doleschall, 1858) (Euprosopia) . .. 2. .. s+ oe 66

albolineata de Meijere, 1913 (Euprosopia) .. .. -. 1+ +5 oe 66

alhopilosus de Meijere, 1915 (Elassogaster) . 1. 2. ue ee oe 16

albovittata (Rondani, 1875) (Neohemigaster) .. 2. «+ «2 +. 65

alticeps Malloch, 1940 (Euprosopia) . .. 1. 1. ee ee ne ve 66

amabalis Osten-Sacken, 1881 (Cleitamia) .. .. .. 2. ++ +e 28

amitina Frey, 1930 (Lamprogaster) .. 6. 6. ce ee ee ee oe 42

amplividens Walker, 1859 (Achias) .. 2. 6. 46 ee ve ee ne 39

angulata Hendel, 1914 (Pseudepicausta) . .. 6. 62 ee ee we 23

angusta Enderlein, 1924 (Lamprogaster) . .. .. «+ s+ ++ = 42

angustifrons Hendel, 1914 (Dasyortalis) . 2. 0. 61 ee ee ee 35

angustilimbata de Meijere, 1915 (Brea) .. .. 1 7. ee ee ee 54

annulipes Hendel, 1914 (Duomiyia) .. .. 6. 2. ee ee ee ee 19

antica Walker, 1857 (Xiria) .. 6. 6. ee ee ee ee ee 4

aperta Steyskal, 1952 (Physiphora) .. 2. 6. 2. 6 ee ee ee 76

apicaliy Malloch, 1939 (Pseudepicausta) . 2. 6. 2. 22 ea ne 23

apicalis (Walker, 1849) (Lamiprogaster) . .. 2. 2. 2. ss ee 42

apicalis (Williston, 1896) (Acrosticta) 2. 6. 6. ee ee ee ve 78

apictipennis Hennig, 1940 (Achias) . 1. 1. 4. ye ee ee ae 40

aristalis Bezzi, 1928 (Pseudorichardia) .. 2. 6. 2. 6 ee we 53

armata Malloch, 1931 (Plagiastenopterina) . 1... 6. we ws i

aspiciens (Walker, 1864) (Achiosoma) .. 4. 2.0.60 44 46 ae 39

astrolabei (Boisduval, 1833) (Cleitamia) . 2... 2. 2. oe ee 28

atomaria (Walker, 1861) (Euprosopia) .. 2. 2. 6. ce oe ee 66

attenuata (Malloch, 1931) (Scetinosoma) .. .. .. se ee ae 26

aureovitta Malloch, 1939 (Euprosopia) .. vt aches wey és

austeni Sharp, 1900 (Lamprogaster) . 6.06. 6. ee ee ee a 43

australis Malloch, 1939 (Achias) .. 2... 2. - 6 40

australis (Walker, 1849) (Euprosopia) . sete tes ate Sa 66

azurea Hendel, 1914 (Duomyiu) .. .. 2. 2. 2. 2 we we ee 19

harhata Hendel, 1914 (Dasyortalis) 2.06. 66 pe we ee ee ee 55

basalis (Walker, 1849) (Plagiostenopterina) .. 2. 2. .2 4s 11

basaliy (Walker, 1860) (Euprosopia) . .. 6. 6. we ee ee ee 66

hasalis (Walker, 1861 (Lamprogaster) .. 2. 2.02. ee ne es 43

LOWER—AUSTRALIAN REGION OTITIDAE

basalis Enderlein, 1924 (Brea) .

basilaris (Wiedemann, 1830) (Rivellia) — : re? .

basilutea (Walker, 1857) (Pterogenia) .. .. ..

hataviensis (Schiner, 1868) (Pseudepicausta) ..

bezziana Steyskal, 1952 (Pseudorichardia) .. ..

hbiarcuata (Walker, 1865) (Cleitamia)

hbiarmata Malloch, 1929. (Euprosopia)

hicolor Macquart, 1847 (Lampregaster) Pa

bicolor Frey, 1930 (Picrometopus) .. 2. .. 2...

bifasciata Enderlein, 1924 (Pterogenia) .. .. .. .. .

bilineata de Meijere, 1906 (Euprosepia) . .. ..

bimaculatuy Hendel, 1914 (Scholastes) ..

bipunctata Hendel, 1914 (Euxestomoea) .

biroi de Meijere, 1906 (Antineura) .. ..

biroi Hendel, 1914 (Laglaisia) . ..

bistrigata Hendel, 1914 (Scotinosoma)

hoettcheri Frey, 1930 (Pterogenia) .. .. .. 00. se ue

brachyophthalmus Walker, 1865 (Achias) .. .. .. ..

brevicornis Hendel, 1914 (Euprosopia) .. ..

brevis (Walker, 1865) (Pterogenia) .

bryani Malloch, 1930 (Xenognathus) . .. “$ iY a f : 4: :

burgersiana Enderlein, 1924 (Tetrachaetina) . .

caerulea (Macquart, 1846) (Lenophila) . .. ..

calearata (Macquart, 1843) Magiotenaglarina]

caloptera Bigot, 1880 (Laglaisia) .. .. .

calypterata Enderlein, 1912 (Euprosopia)

carolinensis Enderlein, 1924 (Scholastes) . 6. 16 6s 64 os

catharinae de Meijere, 1913 (Cleitamia) .

cavenda Bezzi, 1917 (Lamprophthalma) .

celebensis Enderlein, 1924 (Eamiprogaster)

celyphoides Walker, 1859 (Mesoctenia) (See Zygaenula) ;

centralis Bezzi, 1917 (Pterogenia) . .. 7 eh 8

chalybea (Doleschall, 1858) (Pseiidepicausta) .

chalyhea Frey, 1930 (Eupresopia) .

chalyheiventriy (Wiedemann, 1830) ticteracuntha}-,

cheesmanae Malloch, 1939 (Cleitamia) .. .. ..

cimiforme Loew, 1867 (Notogramma) .. .. +.

cinctus Guérin, 1830 (Scholastes) .

clarissa Frey, 1930 (Xiria) .

coalescens Hendel, 1914 (Zpeaenula)

commoni Paramonoy, 1957 (Pogenortalis) .

complens (Walker, 1859) (Dasyortalis) .

completa (Malloch, 1931) (Scotinoesoma)

complicata Walker, 1857 (Vadlonia) ..

concisivitta (Walker, 1862) (Rivellia)

conjuncta Hendel, 1914 (Euprosopia) . 2. 6. 2. ee ee ee oe

connata (Thomson, 1868) (Rivellia) .

connexa Hendel, 1914 (Rivellia) ..

REC. S. AUST. MUS., 16 (2): 1-93 March, 1970

Page

connexa Malloch, 1940 (Euprosopia) . Ba kat) aah bale 67

contrahens (Walker, 1860) (Pseudepicausta) pe thre +5 23

contraria Walker, 1859 (Brea)... .. ae eee 54

costalis Walker, 1861 (Lamprogaster) . 1.0.5 25 25 ce ee ae 43

costalis Malloch, 1939 (Achiosoma) .. .. has 24 25 39

cristiventris (Gerstaicker, 1860) (Tropidogastrellay sp eh pha tas 57

cupidus Osten-Sacken, 1882 (Philocompus) . 2. 6. ss ue a 6

curta (Osten-Sacken, 1882) (Euprosopia) .. 2. 66 ee ee ee 67

cyanea Frey, 1930 (Xenaspoides) . .. wie qonie “a; ta wile 9

cyaneiventris (y. d. Wulp, 1881 (Icteracartha) . ast “Ose ease 19

cyclops Malloch, 1939 (Cleitamia) .. 6. 6. ce ce ee ee ee 29

dacoides (Walker, 1865) (Achivsonia) .. 6. 66 ee ee ee 39

dayak Bigot, 1859 (Pterogenia) .. . a ate Se xe ite 6l

decatomoides (Walker, 1862) (Riv eli) £4 bf fe wy we 34

decolor Malloch, 1939 (Lamprogaster) . oe wes ee Ph ols 43

decora (Fabricius, 1805) (Loxonevra) .. 6. 26 ee ae eee 38

decora (Macquart, 1846) (Duomyia) .. . ae die te Ll 20

decora (de Meijere, 1911) (Tropidogastrelia) wey Ngee eae fe 57

delandi Malloch, 1939 (Cleitamia) . wk re get sd ee 29

demandata (Fabricius, 1798) (Phy phe. te ite pet ek 76

dentipes (Macquart, 1843) (Lenophila) . 1. 1. 6. we ee ee 50

depressifrons Malloch, 1940 (Euprosopia) .. 1... 2. ee ee 68

detrudens (Walker, 1865) (Pseudepicausta) . 2. 6. 6. ee ee 24

devia (Walker, 1861) (Antineura) .. .. 26.0 2. 24 ee ee es 7

didymoides Hendel, 1914 (Elaysogayter) . 2. 2. 2. 1. we ee 16

didymus (Osten-Sacken, 1881) (Elassogaster) .. 2. 6. eee 16

dimidiaia de Meijere, 1908 (Rivellia) . tel ats be ate aie 34

diptera Malloch, 1931 (Plagiastenopterina) i. ax wa le ge. aa 11

discalis Walker, 1859 (Brea) 2. 6. 2 4. ce ee ee ee ee ee 54

discifera Hendel, 1914 (Brea) . 6. 6. ce ce ce ee ee te ee 54

discifera de Meijere, 1913 (Euxestomvea) .. 2. 2. 2. ee ee 37

discolor Malloch, 1931 (Plagiostenopterina) . 6... 6. we ee IL

dispila Thomson, 1868 (?Zygaenula) (see note). .. .. 2. a. 58

distobasalis Hardy, 1959 (Rivellia) . oo. 6. ce ea ce ee ee ee 34

divergens (Walker, 1860) (Philicompus) . .. 2. 2. 2. 5. ee 6

diversifrony de Meijere, 1913 (Achias) .. 2. 6. ee ee ee ee 40

diversipennis Malloch, 1932 (Perissoneura) . 2. 2. 2. 2. ee sl

diviva (Walker, 1857) (Pterogenia) .. 1... 6. 2. ee ee ee 61

doclea (Walker, 1849) (Pogeonortalis) . .. ee se bs 32

doleschallt (Enderlein, 1912) (Lamprophthalma) . al ot 10

dorsata Hendel, 1914 (Euprosopia) . . ad core 68

dubiosa Malloch, 1931 (Plagiostenopterina) . vay ee tes 11

dubitalis Malloch, 1939 (Euprosopia) . 2. 2. 2. 1. ee ee 68

egregia de Meijere, 1924 (Plagiostenopterina) .. 6. 1. we ee 12

elongata v. d. Wulp, 1885 (Lanipregaster) .. 2. 6 ee ee ne 44

enderleini Hendel, 1914 (Plagiostenopterina) . 2. 2... 2... 12

LOWER—AUSTRALIAN REGION OTITIDAE

erasa Malloch, 1939 (Scotinosoma) . .

evitta Malloch, 1939 (Elassogaster) .

excepta Malloch, 1939 (Cleitamia) ~

exigeny (Walker, 1860) (Poewile picalusta)

experta (Walker, 1862) (Pseudepicausta) ,.

extranea Bezzi, 1917 (Xenaspis) .. 2. 1. ce ke ee ee we we

farinosa Hendel, 1914 (Plagiostenopterina) .. .. .. «+ +5 «+

fascipennis Hendel, 1914 (Loxonevra) .. 6. 2 ee ee ee

fascipennis de Meijere, 1915 (Laglaisia) . .. .. 1 +s

fascipennis Malloch, 1942 (Guamomyia) . 1. 6. we ue ae

fatuhivae Malloch, 1932 (Heterodoxa) .. .. .. 2. ee ee

femorata Hendel, 1914 (Scelostenopterina) .. .. .. -

ferruginea Hendel, 1914 (Rivellia) . .. 6. 6. ae eee

fidschiensis Enderlein, 1924 (Duomyia) 2. 6. 6. 6. ee

flaviceps Hendel, 1914 (Asyntona) , .. 2. ee ae pe ee ee ee

flavicornis Malloch, 1930 (Apactoneura) . 1. 2. 66 ee ee ae

flavicornis Frey, 1930 (Plerogenia) .. .. ap ot tte ated

flavipennis Macquart, 1843 (Lamprogaster) py ae! Btn “aps

flavipes (Schiner, 1868) (Elassoguster) .. 2. 6. 2. ca we we

flavipes de Meijere, 1913 (Brea) .. .. a

flavitarsis (Macquart, 1850) (Pseudorichardia) A

frauenfeldi (Schiner, 1868) (Scholastes) . 6. 2. 2. 16 ee ee

frontalis (Walker, 1861) (Euprosopia) .. .~

fuliginosa Hendel, 1914 (Plterogenia) .

fulvescens Malloch, 1931 (Lamprogaster)

fulvescens Malloch, 1940 (Rivellia) ..

fulviceps de Meijere, 1913 (Achias) .. 6. 64 we ce ee ee es

fulvipes Malloch, 1939 (Lampropaster): | Phe i ae

fulvofemoralis Malloch, 1942 (Pogonortalis)’ te ve 4

Jumicosta Malloch, 1930 (Neoeuxesta) .. 2...

furcatifascia Enderlein, 1924 (Schelastes)

furcatus Hendel, 1914 (Achiay) .. Bie

fusca (Thomson, 1868) (Rivellia) . ..

fuscibasis Malloch, 1930 (Lamprogaster) . 2. 2.02. ee ee a

fusifacies (Walker, 1859) (Euprosopia) .. 2. 6. 2. 2. 2s ee

geniculata (vy. d. Wulp), 1898 Sartidatiia ies ty a a ale

gestrai Kertész, 1899 (Cleitamia) . Ay Snatgw tated ten tas det

gigas Bezzi, 1917 (Euprosopia) . .. 64 62 26 we ve ae 2 ee

gjellerupi de Meijere, 1915 (Achiay) . 1. 1. 6. ee ee ee ae

glabra (Walker, 1857) (Pterogenia) . deitnens Be aie, jhe Vor

glabrella Hendel, 1914 (Pterogenia) .. ..

glabrina Hendel, 1914 (Pterogenia) ..

goniceps Hendel, 1914 (Dasyortalis) . 2. .. 1. 12 ee ee ee

gracilis Hendel, 1914 (Microepicausta) .. ..

grandis (Doleschall, 1858) (Antineura)

REC, S. AUST. MUS., 16 (2): 1-93 March, 1970

Page

erandis (Schiner, 1868) (Duomyia) .. 1. 22 ce se ae ae oe 20

grossa Malloch, 1939 (Lamprogaster) . et Peat 3 44

guamana Steyskal, 1952 (Neoeuxesta) . eee ae 81

guttata (No. 1) (Walker, 1857) (Prevoueria) -. ae rar 62

guttata (No, 2) (Walker, 1857) (Pterogenia) .. ~. 24 os 2. 62

guttipennis Kertész, 1899 (Loriomyia) .. .. -. «1 «e+ + 31

hamifera Frey, 1930 (Pterogenia) . ha Get Be sb rate on 63

hebes Hendel, 1914 (Pliptcstenopierina).-. Hieiathy steed “std as 12

hendeliana Bezzi, 1917 (Rivellia) . 2. 6. 6. ee ee ea ee ee 35

hilaris (Walker, 1849) (Lamprogaster) -. -. 5. 6 24 25 ee 44

hirsuta Hendel, 1914 (Lavsioxiria) . 6. 6. 6. ee ce ee ae ee 5

hirtiventris Malloch, 1942 (Scholastes) 2. .. 2. 22 ce ea ae 52

hivaeae Malloch, 1932 (Heterodoxa) . 2. 6. 2. ee ee ae oe 80

homichlodes Hendel, 1914 (Xenaspis) . 2. 2. 6. 4. we ee ee 8

howei Paramonov, 1957 (Pogonortalis) .. 2 6. 6. ce ee ae 32

hyalipennis Malloch, 1931 (Elassogaster) . 1. 2. 2. ee ea ee 16

hyalipennis Malloch, 1932 (Euxesta) . .. 2. 2. ce ee ce ae 77

hylaris Hendel, 1914 (Zygaenula) . 2. 6. ck ee ce ee ne we 58

ichneumonea Frey, 1930 (Xenaspoides) . 2. 12 ue ce we we 9

imitans (Walker, 1860) (Plagiostenopterinad) .~, .. .. ss «2. 12

imitans Malloch, 1930 (Rivellia) .. .. .. . 35

impingens (Walker, 1865) (Lepidocompsiay ( See Euprosopia) 68

inapta (Walker, 1860) (Plagiostenopterina) . .. . 12

indistincta Malloch, 1928 (Lamprogaster) .. .. -. se +e oe 45

inermis Malloch, 1930 (Xenognathus) .. 6. 6. ue ee ee oe 30

inermis Frey, 1930 (Peronotrochus) 2... 2. 22 24 ae ve ee 15

inermis Malloch, 1931 (Plagiostenopterina) . 2. 2. 2. we ee 12

innocua Malloch, 1939 (Euprosopia) . .. 6. 66 oe ee ee 69

insignis de Meijere, 1915 (Cleitamia) . .. 2. 2. 26 ee ee ae 29

instabalis (Walker, 1861) (Lamprogaster) .. 6. 2. 0. 15 ae 45

insulariy Paramonov, 1957 (Naupoda) .. .- 2. 2. ee ve ee 59

insulicola Malloch, 1940 (Euprosopia) .. 1. 6. 6c ce ae ae 69

interrupta Bezzi, 1928 (Pseudorichardia) . .. .. 2. 2. 1. ae 53

interrupta Malloch, 1930 (Zealandortalis) .. 2, 2. 24 ue ae 37

irregularis Malloch, 1929 (Duomyia) . .. 22 2. cu ae ce ae 20

isolata Malloch, 1930 (Rivellia) .. 2. 6. 6k ee ee ne ee 35

jucunda (Walker, 1849) (Lamprogaster) 2 2. 6. 1. ee ee ee 45

kerteszi de Meijere, 1906 (Antineura) .. 1. 22 ce ce ae oe 7

kerteszi (Hendel, 1914) (Cleitamoides) . 1. 6. 1. ne ce we 27

kochi de Meijere, 1907 (Laglaisia) .. 6. 6. ek ee ee ee 31

kurandana Hennig, 1950 (Achias) .. 2. 2. ce ca ne ce 40

LOWER—AUSTRALIAN REGION OTITIDAE

laeta (Guérin, 1830) (Lamprogaster) .

laeia (Walker, 1849) (Duomyia)

lafoont Steyskal, 1952 (Euxesta) .

latericia Hendel, 1914 (Pterogenia)

laticeps Bezzi, 1917 (Pterogenia)

latifascia (Walker, 1859) (Cleitamoides) aoa

latifasciata Malloch, 1931 (Mindanaia) ..

latiuscula (Walker, 1857) (Rhegmatosaga) .. 2. 2. 2. 2. ee

latiuscula (Loew, 1873) (Ceratoxys) . rhe os

lativentris (Walker, 1859) (Plagiostenopterina)

latividens Walker, 1859 (Achias) .. 2. 2. 0. ce ee ee we ee

lavata Hendel, 1914 (Rivellia) .

lavinia v. d. Wulp, 1898 (Xiria) as

lepida Walker, 1857 (Lamprogaster) .. ..

lepida Curran, 1936 (Euprosopia) . 2. 2. 1. ee ce ee ee

lepidophora Bezzi, 1917 (Euprosopia) .

leucomera (Walker, 1863) yeayartalisy

lifua Bigot, 1860 (Montrouziera) .

limbata de Meijere, 1924 (NXiria) .. ..

limpidipennis (Doleschall, 1858) (Pseudepicausta)

lineatus de Meijere, 1915 (Elassogaster) . sb gs

liturata (Walker, 1861) (Cleitamoides) ..

lonchifera Hendel, 1914 (Scholastes) .

longicornis Bezzi, 1917 (Euprosopia) .

longivideny Walker, 1859 (Achias) .

longivitta (Walker, 1859) (Plagiostenopterina) .

luteipennis Bezzi, 1917 (Pterogenia) . .3

macrocephala Hendel, 1914 (Lamiprogaster) .

macrocephalus Hendel, 1914 (Achias) .. ,.

macrotegularia Malloch, 1928 (Eupresepia) .

maculipennis (Guérin, 1831) (Euprosopia) .. .

maculipennis aromas 1847 (Lamprogaster) ..

maculipennis Hendel, 1914 (Duomyia) ..

magnifica Hendel, 1914 (Brea) .

marginalis Malloch, 1940 (Elassogaster). .

marginata (vy. d. Wulp, 1880) (Plagiostenopterina) Fe

marina Malloch, 1940 (Rivellia) ..

medionotata de Meijere, 1924 ( Plagiostenoplerina) .

metjereana (Enderlein, 1912) (Lamprophthalma)

metallica Malloch, 1931 (Euprosopia) . .. .. ..

metallica Enderlein, 1937 (Imugana) .

miliaria Hendel, 1914 (Euprosopia) .. ..

millepuncta Bezzi, 1917 (Euprosopia) . .. .. .. -.

minor Malloch, 1940 (Euprosopia) ws |.

minuta Malloch, 1939 (Euprosopia) .. .. ..

mithrax Hendel, 1914 (Duomyia) . .. .. 2. 2...

REC. S, AUST. MUS., 16 (2): 1-93

mohnikei (Doleschall, 1858) (Notopsila) . ..

monstruosum Bezzi, 1917 (Tylopterna) .. .. ~. -+ ++

multivitta (Walker, 1859) (Euprosopia) .

mutilloides (Walker, 1859) (Pseudepicausta)

neurostigma Bezzi, 1928 (Plagiostenopterina)

nigricosta Mallo

nigricostata (Doleschall, 1858) (Plagiostenopterina)

ch, 1929 (Duonryia) .

nigrifacies Malloch, 1939 (Achiosoma) .. -.

nigripes (Macquart, 1850) (Duomyia) .. .. 1. ee

nigripes Malloch, 1940 (Elassogaster) .. 6. 24 se oe

niveitarsis (Bigot, 1859) (Prerogenia) .

nobilis Frey, 1930 (Euprosopia) .. 6. 66 ve ue ce oe

ijére, 1913 (Brea) + we se wee bd eb Fes

1, 1914 (Pterogenia) .. 6. 2. ee we we ees

1928 (Pterogenia) . .. .. 2. 2+ se ee

nouhuysi de Me

nubecula Hende

nudiseta Bezzi,

obliqua (Walker, 1861) (Rivellia) 2 6. 66 ee ee we ee ee

obliqua Osten-Sacken, 1881 (Xiria) ©. 1. 6. ee ee

obliqua Frey, 1930 (Lamprogaster) 2. 6. 21.6 5

obscura Walker, eels ede eb gat oe

8; 805. (AGHIOS) 4 ean we ae 2

oculatus Fabrici

1849 (Duomyia) .

orthocephala Hendel, 1914 (Cleitamia)

ostensackeni Ke

pallidipes Frey,

palmyra Curran,

papuana Hennig,

rtész, 1898 (Cleitamia) ..

1930 (Plagiostenopterina) .. .-

1936 (Scholastes)

1940 (Xiria) . .. . Saks ab -Pe

paradoxa Doleschall, 1858 (Zypaerula)

parva Bezzi, 1917 (Pterogenia) . tte ee a ice ot: Sit OF

parva Malloch,

patula Walker,

1931 ( Plagiostenopterina)

1861 (Lamprogaster) .

pectoralis Hendel, 1914 (Pterogenia) . .. -. .- -« «

penicillata Hendel, 1914 (Euprosopia) .. .. .. «.

perilampoides Walker, 1858 (Loxonevra) .. .- «+

philpotti Harrison, 1959 (Zealandortalis) . .. «+

pictipennis (Wa

pipartita Hendel,

placida Walker,

plagiata (Bezzi,

planidorsum (Walker, 1860) (Plagidstenopterina). nem

platessa Osten-S

platychirus Heridel, 1914: CAchias): 4 ca ex crea p08:

Iker, 1849) (Xenaspis) ., .. ..

1849 (Lampregaster) .

1917) (Plagio stenopterina)

acken, 1882 (Naupoda) . .. «+ os

plumatum Hendel, 1914 (Euthyplatystoma) .

poecila Hendel,

1914 (Lamprogaster) .

1933 (Rivellia) .. 6. 66 1. ae ae ee

March, 1970

Page

LOWER—AUSTRALIAN REGION OTITIDAE

polistes Osten-Sacken, 1882 (Xenaspis) .

polita Hendel, 1932 (Rivellia) .

pompiliformis. Enderlein, 1937 (Imugana),

pompiloides (Walker, 1859) (Pseudepicausta)

potens (Walker, 1862) (Euprosopia) . ;

potens Frey, 1930 (Elassogaster) .

prima (Osten-Sacken, 1881) (Psendeuxesta) e

producta Walker, 1861 (Euprosopia)

prompta (Walker, 1859) (Euxestomoea) .

protensa (Walker, 1864) (Euprosopia) ..

pruinosa Malloch, 1932 (Euxesta) .

pseudelongata Malloch, 1930 (Laniprogaster)

pubiseta (Walker, 1861) (Antineura) . aM

pumicata vy. d. Wulp, 1885 (Lamprogaster)

punctata (Walker, 1857) (Prerogenia)

punctifacies Malloch, 1928 (Euprosopia) .

punctifrons Malloch, 1929 (Duomyia)

punctulatus de Meijere, 1913 (Achias)

quadrilinea Walker, 1859 (Lamprogaster)

quadrisetosa (de Meijere, 1911) (Pseudepicausta) ;

quadrivittata (Macquart, 1835) (Euxesta) .

radiata Hendel, 1914 (Rivellia) . 2. 2. 2. ee,

ralumensis Enderlein, 1924 (Brea)... ..

ralumensis Enderlein, 1924 (Mesoctenia) ,

regina Hendel, 1914 (Naupoda) .. ..

rigidum (Walker, 1857) (Euthyplatystoma) <

rivellioides Osten-Sacken, 1881 (Clettamia)

roderi Kertész, 1899 (Cleitamia) . f,

rothschildi Austen, 1910 (Achias) . ..

rufibasis Malloch, 1939 (Rivellia) .

ruficrus Hendel, 1914 (Pterogenia)

rufipes Hendel, 1914 (Lamprogaster) . .. 2...

rufiventrixs Hendel, 1914 (Euprosopia) .

rugifrons (Thomson, 1868) (Rhytidortalis) -

rutila Hendel, 1914 (Plagiostenopterina) .

samouensis Malloch, 1930 (Plagiostenopterina)

sauter? Hendel, 1914 (Rivellia) . 2. .

scatophaga Malloch, 1930 (Euprosopia)

scutellaris (Macquart, 1850) (Duomyia) .

scutellariy (Walker, 1859) (Pterogenia) .. .. ..

semiarmata Malloch, 1931 (Euprosopia) .

semifasciata Malloch, 1930 (Euxesta) .

separata Hendel, 1914 (Luwprosopia)

REC. S. AUST. MUS., 16 (2): 1-93

sepedonoides (Walker, 1864) Leipeprapnthalene) -

sepsoides (Walker, 1861) (Elasyogaster) . ..

sericata Osten-Sacken, 1882 (Antineura) -

sericea Hendel, 1914 (Duomyia) .. .

setigera Malloch, 1939 (Powiddoleitamiay ;

setinervis Malloch, 1939 (Euprosupia) .

severa Hendel, 1914 (Lamprogaster)

sexpunctata Osten-Sacken, 1882 (Fiuprosapiay: .

sexvittatus (Walker, 1861) (Scholastes) . ..

signatipes (Walker, 1861) (Elasyegaster) .

signifacies (Walker, 1861) (Dasyortalis) .

similis Kertész, 1899 (Cleitamia) .. .. ..

similis Hendel, 1914 (Pogonortalis) ..

Similis Hendel, 1914 (Rivellia) .

similis Malloch, 1939 (Prerogenic) .

simmondsi Bezzi, 1928 (Naupeda) , .. 4. 2 4 ni $2

simplex Frey, 1930 (Elassoegaster) .

singularis Bigot, 1859 (Pteregenia)

solocifemur Enderlein, 1924 (Pseudepicausta) {

sordidus (Walker, 1861) (Elassogaster) .

spinifemorata Malloch, 1929 (Duaymia) .

stenoparia Hendel, 1914 (Lamprogaster) . .,

Stigma Enderlein, 1912 (Xenaspis)

stolata Osten-Sacken, 1882 ( us chaeR

strigata Hennig, 1940 (Xiria) .

strigatus de Meijere, 1913 (Achias) ..

strigifery (Walker, 1862) (4aiineura’ }

strigifera de Meijere, 1919 (Naupoda) . .. 2. 1. 6. ea ee

stylops Enderlein, 1924. (Laglaisia)

subcruciata Frey, 1930 (Pterogenia)

subnudus Malloch, 1939 (Achias) .

suimbawana Hennig, 194] (Rivellia) ..

superbum (v. d. Wulp, 1881) (Euthyplatystoma)

superna Walker, 1862 (Lampragaster) ..

taeniata (Macquart, 1843) (Poecilotraphera)

taeniata y. d. Wulp, 1885 (Laniprogaster)

tarsalis (Walker, 1861) (Dasyortalis) . 2. ..

tarsalis (Walker, 1864) (Euprosopia) .

taylori Malloch, 1939 (Scholastes) . .. .. ..

tegularia Malloch, 1928 (Euprosopia) . .. ..

telescopica Enderlein, 1924 (Laglaisia)

tenuicorniy Macquart, 1847 (Euprosepia) ,

terrae-reginae Malloch, 1928 (Elaysoguster) .

tetyroides (Walker, 1859) (Asyntone)

thalassina Walker, 1849 (Duomyia) ..

theracalis Hendel, 1914 (Achias) .. ..

tigrina Osten-Sacken, 1881 (Euprosopiay. i

tomentosa Hendel, 1914 (Duonryia) .. ..

March, 1970

LOWER—AUSTRALIAN REGION

tricurvata (Walker, 1864) (Cleitamia) .

trifasciata (Doleschall, 1859) (Chaetorivelliay .

trifasciatus Enderlein, 1924 (Scholastes)

trigonalis de Meijere, 1913 (Cleitamia) .

trisignata v. d, Wulp, 1885 (Lamprogaster)

tristis Bezzi, 1917 (Pterogenia) .

trivittata (Walker, 1849) (Plagidstenopterinay:

trivittata Bezzi, 1917 (Euprosopia) .

trivittigera Malloch, 1931 (Plagiostenopterind). .

truncata (Enderlein, 1924) (Notopsila) ..

truncatula (Walker, 1857) (Pterogenia) .

tuberculifrons de Meijere, 1933 (Lamprophthaima) Aa

typicum Bezzi, 1917 (Scotinosoma)

uahukae Malloch, 1932 (Helterodoxa) . ..

uapouae Malloch, 1932 (Heterodoxa) .

uncinata de Meijere, 1911 (Pogonortalis) .

unifasciata Bezzi, 1917 (Naupoda) . .. .. ..

unimacula Hendel, 1914 (Lamprogaster) .

vacillans (Walker, 1860) (Rivellia)

valida Bezzi, 1917 (Pterogenia)

vartipennis Walker, 1861 (Pterogenia)

varipennis Hendel, 1914 (Loxoneuroides)

vella (Walker, 1849) (Lamprogaster) .

ventralis (Walker, 1859) (Euprosopia) ..

ventralis Curran, 1936 (Naupoda) .

venustulus Walker, 1865 (Achias) .

viola Malloch, 1929 (Lamprogaster) .. .. 2... 0.

violacea (Wiedemann, 1830) (Xiria) .

violacea (Macquart, 1843) (Lamprogaster) .

virgo Hendel, 1914 (Rivellia)

viridis Hendel, 1914 (Rivellia) . Si

vittata (Walker, 1857) (Pterogenia) .,

vitlifinis Walker, 1861 (Pterogenia) ..

WwW

walkeri (Enderlein, 1912) (Xenaspis) .

wallacei Hendel, 1914 (Pseudepicausta) ,

ypsilon vy. d. Wulp, 1898 (Naupoda) .

ypsilonoides de Meijere, 1924 (Naupoda)

zelotypa Hendel, 1914 (Lamiprogaster) ..

zonata (Walker, 1857) (Plerogenia) .

RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

THE RHODACARIDAE (ACARI :

MESOSTIGMATA); = CLASSIFICA-

TION, EXTERNAL MORPHOLOGY

AND DISTRIBUTION OF GENERA

By DAVID C. LEE

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 3

22 July, 1970

THE RHODACARIDAE (ACARI: MESOSTIGMATA); CLASSIFICATION,

EXTERNAL MORPHOLOGY AND DISTRIBUTION OF GENERA

BY DAVID C. LEE

Summary

The Rhodacaridae is redefined and considered to contain six subfamilies (Rhodacarinae,

Gamasiphinae, Laelaptonyssinae, Olgamasinae, Sessiluncinae and Tangaroellinae), of which one

(Ologamasinae) is divided into two tribes (Ologamasini and Gamasellini), and 45 genera, of which

six are divided into subgenera or species-complexes. Keys to the adults, diagnoses, morphology and

distribution data are given for all the above taxa. Separate descriptions, mainly in the form of

figures, are given for 63 of the 223 nominal species of Rhodacaridae. Three species are not placed

in genera but are considered as incertae sedis.

(LEE—RHODACARIDAE)

THE RHODACARIDAE (ACARI : MESOSTIGMATA); CLASSIFICA-

TION, EXTERNAL MORPHOLOGY AND DISTRIBUTION OF

GENERA

By DAVID C. LEE

CONTENTS

SYNOPSIS. cas eS ee pee aed aE ee ee ep

Introduction ,

Material and Methods gee pe ee rele lt mt

History of Classification . .. .. .. 6. ee ee ee

Morphology . Ae

1. Ventral Sclerotization ,

2. Leg Chaetotaxy .

3. Dorsal Sclerotization and “Chaetomorphy A

4, Spermathecal System .

Classification

Rhodacaridae .. . .

Key to Subfamilies a,

Rhodacarinae . . 4

Key to Genera ..

Gamasiphinae .. .. .. 2... we ee

Key to Genera ..

Laelaptonyssinae . .

Ologamasinae . . mn Ee

Key to Tribes and Genera ‘ui

Ologamasini . . ee: oe

Gamasellini . ..

Sessiluncinae .. on

Key to Genera ..

Tangaroellinae - 2 ress

Species Incertae Sedis . Aa

Supraspecific Taxa Sizes and Distribution ..

Changes in Nomenclature .. .. . ~~

Included Available Genus-Group Rares: :

Excluded Genera ..

Acknowledgments .. ..

References ..

208

210

211

212

22 July, 1970

REC, S. AUST. MUS. 16 (3): 1-219 July, 1970

SYNOPSIS

The Rhodacaridae is redefined and considered to contain six subfamilies

(Rhodacarinse, Gamasiphinae, Laelaptonyssinac, Ologamusinae, Sessilun-

cinae und Tangaroellinae), of which one (Ologamasinae ) ts divided into two

tribes (Ologamasini and Gamasellini), and 45 genera, of which six are

divided into subgenera or species-complexes. Keys lo the adults, diagnoses,

morphology and distribution data are given for all the above taxa. Separate

descriptions. mainly in the form of figures, are given for 63 of the 223

nominal species of Rhodacaridae. Three species are not placed in genera

but are considered as incertae seilis.

INTRODUCTION

The Rhodacaridae is a group of mainly free-living, predatory mites

occurring in ground habitats and, although the family as a whole is

cosmopolitan, the majority of its genera and species have only been found

in the Southern Hemisphere.

A number of concepts of the family have been put forward during the

last 15 years. The most similar to that delineated by the diagnosis given

here would either be that of the “Rhodacarus-group” genera proposed by

Evans (1963), being distinguished from another group of genera (including

Asea, Digamasellus and Halolaelaps) within the Rhodacaridae sensu Ryke,

1962b by their leg chaetotaxy, or a combination of the two families

Rhodacaridae and Cyrtolaelapidae as recognized by Johnston (1968).

Oudemans (1902a) established the Rhodacarinae with one, mono-

specific genus, Rhodacarus, while Halbert (1915) was the first to regard

the taxon as a family. Only a few genera were added to the family before

1955, but since then the number of genera referred to the family, as defined

below. has risen rapidly, initially by transferring established genera {rom

other families and more recently by establishing new genera within the family,

In the present study, as in the past, the taxonomic characters used to

classify categories within the Rhodacaridae are mostly external morphological

structures of the adults, The external morphology of the immature stages

has been used in gamasine classification, but for rhodacarids there is not

usually enough of this kind of data for it to be useful,

Soon after I started to study the Australian members of the Rboda-

caridue it became clear that this could not be done satisfactorily without a

more general, prior survey of the family, because the original descriptions

of the type-specics of many nominal rhodacarid genera and subgenera ure

meagre by present standards and only in some cases have they since been

improved, As a consequence, it is often difficult to decide how to upply

many of the available genus-group names; the placement of new or even

LEF—RHODAC ARIDAE J

long since established species within a genus is often uncertain. or the validity

of a new genus recognized to contain such a species is often dubious. Th an

attempt to overcome this difficulty, the type-species of nearly all nominal

genera or subgenera have been examined. As a result it is possible to make

a conservative decision as to which genus-group names can be regarded as

valid and to base the diagnosis of the taxa to which they are applicable on

the characters of the type-species.

Having based the definitions of rhodacarid genera and subgenera on

their types, [ considered it desirable to extend these definitions without

embarking upon a project that would too long delay my study of the

Australian rhodacarid fauna. Therefore, although care has been taken to

try and ensure that most nominal species are placed within the correct genus

and that any consequent adjustments to the generic diagnosis are made,

this survey is not sufficiently extensive to ensure the validity of all such

species. The conservative approach of trying to refer all nominal species

to previously established genera has usually been possible, sometimes by

recognizing species-complexes within a genus and sometimes by considering

Species as incertae sedis, but some new genera have been established. Also,

from amongst the many unnamed species scen only a few are established as

new species) usually, either so that their descriptions. will illustrate the range

of characters within a genus, or in order to rename rhodacarid mites that

have been misidentified in the literature. But, in order to extend the general

usefulness of this work, | have referred several of these unnamed species to

genera so that their locality data can be used to give a more corniplete

indication of the geographical occurrence ot the supraspecific taxa,

As a result 45 genera are recognized as valid within the Rhodacaridae

und are placed in a framework of suprageneric taxa in order to facilitate the

further study of the family. The genera are grouped in six subfumilics

and the genera of one subfamily, the Ologamasinaec, are further grouped

in two tribes. Although the delimitation of the subfamilies should not be

considered final and some genera are only provisionally referred (to them, |

have confidence in the viability of the basic concepts of these groupings und

would regard them as an indication of the phylogeny of rhodacarid mites.

In contrast, even if the tribal names used within the Ologaumasinae contmue

in use in the fulure, the present concepts of these two taxa ure used here

largely for convenience, particularly as they reflect a previous, widely held,

concept for which a reliable alternative has not yel been found.

MATERIAL AND METHODS

Some mites have been mounted in lactic acid, before being drawn with

the aid of a camera lucida, while others were mounted in a gum chloral

medium and may have been more squashed when drawn,

4 REC. S. AUST. MUS., 16 (3); 1-219 July, 1970

The material described or examined is deposited as listed after the

locality data of the species, using the following abbreviations. BBM:

Bernice P. Bishop Museum, Honolulu, Hawaii. BM(NH): British Museum

(Natural History). London, England. LAEHE: Laboratoire d’Acarologie,

Ecole Practique des Hautes Etudes, Paris, France. NRS: Naturhistoriska

Riksmuseum, Stockholm, Sweden. PUSA: Potchefstroom University,

South Africa. SAM: South Australian Museum. SEAF: Stazione di

Entomologia Agraria, Florence, Italy. ZMH: Zoologisches Museum,

Hamburg, Germany.

Generally, the terminology follows that used by Evans and Till (1965),

but using a change in nomenclature for tarsal chaetotaxy (Evans, in press)

which regards this segment as carrying four instead of three whorls of setae

(see Fig. | for abbreviations used). The names for parts of the female

spermathecal system are Anglicized terms derived from those used by

Michael (1892). Under the heading “Diagnosis” the following terms are

used for categories of adult mites having an idiosomal length within the

particular ranges given; “minute” for less than 400y, “small” for 400, to

600., “average sized” for 601 to 800., “large” for 801 to 1,200 and

“gigantic” for more than 1,2002, Under the heading “Sclerotization” the

word “shield” is omitted, as is “seta” under “Chaetotaxy”. A map (Fig. 427)

of Zoogeographical regions of the world is given as a key to abbreviations

used for the distribution of the members of each genus.

TARSUS TTT

-—pe—-xOwv

rHPri4HM-—oO

POSTERIOR

O A oO = d= dorsal seta a= anterior

=) A fad = |= lateral seta Pp = posterior

e& A || = v= ventral seta

Seta first appears in: Larva, Protonymph, Deutonymph

Fig, |. Setal nomenclature for tarsus TEL.

LEE—RHODACARIDAE 5

Previously (Lee, 1966), I used the term “species-group” for a supra-

specific category within a genus, but in the text below the alternative term

“species-complex” is used in order to avoid confusion with the use of

“species-group” for a species and infraspecific category in the Internation

Code of Zoological Nomenclature, 1961. Particular complexes of species,

named after characteristic species, as for example in falciger-complex, follow

this system.

HISTORY OF CLASSIFICATION

Oudemans (1902a) exaggerated the importance of some characters of

Rhodacarus roseus when he established the species in a new subfamily—the

Rhodacarinae—within the Parasitidae, a family then equivalent to the present

Mesostigmata. As a result in 1923 he incorrectly allied the Rhodacaridae

(still containing only one genus) by placing it in the Sejina and not the

Gamasina, and also maintained an inadequately narrow concept of the

family when he (Oudemans, 1939a) transferred Rhodacarellus to the

Gamasolaelaptidae leaving only Rhodacarus and Rhodacaropsis as rhodacarid

genera.

Berlese (1913b) used a systematic framework in which the arrangement

of the genera, considered here as belonging to the Rhodacaridae, fits the

concept of the family used here much better than that of Oudemans. Berlese

referred nearly all the then nominal rhodacarid genera (exceptions being

Heydeniella, then considered a junior synonym of Gamasiphis, and

Stylochirus) to either his tribe Cyrtolaelaptini or to the tribe Gamasini.

Although neither of these family-group names are regarded as available here,

Cyrtolaelaptini can be regarded as equivalent to the Veigaiidae, while

Gamasini can be regarded as equivalent to the Parasitidae. Gamasellus,

Heterogamasus, Protolaelaps (junior objective synonym of Cyrtolaelaps),

Rhodacarus and Sessiluncus were grouped with what are now considered to

be veigaiid genera and Euryparasitus, Gamasiphis, Hydrogamasus, Laelo-

gamasus and Ologamasus were grouped with what are now considered to be

parasitid genera.

Vitzthum (1941) maintained nearly as narrow a concept of the family

as that of Oudemans, but included Rhodacarellus, and was followed in

this by Baker and Wharton (1952). In the latter publication, genera

that are here grouped in the Rhodacaridae were referred to five other

families (Ascaidae, Gamasolaelaptidae, Macrochelidae, Neoparasitidae and

Pseudoparasitidae ).

The concept of the Rhodacaridae was broadened by Evans (1955 and

1957) in his classification of the British Mesostigmata, when he included in

the family those members of the Gamasina with a three-pronged apotele

(without an associated hyaline flap) and a divided dorsal shield. By

6 REC. 8, AUST. MUS., 16 (3): 1-219 July, 1970

including Sessiluncus (entire dorsal shield) and Digamasellus (two-pronged

apotele) in the Rhodacaridae, Ryke (1958) implied a further extension of

the limits of the family. When he later redefined the family (Ryke, 1962b),

he recognized 37 genera or subgenera within it, of which 11 are not regarded

here as belonging to the Rhodacaridae. His broad definition appears to only

exclude, with certainty, those members of the Gamasina without a divided

dorsal shield in the deutonymph and a ventro-anal shield in the female. In

the latter publication, Ryke recognized two subfamilies within the Rhoda-

caridae; the Rhodacarinae with a divided dorsal shield in the adult and the

Ologamasinae with an entire dorsal shield in the adult. A grouping of

genera within the Rhodacaridae similar to that for the Rhodacaridae sensu

Evans, 1957, was used by Karg (1961), while Athias-Henriot (1961b)

apparently used a grouping similar to that for the Rhodacarinae sensu

Ryke, 1962b.

During the last decade, two main schools of thought have developed on

the classification of the Mesostigmata, originating either from the work of

Hirschmann (1957 and 1959) or that of Evans (1963). Hirschmann (1957

and 1959) based his “Gangsystematic” studies on morphological characters

which are preferred if they are similarly expressed at all stages of their

ontogenetic development, but which must be distinctive throughout this

development so that any postembryonic instar of a species can be correctly

placed in a supraspecific category. The characters Hirschmann used are

the nature of the dorsal sclerotization, the chaetotaxy of the idiosoma, and

the structure of the appendages and ventral surface of the gnathosoma.

Evans (1963) introduced a new diagnostic character, the leg chaetotaxy,

which is not similarly expressed during ontogenetic development and is

usually only distinctive for a taxon in the later-stages of development

(7.e., in the deutonymph and adult). Although Evans rightly emphasized

that the leg chaetotaxy must not be used in isolation from other characters,

it has proved of considerable importance because of its usefulness. Since

I follow Evan’s usage of morphology, I will only consider the further history

of the classification of the Rhodacaridae as it has been affected by acarologists

using leg chaetotaxy as an important taxonomic character. After which J

will indicate the approximately equivalent groupings of genera by other

acarologists.

Evans (1963), after pointing out that his analysis of the leg chaetotaxy

of free-living Gamasina supported familial concepts based on other morpho-

logical criteria, noted that the Rhodacaridae (sensu Ryke, 1962b) was an

exception in containing two groups of genera, one of which he called the

“Rhodacarus-group”, which could be accommodated in the Rhodacaridae,

while the other group of genera, containing Asca, Digamasellus and

Halolaelaps, did not appear to be confamilial with the first group. This

gave rise to the concept of a “Rhodacaridae sens. lat.” equivalent to the

LEE—RHODACARIDAE 7

Rhodacaridae sensi Ryke, 1962b, and a “Rhodacaridae sens, str” equivalent

to the “Rhodacarus-group” genera of Evans (1963),

The above concept of “Rhodacaridae sens. lat” with uncertain limits

was retained longer than necessary. When genera not belonging to the

“Rhodacarus-group” had been transferred either to the Ascidae or Diga-

masellidue (Lindquist and Evans, 1965) or to the Halolaclapidae (Karg,

1965), the Rhodacuridue could be defined on the basis of other morphological

criteria as well as the leg chaetotaxy, The unnecessay reference to the

“Rhodacarus-group" genera as if they Were a category within an undefinable

Rhodacaridae sens, lat, can be seen in my own work (Lee 1966 and 1967).

I (Lee, 1966) did not accept the division of genera into two sub-

families within the Rhodacaridae as established by Ryke (1962b). Instead,

as a temporary measure recognized as unsatisfactory at the time, T used two

unnamed groups of genera, the members of which were distinguished by the

structure of the dorsal setae.

Johnston (1968) transferred all those genera containing species with

seta pl4 present on tarsus TV from the Rhodacaridae to u new family, the

Cyrtolaelapidae, The resulting narrow concept of the Rhodacaridae is

equivalent to that for the Rhoducarinae as defined below,

Within Hirschmann’s (1962) classification. the reference of genera to

higher taxa indicates that his concept of the Gamasellini: within the

Bugamasinae closely approximates to thut used here for the Rhodacandae,

The other tribe in his Eugarmasinae, the Eugamasini, contains genera that

were referred lo the Parasitidae and Veigatidue by Vitzthum (1941), a

reference accepted by most acarologists since then. On the other hand Karg

(1965) places most rhodacarid genera, genera that Johnston (1968) has

grouped in the Cyrtoluelapidac, in the Gamasellinae, a subfamily within the

Euvamasidae, while Rhedacarus, Rhodacarellus, Dendrolaelaps (syn. Diga-

masellus), and Protogamasellus are grouped in the Rhodacaridae. The other

subfamilies in Kary’s Bugamasidae are the Parasitinue and Veigaiinae.

Bregetova (1967), on the basis of the ontogenesis of the dorsal sclerotization,

groups m the Rhodacaridae approximately those genera that were grouped

by Ryke (1962b) in the Rhodacarinae and. presumably. genera from Ryke's

Ologamasinae would be referred to the Parasitidue,

MORPHOLOGY

A comparative study of all the morphological characters used in the

classification given below has not been made. But a study has been made of

the following four sets of characters that are important in the diagnoses

of supragenerte rhodacurid taxa, The ventral sclerotization of the adult

idiosoma which is used in diagnosing subfamilies, as is the deutonymph and

aduit lez chactotuxy., The dorsal sclerotization of the adult idiosoma and the

8 REC. S. AUST. MUS., [fh (3): 1-219 duly, 1970

form of the dorsal setae which are used in diagnosing the two tribes in the

Ologamasinae because of precedence and convenience, and the location of

the external aperture of the spermathecal system which may in the future

be more important in the diagnoses of these two tribes.

1, VENTRAL SCLEROTIZATION

In the past, the ventral sclerotization has been used to distinguish

genera, for example the fusion of the ventro-anal shield with the opisthonotal

shicld has been considered diagnostic for Gamasiphis and Ologamasus, but

I use it here for diagnosing suprageneric taxa. The following two characters

of the ventral sclerotization are considered of value; the fusion of the ventro-

anal shield with other shields and the fusion of the posterior end of the

peritrematal shield with other shields. Two monogeneric subfamilies,

Laelaptonyssinae and Tangaroellinae, are not considered in this section,

because, although their ventral sclerotization is somewhat atypical, alternative

characters are more useful in distinguishing them from other rhodacarid

subfamilies,

The four major groups of genera which | treat as subfamilies are largely

characterized by the fusion or lack of fusion of the ventro-anal shield to

other shields and, ignoring the exceptions, they can be distinguished as

follows; Rhodacarinae, female ventro-anal shield discrete, male ventro-anal

shield fused to opisthonotal shield but not to peritrematal shield:

Gamastphinae, female and male ventro-anal shield fused to opisthonotal

shield but not to peritrematal shield; Ologamasinae, female ventro-anal

shield discrete, male ventro-anal shield fused to opisthonotal and peritrematal

shield, Sessiluncinae, female and male ventro-anal shield not fused to

opisthonotal or peritrematal shield,

These differences are illustrated in Figure 2, and the importance of

Knowing both sexes of a species in the use of this character for diagnosis

should be noted. However, even if only one sex of a species 1s known,

members of the Rhodacarinae can be distinguished from other rhodacaridl

species by the chaetotaxy of tarsus PV and females of the Ologamasinae can

be distinguished trom females of the Sessiluncinue by the fusion of the

peritrematal shield to other shields. As a result, diagnoses of the subfamilies

would be relatively easy if there were no exceptions to the characteristic

fusion of the yentro-anal shield to other shields, but this is not so.

Within the subfamilies Rhodacarinae, Gamasiphinae and Sessiluncinae.

the extent of the fusion of the ventro-anal shield to the opisthonotal and

peritrematal shields shows only a few, simple exceptions to the above

characteristic forms, thus: some males of the Rhoducarinae have a discrete

ventro-anal shield; some species of Gamasiphoides within the Gamasiphinae

have females and males with a diserete ventro-anal shield; all species of

LEE—RHODACARIDAE 9

m>Z—DPOFUOCLIA

mPpZ—-LVU—MNPZPO

L bs

A Pong ot oO

E \ L

L 4

. G

F A

O AN M

N A

Y i

; f «§

N 7 E

mPZ—OZCrH—MOMMNW

mP>2Z—rrmOowrPonz>yHa

J ff

Do” PA

Fig. 2. Ventral sclerotization of the adults of the lype-species of the type-genera of

rhodacarid subfamilies, excepting Laelaptonyssinae for which Laclaptonyssus chinensis

is illustrated and Ologamasinae for which the type of Cyrtolaclaps is illustrated.

1) REC, 8. AUST, MUS., I@ (3); 1-219 Jub, (970

Stylochirus within the Sessiluncinae have females and males with the ventro-

anal shield fused to the notal shield. On the other hand. within the

Ologamasinae there dre many exceptions to the characteristic fusion of the

ventro-anal shield with other shields, and in some lightly sclerotized (e.g..

Acugamasus cursor) or heavily sclerotized (e.g.. Hiniphis hinnus) species

this character is not sexually dimorphic,

Two general statements without exceptions, to which | attach consider-

ble importance, can be made about the fusion of the ventro-anal shield to

other shields. Only members of the Rhodacarinae and Ologamasinae ever

have sexual dimorphism: of the fusion of the ventro-anal shield to the

opisthonotal shield, and only metnbers of the Ologamasinae ever have the

ventro-anal shield fused to the peritrematal shield and when this is the case

it usually only occurs on the male,

Members of the Sessiluncinae can usually be further diagnosed by the

peritrematal shield not being fused to exopodal shield [V, bur species of

Gamasellopsis, Onchogamasus communis and Queenslandolaelaps vitethumi

ure exceptions. It is possible that the fusion in these latter cases is different

in origin from the apparently similar fusion in most members of the other

three large subfamilies. A visible difference in such a fusion on members

of the Sessiluncinae is that there is no fissure between the peritrematal and

exopodal shields posterior to the stigma. This is true also for a few members

of the other subfamilies (some Gamasiphis species and Hiniphis hinnus),

but these particular rhodacarid mites are usually heavily sclerotized and 1

consider them to have relatively recent ancestors in which the above fissures

occurred behind the stigma.

2, LEG CHAETOTAXY

The first analysis of the leg chuetotaxy of the free-living Gamasina, as

well as a system of setal nomenclature for individual leg segments, was

introduced by Evans (1963) who considered the setue as being in patterns

based on the numbers distributed amongst six zones on the surface of a leg

segment. According to which zone they are in, the setae are named cither

antero-lateral, antero-dorsal, postero-dorsal, antero-ventral, postero-ventral

or postero-lateral as indicated in the accompanying diagram (Fig. 1). For

inost leg segments, the setal pattern or chaetotaxy varies in the deutonymphs

and adults of the Gamusina, Evans (1963) showed that this variation

provides a valuable taxonomic criterion for the classification of these mites

and he listed the types of chaetotaxy that he had observed on each leg

segment excepting tarsus 1, which, because of its large number of setae, has

not yet been described for any gamasine miles. Although Evans refers to

“types of chactotaxy” and Costa (1968) uses the term “chaetotiuctic variants”,

here T refer to “kinds of chaetotaxy”,

LER —RHODACARIDAE ‘|

| have confirmed that for the majority of species the leg chaetotaxy Is

the same as that described by Evans (1963) for the “Rhodacarus-group”

venera. This kind of leg chuetotaxy has been illustrated in a previous paper

(Lee, 1966, Fig. 1) for Gamasellus diseutatus and ts termed here the

“commonest kind of rhodacarid leg chaetotaxy” or as being “as Gamasellus”

or “us Gamasellus discutatus”.

While Evans (1963) mainly demonstrated that the chactotaxy of legs

[and tare valuable in the diagnosis of gamasine families, later publications

show that leg chuetotaxy, especially of legs HT and TV, 1s a valuable diagnostic

character for genera; for example in the Ascidae (Lindquist and Evans,

1965) and Dermanyssidae (Bvans and Till, 1965). Leg chactotaxy has

also proved useful in the diagnosis of rhodacarid taxa. There are a fair

number of rhodacarid species with a different kind of leg chactotaxy to the

commonest One, und amongst them L have observed 18 kinds of leg

chactoluxy, if a kind of leg chaetotaxy is recognized by differences as small}

as the presence or absence of one seta from amongst all the leg setae.

To give a veneral indication of the variability of rhodacarid leg

chuetolaxy, | have made u visual presentation of the data (Fig. 3) that

provides a simple measure of the variability of particular setae in different

subfamilies and tribes. Labels are attached to all the setae (Fig. 3) that

sometimes do not occur (present or ubsent) as in the commonest kind of

lev chaetotaxy,

The symbols used on the diagram (Fig. 3) have the following meunings,

A “--ve'! indicates that the seta is sometimes present, but is absent in the

commonest leg chaetotaxy, and “—ve" indicates that the scta is sometimes

absent but is present in the commonest leg chaetotaxy, The letters indicate

the subfamilies or tribes in which the particular seta’s occurrence differs from

that in the commonest leg chactotaxy. The figures associated with these

letters indicate the number of kinds of leg chactotaxy, found in these taxa,

which inelude an uncommon occurrence of the particular seta, and this gives

a measure of the seta’s variability. The actual kinds of total leg chaetotaxy

are viven under the different subfamily headings later in this piper.

An example of the use of Figure 3 is given as follows. Ifa rhodacarid

species is examined and found to lack seta pv on genu TIT, Figure 3 is

referred to und this seta is seen to be labelled * 4G:38:1G.S". This means

that the seta is present on most nominul rhodacarid species but absent in at

least some members of both the Gamasiphinae and Sessiluncinae and it is

quile often absent since it is missing in & of the 18 kinds of leg chaetotaxy

differing trom the commonest rhoducarid kind, Also, since 4 of these

kinds of chaetotaxy occur on some members of the Gamasiphinae, 3 on

some inembers of the Sessiluncinae and | on some members of both the

Ganasiphinue und Sessiluncinae, the seta tends to equally variible in both

subfamilies.

12 REC, S. AUST. MUS., 16 (3): 1-219 July, 1970

Leg L

Leg IL

Leg OIL

Leg IV

[proximal part |

7 1

4G:101 3G:3S

Tarsus [TV

7 -2R*

1G:2S:1T iT

R = RHODACARINAE

G = GAMASIPHINAE S = SESSILUNCINAE

Ol = OLOGAMASINI T = TANGAROELLINAE

OLOGAMASINAE

Gl = GAMASELLINI

LAELAPTONYSSINAE OMITTED

Fig. 3. Setation of adult rhodacarid appendages. Unlabelled setae and labelled setae

accompanied by a “~ ve” sign indicate chaetotaxy of Gamasellus discutatus, Labelled

setae accompanied by a “+ve” sign occur in certain rhodacarid taxa but not in

G, discutatus. The labels to setae are explained in the text (p. 11). The symbols

used for setal position and first appearance are as in Fig. 1.

LEE~-RHODACARIDAE 13

Costa (1968) compared the number of kinds of chaetotaxy for different

leg segments on free-living Gamasina (using the data of Evans, 1963) and

on 4 gamasine parasite (Hemipreroseius adleri) which has imtraspecitic

variation of leg chaetotaxy, and showed that in both cases there ts a similar

eradient of variability depending on the location of the leg segments. The

setal patterns are more stable on the segments of legs | and I than on legs

If and IV, and on the proximal leg segments and tarsi than on the genua

und tibiae, The numbers of different kinds of chactotaxy for particular

udult rhodacarid leg segments usually support Costa’s observation if the

aberrant leg chaetotaxy of Laelaptonyssus is ignored, and there also appears

to be a gradient around the surface of the leg segments. For example.

although there are 34 anterolateral, 69 dorsal, 30 postero-lateral and 62

ventral setae on the adult Gamasellus legs (excluding tarsus 1) of one side,

the variability of setae from these zones, as measured by the addition ot the

number of kinds of leg chaetotaxy in which a particular seta does not occur

as on Gamasellus, is 1 for the antero-lateral, 9 for the dorsal, 17 for the

postero-lateral und 23 for the ventral setae,

The point, emphasized by Lindquist and Evans (1965), that leg

chaetotaxy is not a taxonomic panacea, but is another set of characters which,

when regarded alone, may have exceptions among related species, is well

illustrated in the Rhodacaridae. For example. the setal pattern on tarsus 1'V

has been regarded as stable amongst free-living Gamasina, but vartations

in the occurrence of setae on the tarsi IV of rhodacarids can be used in the

diagnosis of the Rhodacarinae (45 nominal species), Tangaroellinae (1

nominal species), Gamasellopsis (4 nominal species), Gamasiphoides

propingua and Laélaptonyssus mitis,

3. DORSAL SCLEROTIZATION AND CHAETOMORPHY

Amonest members of the Rhodacaridae, the attributes of three

characters are often associated as follows: ‘some complex dorsal setue—

divided dorsal shield—dull sclerotization’, or ‘all simple dorsal setae—entire

dorsal shield—shiny sclerotization’. Two groups of mites, distinguished by

this association of attributes, were placed in either Garnayellus or Hydro-

gamasus by carlier acarologists, or the subgenera Gamasellus or Ilydro-

vamasellus by Hirschmann (1966). From amongst these three characters,

the presence or absence of 4 division of the dorsal shield was used by Ryke

(1962b) in dividing the family into the Rhodacarinae and Ologamasinue,

and latter the dorsal chaetomorphy was used by Lee (1966) in a temporary

division of the family into two unnamed groups of genera,

A comparison between species of Evepicrius and characteristic species

of Gamasellus and Gamasiphis indicates that, if the classification given below

is followed, the above associations of attributes should be regarded as

14 REC. S. AUST. MUS., lo (3): 1-219 July, 1970

relatively superficial within this family. Despite this, I recognize the two

groups of genera within the Ologamasinae, referred to as the tribes

Ologamasini and Gamasellini, by such attributes. The character given the

most importance here is the dorsal sclerotization as used by Ryke (1962b)

in his division of the whole family: most adults of the Ologamasini have an

entire dorsal shield, while on adults of the Gamasellini the dorsal shield is

usually divided. Species in the Ologamasinae which do not have the dorsal

sclerotization characteristic of their tribe, are members of genera

(Geogamasus, Heydeniella, Hydrogamasellus and Gamasellus discutatus-

complex) in which this character varies between species or sexes of the

same species and occasionally males of two of these genera (Geogamasus

and Hydrogamasellus) have a partially divided dorsal shield, a rare attribute

in this family. Of these genera with varying adult dorsal sclerotization,

Geogamasus, Heydeniella and Hydrogamasellus have simple dorsal setae and

are placed in the Ologamasini, while the Gamasellus discutatus-complex

species have some pilose dorsal setae and are placed in the Gamasellini.

The concepts used here for the two Ologamasinae tribes are mainly to provide

a convenient division of this large subfamily that follows some precedence.

But it is interesting that the resulting grouping of genera is similar to one

based on the location of the external spermathecal aperture (see below).

Therefore, although the present tribal taxa are tentative, a similar, stable

grouping may be made later which is an indication of the phylogeny of the

included mites.

4. SPERMATHECAL SYSTEM

In the Gamasina, male gametes are transferred to the female in a

spermatophore, and Michael (1892) considered it likely that these gametes

reached the spermatheca and ova through the vagina in some species, and

through special tubes, the “tubuli annulati” (here termed “spermathecal

ringed tubes”) with an external aperture distinct from that of the vagina, in

other species. Michael’s evidence included the presence of a_ possible

spermathecal sacculus and ringed tubes in the female only, the connection

of these structures with the ovaries, the similarity of their contents to the

contents of the spermatophore and observations of mites copulating.

Although evidence supporting Michael’s (1892) work on gamasine

reproduction is meagre, Camin’s (1953) description of vaginal insemination

for Ophionyssus natricis and Dosse’s (1958 and 1959) description of direct

spermathecal insemination for Typhlodromus zwolferi, as well as some more

recent work by others, establish that the two kinds of insemination suggested

by Michael do occur in different members of the Gamasina. Costa (1966)

implies that the members of a particular family would all have the same kind

of insemination, but this may not be true for this family or for the included

LEE—RHODACARIDAF |

Ologamasinue. Spermathecal ringed tubes are present in many species

grouped in these taxa, but Michael (1892) presents evidence that the

insenination of a member of the Ologamasinae, Euryparasitus emuarginats,

is through the vagina and | have been unable to find spermathecal ringed

tubes in Acitzamasus females despite my examination of many carefully

cleared specimens,

By describing the spermathecal ringed tube and sacculus in a large

number of gamasine mites, Athias-Henriot (in press) demonstrates the

variety of their fony and location of access aperture, especially amongst

rhodacarids, One, two or, ina single instance, three locations of the sperma-

theeal ringed tube aperture are described on the females of unnamed species

that Athias-Henriol considers us belonging to the same rhodacarid genus.

From my own work there usually appears, for any particular rhodacarid

genus. Lo be only one locity for the external aperture of the spermatliecal

ringed tube, Therefore, | suggest that Athias-Henriot (in press) allots some

Southern Hemisphere rhodacarids to the wrong genera. I[f the assumption

that there is one locality for the external aperture of the spermatheeal ringed

lube in most rhodacarid ygeneru is substantiated in the future, then the

considerable variety of this character will make it an injportant taxonomic

criterion.

Because of the possibility of confusing a structure (e¢.g., duets of coxal

and femoral glands, described by Pain, 1966, or even the gemital apodeme

and muscles. described by Treat. 1965) with the spermathecal ringed (ube

it is necessary to have some indication of the reliability of such an identifica-

tion. When there is 4 connection between the tube in question and a central

sacculus, | regard it as certain that this tube is a spermathecal ringed tube.

On this basis a spermathecal ringed tube opens near the posterior paraxial

edge of acetabulum TV just ventral to the genital apodeme in the following

genera: Gamasiphis, Caliphis, Euepicrius, Gamaselliphis, Gamasiphaides,

Laelaptiella, Parasitiphis, Gaumasellus falciger-complex, Perisetus (Psam-

monsella), Attennolaelaps and Queenslandolaelaps. Observations (to be

published later) of copulation in Evepicrius filamentosus and Gamasellus

travardhi endorse this, and similar observations for Meydenfella dentata

endorse the identification of a tube opening near the dorsal distal margin of

trochanter MH as a spermathecal ringed tube. In other cases | am, for a

number of reasons, fairly sure that I have correctly identified the sperma-

thecal ringed tube and in the text below the “probable” location of its

external aperture is referred to. ‘This is true for the following genera with

the ringed tube probably opening near the posterior paraxial edge of

acetabulum IV: Rhodacarellus, Hydrogamasus, Gamuasellus discutaius-

complex, Periseius (Periveius), Pilellus and Paragamasellevans. This 1s

ulso true for some species of Afrogamusellus where the opening is probably

on the metapodal shield and for the following genera in which the ringed

It REC. S. AUST. MUS., 1f (3): 1-219 July, 1970

Hydrogamasellus

and Laelogamasus

Geogamasus

: "~~ Heydeniella

and Pyriphis

some

Afrogamasellus

Ologamasus

many

rhodacarid genera Rykellus

Fie. 4. Probable location of access to spermatheca on rhodarid females,

tube probably opens on the proximal segments of legs [II] and IV:

Ologamasus, Cymiphis, Geogamasus, Hydrogamasellus, Rykellus, Pyriphis

and Laelogamasus. On female Cyrtolaelaps and Euryparasitus there is a

tube opening on the sternal shield between seta sf3 and acetabulum III,

other than the sternal pores which also occur on the male. But, because of

Michael’s (1892) observation that the spermatophore is applied to the

vagina of Euryparasitus emarginatus (as Gamasus terriblis) and spermatozoa

are found in a domed recess in the vagina, [ regard the identification of the

above sternal tubes of Cyrtolaelaps and Euryparasitus as spermathecal ringed

tubes as dubious. Also, on one of the three known females of Neogama-

sellevans berlesei (described under species incertae sedis), a tube apparently

opening near the genital apodeme has been blown partly out of the idiosoma

by the clearing process and may be a spermathecal ringed tube. Therefore,

there are 23 rhodacarid genera, subgenera or species-complexes of which I

have not seen members or in which I have been unable to locate a possible

spermathecal ringed tube. Amongst these latter genera, Athias-Henriot

(in press) has located a possible spermathecal ringed tube opening on femur

IIT of a Rhedacarus species and on coxa II] and coxa TV of two Sessiluncus

species,

LEE—RHODACARIDAE \7

On the assumption that only one location for the aperture of the

spermathecal ringed tube usually occurs in a particular genus, [ summarize

my observations in Figure 4. All genera with the spermathecal ringed tube

probably opening on the proximal segments of legs IT and IV are listed on

the right hand side of the diagram and, excepting Laelogamasus, they all

belong to the Ologamasini, Therefore, it is possible that the diagnosis of

the Ologamasini may later be based on the location of the aperture of the

spermathecal ringed tube.

CLASSIFICATION

Family RHODACARIDAE Oudemans

Rhodacarinae Oudemans, 1902a, p. 48.

Rhodacaridae Oudemans, Halbert, 1915, p. 81.

Cyrtolaclaptini Berlese, L913b, p, 11,

(This name is considered to be based on a misidentified type-genus

and therefore is not available by the terms of the International Code of

Zoological Nomenclature (1961), The evidence put forward for mis-

identification is that later (p. 86), in the paper where this family-group

name was first published, Cyrrolaelaps is redefined with “species typica

== Gamasus nemorensis K", indicating that Berlese had overlooked the

type-designation by monotypy (Berlese, 1887b) of Gamasus mucro-

natus G. and R. Canestrini for Cyrtolaelaps and the type-designation

by Oudemans (1905) of Gamasus nemorensis Koch for Veigaia.)

Cyrtolaelaptinae Berlese, 1920, p, 166.

221.

Rhodacaridae Oudemans, sensu Evans (in part), 1957, 2

155.

Rhodacaridue Oudemans, sensu Ryke (in part). 1962b, p.

Gamasellini Hirschmann, 1962, p. 39.

Rhoedacarus-group Evans, 1963, p. 302.

Rhodacaridae Oudemans, sensu Karg (in part), 1965, p, 295,

Rhodacaridae Oudemans, sensu Bregetova (in part), 1967, p. 472.

Type-genus: Rhodacafus Oudemans, 1902a.

DIAGNOSIS. — Sclerotization very variable, but always separate

podonotal and opisthonotal shields on deutonymph and a_ posteriorly

truncated female genital shicld separated from a conspicuous ventro-anal

shield. Female metasternal seta, »/4, usually on a sterno-metasternal shield.

Leg chaetotaxy usually as for the “Rhodacarus-group” genera referred to by

Evans (1963), Apotele usually three-pronged, never with associated hyaline

flap, and if two-pronged then there ure four ventral setae on tibia I. Male

18 REC. S. AUST. MUS. 16 (4): 1-219 July, 1970

with distally free spermadactyl, presternal genital orifice and seta av on

femur Tl larger than that of female and usually considerably modified to a

conspicuous spur,

MORPHOLOGY.

SCLEROTIZATION, There are always separate podonotal and opis-

thonotal shields on the deutonymph, and on the adults they may or may not

be fused together. On the male of Geogamasus delamarei and un unnamed

Hydregamasellus male there are lateral incisions partially separating the

podonotal and opisthonotal shields, The ventro-anal shield and the posterior

half of the peritrematal shicld may or may not be fused to other shields (see

“Morphology”, p. $8). ‘The female genital shield is rounded apteriorly or

has a pointed hyaline flap. and is truncated posteriorly, There are no

associated parasternal shields, but there is usually a central vaginal shield.

und sometimes small intergenito-ventral shields posteriorly. There is always

a continuous inale sternito-genital shield, The extent of the fusion between

the pre-endopodal, jugular, sternal, metasternal, endopodal and expodal

shields varies. In some subfamilies there is conspicuous sexual dimorphism

of the sclerotization.

CHAETOTAXY, Idiosoma! There are usually 20, 21 and 22 pairs of

setae on the podonotum, But there may be 23 pairs (Rhodacarus and

Rhodacaropsis) or there may be hypertrichy of the podonotal setae as in some

members of the Gamasiphinae or Gamasellini, The two monogeneric sub-

families have an unusually low number of podonotal sctue; Laclaptonyssinae

having 13. 19 or 20 pairs and Tangaroellinae having 16 pairs. The

setation of the opisthonotum is very variable. There ure usually four pairs

of setae On a sterno-metasternal shield and one pair on the genital shield,

but setae srl and sf4 may be on separate shields, and seta sé may be on

striated cuticle or it may be absent as in’ Evanssellus medusa and

Laelaptonyssus mitis. Setae Jv2, Jv3, 4v2, a pair of paranals and an

unpaired postanal are always on the ventro-anal shield. There are no cuanal

setae on the adult anal valves,

Legs: The commonest chactotuctic pattern is that found on the legs of

Gamasellus species where the coniplement of setae on each leg segment is:

coxae I to IV, 2-2-2-1: trochantera I to TV, 6-5-5-5: femora I to TV,

13-11-6-6: genua I to TV, 13-11-9-10: tibiae T to IV, 14-10-8-10: tarsi U

to IV. 18-18-18. Eighteen variants from this pattern ure known, usually

with fewer setae. Only two genera huye less than the maximum complement

of 14 setae on tibia I: Aydregamasus has 13 setae (2-6/3-2) and Laelap-

ronyssus has 9 setae (1-4/3-1) or 11] setae (2-4/3-2).

OTHER CHARACTERS. Apotele three-pronged, except for the two-

pronged apotele of Tungarvellus porosus, and without an associated hyaline

flap, Conspicuous salivary styli lie ventro-laterally to chelicerae. Movable

LEE—RHODACARIDAE 19

digit on male chelicera carries u distally free spermadactyl, although the two

are fused into what appears to be a single process in some species of

Afrogamasellus. Chaetomorphy of most setae varies through a wide range

of shapes, and there are thin walled sensory setae at the distal tip of tarsus T,

strong, grooming setae on the antero-lateral margin of the palp genau and

enlarged, spur-like setae on the male leg IL The spermathecal system i the

female is either closely associated with the vagina or there ure y pair of

ringed tubes opening on the idiosoma or proximal leg segments, level or

nearly level with genital shield, and leading to a single, central sacculus.

The male genital orilice is presternal and level with the anterior edge of

acetubulum [, although it may be enclosed by the fused sternal and pre-

endopodal shields.

DISTRIBUTION. The family as a whole is cosmopolitan, having been

found in every major zoogeographical region, although none of the individual

genera have this wide distribution and their geogriphical range varies

considerably. Taxa within the family are more numerous and morpho-

logically diverse in the extra-holarctic regions, For example, 5 genera

we known only from the holaretic region, while there ure 31 genera from

only the extra-holaretic regions, with 9 found in both holarctic and extra-

holarctic regions.

This distribution of the Rhodacaridae has resulted in it being considered

as replacing the Parasitidae, which is more numerous in the Northern

Hemisphere, as free-living predators in the ground habitats of the Southern

Hemisphere (Athias-Henriol, 1968 and Balogh, 1963a) or the extra-

palaearctic regions (Sheals, 1962). Meinbers of the Rhodacaridue are very

much rarer than those of the Parasitidae in Palaearctic regions but sre

probably as common in Nearctic regions, commoner in Tropical regions and

very much commoner in Southern Temperate regions. But, the enormous

diversily in form of rhodacarid mites in Southern Temperate regions suggests

that they replace more than the morphologically conservative Parasitidae.

The rhodacarid fauna of the Northern Temperate regions is similar

throughout, except that Hydrogamasus and Sessiluncus appear to be conlined

to the Palauearctic region. The Tropical regions show some of the differences

that are found between the rhodacarid faunas of the major Southern

Temperate regions, but because two well represented rhodacurid genera,

Rhodacarusy and Garasiphis, have a pantropical distribution and may be

the only rhadacarid genera present in some loculities, the rhodacarid faunas

of these regions are fairly similar throughout. In the Southern ‘Temperate

regions there are considerable differences in the rhoducarid faunas of the

Neotropical, Ethiopian and Australian regions indicating that they have been

evolving in isolation from cach other for some time. On the other hand,

in the Subantaretic and Antarctic regions, there appears to huye been a

2) REC. 8. AUST. MUS., 16 (3); 1-219 July, 1970

relatively recent circumpolar distribution because of the similarity between

the rhodacarid faunas,

A table summarizing the size and distribution of supraspecific taxa, as

well as a map of the zoogeographical regions used, is given at the end of

this paper (p. 204 and Fig. 427).

REMARKS, The concept of the Rhodacaridae used here is based on

the Rhodacarus-group of Evans (1963), The approximately equivalent taxa

in other systematic frameworks would be Hirschmann’s (1962) Gamasellini,

Johnston’s (1968) Rhodacaridae and Cyrtolaelapidae, Karg’s (1965)

Rhodacaridae (in part, excluding the Digamasellidac) and Gamasellinac.

and Ryke’s (1962b) Rhodacaridae (in part, excluding some Ascidae, the

Digamasellidae and Halolaelapidae). Bregetova (1967) places Rhodacarus,

Asca, Digamasellus, Halolaelaps and Cyrtelaelaps in the Rhodacaridae, and

appears to exclude by definition all species with a holonotal shield in the

adult.

The Rhodacaridae is probably closely allied to both the Parasitidae

and Veigaiidae, with the same type of leg chaetotaxy, and the Digamasellidae

and Halolaelapidae, with a reduced leg chaetotaxy. Although the Rhoda-

caridae is almost certainly not similarly allied to the Ascidae, some species.

which would be atypical of either of these families, are placed in one of

them with little certainty, Finally, on superficial examination, some members

of the Pachylaelapidae, Parholaspidae and Zerconidae (females only) muy

be thought to be rhodacarids.

Within the family, as recognized here, there are six major subgroups,

which I recognize formally as subfamilies. The names Rhodacarinae

Oudemans, 1902a, Luelaptonyssinae Womersley, 19564 und Ologamasinae

Ryke, 1962b are already available for three or these groups and below 1

propose the names Gamasiphinae, Sessiluncinue and Tangaroellinae for the

other three groups. Adults of these groups can be distinguished by using the

following key.

KEY TO SUBFAMILIES OF RHODACARIDAE

|. Femur I with less than 13 setae, having only

2 or 3 ventral setae .

tnd

Femur | with 13 setae, having 4 ventral setae

2, Apotele 2-pronged. Ley chaetotaxy as for

Gamasellus, except femur [ with 12 setae

(one ventral seta less), femur 1V with 7

setae (one ventral seta more) and tarsus

IV with 16 setae (two dorsal setae less) ., Tangaroellinae

LEE—RHODACARIDAE

Apotele 3-pronged. Chaetotaxy of many leg

segments differs from Gamasellus, for

example tibia I has only 9 or I] setae ..

3. Tarsus [V with 17 setae, seta p/4 absent. An

exception, with 18 setae because seta p/4

is present on tarsus IV, is one Afrogama-

sellus sp. which has seta stl on an area of

sternal shield with punctate sclerotization

Tarsus 1V with 18 setae or, if 17 setae, seta

pd4 absent. Seta stl on an evenly sclero-

tized area of sternal shield ..

4, Peritrematal shield not fused to exopodal 1V

shield or, if fused, then completely merged

with exopodal shields behind — stigma.

Ventro-anal shield rarely fused to notal

shield and when it is (as in Stylochirus),

seta al on palp femur is in central third.

Peritreme extends in front of anterior edge

of acetabulum IIIf (except in some species

of Stvylochirus) and setae on male tarsus

II are never enlarged into tubercles ..

Peritrematal shield generally fused to exopodal

[V shield, but usually it is also’ partly

separated from the exopodal shields behind

the stigma by a fissure running backwards

from between the anterior parts of the peri-

trematal and exopodal shields. On_ palp

femur, seta a/ is in proximal third (except

in some species of Litogamasus). It

peritrematal shield is completely merged

with exopodal shields behind stigma (as in

some species of Gamasiphis and Hiniphis),

yentro-anal shield is fused to notal shield.

Or, if peritrematal shield not fused at all

to exopodal shields then ventro-anal shield

is either fused to notal shield (as in some

species of Caliphis, Hydrogamasus and

Geogamasus males), or the mite is minute,

with a short peritreme not extending in

front of anterior edge of acetabulum TIT (as

in Neogamasellevans and Notogamasellus),

or it is the male of a littoral mite with an

Laelaptonyssinae

Rhodacarinae

Sessiluncinae

22 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

enlarged leg TV and seta av2 on tarsus IT

enlarged into tubercle (as in Litogamasus

and Parasitiphis) .. .. 2. 1. .. 2. .. 5

5. Discrete genital shield covering genital orifice

which is posterior to sternal shield. Female 6

Sternito-genital shield with genital orifice close

to its anterior margin. Male .. .. .... 7

6. Ventro-anal shield fused to notal shield except

on some females of Gamasiphoides. On

sterno-metasternal shield a line joining

setae sf2, sf3 and st4 would enclose an

angle of less than 95°, except in Hydro-

gamasus which has 13 setae (2-6/3-2) on

tibia I. Some females of Gamasiphis and

Caliphis (e.g.. G. concilator and C. calvus)

have seta st3 so far forward that it could

be mistaken for seta s/2, so that they may

be confused with Ologamasinae females on

which the ventro-anal shield is fused to the

notal shield (compare Fig. 64 and 233).

But, the absence of a split in exopodal III

shield and having the combination of less

than two lateral prongs on palp genu seta

all, a peritrematal shield not fused to the

ventro-anal shield and the sternal shield

continuous with endopodal IV shield dis-

tinguishes Gamasiphis and Caliphis females

from such Ologamasinae females .. .. .. Gamasiphinae

Generally ventro-anal shield not fused to notal

shield, but there are various exceptions.

On sterno-metasternal shield a line joining

setae sf2, sf3 and st4 is usually nearly

straight, and never encloses an angle of less

than 95°, except on some females of the

Gamasellus discutatus-complex, which can

be distinguished by having complex notal

setae and a ventro-anal shield that is not

fused to notal shield. Chaetotaxy of leg I

always as for Gamasellus, with 14 setae

(2-6/4-2) on tibia .. 2. 2. 2. 2. 2... ~ Ologamasinae

LEE—RHODACARIDAE 23

7. Ventro-anal shield fused to notal shield except

on some males of Gamasiphoides, but not

to peritrematal shield. © Gamasiphoides

males with ventro-anal shield not fused to

notal shield have simple idiosomal setae, no

lateral prongs on palp genu seta all, 2 pairs

of pre-endopodal shields and a holonotal

shield. Idiosomal sclerotization always

similar to that of female .. .. .. .. .. Gamasiphinae

Ventro-anal shield cither not fused to notal

shield or, if fused, also fused to peritrematal

shield. Exceptions with «a ventro-anal

shield fused to notal shield and not to peri-

trematal shield are found in the Gamasellus

discutatus-complex and pyriformis-complex

which can be distinguished by having both

the podonotal seta /4, <5, and r3 obviously

pilose amongst simple or lanceolate setae

and 2 or 3 pairs of pre-endopodal shields.

Also some males of Parasitiphis are similar

exceptions but can be distinguished by

having seta av2 on tarsus Il modified into

a tubercle. If ventro-anal shield not fused

to notal shield or peritrematal shield then,

cither some idiosomal setae are complex,

or palp genu seta a/l has more than 2

lateral prongs, or there is a single pair of

pre-endopodal shields or there are separate

podonotal and opisthonotal shields. Idioso-

mal sclerotization often differing consider-

ably from that of female .. .. .. .. .. Ologamasinae

Subfamily RHODACARINAE Oudemans

Rhodacarinae Oudemans, 1902a, p. 48. Type-genus: Rhodacarus

Oudemans, |902a.

DIAGNOSIS: Minute or small mites which are generally lightly

sclerotized, except for Afrogamasellus, a genus restricted to the Ethiopian

region, which contains a number of average sized, heavily sclerotized species.

The sclerotization is often conspicuously sexually dimorphic. Only very

rarely are opisthosomal shields fused to those of the podosoma. Seta p/4

is absent on tarsus [V, except on one unnamed species.

24 REC. 8. AUST. MUS. 16 (3): L210 July, 1970

MORPHOLOGY.

SCLEROTIZATION. The shields on the podosoma ure not usually

fused to those on the opisthosoma, but on Afrogamasellus luberoensis the

podonotal is fused to the opisthonotal (although a distinct groove marks the

line of fusion), and in an unnamed species described by Loots (thesis, 1967)

the female (the male is unknown) metapodal is completely merged with the

peritrematal. In most species, except the heavily sclerotized 4 fregamusellus

species, parts of the shields have punctate sclerotization, Rarely are there

free pre-endopodals as in Rhadacaropsis, although the jugular may be discrete

from the sternal. The male sclerotization is usually more extensive than that

of the female (except in some species of Afregamasellus and Rhodacaropsis

where it 1s similar in both sexes) with the ventro-anal fused posteriorly to

the opisthonotal, the metapodal sometimes enlarged or in lightly sclerotized

species fused to the opisthonotal by punctate sclerotization and, in some

Afrogamasellus species, the peritrematal is enlarged. Unlike the Ologama-

sinae, the other subfamily with many species having sexually dimorphic

sclerotization, the male ventro-anal is never fused to the peritrematal.

CHALTOTAXY. Idiosoma: There are 22 or 23 pairs of podonotal

selae, except on one species of Afrogamasellus where there are 18 pairs, If

there are 23 pairs they are regarded as being in rows with 6/, 6, 6s, 5r,

unlike Lindquist and Evans (1965, p, 14) who would regard the podonotal

setae of Rhedacarus as being in rows with 6j, 62, Ss, 6r, naming the four

pairs of setae on the anterior edge of the podonotal shield jl, zl, sl. rl.

Here, row j is considered to be compressed forward in Rhodacarus (compare

Figs. 5, 14 and 20) so that /l, /2, zl, sl are on the anterior edge of the

podonotal shield, and therefore, as in other Gamasina, rl does not exist.

6/, 5 or 6z, 3, 5 or 6s, 4 or Sri SV, SZ, SS. 2, 4 or SR: Sst B4v, 3Zv,

O or Sv.

Legs: Unique amongst rhodacarids. and Gamasina, is the absence of

pl4 on tarsus 1V, except for one unnamed species of 4 frogamasellus described

by Loots (thesis, 1967) where it is present on the adult but absent on the

deutonymph. The only other variant from the Gamasellus leg chaetolaxy

is the absence of some dorsal sctac on femur II and IIL on at least one species

of Rhodacarapsis.

OTHER CHARACTERS, On palp femur, seta al is on central third, as in

Laeloptonyssus and a number of Sessiluncinae genera, in contrast to nearly all

members of the Ologamasinae, Gamasiphinue and Tangaroellus where it is

on the proximal third. Dorsal sctae are usually simple and setose but in

some 4 frogamaselluy species they are spatulate and/or pilose. The sperma-

dactyl is unusual in either being recurved (Rhedacarus, Rhedacarellus and

1 KE—RHODACARIDAF 25

Rhodacaropsis) or being merged with a reduced movable digit (4Afrogama-

sellus), A recurved spermadactyl does occur on members of other sub-

families (Pyriphis and Sessiluncus). Pulvilli I-fY with paired lateral, short

lanceolate lobes and paired central circular lobes.

DISTRIBUTION, = Rhedacarus (and probably R/odacarellus and

Rhodacaropsis when more collecting is done) is cosmopolitan except that tt

has not yet been collected from the polar regions or the southern Neotropical

regions; is commonest in the deeper soil layers and the littoral zone and is

constant in form as well as being small and lightly sclerotized. 4 frogamasellus

is confined to the Ethiopian region, is more often collected from plant litter

or upper soil layers and is variable in form, as well as sometimes being

larger and heavily sclerotized.

REMARKS. Although the concept of this subfamily is similar to that

of the Rhodacaridae up until 1955, the important diagnostic characters used

here are largely those which will be used by Loots (in press), as are the

characters used to distinguish the genera it contains, The addition of a new

genus, Afrogamasellus Loots and Ryke (1968), is important in clarifying

which characters of the previous nominal members of the Rhodacarinae are

absent when a species is not specialized for living in the high humidity and

small spaces between soil particles; for example, small size, light sclerotiza-

lion and products of an articulated narrow idiosoma that allows the separate

movements of the opisthosoma needed to circumambulate narrow soil pore

spaces. But, although the larger species of Afrogamuasellus are heavily

sclerotized and probably hemiedaphic, the opisthosomal shields are rarely

fused to those of the podosoma, and when they are there is a line of

demarcation suggesting that the fusion is secondary, This indicates that the

ancestors of these Afrogamasellus species may have been euedaphic like

Rhodacarus and had an articulated idiosoma.

The removal of Rhedacaroides trom the Rhodacarmae to the

Gamasellini depends partly on the characters of unnamed species that |

consider to be congeneric with the type species (which T have not seen).

But on Willmann’s (1959) original description there can be little doubt that

this genus is not closely allied to Rhoedacarus.

| recognize 4 genera within this subfamily, Rhedacarus, Afrogamasellus,

Rhodacarellus and Rhedacarepsis, Adults of these genera can be dis-

tinguished by using the following key,

KEY TO GENERA OF RHODACARINAE

1. Four pairs (jl, j2, <1 and yl) of setae on anterior

edge of podonotal shield. Third hypostomal seta

approximately halfway between Ayp2 and /iyp4

26 REC. S. AUST. MUS.,, 16 (3): 1-219 July, 1970

(capitular) setae. On ventral surface of podo-

notal shield, 3 light-refractile structures between

setae /5 and /6

Two or 3 pairs (always jl and <1, sometimes j2)

of setae on anterior edge of podonotal shield.

Third hypostomal seta approximately level with

hyp2. On ventral surface of podonotal shield,

4 or more light-refractile structures between

setae 75: and jG 2262 2.0c ne be ke ee ew OS

2. No pretarsus I. No pre-endopodal shields of

normal sclerotization, although anterior edge of

sternal shield may be separated from the rest by

punctate sclerotization and there may be separ-

ate shields of punctate sclerotization .. .. .. Rhodacarus

Pretarsus I present, consisting of a pair of strong,

sessile — claws. Normally sclerotized _ pre-

endopodal shields separated from sternum by

Striated cuticle .. 2. 2. 2. 0. 2. 2. 2. .. ~=Rhodacaropsiy

3. Arthrodial process at base of movable cheliceral

digit is a simple coronet. Spermadactyl separates

from normal movable cheliceral digit at level of

single tooth and is recurved, enclosing an angle

of less than 100° 2. 2. 2... 0. ee 2 2. 2.) Rhodacarellus

Arthrodial process at base of movable cheliceral

digit is produced into a conspicuous brush.

Spermadactyl curves in the same way as movable

cheliceral digit, which is reduced and may be

entirely merged with it... 2. 2. 2... 2...) Afrogamasellus

Genus RHODACARUS Oudemans

Rhodacarus Oudemans, 1902a, p. 50. Type-species: Rhodacarus roseus

Oudemans, 1902a, by monotypy.

DIAGNOSIS. Minute or small mites, with lightly sclerotized, slim,

centrally articulated idiosoma and a relatively large, well sclerotized

gnathosoma. Third hypostomal seta approximately halfway between second

and fourth hypostomal setae. Podonotal shield has 4 pairs of setae on

anterior margin and 3 light-refractile structures, Pre-endopodal shields

absent. Spermadactyl recurved. Pretarsus I absent.

1. EE—RHODACARIDAE 27

MORPHOLOGY,

ScLEROTIZALION, Female: Separate podonotal (may be fragmented

into three or four smaller shields) and opisthonotal. On ventral surface

of podonotal, 3 light-refractile structures between setae j5 and /6 (usually

on rhodacarids there are 4 such structures, if present, but Rhodacaropsis

also. has 3). Discrete ventro-anal (bearing ariculae posterior to

anus) widely separated from genital and exopodal TV, Discrete metapodal

which may be fragmented and have punctate sclerotization. Intergenito-

ventral shield present. Peritrematal reduced and may or may not be narrowly

fused to podonotal and exopodal IV. If present, exopodals only between of

al posterior edge acetabula. Sterno-metasternal fused to endopodal IL and

part of Hl. Pre-endopodals absent, although anterior edge of sternal may

be normally sclerotized and separated from similarly sclerotized areas by

punctate sclerotization. Areas of punctate sclerotization on edges of many

shields, seta stl usually on such an area,

Male: Ventro-anal fused to opisthonotal, but widely separated from

sternito-genital, exopodal TV and peritrematal. Metapodal may be broadly

fused to opisthonotal by punctate sclerotization.

CHAETOTAXY, Idiosoma: 6j, 62, 6s, Sr: SJ, 5Z. 58, SR: Set:

3Jv, 3Zy,

Legs: As Gamasellus, except tarsus LV lacks p/4.

OTHER CHARACTERS. Female: Size of gnathosoma relative to

idiosoma is unusually large and chelicerae are long and heavily sclerotized.

Deutosternal denticles in 7 horizontal rows with a longitudinal und a

horizontal deutosternal ridge anterior to denticles. Tectum anterior margin

basically trispinate, but the large central spine, and more rarely the lateral

spines, may carry spinules distally. Hypostomal setae almost in a longi-

tudinal line (unique to this rhodacarid genus and Rhadacaropsis). Movable

cheliceral digit with 3 teeth. On palp genu, setae a/l and al2 almost level

and very slightly spatulate. On palp femur, seta al in central third.

Sclerotization is pale, and may be pinkish with a red gnathosoma. Idiosoma

is constricted between podosoma and opisthosoma. Dorsal setae simple,

setose. Pretarsus T absent. Spermathecal ringed tube not located in this

study, but Athias-Henriot (in press) states that it opens on femur TI,

Pretarsus | absent. Amongst dorsal setae on tarsus TV, seta pd3 is the

longest.

Male: Movable cheliceral digit with one tooth and fused at base to

recurved, spatulate spermadactyl, On leg HH. femur setae av and pvi, genu

seta av, tibia seta av, enlarged into spurs.

28 REC. S. AUST, MUS., 16 (3): 1-219 July, 1970

Ss _

FOSS ‘

f Se

22 =

cai _—o—_

o—o 3.

\aenes

i‘

XO. F

an li

uw —

gs”

Seah eerie: th

poss

—=<—

Eee:

Z f a4 4 ‘\ Pp

13 ° a eared 2

-_ | * A hes

1004 “a St

11 ff

9 “hy

Figs. 5-13. Rhodacarus roseus Oudemans.

5-9. female: 5, soma, dorsum; 6, idiosoma, venter; 7, gnathosoma, yenter: 8. leg IV

(part), dorsal setae only; 9, tarsus I, distal tip. 10-13, male: 10, leg Il (part),

antero-latus; 11, chelicera; 12, opisthosoma, latus; 13, idiosoma, venter.

LEF—RHODACARIDAE 29

DISTRIBUTION, Nn, Na; NTa, NTb: Ew, Ee, Es: Pe, Pm: Oi, Om:

Am. Aa. Besides the locality records published with the original descriptions

of nominal species or forms there are records of Rhodacarus species from

North America (Emberson, thesis, 1968; Fox, 1967; Haq, 1965; Rodriguez

and Ibarra, 1967), Africa (Loots, in press), Israel (Costa, 1966a) and

numicrous records from the european Palaearetic region. T regard as dubious

the record of u Rhedacarus sp. from the Crozet Islands (Sk) by Richters

(1907) and it should be noted that Rhodacarus costar from Argenuna has

heen removed from this genus. The specimens from Neotropical regions

(from Jamaica, Trinidad and British Guiana) and the Oriental and malayan

Australian regions (from Nepal, Malaya and Solomon Islands) are before

me and to be dep, BM(NH). It will be interesting if Rkodacarus continues

to be absent from collections taken from southern Neotropical regions,

Found amongst plant litter and soil (particularly deeper layers below

3 inches) and in littoral zone

REMARKS, The concept of Rhodacarus was broader when Ryke

(1962b) included Rhodacarellus, Rhodacaropsis and Rhodacaroides as sub-

genera. It is possible that Rhodacaropsis should be synonymized with

Rhodacarus. Previously species have usually been correctly placed in this

genus, but I consider Rhodacarus costal a species incertae sedis and not a

member of the Rhodacarinae, while the transfer of Rhodacarellus minimus

to this genus by Hirschmann, 1962, is incorrect, The tollowing 17 nominal

species are, therefore. included in this genus: R. roses Oudemans, 1902a;

R. ananasi Ryke, 1962a; R. angustiformis Willmann, 1951. R. calcarilatus

Berlese, 1920 (syn. R. pallidus in Sheals, 1958); R, clavularus Athias-

Hewriot, 196lb; R. corenatus Berlese, 1920; R, cuneatus Athias-Henriot,

l96lb; R. denriculatus Berlese, 1920; R. lauret? Athias-Henriot. 19616;

R. mandibularis Berlese, 1920 (syn. R. roseuy in Sheals, 1958); R, marksae

Domrow, 1957; R. pallidus Hull, 1918; R. reconditus Athias-Henriot, 1961b;

R. rhodacaropsis Ryke, 1962a; R. stenzkei Willmann, 1957; R. sublapideus

Ryke, 1962a; R. tribaculatus Athias-Henriot, 1961b. Two new species are

described by Loots (in press). Also there are s/mplex forms of R. roseus

(in Sheals, 1958), R, corenatus (in Athias-Henriot, 19Y61b) and R. pallidus

(in Sheals, |958, referred to us ecalcarulatus torn) in which the podonotal

shi¢ld is not split into anterior and posterior sections by a V-shaped fissure,

Rhodacarus roseus Oudemans

Rhedacarus roseus Oudemans, 1902a, p. 50,

FEMALE, Fig. 5-9. Idiosomal length, 450.. The punctate sclerotiza-

tion is usually an area where thicker sclerotization is limited to spots on a

thin shield, but at the anterior of the ventro-anal shield there are areas where

the thicker sclerotization has spots of thin sclerotization in it. The anterior

30 REC, S. AUST, MUS, 16 (4); 1-219 July, 1970

end of the peritreme is joined to the small humeral section of the podonotal

shield by a thin strip of thickened sclerouzation.

MALE. Fig. 10-13. Idiosomal length, 410». In Fig. Tl, tooth on

movable cheliceral digit is mainly obscured by spermadacty|,

LOCALITY, Three females (N196879-N196881) and 4 males

(N196882-N196885) drawn or examined: Australia; LF5S8, grass und

leaf litter under Eucalyptus camatlidulensis, Heywood Purk, Adelaide, South

Australia, 6.6.1965, col. R. V. Southcotl, dep. SAM.

REMARKS. R. roseus is the best name for the above specimens,

although there are slight differences between them and the previous descrip-

tions of this species, particularly in the extent of the punctate sclerotization.

Genus AFROGAMASELLUS Loots and Ryke

Afrogamasellus Loots and Ryke, 1968, p. 2. Type-species: Cyriolaclaps

(Gamasellus) franzi, Loots and Ryke, 1966, by original designation,

DLAGNOSIS. Minute to average sized mites, with light to heavy

sclerotization. Idiosoma is often ellipsoidal without central articulation.

Second and third hypostomal setae nearly level, both being a similar distance

from hypostomual seta 4, Podonotal shield hus 2 or 3 pairs of setae on

anterior margin and 4 light-refractile structures. Pre-endopodal shields

absent. Spermadactyl not recurved but broadly merged with reduced

inovable digit of chelicera. Pretarsus | present,

MORPHOLOGY,

SCLEROTIZATION, Female: Usually separate podonotul and opis-

thonotal, but may be holonotal with groove across shield where split is

normally. On ventral surface of podonotal, 4 light-refractile structures

between setae /5 and j/6. Discrete ventro-anal (sometimes bearing aciculae

posterior Lo anus) sometimes fused to metapodal and anterior margin may

or may not lie close to genital and exopodal TV. Usually discrete metapodal.

Intergenito-ventral shicld absent. Peritrematal usually fused to exopodal

IV, and rarely to metapodal us well, but may be reduced and not fused to

any shields posteriorly. Split in exopodals 1, WT and IV. Sterno-metasternal

tused to endopodals If and HI, and rarely to endopodal TV as well, May be

separate jugular, or punctate anterior area of sternum which may or may not

carry seta wl. Pre-endopodals absent, although anterior edge of sterno-

metasternal may be normally sclerotized and separated from similarly

sclerotized areas by punctate sclerotization.

Male: May be similar to female, or metapodal is enlarged, or metapodal

and peritrematal are enlarged with the peritrematal and ventro-anal fused

to the notals and in one species the metapodal is joined to the opisthonotal

by punctate sclerotization. Usually an entire sternito-genital, but section

1. FE—RHODACARIDAE ul

carrying seta yfS may be separated from rest and fused to endopodul IV.

Although ventro-anal shield is always separate from sternito-genital it ts

usually clase to it.

CHAETOTAXY, Idiosoma: 6j, 5 or 62, 3, 5 or 6y, 4 or Ses SV, 52. 55,

dor 5R: 5st: 3/v, 3Zv, 18v-

Legs: As Gamasellus, except tarsus TV Jacks p/4 (other than in one

unnamed species where it is present on the adult but missing on the

deutonymph ),

OTHER CHARACTERS. Female: Relative size of gnathosoma com-

pared to idiosoma is average. Deutosternal denticles i 7 horizontal

rows with a horizontal deutosternal ridge anterior to denticles. Tectum

anterior margin may be basically unispinate or trispinate, and usually also

has spinules and sometimes the central spine broadens out at its tip, Third

hypostomal seta nearly level with seta /ryp2, both being a similar distance

from seta Ayp4. Movable cheliceral digit with 3 teeth. Cheliceral arthrodial

process at base of movable digit is extended into short brush. usvally about

half the length of the movable digit, while in most other genera it is a simple

coronel. On palp venu, seta all and seta al2 may both be spine-like or

spatulate or seta a/| may be spatulate and seta a/2 slightly pilose. On palp

femur, seta a/ in central third, Sclerotization usually shiny, Dorsal setae

usually tapering and simple, may be slightly pilose. or pilose and spatulate.

On the basis of my examination of female A, retrastigma and A. mitigatus,

I consider it probable that the pore at the anterior end of the metapodal

shield (see Figs. 17 ancl 22) is the opening of the spermathecal ringed tube,

This pore is referred to as the “quasi-stigma” by Loots (1969), The

strongest evidence supporting my homology is the description of the female

and male of an unnamed species by Loots (thesis, 1967): the female has a

long, internal ehitinized tube attached to the pore, which is absent on the

male und the male spermadactyl is unusually long and narrow, [lf this

homology is correct, then the pore drawn on the male metapodal shield of

A, tefrastigema by Loots (1969) would be homologous to the pore near the

posterior, paraxial margin of the female metapodal shield (see Fig, 17).

Further evidence is required before the location of the aperture of the

spermathecal ringed tube is considered to be probably on the female

metapodal shield of wffregamaselluy species other than those belonging to

the tefrastigma-complex or the unnamed species mentioned above. Pretarsus

I may be pedunculate with smaller claws than other pretarsi or broadly fused

to tarsus. Leg | often thicker than other legs. Femur Il may have ventral

process, Amongst dorsal setae on tarsus IV, setae pd3 and pd4 are the

longest being subequal in length.

Male: Movable cheliceral digit reduced and merged into spermadactyl,

sometimes to the extent that only the grooved spermudactyl appears present,

but often there is a conspicuous dorsal tooth which could be homologous

32 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

—— 0— 90 —_

9? ro)

—_°?

No Scale— 20, 21, 22

14,17 .

15 eens

Figs. 14-22. Afrogamasellus Loots and Ryke.

14-19, A. fetrastigma (Berlese), female: 14, soma, dorsum; 15, pretarsus 1 and

tarsus [ (part): 16, leg [V (part), dorsal setae only; 17, idiosoma, venter: 18, tibia I.

11, IT and lV, ventral setae only; 19, gnathosoma, venter. 20, nominate species of

camaxiloensis-complex Loots (1969), podonotal chaetotaxy. 21, A. siecincris

(Berlese), parts of idiosoma, yenter. 22. 4. mitigatuy (Berlese). acetabulum IV

and surroundings.

LEE—RHODACARIDAE as)

with the female teeth or could represent the tip of the movable digit. On

tarsus I, in one species, one of the distal, hollow setae is ovoid, being swollen

compared with the equivalent female seta, On leg Il, seta av on feniur and

venu enlarged into spurs, May be other spurs on these segments and on

libia I, but their homologies are not known.

DISTRIBUTION. Ew, Ee, Es. Besides the locality records published

with the original descriptions of nominal species, there are other records of

linnamed species described by Loots (thesis, 1967), Members of this genus

have not been collected south of 30°S.

Found in plant litter and soil,

REMARKS. Before 1968 the species now included in A fregamasellus

were placed in Gumasellus (as subgenus of Cyrtolaelaps). Loots (1969)

states that this is a heterogenous taxon and divides it into the following

4 species-complexes, which contain 18 nominal species and 7 unnamed

species (Loots, in press),

|. quadrisigillatus-succinetus-complex (includes the type-species of

the genus).

The following 9 nominal species, plus a subspecies distinguished from a

nominule subspecies, are included in this species-complex: A. quadrisigillatus

(Berlese. 19l6a); A, svecinctus (Berlese, 1916a); A. franzi (Ryke and

Loots, 1966); A, kilimanjarvensiy (Ryke and Loots, 1966); 4, kivuensiy

(Ryke and Loots. 1966); 4. leleupi (Ryke and Loots, 1966); A. /unguensis

(Ryke and Loots, 1966); A. maskamensis (Ryke and Loots, 1960);

A, uviraensis (Ryke and Loots, 1966); A. uviraensis congoensis (Ryke and

Loots, 1966).

Jugular shield may be present, or seta s¢#] may be on the sternal shield

or on punctate sclerotization, If punctate sclerotization is present in region

of seta stl, it either bears this setae when it may or may not separate a

normally sclerotized anterior edge of the sternal shield from the rest (Fig.

21), or it separates off a jugular shield, FEMALE. Metapodal shield

smaller than anus, sometimes with one or two small elongate shields between

it and the ventro-anal shield. Striated cuticle between ventro-anal shield

and podal shields never more than moderately extensive so that anterior edge

of ventro-anal shield is never much further (more than 10,) from sternal

seta sf4 than the distance between both setae st4. MALE, Ventro-anal

shield fused to notal shield posteriorly, and the metapodal shield is much

larger than the anus, with a triangular outline wedged between the lateral

margins of the ventro-anal and notal shields. Sternito-genital shield

undivided and broadly fused to endopodal IV shield,

34 REC. S. AUST, MUS. 16 (3); 1-219 July, 1970

2. camaxiloensis-complex.

I have only seen the descriptions of 7 unnamed species belonging to

this species-complex by Loots (thesis, 1967), although, since a diagnosis is

given for this species-complex by Loots (1969), names for these species

must now be formally recognized and the descriptions by Loots (in press)

published.

Jugular shield absent, but always an area of punctate sclerotization in

this region, which may or may not bear seta yl, and which never separates

a normally sclerotized anterior cdge of the sternal shield from the rest,

FEMALE. Metapodal shield smaller than anus, and with a small elongate

shield between it and ventro-anal shield, Extensive area of striated cuticle

between ventro-anal shield and podal shields; the anterior margin of ventro-

anal shield is much further from sternal seta sf4 than the distance between

both seta s#4. Posterior margin of genital shield usually bordered by punctate

sclerotization. MALE, Ventro-anal shield fused to notal shield posteriorly.

Metapodal shield larger than anus with a triangular outline wedged between

the lateral margins of the ventro-anal and notal shields. Sternito-genital

shicld around seta st5 is separate from the rest of the shield and fused to

endopodal TV shield, while the rest of the sternito-genital shield is separated

by punctate sclerotization or striated cuticle from endopodal TV shield.

3. luberoensis-complex.

The following single nominal species, plus a subspecies distinguished

from the nominate subspecies, is included in this species-complex: A. lubero-

ensis Loots, 1968; A. luberoensis kalibuensiy Loots, 1968,

This is the only species in the Rhodacarinae on which adult podosomal

and opisthosomal shields are fused together; the podonotal and opisthonotal

shields are fused to form a holonotal shield and the metapodal is fused to

the peritrematal shield, and only on the female to the ventro-anal shield as

well, Otherwise this species is similar to members of the fetrastigma-complex,

except that in the nominate subspecies the metapodal is smaller than the

anus. The male ventro-anal shield is not fused to the notal shield.

4, terrastigma-complex.

The following & nominal species are included in this species-complex:

A. tetrastizma (Berlese, 1916a): A. celisi Loots, 1969; A. evansi Loots,

1969; A. kahusiensis Loots, 1969; A. mitigatus (Berlese, 1923): A.

muhiensis Loots, 1969; A, rugegensis Loots, 1969; A. nyinabitabaensis

Lools, 1969,

Jugular shicld present and discrete. FEMALE. Metapodal shield

larger than anus, usually discrete, but narrowly fused to ventro-anal shield

on A. mitigafus, Usually striated cuticle inconspicuous between ventro-anal

LEE—RHODACARIDAF 45

shield and podal shields and never more extensive than on the females of

the quadrisigillatus-succinetus-complex. MALE. Ventro-anal shield may

or may not be fused to notal shield posteriorly; if it is not fused then the

inetapodal shield is smaller than the anus, but if it is fused then the metapodal

shield is much larger than the anus with a triangular outline wedged between

the lateral margins of the ventro-anal and notal shields. Sternito-genital

shield undivided and broadly fused to endopodal IV shield.

Two further species-complexes, each including a single unnamed

species, were recognized by Loots (thesis, 1967), One of these unnamed

species has a conspicuous spermathecal ringed tube as mentioned above.

Afrogamasellus mitigatuy ( Berlese)

Gamasellas mitigatus Berlese, 1923. p. 250.

FEMALE. Fig, 22. Idiosomal length, unknown. The sketch (Fig.

22) shows that the metapodal shield bears a pore leading to an internal sac

and not an external process as in the drawing (Fig. 125) in Ryke, 1962c.

This internal sac is probably part of the spermathecal ringed tube.

MALE. Not known.

LOCALITY. The ‘tipico’ female (221/11), sketched and examined,

East Africa; col. Alluaud and Jeannell, dep. SEAP.

Afrogamasellus succinctus ( Berlese )

Gamasellus succinetus Berlese, 1916a, p. 160.

FEMALE. Fig. 21. Idiosomal length, unknown. The 3 sketches

(Fig. 21) given, show that this species is more like Afrogamasellus franzi

than the drawing (Fig. 3) of it (named C. (Gamasellus) succinctus) given

by Loots and Ryke (1966) indicates. The pore on the metapodal shield

may be homologous to the pore near the posterior, paraxial margin of the

metapodal shield of A. tefrastigma (see Fig. 17). or it may homologous

with the pore considered to be the aperture of the spermathecal ringed tube,

MALE. Not known,

LOCALITY. A ‘tipico’ female (175/35), and a ‘cotipico” female

(197/46), sketched or examined: East Africa: col Alluaud and Jeannell.

dep. SEAF.

Afrogamasellus tetrastigma (Berlese)

Gamasellus tetrastigma Berlese, 19] 6a, p. 161.

FEMALE, Fig. 14-19. Idiosomal length, 520e. The pore at the

anterior end of the metapodal shield, here considered to be the aperture of

the spermathecal ringed tube, is drawn by Loots (1969) on similar females

referred to this species. On the male the pore that is drawn by Loots (1969)

on the metapodal shield is probably homologous to the pore near the posterior,

36 REC. 8S. AUST. MUS., 16 (3); 1-219 July, 1970

paraxial margin of the female metapodal shield. Pretarsus | has claws

subequal in size to those of other pretarsi, and in a sclerotized sheath that

is broadly fused to the tarsus. Leg I is relatively large as drawing (Fig, 18)

of tibiae shows.

MALE. Specimens referred to this species described by Loots (1969).

LOCALITY. Two ‘tipico’ females (175/38) and a ‘cotipico’ female

(197/45) drawn or examined: East Africa; col. Alluaud and Jeannell,

dep, SEAF.

Genus RHODACARELLUS Willmann

Rhodacarellus Willmann, 1935, p. 429. Type-species: Rhodacarellus

subterraneus Willmann, 1935, by original designation.

DIAGNOSIS, Minute or small mites, with lightly sclerotized, centrally

articulated idiosoma and a relatively average sized gnathosoma. Second

and third hypostomal setae nearly level, both being a similar distance from

hypostomal seta 4, Podonotal shield has 2 or 3 pairs of setae on anterior

margin and 4 light-refractile structures. Pre-endopodal shields absent.

Spermadactyl recurved. Pretarsus | present,

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal (may be fragmented

into four smaller shields) and opisthonotal, On ventral surface of

podonotal, 4 light-refractile structures between setae (5 and j6. Discrete

ventro-anal (bearing aciculae posterior to anus) widely separated from

genital and exopodal IV. Discrete metapodal, Intergenito-ventral shield or

shields present, Peritrematal reduced but may or may not be narrowly fused

to podonotal and exopodal TV. If present, exopodals only between or at

posterior edge of ucelabula. Sterno-metasternal fused to endopodals IT and

Il, Pre-endopodals absent, seta stl on punctate sclerotization at anterior

end of sternal,

Male: Ventro-anal fused to opisthonotal, and metapodal enlarged and

fused to anterior end of this line of fusion. Ventro-anal widely separated

from sternito-genital and peritrematal.

CHAETOTAXY. Idiosoma: 6j, 6z, 5s, 5r: S5/, 57, 58, SR: Syste Av,

3Zv, 0 or 1SV.

Legs; As Gamasellus, except tarsus IV lacks p/4,

OTHER CHARACTERS. Female: Relative size of gnathosoma com-

pared to idiosoma is average or smaller. Deutosternal denticles in 7

horizontal rows, with a horizontal deutosternal ridge anterior to denticles,

Tectum anterior margin basically trispinate or quinquispinate, with the longer

spines nearer the centre, and spinules on or lateral to spines. Third

hypostomal seta nearly level with seta /yp2, both being a similar distance

LEE—RHODACARIDAE 37

from seta Ayp4. Movable cheliceral digit with 3 teeth. On palp genu, setae

all and al2 almost level and slightly spine-like, On palp femur, seta a/ in

central third, Idiosoma is constricted between podosoma and opisthosoma.,

Sclerotizution is pale. Dorsal setae simple, setose, Usually setae /vl and

Zv1 on striated cuticle anterior to ventro-anal shield but either both of them

(e.g., R. corniculatus), or just seta Zvl (e.g. R, kreuzi) may be on shield,

Spermathecal ringed tube probably opens near posterior paraxial edge of

acetabuluny IV. Pretarsus 1 pedunculate or broadly attached to tarsus, and

smaller than other pretarsi, Amongst dorsal setae on tarsus IV, seta pd3

is the longest.

Male: Movable cheliceral digit with one tooth and fused at base to

recurved, pointed spermadactyl. On leg IL, femur seta av, genu seta ay,

tibia seta av, and distal setae on tarsus enlarged into spurs.

DISTRIBUTION, Nn: Pe, Pm: Aa, The distribution of this genus

may be considerably more extensive than the records indicate. There are

“ number of Palacarctic records besides those published with the original

descriptions of nominal species, but the record from Israel (Costa, 1966a)

is outside this established range. The records from North America are from

Fox (1967) and Emberson (thesis, 1968) and a female (N1968280) from

South Australia is dep. SAM.

Found umongst plant litter and soil (particularly deeper layers below

3 inches).

REMARKS. The appearance of Rhedacarellus species is similar to

the lightly sclerotized species of Afrogamasellus, but the cheliceral arthrodial

process and spermadactyl are like those of Rhedacaruy and Rhodaearopsis.

The following 9 nominal species are included in this genus: RK. arcanus

(Athias-Henriot, 1961b); R. corniculatus Willmann, 1935; R. epigynialis

Sheals, 1956; R. francescae Athias-Henriot, L961b; R. krenzi Karg, 1965;

R. minimus Kary, (961: R. silesiacus Willmann, 1936: R. subterranneus

Willmann, 1935; R. vervacti (Athias-Henriot, 1961b). There is one sub-

species! R, epigynialis perspicuus Halaskova, 1959,

Genus RHODACAROPSIS Willmann

Rhodacaropsis Wilmann, 1935, p. 426. Type-species: Rhodacaropsis

inexpectatus Willman, 1935, by original designation,

DIAGNOSIS, Minute or small mites, with lightly sclerotized, slim,

centrally articulated idiosoma and a relatively large gnathosoma, Third

hypostomal seta approximately half-way between second and fourth hypo-

stomal setac. Podonotal shield has 4 pairs of setae on anterior margin and

3 lightrefractile structures. Pre-endopodal shields present. Spermadacty!

recurved, Pretarsus | present.

3S REC, 8. AUST. MUS., 16 (3): 1-219 July, J970

MORPHOLOGY,

SCLEROTIZATION. Female: Separate podonotal (may be fragmented

into smaller shields) and opisthonotal. On ventral surface of podonotal,

3 light-refractile structures between setae j/5 and /6. Discrete ventro-anal

(bearing aciculae posterior to anus) widely separated from genital and

exopodal IV, Discrete metapodal. Intergenito-ventral shields absent.

Peritreme reduced and peritrematal absent. If present, exopodals only

between or at posterior edge of acetabula. Sterno-metusternal fused to

endopodals I and Wl. Two pairs of pre-endopodals, but posterior pair

may be connected to sternal by punctate sclerotization. although seta srl is

never on such an area.

Male: Ventro-anal discrete. Sternito-genital is divided, a single

discrete posterior section carrying both setae s/5.

CHAETOTAXY, Idiosoma: 6/, 62, 6s, Sr: SJ, 52, 58, 4R: Sst: By,

37, LS.

Legs; As Gamasellus, except tarsus 1V lacks p/4 and femur I lacks

one dorsal seta (2, 4/3, 1) and femur If lacks 2 dorsal setae (1, 2/1, 0)

on an unnamed species for which the leg chaetotaxy was examined by Loots

(in press),

OTHER CHARACTERS. Female: Relative size of gnathosoma_ to

idiosoma is unusually large and the chelicerae are long and heavily sclerotized.

Nature of deutosternal denticles or ridges unknown. Tectum anterior margin

basically unispinate, but may be spinules at tip or base of spine and if the

latter are large enough, the tectum is similar to that of Rhodacarus species,

Hypostomal setae almost in a longitudinal line, seta Ayp3 being approximately

half-way between seta Ayp2 and hyp4, Movable cheliceral digit with 3 teeth.

Shape of setae on palp femur and genu unknown. Sclerotization pale.

Idiosoma constricted between podosoma and opisthosoma, Dorsal setae

simple, setose. Position of spermathecal ringed tube not known. Pretarsus

I reduced except for Jarge claws that are attached almost directly on to tarsus.

Male: Movable cheliceral digit with one tooth and fused at base to

recurved, spatulate spermadactyl. On leg If, femur seta av enlarged into a

spur and genu seta av may be spine-like,

DISTRIBUTION. Na: Es: Pe. The single palaearctic record was

published with the original description of the type-species. The record from

North America is of the type-specics and is from Haq (1965), and that from

South Africa is of an unnamed species (Loots, in press).

Found in the littoral zone in sandy rather rocky areas.

REMARKS. Rhodacaropsiy is closely allied to Rhoducarny. The

following single nominal species is included in this genus: R. inexpeetatus

Willmann, 1935, A new species is described by Loots (in press).

LEE—RHODACARIDAE 39

The 5 Rhodacaropsis species listed by Athias-Henriot (1961b) belong

elsewhere; 3 species were transferred to Protogamasellus by Lindquist and

Evans (1965) and the 2 species listed above under Rhodacérellus are

referred to that genus by Loots (in press).

Subfamily GAMASIPHINAE subf.n.

Type-genus: Gamuasiphis Berlese, 19044,

DIAGNOSIS. Minute to large mites, usually with extensive, well

sclerotized shields and often the idiosoma is strongly convex dorsally.

Holonotal shield or separate podonotal and opisthonotal shields, Ventro-anal

shield fused to the notal shield except on some Gamasiphoides species,

Peritrematal shield usually fused to exopodal LY shteld and never fused to

ventro-anal shield. Sclerotization not conspicuously sexually dimorphic.

On the female sterno-metasternal shield a line joining setae sf2, st3 and st4

would enclose an angle of less than 95° except in Hydrogamasus,

MORPHOLOGY.

SCLEROTIZATION. The extent of the fusion between shields is fairly

constant amongst species of this subfamily, and except for the fusion of

the genital with other shields it is the same amongst females and males

of the same species. In five genera there is always a holonotal, but in

Euepicriuy and Gamaselliphis there 1s a separate podonotal and opisthonotal.

The ventro-anal is fused to the notal, except on some Gumasiphoides species,

and is either fused to exopodal IV or lies very close to it, Intergenito-

ventral shields are only present on [ydrogamasus species. The peritrematal

is fused to the notal anteriorly and, although posteriorly it is never fused to

the ventro-anal, it is usually fused to exopodal TV, except on Caliphis, Laelap-

fella and Hydrogamasus, In many species there is a triangular posterior

extension of the apparent peritrematal, posterior to its fusion with exopodal

lV, which may be homologous with the metapodal and is referred to here as

the metapodal. The exopodals tend to be fused together; the exopodals

completely encasing the peraxial margin of acetabulum TH, except on

Gamaselliphis and Gamasiphoides species. The sterno-metasternal ts fused

to endopodal IV on Gamasiphis, Caliphis and Evepicrius species, and one

Gamasiphoides species. The pre-endopodal may be fused to the sterno-

metasiernal or it may be one or two pairs of discrete shields.

CHAE TOTAXY. The number of setae on the notum is variable, although

there are never less than 21 pairs on the podonotum, Caliphis and

Euepierius species may be hypertrichous on the notum, and Gamusiphis

species are sometimes hypotrichous on the opisthonotum. The leg chactotaxy

is as Gamayellus in Caliphis, Gamaselliphis and Laelaptiella, but amongst the

other four genera there are eight kinds of chaetotaxy differing from this

au REC. S&S. AUST. MUS. 16 (3): 1-219 July, LOT)

pattern, On Gamasiphis, Euepierius and Gamaysiphoides species there is

always at least one ventral less on genu TV (2, 5/1, 1) than in Gamraselliss,

OMeER CHARACTERS, Tectum anterior margin always more complex

than unispinate. Setae all and al2 on palp genu usually simple, although

muy be spine-like or spatulate, and in Caliphis, Gamaselliphis and Caliphis

seta a/l may have one or two small lateral prongs, On palp femur,

seta al always in proximal third unlike Rhodacarinae, Tangaroellinae and

some Sessiluncinae species, Setae are Usually simple and sclerotization shiny

on species with a holonotal shield, while in Euepicrius and Gamaselliphis

species the sclerotization is dull, being covered by a thick exudate, and always

some dorsal setae are complex. On female sterno-metasternal shield a line

joining seta sf2, 9/3 and st4 would enclose an angle of less than 95°, except

in Hydrogamasus species, while in some Gamayiphis (e.g. G. concilator )

and Caliphis (é.v., C. calvus) species seta sf3 is positioned so far forward

that it can be mistaken for seta s?2 und these sternal setae appear to be ina

straight line, Spermathecal ringed tube opens near posterior paraxial edge

of acetabulum TV and its junction with a single central sacculus is often quite

clear, If pretarsus [ present it is often pedunculate and similar in size to

other pretarsi, Pulvilli 1-[VY with paired lateral, short lanceolate lobes and

paired central, circular lobes except in Aydrogamasus (see Fig. 96),

DISTRIBUTION, None of the genera are restricted to the Northern

Hemisphere, while five of the genera are only found in the Southern Hemis-

phere. Excluding the discontinuous distribution of Hydregamasits, contain-

ing species which are found in the littoral zone, only Gamasiphis and

Gamasiphoides have a wide distribution and they have been collected from

all three of the Southern Hemisphere temperate regions (Neotropical,

Ethiopian and Australian),

REMARKS. | have confidence in my grouping of Gamasiphis, Caliphis,

Fuepicerius, Gamasiphoides and Laelaptiella as closely allied genera, although

a superficial examination can lead to some species of Gemasiphoides with

a discrete ventro-anal shield being confused with some species of Ologamasini.

Gamaselliphis species have characters that are unusually similar to those of

Gamaselluy species and, since the Gamasellus discutafus-complex contains

females on which the sternal setae are located us in Gamasiphinae females.

the placing of this genus as closely ullied to Gamasiphis is tentative, Hydro-

gamasus is not closely allied to the above genera, and is placed in the

Gamasiphinae in preference to any other subfamily.

| recognize 7 genera within this subfamily; Gamasiphiy, Caliphis,

Euepicrius, Gamaselliphis, Gamasiphoides, Hydrogamasuy and Laelaptiella.

Members of these genera can be distinguished by using the fotlowing key.

LEE—RHODACARIDAE

KEY TO GENERA OF GAMASIPHINAE

_ Tibia I with 3 ventral setae (2, 6/3, 2). On female

sterno-metasternal shield a line joining setae sf2,

st3 and st4 would be nearly straight and sterno-

metasternal shield not fused to endopodal IV

shield. Male seta st5 on ventro-anal and not

sternito-genital shield .

Tibia I with 4 ventral setae (2, 6/4, 2), On female

sterno-metasternal shield a line joining setae sf2,

st3 and st4 would enclose an angle of less than

95°, and if seta st3 positioned so far forward

that it can be mistaken for seta st2, then sterno-

metasternal shield is fused to endopodal IV

shield, Male seta st5 is on_ sternito-genital

shield ..

Separate podonotal and opisthonotal shields.

Always some complex dorsal setae; notal shields

dull set ie Migigint ae mpa® ahaa’ tw

Holonotal shield. seine! dorsal setae simple and

notal shield shiny .

Pretarsus I absent. Pre- sadopedeh shield fused | to

sterno-metasternal shield. Behind stigma, there

are 3 conspicuous pits opening along peritre-

matal ridge. Genu [V with I ventral seta

(2, 5/1, 1) 9 ei ebb tee -Pemcece BE

Pretarsus I present. Pre-endopodal shields in

single, discrete pair. Behind stigma, only one

conspicuous pit opening beside peritrematal

ridge near stigma. Genu IV with 2 ventral setae

(2, 5/2, 1). :

Pre-endopodal shields in 2 pairs (anterior one may

be fragmented). Either peritrematal shield

separate from exopodal IV shield or exopodal

ITI shield is split ..

Pre-endopodal shields in single pair, or if 2 pairs,

peritrematal shield fused to exopodal IV shield

and exopodal HI shield is not split . .

Palp genu three-times longer than broad. Peritre-

matal shield separate from exopodal IV shield

and exopodal III shield not split. Genu IV with

2 ventral seta (2, 5/2, 1)

Hydrogamasus

Euepicrius

Gamaselliphis

Laelaptiella

42 REC. S. AUST. MUS., 16 (3): L219 July, 1970

Palp genu less than two-times longer than broad.

Peritrematal shield fused to exopodal IV shield

and exopodal III shield split. Genu IV with 1

ventral seta (2, 5/1, 1) .. 2. 2. 2. 2... 2. Gamasiphoides

6, Peritrematal shield either separate from fused meta-

podal and exopodal IV shields or narrows behind

stigma before its fusion with enlarged triangular

metapodal which is usually fused to exopodal IV

shield. Exopodal II shield undivided. Genu TV

with 2 ventral setae (2,5/2, 1) ...... 0... Caliphis

Peritrematal shield broadens behind stigma to fuse

with the fused metapodal and exopodal IV

shields. Exopodal II shield split. Genu IV with

| ventral seta (2, 5/1, | or 2,5/1,0) .. 2. 2. ~=Gamasiphis

Genus GAMASIPHIS Berlese

Gamasiphis Berlese, 1904a, p. 261. Type-species: Gamasus pulchellus

Berlese, 1887, by original designation.

Micriphis Berlese, 1914, p. 140, syn.n. Type-species: Gamasiphis gama-

sellus Berlese, 19134, by monotypy.

Heteroiphis Tragardh, 1952, p. 55. Type-species: Gamasiphis (Heteraiphis)

arcuatus Tragardh, 1952, by original designation,

Neogamasiphis Tragardh, 1952, p. 57, syn.n. Type-species: Neogamasiphis

hamifer Trigardh, 1952, by original designation,

DIAGNOSIS. Minute to large mites, always with extensive. well

sclerotized shields and often the idiosoma strongly convex dorsally. Ventro-

anal shield fused to a holonotal shield. Peritrematal shield fused to exopodal

IV shield. Exopodal III shield never split. Sterno-metasternal shield fused

to endopodal [V shield. Female movable cheliceral digit has 4 teeth,

Pretarsus | usually subequal in size to other pretarsi, but may be smaller.

Genu IV with one ventral seta.

MORPHOLOGY.

SCLEROTIZATION, Female: Holonotal fused to ventro-anal, which

bears aciculae posterior to anus. Peritrematal fused to exopodal IV and to

a triangular metapodal posteriorly, Always split in exopodal II, sometimes

split in exopodal TV, but never split in exopodal III. Sterno-metasternal

fused to endopodals Il, HII and 1V. One or 2 pairs of pre-endopodals,

Male: Sternito-genital, otherwise as female.

CHAFTOTAXY. Idiosoma: 6), 6z, 4-Ss, Sr: 3-5/, 3-5Z, 3-SS, 3-5R,

O0-2UR: Sst: 34v, 3Zv, Sv.

LER—RHODACARIDAE 44

Leys: Following segments may (genu IV always differs) differ from

Gamasellus; genu HL (2, 4,1 or 2 as for Gumasellus, 1); genu iV (2, 5/1,

0 or | as tor Gamasellus): tibia [IV (1 ot 2 as for Gamasellus, 4/2, 1 or 2

as for Gamasellus). The species with only one antero-lateral seta on tibia

IV (lt, 4/2, 1) is unnamed and from the Nearetic region (Emberson,

thesis, 1968),

OTNER CHARACTERS, Female: Deutosternal denticles in 7 or fewer

horizontal rows, Tectum anterior margin basically trispinate, but it may

be complicated by spinules and central spine may also be spatulate,

Movable cheliceral digit with 4 teeth. On palp genu, setae al] and al2

spine-like, Dorsal setae simple or rarely with hyaline flaps making them

spatulate or lanceolate. Shields shiny and often reticulated, Spermathecal

ringed tube opens near posterior paraxial edge of acetabulum IV, passing

back to single central sacculus, Pretarsus I usually similar in size to other

pretarsi, but may be conspicuously smaller. Amongst dorsal setae on tarsus

IV, setae ad2, pd or pd4 may be the longest.

Male: Corniculi attenuated and long compared with female. Movable

cheliceral digit with one tooth, fused at base to spatulate spermadactyl that

is subequal in length, or up four times as long. On leg IL, femur seta av

and sometimes seta pvl, genu seta av and sometimes seta py, tibia seta ay,

enlarged into spur or stout spine, On tarsus IT, seta av3 may be on sinall

tubercle.

DISTRIBUTION, Ne, Na: NTa, NTb, NTc: Ew, Es, Em: Pm, Pe:

Oi, Os, Om: Am, Aa, Ap, An. Besides the locality records published with

the original descriptions of nominal species there are records of Gamasiphis

species from North America (Emberson, thesis, 1968; Pearse, 1946:

Rodriguez and Ibarra, 1967), South Africa (Loots, thesis, 1967), Pyrenees

in France (Triigirdh, 1912), Israel (Costa, 1966a), India (Bhattacharyya,

1968) and Hawaii (Butler and Usinger, 1963). Specimens from many

other regions, Jamaica (NTa), Trinidad (NTb), British Guiana (NTD),

Argentina (NTe), Annobon Island (Ew), Seychelle Islunds (Em), Nepal

(Os), Malaya (Om), Borneo (Om) and Solomon Islands (Am), are before

me und to be dep. BM(NH), while yet others dep. SEAF are listed under

G, pulchellus. The genus is distributed right round the tropics, is less

common in temperate regions, and has not been found north of 50°N, or

south of 50'S,

Found amongst moss, plant litter and upper soil layers. A single female

was found on a rat in Puerto Rico (Fox, 1949).

REMARKS. Gamasiphis has sometimes been incorrectly associated

with Ologamasus as Mentioned below under that genus. Three subgenera,

Periphis, Epiphis and Megaliphis, which were established within Gamasiphis,

| here transfer to Sp/ochirus, a genus in the Sessiluncinac. The synonymy

44 REC. 8. AUST. MUS., 16 (3); 1-219 July, 1970

Figs. 23-31. Garmasiphis pulchelluy (Berlese).

23-27, female: 23, idiosoma, venter; 24, leg IV (part), dorsum: 25, tectum: 26.

gnathosoma, yenter; 27, pretarsus I and tarsus 1 (part). 28-31. male: 28, corniculus:

29, leg IL (part), antero-latus; 30, chelicera: 31, idiosoma, venter.

LEE—RHODACARIDAE 45

Vie kg

Figs. 32-39. Gamasiphis Berlese.

32-34, G. gamasellus Berlese, males 32, idiosoma, venter; 33, gnathosoma, venter:

34, leg IL (part), antero-latus. 35-37, G. concilator Berlese, female: 35, tectum:

36, idiosoma, venter; 37, setae all and alZ on palp genu. 38 and 39, G. pilosellus

Berlese, female: 38, stigma and surroundings; 39, anus and surroundings. 40 and

41, G. “pulchellus” from Java: 40, stigma and surroundings; 41, seta SS,

REC. 8S. AUST. MUS., 16 (3): 1-219

July, 1970

0-20,

is]

—

“9—__» 0-0,

w_—

(

Figs. 42-50. Gamasiplis australicus Womersley.

42-45. female: 42, soma, dorsum: 43, idiosoma, yenter; 44. gnathosama. venter: 45.

leg IV (part), dorsum, 46-50, male: 46, idiosoma. venter; 47, idiosoma. latus:

48, leg IT (part); 49, chelicera; 50, corniculus,

LEE—RHODACARIDAE

i=}

“

tH.

ont

u part), dorsum; 53, gnathosoma, venter:

1 soma, venter.

48 REC, S. AUST. MUS., 16 (3): 1-219 July, 1970

55-58, male: 55, idiosoma, venter; 546, Jeg IT (part), antero-latus; 57, chelicera:

58, corniculus. 59, female, idiosoma, latus. 60 and 61, deutonymph: 60, idiosoma,

dorsum; 61, idiosoma, latus.

LERE—RHODACARIDAE 49

of Heteroiphis with Gamasiphis was established by Ryke (1962b). and

agreed with here, since G. arcuatus is very like G. setosus, although consider-

ably larger. The synonymy of Micriphis and Neogamasiphis with Gamasiphis

is new, and made because the characters claimed by Berlese (1914) and

Trigardh (1952) as distinguishing the type-species of these genera from

the type-species of Gamasiphis are in fact similar and no other suituble

distinguishing characters have been found. Womersley (1956a) treated

Neogamasiplis as a subgenus of Gamasiphis and placed species in CG,

(Neogamasiphis) that are here either retained in Gamasiphis or transferred

lo a new genus, Caliphis, The following 16 nominal species are, therefore,

included in this genus: G. pulchellus (Berlese, 1887a); G. arcuatits

Trigardh, 1952; G. australicus Womersley, 1956a; G. bengalensis

Bhattacharyya, 1966; G, conciliator Berlese, 1916a; G. elegantellus Berlese,

1910a: G, elongatellus Berlese, 1910b; G. femeralis (Banks, 1916): G, forni-

cals spam, Gr. gamasellus Berlese, 1913a; G. hamifer (Triighrdh, 1952):

G. illotus Fox, 1949, G, pilosellus Berlese, 1913a; G. preductelluy Berlese,

1923; G. setosus Womersley, 1956a; G, wncifer Trigardh, 1931. Two

unnamed species are described by Loots (thesis, 1967), Two unnamed

species are listed by Emberson (thesis, 1968) and the unusual leg setution

of one of these species is described.

Gamasiphis pulchellus (Berlese)

Gamasus pulehellus Berlese, 1887a, XXXIX, 4.

FEMALE. Fig. 23-27. Idiosomal length, 410». Dorsal idiosoma

not clear enough to be drawn. Most dorsal setae seen, but not drawn, appear

to be similar to seta Zv3, except the humeral setae which ure more like seta

R5. Luteral suture (using Triigardh’s (1952) term in describing Heteroiphis

arcuafus, although this line is not a break in the shield, but a thickened ridge

beside a line of weak sclerotization, similar to a strong reticulation line)

follows fissure at edge of holonotal shield, Chuctotaxy differs from

Gamasellus on the following leg segments: genu HE (2, 4/1, 1): genu TV

(2, 5/1, O); tibia IV (2.4/2, 1), On some specimens, setic ad! and pd

on tarsi I-IV appear to have hyaline flap.

MALE. Fig. 28-31. Idiosomal length, 360z.

LOCALITY. Female (29/39) and male (29/38), that appear to be

conspecific with the type specimens, were drawn! Italy; leaves, Castions di

Strada, dep. SEAP.

Female, 2 males and a deutonymph on one slide (8/10. ‘tipico’) which,

because some of their characters were obscured, were only examined: ltaly;

humus, Botanic Gardens, Padova, dep. SEAF.

Gamasiphis specimens labelled G. pulehellus from North America

(103/17), South America (103/18, 103/19 and 151/34) and South Africa

SO REC, 8, AUST, MUS... 16 (3): 1-219 July. 1970

(103/20, 103/21, 136/24, 136/35); specimens labelled ‘speciosellus’ (64/7,

140/18, 147/14, 221/12) from Japan; and G. elongatellus (130/6, 136/20,

136/23, 148/21) from Java and G. productellus (221/14) from China,

dep. SEAF, examined and all considered here to be similar to G. pulchellus.

A careful study of these specimens is needed before making a decision us to

which of them belong to G, pulehellus and which, if any, should belong to

different species.

Gamasiphis australicus Womersley

Gamasiphis (Heteroiphis) australicus Womersley, 1956a, p. 321.

FEMALE. Fig. 42-45. Idiosomal length, 360. The lateral suture

is longer than as described by Womersley (1956a), since it contimues

posteriorly beside the ventro-holonotal fissure, and, after the latter stops just

behind seta Svl, it continues on to behind seta Zv3. In the drawing of the

dorsum (Fig. 42) the lateral suture is shown merging with the only reticula-

tion line drawn from amongst many such lines; this is not a fissure separating

the holonotal shield into two parts, Chaetotaxy differs from Gamasellus on

the following leg segments: genu IIL (2, 4/1, 1); genu TV (2. 5/1, 0),

MALE. Fig. 46-50. Idiosomal length, 350...

LOCALITY. ‘Three females (NI9G886-NI96888) and 2 males

(N196889-NI96890) drawn or examined: Australia; LFSS8, grass and leaf

litter under Eucalyptus camaldulensis, Heywood Park, South Australia,

6.6.1965, col. R. V. Southcott, dep. SAM.

The holotype female (NI96891) examined: Australia; moss, Mylor,

South Australia, 27,6,1948, col, G, F. Gross, dep. SAM.

REMARKS. Gamasiphis australicus: Domrow, 1957 is more like the

female G. setosus (male is a dermanyssid) than this species, the laterai

sulures being similarly placed, although Womersley (1956a) does not record

their presence. Therefore, | regard the Gamasiphisy specimens described by

Domrow (1957) as being G. setosus in preference to any other nominal

species.

Gamasiphis concilator Berlese

Gamasiphis (Periphis) concilator Berlese, 1916a, p. 159.

FEMALE. Fig. 35-37. Idiosomal length, 910». In this species seta

st3 is so far forward that it is anterior to seta s72 and may be mistaken for it,

MALE. Not known.

LOCALITY. The ‘tipico’ female (175/42) drawn: New Caledonia;

500 m.. on Mt, Panié, col. Sarrasin and Roux, dep. SEAF.

REMARKS. This species is a typical Gamasiphis and clearly not

congeneric with the type-species of Periphis which is here transferred to

Stvlochirus in the Sessiluncinae.

LEE—RHODACARIDAE Sl

Gamasiphis fornicatus sp.n.

FEMALE, Fig. 51-54 and 59, Idiosomal length, 500p. Genital

shield has a group of 4 oval depressions on it and the ventro-anal shield is

reticulated. There is a large pore-like structure, with a raised flange on its

posterior rim, which may be a modified seta S4 (Fig. 59). The drawings

of the lateral surface of the idiosoma (Fig. 59 and the deutonymph in Fig.

61) are of squashed specimens and what is drawn ventral to setal rows

rand § would not be visible normally since the venter is flat. Chaetotaxy

differs from Gamasellus on genu IV (2, 5/1, 1).

MALE. Fig. 55-58. Idiosomal length, 490,.

DEUTONYMPH. Fig. 60 and 61. Idiosomal length, 450». The

idiosomal setation has been drawn in attempt to make the labelling of the

lateral notal setae more accurate. It supports the above suggestion about

the modification of seta 84, but the labelling of setal rows, r, R and UR is

sull uncertain.

LOCALITY. The holotype female (N196892), allotype mule

(N196893) and morphotype deutonymph (N196894) drawn, and 6 para-

type females (N196895-N1968100), 3 paratype males (N1968101-

N1968103) and one paratype deutonymph (N1968104) examined:

Australia: LF 142, moss amongst bracken and gum trees in gully, Mt.

Remarkable, South Australia, 9.8.1966, col. H. M. Cooper, dep. SAM.

REMARKS. This unusually globular species with its long dorsal setae

and long spermadactyl is easily distinguishable from other nominal species,

Gamasiphis gamasellus Berlese

Gamasiphis gamasellus Berlese, 1913a, p. 80.

FEMALE, Not known.

MALE. Fig. 32-34. Idiosomal length, 3102, Leg chaetotaxy as

G. pulchellus. Dorsal setae on leg TV with similar lengths relative to each

other as the equivalent setae on G. australicus (Fig. 45), except that seta

ad2 on genu IV is as long as the genu.

LOCALITY. The ‘tipico’ male (144/11) drawn: Java; humus.

Samarang, dep. SEAP.

REMARKS. This species is the type. by monotypy, of Micriphis

(initially a subgenus of Ologamasus, then Ologamasellus), but is clearly

congeneric with G. pulchellus.

Gamasiphis pilosellus Berlese

Gamasiphis pilosellus Berlese, 1913a, p. 81.

FEMALE, Fig. 38 and 39. Idiosomal length, 590. As on

Gamasiphis elegantellus Berlese, 1910a, the peritrematal shield is not

separated from the exopodal shields posterior to stigma, in contrast to

52 REC. $8. AUST. MUS, I@ (3); 1-219 July, 1970

Berlese’s drawing (1913a, Fig. 27) of the latter. Other similarities between

these species are also closer than suggested by Berlese’s drawings. The lateral

suture nearly reaches to the anterior end of the peritrematal-holonotal fissure

(Fig. 38) and seta R5 is nearly as short as the paranal setae (Fig. 39).

An unnamed species (labelled G. pulchellus, male (37/34), from Java, dep.

SEAF) has a lateral suture which extends anteriorly almost as far as in this

species and also a dorsal branch (Fig. 40) similar to the anterior end of

the lateral suture on G. australicus. This unnamed species also has a short

seta R5 and those dorsal setae which are relatively long (e.g,., seta S5,

Fig, 41) have a hyaline flap at the tip. Leg chaetotaxy as G. pulchellus,

MALE. Not figured, Idiosomal length, 490, Corniculi, sperma-

dactyl and leg IT (even seta av3 of tarsus on small tubercle) are similar to

those of G, pulehellus,

LOCALITY. The ‘tipico’ female (136/17), and “tipico’ male

(136/14) drawn or examined (other four specimens in collection are

difficult to observe): Java; humus, Samarang, dep, SEAPF.

Genus CALIPHIS gen.n,

Type-species: Caliphis calvus spn.

DIAGNOSIS, Small to large mites, always with extensive, well

sclerotized shields and idiosoma which is strongly convex dorsally, Ventro-

anal shield fused to a holonotal shield, Peritrematal shield either separate

from fused metapodal and exopodal [V shields or narrows behind stigma

before its fusion with an enlarged triangular metapodal, which may or may

not be fused to exopodal IV shield. Exopodal LI! shield never split. Sterno-

metasternal shield fused to endopodal IV shield. Fetmale movable cheliccral

digit has 3 teeth, Pretarsus I subequal in size to other pretarsi. Genu IV

with 2 ventral setae,

MORPHOLOGY,

SCLEROTIZATION, Female: Holonotal fused to ventro-anal, which

bears aciculae posterior to anus. Peritrematal usually fused to combined

exopodal IV and metapodal, but peritrematal may be separate (C. hickmani

and C. queenslandicus) or exopodal TV may be separate (C. tamborinensis).

Exopoduls H, HT and 1V fused into continuous strip. Sterno-metasternal has

a short axis, but laterally a narrow strip extends a long way backwards

fusing with endopodals II, I] and IV. Single pair of pre-endopodals.

Male: Sternito-genital, otherwise as female.

CHARTOTAXY. Idiosoma: 6j, 6z, Ss, Sr or, as for the new species

described here, podonotum is hypertrichous; opisthonotum apparently always

hypertrichous although there may be us few as 48 setae: Sst: 3/v, 3Zv, 2Sv,

Legs: As Gamasellus.

LEE—RHODACARIDAF 3

OTHER CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows. Tectum anterior margin basically trispinate but there may be

spinules between the spines. Movable cheliceral digit with 3 teeth. On

palp genu, seta a/l with single lateral prong. both setae a/] and al2 slightly

lanceolate or spatulate. Dorsal setae tapering and simple. Shields shiny

and covered with reticulations. On sternal shield. seta st3 may be placed

anterior to sf2. Spermathecal ringed tube opens near posterior paraxial edge

of acetabulum TV, passing back to single central sacculus, Pretarsus 1 sub-

equal in size to other pretarsi. Amongst dorsal setae on tarsus TV, seta pd3

is the longest.

Male: Corniculi proportionately slightly longer than in female.

Movable cheliceral digit with single tooth and fused along proximal half

with sinuous spermaductyl which is subequal in length or up to twice as long,

On leg I, femur setae av and pvl, genu seta av and tibia seta av enlarged

intO 4 spur or spine.

DISTRIBUTION. Aa, An: Sa. All published records are with the

original descriptions of nominal species. Specimens have been collected

from Lord Howe Island, dep. SAM. Found in plant litter and moss.

REMARKS. Caliphis is similar to Gamasiphis, but can be distinguished

from it as indicated in the key to Gamasiphinac. The following 6

nominal species are included in this genus: Caliphis calvas spsn., Caliphis

hickmani (Womersley) comb.n. for Gamasiphis (Neogamasiphis) hickmant

Womersley, 1956a, Caliphis novaezelandiae (Womersley) comb.n, for

Gamasiphis (Neogamasiphis) novae-zelandiae Womersley, 1956a, Caliphiy

queenslanidicus (Womersley) comb.n. for Gamasiphis (Neogamasiphis)

queenslandicusy Womersley, 1956a: Caliphiy sehusteri (Hirschmann)

comb.n. for Gamasellus (Hydrogamasellus) schusteri Hirschmann, 1966;

Caliphis tamborinensis (Womersley) comb.n, for Garnasiphis (Neogama-

viphts) hickmani var. tamborinensis Womersley, 19564,

Caliphis calvus spn,

FEMALE, Fig. 62-64, 69. [diosomal length, 430». — Idiosoma

covered in reticulations as figured for male (Fig, 65). The spermathecal

ringed tube and anterior edge of sacculus can be clearly seen and are shown

by dotted lines in drawing of venter (Fig. 64),

MALE, Fig, 65-68, Idiosomal length, 410).

LOCALITY. The holotype female (N1968231), allotype male

(N1968232), 19 paratype females (N1968233 and N1968235-N1968252)

and 3 paratype males (N1L968234, N1968253, N1968254) drawn or

examined: Australias LF 191, moss, Lena Valley Track, 8OOft. Mt.

Wellington, near Hobart, Tasmania, 14.12.1966, col. L. W. Miller, dep.

SAM.

54 REC, 8S. AUST. MUS., 16 (3): 1-219 July, 1970

Figs. 62-69. Caltplis calvus sp.n..

62-64 and 69, female; 62, soma. dorsum; 63, leg 1V (part), dorsal selae only; 64,

idiosoma. venter: 69, gnathosoma, yenter, 65-68. male: 65, idiosoma, yenter;

66, leg Ul (part), antero-latus; 67, chelicera; 68, corniculus.

LEE—RHODACARIDAR 55

REMARKS, The large number (more than 150) of uniformly short

dorsal setae distinguishes this species from other nominal species, and it

also differs from C. hickmani, C. queenslandicus and C. tamborineusis iw

having a peritrematal shield fused to a combined exopodal [V and metapodal

shield.

Genus EUEPICRIUS Womersley

Evepicriuy Womersley. 1942, p. 170. Type-species: Euepicrius filamentosus

Womersiey, 1942, by original designation.

DIAGNOSIS. Small to average sized mites, always with extensive, well

sclerotized shields and idiosoma which is strongly convex dorsally. Ventro-

unal shield fused to the opisthonotal shield which is separated from the

podonotal shield. Peritrematal shield is fused to all the exopodal shields,

which are themselves fused into a strip of undivided shield. Sterno-

metasternal shield fused to both endopodal shield 1V and the pre-endopodal

shields. Leg I long and slender with long setae distally on tarsus. Female

movable cheliceral digit has 3 teeth. Pretarsus I is absent, Genu IV with

one ventral seta.

MORPHOLOGY,

SCLPROTIZATION. Female: Separate podonotal and opisthonotal,

ihe latter is fused to ventro-anal which bears aciculae posterior to anus.

Peritrematal appears to be fused to the complete strip of undivided exopodals

and to a triangular metapodal posteriorly. Sterno-metasternal fused to

endopodals IH, IIT and ['V as well as. to. pre-endopodals.

Male: Sternito-genital, otherwise as female.

CHAETOTAXY. Idiosoma: Hypertrichous on notum: Sst: 34, 324,

38\',

Lees: As Gamuasellus except that there is only one ventral on genu IV

(2,5/1. 1).

Orner CHARACTERS. Female: Deutosternal denticles in § horizontal

rows. Tectum anterior margin basically trispinate with spinules between

lateral spines. Movable cheliceral digit with 3 teeth. On palp genu,

setae a/l und a/2 spine-like or slightly lanceolate. Gnathosoma has distinct

lateral notch and tubercle. Dorsal setae may be simple and tapering, but

usually, either sinuous and slightly spatulate or straight and pilose with clearly

spatulate tip. Shields ridged, pitted and dull. being covered by a thick

exudate. Peritreme with lateral pockets and corrugated ridge outside it,

which runs to posterior edge of peritrematal shield and borders 3 conspicuous

pores behind stigma. Spermathecal ringed tube opens near posterior paraxial

edge of acetabulum IV, passing back to a single central sacculus. Leg I

long and slender, tibia may be up to 16-times longer than its breadth. Tip

56 REC. S. AUST, MUS,. 16 (3): 1-219 July. 1970

Figs. 70-78. Euepicrius lootsi sp.n..

70-72. 74, 77 and 78, female: 70, soma, dorsum: 71, idiosoma, latus: 72, idiosoma.

venter; 74, tarsus [V, dorsum; 77, tarsus 1; 78. gnathosoma. venter. 73, 75 and 76,

male: 73, idiasoma, venter: 75, chelicera: 76. leg IL (part), antero-latus.

LEE—RHODACARIDAE 57

of tarsus 1 with long setae but no pretarsus, Amongst dorsal setae on tarsus

IV, the slighdly spatulate seta pd3 is the longest.

Male; Movable cheliceral digit with one tooth, fused along its entire

length to spatulate spermadactyl which is longer, and may be up to 4-times

the length of the digit. On leg HH, femur seta av, genu seta ay, tibia seta av

enlarged into spurs,

DISTRIBUTION, Aa, An: Sa. Besides the locality records published

with the original descriptions of nominal species there are specimens before

me from Auckland and Campbell Islands (Sa) to be dep, BBM, and Lord

Howe Island and New Caledonia, dep. SAM.

Found in moss and plant litter,

REMARKS. Euepicrius was originally placed in the Macrochelidae,

1 (Lee, 1966) transferred it to the Rhodacaridae, but was uncertain as to

which genera it was related. I now consider it to be quite a close ally to

Gamasiphis despite the differences in superficial appearance, Members of

Enepicrius have a very characteristic appearance, so there ts no doubt that

the new species described here is congeneric with the type-species. The

following 3 nominal species are included in this genus: E&. filamentosus

Woniersley, 1942; &, looisi span; E, queenslandicus Womersley, 1956b.

Euepicrius lootsi spn.

FEMALE. Fig. 70-72, 74, 77, 78. Idiosomal length, 610n. This

species has an unusually large number of idiosomal setae, the number on

the podonotum and opisthonotum in Fig. 70 indicates the number of setae

in one half of these regions, excluding those on the centre line, Only part

of three long setae on distal end of tarsus I are drawn (Pig. 77).

MALE. Fig. 73, 75, 76. Idiosomal length, 550).

LOCALITY. The holotype female (N1968138), allotype male

(N1968139) and 4 paratype females (N1968140-143) and 3 males

(N1968144-N1968146) drawn or examined: Australia; LF127, moss and

grass beside Wannon River, near Yarram Gap, Grampians, Victoria,

14,5.1966, col. D. C, Lee, dep. SAM.

REMARKS. This species can be distinguished from the other two

nominal species by the shortness of the two terminal pairs of spatulate setac

on the opisthosoma, which are Jess than three-times the length of the anal

valves, and the relatively short spermadactyl, the free part of which is less

than two-times the length of the movable digit and is straight, having no

bends us in the other nominal species.

Genus GAMASELLIPHIS Ryke

Gamaselliphiy Ryke, 1961a, p. 99. Type-species: Cyrtolaelaps (Gama-

selliphis) potchefstroomensis, Ryke, 1961a, by original designation.

58 REC. & AUST. MUS., 16 (3): 1-219 July, VY70

DIAGNOSIS. Minute to average sized mites, always with extensive,

well sclerotized shields and idiosoma which is fairly convex dorsally. Ventro-

anal shield fused to opisthonotal shield which is separated from the podonotal

shield. Peritrematal shield fused to exopodal TV shield. Exopodal II shield

split. Sterno-metasternal shield not fused to endopodal TV shield. Female

movable cheliceral digit has 3 teeth. Pretarsus | smaller than other pretarsi.

Genu IV with 2 ventral setae.

MORPHOLOGY.

SCLEROTIZATION. Female; Separate podonotal and opisthonotal, the

latter is fused to yentro-anal, which bears aciculae posterior to anus.

Peritrematal fused to exopodal IV and to a triangular metapodal posteriorly.

Always a split in exopodals Uf, II and ITV. Sterno-metasternal fused to

endopodals IT and III. One pair of pre-endopodals.

Male; Sternito-genital, otherwise as female,

CHAETOTAXY, Idiosomu: 6j, 62, 5s, 5h: SJ, 57, 5S, SR, 0-2UR:

Sst: 3Jv, 3Zy, 28v.

Legs: As Gamasellus,

OTHER CHARACTERS, Female: Deutosternal denticles in 7 horizontal

rows, Tectum anterior margin is trispinate. Movable cheliceral digit

with 3 teeth. On palp genu, seta a/l spatulate with 2 small lateral prongs

and seta a/2 is lanceolate. Dorsal setae on idiosoma and legs rarely

simple, usually at least lanceolate, may be pilose as well, or spatulate and

pilose. Shields dull being covered by thick exudate. Single pore, which is

conspicuous in one specimen (N1968275), opens beside peritrematal ridge,

level with stigma, and may homologous with one of the three similarly

placed pores on Evepicrius species. Spermathecal ringed tube opens near

posterior paraxial edge of acetabulum IV, passing back to single central

sacculus. Pretarsus I smaller than other pretarsi especially in one species

(G. grahamstowni) which has an unusually long leg I, with long distal setae.

Amongst dorsal setae on tarsus [V, seta pd3, which is setose or lanceolate,

is the longest.

Male: Movable cheliceral digit with one tooth and fused at base to

sinuous spermadactyl, which is slightly longer or up to four times as long as

digit. On leg Ul, femur seta av, genu seta av, tibia seta av enlarged into

spurs, Genu and tibia II setae py may be stubby spines,

DISTRIBUTION. Ew, Ee. Es, All the published records are with

the original descriptions of nominal species from South Africa. Records

from Congo, Kenya, Rwanda and Tanzania are from Loots (thesis, 1967).

LEE—RHODACARIDAE 59

The males of Gamaselliphis recorded from Nobska Beach, Massachusetts,

U.S.A. (Na) by Hag (1965) are Parasitus species according to Emiberson

(thesis, 1968),

Found in plant litter.

REMARKS. Gamaselliphis was established as a subgenus of Cyrto-

luelaps by Ryke (1961a), Because of the extensive sclerotization without

fusion of the broad peritrematal to the ventro-anal shield on Gamaselliphis

males, as well as other characters, | am now treating this taxon as a genus

and consider it most like Evepicrius. Species in this genus have a lot of

churucters similar to those of some Gamasellus discutatus-complex species

whose females have the sternal setae located as in Gamasiphinae females.

Therefore, the alliances I have indicated for this genus depend on the

importance | have given to the lack of sexual dimorphism of the ventral

sclerotization. L have examined three non-type females of this genus dep,

SAM,: one G, potchefstroamensis (N1968275) and two similar to G.

montanellus (N1968276 and N1968277) from South Africa. The following

5 nominal species are included in this genus; G. potchefstroomensis Ryke,

1961a; G. cathkini Ryke, 196la; G. grahamstowni Ryke, 1961a; G.

lawrencei Ryke. 19614; G. montanelluy Ryke, 1961a. Also, | have examined

the specimen with the manuseript name “Laelogamasus inguinalis”, slide

222/16, Eust Africa, dep, SEAF, and confirm Ryke’s (1962c, p. 51)

placing of it in this genus but have not established whether or not it belongs

to &@ noniinal species,

Genus GAMASIPHOIDES Womersley

Gamasiphoides Womersley, 1956a, p. 528. Type-species: Gamasiphis

(Gamasiphoides) prepingua Womersley, 1956a, by original designation,

PIAGNOSIS. Small to average sized mites, often with extensive, well

sclerotized shields. The ventro-anal shield may or may not be fused to a

holonotul shield. The peritrematal shield fused to exopodal IV shield,

The exopodal II] shield is split. The sterno-metasternal shield rarely fused

to endopodal 1V shield. Female movable cheliceral digit has 3 teeth,

Pretarsus I usually subequal in size to other pretarsi, but may be smaller,

Genu IV with one ventral seta,

MORPHOLOGY,

SCLEROTIZATION. Female: Holonotal, which may or may not be fused

io ventro-unal that bears aciculae posterior to the anus. —Peritrematal

fused to exopodal TY and only sometimes is there a posterior triangular

metapodal extension of this merged shield, There is often a discrete, sub-

circular shield posterior to acetabulum TV which Womersley (1956a) terms

Ag REC, S&S. AUST. MUS., 16 (3): 1-219 July, [970

the metapodal, Exopodal 1, ILL and IV split. Sterne-metasternal fused to

endopodals IH and Il, and rarely to TV. Two pairs of pre-endopodals,

anterior pair may be fragmented,

Male! Sternito-genital. Subcircular shield posterior to acetabulum IV

is merged in ventro-anal. Otherwise similar to female of same species.

CHAETOTAXY. Idiosoma: Variable, usually setae on striated cuticle

between holonotal and ventral shields, and between levels of setae yt4 and

Zy2, and these are included in peripheral dorsal rows, although they might

be better regarded as a separate row. 6/, 5 or 6z, 58, 5 or 6; 4 or 5J, 4 oF

54, 4or 58, 4, 5 or 8R: Sst) 3/v, 327, 1Sv.

Leys: Most Australian species differ from Gamasellus only by having

one less ventral on genu TV (2, 5/1, 1). But, G, propingqua also has no

pd4 on tarsus LV, and an unnamed Gamusiphoides female from Tasmania

(dep. SAM, N1968280) differs on genu ITI (2, 4/1, 2) and tibia ILI (2, 3/2,

2) in having an extra postero-lateral, African and South American species

differ from Gamasellus in having one less ventral on genu HI (2, 4/1, 1)

and genu [IV (2, 5/1, 1), The species from the Subantarctic regions may

have a ventral seta less on genu IV or on genu IIL and IV,

OTHER CHARACTERS. Female: Deutosternal denticles in 8, 9 or 10

rows, Tectum anterior margin is trispinate, Movable cheliceral digit with

3 teeth, On palp genu, setae a/l and a/2 spatulate. Dorsal setac simple

and tapering. Shields shiny and often reticulated, Spermathecal ringed tube

Opens near posterior paraxial edge of acetabulum IV, passing back to single

central sacculus, Pretarsus | usually similar in size to other pretarsi, but

may be conspicuously smaller. Amongst dorsal setae on tarsus [V, seta

pd3 is the longest.

Male; Palp femur has small tubercle in one species (Fig. 91),

Movable cheliceral digit with one tooth, and fused along proximal half to

slightly longer, spermadactyl. On leg If femur seta av and sometimes seta

pv, genu seta av, tibia seta av and tarsus seta av2 enlarged to spines or spurs,

DISTRIBUTION, NTc: Es: Aa; Sim. Sa. Besides the locality records

published with the original descriptions of nominal species, two unnamed

species from South Africa were described by Loots (thesis, 1967), and

specimens have been collected from Tierra del Fuego, dep. BM(NH) and

from Lord Howe Island and New Caledonia, dep. SAM.

Found in moss and plant litter,

REMARKS. ~~ Gamasiphoides was originally a subgenus within

Gamasiphis but it was given the rank of genus by Ryke (1962b). An

unnamed species from Victoria, Australia, dep, SAM (female, N1968282:

male, N1968283), has more extensive shields and is similar to Gamasiphis

species in having a female metasternal shield that is fused to endopodal 1V

LEE—RHODACARIDAE 61

o— O05. .

Figs, 79-85. Gamasiphoides propingua Womersley.

7U¥-81 and 85, female: 79, soma, dorsum; 80, leg ITV (part), dorsum: &1, idiosama,

venler; 85, enathosoma, venter. 82-84, male: 82. idiosoma, venter; 83, leg IL (part);

84, chelicera.

62 REC. S. AUST. MUS.. 16 (3): 1-219 July, 1970

(eo)

S oO %

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os aes

Figs. 86-92. Guimasiphoides aitkeni sp.

86-88 and 92, female: 8&6, soma, dorsum; 87, leg iv Coats dois sum; 88, idioso soma,

venter; 92, guathescyies venter, 89-91, male: 89, idiosoma, venter; 90, leg IL

(part); 91, chelicera, plus palp trochanter and femur, venter,

LEE—RHODAC ARIDAE, 63

shield, but because of the split exopodal IV shield, the three teeth on the

movable digit of the female chelicera and the short, non-spatulate sperma-

dactyl, it is placed in this genus. Another unnamed species from Tasmania,

Australia, dep, SAM (female, N1968281) is mentioned above because of

its unusual leg chaetotaxy. Gamasiphoides species with a reduced sclerotiza-

tion may be mistaken for members of the Ologamasini when superficially

examined. The following 4 nominal species are included in this genus:

G. propingua Womersley, 1956a; G. aitkeni sp.n.; G. gamasiphioides (Sheals )

comb.n. for Hydrogamasus gamasiphioides Sheals, 1962; G. macquariensis

(Hirschmann, 1966) comb.n. for Gamasellus (Hydrogamasellus) macquart-

ensis Hirschmann, 1966. Two unnamed species are described by Loots

(thesis, 1967),

Gamasiphoides propingua Womersley

Gamasiphis (Gamasiphoides) propinqua Womersiey, 1956a, p. 528.

FEMALE. Fig, 79-81, 85. [diosomal Jength, 780), On dorsum, seta

j\ ts so sniall that it is difficult to discern. On tarsus IV, seta pd4 is absent

(Fig. 80), and leg chaeotaxy also differs from Gumasellus on genu IV

(2, 9/1, 1)

MALE. Fig. 82-84. Idiosomal length, 760..

LOCALITY, Female (N1968151) and male (NL968152) drawn:

Australia; LFI6, rotting wood and lichen, near Mt, Lofty summit, South

Australia, 20.11.1964, col. G. F. Gross, dep. SAM. The paratype female

(N1968153) and paratype male (N1I968154) examined: Australia; moss,

National Park, Belair, South Australia, 9.1938, col. H. Womersley, dep. SAM.

Gumusiphoides aitkeni sp.n.

FEMALE. Fig. 86-88, 92, Idiosomal length, 420», Leg chaetotaxy

differs from Gamasellus on genu TY (2, 5/1, 1).

MALE, Fig. 89-91. Idiosomal length, 400¢. Small process on

antero-ventral edge of palp femur (Fig. 91). Ventro-anal shield not fused

to notal shield although this is not clear in Fig. 89.

LOCALITY. The holotype female (N1968155), allotype male

(NI968156), paratype female (NI968157) and paratype male

(N1968158) drawn or eXamined; Australia; LF36, moss and litter, under

shrubs at top of LOOft, elf near Glenelg River, near Nelson, Victoria,

28.1,1965, col. P. F. Aitken and N, B. Tindale, dep. SAM.

REMARKS. The only other nominal species in this genus on which

the ventro-anal shield is separate from the holonotal shield is G. macguariensis,

and specimens to be dep. BBM and apparently belonging to that species have

64 REC, 8. AUST, MUS., [6 (3): 1-219 July, 1970

only one ventral seta on genu ill, while on this species there ure two ventral

setae on genu III, Also G, macquariensis is a much bigger species, the

female idiosomal length being about 760,,

Genus HYDROGAMASUS Berlese

Hydrogamasus Berlese, 1892b, LXV, 5. Type-species: Gamasus littoralis

G, & R, Canestrini, 1881, designated by Vitzthum, 1941, p. 756 as the

junior synonym of Gamasus salinus Laboulbéne, 1851, a synonymy which

is revoked below.

DIAGNOSIS. Small to large mites with shields not so extensive and

dorsal idiosomal surface not so convex as most members of this subfamily.

Ventro-anal shield fused to holonotal shield. Peritrematal shield usually

fused to exopodal IV shield. Exopodal II shield continuous, but has a

longitudinal split. Sterno-metasternal shield not fused to endopodal IV

shield, Female moyable cheliceral digit has 3 teeth. Pretarsus [ smaller

than other pretarsi, Setation of tibia | (2, 6/3, 2) differs from Gamaselltis

in having only three ventral setae.

MORPHOLOGY,

SCLEROTIZATION, Female: Holonotal, fused to ventro-anal posterior

to seta Svl. Peritrematal usually fused to exopodal IV in varying

degrees (Hirschmann, 1966, draws H, /ittoralis, using the revoked synonymy

H. salinus, without such a fusion) and to a triangular metapodal posteriorly,

Exopodals HW and IIL not split, but in Palaearctic species the broad shield

is incised parallel to part of the edge of acetabulum TI]. Exopodal IV is

split. Intergenito-ventral is present. Sterno-metasternal fused to endopodal

Hand part of If, Two pairs of pre-endopodals.

Male; Sternito-genital fused to endopodals Il, U1] and IV or only to

endopodal If und part of III. Since seta sf5 is on ventro-anal, part of

sternito-genital might be regarded as being represented in this shield.

Otherwise similar to female.

CHAETOTAXY, Idiosoma: 6/, Sz, 48, Sr: 5/, 52, 5S, 1-3R! Sor:

a/v, 3Zv, 1Sv.

Legs: Following segments differ from Gamasellus in having a postero-

lateral or ventral seta missing: tibia T (2, 6/3, 2): genu IV (2, 5/2, 0);

tibia IV (2, 4/2, 1),

OTHER CHARACTERS, Female: Deutosternal denticles in & horizontal

rows. Tectum anterior murgin trispinate, with stout central spine and 2

small lateral spines, Movable, cheliceral digit with 4 conspicuous teeth

und sometimes with series of small teeth distal to these. On palp genu, seta

al\ slightly lanceolate with one lateral prong and seta a/2 slightly spatulate.

LEE—RHODACARIDAE 65

Figs. 93-99.) Hydrogamasus littoralis (G. & R. Canestrini).

93-97 and 99, female: 93, soma, dorsum; 94, idiosoma, venter; 95, leg LV (part).

dorsum; 96, pretarsi | and IV; 97, gnathosoma, yventer. 99, distal tip of tarsus 1,

98, male, chelicera,

66 REC, 8, AUST. MUS.. 16 (3): 1-219 July, 1970

100

101

103

4 104

Ors.

Figs, 100-106. Hydrogemasus littoralis (G. & R, Canestrini).

100-102, male: 100, idiosoma, yventer; 101, idiosoma, latus; 102. leg Il (part).

103-105, deutonymph: 103, idiosoma, dorsum; 104, idiosoma, venter; 10S, pretarsi

Tand TY. 106, protanymph, idiosoma, venter.

LEE—RHODACARIDAE 67

% re, te fp

or,

Figs. 107-L15. Hydrogamasus littoralis (G. & R. Canestrini).

107-109, protonymph: 107, soma, dorsum: 108, gnathosoma, yenter; 109, leg IV

(part), dorsum. 110-114, larva: 110, soma, dorsum; 111. idiosoma, venter; 112, leg

HL (part), dorsum; 113, gnathosoma. venter; 114, pretarsus I. 115, egg.

68 REC, 5S. AUST. MUS., 16 (3): 1-219 July, 1970

Dorsal setae long and tapering, may be slightly pilose. Shields with reticula-

tions (mainly on ventro-anal shield, but also around edges of other shields).

Spermathecal ringed tube probably opens near posterior paraxial edge of

acetabulum TV. Pretarsus | pedunculate and smaller than other pretarsi,

Lobes of pulvillus [l-IV attenuated and long with extra, skirt-like lobe

around base of claws. On tarsus IV all four dorsal setae in whorls 3 and 4

are subequal in length.

Male: Tectum may differ slightly from female. Movable cheliceral

digit with series of more than 3 teeth, and fused at base to slender sperma-

dactyl, which loops over back on itself level with digit tip. Seta st5 on

ventro-anal shield. On leg U1, femur seta av, genu seta av, and tibia seta av

enlarged into spurs. [n Hirschmann’s (1966) drawings, H. vitcthumi

appeurs to have ventral process on tarsus LI,

DISTRIBUTION. Pe, Pm: An: Sa. Besides the locality records

published here and with the original descriptions of nominal species there

are records of Hydrogamasus species from the North Atlantic coastline of

Europe, the furthest north being from Anglesey, North Wales (Glynne-

Williains and Hobart, 1952). Also, specimens have been found as far east

in the Mediterranean as Israel (Costa, 19662).

Usually found in rock crevices of the upper littoral zone, but may be

found elsewhere on the marine shore.

REMARKS. Hydrogamasus has in the past included many rhodacarids

which have a holonotal shield that is not fused to the ventro-anal shield of

the female, but they have been transferred to Gamasiphoides, Geogamasus,

Heydeniella and Hydrogamasellus by Hirschmann (1966) or myself (see

below). Hirschmann (1966) restricted the genus and divided it into 2

subgenera, One subgenus, 4ustrohydrogamasus, is here newly regarded as

synonymous with Parasitiphis, and the genus is considered restricted to his

other subgenus. The resulting genus is cusy to diagnose, und I consider its

closest relauionship is with the Gamasiphinae, although it is atypical within

the subfamily. The following 4 nominal species are included in the genus:

H. littoralis (G, & R, Canestrini, 1881); AH. efard’ (Trouessart, 1888);

H. kensleri Luxton, 1967; H. vitzihumi Hirschmann, 1966.

Hyvdrogamasus littoralis (G, & R, Canestrini)

Gamasus littoraliy G. and R. Canestrini, 1881, p. 1078.

FEMALE. Fig. 93-97 and 99, Idiosomal length, 860),

MALE, Fig. 98 and 100-102. Ldiosomal length, 7800p.

DEUTONYMPH. Fig. 103-105. [diosomal length, 660,,

PROTONYMPH., Fig, 106-109, [diosomal length 630..

1.EP—RHODACARIDAE 69

LARVA. Fig. 110-114. Idiosomal length, 450.

EGG. Fig. 115. Longest axis length, 390,,

LOCALITY, Female (N196859), male (N1I96860), deutonymph

(N196861), protonymph (N196862), larva (N196863), and unlaid egg

(N196864) drawn: Channel Islands (English Channel): rocks below high

tide, Belerouts Bay, Jersey, 18,7,1950, col. E.B,, dep. SAM,

The ‘tipico’ female (34/25) and ‘tipico’ male (34/26) examined:

Italy; the Lido, Venice, dep, SEAF,

REMARKS, In figured chaetotaxy patterns, filled in circles represent

setae considered to have been added at the previous moult. Characters of

the immature stages have not been used in the classification given here. The

extensive drawings of these stages are given only to increase this type of

data, which ts too limited from this family for satisfactory comparative

studies. Similar drawings are made in this paper of the immature stages of

Hevdeniella dentatu and Gamasellus tragardhi.

The adult HW, Jitteralis specimens in the Berlese collection agree with

the figures here, but disagree with Hirschmann’s (1966) drawings which show

a peritrematal shield completely free of the exopodal shields.

Gamasus litroralis is usually regarded as a junior synonym of Gamasus

salinus Laboulbéne after Oudemans (1902). Not many characters of

G. salinus can be gleaned from Laboulbéne’s (1851) description, but the

a setae on the palp femur and genu are drawn (enlarged copy, Fig, 317)

and commented on in the text. The position and shape of seta a/ on the

palp femur in this deseription is not similar to that of any rhodacarid, but it

is Similar to seta a/ on the palp femur (Fig. 316) of Hydrogamasus silvestri

Berlese, 1904b which Berlese, in the catalogue of his collection, has placed

under Pergamasus hamatus (K), 1 have, therefore, revoked Oudemuns'

(1902, p. 286) synonymy of G. salinus with G. Jitteralis and made a new

combination, Pergamasus salinuy (Laboulbene. 1851), and regarded the

Original description as being of the deutonymph and G, maritimus

Laboulbene, 1851 as being the adult female.

Hydrogamasus kensleri Luxton

Hydrogamasus kensleri Laxton, 1967. p. 76,

LOCALITY. Three females (N1I96865-N196867). two males

(NI96868 und N196869), two protonymphs (N196870 and N196871)

and a larva (N196872) examined: Macquarie [sland; rotting kelp, between

ude marks, Lusitania Bay, 4,12,1931, col. B,A.N,.Z.A.R, Expedition, no.

1668, station 82, dep. SAM.

REMARKS. Adults, protonymphs and a larva considered as belonging

lo this species. which was originally described from New Zealand, have been

70 REC, &, AUST, MUS., It (3); 1-219 July, 1970

collected as above. The larva appears to be conspecific with the larva which

was described as Hydrogamasus ( Austrohydrogamasus) watsont Hirschmann,

1966 (syn. Parasitiphis jeanneli (André) ),

Genus LAELAPTIELLA Womersley

Laelaptiella Womersley, 1956a, p. 512. Type-species: Laelaptiella anomala

Womersley, 1956a. by original designation.

DIAGNOSIS. Minute to small mites with shields not so extensive and

dorsal idiosomal surface not so convex as most members of this subfamily.

Ventro-anal shield fused to holonotal shield. Peritrematal shield not fused

to exopodal IV shield. Exopodal II shield not split. Sterno-metasternal

shield not fused to endopodal IV shield. Female movable cheliceral digit

has 3 teeth. Pretarsus | subequal in size to other pretarsi. Genu TV with

two yentral setae,

MORPHOLOGY

SCLEROTIZATION, Female: Holonotal fused to ventro-anal posterior

to sela Zv3, anterior to which holonotal is widely separated from

the ventro-anal, Diserete subcircular shield posterior to acetabulum IV.

Peritrematal reduced, narrowly boardering peritreme and not fused to

exopodal IV. Metapodal may be represented as part of small triangular

posterior extension of exopodal TV. Exopodals without any splits. Sterno-

metasternal fused to endopodals Il and part of III. Two pairs of

pre-endopodals,

Male: Sternito-genital. Subcircular shield posterior to acetabulum ['V

is meryed into ventro-anal. Otherwise as female.

CHALTOTAXY. Idiosoma: 6/, 6z, 5x, Or: 5/, 57, 5S, SR: Ssts 37 y,

34, SV,

Legs: As Gamasellus.

OruER CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows of what are either single, large denticles or ridges without denticles

that are acutely bent in the middle. Teectum anterior margin has a

central process, with spinules along its broad, conyex distal edge which is as

wide as its base. Movable digit with 3 teeth. Palp attenuated and long.

On palp genu, setae a/l and al2 spine-like. Dorsal setae simple, slightly

lanceolate, Holonotal shield dimpled but not reticulated, while venter is

reticulated, and all shields are shiny, Spermathecal ringed tube opens near

posterior paraxial edge of acetabulum IV, passing back to single, central

sacculus. Pretarsus [ similar in size to other pretarsi. Amongst dorsal setae

on tarsus TV, seta pe3 is only just the longest.

Male: Movable cheliceral digit with one tooth, fused at base to longer

spermadactyl On leg IT, femur seta av, genu seta av, tibia seta av enlarged

into spurs.

LEE—RHODACARIDAE 7

o—o-—?—~

}

0 |

Z 5s

Figs. 116-122. Laelapticlla anomala Womersley.

116-118, 122, female: 116, soma, dorsum; 117, leg TV (part), dorsum: 118, idiosoma,

venter: 122. gnathosoma, venter. 119-121, male: 119. idiosoma, yenter; 12

chelicera; 121, leg IL (part).

72 REC. S$. AUST, MUS... 16 (3): 1-219 July, 1970

DISTRIBUTION. Aa. All published records are with the original

descriptions of nominal species. Specimens have been collected from Lord

Howe Island and New Caledonia. dep. SAM.

Found in moss and plant litter.

REMARKS. Originally Laelaptiella had genus rank and was placed

in the Ascidae, Domrow (1957b) transferred it to Gamasiphiy as a sub-

genus, while Ryke (1962b) treated it as having genus rank when he included

Gamasiphis and Laelaptiella in the Ologamasinae. The following 2 nominal

species are tncluded in the genus: L. anomala Womersley, 1956a; L.

mackerrasae Domrow, 1957.

Laelaptiella anomala Womersley

Luelaptiella anomala Womersley, 1956a, p. 512.

FEMALE. Fig. 116-118, 122. Idiosomal length, 420.

MALE. Fig. 119-121. Idiosomal length, 370».

LOCALITY. Three females (N1968147-N1968149) and one male

(N1968150) drawn or examined: Australia; LF30, litter under epacridaceous

shrub amongst ‘Bottle Brush’ bushes and Eucalyptus, 8 miles north of Apsley,

Victoria, 17.12.1964, col. D. C. Lee, dep. SAM.

The holotype (N1968278) and paratype (N1968279) females

examined: Australia; Adelaide, South Australia, 6.1935, col. H. Womersley,

dep. SAM.

Subfamily LAELAPTONYSSINAE Womersley

Laelaptonyssidae Womersley, 1956a, p. 543. Type-genus: Laelaptonyssus

Womersley, 1956a.

REMARKS. | Since this subfamily contains only two species grouped

in a single genus, and the main function of this paper is to give diagnoses

for genera, the characters of the known material are listed under the genus

heading,

Genus LAELAPTONYSSUS Womersley

Laelaptonyssus Womersley, 1956a, p. 543. Type-species: Laelaptonyssus

mitis Womersley, 1956a, by original designation.

Puchihlungia Samsinak, 1964, p. 39, syn.n. Type-species: Puchihlungia

chinensis Samsinak, 1964, by original designation.

DIAGNOSIS. Average sized mites. Separate podonotal and opistho-

notal shields. Reduced ventral shields; ventro-anal shield being narrow with

a pair of small discrete shields posterior to metapodal shields and peritrematual

shield reduced and discrete. The setation of the notum and legs reduced,

| FE—RHODACARIDAE Th

All setae are simple, tapered, Cheliceral digits attenuated and spermadacty|

strongly recurved. Location of spermathecal ringed tube, if present, is

unknown. Pretarsus I pedunculate and larger than other pretarsi.

MORPHOLOGY.

SCLEROTIZATION. Female; Separate podonotal and opisthonotal.

Discrete, narrow ventro-anal, from which two small lateral shields have split

off at level of seta Jv3. Discrete metapodal. Inconspicuous. peritrematal

around the peritreme and stigma which are subequal in size. Exopodals do

not completely encase peraxial edges of acetabulu. Sterno-metasternal either

fused to endopodal II and part of IIT or discrete. Single pair of pre-

endopodals, or they may be absent. L. chinensis illustrated in Fig. 2.

Male: Sternito-genital and more extensive peritrematal, otherwise as

female.

CHAETOTAXY. Unusually variable and the homologies are uncertain.

On the legs the variations from the chaetotaxy of Gamasellus do not follow

ihe gradients indicated in the section on Leg Chaetotaxy (p. 13). For the

lez chaetotaxy only numbers of setae on a segment are given below,

Idiosoma. L.. mitis: 67. 7 or 8z, 6s, Or: 4 or 4 plus an unpaired J,

5Z, 48, OR: 4yt: BJ vy, 3Zv, OSv.

L. chinensis: 5j, 52, 3s, Or: SJ, 5Z, 5S, OR: Sst: 340, 32, 28v.

Legs. L, mitiy: palp, Ir, 3f, Sg: leg L 2e, 51, 11f, 12 or 13g, Ot:

leg Il, 2c, 6 or 71, 10f, 10g, 7H, 17ta: leg WN, 2e, St, 8f, 9g, 6t7, I7ta:

lex IV. Le, 51, 7f, lOg, 7, 17ta (seta av3 ubsent).

L, chinensis; palp, 21, 5f, 6g: leg 1, 2c, 5¢, Lf, Vlg, Viti leg TL, 2c,

6f, 10f, 92, 917, 18a: Jeg IL, 2c, 54, 7f, 8g, Bri, W8ta: leg TV, Le, 52, Of, 7g,

&ti, |8fa.

OTHER CHARACTERS. Female: Tectum anterior margin with a long,

narrow, central spine. Cheliceral digits unusually attenuated and movable

digit with only 2 fine teeth or edentate. On palp genu, setae a/l and al2 are

simple, tapered, On palp femur, seta a/ in proximal third or absent, Dorsal

setae simple, tapered, No noticeable spermathecal ringed tube. Pretarsus |

pedunculate and larger than other pretarsi. Pulvilli I-1Y appear to have a

single, large, bell-like lobe and claws are inconspicuous. Dorsul setae on

tarsus [TY subequal in length and setose.

Male: Movable cheliceral digit with one tooth or cdentate und fused at

base 0 strongly recurved spermadactyl that is similar length or more than

three times as long. On leg IJ, femur seta av enlarged into a spur.

DISTRIBUTION. Qs; Aa, Only records from Western Australia

and Kanton Province, China.

74 REC. S. AUST, MUS.. 16 (345 1-219 Juiy, 1970

Found associated with insects; £. mitiy in a fly culture and L. chinensis

on termites or in their nest.

REMARKS. When Womersley (19562) established Laelaptonyssidae,

containing only Laelapionyssus mitis, he referred to it as bemg a “luelaptid”.

Samsindk (1964) described L. chinensis as the only species in Puchi/lungia,

which was regarded as belonging to the Rhodacaridae. Despite the reduced

setation and, in one species, the reduced sternal sclerotization (cf.

Halolaelapidae), | have retained Samsinak’s (1964) concept of this genus

being a rhodacarid on the basis of the apotele, idiosomal sclerotization and

arming of the male leg Il. But, Laelaptonyssiusx is considered unusual

enough to be placed in a separate subfamily. The following 2 nominal

species are included in the genus; L. mitis Womersley, 1956a; L, chinensis

(Samsinak, 1964) comb.n,, Allotype female (N1968284), three paratype

females (N1968285-N1968287). holotype male (N1968288) and three

paratype males (N1968289-N1968291) of L. mitis have been examined:

Australia, from fly culture, Zoology Department, University of Western

Australia, 10.1950, col. E. P. Hodgkin, dep. SAM. Paratype female,

paratype male (N1968269) and paratype protonymph (N1968270) have

been examined: China; off Coprorermes formosanus Shiraki, near Koa-ho,

Kanton Province, 1963, col, K. Samsinaék, dep, male and protonymph SAM,

female in the collection of K. Samsinik.

Subfamily OLOGAMASINAE Ryke

Ologamasinae Ryke, 1962b (Ist August), p. 159.

Gamasellinae Hirschmann, 1962 (August, assumed 3fst by terms of the

International Code of Zoological Nomenclature, 1961), p. 39.

Cyrtolaelapidae Berlese, sensu Johnston (in part), 1968, p. 19.

Type-genus: Ologamasus Berlese, 1888,

DIAGNOSIS. Minute to gigantic mites, with shields that range from

being Limited in extent and lightly sclerotized, to being extensive and heavily

sclerotized, Holonotal shield or separate (or very rarely partially separated )

podonotal and opisthonotal shields, in some cases both attributes of this

character occur in members of the same genus or sexes of the sume species.

Ventro-anal shield may or may not be fused to the notal shield and this also

varies within a genus or species, Peritrematal shield usually fused to the

exopodal IV shield and often to the ventro-anal shield, although in

most cases the latter fusion only occurs on males, Sclerotization often

conspicuously sexually dimorphic. On the female sterno-metasternal shield

a line joining setae 72, s13 and s74 would never enclose an angle of less than

95° except in some Gamasellus discutatus-complex species.

LEE-—RHODACARIDAE 75

MORPHOLOGY,

ScLEROTIZATION, The size of shields and extent of the fusion

between them varies considerably between species and often between the

females and males of the same species. This sexual dimorphism usually

uppeurs as a more extensive sclerotization of the male venter, but in

Gamaselluy discutatus, some Geogamasus species and an unnamed

Heydeniella species the female has a holonotal. while the male has a narrow

fissure between the podonotal and opisthonotal, and on the male of

Geovamayus delamare’ and an unnamed Hydrogamasellus species there ure

lateral incisions partly separating the podonotal and opisthonotal, while in

some males of the Acugamasus punctatus-complex a humeral section of the

podonotal is partly separated from the rest and fused to the peritrematal,

Conspicuous, wide spread sexual dimorphism of this type only occurs in

species of this subfamily and the Rhodacarinue, and an aspect of it most

commonly found in this subfamily, but not in other rhodacarids, is the fusion

of the ventro-anal to the peritrematal of the male, Also the females of some

species of Ologamasinae differ from all the females in the other subfamilies

within the Rhodacaridue by having the ventro-anal fused to the peritrematal

(Cymiphis, Hiniphis, Periseius, and some Gamasellus Jalciger-complex and

discitatus-complex species), The following other sclerotizauion characters

are fairly constant within the subfamily. Although the male ventro-inal is

often fused to the notal, in the female this is not so except for the following

eroups; Oloyamasus, Cymiphis, some Heydeniella crozetensis-complex and

Hydrogamaselluy species, und Hiniphiy and Pyriphiy species, Small

intergenito-ventrals occur on some members of 4 number of genera, The

peritrematul is fused to the notal anteriorly and exopodal TV posteriorly

except for some species with unusually short peritremes (Notegamasellus

and Neogamasellevans species) and for the males of most species of

Geogamasus, Parasitiphis and Litogaumasus. Exopodal IIL always split, The

female metasternal is not fused to endopodal IV except for the following

vroups: Cymiphis, Hiniphis, and some Heydeniella crozetensis-complex

species.

CHAETOTAXY. The number of setae on the idiosoma is very variable

especially on the opisthosoma, which may also be hypertrichous or

hypotrichous. The podosoma usually has between 20 and 22 puirs of setae

dorsally and 5 pairs of setae ventrally, the few exceptions being for some

species of Gamasellini, The leg chactotaxy is fairly constant, but compared

with Gantasellus, some species of Gamasellini may have an extra one or two

setuc on lez HP and some Ologamasini may have one seta less on leg IV.

Ovnek CHARACTERS. 'Tectum anterior margin can have many different

shapes and may be sexually dimorphic. On pulp genu, seta a/l is

often pectinate or spatulate with a single lateral prong and seta al2 is

often) spatulate, as for the antero-loteral setae on the palp genu of some

76 REC, S&S. AUST, MUS, b6 (ads 1-219 July. 1970

species of Sessiluncinae. On palp femur, seta a/ is generally on proximal

third jn contrast to Rhodacarinue, Tangaroellinac and some Sessiluncinae

species, but on an unnamed Liftegamasus species it is on central third.

On the ventral surface of the podonotal shield of some species there are 4

light-refructile structures between seta j/5 and /6. These structures also occur

in the Rhodacarinae and may be strengthened areas of sclerotization to which

muscles are attached, The female sternal seta sf2, st3 and yf4 are usually

nearly in a straight line. and the smallest angle that would be contained by

such a line is 95° except in some species of the Gamtaselluy discutatus-

complex, The method of introduction of sperm into the female may vary

considerably in this subfamily. According to Michael (1886 and 1892) the

sperm is introduced directly into the yagina of Euryparasitus emarginatus

and because of the similarity of the spermadactyls the same could be true

for Cyrtolaelaps. On the other hand many species of Ologamasinae have

structures looking like spermathecal ringed tubes which usually open near

the posterior paraxial edge of acetabulum TV, but on a number of other

species (usually mentbers of the Ologamasini) they open on the proximal

segments of leg If and leg IVY, Since these tubes cannot always be seen to

be attached to a sacculus, data is needed to support their spermathecal

function and this is not yet available in many cases.

As commented on in the general section on Morphology (p. 13),

within the Rhodacaridae there is often an association of three sets of

attributes as follows: ‘some complex dorsal setae—divided dorsal shield—

dull sclerotization’, or ‘all simple dorsal setae—entire dorsal shield—shiny

sclerotization. This is particularly obvious in the Ologamasinae and

although some species do tot have an association of the attributes as

given, these characters are the basis for distinguishing two tribes in this

subfamily, The dullness appears to be caused by a covering of adhesive

exudate with attached detritus and fungal hyphae in older specimens. In

species where the exudate is thick, simple or lanceolate setae run parallel to

the body surface, imbedded in the exudate, while pilose setae stick out

through the exudate at right angles to the body surface, On species with a

shiny sclerotization, the dorsal seta are rarely complex and detritus and fungal

hyphae are not found attached to the mite.

Besides the conspicuous sexual dimorphism in sclerotization many other

structures vary between the sexes in this subfamily, most of which are

probably concerned with assisting the male keep its grip on the female during

sperm transference, The male gnuthosoma may have attenuated corniculi

which are often on u protuberance of the hypostome, also there may be a

ventral process on the palp (Hydrogamasellus and Acugamasus natalensis-

complex) and the tectum may be different (Ologamasus and Geogamasus

species). The anterior edge of the male podonotal shield may extend forward

over the gnathosoma (some Heydeniella and Euryparasitus species), The

LE E—RMHODAC ARIDAE 7

enlargement of setae on leg UL is usually more extensive than in other sub-

families and tarsus I] may have ventral setae (Parasitiphis, Cyrtolaelaps,

Evuryparasitus and Litogumasus species) or dorsal setae (Cymiphis,

Geogamasus and Hydrogamasellus species) enlarged into tubercles. Two

littoral genera (Parasitiphis and Litogamasus) have the male legs IT and

IV enlarged and carrying setous or non-setous processes, Often the cuticular

processes on the distal margins of the proximal segments of female legs HI

and IV are enlarged, but sometimes there are more conspicuous processes

on the female legs (Cymphiy and some Heydeniella and Gamasellus

discutats-complex species ).

DISTRIBUTION. The subfamily is cosmopolitan, with 17 genera

found only in the Southern Hemisphere, Three genera (Cyrtolaelaps,

Allogamasellus and Euryparasitus) are only found in the Northern Hemis-

phere. The Ganrasellus faleiger-complex is the only group of closely related

species found in moss, plant litter and soil that has a large number of species

in both the Northern and Southern Hemispheres, Periseius and probably

Rhodacaraides occur in both of these hemispheres, Heydeniella goei from

the British Isles is the only species belonging to the Ologamasini that is not

restricted to the Southern Hemisphere. Species are usually found in moss,

plant-litter and soil, but some genera (Cvrtolaelaps and Euryparasitus) are

usually found in animal nests or roosting pluces of bats while other genera

(Parasittiplis, Litogamasus and Periseius) are usually found in the littoral

zone and Rhoducarvides is either littoral or probably commoner in deeper

soil layers,

REMARKS. Among the rhodacarids some members of the Ologama-

sinae are the most similar to the Parasitidae and Veigaiidae and therefore I

would regard them as more like the ancestoral rhodacarids. Also, on the

basis of the presence or ubsence of conspicuous sexual dimorphism, | would

regard this subfamily as more closely allied to the Rhodacarinae than to the

Gamasiphinae and Sessiluncinae. Therefore, | disagree with Johnston's

(1968) transter of members of this subfamily to another family, Cyrto-

laclapidae, with the members of the Gumasiphinae and Sessiluninae, leaving

only what are here regarded as members of the Rhodacarinae in the

Rhoducaridae. This subfamily has the most species and genera, and its

genera are the most difficult to delineate. The division of the subfamily

into two tribes as presented here is probably no indication of its phylogeny

but itis convenient and the use of the adult dorsal sclerotization to distinguish

the groups resulting frond such a division has precedent.

I recoynize 2 tribes within this subfamily; Ologamasior and Gamasellini.

Adults of these tribes. and the genera grouped in them, can be distinguished

by the following key. The 3 species incertae sediv commented on ut the end

of this paper belong to the Ologamusinae und key out to their correct tribe,

78 REC. §. AUST. MUS,, 16 (3): 1-219 July, 1970

but not necessarily to the genus which they have been placed in for

convenience.

KEY TO TRIBES AND GENERA OF OLOGAMASINAE

1. Usually holonotal shield. If separate or partially

separate podonotal and opisthonotal shields

(some Geogamasus, Heydeniella and Hydro-

gamasellus species), then single pair of pre-

endopodal shields (as for all other members of

this tribe except for Neogamasellevans berlese/),

all dorsal setae superficially simple (may carry

inconspicuous lateral prongs. but never have

conspicuously pilose podonotal setae j4, 25 and

r3 amongst simple setose or lanceolate setae

which is also the case for all other members of

this tribe except for Pyriphis species), pretarsus

I pedunculate and seta a/l on palp genu has

more than 5 lateral prongs .. .. .. .. -. .- Ologamasini, 2

Usually separate podonotal and opisthonotal shields,

If holonotal shield (some Gamasellus discutatus-

complex species) then 2 or 3 pairs of pre-

endopodal shields and podonotal setae j4, <5

and r3 are conspicuously pilose amongst simple

setose or lanceolate setae. No species in this

tribe has all the following characters: single pair

of pre-endopodal shields; all dorsal setae super-

ficially simple; pretarsus | pedunculate and seta

all on palp genu with more than 5 lateral prongs Gamasellini, 10

Seta a/l on palp genu is spine-like or spatulate and

never has more than 2 lateral prongs .. .. -, 3

Seta a/l on palp genu always has more than 2 lateral

POTS ee ede oe: fee A a a SG

3. Pretarsus I not pedunculate. Peritrematal shield

reduced; on female narrows behind stigma before

fusing with exopodal IV shield, on male

terminates behind stigma without fusing to other

shiglds~.. Se Fe ee Ce ATL BL “Pardsttiphis

Pretarsus I pedunculate. Peritrematal shield sub-

stantial; on female broadens behind stigma

before fusing with exopodal IV shield, on male

extends back behind stigma to fuse with both

exopodal IV and ventro-unal shields .. .. .. 4

LEE—RHODACARIDAE

4, Dorsal setae all simple, Female ventro-anal shield

separate from notal shield and male ventro-anal

shield fused to sternito-genital shield. Sperma-

dactyl straight, running parallel to movable

cheliceral digit oe

Some dorsal setae conspicuously pilose, Female

ventro-anal shield fused to notal shield and if

male yventro-anal shield fused to sternito-genital

shield then spermadactyl recurved from point of

attachment to movable cheliceral digit so that it

nearly points in the opposite direction . .

5. Female sterno-metasternal shield fused to endopodal

IV shield. Male sternito-genital shield not fused

to ventro-anal shield. Spermadactyl approxi-

mately parallel to movable cheliceral digit

Female sterno-metasternal shield not fused to endo-

podal IV shield. Male sternito-genital shield

fused to ventro-anal shield. © Spermadactyl

recurved from point of attachment to movable

cheliceral digit so that it nearly points in the

Opposite direction ws, VF

6. Single pair of female pre- -endopodal shields have

paraxial margins subequal in length to distance

between sternal setae s#l and sf2, and ventro-

anal shield not fused to notal shield. Peritreme

short, not reaching forward to level of anterior

margin of acetabulum III. Peritrematal shield

not conspicuously fused to exopodal shields.

Spermadactyl nearly straight

If female pre-endopodal shield with paraxial margin

subequal in length to distance between sternal

setae stl and st2, then ventro-anal shield fused

to notal shield. Peritreme reaches forward

beyond anterior margin of acetabulum III. If

peritrematal shield not conspicuously fused to

exopodal shields (some Geogamasus males),

then spermadactyl able to coil up around 3

hinges

7. Podonotal setae in 22 pairs.

tube opens on coxa III

trochanter III. On

modified to tubercle

Spermathecal ringed

or proximally on

male tarsus II, seta ad3

Rykellus

Cymiphis

Pyriphis

Neogamasellevans

11.

REC. S. AUST, MUS., 16 (3): 1-219

Podonotal setae in 20 or 21 pairs. Spermathecal

ringed tube opens on coxa IV or distally on

trochanter III]. On male tarsus II, seta ad3 is

setose ..

. Spermathecal ringed tube opening proximally on

posterior surface of trochanter III, often coiling

in coxae JI. Spermadactyl able to coil up

around 3 hinges .. -.

Spermathecal ringed tube opening distally on dorsal

surface of coxa Ill. Spermadactyl unable to

coil up around 3 hinges .. .. 2. 2... ee ee

Spermathecal ringed tube opening distally on dorsal

surface of female trochanter III. If female

yentro-anal shield fused to holonotal shield then

sterno-metasternal shield fused to endopodal 1V

shield. Central spine on anterior edge of tectum

comes to a single point. There are 20 pairs of

podonotal setae. On male tarsus IT, seta av2

may be modified to a spine... .. 2. 2. 2. ee

Spermathecal ringed tube opening distally on ventral

surface of female coxa IV. Female sterno-

metasternal shield not fused to endopodal IV

shield although ventro-anal shield is fused to

holonotal shield. Central spine on anterior edge

of tectum is bifurcate at tip. There are 21 pairs

of podonotal setae. On male tarsus II, seta av2

is setose ..

Podonotal shield conspicuously larger than opistho-

notal shield and bearing 56-65 setae. Unpaired

pre-anal seta about as close to anus as paranal

setae ..

Podonotal shield subequal in size to opisthonotal

shield, or fused to it, and bearing up to 48 setae

although usually 22 pairs of setae or less. No

unpaired pre-anal seta near anus.

Well developed leg I, bearing stout spines (enlarged

ventral and antero-lateral setae) on genu and

tibia ..

Setae on genu and tibia of leg I not enlarged to

stout spines

July, 1970

Geogamasus

Hydrogamasellus

Heydeniella

Ologamasus

Notogamasellus

LEE—RHODACARIDAE

12, Anterior spine on genu I is seta all. Pretarsus |

absent. Single postero-lateral seta on genu III

(2, 4/2, 1) and tibia IN (2, 3/2, 1). Vertical

setae (jl) on prominent protruberances

Anterior spine on genu I is seta av2. Pretarsus |

present, although small compared with other

pretarsi it has 2 claws. Two postero-lateral setae

on genu II (2, 4/2, 2) and tibia III (2, 3/2,

2). Vertical setae (j1) not on prominent

protruberances eb eh Lo he tee

13, Ventro-anal shield separate from opisthonotal shield,

but fused to peritrematal shield in both sexes.

In female, metasternal shield fused to endopodal

IV shield, although it may be separate from

sternal, and in male, sternito-genital shield fused

to ventro-anal shield . . ts gtode

Rarely is ventro-anal shield both fused to peri-

trematal shield and separate from opisthonotal

shield, and then only in females with metasternal

shield separate from endopodal IV shield and in

males with ventro-anal shield separate from

sternito-genital shield Fi nw oy oy

14. Pretarsus I present but not pedunculate. All dorsal

setae with tapering tip. Female sclerotization

reduced ventrally: sterno-metasternal shield

never fused to more than endopodal IL shield

and the anterior half of endopodal III] shield;

discrete metapodal shields: peritrematal shield

usually narrows behind stigma or at least is not

so broad that it has a truncated posterior margin,

Male chelicerae atypical: either fixed digit with

dorsal groove (Fig. 362), or movable digit fused

along its entire length to longer, stout sperma-

dactyl (Fig. 331 and 343) fae ech

Pretarsus I, if present, usually pedunculate, if not,

(Gamasellus pyriformis-complex) then some

dorsal setae with spatulate tip, and female

sclerotization not reduced ventrally as above.

The male chelicerae not atypical as above . .

15. One or more pairs of pre-endopodal shields, if only

one pair then female metasternal seta (st4) is

on small discrete shield. Male fixed cheliceral

Evanssellus

Heterogamasus

Periseius

17.

18.

19,

REC, S. AUST, MUS., 16 (3): 1-219

digit has dorsal groove, and spermadactyl is

slim and fused to the movable digit at its base

0.0 ie Sr SC <a

If pre-endopodal shields present, then one pair only

and metasternal seta (st4) is on sterno-meta-

sternal shield. Male fixed cheliceral digit with-

out dorsal groove, and spermadacty] is stout and

fused to the movable digit along the digit’s entire

length .. 2... ee

. Clearly defined pre-endopodal shields absent.

Paranal seta anterior to centre of anus. Female

setae Jvl and Zv1, or just seta Jvl, are anterior

to yentro-anal shield. Male tibia and tarsus II

with articulated connection PA dee ed hes

Clearly defined pre-endopodal shields _ present.

Paranal setae posterior to centre of anus. Setae

/vl and Zvi are on vyentro-anal shield. Male

tibia and tarsus II are rigidly fused together .

Female ventro-anal shield fused to dorsal shield,

and sterno-metasternal shield fused to endopodal

IV shield. Male ventro-anal shield fused to

sternito-genital shield. Two pairs of pre-

endopodal shields. Peritrematal shield com-

pletely merged with the exopodal shields behind

stigma .. Hane eR 3 a, ye ned, ih

Female ventro-anal shield never fused to dorsal

shield, and sterno-metasternal shield never fused

to endopodal IV shield. If male ventro-anal

shield fused to sternito-genital shield then there

is a single pair of pre-endopodal shields and a

fissure separates peritrematal from part of

exopodal IV shield behind stigma . ,

All dorsal setae simple, setose or spine-like, except

that setae jl and <1 may be slightly pilose .

At least some dorsal setae conspicuously pilose, and

often lanceolate dorsal setae present . ‘

Dorsal setae, short, spine-like, with setae jl and z1

the longest and slightly pilose. Three pairs of

pre-endopodal shields . . ‘

Dorsal setae, fine and setose. One or two pairs of

pre-endopodal shields ,

July, 1970

Litogamasus

Cyrtolaelaps

Eurvparasitus

Hiniphis

Allogamasellus

Rhodacaroides

LEE—RHODACARIDAE 83

20. One pair of pre-endopodal shields .. .. .. .. .. 21

Two or three pairs of pre-endopodal shields .. .. 22

21. On opisthonotal shield there are 60 or more setae,

not all of which are paired, and there is at least

one accessory seta between the J rows of setae.

Male sternito-genital shield fused to ventro-anal

shield 2... a. : Pilellus

On opisthonotal shield here are “58 or : eet setae

in pairs, and no accessory setae between the J

rows of setae. Male sternito-genital shield not

fused to ventro-anal shield .. .. .. . Acugamasus

22. All dorsal setae on idiosoma and genu aang fibia of

legs LH, III and IV are densely poset ne their

entite* length. ign a ee CA ee . .. Laelogamasus

Only some dorsal setae on sdieaatiis are “dees

pilose, and usually for only part of their length,

while the dorsal setae on the genu and tibia of

leg H, HI and IV are not pilose .. .. .. .. Gamasellus

Tribe OLOGAMASINI Berlese

Ologamasinae Ryke, 1962b, p. 159. Type-genus: Ologamasus Berlese,

1888.

DIAGNOSIS. Minute to large mites. The following characters usually

occur on adults of this tribe (exceptions are given in parenthesis). Holonotal

shield (some Geogamasus, Heydeniella and Hydrogamasellus species). One

pair of pre-endopodal shields (Neogamasellevans berlesei). Simple dorsal

setae or, if complex, then single conspicuously complex setae are not

surrounded by simple setae (Pyriphis species). Spermathecal ringed tube

opens on proximal segments of legs HI and IV (Parasitiphis and possibly

Neogamasellevans species ).

MORPHOLOGY.

SCLEROTIZATION. Usually holonotal, but may be separate or partially

separate podonotal and opisthonotal as on some Geogamasus, Heydeniella,

and Hydrogamasellus adults. Rarely more than one pair of pre-endopodals

but may be 2 pairs as on Neogamasellevans berlesei.

CHAETOTAXY. Setation of the legs is as that of Gamasellus, except

for many Heydeniella species and some Hydrogamasellus species which have

only one ventral seta on genu IV (2, 5/1,1).

OTHER CHARACTERS. Usually, all dorsal setae are simple and setose,

but they may be slightly pilose, while in some Ologamasus and_ all

Cymiphis and Pyriphis species a number of dorsal setae are conspicuously

84 REC. 8. AUST, MUS, 16 (3); 1-219 July, 1970

pilose and may be spatulate. The tendency in the former two genera is lor

lateral and posterior dorsal setae to be larger and more pilose, and podonotal

setae 23, 24 und £3 ure never pilose amongst simple setae as in Pyriphis

species. Female sternal setae are usually nearly in a straight line but never

us on most Gamasiphinue species, Spermathecal ringed tubes muy open

near the posterior paraxial edge of female ucetabulum TV (Parasitiphis and

possibly Neogamasellevans species) but usually opening is probably on the

proximal segments of legs III or IV.

DISTRIBUTION. Only one species, Heydeniella goei, belonging to

this tribe has been found in the Northern Hemisphere. Many species have

been found in the Southern Hemisphere. It is common in the Australian

region, while its species constitute the bulk of rhodacarid material collected

from the Neotropical region.

REMARKS, The Ologamasini is intended mainly for species with

adults that have a single holonotal shield and usually simple dorsal setae,

There is a precedent for this concept although it probably gives little indica-

tion of the phylogenetic relationship of the species. It may later prove more

useful to limit this tribe to species in which the spermathecal ringed tube

opens on the proximal segments of female legs HT or IV, At the moment

it is not easy to distinguish the females of three genera, Ologamasus,

Heydeniella and Hydrogamasellus, because their diagnoses ure based on the

location of the external opening of the spermathecal ringed tube, a character

which can be difficull to observe. But if both females and males of a species

belonging to these venera are known, the placing of the species in the correct

genus is much easier,

I recognize 9 genera within this tibe, Ologamusus, Cymiphis,

Geogamasus, Heydeniella, Hydrogamasellus, Neogamasellevans, Parasiti-

phis, Pyriphis, and Rykellus. Also, two species, incertae sedis (Neogama-

sellevans berlesei and Hydrogamasellus ubatubaensis). commented on at the

end of this paper, are placed in this tribe, Members of these genera can be

distinguished by using the above key (p, 78),

Genus OLOGAMASUS Berlese

Ologamasus Berlese. 1888, p. 194 (not Berlese, 1906, p. 242 and Japsus,

Hologamasus Berlese, 1892d, inter page 61 er 62 (bound between page

60 and 61). Type-species: Gamasnx aberrans Berlese, 1888, by

monotypy.

Ologamasellus Berlese, 1914, p. 139. Type-species: Gamasuy cherranys

Berlesc, 1888, by original designation.

DIAGNOSIS. Small to average sized mites, always with extensive.

well sclerotized shields. Always holonotal shield fused to ventro-anal shield,

although the extent of this fusion may be limited, Male ventro-anal shield

LEE—RHODAC ARIDAE 85

always fused to the exopodal IV shield and peritrematal shield. Sterno-

metasternal shield is not fused to endopodal IV shield. Dorsal setae may be

simple, pilose, spatulate or pilose and spatulate, Spermathecal ringed tube

opens distally on ventral surface of coxa LV, Pretarsus I pedunculate and

smaller than other pretarsi, On male tarsus HH, setae all setose.

MORPHOLOGY,

SCLEROTIZATION. Female: Holonotal fused to ventro-anal, although

fusion may be limited to area bearing aciculae posterior to anus. No

discrete metupodal. Peritrematal fused to exopodal TV and may extend

posterior to this as triangular shield which may be homologous with the

metapodal, Split in exopodals LH, Il] and 1V. Sterno metasternal fused to

endopodal Hand part of TI, Single pair of pre-endopodals,

Male: Fusion of ventro-anal to holonotal similar to female or slightly

more extensive. Peritrematal and exopodal TV fused to ventro-anal.

Sternito-genital not fused to ventro-anal,

CHAETOTAXY. Idiosoma: 6, 62, 4s, Sr: 5/4, 524, SS, SR: Syts 3UV,

3Zy, O or Sy,

Legs, As Gamasellus.

Ovrnrr CHARACTERS. Female: Deutosternal denticles in 8 or 9

horizontal rows. Tectum anterior margin basically unispinate, but spine is

bifurcate al lip and has a spinule on each side, Movable, cheliveral digit

with 3 teeth. On palp genu, seta all pectinate with 5 lateral prongs and

seta a/2 spatulate, Dorsal setae may be simple or spatulate or spatulate and

pilose at tip. Shields apparently shiny. Usually genital and sometimes sternal

shicld with longitudinal striae. On coxa TV, just distal to ventral seta, there

is the aperture of what is probably the spermatheeal ringed tube. Pretarsal

sheath I on slim peduncle of similar length, Pulvilli H-PV with lateral lobes

that are attenuated und long. Amongst dorsal setae on tarsus IV, seta pd3

is the longest. being either setose or spatulate.

Male: Corniculi proportionately longer than on female. Tectum may

be more complex than on female. Movable cheliceral digit with one tooth,

and it may have a ventro-lateral groove opposite a spermadactyl that ts

subequal in length to digit. Sternito-genital shield with longitudinal striae.

On leg IT, femur seta av and sometimes setu pyl, genu seta av and sometimes

tibia seta ay enlarged into spur or spine. May be non-setous processes on

these segments.

DISTRIBUTION. NTb, NTc. All records are from a region between

the Mato Grosso in Brazil and La Plata in Argentina, occurring in a range

simular to that covered by the locality records published with the original

descriptions of nominal species in this genus.

Found amongst plant litter on soil, or under stones or the bark of trees.

86 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

om Fe }

ita

123, 124, 127, 129, 132

126, 126, 131

loop

E18)

125

Sy oe

— ==)

ss leo

Figs, 123-133. Ologamastius aberrans (Berlese).

123-128, female: 123, idiosoma, venter: 124. leg [V (part), dorsum: 125, pretarsus L

126, coxae IV, plus trochanter IV (part): 127, tectum; 128, gnathosoma, venter.

129-133, male: 129, podosoma, venter 130, leg He (part). postero-latus; 131,

corniculus; 132, tectum: 133, chelicera.

LEE—RHODACARIDAE 87

Figs. 134-14). Qlogamasus stimplicior (Berlese).

134-137 and 141, female: 134, soma, dorsum; 135, idiosoma, venter: 136. coxue Il

and IV plus trochanters (parts); 137, leg 1V (part). dorsum; 141, gnathosoma, venter,

138-140, male: 138, idiosoma, venter; 139, chelicera; 140. leg IL (part).

aK REC. S$. AUST. MUS., 16 (3): 1-219 July, 1970

Figs. 142-151. Ologamasus striolatuy (Berlese).

142-145 and 151, female: 142, soma, dorsum: 143, idiosoma. venter: 144, coxae III

and [V plus trochanters (parts); 145, leg IV (part), dorsum; 151. gnathosoma.

venter., 146-150, male: 146, idiosoma, venter; 147, chelicera; 148, lez IT (part);

149, carmiculus and palp trochanter: 150, pretarsus I plus tarsus 1 (part).

LEE—RHODACARIDAE ao

REMARKS, Because of the fusion of the female ventro-anal shield to

the notal shield, Ologamasus has been confused with a very different genus,

Gamasiphis, as indicated by a reference to the ‘Ologamasus-Gamasiphis

group’ in Ryke (1961b). Lee (1966) redefined this genus, placing in it

species with a single holonotal shield in the female, complex dorsal setae and

in some cases without fusion between the ventro-anal shield and the notal

shield. This definition is revoked here, und the genus is used to contain

species with females having what is probably a spermathecal ringed tube

opening ventrally on coxa TV. So far the females of all the nominal species

have the ventro-anal shield fused to the notal shield, but the extent of this

fusion varies considerably and may be very limited. The form of the dorsal

setae is also very variable and they may be mostly simple with only a few

setae being inconspicuously pilose. Since some species of Hydrogamasellus

are similar to species in Ologamasus and other //ydrogamasellus species are

similar to Heydeniella species. alloting a species to one of these three genera

can be difficult, especially when only the female is known and the probable

spermatheeal ringed tube is indistinct. The following 3 nominal species are

included in this genus: O. aberrans (Berlese, 1888): O. simplicior ( Berlese,

1914); O, strielatus (Berlese, 1916a).

Ologamasus aberrans (Berlese )

Gamasus aberrans Berlese, 1888, p. 194.

FEMALE, Fig, 123-128. Idiosomal length, 730p, Dorsal setae

difficult to examine, but podonotal setae mostly shaped as seta 42, or

simpler, while opisthonotal setae as seta 75, Proximal segments of leg 1V

drawn (Fig. 124) belong to the right side while the tarsus belongs to the

left side.

MALE, Fig, 129-133. Idiosomal length, 700e, On leg II, only

seta av on femur and genu ure enlarged into spurs. The posterior process

of femur seta av is quite thick although lightly sclerotized. All setae on

tursus IT are setose.

LOCALITY. The ‘tipico’ female (9/16), and ‘tipico” male (9/1),

drawn, also ‘cotipico’ female (215/28), and 2 ‘upico’ males (9/16),

examined: Brazil; under the bark of trees, Mato Grosso, col. A. Balzan,

dep. SEAF.

Ologamasus simplicier ( Berlese )

Ologamasellus simplicior Berlese. 1914, p. 140,

FEMALE. Fig, 134-137 and 141. Idiosomal length, 620», Pretarsus

1 with peduncle that is proportionately shorter and broader than that of

O. aberrans.

90 REC. 8S. AUST. MUS., 16 (3): 1-219 July, 1970

MALE. Fig. 138-140. Idiosomal length, 580. Tectum similar to

that of female.

LOCALITY. Female (N1968255) and male (N1968256) drawn,

and female and 3 males examined: Argentina; in dark place among humus,

fallen leaves and rotten wood, 9 de Julio Park, Tucuman City, 1953, col.

P. Wygodzynsky dep. SAM (N1968255 and N1968256) and BM(NH)

(other 4 specimens).

The ‘tipico’ female (165/45) and ‘tipico” male (165/45) examined:

Argentina; La Plata, col. Bruck, dep, SEAF.

Ologamasus striolatus ( Berlese)

Ologamasellus striolatus Berlese, 1916a, p. 163,

FEMALE. Fig. 142-145 and 151. Idiosomal length, 750». Three

raised, oval areas on podonotal shield anterior to seta r2, appear to have

many fine pores. Berlese’s specimens have peritrematal shield more com-

pletely fused to exopodal shield IV and processes on coxae II and IV are

not so big.

MALE, Fig. 146-150, Idiosomal length, 720%. Tectum similar to

that of female. On Berlese’s specimens, seta pvl on femur II shorter and

fatter.

LOCALITY. Female (N1I968257) and male drawn, and 2 females

examined: Argentina; rotten vegetable matter (chiefly corn stalks, grass and

twigs), vacant lot, Tucuman City, 1.1953, col. P. Wygodzynsky, dep. SAM

(N1968257) and BM(NH) (other 3 specimens).

The ‘tipico’ female (170/16), ‘tipico’ male (170/16), and female

(170/49), and three males (170/49, 170/17 and 215/27) examined:

Argentina; under stones, La Plata, col. Bruck, dep. SEAP.

Genus CYMIPHIS gen.n.

Type-species: Ologamasus cymosus Lee, 1966.

DIAGNOSIS. Minute to large mites, always with extensive, well

sclerotized shields. Holonotal shield always extensively fused to ventro-

anal shield. Male and sometimes female peritrematal shield fused to

ventro-anal shield. Sterno-metasternal shield is fused to endopodal IV shield.

Dorsal setae may be simple, pilose or pilose and slightly spatulate. Sperma-

thecal ringed tube probably opens distally on antero-lateral surface of coxa

IV. Pretarsus [ pedunculate and smaller than other pretarsi. Conical,

non-setous spur on ventral surface of female femur I] and sometimes on

femur HT and, or TV. On male tarsus IH, seta ad3 modified to a short spine

or tubercle.

| FE—RHODACARIDAE 9]

MORPHOLOGY,

SCLEROTIZATION, Female: Holonotal fused to ventro-anal. — Peri-

trematal fused to exopodal IV and it extends to point posterior to this as

triangular shield which may be homologous with the metapodal. Peritrematal

may also be fused to ventro-anal. Split in exopodals II, Il and TV. Sterno-

metasternal fused to endopodals Il, 1 and 1V. Single pair of pre-endopodals,

Male: Peritrematal fused to ventro-anal. Sternito-genital not fused to

ventro-anal,

CHAETOTAXY. Idiosoma; 6], 6z, Ss, 4r: 4 or S/, 52, 48, 4, 6 or

TR: Sst; adv, 324, 1S8v,

Legs: As Gamasellis,

Oruer CHaracters. Female: Deutosternal denticles in 7 horizontal

rows. Tectum anterior margin is basically trispinate but may have

lateral spinules, which may be large enough to make margin quinquispinate,

Movable cheliceral digit with 3 teeth. On palp genu, seta a/l with single

lateral prong and both setae all and al2 slightly spatulate. Some dorsal

setae may be simple, but pilose or pilose and spatulate setae always present.

Shields dull and usually covered with reticulations. Spermathecal ringed

tube is often not easy to locate, but in some species a tube clearly opens

distally on the antero-lateral surface of coxa IV. Pretarsus | pedunculate

and smaller than other pretarsi. Conical, non-setous spur on femur II and

sometimes on fernur TI] and, or IV. Pulvilli I-IV with lateral lobes that are

attenuated and long. Amongst dorsal setae on tarsus TV, seta pd3 is the

longest, and at least slightly lanceolate or pilose.

Male: Corniculi may be proportionately longer than in female.

Movable cheliceral digit with | tooth or 3 teeth as in female, and fused at

base to stout, slightly longer spermadactyl. On leg Il, femur setae ay and

py, genu seta avy and sometimes tibia seta av enlarged into spurs, while on

tarsus, seta ad3 modified to a short spine or tubercle.

DISTRIBUTION, An: Sa. All published records are with the original

descriptions of nominal species.

Found in moss and plant litter.

REMARKS. Six out of the seven Cymiphis species were originally

described as belonging to the ‘eymosus-group’ in Ologamasus by Lee (1966).

Cymiphis is similar to Ologamasus, but there are a number of morphological

differences, besides the absence of lateral prongs on seta a/l on the palp

genu used in the key to Ologamasinae genera, which are found only on these

species [rom New Zealand and near by islands. ‘The following 7 nominal

species are included in this genus: C. ecymosus (Lee, 1966) comb.n.,

C. dumosus (Lee, 1966) comb.n.; C. leptosceles (Lee, 1966) comb.n.;

92 REC, 8S. AUST. MUS., 16 (3); 1-219 July, 1970

C, mansoni (Lee, 1966) comb.n.; C. nucilis (Lee, 1966) comb.n.: C. validus

(Lee, 1966) comb.n.; C. watseni (Hirschmann, 1966) comb.n. for

Gamasiphis watsoni Hirschmann, 1966,

Genus GEOGAMASUS gen.n,

Type-species; Geoagamasus skoshi sp.n.

DIAGNOSIS. Minute or small mites, only the male having extensive

shields, May be holonotal shield or separate or partially separate podonotal

and opisthonotal shields. Female ventro-anal shield never fused to notal

shield and peritrematal shield only fused to notal shield by a narrow anterior

strip. Male ventro-anal shield and peritrematal shield broadly fused to

notal shield. Sterno-metasternal shield is not fused to endopodal IV shield.

Dorsal setae simple or larger setae may be inconspicuously pilose, Sperma-

thecal ringed tube opens at proximal end of trochanter HII. Spermadactyl

able to coil up around three hinges. Pretarsus | pedunculate and smaller

than other pretarsi. On male tarsus LI, seta ad3 modified to a tubercle,

MORPHOLOGY,

SCLEROTIZATION, Female: Holonotal or separate podonotal and

opisthonotal. Discrete ventro-anal, Small, discrete shield only bearing

aciculae, Small shield with pore, which is either discrete or fused to exopodal

IV, may be regarded as metapodal. Peritrematal reduced and narrowly

fused to exopodal LV. Split in exopodal [If and there may be split in

exopodal IV. Sterno-metasternal fused to endopodal II and part of IL.

Single pair of pre-endopodals.

Male; Notal may be as female, or, if holonotal on female, male may

have separate or partially separate podonotal and opisthonotal. Ventro-anal

fused to opisthonotal, exopodal IV, sometimes to peritrematal, but not

sternito-genital. Peritrematal fused to notal along its entire length.

CHAETOTAXY. Idiosoma: 6/, 62s, 4 or 5s, 4 or Sri 5J, 5Z, 5S, SR:

Sst: 3/y, 32Zv, 1S8y,

Legs: As Gamasellus,

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows. Tectum anterior margin basically trispinate with central spine

broader distally, and spinules may make the margin complex. Movable

digit with 3 or 4 teeth, On palp genu, seta a/l with 5 to LO lateral prongs,

und seta a2 spatulate. Dorsal setae tapering, simple or slightly pilose,

Shields shiny or pale. Spermathecal ringed tube probably opens proximally

on postero-lateral surface of trochanter III and usually coils around within

coxa II, Pretarsus | pedunculate and smaller than other pretarsi. Pulvilli

LEE—RHODACARIDAE 93

O20

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152, 153

156

155, 157

158, 161

a 154

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Figs. 152-161. Geogamasis Skoshi sp.n..

152-155 and 159, female: 152, soma, dorsum; 153, idiosoma, venter; 154, leg IV

(part), dorsum: 155, gnathosoma, venter; 159. spermathecal ringed tube in coxa and

trochanter (Il. 156-158, 160 and 161, male: 156, idiosomua, yenter; 157, chelicera;

158, tectum; 160, leg He (part): 161. gnathosoma and palp (part), venter.

94 REC. 8. AUST. MUS., 16 (3): 1-219 July, 1970

162, 163

166

164, 167

165, 169

Figs. 162-171. Geeganiasus howardi sp.n..

162-165 and 169, female: 162, soma, dorsum; 163, idiosoma. yenter: 164, gnathasenia.

venter: 165, leg IV (part). dorsum, 169, spermathecal ringed tube in coxa and

trochanter [Il 166-168, 170 and 171, male: 166, idiosoma, venter, 167, chelicera:

168, gnathosoma and palp (part), venter: 170. leg I (part); 171. tectum,

LEE—RHODACARIDAE 95

H-IV with lateral lobes lanceolate and slightly longer than central circular

lobes, or short and inconspicuous. Amongst dorsal setae on tarsus TV, seta

pal3 is the longest and setose.

Male: Corniculi attenuated and Jong compared with female. Tectum

differs from female. Movable cheliceral digit with single tooth or no teeth

and broadly fused to long sinuous spermadactyl, which can coil up around

3 hinges. Palp setae may be modified compared with female. On leg II,

femur setil av’, genu seta av, tibia seta av and tarsus seta ad3 enlarged to

spurs or lubercles, there are also non-setous processes, and other setae may

be modilied.

DISTRIBUTION, NTc: Aa. All published records are with the

original descriptions of nominal species,

Found i moss and plant litter,

REMARKS, Of the first two Geogamasus species described. the one

with a single adult holonotal shield was placed in Hydrogamasus and the one

with separate podonotal and opisthonotal shields was placed in Gamasellus.

Both species were placed in Gamasellus (Hydrogamasellus) by Hirschmann

(1966) who defined the subgenus Hydrogamasellus as having females and

males with one dorsal shield. Athtas-Henriot (in press) refers to species

of Queenslandolaelaps from South America which probably belong to this

genus, The location of the opening of the spermathecal ringed tube and

the form of the spermadactyl in this genus are unique among rhodacarids,

It these characters are disregarded there are many similarities between this

genus and Neogamasellevuns, The following 4 nominal species are included

in this genus; G. skoshi spn: G, coxalis (Sheals, 1962) comb.n.: G.

delamare? (Sheals, 1962) comb.n.; G. howard? sp.n..

Geogamasus skoshi spn.

FEMALE. Pig. 152-155, 159, Idiosomal length, 450».

MALE. Fig. 156-158, 160, 161. Idiosomal length, 4104. The notal

shield is similar to the female excepting that the podonotal and opisthonotal

shield are separate, the lateral extremity of the fissure being shown on the

drawing of the venter (Fig. 156). Seta ad3 (Fig. 160) is labelled “ad2”.

LOCALITY. The holotype temale, allotype male. paratype female

(N1968215) and paratype male (N1968216) drawn or examined: South

America; University grounds, Tucuman, Argentina, 11.1957, col. P-

Wygodzynsky, dep, holotype and allotype in BM(NH) paratypes in SAM.

REMARKS. The female of this species can be distinguished from

G. coxaliy and G. howardi by its entire dorsal shield, and from G, delamarei

by its sterno-metasternal shield which is more extensive posteriorly, lying

close to endopodal LV shield, The male is very similar in the distribution

6 REC, 8, AUST, MUS,, 16 (3); 1-219 July, 1970

of shields to both of the other two nominal males (paratype male of

G, delamarei, \961,6,20,18, dep, BM(NH), has been examined and it has

the podonotal shield and opisthonetal shield partially separated by lateral

incisions and the peritrematal shield is separate from the cxopodal shields

except posterior to stigma), The male can be distinguished from that of

G. delamarei by the podonotal and opisthonotal shields being completely

separated by a fissure, and also by having a tooth on the movable cheliceral

digit and lucking the process on the spermadacty! and movable digit which

is present on G. delamarei, The male can be distinguished from that of

G. howardi by seta al on femur Il not being enlarged, bent and spatulate.

Geogamasus howardi spn.

FEMALE, Fig. 162-165, 169. Idiosomal length, 300p.

MALE, Fig. 166, 168, 170, 171. Idiosomal length, 2702, The

inset drawing of the region around the stigma in Fig. 166, shows part of a

less sclerotized specimen. Seta ad3 (Fig. 170) is labelled “ad2”.

LOCALITY. The holotype female (N1968217), allotype male

(N1968218), 3 paratype females (N1968219, N1968220 and another) and

2 paratype males (N1968221 and another) drawn or examined; Aastralia;

plant litter and soil, Pinus radiata torest, Mt. Burr, South Australia,

30,5.1966, col. G. W. Howard, dep. SAM (N1968217-N1968221) and

BM(NH) (other 2 specimens).

REMARKS, The female of this species can be distinguished from

G, skeshi and G, delamarei by its having separate podonotal and opisthonotal

shields and from G. coxalis by the complex coiling of its spermathecal ringed

tube. The male is easily recognized by seta a/ on femur IL being enlarged,

bent and spatulate.

Genus HEYDENIELLA Richters

Heydeniella Richters, 1907, p. 281. Type-species: Heydeniella crozetensis

Richters, 1907, by monotypy,

DIAGNOSIS. Small to large mites, with a considerable range in the

extent and sclerotization of shields, Holonotal shield except for the male

of an unnamed species belonging to the crozetensis-complex, which has a

separate podonotal and opisthonotal shield, Usually ventro-anal shield of

female not fused to the notal shield, while on male it is fused to notal shield,

but such u fusion may not occur in either sex or it may occur in both sexes.

If female ventro-anal shield fused to the notal shield, then the sterno-

metasternal shield also fused to the endopodal IV shield, Male ventro-anal

shield always fused to the exopodal IV shield and peritrematal shield, Dorsal

setue ulways simple. Spermathecal ringed tube opens distally on dorsal

LEE—RHODACARIDAE 97

surface of trochanter IH, Pretarsus T pedunculate and conspicuously smaller

than other pretarsi. On male tarsus II, seta av2 may or may not be

spine-like.

REMARKS, Heyvydeniella was established to contain one species,

H, crozetensis, and was regarded as synonymous with Gamasiphis by

Triigardh (1907) when he described the male of a new species as Gamasipliis

loricatus. In fact, neither of these species belong to Gamasiphis, and they

probably both belong to Heydeniella. Unfortunately, the type-species of this

genus has been lost, but the description of it (Richters, 1907) is complete

enough tor it to be likely that a neotype can be designated when further

collections have been made on the Crozet Islands. It is certain that the

type-species belongs to the Ologamasini and it is likely that it is closely allied

lo u group of species with less extensive shields on the female, the dentata-

complex, that are common in Australia and New Zealand. All females of

the dentafa-complex have a spermathecal ringed tube that opens distally on

trochanter III]. Species have been examined from the antipodean Sub-

antarctic Islands that appear to have a spermathecal ringed tube opening

distally on trochanter TIT, are similar to the dentata-complex species, and

in some cases have eXtensive shields on the female similar to those described

for HW. erozetensiy. UL have, therefore, assumed that the spermathecal ringed

tube opens distally on the female trochanter IIL of H. crozetensis, and T am

using Heydeniella to contain only species with this character though it also

occurs in Pyriphis species,

| recognize 2 species-complexes within this genus, crezetensis-complex

and dentata-complex, which can be distinguished by the following key.

The morphology and distribution of what are assumed to be members of

this genus are given under the species-complex headings.

KEY TO SPECTES-COMPLEXES OF HEYDENIELLA

1, On palp genu, if seta a/l2 broadens distally

it is spatulate, broadening out to suboyal

shape, and seta a/l often has less than 12

lateral prongs ,. 00, 0-2 2. ee we ee ee Crezetensis-complex

On palp genu, seta a/2 cuncate, broadening

out in distal half to inverted subtriangular

shape, and seta a/l always has at least 12

lateral prongS .. 6) 6) ve ve ee ee ee © dentata-complex

CROZETENS]S-complex

DIAGNOSIS. As for genus. On palp genu. seta a/2 lanceolate or

spatulate. Spermadactyl always fused to movable cheliceral digit throughout

section level with proximal tooth.

OR REC, S, AUST, MUS., 6 (3): 1-219 July, 1970

MORPHOLOGY,

SCLEROTIZATION, Female: Holonotal which may or may not be fused

to ventro-anal. Tf ventro-anal is fused to holonotal then it is usually

also fused to peritrematal, Aciculae may be on ventro-anal or on small

discrete shield. There is sometimes a small shield lateral to anterior margin

of ventro-anal which may be regarded as a metupodal. Exopodals IH, Ul

and [¥ split. Sterno-metasternal fused to endopodal I and part of HI or,

in species with ventro-anal fused to holonotal, it is fused to endopodal I,

HI and IV. Single pair of pre-endopodals.

Male: Rarely there is a separate podonotal and opisthonotal. Ventro-

anal may or may not be fused to notal, and sternito-genital, but it 1s always

fused to peritrematal and exopodal LV.

CHAETOTAXY. Idiosoma: 6/, 6z, 48, 4r: SJ, 527, 58, SR: Sst) By,

3Zy, Sv.

Legs: As Gamasellus or rarely with one less ventral on yvenu TV

OTHER CHARACTERS, Female: Deutosternal denticles in & horizontal

rows. Tectum anterior margin either basically unispinate with lateral

spinules or trispinate. Movable cheliceral digit with 3 teeth. On palp genu,

seta all pectinate with 3 to 12 lateral prongs and seta a/2 is usually spatulate

but may be lanceolate. Dorsal seta simple, tapering. Shields shiny.

Spermathecal ringed tube is considered to open distally on dorsal surface of

trochanter II], Pretarsus | pedunculate and smaller than other pretarsi.

Pulvilli H-TY with lateral lobes that are usually attenuated and longer than

central circular lobes. Amongst dorsal setae on tarsus IV, setae pd3 is

setose and at least twice as long as any other seta,

Male: Corniculus slightly attenuated compared with female. Movable

cheliceral digit with 1 tooth, or rarely 2 teeth, and fused to similarly shaped

spermadactyl that is subequal in length or slightly longer or shorter. The

spermadactyl is fused to the movable digit along most of its Jength in some

species, and always as far as section level with proximal tooth. On Jeg IT,

femur seta av is always enlarged into a conspicuous spur, but femur seta py,

genu seta av and tibia seta av may be enlarged into a spine or spur. Seta

av2 on tarsus UI is not conspicuously spine-like.

DISTRIBUTION, Sm, Sk, Sa. Besides the locality records published

with the original descriptions of the two nominal species | have examined

unnained species from Atickland and Campbell Islands to be dep, BBM.

Found umongst lichens, moss or plant litter, or under stones.

REMARKS, The crozefensis-complex contains species that may have

much more extensive shields on the female than on the females of the

dentata-complex, while other species may be difficult to distinguish from

LEE—RHODACARIDAE

Figs, 172-181.

172-177, Heydeniella loricata (Triigirdh), male: 172,

No Scale

176

172

177

(78

179

retool

173

174

180

18)

idiosoma, venter; 173,

gnathosoma, venter; 174, femur and genu If; 175, pretarsus I and tarsus I (part);

176, tectum; 177, tarsus TV, antero-latus, 178-181, Neovaniasellevans berlesei

(Womersley), female: 178, soma, dorsum; 179, idiosoma, venter: 180. tarsus TV,

dorsum; I&81, gnathosoma. venter.

100 REC, 8. AUST, MUS., 16 (3): [-219 July, 1970

members of the dentata-complex. The unnamed species trom Auckland and

Campbell Islands, which | have examined and regard here us being members

of the crozetensis-complex, are a morphologically diverse group of species.

When the latter species, the type-species and possibly other unknown species

have been described or redescribed, it may be found that it is not expedient

lo split this genus into the two species-complexes delineated here. The

reason for the division given here is to separate the well known nominal

species, which represent a successfully, Jarge group of species in the

Australian region, from the poorly known species from the Subantarctic

region, which | may have incorrectly allied to H, crozerensis because | know

so few of the latter’s characters. The following 2 nominal species are

included in this species-complex: H. crezentensiy Richters, 1907; H. loricata

(Trigardh) comb.n, for Gamasiphis loricatus Tragirdh, 1907, Six unnamed

species from Auckland and Campbell tstands are described by Hunter and

Lee (manuscript), dep, BBM.

Heydeniella crozetensiy Richters

Hevdeniella crozetensis Richters, 1907, p. 281.

FEMALE. Fig. none. Approximate length, 1,120.2. Tectum anterior

margin basically unispinate with lateral spinules. Movable cheliceral digit

with 3 teeth, Holonotal shield fused to ventro-anal shield, Peritrematal

shield fused to exopodal shield and although it extends posterior to. this

fusion it does not fuse with ventro-anal as in two unnamed species with fused

holonotal and ventro-anul shields from Auckland and Campbell Islands.

Split in exopodals II, I and £V, Sterno-metasternal shicld fused to exopodal

shields Il, IIT and LV. Single pair of pre-endopodal shields. Idiosomal

setae short.

MALE. Fig. none. Approximate length, 1,120». Movable cheliceral

digit with | tooth and fused at base to similarly shaped but slimmer and

shorter spermaduactyl, Ventro-anal shield broadly fused to all surrounding

shields except perhaps the sternito-genital shield. On leg H, feniur seta av is

enlarged into a spur which is not wide ut the distal end as on the /ydro-

gamaselluy males, and some setae On the genu and tibia are spine-like.

LOCALITY, Possession Island, one of the Crozet Islands in’ the

kerguelenian Subantarctic region, col. German Antarctic Expedition,

1901-1903,

REMARKS. The types of this species, and other mites collected by

the same Expedition, were probably destroyed with the rest of Richter’s

collection, by the British Army occupying Germany just after the Second

World War (Sellnick, personal communication, 24.4.1966), The ubove

description is based only on Richter’s original description, which is

accompanied by a photograph of the female venter,

LF R—RHODACARIDAE 11

Heydeniella loricata Tragardh, comb.n.

Gamasiphis lovicatus Tragardh, 1907, p. 10,

FEMALE, Not known.

MALE. Fig. 172-177. Idiosomal length, 810. The sclerotization

is similar to that of H. crozetensis except that the sternito-genital shield is

certainly not fused to the ventro-anal shield. The chaetotaxy of the holo-

notunt has not been reconstructed from the fragments of this part of the

idiosoma. All idiosomal setae are simple and the range of their lengths does

not extend beyond that of those drawn (Fig. 172). Except for the centre

of the podonotal shield most shields bear reticulations. Setae on tibia and

tarsus IT are simple, unmodified.

LOCALITY. The holotype male drawn; Falkland Islands; under

stone, east of Port Stanely, 25.2.1902, col. Swedish South Polar Expedition

of 1901-1903, dep. NRS,

DENT AT A-complex

DIAGNOSIS. As for genus, except there is always a holonotal shield,

and the female ventro-anal shield always discrete and sterno-metasternal

shield never fused to endopodal IV shield. On palp genu, seta a/2 cuneate,

Spermadactyl never fused to movable cheliceral digit throughout section level

with proximal tooth.

MORPHOLOGY.

SCLEROTIZATION. Female: Holonotal, Ventro-anal discrete. Aciculae

on small discrete shield. Metapodal may be discrete or included in ventro-

anal. Peritrematal fused to exopodal TV. Exopodals If, ILL and IV split.

Sterno-metasternal fused to endopodal If and much of TI, Single pair of

pre-endopodals,

Male: Ventro-anal usually fused to holonotal, always fused to. peri-

trematal and exopodal IV, and separate from sternito-genital,

CHALTOLAXY, Idiosoma: 6/, 62, 4s, 4r: 5/, 52, 58, SR: Sst: 34,

3Zy, ISv.

Legs: Only on the single species from the Northern Hemisphere is it

as Gamasellus, usually there is one less ventral on genu TV (2, 5/1. 1).

OTHER CHARACTERS, Female: Deutosternal denticles in 6, 7 or 8

horizontal rows. “Tectum anterior murgin unispinate with prominent, narrow

central spine, and spinules on or near spine. Movable cheliceral digit

with 3 teeth. On palp genu, seta a/l pectinate with at least 12 lateral

prongs and seta a/2 broadly cuneate in distal half. Dorsal setae simple,

tapering. Shields shiny, usually reticulated, but may be smooth centrally

on sternum and podonotum. Spermathecal ringed tube opens distally on

102 REC, $8. AUST. MUS., 16 (3); 1-219 July, 1970

182 183

1004 184 188

165 186

167 189

190 19}

192

Figs. 182-192. HMeydeniella dentata (Womersley ).

182-186, 188-190 and 192, female: 182, soma, dorsum: 183, idiosoma, venter; 184,

leg [V (part), dorsum; 185, coxa and trochanter [Hl and LY; 186, ventral setae on

tibia IV; 188, tarsus I; 189. pretarsus [ and 1V; 190, trochanter TV of specimens from

Wilmington: 192. gnathosoma, venter. 187 and 191, male: 187, trochanter-femur

IV joint; 191, trochanter-femur LY joint of specirnens from Wilmington.

LEE—RHODACARIDAE 103

193, 194

197, 198

195, 196

1004

Figs. 193-199, Meydeniella dentata (Womersley ).

193-196, male: 193, idiosoma, venter; 194, idiosoma, latus: 195, gnathosoma, venter:

196, leg Il (part), 197-199, deutonymph: 197, idiosoma, dorsum; 198, idiasoma,

venter; 199, pretarsus I and LY,

104 REC. S. AUST, MUS., 16 (3); 1-219

200 “4 : 202 _

July, 1970

: j

vu 6&6 @

vee ;

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van

204, 206

202, 203

100p 207, 208

san

Figs. 200-209. Heydeniella dentata (Womersley).

200-203, protonymph: 200, soma, dorsum; 201, idiosoma, venter, 202, gnathosoma

yenter; 203, lez LV (part), dorsum, 204-208, larva: 204, soma, dorsum; 205,

pretarsus [; 206, idiosoma, venter; 207. leg Il (part), dorsum; 208, goathosoma

venter. 209, exg.

LEF—RHODACARIDAE 105

dorsal surface of trochanter [I]. Pretarsus | pedunculate and smaller than

other pretarsi, Pulvilli 11-I'V with lateral lobes always attenuated and longer

than central circular lobes. Proximal segments on legs HL and TV often have

non-setous processes. Amongst dorsal setae on tarsus TV, seta pd3 is setose

and al least twice as long as any other seta.

Male! Corniculus slightly attenuated compared with female. and on a

raised base. Movable digit usually with 1 tooth, but it has 2 tecth on a

single species from the Northern Hemisphere, and fused at base to spermia-

dactyl of similar length, but may be slightly longer or shorter. Spermadacty!

is never fused with movable digit as far as section level with proximal tooth.

On leg IT, femur seta av and pvl, genu seta av, Ubia seta av and tarsus seta

av2 enlarged into spurs or spines, may also be non-setous spurs on femur,

genu and tibia.

DISTRIBUTION, Pe: Aa, An. All records are published with the

origiaal descriptions of the nominal species. Many unnamed species have

been seen from Australia, Lord Howe Island, New Caledonia and New

Zealind, dep, SAM.

Found in moss, plant bitter and soil. The single species from the

Northern Hemisphere was found amongst mould in a chalk mine.

REMARKS. The first four dentata-complex species described were

placed in Hydrogamasus, then Hirschmann (1966) realized that these species

are not congeneric with the type of that genus and placed them in a new

subgenus, Gamasellus (Hydrogamasellus), for which the type was G. (H.)

antarcticus. The species of the dentata-complex differ from H, antarcticus

in having the spermatheeal ringed tube opening distally on trochanter IIT

and the dorsal seta ad3 on the male tarsus If not being enlarged. Since

only a few characters of H. crozetensis (type-species of Heydeniella) are

known, the placing of the dentata-complex in Heydeniella is tentative and

results from the study of unnamed species from Auckland and Campbell

Islands which are here considered as belonging to the crozetensis-complex.

The following 6 nominal species are included in this species-complex:

H. dentata (Womersley. 1942) comb.n., H. australica (Womersley, 1942)

comb: H. gee? span Ho markmitchelli sp.n., H. relata (Womersley. 1942)

comb.n.; J. relicta (Womersley. 1942) comb... The species H, relicta has

# variety majar Womersley, 1942.

Heydeniella dentata (Womersley ) comb.n,

Hydregamasus dentatus Womersley, 1942, p. 149-

FEMALE. Fig. 182-186, 188-190 and 192. Idiosomal length, 700-

The lex chaetotaxy differs from Gamasellus in having only one ventral seta

on genu TV (2, 5/1. 1), Specimens from the Flinders Ranges and the

REC. S. AUST, MUS., 16 (3): 1-219 July, 1970

Figs. 210-219. Heydeniella goei sp.n..

210-214, female; 210, soma, dorsum; 211, idiosoma, venter; 212, gnathosoma,

venter; 213, trochanter and femur IIL; 214, leg IV (part). dorsum. 215-219, male;

215, leg [Il (part); 216. idiosoma, venter; 217. idiosoma, latus; 218. trochanter and

femur JIT; 219, chelicera.

LEE—RHODACARIDAE 107

6 -90—_p 0-9

{

220

22)

{OOu 222

225

Figs. 220-226. Heydeniella markmitehelli sp.n..

220, 221, 224 and 225, female: 220, soma, dorsum: 221, idiosoma, venter; 224.

gnathosoma, venter; 225, leg [V (part), dorsal setae only, 222, 223 and 226. male:

222. idiosoma, venter: 223, leg Il (part): 226. chelicera,

108 REC. S. AUST. MUS.. 16 (3): 1-219 July, 1970

Hummock Ranges (localities which are 80-180 miles north-west of the type

locality) have a shorter ‘tooth’ on trochanter TY (Fig. 190).

MALE. Fig. 187, 191 and 193-196, Idiosomal length, 690". There

is a conspicuous rugose tubercle on the distal edge of the ventral surface of

the palp femur and genu which do not show clearly in the drawing (Fig. 195),

DEUTONYMPH. Fig. 197-199. Idiosomal length, 580».

PROTONYMPH.. Fig. 200-203. Idiosomal length, 480;.

LARVA. Fig. 204-208. Idiosomal length, 360.

EGG. Fig. 209. Longest axis length, 200p.

LOCALITY. Two females (N1968108 and N1968109), males

(NI96SLLO and N1968111), deutonymph (N1968112), 2 protonymphs

(N1I968113 and NI968114), larva (N1968115) and egg (N1968116)

drawn: Australia; LF134, moss and litter, near tree-fern at edge of pond,

Waterfall Gully, Mt. Lofty Ranges, near Adelaide, South Australia,

30.5.1966, col. D. C. Lee, dep SAM (immature stages not collected direct.

but bred from females in this sample).

The holotype female (N1968117), allotype male (N1968118), deuto-

nymph (N1968119) and protonymph (N1968120) examined: Australia:

moss, Long Gully, Mt. Lofty Ranges, near Adelaide, South Australia 8.1938,

col. H, Womersley, dep. SAM.

Female (N1968121) and male (N1968122) examined and base of leg

IV drawn: Australia; LF133, moss off rocks, Stony Creek, Flinders Ranges,

nr. Wilmington, South Australia, 25.5.1966, col. H. M. Cooper, dep. SAM.

REMARKS. In figured chaetotaxy patterns, filled in circles represent

setae considered to have been added at the previous moult. Characters of

the immature stages have not been used in the classification given here. The

extensive drawings of these stages are given only to increase this type of data

which is too limited from this family for satisfactory comparative studies.

Similur drawings are made in this paper of the immature stages of

Fydrogamasus littoralis and Gamasellus tragardhi,

Heydeniella goei spn.

FEMALE. Fig. 210-214. Idiosomal length, 690,.. The spermathecal

ringed tube opens in a similar position to that of Heydeniella dentata

(Fig. 185). Leg chaetotaxy differs from all other dentata-complex species

that | have seen, in being as Gamasellus, having 2 ventral setae on genu IV

(2, 5/2, 1).

LEERE—RHODAC ARIDAE 109

MALE. Fig. 215-219. Idiosomal length, 620», Unusual for

Heydeniella in having 2 teeth on movable, cheliceral digit, the anterior one

being ussociated with a lateral process on the digit, The posterior process

on genu IL is non-setous.

LOCALITY, The holotype female and allotype male drawn: England;

under mats of fur-like mould, on decaying wooden pit props, in chalk-mine,

Chislehurst Caves, Kent, 20,2.1955, col, P. N. Lawrence, dep. BM(NH).

REMARKS. This species can be distinguished from other nominal

species in the dentata-complex by the chactotaxy of genu IV, the proximal

protruberance on the female femur IL and the two teeth on the male movable

cheliveral digit.

Heydeniella markmitchelli spa.

FEMALE. Fig. 220, 221, 224 and 225, Idiosomal length, 500,

The spermathecal tube opens in a similar position to that of Heydeniella

dentata (Fig. 185). The leg chaetotaxy differs from Gumasellus in having

only one yentral seta on genu TV (2,5/1, 1).

MALE, Fig. 222, 223 and 226. Idiosomal length, 400p.

LOCALITY. The holotype female (N1968123) and 2° paratype

females (N1968124 and N1968125) drawn and examined: Australia; moss,

2,000ft. Flinders Ranges, near Wilmington, South Australia, 25.9.1958, col.

H, M, Cooper, dep. SAM.

The allotype male (N1968127) and 2 paratype males (N1968128 and

N1968129) drawn and examined: Australia: moss, 1,064ft, Hammock

Ranges, near Bute, South Australia, 9.9.1958, col, H. M. Cooper, dep. SAM,

A paratype female (N1968126) and paratype male (N1968130)

exumined: Australia; LF133, moss off rocks, Stony Creek, 1.200ft, Flinders

Ranges, near Wilmington, South Australia, 25.5.1966, col. H. M, Cooper,

dep. SAM.

A female (N1I968131) examined; New Zealand; off Uloma

tenebrionoides (tenebrionid beetle), Kaitoke, North Island, 3.7.1960, col.

D.C.M. Manson, dep. SAM.

REMARKS. The gnathosoma of this species is relatively large while

the sclerolization is reduced, This species can be distinguished from other

nominal species in the dentata-complex by setal row R being on the striated

cuticle instead of the holonotal shield and by the male ventro-anal, which is

separate from the holonotal shield. Since making the drawings, I have seen

the female und male specimens from Stony Creek that indicate that at the

locality where the holotype female was collected members of this species

ire larger than at the locality where the allotype male was collected.

110 REC, S, AUST, MUS., 16 (4): 1-219 July, 1970

Genus HYDROGAMASELLUS Hirschmann

Hydrogamasellus Hirschmann, 1966, p. 7. Type-species: Hydrogamasus

antarcticus Tragirdh, 1907, by original designation,

DIAGNOSIS. Small to large mites, with a considerable range in the

extent and sclerotization of shields. Usually holonotal shield, but may be

separate or partially separate podonotal and opisthonotal shields, Usually

ventro-anal shield of the female not fused to the notal shield, while on male

it is fused to the notal shield, but such a fusion may not occur in either sex

or it may occur in both sexes. If female ventro-anal shield is fused to the

notal shield then, as on other females in this genus, sterno-metasternal shield

not fused to endopodal IV shield. Male ventro-anal shield always [used to

exopodal IV shield and peritrematal shield. Dorsal setae always simple.

Spermathecal ringed tube opens at distal end of coxa III. Pretarsus I

pedunculate and conspicuously smaller than other pretarsi. On male tarsus

II, seta ad3 modified, not setose,

MORPHOLOGY.

SCLEROTIZATION, Female; Holonotal or rarely a separate podonotal

and opisthonotal. Ventro-anal may or may not bear aciculae, include

metapodal or be fused to notal, Aciculae may be on small discrete shield.

Peritrematal fused to exopodal TV, and may extend well behind this point.

Exopodals HU, Il and [V split. Sterno-metasternal fused to endopodal I

and part of III. Single pair of pre-endopodals.

Male; Notal as above, except that on the male of an unnamed species

the podonotal and opisthonotal are partially separated. Ventro-anal usually

fused to notal, always fused to peritrematal and exopodal ['V, and separate

from sternito-genital.

CHAETOTAXY. Idiosomu: 6), 62, 5s, 56: SJ, 527, 58, SR: Sst: 3/y,

3Zv, 248v,

Legs: Usually as Gamasellys, but there is one less ventral on genu [V

(2,5/1, 1) in an unnamed species.

OTHER CHARACTERS, Female: Deutosternal denticles in & or 9

horizontal rows. Tectum anterior margin basically unispinate, with broad

base to spine which bears small spinules or rarely (as in A. richtersi) quin-

quispinate with complex, broad tip to central spine. Movable, cheliceral

digit with 3 teeth. On palp genu, seta a/l pectinate with 5 or 6 lateral

prongs, and seta a/2 spatulate. Dorsal setae usually simple, slightly spatulate

or tapered, may be slightly pilose. Shields shiny and usually reticulate, while

LEE

RHODACARIDAE 11

sternal shield may have longitudinal striae. Spermathecal ringed tube

probably opens distally on dorsal surface of coxa HI. Pretarsus [ pedunculate

and smaller than other pretarsi. Pulvilli II-1V with attenuated lateral lobes

which may be shorter or much longer than central circular lobes. Amongst

dorsal setae on tarsus IV, seta pd3 is setose and at least twice as long as

any other seta.

Male: Corniculus attenuated and on raised process which may have

paraxial tubercle carrying seta Aypl. Movable cheliceral digit with one

tooth, attached at base to slim spermadactyl which may be subequal in length

or up five times as long. Tectum anterior margin may differ from female

and be trispinate. Often process on ventral surface of palp trochanter or

femur. On leg H, femur seta av and usually seta pvl, genu seta av, tibia

seta av enlarged into spur or spine, while tarsus seta ad3 and sometimes

seta al2 enlarged into a hollow spine or spur, or a rimmed pit. Non-setous

processes also occur on tibia and tarsus II.

227. A 228

iK \ *

ie \

Sst re ’ \ yous

cH dl ° \ \ AA,

5 [e) | \ } ,

\ | 90 \ f \ oe

i Lt ae

<9 i \

\9- loo a ° NeN 13

i | s

| RP oo PLL gos

(e fe) | / &

90 Woe Fp ay

if g 9 \

oO | \\ \ 4 ° \ cd

fe) \ | a | : Seer

| 'e% \ fe) 1 /

o. \b ?

No aSv \\ | \i/

f —27 1 / He Lo

100x| 230 5R | :

228, 229, 231 55 ¢> Ay

Figs. 227-231. Hydrogamasellus antarciicus (Trigirdh), female.

227. pretarsus | and tarsus [ (part). 228, soma. dorsum. 229, leg IV (part),

dorsum. 230. gnathosoma, venter. 231, idicsoma, venter.

112 REC, 8S, AUST. MUS., 16 (3): 1-219 July, 1970

Figs, 232-238. Hydrogamasellus cavei (Sheals).

232-234 and 238, female: 232, soma, dorsum; 233, idiosoma, venter; 234, leg IV

(part), dorsum; 238, gnathosoma, venter. 235-237, male: 235, idiosoma, venter;

236, leg IT (part); 237, chelicera, and gnathosoma and palp (part), venter,

LEE—RHODACARIDAE

Figs. 239-247. Hydrovamasellus coleopirants (Berlese ).

239-242. female: 239, idiosoma, venter; 240, tectum; 241, leg IV (part), dorsum;

242, gnathosoma, venter. 243-247, male: 243, chelicera: 244, tectum; 245,

gnathosoma and palp (part). venter; 246, lex Il (part): 247, idiosoma, venter.

114 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

248, 249, 250

252, 253

1oou

25!, 254

255, 256

Figs. 248-257. Hydrogamasellus racavitzai (Trouessart).

248-251, 256 and 257, female: 248, soma, dorsum; 249, idiasoma, yenter; 250, leg

IV (part). dorsum; 251, trochanter Iff and 1V, and other segments (parts); 256,

gnathosoma, venter; 257, pretarsus I, 252-255, male; 252, tectum; 253, idiosoma,

venter; 254, chelicera, and gnathosoma and palp (part). venter; 255, leg IT (part).

LEE—RHODAC ARIDAE. 11s

DISTRIBUTION, NTc: Sm, Sk, Sa: ACs. Besides the records

published with the original descriptions of the nominal species, H. racevitzai

is recorded from many other localities in the scotian Antarctic region,

HA. antarcticus is recorded from Macquarie Island (Hirschmann, 1966) and

unnamed males have been collected in Tierra del Fuego, dep BM(NH).

Found in moss, lichens, algae, plant litter and soil, and under stones,

REMARKS, Gamasellus (Hydrogamasellus) was originally diagnosed

(Hirschmann, 1966) us being Gamasellus-like, but having an entire dorsal

shield and simple dorsal setae, As diagnosed here, with generic status,

Hydrogamasellus has members with a considerable range in the extent and

degree of fusion of their idiosomal shields, including an entire or divided

dorsal shield, and with all simple or some slightly spatulate or pilose dorsal

setae but the location of the spermathecal ringed tube aperture and the male

gnathosoma und leg U characters are similar. Therefore, an unusually high

number of species have been removed from (see “Changes in Nomenclature” ),

or transferred to this genus. Because Hydrogamasellus is similar to

Heydeniella (type-species; H. crozetensis) both crozetensix species truns-

ferred lo Mydrogamasellus have been given new names. The following 8

nominal species are included in this genus: HA. antarcticus (Tragirdh, 1907);

H, cavei (Sheals) comb.n. for Ologamasus cavei Sheals, 1962; H. coleop-

fratuy (Berlese) comb.n. for Ologamasus coleoptratus (Berlese, 1888);

H. getussi noma. for Neoparasitus crozetensis Richters, 1907; H, racovitzai

(Troucssart, 1903) comb.n. for Gamasellus racovitza’ (Trouessart, 1903);

H. richtersi nom, for Gamasellus crozetensis Richters, 1907: H. striatus

(Sheals, 1962); HW. tepali (Balogh, 1963b). Two unnamed species (from

Argentina; Province of Neuquen or Tucuman, 3 or 4. 1959, col. C. Delamare

Deboutleville, dep, BM(NH)) which belong to this genus are listed so that

their characters, not found on nominal species of this genus, can be listed.

One Unnamed species (single female) has only one ventral seta on genu LV,

while the other (single male) has the podonotal and opisthonotal shields

partially separated by lateral incisions. AMydregameasellusy ubatubaensis,

incertae sedis, has been left in this genus tor convenience.

Hydrogamasellus antareticus (Triigirdh )

Hydrogamasus antarcticus Triagirdh, 1907. p, 12.

PEMALE. Fig. 227-231. Idiosomul length, 750. On right side,

seta Svl as off ventro-unal shield.

MALE. Fig. none. No males seen from Naturhistoriska Ricksmuseum,

Stockholm, but a full description is given in Hirschmann (1966) of con-

specilic specimens,

116 REC, S. AUST. MUS,, 16 (3): 1-219 Jiily, 1970

LOCALITY. The holotype female with two legs missing drawn:

Antarctica; wet moss, Paulet Island, between South Shetland Islands and

Palmer Peninsula, 15.1.1902, col. Swedish South Polar Expedition of

1901-1903, dep. NRS.

Hydrogamasellus cavei (Sheals) comb.n,

FEMALE. Fig. 232-234, 238. Idiosomal length, 510». Dorsal

surface strongly convex. Lateral longitudinal split in holonotal shield. On

tarsus IV, setae p/2 and p/4 are simple and not spine-like and pilose us in

H, racovitzat.

MALE, Fig. 235-237. Idiosomal length, 510”. Slight thickening on

ventral, distal edge of palp femur which may be equivalent to process in

H. racovitzai. On tarsus I], seta ad3 appears to be a hollow spine.

LOCALITY. A paratype female (1961.6,.20.22) and three males

examined or drawn: Argentina; litter under Nothophagus dombeyi, Nahuel

Huapi Reserve, Puerto Blest, 7.3.1959, col. C. Delamare Deboutteville,

dep, BM(NH).

Hydrogamasellus coleoptratus (Berlese) comb.n.

Hypoaspis coleoptratus Berlese, 1888, p. 199.

FEMALE. Fig. 239-242. Idiosomal length, 7O0Qu. Dorsal setae

difficult to examine, but opisthonotal setae mostly slightly lanceolate and as

long as the peripheral setae that are drawn, while podonotal setae (except

seta r2 which is similar to the peripheral opisthonotal setae) are shorter and

tapering like seta Zvl.

MALE. Fig, 243-247. Idiosomal length, 6202. Small process on

palp trochanter. Tarsus [I has a row of steeply rimmed pits on its dorsal

surface, the larger 2 pits are considered homologous with setae al2 and ad3,

LOCALITY. The ‘tipico’ female (34/32), ‘tipico’ male (34/32) and

female and male (202/15) examined or drawn: Argentina; under the bark

of trees, Buenos Aires, dep. SEAF.

Hydrogamasellus racovitzai (Trouessart) comb.n.

Gamasus racovitzai Trouessart, 1903, p. 8.

Zercon tuberculatus Triigardh, 1907, p. 17.

FEMALE. Fig. 248-251, 256 and 257. Idiosomal length, 960».

Pre-endopodal shields appear to be connected to sternal shield by a lightly

sclerotized and reticulated shield. Seta p/4 (Fig. 250) is labelled “p/3”.

MALE. Fig. 252-255. Idiosomal length, 870«. Large process on

palp femur, Distinct process on hypostome bearing seta Aypl. On tarsus

Il, seta ad3 appears to be a hollow cone, with thick vertical ridge in

posterior wall.

LEE—RHODACARIDAE 117

LOCALITY. Three females (N1968132-1968134) and three males

(N1968135-1968137) examined or drawn: South Orkneys; under stones on

shore, station 1089, Michelsen Island, 3.1.1933, col. Discovery Investigations

(British) of 1931-1933, dep. SAM.

REMARKS. The above locality is close to Laurie Island where

Gamasellus racovitzai neorcadensis Trouessart, 1912, was collected.

Probably this should not be a subspecies, the structure of seta av on the

male femur II (the character used to distinguish the subspecies) would

appear as the nominate subspecies if viewed laterally, and as the other

subspecies if viewed ventrally.

Genus NEOGAMASELLEVANS Loots and Ryke

Neogamasellevans Loots and Ryke, 1967a, p. 13. Type-species: Neogama-

sellevans preendopodalis Loots and Ryke, 1967a, by original designation.

DIAGNOSIS. Minute mites, only the male having extensive shields.

Holonotal shield. Discrete ventro-anal shield on the female, but it is fused

to the holonotal, peritrematal and exopodal IV shields on the male. Female

peritrematal shield joined only by a slight thickening of striated cuticle to

exopodal IV shield. Male peritrematal shield separated from exopodal IV

shield and notal shield at level of acetabulum IV, although all 3 shields are

fused to ventro-anal shield. Female sterno-metasternal shield not fused to

endopodal [V shield. Dorsal setae simple and tapered. Spermathecal ringed

tube opening not known. Spermadactyl short and straight. Pretarsus I

pedunculate and smaller than other pretarsi. On male tarsus II, all setae

are setose.

MORPHOLOGY.

SCLEROTIZATION. Female: Holonotal. Discrete ventro-anal. Posterior

to ventro-anal there is a small discrete shield only bearing aciculae.

Backward extension of exopodal IV, may have included metapodal.

Peritrematal reduced, joined by a slight thickening of striated cuticle to

exopodal IV. Exopodals II, II] and IV split. Sterno-metasternal fused to

endopodals I] and HI. Single pair of large pre-endopodal shields, paraxial

margins being equal in length to distance between setae sfl and s?f2.

Male: Ventro-anal is fused to exopodal IV, peritrematal and holonotal.

Peritrematal is not fused to exopodal IV, nor to holonotal between levels of

setae st4 and Zvl. Sternito-genital shield separate from ventro-anal and

from anterior section of endopodal II.

CHAETOTAXY. Idiosoma: 6/, 6z, 5s, Sr: S5/, 5Z, 5S, SR: Sst: 3/v,

3Zyv, 1Sv.

Legs: As Gamasellus.

OTHER CHARACTERS. Female: Deutosternal denticles in at least 6

horizontal rows. Tectum anterior margin basically trispinate with lateral

l1s REC, S. AUST, MUS., 16 (3): 1-219 duly, 1970

spinules and central spine broader distally with spinules making the margin

complex. Movable cheliceral digit with 3 teeth. On palp genu, seta

ail pectinate with 5 lateral prongs, and seta a/2 slightly lanceolate. Dorsal

setae, simple tapering. Shields slightly shiny and reticulated, Peritreme

short, not extending anterior to mid-acetabulum LIL, Spermathecal ringed

tube opening not known, Pretarsus I pedunculate and smaller than other

pretarsi. Pulvillt H-L1V with attenuated lateral lobes that are slightly longer

than central circular lobes, Amongst dorsal setae on tarsus TV, seta pd3 is

the longest and setose.

Male: Tectum anterior margin with the central spine shorter but

broader and more complex than on female, while the two lateral spines are

reduced to spinules, Movable cheliceral digit with one tooth, fused at base

to shorter similarly shaped spermadactyl, Dorsal setae on palp femur are

enlarged to spines. On leg I, femur, genu and tibia seta av enlarged into

spurs, also seta pvl on femur is a spine and there are non-setous processes

on the tibia,

DISTRIBUTION, NTc. The only published record is with the original

description of the single, nominal species.

Found in plant litter on soil.

REMARKS, Neogamasellevans is not here regarded as closely allied

to Gamasellevans as might be assumed from its name. Hirschmann (1968)

considers N. preendopodalis similar to Gamasellus (Hydrogamasellus)

delamaret (transferred above to Geogamasus) and therefore makes Neogama-

sellevans a junior synonym of Hydrogamasellus. 1 would agree it is similar

to Geogamasus, but regard the probable differences in location of the

aperture of the female spermathecal ringed tube and in the structure of the

male spermadactyl sufficient to refer the species to different genera, and to

suggest that these genera may not in fact be closely allied. Some of the

characters given for the genus were obtained by examination of 2 paratype

females and 3 paratype males (1967.9.7.39-43), dep. BM(NH), of the

single nominal species, N. preendopodalis Loots and Ryke. 1967a, included

in this genus. Queenslandolaelaps berlesei, incertae sedis, has been trans-

ferred to this genus because it should be in the Ologamasini, although it is

improbable that it belongs to this genus.

Genus PARASITIPHIS Womersley

Parasitiphis Womersley, 1956a, p. 535, Type-species; Parasitiphis littoralis

Womersley, 1956a, by original designation.

Austrohydrogamasus Hirschmann, 1966, p. 10, synn.. — Type-species:

Hydrogamasus (Austrohydrogamasus) watsoni Hirschmann, 1966, by

original designation.

LEE—RHODACARIDAE 119

DIAGNOSIS, Average sized to large mites, Holonotal shield. Discrete

ventro-anal shield in both sexes, except on some males where metapodal

shields and/or holonotal shield may be fused to the ventro-anal shield.

Peritrematal shield fused to exopodal TV shield on female but not on male.

Female sterno-metasternal shield not fused to endopodal TV shield. Dorsal

setae simple and tapering. Spermathecal ringed tube opens near posterior

paraxial edge of acetabulum IV. Movable cheliceral digit of male always

with more than one tooth and parallel spermadactyl that is subequal in

length. Pretarsus | not pedunculate and at least as large as the other pretarsi.

On male tarsus Il, seta av2 modified into a spur or tubercle,

MORPHOLOGY.

SCLEROTIZATION. Female: Holonotal, Discrete metapodal and ventro-

anal bearing aciculue posterior to anus. Peritrematal narrowly fused

to exopodal IV, Exopodals do not completely encase peraxial edge of

acetabulum I} nor usually of acetabulum 1, Sterno-metasternal fused to

endopodal If and part of HI. Single pair of pre-endopodals.

Male: Discrete sternito-genital shield, which tends to be unusually

narrow posteriorly, Ventro-anal posterior to seta Zv3 may or may not be

fused to holonotal. Metapodals may or may not be [used to the ventro-anal.

Peritrematal not fused to exopodal LV.

CHAETOTAXY. Idiosoma: 6), 5z, 5s, 4rz SJ, 57, 4 or 5S, 0 or 3R,

IWR: Set: 34, 3Zy, 28yv.

Legs: As Gamasellus or differs in being without a postero-lateral on

penu TV (2, 5/2. 0).

OTHER CHARACTERS, Female: Deutosternal denticles in 8 horizontal

rows. Tectum anterior margin unispinate; the spine being large and

having central ridge. Movable cheliceral digit with 4-7 teeth. On palp

genu, seta a/l and seta a/2 spatulate. Dorsal setae simple, tapered, Shields

shiny and reticulated. Spermathecal ringed tube opens near posterior

paraxial edge of acetabulum IV. Pretarsi sheath I not pedunculate, only

being long enough to contain the retracted claw-complex, which is large,

being subequal in size to other claw-complexes. Pulvilli HI-1V with lateral

lobes attenuated and subequal in length to central circular lobes. Amongst

dorsal setae on tarsus TV, seta pd3 is either just the longest or both setae pd3

and ad3 are the longest, and the former may be slightly pilose.

Male: Movable cheliceral digit with 2-5 teeth and fused at base to

spermadactyl of similar length, On leg IL femur seta av, genu seta av, tibia

seta ay and tarsus seta av2 enlarged into spurs. Bases of leg TTL and IV may

have conspicuous non-setous processes,

120 REC. S. AUST. MUS.. 16 (3); 1-219 July, 1970

a \)

JF 261

—0—_4

/ fe] Nez

a |

— wn

Figs. 258-266. Parasitiphis litturalis Womersley.

258-262, female; 258, soma, dorsum: 259, idiosoma, venter: 260, leg ITV (part),

dorsum; 261, pretarsus | and part of tarsus; 262, gnuthosoma, venter, 263-266,

male: 263, idiosoma, venter; 264, chelicera; 265, leg IT (part); 266, trochanter and

femur Il, and trochanter LV.

LEE—RHODACARIDAE 12]

a) o-— 9° ~o— fo)

267, 270

Figs. 267-271. Parasitiphis aurora sp.n., female.

247. soma, dorsum. 268, pretarsus [| and tarsus | (part). 269, leg EV (part),

dorsum. 270, idiosoma. venter. 271, gnathosoma, venter.

DISTRIBUTION. Aa, An: Sm, Sk, Sa: ACs, Besides the locality

records published with the original or the below descriptions of the nominal

species, Hunter (manuscript) records P. jeanneli (as H. (A.) watson) from

South Georgia. Found in or near the littoral zone.

REMARKS. Previously Parasitiphis contained one species. but I now

include species or misidentified specimens from Hydrogamasus, Gamasellus

und Laelaps. Laelaps brunneus Kramer is referred here on the basis of the

few characters given in the original description, because the only two

specimens (females), collected from the sea shore, Tierra del Feugo, by

Michaelsen, dep. AMH, cannot be found (Rack, personal communication,

21.4.1967). The only genus that can be considered a close ally is

Litogamasus, a new genus in the Gamasellini, which occurs in similar

habitats. Hirschmann (1966) described Parasitiphis jeanneli adults and

nymphs as Hydrogamasus (Austrohydrogamasus) watsoni; a new species

that was the type of the subgenus. Paravyitiphis is quite unlike Hydrogamasus,

but the larva described as H. (A.) watseni was that of Hydrogamasus

kensleri. The following 4 nominal species are included in this genus:

P. litteralis Womersley, 1956a; P. aurora spn; P. brunneus (Kramer)

comb.n, for Laelaps brunneus Kramer, 1898; P. jeanneli (Andre) comb.n.

for Gamasellus jeanneli André, 1947.

122 REC, 5. AUST. MUS., 16 (3): 1-219 July, 1970

Figs. 272-280. Parasitiphiy jeenneli (André).

272-276, femate: 272, soma, dorsum; 273, idiosoma, yenter; 274, lez LV (part),

dorsum; 275, pretarsus | and tarsus | (part): 276, gnathosoma, venter. 277-280,

male: 277, idiosoma, venter; 278, chelicera; 279, leg Il (part); 280, trochanter III

and

LEE—RHODACARIDAE 123

Parasitiphis littoralis Womersley

Parasttiphis litteralis Womersley, 1956a, p. 536.

FEMALE. Fig. 258-262. Idiosomal length, 770u. Claw complex |

(Fig. 261) is shown partly extended, when retracted it appears to be similar

to that of the other nominal species (e.g., Fig. 268). Leg chaetotaxy as

Gamasellus.

MALE. Fig. 263-266, Idiosomal length, 740», Idiosomal length of

allotype male, 760).

LOCALITY. The holotype female (N1968160) drawn, 4 paratype

females (N1968162-N1968165) and allotype male (N1968161) examined:

Australia; littoral zone, mouth of American River, Kangaroo Island, South

Australia, 1.1946, col. H. Womersley, dep. SAM.

Two males (N1968166 and N1968167) drawn or examined: Australia;

on seaweed covered rocks, littoral zone, Myponga Cove, South Australia,

30, 10.1966, col. D. C. Lee, dep. SAM.

Parasitiphis aurora sp.n.

Hydrogamasuys antarcticus: Womersley, 1937 (not Tragiardh, 1907), p. 17,

syn.n,

FEMALE. Fig. 267-271. Idiosomal length, 860». Leg chaetotaxy

as Gamasellus except for genu 1V (2, 5/2, 0). Female from Gough Island

has 4 setae, and not just 3, in row R.

MALE. Not known when drawing made, but since seen from Auckland

and Campbell Islands, to be dep. BBM. Will be described later. The male

of this species is unusual in haying the ventro-anal shield posterior to seta

Zv3 fused to the holonotal shield,

DEUTONYMPH. On the basis of the lengths of the notal setae, form

of the dorsal setae on leg IV and the chaetotaxy of genu TV, the deutonymph

drawn with the description of H. antarcticus: Womersley, 1937, belongs to

this species, but the other deutonymph (N1968184), in the same collection,

belongs to P. jeanneli.

LOCALITY. The holotype female (N1968182) drawn and morpho-

type deutonymph (N1968183) examined; Macquarie Island; between tide-

marks, 28.11.1913, col. T. H. Johnston (on Australasian Antarctic

Expedition, 1911-1914), dep. SAM.

Female examined: Tristan da Cunha Group: with Perforgunia sp.,

Archway Point, Gough Island, col. M. W. Hollgate (Gough Island Scientific

Survey, 1955-56), dep. BM(NH).

124 REC. S. AUST. MUS., 16 (3); 1-219 July, 1970

REMARKS. This species can be distinguished from the other two

nominal species by the lack of a postero-lateral seta on genu IV (2, 5/2, 0),

and by the two antero-dorsal setae on femur IV being spine-like. The male

also differs in having the ventro-anal shield fused to the notal shield and in

lacking conspicuous non-setous processes on the proximal segments of logs

II] and IV,

Parasitiphis jeanneli (André) comb.n.

Gamasellus jeanneli André, 1947, p. 70.

Hydrogamasus (Austrohydrogamasus) watsoni Hirschmann, 1966, Dial

syn.n.

FEMALE. Fig. 272-276. Idiosomal length, 920». Leg chaetotaxy

as Gamasellus.

MALE. Fig. 277-280. Idiosomal length, 890. Dorsal setae 73 and

Z5 longer than on female. Ventro-anal shield in Heard Island males have

an unsclerotized patch, lateral to seta Zv2 (sce Fig. 277), which may be more

extensive than on specimen drawn, nearly separating off antero-lateral corner

as a metapodal shield. In Macquarie Island specimens the metapodal shield

is separated completely from the ventro-anal shield.

LOCALITY. Two females (N196168 and N1968169) and 2 males

(N1968170 and N1968171) drawn or examined: Heard Island, on vegeta-

tion, Rogers Head, 31.1.1952, col. K. Brown (on Australian National

Antarctic Research Expedition, 1947-1955), dep. SAM.

Four females (N1968172-N1968175) examined: Kerguelen Islands;

under stones, between tidemarks, 16.11.1929. col. British-Australian-New

Zealand Antarctic Research Expedition, 1929-31, dep. SAM.

Two females (paratypes of Hydrogamasus (Austrohydrogamasus)

watsoni, N1968176 and N1968177) examined: Macquarie — Island:

Pucinellia, Isthmus or Mid Royal Penguin rookery, North Head, 1961, col.

K. Watson, dep. SAM.

Two females (N1968178 and N1968179) and two males (N1968180

and N1968181) examined: Macquarie Island; Isthmus, 12.1949, col. T.M.

(on Australian National Antarctic Research Expedition, 1947-1955), dep.

SAM.

One deutonymph (N1968184), previously named Aydrogamasus

antarcticus: Womersley, 1937, examined: Macquarie Island: between tide-

marks, 28.11.1913, col. T. H. Johnston (on Australasian Antarctic Expdi-

tion, 1911-1914), dep. SAM.

LEE—RHODACARIDAE 125

REMARKS. Although there are differences in size and slight morpho-

logical differences between specimens collected in different parts of the

Subantarctic region, | consider that all the specimens with their collection

data given above are members of this one species,

Genus PYRIPHIS gen.n.

Type-species: Ologamasus pyrenoides Lee, 1966.

DIAGNOSIS, Average sized mites. Holonotal shield. Ventro-anal

shield broadly fused to notal and exopodal IV shield on female and notal,

peritrematal, exopodal IV and sternito-genital shield on male. Single pair

of pre-endopodal shields. At least two pairs of setae (25 and r3) on

podonotum are conspicuously pilose and surrounded by mainly simple or

lanceolate setae. Spermathecal ringed tube opens distally on dorsal surface

of trochanter HI. Spermadactyl strongly recurved. Pretarsus | pedunculate

and smaller than other pretarsi. On male tarsus II all setae are sctose.

MORPHOLOGY.

SCLEROTIZATION, Female: Holonotal, broadly fused to ventro-anal

which bears aciculae posterior to anus. Ventro-anal and peritrematal fused

to exopodal TV, Split in exopodals II, Hf and IY. Sterno-metasternal

fused to endopodal UH only. Single pair of pre-endopodals.

Male: As female but peritrematal and sternito-genital also fused to

ventro-anal,

CHAETOTAXY. Idiosoma: 6/, 6z, 5», Sr: 5/, 47, 5S, 8R, OUR: Sst:

3Jv, 3Zv, 2S8v,

Legs: As Gamasellius.

OTHER CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows, Movable cheliceral digit with 3 conspicuous teeth and a series of

smaller teeth. On palp genu, seta a/l with single lateral prong while both

setae a/l and a/2 are slightly spatulate. There are three pairs of podonotal

shield protruberances (bearing setae jl, 25 and r3). Some dorsal setae may

be simple, tapering, but some are lanceolate, or pilose and spatulate. Shields

are dull. Spermathecal ringed tube probably opens distally on dorsal surface

of trochanter III. Pretarsus I reduced and pedunculate. Lateral lobes of

pulvilli I-LV attenuated and longer than central oval lobes. Amongst dorsal

setae On tarsus TV, setae ad1 and pdl are the longest.

Male: Movable cheliceral digit with one tooth, fused at base to strongly

recurved spermadactyl. On leg II, femur setae av and py, genu setae av and

py, tibia seta ev enlarged into spurs or spines.

126 REC. S$. AUST. MUS., 16 (3): 1-219 July, 1970

DISTRIBUTION, Aa, Besides the locality published with the original

description of the type-species, members of this genus have been found in

other forests on the coastal ranges of Victoria, southern New South Wales

and Tasmania, dep. SAM.

Found in leaf litter.

REMARKS. Pyriphis is established for one species originally described

(Lee, 1966) as the only species in the Ologamasus pyrenoides-group. At

least two unnamed species have been seen since, dep. SAM, with the same

extensive sclerotization and unusual, strongly recurved spermadactyl. As in

some Gamasellus discutatus-complex species, members of this genus are

unusual in having both a holonotal shield and conspicuously pilose dorsal

setae amongst simple setae. Besides the holonotal shield, the single pair of

pre-endopodal shields and, more important, the location of the spermathecal

ringed tube aperture on the proximal leg segments support the placing of

Pyriphis in the Ologamasini. The following single nominal species is

included in this genus: P. pyrenvides (Lee, 1966) comb.n.

Genus RYKELLUS gen.n.

Type-species: Cyrtolaelaps (Gamasellus) darglensis Ryke, 1962c.

DIAGNOSIS. Large mites. Although there is a clear line on the

dorsal shield dividing it into podonotal and opisthonotal regions, it is more

like a strong reticulation marking than a fissure, and | would consider that

a holonotal shield was present. Ventro-anal shield of the female discrete,

while on male it completely merges with all the surrounding shields. Single

pair of pre-endopodal shields. Dorsal setae all simple. Spermathecal ringed

tube opens distally on postero-lateral surface of coxa [V. Pretarsus |

pedunculate and smaller than other pretarsi. On male tarsus IL all setae

are setose,

MORPHOLOGY.

SCLEROTIZATION. Female: Holonotal, with groove between podonotal

and opisthonotal regions. Discrete ventro-anal bearing aciculae posterior

to anus. No discrete metapodal. Peritrematal fused to exopodal IV,

Exopodals II and IV split. Sterno-metasternal fused to endopodal IU.

Single pair of pre-endopodals.

Male: Ventro-anal is fused to all the surrounding shields.

CHAETOTAXY, Idiosoma: 6/, 6z, 4s, 4r: SJ, 5Z, SS. SR: Sar: 3/v,

aZv, 28y.

Legs: As Gamasellus.

LEE—RHODACARIDAE 127

OTHER CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows, ‘Tectum anterior margin has single central spine with multispinulate

proximal sides, Movable cheliceral digit with 3 teeth. On palp genu,

sela all is spine-like or spatulate and seta a/2 is spatulate. Dorsal

setae simple, tapering. Shields shiny and reticulated, Spermathecal ringed

tube probably opens distally on postero-lateral surface of coxa IV. Pretarsus

I peduncluate and smaller than other pretarsi. Pulvilli I-IV with attenuated

lateral lobes that are longer than central circular lobes. Non-setous spur

inay be present on postero-lateral surface of trochanter 1V, Amongst dorsal

setae on tarsus IV, seta pd3 is the longest and setose,

Male: Corniculi on raised hypostomal process. Movable cheliceral

digit with | tooth and fused at base to slim, similarly directed spermadactyl

that is subequal in length. On leg H, femur seta ay, genu seta av and tibia

seta ay enlarged into spur, while femur seta pyl is spine-like and there may

be small non-setous processes on leg I segments.

DISTRIBUTION. Es. All records are published with the original

descriptions of the nominal species,

Found in plant litter,

REMARKS. Both Rykellus species were originally placed in Cyrto-

laelups (Gamasellus). Hirschmann (1966) transferred them to Gamasellus

(Hydrogamasellus), Because of the holonotal shield, simple dorsal setae

with only 20 podonotal pairs, spermathecal ringed tube aperture on proximal

leg segments and single pair of pre-endopodal shields, they clearly belong to

the Ologamasini in my classification. Because of the simple dorsal setae and

seta a/l on palp genu, the unique location of the spermathecal ringed tube

aperture and the fusion of the male ventro-anal shield to the sternito-genital

shield, T have established a new genus for them, I have examined two

paratype females of R. nkandhlaensis, dep. PUSA. The following 2 nominal

species are included in this genus: R. darglensis (Ryke, 1962c) comb.n.:

R. nkandhlaensis (Ryke, 1962c) comb.n,,

Tribe GAMASELLINI Hirschmann

Cyrtoluclaptini Berlese, 1913b, p. II.

(As indicated with the synonyms of the family nume, this family-group

name is not considered available. )

Gamasellini Hirschmann, 1962, p, 39,

Cyrtoluclapidae Berlese, sensu Johnston (in part), 1968, p. 19.

Type-genus; Gamaysellus Berlese, 1892d

DIAGNOSIS, Minute to gigantic mites. The following characters

usually occur on adults of this tribe (exceptions are given in parenthesis).

128 REC, 8S. AUST. MUS., 16 (3): 1-219 July, 1970

Separate podonotal and opisthonotal shields (some Gamasellus discutatus-

complex species), One or more pairs of pre-endopodal shields. Form ol

dorsal setae very variable, and if only some setue are pilose on podonotum

then usually they include seta /4, 25 and r3 surrounded by simple or

lanceolate setac, Spermathecal ringed tube, if recognized, opens of

idiosoma (Laelogamasus species ),

MORPHOLOGY,

SCLEROVIZATION, Usually podonotal and opisthonotal are separate but

they are fused together on most Gamasellus discutatus-complex adults.

Often there is more than one pair of pre-endopodals as in A//ogamasellus,

Gamasellus, Laelogamasus, Hiniphis, Periseius and Rhodacaroides species.

CHAETOTAXY, Setation of the legs is as that of Gamasellus except for

some Cyrtolaelaps species which have an extra postero-lateral seta on

tibia LI, while other Cyrtolaelaps species and Heterogamasus and Notogama-

sellus (Nofogamaselluy) species have an extra postero-lateral seta on genu HI

(2, 4/2, 2) and tibia ILL (2, 3/2, 2).

OTHER CHARACTERS, Often some dorsal setae are conspicuously pilose,

lanceolate or spatulate. Species which have only setose dorsal setae

(that may sometimes be slightly pilose) occur in the following groups:

Allogamasellus, Euryparasitus, Hiniphis, Litogamasus, Periseius (Psammon-

sella) and Rhodacaroides, When only some podonotal setae are pilose and,

or spatulate, they are often setae jl, /4, 25, and 73, as in Cyrfoluelaps and

Gamasellus species. Some females of the Gamasellus discutatus-complex

have the sternal setae positioned as in the Gamasiphinue, but usually they

are in a pair of nearly straight rows, If spermathecal ringed tubes ure

recognized they usually open on the idiosoma, but in Laelogamasus species

they open on coxa TIL.

DISTRIBUTION, Species of Gamasellini and the Rhodacarinae make

ip most of the rhodacarid fauna of the Northern Hemisphere, but about

two-thirds of those belonging to this tribe were collected in the Southern

Heniisphere,

REMARKS. The Gamasellini is intended mainly for the many species

with adults that have two subequal dorsal shields and often some complex

dorsal setae. There is a precedent for this concept, although it probably

gives little indication of the phylogenetic relationships of the species.

| recognize the following 14 genera within this tribe: Gamrasellus,

Acugamasus, Allogamasellus, Cyrtolaelaps, Euryparasitus, Evanssellus

Heterogamasus, Hiniphis, Laelogamasus, Litogamasus, Notogamasellus,

Periseius, Pilellus and Rhodacaroides. Members of these genera can be

distinguished by using the above key (p. 78).

LEE—RHODACARIDAE 129

Genus GAMASELLUS Berlese

Gamasellus Berlese, 1892d, p. 60. Type-species: Gamasus falciger G. & R,

Canestrini, 1881, designated by Berlese, 1906, p. 101.

DIAGNOSIS. Small to large mites, with a considerable range in the

extent and selerotization of shields. Usually separate podonotal and opistho-

notal shields, but most females atid some males of the discutatus-complex

species have a holonotal shield. Ventro-anal shield on the female rarely

(us in the anal region of pyiformis-complex species) fused to notal shield,

while on male it may be fused to the notal shield. Two or three pairs of

pre-endopodal shields, Some dorsal setae always complex. Spermathecal

ringed tube opens near posterior paraxial edge of acetabulum IV. Pretarsus

1 either pedunculate and conspicuously smaller than other pretarsi, or not

pedunculate and claws subequal in size to claws of other pretarsi,

REMARKS. There is a distinctive group of nominal species, simular

to the type-species of Gamasellus to which I refer to as the falciger-complex.

A very similar pyriformis-complex is recognized from East Africa and a

discutatus-complex is recognized from Australia, Most members of the latter

complex are conspicuously different from the falciger-complex because of

their idiosomal sclerotization while one species is quite similar to the

falciger-complex,

The 3 species-complexes that [ recognize within this genus can be

distinguished by the following key. The morphology and distribution of

members of this genus are given under the species-complex headings,

KEY TO SPECIES-COMPLEXES OF GAMASELLUS

1, Less than 23 pairs of opisthonotal setae, and

fewer than 6 pairs of setae on striated cuticle

between ventro-anal and opisthonotal shield

other than the setae in row R. Male ventro-

anal shield rarely fused to opisthonotal shield,

or, if it is, also fused to peritrematal shield - falciger-complex

With either 25 or more pairs of opisthonotal

setae, or, if 23 or 24 pairs, there are more

than 6 pairs of setae on striated cuticle

between ventro-anal and opisthonotal shield

while setae in row RK are on opisthonotal

shield. Male ventro-anal shield usually fused

to opisthonotal shield but not peritrematal

shield

130 REC. 8. AUST. MUS., 16 (3): 1-219 July, 1970

Seta a/l, on palp genu, is pectinate with at least

5 lateral prongs. Pretarsus I not pedunculate,

sheath being only long enough to contain

retracted claw-complex. Female ventro-anal

shield not fused to exopodal IV shield, but

fused in its anal region to opisthonotal shield — pyriformis-complex

Seta all, on palp genu, is spatulate with a single

lateral prong. Pretarsus I is pedunculate.

Female ventro-anal shield is fused to exopodal

IV shield, but not to opisthonotal shield . .. — discutatus-complex

FALCIGER-complex

DIAGNOSIS. Small to average sized species. Separate podonotal

and opisthonotal shield. Ventro-anal shield never fused to notal shield,

exopodal IV shield or peritrematal shield of female. and rarely fused to notal

shield and peritrematal shield on male, but quite often fused to male exopodal

IV shield. If male ventro-anal shield fused to notal shield it is also fused to

peritrematal shield. Less than 23 pairs of opisthonotal setae. Spermadacty!

subequal in length to movable cheliceral digit of male, or slightly longer.

On palp genu, seta a/l spine-like with 2 to 6 lateral prongs. Pretarsus I

pedunculate.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Ventro-anal usually discrete, but may bear aciculae posterior to anus. May

be small discrete shield only bearing aciculae. No discrete metapodal.

Peritrematal fused to exopodal TV. Split in exopodals I], IT and IV.

Sterno-metasternal fused to endopodal II and part of Il, Three pairs of

pre-endopodals.

Male: Ventro-anal either discrete but extending closer to exopodal 1V

than on female, or more rarely it is fused to exopodal TV, and on G. falciger

and G. nepotulus it is fused to sternito-genital, exopodal IV. peritrematal

and notal.

CHAETOTAXY. Idiosoma: 6j, 6z, 5s, Sr: SJ, 5Z, 58, 5R, O-2UR:

Sst: 34, 3Zy, O-2Sy.

Legs: As Gamasellus discutatus (see Pig. 3).

OTHER CHARACTERS. Female: Deutosternal denticles in 7 or 8

horizontal rows with | or 2 horizontal deutosternal ridges anterior to

denticles. Tectum anterior margin is basically trispinate but usually with a

number of spinules around the two smaller lateral spines, which may be large

LEE—RHODACARIDAE 131

enough to make the margin quinquispinate. Movable cheliceral digit with

3 teeth. On palp genu, seta all spine-like with 2 to 6 lateral prongs, and

seta al2 slightly spatulate, Some dorsal setae on nominal species are always

pilose and often spatulate as well, and usually many are lanceolate, only a

few being setose. Shields usually dull being covered in an adhesive exudate.

Spermathecal ringed tube opens near posterior paraxial edge of acetabulum

IV. Pretarsus [ pedunculate, and smaller than other pretarsi. Lateral lobes

of pulvilli II-[V much shorter than central, circular lobes. Amongst dorsal

setae on tarsus TV, seta pd3 is the longest and is spatulate.

Male: [f there is a ventral protruberance on palp, there is also one on

female palp. Movable cheliceral digit with one tooth, and fused at base to

similarly shaped spermadactyl, that is subequal in length or slightly longer.

On leg Il, femur seta av, and sometimes seta pvl. genu seta ay. and some-

limes tibia seta av and tarsus seta av2 enlarged into spurs or spines.

Sometimes non-setous spurs on leg Il (see Fig. 318),

DISTRIBUTION. Nn, Ne, Nr, Na: Es: Pe, Pm, Ps, Pe: Os: Aa, An.

The locality records with the original descriptions of nominal species are

front the following major regions.

Neurctic region: G. leggetti Ryke, 1962c; G. vihrissatus Emberson,

1967; G. hellavistae Emberson, 1967.

Ethiopian region: G. deepdalensis Ryke, 1962c; G, shongwiensix Ryke,

1962¢,

Palaearctic region: Gamasellus borealis (Koch, 1879) comb.n.;

G. falciver (G. & R. Canestrini, 1881); G. humosus Ishikawa, 1969; CG.

nepotuluy Berlese, 1908; G. montanus (Willmann. 1936); Gy serfatis

(Willmann, 1941) (G. faleiger, syn. Ryke, 1962c): G. alpinus Schweizer,

1949; G. nivalis Schweizer, 1949; G, silvesiriy Halaskova, 1958 (G.

montanus, syn. Hirschmann, 1962); G. spiricornis (G. & R. Canestrini,

1882),

Australian region: G, concinnus (Womersley, 1942); G. tragardhi

(Womersley, 1942): G. coopert (Womersley, 1961).

My new combination for Gamasus borealis from the siberian Palaearctic

region is only based on the original, brief description. ‘en unnamed species

from all four zoogeographical regions of North America are described by

Eimberson (thesis, 1968). The specimens from the Oriental region were

from Nepal and are to be dep. BM(NH). Specimens trom New Zealand

and New Caledonia are dep. SAM,

Found amongst moss, plant litter and upper soil layers. A single female

of G. spiricornis was found on a carabid beetle, Belgium (Cooreman, 1943),

132 REC, 8S. AUST. MUS., 16 (3): 1-219 July, 1970

261, 282

100 283,288

Figs. 281-288. Gamasellus tragardhi (Womersley ).

281-285 and 287, female: 281, soma, dorsum: 282, idiosoma, venter; 283, leg IV

(part), dorsum; 284, pretarsus 1 and 1V; 285, coxae and trochanter IIT and IV;

287, gnathosoma, venter. 286 and 288, male: 286. chelicera: 288. idiosoma, venler.

LEE—RHODACARIDAE 133

Figs. 289-294. Gamasellus tragardhi (Womersley ).

289 and 290, male: 289, idiosoma, latus; 290, leg I] (part), 291-293, deutonymph:

291, idiosoma, dorsum; 292, idiosoma, venter; 293, pretarsus I and IV, 294,

protonymph, soma, dorsum,

July, 1970

296

1004 300

303

297

100yn 30!

[004 304

Figs. 295-304. Gamasellus tragardhi (Womersley ).

295-297, protonymph: 295, idiosoma, venter; 296, gnathosoma, venter: 297, leg IV

(part), dorsum. 298-302, larya: 298. pretarsus I; 299, soma, dorsum; 300,

gnathosoma, venter; 301, leg IIT (part), dorsum; 302, idiosoma, venter, 303, female,

tarsus I, dorsum, 304, egg.

LEE—RHODACARIDAE 135

REMARKS. Of the Cyrtolaelaps (Gamasellus) species listed by Ryke

(1962) but not listed here in this genus, or in Acugamasus, Cyrtolaelaps,

Parasitiphis or Rykellus, 1 have seen those described by Berlese and they

are not rhodacarids. The same is probably true for the Gamasellus species

described by Cooreman and Vitzthum, although I have not seen them, and

for G. grabouwensis Ryke, 1964. The 18 nominal species included in this

species-complex are listed in the ‘Distribution’ section.

Gamasellus tragardhi (Womersley )

Digamasellus tragardhi Womersley, 1942, p. 161.

FEMALE. Fig. 281-285, 287 and 303. Idiosomal length, 610».

MALE, Fig. 286 and 288-290. Idiosomal length, 6004. Note

process on genu II is seta av, while on G. falciger (Fig. 318) and G.

nepotulus in the Berlese Collection, it is non-setous,

DEUTONYMPH. Fig. 291-293. Idiosomal length, 540y.

PROTONYMPH. Fig. 294-297. Idiosomal length, 360».

LARVA. Fig. 298-302. Idiosomal length, 260».

EGG. Fig. 304. Longest axis length, 230p.

LOCALITY. Female (N196843), male (N196844), deutonymph

(N196845), protonymph (N196846), larva (N196847) and egg (N196848 )

drawn: Australia; LF133, moss off rocks, Stony Creek, nr. Wilmington,

South Australia, 25.5.1966, col. H. M. Cooper, dep. SAM (immature stages

not collected direct, but bred from females in this sample except the egg

which was dissected from a female).

Holotype female (N196849) examined: Australia; moss, Adelaide,

South Australia, 6.1935, col. H. Womersley, dep. SAM.

Allotype male (N196850) examined: Australia; moss, Bridgewater,

South Australia, 8.1942, col. J. S. Womersley, dep. SAM.

REMARKS. In figured chaetotactic patterns, filled in circles represent

setae considered to have been added at the previous moult. Characters of

the immature stages have not been used in the classification given here. The

extensive drawings of these stages are given only to increase this type of

data which is too limited from this family for satisfactory comparative studies.

Similar drawings are made in this paper of the immature stages of

Hydrogamasus littoralis and Heydeniella dentata.

136 REC. 8. AUST. MUS, 16 (4): 1-219 July, 1970

DISCUTATUS-complex

DIAGNOSIS. Small to large mites. Both sexes may have either a

holonotal shield or separate podonotal and opisthonotal shields, or female

may have a holonotal shield and male of the same species have separate

podonotal and opisthonotal shields. Ventro-anal of the female fused only

to exopodal TV shield and rarely to peritrematal shield as well, while on

male it is fused to exopodal IV shield and either to part of the peritremutal

shield, or to notal shield. More than 22 pairs of opisthonotal setae. Some

females differ from all other Ologamasinae species in that a line joining

sternal setae sf2, sf3 and s/4 would enclose an angle of just less than 95°.

Spermadacty! sinuous and conspicuously longer than movable cheliceral digit

of male. On palp genu, seta a/l spatulate with a single lateral prong.

Pretarsus | pedunculate.

MORPHOLOGY.

SCLEROTIZATION, Female; Usually holonotal but has separate

podonotal and opisthonotal in one nominal species (G. fasmanicus), Ventro-

anal is fused to the exopodal IV along the posterior rim of acetabulum IV

and may also be fused to peritrematal. Posterior to ventro-anal there 1s

a small, discrete shield bearing aciculae. No discrete metapodal. Peri-

trematal fused to exopodal [V. Split in exopodals IT, If and IV. Sterno-

metasternal fused to endopodal II and Ill. Two or three pairs of

pre-endopodals.

Male: Usually notal is as in the female but in one nominal species

(G. discutatus) the male has separate podonotal and opisthonotal while the

female has a holonotal. Ventro-anal may be fused to notal, in a species

(G. southcotti) where tt is not, its fusion to the podal along the posterior

rim of acetabulum TV may extend to part of the peritrematal. Peritrematal

usually broader and as a result may carry seta r4, which is on striated cuticle

in the female.

CHAETOTAXY. Idiosoma: 6/, 62, 59, 5¢: 6/, 4 or 67, 4 or 6S, 5 or

6R, 4, 6 or OUR: Sst: 3Jv, 3Zv, 38v, 4 or SRv. In some species setae

belonging to row Rv are on the notal shield. The ‘post-lateral’ setae referred

to as present on these species by myself (Lee, 1966), are here regarded as

belonging to row Sv or Rv.

Legs: As Gamasellus discutatus (see Fig. 3).

OTHER CHARACTERS. Female; Deutosternal denticles in 8 hori-

zontal rows with one horizontal deutosternal ridge anterior to denticles,

Tectum anterior margin basically unispinate with lateral spinules, which may

be large enough to make the margin quinquispinate. Movable cheliceral

LEF—RHOBDACARIDAE 137

digit with 4 teeth. On palp genu, seta a/l spatulate with single lateral prong

and seta @/2 slightly spatulate. Most dorsal setae are lanceolate, but always

some setae are pilose or pilose and spatulate. Shields dull being covered

by an adhesive exudate. Spermathecal ringed tube probably opens near

posterior edge of acetabulum IV, Pretarsus | pedunculate and smaller than

other pretarsi, Lateral lobes of pulvilli LI-IV attenuated and conspicuously

longer than central, circular lobes. Amongst dorsal setae on tarsus TV, seta

ad2 is the longest and is setose.

Male: Movable cheliceral digit with one tooth and fused at buse to

slimmer and longer spermadactyl which is sinuous and spatulate at up. On

leg Hl, femur setae av and pvl, genu setae av and py, tibia seta av and tarsus

sela ay2 enlarged to spurs or spines. I (Lee, 1966) incorrectly drew seta

av2 on tarsi IL of males as being setose:

DISTRIBUTION, Aa. All records are from the south eastern states

of Australia including Tasmania and are published with the original deserip-

tions of the nominal species.

Found in moss and plant litter,

REMARKS, Five species trom the discufatus-complex were originally

described in the ‘diseutarus-group’ of Ologamasus (Lee, 1966), Characters

such us the 2 or 3 pairs of pre-endopodal shields, the pilose setae amongst

the simple dorsal setae and the spermathecal ringed tube aperture on the

idiosoma are now regarded as more important than the female holonotal

shield, and, therefore, this species-complex has been transferred to Gamasellus

and G. tasmanicus, which has a separate podonotal and opisthonotal shield

in both adults and was transferred to Gamasellus by Ryke (1962c). has also

been included. The following 6 nominal species are, therefore, included in

this species-complex: G. discutatusy (Lee, 1966) comb.n.; G. litoprethrix

(Lee, 1966) comb.n., G. southcorti (Lee, 1966) comb.n.; G. fasmanicus

(Womersley, 1956a); G. tindalei (Lee, 1966) comb.n.: G_ virgosus (Lee.

1966) comb.n.

PYRIFORMIS-complex

DIAGNOSIS, Small to average sized mites, Separate podonotal and

opisthonotal shields. Ventro-anal fused to notal shield only in anal region

on female but extensively on male. Male ventro-anal shield also fused to

exopodal IV shield, but there is a conspicuous fissure between it and the

peritrematal shield, More than 22 pairs of opisthonotal setae. Sperma-

dactyl straight and subequal in length to movable cheliceral digit of male.

On palp genu. seta a/l spine-like with 5 lateral prongs. Pretarsus [ not

pedunculate, sheath being broadly fused to tarsus and only long enough to

contain retracted claw-complex.

138 REC. S. AUST. MUS., 16 (3): 1-219 July, (970

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal und opisthonotal,

Ventro-anal bearing aciculae posterior to anus and in this limited region

fused to opisthonotal, but not fused to any other shields. No discrete

metapodal. Peritrematal fused to exopodal 1V. Split in exopodals II, U1

and IV, Sterno-metasternal fused to endopodal If and part of HI, Three

pairs of pre-endopodals,

Male: Ventro-anal is fused to opisthonotal and to exopodal IV around

posterior edge of acetabulum IV, but there is a fissure between it and

peritrematial.

CHAETOTAXY, Idiosoma: 67, 62, 58, Sr: 6/, 5.27, 58, 5R, TUR: Sst:

3), 3Zy, LSv.

Legs; As Gamuasellus discutatus (see Fig. 3).

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows, Tectum anterior margin is basically unispinate, but multispinulate

laterally. Movable cheliceral digit with 3 teeth. On palp genu, seta

a/] with 5 lateral prongs and seta a/2 slightly spatulate. Dorsal setae

vary in form as for falciger-complex. Spermathecal ringed tube opening

hot known. Pretarsus I not pedunculate, with sheath that is only long enough

to contain retracted claw-complex.

Male: Movable cheliceral digit has two small denticles (these may

represent the edges of base of broken off single tooth) and is fused at base

to similarly shaped spermadactyl which is slimmer but subequal in length.

On leg I, femur seta av, genu seta av, tibia seta av and tarsus seta av2 are

enlarged into spurs or spines.

DISTRIBUTION. Ee. The only published record is from East Africa

with the description of the nominate species, while an unnamed species from

this region is described by Loots (thesis, 1967).

Found in plant litter and soil.

REMARKS, The members of the pyriformis-complex are very similar

to the falciger-complex, but differ in having a large number of opisthonotal

setae, a short, large clawed pretarsus | and a slightly unusual ventral

sclerotization on both sexes. There is one nominal species (only the female

is known) included in this species-complex, G. pyriformis Berlese, 191 6a,

and Loots (thesis, 1967) describes the female and male of an unnamed

species.

Gamasellus pyrifermis Berlese

Gamasellus pyriformis Berlese, 1916a, p. 161.

FEMALE, Fig. 312-315. Idiosomal length, 590. Pretarsus I

broudly fused with tarsus, so that claws appear retractable into tarsus,

LEE—RHODACARIDAE 139

pulvillus may be present, although not drawn. This specimen contains an

egg, with larva (legs and setae visible) inside.

MALE. Not known.

LOCALITY. Single specimen in SEAF collection. Female (175/37)

drawn: East Africa; soil and litter, col. Alluaud and Jeannell, dep. SEAP.

Genus ACUGAMASUS gen.n.

Type-species: Digamasellus punctatus Womersley, 1942.

DIAGNOSIS. Small to average sized mites. Separate podonotal and

opisthonotal shields. Ventro-anal shield on female never fused to notal

shield, peritrematal shield or exopodal IV shield, but on male may or may

not be fused to these shields. Single pair of pre-endopodal shields. Some

or most dorsal setae pilose and/or spatulate. Spermathecal ringed tube

opening not known. Spermadactyl shorter than movable cheliceral digit of

male. Pretarsus [ pedunculate.

REMARKS. When Ryke (1962c) redefined Gamasellus, as a sub-

genus of Cyrtolaelaps, he included in it a wide variety of species. While

some of these species are conspicuously different from Gamasellus falciger

(e.g., those now referred to Afrogamasellus (Rhodacarinae) and Rykellus

(Ologamasini)), the majority of species not congeneric with the type of

Gamasellus belong to the same tribe (Gamasellini) and have only a single

pre-endopodal shield. [I have established this group of species as a new

genus; Acugamasus.

I recognize two species-complexes within this genus, punctatus-complex

and natalensis-complex, which can be distinguished by the following key.

The morphology and distribution of members of this genus are given under

the species-complex headings.

KEY TO SPECIES-COMPLEX OF ACUGAMASUS

1. Seta of row R off opisthonotal shield. Excluding

anal setae, at least 6 pairs of setae on ventro-

anal shield. No ventral spurs on male palp.

Male ventro-anal shield never fused to opistho-

notal shield . + 22... 12 4 is by ee ee

Setae of row R on opisthonotal shield. Excluding

anal setae, sometimes 6 pairs but often 5 pairs

of setae on ventro-anal shield. Ventral spur

on proximal palp segments of male or at least

1 ventral setae of palp femur enlarged into

spine. Male ventro-anal shield usually fused

to opisthonotal shield .. 2. 2.0.2. 0..0.. .. natalensis-complex

punctatius-complex

140 REC. 8. AUST. MUS., 16 (3): 1-219 July, 1970

PUNCTATUS-complex

DIAGNOSIS. Ventro-anal shield not fused to notal shield and widely

separated from exopodal IV shield on both sexes. Usually most dorsal setae

are pilose and sometimes also spatulate. Between 19 and 24 pairs of

opisthonotal setae, with at least anterior 2 setae of row R on striated cutucle,

and 6 pairs of setae on Ventro-anal shield (excluding anal setae), Male

chelicera and spermadactyl is smooth, Male palp similar to female, being

without ventral spurs or spines.

MORPHOLOGY.

SCLEROTIZATION. Female; Separate podonotal and opisthonotal.

Ventro-anal discrete and bearing aciculae posterior to anus. Discrete

metapodal. Peritrematal fused to exopodal [V. Exopodals not completely

encasing peraxial edges of acetabula. Sterno-metasternal fused to endopodal

Il and a small part of II. Single pair of pre-endopodals.

Male: Except on A, watsoni, a section of the podonotal carrying at

least seta r2 and r4, is split away posteriorly from the rest of shield and

fused to peritrematal. Ventro-anal separate from opisthonotal, but broader

than in female (setae Zvl and Zv2 being well in from edge) probably

including metapodal, Sternito-genital separate from ventro-anal and may

be separate from endopodal TV Jeaving st5 either on striated cuticle, or on

small discrete shield or on shield fused to endopodal IV,

CHAETOTAXY, Idiosoma: 6/, 6z, 5s, Sr: 5/, 5Z, 58, SR, O-3UR: Sst:

3/v, 3Zv, 2Sv.

Legs: As Gamasellus.

OTHER CHARACTERS, Female; Deutosternal denticles in 8 hori-

zontal rows with one deutosternal ridge anterior to denticles. Tectum

anterior margin having a central spine with a multispinulate lateral margin.

Movable cheliceral digit with 3 teeth. On palp genu, seta a/l spatulate, seta

ul2 slightly lanceolate. Three forms of dorsal setae always present, simple,

pilose, and pilose and spatulate (size increases through this series), but at

the most there are only four pairs of simple setae. Spermathecal ringed tube

opening not known. Pretarsus I similar to that of Gamasellus tragardhi.

Pulvilli [-TV may be as A. cursor or with lateral lobes longer than central

circular lobes. Amongst dorsal setae on tarsus IV, seta ad2 is just the longest

and is sctose,

Male; Movable cheliceral digit with one tooth, and fused at base to

shorter (may be only half length) spermadactyl. On leg UH, femur setae av

and pvl, sometimes genu sctae av and pv, and sometimes tibia seta av,

enlarged into spur or spine. May be non-setous process on genu II,

LEE—RHODACARIDAE 141

——o

° ~o

Oo—o——

ce rf

Figs. 305-311. Acugamasus cursor sp.n..

305-307 and 311, female: 305, soma, dorsum; 306, leg TV (part), dorsum; 307.

idiosoma, yventer; 311, gnathosoma, venter. 308-310, male: 308. idiosoma, venter:

309, leg Il, antero-latus; 310, chelicera.

142 REC. 8S. AUST. MUS., 16 (3): 1-219 July, 1970

DISTRIBUTION. Aa, Am, An: Sa. The distribution in the australian

Australian region is wider than the records published with original descrip-

tions of nominal species would indicate. I have seen specimens from the

Australian Capital Territory, Lord Howe Island and New Caledonia, dep.

SAM, and from the Solomon Islands, dep. BM(NH). There is one male,

closely allied to A. semipunctatus, from Wellington, New Zealand, dep.

SAM. A. watsoni has been collected from Auckland and Campbell Islands,

dep. BBM.

Found amongst moss, plant litter and upper soil layers.

REMARKS. The punctatus-complex contains the type of Acugamasus

which is superficially similar to Gamasellus because of its two dorsal shields

and pilose dorsal setae. Members of the punctatus-complex can be

distinguished from Gamasellus by having only one pair of pre-endopodal

shields, a short spermodactyl and a setose seta av2 on the male tarsus II.

The following 4 nominal species are included in this species-complex:

A. cursor sp.n.; A. punctatus (Womersley, 1942) comb.n.; A. semipunctatus

(Womersley, 1942) comb.n.; A. watsoni (Hirschmann, 1966) comb.n..

A cugamasus Cursor Sp.N.

FEMALE. Fig. 305-307 and 311. Idiosomal length, 5104. Opistho-

notal shield has reticulations but neither notal shield has raised punctations.

The cells drawn in Fig. 305, near seta /2, belong to a piece of fungal hypha

in the exudation layer. On one paratype female (N196855), seta st4 is on

striated cuticle.

MALE. Fig. 308-310. Idiosomal length, 460». The lateral edge of

the podonotal shield extends as far forward as seta 53, as a fissure separating

off part of the podonotal shield bearing setae r2, r3 and r4, and this is fused

to peritrematal shield.

LOCALITY. Holotype female (N196851) and allotype male

(N196852) drawn, and 3 paratype females (N196853-N196855) and 3

paratype males (N196856-N196858) examined: Australia; LF99, moss on

rocks and rotting tree stumps, by Beauchamp Falls, near Beech Forest,

Otway Ranges, Victoria, 9.12.1965, col. D. C. Lee, dep. SAM.

REMARKS. Although the number of setae on the opisthonotal shield

appears to vary between members of the same nominal species in this species-

complex, this species is unusual in having at least 18 pairs of setae, including

seta $1, on this shield. This species can also be distinguished from the other

two nominal species from Australia in not having raised punctations covering

most of the podonotal shield.

LEE—RHODACARIDAE 143

NATALENSIS-complex

DIAGNOSIS, Nentro-anal shield discrete on female, but rarely so on

male where it is usually extensively fused to all the surrounding shields

except the sternito-genital. Dorsal setae may be mostly pilose and/or

spatulate as on punctatus-complex species or only a few dorsal setae may be

pilose and/or spatulate as on Gamasellus falciger-complex species. Between

21 and 25 pairs of opisthonotal setae, with all setae of row R on opisthonotal

shield, and usually only 5 pairs of setae on ventro-anal shield (excluding

anal setae). Male chelicera and spermadactyl often gnarled. Male palp

always has a ventral spur or spine on femur or trochanter that are not

present on female.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Discrete ventro-anal. May be small discrete shield only bearing aciculae.

May be discrete metapodal. Peritrematal fused to endopodal IV. Sterno-

inetasternal fused to endopodal HI and part of HI. Single pair of pre-

endopodals,

Male: Ventro-anal may be discrete, but usually fused to all surrounding

shields except the sternito-genital. In the latter case the peritrematal is

brouder und extensively fused to podonotal.

CHARTOTAXY. Idiosoma: Podosomua has 6/, 62, Ss, 5p: Swf. The

setation of the opisthosoma is uncertain, but in most species it probubly

conforms to 5/J, 52, 58, 5R, 2-4UR: 3/v, 3Zv, ISv.

Legs: Unknown.

OTHER CHARACTERS. Female: Teetum anterior margin is trispinate

or quinquispinate, or basically trispinate with spinules on and around

the spines, Movable cheliceral digit with 3 teeth. The dorsal setae

may be either nearly all pilose or all spatulate, or pilose and spatulate, or

there may be pilose setae amongst simple setae. In species with a number

of row UR setae, there ate only 5 pairs of setae on the ventro-anal shield,

excluding unal setae, seta Zy3 being on the striated cuticle.

Mule; Tectum anterior margin is usually more elaborate than in female,

Ventral spur or spine on proximal segments of palp that is not present in the

female. Movable cheliceral digit with | tooth and fused at base to similarly

shaped, but slightly shorter spermadactyl, Both chelicera and spermu-

dactyl may be gnarled. On femur II, seta av enlarged into a spur, usually

other spurs also present on leg II.

DISTRIBUTION, Es. All records are from Cape Province and Natal,

South Africa and are published with the original descriptions of the nominal

species,

144 REC, S, AUST. MUS. 16 (3); 1-219 July, W70

Found in plant litter and soil.

REMARKS, 1 have not seen any members of the natalensis-complex

so that fewer characters are mentioned, Because its members have only a

single pair of pre-endopodal shields, I have transferred this species-complex

from Gamasellus to Acugamasus, but Lam uncertain of its relationship to the

Punctatus-complex in latter genus. The following 8 nominal species are

included in this species-complex; 4. drakensbergensis (Ryke, 1962c)

comb.n.; A, grahami (Ryke, 1962c) comb: A, fluhluwensis (Ryke,

1962c) comb.n.; 4. knysnaensiy (Ryke, 1962c) comb.n.; A. macrosetosus

(Ryke, 1962c) comb.n.: A. natalensis (Ryke, 1962c) comb.n.; A, neotas-

manicus (Ryke, 1962c) comb.n.; 4A. paranatalensis (Ryke, 1962c) comb.m.

An unnamed species was described by Loots (thesis, 1967).

Genus ALLOGAMASELLUS Athias-Henriot

Allogamasellus Athias-Henriot, 1961b, p. 473. Type-species: Allogama-

Sellus aquafortensis Athias-Henriot, 1961b, by original designation.

DIAGNOSIS, Minute mites only known from females. Separate

podonotal and opisthonotal shields. Discrete ventro-anal and metapodal

shields. Three pairs of pre-endopodal shields. Dorsal setae short and spine-

like, except setae jl and zl! which are longer and slightly pilose. Sperma-

thecal ringed tube opening not known. Pretarsus I pedunculate.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal

Aciculae not observed, Discrete ventro-anal and metapodal. Peritrematal

fused to exopodal TV. Exopodals not completely encasing peraxial edges

of acetabula. Metasternal fused, ulthough it may be narrowly, to sternal,

Sternal fused to endopodal IT and Ill. Three pairs of pre-endopodals.

CHAETOTAXY. Idiosoma: 6/, 62, 5s, Sr: SJ, 5Z, 58, SR, 2 or 4UR:

Sst: 3/v, 3Zy, 1Sv.

Legs: As Gamasellus,

OTHER CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows plus a single, anterior, horizontal deutosternal ridge. Movable cheliceral

digit with 3 or 4 teeth. On palp genu, seta a/l and a/2 slightly spatulate.

Small pore-like structure posterior to capitular seta hyp4 on hypostome. Most

dorsal setae, short, spine-like, but setae j] and zl, longer and slightly pilose.

Lurge pore between setae s5 and r5 on podonotal shield. Peritreme short,

less than twice the diameter of the spiracle. Spermathecal ringed tube open-

ing nol known. Pretursus | pedunculate and smaller than other pretarsi,

LEE—RHODACARIDAE 145

No Scale

316

317

318

Figs. 312-322. Gamasellus Berlese, Laelogamasits Berlese and parasitid

(ex-Hydrogamasus) species.

312-315, G. pyriformis Berlese, female: 312, soma, dorsum; 313, idiosoma, venter:

314, palp genu; 315, pretarsus | and tarsus I (part). 316 and 317, ex Hydrogumasus,

al setae on palp femur and genu: 316, H. silvestri Berlese; 317, Hl. salinus

(Laboulbéne). 318, G. faleiger (G. & R. Canestrini), male, genu Hf, 319-322,

L. simplex (Berlese), male: 319, mid-tectum; 320, chelicera; 321, palp trochanter.

femur and genu, antero-latus; 322, leg IL (part).

146 REC. $. AUST. MUS.,. 16 (3): 1-219 July, 1970

DISTRIBUTION. Pm. The only records are with the original

descriptions of the nominal species from Algeria.

Found amongst plant roots and Jitter.

REMARKS. Allogamasellusy males are unknown. The females are

similar to those of the Gamasellus falciger-complex but there would have to

be new intermediate species for it to be preferable to regard this genus as

congeneric with Gamasellus. The following 2 nominal species are included

in this genus: A. aquafortensis Athias-Henriot, 1961b; 4. squalidus Athias-

Henriot 1961b. Two syntype females (C988 and C991) of A. aquafortensis

and the holotype female (C464) of A. squalidus, dep. MNHN, have been

examined.

Genus CYRTOLAELAPS Berlese

Cyrtolaelaps Berlese, 1887b, XLIV, 5 (not Berlese, 1892a, LXIV, 3).

Type-species: Gamasus mucronatus G. & R. Canestrini, L881, by

monotypy.

Protolaelaps Tragardh, 1912, p. 563. Type-species: Gamasellus |) brevi-

spinesus Trigardh, 1910 (syn, Gamasus mucronatus G. & R. Canestrini,

1881), by original designation.

DIAGNOSIS. Average sized to large mites. Separate podonotal and

opisthonotal shields. Ventro-anal shield discrete on female, while on the

male it may or may not be fused to notal, but it is always fused to peritrematal

and exopodal IV shields. Clearly defined pre-endopodal shields absent.

Usually, at least one pair of conspicuously pilose setae on podonotal shield

and on opisthonotal shield. Spermathecal ringed tube may open on sterno-

metasternal shield. Spermadactyl stout and fused to shorter movable

cheliceral digit along its entire length. Pretarsus I not pedunculate, sheath

being broadly fused to tarsus and only long enough to contain retracted

claw complex.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Aciculae may be on separate shield or on ventro-anal. Discrete ventro-anal

and metapodal. Peritrematal fused to exopodal IV. Exopodals not com-

pletely encasing peraxial edges of acetabula. Metasternal discrete or fused

to sternal. Sternal fused to endopodal I and sometimes to part of endopodal

III. Pre-endopodals absent, or are small and ill defined as described for

C. paraster by Costa (1961).

LEF—RHODAC ARIDAER 147

Male: Ventro-anal cither entirely fused to opisthonotal, peritrematal

and exopodal [V, but separate from sternito-genital, or fused only to

peritrematal and exopodal TV as described on C. aster by Schweizer (1961),

CHAETOTAXY, Idiosoma: 7j, 62, 5-65, Sr: 5/, 5Z, SS, 5R, 2UR or

hypertrichous with up to 100 or more setae; Sst; 3/v, 4Zyv, 2-4Sv.

Legs: As Gamasellus with | postero-lateral on tibia HT and genu II,

or 2 postero-laterals on tibia MT (2, 3/2, 2) genu HM (2, 4/2, 2), or an

extra postero-lateral on tibia TIT only,

OrucR CHARACTERS, Female: Deutosternal denticles in 10 or

11 horizontal rows. 'Tectum anterior margin has large central spine with

multispinulate sides. Movable cheliceral digit with 3 or 4 teeth, On palp

venu, seta all and al2 simple or very slightly spatulate. Vertical seta, /1.

usually stout spine. Always, some dorsal setae ure conspicuously pilose on

nominal species, but Emberson (thesis, 1968) records an unnamed species

from the californian Nearctic region, which is probably referable to this

venus, with simple dorsal seta. Shields are dull, Spermathecal ringed tube

may open between sternal seta sf3 and acetabulum HT. But. considering the

similarity of the spermadactyl on males of this genus to the unusually

powerful looking spermadactyl of Euryparasitus emarginatus, which Michael

(1892) claims is used to lift female genital plate so thal the spermatophore

ean be applicd directly to the vagina, it may be that vaginal fertilization

occurs in this genus also, instead of fertilization through spermathecal ringed

tubes. Pretarsus 1 not pedunculate, only being Jong enough to contain

retracted claw-complex, Pulvilli TI-PV have lateral lobes which are smaller

but similar in shape to central lobes, and a pair of streamer-like lobes ventral

to claws, Amongst dorsal setae on tarsus TV. seta ad2 is the longest and

is Setose,

Male; Cormculi proportionately longer than in female, Movable

cheliceral digit with one tooth and fused to longer, stout spermadacty! ulong

ils entire length. On leg Il, at least femur setae av and prl, genu seta ay,

tibia seta av and tarsus seta av2 enlarged into spurs or spines.

DISTRIBUTION. Nn. Ne, Nr: Pe, Pm, Ps, Pe. Besides the locality

records published with the original descriptions of nominal species there are

many published records from the european and mediterranean Palacaretic

regions, ranging from Scotland and the Russian Karelian Isthmus in the

north, to Spain, Sardinia and Israel in the south, There are also records

from the Kemerovo (Tayil’tsev and Men Yun-Tsuy, 1963) and South

Primorye regions (Aksenenko, 1963 and Khudyakov, 1963) of Russia, and

the western Canadian prairies und California (Emberson, thesis, 1968) in

North America showing that this genus has a Holarctic range,

148 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

323, 324, 327,

328

329, 332

330, 331

325, 326

Figs. 323-332. Cyrtelaelaps mnincronatus (G. & R. Canestrini).

323-327 and 332, female: 323, soma, dorsum: 324, leg IV (part), dorsum; 325,

pretarsus IV; 326. pretarsus [ plus tarsus I (part): 327, idiosoma, venter: 332,

gnathosoma, venter. 328-331, male: 328, idiosoma, venter,; 329. leg IL (part):

330, corniculus; 331, chelicera.

LEE—RHODACARIDAE 149

333, 334,337

335

100

Figs. 333-337. Cyrtolaelaps Berlese.

333-335, C. asier (Berlese), female: 333, soma, dorsum; 334, idiosoma, venier:

335, coxa IV. 336 and 337, C. rectus (Berlese). female: 336. idiosoma, dorsum:

337, idiosoma, venter.

TSO REC. S, AUST. MUS., 16 (3): L219 July, 1970

Usually found in the nests of rodents and moles, but also found on

these animals, in the nests of other Insectivora and birds, on ferrets, in caves

(sometimes near bat roosts), under stones or fallen leaves, amongst leaf

mould. grain spill. rotting potatoes and manure.

REMARKS. Cyrtolaelaps is similar to Euryparasitus, and if it is

shown that both are unusual within the Rhodacaridae in having females

that are fertilized either through the vagina or a spermathecal ringed tube

opening on the sterno-metasternal shield, it may be preferable to regard them

as synonymous. Ewryparasitus was included in Cyrtolaelaps as a subgenus

by Ryke (1962c), but he also similarly included Gamasellus, Digamasellus

and Gamaselliphis. The following 6 nominal species are included in this

genus: C. mucronatus (G. & R. Canestrini, 1881); C. aster (Berlese, 1918):

C. miner Willmann, 1952; C. paraster Costa, 1961; C. rectus (Berlese,

1920); ©. spurius (Holzmann, 1969) comb.n.. Gamasellus spalacis

Oudemans, 1912, probably belongs to this genus, but may well be

synonymous with C. mucronatus. An unnamed species from California is

partly described by Emberson (thesis, 1968).

Cyrtolaelaps mucronatus (G. & R. Canestrini).

Gamasus mucronatus G. & R. Canestrini, 1881, p. 1081.

FEMALE. Fig. 323-327 and 332. Idiosomal length, 1,000».

Chaetotaxy of genu IIL (2, 4/2, 2) and tibia I (2, 3/2, 2) differs from

Gamasellus in having two instead of one postero-lateral seta.

MALE. Fig, 328-331. Idiosomal length, 960p.

LOCALITY. Female (N196841) and male (N196842) drawn:

Scotland; mole’s nest (4), Roxburgh, East Lothian, 1964, col. R. M.

Emberson, dep. SAM. Female idiosoma (34/17, designo in A.M.Sc,it,)

examined and considered conspecific with female (N196841) drawn: Maly;

manure, Old College, Padova, dep. SEAF.

Cyrtolaelaps aster (Berlese )

Gamasellus (Protolaelaps) aster Berlese, 1918, p. 137.

FEMALE. Fig. 333-335. Idiosomal length, 700». Note following

differences to Ryke (1962c) Fig. 9 and 11; seta z5 is posterior to /6, seta

Jv3 and Sv2 are pilose, peritrematal shield is fused to exopodal ITY. The

setal characters, as in Ryke (1962c), were used in Costa (1961) to

distinguish this species from C, paraster, but, although they cannot now be

LEE—RHODACARIDAE 151

used, being inaccurate, the difference in stoutness of vertical seta (j1) holds,

and the capitular seta (Ayp4) is not pilose and a posterior accessory seta

(px) is present on the opisthonotal shield in this species. Ventral shields

are reticulated. Broken line in Fig. 334 indicates extent of opisthonotal

shield on right side. There is a small process on coxa IV. Chaetotaxy of

genu IIT (2, 4/2, 1) is as for Gamasellus while on tibia HI (2, 3/2, 2) there

is an extra postero-lateral seta.

MALE. Not figured. Idiosomal length, 680». Corniculi propor-

tionately longer than in female. Spermadactyl viewed ventrally seems similar

to that of C. mucronatus, but shorter. Sclerotization as C. mucronatus (and

not with ventro-anal shield separate from notal shield as drawn by Schweizer,

1961), but sternito-genital shield extends back closer to ventro-anal shield.

Leg I similar to C. mucronatus but on genu, seta pv is a spur half the size

of seta av. Process on coxa IV smaller than in female.

LOCALITY. Only two specimens in SEAF collection. Female

(190/37) examined: Sardinia; field-mouse’s nest, Asuni, col. Krausse,

dep. SEAF.

Cyrtolaelaps rectus (Berlese)

Gamasellus (Protolaelaps) rectus Berlese, 1920, p. 163.

FEMALE. Fig. 336 and 337. Idiosomal length, 850. Gnathosoma

similar to that of C. mucronatus but seta hyp3 proportionately shorter.

Aciculae, unlike those of other species, are on discrete shield posterior to

ventro-anal shield. Vertical seta, 71, setose, not a short spine. Chaetotaxy

of genu III (2, 4/2, 1) and tibia HI (2, 3/2, 1) is as for Gamasellus.

MALE. Not known.

LOCALITY. The single ‘tipico’ female (40/17) drawn: North

America; leaf mould, Columbia, Missouri, U.S.A., 1904-1906, col. C. R.

Crosby, dep. SEAF. (For justification of this data, see Hammen, 1959,

p. 24.)

Genus EURYPARASITUS Oudemans

Euryparasitus Oudemans, 1902a, p. 30. Type-species: Gamasus terriblis

Michael, 1886 (syn. Gamasus emarginatus C. L. Koch, 1839), by

monotypy.

Eurylaelaps Oudemans, 1902a, p. 8. Lapsus for Euryparasitus Oudemans,

1902a.

DIAGNOSIS. Gigantic mites. Separate podonotal and opisthonotal

shields. Ventro-anal shield discrete on female, but fused to peritrematal

and exopodal IV shield on male. Single pair of clearly defined pre-endopodal

152 REC, S. AUST, MUS, 16 (3): 1-219 July, 1970

shields. Dorsal setae mostly simple, tapered, although some setae near

anterior edge of podonotal shield may be slightly pilose. Spermatheca may

Open direct into vagina, Spermadactyl stout and fused to shorter movable

cheliceral digit along its entire length, Pretarsus [ not pedunculate, sheath

being broadly fused to tarsus and only long enough to contain retracted

claw complex,

MORPHOLOGY.

SCLEROTIZATION. Female; Separate podonotal and opisthonotal.

Discrete metapodal and ventro-anal (bearing aciculae posterior to anus),

Peritrematal fused to exopodal IV, Exopodals narrow, inconspicuous.

Sterno-metasternal fused to endopodal II, Single pair of pre-endopodals.

Male: Ventro-anal fused to peritremutal and exopodal TV, but separate

from opisthonotal and sternito-genital,

CHAETOTAXY. Idiosoma; 6), Gz, Sy, Sr: 5/, 57, 5S, SR, TUR: Sot:

34, 3Zv, 3S8v, 3Rv.

Legs: As Gamasellus,

OTHER CHARACTERS. Female: Deutosternal denticles in 16 horizontal

rows. Tectim anterior margin basically trispinate, with central spine

being the largest and with a number of spinules lateral to the 2 lateral

spines, Movable cheliceral digit with 3 teeth. On palp genu, seta a/l

slightly pilose and seta a/2 slightly spatulate. On palp femur, seta a/ in

proximal third, unusually long, being sub-equal in length to ventral setac on

palp trochanter. Dorsal setae simple (or slightly pilose), tapered, Shields

are dull, Peritreme fairly short, not extending anteriorly to mid-acetabulum

I, Michael (1892) claims that the spermatophore is applied directly to the

vagina, Which has a domed recess (spermatheca) in its roof that stores the

sperms, But, there is a tube opening between sternal seta st3 and acetabulum

II, apparently on the female only, as for Cyrrolaelaps, and this may be a

spermathecal ringed tube, Pretarsus | not pedunculate, only being long

enough to contain retracted claw-complex. Pulvilli II-lV with central lobes

enlarged and merged into a single pleated flap with a fimbriated distal margin,

while lateral lobes are inconspicuous. Amongst dorsal setae on tarsus IV,

seta pd3 is the longest and is setose,

Male; Corniculi with paraxial flap. Movable cheliceral digit without

tooth and tused to much longer, stout spermadactyl along its entire length,

Anterior edge of podonotal shield is more heavily sclerotized than in female

and produced forward (in E. calearater it covers the gnathosoma), On

trochanter T, seta d large and knob-like. On leg I, femur seta av. genu seta

av, tibia seta av, tarsus seta ay2 enlarged into spurs; while femur seta pv!

and genu seta py are spines, Tibia and tarsus I are fused together.

LEE—RHODACARIDAE 153

338, 339,34), 344

345, 347

342, 343, 346

ré

frets

Pines

ee)

Yer:

ae i Sa

Figs. 338-347. Euryparasitus emarginatus (Koch),

338-341 and 347, female: 338, soma, dorsum; 339, leg IY (part), dorsum; 340,

pretarsus | and tarsus I (part); 341, idiosoma, venter; 347. gnathosoma. venter.

342-346, male: 342, trochanter I (part), dorsum: 343. chelicera; 344. venter;

345, leg I (part); 346, corniculus.

154 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

DISTRIBUTION. Nc, Na: Pe, Pm, Ps, Pc. Besides the locality

records published with the original descriptions of the two nominal species

there are many published records from the european Palaearctic region,

ranging from Scotland and the Russian Karelian Isthmus in the north to

Spain and Corsica in the south. There are also records from Kemerovo

(Tagil’tsev and Men Yan-Tsuy’, 1963) and South Primorye (Aksenenko,

1963 and Khudyakov, 1963) regions of Russia and from British Columbia

and Oregon in North America (Emberson, thesis, 1968).

Usually found in nests of rodents, moles and sometimes other

Insectivora, but also found on these animals, in caves (sometimes near bat

roosts), in moss on soil and in forest soil.

REMARKS. There are a number of similarities between Euryparasitus

and Cyrtolaelaps, some of which probably reflect a close phylogenetic

relationship as well as the occupation of a similar ecological niche. As

suggested in the remarks on Cyrtolaelaps, it may be preferable to transfer

members of this genus to Cyrtolaelaps. The following two nominal species

are included in this genus: E. emarginatus (C. L. Koch, 1839); E. calcarator

(Banks, 1910).

Euryparasitus emarginatus (C. L. Koch)

Gamasus emarginatus C. L. Koch, 1839, 24.17.

Gamasus terriblis Michael, 1886, p. 265.

FEMALE. Fig. 338-341 and 347. Idiosomal length, 1,800p.

MALE. Fig. 342-346. Idiosomal length, 1,780. The anterior edge

of the podonotal shield is more heavily sclerotized than in female and

produced forward to cover more of the gnathosoma.

LOCALITY. Two females (N196875 and N196876) and two males

(N196877 and N196878) drawn or examined: Scotland; mole’s nest (4),

Roxburgh, East Lothian, 1964, col. R. M. Emberson, dep. SAM.

Subgenus EVANSSELLUS Ryke

Evanssellus Ryke, 1961b, p. 17. Type-species: Evanssellus foliatus Ryke,

1961b, by original designation.

DIAGNOSIS. Small to average sized mites. Separate podonotal and

opisthonotal shields. Ventro-anal shield discrete on female, while on the

male it is fused to the peritrematal shield and exopodal IV shield and

sometimes to the sternito-genital shield. Single pair of pre-endopodal shields.

Dorsal setae sinuous, lanceolate or spatulate and sometimes pilose, rarely

setose. Spermathecal ringed tube opening not known. Pretarsus I absent.

Large leg I bearing stout spines (anterior spine on genu I is seta all).

LEE—RHODACARIDAE 155

MORPHOLOGY.

SCLEROTIZATION. Female; Separate podonotal and opisthonotal.

Discrete ventro-anal, Posterior to ventro-anal there is a small discrete shield

only bearing aciculae. Peritrematal fused to exopodal IV. Exopodals slim

or absent around middle of peraxial margins of acetabula I] and IIT, Sterno-

metasternal fused to endopodal If. Single pair of pre-endopodals,

Mule: Ventro-anal not fused to opisthonotal, but fused to peritrematal

and exopodal ['V, and may or may not be fused to sternito-genital.

CHALTOTAXY. Tdiosoma: 6j, 62, 5s, Sr: 5/, 52, 58, SR: 4 or Sy:

Bv, 3Zy.

Legs: As Gamasellys.

OrneR CHARACTHRS. Female: Deutosternal denticles in 8 horizontal

rows, Tectum anterior margin basically unispinate but with lateral spinules.

Movable cheliceral digit with 2 teeth. On palp genu, seta all pilose with

over | 1 lateral prongs and seta a/2 spine-like. Dorsal setae are sinuous,

lanceolate or spatulate and sometimes pilose as well. Vertical seta, /1, on

prominent protuberance. Sternal seta, st4, may be absent. Shields dull and

covered by an adhesive exudate. Spermathecal ringed tube opening not

known. Pretarsus absent, The following setae on leg | developed into large

spines: setae all and pv on genu, setac ay2 and pv2 on tibia. Lateral lobes of

pulvilli [I-IV attenuated and longer than central oval lobes. Amongst dorsal

setae on tarsus TY, seta pd3 or ad2 is the longest and both are lanceolate

or spatulate.

Mule: Movable cheliceral digit with | tooth, and fused at base to

slightly longer spermadactyl. On femur [, seta @v enlarged into a spur

which may be quite inconspicuous.

DISTRIBUTION. Aa, An: Sa: NTe. Besides the published records

of nominal species which are all from Australia or New Zealand south of

36° South, [ have seen specimens from Auckland Island to be dep. BBM,

and Athias-Henriot (personal communication, 26.3.1969) informs me that

Evanssellus specimens were collected on the Isla de Chiloc, Chile, 1961-62.

during a study directed by Professor F. di Castri.

Found in moss or plant Jitter,

REMARKS, 1 (Lee, 1967) transferred Evanssellus to Heterogamasus

giving it the rank of subgenus. I here revoke this change in rank, once more

regarding Evanyyellus as a genus. The consistent differences between the

two groups of species (Heterogamasus and Evanssellus), the few individual

members of which exhibit a fair range of characters in both cases, suggest that

these taxa have had a separate evolution considerable enough to be com-

parable with that of other rhodacarid genera. The following two nominal

species are included in this genus: E. foliatus Ryke, 1961; E. medusa Lee,

1967.

156 REC, S&, AUST. MUS. 16 (3): 1-219 July, 1970

Genus HETEROGAMASUS Trigiirdh

Heterogamasus Tragardh, 1907, p. 2. Type-species: Heterogamasus

claviger Triigardh, 1907, by monotypy.

DIAGNOSIS. Small to average sized mites. Separate podonotal and

opisthonotal shields. Ventro-anal shield discrete on female, while on male

it is fused to peritrematal and exopodal IV shields and sometimes to sternito-

genital shield. Single pair of pre-endopodal shields. Dorsal setae spatulate

or lanceolate and sometimes pilose, rarely setose. Spermathecal ringed tube

opening not known. Pretarsus I pedunculate and small. Large leg I bearing

stout spines (anterior spine on genu I is seta av2).

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and _ opisthonotal.

Discrete ventro-anal bearing aciculae posterior to anus. Peritrematal fused

to exopodal TV and extends backwards from this point as well as dorsally, to

broadly fuse with podonotal. Split in exopodals I, UI and IV, Sterno-

metasternal fused to endopodal II. Single pair of pre-endopodals.

Male: Ventro-anal not fused to opisthonotal, but fused to peritrematal,

exopodal ITV and sternito-genital.

CHARTOTAXY, Idiosoma: 6/, 62, 58, 5r: 5/, 52, 5S, 6R, O or 3UR:

Sst; 34, 3Zv, 0-2S8y.

Legs; Ditters trom Gamasellus in having an extra postero-lateral on

genu IIT (2, 4/2, 2) and tibia IIL (2, 3/2, 2).

OTHER CHARACTERS, Female: Deutosternal denticles in 7 horizontal

rows, Tectum anterior margin basically unispinate, but spine is bifurcate

at tip and there are a series of lateral spinules. Movable cheliceral

digit with 3 or 4 teeth, On palp genu, seta all slightly pilose over distal

half and seta al2 simple, Most dorsal setae spatulate or lanceolate and may

be pilose. Vertical seta, jl, not on prominent protruberance. Shicids dull

and covered by an adhesive exudate. Spermathecal ringed tube opening not

known, Pretarsus I pedunculate and smaller than other pretarsi. The

following setae on leg I developed into large spines: setae av2 and py on

genu, setae av2 and pv2 on tibia. Lateral lobes of pulvilli I-IV attenuated

and longer than central oval lobes. Amongst dorsal setae on tarsus TV,

seta ad2 is the longest and is setose,

Male: Movable cheliceral digit with | tooth, and fused at base to

spermadactyl of similar length, On leg Il, femur seta av, and genu seta av

enlarged into spurs,

LEE—RHODAC ARIDAF 157

DISTRIBUTION, NTc: Sm. All records are from near the southern

end of South America and are published with original descriptions of nominal

species.

Found amongst plant litter or under stones,

REMARKS. Previously | (Lee, 1967) redefined Heterogamasus and

included Evanssellus as a subgenus. As remarked on under Evanssellits,

that taxon is again regarded as a genus. The following three nominal species

are included in this genus: H. claviger Triigardh, 1907; H. calcarellus Lee,

1967: H, spinosissimus (Balogh, 1963b).

Genus HINIPHIS gen,n.

Type-species: Hiniphiy hinnus spn.

DIAGNOSIS. Minute mites. Separate podonotal and opisthonotal

shields. Ventro-anal shield is broadly fused to opisthonotal shield, peritre-

matal shield and exopodal IV shield on both sexes and to the sternito-genital

shield on the male. Two pairs of pre-endopodal shields. All dorsal setae

ure simple and tapered, Spermathecal ringed tube opening not known.

Pretarsus | pedunculate.

MORPHOLOGY,

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Ventro-anal fused to opisthonotal, peritrematal and exopodal TV, and bearing

aciculae posterior to anus. Peritrematal fused to exopodal Hl and IV, Split

in exopodals [1] and II. Sterno-metasternal fused to endopodals Il, IT and

IV. Two pairs of pre-endopodals.

Male: Sternito-genital fused to ventro-anal, otherwise as female.

CHAETOTAXY. Idiosoma: 6), 6z, 5s, 5r: S5/, 57, 58, SR: Sst: SJy,

3Zy, 2S8v,

Legs: As Gamasellus.

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows, Tectum anterior margin is unispinate. Movable cheliceral digit

with 3 teeth. On palp genu, seta a/l tapering spine and seta al? is slightly

spatulate. Dorsal setae are simple und tapered. Shields are slightly shiny.

Spermathecal ringed tube opening not known. Pretarsus | pedunculate and

smaller than other pretarsi. Lateral lobes of pulvilli II-IV attenuated and

longer than central oval lobes. Amongst dorsal setae on tarsus IV, seta pd3

is the longest and is setose.

Male: Movable cheliceral digit with | tooth and fused along proximal

half with spermadactyl of similar length. On leg Il, femur seta av, tibia seta

av enlarged into spurs, and femur and genu setae py are small spines, while

tarsus seta av2 is a large spine.

July, 1970

REC, S. AUST. MUS.. 16 (4): 1-219

ae }

: % hf ys

x a 353 ey

348-351 and 355, female: 348. soma. dorsum; 349. leg IV (part), dorsum; 350,

trochanter IV. venter; 351, idiosoma, venter: 355, gnathosoma, venter. 352-354,

male: 352, idiosoma, venter; 353, leg IL (part); 354, chelicera,

LEE—RHODACARIDAE 159

DISTRIBUTION. Aa. Collected from a number of locations in

southern Victoria, Australia, besides the locality given below with the

description of the type-species.

Found in moss.

REMARKS. Hiniphis is unusual in having a divided dorsal shield but

considerable fusion of other shields on the female as well as the male. The

following single nominal species is included in this genus: H. /innus sp.n.

Hiniphis hinnus spn.

FEMALE. Fig. 348-351, 355. Idiosomal length, 390. Trochanter

IV has a non-setous spur.

MALE. Fig. 352-354. Idiosomal length, 360”. Trochanter IV has

a spur that is about half as long as that figured for the female.

LOCALITY. Holotype female (N1968222), allotype male

(N1968223), 4 paratype females (N1968225-N1968228) and 3 paratype

inales (N1968224, N1968229, N1968230) drawn or examined: Australia;

LF104, moss, Pinus radiata plantation, near Beech Forest, Otway Ranges,

Victoria, 9.12.1965, col. D. C, Lee. dep. SAM.

Genus LAELOGAMASUS Berlese

Laelogamasus Berlese, 1905, p. 167. Type-species: Gamuasits (Laeloga-

masus) simplex Berlese, 1905, by original designation.

DIAGNOSIS. Small mites. Separate podonotal and opisthonotal

shields, Ventro-anal shield discrete on female, but fused to exopodal IV

and peritrematal shield on male. Two or 3 pairs of pre-endopodal shields,

Most dorsal setae on idiosoma, and genu and tibia II, Ill and IV, are densely

pilose along their entire length. Spermathecal ringed tube opens distally on

dorsal surface of coxa II]. Pretarsus | pedunculate or absent.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Discrete ventro-anal bearing aciculae posterior to anus. No discrete

imetapodal. Peritrematal fused to exopodal IV. Split in exopodals IH, III

und TY. Sterno-metasternal fused to entire endopodal [1 and part of IIL.

Two or three pairs of pre-endopodals.

Male: Ventro-anal is not fused to sternito-genital or notal but it is

fused to peritrematal and exopodal IV.

CHAETOTAXY. Idiosoma: 6/, 62, 5s, Sr: 5/, 57, 58, 5R, SUR: Sst:

3/y, 3Zv, 3Sv, 2 or 3Ry.

Legs: As Gamasellus,

160 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

OTURR CHARACTERS, Female: Deutosternal denticles in 8 horizontal

rows. Tectum anterior margin having a central spine with multi-

spinulate lateral margin. On palp genu, seta a/l pilose with more than

5 lateral prongs and seta al2 setose or spine-like. Dorsal idiosomal setae all

densely pilose. Shields are dull, Spermathecal ringed tube probably opens

distally on dorsal surface of coxa II]. Pretarsus I pedunculate and smaller

than other pretarsi or absent, Lateral lobes of pulvilli I-IV are attenuated

and shorter or subequal in length to central circular lobes.

Male: Movable cheliceral digit with | tooth, and fused near base to

similarly curved spermadactyl which is subequal in length or slightly longer,

On leg Il, femur seta av enlarged into a spur while most ventral setae on

more distal segments are enlarged into spines.

DISTRIBUTION. Om. Published record of single nominal species

is from Java, but unnamed specimens have been seen from Borneo, Malaya

and Sumatra, dep. BM(NH).

Found amongst plant litter on soil.

REMARKS. Having seen at least 3 conspicuously different species,

with the unusually pilose dorsal setae on the legs as well as the idiosoma,

with the probable, unusually location of the spermathecal ringed tube for

Gamasellus-like mites, and from a limited geographical region, I consider

that Laelogamasus is a distinct genus, possibly relatively unrelated to

Gamasellus. There is | nominal species in this genus, L. simplex Berlese,

1905, but at least 2 unnamed species have been collected from the localities

listed above, dep. BM(NH),

Laelogamasus simplex ( Berlese)

Gamasellus (Laelogamasus) simplex Berlese, 1905, p. 167.

FEMALE. Redescribed by Berlese, (1906, p. L115).

MALE. Fig. 319-322. Idiosomal length, 440. Apparently all that

remains of the types is on the slide listed below. Most of this material (that

can be seen without remounting) is drawn.

LOCALITY. Some of the ‘cotipico’ male (33/4), slide has

gnathosoma and parts of legs only, drawn: Java; Island of aba, Buitenzorg,

Genus LITOGAMASUS gen.n.

Type-species: Cyrtolaelaps setosus Kramer, 1898,

DIAGNOSIS. Small to pigantic mites, Separate podonotal and

opisthonotal shields. Ventro-unal shield discrete on female and mate.

Peritrematal shield always fused to exopodal IV shield on female but inay

LERF—RHODACARIDAFR lo}

not be on male. May be single pair of pre-endopodal shields, but more

often there are a number of small shields posterior to the main pair of shields.

Dorsal setae tapered and may or may not be slightly pilose. Spermathecal

ringed tube opening not known. Male fixed cheliceral digit has conspicuous

ridges dorsally, each side of a groove in which the pilus dentilis is located.

Pretarsus | not pedunculate, sheath being broadly fused to tarsus and only

long enough to contain retracted claw complex,

MORPHOLOGY,

SCLEROTIZATION, Female; Separate podonotal and opisthonotal.

Ventro-anul discrete, may be narrow (beuring less than the 6 pairs of setac

in setal rows Jy and Zv) and may or may not bear aciculae. Posterior to

ventro-anal there may be a small discrete shield only bearing aciculae.

Discrete metapodal, Peritrematal reduced and narrowly joined to exopedal

TV. Exopodals slight and not split around peraxial edge of acetabula [1

and IY, Sterno-metasternal fused to endopodal IT. Usually 2 pairs of

pre-endopodals with the posterior or both pairs fragmented, but may be only

single pair,

Male: Ventro-anal discrete. Sternito-genital not fused to ventro-anal

or endepodal IV. Peritrematal may or may not be fused to exopodal IV.

CHAE TOTAXY, Idiosoma: 6), 6c, 5s, Sr: SJ, 527, 4 or 5S, 3 or 5R:

Sst: 34/y, 3Zv, 1 or 28v,

Legs: As Gamavellus.

OTHER CHARACTERS. Female: Deutosternal denticles in 7 or 8 hori-

vontal rows. Tectum anterior margin basically unispinate, but may have

a few lateral spinules. Movable cheliceral digit with 3 teeth. On palp

genu, seta a/| tapered and slightly pilose, and seta a/2 spatulate or spine-

like. On pulp femur, seta a/ usually in proximal third, but in one unnamed

species fron South Georgia it is in central third (the only species of

Ologamasinae for which this is true). Dorsal setae tapered and sometimes

incouspicuously pilose, Shields shiny, Spermathecal ringed tube opening

not known. Pretarsi sheath | not pedunculate only being large enough to

contain retracted claw-complex. Pulvilli J[-LV have a pair of streamer-like

lobes ventral to claws, while dorsally there may be attenuated central and

lateral lobes or a single pair of large circular lobes. Amongst dorsal setae

on tarsus IV, setae ad4 and pd4 may be the longest, or they may be short

and spine-like and seta pd3 the longest,

Male: Movable cheliceral digit with single tooth and fused at base to

attenuated spermadactyl or similar length. Fixed cheliceral digit ridged

dorsally, On leg Il, femur setae av and py, genu seta av, tibia seta av, tarsus

seta @v2 modified into spurs or spines. Other legs may have modified sctous

or non-setous processes,

AUST. MUS., 16 (3): 1-219 July, 1970

Figs. 356-364. Litogamasus setosus (Kramer).

356-358 and 364, female: 356, soma, dorsum; 357, gnathosoma, yenter; 358, idiosoma,

venter: 364, pretarsus | and HI. 359-363, male: 359, leg If (part). antero-latus:

360. tarsus IH: 361, tibia and tarsus IV: 362, chelicera; 363. idiosoma, venter.

LEE—RHODACARIDAE 163

DISTRIBUTION. Sm, Sa: ACs. Besides the published record of the

single nominal species from Tierra del Feugo, I have seen an unnamed species

and L. setosus trom Auckland and Campbell Islands to be dep. BBM, and

another unnamed species from South Georgia, dep. SAM.

Found in or near to littoral zone.

REMARKS, Excepting the division of the dorsal shield Litogamasus

has a number of characters in common with Parasitiphis, but these may

only be an indication that they occupy a similar habitat. The following

single nominal species, previously in Cyrtolaelaps, is included in this genus;

L. setosus (Kramer, 1898) comb.n.. Two unnamed species have been

described, one from South Georgia by Hunter (manuscript), some dep.

SAM, and the other from Auckland and Campbell Islands by Hunter and

Lee (manuscript) dep, BBM,

Litogamasus setosus (Kramer) comb.n,

Cyrtolaelaps setosus Kramer, 1898, p. 22.

FEMALE, Fig. 356-358, 364. I[diosomal length, 1,410p.

MALE, Fig. 359-363. Idiosomal length, 1,420”. Tectum is slightly

broader and more heavily sclerotized than on female. The processes on

tarsus IIL are non-setous. On leg [V, genu ventral setae ure spine-like as

are those on the tibia, Leg IV is enlarged to u similar extent to that of

leg VW,

LOCALITY. Type female (137) and type male (117) drawn: South

America; under stone, at mouth of brook above high tide mark or seashore,

Uschuaia, Tierra del Fuego, Argentina, 27.10.1892, col. Michaelsen, dep,

ZMH.

Genus NOTOGAMASELLUS Loots and Ryke

Notogamasellus Loots and Ryke, 1966b, p. 30. Type-species: Notogama-

sellus (Notogamasellus) vandenbergi Loots and Ryke, 1966b, by original

designation.

DIAGNOSIS. Minute mites. Separate podonotal and opisthonotal

shields, podonotal shield being conspicuously larger than opisthonotal shield,

Ventro-anal shield discrete on the female and male, but on male it ts larger

extending anteriorly to carrying more setae. Peritrematal shield reduced

and may or may not be narrowly connected to exopodal TV shield. Setation

of podonotum hypertrichous. Single pair of pre-endopodal shields. The

short dorsal setae may all be pilose or most of them are peg-like with a central

spherical dilation. Location of spermathecal ringed tube, if present, unknown,

Pretarsus 1] absent,

164 REC. S$. AUST. MUS., 16 (3): 1-219 July, 1970

REMARKS. Notogamasellus was originally defined with two sub-

genera, Notogamasellus and Podonotogamasellus, which may be distinguished

by the following key. The morphology and distribution of the members of

this genus are given under the subgeneric headings.

KEY TO SUBGENERA OF NOTOGAMASELLUS

1. Two postero-lateral setae on genu LIL (2,

4/2, 2) and tibia III (2, 3/2, 2).

Podonotum with 32 pairs of setae and

One unpaired seta... .. .. .. .. .. N. (Notogamasellus)

One postero-lateral seta on genu III (2.

4/2, 1) and tibia HI] (2, 3/2, 1).

Podonotum with 28 pairs of setae .. N. (Podonotogamasellus )

Subgenus NOTOGAMASELLUS Loots and Ryke

Notogamasellus Loots and Ryke, 1966b, p. 30. Type-species: Notogama-

sellus (Notogamasellus) vandenbergi Loots and Ryke, 1966b, by original

designation.

DIAGNOSIS. As for genus with the addition of the characters given

in the relevant half of the couplet in the key to subgenera of Notogamasellus.

MORPHOLOGY.

SCLEROTIZATION, Female: Podonotal and opisthonotal, Discrete

ventro-anal bearing aciculae posterior to anus. Peritrematal reduced,

narrowly connected to exopodal IV. Exopodals inconspicuous. Sterno-

metasternal fused to endopodal If and part of III. Single pair of

pre-endopodals.

Male: Discrete sternito-genital and ventro-anal, but the latter extends

further anterior than in the female so that it also carries setae Jvl, J/v2

and Zyl.

CHAETOTAXY. Idiosoma: | unpaired, 9j, 82, 8s, 7r: SJ, 52, 5S, SR:

Sst: 1 or 2 unpaired, 3/v, 3Zy, LSy.

Legs: Differs from Gamasellus in having an extra postero-lateral on

genu III (2, 4/2, 2) and tibia Il (2, 3/2, 2).

OTHER CHARACTERS, Female: Deutosternal denticles in 7 horizontal

rows. Tectum anterior margin is basically unispinate, the spine being

narrow and parallel sided for most of its length, but most of the margin,

including the tip of the spine, is multispinulate. Movable cheliceral digit

LEE—RHODACARIDAE 165

with 3 teeth. Form of setae on palp genu unknown. All dorsal setae are

pilose. Shields dull with adhesive exudate so that fragments of the nymphal

exoskeleton and other detritus cover the idiosoma of the specimens described.

Pretarsus [ absent.

Male: Movable cheliceral digit with 1 tooth, and fused at base to

attenuated spermadacty! of similar length. On leg Il only femur seta av is

enlarged into spur.

DISTRIBUTION. Es. The single nominal species has only been

recorded from the Transvaal, South Africa.

Found in plant litter on forest floor.

REMARKS. The following single nominal species is included in this

subgenus: N. (N.) vandenbergi Loots and Ryke, 1966b.

Subgenus PODONOTOGAMASELLUS Loots and Ryke

Podonotogamasellus Loots and Ryke, 1966, p. 471. Type-species: Noto-

gamasellus (Podonotogamasellus) magoebaensis Loots and Ryke, 1966,

by original designation.

DIAGNOSIS. As for genus with the addition of the characters given

in the relevant half of the couplet in the key to subgenera of Notogamasellus.

Males of this subgenus are unknown.

MORPHOLOGY.

SCLRROTIZATION, Female: Similar to that of the subgenus Notegama-

sellus, except that peritrematal is not narrowly connected to exopodal IV.

CHAETOTAXY. Idiosoma: 7/, 7z, 7s, Tr: S5/, 5Z, 58, 5R: Syts |]

unpaired, 3/1, 3Zv, 1Sv.

Legs: As Gamasellus,

OTHER CHARACTERS, Female; Similar to that of the subgenus

Notogamasellus, except that the form of setae on palp genu is known and

the form of the dorsal setae is different. On palp genu, seta a/l is pilose

with 6 short lateral prongs and seta a/2 is simple, setose. The dorsal setae

are mostly short and peg-like with a central spherical dilation, and the

proximal half of the seta is covered by a membranous flap. The vertical

seta jl is lanceolate with a serrate edge and with | proximal tooth enlarged

so that it is nearly as long as the whole seta.

DISTRIBUTION, Es. Transvaal.

Found in plant litter on forest floor.

REMARKS. The following single nominal species is included in this

subgenus: N. (P.) magoebaensis Loots and Ryke, 1966b.

166 REC. S. AUST. MUS., [6 (3): 1-219 July, 1970

Genus PERISEIUS Womersley

Periseius Womersley, 1961, p. 198. Type-species: Periseius littorale

Womersley, 1961 (syn. Cyrtolaelaps hammeni Womersley, 1961), by

original designation.

Psammonsella Hag, 1965, p. 413. Type-species: Psammonsella nobskae

Hag, 1965, by monotypy.

DIAGNOSIS. Minute to small mites. Separate podonotal and

opisthonotal shields. Ventro-anal shield separate from notal shield but fused

to peritrematal and exopodal IV shields on both female and male, and also

fused to sternito-genital shield on male. ‘Three pairs of pre-endopodal

shields, Some dorsal setae may be pilose or all dorsal setae may be simple,

tapered. Spermathecal ringed tube opens near posterior paraxial edge of

acetabulum IV. Pretarsus I pedunculate and smaller than other pretarsi,

REMARKS. Hirschmann (1966) redefined Periseius and transferred

Psammonsella to it with subgeneric status. This concept is followed here,

and members of the two subgenera can be distinguished by the following

key. The morphology and distribution of members of this genus are given

under the subgeneric headings.

KEY TO SUBGENERA OF PERISEIUS

I. Dorsal setae r3 and 275, and postanal seta, are

pilose and spatulate. Female metasternal

shield is fused to sternal shield .. .. .. .. P. (Periseius)

Dorsal setae r3 and Z5, and postanal seta, are

simple and tapered. Female metasternal

shield is not fused to sternal shield .. .. .. P. (Psammonsella)

Subgenus PERISEIUS Womersley

Periseius Womersley, 1961, p. 198. Type-species: Periseius littorale

Womersley, 1961 (syn. Cyrtolaelaps hammeni Womersley, 1961), by

original designation.

DIAGNOSIS. Small mites with some pilose dorsal setae and female

metasternal shield fused to sternal shield.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Ventro-anal fused to peritrematal, and both are fused to exopodal IV.

Aciculae on ventro-anal, posterior to anus. Exopodals II, HI and IV are

split. Sterno-metasternal fused to endopodals II, I] and IV. Three pairs

of pre-endopodals.

LEE—RHODACARIDAE 167

Male; Ventro-anal| fused to peritrematal, exopodal [VY and _sternito-

genital,

CHAWHTOTAXY. Tdiosoma: 6j, 5 of 6z, 5s, Se: SJ, 5Z, 5 or 6S, OR:

Sst: 3/v, 32¥, 1Sy.

Legs: As Gamasellus on type-species but an extra postero-lateral on

genu HI (2, 4/2, 2) and tibia IIE (2, 3/2, 2) of an unnamed species from

North America.

Orne CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows, Tectum anterior margin basically uninspinate with spinules on

sides, Movable cheliceral digit with 3 teeth. On palp genu, seta all

spatulute with single prong each side and seta a/2 slightly spine-like. Some

dorsal setue muy be simple and tapering, but there are always pilose, and

pilose and spatulate setae present. Spermathecal ringed tube probably opens

nea’ posterior paraxial margin of acetabulum TV. Pretarsus I pedunculate

and smaller than other pretarsi. Lateral lobes of pulvillii H-IV attenuated

und longer thin central oval Jobes. Amongst dorsal setae on tarsus IV, seta

pd3 is the longest and ts lanceolate.

Male: Movable cheliceral digit, with | tooth, and fused at base to

slightly longer, but attenuated spermadactyl. On leg I, seta femur av,

sometimes seta pvl, and genu seta av enlarged into spur, Sometimes non-

setaus processes on leg I.

DISTRIBUTION. Na: NTb: Am. The three known records are from

between the latitudes 25°N and 10°S, being from Biak Island off the north

coast of New Guinea, Recife on the Brazilian coast and a record (Emberson,

thesis, 1968) of an unnamed species from the Florida Keys, only just north

of the tropic of Cancer.

Found in or near littoral zone,

REMARKS. This taxon, when originally established as a genus,

contained only the type-species, P, littorale, apparently only known from the

deutonymphs, but Cyrtolaelaps hammeni adults described in the same paper

ure now recognized as conspecific with those deutonymphs and hammeni

has been chosen as the valid name. The following two nominal species are

included in this subgenus: P. (P.) hammeni (Womersley, 1961) comb.n.;

P. (P.) brasiliensis Hirschmann, 1966, A unnamed species is partly

described by Emberson (thesis, 1968).

Periseius hammeni (Womersley) comb.n.

Cyrtolaelaps hammeni Womersley, 1961, p. 190,

Periseius litterale Womersley, 1961, p. 198, synn.

SCLEROTIZATION. Womersley's (1961) drawings of the distribu-

tion und fusion of the shields on the adults and deutonymphs are inaccurate.

168 REC. S. AUST. MUS., 16 (3): 1-219 July, 1970

The drawings by Hirschmann (1966) of P. brasiliensis are very similar to

the actual sclerotization on P. hammeni. The main differences are to

Hirschmann’s drawing (Fig. 22VW, 1966) of the female venter. On the

venter of the female of P. hammeni there is a narrow fissure of striated

cuticle between the genital and the ventro-anal, and there is no indentation

on the posterior edge of sterno-metasternal running in behind pore 4, just

anterior to seta st4. The important differences between P. hammeni and

Womersley’s (1961) drawings are that the ventro-anal is fused to the

peritrematal in both sexes, the female metasternal is fused to the sternal and

the male sternito-genital is fused to the ventro-anal. Also, the deutonymph

has a widely separated podonotal and opisthonotal.

LOCALITY. Paratype female (N1968212), paratype male

(N1968213) and paratype deutonymph (N1968214) examined: New

Guinea; Cladophora socialis and red algae, on stones, intertidal zone, Biak

Island, 10.12.1953, col. L. V. D. Hammen, dep. SAM.

REMARKS. A comparison between C. hammeni described only from

adults, and P. littorale (should be changed to littoralis because Seius is

masculine) described only from deutonymphs in the same collection, as well

as a comparison between this material and the drawings of the adults and

deutonymph of P. brasiliensis, leaves no doubt about the synonymy of the

first two species names. I have chosen hammeni as the valid name for three

reasons: it is the name for the adult; the name Jittoralis is valid for the type

species of two other rhodacarid genera (Hydrogamasus and Parasitiphis) ;

the name hammeni has precedence of position in the text. Therefore, despite

the name Jittorale being the first name used for specimens on which the

genus was originally based, I have made it invalid.

Sugenus PSAMMONSELLA Haq

Psammonsella Haq, 1965, p. 413. Type-species: Psammonsella nobskae

Haq, 1965, by monotypy.

DIAGNOSIS. Minute mites with no pilose dorsal setae and female

metasternal shield separate from the sternal shield.

MORPHOLOGY.

SCLEROTIZATION. Similar to that of the subgenus Periseius, except

that female ventro-anal is more narrowly fused to the peritrematal and the

metasternal is entirely separate from the sternal.

CHAETOTAXY. Idiosoma: 6j, 6z, 5s, 5r: 5/, 5Z, 58, 5R: Sst: 3/y,

3Zyv, 1Sv.

Legs: As Gamasellus.

LEE—RHODAC ARIDAE 169

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows. Tectum anterior margin basically unispinate with spinules on

sides, Movable cheliceral digit with 3 teeth, On palp genu, seta all

spatulate with single prong on each side and seta a/2 slightly lanceolate.

Dorsal setae simple, tapered, and podonotal setae sl, .»2 and r3 much longer

and stouter than other notal setae. Spermathecal ringed tube opens near

posterior paraxial margin of acetabulum TV, Pretarsus | peduncuiate and

smaller than other pretarsi. Lateral lobes of pulvilli II-[V attenuated and

more than twice as long as central oval lobes. Dorsal setae on tarsus TV

short except for seta ad2 which is long and setose.

Male: Movable cheliceral digit with | tooth (may be small second

tooth) and fused at base to spermadactyl of similar length. On leg I,

femur seta ay, genu seta ay, tibia seta av enlarged into spurs.

DISTRIBUTION, Na: Pm, All records are from the northern

mediterranean coastline or from a similar latitude on the east coast of North

America (Massachusetts ).

Found in littoral zone.

REMARKS, — Hirschmann (1966) transferred Psammonsella to

Periseius with subgeneric status. It may later be preferable not to recognize

subgenera within this genus. I have seen paratypes of the two species

(P. nobskue dep. Mrs. Haq’s personal collection, P, sehusteri dep. SAM,

N1968258-N1968268) and Hirschmann’s key (1966, p. 5) does fot

distinguish between them, so they may be conspecific. The following 2

nominal species are included in this subgenus: P. (P.) nobskae Haq, 1965;

P. (P.) sehusteri Hirschmann, 1966.

Genus PILELLUS gen.n.

Type-species: Cyrtolaelaps (Gamasellus) rykei Hunter, 1967,

DIAGNOSIS. Small to average sized mites, Separate podonotal and

opisthonotal shields. Ventro-anal shield discrete on female, but fused to

sternito-genital, exopodal LV and peritrematal shields on male, and may also

be fused to notal shield. One pair of pre-endopodal shields. Most dorsal

setae are short and lanceolate, and at least one pair of dorsal setae are pilose.

Setation of both podonotum and opisthonotum is hypertrichous. Sperma-

thecul ringed tube opens near posterior paraxial edge of acetabulum IV.

Pretarsus | pedunculate,

MORPHOLOGY,

SCLEROTIZATION, Female: Separate podonotal and opisthonotal.

Discrete ventro-anal bearing aciculae posterior to anus. No discrete

metapodal. Peritrematal fused to exopodal TV. Split in exopodals TT, IL

V7 REC. S$. AUST. MUS., Ih (35: 1-219 July. 1970

and [V, Sterno-metasternal fused to endopodal If and part of Hl. Single

pair of pre-endopodals,

Male: Ventro-anal may be either separate from or fused to opistho-

notal, but always fused lo sternito-genital, exopodal IV and peritrematal,

CHAETOTAXY. Idiosoma: 6j, 6z plus 16 or 17: over 60 opisthonotal

setae, some paired, some single accessory setae: Sst: 3/y, 3Zy, plus 0 or 9

or more.

Lees: As Gamavyellus,

OTHER CHARACTERS. Female; Deutosternal denticles in 7 horizontal

rows, Tectum anterior margin with central spine with uneven, lateral

spinules. Movable cheliceral digit with 3 teeth. On palp genu, seta

all with | or 4 lateral prongs and seta a/2 spine-like or lanceolate.

Dorsal setae either lanceolate, pilose or pilose-lanceolate. Shields are dull.

Spermathecal ringed tube probably opens near posterior paraxial edge of

acetabulum IV, Pretarsus I pedunculate and smaller than other pretarsi.

Lateral lobes of pulvilli I-IV are attenuated and longer than central, circular

lobes. Amongst dorsal setae on tarsus IV, seta ad2 is the longest and is

setose,

Male: Movable cheliceral digit with 1 tooth, und fused at base to

similarly shaped spermadactyl, that is subequal in length or longer. On

femur IT, seta av is enlarged into a spur, while pvl is spine-like. Setae of

other leg I] segments are similar to those on female,

DISTRIBUTION. Sa: ACs. The single nominal species is recorded

from Candlemas Island and South Sandwich Islands, but an unnamed species

is recorded from Auckland and Campbell Islands by Hunter and Lee

(manuscript), dep. BBM.

Found in moss mats.

REMARKS. The only nominal species in Pilellus was previously in

Gamasellus. Although it is not congeneric with G. falciger, | could have

transferred this species to a special species-complex in Acwgamasus. Instead,

a new venus is established for it, distinguishable from Acugamasus by the

chaetotaxy and form of its dorsal setae. There is one nominal species

included in this genus, P. rykei (Hunter, 1967) comb.n., and an unnamed

species is described by Hunter and Lee (manuscript), dep. BBM.

Genus RHODACAROIDES Willmann

Rhodacararoides Willmann, 1959, p, 97, Type-species: Rhodacaroides

degyptiacus Willmann, 1959, by original designation,

DIAGNOSIS, Small mites, Separate podonotal and opisthonotal

shields, Ventro-anal shield discrete on both sexes. Two pairs of pre-

endopodal shields. Dorsal setae all simple. Location of the spermathecal

LEE—RHODACARIDAF 71

ringed tube, if present. is unknown. Spermadactyl fused to base of movable

cheliceral digit and runs parallel to it as on most Gamiasellini males.

Pretarsus 1 pedunculate.

MORPHOLOGY.

SCLEROTIZATION. Female: Separate podonotal and opisthonotal.

Discrete ventro-anal. Peritrematal fused to exopodal TV by a narrow strip.

Exopodals do not completely encase the peraxial edges of the acetabula.

Sterno-metasternal fused to endopodals Il and [ll Two pairs ol

pre-endopodals,

Male: Sternito-genital shortened posteriorly so that it does not carry

seta sf5,

CHAETOVAXY. Idiosoma: 6/, 6z, 6s, Sr: 5J, SZ, 58, OR: Sats By,

3Zv, ISv.

Legs: Not known, but assumed to be as Gamasellus.

Orner CHARACTERS. Female: Nature of deutosternal denticles 1s

unknown, Tectum anterior margin trispinate with large central spine.

Movuble cheliceral digit with 4 teeth. Although the rows of podonotal setae

contain the same numbers as Rhodacarus, their positions are more like some

Afrogamasellus, especially in seta sl being well posterior to seta <1 thereby

leaving only 3 pairs of seta on the anterior margin of the podonotal shield,

Dorsal setae simple, tapered. Spermathecal ringed tube aperture not known,

Pretarsus 1 reduced, but pedunculate and with pulvillus.

Male: Movable cheliceral digit with 1 tooth and fused at base to

similarly shaped, but attenuated and longer, spermadactyl, On leg LH, femur

seta av, venu seta av and tibia seta av enlarged into spurs,

DISTRIBUTION. Pm. Single record with the original description

of the type-species is from the Red Sea coast line, Egypt.

Found in moist soil bordering sea.

REMARKS. Because of a number of characters on the type-species

not being known, the diagnosis of Rhodacarvides is difficult. Neither

Willmano nor Schulz (personal communications, 8.1967) were able to

locate the types of R. aegyptiacus, but they are now in the possession of

Dr. W. Hirschmann who will soon be publishing a redescription of this

species. The placing of Rhodacaroides in the Ologamasinae rather than the

Rhodacarinae (the members of which it was previously allied to) is specula-

tive. The speculation is made partly because of a number of unnamed

species (two partly described by Emberson, thesis, 1968, and two collected

by myself in South Australia) which are Rhodacarus-like, but clearly belong

172 REC, S. AUST, MUS., 16 (3); 1219 July, V70

in the Ologamasinae, and do have characters similar to those of R.

aegyptiacus. The following single nominal genus is included in this genus:

R. aegyptiacus Willmann, 1959. Rhodacarus costai, incertae sedis, has been

transferred to this genus because it should be in the Ologamasinae, but it is

improbable that it belongs in this genus,

Subfamily SESSILUNCINAE subf.n.

Type-genus! Sessiluncus G. Canestrini, 1898,

DIAGNOSIS. Minute to gigantic mites, usually with moderately well

sclerotized and extensive shields. Holonotal shield or more rarely separate

podonotal and opisthonotal shields. Ventro-anal shield only fused to notal

in one atypical genus; Stylochiruy. Peritrematal shield often separate from

exopodal TY shield but if fused to this shield, then it is completely merged

with it from level of stigma back. Sclerotization not conspicuously sexually

dimorphic. On the female sterno-metasternal shield a line joining setae sf2.

st3_ and yt4 would or would not enclose an angle of less than 95°.

MORPHOLOGY.

SCLUROTIZATION, The extent of the fusion between shields is fairly

constant amongst species of this subfamily, and except for the fusion of

the genital with other shields it is nearly always constant amongst females

and males of the same species. In cight genera there is a holonotal, although

Gamasellevans has a line of demarcation between the podonotum and

opisthonotum. Paragamasellevans is the only genus with a separate

podonotal and opisthonotal, and the opposing edges of these shields are

touching. Except in S/y/ochirus the ventro-anal is separate from the notal,

but in some undescribed males of Antennolaelaps, where the aciculae are on

the ventro-anal. it lies so close to the notal in the region posterior to the anus

that it appears to be fused to it on superficial examination, Usually the

ventro-anal is separate from the exopodals, but in Gamasellevans, where the

peritrematal is always reduced and free posteriorly, & posterior extension of

exopodal [TV (which may represent the metapodal) may be broadly or

narowly fused to the ventro-anal. The peritrematal is free posteriorly in

Sessduncus, Antennolaelaps, Gamasellevans, Gamasitus, the Onchogamasus

pumilio-complex. and Paragamasellevans, \n Styvlochirus the peritrematal

may be free posteriorly or fused to the notal along its entire length, In

Gamasellopsis, Onchogamasus communis and Queenslandolaelaps the peri-

trematal is fused to at least exopodal TV, The exopodals may form a

continuous strip, but in Gamasellopsis, Onchogamasus communis and

Queenslandolaelups there is a split in exopodal If. and in Gamuasellevans,

Gamasitus, the Onchogamasusy pumtlio-complex and Paragamasellevans there

LFE—RHODAC ARIDAF 173

is a split in exopodal Il and Ill, The sterno-metasternal is separate [rom

endopodal IV in most Gamasellopsis species, Paragamasellevans and

Stylochirus, but usually it is fused to endopodal 1V. There may be a single

pair of pre-endopodals or they may be fused to the sterno-metasternal.

CHAETOTAXY. Idiosoma: 67, 6z, Ss, 4 or Sr: SJ, 524, 58, 4 or 5R,

Nor 2UR: Syst) 3/y, 3Zv, 1 or 28v. Stylochirus species ure exceptional in

that they have a hypertrichous opisthosoma.

Legs: As Gamasellus except in Sessiluncus, Gamasellopsis and

Gamasilus where there may be between 2 and 7 setae less, and there is

always only | ventral on genu UT and [V.

OTHER CHARACTERS. Tectum anterior margin varies from basically

ubispinate with lateral spinules to trispinate with spinules making structure

complex. Fixed cheliceral digit of males of Sessiluneus, Gamasellevans,

Paragamasellevans and Queenslandolaelaps carries tubercle of process

dorsally, Usually on palp genu seta all is pectinate and seta a/2 is setose

or spatulate, @ combination often found in Ologamasinac, especially

Ologamasini. On palp femur. seta a/ is on central third in Sessiluncus,

Gamasellevans, Gamasellapsis, Qnchogamasuy communis and Stylochirus

as in the Rhodacarinne and Tangaroellinae, but otherwise it is on the

proximal third as in most Rhodacaridae. The female sternal seta sr2, sf3

and s/4 are usually nearly in a straight line as in most Ologamasinae, but in

Antennolaelaps, Onchogamasus communis and Queenslandolaelaps & line

joining these setae would enclose an angle of less than 95° as in the

Gamasiphinae. Usually idiosomal and leg setae are simple, setose, but

although they ure never spatulate they may be pilose. Some Antennolaelaps

species ure the only rhodacarids with pilose sternal setae. Spermathecal

ringed tube opens near posterior paraxial edye of acetabulum TV in

Antennolaelaps and Queenslandolaelaps and probably so in Paragama-

sellevans species. If pretarsus L present it may be pedunculate or in

Sessiluncus, Gamasellevans and Paragamasellevans the pretarsal sheath T Is

only long enough to contain retracted claw-complex and is often broudly

altached at base so that it merges into tarsus.

DISTRIBUTION. Only one genus, Stylechirus, has species occurring

in the Nearetic region and no members of the subfamily have been collected

from the Neotropical region, Except for Sessi/uncus, the other genera are

contined to the Southern Hemisphere and upparently only occur in either the

Ethiopian or the Australian region.

REMARKS. | have contidence in my grouping together of all but one

of the genera in the Sessi/uncinae, and that is Stylochirus, which is not

closely allied to the other genera and is placed in the Sessiluncinue in

preference to any other subfamily.

174

Gamasellevans,

sellevans, Queenslandolaelaps and Stylochirus.

REC. 8S. AUST. MUS., 16 (3):

1-219

July, 1970

I recognize 9 genera within this subfamily; Sessiluncus, Antennolaelaps,

Gamasellopsis, _Gamasitus,

be distinguished by the following key.

. Pretarsus I pedunculate.

. Peritrematal

. Peritreme with a series of lateral pockets.

Onchogamasus,

Adults of these genera can

Paragama-

KEY TO GENERA OF SESSILUNCINAE

. Ventro-anal shield fused to posterior end of notal

shield . ort hilar ge ie ee word

Ventro-anal shield separate from notal shield ..

. One ventral seta on genu III (2, 4/1, 1) and

genu IV (2,4 0r 5/1, 1) .. ..

Two yentral setae on genu HI (2, 4/2,

1) and

genu IV (2, 5/2, 1) ob

. Two ventral setae on genu T (2, 6/2, 2) and

peritrematal shield fused to exopodal

shield .. 1 Mpsteal cae” de aheleae KD SS

Three ventral setae on genu I (2, 6/3, 2) and

peritrematal shield separate from exopodal IV

shield ..

Exopodal III shield

split level with centre or acetabulum

Pretarsus I not pedunculate, having sheath only

long enough to contain retracted claw-

complex. Exopodal shield Hf continuous

around peraxial edge of acetabulum

shield never extends backwards

posterior to level of mid-acetabulum IV .

Peritrematal shield extends backwards to level of

posterior margin of acetabulum IV ..

Line

joining female sternal setae s/2, st3, and st4

would enclose an angle of less than 95°.

Male chelicerae normal with spermadactyl

running parallel to digits from point of attach-

ment near base of movable digit .

Stylochirus

Gamasellopsis

Gamasitus

Sessiluneus

Antennolaelaps

LEE—RHODACARIDAE 175

Peritreme without a series of lateral pockets.

Line joining female sternal setae sf2, st3 and

st4 would enclose an angle of more than 110°.

Male cheliceral fixed digit with tubercle or

process and spermadactyl curving away from

movable digit at point of attachment near

distal: tip “sf 4. ey igs bat Ges

7. Separate podonotal and _ opisthonotal shields.

Seta st5 on female genital shield is posterior

to acetabulum IV. Male seta st5 off sternito-

genital shield .. 2. 2. 2. 2. 2. 2. 2... =Paragamasellevans

Holonotal shield, but usually with clear demarca-

tion line between podonotal and opisthonotal

areas. Seta st5 on female genital shield level

with posterior edge of acetabulum IV .. .. Gamasellevans

8. Single conspicuous pore posterior to stigma on

peritrematal shield. Tectum anterior margin

basically trispinate, with central spine narrow

at base and spreading to broader, fimbriate

tip. Male cheliceral fixed digit with dorsal

process and spermadactyl curving away from

movable digit at point of attachment near

distal tip .. 2. 2. 2. 0. ee we ee a ~~ Queenslandolaelaps

Two conspicuous pores posterior to stigma on

peritrematal shield. Tectum anterior margin

basically unispinate although there may be

lateral spinules on spine which is broad at

base, narrowing to a pointed tip. Male

unknown .. .. .. 2. 2. 2. ee ee ee ee) )© Onchogamasus

Genus SESSILUNCUS G. Canestrini

Sessiluncus G,. Canestrini, 1898, p. 486. Type-species: Gamasus hetero-

tarsus G. Canestrini, 1897, by original designation.

DIAGNOSIS. Small to large mites. Holonotal shield. Ventro-anal

shield separate from notal shield but apparently fused to exopodal IV shield

on both sexes. Peritrematal shield not fused to exopodal shields. Sterno-

metasternal shield fused to endopodal IV shield and a line joining sternal

setae st2, st3 and st4 would enclose an angle of more than 110°. Sperma-

dactyl fused to base of male movable cheliceral digit and strongly recurved.

176 REC, 8S, AUST, MUS. 16 (3); 1-219 July, 1970

One ventral seta on genua IT and PV. Pretursus I not pedunculate, sheath

being broadly fused to tarsus and only long enough to contain retracted

claw complex.

MORPHOLOGY.

SCLEROTIZATION. Female; Holonotal. Ventro-anal separate from

notal, bearing aciculae posterior to anus and apparently fused to exopodal

IV. No discrete metapodal, may be represented in posterior extension of

peritrematal, Peritrematal not fused to exopodals. Exopodals I, HI and

IV form continuous strip, Sterno-metasternal usually fused to endopodals

Hl, If and IV, but may be separate from endopodal [V (as on an unnamed

species from Nepal, dep. BM(NH)), and sometimes fused to the single pair

of pre-endopodals.

Male: Sternito-genital, otherwise as female, except perilrematal may

extend further back,

CHAETOTAXY. Idiosoma: 6), 62, 5s, 4r: 5/, 52, 5S, 5R: Syrs

3/v, 3Zy.

Leys: Following segments differ from Gamasellus in having ventral or

sometimes a postero-lateral seta missing; genu IL, (2. 4/1. 1): genu TV,

(2, 5/1, L); tibia TY, (2, 4/2, | or 2 as for Gamasellus).

OTHER CHARACTERS, Female: Deutosternal denticles in 8 or 9

horizontal rows, Corniculus may be simple or it may have a paraxial

hyaline flange. Tectum anterior margin is trispinate, the central spine being

the largest. Movable cheliceral digit with 3 teeth. On palp genu, seta a/!

finely pilose and nearly level with a simple or slightly spatulate seta a2,

On palp femur, a@/ in central third, Dorsal setae, simple or slightly pilose

and tapering tip may be coiled, Shields often strongly reticulated or pitted

but are shiny, Peritreme may be sinuous and also have lateral pockets, On

sternum a line joining setae s72, s73 and st4 would enclose un angle of more

than 110°. Spermathecal ringed tube openiny not located in this study, but

Athias-Henriot (in press) states that it opens on coxa HIE or coxa TV.

Pretarsus IT not pedunculate but broadly fused to tarsus and sclerotized, so

that the large claws appear retractable into tarsus, Lateral lobes of pulvilli

I-IV attenuated and shorter than central circular lobes. Amongst dorsal

setae On tarsus IV, seta ad2 is the longest. There may be spurs on the femur

of leg ILE and the trochanter and femur of leg IV.

Male: Corniculus with hyaline paraxial flange similar to that of female,

Movable, cheliceral digit with | or 2 tecth, and fused at base to strongly

recurved spermadactyl. Fixed cheliceral digit may have dorsal process, On

leg I, setae av on femur, genu and tibia enlarged into spurs, Postero-ventral

setae on femur und genu may be spines. There may also be spurs on the

femur, genu and tibia of leg HL and the trochanter and feniur of leg [V.

LEE—RHODACARIDAE 177

Figs. 365-372. Sessiluncus heterotarsus (Canestrini).

365-368 and 372. female: 365, soma, dorsum; 366, leg 1V (part), dorsum; 367.

pretarsus | and tarsus L (part): 368. idiosoma, venter; 372. gnathosoma, venter.

369-371, male: 369, idiosoma, venter; 370, leg IT (part); 371, chelicera,

178 REC, S. AUST, MUS., 16 (3): 1-219 July, 1970

DISTRIBUTION, Em: Pe, Pm: Oi, Om: Am, Ap. From the records

of nominal species the distribution of this genus appears to lie around a line

running south eastwards from Karg’s (1964) record from Hungary (Pe)

to Vitzthum’s (1935) record from the Society Islands (Ap), Unnamed

species from Nepal, Seychelles, Malaya, Sumatra, Borneo and the Solomon

Islands are to be dep. BM(NH), while Bhattacharyya (1965 and manu-

script) has recorded §. vculatus and four unnamed species of Sessifuncus

from West Bengal, India.

Found amongst plant litter and upper soil layers. One species from

cave in the Balkans (Pm),

REMARKS. Sessiluncus is perhaps most like the two other genera,

Gamasellopsis and Gamasitus, that have a reduced leg setation compared to

Gamasellus, but it is also similar to Onchogamasus, The following 4 nominal

species are included in this genus: S$, heterotarsus (Canestrim, 1897); S.

eculatus Vitzthum, 1935; 8S. cavensis Willmann, 1940; 8. iungaricus Karg,

1964, Berlese’s S. eremita, S. solitarius and S, latus are not rhodacarids.

Four unnamed species are described by Bhattacharyya (manuscript).

Sessiluncus heterotarsus (Canestrini)

Gamuasus heterotarsus Canestrini, 1897, p. 473.

FEMALE. Fig. 365-368 and 372. Idiosomal length, 620. Some

of the longer dorsal setae are very slightly pilose, although not figured as

such. Leg chaetotaxy differs from Gamusellus on the following leg segments;

genu IIT, (2, 4/1, 1); genu TV, (2, 5/1, 1): tibia [V, (2, 4/2, 1).

MALE, Fig. 369-371, Idiosomal length, 620,.

LOCALITY, Female (1956.1.22.1) and male (1956.1.22.2) drawn:

Indonesia; Bogor, 10-12.1.1954, col. A. H. G. Alston, dep. BM(NH).

Male (37/28 and 37/29) examined: Java, dep. SEAF.

Sessiuncus hungaricus Karg

Sessiluncus hungaricus Karg, 1964, p. 73.

MALE. = Spermadactyl recurved. Leg chaetotaxy differs from

Gamasellus on the following leg segments: genu IMT, (2, 4/1, 1); genu TV,

(2,.5/1, 1).

LOCALITY, Two males (N196873 and N196874) (M, Costa deter-

mined associated females as being this species or very similar) examined:

Israel, forest litter (Wadi), Mishmar Haemek, 24.4.1967, col, M. Costa,

dep. SAM.

Genus ANTENNOLAELAPS Womersley

Antennolaclaps Womersley, 1956b, p, 112. Type-species: Antennolaelaps

affinis Womersley, 1956b, by original designation.

Stylogamasus Womersley, 1956b, p, 113, symm. Type-species: Stylo-

gamasus convexa Womersley, 1956b, by original designation,

LEE—RHODACARIDAE 179

DLAGNOSIS. Small or average sized mites. Holonotal shield, Ventro-

anal shield separate from notal shield, but closely abuts on exopodal IV

shield or on some males fused to this shield. Peritrematal shield limited in

extent and not fused to exopodal shields. Sterno-metasternal shield fused to

endopodal IV shield and a line joining sternal setae sf2, sf3 and sf4 would

enclose an angle of less than 95°. Spermadactyl fused to base of male

movable cheliceral digit, and if strongly recurved then only distally near tip.

Two ventral setae on genua HI and IV. Pretarsus | pedunculate or absent,

MORPHOLOGY,

SCLEROTIZATION. Female: Holonotal, Discrete ventro-anal which

closely abuts on genital and exopodal IV shield. Small discrete shield

bearing only aciculae lies posterior to anus. Peritrematal free posteriorly

and limited in extent, not reaching back to level of mid acetabulum IV,

All exopodals fused into continuous strip. Sterno-metasternal fused to

endopodals If, If and IV and sometimes to pre-endopodals. which are a

single pair if separate.

Male: Sternito-genital and exopodal TV may be fused or not fused to

ventro-wnal, Otherwise as female.

CHAETOTAXY. Idiosoma: 6), 6z, 5s, 5r: SJ, 5Z. 4-58, 1-SR: Syt:

3Jy, 3Zy, O-28y.

Legs: As Gamasellus.

OTHER CHARACTERS, Female: Deutosternal denticles in 6 to 9 hori-

zontal rows. Corniculus has paraxial flange. Tectum anterior margin

is either trispinate or there may be spinules on lateral spines and central

spine may be trident-like. Movable cheliceral digit with 3 teeth. On palp

genu, seta a/l pectinate with 8 lateral prongs and seta a/2 spatulate, On

palp femur, seta a/ in proximal third, Dorsal setae tapering may be simple

or pilose. Shields may be pitted or reticulated and usually there is a strongly

raised central ridge and peripheral rim on sternum, also shields may be either

shiny or dull, being covered by an adhesive exudate. Peritreme with Jateral

pockets. On sternum, a line joining setae sf2, sf3 and st4 would enclose

an angle of less than 95°. Spermathecal ringed tube opens near posterior

paraxial edge of acetabulum IV. Pretarsus | pedunculate or absent. Leg

setae may have strong spines and av on femur II und one species is a spine

set on a tubercle. Lateral lobes of pulvilli I-IV attenuated and shorter than

central circular lobes. Amongst dorsal setae on tarsus TV, setae ad2, ad3.

pd3 and ad4 may be subequal in length, or seta ad2 or seta pd3 may be

the longest and these seta may be simple or pilose.

TRO REC, S, AUST, MUS., 16 (3): 1-219 July, 1970

373, 375

377, 379

374, 376 381

378, 360

2 382 tt ‘ }

Figs. 373-382.

373-378, Antennolaelaps affinis Womersley, male: 373, soma, dorsum; 374. tarsus 1,

distal tip; 375, idiosoma, venter; 376, tarsus IV, dorsum; 377, leg IL (part), antero-

latus; 37%, gnathosoma, venter, 379 and 380, Gamasitus obscurus Womersley.

female; 379, stigma and surroundings; 380, seta pv! on femur Il and setae afl and

a2 on palp genu. 381 and 382, Onchogamasusy communis Womersley, female:

381. stigma and surroundings; 382. corniculus and setae all and a/2 on palp genu,

LEE—RHODACARIDAE

383, 384

385, 386

Figs. 383-390. Antennolaclaps canyvexuy Womersley.

383-385. 388 and 390. female: 383, soma, dorsum; 384, leg IV (part), dorsum:

385, idiosoma, venter; 388, femur Il, antero-latus; 390, gnathosoma, venter. 386,

387 and 389, mule: 386, idiosoma, venter; 387, leg Il (part); 389, chelicera,

182 REC, 8. AUST, MUS., 16 (3): 1-219 July, 1970

395, 397

Figs. 391-398. Antennelaelaps festiido sp.n..

391-393, 396 and 398, female: 391, soma, dorsum; 392, leg LV (part), dorsum;

393, idiosoma, venter, 396, pretarsus I and tarsus [ (part); 398, gnathosoma., venter.

394. 395 and 397. male: 394, idiosoma, venter; 395. lee I] (part); 397, chelicera,

LEE—RHODACARIDAE 183

Male: Corniculus with paraxial flange or on A. affinis (female not

known) it is attenuated and without flange. Movable cheliceral digit with

| tooth, fused at base to longer spermadactyl which may be hook-like at tip.

On leg II femur seta av, and sometimes pvl, and genu seta av clearly enlarged

into spurs. Tibia seta av may be a spine but not so stout as some of the

dorsal setae on that segment.

DISTRIBUTION. Aa, An. Besides the records of nominal species

there are two females of a possibly unnamed species from near Auckland,

New Zealand, and specimens from Lord Howe Island and New Caledonia,

lo be dep. SAM.

Found in moss and plant litter.

REMARKS. Antennolaelaps is a distinctive genus, rather different

from its most closely allied genera. Although the type-species is unusual

compared to the species that was previously placed in Stvlogamasus and to

the new species, | have preferred to group them all in one genus, because

of the similarities in the structure of the peritreme and the ventral sclerotiza-

lion. The following three nominal species are included in this genus:

A. affinis Womersley, 1956; A. convexus (Womersley) comb.n. for Stylo-

gamasus convexa Womersley, 1956b: A. testudo sp.n.

Antennolaelaps affinis Womersley

Antennolaelaps affinis Womersley, 1956b, p. 112.

FEMALE. Not known.

MALE. Fig. 373-378. Idiosomal length, 470».

LOCALITY. Male (N1968193) drawn: Australia; LF172, moss and

plant litter, Brookfield, near Brisbane, Queensland, 7.9.1966, col. D. C. Lee,

dep. SAM.

The holotype male (N1968194) and paratype male (N1968195)

examined: Australia: litter, Brookfield, near Brisbane, Queensland, 31.5-

10.6.1949, col. BE. H. Derrick, dep. SAM.

Antennolaelaps convexus (Womersiey) comb,n.

Stylogamasus convexa Womersley, 1956b, p. 113.

FEMALE. Fig. 383-385, 388, 390. Idiosomal length, 5504. There

is a conspicuous adhesive exudate on the idiosoma and legs of this species.

Also the legs bear reticulations dorsally.

MALE. Fig. 386, 387, 389, Idiosomal length 540,.

184 REC. S. AUST, MUS., 16 (3): 1-219 July, 1970

LOCALITY, One female (N1968196) and two males (N1968197

and N1968198) drawn or examined: Australia; LFI71, moss. Brookfield,

near Brisbane, Queensland, 7.9.1966, col. D. C, Lee, dep. SAM.

The holotype female (1968199) and allotype male (1968200)

examined; Australia; soil litter, Brookfield, near Brisbane, Queensland,

31.5-1.6.1949, col, E. H. Derrick, dep. SAM.

Antennolaelaps testudo span.

Sessiluncus heterotarsus:! Domrow, 1957, p., 204, syn.

FEMALE, Fig. 391-393, 396, 398. Idiosomal length, 550».

MALE, Fig, 394, 395, 397. Idiosomal length, 450». Movable

cheliceral digit not longer than fixed digit, but has been drawn (Fig. 397)

pushed forward by clearing process.

LOCALITY. The holotype female (1968201), allotype male

(N1968202), 6 paratype females (N1968203-N1968208) and 3 paratype

males (N1968209-N1968211) drawn or examined: Australia; LFI71,

moss, Brookheld, near Brisbane, Queensland, 7,.9.1966, col. D. C. Lee,

dep, SAM.

REMARKS, Note that the new synonymy above is only based on a

comparison of this material with Domrow’s (1957) drawings, but I have

little doubt that the specimens are conspecific and no doubt that they are

congeneric.

Genus GAMASELLEVANS Loots and Ryke

Gamasellevans Loots and Ryke, 1967b, p. 212. Type-species: Gama-

sellevans epigynialis Loots and Ryke, 1967b, by original designation.

DIAGNOSIS. Minute or small mites. Holonotal shield (but there

is a complete or broken demarcation line between podonotal and opisthonotal

parts). Ventro-anal shield separate from notal shield, and may be cither

separate from all other shields or narrowly fused to a posterior extension olf

exopodal IV shield (that may in fact be cither homologous with the meta-

podal shield or the posterior extension of the peritrematal on, for example,

Gamasellopsis, since it bears a pore) or broadly fused to exopodal TV shield.

Peritrematal shield reduced and free posteriorly, Sterno-metasternal shield

fused to endopodal IV shield and a line joining sternal setae sf2, sf3 and st4

would enclose an angle of more than 110°, Spermadactyl fused to male

movable cheliceral digit on distal third, where it points away from the tip

of digit as an attenuated, curving process. The arthrodial process on the

male cheliceral enlarged and spoon-like, being subequal or longer than

movable digit, Two ventral setae on genua Hl and TV. Pretarsus I not

pedunculate, sheath being only long enough to contain retracted claw

coniplex.

LFE—RHODACARIDAL 185

MORPHOLOGY

SCLEROTIZATION, Female: Holonotal (there is complete or broken

line across middle of shield between podonotal and opisthonotal parts,

but holonotal does not break along it), Ventro-anal may be either discrete

or narrowly fused to a posterior extension of exopodal TV (since this

extension bears a pore, it may in fact be homologous with the metapodal

or part of the peritrematal) or broadly fused to exopodal IV. Peritrematal

reduced and {ree posteriorly, Exopodals Il and Il may or may not be split.

Sterno-metasternal fused to endopedals I, Wl and IV, Metasternal region

may be partially separated from sternal region by split from posterior margin

lo near seta st4. Single pair of pre-endopodals.

Male: Sternito-genital not fused to ventro-anal, Ventro-anal fused or

not fused to exopodal IV as in female.

CHAETOTAXY, Tdiosoma; 6j, 6z, Ss, Sr: 5/4, 57, 5S, OR: Sst: Sly,

3Zy, 2S8y.

Legs: As Gamasellus.

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows, Tectum anterior margin usually basically unispinate with lateral

spinules but may be trispinate, with spinules on lateral spines. Movable

cheliceral digit with 3 teeth. On palp genu, chaetomorphy unknown, On

palp femur, seta a/ in central third. Dorsal setae simple and tapering.

Peritreme without conspicuous lateral pockets. On sternum, a line joining

setae sf2. sf3 and st4 would enclose an angle of more than 110°. Anterior

hyaline flap on genital shield extends forward, with point reaching anterior

to seta sf3 and sometimes nearly reaching seta st], Spermathecal ringed

tube opening not known. Pretarsus | not pedunculate, sheath only long

enough to contain retracted claw-complex. Often small processes on

femur WH,

Male: Corniculus attenuated and longer than on female. Movable

cheliceral digit with | tooth or no teeth, and fused for most of its length with

long curving spermadactyl. Dorsal process distally on fixed cheliceral digit.

Arthrodial membrane long and spoon-like (this may be homologous with the

extra, fimbriated flap in Queenslandolaelaps vitzthumi). On femur, genu and

tibia of lee Tf there are spurs which are probably modified antero-ventral

setae.

DISTRIBUTION, Es, All nominal species recorded from the

Transvaal, South Africa.

Found in pasture soil and evergreen forest litter,

REMARKS. The long anterior hyaline flap on the genital shield and

the reduced peritrematal shield in all species, as well as the extensive fusion

of the exopodal IV shield to the ventro-anal shield in some species would

186 REC, §, AUST. MUS, 16 (3): 1219 July, (970

seem to make Gamasellevans quite distinctive, but two of these three

attributes do not occur in the closely alhed Paragamasellevans, while the

unusual male chelicera is very similar to that of Paragamasellevans and

Queenslandolaelaps. Therefore Gamasellevans may be more closely allied

to Queenslandolaelaps than the characters of the idiosomal venter would

suggest. The following 7 nominal species are included in the genus;

G, epigynialis Loots and Ryke, 1967b; G, bispermadactylus Loots and Ryke,

1967b; G. evansi Loots and Ryke, 1967b; G. magoebaensis Loots and Ryke,

1967b:; G. reticulatus Loots and Ryke, 1967b; G. spermadactylus Loots and

Ryke, 1967b; G, vandenbergi Loots and Ryke, 1967b,

Genus GAMASELLOPSIS Loots und Ryke

Gamasellapsis Loots and Ryke, 1966a, p. 551, Type-species: Gamasellopsis

curtipilus Loots and Ryke, 1966a, by original designation.

DIAGNOSIS, Minute or small mites. Holonotal shield. Ventro-anal

shield discrete, being separate from notal shield, although it may closely abut

on exopodal IV shield. Peritrematal fused to exopodal IV shield. Sterno-

metasternal shield may or may not be fused to endopodal IV shield and a

line joining seta s/2, 9f3 and st4 would enclose an angle of more than 110°,

The spermadactyl fused to the male movable cheliceral digit for at least

two-thirds of its length, and is longer than the digit but similarly shaped.

One ventral seta on genua III and [V¥. Pretarsus I pedunculate.

MORPHOLOGY.

SCLEROTIZATION, Female: Holonotal. Discrete ventro-anal bearing

aciculae posterior to anus, Peritrematal fused to exopodal TV, No split in

exopodal III or [V,. Sterno-metasternal either fused to endopodal TV or to

single pair of pre-endopodals but not to both.

Male; Sternito-genital not fused to ventro-anal.

CHAETOTAXY. Idiosoma: 6/, 4 or 5z, 5s, 4r: 5/, 5Z, 5S, | or 2R:

Sst; 3/y, 3Zv, 1Sy.

Legs; There are 6 or 7 setuc less than in Gamasellus, the following

segments being different: genu I (2, 6/2, 2); genu III (2, 4/1, 1); genu IV

(2, 4 or 5/1, 1); tibia TV (2, 4/2, 1); tarsus IV (3, 6/5, 3). The seta

missing on tarsus TV is pd4.

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows, Tectum anterior margin quinquispinate. Movable cheliceral digit

with 3 teeth. On palp genu, seta all pilose with over 10 short, lateral

prongs and seta q@/2 simple, tapering. On palp femur, a/ in central third.

Dorsal setae simple, tapering. Peritreme without conspicuous lateral pockets.

L.EE—RHODACARIDAE 187

On sternum, a line joining setae sf2, s¢3 and st4 would enclose an angle of

more than 110°. Tibia I is swollen ventrally and seta pyl may be on a

small protuberance. Spermathecal ringed tube opening not known, — Pre-

tarsus I pedunculate and smaller than other pretarsi.

Male: Movable digit with 1 tooth, may be considerably shorter than

fixed digit, fused along two-thirds of its length to spermadactyl which is at

least half as long again, On leg IL, femur seta av enlatged into spur, may

he other protuberances on femur and genu, but their homologies are

unknown.

DISTRIBUTION. fs. All nominal species recorded [rom Transvaal,

South Africa.

Found in evergreen forest litter.

REMARKS. Members of Gamasellopsis are very similar to. the

Sessiluncus species of the Palaearctic region although there are a number of

conspicuous differences such as the spermadactyl shape. the fusion of the

peritrematal shield to the exopodal shield IV, the pedunculate pretarsus I

and the leg chaetotaxy. There are also similarities to Gamuasitus and

Onchogamasus, but the males of those genera are not yet known. The

following 4 nominal species are included in this genus: G. curtipilus Loots

and Ryke, 1966a; G. longipilus Loots and Ryke, 1966a,; G. magoebaensis

Loots and Ryke, 1966a; G. vandenbergi Loots and Ryke, 1966a.

Genus GAMASITUS Womersicy

Gamasitus Womersley, 1956a, p. 531. Type-species: Gamasitus obscurus

Womersley, 1956a, p. 531.

DIAGNOSIS. Large mite. A single female is the only known member

of this genus. Holonotal shield. Ventro-anal shield separate from notal

shield, but closely abuts on exopodal IV shield. Peritrematal shield not

fused to exopodal [V shield and extends as far posteriorly as that shield.

Sterno-metasternal shield fused to endopodal IV shicld and pre-endopodal

shields, and a line joining sternal setae s72, st3 and st4 would enclose an

angle of more than 110°, One ventral seta on genua HI] and IV. Pretursus

I pedunculate and subequal in size to other pretarsi,

MORPHOLOGY.

ScLEROTIZATION. Female: Holonotal, Discrete ventro-anal, Small

discrete shield bearing only aciculae lies posterior to anus. Peritrematal

lree posteriorly and reaching back to level of posterior edge of acetabulum

IV. Exopodals If and IM split, while TV is not split. Sterno-metasternal

fused to endopodal II, I and IV, and to pre-endopodals.

188 REC, S. AUST, MUS. 16 (3); 1-219 daly, L970

CHAETOTAXY. Idiosoma: Notum too fragmented to recognize chaeto-

laxy: Sst: 3/¥, 2Z2y.

Legs: Following segments differ from Gamasellus in having a ventral

seta missing: genu IIT (2, 4/1, 1); genu IV (2, 5/1, 1).

OTHER CHARACTERS. Female; Movable cheliceral digit with 4 tecth.

Tectum anterior margin trispinate, the lateral spines being quite small. On

palp genu, seta a/1 with 7 lateral prongs and seta a/2 slightly spatulate. On

palp femur, seta a/ in proximal third. Dorsal setae simple. tapering. Shields

shiny. Peritreme without conspicuous lateral pockets. On sternum, a line

joining setae sf2, st3 and st4 would enclose an angle of more than 110°.

Spermathecal ringed tube opening not known. Pretarsus I pedunculate and

subequal in size to other pretarsi. Femur IT with seta pyl spine-like and on

process. Lateral lobes of pulvilli I-IV attenuated and longer than central,

circular lobes. Amongst dorsal setae on tarsus IV setae pd3 and pd4 are

the longest and subequal in length.

DISTRIBUTION, Aa. The only record is of the single female from

Tasmania, Australia.

Found in moss,

REMARKS. Male Gamasitus are unknown and only one incomplete

and fragmented female is known. As observed by Womersley when he

established this genus, it is similar to Sessiluncus. The following single

nominal species is included in this genus: G, ebsewrus Womersley, 1956a.

Gamasitus obscurus Womersley, 1956a

Gamasitus obscurus Womersley, 1956a, p. 531.

FEMALE. Fig. 379, 380. Idiosomal length, 940),

MALE. Not known,

LOCALITY. The holotype female (N1968159) drawn and examined:

Australia; moss, Mt. Wellington, Tasmania, 2.12.1934, col. V. V. Hickmann,

dep. SAM.

Genus ONCHOGAMASUS Womersley

Onchogamasus Womersley, 1956b, p. 108. Type-species: Onchogamasus

communis Womersley, 1956b, by original designation.

DIAGNOSIS. Minute or small mites, only known from the females.

Holonotal shield. Ventro-anal shield separate from the notal shield, but

fused to exopodal IV shield. Peritrematal shield may or may not be fused

to exopodal IV shield and a line joining sf2, st3 or st4 would or would not

enclose an angle of less than 95°, Two ventral setae on genua HI and IV,

Pretarsus I pedunculate and subequal in size to other pretarsi.

LEE—RHODACARIDAE 189

REMARKS. Onchogamasus was originally established for a single

female specimen, similar to members of Gamasellopsis and Gamasitus, and

also to the Palaearctic members of Sessiluncus, but, unlike those three genera,

its leg chaetotaxy is as for Gamasellus. Unfortunately no males of the two

Australian genera (Gamasitus and Onchogamasus) are known and therefore

the relationships within this group of genera are unusually uncertain, When

trying to place two species, pumilio and quasicurtipilus, in a genus, | decided

to put them in Onchogamasus mainly because of their leg chaetotaxy, but as

a separate species-complex because of their characters which are different to

those of Onchogamasus and similar to those of other genera. Therefore, |

recognize the following two species-complexes within this genus, communis-

complex and pumilio-complex, which can be distinguished by the following

key. The morphology and distribution of members of this genus are given

under the species-complex headings.

KEY TO SPECIES COMPLEX OF ONCHOGAMASUS

|. Peritrematal shield is fused to exopodal shields,

which are unsplit. Line joining setae s?f2,

st3 and st4 would enclose an angle of less

than 95° 2. 0 oe ee ee ee ee ee © COMMUNis-COMplex

Peritrematal shield is not fused to exopodal

shields, of which Hl and II are split. Line

joining setae sf2, st3 and st4 would enclose

an angle of more than 110° .. .. .. 2. pumilio-complex

COMMUNIS-complex

DIAGNOSIS. Limited within generic diagnosis to small mites, with

peritrematal shield fused with all exopodal shields into a single shield, and a

line joining sternal setae sf2, st3 and st4 would enclose an angle of less

than 95°.

MORPHOLOGY.

SCLEROTIZATION. Female: Holonotal. Ventro-anal separate from

notal but fused to exopodal IV. Aciculae are on small, discrete shield

posterior to anus. Peritrematal fused to all exopodals forming a single

shield. Sterno-metasternal fused to endopodal Il, HI and IV. One pair of

pre-endopodals.

CHAETOTAXY. Idiosoma: Notum too fragmented to recognize chaeto-

taxy: Sgr: 3/v, 3Zv, LSv,

Legs: As Gamasellus.

190 REC, S$, AUST, MUS., 16 (3): 1-219 July, 1970

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows. Tectum anterior margin is unispinate. Movable cheliceral digit

with 3 teeth. On palp genu, seta all pectinate with 7 lateral prongs

and seta al2 slightly spatulate. On palp femur, seta a/ in proximal third.

Dorsal seta tapering, may be simple or slightly pilose. Shields slightly shiny.

Exopodal process between coxae Il and III has dorsal flap. Peritreme

without lateral pockets, On sternum a line joining seta sf2, st3 and st4

would enclose an angle of less than 95°. Spermathecal ringed tube opening

not known. Pretarsus | pedunculate and subequal in size to other pretarsi.

Lateral lobes of pulvilli II-LV attenuated and longer than central oval lobes.

Amongst dorsal setae on tarsus TV, seta pd3 is the longest and slightly pilose.

DISTRIBUTION. Aa. The only record is of the single female from

Queensland, Australia.

Found amongst plant debris and soil.

REMARKS. Males of the communis-complex are unknown, Only

one squashed and fragmented female is known. This female has similarities

to Sessiluncus, Gamasellopsis and Gamasitus. The following single nominal

species is included in this species-complex; O, communis Womersley, 1956b.

Onchogamasus communis Womersley

Onchogaumasus communis Womersley, 1956b, p, 108.

FEMALE. Fig. 381. 382. Idiosomal length, 560,. Idiosoma

covered in reticulations which often consists of wavy lines because of puncta-

tions beside them. Tarsus II bears conspicuous spines, one is seta av2, while

the two at the distal tip are non-setous. Some leg setae are on slight

protruberances, seta ad2 on genu II is on a conspicuous protruberance,

MALE. Not known.

LOCALITY. The holotype female (N1968185) drawn and examined:

Australia; soil and plant debris, Brookfield, near Brisbane, Queensland,

21.5.1949-2.6.1949, col. E. H. Derrick, dep. SAM.

PUMILIO-complex

DIAGNOSIS, Limited within generic diagnosis to minute mites, with

peritrematal shield separate from all exopodals, and a line joining sternal

setae sf2, st3 and st4 would enclose an angle of more than 110°.

MORPHOLOGY.

SCLEROTIZATION. Female: Holonotal. Ventro-anal separate from

notal but fused to exopodal IV. Aciculae on small discrete shield posterior

to anus, Peritrematal not fused to exopodals or any other shields posteriorly.

LEE—RHODACARIDAE 191

Exopodals I] and IIT are split. Sterno-metasternal fused to endopodal II,

HW! and TV. One pair of pre-endopodals which may be fused to sternal.

CHAETOTAXY. Idiosoma: 6), 6z, 58, 5r: SJ, 5Z, SS, SR, 2-3UR:

5st; 3/v, 3Zy, ISv.

Legs: As Gamasellus.

Orner CHARACTERS. Female: Deutosternal denticles in 6 hori-

zontal rows. Tectum anterior margin basically unispinate with lateral

spinules. Movable cheliceral digit with 2 or 3 teeth. On palp genu, seta all]

pectinate with 7 lateral prongs and seta a/2 slightly spatulate. On palp

femur, seta al in proximal third. Dorsal setae simple and tapering. Shields

pale. Peritreme without lateral pockets and does not extend anterior to

mid acetabulum II. On sternum a line joining setae sf2, st3 and s?4 would

enclose an angle of more than 110°. Spermathecal ringed tube opening not

known. Pretarsus I pedunculate and subequal in size to other pretarst.

Tibia I is dilated distally in area of ventral setae. Lateral lobes of pulvilli

II-!V attenuated and shorter than central oval lobes. Amongst dorsal setae

on tarsus IV, setae ad2, ad3 and pd3 subequal in length.

DISTRIBUTION. Aa. As recorded below the nominal species are

from South Australia and Victoria, Australia.

Found in moss and plant litter.

REMARKS, Males of the pumilio-complex are unknown. The females

are similar to those of the Onchogamasus communis-complex, but there are

enough differences, including characters which are similar to those of

Gamasitus, for me to consider it worth while to keep them in a separate

complex until more is known about these mites. The following two

nominal species are included in this species-complex: O. pumilio sp.n.:

QO. quasicurtipilus spn.

Onchogamasus pumilio spn.

FEMALE. Fig. 399-404. Idiosomal length, 320». The chelicera

(Fig. 402) is drawn from a postero-lateral angle, while viewed laterally it

would probably appear similar in shape to the chelicera (Fig. 408) of

O. quasicurtipilus. Seta Zv3 of the right side is on ventro-anal shield.

MALE. Not known.

LOCALITY. The holonotal female (N1968186) drawn: Australia;

LF97 moss beneath White Mallee scrub, overlooking Formby Bay, Yorke

Peninsula, South Australia, 3.11.1965, col. N. McFarland and N. B. Tindale,

dep. SAM.

192 REC, 8S. AUST. MUS., 16 (3): 1-219 July, 1970

sr \ oad

*o —_4

So”

oO—oO —9-0.

re)

“

o-?

o—_9g

fe)

=o 9-0-0

)

™o

pO—— 9g

-O

rae

406, 407 |

410 oO

399, 40)

403

kes

~— se

402 ~~

408

o—o——p9 |

SY ; se

404 409 in ha

Figs. 399-410, Onchogarmnasus Womersley,

399-404, puniilio sp.n.. female: 399, soma, dorsum: 400, pretarsus TL and tarsus 11

(part); 401, leg IV (part). dorsum; 402, gnathosoma, venter: 403, idiasoma, venter:

404, genu 1, ventral setae only. 405-410, quasicurtipilus sp.n.. female: 405, pretarsus

Uo und tarsus IT (part); 406, soma, dorsum; 407, lez IVY (part), dorsum; 408,

gnathosoma, yenter; 409, genu IL ventral setae only; 410, idiosoma, venter,

LEE—RHODACARIDAE 193

Onchogamasus quasicurtipilus spm.

FEMALE. Fig. 405-410. Idiosomal length, 430». Instead of the

three non-setous spurs at distal tip of tarsus Il being equally increased in

size, as on O. pumilio, only the central one is enlarged (Fig, 405).

MALE Not known.

LOCALITY. The holotype female (N1968187) and paratype female

(N1968188) drawn and examined: Australia; LF79, moss and litter beneath

tree ferns and Eucalyptus, Otway Ranges, Victoria, 28.8.1965, col. F. J.

Mitchell, dep, SAM.

Genus PARAGAMASELLEVANS Loots and Ryke

Paragamasellevans Loots and Ryke, 1968, p. 3. Type-species: Paragama-

sellevans michael; Loots and Ryke, 1968, by original designation.

DIAGNOSIS. Minute mites. Separate podonotal and opisthonotal

shields. Ventro-anal shield widely separated from all other shields in both

sexes, but broader on male where it may include metapodal shields.

Peritrematal shields reduced to slim strip between anterior end of peritreme

and podonotal shield and therefore not fused to exopodal IV shield. Sterno-

metasternal shield not fused to endopodal IV shield and a line joining sternal

setae yt2, sf3 and st4 would enclose an angle of more than 110°. Sperma-

dactyl fused to male movable cheliceral digit on distal third, where it points

away from the tip of digit as an extremely long, attenuated process. The

arthrodial process on the male chelicera similar to that on female. Two

ventral setae on genua HIT and IV. Pretarsus I not pedunculate.

MORPHOLOGY,

SCLEROTIZATION. Female: Podonotal and opisthonotal with facing

edges closely apposed. Discrete ventro-anal bearing aciculae posterior to

nus. Discrete metapodal. Peritrematal reduced to slim strip between

anterior end of peritreme and podonotal. Exopodals do not completely

encase peraxial edges of acetabula. Sterno-metasternal fused to endopodal

IT and part of IIL. Single pair of pre-endopodals.

Male: Sternito-genital not fused to endopodal [V and not broad enough

posteriorly to bear seta sf5,

CHAETOTAXY. Idiosoma: 6/, Sz, 5s, Sr: SJ, 57, 58, 3 or SR: Sgt:

3Jyv, 3Zy, 1Syv,

Legs: As Gamasellus.

194 REC, S. AUST. MUS., 16 (3): 1-219 July, 1970

Sry ae 416

¥ 4u

F 418 aia

4|3

4l4

415

aiz

Ps;

ae +3 415

Figs. 411-418. Stylochirus G. and R. Canestrini.

411 and 412, S. haemisphaericus (Koch), female: 411, idiosoma, venter; 412,

gnathosoma, venter. 413-415, $8, rarior (Berlese), female: 413, tectum; 414,

gnathosoma, venter; 415, idiosoma. yenter. 416-418, S. ampulliger Berlese, male:

416, tarsus 1. distal tip: 417, femur II, venter; 418, idiosoma, venter.

LEE—RHODACARIDAE 195

OTHER CHARACTERS. Female: Deutosternal denticles in 7 horizontal

rows, Tectum anterior margin is basically unispinate with lateral spinules.

Movable cheliceral digit with 3 teeth, Shape and position of antero-

lateral setae on palp femur and genu unknown. Dorsal setae tapering,

simple or slightly pilose. Peritreme without lateral pockets. On sternum,

a line joining setae sf2, st3 and st4 would enclose an angle of more than

110°, What is probably a long, conspicuous spermathecal ringed tube opens

near posterior paraxial edge of acetabulum TV. Pretarsus I not pedunculate.

Male: ‘Tectum anterior margin is without lateral spinules as in female.

Movable cheliceral digit with | tooth, fused to extremely long spermadactyl

along its entire length. When at rest, long spermadactyl loops back into

pouch lying immediately dorsal to sternum, and reaching back to just

posterior to sternal seta st4. Corniculi are longer than on female. On leg

IT, seta av on femur, genu and tibia is enlarged into spur.

DISTRIBUTION. Es. All nominal species recorded from Transvaal,

South Africa.

Found in forest soil.

REMARKS. Paragamasellevans is closely allied to Gamasellevans,

but with a morphology that is better adapted for life in the lower soil layers.

The following two nominal species are included in this genus: P. michaeli

Loots und Ryke, 1968; P, vandenbergi Loots and Ryke, 1968,

Genus QUEENSLANDOLAELAPS Womersley

Queenslandolaclaps Womersley, 1956b, p. 109. Type-species: Queenslando-

laelaps vitzthumi Womersley, 1956b, by original designation.

DIAGNOSIS. Small or average sized mites. Holonotal shield. Ventro-

anal shield separate from notal shield and fused to exopodal IV shield on

both sexes. Peritrematal shield fused to exopodal IV shield. Sterno-

metasternal shield fused to endopodal IV shield and line joining sternal setae

sf2, st3 and sf4 would enclose an angle of less than 95°. Spermadacty|

fused to male movable cheliceral digit on distal third, where it points away

from the tip of digit as an attenuated, curving process with lateral hyaline

flaps. Hyaline process with fimbriated border between arthrodial process

and movable cheliceral digit male. Two ventral setae on genua IIT and TY.

Pretarsus [ pedunculate and smaller than other tarsi.

MORPHOLOGY,

SCLEROTIZATION, Female: Holonotal. Ventro-anal fused only to

exopodal IV. Aciculac on small discrete shield posterior to anus. No

196 REC. S. AUST, MUS... 16 (3)2 1-219 July, 1970

separate metapodal, Peritrematal fused to exopodal IV, Exopodal II and

HL not split. Sterno-metasternal fused to endopodal If, II and TV. Single

pair of pre-endopodals.

Male: Separate sternito-genital,

CHAETOTAXY, Idiosoma: 6/, 62, 5s, 5r: SJ, 5Z, 5S, SR: Sst: 3Jv,

3Zy, 28v.

Legs: As Gamasellus.

OTHER CHARACTERS. Female: Deutosternal denticles in 8 horizontal

rows. Tectum anterior margin trispinate with central process narrow at

base, spreading out to broader fimbriated tip. Movable cheliceral digit

with three teeth. On palp genu, seta a/l pectinate with 7 lateral prongs and

seta al2 slightly lanceolate. On palp femur, seta al in proximal third,

Dorsal setae tapering, simple or slightly pilose. Exopodal process between

coxae IIT and III has dorsal flap. Peritreme without lateral pockets. On

sternum, a line joining setae sf2, st3 and st4 would enclose an angle of less

than 95", Spermathecal ringed tube opens near posterior paraxial edge of

acetabulum IV, and its junction with a single central sacculus is clear.

Pretarsus I pedunculate and smaller than other pretarsi, Lateral lobes of

pulvilli I-IV attenuated and shorter than central circular lobes. Amongst

dorsal setae on tarsus IV, set pd3 is the longest and setose.

Male: Movable cheliceral digit with | tooth, and fused for much of

its length with a long, sinuous spermadactyl with lateral flaps. Extra,

fimbriated flap between arthrodial process and movable cheliceral digit, and

there is a dorsal process distally on fixed cheliceral digit. On leg I, femur

setae av and pvl, genus setae av and pv, tibia seta ay, modified into spurs

or spines. On tarsus II, seta av2 is slightly enlarged into a spine.

DISTRIBUTION, Aa. The only records are from Queensland,

Australia.

Found in plant debris and soil.

REMARKS. Although there are differences between the ventral

sclerotization of the idiosoma of the single species in Queenslandolaelaps

and Gamasellevans, the similarities between the unusual male chelicera

suggest that the two genera may be closely allied. The following single

nominal species is included in this genus: Q, vitzthumi Womersley, 1956b.

419, 421

422, 426

/ .

} j 420, 423

A 424, 425

Figs. 419-426. Queenslandolaelaps vitzthumi Womersley.

419-421, 425. 426, female: 419, soma, dorsum; 420, gnathosoma, venter; 421,

idiosoma, venter; 425, spermatheca; 426, tarsus IV, dorsum, 422-424, male: 422.

idiosoma, venter; 423 (two parts), leg Il: 424. chelicera.

198 REC. S. AUST. MUS., 16 (3); 1-219 July, 1970

Queenslandolaelaps vitzthumi Womersley

Queenslandolaelaps vitzthumi Womersley, 1956b, p, LOY,

FEMALE. Fig. 419-421, 425, 426. Idiosomal length, 630».

MALE. Fig. 422-424. Idiosomal length, 550p.

LOCALITY. Females (N1968189) and male (N1968190) drawn:

Australia; LF200, leaf litter, Brookfield, near Brisbane, Queensland.

10,5.1967, col. J. A. Herridge, dep. SAM.

The holotype female (N1968191) and allotype male (1968192)

examined: Australia: soil debris, Brookfield. near Brisbane, Queensland,

31.5-10.6,1949, col. E, H. Derrick, dep. SAM.

Genus STYLOCHIRUS G. and R, Canestrini

Stylochirus G. and R. Canestrini, 1882, p. 56. Type-species: Stilochirus

rovennensis G. and R, Canestrini, 1882, by monotypy.

Physallolaclaps Berlese, 1908, p. 13, syn.n., Type-species: P/iysallolaelaps

ampulliger Berlese, 1908, by original designation.

Periplis Berlese, 1914, p. 142, sym.n.. Type-species: /phiy haemisphaericus

Koch, 1839, by original designation,

Epiphis Berlese, 1916b, p. 302, syn.n.. Type-species: Gamasiphis (Epiphis)

rarior Berlese, 1916b, by original designation,

Megaliphis Willmann, 1938, p. 161, syn... Type-species: Gamasiphis

(Megaliphis) giganteus Willmann, 1938, by monotypy.

DIAGNOSIS. Average sized to gigantic mites. Holonotal shield.

Ventro-anal shield fused to notal shield on both sexes, Peritrematal extend-

ing well posterior to acetabulum IV, and either free posteriorly or fused to

notal shield. Sterno-metasternal shield not fused to endopodal IV shield

and a line joining sternal setae st2, sf3 and sr4 would enclose an angle ot

more than 110°. Male cheliceral digits reduced and edentate, the movable

digit being fused along its entire length to a much longer, tapering sperma-

dacty!. Two ventral setae on genua If and IV. Pretarsus I is pedunculite.

MORPHOLOGY.

ScLeroTizATION, Female; Holonotal fused to ventro-anal, which is

otherwise discrete but closely abuts on exopodal [V shield. Peritrematal

only fused to holonotal, sometimes along its entire length. Usually no splits

in exopodals but the female of §. minor is drawn with exopodals not

completely encasing the peraxial edges of the acetabula, Metasternal may

be discrete or narrowly jomed to sternal which is only fused to endopodal 11.

Single pair of pre-endopodals.

LEE—RHODACARIDAE 199

Male; Discrete sternito-genital.

CHAETOTAXY. Idiosoma; Not known. other than 5st and opisthosoma

is hypertrichous.

Lees! As Gamasellus.

OVHER CHARACTERS. Female: Globular mites, pointed posteriorly,

wand with a relatively small gnathosoma and = attenuated chelicerae,

the latter being about as long as the distance from anterior edge of

acetabuluin IE to posterior edge of acetabulum [V, Movable cheliceral digit

with 3 oy 4 teeth. Tectum anterior margin with triangular shaped central

process. On palp genu, seta a/l and al2 slightly pilose or lanceolate but not

simple. On palp femur, seta a/ in central third. Dorsal setae simple and

short. Shields extensively reticulated. Peritreme without lateral pockets

and short ool extending anterior to mid-acetabulum I] and may be only as

long as stigma width. On sternum 4 line joining setae sf2, 973 and sr4 would

enclose an angle of more than 110°. Spermathecal ringed tubes not

recognized. Pretarsus I is pedunculate and may be reduced compared to

ollier pretarsi,

Male: The single well described male is that of S. ampulliger for which

the female is not known (unless §. revennensis is conspecific), so it is ditticult

fo conmpare the sexes. The dilated, bulbous seta at the distal end of tarsus |

(Fig. 416) may occur only in the male since this is true for the male of an

unnamed 4frogamasellus species described by Loots (thesis, 1967). The

male cheliveral digits are reduced and edentate, and the movable digit is

fused along its entire length to a stout, tapered spermadacty! which may be

longer than palps, On leg HH. femur seta av and tibia seta av are enlarged

into a spur while all the other setae appear setose, and seta av on the genu

is certainly setose although slightly stouter and blunter than seta py,

DISTRIBUTION. Na: Pe, Pm. Beside the locality records published

with the original descriptions of nominal species. Athias-Henriot (1961a)

records S, ampu/liger trom the Appennino Ligure Mountains, Italy and the

Pyrenees Mountuins, France,

Found in moss and plant litter,

REMARKS. There is little doubt about the close relationship between

all the members of this genus except for one. Unfortunately, the exception

is the type associated with the oldest available generic name for this taxon,

Le, 8. rovennensis. Athias-Henriot (19614) has pointed out the resemblance

between the meagre description of S. rovennensiy and her thorough redescrip-

lion of Physallolaelaps ampulliger, suggesting that they might be the same

species, but not formally recognizing their synonymy. It can be argued that

Berlese should have noticed this resemblance, but the male §$, rovennensis

that Professor Canestrini sent to him was obscure and fragmented (Berlese,

200 REC. S. AUST. MUS. 16 (4): 1-219 Jily, 1970

1892c), | have here formally recognized S, rovennensis and P. ampulliger

as congeneric and therefore Stylochirus is the valid name for this taxon, If

any division into subgenera was used, then the ex-members of Megaliphis,

the two largest species with the peritreme no bigger than the stigma, would

be the best to group into a separate subgenus. This genus is not closely

allied to the other Sessiluncinue genera and is placed here for convenience,

The following 6 nominal species are included in this genus: S. revennensis

G, and R. Canestrini, 1882; §. ampulliger (Berlese) comb.n, for Physallo-

laelaps ampulliger Berlese, 1908; S. giganteus (Willmann) comb.n. for

Mevaliphis viganteus Willmann, 1938; S. haemisphaericus (Koch) comb.n.

for Periphis haemisphaericus (Koch, 1839); 8. miner (Willmann) comb.n,

for Megaliphis minor Willmann, 1953; 8. rarior (Berlese) comb.n, tor

Epiphis rarior Berlese, 1916b.

Stylochirus rovennensis G. and R. Canestrini

Stilochirus rovennensis G. and R, Canestrini, 1882, p. 56.

FEMALE, Not figured. Idiosomal length, 940. Cheliceral fixed

digit is multidenticulate and the movable digit is tridentate, the posterior

tooth being the largest, Tectum anterior margin with single triangular

process, Palps close together, the hypostome being constricted and clearly

separated from the base of the palps. Corniculi small (“absent”), Idiosoma

strongly convex and pointed posteriorly, with smooth surface and short setae

(“unprovided with setae”), Holonotal shield. Ventro-anal shield semi-

circular, Peritreme short. Reddish brown.

MALE. Not figured. Idiosomal length, 8004, Cheliceral digits

reduced and with elongated spermadactyl directed forward past palps. Leg

IT with medium sized process on femur and a small process on tibia.

LOCALITY, — Italy; moss, fields of Tridentini, Non Valley, Mt.

Rovenna, Trentino District, Alps.

REMARKS. This description is culled from G, and R, Canestrini’s

(1882) and Berlese’s (1892c) descriptions of this species,

Stylochirus ampulliger Berlese

Physallolaelaps ampulliger Berlese, 1908, p. 13.

FEMALE. Not known.

MALE. Fig. 416-418. Idiosomal length, 8904. From the single

specimen in Berlese Collection it is difficult to observe any finer detail than

what is drawn here. Athias-Henriot (1961a) gives a detailed description of

conspecific males.

LOCALITY. The ‘tipico’ male (75/9, two slides, one leg | being

mounted separately) drawn: Italy; Vallombrosa, dep. SEAF.

LEE—RHODACARIDAE 20)

Stylochirus haemisphaericus (Koch) comb.n.

Iphis haemisphaericus Koch, 1938, 27.16.

FEMALE. Fig. 411, 412. Idiosomal length, 910. Gnathosoma

bent ventrally, probably obscuring seta hyp4. Part of ventro-anal shield

obscured (shown by hatched shading on Fig. 411).

MALE, Not known.

LOCALITY. Two females (148/23 and 151/35, legs, palps and

gnathosoma fragments) drawn: Italy; moss in pasture, Cansiglio, dep. SEAF.

Stylochirus rarior (Berlese) comb.n.

Gamasiphis (Epiphis) rarior Berlese, 1916b, p. 303.

FEMALE, Fig. 413-415. Idiosomal length, 950». Holonotal and

ventro-anal seta may be hypertrichous. Some dorsal setae on legs as spiny

as sternal seta sf2 and slightly pilose at tip.

MALE. Not known.

LOCALITY. The ‘tipico’ female (175/43) drawn: North America;

leaf mould, Columbia, Missouri, U.S.A., 1904-1906, col. C. R. Crosby, dep.

SEAF (for justification of this data, see Hammen, 1959, p. 24).

Subfamily TANGAROELLINAE subf.n.

Type-genus: Yangaroellus Luxton, 1968.

REMARKS. Since this subfamily contains only a single species and

the main function of this paper is to give diagnosis for genera, the characters

of the species are listed under the genus heading.

Genus TANGAROELLUS Luxton

Tangaroellus Luxton, 1968, p. 497. Type-species: Tangaroellus porosus

Luxton, 1968, by original designation.

DIAGNOSIS. Small mites. Separate podonotal and opisthonotal

shields. Discrete ventro-anal shields on both sexes, although on male there

is no discrete pair of metapodal shields so they may be merged into ventro-

anal shield. Peritrematal shield not fused to exopodal IV shield. Setation

of legs unique within Rhodacaridae in having one ventral seta less than

Gamasellus on femur I, one ventral seta more on femur ITV, and two dorsal

setae less on tarsus TV. Only rhodacarid genus with 2-pronged apotele.

Location of spermathecal ringed tube, if present, unknown. Pretarsus [ not

pedunculate.

2()2 REC, S, AUST, MUS, 16 (3): }-219 July, 170

MORPHOLOGY.

SCLEROTIZATION, Female; Separate podonotal and opisthonotal,

Discrete veniro-anal and metapodual, Peritrematal not fused to any other

shields posteriorly. Exopodials do not completely encuse peraxial edges of

acetabula. Sterno-metasternal fused to endopodal Il. Single pair of

pre-endopodals,

Male: Sternito-genital. Ventro-anal larger and probably including

netapodals. Fusion of peritrematal to podonotal anteriorly is more

extensive,

CHAETOTAXY, Idiosoma: 6/, 42, 38, 3r: 47, 42, 45. OR) Syste 3d v4

2Z\, OSv.

Legs: Differs from Gamasellus in having one less ventral on femur |

(2, 5/3, 2) and one more ventral on femur IV (1. 4/2, 0), and in having

two dorsal setae (setae ad3 and pd4) less on tarsus TY.

OTHER CHARACTERS. Female: Tectum anterior margin unispinate.

Movable cheliceral digit with 3 teeth. On palp genu, setae a/| and w/2 are

simple, setose. On palp femur, seta a/ in proximal third. Apotele 2-pronged

in cUntrast to all other rhodacarid mites which have a 3-pronged apotele.

Most dorsal setae are simple, setose, but some are spatulate, No noticeable

spermathecal ringed tube, Pretarsus I not pedunculate, sheath only being

long enough to contain the retracted claw-complex. Pulvilli IL-TV have two

oval lobes. Amongst dorsal setae on tarsus TV, seta pd3 is the longest and

is spatulate,

Male: Movable cheliceral digit with | tooth and fused at base to longer,

tapered spermadactyl. Seta p/ on palp genu is modified into a spur. On

leg IH, setae av on femur, genu and tibia are spurs and other setae may be

robust spines, larger than the same setae on the female.

DISTRIBUTION, An. Only records are from New Zealand. Found

in littoral zone, usually in the crevices between barnacle carapaces.

REMARKS. Tangaroellus is unusual in having both a 2-pronged

apotele and 4 ventral setae on tibia 1, The chaetotaxy of tibia T is given

more weight and the genus is retained in the Rhodacaridae. Because of the

structure of the apotele, the unusual ley setation, the reduced idiosomal

setation and the ventral sclerotization of the idiosoma this genus is considered

unusual enough to be placed in a separate subfamily. Luxton (1968)

pointed out that there was a good case for establishing a new family group

for Tangaroellus, The following single nominal species is included in this

genus} 7, porayus Luxton, 1968. Two paratype females (N1968271 and

N1968272) and two paratype males (N1968273 and N1968274) examined:

New Zealand; among barnacles, littoral zone, Magazine Point, near Nelson,

5.6.1967, col. G. W. Ramsay, dep. SAM,

LEE—RHODACARIDAE 205

SPECIES INCERTAE SEDIS

Neogamasellevans berlesei (Womersley) comb.n.

Oueenslandolaelaps berlesei Womersley, 1956b, p. 111.

FEMALE. Fig. 178-181. Idiosomal length, 520). All shields

strongly reticulated. Spermathecal ringed tube possibly opens near the

posterior paraxial edge of acetabulum IV.

MALE. Not known.

LOCALITY. Two females (N1968105 and N1968106) drawn or

examined: Australia; LF1I71, moss, beside Gold Creek, Brookfield, near

Brisbane, Queensland, 7.9.1966, col. D. C. Lee, dep. SAM.

The holotype female (N1968107) examined: Australia; soil litter,

Brookfield, near Brisbane, Queensland, 31.5,1949-10.6.1949, col. E. H.

Derrick, dep. SAM.

REMARKS. This species belongs to the Ologamasinae and not the

Sessiluncinae which contains Queenslandolaelaps, the genus it was originally

placed in, Beyond this | am uncertain of its relationships, but I have referred

it to Neogamasellevans as a temporary measure until more data, especially

the male morphology, is available.

Rhodacaroides costai (Sheals) comb.n.

Rhodacarus costai Sheals, 1962, p. 85.

FEMALE. The following characters not noted in the original descrip-

tion are listed here. Peritrematal shield is connected by a thickened striation

on cuticle to exopodal TV shield. Both tarsi [V are missing, so the setation

of this segment is unknown, but otherwise the leg chaetotaxy is as for

Gamasellus. Third hypostomal seta nearly level with seta hyp2, both being

a similar distance from seta hyp4. On palp genu, seta a/2 has a number of

lateral prongs. On palp femur, seta af in proximal third. Spermathecal

tinged tube opening not located,

MALE. Not known.

LOCALITY. Holotype female (1961-6-20-2) examined: South

America: surface soil, Los Arrayanes, Nahuel Huapi Reserve, Andes

Mountains, Argentina, 2-5.1959, col. C, Delamare Deboutteville, dep.

BM(NH).

REMARKS. This species belongs to the Ologamasinae and not the

Rhodacarinae which contains Rhodacarus, the genus it was originally placed

in, Beyond this | am uncertain of its relationships, but [ have referred it to

Rhodacaroides as a temporary measure until more data is available.

204 REC. S. AUST. MUS.. 16 (3): 1-219 July, 1970

Hydrogamasellus ubatubaensis Hirschmann, 1966

Gamasellus (Hydrogamasellus) ubatubaensis Hirschmann, 1966, p. 25.

REMARKS. This species from the rocky shoreline, Recife, Brazil,

cannot be placed in any established genus with confidence and | have been

unable to borrow the type material in order to decide whether or not a new

genus should be erected for it. Despite the fact that the peritrematal shield

is not fused to exopodal IV shield, I would regard this species as belonging

to the Ologamasinae and therefore have left it in Hydrogamasellus as a

temporary measure until it is more fully described.

TAXA SIZES AND DISTRIBUTION

Table | summarizes the number of species in the supraspecific taxa

and the distribution of those taxa amongst the major zoogeographical regions.

In the map (Fig. 427) of the zoogeographical regions the minor regions and

the abbreviations used for them are given, but these are only referred to in

the text under the genera headings. Kerguelenian is mis-spelt in Fig. 427.

It should be remembered that the rhodacarid fauna of South America is

probably as morphologically diverse as that of Australia, rather than that of

North America as suggested by the table. The unnamed species enumerated

include only species that have been described but not formally named.

TABLE |

Sizes and distribution or rhodacarid supraspecific taxa

Taxa Number of known | Major Zoogeographical Regions

species |

Nominal Unnamed) No PONT) E-

O A/S AC) Page

~ 1 | |

| | |

RHODACARIDAE = (52 |

distinct genus-group taxa) 220 49 N| Po NT'E,O A’ S |AC 17

(10), (14) (10) (16)| (6) (27)/ 13) (4)

RHODACARINAE .... 45 12 N PINT|E|O A — | S| 23

Rhodacarus ...5..,..... 17 2 | N| PINT) £E| oO} A|—|— 26

Afrozamasellus.......... 18 ] — —|—|E — — | — 30

Rhodacarellus .......... 9 — N | P A / 36

Rhodacaropsis ..... 2... | | 1 N| P E | 37

{ | |

| |

GAMASIPHINAE ..... 40 6 N| PINT E/O/;/A Sj) — | #

Gamasiphis o. 000005000. 16 4 N!| PIN?) £)O)}Al—, = 42

Caliphis 20.200... 0.4. 6 ~ -~;—|—}/—!—|/ Al Ss | — 52

Euepicrius ......0...44. 3 | | —|\A | Ss 5

LEE—RHODACARIDAE

TABLE |—continued

Sizes and distribution or rhodacarid supraspecific taxa—continued

Number of known

Major Zoogeographical Regions

205

Taxa species

| Nominal | Unnamed | N|P NT|E O|A!/S | AC Page

Gamaselliphis 2.02.02... | 5 | E | | 57

Gamasiphoides ......... 4 2 —|—|NT| EB A) S | — 59

Hydrogamasus ......... 4 P Al| Ss | — 64

Laelaptiella,... 00.5... 2 | A | 70

| |

LAELAPTONYSSINAE. 2 | | A }—|—]) 7

Laelaptonyssus ... 54.5 .- 2 -- | Aj} —}| — 72

OLOGAMASINAE 101 20 N P JNT| E A | S |AC 74

OLOGAMASINI ...... 38 8 — PNT E |; A} 8 |AC 83

Ologamasus ©... 2.52245. 3 | NT 84

Cyrniphis. 2G 5% Mahe, 7 A| S| — 90

Geogamasus .... 0000066. 4 NT A 92

Heydeniella— |

crozetensis-complex.... 2 6 97

dentatus-complex,... .. 6 P | A 101

Hydrogamasellus ....,.. 8 2 NT AC, 110

Neogamasellevans ....... | NT 117

Parasitiphis 2... .....-. 4 A AC 118

Pyriphis .. 00.0.0. cee | A 125

Rykellus .. ccc cee 2 126

GAMASELLINI ....... 63 19 /N|P ONT E A!) S AC) 127

Gamasellus— |

faleiger-complex ...... 18 10 N| P —|E A |— | — 130

discutatus-complex .. . . 6 | A 136

pyriformis-complex .... 1 1 E 137

Acugamasus- | |

punctatus-complex..... 4 A} S |— 140

natalensis-complex, .... | 8 1 E 143

Allogamasellus ......... 2 P 144

Cyrtolaelaps oo... 06... 6 1 N P 146

Euryparasitus 22.0.2... . 2 — N | P } |— 151

Evanssellus 0.0 0000545 2 NT | Ss | — 154

Heterogamasus... 0.4... 3 NT. Ss | 156

Hiniphis 2.0.0 05006 0544, | I | 157

Laelogamasus ......,.-, | 1 2 — 159

Litogamasus .. 60.64.65. l 2 S |AC. 160

Notogamasellus—

(Notogamasellus) ..... | E 164

(Podonotogamasellus) . . l | E | 165

Periseius—

(Periseius) coc ccece cas | 2 I N NT | A 166

(Psammosella) oo... 4. 2 — NP | 168

Pilates cries acta ae ne ere | 1 I | | S |AC! 169

Rhodacaroides .......... | l | 170

206 REC, S. AUST. MUS., 16 (3): 1-219 July, 1970

TABLE 1—continued

Sizes and distribution or rhodacarid supraspecific taxa—continued

Number of known Major Zoogeographical Regions

Taxa | species

Nominal Unnamed) NP NT! EO. A. S$ AC. Page

SESSILUNCINAE ..... 31 4 IN/P/—|EBE/O/;A/—]— } 172

| |

Sessiluncus .. 0.0000 ceee 4 4 —|/P/—/]E|o|}Al—|—I| 195

Antennolaelaps ......... 3 A | 178

Gamasellevans .......... 7 E 184

Gamasellopsis .......... 4 E 186

Gamasitus .. 0... 1 | A 187

Onchogamasus— |

communis-complex .... I | A 189

pumilio-complex ...... 2 A 190

Paragamasellevans ...... 2 E 193

Queenslandolaelaps ..... 1 A 195

Stylochirus 6.0.00. ee 6 NP 198

TANGAROELLINAE .. 1 | A 201

Tangaroellus oo. 0.006545 I | | A 201

| | |

207

LEE—RHODACARIDAE

PuBjeaz—MauU —Uy UeAejeW—opul —wCy uzunyoueW — dq uesehejgyul —Wwg ueiyius —lN ueiueyGajje —eN

Ueapodiyue —Bg uelsaudjod —dy asauiys—opy) —so VELagIS Sd yings =—83 ueli22q —GLN UlBzuNOW A490) IN

ue}jo9s —sOY ubivaljanilaxy 3g ueijesjsne —ey asaugyAsa —nQ UealPualips —WY isea ag upsjjijue —eLN UBIO /89 —IN

ugisso# —390y vejue;jabew —ws uedejew —wy veripur —ig uradoina —aq sam —Mg ura ly Wayyipu UN)

DILIYVING DILIYVLNVaNS NvIiTveisny AWLIN3SIHO DlLoOeVvavIve NVIdOIHLS TWWS|dOuLOaN DILDYVAN

DILOYVING

DILDYVLINVANS

ra)

’

‘

a ay i

Grandjean, 1966).

ammen-

after Vere

Zooygeographical Regions (

ig. 427,

208 REC. 8S. AUST. MUS., 16 (3): 1-219 July, 1970

CHANGES IN NOMENCLATURE

The author and date of publication are only given for names that were

misidentifications or are preoccupied,

NEW TAXA.

Subfamilies: Gamasiphinae; Sessiluncinae; Tangaroellinae,

Genera: Acugamasus; Caliphis; Cymiphis; Geogamasus; Hiniphis,

Litogamasus; Pilellus; Pyriphis; Rykellus.

Species: Gamasiphoides aitkeni; Parasitiphis aurora; Caliphis

calvus; Acugamasus cursor; Gamasiphis fornicatus: Heydeniella

goei, Hiniphis hinnus; Geogamasus howardi; Euepicrius lootsi;

Heydeniella| markmitchelli;, Onchogamasus pumilio; Oncho-

gamasus quasicurtipilus, Geogamasus skoshi; Antennolaelaps

testudo.

NOMINA NOVA,

Species: Hydrogamasellus gaussi for Neoparasitus crozetensis

Richters 1907; Hydrogamasellus richtersi for Gamasellus

crozetensis Richters, 1907.

NEW RANK.

Subfamily: Laelaptonyssinae.

Tribe: Ologamasini.

Genera: Gamaselliphis, Hydrogamasellus; Laelogamasus.

Species: Caliphis tamborinensiy.

NEW SYNONYMS.

Genera: Austrohydrogamasus under Parasitiphis; Epiphis under

Stylochirus; Megaliphis under Stylochirus: Micriphis under

Gamasiphis; Neogamasiphis under Gamasiphis; Periphis under

Stylochirus; Physallolaelaps under Stylochirus; Puchihlungia

under Laelaptonyssus; Stylagamasus under Antennolaelaps.

Species: Hydrogamasus antarcticus sensu Womersley, 1937 under

Parasitiphis aurora; Gamasiphis australicus sensu. Domrow,

1957 under Gamasiphis setosus; Sessiluncus heterotarsus sensu

Domrow, 1957 under 4Antennolaelaps testudo; Periseius littorale

under Periseius hamment; Hydrogamasus (Austrohydrogamasus )

watsoni under Parasitiphis jeanneli,

REVOKED SYNONYMS.

Genus: Heydeniella not under Gamasiphis; Laelogamasus not

under Gamasellus; Neogamasellevans not under Hydrogamasellus.

Species: Hydrogamasus littoralis not under Gamasus salinus.

LER—RHODAC ARIDAE 209

NEW COMBINATIONS. — ampulliger (ex Physallolaelaps) with

stvlochirus, ausiralica (ex Hydrogamasellus) with Heydeniella:

herlesei (ex Queenslandolaelaps) with Neovamasellevans (undet

species tacertue sedis); beorealiy (ex Gamasus) with Gamasellis;

hrunneus (ex Laelaps) with Parasitiphis; cavei (ex Qlogamasus) with

Hydrogamasellus, coleoptratus (ex Ologamasus) with Hydrogama-

sellus; convexus (ex Stylogamasus) with Antennolaelaps; costai (ex

Rhodacarus) with Rhoedacaroviies. (under species incertae sedis);

covdlis (ex Hydrogamasellus) with Geogameasus: erocetensis (ex

(iamasellus) with Hydrogamasellus; crozetensis (ex Neoparasitits)

with Hydrogamasellus; cymosus (ex Ologamasus) with Cymiphis:

darglenysis (ex Gamasellus) with Rykellus: delamarei (ex Hydre-

ramasellus) with Geegamasus: dentata (ex Hydrogametyellis) with

Hevdeniella, discutatus (ex Ologamasus) with Gamasellus, dvakens-

bergensis (ex Gamavsellus) with Acugamasus; dumosus (ex

Ologamasus) with Cymiphis; gamasiphivides (ex Mydrogamaselluy)

with Gamasiphoides, giganteus (ex Megaliphis) with Srylochleius:

erahami (ex Gamasellus) with Acugamasus, haemisphaericus (ex

Periphis) with Stylochirus; hammeni (ex Cyrtolaelaps) with Periseitis:

hickmani (ex Neogamasiphis) with Caliphis; hluhluwensis (ex

Gamasellus) with Acugamasns; feanneli (ex Gamasellus) with Para-

sitiphis: knysnaensis (ex Gamasellus) with Acugdmasus;, leptosceles

(ex Ologamasus) with Cymiphis; litoprethrix (ex Ologamasus) with

Gamusellus, loricata (ex Gamasiphis) with Heydeniella: ititcyuari-

ensis (eX Hydrogamasellus) with Gamasiplioides, macrosetosus (ex

Clamasellus) with Acugamasus; mansoni (ex Ologamasus) with

Cymiphis: minor (ex Mevaliphis) with Stylochirus: natalensis (ex

Gamuisellus) with Acugamasus: neolasmanicus (ex Gamasellits)

with Acugumasus; nkandhlaensis (ex Gamasellus) with Rvkellits;

novaezelandiae (ex Neogamasiphis) with Caliphis; nucilis (ex

Qlogamasis) with Cymiphis; paranatalensis (ex Gamasellus) with

Acugamasus, punctatus (ex Gamasellus) with Aciugamasnsy

pyrenvides (ex QOlogamasus) with Pyriphis; queenslandicus (ex

Neogvamasiphis) with Caliphis: rucovitzai (ex Gamasellus) with

Hydrogamasellus; rariar (ex Fpiphis) with Stylochirus, relata (ex

Hydrogamasellus) with Heydeniella: relicta (ex Hydrogamutsellus )

with Hevdeniella, rykei (ex Gamasellus) with Pilellus: salinus (ex

Hydrogamasus) with Pergamasus (Parasitidae); schustert (ex

Tydrogamasellus) with Caliphis; semipunctatus (ex Gamasellus)

with Aewgamasus, setosus (ex Cyrfolaelaps) with Litogamasus:

southeotti (ex Ologamasus) with Gamasellus; spurius (ex Eury-

parasitus) with Cyrtolaelaps; tamberinesis (ex Neogamasiphis) with

Caliphis; tindalei: (ex Ologamasus) with Gamasellus; validus (ex

Ologamasus) with Cymiphis; virgosus (ex Ologamasus) with

Gamaselluy; watson? (ex Gamasellus) with Actigamasus, watsoni

(ex Gamasiphis) with Cymiphis.

210 REC. 8. AUST. MUS., 16 (3): 1-219 July, 1970

INCLUDED AVAILABLE GENUS-GROUP NAMES

The names are in alphabetical order. The reference in parenthesis

indicates when the genus was first placed in this family if this was after the

name was made available,

Page

Acugamasus new genus .. . sw san! hel heres ache 139

Afrogamasellus Loots and Ryke, 1968 net asteert each dy} aos 30

Allogamasellus Athias-Henriot, L961b .. .. ec 4h 5% 144

Antennolaelaps Womersley, 1956 (Ryke, 1962b) ape od 4 178

Austrohydrogamasus Hirschmann, 1966 apy inchided) . 118

Caliphis new genus . .. 1. 1... . yeas ple tat at's 52

Cymiphis new genus .. .. Mee el tard wt tee att 90

Cyrtolaelaps Berlese, 1887 (Byans, 1957) a ee ae 146

Epiphis Berlese, 1916 (Ryke, 1962b) .. 2. ..00. .. .- 198

Euepicrius Womersley, 1942 (Lee, 1966) . .. .. 2. ©. 55

Euryparasitus Oudemans, 1902 (Evans, 1957) . .. 0. .. 151

Evanssellus Ryke, 1961b .. 0. 6p ce ee ee eee ees 154

Gamasellevans Loots and Ryke, 1967 .. .. 1... 0... 184

Gamasellopsis Loots and Ryke, 1966. .. .. .. 5. oe. 186

Gamaselliphis Ryke, 1961la. 2. 2. 6k ee ee ee 57

Gamasellus Berlese, 1892 (Ryke, 1958) .. 2. 2... 129

Gamasiphis Berlese, 1904 (Ryke, 1962b) .. .. «2. 42

Gamasiphoides Womersley, 1956 (Ryke, 1962b) .... .. 59

Gamasitus Womersley, 1956 (Ryke, 1962b) .. .. .. .. 187

Geogamasus new genus . 6. ee ee ee ee 2

Heterogamasus Trigardh, 1907 (Lee, 1966) .. .. .. .. 156

Heteroiphis Tragardh, 1952 (Ryke, 1962b) . .. 2. 2... 42

Heydeniella Richters, 1907 ye POBZBY oF yc koh. ap 96

Hiniphis new genus . .. ees 157

Hydrogamasellus Hitschmeann, 1966 ‘toewly included) 4 110

Hydrogamasus Berlese, 1892 (Ryke, 1962b) .. .. .. .. 64

Laelaptiella Womersley, 1956 (Ryke, 1962b) .. .. 2... 70

Laelaptonyssus Womersley, 1956 (newly included) .. .. 72

Laelogamasus Berlese, 1905 (Ryke, 1962b) ... .. 2... 159

Litogamasus new genus . .. Se, ene 160

Megaliphis Willmann, 1938 (Ryke, 1962b) . 4688 RF 198

Micriphis Berlese, 1914 (Ryke, 1962b) .. 2. 1. 0... 42

Neogamasellevans Loots and Ryke, 1967 .. .. 2... .. 117

Neogamasiphis Tragardh, 1952 (Ryke, 1962b). .. 2... 42

Notogamasellus Loots and Ryke, 1966 .. 2... .. 2... 163

Ologamasellus Berlese, 1914 (Ryke, 1962b) ay Agreed et 84

Ologamasus Berlese, 1888 (Ryke, 1962b) . .. .. 2... 84

LEE—RHODACARIDAE

Onchogamasus Womersley, 1956 (Ryke, 1962b)

Paragamasellevans Loots and Ryke, 1968 .. 3.

Parasitiphis Womersley, 1956 (Ryke, 1962b) ..

Periphis Berlese, 1914 (Ryke, 1962b)

Periseius Womersley, 1961 (Lee, 1966)... ..

Physallolaelaps Berlese, 1908 (Ryke, 1962b) ..

Podonotogamasellus Loots and Ryke, 1966 . .

Protolaelaps Trigardh, 1912 (Evans, 1957) ..

Psammonsella Haq, 1965 .. er Ge

Puchihlungia Samsinak, 1964 .

Pilellus new genus ..

Pyriphis new genus .. .

Queenslandolaelaps Wotnersley, 1956 ( Ryke, 1962b) .

Rhodacarellus Willmann, 1935 ..

Rhodacaroides Willmann, 1959 ,

Rhodacaropsis Willmann, 1935 ..

Rhodacarus Oudemans, 1902 .

Rykellus new genus . ..

Sessiluncus Canestrini, 1898 ( Ryke, 1958) .

Stylochirus G. and R. Canestrini, 1882 (newly included) ‘

Stylogamasus Womersley, 1956 (Lee, 1966) .

Tangaroellus Luxton, 1968 . one

EXCLUDED GENERA

The following genera were placed in the Rhodacaridae when made

available or in the publication referred to in parenthesis, but have since been

regarded as belonging to other families, a transfer followed here.

is the only genus newly regarded as belonging to another family.

Antennoseius Berlese, 1916 (Ryke, 1962b), to Ascidae.

Asca Heyden, 1826 (Ryke, 1961c), to Ascidae.

Panteniphis

Digamasellus Berlese, 1905 (Ryke, 1958), to Digamasellidae.

Gamasellodes Athias-Henriot, 1961b, to Ascidae.

Halolaelaps Berlese and Trouessart, [889 (Evans, 1955),

Halolaelapidae.

Leitneria Evans, 1957, to Halolaelapidae.

Longoseius Chant, 1961 (Ryke, 1962b), to Digamasellidae.

Pachyseius Berlese, 1910a (Ryke, 1962b), to Pachylaelapi

dae.

2)2 REC. 8S. AUST. MUS,, 16 (3); 1-219 July, 1970

Pantheniphis Willmann, 1949 (Athias-Henriot, 1968) to Ascidae.

This genus was not placed in any family when originally described,

but has since been allotted to the Rhodacaridae by Athias-Henriot

(1968). It contains one species, P. mirandus, only the female being

described when the name was made available (Willmann, 1949),

but the male has since been thoroughly described by Athias-Henriot

(1969), I have preferred to exclude this genus from the Rhoda-

caridae, transferring it to the Ascidae, mainly on the basis of three

characters (reduced leg setation with only |2 setae on tibia I,

2-pronged apotele, inconspicuous enlarging of setae on male leg UI

with only seta av on femur slightly enlarged to a short, blunt spine)

which all occur on members of the Rhodacaridae, but rarely, and

never together,

Protogamasellus Karg, 1962 (Karg, 1965), to Ascidae.

Saintdidieria Oudemans, 1939b (Ryke, 1961), to Halolaelapidac.

Saprolaelaps Leitner, 1946 (Evans, 1957), to Halolaelapidae.

Trachygamasus Berlese, 1904 (Ryke, 1962b), to Parasitidae.

ACKNOWLEDGMENTS

Lum indebted to Dr. R. V. Southcott, Messrs. P. F. Aitken, H. M. Cooper, G, F, Gross,

G. W. Howard, B. J. Martyn, L. W. Miller, N. MeParland, P. J, Mitchell, N. B. ‘Tindale

and Miss J. A, Herridge for kindly collecting litter and moss containing mites, and to

Drs. S. K. Bhattacharyya, M. Costa, J. Haq. W. Hirschmann, P. E. Hunter, K. H. Hyatt,

D. E. Johnston, F. Kjellander, M. Luxton, D, Macfarlane, M. Mrviak, F. Pegazzano, G. Rack,

K. Samsinak. E. Schulz, J, G, Sheals, M. Sellnick, C. Willmann and T. G. Wood for their

venerous assistance and in some cases for either making available mites for study or for

their gifts of miles.

1 un) also grateful to the Science and Industry Endowment Fund of the Commonwealth

Scientific and Industrial Research Organization and to the Mark Mitchell Research Foundation

for putting at my disposal funds which allowed me to travel to Europe, and to Professor

R. Zocchi. Director of the Stazione de Entomologia Agraria, Florence and The Trustees of

the British Museum (Natural History), London for permitting me to work in their respective

establishments.

Special thanks for their encouragement, assistance, discussion and criticism during the

prepuralion of this paper are due to Drs. C. Athins-Henriot, W, P, Croweroft, R. M. Emberson,

G. O. Evans, W, G, Inglis. G. C. Loots and Mr. G. F, Gross.

My greatest debt of gratitude is to Mrs. Brenda Head for inking in all my drawings of

particular species and for drawing Figs, |. 2, 3, 4 and 427, and also to Mrs, Joan Murphy for

patiently typing this manuscript.

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Tagil’tsey, A. A. and Men Yan-Tsuy’, 1963: Fauna and ecology of Gamasid mites, animals

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375-586.

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LEE—RHODACARIDAE 219

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RECORDS oF THE

SOUTH AUSTRALIAN /

MUSEUM Ky

A REVISION OF THE AUSTRALIAN

PENTATOMID BUGS OF THE

GENUS CEPHALOPLATUS WHITE

(Hemiptera—Pentatomidae—

Pentatominea)

By GORDON F. GROSS

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 4

23 October, 1970

A REVISION OF THE AUSTRALIAN PENTATOMID BUGS OF THE

GENUS CEPHALOPLATUS WHITE (HEMIPTERA-PENTATOMIDAE-

PENTATOMINEA)

BY GORDON F.. GROSS

Summary

The genus Cephaloplatus White 1842 as now understood includes 15 species. The genus is easily

recognized because, excepting C. minor Distant which is blackish brown, the species are yellowish,

yellowish brown or reddish brown moderately sized Pentatomine bugs. They have the juga of the

head broad and flattened and extending forward well past the apex of the anteclypeus. The

anterolateral margins of the pronotum are explanate and are usually crenulate or dentate. The

anterolateral margins exterior to the true anterior angles of the pronotum are usually produced

forward as angulate flattened lobes. These processes are indicated in the descriptions as the “regions

of the anterior angles”. The species are found in the tropical north of Australia and southwards of

this into the arid desert areas and the semi arid desert fringes, they rarely occue in the moister

regions of the south. The genus is known only from Australia.

A REVISION OF THE AUSTRALIAN PENTATOMID BUGS OF THE

GENUS CEPHALOPLATUS WHITE

( Hemiptera — Pentatomidae

By GORDON F. GROSS

Pentatominae )

INTRODUCTION

The genus Cephaloplatus White 1842 as now understood includes 15

species. The genus is easily recognized because, excepting C. minor Distant

which is blackish brown, the species are yellowish, yellowish brown or

reddish brown moderately sized Pentatomine bugs. They have the juga of

the head broad and flattened and extending forward well past the apex of

the anteclypeus. The anterolateral margins of the pronotum are explanate

and are usually crenulate or dentate. The anterolateral margins exterior to

the true unterior angles of the pronotum are usually produced forward as

angulate flattened lobes. These processes are indicated in the descriptions

as the “regions of the anterior angles”, The species are found in the tropical

north of Australia and southwards of this into the arid desert areas and the

semi arid desert fringes, they rarely occur in the moister regions of the south,

The genus is known only from Australia.

The species are quite difficult to separate if not illustrated, Untor-

tunately the previous descriptions are all verbal with the consequence that

only one or two of the more distinctive species could be named confidently

by a worker in this country who did not have recourse to the types,

With the assistance of the Sir Mark Mitchell Trust and the C.S.1.R.O.

Science and Industry Fund, I was able recently to visit Europe to help com-

plete the first part of a projected descriptive Handbook of the South Aus-

tralian Heteroptera and in so doing was able to see the Cephaloplatus types

lodged there. The type of Cephaloplatus nubifer Bergroth was found un-

marked as such in Helsinki and has been selected and labelled as a lectotype,

the types of Cephaloplatus granulatus Bergroth and Cephaloplatus reticulatus

Bergroth were located on my return amongst material lent for the purpose of

this study by the National Museum, Melbourne, similarly not marked as

types, these also haye been selected and labelled as lectotypes.

I am indebted to the Directors of entomological staffs of the following

institutions who permitted me to examine their collections, to make notes

on type material and in many cases lent critical and unsorted material to

Issued 23rd Qetaher, 1970]

2 REC. & AUST. MUS, 16 (4): 1-58 October, 1970

the completion of this project. The letters in brackets before each Institution

are the abbreviation used to designate the respective Institution in which

material is lodged in the “Distribution” section under each species.

(Q.M.) The Queensland Museum, Brisbane.

(U.Q.) The collections of the Department of Entomology,

the University of Queensland, Brisbane.

(A.M.) The Australian Museum, Sydney.

(A.N.LC, ) The Australian National Insect Collection. c/o

C,S.1.R.0., Canberra.

(N.M.) The National Museum, Melbourne.

(S.A.M.) The South Australian Museum, Adelaide.

(Helsinki) Universitetets Zoologiska Museum, Helsingfors.

(Stockholm) Naturhistoriska Riksmuseum, Stockholm.

(Paris ) Museum d'Histoire Naturelle, Paris.

(B.M.) The British Museum (Natural History), London.

(A.M.N.H.) = American Museum of Natural History, New York.

(U.S.N.M.) The Smithsonian Institution, The United States

National Museum, Washington.

(Snow) The Snow Entomological Collection, The University

of Kansas, Lawrence.

(C.A.) The California Academy of Sciences, San Francisco,

( Bishop) The Bernice P. Bishop Museum, Honolulu.

My especial thanks go to Drs, W. Hackman and M. Meinander

(Helsinki), Professor L. Brundin and Dr. P. [. Persson (Stockholm), Dr.

Knight, Mrs. J. M. Black, and Mr. L. Mound (B.M.), Dr. J. Rozen

(A.M.N.H.), Drs. R. Froeschner and J. Herring (U.S.N.M.), Dr. P. Ashlock

(Kansas), Drs. E. Ross and P. Arnaud (C.A.), and Miss S. Nakata (Bishop)

for the very special efforts on their part to see that all material relevant to

this project in particular was available for me to see and/or borrow at the

time | passed through.

SYSTEMATIC TREATMENT

Genus Cephaloplatus White 1842

Dryptocephalus?( Cephaloplatus) White, 1842, Trans. ent. Soc. Lond., 3:91.

Cephaloplatus Dallas, 1851, List ef the Specimens of Hemipterous Insects

in the Collection of the British Museum 1:131, 148.

Cephaloplatys Stil, 1867, Ofvers. Kongl. svenska Vetensk Akad Férh., 507.

1876, Kongl. svenska Vetensk Akad Handl., 14(4):55, 71. Lethierry &

Severin, 1893, Catalogue général des Hémiptéres 1:113.

GROSS-AUSTRALIAN CEPHALOPLATUS 3

Rather oval moderate sized Pentatominac, reddish brown, yellowish

brown, or blackish brown in colour.

Head coniparatively large, basally slightly swollen, apically flattened

or even concave, Eyes moderately prominent, very close to the anterior

margin of the pronotum in the subgenus Cephaloplatus and Delichoplatus

subgen. noy., well separated in Melanoplatus stbgen. nov. Juga laterally

expanded, produced well in front of the anteclypeus and their apical and

lateral margins sometimes strongly reflexed. Anteriorly their margins may

be truncate or rounded, laterally usually sinuate, in some species thrown

into a flattened spine or process in front of the eyes. Antennae very slender,

first segment not reaching the apices of the juga, antennophore usually with

a short blunt hooked process laterally.

Pronotum fairly flattish, the anterior margin strongly concave, the

lateral margins explanate and the anterior angles produced angulately und in

the subgenera Cephaloplarus and Dolichoplatus reaching forward to in front

of the eyes, in some species of Cephaloplatus sens, str, markedly so

(C, (C.) explanatus sp. nov.) The lateral angles are acute or ungulate,

the posterolateral margins ure sinuate with a low tumescence just behind the

lateral angles, the posterior margin is feebly concave,

Sctitellum more or less triangular and a little larger than usual in

Pentalominac, only very feebly raised basally and its apex rounded, often

broadly so,

Hemelytra always wider than the abdomen in their basal half, in most

but not all species narrowing in their apical half to expose some or all of

the connexivum, Veins of the membrane straightish, or apically forked, or

reticulate. ‘lhe laterotergites are unarmed anteriorly and posteriorly,

Beneath the bucculae are sinuously elevated and do not reach the base

of the head, the rostrum reaches about the hind coxae. The prosternum

is only shallowly depressed, the mesosternum ts very feebly convex and has

the median longitudinal ridge or raised line characteristic of groups of

Pentatominae from more or less the Ha/yy group onwards in arrangements

of genera such as Lethierry and Severin. The metasternum appears to be

almost flat. The embolium of the hemelytra ts well developed over the

thorax but is concealed behind this point. The legs are normal and the

ubdomen fairly convex.

Type: Dryptocephalus? (Cephaloplatus) pertyi White 1842, mono-

basic.

Remarks: White did not give a generic diagnosis and the only deserip-

tions in Dallas and Stil form part of their respective Keys to genera. The

often used spelling Cephaloplatys dates [rom Stil but is not valid, it is

not striclly a dupsuy calami as Stal knew very well what he was doing—vide

his 1876 citation of references under “Cephaloplarys’’.

4 REC, 8S. AUST. MUS. 16 (4): 1-58 October, 1970

Fifteen species can be recognized as belonging to this genus and they

may be separated by the following Key.

Key to subgenera and species of Cephaloplatus White.

1. Smaller (7-8 mm.), blackish brown; antero-

lateral explanate margins of pronotum con-

cave, only minutely crenulated, anterior

angle produced into a rather curved triangu-

lar outwardly directed flattened process

which does not reach in front of the eye . .

Larger or smaller, yellowish brown or reddish

brown. Anterolateral margins of pronotum

straight, angulately concave (spurcatus

Walker) or convex, generally strongly

crenulated, at least anteriorly, the anterior

angles rather lobately or triangularly pro-

duced forward, reaching to, or well in

front of, the eyes ..

2, Smallish (7-8 mm.), elongate; anterior angles

of pronotum produced into a lobate anter-

iorly directed flattened process which

reaches the anterior margins of the eyes,

anterolateral margins of the pronotum not

explanate or depressed behind these lobes .

Larger (more than 8 mm.), obovate; anterior

angles of pronotum produced into an angu-

late process which reaches or surpasses the

anterior margins of the eyes, anterolateral

margins explanate and depressed (at least

anteriorly) behind these lobes . .

Subgenus Melanoplatus

nov. (one included

species C. (M.)

minor Distant)

Subgenus Dolichoplatus

noy. (One included

species C. (D.) elon-

gatus Distant)

(Subgenus Cephalo-

platus White)

GROSS—AUSTRALIAN CEPHALOPLATUS

Juga relatively narrow across at the level of

the apex of the anteclypeus, not as wide as

width of head across eyes; lateral margins

of head produced into an obvious laterally

directed flattened tooth or triangular pro-

cess just in front of the eyes . ,

Either juga relatively wide across at the level

of the apex of the anteclypeus, as wide as

width of head across eyes and/or lateral

margins of head not produced into an

obvious flattened tooth like or spinous pro-

cess in front of eyes [C. (C.) pertyi has

expanded juga and something of a produced

tooth in front of the eyes] .. ..

Membrane reticulate; generally over 11 mm.

in Tength oo) 66. 83 Ah Talay wae a

Membrane not reticulate; generally under

11 mm. in length ..

Margins of head just in front of eyes produced

into a broad triangular process which is

strongly convex above

Margins of head just in front of eyes produced

into a spine or tooth like process . .

. Anterior angles of pronotum exteriorly pro-

duced forward to about anterior margins

of eyes ..

Anterior angles of pronotum exteriorly pro-

duced forward well in front of anterior

margins of eyes, reaching to anterior margin

of produced triangular process on lateral

margin of head . .

Small (8.0-9.5 mm.); greyish; occurring in

the gulf of Carpentaria region . .

Larger (9.75-11.0 mm.); reddish or greyish;

occurring in Western and Central Australia

C. (C.) reticulatus

Bergroth

C. (C.) clementi

Distant

C. (C.) pellewensis

sp. nov.

10.

ll.

REC. S. AUST. MUS. 16 (4): 1-58

Greyish; juga at their apices rather widely

separated; anterolateral margins of pro-

notum rather strongly angulately concave

just behind the middle . . Dee 4d

Reddish; juga at their apices contiguous;

anterolateral margins of pronotum only

slightly concave near their middle . .

Anterolateral margins of pronotum almost

straight, slightly convex anteriorly, if denti-

culate or serrate then so very minutely

that the margin appears to be entire , .

Anterolateral margins of pronotum angulately

concave, the angulation nearer the lateral

angle than the anterior; the anterolateral

margins strongly denticulate or crenulate . .

Anterolateral margins of pronotum strongly

angulately concave just behind the middle,

anteriorly to this angulation with four or

more strong denticulations; juga apically

only shortly surpassing the apex of the

anteclypeus and usually their apices clearly

divaricate; occurring in the North of the

Northern Territory and Western Australia,

and in Cape York Peninsula and the Torres

Strait Islands -.

Anterolateral margins of pronotum not so

strongly and angulately concave behind the

middle, anteriorly to this angulation crenu-

late, the crenulations all of the same size;

juga apically more longly surpassing the

apex of the anteclypeus and usually their

apices not divaricate (although they may

not be contiguous); occurring in Queens-

land south of Cape York Peninsula and

in New South Wales - .

Juga markedly expanded and foliaceous, apic-

ally widely divaricate; near the apex of the

anteclypeus the width of the head across

the juga is as wide or wider than the width

across the eyes -.

October, 1970

C. (C.) australis Dallas

C. (C.) nubifer Ber-

groth

C. (C.) bellus sp. nev.

(C.)

darwini

Distant

C. (C.) spurcatus

Walker

GROSS

Juga not markedly expanded and foliaceous,

apically widely divaricate or not; al the level

of the apex of the anteclypeus the width

of the head across the juga not equal to or

wider than its width across the eyes .. ..

Anterolateral margins of the pronotum and the

basal third of the exterior margin of the

corium strongly serrate or denticulate . .

Anterolateral margins of the pronotum and

the basal third of the exterior margin of

the corium nearly entire, only very minutely

denticulate or crenulated .. 2. 2. 1. 4.

Anterior angles of the pronotum produced well

in front of the eyes; a pair or more longi-

tudinal ochraceous or luteous stripes on

the-scutellum 2. 4. 2.6 wi ia we ee ae ca

Anterior angles of the pronotum produced

about to the front of the eyes, no longitudi-

nal paler stripes on the scutellum . .

Three longitudinal ochraceous or luteous

stripes on scutellum composed of a broad

one on each side not reaching apex (and

margined with dark) and an_ obsolete

median one; membrane brownish hyaline

with piceous veins; no transverse ridge

between the lateral angles of the pronotum

More than three longitudinal ochraceous or

luteous stripes on scutellum composed of a

narrow one on each side not reaching apex

(and margined with dark) and two or

three median ones divaricating basad;

membrane milky hyaline with brown veins:

a conspicuous transverse ridge running

between the lateral angles of the pronotum

Subgenus Melanoplatus nov.

AUSTRALIAN CEPHALOPLATUS 7

C. (C.) pertyi (White)

C. (C.) explanatus sp.

nov.

Cc. (C.) pallipes

Walker

C. (C.) granulatus

Bergroth

C, (C.) fasciatus

Distant

This subgenus is being erected for one species, Cephaloplatus minor

Distant, which shows a number of differences from the species which are

placed here in the subgenus Cephaloplatus sensu stricto. Cephaloplatus

8 REC, & AUST. MUS. 14 (4): 1-58 Qetober, 1970

ninor is a piceous colour whereas these species of Cephaloplatus sensu

stricto are reddish or yellowish in appearance, the lateral margins of the

pronotum although produced are shaped quite a lot differently. The lateral

margins are very little explanate trom the lateral angles forward until a point

just in front of the calli where they suddenly are produced as explanate

triangular processes which are directed forward and outwards, the processes

are strictly processes of the anterolateral margins because the anterior angles

are distincly evident lying behind the eyes and interiorly to these structures.

In Melanoplatus the anterior lobe of the pronotum is also raised and very

rugulose whereas in Cephaloplatus senyu stricto it is declivous. Otherwise

the species seems to belong in the one genus with other Cephaloplatus species

and does not merit the erection of a new genus purely to accommodate it.

Type: Cephaloplatus miner Distant 1910,

Cephaloplatus (Melanoplatus) minor Distant

Cephaloplatys minor Distant, 1910, Ann. Mag, nat. Hist., (8)6; 474,

Fig. |

Smallish, suboval, strongly piceous in dorsal view with one or two

lighter areas, Densely and finely punctate, the punctations concolorous.

Head with juga somewhat foliaceously developed and produced forward

past the upex of the anteclypeus, though not much. Luteral margins of the

juga in front of the eyes produced into a blunt triangular process which is

strongly concave above. Lateral margins of juga in front of this process at

first incised then broadly and semicircularly convex to their shortly rounded

anterior angles, Inner margins of juga shortly truncate, divaricate. Lateral

margins of the juga somewhat raised so that the head in front of the eyes

is somewhat convex. Head between the ocelli somewhat raised, eyes moder-

ately prominent, in Most specimens fairly well separated from the anterior

margin of the pronotum (this does not appear in the drawing), facets

occupying most of the anterior surface. That part of the juga lying in front

of the insertions of the antennae tending to be somewhat paler than the rest

of the body, sometimes also some pale spots on the anteclypeus. Antenno-

phore with a short reflexed spine exteriorly; antennae moderate sized, five

segmented, the segments progressively longer from one through to five, the

first not reaching the apices of the juga.

Pronotum with the anterior and posterior lobes slightly raised with a faint

depression between the two, Anterior margin concave behind the collum,

the anterior angles subacute and located just behind the inner margins of

the eyes. Anteroluteral margins beginning at the lateral angles, only slightly

produced and reflexed concave as they run forward to just in front of the

calli where they abruptly turn outwatds to form an outwardly and forwardly

CEPHALOPLATUS

GROSS—AUSTRALIAN

Brenda K. HERO

Cephaloplatus (M.) miner Distant.

Fig.

10 REC, S, AUST, MUS. 16 (4): 1-58 Oclaber, 1970

directed, somewhat recurved, spinous process which is flat on top, between

this process and the anterior angles truncate, Lateral angles subacute,

posterolateral margin angulately convex, posterior margin nearly truncate.

The raised portion of the anterior lobe of the pronotum and to a certain

extent some of the posterior lobe rather rugulose.

The scutellum shaped very much as other members of the genus, the

concave angulation of the lateral margins a little nearer the middle than

in some of the others, tip broadly rounded, In each basal angle a somewhat

clongate concolorous fascia, interiorly to this margined by a callous or

smooth raised granule, a light point at the base of the scutellum medially

and in some specimens a short streak on the ventral margins on either side

just before the apex. Base of the scutellum rather triangularly raised, apical

half flatter, somewhat rugulose.

Hemelytra wider than the body only in the anterior quarter, the exterior

margin of the corium forming a smooth gently convex curve, entire, not

irregular. This curve leaves about half of the laterotergites exposed. Hind

margin of the corium somewhat convexly produced exteriorly, then more or

less truncate becoming broadly convex interiorly, Membrane with slightly

darker veins, several of the veins forked, one or two closed cells basally,

Laterotergites yellowish or yellowish brown with a transverse bar immediately

behind each incisure which is piceous and which runs into a narrow longi-

tudinal piceous area along the inner edge of the connexivum,

Beneath concolorous with above, if anything somewhat durker, The

following yellowish areas are present: the anterior underside of the head

(and the bucculac) save for a longitudinal piceous stripe immediately in

front of the insertion of the antennae; the base of the head; the rostrum; all

legs; the extreme exterior of the abdominal ventrites making the edge of the

abdomen appear us if it has a yellowish stripe: the mule terminalia. On the

thorax the epimera and episterna and most of the pleura are somewhat

lightened.

Length, 7,5-8.5 mm.

Distribution: Queensland Holotype & and | other, Peake Downs

(B.M.); 1 6, 2 29%, Peake Downs (Stockholm): 1 4, Flinders Island.

Jan, 1927, coll, Hale & Tindale; 3 49, 2 9%, Gladstone, coll, Lea

(S.A.M.); 26 6, 3992, Clermont, coll. Dr, K. K. Spence; 29 9.

no precise locality, 25 August 1929, coll. Dr. K. K. Spenee (A,M.):

329, Gin Gin, 13 October 1901, coll. W. W, Froggatt (A.N.LC.):

19, Mutchilba, Feb, 1933, coll, A, D. Selby (N.M.); 14 12, Mount

Carbine, 20 July, 1932, coll. Darlington on Harvard Expd, (A.M.N.H.).

Northern Territory \ (abdonien missing), Horn Islet in Sir Edward Pellew

Group, [5-21 November 1968, coll. B. Cantrell (U.Q.). 9 Western

Australia le. Derby, coll, W. D. Dodd (S,A.M.).

GROSS—AUSTRALIAN CEPHALOPLATUS 1

Remarks; The species is very easy to recognize in the genus Cephalo-

platusy sensu lato. Its conspicuous piceous colouration, its relatively small

sive and the fact that the anterolateral margins of the pronotum ure formed

into an outwardly directed triangular process which dees not reach forward

much behind the hind margin of the eye distinguishes it from all members of

the subgenus Cephaloplatus sensu stricto.

Subgenus Dolichoplatus nov.

This subgenus is being erected to accommodate the one species

Cephaloplatus elongatus Distant, which shows certain differences to those

species placed here in the subgenus Cephaloplatus sensu stricto. Species

placed in the last mentioned subgenus are all rather oval whereas C. elongatus

is a lot more elongate; the anterolateral margin of the pronotum in C,

elongarus is not expanded and laminate, only the anterolateral margin just

exterior to the anterior angle is produced as a roundish lobe projecting to in

front of the eyes whereas in species of Cephaloplatus sensu stricto the

anterolateral margins of the pronotum are expanded and more or less laminate

at least in their anterior half and the produced portions outside of the

anterior angles are always acute at their anterior apices. The pronotum of

C’. elongatus has about 12-20 large granules on the disc, in other Cephalo-

platus species these are either very much smaller and very much more

numerous or absent,

Type: Cephaloplatus elongatus Distant 1899,

Cephaloplatus (Dolichoplatus) elongatus Distant

Cephaloplarys elongatus Distant, 1899, Ann, Mag. nat. Hist., 7(4):433.

Fig. 2

Smallish, elongate ovate, rather pale brownish in macroscopic dorsal

view, The ground colour in fact is light brownish yellow in front of about

the middle of the pronotum and yellowish behind this point but this is

obscured somewhat by a moderately dense and fairly regular brown puncta-

tion.

Head with juga moderately follaceously developed, produced to about

the level of the apex of the anteclypeus, Lateral margins of the head in

front of the eyes produced into a blunt triangular process which is only

faintly concave above and then only anteriorly. Lateral margins of juga in

front of this forming a segment of a convex curve to their comparatively

broadly rounded apices, the latter fairly widely separated and the inner

margins of the juga tending to be a little obliquely rounded in front of the

point where they touch the unteclypeus. The anteclypeus is rather declivous

from a point just in front of the produced triangular lobes of the margin of

12 REC. S. AUST. MUS. 16 (4): I-58 October, 1970

Fig. 2: Cephaloplatus (D.) elongatus Distant.

{Photograph by courtesy of the Trustees of the British Museum (Natural History).

CROSS—AUSTRALIAN CEPHALOPLATUS 13

the head forwards, the inter part of each jugum tends to be depressed com-

mesurately on either side so that the head is anteriorly depressed in the

middle, apex of anteclypeus Well below the apices of the juga. Head between

the ocelli feebly raised, cyes moderately prominent and very close to the

anterior margin of the pronotunt, facets occupying all but the flattened

posterior surface. Antennophore with a short reflexed spine exteriorly,

antennae five segmented and rather shorter than in most other species of

the genus, second third and fourth segments nearly subequal and each one

longer than the first, fifth the longest, first not reaching apex of jugum,

Third and fourth brown in apical one half, fifth brown except in basal fifth,

Pronotum with the posterior lobe raised and sloping declivously

anteriorly (o the anterior margin. Anterior margin concave behind the eyes

and collum, anterior angles and part of the anterolateral margin of the

pronotum produced into an apical rounded anteriorly produced lobe which

extends forward to nearly the level of the anterior margin of the eyes,

Anterolateral murgins behind this structure hardly or not explanate or

laminate somewhat concavely excavate and somewhat irregular though

not denticulate or crenulate, produced a little in front of the lateral angles.

Lateral angle comparatively broadly rounded, posterolateral margin some-

what concave; posterior margin almost straight, Dise of pronotum with

between 12 and 20 callous raised areas or very large granulations.

Scutellam comparatively longer than in other members of the genus, the

concave angulation of the lateral margins very nearly at the mid point

of the length of the scutellum. Apex tending to be acuminately rounded, the

dise from somewhat in front of the upex to the base progressively though

not very raised, depressed on either side just in front of the angulation of the

lateral margin and again in each basal angle where there is a brownish

impression bordered on its inner side by a luteous callous area, traces of other

such ureas on the scultellum.

Hemelytra wider than ihe body only in their anterior quarter, the exterior

margin of the corium anteriorly rather concave then forming a smooth convex

curve lo ils apex, the margin a little irregular anteriorly, Hind margin of

corium almost straight or only very slizhtly concave, its inner angle broadly

rounded. Membrane whitish with only slightly darker veins, the latter (at

least distally) apparently mainly parallel, Laterotergites concolorous except

for a submurginal fine brown stripe, their posterior angles somewhat bluntly

projecting.

As the unique specimen is carded not a great deal is visible of the

underside. The punctation and colouration seems much the same as that

above. A short brown stripe in front of each untennifer, the usual somewhat

curved brown stripe on the propleuron on the muscle scar, on the meso-

pleuron, metapleuron, and the sides of abdominal segments II-VI the

14 REC, & AUST, MUS, l@ (4)2 1-58 October, 1970

punctations tending to be concentrated to form a denser and darker sublateral

line, a brown spot anteriorly and laterally on each abdominal sternite.

Femora and tibiae coarsely maculated with brown.

Length: 7 mm.

Distribution: North Western Australia Holotype @ S. Heywood L.,

90-126 (B.M.).

Remarks; The holotype is the only specimen of this species to hand;

it came from a very poorly collected area and it is not practicable to predict

its possible range,

Subgenus Cephaloplatus White

To include species which are are elongate oval or ovate but which have

the anterior angles and the anterior part of the anterolateral margins of

the pronotum extending forward us angulate processes to at least the level of

the anterior margins of the eyes, The anterolateral margins of the pronotum

are also rather explanate, particularly anteriorly,

Type: Pryptocephalus? (Cephaloplatus) pertyi White 1842, monobasic.

Cephaloplatus (C.) pallipes Walker

Cephaloplatus pallipes Walker, 1868, Cat, Hem. Het Brit. Mus., 3:541.

Cephaloplatys pallipes Distant, 1910, Ann. Mag, nat, Hist., (8)6:472.

Fig, 3.

Rather smaller than average for the genus, strongly greyish in macro-

scopic appearance although the ground colour is actually luteous or light

yellowish with numerous brown or blackish brown punctations. The hinder

parts of the pronotum, scutellum, and the coriaceous parts of the hemelytra

densely covered with small luteous or light yellowish granules.

On the head the juga are expanded somewhat though not as foliaceous

as in some of the succeeding species, extending somewhat in front of the

apex of the anteclypeus. Anterior margins of juga oblique. shortly rounded

to join the parallel lateral margins which immediately in front of each eye

are convex, inner margins divaricate, short. Disc of head rather rugulose

and coarsely punctate, head slightly raised behind. Eyes moderately promi-

nent and the facets occupying nearly all of the optic process save the oblique

and flattened hind margin, Antennae moderately slender, first segment the

shortest, third segment the second shortest, second and fourth segments

subequal, fifth segment slightly longer, first segment not reaching the apices

of the juga,

Pronotum rather strongly raised posteriorly, flatter and more declivous

anteriorly, the anterolateral margins explanately produced progressively from

posteriorly to anteriorly. Region of the anterior angles shortly triangular but

GROSS—AUSTRALIAN CEPHALOPLATUS

3mm, od

Les

Jee ety,

“Be

€

aaa ee

bites Ar Oe A

by Oat

a4,

Cephaloplaius (C.) pallipes Walker.

Fig.

16 REC. 8S. AUST. MUS, 16 (4); 1-58 October, 1970

only extending to aboul the level of the anterior margin of the eyes. Anterior

margin convex behind the collum, shortly truncate behind each eye, then

slightly diverging to the apices of the produced angles. Anterolateral margins

almost straight or perhaps very slightly concave, trregular and tending some-

what crenulate in their anterior third. Lateral angles irregularly truncate,

posterolateral margins nearly straight, posterior murgin very slightly concave,

Hinder portion of the disc of the pronotum densely covered with small

luteous granules, these granules between the lateral angles tending to be

more concentrated und to form a more or less continuous sinuate line.

Scutellum substantially triangular, with the usual slight concavity some-

what behind the middle. Basal half rather raised, apical half rather flat,

lip broadly rounded, In each basal angle a somewhat elongate black fovea

inward of which lies a rather prominent Juteous large granule or small

lumescence, Apex of scutellum sometimes with a darkened medial longitudi-

nal line. Scutellum densely covered with small luteous granules,

Hemelytra wider than the body in their basal quarter, this portion of

the corial margin slightly irregular. Behind this point the lateral margins of

the corium smoother and gradually converging posteriorly leaving some of the

connexivum exposed, the degree of convergence is not so marked as in any

of the preceding species. Hind inargin of corium somewhat sinuate, Cortum

and parts of the clavus covered with small Juteous granules. Membrane

milky hyaline, veins the same colour, several of them forked. Luaterotergites

with their posterior angles not prominent, incisures between the laterotergites

infuscated as is the inner half of each laterotergite,

Beneath mainly concolorous with the dorsal surface, punctations larger

and darker on the head and prothorax, finer and much denser on the

abdomen, Head just inward of the lateral margin infuscated, all the thoracic

sterna brown. Femora with some spaltered brown granules, sometimes the

tibiae also, Male genitalia with the posterior ridge convexly curyed rather

as in Fig. 6B, its lateral angles stronyly prominent and directed posteriorly.

Length: 9.5-10.75 mm.

Distribution: Unlocalized. Holotype 2 (B.M.), Western Australia,

1 4, Junction of Fitzroy and Margaret Rivers, 1896, coll, Calvert Expedition

(S.A.M.). Nerthern Territery. | 2°, Horn Islet in Sir Edward Pellew

Group, 15-21 November 1968, coll, B. Cantrell (U.O,); a series, Alexandra

(B.M.); 2 9 9%, Newcastle Waters, 2 June 1929, coll, T. G, Campbell

(A.M,); 3 48, 2 99, Neweustle Waters, 2-5 June 1929, coll. T. G.

Campbell; 2 ¢ ¢. Brunette Downs, 20 March 1968, coll, L. Hall; | @, 1 ?,

2 miles South of Barrow Creek, 13 February 1966, coll. Britton, Upton &

McInnes; | @, 22 miles South of Alice Springs, at light on railway, 15

February 1966, coll. E. Britton (A.N,L.C.); large series, 28 miles South of

GROSS—AUSTRALIAN CEPIHALOPLATUS 17

Renner Springs, at ultraviolet light, coll, N. McFarland; | 4, unlocalized,

coll, S.A. White (S.A.M,), South Australia. 1 4, 4 2% 9, Oodnadatta;

1 ¢, North East Corner of state, coll. F, Parsons (S.A.M.). Queensland,

| 4,2 2 2%, Winton—Longreach, 15-24 August 1963, coll, T. BE. Wood-

ward: | ¢, Mt. Isa, 23 January 1968, coll. B, Cantrell: | 4,2 92, 1,

Cloncurry, 15 April 1947, coll. H. Bell (U.Q,); 1 ¢, 1 2, Cairo Station,

15-31 January 1951, coll. C. McC. (N.M,);9 ¢ 4,6 2 ®,at light, Norman-

ton, 3-4 May 1963, coll. P, Aitken & N. B. Tindale: 1 9, at light Mornington

Island Mission, 12 May 1963, coll. P. Aitken & N. B. Tindale (S.A.M.).

Remarks: This is the first of the species of the Subgenus Cephaloplatus

sensu stricto to be treated and the first of a section within the subgenus which

does not have a prominent and acute spine on the lateral margins of the

juga in front of the eyes. This species can be recognized quite easily from the

other members of its section in that the anteriorly produced angulate portions

of the pronotum do not protrude in front, or only very little in front of the

anterior margin of the eyes. Its greyish colouration is similar to that of

Cc. (C.) granulatus Bergroth but it lacks the strong luteous fasciae on the

scutellum and is also rather smaller, All of the localities cited are in

moderately uwrid to very arid regions,

Cephaloplatus (C.) fasciatus Distant

Cephaloplatys fasciatus Distant, 1881, Trans. ent. Soc, Lond,, 212.

Fig. 4, 6A

Average size for the genus and rather elongate, ground colour luteous

with brown and some patches of blackish punctations, the latter along the

anterior half of the lateral margin of the scutellum, and a number of small

luleous pranulations on the hind margin of the pronotum, the scutellum

and the coriaceous parts of the hemelytra.

Head with juga as flattened or foliaceous structures which however,

are not unusually extensive in relation to the area of the rest of the head, at

their widest they do not extend much past the inner margin of the eyes nor

anteriorly much in front of the apex of the anteclypeus. Anterior margins

of the juga at first oblique then rounding broadly onto the virtually straight

lateral margins, latter incised just in front of anterior margin of the eyes,

somewhat reflexed. Inner margins short, sometimes touching, sometimes

not, Eyes moderately prominent and the facets occupying nearly all of the

optic process save the oblique and flattened hind margin. Antennae slender

and five segmented, fourth and fifth segments the longest and subequal in

length, brownish or yellowish, apices of the fourth and fifth segments some-

times infuscated.,

‘

wa <e VM KITSON,

Fig. 4: Cephaloplatus (C.) faseiatuy Distant.

GROSS—AUSTRALIAN CEPHALOPLATUS 19

Pronotum somewhat raised posteriorly, flatter and more declivous

anteriorly, between the lateral angles a continuous line of raised contiguous

granules. Anterolateral margins progressively more explinate from post-

eriorly to anteriorly, the region of the anterior angles forming a curved

inangular process which extends somewhat in front of the eyes. Anterior

margin of the pronotum cancavely excavate behind collum, shortly truncate

behind the eyes, margins then virtually parallel anteriorly to the apices of

the anterior processes. Anterolateral margins somewhat crenulate, at first

convex unteriorly then becoming slightly and angulately concave at about

half their length back, Lateral angles irregular. posterolateral margins almost

straight, posterior margins straight.

Scutellum substantially triangular with the lateral margins gently

angulutely concave, the tip more or less rounded, Basally somewhat raised,

flattened in the posterior half, Dise with five rather diffuse luteous stripes.

outer pair reaching only to the angle of the literal margins, the median one

reaching to only about the same level, thereafter obsolete, the exterior pair

murgined on their outside by a dense streak of black punctations, this streak

becoming more obsolete as it approaches the angle of the lateral margins.

Hemelytra wider than the body in their basal quarter, this portion of

the corial margin slightly irregular. Behind this point the lateral margins

of the corium smoother and gradually converging posteriorly leaving pro-

vressively more of the connexivum exposed, hind margin of corium sinuiate,

convex interiorly, Membrane subhyaline with veins and spots between the

veins only slightly darker, several of the veins forked. Laterotergites with

their posterior angles somewhat prominent.

Beneath tainly concolorous with the dorsal surface, covered with dense

lurk punctations, these punctations finer on expanded lateral margins of the

pronotum and on the abdomen. Head beneath with a short black streak

beginning on the upper insertion of the aptennae and proceeding forward

for a short distance on either side. Prothorax on either side with a black

or picecous longitudinal bar extending from the anterior margin three quarters

of the way to the posterior margin, ut the base of the epimeron and episternum

sometimes a patch of darker punctations. The dark line continued though

much more narrowly onte the lateral areas of the mesothorax and metathorax

and then beconmng wider and extending to about the middle of the sixth

ventrite, A small black patch at the extreme base of the head and all

thoracic sterna black or piceous, two semicircular piccous areas On the third

abdominal seginent immediately behind each metacoxa. a small median

spot neur the base of both the fifth and sixth abdominal segments and a

median longitudinal brown stripe running from the base of the seventh to

three quarters af the distance posteriorad piceous.

Length: |0.0-12.3 mm.

REC. & AUST. MUS. 16 (4): 1-58 Oetaher, 1970

Brenna K, Husearo,

Fig. 5: Cephaloplatus (C.) granulains Bergroth

GROSS—AUSTRALIAN CEPHALOPLATUS 2I

Distribution: Queensland. Holotype 2, Rockhampton (B.M.); 4

“Austral. bor.”, coll, Damel and Rockhampton (Stockholm); 1 4,5 2 2,

Cairns (1918), at light (1920), coll. J. F. Ulingworth (Bishop); 14, 12,

Kuranda, coll. F. P. Dodd: 22 %, Bowen, coll. A. Simpson; 12, 2? %, at

light, Normanton, 4 May 1963, coll. P. Aitken & N. B, Tindale; | 4,3 2 &,

Mornington Island (Birri), 8 & 12 May 1960, coll, P. Aitken & N. B.

Tindale; | ¢ 2% 2, at light, Mornington Island, May 1963, coll. P, Aitken &

N. B. Tindale (S.A,M.):; 14, Coen, 14-28 May, 1951, coll. C. Oke; 1¢,

unlocalized, donated F. P, Spry, 5 October 1922 (N.M.). Northern

Territory. Series Adelaide River and Stapleton (B.M.); 1 2, Darwin, coll.

W. K. Hunt: | ¢, Batchelor, coll. G. F. Hill; | 2, Daly River, coll. H.

Wesselman; | @, unlocalized, 1875, coll. Tepper (S.A.M,); 1 @, Pt. Denison

(A.M.); 34 6, 49 @, Sixty Mile, on rice (Oryza sativa L.), 23 March

1956, coll. L. D. Crawford (A.N.LC.), Western Australia, 14, Leopold

Downs, coll, W. R. Richardson; | &, Junction of the Fitzroy and Margaret

Rivers, 1896, coll, Calvert Expedition (S.A.M.), New South Wales. 12%,

No. 393 (A.M.N.H.).

Remarks: This species is more elongate and browner than C, granulatus

Bergroth, the lateral margins of the pronotum are yaguely excavate, and

there are five pale luteous bars on the pronotum.,

Cephaloplatus (C.) granulatus Bergroth

Cephaloplatys granulatus Bergroth, 1895, Proc. R. Soc. Vict., 7:288.

Cephaloplatus granulatus Tay, 1966, Pap. Dep, Ent. Univ. Qd., 2(4):76,

figs,

Average sized for the genus, ground colour luteous with brown and

blackish punctations, the latter finer and denser on the hind lobe of the

pronotum, and a number of small Juteous granulations on the hind portion

of the pronotuny and on the scutellum and coriaceous parts of the hemelytra.

Head with juga as llattened or foliaceous structures which however, are

not unusually extensive in relation to the area of the rest of the head, at their

widest they do not extend much past the inner margin of the eye nor

anteriorly much in front of the apex of the anteclypeus. Anterior margins

of the juga at first slightly oblique then rounding broadly onto the lateral

margins, latter at first straight then gently convex in front of the eyes, some-

what reflexed. Inner margins short, sometimes touching. sometimes not.

Eyes moderately prominent and the facets occupying nearly all of the

optic process save the oblique and flattened hind margin. Antennae slender

and five segmented, second, fourth and filth segments subequal in length, first

not reaching the apex of the jugum, light brownish in colour.

REC, $ AUST, MUS, 16 (4) 1-58 Oefeber, 1970

Pronotum rather strongly raised posteriorly, Hatter and rather declivous

anteriorly, the anterolateral margins explanately produced, at least on the

anterior half, and the region of the anterior angles produced forward as a

rather triangular process Which does not reach much in front of the eyes.

From the apices of the anterior angles the anterior margin of the pronotum ts

convexly excavate, though shortly truncate just behind each eye. Antero-

lateral margins distinctly crenulate, the lateral angles obtuse. Posterolateral

margins rather convexly angulate, the hind margin almost straight.

Scutellum substantially triangular with the lateral margins gently angu-

lately concave, the tip more or Jess rounded. Basally somewhat elevated and

on each side an oblique broad, luteous, impunctate (but granulate) streak

running from the base to about the angle of the lateral margins. Punctations

bordering this streak much darker and denser than elsewhere, particularly

basally.

Hemelytra wider than the body in their basal quarter, this portion of

the corial margin slightly irregular. Behind this point the lateral margins

of the corium smoother and gradually converging posteriorly leaving more

of the connexivum exposed, hind margin of corium more or less straight,

Membrane milky with brownish veins, a number of the latter forked, and

brown maculae between the veins, Laterotergites with their posterior angles

only slightly produced.

Beneath mainly concolorous with the dorsal surface, the punctations

larger and darker on the head and prothorax, Thoracic sterna and last

segment of rostrum piceous or dark brown as is also the base of the third

abdominal segment. Fourth and fifth also may be darker along the incisures.

Male genitalia trom below Fig, 6B,

Length: 10.5-13.5 mm.

Distribution: Queensland, Lectotype) (Reg. No. T4117), unloculized,

numbered 63, donated C, French Jun, 15 November 19/1; | 4, Mulgowie

Well, 8 September 1953, coll. Smith; | &, Rymple, 16 November 1955

(N.M.). South Australia, 22 2, Lake Calabonna, coll, A. Zietz (A.M. &

S.A.M.); |? Innaminka, at light. 18 October 1962, coll. J. Findley; | 4

Trouda Bore (Murnpeowie Station), at light, 24 August 1967, coll. G, F,

Gross: 18,59 2, Cooper Crossing, at light, 12 November 1955, coll, E, T.

Giles, 2 ¢¢, 3 %%, Coopers Crossing, at light, 21 February 1956,

coll. G. F. Gross; I¢, 29 9, Clayton Crossing, at light, 13> November

1955, coll, EB. T, Giles: 2 6 #@, 1 9, Marree, coll. L. Reese; |,

1 ?, Muloorina Station, 18-19 February 1956, coll. G. F. Gross;

Ia, 699, Lake Byre, 22 April 1955, coll. G. F. Gross; large

series, Madigan Gulf area of Luke Eyre, at light, 3-6 November 1955,

coll. E. T. Giles; 1 ¢, 1 2, found dead on salt surface of Lake Eyre (Madi-

gan Gulf), & November 1966, coll. G. F, Gross: | ?, attracted to light,

GROSS—AUSTRALIAN CEPHALOPLATUS 23

° sis.tit8 |— |mm. —

Fig. 6: Ventral (and slighlly posterior) aspect of the male pygophore of A. Cephalo-

platus (C.) fasciatus Distant; B. Cephaloplatus (C.) granulatus Bergroth; (C.)

Cephaloplaus darwini Distant: D. Cephaloplatus (C.) explanatus sp. nov.;

E. Cephaloplathsy (C.) reticulatus Bergroth.

24 REC_ S. AUST, MUS. | (4); 1-58 October, L970

Prescott Point, Lake Eyre, 30 September 1967, coll. G. FP, Gross; 1 @,

Prescott Point on Madigan Gulf Lake Eyre, at light, 22 February 1968,

coll. G. F. Gross; 4 § 4, | %, Wirraminna, 27 October 1953, coll. N. B.

Tindale (S.A.M. ) 3 94,2 9%, Coopers Creek, 12 August 1958, coll,

R, A, Stirton, 2¢ 4, 32 2, Lake Palankarinna, 28 July 1953, coll. R. A.

Stirton; 14, Neales River at Agebuckina, 10 May 1953, coll. R. A.

Stirton & R, H. Tedford . A.) New South Wales. J ¢, Broken Hill,

22 November 1943, coll, C, E. Chadwick (A,M,),

Remarks; An examination of the material in the National Museum,

Melbourne where much of the C. French Jun. collection is to be found or

the collections of the Zoological Institute in Helsinki where a large part of

Beryrowth’s own collection is lodged failed to reveal any of the material

described by Bergroth in this genus marked as types. Drs. Hackman and

Meinander were able to inform) me that on the basis of their experience

with the Mascarene material of the Bergroth collection that the latter had

frequently returned the material he described to whoever had sent it to him,

unlabelled but possibly accompanied by some sort of list, No specimen in

Helsinki could be located to fit Bergroth's description of this species or

having French as a collector on it (although—vide infra—some other

material considered to be types of other Bergroth species was found), How-

ever, u single male specimen in the National Museum of Victoria fits Ber-

groth’s description (even to having only the first antennal segment remaining )

and has 4 Queensland-French indication on the label. [ have every reason

to believe that this specimen is in fact the type of “Cephaloplatys granulatus”

Bergroth described from Queensland from a collection sent to Bergroth by

French. Accordingly this specimen is here selected as the lectotype male of

“Cephaloplatys granulatus” Bergroth,

Although described from Queensland the species appears to be rather

rare there and also in New South Wales. Its main centre of occurrence

uppears to be in the region of the Lake Eyre drainage basin in South

Australia. Many of the specimens to hand were collected at lights.

Cephaloplatus (C.) clementi Distant

Cephaloplatys clementi Distant, 1910, Ann. Mag, nat. Hist,, (8)6:473,

Fig. 7

Average sized for the genus, rather reddish in macroscopic appearance,

the ground colour reddish yellow with numerous brown punctations and

rather sparse small concolorous granules on the hinder part of the pronotum,

scutellum, and the coriaceous parts of the hemelytra.

Head with juga strongly and foliaceously developed, extending well in

front of the anteclypeus, at their widest, however, not wider than the width

of the head across the eyes, Lateral margins of the juga just in front

ie Caato®

aes

& *

ew * 586

Se "he € +

oe Spek ie ae

: ee

Fig. 7: Cephaloplatus CC.) clement Distant,

26 REC. 8S. AUST. MUS. lo (4): 1-58 October, 1970

of the eyes produced into a blunt conyex process which is distinctly concave

above, lateral margins in front of the convexity almost semicircular, forming

a broad sweep to the shortly rounded interior angles of the apices of the

juga, interior murgins of juga more or less straight, separated or not, Disc

of the juga somewhat concave, particularly anteriorly. Base of head between

the ocelli somewhat elevated. Eyes moderately prominent, facets occupying

most of the anterior portion. Antennae slender and five segmented, though

none of the specimens to hand has the fifth segment, First segment not

reaching apices of the juga, third and fourth segments subequal, second

somewhat shorter than either, brownish or reddish brown.

Pronotum only slightly raised posteriorly, declivous anteriorly, antero-

lateral margins progressively more explanate from posteriorly to anteriorly,

regions of the atilerior angles projecting forward us somewhat recurved

processes extending well in front of the eyes and to in front of the convex

process on the lateral margins of the juga. Anterior margin truncate behind

the collum, oblique behind each eye then somewhat convergent to the

apices of the anterior processes. Anterolateral margins slightly convergent

anteriorly, thence nearly straight to the lateral angles bul with a. slight

more or less angular concavity behind the midline of the pronotum, finely

denticulate or crenulate. Luteral angles irregular or irregularly rounded, pos-

terolateral margins somewhat sinuate, posterior margin truncate.

Scutellum very similar in shape to most other Cephaluplatus with the

slight angulate concavity just behind the middle and the tip broadly rounded,

Basally slightly raised, apically more flattened, in each basal angle a somewhat

elongate black fovea inward of which lies a concolorous callous area or

point.

Hemelytra in their basal quarter wider than the abdomen, this margin of

the corium slightly sinuate, not denticulate or erenulate. Behind this

point hemelytra gradually narrowing and leaving more or less half of the

connexivum exposed. Outer and inner parts of the hind margin of the corium

convex, the outer angle of the corium somewhat produced. Membrane

opaque with only slightly darker light brown veins and odd scattered brown

maculae in the cells between the veins, some of the veins forked and some

closed cells at the base of the menibrane. Laterotergites concolorous, not

infuscated, their posterior angles not produced.

Beneath rather darker than the dorsal surface, the area around the

opening of the scent glands brown with concolorous punetations, punctations

on the rest of the thorax aad on the underside of the head course and piceous,

on the abdomen finer, denser, piceous. On the head there is a short piceous

bar in the front of the antennifers and a piceous patch ut the base of the head

behind the bucculac, On the pronotum all thoracic sterna are piceous and

there is a semicircular piceous bar in line with the eyes on the propleuron and

GROSS—AUSTRALIAN CEPHALOPLATUS 27

a shorter one on the metapleuron behind the evaporative area. On the

abdomen there are piceous or infuscated areas as follows; a pair of semi-

circular patches at the base of the abdomen behind each metacoxa, a rather

diffuse lateral line on either side extending from the base of the abdomen

to nearly the apex of the fifth abdominal segment, and a broken line ventrally

made up of basal bars on the fifth, sixth and seyenth abdominal segments.

Rostrum and leys yellowish, the latter with scattered brown raised maculae

on the femora and tibiae. Male pygophore from below similar to fig. 6B,

Length: 10.25-11.5 mm,

Distribution: Western Australia. Holotype @, Nicol Bay District,

coll. Dr. Clement; 1 @, Onslow, donated C. French Jun. 15 November

1911 (BM): 3 @ 32, 7 9%, Onslow, some donated C, French Jun, 15

November 1911, others F. P. Spry, 5 October 1922, (N.M,): | 2%, Onslow;

| ¢, Onslow, donated F, P, Spry, 5 October 1922 (A.M.); 1 ¢, Wittenoom

Gorge, 475 m, 9 October 1962, coll. E. 8. Ross & D. Q. Cavagnaro; 14.

Milly Milly, 600 m, 6 October 1962, coll. E. S. Ross & D, Q. Cavagnaro

(C.A,),

Remarks: This species appears restricted to Western Australia, in

fact possibly only the Central West of Western Australia, It is a ruddy

brown species with a remarkably even colour pattern on the dorsal surface.

It is the first of 4 series of species (or taxa) in the subgenus Cephaloplatus

which have the lateral margins of the head produced into a small but con-

spicuous triangular process just in front of the eyes.

Cephaloplatus (C.) australis Dallas

Cephaloplatus australis Dallas, 1851, List of specs. Hem, Ins, Coll, Brit.

Mus. 1:138,

Fig. &

Average size for the genus, appearing rather greyish and variegated in

macroscopic view. Ground colour yellowish, punctations brown, in some

parts of the pronotum and scutellum rather blackish. Some fine lutcous

granules present on the dorsum particularly in the hind portion of the pro-

notum between the lateral angles, Three fuscous maculae across the base

of the head and one shaped rather like a U on its side around the inner margin

of each callus on the pronotum.

Head with juga foliaceously expanded and rather acutely produced

in front of the anteclypeus like the next taxon (nubifer Bergroth). Lateral

margins of juga immediately in front of the eyes produced into an angulaie

process which is vaguely concave dorsally and directed somewhat upwards,

(his process not so strongly developed as to constitute a spine but is rather

8 REC. $8 AUST. MUS. 16 (4): 1-58 October, 1970

Fig. &: Cephaloplatuy (C.) australis Dallas,

[Mhetagraph by courtesy of Uhe Troaslees of the Mritieh Musemmn (Natural History),

an equilateral triangle. Lateral margins of juga in front of this point broadly

curved to their relatively acute apices, this part of juga not extending much

in width beyond the inner margin of the eyes, Juga apically shortly and

GROSS -AUSTRALIAN CEPHALOPLATUS 2“

roundedly acute, inner margins parallel and relatively widely separated, (con-

tiguous in nubifer). Dise of head slightly raised between the ocelli, behind

each ocellus and medially, on the hind margin a black patch, punctations in

the basal balf of the anteclypeus strongly blackish, behind them a glabrous

patch, Antennae in type represented by only the first three segments, appar-

ently slender, first segment somewhat orange and not reaching the apex of the

jugum, second longer than the third, yellowish in its basal 2/3 then with a

preapical brown ring, third segment blackish brown except basally.

Pronotum rather more raised posteriorly than nuvbifer and anteriorly to

this dechvous und with a shallow transverse depression just behind the

collum. Anterolateral margins progressively more explanate from posteriorly

to anteriorly and rather depressed. The regions of the unterior angles are

produced forward as angular processes which reach about the anterior

iargins of the eyes, Anterior margin strongly rectangular behind the collum

and behind the eyes shortly truncate (ner ebliquely so as in nubifer) and

then diverging to the apical processes, Anteroluteral margins almost straight,

or only very slightly concave anteriorly, a lite behind the middle turning

out rather angulately (this concave angulation much more conspicuous than

in muvbijer) to the shortly and triangulately produced paler lateral angles.

The true lateral angle lies behind this process und is rounded, The antero-

lateral angles in front of the concave angulation are conspicuously (though

the serrations are rather small) serrate. (In wihifer they are virtually entire).

Posterolateral margin sinuate, posterior margin entire, On the dise of the

pronotum anteriorly the inner margins of the calli outlined in black, on the

posterior labe a siniious whitish line made up of more than usually dense

vranules running between the lateral angles, behind this line a similar line

made up of black punctutions, a dense patch of black punctations also in

the anterolateral margins at the angulation,

Scutellum very much as in other species of Cephaloplatus with the

usual slight concavity of the lateral margins just behind the middle.

Anteriorly somewhat raised (in vubifer a lot more strongly raised). In each

basal angle of the seutellum the usual bluck fovea margined on its inner

side by a glabrous point,

The hemelytra wider than the body in their basal quarter, this portion

of the corial margin vearly entire, Behind the basal quarter the lateral

margins of the coria rather strongly converging to leave most of the con-

nexivum exposed, apical margin of the corium ulmost straight, rounded

broadly at each end (in mubifer rather concave medially). Membrane milky

white or opaque with light brown veins and light brown maculae between

the veins. Laterotergites mainly concolorous but with a black spot interiorly

just in front of each incisure and a large Wiangular one exteriorly just behind

each incisure,

au REC. S. AUST, MUS, 16 (4): 1-58 Ocraher, 1970

Beneath concolorous with above, punctalion coarse and evenly distri-

buted, areas more infuscated or blackish are, a patch on the head beneath

behind the bucculae, the thoracic sterna, a short shining bar anteriorly and

exteriorly on the propleuron and the mesopleuron and metapleuron obscurely,

a sinall patch on the lateral margin of each abdominal segment just behind

each incisure and a medial streak in the basal half of the ninth abdominal

segment; the femora and tibiae are coarsely maculated with black, the latter

only exteriorly.

Length: 11 mm. (approx.),

Distribution: Western Australia, Holotype 2 (B.M.)

Remarks: The type does not now bear any indication of provenance

but Dallas in his original description indicated the North West Coast of the

continent,

The status of this taxon and the three which follow (nubifer Bergroth,

pellewensis sp. nov. and bellus sp. nov.) is still rather uncertain. Nubifer

could be an arid region subspecies of australis; it is about the same size

though rather reddish but has many of the same dark markings. The

anterolateral margins of the pronotum are not so incised but this probably

a rather variable character and could be subspecific. Pellewensis on descrip-

tion fits australis rather well although it is conspicuously smaller but could

be the subspecies found in the Gulf of Carpentaria region. Bellus looks very

like pellewensis and is about the same size and could be a subspecies of

australis from Central Australia closest to pellewensis, although it is con-

spicuously reddish it has the same colour patterns as pellewensis, however

the lateral margins of the pronotum are hardly serrate (in pellewensis they

are conspicuously so), and in bellus the triangular lobe in front of the eye

has become a distinct spine like process. The relative lengths of the

antennal segments and their almost identical colouration in the four taxa

would tend to support the same hypothesis as does the structure of the male

pysophore from both dorsal and ventral aspects (the australis type unfor-

tunately is a female). The female external genitalia also appear not to

differ on shape or colour pattern.

However, the above cannot be the correct explanation for mubifer and

bellus coexist in the Alice Springs region of Central Australia.

Alternatively a case could be made out that australis Dallas and nubifer

Bergroth are the Northern and more Southern subspecies of one single

species in the 9.75-11.0 mm. range and that pellewensis sp, nov, and

bellus sp. noy, are subspecies, Northern and Southern respectively of a more

Eastern species in the 8.0-9.5 mm, size range.

GROSS—AUSTRALIAN CEPHALOPLATUS 31

For the moment the only really satisfactory method appears to be to

regard these taxa as distinct species until more material, especially from the

Northern coastal regions of the continent, is available,

Cephaloplatus (C,) nubifer Bergroth

Cephaloplatus nubifer Bergroth. 1916, Proc. R. Soc, Vict., 29:25,

Fig. 9

Average size for the genus, appearing rather reddish and variegated in

macroscopic view, Ground colour reddish yellow or yellow, punctations

brown, tending to be rather finer and sparser on the head and pronotum than

in some of the other species. Some fine concolorous granulations present

on the dorsum particularly in the extreme hind portion of the pronotum, the

hind part of the scutellum and the hind quarter of the coriaceous parts of the

hemelytra.

Head with juga expanded somewhat foliaceously, but rather more

acutely produced in front of the apex of the anteclypeus than in other species

of this genus, Lateral margins of juga immediately in front of the cyes

produced into an angulate process which is concave dorsally, this process is

not so strongly developed as to constitute a spine, it is more in the shape of

an equilateral triangle. Lateral margins of juga in front of this point

broadly curved to their relatively acute apices. this part of juga not extending

much beyond the inner margins of the eyes. Anterior avigles of jugae more

acute than other species, inner margins convex contiguous anteriorly but

leaving a small gap immediately in front of the anteclypeus in the three

specimens availuble. Head slightly raised basally between the ocelli, Eyes

prominent, facets occupying only the anterior facing surfaces, hind surface

flattened and oblique, Antennae relatively long and slender. five segmented,

first Se~ment not reaching the apices of the juga, second, third and fourth

segments subequal, fifth segment somewhat longer, Antennae brownish or

reddish brown in colour, the apical halves of the third. fourth and fifth seg-

ments infuscated.

Pronotum only very slightly raised posteriorly, anteriorly to this some-

what declivous but with a slight transverse convexity between the anterior

margin and the hinder portion. Anterolateral margins progressively more

explanate from posteriorly to anteriorly. The region of the anterior angles

shortly produced as angular processes but reaching only to about the anterior

margins of the eyes. Anterior margin shallowly concave behind the collum,

obliquely truncate behind each eye and diverging to the apices of the

anterior processes. Anteroluteral margins somewhat curved anteriorly

thence more or less straight with perhaps a slight concavity between this

point and the lateral angles, almost entire. with only very fine irregularities,

1970

October,

MUS. If (4):

S. AUST.

REC.

lat’

mm.

Brenpa K Heap

Cephaloplaius (C.) nubifer Bergroth,

Fig. 9;

rs)

GROSS—AUSTRALIAN CEPHALOMLATUS

Lateral angles obliquely truncate, posterolateral margin sinuate, posterior

margin truncate, On the disc of the anterior lobe of the pronotum the

calli relatively conspicuous, on the hind lobe a trace of a sinuous line

running between the lateral angles made up of rather more than usually

densely concentrated granules,

Scutellum very much as in other species of Cephaloplatus with the

usual slight concavity of the lateral margins just behind the middle,

Anteriorly somewhat raised, this anterior portion continuing as a slightly

raised area to the tip thus giving the impression of a flask shaped raised area

on the scutellum. In each basal angle of the scutellum the usual rather

elongate black fovea margined on its inner side by a concolorous glabrous

area Or point,

The hemelytra wider than the body in their basal quarter, this portion

of the corial margin nearly entire. Behind this point the lateral margin of

the corium rather more strongly converging posteriorly than in some other

species and leaving most of the connexivum exposed, apical margin of

corium slightly concave exteriorly, broadly rounded interiorly, the outer angle

of the corium very slightly prominent. Membrane milky while or opaque

with almost concolorous veins and some light brown maculae in the cells

between the veins. veins apparently not forked apically. Laterotergites

concolorous but with an infuscation on each just behind the incisure.

Underside concolorous with above, punctations comparatively sparse

und course. On the underside of the head a piceous short bar just in front

of the insertion of the antennae and a piceous spot on either side of the base

of the head just behind the bucculae, in some specimens the bucculae infus-

cated, in others not, Thoracic sterna piceous, a piceous somewhat irregular

line present near the exterior of the propleuron and thinner and shorter

lines present in much the same position on the metapleuron and mesopleuron,

Abdomen at base With a small semicircular piceous area behind each meta-

coxa, Incisures somewhat darkened and a distinct fuscous spot exteriorly

immediately behind where each reaches the lateral margin. On the venter of

the abdomen faint traces of an interrupted series of fuscous spots or short

small bars. Male genitalia from below similar to Fig. 6B.

Length: 9.75-10.0 mm.

Diswibution: Northern Territory, Lectearype &, Medonnell Ranges;

1%, Alice Springs (Helsinki); | @, Alice Springs. 31 August 1936, coll.

H, O, Fletcher & W. Barnes; | 5, 1 2, 14 miles North of Tyon Station,

August 1936, coll, H. O, Fletcher & W. Barnes (A.M.): | @, Hermannsburg,

1908, coll, Ho J. Hillier (B.M.), Western Australia. 1, Hermite Is., coll.

H.M.S. Campania, accessed 1952 (B.M.): | 4, Fitzroy Crossing, 4 Septem-

ber 1953, coll N. B. Tindale: | @. Derby, coll. W. D. Dodd. (S.A.M.).

Unlocalized, | @ (N.ML),

34 REC, §. AUST. MUS. I@ (4); 1-58 Ocraber, \970

Remarks: Bergroth in his original description of subifer mentions

two localities “Mcdonnell Ranges” and “Alice Springs” the material being

collected on the Horn Expedition into Central Australia, In Helsinki | was

able to find a section of Horn Expedition material which Bergroth had

worked over but no types were marked. Amongst it were a male from

Medonnell Ranges and one other from Alice Springs of a species of

Cephaloplatus which fit the description of nubifer. The Mcdonnell Range

male fits the description very well and there can be no doubt that this is the

type, this specimen is accordingly selected here as the lectotype of Cephalo-

platus nubifer Bergroth,

Cephaloplatus nubifer appears to be a fairly rare species probably

ditributed over a fairly narrow belt of arid country stretching from about

Alice Springs to approximately the same latitude on the Western Australian

coast and off shore islands,

The possibility that nubifer is a subspecies of australis Dallas is discussed

in the remarks section under the latter.

Cephaloplatus (C.) pellewensis sp. nov.

Fig. 10

A little smaller than the average size for the genus, usually appearing

greyish and variegated in macroscopic view. The ground coloration is

actually rather whitish with coarse brownish black punctations, many of

them forming clusters, The hind part of the pronotum, scutellum and the

coriaceous parts of the hemelytra with sparse concolorous small granules

Head with juga strongly and foliaceously developed, produced well in

front of the apex of the anteclypeus, Anterior margins of juga oblique, then

rounded smoothly into the lateral margins of the head which are produced

into a prominent slightly curved triangular process in front of each eye, inner

inargins straight or curved, contiguous or not. Eyes moderately prominent,

facets occupying ull but the posterior flattened surfaces. Antennac moderately

slender, first segment not reaching apex of head, third somewhat shorter

than second; second, fourth und fifth subequal, yellowish, on the second

segment a subapical darkened ring, apical two thirds of third, fourth, and

fifth segments darkened.

Pronotum somewhat raised posteriorly, more depressed anterolaterally,

the anterolateral margins progressively more explanate from the lateral angles

forward, the region of the anterior angles produced forward as a triangular

process to about the level of the convex spine in front of the eyes. Anterior

margin of pronotum truncate behind the collum and again behind each eye

then diverging forward to the apex of each anterior process, Anterolateral

margins at first rather convex then concave, distinctly finely serrate. Lateral

PHALOPLATUS

GROSS—AUSTRALIAN C

‘

Aha

BRenorm K. HEAD

Cephaloplats (C.) pellewensis sp. nov.

Fig. 10

36 REC. S. AUST, MUS, I (4)! 1-58 October, VI

angles distinctly prominent as small triangular flattened lobes, posterolateral

margin strongly sinuate, posterior margin truncate. The hind quarter of

the pronotum with moderately sparse small concolorous granules. Just in

front of these a sinuous thinnish line of more than usually concentrated

blackish punctations and in front of this u broad sinuous transverse bar, in

front of this again rather darker with odd patches of denser punctations,

especially at the inner edges of the calli, punctations on the expanded

anterolateral margins quile sparse.

Scutellum substantially triangular with the usual angulate concavity

somewhat behind the middle. Basal half somewhat raised, apex depressed,

Dise only moderately densely covered with small granules, in each apical

angle an elongate black fovea margined interiorly by a small glabrous pateh,

another glabrous patch medially on the base, a paler less punctate broad

longitudinal streak medially, lateral areas with symmetrically placed darkish

areas of more densely concentrated punctations. Tip rounded.

Hemelytra wider than the body in their basal quarter, behind this point

converging rather strongly to leave the more posterior laterotergites com-

pletely exposed, whole corial exterior margin rather irregular. Hind margin

of corium straight exteriorly, broadly curved at its inner angle, On the dise

of the corium patches of impunctate areas and areas of dense punctation.

Membrane whitish hyaline with light brownish veins and a number of brown

maculae in the cells between the veins (which do not appear to be forked).

Laterotergites with a brown patch at each posterior inner angle and each

anterior outer angle.

Beneath mainly concolorous with the dorsal surface, punctations on the

metapleura rather finer, Strongly infuscated areas are present to a greater

or lesser degree, in the darkest specimens they occur as follows, a narrow line

on the side of the head imnrediately in front of the insertion of the antennae,

a large spot at the base of the buccalae, all thoracic sternites, lateral bars on

the thoracic pleura, on the third abdominal sternite immediately behind each

metacoxu, most of the abdominal incisures, medially at the base of at least

the sixth and seventh sternites, exteriorly at the anterior angle of cach sternite

and on the apices of the first valvifers of the female genitalia, Male pygo-

phore from below similar to fig. 6B. Legs with coarse brownish maculations.

Length: §8.5-9.5 mm.

Distribution: Northern Territory, Holotype 6 (T6908) Horn Islet, Pellew

Group, 16-23 May 1968, coll. B. Cantrell (Q.M.); allotype 2? paraivpe &,

paratype °, Horn Islet, Sir Edward Pellew Group, 25-31 January 1968, coll,

B. Cantrell; | ¢ doubtful, Horn Islet, Sir Edward Pellew Group, 22-28

February 1968, coll. B. Cantrell; | ¢ doubtful, Horn Islet, Sir Edward

Pellew Group, 15-2] February 1968, coll. B. Cantrell (U.Q.).

GROSS—AUSTRALIAN CEPHALOPLATUS 37

Remarks: The possibility of a subspecific relationship existing between

pellewensis and either australis Dallas or belluy sp. nov, is discussed in the

remarks section under australis (p. 30). The last two males mentioned

under the distribution preceding above are only provisionally referred to

this species for the moment. They are very similar to each other but show

a rather different facies to the type series of pellewensis, They are both in

the same size range as pellewensis, have the triangular lobe in front of the

eyes, the sume structure and coloration of the antennae and the male and

female genitalia appear to be identical and the same general colour pattern.

However, they are both very much smoother, the anterolateral margins of the

pronotum are Virtually straight and they lack all but a few of the dense

patches of black punctations above. They may represent an extreme of the

variation to be found in pellewensis or alternatively yet another species in

the complex from australiy through to bellus, It is worth noting in the

latter context that three other species of Cephaloplatus also occur on Horn

Isict, namely C, (M.) miner Distant C, (C.) pallipes Walker and C, (C.)

pertyi (White),

Cephaloplatus (C.) bellus sp, nov.

Fig. 11

A little smaller than the average size for the genus, often appearing

rather reddish and variegated in macroscopic view, The variegation in some

parts is lutcous or even almost whitish, in other parts reddish, in other parts

pinkish or even ferrugincous. Punctations in the main brownish but small

patches on the head, pronotum and scutellum blackish. The hind parts of

the pronoetum, scutellum and the coriaceous parts of the hemelytra with

numerous concolorous small granules.

Head with juga strongly and foliaceously developed, produced well in

front of the apex of the anteclypeus. Anterior margins of the juga oblique,

turning shortly into the sinuous lateral margins which are produced into a

prominent slightly curved elongate triangular or spinous process in front

ol cach eye, inner margins of the juga curved, sometimes contiguous, some-

limes not, Eyes moderately prominent. facets occupying only the anterior

half, Antennae moderately slender, first segment not reaching the apex of

the head, third segment relatively short, second, fourth, and fifth segments

subequal, on the second segment a subapical darkened ring, apical two thirds

of third segment infuscated, apical half of fourth infuscated, apical two thirds

of fifth infuseated.

Pronotum somewhat raised posteriorly, more depressed anterolaterally,

the anterolateral margins progressively more explanate from the lateral

angles forward, the region of the anterior angles produced as u triangular

process forward to ubout the level of the apex of the spine in front of the eves.

Cepluloplatus (Co) bellus sp. nov.

uid

Fig.

GROSS—AUSTRALIAN CEPHALOPLATUS 399

Anterior margin of pronotum concave behind the collum, truncate behind

each eye, then parallel forward to the apex of each anterior process, Antero-

lateral margins almost straight or very slightly convex, irregular but definitely

not crenulate or serrate. Lateral angles shortly truncate, posterolateral mar-

gin somewhat sinuate, posterior margin truncate, Hind area of the pronotum

fairly densely covered with small concolorous granules. The areas of darker

punctations lending to Vary in position from specimen to specimen but always

syinmetrically placed on either side of the midline,

Scutellum substantially triangular with the usual angulate concavity well

behind the middle. Basal half somewhat raised, apical half somewhat

depressed. Dise fairly densely covered with small granules, in each apical

angle a somewhat elongate black fovea margined interiorly by a small

glabrous area, Patches of dark punctations on the scutellum likewise ditfer-

ently placed in different specimens but tending to be symmetrical about the

middle. Tip broadly rounded.

Hemelytra wider than the body in their basal quarter, this portion of

the corial margin smooth, Behind this point the lateral margins of the

corium smoother and gradually converging posteriorly leaving more of the

connexivum exposed, hind margin of corium more or less straight extertorly,

broadly curved at its inner angle. On the disc of the corium and clavus

various patches of concolorous impunctate areas. Membrane milky white

with light brownish veins and a number of brown maculae in the cells

between the veins, some of the veins forked. Laterotergites with a brown

patch at each anterior angle, this brown patch sometimes extending inwards

further towards the inner margin of the connexivum,

Bencath mainly concolorous with the dorsal surface. the punctations

tending to be finer only on the metapleura. Strongly intuseated areas are

present to a greater or lesser degree; in the darkest specimens they occur

us follows: on the sides of the head immediately in front of the insertions

of the antennae, a spot on either side of the basal ends of the bucculac, all

thoracic sternites, Jateral bars on the thoracic pleura, and the meisures of

ull abdominal sternites, In other specimens there ure only the thoracic

sternites, a diffuse lateral line on either side of the abdomen and a broken line

of infuscations along the midline of the venter. Legs with brownish granula-

tans or maculations. Male pygophore somewhat like fig. 6A.

Length: 9.5-10.0 mm

Distribution: Northern Territory. Holotype & (Reg. No. 120,549)

2 paratype ° 9, | paratype ? sex (as abdomen is missing) (Reg. Nos,

120,550-2), Haast Bluff Station, 2,000 feet 69> Fahrenheit at Mercury

Vapor Light, 4 September 1957, coll, N. B. Tindale; 1 paratype & , 3 paratype

® & (Reg. Nos. 120,553-6) Haast Bluft Station, 2,000 feet, 62° Fahrenheit

at Mercury Vapor Light, 5 September 1957, coll. N. B. Tindale: 2 paratype

40 REC. S. AUST. MUS. 16 (4): 1-58 Ovlober, 1970

® @ (Reg. Nos, 120,557-8), near Ayers Rock, 20 August 1960, coll. R.

Garnetl; paratype @ (Reg. No. 120,562), Yuendemu, February 1968; 1 para-

type ¢,2 paratype ® & (Reg. Nos, 120,569-71), Yuendemu, February 1968,

coll. C. Watts; paratype § (Reg, No, 120,563), Devil’s Marbles 7 miles North

of Wauchope, 22 April 1966, coll. N. McFarland (S.A.M.); alletype 2 2

paratype & 2 (1 at light) 4 paratype 2? @ (2 at light), Stanley Chasm,

31 miles West of Alice Springs, 9 February 1966, coll. Britton, Upton &

Melnnes; | paratype &, 48 miles WSW of Alice Springs, 10 February 1966,

coll, Britton, Upton & MelInnes; 2 paratype § 4, near Reedy Rockhole,

Amadeus Basin at 24°20°S 131 '35’E, 6 June & 8 September 1962, coll.

R, Ranford; | paratype ?, Amadeus Basin, 15-16 September 1962, coll.

R. Ranford (A.N.LC.); 1 paratype 2%. Mt. Gillen near Alice Springs,

1,500 feet, 31 August 1936, coll, H, O, Fletcher & W. Barnes (A.M.);

| paratype é, 11 miles North of Alice Springs 625 m, at 15 watt ultra violet

(black) light, 28 October 1962, coll. E. S. Ross & D. Q. Cavagnaro,

| paratype 2, Devil's Marbles, 350 m, 27 October 1962, coll. EB, S, Ross &

D. QO, Cavagnaro (C.A.). Western Australia. 2 paratype ° @ (Reg. Nos.

120,560-1), Pilgangoora Well, Pilbara District, 6 & 7 June 1953, coll,

N. B. Tindale; 1 paratype @ (Reg. No, 120,559), Coolgardic (S,A.M.).

Queensland. 1 paratype é&, Split Rock 30 miles East of Camooweal,

September 1936, coll, H, O, Fletcher & W, Barnes (A.M.), South Avstralia.

| paratype $ and | paratype °, (Reg. Nos. 120,564-5), found dead on

salt of Lake Eyre North (Madigan Gulf), 27 October and 4 November

1963, coll, G, F, Gross (S,A,M,),

Remarks: The possible relationship of this species with the series

beginning with C. (C.) australis is discussed on page 30. Its centre of

distribution is clearly in the southern part of the Northern Territory extending

out East and West into Oucensland and Western Australia and South into

South Australia.

It is the first of a series of species in which the lateral margin of the head

is produced into a distinct spine like process just in front of each eye.

Cephaloplatus (C.) spurcatus Walker

Cephaloplatus spurcatus Walker, 1867 Cat. spec. Het.-Hem. Coll. Brit. Mus..

119d,

Cephaloplatys spurcatus Van Duzee, 1905, Bull. Amer, Mus. nat, Hist.,

21:201. - .

Fig, 12

Average size of the genus, in macroscopic view tending to appear grey,

light or dark. Ground colour light yellowish or luteous with numerous

coarse brown punctations which are not evenly disposed over the dorsum

of the animal. Pronotum, scutellum and coriaceous parts of the hemelytra

covered with sparse luteous granules.

GROSS—AUSTRALIAN CEPHALOPLATUS 4)

Smm.

Fig. 12: Cephaloplatus (C.) spurcatus Walker.

42 REC. §& AUST, MUS. [6 (4): 1-58 Oclober, 1970

Head with juga rather foliaceously developed and produced forward

in front of the apex of the anteclypeus, their apices broadly rounded, running

back to a prominent flattened spine between the base of the antennifer and

the eye which projects forwards and outwards to about the level of the outer

margin of the eye, Just in front of this spine the lateral margins of the

juga strongly incised, their inner margins in front of the apex of the uante-

clypeus rather convex, contiguous or not. Lateral and anterior margins of

juga somewhat raised so that the anterior portion of the head is somewhat

concave, Eyes moderately prominent, facets occupying only the anterior

half to two thirds. Antennae moderately slender, first segment not reaching

the apices of the juga, second and third segments subequal, fourth about

30% longer than them and filth about 25° longer than fourth, Distal

antennal segments brownish with segments basally yellowish, the first

segment yellowish and sometimes also the second,

Pronotum only somewhat raised posteriorly, depressed atiterolaterally.

the anterolateral margins progressively more explanate from the lateral angles

forward, the region of the anterior angles produced anteriorly us a triangular

process to about the level of the apex of the spine in front of the eyes,

Anterior margin of pronotum truncate behind collum, rather oblique behind

each eye, rounded, straight or oblique to the apex of each anterior process,

Anterolateral margins feebly to strongly angulately concave a little behind the

middle, clearly crenulate or serrate in front of the angle of the concavity,

the crenulations or serrations all the sume length, lateral angles subacute

(strictly speaking the acute projection lies a little in front of the true lateral

angles which are obtusangulately rounded), posterolateral margin almost

straight behind the true lateral angles, posterior margin straight to rather

concave, Luteous granules on the pronotum rather sparse and mainly con-

fined to the posterior half. Punctations quite sparse on the expanded antero-

lateral margins, on the disc sometimes more concentrated in a sinuous line

extending between the luteral angles,

Scutellum substantially triangular with the usual angulate concavity

behind the middle, Basal half raised only medially, apical angle very broadly

rounded, Disc with scattered small Juteous granules, punctations not evenly

disposed, in each apical angle a black fovea margined interiorly by a small

glabrous point,

Hemelytra wider than the body ih their basal third, this part of the

corial margin clearly crenulate. Behind this point the lateral margins of

the corium smoother and gradually converging posteriorly leaving about half

or more of the connexivum exposed, hind margin of corium extertorly rather

concave, rounded at its inner angle. On the dise of the coriaceous portion

of the hemelytra small scattered luteous granules and the punctations not

evenly disposed. Membrane a very pale brown with light brown veins and

GROSS—AUSTRALIAN CEPHALOPLATUS 44

some faint brown maculae in the cells between the veins, some of the veins

forked and near the base of the membrane some closed cells. Laterotergites

concolorous and punctate only interiorly, black in each anterior angle and

along their hind margins, their hind angles a little protuberant.

Beneath much darker than the dorsal surface, the punctuation much

denser, the juga, the explanate anterolateral margins of the pronotum, the

episterna and epimera, the embolium and the sides of the abdomen pale with

relatively Few punctations. A large brownish black spot on the head beneath

at the base of the bucculae, in some specimens a line on the side of the head

in front ef the antennifers, the thoracic sterna, a smooth somewhat curved

longitudinal line exteriorly on the propleuron, another on the metapleuron,

a spot near the lateral angles, a large area at the base of the abdomen at each

side behind each metacoxa, a longitudinal broad line on either side of the

abdomen on segments TL-V and a medial spot at the base of segments II-VI

all blackish or piccous. The anterior half of the lateral margin of cach

abdominal segment and u spot near the edge of each incisure immediately

in front of the former blackish, Femora and tibiae with course brown or

piceous maculae, sparsely distributed, Male pygophore with apical angles

prominent, otherwise similar in appearance to fig 6C,

Length: 8.5-11.0) mm.

Distribution: North Australia. Holotype 4, 4a ete. (B.M.);

29 ¢, coll, Diimel (Stockholm). Queensland. | 9. Brisbane, 2 December

1961, coll, A. J. Hiley; | ?, Brsbane, 10 March 1962, coll. D. Schoorl;

1 &. Toowoomba, 3 February 1963. coll, J. C, Cardale, | ¢, 2 2 2,

Carnarvon, 29 May 1959, coll, T. E, Woodward; |) °. Sunnybank,

6 February 1966, coll, F, R. Wylie: 1 @, Mt. Elliot via Townsville, 24 May

1968, coll, G, Monteith; | @, Mt. Etna, 29 April 1967, coll. J. Bluhdorn;

1 9. Mt. Isa, 23 January 1968, coll. B, Cantrell; | ¢, Highvale, 28 May

1958, coll. T, BE, Woodward (U.QO.); 1 42, 3 29, Clermont, April and

December 1928, coll. Dr. K. BK. a sats 24 4, 12. Almaden, March 1928

& 1929, coll. W. D. Campbell} | &. Gayndah, coll, Masters (A.M.); 1 6,

Brisbane, 15 March 1908, coll. Ww. W, Frog pgatt, 1 2,50 miles SW aay 9

October 1950, coll. B, FP. Riek (A.N.LC.1; | 2, Toowong, 3 May 1908,

(N.M.); 14, 1%, Bowen, coll, A, Simpson; 1 ® , Normanton, coll. R. Kemp:

12, Maryborough, coll. BE. W. Fischer (S.A.M.): 12, Brisbane, 23. Novem-

ber 1915, coll, H, Hacker; | @, aR i July 1926; | 4, Brisbane, 6

May 1937, coll. R. F. Langdon (B.M.); » Rockhampton (Stockholm):

2 24,2 9%, Townsville, January ‘on6: é Malkin ee SN. M.); 1 ¢,

32 Km South ot Ayr, 6 November 1964, coll J. Sedlacek; , 8&8 Km East

Charters Towers, 18 January, 1969, coll. J. Sealacek (Bishop).

Northern Territory. 5 4 4. 10 2 @, Borroloola, November 1911, coll.

G. F. Hill on Barclay Expedition (N.M.), 1¢, | 9. Borroloola, 2& February

44 REC. So AUST. MUS. 16 (4): 1-58 Ovtaher, 1970

1968, coll, B. Cantrell (U.Q.). New South Wales, 1 &, (BL.M,); 1 8,

from E. P. Yan Duzee collection (€.A.). Unlocalized, 2 29,17 (A.M.).

Remarks: This species is very close to the one following (darwins),

The salient features distinguishing the two and a possible subspecies relation-

ship between them is discussed under the remarks section of the latter.

C. (C.) spurcatus occurs over most of Eastern Queensland except the Cape

York areas, part of the Gulf of Carpentaria region of both Queensland and

the Northern Territory, and some part of New South Wales, possibly the

North East Coast of that state. [ts distribution therefore, hes in the centre

then East and South of the darwini distribution with areas of probable con-

tact with the range of the latter across the base of Cape York Peninsula, near

the Queensland-Northern Territory border, and in the region of the southern

section of the shores of the Gulf of Carpentaria.

Cephaloplatus (C.) darwini Distant,

Cephaloplatus spurcatus (non Walker 1867) Carpenter, 1891, Sei. Proc.

R. Dublin Soc, 7:139,.

Cephaloplatus darwini Distant 1910, Ann. Mag. nat, Hist., (8)6:473,

Fig. 13, 6C,

Rather variable in size but large specimens average sized for the

genus, if macroscopic view tending to appear rather greyish. Ground

colour light yellowish or luteous with fumerous moderately coarse brown

punctations which are not evenly disposed over the dorsum of the animal.

Pronotum, scutellum and coriaceous parts of the hemalytra covered with

sparse luteous yranulations,

Head with juga rather foliaceously developed and produced forward

in front of the apex of the anteclypeus, their apices oblique, then broadly

rounded just in front of the antennifer then running back to a prominent

spine between the base of the antennifer and the eye which projects outwards

and forwards to about the level of the outer margin of the eye. In front of

this spine lateral margins of the juga strongly incised, their inner margins in

front of the apex of the anteclypeus rather straight, usually well separated

from each other and in many cases divaricate, the apex of the anteclypeus

therefore free. Lateral and anterior margins of juga somewhat raised so that

the anterior portion of the head is rather concave, Eyes moderately promi-

nent, facets occupying only the anterior half. Antennae moderately slender,

first segment not reaching the apices of the juga, the second and fourth

segments subequal, the fifth segment longer than either, the third segment

shorter than either. In some specimens the antennae in the main reddish, in

others brownish or brownish with the apical halves of the third, fourth and

fifth segments infuscated,

GROSS—AUSTRALIAN CEPHALOPLATUS 45

L a /

BRENDA K. HEAo

Fig. 13; Head and pronotum of Cephaloplatus (C.) darwini Distant.

Pronotum only somewhat raised posteriorly. depressed anterolaterally,

the anterolateral margins progressively more explanate from the lateral

angles forward, the region of the anterior angles produced as a triangular

process forward to about the level of the apex of the spine in front of the

46 REC. 8. AUST, MUS. 16 (4)+ 1-58 October, L970

eyes, Anterior margin of pronotum truncate behind the collum, truncate

behind each eye. then diverging somewhat forward to the apex of each

anterior process. Anterolateral margins strongly angulately concave a

little behind the middle, with four to seven very marked denticulations in

front of the angle of this concavity, in front of the lateral angles subacutely

produced, true lateral angles rounded, posterolateral margin at first rather

convex then obliquely concave, posterior margin truncate. Luteous granules

on the pronotum yery sparse and mainly confined to the posterior half.

Between the lateral angle is a vestige of a sinuous transverse line made

Up of rather more than usually densely concentrated luteous granules.

Scutellum substantially triangular with the usual concavity somewhat

behind the middle. Basal half somewhat raised, apical half somewhat

depressed, apical angle rounded and narrower than that of ©. (C.) spurcatus.

Disc with scattered small luteous granules, in each apical angle a somewhat

elongate black fovea margined interiorly by a small glabrous area or point,

another glabrous point medially on the base.

Hemelytra wider than the body in their basal quarter, this portion of

the corial margin finely denticulate. Behind this point the lateral margins

of the corium smoother and gradually converging posteriorly leaving

about half of the connexivum exposed, hind margin of corium exteriorly

inore or less straight, broadly curved at its inner angle. On the

disc of the coriaceous parts of the hemelytra small scattered luteous granules

und the punctations not disposed evenly. Membrane opaque with light

brownish veins and a number of light brown maculae in the cells between the

veins, some of the veins forked and near the base of the membrane some

closed cells. Laterotergites concolorous, generally only lightly infuscated at

the incisures, their hind angles slightly protruding, lightly punctate and that

only in their inner portion,

Beneath tending to be darker and more reddish than the dorsal surface,

the punctations on the underside of the head and on the thorax coarse and

dense, Piccous are: a longitudinal streak in front of each antennifer, a large

spot at the base of the bucculae, the thoracic sterna (with the exception of the

yellowish keel on the mesosternum), sundry darker patches above the epi-

meron and episternum, two lurge semi-circular areas at the base of the

abdomen behind each metacoxa, an interrupted line on the abdomen made

up of triangular median bars on the fourth, fifth, sixth and seventh ventrites,

a broad to very broad lateral band on either side extending from the base of

the abdomen to the apex of the fifth ventral segment. All fernora and tibiae

with course brown or piceous raised coarse maculae, fairly sparse in distribu-

tion, these spots not present on the Murray Island specimen. Male pygopore

with apical angles prominent (Fig, 6C),

Length: &.011,0 1m

GROSS—AUSTRALIAN CEPHALOPLATUS 47

Distribution: Northern Territory. Holotype 9, Port Darwin, coll.

Walker: | ¢, Darwin 7 April 1913, coll, G. F. Hill (B.M.); 1 ?. Darwin,

January, 1939, coll, M. Kamper; | 2, 1 9, 1 ?, Burnside, 23 March 1929,

coll, 'T, G. Campbell (A.M.); 12%. same data; | @, Brocks Creek, Mareh

1933, coll. T. Campbell (A.N.-C.); 1 4, 1 &, Darwin, coll. G. F. Hill;

222, 30 miles East of Darwin, coll. G. F. Hill; 14, Croker Island

Mission, 28 March-5 April 1961; 1 4. Oenpelli, December 1918, coll. D. P.

Cahill (N.M.); 14. 19, Darwin, 12 March 1913, coll. G. F. Hill; Le,

49 9, Darwin, coll, G. F. Hill; 1 2, Darwin: 1 ¢, Darwin March 1956, coll,

D. Groves; |} 2, 3 9 9°, Darwin by sweeping in cleared monsoon forest,

7 January 1961, coll. G. F. Gross; | @, Melu Island near Darwin, coll.

G. F, Hill; 2 4 6, @ @ 2, 30 miles Bast of Darwin, coll. G. F. Hill: | 2,

Batchelor, coll. GF. Hill, | &, Roper River, coll. N. B. Tindale (S.A.M.):

2 ¢¢,2 22, Darwin, 11 February 1945, coll, B, Malking 3 4 6,398,

Darwin, 25 March 1945, coll. B. Malkin; | ¢, Batchelor, coll, G. F. Hill

(U.S.N.M.): 19,22 2, 3-7 December 1963, 2¢ ¢.1 2, 1-8 January 1964,

Darwin, at m.v. light trap, coll. J, Sedlacek; 2 ¢ 4, 1 2@, Darwin, coll. J.

Sedlacek; 2% @, Berry Springs, 50 Km South-East of Darwin, 12 March

1966, coll. J. L. & M. Gressitt (Bishop). Seu Australia, 1 /, donated

C, French Jun., 15 November 1911 (N.M,). (The Northern Territory was

formerly part of South Australia politically and it is reasonably certain that

this specimen would have come from what is now a Northern Territory

locality.) Western Australia. 19, Wyndham, 15 December 1930-8 Jan-

vary [931.24 6, Chumooli near Wyndham, 18% January 1930-12 February

1931, coll H, J. Willings (A.N.LC.); 1, Junction of Fitzroy and

Marguret Rivers, 1896, coll, Calvert Expedition; 12, Derby (S.A.M.),

Queenslund. 2% 9, Lockerbie, Cape York Pen., LO-15 June, 1969, coll,

G,. B. Monteith. | @, Homestead. Silver Plains via Coen, 11 December, 1964,

coll, G, Monteith; | ?, Bamaga, Cape York Peninsula, May 1968, coll. R.

Trundle (U.0.);2 ¢@¢, 1 %, Bamaga, Cape York Peninsula, 5 April 1964,

coll, I, F, B, Common & M, S, Upton (A,N.L-C,); 2 ¢ 4, Stewart River, Jan-

uary-Pebruury 1927, coll. Hale & Tindale (S.A.M.); 1 &@. Weipa, 15 June

1969, coll. C. MeCubbin (N.M.): 12%, Rocky Scrub, Mcilwraith Range,

Cape York Pen., 28 June 1932, coll, Darlington on Harvard Expd.

(A.M.N,H.), Yorres Strait Islands. 14. Murray Island (A.M.); 32 #,

Murray Island, coll, A. M. Lea (S.A.M.): | neanide, Thursday Island,

5 June 1969, coll, A, Neboiss! 19. Prince of Wales Island, 29 May 1969,

coll. A. Neboiss (N.M.); 12. Prince of Wales Island, 27 January 1939,

coll, R. G. Wind (C.A.).

Remarks: “Chis species is to be found in North Western Australia, north

of approximately the latitude of the Fitzroy River, in the Darwin and

Arnhem Land regions of the Northern Territory, in the Northern part of the

Cape Yorke Peninsula and Torres Straight Islands areas of North Queenslind.

48 REC. 8. AUST, MUS. 16 (4): 1-58 Oclober, 1970

The Northern Territory and Queensland populations appear to be separated

by an extension of C. (C.) spurcatus which runs up through Mt. Isa to the

south coast of the Gulf of Carpentaria. ©. (C,) spurcatus and C, (C.)

darwini are very close to each other in general appearance, distinctions are

(1) in spurcatus the anterolateral margins of the pronotum are fairly finely

crenulate with many small teeth, in darwini this margin is coarsely serrate

with about 4-7 much larger teeth, the latter of unequal length and some are

doubled. (2) the second segment of the antennae is very nearly the same

length as the third (15:15) in spurcatus whereas in darwini the second is

usually about 30% longer than the third (20:15) although some variation

does exist. (3) the dark bands on the sides of the abdomen in spurcatus

tend to be narrow but very broad in darwini. The male pygophore is appar-

ently virtually the same in both species.

It could be argued that darwini is u subspecies of spurcatus. Evidence

to the contrary is that darwini is made up of two populations west and east

divided by an extension of spurcatus (which is hardly to be expected of

subspecies) and that transitional forms are very rare, only the Derby and

one or two (not all) of the Murray [sland specimens of darwini approach

spurcatus, these being from localities quite remote from any areas of possible

spurcatus-darwini contact or overlap whereas if they were subspecies these

forms would be expected ulong the zones of contact, ie., the region of the

Gulf of Carpentaria. As far as can be determined from the material at hand

the two species ure allopatric, It could also be argued that darwini is an

impressed variation of spurcatus to be found in hotter and/or more humid

areas. It can hardly be the more humid factor as many of the localities for

spurcatus are from quite wet areas of Eastern Queensland, On the evidence

presented here the series of spurcafus referred to by Curpenter from the

Torres Strait region are more likely to be darwini, unfortunately I did not

see this series when in Europe.

Cephaloplatus (C.) pertyi (White)

Dryptocephalus? (Cephaloplatis) pertyi White, 1842, Trans. ent. Soc, Lond.,

3:90.

Cephaloplatus pertyi Dallas, 1851. List spec. Hem. Ins. coll. Brit. Mus.,

1:148.

Cephaloplarys pertyi Distant, 1899, Ann, Mag, nat, Hist., (7)4:433.

Fig. 14

Average sized for the genus, light yellowish or luteous with light brown

punctations, sometimes with a reddish tinge. Dorsal surface with scattered

granulations,

GROSS—AUSTRALIAN CEPHALOPLATUS 49

Smm.

Vv. M. KITSON

Fig. 14: Cethaloplatus (C\) pertyi (While),

st REC. S, AUST, MUS. 16 (4): 1-58 October, 170

Head with juga developed into foliaceous structures though these at

their widest do not extend outwards past the level of the eyes. Lateral

margins of the expanded juga slightly concave and convexly flexed laterally,

then convexly rounded in turn becoming the relatively long oblique anterior

margins, these anterior angles shortly rounded and the inner margins of the

produced portion more or Jess straight and well separated from each other,

though not divaricate, Eyes not very prominent and the facets occupying

only the anterior half of the optic process which is inclined posteriorly.

Antennae slender and five segmented, the fifth segment the longest. the first

segment not reaching the apex of the jugum, yellowish but the third and

fourth segments in their apical half and the whole of the fifth somewhat

infuscated.

Pronotum yery moderately convex on the disc, the lateral margins

strongly explanately produced and the region of the anterior angles produced

forward as a somewhat triangular process to almost the apex of the dorsal

aspect of the anteclypeus. Anterior margin of the pronotum convexly

excavate, anterolateral margios coarsely dentate or serrate, lateral angles

irregularly rounded, posterolateral margins almost straight with a prominent

but small tumescence just inward of the lateral angles, hind margin nearly

straight.

Scutellum substaotially triangular, the lateral margins rather angulately

concaye, the maximum concavity occurring behind the mid point, apex

narrowly rounded, the dise basally somewhat raised.

Hemelytra wider than the body in their basal quarter, this portion of the

corial margin denticulate Or coarsely crenulate. Behind this point the lateral

margins of the corium smooth and gradually converging posteriorly leaving

progressively more of the connexivum exposed, hind marvin of corium more

or less straight. Membrane greyish hyaline with brown, mostly longitudinal,

veins and some brown spots between them, Laterotergites with their

posterior angles rather angulate and appearing to be somewhat produced.

Beneath mostly concolorous with the upper surface, punctations an the

head and thorax coarser and sparser, those on the abdomen finer and

denser, Mesosternum, metasternum, two patches at the base of the abdomen,

one just behind each metacoxa, and a narrowing Jateral band on the first few

visible abdominal sternites infuscated. Male pygophore similar to fig, 6C,

Length: 9.75-11.0 mm,

Distribution: Unlocalized. Holotype 2 (B.M.): 1 9, trom C, French

Jun. donated 15 November 1911 (N,M.):; 1 2 (S,A.M,). Northern

Territory, 1 %, 25-31 January 1968.2 ¢ 6, 4 2%, 1-7 February 1968,

1 2, 8-14 February 1968, Horn Islet, Sir Edward Pellew Group, coll. B.

Cantrell (U.Q,); | %, Port Denison; 2 4 4, Dip Yard Lagoon via Brock’s

GROSS—AUSTRALIAN CEPHALOPLATUS 51

Creek, 8 April 1929, coll. T. G. Campbell; 1%, Roper River, September-

December 1910, coll. E. D. Frizelle; | ¢, Pt. Keats, 26 June 1952, coll

Australian Museum NW Australian Party (A.M.); 2 ¢ ¢, Manbullo Station,

Katherine, 8 July 1929, coll. T. G. Campbell; 1 ¢, Marrakai Station,

coll. 28-31 July 1929, coll. I. M. Mackerras & T. G. Campbell; 1 ¢, Green

Ant Creek near Dip Yard via Brock’s Creek, 9 April 1929, coll. T. G.

Campbell; 1 2, Ladysmith Yard, Turn-off Lagoon, 17 September 1930,

coll. T. G. Campbell (A.N.LC.); 1 @, 30 miles East of Darwin, coll.

G. F. Hill; 1 ¢,1 2, Borroloola, 2 October 1911, coll. G. F. Hill on Barclay

Expedition; 1 ?, King River, 24 December 1915 (N.M.); 12, Port Darwin;

1 4,2 29, Darwin, coll. G. F. Hill; 1 ¢, 30 miles East of Darwin, coll.

G. F. Hill; 2 ¢ 4, 1 9%, Daly River, coll. H. Wesselman; 1 ¢, 2 22,

Groote Eylandt, coll. N. B. Tindale; | ¢, Finke River, coll. Dr. H. Basedow;

1 ?, unlocalized, coll. S. A. White; 1 ¢, 1 2%, unlocalized (S.A.M.); 1 ?,

Darwin (Paris); series, Darwin; series, Adelaide River; 1 ¢ Stapleton, coll.

G. F. Hill (B.M.); 2 6 4 on grass, 7 March 1961, 1 ¢, 21 March 1961,

Maningrida, 5 m, Arnhem Land, coll. J. L. & M. Gressitt; | @, 128 Km

South of Darwin, 11 January 1964, coll. J. Sedlacek; 1 4, 30 miles East of

Darwin (Bishop). Western Australia. 19, Lulingui Station near Derby,

West Kimberly, 26 August 1929, coll. I. M. MacKerras & T. G. Campbell

(A.N.1.C.); 14, Roebuck Bay, 19 April 1892, coll. Kraptan (S.A.M.); 1°,

Wyndham, coll. Helms (Bishop). Queensland. 146, Mcllwraith Range,

31 May 1932, coll. Darlington on Harvard Expedition; | ? Coen, Cape York

Peninsula, 6 July 1932, coll. Darlington on Harvard Expedition (A.M.N.H.);

1@, Yarra, June 1959, coll. J. Guerassimoff; 1 ¢, 2? 9, Breakfast Creek,

Silver Plains, East Coast of Cape York Peninsula, 10 August 1963, coll.

J. L. Wassel; 1 6, 22 2, Charters Towers, 15 January 1964, coll. G. Mon-

teith (U.Q.); 1 4, Retro Station near Capella, August 1929, coll. J. R. Sleven

(A.M.); 24 46, 19%, Eidsvold, 28 September 1929; 12, 8 miles WNW of

Mt. Carbine, 20 September 1956, coll. M. J. D. White; 26 ¢,52 2, Bamaga,

Cape York Peninsula, 27 March-4 April 1964, coll. I. F. B. Commen &

M. S. Upton (A.N.LC,); 22 2, Bluff, coll. A. M. Lea; 2? 2, Mutchilba,

February 1933, coll, coll. A. D. Selby (S.A.M.); 346 6, Rockhampton

(Stockholm); 1 ?, Somerset (Paris); 1¢, 22? 2, at light, Mornington Island,

May 1963, coll. P. Aitken & N. B. Tindale; 14, 22 2, Appel Channel.

Mornington Island, 2 June 1960, coll. P. Aitken & N. B. Tindale (S.A.M.).

Torres Straits Islands. 1 6, Prince of Wales Island, August 1920, coll. J. A.

Kusche (Bishop).

Remarks: The species appears to occur in the Kimberly Division of

North Western Australia, the whole of the Northern Territory and in those

parts of Queensland North of the latitude of the Northern Territory-South

Australian border and in the Torres Straits Islands. It is much more

abundant in the North of its distribution than in the South. The Finke

$2 REC. 8, AUST, MUS, 16 (4); 1-38 Oelober, 1970

River specimen clearly belongs to this species but the locality label unfor-

tunately does not indicate which state, it could have come from the North

of South Australia as the Finke River runs from the Northern Territory

into South Australia. It has been assumed to have come from the Northern

Territory section of the river’s course,

This species and the next (explanatuy n. sp.) belong to a small section

of the subgenus Cephaloplatus in which the juga are quite markedly expanded

laterally in comparison to the preceding groups of species, reaching laterally

to at least the level of the outer surfaces of the eyes. Pertyi is easily dis-

tinguished from explanatus in that the anterolateral margins of the pronotum

are strongly crenulate or denticulate (finely crenulate in explanatus) and

that there is a triangular lobe in front of the eyes (absent in explanatus).

Cephaloplatus (C.) explanatus sp. noy,

hig. 15, 6D

Moderate to rather large in size, yellowish with brown punctations

above and below.

Head with juga strongly developed into foliaceous structures which at

their widest extend at least to the level of outer margins of the eyes, the

lateral margins vaguely semicircular in outline and raised above the general

level of the head so that the juga are rather inclined towards the midline.

The apical angles of the juga sharply rounded, the inner margins in front of

the anteclypeus almost straight, contiguous or not. Eyes not very prominent

and the facets occupying only the anterior half of the optic process which

is inclined posteriorly. Antennae slender and five segmented, the fourth

and fifth segments the longest and subequal, the first segment not reaching

the apex of the jugum, yellowish, but the fourth and fifth segments infus-

cated apically.

Pronotum fairly flattish, raised only a little posteriorly, the lateral

margins strongly explanatory produced and the region of the anterior

angles produced forward as a broad triangular process almost to the level

of the apex of the anteclypeus (as seen from above). Anterior margin of

the pronotum rather rectangularly excavate, anterolateral margins finely

crenulate, almost entire. Lateral angles broadly rounded, posterolateral

margins almost straight, hind margin also.

Scutellum substantially triangular, the lateral margins rather angulately

concave, the maximum concavity occurring about the midpoint or a little

behind, the apex tending a little towards being lanceolate. Scutellum basally

somewhat raised and somewhat granulate on the disc.

GROSS—AUSTRALIAN CEPHALOPLATUS

es, a |

Beaenon K. Head

Fig. 1S: Cephaloplatis (C.) explanatus sp. nov.

S4 REC, S, AUST, MUS, I& (4): 1-58 Ocloher, 1970

Hemelytra in their basal quarter wider than the body, this part of the

corial margin somewhat produced and feebly crenulate. Behind this point the

lateral margins of the corium pradually converging posteriorad leaving

progressively more of the connexivum exposed, hind margin of corium

rather convex. Membrane hyaline with veins almost the same color and

mostly longitudinal. Laterotergites rectangular, posteriorly coarsely punc-

tate.

Beneath concolorous with brown punctations, coarser and denser on the

thorax, fine and sparse on the underside of the head, same on the underside

of the juga where they are coarser and denser, Femora with a few brown

granulations, Male pygophore with the apical angles produced (Fig. 6D).

Length: 10,5-13.5 mm.

Distribution; Northern Territory. Holotype (Reg. No, ANIC9421),

Katherine, 17 April 1962, coll. I. F. B. Common (A.N.LC.): allotype ®,

Daly Waters, 26 October 1962, coll. E. §. Ross & D. Q. Cavagnaro (C.A.S.).

Western Australia, 2 paratype ? %, Upper Ord River, East Kimberley,

coll, Helms (B.M.); | paratype 4 (Reg. No. T4116), North West Australia

(N.M.). Unlocalized. 1 paratype & , presented by Commonwealth Institute

of Entomology 1948 (B.M.).

Remarks; The species seems rather rare and is easily recognized by the

very foliaceously produced juga and the strongly developed laminate lateral

Margins of the pronotum which are strongly produced forward but not con-

spicuously crenulate,

Cephaloplatus (C.) reticulatus Bereroth

Cephaloplatys reticulatus Bergroth, 1895, Proc, R. Soc. Vict. 7:288. Distant,

1910, Ann, Mag, nat. Hist., (8)6:472,

Cephaleplatus reticulatus Tay, 1966, Pap, Dep. Ent. Univ. Qd, 2(4):76-77.

figs.

Fig. 16, 6E.

A very distinctive and rather large species of the genus. In macro-

scopic appearance reddish or orange, the ground colour is actually pinkish

yellow or orange, and the punctations black or dark brown. There ure

small to moderate sized granular structures on the hind lobe of the pronotum,

the scutellum, and the hemelytra; these are rather different in appearance und

form to those of the species we have treated previously, they may be better

described as rugulosities.

Head with juga conspicuously and rather foliaceously expanded, pro-

duced forward well in front of the anteclypeus, Lateral Margins of juga

immediately in front of eye formed into a broadly triangular lobe which

reaches outwards to about level of the outer margin of the eyes, this lobe

GROSS—AUSTRALIAN CEPHALOPLATUS 34

56 REC. 8. AUST. MUS. 16 (4): 1-58 October, 1970

rather strongly convex above. Lateral margins of the juga anterior to this

process broadly semicircularly curved to the shortly rounded apices, this cur-

vature sometimes somewhat sinuate, interior margins of juga only slightly con-

vex widely separated or contiguous. Lateral and anterior margins of juga

somewhat raised so that the anterior portion of the head in front of the eyes is

rather concave. Head moderately raised between the ocelli. Eyes fairly

prominent, facets occupying all of the anterior facing surface, only the hind

rather concave surface bare and somewhat flattened. Antennae a little more

robust than in the other species, first segment not reaching the apices of the

juga, second, fourth, and fifth segments subequal, third segment somewhat

shorter. Antennae reddish or yellowish in colour all but the apex and base

of the third segment and the apical half of the fourth and fifth infuscated.

Pronotum raised posteriorly, more depressed anteriorly, the anterolateral

margins progressively more explanate from the lateral angles forward, the

region of the anterior angles produced as a triangular process forward to

about the anterior portion of the convex process on the margins of the

juga (just in front of the eyes). Anterior margin of pronotum strongly con-

cave behind the collum, just behind the eyes forming a slight angle then

concave again and diverging to the apices of the anterior processes. Antero-

lateral margins rather sinuate, strongly denticulate, crenulate, or serrate, in

many specimens tending to be somewhat flexed a little in front of the

lateral angles. Punctations on the explanate lateral margins of the pronotum

much larger and much less dense than on most other parts of the body.

Lateral angles produced as a rather blunt lobe, posterolateral margins

angulately convex, posterior margin straight or somewhat concave. Disc

of the posterior part of the pronotum with scattered rugulosities, these

rugulosities tending to appear in the form of short raised lines and some of

these lines tending to be raised as an incipient sinuous transverse line extend-

ing across the pronotum between the lateral angles,

Scutellum much the same shape as in the other species of the genus

but the angulation in the lateral margins behind the middle somewhat more

pronounced, tip broadly rounded. In each basal angle of the scutellum

the usual elongate black fovea, but this fovea not margined on its interior

side by a smooth callous area. Whole disc of the pronotum rather irregularly

rugulose in contrast to all preceding species which have been in the main

granulate.

Hemelytra in their basal quarter wider than the abdomen, this portion

of the corial margin formed as a convex angle in contrast to other species

and interiorly and forward of this angle and between the first and outer visible

vein strongly concave. Behind this point the corial margins relatively

straight then strongly converging leaving almost all of the connexivum

exposed. Disc of the corium covered with rugulose and raised glabrous

GROSS-AUSTRALIAN CEPI ALOPLATUS 57

areas. Hind margin of corium exteriorly somewhat convexly prominent,

thence slightly concave interiorly, broadly convex. Membrane with veins

strongly reticulate, hyaline, the veins brownish, no trace of raised brownish

spots or maculae between the veins, Abdomen relatively much wider than

in the other species, laterotergites concolorous but with a prominent infus-

cated marking exteriorly behind each incisure. Hind angle of the latero-

tergites strongly prominent,

Beneath concolorous with the dorsal surface, punctations more evenly

distributed and somewhat denser, On the underside of the head there

is a short infuscated bar immediately in front of the insertion of each

antenna und a fuscous area at the base of the head behind the bucculue.

All thoracic sterna are brown or piceous save the keel on the mesosternum

which is yellow, A small fuscous point just above the junction between

the epimeron and Lhe episternum on each thoracic segment. On the abdomen

the base of the third segment narrowly and transversely darkened, strongly

punctate, medially at the base of the fourth, fifth, sixth and seventh abdonninal

sternites a short brownish line or bar, running longitudinally, and giving the

impression Of an interrupted line running the length of the abdomen

inedially. There is no sign of any areas of lateral infuscation. Legs reddish

yellow or yellowish with on the the femora and tibia lurgish somewhat

raised brown spots, some of those on the femora bearing a hair. Male

pygophore figured (fig. 6B).

Length: 1(2.0-14.0 mn.

Distribution: Queensland. Lectotype é (Rey. No. T4115), donated C,

French Jun,, 15 November 1911 (N.M.). New South Wales. Tibooburra, 23

May 1949, coll, K, H, L. Key (A.N.LC.), Fietoria, 1 2, Hattah, 16 June

1948, coll. C, Oke (N.M.). South Australia, 3° °, Leigh Creek: 3 2 @,

South of Ediacara, 5 November 1959, coll. P. Aitken (S.A.M.). Western

Australia, 3 2 2, Lennaville; | ¢, Wiluna, 29 August 1968, coll. F, H,

Uther Baker; | °, Mullewa, coll Miss F. May (S.A.M,); series, Violet Range,

E. Murchison, uccessed 1904 (B.M.). Unlocalized. 1 2, donated by

F. P, Spry, 5 October 1922 (N.M.).

Remarks: As for C, (C.) granulatus Bergroth the type of this species

was not found in Helsinki, nor indeed any examples of this species. However,

there was again in the collection of the National Museum of Victoria a male

specimen from the C, French collection which fits Bergroth’s description of

Cephaloplatys reticulatus even to the missing fifth antennal segment, although

the specimen no longer carries dn indication that it came from Queensland,

This male specimen is accordingly selected as the lectotype of “Cephaloplartys

reliculatus” Bergroth and marked accordingly.

38 REC. S. AUST. MUS. 16 (4); 1-58 October, 1970

Because of its larger size and conspicuously reticulate membrane the

species is one of the easiest of Cephaloplatus species to recognize. From the

data available it appears to be restricted to drier habitats but with a rather

more southern distribution than the other species.

The male and female genitalia were discussed by Tay (1966).

BIBLIOGRAPHY

Bergroth, E., 1895: Contributions to a knowledge of the Rhynchota of Australia. Proc. R.

Soe. Viet., 7:287-302.

1916: Heteropterous Hemiptera collected by Professor W. Baldwin Spencer

during the Horn Expedition into Central Australia. Proc. R. Soc. Vict.,

(N.S.) 29(1):19-39.

Carpenter, G. H., 1891: 15. Reports on the Zoological Collections made in Torres Straits

by Professor A. C. Haddon, 1888-1889. Rhynchota from Murray Island and

Mabuiag. Sci, Proc. R. Dublin Suc. 7(3):137-146, 2 plates.

Dallas, W. S., 1851: List of the Specimens of Hemipterous Insects in the Collection of the

British Museum. Brit. Mus. Pub,, Pt. 121-368, 11 plates.

Distant, W. L., 1881: Rhynchota from the Australian and Pacific Regions. Trans. R. ent.

Soe. Lond., 211-18,

1899: Rhynchotal Notes—III. Heteroptera: Discocephalinae and Pentatominae

(part), Ann. Mag. nat. Hist, (7) 4: 421-445.

1910: Rhynchotal notes—S2. Australasian Pentatomidae, Ann. Mag. nat. Hist,

(8)6:369-386, 465-481, 585-603.

Lethierry, L. & Severin, G., 1893: Catalogue general des Hemipteres, Bruxelles. 1:i-x,

1-286.

Stil, C.. 1867: Ofvers. Konel. svenska Vetenska Akad. Foérh., 491-560.

1876: Enumeratio Hemipterorum 5. K, svenska Vetensk Akad. Handl. 14(4):

1-162,

Tay, E. B., 1966; Morphology of the External Genitalia of Some Pentatominae (Heterop-

tera). Pap, Dep. Ent, Univ. Qd. 2(4):75-84, 5 plates.

Van Duzee, E. Pp. 1905: Notes on Australian Pentatomidae with descriptions of a few new

species, Bull. Amer. Mus. nat. Hist., 21:187-214. PL 8.

Walker, F., 1867: Catalogue of the Specimens of Heteropterous—Hemiptera in the Collec-

tion of the British Museum. Brit. Mus. Pith., Pt. 1:1-240,

1868: Catalogue of the Specimens of Hemiptera Heteroptera in the Collection of

the British Museum. Brit. Mus. Pub. Pt, 1111-599,

White, A., 1842: Description of Hemipterous Insects of the Section Heteroptera, Trans.

R. ent. Soc, Lond, 3:84-94.

RECORDS OF THE

SOUTH AUSTRALIAN

MUSEUM

CYATHOLAIMIDAE (NEMATODA)

FROM THE COAST OF WESTERN

AUSTRALIA

By W. GRANT INGLIS

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 5

1 December 1970

CYATHOLAIMIDAE (NEMATODA) FROM THE COAST OF WESTERN

AUSTRALIA

BY W. GRANT INGLIS

Summary

The five species described here were collected on the coast of Western Australia and represent on

new species of Praeacanthonchus, and four species of Paracanthonchus, of which three are new.

Both genera are referable to the family Cyatholaimidae of the Order Chromadorida, an Order in

which most of the classification is based on the structure of the cuticle, the shape of the amphids

and the form of the dorsal onchium in the oesophastome. However a study of Euchromadora de

Man, 1886 and apparently similar species, all referable to the Chromadorida, has raised doubts as to

the value of some of the characters, and to the way in which they have been used (Inglis, 1969).

CYATILOLAIMIDAE (NEMATODA) FROM THE COAST OF

WESTERN AUSTRALIA

By W, GRANT INGLIS

South Australian Museum, Adelaide, S.A. 3000

The five species described here were collected on the coast of Western

Australia and represent one new species of Praeacdnthonchus, and four

species of Paracanthonchus, of which three are new. Both genera are

referable to the family Cyatholaimidae of the Order Chromadorida, an

Order in which most of the classification is based on the structure of the

cuticle, the shape of the amphids and the form of the dorsal onchium in

the oesophastome. However a study of Euchromadora de Man, 1886 and

apparently similar species, all referable to the Chromuadorida, has raised

doubts as to the value of some of the characters, and to the way in which

they have been used (Inglis, 1969),

Brictly, the difficulty arises because much of the classification of the

family Chromadoridae, of which Euchromadora is a member, has been based

on the distinction between u solid or a hollow dorsal onchium in association

with features of the cuticle and the shape of the gubernaculum and on

assumed co-variances between these characters. The previous study (Inglis,

1969) has shown that a solid dorsal onchium is associated with a number of

other characters, particularly with the presence of small denticles along the

anterior edge of the dorsal wall of the ocsophastome and a series of transverse

rows of denticles on the lateral and ventral walls of the oesophastome.

In contrast, when the dorsal onchiuin is hollow there are no dorsal denticles

and instead of transverse rows of lateral and ventral denticles there ure a

small number of conical denticles laterally, in one row, and two or four

similar ventral denticles. Other features, particularly of the cuticle, are

associated with the two groups while the shape of the gubernaculum ts not

convariant with these characters.

In some specimens it was difficult to decide if the dorsal onchium was

unequivocably solid or hollow but in some in which it appeared solid the

dentition and characters of the cuticle were similar to those otherwise only

found in forms with hollow onchia. It is, therefore, likely that there are

several convergent groups within the Chromadorida which culminate in

superficially similar forms with complicated cuticles, solid dorsal onchia

and similar gubernacula. This implies the possibility that forms with

simple punctate cuticles, which are currently classified together, may also

be grouped on convergent characters. All the species described here have

such simple punctate cuticles and it appeared possible that their head structure

would help to resolve this problem.

Isvtwd Lo Deeceniber 1970

REC. 8S. AUST. MUS.. 16 (5); 1-13 December, (970

COMPARATIVE ANATOMY

CUTICLE

In all the species described below, with one exception, the cuticle is

marked by distinct, round punctations but the occurrence and kind of

lateral differentiation is variable. The amphids are spiral and “‘pore-

complexes” are present in some species. As a result of Wright and Hope’s

(1968) study of the “campaniform-type” organ L use their term “pore-

complex” while stressing that two types exist (Inglis, 1963). The Type-1,

which Wright and Hope studied appears as a pit in the cuticle transversed

by an elongate opening while the Type-2, which is generally a larger form,

is similarly a pit but with a circular opening (or dome?) in the centre,

The distribution of these two types still appears to be of significance.

Type-2 pores have only been found in files posterior to the amphids or

on the lateral surface of the body at the level of the cloacal opening. In

P, margaretae, which is described below, there is a file of Type-l pores

running posteriorly to the amphids. This raises the possibility that Types-1

and -2 are two expressions of the same type. This must await confirmation

as such an implied distortion has never been seen when specimens of the

same species ure compared.

HEAD

The mouth opening is bounded by the usual twelve rugae and leads

into an oesophastome through a cheilostome which appears to be circular

in transverse optical section. There is a hollow dorsal onchium in all species,

In the three species of Paracanthonchus in which en face preparations were

studied the oesophastome is triangular in transverse section with two small

onchia borne on a muscular pad on each ventro-luteral oesopharhabdion

(Figs. 2, 8 and 14). The apparent differences are probably due to the

degree to which the oesophastome is open. If this is so the condition shown

for P. margaretae (Fig. 14) may be interpreted as the fully open condition,

that for P. cheynei (Fig. 8) as fully closed while that for P. kartanum

(Fig. 2) is intermediate.

The structure of the oesophastome in the three species studied corres-

ponds to the grouping formed on other characters and the conditions are

different from those described for the Euehromadora and Euchromadora-

like species studied before (Inglis, 1969), particularly in the triangular

oesophastome in transverse section. Nevertheless the conditions correspond

more closely to those in the “hollow-toothed” forms than those in the

“solid-toothed”. Further work is certainly warranted and likely to be of

value in clarifying the classification of the Chromadorida.,

INGLIS—WESTERN AUSTRALIAN CYATHOLAIMIDAE

MALE

In all the species there is a series of pre-cloacal supplements on the

mid-ventral surface anterior to the cloacal opening. In all four species of

Paraconthonchus the supplements are rod-like while in the Praeacanthonchus

species they are indistinct and tube-like. However in all cases they do not

appear to open on the surface of the body, in so far as [ can tell with the

light microscope.

In addition to these supplements, in all species, two other structures lie

close together immediately anterior to the cloacal opening which are usually

called supplements but which are structurally different. In all five species

these are in the form of pits in the cuticle into which a cuticular process

projects from the anterior edge (Figs. 5, 20 and 23). That these

“supplements” are different from those more anteriorly is not, of course, a

new discovery and they occur in a wide range of Chromadorida, It appears

likely that they are of taxonomic importance as they differ in shape and

their function must be different from the other more anterior supplements.

| propose, therefore, to call them “cloacids” about which more information

is needed.

FEMALE

In two of the species described below there is a distinct pit in the cuticle

both anterior and posterior to the vulvar opening. They are probably sense

organs as a nerve process appears to project into the bottom of each,

DESCRIPTIVE SECTION

Holotype males will be deposited in the Western Australian Museum

and paratypes in that institution and in the British Museum (Natural History )

where unsorted samples are also held. All localities are in Western Australia,

In agreement with previous policy (Wieser and Hopper, 1967: Inglis,

(1969) only absolute measurements are given in the descriptions.

Paracanthonchus kartanum (Mawson, 1953)

(Figs. 1-5)

SYNONOMY

Harveyjohnstonia kartanum Mawson, 1953: Pennington Bay, Kangaroo

Island, South Australia.

4 REC. S. AUST. MUS., 16 (5): 1-13 December, 1970

Figs. 1-5. Paracanthonchus kartanum. Fig. 1. Lateral view of head, dorsal to right.

Fig. 2. Oesophastomal dentition, sketch. Fig. 3. Cloacal region. Fig. 4. Spicules and

gubernaculum, ventral view. Fig. 5. Cloacid, sketch. Figs. 6-11. P. cheynei.

Fig. 6. Lateral view of head, dorsal to left. Fig. 7. Head, en face. Fig. 8.

Oesophastomal dentition, sketch. Figs. 9-10. Spicules and gubernaculum. Fig. 11.

Lateral view, male tail.

INGLIS—WESTERN AUSTRALIAN CYATHOLAIMIDAE 5

LOCALITIES

Among weed and associated sand in hold fasts on exposed rocks, Goode

Beach, Albany.

Among weed and mussels in rock pools exposed at low tide, Hall's

Head, Mandurah.

Among weed and associated hold fasts and sand on a silty sheltered

beach, Cheyne Beach, near Albany.

Among sea-weeds and associated sand in rock pools, Bunker Bay,

Geoyraphe Bay.

MEASUREMENTS (mm).

MALE. Body length: 0,87: O91; 0,99; 1.04, Body breadth:

0.043; 0,043; 0.050; 0.044. Ocsophagus length: 0.152; 0.159; 0.152;

0.155, Diameter of head: 0.017; 0.017; 0.018; 0.017. Lengths of cephalic

setae: 0.005; 0.005; 0.004; 0.004. Excretory pore from anterior end of

body: 0.068; 0.066; 0.070; 0.069. Diameter of body at amphid: 0.024;

0.022; 0.023; 0.024. Diameter of amphid: 0.009; 0.009; 0.009; 0.009.

Length of spicules: 0.027; 0.023; 0,025; 0,029, Length of gubernaculum:

0,033; 0.026; 0.033; 0.036. Length of tail: 0.078; 0.097; 0.079; 0.086.

Cloacal diameter: 0.043; 0.039; 0.040; 0.036.

The cuticle is marked by fine punctations and there is no lateral

differentiation, There are two lateral files of setae running the length of

the body but no “pores” were seen. No dome-like Type-2 “pores” occur

behind the amphids or lateral to the cloacal opening.

The six inner sense organs are setose, and there are twelve equal setae

in the outer circle. The amphids are spiral with 44 to 44 turns. The dorsal

onchium is very small and there are two similar onchia, which arise from a

raised muscular pad on each yentro-lateral wall of the oesophastome. The

cheilostome and oesophastome are triangular in transverse section although

Lam not certain whether this reflects a constant feature or simply the degree

of muscle contraction,

The tail is fairly slim. There are four well cuticularized, tubular

pre-cloacal supplements on the mid-ventral surface of the body anterior to

the cloacal opening which increase in size anteriorly, In addition there are

two circular cloacids immediately anterior to the cloacal opening into each

of which projects a small cuticular process.

The spicules are fairly massive with narrow alae. The gubernaculum

is ubout the same width along its length with a rounded slightly swollen region

posteriorly, which bears a series of four small denticles and from which arises

a pair of obvious finger-like postero-dorsally directed processes.

6 REC. S. AUST. MUS., 16 (5): 1-13 December, 1970

DISCUSSION

The only marked difference between the specimens from Western

Australia and those described by Mawson (1953) is in the number of turns

in the spiral amphids. Mawson refers to 34 while I count 44 to 42, on

different specimens. The difference is probably due to the method of

counting.

It is worth noting that Mawson comments specifically on the increase

in size of the pre-cloacal supplements anteriorly and this feature is very

Obvious in the Western Australian specimens.

Paracanthonchus cheynei sp. nov.

(Figs. 6-11)

LOCALITIES

Among sea-weed and hold-fasts with associated sand in 20 cm of silty,

sheltered water. Cheyne Beach, near Albany. Type locality.

Among sea-weed on exposed reef in 10 cm of clear water. Radar Reef,

Rottnest Island.

MEASUREMENTS (mm).

MALE. Body length: 1.24; 1.28. Body breadth: 0.052; 0.052.

Oesophagus length: 0.198; 0.203. Length of cephalic setae: 0.005; 0.005.

Diameter of head: 0,021; 0,023. Distance of excretory pore from anterior

end of body: 0.057; 0.059, Amphid diameter: 0.010; 0.009, Length of

spicules; 0.046; 0.048. Length of gubernaculum: 0,039; 0,042. Cloacal

diameter: 0.046; 0.049. Length of tail: 0.125; 0.131.

FEMALE. Body length: 1.09. Body breadth: 0.059. Ocsophagus

length: 0.173. Length of cephalic setae: 0.005. Diameter of head: 0.025.

Distance of excretory pore from anterior end of body: 0.057. Amphid

diameter: 0.009. Anal diameter: 0.038. Length of tail: 0.103. Distance

of vulva from anterior end of body: 0.60. Eggs: 0.062 x 0.036.

The cuticle is marked by fine punctations without lateral differentiation.

There are two files of transverse Type-I “pores” on each lateral surface of

the body and four circular Type-2 “pores” in a file posterior to each amphid.

The mouth is bounded with the usual twelve rugae, the inner sense

Organs are six papillae and the twelve members of the outer circle are short

setae all of the same length. The amphids have 4¢ spirals. The dorsal

onchium is small and there are two smaller, sharp onchia on each ventro-

lateral wall of the oesophastome, No ocelli were seen.

The tail is relatively stout and bears a pair of Jong ventral setae about

one quarter tail length posterior to the cloacal opening.

INGLIS—WESTERN AUSTRALIAN CYATHOLAIMIDAE 7

or ce; .

WS eA Me Z

‘ ®S SF

a NA \

a) [ vans S

( Sy ee N\@/S5)7 \

— a as G / —— J : \

«we celllescces\ct WANS ~\

Sa AR anecossey | Non: EK (( \

LD “NS KAN

Awe So 21

Figs. 12-18. Paracanthonchus margaretae. Fig. 12. Lateral view of head, dorsal to

left. Fig. 13. Head, en face. Fig. 14. Oesophastomal dentition, sketch. Fig. 15,

Oesophastomal dentition, lateral view, sketch. Fig. 16. Spicules and gubernaculum,

lateral view. Fig. 17. Gubernaculum, ventral view. Fig. 18. Detail of spicule.

Figs. 19-21. P. hartogi. Fig. 19. Male tail, lateral view. Fig. 20. Ventral view of

gubernaculum and cloacids. Fig, 21. Lateral view of head, dorsal to right.

8 REC. S. AUST. MUS., 16 (5): 1-13 December, 1970

MALk, There are four very prominent, strongly cuticularized pre-

cloacal supplements and two cloacids with the usual cuticular processes

projecting into them. The spicules are slightly bowed with sharp posterior

ends and broad alae, confirmed by dissection, The gubernaculum is massive

und slightly L-shaped. The posterior end is swollen with small denticles

along the postero-ventral edge. In ventral view this region curves laterally

and the denticles run along the posterior edge.

FEMALE, The reproductive system is double, opposed and reflexed.

There are small vulvar pits in the cuticle, into which sensory processes appear

to project through the base.

DISCUSSION

This species is very similar to P. platypus Wieser and Hopper, 1967,

particularly in the shape of the gubernaculum. It differs in the series of

small denticles on the posterior edge of the gubernaculum, the strongly alate

spicules, the greater number of spirals in the amphids, and the absence of a

spine on the anterior lip of the cloacal opening.

Paracanthonchus margaretae sp. nov,

(Figs. 12-18)

LOCALITIES

Among weed and associated hold-fasts and sand in 20 cm of silty,

sheltered water, Cheyne Beach, near Albany, Type locality.

Among weed and associated sand and silt on partly exposed, fairly

sheltered rocks, Windy Harbour.

Among weed and hold-fasts on rocks in 10 cm of water. Bunker Bay,

Cape Naturaliste.

MEASUREMENTS (mm).

MALE, Body length: 1.28; 1.48. Body breadth: 0.048: 0.055.

Oesophagus length: 0.25; 0.34. Length of cephalic setae, long/short/inner;

0,012/0.009/0.003; 0.011/0.008/0.003, Amphid diameter: 0.010; 0.010.

Distance of excretory pore from anterior end; 0.069; 0.071. Spicule length:

0.048; 0.047. Gubernaculum length: 0.044; 0.045. Tail length: 0,100;

0,102. Cloacal diameter: 0.039; 0.046,

FEMALE. Body length: 1.21. Body breadth: 0.059. Oesophagus

length: 0.30, Length of cephalic setae, long/short/inner: 0.013/0.011/

0,004. Amphid diameter: 0.011. Distance of excretory pore from anterior

end: 0,069. Tail length: O.101. Anal diameter: 0.036. Distance of

vulva from anterior end: 0.64.

The cuticle is marked by fairly strong punctations and the lateral

differentiation is well developed as large, regularly arranged dots anteriorly

INGLIS—WESTERN AUSTRALIAN CYATHGLAIMIDAE 9

which become less obvious posteriorly. There are eight files of transversely

ridged Type-1 pores, two on the lateral, two on the dorsal and two on each

ventral surface. There are three very prominent, elongate Type-| pores, with

antero-posterior openings, at the level of the cloacal opening on the males,

The cephulic sense organs of the inner circle are setae and the outer

circle of ten setae consists of six long and four short. The amphids have

4} spirals. The oesophastome is a prominent, deep cavity which ts triangular

in transverse section. The dorsal onchium is large and hollow and there are

two small onchia on each ventro-lateral wall of the oesophastome arising from

ai muscular pad, The tail is short and stout,

MALE. The four pre-cloacal supplements are relatively short and

rod-like while the two cloacids have somewhat arrow-shaped processes

projecting into them, The spicules bear fairly broad alae, are massive and

even in width, The anterior ends are swollen and rounded while the postetior

ends bear distinct rounded swellings before narrowing rapidly to a sharp

posterior tip, The gubernaculum is also massive and apparently ends

posteriorly in a straight dentate edge from the centre of which arises a

laterally directed tooth. In ventral view however each half of the guber-

naculum folds laterally at the posterior end as a triangular curved portion

which has small denticles along both outer edges. The impression of a large

median tooth in lateral view is due to studying the folded region from the side.

FEMALE. The reproduction system is opposed and reflexed with sensory

pits anterior and posterior to the vulva.

DISCUSSION

This species is similar to P. calcus (Bastian, 1865) and P. tyrrhenicus

(Brunetti, 1949) but differs from them, and all others in the genus, in the

posterior end of the gubernaculum, the small number of spirals in the amphid

and in the lateral differentiation.

Paracanthonchus hartogi sp, nov.

(Figs. 19-22)

LOCALITY

Among mud and weed from trawl in 35 metres. Shark Bay,

MEASUREMENTS (mm).

MALE. Body length: 1,24; 1.42. Body breadth; 0.066; 0.072.

Oesophagus length: 0.22; 0,20, Length of cephalic setae, long/ short:

0.0038/0.0026; 0.0039/0,0026. Diameter of amphid: 0.013; 0.012.

Distance of excretory pore from anterior end: 0.056; 0.049. Spicule length:

0,049; 0.044. Gubernaculum length: 0.044; 0.039, Length of pre-cloacal

supplements; 0,026; 0.024. Length of tail: 0.010; 0.011, Cloacal

diameter: 0.049; 0,049,

Ma REC. 8. AUST. MUS., 16 (4): 1-13 December, 1970

The cuticle is marked by fine striations and there is no lateral

differentiation or indication of punctations. Two files of stout setae run

the length of each lateral surface of the body but no “pores” were seen.

The inner circle of cephalic sense organs was not seen and the outer

circle of twelye setae consists of six long and four short. The spiral amphids

consist of 32 spirals. No en face view was prepared and from the lateral

aspect only a small dorsal onchium could be seen. The oesophagus is simple

club-shaped and expands evenly and slightly posteriorly.

The short stout tail is curved ventrally and the four pre-cloacal

supplements are prominent, well sclerotized tubes. The two cloacids are

circular pits into which project arrow-head-like processes. The spicules are

relatively small with narrow alae and distinctly set off semi-spherical

proximal ends. ‘The gubernaculum is L-shaped with a massive distal end

which bears four prominent teeth.

DISCUSSION

The relationships of this species are a little uncertain because the

cuticle is marked by fine striations with no indication of punctations. On

the other hand the shape of the gubernaculum and the form of the pre-cloacal

supplements is so similar to that found in some species referred to

Paracanthonchus, that [ have little hesitation in referring this Western

Australian species to that genus,

In particular P. hartogi resembles P. serratus Wieser 1959 but differs

from it in the square distal end of the gubernaculum and the prominent teeth

on the same region, in addition to the form of the cuticle.

Praeacanthonchus cygnis sp, nov.

(Figs, 23-28)

LOCALITY

Among rocks and sea-weed in 10 em water, Woodman’s Point,

Fremantle,

MEASUREMENTS (inm).

MALE. Body length: 1.60; 1.69; 1.85. Body breadth: 0.079;

0.075; 0.068, Oesophagus length: 0.22; 0.23; 0.23. Length cephalic

setae, long/short: 0.008/0.009; 0.009/0.011; 0.010/0.012. Distance of

excretory pore from anterior end: 0.12; 0.13; 0.12. Diameter of amphid:

0.013; 0.012; 0.013. Diameter at amphid: 0.039; 0.039; 0.035. Spicule

length: 0.047; 0.048; 0.046. Gubernaculum length: 0.057; 0.061: 0.056.

Length of tail: 0.15; 0.17; 0.19. Cloacal diameter: 0.073; 0.066; 0.057.

The cuticle is marked by regular punctations with a slight lateral

differentiation where the punctations are irregularly arranged. The body

INGLIS—WESTERN AUSTRALIAN CYATHOLAIMIDAE at

Fig. 22. Paracanthonchus hartogi. Detail of spicules and gubernaculum. Figs.

23-28, Praeacanthonchus cygnis. Figs. 23-24. Ventral and lateral views of cloacid,

sketch. Fig, 25. Lateral view of head, dorsal to right. Figs. 26-27. Detail of spicules

and gubernaculum. Fig. 28. Male tail, lateral view.

12 REL, S. AUST. MUS. 16 (44) 1-13 December, 1970

bears a number of short setae, and eight files of transverse Type-1 pores.

A wholly lateral file of four circular Type-2 pores runs posterior to each

amphid and there is 4 file of six such pores on each lateral surface of the

body at the level of the cloacal opening.

The head is typical with an inner circle of six papillate sense organs

and an outer circle of ten setae of which six are short and four are long.

The amphids are spiral with 34 to 34 turns each and lie relatively far

posterior to the anterior end of the head, TI have been unable to prepare a

satisfactory en face view of the head but there is a small dorsal onchium and

no indication of any lateral or ventral onchia in lateral view.

The tail is short and stout, There are six indistinct midventral, tubular

pre-cloacal supplements. The two cloacids lying just anterior to the cloacal

opening are small ovoid depressions into each of which projects a small

finger-like process.

The slim spicules are about three-quarters the length of the massive

gubernaculum and bear slight alae. The gubernaculum is massive distally

with four strongly developed teeth. Proximally the guberculum is slim in

lateral view. The spicules run in a groove between the four-toothed posterior

heads and a pair of lateral processes which arise from the main body of the

gubernaculum., The body of the gubernaculum is split for almost its whole

length and consists of a well sclerotized rod which bears a thin, less well

sclerotized region dorsally and internally.

DISCUSSION

The combination of a typically Cyatholaimus-type gubernaculum, slight

lateral differentiation on the cuticle, and indistinct tubular pre-cloacal

supplements ts diagnostic of the genus Praeacanthonchus Micoletzky, 1924b

according to Wieser (1954). The only previously known species of this

genus is P, punctatus (Bastian, 1865) if De Man’s (1890) redescription

is accepted.

P. eygnis diflers from P. punctatus in several ways, particularly in the

shape of the proximal end of the gubernaculum which is slim in P, cygnis

but stout in P. punctatus; the spicules of the latter species have very wide

alae while those of P. cyenis are narrow und the spicules of P. punctatus

ure longer than those of P. cygnis relative to the length of the gubernaculum,

ACKNOWLEDGMENTS

The specimens deseribed here were collected in Western Australia while 1 was an

exchange curator al (he Western Australian Muscum. 1 thank the Director and stall! of that

Museum for their assistince during my stay. Miss B. Mitchell assisted in sorting material

in the British Museum (Natural History) and Mrs, &. K, Head, South Australian Museum,

prepared the illustrations.

INGLIS—WESTERN AUSTRALIAN CYATHOLAIMIDAE 13

REFERENCES

Bastian, C. H., 1865: Monograph of the Anguillulidae or free nematoides, marine, land and

fresh-water: with descriptions of 100 new species. Trans. Linn. Soc. Lend.,

25: 73-184.

Brunetti, B., 1949: Contributo alla conoscenza dei Nematodi del M. Tirreno Il. Alcune

specie appartenenti alle famiglie: Enoplidae, Cyatholaimidae, Chromadoridae,

Axonolaimidae. Monitore zool. italiano, 57: 41-59.

Inglis, W. G., 1963: “Campaniform-type” organs in the Nematoda. Nature, 197: 618.

Inglis, W. G., 1969: Convergence in the structure of the head and cuticle of Euchromadora

and apparently similar nematodes. Bull. Brit. Mus, nat. Hist., 17 (5): 149-204,

de Man, J. G., 1886: Anatomische untersuchungen iiber freilebende Nordseenematoden,

Leipzig.

de Man, J. G., 1890: Quatriéme note sur les nématodes libres de la Mer du Nord et de la

Manche. Mém. Soc. zool. Fr., 3: 169-194.

Mawson, P. M., 1953: Some marine freeliving nematodes from the Australian coast. Trans.

roy. Soc. S. Aust., 76: 34-40,

Micoletzky, H., 1924: Letzter Bericht tiber freilebende Nematoden aus Suez. Sitzungsber.

Akad. Wiss. Wien. Math.—Naturw. Kl., 133: 137-179.

Wieser, W., 1954: Reports of the Lund University Chile Expedition 1948-49. 17. Free-

living marine nematodes. II. Chromadoroidea. Acta, Univ. lund. N.F.,

50 (16): 1-148.

Wieser, W., 1959: Free-living nematodes and other small invertebrates of Puget Sound

beaches. Univ. Washington Publ. Biol., 19: 1-179.

Wieser, W. and Hopper, B., 1967: Marine nematodes of the East Coast of North America.

I. Florida. Bull, Mus. comp. Zool. Harv., 135: 239-344,

Wright, K, A. and Hope, W. D., 1968: Elaborations of the cuticle of Acanthonchus duplicatus

Wieser, 1959 (Nematoda : Cyatholaimidae) as revealed by light and electron

microscopy. Canad. J. Zool., 46: 1005-1011.

A. B. JAMES, GOVERNMENT PRINTER, ADELAIDE

RECORDS oF THE O

SOUTH AUSTRALIAN pA Q

MUSEUM By:

THE GENUS CTENOTUS (LACERTILIA,

SCINCIDAE) IN SOUTH AUSTRALIA

By G. M. STORR

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 6

18 February 1971

THE GENUS CTENOTUS (LACERTILIA, SCINCIDAE) IN SOUTH

AUSTRALIA

BY G. M. STORR

Summary

Examination of the material in Australian museums reveals that 17 species and subspecies of

Ctenotus occue in South Australia, including two new species, C. brachyonyx and C. regius, and

two new subspecies, C. uber orientalis and C. brooksi euclae. A key is provided.

THE GENUS CTENOTUS (LACERTILIA, SCINCIDAE) IN

SOUTH AUSTRALIA

By G. M. S5TORR

Curator or Biraps AND REPTILES, WESTERN AUSTRALIAN MUSEUM

SYNOPSIS

Examination of the material in Australian museums reveals that 17

species and subspecies of Cfenotus occur in South Australia, including two

new species, C. brachyonyx and C. regius, and two new subspecies, C. uber

orientalis and C. brooksi euclae. A key is provided,

INTRODUCTION

Of the 14 species of Crenotus known from South Australia, all but two

occur in neighbouring parts of Western Australia or the Northern Territory.

My papers on the Crenotus of the Eastern Division of Western Australia

(Storr 1969) and of the Northern Territory (Storr 1970) thus serve as an

introduction to the South Australian fauna. The reader is also referred to

those papers for descriptions of the seven species and subspecies restricted

in South Australia to the far north and west; they are represented by too

few South Australian specimens to warrant a local description.

As the foregoing remarks imply, the arid northwest of South Australia

is much richer in Cfenotus than the humid southeast. The differences are

probably still greater than is apparent here, for much of western South

Australia is virtually unexplored, and a further six taxa could eventually be

found there: dux and colletti nasutus in the far northwest: grandis, piankai

and calurus in the Great Victoria Desert; and w. uber on the Nullarbor

Plain, Users of the key will need to bear this in mind when identifying

western specimens.

C. impar has been omitted from this paper. This species is certainly

known only from southwestern Australia, and the provenance of a specimen

from Ooldea (SAM 9980) requires confirmation.

For the loan of specimens in the South Australian Museum (SAM),

National Museum of Victoria (NMV), Australian Museum, Sydney (AM).

Queensland Museum (QM), Arid Zone Research Institute, Alice Springs

(NTM), and the Pianka Collection (ERP), I am grateful respectively to

Mr. F. J. Mitchell, Miss J. M. Dixon, Dr. H, G. Cogger, Miss J. Covacevich,

Mr. B. L. Bolton, and Dr. E. R. Pianka,

I dedicate this paper to the memory of Francis John Mitchell, Jute

Curator of Reptiles in the South Australian Museum.

Lysued W& February, 1971

REC. 8. AUST. MUS., 16 (6): I-15 February, 1971

Key to Species and Subspecies

. Pattern consisting of longitudinal stripes and/

or series of spots, but no ocelli; nasal weakly

or not grooved; midbody scale rows 22-34

Pattern consisting solely of black-and-white

ocelli; nasal strongly grooved; midbody

scale rows 32-38 .. 1. ., nae :

Adults large (SVL = snout-vent length up to

95 mm.); toes slightly or not compressed;

subdigital lamellae smooth or broadly

callose; second supraocular longer than

first; nasals separated; prefrontals usually

forming median suture—lesueurii group ..

Adults small to moderately large (SVL up to

80); toes moderately or strongly com-

pressed; subdigital lamellae keeled or

narrowly callose . .

. Vertebral stripe distinctly pale-edged .

Vertebral stripe with little or no indication of

pale edge .. =

White midlateral stripe extending forward to

lores; vertebral stripe nearly as wide as a

paravertebral scale; brow subacute .

White midlateral stripe extending forward only

to arm; vertebral stripe much narrower than

a paravertebral scale; brow obtuse .

- White dorsolateral and midlateral stripes

moderately well developed; claws very short

White dorsolateral and midlateral stripes

absent or barely discernible; claws moder-

ately long

- Adults moderately large (SVL up to 80):

subdigital lamellae obtusely keeled or

narrowly callose

Adults small (SVL up to 62); subdigital

lamellae sharply keeled and mucronate ..

pantherinus ocellifer

ies)

robustus

saxatilis

brachyonyx

helenae

STORR—GENUS CTENOTUS IN SOUTH

7. Ground colour brownish; pattern consisting of

dark and pale stripes and longitudinal series

of pale spots—leonhardii group

Ground colour blackish; pattern consisting

solely of pale stripes—taeniolatus group -.

8. White midlateral stripe well developed (broad,

straight, and extending forward to ear);

blackish vertebral stripe moderately wide

and narrowly margined with white; mid-

body scale rows 24-28 ©. 2. e+ ee ee es

White midlateral stripe absent or poorly

developed (not extending forward to arm);

dark brown vertebral stripe narrow, with

or without pale brown margin; midbody

scale rows 26-34 . fhe ade

9. White midlateral stripe absent or scarcely

indicated; dark laterodorsal stripe enclosing

a series of pale spots; nasals usually

separated; prefrontals contiguous or sep-

arated ..

White midlateral stripe well defined posteriorly;

dark laterodorsal stripe not enclosing pale

spots; nasals usually forming median suture,

prefrontals usually separated .. .. -.

10. Pale lines and stripes totalling 8 or LO, with

only one line on each side between mid-

lateral and dorsolateral stripes

Pale lines and stripes totalling 14 or 16, with

at least 2 lines on each side between mid-

jateral and dorsolateral stripes .

11. Pattern consisting of stripes, spots, blotches

and variegations: tail less than twice as

Jong as SVL—schomburgkii group .

Pattern consisting solely of alternating dark

and pale stripes; tail bright red, more than

2.3 times as long as SVL

AUSTRALIA

regius

uber orientalis

leonhardii

atlas

guattuordecimlineatus

leae

REC. S. AUST. MUS., 16 (6): 1-15

12. Nasals usually separated ( occasionally in short

16.

. Head and back bright red in

. Blackish vertebral

contact); prefrontals in contact or narrowly

separated; ear lobules short to moderately

long; lamellae under fourth toe 19-28;

hindleg 41-56% of SVL .. ye,

Nasals usually forming a median suture; pre-

frontals widely separated; ear lobules very

short; lamellae under fourth toe 16-20;

hindleg 33-41% of SVL ..

. Usually only one presubocular; plantars oppo-

site fourth toe enlarged and keeled; pre-

frontals usually in contact... .. 2... ..

Presuboculars 2; plantars uniformly granular;

prefrontals usually separated . .

life (fading to

pale green in alcohol) .. .. .. .. .. .,

Head and back pale pink or brown .. ..

stripe and laterodorsal

variegations persistent in adults: lamellae

under fourth toe usually fewer than 23:

hindleg usually less than 47% of SVL . ..

Dorsal pattern obsolescent; lamellae under

fourth toe usually more than 23; hindleg

usually more than 47% of SVL ..

Head and back pinkish white; blackish upper

lateral stripe enclosing a series of pale spots

Head and back pale brown; blackish upper

lateral stripe reduced to a series of vertically

elongate, rectangular blotches .. .. .. ..

February, 1971

strauchii

schomburgkii

brooksi aranda

brooksi. brooksi

brooksi euclae

brooksi taeniatus

Ctenotus pantherinus ocellifer ( Boulenger)

Synonymy, diagnosis and description: See Storr (1969, 1970).

Distribution: Northwestern interior.

and Northern Territory.]

Musgrave Range (AM 17245); 98 mi

[Extralimital in Western Australia

South Australian material: Mt. Davies (AM 17316); Erliwunyawunya,

. N of Cook (WAM 34522).

STORR—GENUS CTENOTUS IN SOUTH AUSTRALIA 5

Ctenotus robustus Storr

Diavnosis; A member of the Jesveurii group, distinguishable from

saxatilis and helenae by sharper brow and more strongly developed pattern,

e.#., white midlateral stripe extending forward unbroken to car aperture,

whence if continues narrowly forward to lores.

Distribution: Southeastern and central districts, north and west lo

Bute. [Extralimital in eastern Australia from northeast Queensland to

Victoria and in Northern Territory. |

Peseription: Snout-vent length (mm): 40-82 (62.7). Length of

appendages (% SVL): tail 167-204 (188), foreleg 24-28 (26.2), hindleg

37-47 (42.6).

Nasals separated, not grooved, Prefrontals in contact (except in two

specimens where separated by azygous scale), Supraoculars 4, first 3 in

contact with frontal, first much smaller than second, Supraciliaries 8-11]

(8.8), fourth to penultimate very small and tending to be concealed by

moderately sharp brow. Palpebrals 9-14 (10.7). Second loreal 1.2-2.1

(1.44) times as wide as high, Upper labials 8 (rarely 9), second to fourth

about twice as high as wide. Ear lobules 3-5 (3.6), acute in adults, first or

second largest. Nuchals 3 or 4. Midbody scale rows 28-34 (30.4).

Laumellae under fourth toe 19-23 (20.9); proximal lamellae divided and

subtubercular, remainder smooth or widely callose.

Dorsally olive brown, darker on head, paler on tail. Black vertebral

stripe from nape to proximal part of tail, nearly as wide as a paravertebral

scale, narrowly edged with creamy white. Whitish dorsolateral line from

brow to middle of tail (on which it is wider and suffused with brown),

murgined above by black laterodorsal stripe. Upper lateral zone blackish

brown, enclosing a series of moderately large brownish-white spots;

represented on tail by pale, dark-edged stripe. Whitish midlateral stripe

from ear aperture to middle of tail, partly interrupted by thigh and sending

down brunch to insertion of arm: anteriorly represented by fine line curving

under eye to lores. Lower lateral zone narrow, greyish, flecked with

whitish marks,

South Australian material: Bute (SAM 1705); Mt. Osmond (SAM

10006); Waterfall Gully (SAM 9983-6, 9992, 9994-6); Encounter Bay

(SAM 1697, 10010); Avenue Range (SAM 3789).

Ctenotus saxatilis Storr

Diagnosis: A member of the /esuewrii group with dark yertebral stripe

considerably narrower than a paravertebral scale and yery narrowly pale-

edged; white dorsolateral line; dark upper lateral zone with indistinct whitish

mottling; white midlateral stripe seldom extending anteriorly as far as arm,

6 REC. S. AUST. MUS, 16 (63) 1-18 Fehvitary, (971

Distribution: Far north. [Extralimital in the Northern Territory. |

South Australian material; Erliwunyawunya, Musgrave Range (AM

7246-7); Oodnadatta (SAM 43),

Ctenotus brachyonyx sp. noy.

Holotype: D 13944 in National Museum of Victoria, collected by

J. L. Hayward in June, 1969, at Annuello, Victoria (34 °52’S, 142°49’E).

Diagnosis; A member of the Jesweurit group with reduced colour

pattern and extremely short claws, Further distinguishable from saxatilis

by unmargined vertebral stripe and by some indication of pale subocular

line, from fielenae by presence of dark laterodorsal stripe and pale dorso-

lateral and midlateral stripes, and from rehustuy by number and nature of

supraciliaries (fewer than 8, fourth to penultimate not greatly smaller than

others and not tending to be hidden by brow).

Distribution: Valley of the Murray River. [Extralimital in north-

western Victoria.]

Description (based on all specimens); Snout-vent length (mm);

73-83 (79.4), Length of appendages (“% SVL): tail 181-214 (194),

foreleg 23-25 (24,3), hindleg 38-40 (39.3).

Nasals narrowly separated, not grooved. Prefrontals usually in contact

(very narrowly separated in one specimen), Supraoculars 4, first 3 in

contact with frontal. Supraciliaries 6 or 7. Palpebrals 8-12 (10.5).

Second loreal 1.4-2.0 (1.62) times as wide as high. Upper labials 8 or 9.

Ear lobules 2-5 (3.8), subacute or truncate, rather small. Nuchals 2 or 3.

Midbody scale rows 30-32 (30.6), Lamellae under fourth toe 22-24

(23,2), widely callose.

Dorsally (South Australia) brown or (Victoria) olive grey. Black

vertebral stripe moderately wide, beginning narrowly on nape and ending

abruptly at base of tail; litte or no indication of pale edge, Indistinct

dorsolateral stripe individually varying from whitish to nearly as dark as

ground colour, margined above by narrow black laterodorsal stripe beginning

well behind vertebral stripe and usually ending a little before it. Upper

lateral zone a little darker than ground colour, with or without blackish

dots or whitish flecks. Pale midlateral stripe, indistinct owing to suffusion

with ground colour, extending back on to base of tail after being wholly or

almost wholly interrupted by thigh; represented anteriorly by short fine line

curving below eye. Lower lateral zone paler than ground colour.

Paratypes: South Australia—Purnong (NMV D 3074, 5295-6),

Victoria—Cowangie (NMV D 12026); Hattah Lakes (NMV D [4213).

STORR—GENUS CTENOTUS IN SOUTH AUSTRALIA 7

Ctenotus helenae Storr

Diagnosis: A greenish member of the Jesueurii group with dark

vertebral stripe very narrow and not pale-edged, and little or no indication

of white dorsolateral and midlateral stripes.

Distribution: Extreme northwest (Tomkinson Range). [Extralimital

in Western Australia and Northern Territory. |

South Australian material: Mt. Davies (SAM 5317, 10016).

Ctenotus regius sp. nov.

Holotype: R 24492 in Western Australian Museum, collected by

G. M. Storr and A. M. Douglas on 4 October 1964 at Kingoonya, South

Australia, in 30°54’S, 135°19’E.

Diagnosis: A member of the leonhardii group differing from C.

leonhardii and uber by fully developed white midlateral stripe (extending

forward to lores) and low number of midbody scale rows (28 or fewer ).

Distinguishable from the superficially similar robustus by grooved nasal and

nature and number of subdigital lamellae.

Distribution: Interior of South Australia and northwestern Victoria.

Description (based on all specimens): Snout-vent length (mm):

37-74 (60). Length of appendages (% SVL): tail 191-252 (219);

foreleg 24-30 (26.9); hindleg 44-58 (51.3).

Nasals separated (rarely in short contact), weakly grooved, Pre-

frontals in contact. Supraoculars 4, first 3 in contact with frontal.

Supraciliaries 7 (rarely 8). Palpebrals 9-14 (11.3). Second loreal 1.0-2.1

(1.64) times as wide as high. Upper labials 8 (rarely 7 or 9). Ear lobules

3-6 (4.7), acute in adults, obtuse in juveniles, second or third largest.

Nuchals 3-6 (4.0). Midbody scale rows 26 or 28 (rarely 24). Subdigital

lamellae slightly compressed, 23-30 (26.0) under fourth toe, each with a

dark brown obtuse keel or narrow callus.

Dorsally brown, darkest on back, more olive on head, paler and reddish

on tail. Narrow blackish-brown vertebral stripe from nape to base of tail,

narrowly margined with white. Narrow but conspicuous white dorsolateral

line from brow to about middle of tail (on which it gradually merges with

background), widely or narrowly margined above with blackish brown (this

margin or laterodorsal stripe rarely enclosing a series of pale dots or short

dashes). Upper lateral zone dark brown or reddish brown, enclosing two

(occasionally one) longitudinal series of whitish dots or small longitudinally

elongate spots; represented on tail by pale, dark-edged stripe. White

8g REC, S, AUST. MUS., 16 (6): 1-15 February, 1971

midlateral stripe from lores to middle of tail, straight and moderately broad,

interrupted by ear aperture and partly by thigh; margined below (on body)

by a brown stripe which occasionally encloses some spots.

Paratypes: South Australia—Lambina (NTM 1552); Lake Eyre

(NMV D31L15, 3118); Killalpaninna (SAM 759); Goyders Lagoon (SAM

10342); Lakes Mulligan and Callabonna (SAM 9988-91); 6 mi. E of

Vokes Hill (WAM 36605); Ooldea (NMV D352); “Overland Railway”

(NMV D3045); Kingoonya (WAM 24493); Andamooka Ranges (SAM

2788, 10024-6); Yudna Swamp, Moralana (SAM 3177, 10013-4); Mern

Merna (SAM 2657. 10028-9, 10031-3); “Waterfall Gully” (SAM 9993).

Victoria—Cowangie (NMY D12027); Irymple (NMV D&8406): Karawinna

(NMV DS5651); Mildura (NMV D8138); Red Cliffs (NMV D8546).

Ctenotus leonhardii (Sternfcld)

Diagnosis: A member of the leonhardii group with nasals usually in

contact and prefrontals usually separated. Further distinguishable from

regius by failure of white midlateral stripe to extend forward to level of

arm (let alone to ear aperture); and from wher by absence of laterodorsal

series of pale spots,

Distribution: Far north. [Extralimital in| Northern Territory and

Western Australia. |

South Australian material: Erliwunyawunya, Musgrave Range (AM

17248); "50 miles south of South Australian border” [presumably between

Granite Downs and De Rose Hill] (SAM 9956).

Ctenotus uber orientalis subsp. nov.

Holotype: D825 in National Museum of Victoria, collected by

W. A. Hall in 1911 at Ouyen, Victoria, in 35°05’S, 142°19’E.

Diagnosis: Differing from C. u. uber of Western Australia (Storr

1969) in its well-developed vertebral stripe and shorter tail. Distinguishable

from leonhardii by presence of laterodorsal series of pale spots and absence

or feeble development of white midlateral stripe.

Distribution: Northern interior, east of the Nullarbor Plam and Great

Victoria Desert, south to central Eyre Peninsula and the Murray Mallee.

[Extralimital in northern Victoria, western New South Wales and south of

Northern Territory. ]

Description (based on all specimens): Snout-vent length (mm); 36-80

58.5). Length of appendages (% SVL): tail 151-214 (186): foreleg

3-30 (27.1); hindleg 39-55 (48.8).

(

STORR—GENUS CTENOTUS IN SOUTH AUSTRALIA 9

Nasals separated, Prefrontals separated or in contact. Supraoculars

4, first 3 in contact with frontal. Supraciharies 7 or 8 (6 in two Eyre

Peninsula specimens). Palpebrals 9-13 (10.4). Second loreal 1,0-1.8

(1,42) times as wide as high, Labials 8 (occasionally 7 or 9), Ear lobules

3-7 (4.6), acute in adults, obtuse in juveniles, second or third usually

largest. Nuchals J-7 (3.9). Midbody scale rows 28-34 (31.3). Lamellac

under fourth toc 19-29 (24.1), slightly to moderately compressed, cach with

a narrow dark callus.

Head and back olive grey, olive brown or dark brown; tail pale brown.

Blackish vertebral stripe from nape to base of tail, narrowly edged with

greyish White or pale brown. A laterodorsal series of pale dots or short

dashes. White dorsolateral line extending from brow back for varying

distances before breaking up into a series of short dashes or small spots.

Broad upper lateral zone dark brown or blackish, enclosing white flecks,

dots or short dashes which tend to align in three longitudinal series; upper

lateral zone represented on tail by dark stippling. White midlateral stripe

absent or indistinct on body (resulting from coalescence of fourth series of

dashes). Lower lateral zone greyish brown, enclosing one or two series of

whilish spots, dots or dashes, or flecked with white and pale prey,

Paratypes: Northern Territory—Macdonnell Ranges (SAM_ 10055),

South Australia—Dalhousie Springs (SAM 9735): between Pernatly and

South Gap Homesteads, Andamooka Ranges (SAM 2789, 9466-9); Lake

Gilles (NMV D9499); Blue Range Creck, Eyre Peninsula (SAM. 10122);

Lake Palankarinna, 62 mi. N of Marree (SAM 3618); Mern Merna (SAM

10017, 10027, 10030): Panaramitee, near Yunta (SAM 5738); “Buchsfeld,

near Adelaide” (Berlin 4719¢-d); Turners Well (SAM 23-4); Pinnaroo

(SAM 1507). New South Wales—Milparinka (SAM 10044), Victoria—

Red Clifls (NMV D2681, 2734); Ouyen (NMV D283-4, 826. 10032):

Mangalore (NMV D3433); Bright (NMV 15578),

Ctenotus atlas Storr

Diagnosis: A member of the faeniolatus group with a total of 10 whitish

dorsal and lateral stripes and lines on a blackish ground, Distinguishable

from quattuordecimlineatus by single pale line between midlateral stripe and

dorsolateral line, avd no pale dorsal line between puravertebral and dorso-

lateral lines,

Distribution: Southern sector of Great Victoria Desert (maflee-spinifex

yone) southeast through similar country round Lake Everard to eastern

interior of Eyre Peninsula. [Extralimital in Western Australia and New

South Wales (central-west), |

10 REC, 8. AUST. MUS,, 16 (6): 1-15 February, 1971

Description: Snout-vent length (mm); 38-69 (58). Length of

appendages (% SVL): tail 160-221 (204), foreleg 26-30 (27.9), hindleg

35-51 (42.7),

Nasals in short contact. Prefrontals in long contact. Supraoculars

normally 4, with first 3 contacting frontal (5 with 4 in one specimen).

Supraciliaries 7 (occasionally 8), Palpebrals 10-12 (11,0), Second loreal

1.2-2,0 (1.52) times as wide as high. Labials 8 or 9. Ear lobules 3-7

(5.5), first 2 or 3 usually very small and acute, Nuchals 1-3 (2.2),

Midbody scale rows 28-34 (30.3). Lamellae under fourth toe 20-28

(23.7), compressed, cach with a narrow dark brown callus or obtuse keel.

Head dark brown; tail brown, Back and sides blackish brown with a

total of 10 white or brownish stripes and lines: on each side a paravertebral

line from occiput to base of tail; dorsolateral line from brow to proximal

quarter of tail; upper lateral line from orbit to proximal quarter of tail;

narrow midlateral stripe from ear nearly to end of tail, partly interrupted

by thigh, extending forward indistinctly below eye to lores; ventrolateral

stripe from below and behind ear to groin, partly interrupted by arm.

Remarks: Apart from minor differences in coloration and number of

midbody scales, C. atlas is yery similar to C. guattuordecimlineatus, which

it replaces in slightly less arid regions. The two forms could therefore prove

to be conspecific. The specimen of atlas from 100 miles north of Cook

has the beginning of a pale line between the midlateral and dorsolateral

stripes, and the beginning of one between the midlateral and ventrolateral

Stripes; it is thus an intergrade with qualtuordecimlineatus which was

collected only 25 miles further north.

South Australian material; 100 mi. N of Cook (WAM 31863); 73 mi.

N of Cook (WAM 36603); 28 mi. NE of Maralinga (WAM 36645): 10 mi.

S of Everard (WAM 24515); Mt Wedge (SAM 9979): Hincks National

Park, Blue Range and Verran Hill (SAM 10123-8).

Ctenotus quattuordecimlineatus (Sternfeld)

Diagnosis; A member of the taenjolatus group with a total of 14 pale

dorsal and lateral stripes and lines on a blackish ground,

Distribution; Great Victoria Desert. [Extralimital in Western Australia

and the Northern Territory. ]

South Australian material: Near the Serpentine Lakes (WAM 34521 );

125 mi. N of Cook (WAM 31856-7).

STORR—GENUS CTENOTUS IN SOUTH AUSTRALIA i

Ctenotus leae (Boulenger)

Didenosis: A moderately small Crenotus with dorsal and lateral pattern

consisting of alternating dark and pale stripes; tail bright red and very long;

midbody scale rows usually 22 or 24. Distinguishable from all members of

the taeniolatus group by its sharply keeled, mucronate subdigital lamellac.

Distribution: Great Victoria Desert. [Extralimital in Western

Australia. ]

South Australian material: 10 mi. E of Golden Well (NMV D1565);

“Adelaide” (holotype).

Ctenotus schomburgkii schomburgkii (Peters)

Diagnosis: A member of the schomburgkii group with nasals usually

separated and back strongly striped with black. Further distinguishable

from strauchii by its longer appendages and less widely separated

prefrontals, and from brooksi by its two presuboculars and homogeneous

plantars,

Distribution: Northern interior. [Extralimital in Western Australia,

Northern Territory and western New South Wales. |

Description: Snout-vent length (mm): 25-50 (41). Length of

appendages (% SVL): tail 165-213 (196), foreleg 25-33 (27.8), hindleg

44-55 (49.4).

Nasals narrowly separated (occasionally widely separated or in very

short contact). Prefrontals narrowly separated or in short contact. Supra-

oculars 4, first 3 in contact with frontal. Supraciliaries usually 7, often 6,

Palpebrals 7-11 (9.3). Second loreal 1,.5-2.3 (1.98) times as wide as high.

Labialsy 7 (occasionally 8). Ear lobules 2-4 (3.2), short and obtuse, first

usually much the largest. Nuchals 3-5 (3.7). Midbody scale rows 24-28

(25.9), Lamellae under fourth toe 20-26 (22.5), each with a fine, sharp,

dark, mucronate keel.

Head, foreback and forelegs bright reddish brown (changing to greenish

in alcohol). Hindback, tail and hindlegs yellowish brown. Black vertebral

line from nape to proximal quarter of tail. Black dorsal line from nape to

base of tail, often breaking up into a series of dots or dashes. While

dorsolateral line from brow to base of tail, margined above with black,

Black upper lateral zone enclosing an irregular series of spots similar in

colour to corresponding part of dorsum: represented anteriorly by a narrow

loreal streak. White midlateral stripe from snout to base of tail, interrupted

by car aperture and partly by thigh. Lower lateral zone like upper but

narrower and less distinctly spotted.

12 REC, S, AUST. MUS, 16 (6): 1-15 February, V7

South Australian material: Mt, Davies, Tomkinson Range (SAM 5318,

10015); Ernabella (AM 17526); 6mi, E of Vokes Hill (WAM 31827-8);

15 mi, E of Emu (WAM 36609): 24 mi, NE of Maralinga (WAM 36640);

150 mi. N of Cook (WAM 36607-8): 144 mi, N of Cook (WAM 36604):

103 mi, N of Cook (WAM 36661); Ooldea (SAM 679, 10012); “Overland

Railway” (NMV D2667, 3063, 3065); 10 mi. of Golden Well, Overland

Ruilway (NMV 1995, 1567); Kychering Soak (NMV D2671); Kingoonya

(SAM 994, L0O341); Birthday Well (SAM 3050); Mullaroo Peninsula

(SAM 3054); Moralana (SAM 3182): Blesing Reserve (SAM 9190);

“Buchsfeld, near Adelaide” (lectotype and paralectotype, Berlin 4713a-h).

Ctenotus strauchii (Boulenger)

Diagnosis: A member of the schomburgkii group with extremely short

appendages and car lobules, and little or no dorsal pattern.

Distribution; Northeastern interior, west to Lambina and south to

Mern Merna. |Extralimital in Northern Territory, Queensland and New

South Wales. |

Description (based on all specimens cited below): Snout-vent length

(mm): 41-52. (46). Length of appendages (%@SVL); tail 126-160

(148), foreleg 21-27 (23.9). hindleg 33-41 (37.3).

Nasals forming a median suture (narrowly separated in one specimen).

Prefrontals moderately to widely separated. Supraoculars normally 4, with

first 3 in contact with frontal (5 with 4 in one specimen), Supraciliaries

6 or 7 (9 in one specimen). Palpebrals 8-11 (9.2). Second loreal 1.4-1.9

(1.59) times as wide as high. Labials 8 (occasionally 7 or 9). Ear lobules

1-4 (usually 3: mean 2.9), very short and obtuse, first or second basally

much wider than others. Nuchals 2-4 (3,2), Midbody scale rows 26-30

(27.9), Lamellae under fourth toe 16-20 (18.0), euch with a sharp,

mucronate keel,

Dorsally reddish brown. Vertebral stripe faint, narrow and pale-edged;

or absent, Dark laterodorsal markings variable: sometimes arranged as

small blotches or cross-bars, sometimes as a stripe enclosing a series of pale

spots, Whitish dorsolateral line from brow to base of tail, Upper lateral

zone variable: essentially consisting of narrow vertical or oblique, alternating

dark and pale blotches; but fine transverse connections between dark blotches

so frequent that zone could often be described as a dark stripe enclosing two

series of pale spots. White midlateral stripe extending narrowly forward

to below eye after bending sharply up and down around top of ear aperture.

Lower lateral zone reduced to a narrow faint streak or absent.

STORR—GENUS CTENOTUS IN SOUTH AUSTRALIA 13

Material; Northern Territory—Tempe Downs (NMV_ D280);

Charlotte Waters (NMV D946). Queensland—Birdsville (QM J9743).

South Australia—Lambina (NTM 1548); 10 mi. N of Clifton Hills (SAM

10345); Mern Merna (SAM 2611, 2646, 10018-23). New South Wales—

Milparinka (SAM 9930, 10042-3).

Ctenotus brooksi brooksi (Loveridge)

Diagnosis: Species breoksi distinguishable from all other members of

schomburgkii group by single presubocular and enlarged, keeled plantars

opposite fourth toe. Subspecies broeksi distinguishable by red head and

back, and little or no indication of dorsal and lateral pattern apart from an

upper lateral series of small, blackish, rectangular blotches.

Distribution: Great Victoria Desert. [Extralimital in Western Australia

and Northern Territory.|

South Australian material: Ooldea (SAM 3217); 10 mi. E of Golden

Well, Trans-Australian Railway (NMV D1566),

Ctenotus brooksi aranda Storr

Diagnosis: Agreeing with C. b. brooksi in red head and back, but

differing in fewer subdigital lamellae and stronger pattern (including pale-

edged vertebral stripe and blackish laterodorsal variegations ).

Distribution: Sand dunes of the Lake Eyre Basin, west to Dalhousie

and south to Marree, [Extralimital in adjacent deserts of Queensland and

Northern Territory. |

Description: Snout-vent length (mm): 28-55 (46). Length of

uppendages (% SVL): tail 140-163 (150), foreleg 24-32 (27.1), hindleg

41-50 (44.4).

Nasals in short contact or narrowly separated. Prefrontals forming a

median suture. Supraoculars 4, first 3 in contact with frontal. Supra-

ciliaries 6 or 7 (6.3). Palpebrals 9-11 (9.6). Second loreal 1.6-2.5 (2.15)

times as wide as high, Usually only one presubocular, occasionally 2.

Labials 7 or 8. Ear lobules 3 or 4, obtuse or subacute, second usually

largest. Nuchals 2-8 (mostly 3 or 4; mean 3.7). Midbody scale rows 26.

Lamellae under fourth toe 19-22 (20,2).

Seuth Australian material: Dalhousie (NMY D207); Lake Eyre shore-

line, including Hunt Peninsula and Madigan Gulf (SAM 3338, 3732, 3769,

9921, 10007, 10041, 10051, L0058-9); Marree (SAM 4704).

14 REC. S. AUST. MUS.,. 16 (@): 1-15 February, 1971

Ctenotus brooksi taeniatus (Mitchell)

Diagnosis: Generally similar to C. b. aranda, including relatively short

appendages, low number of subdigital lamellae, and colour pattern; but

differing m brown (rather than red) dorsal ground colour, separated (rather

than contiguous) prefrontals, and weakly keeled subdigital lamellae.

Distribution: Lake Torrens Basin,

Remarks: More material is required for determining whether the high

number of supraoculars (5) and supraciliaries (8) in the only known

specimen (SAM 2803) is diagnostic for this taxon. Each of these counts

can occur in other subspecies of brooksi, but only rarely and singly,

A specimen of C. schomburgkii from Alice Springs (NMV D166)

with the unique combination of 5 supraoculars, 8 supraciliaries and 8

labials was wrongly identified as taeniatus by Storr (1970: 107); in so doing

taeniatus was wrongly described as having two presuboculars.

Ctenotus brooksi euclae subsp. noy,

Holotype: R287 in Western Australian Museum, collected in 1914 by

W. B. Alexander at Eucla, Western Australia, in 31°43’S, 128°53’E.

Diagnosis; Differing from all other subspecies of brooksi by whitish

back strongly patterned with black, upper lateral zone a black stripe

enclosing whitish spots, and broad white midlateral stripe.

Distribution: White sand dunes bordering the Great Australian Bight

between Eyre, Western Australia, and Point Sinclair, South Australia.

Description: Snout-vent length (mm): 26-51 (43). Length of

appendages (% SVL): tail 153-193 (173); foreleg 24-31 (26.8); hindleg

45-54 (48.3).

Nasals narrowly separated (occasionally in short contact). Prefrontals

in contact or narrowly separated (sometimes by a small azygous scale).

Supraoculars 4, with first 3 contacting frontal (5 with 4 in one specimen).

Supraciliaries usually 7, occasionally 6, rarely 8. Palpebrals 9 or 10

(occasionally 11). Second loreal 1.4-2.3 (1,84) times as wide as high.

Labials 7 or 8. Ear lobules 2-5 (3.6): obtuse in juveniles; acute, subacute

and truncate in adults; second or third usually largest. Nuchals usually 3;

oceasionally 0, 2 or 4. Midbody scale rows 26 or 28, rarely 24 or 30).

Lamellae under fourth toe 19-25, each with a fine, sharp, dark, mucronate

keel.

STORR—GENUS CTENOTUS IN SOUTH AUSTRALIA 15

Dorsally a very pale flesh colour in life (E. R. Pianka, pers. comm.),

fading in alcohol to creamy white. Narrow, black, white-edged vertebral

stripe. On each side of dorsum two more blackish stripes, as wide as

vertebral but anteriorly wavy and posteriorly breaking up into variegations.

Upper lateral zone blackish with an irregular series of pale spots variable

in size and shape; upper edge of zone continuing forward through orbit to

nostril as a dark narrow streak. Broad white midlateral stripe from ear

aperture to hindleg. Lower lateral zone narrow, variably marked with pale

grey. Upper surface of limbs boldly streaked with black.

Paratypes: Western Australia—4 mi. N of Eyre (WAM 34478-80);

Eucla (WAM 288; ERP 13702, 13704, 13713-4, 13716, 13721-2,

13729-33, 13735-6, 13738-40, 13745, 13750-1, 13753, 13761, 13763-6,

13788). South Australia—Point Sinclair, 13 mi. S of Penong (WAM

36559).

REFERENCES

Storr, G. M., 1969: The genus Ctenotus (Lacertilia, Scincidae) in the Eastern Division of

Western Australia. J. Roy. Soc. W. Ausi., 51: 97-109.

Storr, G. M., 1970: The genus Ctenotus (Lacertilia, Scincidae) in the Northern Territory.

J. Roy Soc. W. Aust., 52: 97-108.

RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

ACARINE AND OTHER MICROFOSSILS

FROM THE MASLIN EOCENE, SOUTH

AUSTRALIA

By R. V. SOUTHCOTT and R. T. LANGE

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 7

1 June 1971

ACARINE AND OTHER MICROFOSSILS FROM THE MASLIN EOCENE,

SOUTH AUSTRALIA

BY R. V. SOUTHCOTT AND R. T. LANGE

Summary

The animal (invertebrate) component of a microfossil assemblage from a carbonaceous clay

deposit, North Maslin Sands, South Australia, is described. The deposit is placed in the Lower

Middle Eocene.

The assemblage contained, among numerous fungal, pollen and leaf cuticular remains, many setae

similar to those of modern Acarina and other arthropods. Also there were a number of claws,

probably mainly tarsal, from the same fauna. The deposit contained a fossil eriophyid mite, the first

fossil of this family to be described. This rust mite is placed in a modern genus, and described as

Aculops keiferi sp. nov. A fossil rotifer is also recorded from the same assemblage, and briefly

described as ?Keratella sp.

ACARINE AND OTHER MICROFOSSILS FROM THE MASLIN

EOCENE, SOUTH AUSTRALIA

By R. V. SOUTHCOTT anp R. T. LANGE

SUMMARY

The animal (invertebrate) component of a microfossil assemblage from

a carbonaceous clay deposit, North Maslin Sands, South Australia, 1s

described, The deposit is placed in the Lower Middle Eocene,

The assemblage contained, among numerous fungal, pollen and leaf

culicular remains, many setae similar to those of modern Acarina and other

arthropods, Also there were a number of claws, probably mainly tarsal,

from the same fauna. The deposit contained a fossil eriophyid mite, the

first fossil of this family to be described. This rust mite is placed in a

modern genus, and described as Aculops keiferi sp. nov. A fossil rotifer

is also recorded from the same assemblage, and briefly described as

?Keratella sp.

INTRODUCTION

The first part of this paper, by R. T. Lange, reports the discovery,

isolation and preparation of the fossils, and their palaeontological associations

and age. The second part, by R. V. Southcott, deals with the systemutics

of the fossils and related issues.

PART I—DISCOVERY, ISOLATION AND AGE OF THE FOSSILS

The present microfossils were included in a lens of carbonaceous clay

which was uncovered during 1966 by excavating machinery in a deep sand

quarry of the A.B.M, Company at Maslin Beach, South Australia. The

clay, discarded as dross, was observed by visiting geologists and brought to

the attention of Adelaide University palaeontologists. Collections were then

bagged in polythene for storage at Adelaide University, where they remain

available for study.

Invesigaultions of the clay were immediately undertaken, and are con-

tinuing. Research collections were assembled in the Botany Department,

where most research on the lens is proceeding. Different fossils are revealed

as different techniques are applied to the clay. When particular fossils are

revealed which are within the field of local specialists, such fossils are

submitted for specialist attention, Thus pollens from the lens have been

dealt with by the palynological section of the South Australian Department

of Mines. The present fossils, in view of their main affinity with mites,

were submitted to Dr. R. V. Southcott,

REC, 5, AUST, MUS,, 16 (7): 1-21 June, 1971

The geological section in which the lens occurred is not yet the subject

of publication, but is discussed in a Ph.D, thesis of the University of Adelaide

(Stuart, 1969), The lens was within the North Maslin Sands, which are

inferred to be of Eocene age by downward extrapolation from the Tortachilla

limestone (Lindsay, 1969), Palynological studies of the lens itself (Harris,

1966) indicate a correlation with the Burrungule Member of the Knight

Formation in the Gambier Embayment of the Otway Basin, southeast South

Australia, The Burrungule Member in turn correlates with the Hanikenina

aragonensis zone of the lowest middle Eocene, on the busis of planktonic

foraminifera (McGowran, Harris and Lindsay, 1969) lence good evidence

is provided for eatly middle Eocene age of the present microfossils,

Palaeontologically the present microfossils are a minor but significant

part of a complex microfossil assemblage which is itself intimately associated

with plant macrofossil beds. Many components of the total assemblage are

demonstrably of close ecological association, The present microfossils are

significant as the only animal remains thus far detected, and beeause they

correlate in part with fossils in the Moorlands coal,

The assemblage to which they belong is dominated by a mixed

angiosperm—gymnosperm flora represented by inflorescences, [ruiis and

seeds, anthers, pollen sacs, i sita and dispersed pollen, perianth parts. twigs

and leaves. There is no wood, and pteridophyte mucrofossils ate not yet

recovered, Angiosperm leaves are mostly simple and net-veined, and the

spectrum of their morphologies is not comparable with that of any present

day South Australian communities; similarly gymnosperm leaves are not

comparable with those in the South Australian region. Notably, there ts a

rich assemblage of leaf-inhabiling cryptogams (Lange, 1969) compurable

with that described by Ruinen (1961) from the phyllosphere of Surinam

jungle, and with that described from Eocene leaf beds in Tennessee by

Dilcher (1965). This in site microflora is dominated by foliicolous

ascomycetes of a nature described from present-day material by Hansford

(1946), Microthyriaceous germlings are almost ubiquitous, and palyno-

lovical preparations are almost dominated by dispersed fungal spores,

particularly of Demuatiaceae,

It is premature to propose definite biographical allinities and ecological

parallels for the assemblage, but some important points seem well established,

In the independent major respects (spectrum of leaf morphologies, nature

and mixture of angiosperm and gymnosperm remains, development and

morphology of the phyllosphere flora) the assemblage most resembles floor

litter from wet forest in which “typical” Australian elements such as

Eucalyptus and Callitris are poorly represented in the angiosperm and

gymnosperm floras respectively, yet where taxa like Casuarina are juxtaposed

Sourncore AND LANGHE—MASsLIN EOCENE MICROFOSSILS

with plants of ranalian affinity. Of the comparisons thus far made with

forest floor litters of the Australian region, the assemblage most resembles

litters from wet forest such as is found in Queensland north of Brisbane.

The present suite of microfossils was revealed by palynological methods,

and was observed to be almost ubiquitous in spore and pollen preparations,

lo which it gives a characteristic appearance. Fragments of clay about 2 cm.

square were pared to expose the uncontaminated core. Cores were

submerged for 24 hours in a mixture of equal volumes of HNO: and HCI

(commercial strength) sprinkled with KC1Os. The resulting sludge was

washed three times by centrifugation in distilled water. Sludge samples of

! cc, were dispersed by agitation in saturated ZnCl» solution and centrifuged

to partition organic particles from the clay and float them to the surface,

Small samples of the surface layer were transferred to slightly acidified water

and washed three times by centrifugation, satranin being udded before the

final centrifugation. Very small samples of the final precipitate, which was

entircly of organic fragments, were dispersed in phenol glycerine jelly for

microscopy. As a precaution against contamination, laboratory space was

inist-sprayed before use. As a check on contamination, the isolation

procedure was run blank, and slides filmed with paraflin oil were exposed

and observed microscopically. Occasional winged pollen were contaminants

on such slides, but no spores were detected in blank runs.

The entire procedure has been repeated in laboratories in South

Australia and in Germany, with very different environments for contamina-

tion. No setae were ever detected in blank runs; indeed, deliberate search

for such structures in preparations from nearby gardens and leaf litter have

failed to reveal similar structures.

REFERENCES

Dileher, D. 1. (1965): Epiphyllous fungi from eocene deposits in Western Tennessee, U.S.A,

Palavontographica B., We: 1-54.

Hansford, ©, Gi, (1946): The foliicolous Ascomycetes, their parasites and associated fungi,

Imp. Mycol. Inst. Kew, Mycol. Pap. 15, 240 pp.

Harris, W, K. (1966); Quart. geol. Notes geol. Sury. S. Aust, No, 20, 1.

Lange, R. T, (1969): Recent and fossil fungi of the Manginula-Sheriensix group, Aust J.

Bat, V7; 565-574.

Lindsay, FM, (1969): Bull, Geol, Surv. 8S. Aust, 42,

McGowran, Bo. barcis, W. K., and Lindsay, J. M. (1969): Early Middle Eocene Age of the

Masiin Bay Flora South Australia. S.A. Dept. Mines.

Ruinen, J, (1961). ‘The Phyllosphere I. An ecologically neglected milieu. Plant & Soil, 15:

81-109,

Stuart, Wo J. (1969): Stratigraphic and structural development of (he St. Vincent Tertiary

Basin, South Australia, Ph.D, Thesis, University of Adelaide,

4 REC. 8. AUST. MUS, 16 (7): [-21 Jane, V97)

PART U—SYSTEMATIC DESCRIPTION OF THE FOSSILS, AND

RELATED ISSUES

A, INTRODUCTION

The present study deals with a number of microfossils, some of them

recognizably acurine,

The finest specimen was a minute eriophyid mite, one of a specialized

family of mites which commonly cause rusts and blisters among present-day

plants. The state of preservation of this minute fossil (less than LOO» long)

allows generic and specific placing. It is recorded as a new species of the

genus Aculops Keifer.

Among the assemblage was a large number of setae, so similar in form

to those of modern Acarina that there does not appear to be any reason why

they should not be allotted to this Order with confidence. Some of them in

fact are so similar to setae of the modern superfamilies Erythraeoidea and

Trombidioidea of the superfamily Trombidiformes that it is considered that

this is the correct placement of these structures. In others however, it Is

not possible to allot them with the same degree of precision, and some appear

to belong elsewhere among the arthropods, e.g., possibly Insecta. A number

of varieties of setae are present, ¢.g., clavate setae, plumose setae, a squame,

sensory hairs and others.

In addition to the specimens mentioned, the assemblage included a

number of hairs which, as far as the present author is aware, do not

correspond to the hairs of modern arthropods, nor do they seem to be of

botanical origin, on the advice of botanical colleagues. It 1s proposed to

describe them also in this paper, so that they may be brought to the attention

of other workers.

B. REMARKS ON FOSSILIZATION OF ACARINA

It is unusual for Acarina and other small arthropods to be found us

fossils, and this may be attributed to their small size, fragile nature, and

presumably to their residual food content making them acceptable to micro-

scavengers, or susceptible to degradation by moulds or bacteria. In the cuse

of the Acarina the most extensive fossil fauna known is for the Baltic amber

(Oligocene), the mites and other small arthropods being preserved by

envelopment in the aromatic and indigestible resin of the pines, with its low

solubility in water. There is also one fossil erythraeoid larval mite recorded

for the Cretaceous amber of Canada (see Ewing, 1937; Southcott, 1961a,

pp. 373-4).

Sourneort AND LANGrE—Masiin Eocrsr MICROVOSSILS s

Alternative methods of preservation against destruction by scavenging

or degradation are believed to have occurred. Thus for the Australian

Permian insect fauna the excellent preservation of a number of priniitive

Winged insects has been attributed to suffocation and preservation in fine

voleanic dust (Tillyard, 1926, p. 470). However, these fossils are wings

only, and so far no acarine fossils have been reported from this Permian

fuuna, Some small arthropods have been discovered in the Rhynie chert

deposit of the Middle Devonian Old Red Sandstone, and among these was

the earliest mite so far described, Profacarus crani Hirst, 1923. The mode

of fossilization of these arachnids is speculative, but presumably there was

originally a carbonaceous deposit, later silicified. Nevertheless, it is not

unworthy of notice that very few acarine fossils have been recorded for the

coal deposits of the Carboniferous period, this being possibly partly due to

the conditions preceding fossilization not being conducive to the entombment

and ultimate preservation of these primarily terrestrial animals, or possibly,

simply because they have not been adequately looked for. Overall. the

known records of fossil Acarina are not numerous, making phylogenctic

speculation ubout the Acarina a somewhat fruitless subject (see André and

Lamy, 1937, pp. 34-5; Southcott, 196la, pp. 373-4).

The material submitted contained many specimens, in fact too many

for it to be possible to describe every one, and where types of setae are

extensively replicated, the paper has been limited to representative specimers

of cach kind.

I) addition there was present a fossil rotiter, identified as ?Keratella sp.,

which is illustrated and briefly described.

All specimens have been deposited in the South Australian Museuin

(section of Acarology, as all slides contain setae).

C. SYSTEMATIC DESCRIPTION OF A FOSSIL ERIOPHYID MITE

ACARINA

Family ERIOPHYIDAE Nalepa 1898

Genus ACULOPS Keifer 1966

Aculopy kelferi sp, nov.

Pigs, |, 2: Plate |

Description of deutogyne &

Length 97, width 38. Colourless, in safranin-stained mountunt on

receipt. Rostrum not clearly visible. Antapical rostral setae 5p long.

Shield 39» long by 17» wide, more or less triangular in outline but somewhat

distorted in the preparation: shield lobes rather long and sharp. Median line

“tH

6 REC. 8, AUST, MUS.. 16 (7): 1-21 June, 1971

of shield not discernible, but there is some indication of an admedian or

submedian line. Dorsal tubercles 22» apart, dorsal setae 30» long. Foreleg

present, length uncertain, Patellar seta (genuala) 20 long; tibia 5 long,

with tibial seta obscured; tarsus ca 10» long; featherclaw identifiable, 4.

long. Hindleg obscured. Coxae obscured. Abdominal thanosome with

about 18 strong resistant tergites, lacking granules or microtubercles.

Sternites possess ventral granules (microtubercles), which are pointed.

Lateral seta ca 10 long; first ventral seta ca 30» long, second ventral seta

5» long; third ventral seta ca 10» long. Accessory seta ca 32 long.

Genitalia not recognizable. Genital setae 14 apart, 16. long.

Locality

Maslin fossil beds, North Maslin Sands, South Australia.

Age

Lower middle Eocene.

Remarks

Although certain parts of this rust mite are lost, sufficient of its structure

has been preserved, for classification to the species level.

dorsal

oN te te

yenwel

Sebo

baller

Drone

accegsery

Orta

100

Figs. 1, 2. Aculops keifert sp. nov., Holotype. 1. Dorsal view, entire. 2. Ventral

view, entire. Two pollen grains are also illustrated,

SouTHcorTT awD LANGE—MASLIN Focene MICROFOSSILS 7

The specimen, by virtue of the fact that it lacks granules (microtubercles )

on the tergites, is considered as a deutogyne °. The deutogynes (which ure

females) lack the more specialized processes of the protogynes, and are

thereby less suitable for taxonomic description. However, as only one

specimen is available and as not only is it the first fossil eriophyid to be

discovered, but is also recognizably different from any modern known species

of eriophyid, it requires description and appropriate taxonomic status. On

the advice of Mr. Keifer it is placed in the genus Aculops and described as

a new species. This is a genus whose affinities within the Eriophyidae are

somewhat uncertain, and which may later receive further subdivision,

The species differs from other known species of Aculops in the follow-

ing features: long rather sharp anterior shield lobe, rather long dorsal setae,

tergites strong and resistant (without granules). and with pointed veniral

granules,

The fossil came from a mixed gymnosperm and angiosperm deposit, and

as Mr. Keifer advises that deuterogyny is unknown from conifers it is deduced

it is a rust mile wssociated with angiosperms.

The present author wishes to place on record his indebtedness to

Mr. H. H, Keifer, of San Francisco, the great authority on eriophyid mites,

for very considerable help with the taxonomy and other aspects of the

eriophyid specimen, and it is with pleasure that the species is dedicated to him.

D. OTHER ARTHROPOD REMNANTS, INCLUDING ACARINE

Figs. 3-56

The preparations made from the deposit include a large number of

arthropod remnants, many of them appearing to be recognizably acurine.

Most of then are of setae, but a few are of claws, or of other structures.

Reference to Figs. 3-56 shows that most of these are sctue, many appearing

to be normal acarine body setae with varying degrees of ciliation, These

normal body setae include forms which are tapering and sword-like, while

olhiers are parallel-sided and terminally blunted while others show varying

degrees of terminal expansion. Some of the setae have a distinet angulation

near their origin. Some of the setae, however, are more specialized. Setae

figured in Pigs. 29-34, 47-53 are possibly varying types of sensory sctae, not

all of them corresponding to forms known to the present author from Acarina

or other arthropods, ‘The assemblage contains several specimens of tarsal

claws, resembling in some degree those of modern Acurina. although not

certamly placeable in that Order, One specimen shows a claw in a normal

Orientation to a set of tarsal setae, and some of the tarsal cuticle is faintly

visible (Fig. 38).

1971

June,

1-2]

16 (7):

MUS..

REC. 8, AUST.

pS SESE STG

SSNS

ee SS EEE,

Sa Se —-

TSS

ies

assemblage, some of them appearing to be of

Setae from the Maslin

-26.

Figs. 3

For detailed description see text.

acarine origin.

SouTrHeort AND LANGE—MASLIN EocenNr Mrcrorossits i]

The author's familiarity with certain groups of Acarina possibly tends

to bias him towards an acarine origin of many of these setae. Thus the setae

figured in Figs, 3-26 resemble so strongly those of modern Erythracoidea

and Trombidividea (Trombidiformes) that no particular difficulty is seen

in attributing them to mites of these superfamilies. The other structures

depicted, however, cannot be placed so readily (assuming the former attribu-

lion is correct), and probably these figures should be examined by a large

number of specialists of various groups of organisms in attempt to place

them, Whereas, for example, Fig. 46 represents a squame or squamous seta

similar to those of certain Acarina, it is nevertheless a distinct possibility

that it has come from some other arthropod group, ¢.g., insects such és

Coleoptera or Lepidoptera. On the other hand, the seta shown in Fig, 36

does not correspond to anything the author is familiar with among the

Acarina, but is matched by setae the author has noted on a coleopteron

(family Curculionidae, see later in the present paper and Fig. 66).

As the placement of these setae, or structures, is to a large extent a

matter Of opinion, it has been decided to deal with them in a purely

descriptive fashion, until more definitive remnants are obtained. Thus those

fizured will be considered seriatim:

(1) Description oF SETAE OR SETAE-LIKE STRUCTURES FIGURED IN THE

PRESENT PAPER

Figs. 3, 4. These are parts of the shafts of scobalae (sce Southcott 1961a.

b; 1963 for terminology employed), more or less cylindrical, with

lightly outstanding or adnate barbs.

Fig. 5. Scobala, 191, long, sword-shaped, angled at base, lightly barbed.

Fig. 6. Scobala, 153 long, complete, barbed, blunt-tipped.

Fig, 7, Tapering blunted scobala, 132» long, with blunt barbs, set slightly

swollen near base.

Figs. 8-11. Barbed scobalae, Nos, 10 and 11 being incomplete. In Fig. 11]

the shading shows the degree of staining with the safranin.

Pig. 12, Blunted strongly barbed scobala, 215p long.

Figs, 13-16, 18. 19. Barbed scobalae with angled bases.

Fig. 17, Similar to No. 7,

Pigs, 20-26, Clavate scobalae, of varying lengths and degree of clavation.

In some, e.g., 23, 26 there is some tendency to angulation near the

origin of the seta.

Figs, 27, 28. Barbed scobalae, somewhat unusual in appearance.

Figs, 29-31. Sensory setae (sensalue) with terminal constricted picce of

presumed specialized function,

10 REC. S. AUST. MUS., 16 (7): 1-21 June, 1971

Figs. 27-56. Further microfossils from the Maslin assemblage, comprising setae,

claws and other, not fully identified, structures. For detailed description see text,

Fig. 32. Seta, possibly of sensory function, with swollen proximal part.

Seta 96, long.

Fig. 33. Clavate sensilla-type of seta.

Fig. 34. Another clavate seta, possibly a modified sensilla; 54» long.

SouTucotr AND LANGE—MASLIN EOCENE MICROFOSSILS i

g. 35. Two plumose scobalae, similar to those commonly seen in modern

Trombidudae.

g. 36. Burr-like seta, 48» long. This seta is similar to one figured later

(see Fig. 66). from a modern coleopteron.

. 37. Scobala with few ciliations, 46p long.

. 38. Group of scobalae outlining a tarsus of a leg, with a tarsal claw

in situ (also figured in Plate 4).

#. 39. Tarsal claw, 41 long, with two ventral spurs.

. 40. Tarsal claw 31, long, with 8 lateral ciliations.

. 41. Simple tarsal claw.

. 42, Tarsal claw 14» long.

. 43, Tarsal claw 38 long.

. 44. Tarsal claw 30p. long.

. 45. Tarsal claw 39» long.

. 46. Scobala, a squame or squamala, 38» long (also figured in Plate 3).

. 47. Spinous presumably sensory seta, 102” long. Four-fifths of the

seta is optically active.

ge. 48, As 47, 50 long. One-third of the seta is optically active.

. 49. As 47, 68, long.

. 50, As 47, 57» long (tip apparently incomplete).

ig. 51. As 47, curved, 64» long. About four-fifths of its length is optically

active.

‘ig. 52. A more strongly curved seta, 81 long. About two-thirds of the

seta 1s optically active.

. 53. Similar seta, apparently broken off. Specimen is 66p long, and

the whole of the specimen figured is optically active.

. 54. Robust seta with light protuberances, classified as a scobala; 61p

Jong.

g. 55. Apparently similar to last, 45» long.

g. 56. Structure of uncertain affinity, ?seta, 75yp. long.

(2) OTHER Microrossit HAIRS OF UNCERTAIN ORIGIN

Bicorned structures

Figs. 57-59 show some peculiar bicorned optically active hairs, which

are not certainly identifiable as to origin. Some variety of structure is seen

within these. Thus Fig. 59 shows a more or less symmetrical hair, rather

like a pickhead in shape, and 121, across. In Fig. 58 is shown a similar

12 REC, S, AUST, MUS,, 16 (7): 121 Jie, VYTI

structure but with one of the branches extended to about three times the

length of the other, so as to be filiform. In Fig. 57 another variation ts

shown. In this damaged specimen only one arm is present, this residual arm

being attenuated, but ending in a small projection like an clonguted acorn.

It is not known where these structures have originated nor have several

biologists to whom they have been shown been able to place them. For the

present the non-committal term “birema” is proposed for them, for the

purpose of reference, pending more precise placement,

Linear hair-like structures

Another group of unidentified structures from the Maslin assemblage

is shown in Figs. 60-65. These are all strongly optically active, They are

linear, and ciliated or ornamented, and some are segmented,

Figs. 60-62 show unsegmented barbed hairs (Fig, 61 1s shown as

broken for the purposes of the illustration), Fig, 60 shows a form 375.

long; Fig, 61 represents a specimen 710 long; Tig. 62, 325, long.

Figs, 63-64 represent segmented linear hairs, the joints being expanded

in some cases to a head of three projecting angled lobes on one side (the

proximal side), but more distally to pointed spurs or ciliations at the distal

end of the segments (= proximal side of the joints), Fig. 63 shows a

hair 670» long; Fig. 64 shows one 184, long.

Fig. 65 shows a fine linear structure with enlargements, but without

ciliations or angulations, 200 long.

(3) COMMENTS ON THE IDENTIFICATION OF THE SETAE, AND COMPARISON

Witl MODERN MATERIAL

It would appear unlikely that a fossil deposit originating from terrestrial

arthropods would be restricted to a narrow taxonomic range, such as, for

example, a single order or even class of these animals. Since the material

submitted contained a good deal of vegetable material which had survived

the treatment in Schultz solution—pollens, fungul hyphae and spores, as

well as leaf cuticular remnants (many with stomata), ete—it would be

reasonable to expect it to contain chitinous remnants of a diversity of animals

such us are found on leaves of tropical and temperate angiosperms as well

as of the leaf and other litter below such a flora. Any worker who has

examined the leaf litter of temperate and tropical angiosperm forests must

have been struck by the variey of small animal life present, ¢.g.. mites,

collembola, other insects, small spiders (or even larger ones), centipedes,

millipedes, also polychaetes, not to mention small molluscs, leeches,

opilionids, and a variety of other creatures. Among the living leaves of the

forest the invertebrate fauna tends to be more restricted in its taxonomic

SOUTHCOTT AND LANGE—MASLIN EOCENE MICROFOSSILS 13

Figs. 57-59. Bicornuate hairlike microfossils or “biremae” (sing.: birema) from

the Maslin assemblage. For description see text.

Figs. 60-65.

Linear hair-like structures, of uncertain origin, from the Maslin

assemblage. For description see text.

100

SouTHCOTT AND LANGE—MASLIN EOCENE MICROFOSSILS 1S

range, and perhaps insects, mites and small spiders are the commonest small

animals present. While it is impossible to guess with any accuracy as to

what microfauna may have been present in a particular tropical Eocene

Australian forest, nevertheless it may have been expected that by the Eocene

all modern groups of arthropods favouring a similar habitat would have been

represented,

As most of the animal remnants among the assemblage were of setae,

the author has attempted to widen his knowledge of setal forms among the

arthropods by examining, for this paper, various small modern arthropods

and other possible sources of such hairs. Since rather similar setae may be

found in widely differing groups of arthropods, it would be unjustifiable to

erect new taxa where all that is available is a seta or claw, or some cuticular

fragments. Thus although a number of the setae depicted have the

appearances of acarine setae, this does not identify them as such for certain,

and even if so, it does not allow generic or specific placement. Squames,

for example, are found in various insect groups, such as Coleoptera and

Fig. 66. Setae from trochanter of Jeg of modern curculionid beetle

(family Curculionidae, sf. Cryptorhynehinae), 7Diethusa sp.. from

Mitcham, South Australia. Compare wilh Fig. 36 of the Maslin

assemblage (see further in text).

16 REC, 8S. AUST. MUS., 16 (7): 121 June, 197

Lepidoptera, as well as Acarina. In Fig. 36 was drawn a seta from some

unidentified group of arthropods. Examination of various modern Australian

insects has revealed a rather similar set of setae upon the trochanters of a

weevil, ?Diethusa sp, (identified G. F. Gross), specimen RVS:A1178, from

Mitcham, South Australia, 81.1970, R. V. Southcott (family Curculionidae,

subfamily Cryptorhynchinae), of which some are drawn in Fig. 66.

The student of these remnants cannot but be struck by the faithful

preservation of the physical structure of these minute fossils over this immense

period of time, these remarks applying not only to the external shape, but

wlso to such a physical characteristic as the preservation of the optical

activity of the specimen. Such a finding immediately prompts the question

as to what extent the chemical structure of these fossils is also preserved.

E. DESCRIPTION OF A FOSSIL ROTIFER, ?KERATELLA sp.

Text Fig. 67; Plate 4

In the Maslin assemblage was a structure accepting the safranin stain,

identifiable as a loricate rotifer and provisionally allotted to the genus

Keratella.

Fig. 67. ?Keratella sp., a fossil rotifer

(Order Monogonata, superfamily

Brachionoidea) from the Maslin assem-

blage. The structure e appears to be

a parthenogenelic egg at the posterior

pole of the animal, and possibly there

is a larger one in a lower plane (see

stippled outline, f).

SoOUTHCOTT AND LANGE—MASLIN BoceNe MICROFOSSILS 7

Description: Length (overall) 75, width 32, The specimen consists

of a central part which is a prolate spheroid, about 36» long, and with spines

projecting from each end. The structure is thus a horny lorica, smooth

externally, no sculpturing being identifiable. Five spines occur at the

anterior end (uppermost in Fig. 67), and there are three lorical spines ut

the posterior end (lowermost in Figure). The spines are strong, curved

inwardly, and more or less pointed toward the tip. No corona is visible.

Between the three posterior spines is a clear rounded structure, which appears

to be a parthenogenetic egg (e@), or possibly there are even two of these,

one larger (/) than the other (compare with e.g. Hyman, 1951, p. 142

(her Fig, 67F)). A fracture runs transversely across the main part of

the lorica,

Remarks

The specimen described appears to belong to Keratella or to some

allied genus of the superfamily Brachionoidea (order Monogonata), The

genus Keratella has been recorded as having the tendency for the lorica to

show great diversity of form, even within a single species, and in the past

many of such divergent forms have been described as different species.

Many members of the Keratella are marine species, The material studied

in the Maslin assemblage does not have a marine facies, even if we omit

such characteristic elements as pollens and angiosperm leaf stomata. How-

ever loricate rotifers not unlike Kerafella, speaking in a general sense, are

not uncommon in fresh water.

REFERENCES

Andre, M., and Lamy, FE. (1937): “Les idées actuelles sur li phylogénie des Acariens”

(Authors: Paris).

Raker, &, W.. and Wharton, G. W. (1952): An Introduction to Acarelogy (Miuemillan:

New York)

Ewing, H. EB. (1937): Insects and arachnids from Canadian amber: Arachnida, Order

Acarina. (/niv, Toronta Stud. Geol. 40: 56-62,

Hirst, S. (1923); On some arachnid remains from the Old Red Sandstone (Rhynie Chert

Bed, Aberdeenshire). dan, Mag. nat, Mist, (9)12: 455-24,

Hyman, Libbie NM, (1951): “The Invertebrates: Acanthocephala, Aschelminthes and

Entoprocia, ‘The pscudocoelomate Bilateria’ (Vol WL) (MeGraw Hill: New

York)

Keifer, Ho HW. (1952): The eriophyi mites of California (Acarina: Eriophyidae. Bull. Calif,

Insect Survey 21). 1-123,

Keifer, H. H, (1953): Eriophyid studies—XXI. Bull. Calif. Dept. Agric., 42: 68 (feste Keifer,

1966, p. 9),

Keifer, H. H. (1966); Briophyid studies B—21. Special Publ., Bureau Entom., Calif. Dept.

Agric.

Southeott, R, V. (196la); Studies on the systematics and biology of the Eryihracoidea

(Acarina). with a critical revision of the genera and subfamilies, Aust. J.

Foal,, 93): 367-610.

Southeott, R. V. (1961b): Description of Iwo new Australian Smirididae (Acarina), with

remarks on chaetotaxy and geographical distribution. Trans. Ray, See, §. Anse,

85: 133-53,

18 REC, S, AUST. MUS., 16 (7): 1-21 June, 1971

Southcott, R. V. (1963): The Smarididae (Acarina) of North and Central America and some

other countries. Trans, Roy. Soc. S. Aust., 86; 159-245,

Tillyard, R. J. (1926): The Insects of Australia and New Zealand (Angus and Robertson:

Sydney).

Plate 1.) Aculops heiferi sp. nov, Holotype, Photographed in three optical

planes by phase contrast (1), and normal illumination (R). SOOX.

SoutucoTr AND LANGE—MASLIN EOCENE MICROFOSSILS 19

Plate 2, Various setae from the Maslin assemblage, by phase contrast. S00X

Same of these setae are also shown in the text illustrations

20 REC. S, AUST. MUS., 16 (7): 1-21 June, (971

Plate 3. Various setae, a claw and other structures from the Maslin assemblage,

by phase contrast. S00X. Some ot these are figured also in the text-lgures,

MASLIN EOCENE MiICROFOSSILS 2

SOUTHCOTT AND LANGE

fF %

Fe BR +

pM)

Plate 4. Above: ?Keratella sp. a Joricate rotifer from the Maslin

assembjage, See further in teat. By phase contrast, 500X. Three

optical planes are shown. Below: Tarsal setae and claw, by phase

contrast, at differing contrasts. SOOX. (Also figured in Fig. 38, and

commented on in text.)

RECORDS OF THE

SOUTH AUSTRALIAN

MUSEUM

NEMATODE PARASITES OF OCEANICA.

Xll. A REVIEW OF HETERAKIS

SPECIES, PARTICULARLY FROM BIRDS

OF TAIWAN AND PALAWAN

W. GRANT INGLIS, GERALD D. SCHMIDT

and ROBERT E. KUNTZ

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 8

7th October, 1971

NEMATODE PARASITES OF OCEANICA.

XII. A REVIEW OF HETERAKIS SPECIES, PARTICULARLY FROM

BIRDS OF TAIWAN AND PALAWAN

BY W. GRANT INGLIS, GERALD D. SCHMIDT AND ROBERT E. KUNTZ

Summary

The following species of Heterakis are recorded from Taiwan and Palawan: H. variabilis Chandler,

1926, from Lophyra swinhoii, Bambusicola thoracica, and Syrmaticus mikado, Taiwan; H.

vulvolabiata Chandler, 1926, from Arborophila crudigularis, Taiwan; H. isolonche Linstow, 1906,

from Lophura swinhoii, Taiwan; H. beramporia Lane, 1914, from Gallus gallus, Taiwan; H.

spumosa Schneider, 1866, from (?) Rallina eurizonoides, Tatwan; and H. indica Maplestone, 1932,

from Surniculus lugubris minimus, Palawan; H. vexans sp. nov. is described and H. variabilis and

H. vulvolabiata are redescribed.

NEMATODE PARASITES OF OCEANICA. XII A REVIEW OF

HETERAKIS SPECIES, PARTICULARLY FROM BIRDS OF

TAIWAN AND PALAWAN

W. GRANT INGLIS, THe South AUSTRALIAN MUSEUM, ADELAIDE, SOUTH

AUSTRALIA

GERALD D. SCHMIDT. DEPARTMENT OF BIOLOGY, UNIVERSITY OF

NORTHERN COLORADO, GREELEY, COLORADO 80631, AND Nato

FELLOW, SOUTH AUSTRALIAN MUSEUM

and

ROBERT E. KUNTZ, DEPARTMENT OF PARASITOLOGY, SOUTHWEST

FOUNDATION FOR RESEARCH AND EDUCATION, SAN ANTONIO,

TEXAS AND U.S. NAvy MEDICAL RESEARCH UNit No. 2, TalPet,

TAIWAN, REPUBLIC OF CHINA

ABSTRACT

The following species of Heterakis are recorded from Taiwan and

Palawan: AH. variabilis Chandler, 1926, from Lophura swinhoii, Bambusicola

theracica, and Syrmaticus mikado, Taiwan; H. vulvolabiata Chandler, 1926,

from Arberephila crudigularis, Taiwan; H. isolonche Linstow, 1906, from

Lophura swinhoii, Taiwan; H. beramporia Lane, 1914, from Gallus gallus,

Taiwan; H. spumosa Schneider, 1866, from (?) Rallina eurizonoides,

Taiwan; and H. indica Maplestone, 1932, from Surniculus lugubris minimus,

Palawan; H. vexans sp. noy. is described and H. variabilis and H.

vulvolabiata are redescribed.

INTRODUCTION

The limits of the superfamily Heterakoidea are fairly well established.

The genera, with minor exceptions, are reasonably stable while even the

families and subfamilies are recognizably the same groups in the various

treatments of the superfamily. A major area of disagreement, however, con-

tinues to be the division of the genus Heterakis s.J. into two genera on the

equality or otherwise of the spicules.

Lopez-Néyra (1947) first used equality and inequality of the spicules

as a way of delimiting groups which he called Heterakis and Ganguleterakis

and has been followed in this by Skrjabin and his co-workers in several

publications (most recently Skrjabin, Schikhobalova, and Lagodovskaja,

1961). Because of criticism of the artificiality of this simple criterion,

Issued 7th October, 1971]

REC, 8. AUST. MUS. 16 (8)2 1-14 Oecvroher, 1971

Freitas (1956) attempted to refine the distinction by defining uncqual

spicules as those in which the difference m length of the spicules is at least

one third the length of the shorter and on this basis introduced a new name,

Raillietakis, for species with “equal” spicules, an urgument accepted by

Yamaguti (1961), Madsen (1950), Inglis (1958, 1967) and Chabaud

(1965) do not consider inequality of spicule length suflicient for the delimita-

tion of genera, even as refined by Freitas, although all recognize that the

structure and relative lengths of the spicules can be valuable in delimiting

species,

The disagreement is partly because of the uniformity of the structures

of the male tail in Heterakis, which supply almost all the characters useful in

delimiting species, and partly, we suspect, because of a feeling that the

genus is too diverse to be left as one taxon, However the inequality of

spicule length is so obviously a bibliographic character that its value must be

considered dubious. However, those who, like us, oppose its use have not

established their case in detail, and the opportunity to do so has now arisen

following the study of specimens of Heterakis, particularly from birds in

Taiwan and Palawan, as well us specimens, particularly type material, from

other hosts from other localities.

The former specimens were collected by R.E.K. and his associates of

NAMRU—No. 2 during investigations of the parasite faunas of Taiwan and

Palawan, The speciniens were fixed in hot 70 per cent alcohol, stored in

alcohol and glycerine, and cleared in glycerine or lactophenol.

MORPHOLOGICAL GROUPS

The posterior ends of male Heterakis are very similar, with an obvious

circular pre-cloacal sucker on the mid-ventral surface of the male body

anterior to the cloacal opening. The sucker has a distinct cuticular rim, with

a papilliform sense-organ on the posterior margin. There are broad caudal

alae supported by two pairs (sometimes three) of long pedunculate papillae

lateral to the sucker (parasuctorials), four pairs of such papillae lateral to

the cloacal opening (paracloacals) and one pair roughly half-way between

the cloacal opening and the tip of the tail (postcloacals). In addition there

are two pairs of sessile papillae around the cloacal opening (pericloacals) and

a group of two pairs at the posterior end of the tail, with the phasmids

immediately anterior to them, Variation in these papillae is usually restricted

to a reduction in the number of para-cloacals or to the presence of un

additional pair of peduneculate papillae between the para- and postcloacal

papillae.

Attempts have been made to delimit species on the basis of the dis-

tribution of these papillae and on the presence or absence of an additional

pair. but i Most cases this has later been shown to be unsound.

NEMATODE PARASITES OF OCEANICA |

In contrast, the structure of the spicules supplies the most obvious

characters which delimit species and there has never been any doubt about its

value. Similarly, there is no doubt that groups of species exist in which the

spicules. are either equal or unequal; the disagreement is about the value of

separating such groups and the way in which the groups should be diagnosed.

The major argument about such groupings, other than simple belicf state-

ments, is that other groups can be recognized on other features of the spicules,

as was pointed out very briefly by Inglis (1967). This is now considered in

detail and three major groups can be recognized.

Heterakis dispar—Group

This group consists of the nominal species H. altaica Spaul, 1929;

A, brevispiculum Gendre, 1911; H. caudata Linstow, 1906; H. circumvallata

Linstow, 1906; H, dispar (Schrank, 1790) Dujardin, 1845; H. hyperborea

Swinyard, 1931; H, papillosa (Bloch, 1782) Cram, 1927: A. silindae Sand-

ground, 1933; H. skarbilowitschii Kassimov, 1946; A, stvlosa Linstow, 1907;

H. tenuicauda Linstow, 1883: H. travassesi Khalil, 1932.

In all these nominal species the spicules are nonalate, equal in length

and identical in structure with spiral flanges on their posterior ends (Figs.

1-3: 5-7). These spiral flanges, which are difficult to see at low magnifica-

lion, have been overlooked im some descriptions and have been described as

hooks or barbs in others. In all cases, however, they are cuticular expansions

of the spicules which spiral round the tip.

Within this group of twelve names there appear to be only three actual

species: one characterized by a large goblet-shaped pre-cloucal sucker which

lies relatively close to the cloacal opening, and by a relatively long tail; a

second characterized by a similar sucker lying relatively far anterior to the

cloacal opening, and by a short tail so that the para-cloacal papillae appear

bunched together; and a third characterized by a long, narrow tail, relatively

long spicules and by a small pre-cloacal sucker which lies low on the surface

of the body,

The earliest names for these species are H. altaica, A. brevispiculum and

H. papillosa respectively with the synonyms listed below. However the

problem is by no means resolved and it is possible that more species can be

recognized or even that there is only one, particularly as “a/taica’’ is some-

what intermediate in form between the other two,

Provisional synonomies are (the name of an institution in parentheses

indicates that type specimens are lodged there and have been studied ):

Heterakis altaica Spaul, 1929, (British Museum (Nat. Hist.))

Synonymy: H. skarbilowitschii Kassimow, 1946,

4 REC. S. AUST. MUS.,, 16 (8): 1-14 October, 1971

Spaul overlooked the spiculal flanges, which are well figured by

Kassimow: it is possible that H, macrouwra Linstow, 1883, is this species,

Freitus (1956) refers H. altaica to Odontoterakis but this is completely

wrong.

Heterakis brevispiculum Gendre, 191 |

Synonymy: H. travassosi Khalil, 1932 (Tropeninstitut, Hamburg). H.

silindae Sandground, 1933 (Museum of Comparative Zoology, Harvard).

Heterakis dispar (Schrank, 1790) (Naturhistorisches Museum, Vienna)

Synonymy; ?Ascaris papillosa Bloch, 1782, H, caudata Linstow, 1906; H,

cireumvallata Linstow, 1906; H, /iyperborea Swinyard, 1931; (2?) H,

monticelliana Stossich, 1892; H. stylosa Linstow, 1907 (Instit. fiir Spez.

Zoo., Berlin): H. tenuicauda Linstow, 1883 (Brit. Mus. (Nat. Hist.) ).

In the most recent redescription of A. papillesa to be published, Madsen

(1950) failed to see the spiral flanges on the posterior ends of the spicules.

However one of us (W.G.I.) has examined Madsen’s material, as well as

specimens from the type hast (Oris tarda) in captivity and in the wild, and

flanges ure present on the spicules of all males studied. As there are no other

obvious differences between the two nominal species H. dispar is almost

certainly indistinguishable from H. papillosa. However, H. dispar is a name

very widely used, particularly in veterinary literature and it is preferable that

it remains unchanged. Further, the problem of delimiting the species of this

group is not fully resolved. We therefore prefer to retain H. dispar and to

treat H, papillosa only as a probable synonym,

In addition to these species we later describe a fourth, H. vexans sp. nov,,

which is most appropriately accommodated within this H. dispar-group,

although the flanges on the posterior ends of the identical spicules are non-

spiral,

Heterakis gallinarum—Group

This group contains H. beramporia Lane, 1914; H, bonasae Cram,

1927: Hl. bosia Lane, 1914: H. caudebrevis Popova, 1949; H. gallinarum

(Schrank, 1788) Madsen, 1949 (—-H, pediocetes Mawson, 1956 (Institute

of Parasitology, McDonald College)); H. indica Maplestone, 1932; H,

isolonche Linstow, 1906; H, paveris Maplestone, 1932 (— H. yamadori

Yamaguti, 1941); H. putaustralis Lane, 1914; H. variabilis Chandler, 1926;

H, valvolabiata Chandler, 1926,

In all these species the left spicule at least, and in some both, is alate or

obviously different from the right. The most diagnostic character of the

various species is the shape of the left spicule, of which the tip can be elabor-

ute.

NEMATODE PARASITES OF OELANICA 5

The major problem in this species group is the relationships between

and the delimitation of those species in which the right spicule is long and

slim while the left is usually short but always with broad alae and a relatively

simple posterior end. These worms appear to form a cline from species with

markedly unequal spicules at One extreme to species with equal spicules at

ihe other, At one extreme is H. gallinarum in which the left, terminally

hooked, spicule is much shorter than the right, and at the other extreme is

H. isolonche in which the spicules are about the same length, Intermediate

between the extremes are H. variabilis Chandler, 1926 (redescribed below),

H. putaustralis Lane, 1914 and H, benasae Cram, 1927. Basically similar to

H. gallinarum in having a short, left spicule are H. pavenis Maplestone, 1932

and H, indica Maplestone, 1932, but in both species the left spicule has an

elaborate tip.

This problem still awaits resolution but 1s the major reason we do not

accept a split of the genus Heterakis on the basis of the relative lengths of the

spicules, The remaining species listed are less happily included in this group

und might warrant treatment as a fourth species-group, Nevertheless we leave

them here ut present as a matter of convenience,

Heterakis alata

This group contains H. banecrofti Johnston, 1912; H. alata Schneider

1866 (— H, arquata Schneider, 1866 — H. skrjabini Cram 1927): H.

brasiliana Linstow, 1899 (Instit. fir Spez. Zoo., Berlin); H. nattereri

Travassos, 1913.

Group

In ull the species of this group the spicules are without alae and, in

most species, are slim, In none is there an claboration of the tip of the left

spicule and all occur in hosts in South America, except H. bancroefti which

was found in an Australian host.

REMAINING SPECIES

Among the remaining species usually referred to Heterakis, H, spumosa

Schneider, 1886, is probably the most widespread and best known. It is the

only species of Heterakis which occurs in mammals and is widespread in rats

throughout the World. It is characterised by equal and identical needle-like

spicules, three pairs of para-cloacal papillae and no “post cloacal” pair or,

another interpretation, has only two pairs of para-cloacal papillae of which

ihe more anterior pair is very large and may represent three fused papillae.

Because of these differences Lane (1914) described this species as

Ganvguleterakis gangula, having over-looked Schneider's earlier description.

Since then the generic name Ganguleterakiy has been used by some authors

for 4 group containing so-called “equal spicule” species of Heterakiy, This,

6 REC. & AUST, MUS., 16 (8): I-14 Ovrober, 1971

as argued above, we do not accept but recognize that H, spumosa is very

different from the typical Heterakis of birds and the recognition of a genus

for it alone could be accepted if one so desired.

H. macrospiculum Ortlepp, 1939, H. spalacis Margu, 1930) and

Ganguleterakis spalaxi Kozlov and Yangolenko, 1967, do not appear to be

species of Hererakis, It is possible that H. macrespiculum is an Africana

species and that the other two, which are probably indistinguishable, are

Ascaridia,

HOST AND GEOGRAPHICAL DISTRIBUTIONS

Species of the genus Herterakis occur widely in the cuecum of ground-

feeding, grain-eating birds throughout the world, with some species in water

fowl and a few other hosts. The members of the HM. alata-group are restricted

to birds in South America while the other two groups occur in birds in the

rest of the world, The sole exception is H, hancrofti Johnston, 1912, which

occurs in Australia but is morphologically a member of the South American

aluta-group. Such a geographical relationship is interesting in reinforcing

other Australian-Neotropical faunal relationships and so does not destray the

primarily South American relationships of the alata-group. It would, there-

fore, appear that the groups of species have arisen in response to geographical

separation,

Support for this is given by the genus Odontoterakis, restricted to South

America, in which the spicules are always simple, needle-like, and identical

in structure. In this its members resemble the South American H, alata-

group. The genus Pseudospidodera, in contrast, is restricted to India and

related countries and the spicules are unequal with the right long and thin and

the left short with broad alae (except in P. jnanendrae Chakravarty, 1938:

but this species is in need of redescription), as in the H. gallinarum-group,.

It 1s probable that Odontoterakis arose from the Heterakis species found

in South American hosts while Pseudaspidedera arose from the H. gallinarum-

group of species in South Asia. To this extent the species groups recognized

in Heterakis are supported not only by the morphological and geographical

data within the genus itself but also by the similar evidence supplied by the

species grouped in Pseudaspidodera and Odontoterakis.

If the genus Heterakis is to be fragmented on the basis of spicule

structure, rather than on the relative lengths of the spicules, part of it (the

gallinarum-group) could be grouped with Pseudaspidodera species, part of it

(the alata-group) with Odentoterakis while the remaining part (the dispar-

group) would be left as a distinet genus, Such groupings might reflect the

evolution of the group more accurately than the classification of Inglis

NEMATODE PARASITES OF OCEANICA 7

(1967), since Odontaterakis and Psendaspidodera probably evolved from

Heterakis species. Nevertheless, it is more reasonable and convenient to leave

the generic groupings as they stand with Hererakis as one cosmopolitan genus.

The relationships of the brevispiculum-group remain uncertain as they

could have arisen from either of the other groups. No decision on this is

possible at this time although the new species described below (H. vexans)

suggests an intermediate between the dispar- and alata-groups.

DESCRIPTIVE SECTION

All measurements are in microns unless otherwise stated and all

specimens have been deposited in the collections of the U.S, National

Museum Helminthological Collection, Beltsville, Maryland.

Heterakis vexans sp. nov.

(Figs. 8 and 14)

A single male was found among several H. vulvelabiata Chandler, 1926,

obtained from the caecum (?) of a Formosan hill partridge. and appears to

represent a new species.

DESCRIPTION

Morphology typical for genus. Lips (Fig. 8) rather small, with con-

spicuous papillae. Lateral alae prominent. Anterior end curved dorsad.

Male: 5.0 mm, long, 280 greatest width (1.0 mm. posterior to anterior

end). QOcsophagus (excluding pharynx) 665 long, with posterior bulb 112

vreatest width. Pharynx (measured from flange tooth to junction with

oesophagus) 48 long. Excretory pore 330 from anterior end, Nerve ring

216 from anterior end. Precloacal sucker (Fig. 14) 51 long, 48 wide

(ineasured across outer margins of sclerotized wall), posterior edge 64 from

unus, Tail 340 long. Spicules (Fig. 14) subequal, very stout, with similar

tips, each with sharp, recurved hook at tip and subterminal inflation in lateral

view, Spicules lacking alae, but each with narrow longitudinal flange along

subterminal swelling. Right spicule 450 long, left spicule 380 long. Caudal

papillae Lypical of genus but with only three pairs of paracloacals.

Female: Unknown.

Type host: Formosan hill partridge, Arborophila crudigularis

(Swinhoe, 1864). (Phasianidae: Galliformes. )

Location: Probably caecum.

Type locality: Sun-Moon Lake, Nan-tou Hsien, Taiwan.

Type specimen: USNM Helm. Coll. holotype male no. 63228,

CT / os)

E4 ae AN

ize: Sain, (Se

NEMATODE PARASITES OF OCEANICA 9

716 | 47

Figs. 1 and 2. H. altaica; 3 and 7. H, dispar; 4. H. variabilis; 5 and 6. H, brevispiculum;

8 and 14. A. vexans; 9 and 13. AH. vulvolabiata; 10. H. alata; 11 and 12. H. indica;

15 and 16. H. beramporia; 17 and 18. H. isolonche.

10 REC. 8S, AUST. MUS., 16 (8); I-14 Octoher, WTI

Remarks; (n spicule length, Heterakis vexans sp.n. is most similar to

Heterakis parva Maplestone, 1931, but the spicules of that species are more

slender and lack the terminal flanges characteristic of the H. brevispiculum-

group found on H. vexans, The shapes of the spicules are closest to Heterakis

altaica Spaul, 1929, from Asian galliform birds. Syntypes of this latter

species were studied by one of us (W.G.I.) and the spicules are not as robust

as those of H. vexans and they also lack the terminal hooks. Neither H. parva

nor H. altaica are known to have three lateral paracloacal papillae. These

papillae are variable in other species; it is common for one pair to be lost

through fusion with an adjacent pair, On the other hand, AH. psophiae

Travassos, 1913, consistently has only three pairs. Lt will remain for sub-

sequent discoveries of H. vexans to prove if this is a consistent feature of the

species,

Heterakis variabilis Chandler, 1926

(Fig. 4)

The following redescription is based on twenty males and nine females

from three species of gallinaceous hosts, All specimens are in excellent con-

dition,

DESCRIPTION

Morphology typical for genus. Lips large, with characteristic papillae,

Additional pair of inconspicious papillae occasionally present on each side,

posterior to lips. Lateral alae prominent. Anterior end usually curved

dorsad.

Male: 6.0 to 9.4 mm. long, 290 to 335 greatest width (1.0 mm.

posterior to oesophagus). Oesophagus (excluding pharynx) 0.850 to

1.15 mm. long, with posterior bulb 150 to 180 greatest width. Pharynx

(measured from anterior tooth to junction with oesophagus) 50 to 65 long.

Excretory pore 320 to 400 from anterior end. Nerve ring 230 to 325 from

anterior end. Precloacal sucker (Fig. 4) 70 to 85 long, 60 ta 90 wide

(measured across outer margins of sclerotized wall), posterior edge 105 to

150 from anus. Tail 410 to 670 long, Right spicule 1.20 to 1.97 mm. long,

lacking alae. Left spicule 410 to 900 long, with well-developed alae.

Caudal papillae typical of genus, but variable in number and location.

Occasionally, a supernumerary single or pair of small, sessile papillae is

present anterior to sucker,

Female: (all specimens gravid), 7.0 to 11.0 mm, long, 265 to 360

greatest width (1.0 mm. posterior to oesophagus). Ocesophagus (excluding

pharynx) 0,960 to 1.15 mm. long, with posterior bulb 150 to 180 greatest

NeMATODE PARASITES OF OCEANICA in|

width, Pharynx 65 to 80 long. Excretory pore 350 to 480 from anterior

end, Nerve ring 240 to 325 from anterior end. Tail 0.912 to 1,3 mm. long.

Vulva salient or not. 3.35 to 5.57 mm. from posterior end. Ovijector with

distal loop. Eggs (measured in ovijector) 60 to 76 by 35 to 40. Two

(rarely one or three) tandem, large, postvulvar papillae,

Hosts: Swinhoe’s blue pheasant, Lophura swinhoii (Gould, 1863);

bamboo partridge, Bambusicola theracica sonoricox Gould, 1862; mikado

pheasant, Syrmaticus mikado (Ogilvie-Grant, 1906). (Phasianidae! Galli-

formes. )

Location: Caecum and large intestine.

Localities: Wu-lai, Tai-pei Hsien; Pu-li and Wu-sheh, Nan-tou Hsien:

Ta-fu, Hua-lien Hsien; Hsin-sheh, Tai-chung Hsien: I-lan. I-lan Hsien:

Taiwan,

Specimens deposited; U.S.N.M. Helm. Coll. nos. 63221-63223.

REMARKS

Chandler (1926) described this species from specimens recovered from

three peacock pheasants, Polyplectrum bicalcaratum (L), which had died in

the Caleutta Zoological Gardens. Maplestone (1932) recorded this species

from the type host in India, and Inglis (1958) found it in the type host in the

London Zoological Gardens. Baylis (1936) and Madsen (1950) consider

this species to be a synonym of Heterakis isolonche Linstow, 1906, but that

species has spicules roughly equal in length while they are markedly unequal

in //. variabilis. Inglis (1958) suggested that Heterakis parva Maplestone,

1931, may be a synonym of 1. variahilis, but the present study shows this not

to be the case (see below). Heterakiy variabilis can easily be recognized by

the sizes and shapes of the spicules.

The tiny papillae behind the head have not been reported previously for

this species although we have seen them repeatedly in other species. Their

structure and function remain problematical. Postvulvar papillae are present

on every specimen of A. variabilis that we have studied, including subadults

where they are small and appear to be developing. ‘They always appear in

tandem, are transversely elongate, and have hypodermis intruding into them,

No nervous element could be seen. It seems to us unlikely that they were

formed by the sucker of the male during copulation, because of their size,

shape, and location. Possibly they aid in locating the male genital pore.

Similar structures are known in the hookworm genus 4rthrostoma Cameron,

1926,

(2 REC. S. AUST, MUS,, 16 (8): 1-14 October, 197]

Heterakis yulvolabiata Chandler, 1926

(Figs. 9 and 13)

The following redescription is based on fifteen males and ten females

from six Formosan hill partridges. All specimens are in good to excellent

condition.

DESCRIPTION

A small, slender heterakid, with morphology typical for genus. Lateral

alae narrow. Anterior end usually curved dorsad.

Male: 4.3 to 5.1 mm, long, 168 to 196 maximum width (1.00 mm.

posterior to oesophagus). Oesophagus (excluding pharynx) 530 to 650

long, with posterior bulb 80 to 120 greatest width. Pharynx 40 to 50 long.

Excretory pore 240 to 300 from anterior end. Nerve ring 200 to 235 from

anterior end. Precloacal sucker (Fig. 9) 31 to 40 long, 32 to 40 wide;

posterior edge 30 to 50 from anus. Tail 200 to 230 long. Right spicule

460 to 570 long, very slender, needle-like, lacking alae, with simple tip. Left

spicule (Fig. 13) 290 to 335 long, stout, alate, with blunt, slightly bifid tip.

Caudal papillae typical of genus, but variable in number and location.

Supernumerary papillae anterior to sucker not observed.

Female: (All specimens gravid). 5.0 to 7.0 mm. long, 190 to 265

maximum width (1.0 mm. posterior to oesophagus). Oesophagus (exclud-

ing pharynx) 575 to 865 long, with posterior bulb 100 to 140 greatest width.

Pharynx 40 to 64 long. Excretory pore 260 to 388 from anterior end,

Nerve ring 205 to 330 from anterior end. Tail 335 to 695 long. Vulva

salient or not, 2.4 to 3.4 mm. from posterior end. Post vulvar papillae absent.

Ovijector with tight, distal loop. Eggs (measured in uterus) 66 to 70 by 36

to 40.

Host: Formosan hill partridge, Arborephila crudigularis. (Phasianidae:

Galliformes).

Location: Caecum and large intestine.

Localities: Pu-li, Sun-Moon-Lake, Nan-tou Hsien; Shih-men, Ping-tung

Hsien, Taiwan.

Specimens deposited: U.S.N.M. Helm. Coll. no. 63220.

REMARKS

Chandler (1926) described this species from specimens recovered from

Arborophila torqueola (Valenc.) which had died in the Calcutta Zoological

Gardens, Our specimens are somewhat smaller than those of Chandler,

although the spicule sizes are similar. Chandler apparently reversed the left

NEMATODE PARASITES OF OCEANICA 13

and right spicules, for he stated the left was longer. Maplestone (1932)

simply referred to a longer and shorter spicule, but he did figure the

characteristically-tipped short spicule for the first time. Baylis (1936)

described the right spicule as longer.

The present record is the first outside India, and the host record is also

new. The species is readily recognized by its small size and the very

characteristic delicate right spicule.

Heterakis beramporia Lane, 1914

(Figs. 15 and 16)

Several specimens were found in a domestic fowl, Gallus gallus (L.)

from Tai-pei, Tai-pei Hsien, Taiwan. This is a common parasite of fowls

in Asia, but appears not to have been reported previously from Taiwan.

These specimens were overlooked in a previous report (Schmidt and Kuntz,

1970).

Specimens deposited: U.S.N.M. Helm. Coll. no, 63225.

Heterakis spumosa Schneider, 1866

Our data record this species from a banded crake, Rallina eurizonoides

formosana Seebohm, 1894 (Rallidae) from Chiao-chi, I-lan Hsien, Taiwan.

Since this is a cosmopolitan parasite of domestic rats and other rodents, the

record seems dubious and should be viewed with suspicion, The occurrence

of this parasite on Taiwan should be noted, however.

Specimens deposited: U.S.N.M. Helm, Coll. no, 63226.

Heterakis indica Maplestone, 1932

(Figs. 11 and 12)

Two males, one incomplete. were found in a drongo cuckoo, Surniculus

lugubris minimus Baker (Cuculidae), at Terabanan Concepcion, Palawan,

Republic of the Philippines. This is a new host record, although it has been

reported from domestic fowls from Palawan by Schmidt and Kuntz (1970).

The species was adequately described by Maplestone (1932) and by Li

(1933) (as AH. lingnamensis). Inglis (1958) hesitated to recognize this

species, since specimens were not available for study. The present study

fully supports the status of the species.

Specimens deposited: U.S.N.M. Helm. Coll. no. 63227.

14 REC, 8S. AUST, MUS., 16 (8); 1-14 October, 197)

Heterakis isolonche Linstow, 1906

(Figs. 17 and 18)

(Syn. H. putaustralis Maplestone, 1922; H. tragopanis Lal, 1942.)

Several specimens were found in the caecum and large intestine of a

Swinhoe’s blue pheasant, Lophura swinhoii, from Chun-yeh, Ping-tung Hsien,

Taiwan. These are new host and locality records. This well-known parasite

has been recorded from a wide range of galliform birds in Asia, Europe and

North America and was adequately redescribed by Li (1933).

Specimens deposited: U.S.N.M. Helm. Coll. no. 63224.

ACKNOWLEDGEMENTS

The authors wish lo acknowledge the field support by Dr. D. 8, Rabor, Department of

Biology, Silliman University, Dumaguete City, Negros Onental, Republic of the Philippines,

and the technicians of the Parasitology Department of Naval Medical Research Unit No, 2

for general assistance in obtaining und examining hosts. Dr. B. J, Myers aided in pre-

liminary processing of specimens,

Initial work for this study was supported by funding under Public Law 480, Section

104 (c¢), by funds provided by the United States Bureau of Medicine and Surgery, Navy

Department Work Unit MR 005,20-0098, and by Contract No. NR 103-690/N0014-66-C0094,

between the United States ollice of Naval Research, Department of the Navy, and the

Southwest Foundation for Research and Education.

Final efforts were sponsored by United States Department of the Army, Contract No,

DADA 17-68-C-8094.

Work in South Australia was completed while one of us (G.D.S.) was a Visiting NATO

Fellow and was supported by funds to W.G.1, from the Mark Mitchell Foundation and the

Australian Grants Committee.

REFERENCES

Baylis, H, A. 1936: The fauna ef British India. Nematoda 1. (Asearidoidea and

Strongyloidea), Taylor and Francis, London,

Chabuud, A. G., 1965: Superfamille des HMeterakoidea, in Traitée de Zovlogie 4 (3): L010:

1016.

Chandler. A. C.. 1926: New Heterakidae from Indian galliform birds, Jnedian J. med. Res

13: 617-623.

Freitas, J, F.. Teixeria de., 1956: Notas sdbre “Heterakidae”, Railliet and Henry. 1914.

(Nematoda, Subuluroidea). Rev. bras. Biol, 16: 461-482.

liglis, W. G., 1958: A review of the nemalode superfamily Heterakoidea. Ann. Mag. nat.

Hist. (12) 10; 905-912. (NB, This paper is misdated 1957 as printed but was

published 4th June, 195%.)

Inglis, W, G., 1967: The evolution, host relationships and classification of the nematode

superfamily Heterakoidea, Bull, Brit, Mus, nat, Hist, (Zeolegy) VW (1): 1-28,

Li, H. C., 1933: Parasitic nematodes mitinly from North China. Part TV. Oxyuroidei.

Chinese Med, J, 47, 1307-1325.

Lépez-Néyra, C. R.. 1947: Helniintos de los vertebradoy Théricos IH, Granada.

Madsen, 1., 1950: Studies on species of Hererakiy (Nematoda) in birds. Danish Rev, Game

Bial, 1: 1-42,

Muplestone, P. A., 1932: The genera Meterukiv and Pseudaspidodera in Indian hosts. Rec.

Indian Muy, 20; 403-420.

Sehmilt, G, D. and R. E. Kuntz, 1970: Nemiutode parasites of Oceanica Vil. New records

from wild and domestic chickens (Gullus gallus) from Palawan (Philippine

Islands), Subah (Malaysia), and Taiwan, Avian Diseases 142 184-187.

SOUTH AUSTRALIAN BAK

MUSEUM

SUPERFICIAL MANDIBULAR MUSCU-

LATURE, VOCAL SACS AND THE

PHYLOGENY OF AUSTRALO-PAPUAN

LEPTODACTYLID FROGS

By MICHAEL J. TYLER

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 9

12th May, 1972

SUPERFICIAL MANDIBULAR MUSCULATURE, VOCAL SACS AND THE

PHYLOGENY OF AUSTRALO-PAPUAN LEPTODACTYLID FROGS

BY MICHAEL J. TYLER

Summary

Two major groups of genera can be distinguished among Australo-Papuan leptodactylid frogs on

the basis of the presence or absence of attachments of the Musculus intermandibularis upon the M.

submentalis. These groups correspond to the Cycloraninae and Myobatrachinae respectively except

that Cyclorana cannot be associated with either. Cyclorana lacks the attachments of the

intermandibularis upon the submentalis, characteristic of the Cycloraninae, and differs from the

Myobatrachinae and Cycloraninae in have the intermandibularis differentiated into separate

elements at the apex of the mandibles.

SUPERFICIAL MANDIBULAR MUSCULATURE, VOCAL

SACS AND THE PHYLOGENY OF AUSTRALO-PAPUAN

LEPTODACTYLID FROGS

By MICHAEL J, TYLER

SYNOPSIS

Two major groups of genera can be distinguished among Australo-

Papuan leptodactylid frogs on the basis of the presence or absence of attuch-

ments of the Musculus intermandibularis upon the M. submentalis. These

groups correspond to the Cycloraninae and Myobatrachinae respectively

except that Cyclorana cannot be associated with either. Cyclorana lacks the

attachments of the intermandibularis upon the submentalis, characteristic of

the Cycloraninaec, and differs from the Myobatrachinae and Cycloraninue in

having the intermandibularis differentiated into separate elements at the

apex of the mandibles,

In the majority of genera the vocal sac is a unilobular submandibular

structure but in Nofaden it is bilobular and supramandibular. Heleieporus

and Neohatrachus lack vocal sacs, but it is concluded that the buccal cavity is

an effective resonance chamber because in these genera the floor of the mouth

can be depressed to further increase the capacity of a Jarge buccal cavity. It

is argued that the ancestral stock from which the Australo-Papuan genera are

derived lacked a vocal sac,

Morphological divergence in hyomandibular structures currently used to

distinguish leptodactylid sub-families is reported, and the different evolu-

tionary mechanisms by which such divergence may have arisen are discussed.

INTRODUCTION

Parker (1940), in his monograph of the leptodactylid frogs of the

Australian Geographical Region, recognises two sub-families, the Myobatra-

chinae and the Cycloraninae, largely from a study of the thigh musculature,

hyoid und tongue, supplemented by data from Trewavas (1933) on the

hyoid and larynx. Although features of the hyoid, larynx, and tongue

supported such a clear-cut division Parker nevertheless acknowledged that

the Myobatrachinae might be, “a group of frogs derived from the

Cycloraninae by the evolution of the same type of feeding mechanism on

more than one occasion”.

Because the superficial mandibular musculature often provides a reliable

index of generic and sub-familtal relationships in the Hylidae (Tyler, 19714),

| made similar studies to establish the extent of divergence and taxonomic and

phylogenetic significance of such structures in the Australian leptodactylid

sub-families.

| 12th Muay, 1972

REC. S, AUST. MUS., !6 (9): 1-20 Muay, 1972

In the course of these studies data were also recorded for a few genera

on some of the characters considered by Parker (1940) to be, “possibly of

considerable taxonomic value” (e.g., site of attachment of the Mm,

sternohyoideus and petrohyoidei upon the hyoid plate, and the shape of the

hyoid alary processes). These data permit a reassessment of Parker's

suggestion.

MATERIALS, METHODS AND TERMINOLOGY

Representatives of the following genera and species were examined:

Adelotus brevis.

Crinia georgiana, C. glauerti, C. haswelli, C, laevis, C. leai, C. parinsignifera,

C, riparia, C. signifera, C. victoriana.

Cyclorana alboguttatus, C. australis, C. brevipes, C. cultripes, C. dahli, C.

platycephalus.

Glauertia orientalis, G. russelli.

Heleioporus albopunctatus, H. barycragus, H. eyrei, H. inornatus, H,

psammophilus.

Kyarranus sphagnicolus,

Lechriodus papuanus, L. platyceps.

Limnodynastes convexiusculus, L. dorsalis, L. fletcheri, L. ornatus, L.

peroni, L. spenceri, L. tasmaniensis,

Metacrinia nichollsi.

Mixophyes fasciolatus.

Myobatrachus gouldi.

Neobatrachus centralis, N. pelobatoides, N. pictus, N. sutor, N. wilsmorei.

Notaden bennetti, N. melanoscaphus, N. nichollsi.

Philoria frosti.

Pseudophryne bibroni, P. corroboree, P. dendyi, P. semimarmorata.

Taudactylus acutirostris, T. diurnus,

Uperoleia rugosa.

The methods employed, muscle terminology, descriptive synonymy and

a description of the generalised anuran superficial mandibular musculature

and yocal sac structure are presented in detail elsewhere (Tyler, 1971a).

That terminology is therefore only summarised here so that this paper can be

considered alone.

The anuran superficial mandibular muscles consist of (1) the M.

submentalis which is an ovoid and customarily araphic muscle at the apex of

the mandibles, so described in the following text as “normal”. (2) The M.

Micuarn J. Tyrer—On AusTRALO-PAPUAN LBPTODACTYLID FROGS 3

intermandibulavis which arises as a thin flat sheet from the lateral lingual

surface of the mandibles between the submentalis and the jaw articulation,

and (3) the M, interhyoideus which arises principally from the anterior

cornu of the hyoid and unites anteriorly with the posterior border of the

intermandibularis.

In some genera the intermandibularis is differentiated by the presence of

supplementary elements which lie ventral to the customary sheet of muscle.

The nomenclature of the supplementary elements is derived from the

position on the mandible from which they arise, ¢.g., “apical” or “lateral”.

The term “vocal sac” is here restricted to the epithelium lined chamber

developed as an extension of the floor of the mouth, and customarily lying

dorsal to the superficial mandibular muscles and ventral to the hyoid plate

(see Fig. 1). The vocal sac and the superficial mandibular muscles comprise

the “yocal sac structure”.

COMPARATIVE MORPHOLOGY OF SUPERFICIAL MANDIBULAR

MUSCULATURE AND VOCAL SACS

Adelotus (Fig. 2A)

The M. submentalis is normal but partially obscured by the interman-

dibularis which attaches upon ils ventral surface. The interhyoideus is poorly

developed and anteriorly underlies the intermandibularis. The interhyoideus

arises from the anterior cornu and there is a slender ligamentous attachment

to the posterior face of the squamosal.

The vocal sac apertures extend along the lingual margin of the mandibles

for almost the entire length of the intermandibularis. The vocal sac lies

almost completely aboye the intermandibularis.

Crinia

The submentalis and intermandibularis are normal except that the

latter is reduced postero-medially in association with an increased develop-

ment of the interhyoideus. During the embryology of C. georgiana and C.

haswelli the anterior development of the intermandibularis is arrested so that

this muscle remains separate from the posterior border of the submentalis.

The interhyoideus arises partly from the anterior cornua and partly from the

squamosal; it is well developed posteriorly with a large median lobe extending

far beyond the post-articular extremities of the mandibles.

The vocal sac lies above the intermandibularis and imterhyoideus (Fig.

1). The apertures are relatively long, extending approximately one half of

the total length of the mandibles.

=“)

4 REC. S. AUST. MUS., 16 (9): 1-20 May, 1972

Fig. 1. Mandibular region of male Crinia signifera in sagital section.

Key to abbreviations—A = heart (auricle); genhy. — Musculus geniohyoideus;

genioglo. — Musculus genioglossus; hyoglo. — Musculus hyoglossus; hyoid pl. =

hyoid plate; imand. — Musculus intermandibularis; interhy. — Musculus inter-

hyoideus; L. = larynx; Li. — liver; mand. — mandible at symphysis; O. —

oesophagus: P.L.S. — pectoral lymphatic sac; rect. — Musculus rectus abdominus;

S. = stomach; S.L.S. = submandibular lymphatic sac; st. —= sternum; submen. —

Musculus submentalis; T. — tongue; V. — heart (ventricle); V. S. = Vocal sac.

Cyclorana

The submentalis is small in all members of the genus and normally

araphic. (A distinct median raphe occurred in a single individual of C.

cultripes.) The intermandibularis is differentiated by the development of an

apical element arising on each side of the submentalis, an element similar to

that characterising the Australian hylids (Tyler, 1971la, Fig. 4A). In C.

alboguttatus and C. dahli, a small group of the most anterior fibres of this

apical element are directed forwards and attach medially upon the ventral

surface of the submentalis, but in the remaining species there is neither over-

lap nor attachment.

The interhyoideus is extremely well developed posteriorly, particularly in

C. platycephalus where there is a conspicuously large posterior lobe. There

is, however, considerable variation in the site of origin of that muscle. In

C. australis, C. brevipes and C. dahli this muscle arises entirely from the

anterior cornua with an extensive region of attachment extending to a point

beneath the eustachian tubes. In C. alboguttatus and C. cultripes although

the majority of the fibres are attached to the anterior cornua, approximately

MicHacn J. TyteR—On AUSTRALO-PAPUAN LEPTODACTYLID FROGS §

ten per cent of them attach on the posterior arm of the squamosal. The

extreme condition is exhibited by C. platycephalus where such fibres arise in

equal proportions from the squamosal and anterior cornua.

The slit-like vocal sac apertures are short, extending for less than one-

quarter of the length of the mandibles, with their anterior margins almost on

a level with the posterior limit of the apical element of the intermandibularis,

and the posterior margins near the fusion of the interhyoideus with the

customary element of the intermandibularis.

In male C. platycephalus the corium of the skin in contact with the super-

ficial mandibular muscles is frequently intensely pigmented black, and the

dull greyish external appearance of the submandibular skin would appear to

be entirely caused by this deep pigmentation.

Examination of a series ot tadpoles and transforming juvenile specimens

of C. australiy (stages 41-46), revealed that the sequence of ontogeny of the

mandibular muscles is similar in all respects, to that reported from Australo-

Papuan hylids by Tyler (1971a).

Glauertia

The submentalis is normal but the intermandibularis does not extend

anteriorly to mect it, so that small portions of the geniohyoideus are exposed.

The interhyoideus is well developed extending posieriorly beyond the post-

articular portions of the mandibles; this muscle arises in a broad sheet from

the squamosal and anterior cornua,

The vocal sac apertures are long and broad, bordered by the anterior

cornua on one side and the geniohyoideus on the other, and extending for a

distance equivalent to one half of the total length of the mandibles. The

vocal sac is extensive and occupies almost the entire area above the inter-

mandibularis and interhyoideus,

Heleiaporus (Fig. 2D)

The submentalis is extremely large but completely covered ventrally by

the intermandibularis which is firmly attached to it medially. The inter-

hyoideus arises from the anterior cornua with an intimate proximal attach-

ment to the inferior surface of the eustachian tubes.

Members of this genus lack vocal sacs. The lining of the mouth on each

side of the tongue is markedly pleated and the geniohyoideus extremely loose,

REC. 8S. AUST, MUS,, 16 (9): 1-20 May, 1972

submen.

Fig. 2. Superficial mandibular musculature.

A = Adelotus brevis; B = Myobatrachus gouldi; C = Nataden nichollsi; D =

Neobatrachus centralis. Key to abbreviations—imand, — Musculus_ interman-

dibularis; interhy. — Musculus interhyoideus; submen. — Musculus submentalis.

Micttace J. TyLeR—ON AUuSTRALO-PAPUAN LeEPTODACTYLID PRoGS 7

Kyarranus

The submentalis is of moderate size and partially obscured by the

attachment of the anterior fibres of the intermandibularis. Direct attachment

of the intermandibularis fibres to the mandible is confined to the posterior half

of the latter. Anteriorly the fibres run initially parallel to and then diverge

from the mandible, and the attachment to the mandible is via transparent

connective tissue. The interhyoideus is well developed.

The vocal sac apertures lie parallel to the posterior half of the pre-

articular portion of the mandible. The vocal sac lies above the entire

surface of intermandibularis and interhyoideus.

Lechriodus

The submentalis is of moderate size and its posterior border is partially

obscured by the attachment of fibres of the intermandibularis upon it, The

intermandibularis bears a median aponeurosis. The interhyoideus is not well

developed.

Vocul sac and vocal sac apertures are present in L, fletcheri and L.

papuanus, but not in L. platyceps. The vocal sac is extensive, occupying

most of the area aboye the interhyoideus and intermandibularis,

Limnodynastes

The submentalis is broad and partially obscured by the inter-

mandibularis. The extent to which the intermandibularis intrudes upon the

ventral surface of the submentalis varies from species to species, being most

extensive in L. dorsalis where only a V-shaped portion of the submentalis is

visible behind the apex of the mandibles. The other extreme is exhibited by

L. oernatus and L. spenceri, in which the anterior fibres of the inter-

mundibularis are transversely directed so that the visible segment of the

submentalis is larger and almost semicircular.

In most species the interhyoideus arises solely from the anterior cornua

but in a few individuals of L. dorsalis and L. tasmaniensis slender attachments

to the squamosal occur. Postero-medial development of the interhyoideus is

most pronounced in L. ornatus and L. spenceri, and overlies a portion of the

pectoral musculature.

Vocal sac apertures extend for almost the entire length of the inter-

mandibularis. The vocal sac lies above the intermandibularis and inter-

hyoideus.

For illustrations of the superficial mandibular musculature and inflated

vocal sac structure of L. tasmaniensis see Tyler (1971b).

8 REC. S$. AUST. MUS., 16 (9): 1-20 May, 1972

Metacrinia

The submentalis is normal and completely visible. The inter-

mandibularis is not differentiated into separate elements but individual super-

ficial fibres, particularly those arising at the anterior ends of the mandibles,

are directed obliquely and posteriorly and thus overly those directed trans-

versely. The interhyoideus arises solely from the anterior cornua. In the

mid-line there is a bell-shaped aponeurosis at the junction of the inter-

mandibularis and interhyoideus.

On their dorsal surfaces the intermandibularis and interhyoideus are

directly attached to the broad geniohyoideus. The vocal sac apertures are

short and located close to the mandible directly anterior to the posterior

limit of the intermandibularis. The vocal sac is confined to the area above

the interhyoideus.

Mixophyes

The submentalis is large, its posterior border hidden by transversely

directed fibres of the intermandibularis. The intermandibularis bears a

median aponeurosis near the junction with the narrow interhyoideus. The

interhyoideus arises partly from the anterior cornua and partly from the

squamosal.

The vocal sac apertures are long, equivalent to more than one third of

the length of the mandible. The vocal sac is large, laterally extending along

the entire length of the intermandibularis and interhyoideus, medially to the

apex of the aponeurosis.

Myobatrachus (Fig. 2B)

The submentalis is small and normal. The most anterior fibres of the

intermandibularis are directed anteriorly but do not appear to underly the

submentalis. The interhyoideus is extremely thin and extends posteriorly

into a large median lobe. The segment of the interhyoideus lying behind

the post-articular portion of the lower jaws is not entirely muscular but consists

of muscle fibres broadly separated by connective tissue.

The vocal sac apertures are approximately one-quarter of the total length

of the mandibles. The posterior border of the aperture is on a level with the

posterior limit of the intermandibularis. The vocal sac lies above almost the

entire intermandibularis and interhyoideus and there are slight lateral exten-

sions following the anterior cornua behind the jaws.

Neobatrachus

The submentalis is large and, in those species in which the tip of the

mandibles is particularly blunt (e.g., N. wilsmorei), it is almost semi-circular

in shape.

Anteriorly the intermandibularis overlies the submentalis and attaches

to it in the midline. The interhyoideus arises entirely from the anterior

Mrow\nn J. Pyier—On AUSTRALO-PAPUAN LEPTODACTYLID FROGS 4)

cornua, On their contiguous border the only evidence of interpenetration of

fibres of the interhyoideus and intermandibularis occurs medially.

Vocal sacs are absent in all members of the genus, and the inter-

mandibularis and interhyoideus closely adhere to the muscles lying dorsal to

them. The lining of the mouth between the mandible and lateral border of

the tongue 1s extensively folded (Fig. 3) as in Heleiopoerus,

Nataden (Fig, 2C)

The submentalis is elongated, compressed and partly obscured by the

attachment of the intermandibularis upon its ventral surface, In N. bennetti

and N. niehollsi the interhyoideus arises solely from the anterior cornua but

in N. melanoscarhus the muscle arises in equal portions from the anterior

comua and the tympanic annulus. The interhyoideus is unusually well

developed proximally, the fibres being directed posteriorly from the anterior

corua creating a lateral extension to the vocal suc. In adult males the

interhyoideus is markedly convoluted and the giant bundles of fibres are not

always contiguous being separated, particularly in the median portion, by

connective tissue,

The vocal sac appertures are bounded by the anterior cornua, commenc-

ing at the point where these pass above the geniohyoideus, und extending for

u distance equivalent to one third of the total length of the mandibles. The

development of bilobular supramandibular pouches to the vocal sac in

members of this genus is unique amongst Australo-Papuan leptodactylids, and

may be associated with the extremely obtuse mandibles und the associated

reduction of the site of attachment for the superficial mandibular muscles.

Philoria

The submentalis is large and composed of two separate transverse

segments of which the smaller anterior js attached solely to the prominent

mento-meckelian bones, whilst the larger posterior segment arises from the

dentaries,

The intermandibularis is an extensive muscle whose anterior fibres

diverge from the transverse path, passing forwards to attach upon and almost

completely obscure the posterior segment of the submentalis. On the

anterior one-third of the jaw there is thus no direct attachment of muscle fibres

to the mandibles, and contact 1s maintained by connective tissue. The inter-

hyoideus arises solely from a particularly extensive region of the anterior

cornua and terminates distally at a point midway between the mandible and

the geniohyoideus lateralis. The interhyoideus has a prominent posterior

lobe.

The vocal sac apertures are rather oblique, extending anteriorly from the

point where the anterior cornua pass above the geniohyoidcus lateralis, and

posteriorly to the posterior Limit of the intermandibularis on the muandible.

10 REC. S. AUST. MUS., 16 (9): 1-20 May, 1972

Fig. 3. Mandibular region of male Neobatrachus pictus in transverse

section.

Sectioned posterior to tongue; skin removed. A = entire mandibular

region; B. = enlargement of floor of mouth lateral to mandible (position

indicated by arrow in figure 3A).

MICHAEL J. TYLER—ON AUSTRALO-PAPUAN LePTODACTYLID FRoGS 11

The vocal sac is large, occupying the posterior half of the intermandibularis

and the entire interhyoideus.

Pseudophryne

The submentalis is normal. The intermandibularis is not differentiated

but anteriorly the superficial ventral fibres are directed obliquely and

posteriorly, so that they overly the deeper and transversely directed fibres.

The interhyoideus arises entirely from the anterior cornua and is well

developed posteriorly.

The vocal sac apertures are long, extending for a distance equivalent to

approximately one-half of the total length of the mandible and are obliquely

situated. The vocal sac is rather posterior in position and largely confined to

the vicinity of the interhyoideus.

Taudactylus

The submentalis is composed of two segments in T. acutirostris, a small

anterior portion arising solely from the mento-meckelian bones, which partly

overlies the larger segment arising from the dentaries. No such separation

occurs in 7. diurnus.

There is an elongate, median, longitudinal aponeurosis to the anterior

half of the intermandibularis adhering completely to the deep musculature.

The interhyoideus arises partly from the anterior cornua and partly from the

posterior arm of the squamosal.

In T. acutirostris the vocal sac apertures are rather oblique and posterior

in position, being bounded by the anterior cornua and extending medially to

the point where the cornua pass above the geniohyoideus. The vocal sac is

largely confined to the interhyoideus. Neither vocal sac apertures nor vocal

sac were detected in the 7. diurnus examined.

Uperoleia

The superficial mandibular musculature is similar to that of Glauertia.

The only difference of note is that in Uperoleia the yocal sac apertures are

slit-like and bounded solely by the anterior cornua.

|2 REC. 5S. AUST. MUS., 16 (9): 1-20 Muay, 1972

HYOMANDIBULAR FEATURES AND LEPTODACTYLID

PHYLOGENY

Although Parker (1940) considered a wide variety of morphological

characters, the only ones that supported the recognition of two major groups

of genera were features of the larynx, tongue and hyoid, Laryngeal

characters however were not examined by Parker who relied on the observa-

tions of Trewavas (1933), As the latter examined only single representatives

of one myobatrachine and three cycloranine genera, the value of laryngeal

anatomy as a source of taxonomic criteria is unknown in this group.

The tongue of the Myobatrachinae was found by Parker to be a slender

organ in contrast to the broad cycloranine tongue. The characteristic features

of the hyoid noted by Trewavas (1933) were summarised by Parker as

follows:

“Cycloruninae (Heleioporus, Limnodynastes and Mixephyes).

(1) Alary process narrow proximally, but expanded distally

(pedunculate).

(2) M. omohyoideus present.

(3) Mm. sternohyoideus and petrohyoidei attached at the lateral

edges of the hyoid plate.

Myobatrachinae (Crinia).

(1) Alary process of the hyoid a wing-like expansion of almost the

whole lateral margin of the hyoid plate, without narrow stalk.

(2) M, omohyoideus absent.

(3) Mm. sternohyoideus and petrohyoideus anterior inserted on the

ventral surface of the hyoid, reaching the middle line in the

posterior part of this insertion.”

In this summary Parker employed a certain amount of licence because

the Mixophyes differed from the above cycloranine definition in the following

respects: the alary process is simply a small, triangular protuberance in the

figure (Trewavas, 1933, Fig. 28), and the sternohyoideus is attached to the

body of the hyoid plate; at no point does that muscle reach the lateral borders

of that plate. Parker’s justification for ignoring such discrepancies was that

Trewavas examined a juvenile specimen,

Parker noted that loss of the omohyoideus is believed to have occurred

many times in the Anura. Accordingly he considered it to be of little

taxonomic significance, and did not report its presence or absence in genera

other than those examined by Trewavas. He did examine the shape of the

alary processes and the sites of attachment of the petrohyoideus and sterno-

hyoideus on the hyoid, and reported that their conditions in the different

MicHaAet J. TyLER—ON AUSTRALO-PAPUAN LEPTODACTYLID FROGS 13

genera corresponded to the above subfamilial definitions, Current delimita-

tion of these subfamilies therefore really depends upon these two characters

in conjunction with that of the tongue.

In the study reported here I examined these characters in a few genera,

and in Notaden and Philoria my observations conflict with those of Parker,

Subsequently, I analysed the structures in terms of their function, and con-

sidered the various ways in which such morphological divergence could have

arisen, before attempting to re-evaluate the phylogenetic significance of the

characters,

ant. cornu

alary proc.

hyoglo. membr.

hyoid plate

post.-lat. proc.

post.-med. proc.

Fig. 4. Hyoid plate and variation in alary processes.

A. generalised hyoid plate, B-F alary processes, for explanations see text on page 14,

Key to abbreviations—alary proc, — alary process; ant. cornu = anterior cornu;

hyoglo. membr. = hyoglossal membrane; post.-lat. proc. — postero-lateral process;

post.-med. proc. — postero-medial process.

14 REC. S. AUST. MUS., 16 (9): 1-20 May, 1972

The hyoid plate is a thin sheet lying in a horizontal plane between and

posterior to the mandibles, [t is customarily entirely curtilaginous and

extremely pliable in hfe. Occasionally however it may be partly or entirely

calcified, or partly ossified. Up to four pairs of distinct processes arise from

the borders of the plate! anterior cornua from the anterior surface, alary and

postero-medial processes from the lateral margin, and postero-lateral from the

postero-lateral corner (Fig. 4A). Loss of one or more pairs of the first three

of these has occurred in many anuran genera.

In the literature the term “alary process” is applied to two quite

different hyoid structures: (1) a broad “wing-like” extension of the entire

lateral edge of the plate such as occurs in myobatrachine genera (Fig. 4B), or

(2) a more restricted, discrete process extending from part of the lateral

border on a level with the superior margin of the plate, This latter type can

be a simple extension rounded terminally, which may be as broad proximally

us distally (Fig. 4C), or may be pedunculate with a circular (Fig. 4D), oval

(Fig. 4E) or even irregular (Fig. 4F) terminal margin,

The Mim. petrohyoideus and sternohyoideus attach upon the hyoid plate

at one of two positions; either along the lateral border of the plate or upon

the body of the plate.

In both subfamilies the lateral, lingual margin of the muscles follows the

axis of the proximal portion of the anterior cornua, If the wing-like alary

processes in the Myobatrachinae are regarded as “optional extras”, the lateral

lingual borders of the sites of muscle attachment are seen to be similar to

thuse in the Cycloraninae,

There are no muscles attached to wing-like alary processes whilst, i

contrast, the petrohyoideus attaches to the proximal portion of the peduncu-

late type which thus provides an increase in the site available for attachment.

In contrast | think that the wing-like processes only support the floor of the

mouth, | only arrived ut this conclusion by the elimination of all other

possibilities, T have, however been unable to demonstrate a consistent associa-

tion between the presence of this process and broad mandibles: a feature

which might conceivably influence the evolution of such a structure, while

remembering Gans (1966) stricture that the present architecture of a struc-

ture is not necessarily moulded by its present function,

In cycloranine genera such as Heleioporus in which the distal portion of

the pedunculate alary process is extremely large, a structurally supporting

role similar to that proposed for the myobatrachine alary process is indicated,

Where it is small its ability to perform a similar function is considered

unlikely. From a consideration of the available data the possibility that

wing-like and pedunculate alary processes are not homologous structures

cannot be dismissed. However, to regard the presence or absence of wing-

Micrare J. TyLeER—On AuSTRALO-PAPUAN LEPTODACTYLID FROGS 15

like alary processes a character of taxonomic value appears justified. Thus

although the fact that I have not seen distal dilations to the discrete processes

in Notaden and Philoria necessitates a change in the diagnosis of the sub-

family Cycloraninae to accommodate the variation in the alary processes, it

does not materially affect the taxonomic value of the character.

The differences in the positions occupied by hyoid muscles, as described

here, may have been the result of two independent evolutionary mechanisms.

One, a major alteration of a skeletal component providing attachment, as

exhibited by the gross structural change of the mandibles of Notaden, and the

other ontogenetic heterochrony.

mand.

submen.

genhy. I.

genhy. m.

Fig. 5. Anterior mandibular musculature showing association between Musculus

submentalis and Musculus geniohyoideus,

A. generalised anuran in ventral aspect; B, Nefaden nichollsi. Musculus intermandi-

bularis removed, Key to abbreviations: genhy. 1. = Musculus geniohyoideus

lateralis: genhy. m. = Musculus geniohyoideus medialis; mand. — mandible;

submen. — musculus submentalis, The ligamentous portion of the geniohyoideus

Jateralis of N. nichollyi is represented by broken lines.

16 REC. S. AUST, MUS, 16 (9); 1-20 May, 1972

The M. geniohyoideus lateralis of all anurans previously reported in the

literature is attached upon the lateral lingual surface of the mandible immedi-

ately posterior to or dorsal to the submentalis (Fig, 5A). Bul for a very

slight, initial mediad inclination the fibres of this muscle follow the longi-

tudinal axis. In Neftaden the mandibles are exceptionally blunt anteriorly,

and the submentalis is extremely elongate, If in this genus the yeniohyoideus

lateralis remained in its customary position in relation to the lateral end of

the submentalis, the fibres of the muscle would follow a path at an acute angle

to the longitudinal axis,

lt is evident that Netaden has retained the ancestral condition: the

muscle remains attached to the mandible near the mandibular symphysis, bul

by means of a membrane, and the submentalis now passes partly beneath this

membrane, und partly beneath the geniohyoideus lateralis (Fig. SB). The

major influence here has evidently been that of retention of direct opposition

of sites of muscles origin and attachment, suggestive of un inevitability in

these sites to maintain functional efficiency.

Ontogenetic heterochrony must be regarded as a potential source of

inodification to the positions occupied by adjacent muscles. During ontogeny

the retardation of the development of one or the acceleration of another

oecupying an adjacent site, could clearly permit fibres of one to intrude upon

and occupy a portion of the site normally occupied by the other. In genera

such as Glavertia such a process could be responsible for the more inferior

margin to and exposure of the mandibular site of the geniohyoideus, associated

with the arrested development of the intermandibularis,

Parker (1940) noted that in the Myobatrachinae the sternohyoideus

teaches the midline in the posterior portion of the msertion on the hyoid

plate, contrasting with the cycloranine condition where it is a more slender

muscle and does not, My observations on Nofaden do not agree with those

of Parker. In all specimens examined the sternohyoideus is a massive muscle

which occupies almost the entire ventral surface of the hyoid plate ( reaching

the midline throughout its length). In the face of such marked variation

between the material we have examined, the medial limit of the site of the

sternohyoideus upon the hyoid plate does not constitute a character by which

the two subfamilies can be distinguished,

SUPERFICIAL MANDIBULAR MUSCULATURE AND VOCAL SACS

AS INDICES OF LEPTODACTYLID PHYLOGENY

The two subfamilies of Australo-Papuan leptodactylid frogs currently

recognised can now be distinguished on the basis of one consistent difference

in the superficial mandibular musculature. In the Cycloraninae the most

anterior fibres of the intermandibularis attach apon the ventral surface of the

submentalis, while such attachments are absent in the Myobatrachinue (Table

Micuaen J. Tyter—Ow AustrRaco-PaPuan Lerropactyiip FRocs

1). Unfortunately Cyclorana differs from other cycloranine genera in lacking

the attachments otherwise characteristic of the subfamily. The genus differs

further from both cycloranine and myobatrachine genera by having the

intermandibularis differentiated into separate elements.

In all the cycloranine genera except Cyclorana the nature of the attach-

ments is similar, involving transversely oriented fibres of the intermandibularis.

The differences existing between genera are the extent to which these fibres

underly the submentalis, but it is difficult to express quantitatively the pro-

portion of the submentalis so hidden from view. This is unfortunate because

the data presented here indicate that the extent of such attachment is

taxonomically significant, because there is a close similarity in this character

in genera currently considered to have a close phylogenetic relationship on the

basis of other evidence. For example, Neobatrachus is recognized as a valid

TABLE |

itermandibularis| | Supplementary |

Subfamily Genus Submentalis elements of Vocal Sac

| connection Intermandibularis |

_ _ . ; _ —

Myobatrachinae.. | Crinia ...,.,,. | Absent ......... Absent ........0. Unilobular and

submandibular

Myobatrachinae. Glauertia ..... Absent ......... Absent , 22.2.0. Unilobular and

submandibular

Myobatrachinae.. | Meracrinia.... | Absent ..,...... Absent ......,,. Unilobular and

submandibular

Myobatrachinae.. | Myohatrachis.. | Absent ......... Absent ......... Unilobular and

submandibular

Myobatrachinae.. | Pseudophryne.. | Absent ......... Absent ......... Unilobular and

submandibular

Myobatrachinae.. | Taudactyluy ... | Absent ......-.. Absent .......0. Unilobular and

submandibular

or absent

Myobatrachinac.. | Uperuleia ..... Absent ......... Absent 2, ¢.¢6004 Unilobular and

submandibular

Cycloraninae ..., , Adelotus ....., Present ........, Absent ......... | Unilobular and

submandibular

Cycloraninae .,.. | Cyclorana .... | Absent ......... Present ......... Unilobular and

submandibular

Cycloraninae .... | Heleioporus ... | Present .......-. Absent ......... Absent

Cycloraninae ._.. | Kyarraius } Present ......... Absent ......... Unilobular and

submandibular

Cycloraninae ... Lechriodus .... | Present ......... Absent ......... Unilobular and

submandibular

or absent

Cycloraninae .... | Limnodyrastes . | Present .,....... Absent ......... Unilobular and

submandibular

Cycloraninae .... Mixophyes .... Present ....... . Absent ......... Unilobular and

submandibular

Cycloraninae .... | Neohatrachus Present 22.0.5 5.04. Absent ....,.... | Absent

Cycloraninae .... | Nofaden ....,. Present ......... Absent ......... Bilobular and

supramandi-

bular

Cyeloraninae .... | Philoria ....., Present ......... Absent .....0.45. Unilobular and

submandibular

18 REC, 8. AUST. MUS. 16 (9): 1-20 May, 1972

genus largely on biological data, and Parker (1940) did not distinguish it

morphologically from Heleioperus, Correspondingly, the ventral surface

of the submentalis is completely covered by the intermandibularis in both

genera, a feature nol shared by any of the others.

Vocal sacs are present in all genera except Heleioporus and

Neobarrachus, one species of Lechriodus (L. platyceps) and one species of

Tandactylus (T. diurnus). Absence of voeal sac may be considered a primaty

feature when it is characteristic of a family (e.g., Ascaphidae and Discoglos-

sive) or secondary in genera where the vast majority of species possess vocal

sacs, but one or two do not (Hyla and Litoria, see Tyler, 1971a), In the

former case there ts no evidence that vocal sacs haye ever evolved, whereas in

the latter the interpretation is one of loss,

The vocal sac structures of the other Australo-Papuan leptodactylid

genera are unilobular in form and submandibular in position, except in

Notaden, where they are bilobular und supramandibular.

Heleioporus and Neobatrachus in which there are no yocal sacs, have large

domed buccal cavities which can evidently be further enlarged by depression

of the floor of the mouth, as indicated by the presence of longitudinal pleating

of the floor. The capacity of the buccal cavity is therefore adequate to act

us an effective resonance chamber and so perfornis the role of the vocal sac,

Both groups of evidence suggest that resonance chambers have evolved

independently at least twice amongst Australo-Papuan leptodactylids, and

thal the ancestral stock from which this family is derived lacked them.

In stmmary the myological data demonstrates that the superficial

mandibular musculature of Australo-Papuan leptodactylids indicate the exist-

ence of two major groups of genera, which correspond to the two subfamilies

recognised by Parker (1940), but with neither of which Cyclorana is readily

associated. Within these groups the muscle architecture of the Myoba-

trachinae represents the primitive condition and that of the Cycloraninac. a

derived state in which the path of some of the fibres has been modified.

Thus this evidence does not support Parker's (1940) tentative suggestion

that the Myobatrachinae could have been derived from the Cycloraninae.

It has been demonstrated elsewhere (Tyler, 1971a), (i) that superficial

mandibular muscles are conservative in the evolutionary sense; (1i) that

differentiation of the intermandibularis is a derived state constituting a major

evolutionary development, and (iii) that some hylid sub-families are

characterised by possession of similar supplementary elements of the inter-

mandibuluris.

Therefore the presence of supplementary elements of the inter-

mandibularis in Cyclorana and their absence from other cycloranine genera

are grounds for questioning the sub-familial disposition of Cyclorana. In

contrast the form of the supplementary elements in Cyelorana is identical to

MICHAEL J. TYLER—ON AUSTRALO-PAPUAN LEPTODACTYLID FROGS 19

that possessed by all Australo-Papuan hylids with which it shares other

morphological and behavioural characteristics (Tyler, 1970). Further

comparative studies of leptodactylids and hylids are currently being under-

taken to clarify the familial disposition of Cyclorana, and redefinition of the

Cycloraninae and the Myobatrachinae is deferred pending completion of

these studies.

Note added in proof.

Since the above was written a reassessment of the evolutionary relation-

ships of the Australo-Papuan genera has appeared in a revision of the

Leptodactylidae by Lynch (Misc, Publ. Univ. Kansas Mus. Nat. Hist. (53):

1-238 (1971)). Lynch supports recognition of the sub-families proposed by

Parker (1940) and maintained here, but has proceeded a stage further in

recognizing two tribes within the Cycloraninae. He associates Cyclorana with

Heleioporus, Mixophyes, Neobatrachus and. Notaden as the members of one,

and considers this tribe the most primitive. Therefore our conclusions differ

principally in that I dispute the sub-familial disposition of Cyclorana, but

Lynch does not.

20 REC. S. AUST. MUS., 16 (9): 1-20 May, 1972

ACKNOWLEDGMENTS

For the provision of species not represented in the South Australian Museum collections

I wish to express my thanks to Dr, M. J. Littlejohn, Dr. A. A. Martin and Mr. J. Loftus-Hills

(University of Melbourne), Dr. R, Endean (University of Queensland), Mr. J. M. de Bavay

(University of New England) and Dr. J, L. Hickman (University of Tasmania).

I am also indebted to the Director of the South Australian Museum, Dr. W. G. Inglis,

for his encouragement throughout the project and criticism of the manuscript, to Miss T.

Mannik (University of Adelaide) who prepared the histological sections, and Dr. J. Fanning

for the photomicrograph used in Fig. 3.

REFERENCES

Gans, C., 1966: Some limitations and approaches to problems in functional anatomy.

Folia Biotheoretica 6: 41-50.

Parker, H. W., 1940: The Australasian frogs of the family Leptodactylidae. Novit, Zool. 42

(1): 1-106.

Trewavas, E., 1933: The hyoid and larynx of the Anura. Phil, Trans. Roy. Soc. Lond.,

Ser. B 222 (10): 401-507.

Tyler, M. J., 1970: Patterns of distribution and the origins of the Papuan hylid frog fauna.

Search 1 (4): 246-247.

Tyler, M. J., 1971a: The phylogenetic significance of vocal sac structure in hylid frogs.

Univ. Kans, Publs. Mus. nat, Hist. 19 (4): 319-360.

Tyler, M. J,, 1971b: Voluntary control of the shape of the inflated vocal sac by the

Australian leptodactylid frog Limnodynastes tasmaniensis. Trans. Roy. Soc.

S, Aust. 95 (1): 49-52.

A. B, JAMES, GOVERNMENT PRINTER, ADELAIDE

RECORDS OF THE

SOUTH AUSTRALIAN / o>.

MUSEUM By,

PLANIGALE GILES! (MARSUPIALIA,

DASYURIDAE); A NEW SPECIES

FROM THE INTERIOR OF SOUTH

EASTERN AUSTRALIA

By PETER F. AITKEN

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 10

12th June, 1972

PLANIGALE GILESI (MARSUPIALIA, DASYURIDAE); A NEW

SPECIES FROM THE INTERIOR OF SOUTH EASTERN

AUSTRALIA

By PETER F. AITKEN

South Australian Museum

SUMMARY

Planigale gilesi a new species of dasyurid marsupial is described and

figured. Field notes on the specimens examined are included.

PLANIGALE GILESI*

Diagnosis: A robust Planigale (plate 1) differing from all other

species of Planigale by the possession of only two premolar teeth in each

upper and lower jaw.

Holotype: South Australian Museum no. M8406; adult male puppet

skin and skull with torso in spirit, collected on 29 June 1969 by Messrs.

P. Aitken, A. Kowanko, J. Forrest and J. Howard.

Type Locality: No. 3 Bore, Pastoral Property of Anna Creek, South

Australia (lat. 28° 18/’S., long. 136° 29’ 40”E.).

Paratypes: No. 3 Bore, Anna Creek, South Australia. Male puppet

skins and skulls: South Australian Museum nos. M8407, 25 July, 1969,

P. Aitken, A. Robinson and M. Stanley; M8408 and M8409, 26 November

1969, P. Aitken, J. Forrest and J. Glover. Male in spirit: SAM no. M8410,

27 July 1969, P. Aitken, A. Robinson and M. Stanley. Female in spirit

with skull extracted: SAM no. M8411, 25 August 1970, A. Kowanko and

J. Glover.

Bellata, New South Wales (lat. 29° 55’S., long. 149° 47’E.). Female

in spirit with skull extracted; Australian Museum no. M7033, 27 February

1945, J. Kirkby. Male in spirit with skull extracted: AM no. M7393, May

1948.

Brewarrina, New South Wales (lat. 29° 57’S., long. 146° 51’E.).

Male in spirit with skull extracted: AM no. M7819, 1954, K. Turnbull.

Female in spirit with skull extracted: AM no. M7820, 1954, K. Turnbull.

Lake Cawndilla, Kinchega National Park, New South Wales (lat.

32° 30’S., long. 142° 18’E.). Male in spirit: AM no. M9190, 20 May 1969,

M. Gray.

Descriptive Methods: Pelage colour nomenclature follows the standards

of Ridgway (1912). All body weights are in grams and all anatomical

measurements are in millimetres with the terminology, unless otherwise stated,

after Cockrum (1955). Skull measurements were taken with Helios dial

*JIn honour of Ernest Giles (1835-97) the most intrepid of Australian explorers and,

like this planigale, an accomplished survivor in deserts.

12th June, 1972

2 REC. S. AUST. MUS., 16 (10): 1-14 June, 1972

calipers under a binocular microscope at various magnifications up to x20.

Flesh dimensions were obtained from freshly killed material in the case of

South Australian Museum specimens and from spirit preserved material in

the case of Australian Museum specimens. All measurements were made by

the author.

External Features: Dorsally the soft, dense fur is 8 mm long on the

rump where the basal 5 mm are sooty black, the median 2.5 mm are pinkish

cinnamon and the apical 0.5 mm are fuscous. The fur is 4 mm long on

the crown and 1 mm long on the muzzle where it becomes more bristly with

virtually no sooty black base. Interspersed with the fur are medially-

thickened fuscous black spines 10.5 mm long on the rump reducing to

2 mm on the muzzle. The back is thus a brindled cinnamon colour

from rump to rhinarium, although two of the skins examined (SAM nos.

M8408 and M8409) have much paler spines and, in consequence, less

brindled appearances. A single ring of fuscous black hairs is present around

each eye and there is a dark patch in front of each eye where the tips of

the fur are fuscous black. Mysticial vibrissae number approximately 16 on

each side, are up to 12 mm long and are predominantly fuscous black. Other

vibrissae per side are: supra-orbital, 2, fuscous black; genal, 8, some fuscous

black, others white; ulnar carpal, 3, white; anconeal, 1, white; median

antebrachial, 1, white; submental, 4, white. In addition there are 4 white,

interramal vibrissae.

On the cheeks, flanks and shoulders, behind the ears and inside the

ears, the fuscous tips of the fur are markedly reduced and spines are virtually

absent. These areas are thus pinkish cinnamon in colour.

Ears rounded and slightly fleshy, each with a weak ventral lobe defined

by a shallow notch in the posterior margin, an antero-dorsal overfold and a

posteriorly concave, anteriorly pointed, untwisted supratragus 3.5 mm long

by 2.5 mm broad. Externally, the basal segment of each ear is naked with

a surrounding apical area thinly covered by short pinkish cinnamon hairs.

Internally, the concha and supratragus of each ear carry irregularly spaced,

fine white bristles and each pinna has two thick tufts of pinkish cinnamon

hairs originating on the postero-internal margin, one immediately above the

ventral lobe the other at crown level. The remainder of each pinna is more

thinly covered with short, pinkish cinnamon hairs.

Rhinarium naked, fuscous black with a complete median groove, no

philtrum and semicircular nostrils directed antero-laterally.

Ventrally the fur is soft with no spines and is 5 mm long on the belly

where the basal half is dark mouse gray and the apical half is pinkish buff.

On the interramal region the fur is 1.5 mm long and pinkish buff throughout.

Two of the skins examined (SAM nos. M8408 and M8409) have paler

ventral fur, basally deep mouse gray and apically pale olive buff. The fur

on their interramal regions is also pale olive buff.

PLANIGALE GILESI—A New AustTRALIAN DaAsyuRID 3

Scrotum pendulous with fuscous black pigmented skin and a thick

covering of pinkish buff hairs (pale olive buff on SAM nos. M8408 and

M8409).

Pouch equipped with 12 teats and filled with fine, white hairs up to

6 mm long. It varies in development amongst the three females examined.

In one (SAM no. M8411) it is an oval depression approximately 4 mm

long by 3 mm wide with scarcely perceptible antero-lateral lips. In another

(AM no. M7033) it is an anteriorly deepened pocket with a transverse

postero-ventral opening 3.5 mm wide. In the third (AM no. M7820) it

is an irregular, mammary area up to 18 mm in diameter with a fleshy,

peripheral lip 2.5 mm deep.

Tail slightly incrassated, tapering towards the tip and sometimes

fattened for two-thirds of its length, in which cases basal incrassation is

more obvious (SAM nos. M8407, M8408 and M8410). It is bicoloured

with a thin covering of stiff hairs 3.5 mm long, increasing to 4.5 mm on

the distal third of the underside, but with no brush. Dorsally the hairs

are pinkish cinnamon with fuscous tips, although fuscous tipping may be

much reduced or absent (SAM nos. M8407 and M8408) and some totally

fuscous hairs occur at the distal extremity. Ventrally the hairs are the same

colour as the apical half of the belly fur, either pinkish buff or pale olive

buff throughout. The hairs of the tail tend to grow in ragged whorls based

between 4 and 5 mm apart and in those specimens with fattened tails the

whorl hairs become very divergent, producing a rather sparse covering at

their apices through which the fuscous black pigmented tail skin can clearly

be seen. In such specimens, therefore, the tails exhibit faint patterns of

alternating light and dark transverse bands (SAM nos. M8407 and M8408).

Both fore and hind feet well covered above by short pinkish cinnamon

hairs with a fringe of silvery hairs bordering each naked sole. Soles of

fore feet fuscous, soles of hind feet fuscous black (colours fade in spirit).

Fore feet each 4 mm broad with 5 strongly clawed digits (formula:

3>4>2>5>1) and 6 well developed pads. Soles and pad-bases coarsely

granular with granules up to 0.3 mm across. Each pad-base is surmounted by

a finely striated apical pad: 1st interdigital round, 0.6 mm in diameter; 2nd

and 3rd interdigitals oval, 0.5 x 0.9 mm; 4th interdigital oval, 0.6 x 0.8 mm;

hypothenar proximo-internally flattened and shaped like a bulbous hook

with an external shaft, greatest breadth 0.8 mm by 1.6 mm long; thenar

proximally flattened, demi-oblong, 0.8 mm broad. Hind feet (plate 2)

each 3.5 mm broad across the bases of digits 2-5 with 4 clawed digits

and a clawless hallux (formula: 3>4>2>5>1) plus 6 well developed

pads. Soles and pad-bases coarsely granular with granules up to 0.4 mm

across, although one larger granule, 0.5 mm in diameter, is usually present

near the external margin of each sole between the 3rd interdigital and

hypothenar pads and a few others of similar size occur on each heel, Hach

4 REC. S. AUST. MUS., 16 (10): 1-14 June, 1972

pad-base is surmounted by a finely striated apical pad. st (hallucal),

3rd and 4th interdigitals oval, 0.7 x 1 mm; 2nd interdigital oval, 0.7 x 1.1 mm;

hypothenar oval, 1.4 x 0.8 mm; thenar disto-internally bulbous crescentic,

greatest breadth 0.9 mm by 1.5 mm long.

Dimensions of the soles and pads were obtained from a specimen that

had been preserved in 70% spirit for approximately 6 months (SAM no.

M8410). Some other specimens, examined after much longer immersion

in spirit, had apparently narrower hind feet and shrunken pad-bases that had

become confluent with the soles.

Flesh dimensions of selected specimens are presented in Table 1.

Spirit preserved specimens with extracted skulls were not measured.

Table 1. Flesh Dimensions of Planigale gilesi

3 3 3 ox 2

| Holotype

Measurement M8409 M8410 M9190 M8411

(SAM) (SAM) (AM)

Anna Anna Kinchega

Creek Creek

Body length .. ‘ 73

Tail length. ... : =

Length of hind FE ; 10.5

foot (without

claws)

Height of ear : i 10.4

(from notch)

Weight ....... | 9

(starved)

* Measured from spirit

Skull Characters (plates 3a, b, c, d): Dorsal aspect of cranium flat

with extremely thin bones, a minute sagittal crest and insignificant lambdoidal

crests. Zygomatic arches evenly convex. Interorbital region flat and un-

ridged with bevelled edges, a slight median depression at the posterior

extremities of the nasals and the anterior half of the median frontal suture

irregularly dentate. Post-orbital processes barely discernible or absent with

no pronounced post-orbital constriction. Anterior halves of nasals either

parallel sided or marginally expanded in front, posterior halves flared, the

greatest width across the nasals being at their points of contact with the

fronto-maxillary sutures. Postero-dorsal tips of premaxillae truncated.

A pair of slim, anteriorly pointed, incisive foramina pierce the palate with

their posterior extremities between the canines. Two narrow, sometimes

dissimilar, posterior palatal vacuities are also present with their anterior

margins between M?~ and their posterior margins between M**. Pterygoid

hamulae slender and deeply hooked behind. Alisphenoid bullae bulbous,

the greatest breadth of each bulla being approximately equal to the minimum

PLANIGALE GILESI—A Ne&W AUSTRALIAN DASYURID 3

distance between both bullae. Each periotic bulla with a markedly inflated

distal crescent separated by a shallow sulcus from a less inflated proximal

segment and abutted on its postero-mesial wall by the anteriorly flanged,

blunt para-occipital process.

4 1 2 4

Dental formula: I 3: C 1: P 2: M 4. Teeth, upper jaw: I

prominent, set apart from I°+, which are smaller with I? equal in size to I*

and both larger than I* by crown height but not crown length. I?“ each

with a buccal and a lingual cingulum and a minute talon that is most

marked on I*. Canine bucco-lingually flattened with a complete cingulum

and three times as high as I', from which it is separated by a diastema

of 0.5 mm. In three specimens examined (SAM no. M8501, AM nos.

M7033 and M7819) a distinct talon is also present on each canine. Pre-

molars bucco-lingually flattened, each with a buccal and a lingual cingulum,

a minute anterior cusp and a relatively longer talon. The anterior cusp

is most obvious on the first premolar and the talon is largest on the second

premolar. First premolar set apart from the canine and two thirds as

large as the second premolar by both crown height and crown length, second

premolar offset longitudinally in line with the posteriorly broadened rostrum.

Molars moderately high cusped with M* lacking the entire posterior half.

M?* each with a dorsally displaced protocone. M'“* each with a postero-

lingual cingulum, a dominant metacone, a relatively much lower paracone

and a parastyle, mesostyle and metastyle, of which the parastyle in M! is

antero-dorsally displaced and the mesostyle in M® is reduced in height

compared with the same cusp in either M' or M*. M?~* each with an

anterior cingulum.

Teeth, lower jaw: Incisors slightly procumbent, each with a bucco-

lingually flattened incisal edge and a lingual cingulum. I; not set apart

from and twice as high as Iz, which in turn is a little larger than Is. by

both crown height and crown length. A minute talonid is present on I;.

Canine twice as high and three times as long as Is with a lingual and a

buccal cingulum plus a talonid. Pre-molars bucco-lingually flattened, the

first two-thirds as large as the second by both crown height and crown

length, each with a talonid and a buccal and a lingual cingulum. Moolars

high cusped with M; lacking the postero-buccal quarter. Mu-4 each with

an antero-buccal cingulum, a posterior cingulum and a dominant protoconid,

plus a metaconid and a paraconid, both of which are relatively much lower,

the latter being particularly low in Mi. Each also with a minute entoconid

plus a hypoconid and a hypoconulid, of which the hypoconid is almost totally

suppressed in Mg, is highest in Mi and decreases evenly in height through

Mb-3, whereas the hypoconulid is highest in Ms and equal and lower in

height through M:-s.

Skull and tooth dimensions are presented in Table 2.

June, 1972

1-14

REC. S. AUST. MUS., 16 (10)

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PLANIGALE GILESH—A New AUSTRALIAN DASYURID 7

Field Notes: No. 3 Bore, Anna Creek, is situated at the north-eastern

tip of an isolated belt of sandridges, which transect the stony desert tableland

south of the Neales River on the western side of Lake Eyre North, approxi-

mately 70 km north-east of Anna Creek homestead. The bore was originally

sunk in 1917, but was redrilled in 1966 when an unsuccessful attempt was

made to control its output with casing. No mound has formed around the

bore-head through which 3,337,000 litres of water erupt daily via a circular

hole 7 m in diameter. On leaving the bore-head the water flows down a mean-

dering bore-drain for about 300 m before spreading out to form a shallow

swamp. Beyond the swamp the water continues to trickle across the table-

land for up to 600 m through an expanding system of branched, attenuating

channels, in which the water depths are variable. In consequence, the

extremities of the channels are often dry, but on such occasions the water

table has never been found more than 15 cm below their beds, Water

temperature at the bore-head is 48°C cooling to below 40°C at the entrance

to the swamp. Water salinity is 3,614 parts per million in the bore-drain,

which is flanked by a thin, white mineral crust deposited on its banks.

Desert gobies (Chlamydogobius eremius) and hardyheads (Craterocephalus

eyresii) abound in both the bore-drain and the swamp.

The tableland adjacent to the bore, where the average rainfall is less

than 125 mm per annum, is vegetated by a well spaced, shrub steppe com-

munity dominated by nitre-bush (Nitraria schoberii) with salt-bushes

(Atriplex velutinella and A. angulata) plus another chenopod (Babbagia

dipterocarpa).

The area influenced by the bore is vegetated by a separate community

comprising three distinct plant associations.

1. A bulrush association (plate 4), not found more than 1 m

from surface water, growing in mud along the edges of the

bore-drain and both in and around the swamp. This associa-

tion is dominated by tall, dense stands of bulrush (Typha

angustifolia) basally augmented with tussocks of a small

sedge (Cyperus laevigatus).

2. A sand-spurry association growing in periodically inundated

clearings along the channels. This association is dominated

by low mats of sand-spurry (Spergularia marina) admixed

with love grass (Eragrostis dielsii) and scattered examples of

a small sedge (Scirpus maritimus), pop salt-bush (Atriplex

spongiosa) and an “everlasting” (Helipterum floribundum).

3. A sedge association (plate 5) growing around the bore-head,

over the moist ground between the channels and flanking

the bullrushes along the bore-drain. This association is

8 REC. S, AUST. MUS., 16 (10): 1-14 June, 1972

dominated by thick, interwoven clumps of umbrella sedge

(Cyperus gymnocaulus) occasionally entangled with bushes

of a samphire (Arthrocnemum leiostachyum). Odd exam-

ples of a bindyi (Bassia ventricosa) and ruby salt-bush

(Enchlaena tomentosa var. glabra) are also found through-

out the association.

All specimens of Planigale gilesi collected at No. 3 Bore were trapped

in Sherman live animal traps baited with a mixture of rolled oats, honey,

peanut paste and beef dripping. Two captures were made in the bulrush

association along the bore-drain and four others in the sedge association, two

within 2 m of the bore-head and two between the channels less than 3 m

from an inundated sand-spurry clearing. Both the bulrush and sedge

associations offer superb sanctuaries for small mammals since each provides a

formidable barrier against aerial and terrestrial predators, an internal supply

of food and nesting materials and an effective insulation against extremes of

temperature and humidity. In August, 1969, a comparison of ambient

temperatures with those in a natural runway under the sedge association

revealed that over a 24 hour period temperatures under the sedge remained

fairly constant between 9°C. (0630 h) and 15°C. (1400 h), whereas

ambient temperatures fluctuated between 1°C, (0630 h) and 21°C, (1100 h)

(M. Stanley, unpublished). It is probable that the high humidity resulting

from moist conditions under both the bulrush and sedge associations

remains equally constant.

In captivity examples of Planigale gilesi do not display either distinct

nocturnal or distinct diurnal rhythms, but exhibit short bursts of acitvity

spread throughout each 24 hour period and in view of the protection and

insulation afforded by their habitat it is feasible that their activity patterns

might be similar in the wild. Brown desert mice (Pseudomys desertor),

which are abundant throughout the sedge association at No. 3 Bore, appear

to be most active at night, but can also be trapped quite readily during the

day. Thus indicating that in this habitat they too are not strictly orientated

to either nocturnal or diurnal activity.

Other mammals inhabiting No. 3 Bore are stripe-headed sminthopses

(Sminthopsis frogatti), which are fairly common throughout the sedge

association; long-haired rats (Rattus villosissimus), which have a permanent

breeding colony confined to the bulrush association; house mice (Mus

musculus) of which a few occur in the sedge association around the bore-head;

and rabbits (Oryctolagus cunicudus), which fluctuate in numbers and burrow

on the fringes of the sedge association and around the nitre-bushes in the

surrounding tableland. Visiting mammals observed at or near the bore have

been numerous dingoes (Canis familiaris dingo), some red foxes (Vulpes

PLANIGALE GILESI—A NeW AUSTRALIAN DasyurRID 9

vulpes), occasional feral domestic cats (Felis catus) and small mobs of feral

Arabian camels (Camelus dromedarius), feral donkeys (Equus asinus) and

brumbies (Equus caballus). Domestic cattle (Bos taurus) also water at

the bore.

Damage to the bore vegetation by rabbits appears to be insignificant

and damage to the ground by the hooves of the other introduced herbivores

is slight and limited to the edges of the swamp, which is the only place where

the water is cool enough and still of sufficient depth for comfortable drinking.

Other bores and springs in the surrounding region are not so free of hoof

damage, their vegetated banks being stamped regularly into inhospitable

bogs of mud and dung. No small native mammals have been captured at any

of these sites despite intensive trapping. The introduced carnivores probably

prey on Planigale gilesi occasionally, but in all the dingo and red fox scats

examined rabbits remains were the only mammalian residue. No cat scats

were found, Possible avian predators observed over the bore have been

whistling eagles (Haliastur sphenurus), fork-tailed kites (Milvus migrans),

brown hawks (Falco berigora) and barn owls (Tyto alba). Possible reptilian

predators caught in the sedge association have been King brown snakes

(Pseudechis australis) and sharp-snouted snakes (Pseudonaja acutirostris).

Scats of Planigale gilesi removed from traps in which their producers

had been captured contained fragments of insects exoskeletons, indicating

that these planigales are at least partially insectivorous. In captivity they

consumed meal beetle larvae, early instar locusts and small moths with avidity.

Chopped lambs liver was also accepted, but with less enthusiasm. Habitat

data for examples of Planigale gilesi captured other than at No. 3 Bore are

confined to the Kinchega specimen (AM no. M9190), which according to

the collector was also taken from a sedge association adjacent to water.

The breeding period for members of Planigale gilesi has yet to be

determined, Of the three females examined two have undeveloped pouches.

One of the latter (SAM no. M8411) was collected in winter (25 August)

and the other (AM no. M7033) was obtained in late summer (27 February).

The third female (AM no, M7820) has a lactating pouch, but the exact

date of her collection is unknown. Males in the series examined show

insufficient variation in scrotal size for this character to be used as a reliable

guide to breeding activity.

Relationships: Four species of Planigale have been described previously:

P. ingrami (Thomas), 1906, from the north of the Northern Territory, but

known also from eastern and northern Queensland (Troughton, 1928);

P. subtillissima (Lénnberg), 1913, from the Kimberly district of Western

Australia; P. tenuirostris Troughton, 1928, from north-central and north-

western New South Wales and south-western Queensland, but known also

10 REC. S. AUST. MUS., 16 (10): 1-14 June, 1972

from east-central South Australia (SAM no. M8405); and P. novaeguineae

Tate and Archbold, 1941, from Papua. The range of Planigale gilesi over-

laps that of P. fenuirostris in north-western New South Wales. All four

species differ from Planigale gilesi by the possession of 3 premolar teeth in

each upper and lower jaw. Lénnberg (1913) actually described P. sub-

tillissima as lacking P4-s, but Tate (1947) in a review of the genus corrected

this error after re-examining the holotype.

ACKNOWLEDGEMENTS

I am indebted to the Director and staff of the South Australian State Herbarium for plant

identification; to Miss Meredith Stanley of the Department of Zoology, Monash University

for ecological data pertaining to No. 3 Bore; to Messrs. Alex Kowanko and John Glover

of the South Australian Museum, James Forrest of the C.S.I.R.O. Soils Division, Tony

Robinson of Monash University and John Howard of Wirrula, South Australia for assistance

in the field; to Mr. Roman Ruehle of the South Australian Museum for the photographs;

and to Mrs. Joan Murphy of the South Australian Museum for typing the manuscript.

I am particularly indebted to Mr. Michael Archer of the Western Australian Museum for

supplying me with unpublished information from his current research on the taxonomic

characters defining the genus Planigale; and to Mr. Basil Marlow of the Australian Museum

for the loan of specimens in his care.

REFERENCES

Cockrum, E. L., 1955: “Manual of Mammalogy”. (Burgess: Minneapolis.)

Lénnberg, E., 1913: Results of Dr. E. Mjobergs Swedish Scientific Expeditions to Australia

1910-13. K. svenska Vetensk-Akad. Handl. 52 (1): 9-10.

Ridgway, R., 1912: “Colour standards and Colour Nomenclature”. (Author’s publication:

Washington. )

Tate, G. H. H., 1947: On the Anatomy and Classification of the Dasyuridae (Marsupialia).

Bull. Am. Mus. nat, Hist. 88 (3): 133-135.

Troughton, E. LeG., 1928: A New Genus, Species and Subspecies of Marsupial Mice

(Family Dasyuridae). Rec. Aust. Mus. 16 (6) 281-288, pl XXXIX.

Plate 1.

Planigale gilesié; M8410 (SAM), Anna Creek, South Australia. (Photo by

Roman Ruehle.)

Plate 2. Hind foot of Planigale

gilest, M8410 (SAM). (Photo

by Roman Ruehle.)

12 REC, S. AUST. MUS., 16 (10): 1-14 June, 1972

Plate 3. (a) Skull of Planigale gilesi; dorsal view of cranium and upper

jaws (holotype). (Photo by Roman Ruehle.)

Plate 3. (b) Skull of Planigale gilesi; ventral view of cranium and upper

jaws (holotype). (Photo by Roman Ruehle.)

Plate 3.

PLANIGALE GILESI—A NEW AUSTRALIAN DasyuRID

jaws (holotype). (Photo by Roman Ruehle.)

(d) Mandible of Planigale gilesi; lateral view (holotype).

(Photo by Roman Ruehle.)

14 REC. S. AUST. MUS., 16 (10): 1-14 June, 1972

Plate 4. Bore-drain and bullrush association; No. 3 Bore, Anna Creek, South Australia.

(Photo by the author.)

Plate 5. Sedge association; No. 3 Bore, Anna Creek, South Australia, (Photo by the

author.)

RECORDS oF THE ZQ

SOUTH AUSTRALIAN &.

MUSEUM los

AN ANNOTATED CHECKLIST OF THE

NATIVE LAND MAMMALS OF THE

NORTHERN TERRITORY

By SHANE A. PARKER

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER II

8 January 1973

AN ANNOTATED CHECKLIST OF THE NATIVE LAND MAMMALS OF

THE NORTHERN TERRITORY

BY SHANE A. PARKER

Summary

Until now, the Northern Territory has lacked a checklist of its native land-mammals. To fill this gap

I offer the present compilation. It is modelled on Dr. G. M. Storr’s List of Northern Territory Birds,

and based mainly on records derived from the literature and from unreported material in museums,

especially that in the Northern Territory Museum, Alice Springs. All records used have been

carefully considered, and a number of critical and seemingly anomalous specimens re-examined.

Those responsible for the identification of material are gratefully acknowledged below.

AN ANNOTATED CHECKLIST OF THE NATIVE LAND

MAMMALS OF THE NORTHERN TERRITORY

By SHANE A. PARKER

Hon. Research Worker, South Australian Museum, Adelaide 5000

(formerly Arid Zone Research Institute, Alice Springs)

INTRODUCTION

Until now, the Northern Territory has lacked a checklist of its native

land-mammals. To fill this gap I offer the present compilation. It is

modelled on Dr. G. M. Storr’s List of Northern Territory Birds, and based

mainly on records derived from the literature and from unreported material

in museums, especially that in the Northern Territory Museum, Alice Springs.

All records used have been carefully considered, and a number of critical

and seemingly anomalous specimens re-examined. Those responsible for the

identification of material are gratefully acknowledged below.

The text is straightforward and needs few words of introduction. Under

Range all known localities are listed unless the species is common and wide-

spread. when range limits only are detailed. The distribution maps, however,

show us accurately as possible all records used; records from Bathurst and

Melville islands and a few from Groote Eylandt. lacking further localization,

have merely been plotted within the boundaries given. Under Status appear

what few remarks on abundance and habitat seem justified. Under Taxonomy

will be found notices of recent taxonomic changes and a perfunctory coverage

of trinomial nomenclature.

My purpose throughout has been to provide a summary of the available

information, by which further records may be readily evaluated.

LIST OF SPECIES

With a few exceptions the scientific nomenclature follows Ride (1970).

Where a locality record is unqualified by a bibliographical reference or the

abbreviated name of an institution, it has been taken from one of the two

main references, Finlayson (1961) and Johnson (1964).

Abbreviations ef Institutions—

AM; Australian Museum, Sydney.

AMNH: American Museum of Natural History, New York.

BMNH: British Museum (Natural History), London.

CSIRO: CSIRO Division of Wildlife Research, Canberra.

MCZ: Museum of Comparative Zoology, Harvard University,

Cambridge, Mass.

Iysued & Juniwiity, 1973

2 REC, S. AUST. MUS.. 16 (11): 1-57 January, 1973

MVZ: Museum of Vertebrate Zoology, University of California,

Berkeley.

NMV: National Museum of Victoria, Melbourne.

NTM: Northern Territory Museum (Zoology), Alice Springs (at

present housed at the Arid Zone Research Institute).

QM: Queensland Museum, Brisbane.

SAM: South Ausiralian Museum, Adelaide.

USNM: United States National Museum, Smithsonian Institution,

Washington DC.

WAM: Western Australian Museum, Perth.

MONOTREMES

Tachyglossus aculeatus (Shaw, 1792). Spiny Ant-eater

Range and Status: Rocky areas and isolated outcrops throughout, — In

the Tanami Desert, recorded amongst the giant mounds of Nasutitermes

triodiae (N'TM files, 1965).

Taxonomy: The northern populations were referred to the race

V.a.acanthion (Collett, 1884) by Johnson (1964); see also Griffiths (1968).

MARSUPIALS

Antechinus bellus (Thomas, 1904). Fawn Antechinus

Map |

Range: South Alligator district; Coburg Peninsula (NTM 1961, CSIRO

1965); between Mount Borradaile and Oenpelli (NTM 1968); Cape Arnhem

Peninsula.

Status: Fairly common in some areas, inhabiting woodland.

Because Antechinus flavipes is similar to A. bellus, the specimen from

Port Essington recorded by Thomas (1888 : 292) and a second from the

Daly River recorded by Collett (1897; 334) as “Phascogale flavipes

leucogaster” may be referable to A. bellus, and not to the quite different

A. bilarni as suggested by Johnson (1954 : 78).

Antechinus maculatus Gould, 1851. Pigmy Antechinus

Map 2

Range: King River, north-east of Oenpelli (NMV 1915); Humpty Doo

(AM. WAM); Groote Eylandt. [Bentinck Island, north-western Queensland

(SAM 1962).]

Status: Unknown. Of the five specimens from Groote Eylandt only

one is accompanied by habitat notes: “[Umbakumba] . . . near the crest

PARKER—NORTHERN TERRITORY MAMMALS 3

of a sandy ridge paralleling the shore, where wattle scrub is the dominant

vegetation” (Johnson 1964: 440). P. F. Aitken, in litt., reports that he

secured the Bentinck Island specimen in conditions similar to those recorded

by Johnson.

The specimens collected in pandanus fringes, paperbark fringes and

thorn scrub on the floodplain of the Adelaide River at Humpty Doo, listed

as Planigale ingrami by Davies (1960) are referable to the present species

(M. Archer, in Jitt.).

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Map 1 Map 2

Taxonomy: The race A. m. sinualis (Thomas, 1926) was described

from Groote Eylandt. Archer, pers. comm., plans to transfer this species

to the genus Planigale, in which case the present vernacular name will be

inappropriate.

Antechinus macdonnellensis (Spencer, 1896). Red-eared Antechinus

Map 1

Range: North to The Granites (AM 1952), Helen Springs (AM, reg’d

1928), Alexandria and North Island, Sir Edward Pellew Group (Keith

1968), east in the south to Ringwood (NTM 1959).

Status: Widespread, and in some areas, e.g., the Petermann Ranges,

common, becoming rarer and more scattered in “the northern part of its

4 REC. S. AUST. MUS., 16 (11): 1-57 January, L973

range; mainly amongst rocks, frequenting major ranges and isolated outcrops

alike. Two specimens from the Tanami Desert, however (NTM 1964, 1970)

were trapped at the bases of the giant mounds of Nasutitermes triodiue in

belts of Melaleuca glomerata.

Taxonomy: Ride (1964: 62) placed A. mimulus (Thomas, 1906,

type-locality Alexandria) in the synonymy of A, macdonnellensis.

Antechinus bilarni Johnson, 1954. Harney’s Antechinus

Map 1

Range and Status: Uncommon, amongst rocks in rugged sandstone

country of the western escarpment of Arnhem Land: Oenpelli; Mount

Borradaile (NTM 1968); Deaf Adder Creek (NTM 1969).

Taxonomy: Ride (1970) placed bilarni in the synonymy of 4.

macdonnellensis, but Michael Archer (pers. comm.) on the basis of further

studies, considers that for the present it is best maintained as a separate

species.

Planigale ingrami (Thomas, 1906). Ingram’s Planigale

Map 2

Range: Gulf of Carpentaria drainage and the Barkly Tableland; [Lawn

Hull Creek area, north-western Queensland, vide Troughton 1965 : 29-30

and remarks under A. de Lestang in Appendix I] Red Bank Mine (AMNH

1959): upper Nicholson River (NTM-CSIRO 1967); Alexandria;

Brunette Downs (NTM 1970); Newcastle Waters (AMNH 1960).

Status: Apparently not uncommon; blacksoil plains, dry swamps and

the perennial watercourses of the Gulf drainage. One of the Nicholson

River specimens was trapped amongst rocks at Block Waterhole. Remains

of this species were frequent in bird of prey pellets collected on the cracking

blacksoil plains of Brunette Downs in 1970, See Van Deusen (1969).

Phascogale calura Gould, 1844. Red-tailed Phascogale

Map 5

Range and Status: Alice Springs (Spencer 1896b : 30); Barrow Creek

and Tennant Creek (NMV 1901, two specimens obtained by Spencer &

Gillen). No other records.

Phascogale tapoatafa (Meyer, 1793), Grey-tailed Phascogale

Map 5

Range: Upper South Alligator River; Oenpeli (NTM 1957); near

Patonga at 13° 00’, 132° 22’, one dead on road (NTM); Patonga, others

inside buildings at night, June 1969 (D. Egan, pers. comm.): Yirrkala

(AM 1946),

PARKER—NORTHERN TERRITORY MAMMALS 5

Status: Widespread but not common. The Oenpelli specimen was

caught by a cat. The Yirrkala specimen, collected by W. Harney, bears the

comment “climbs and sleeps in trees”. The Patonga district is one of

lightly-wooded country with laterite ridges.

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( Pn me , inner a,

i | avail ¢- nN VN PS K

pobarne xi AGS Darwin vay "2

o Newcastle Waters o Newcastle Walters

@ Tennant Creek | o Tennant Creek

a a

a | a

bad 4, A alice Springs | A a aaAlce,Springs

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Map 3 Map 4

Taxonomy: The NT populations are generally referred to P. ¢. pirata

Thomas, 1904,

Sminthopsis crassicaudata (Gould, 1844). Fat-tailed Sminthopsis

Map 4

Range: North to Willowra and the Bundey River drainage.

Status: Widespread, and following good seasons locally common.

Prefers more densely vegetated areas and softer soils than the next species

(sandhills, seasonal swamps, bore overflows and creeks).

Taxonomy: The NT populations are generally referred to S. c. centralis

Thomas, 1902.

Sminthopsis froggatti (Ramsay, 1887). Stripe-faced Sminthopsis

Map 3

Range: North to Tanamt (Glauert 1933: 21) and Brunette Downs

(NMV 1968, NTM 1970).

6 REC, S. AUST. MUS,, 16 (ft): 1-57 January. 1973

Status; Widespread and seasonally common; analysis of a large number

of pellets of Letter-winged Kite, Elanus scriptus and Barn Owl, Tyto alba

collected on Brunette Downs in April 1970 showed that S. froggatti had been

an abundant prey species, It appears to prefer more open habitats and

harder ground than does S. crassicaudata. On the Barkly Tableland in April

1970 it was found on treeless blacksoil plains, living in cracks in the carth

(P. K. Latz and D. Howe, pers. comm,); P. Byrne (in Spencer 1896b : 34)

wrote that around Charlotte Waters S$. froggatti seemed to prefer the stony

tablelands whilst §. crassicaudata preferred the softer ground near the creeks

and amongst the sandhills.

Taxonomy: Ride (1970) placed Sminthopsis larapinta Spencer, 1896

in the synonymy of S. froggatti. Troughton (1965 : 36; see also 1964 : 318)

placed Sminthopsis stalkeri, Thomas, 1906 (based on two specimens from

Alexandria and Alroy Downs) in the synonymy of S. crassicawdata. Dr.

Ride, however (in /itt.), regards stalkeri as a synonym of S. froggatti.

Sminthopsis hirtipes Thomas, 1898, Hairy-footed Sminthopsis

Map 5

Range; Station Point, Charlotte Waters; Lake Mackay area.

Stratus; Unknown, No recent localized material from the NT, There

is in the SAM a specimen collected 10 miles north of Warburton Mission,

eastern WA, in May 1968.

Sminthopsis psammophila Spencer, 1895, Large Desert Sminthopsis

Map 3

Range and Status: The holotype (NMYV) was collected by the Horn

Expedition on 18 June 1894 between Kurtitina Well and Ayers Rock, in

sandhill-Triodia country with groves of desert oak Casuarina decaisneana

in the swales (Spencer 1896a: $4). E. Cowle, one of the collectors,

professed to having seen the same species in the James Range, though on

habitat this may be doubted,

Remains of this species haye recently been identified by Michael Archer

in owl pellets (NTM) collected in August 1962 from a cave at Ayers Rock.

In 1969 P. F. Aitken obtained four live specimens from two localities in the

Lock and Kyancutta districts of the Eyre Peninsula, SA, in sandhill country

with Triedia and mallee (Aitken 1971).

Sminthopsis murina ooldea Troughton, 1964. Troughton’s Sminthopsis

Map 4

Michael Archer, in /itr., has referred specimens from the Petermann

Ranges (WAM 1970), Ayers Rock (NTM 1962), Curtin Springs (NTM

PARKER—NORTHERN TERRITORY MAMMALS 7

1970) and Charlotte Waters (NMV) to this taxon, which he is elevating

to specific rank. The possibility that Sminthopsis murina constricta Spencer,

1896 is referable to this taxon should be investigated.

Sminthopsis rufigenis Thomas, 1922. Red-cheeked Sminthopsis

Map 3

Range and Status: Specimens known from Oenpelli (WAM, donated

1960), and Melville Island (SAM 1913) (Michael Archer, in Jitt.).

Troughton (1964 : 319), under S. lumholtzi, listed specimens from the

open blacksoil plains of the Barkly Tableland, NT, and western-central

Queensland. In view of the fact that all Australian specimens of S. rufigenis

so far seen by Archer are from wooded localities in the coastal drainage,

Troughton’s material should be re-examined.

Taxonomy: Ride (1970) relegated Sminthopsis lumholtzi Troughton

& Iredale, 1934 to the synonymy of S. rufigenis, which was described from

the Aru Islands. The type-locality of lumholtzi is Herbert Vale on the

Herbert River of eastern Queensland, not to be confused with Herbert Vale

on the Herbert River north-west of Camooweal, north-western Queensland.

BO moth ROMA

yO a 8

fies aT post carwin "PRG

oo pay

a ° vr ry { A iy ¥

JUG a re

Any at hing aus

a 2 1 ar “ay

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| ~ | on

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| | |

| | | |

Pe Tennant Creek | @ Tennant Creek |

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| | |

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| Peiiice Springs | La . @ Alice Springs |

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| | | |

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Map 5 Map 6

& REC. 8S. AUST. MUS., 16 (11): 1-57 January, 1973

Sminthopsis nitela Collett. 1897, Daly River Sminthopsis.

Map 3

Range and Status; Described from the Daly River. Michael Archer,

in litt., has provisionally referred to this taxon a specimen from the Four-mile

Hole, Wildman River (NTM 1968).

Antechinomys laniger (Gould, 1856). Eastern Antechinomys

Map 6

Range: Lidicker & Marlow (1970) gave the range of this species as

south-central NSW north to the Cunnamulla district of south-central

Queensland, with an isolated record from Cedar Bay, north Queensland.

Material in NMV and SAM not examined by these authors shows that

A. laniger occurs also in the NT: Tarlton Downs (NMV, 18 November

1955); Tobermorey (SAM, coll, C. Watts, 21 September 1968); between

Mangajera and Wajelai billabongs, Roper River (SAM, 4 specimens, coll.

N. Tindale, 1921). In the OM there are specimens from Cheepie and

Charleville, Queensland; a further example in the QM labelled A. laniger,

from Malbon, north-western Queensland, is mounted and not available for

study.

Status: In the NT, unknown. The Roper River specimens were

apparently from a salt-marsh near the river’s mouth (see Gazetteer). These

specimens and the Cedar Bay example, which represents an equally isolated

occurrence, should be critically compared with material from further south

to see whether they really belong to A. laniger or merely resemble that species

in the small size of the auditory bullae.

Antechinomys spenceri Thomas, 1906. Western Antechinomys

Map 6

Range: North to [Canning Stock Route near Sturt Creek, WA] Tennant

Creek, east to Tarlton Downs, [Sandringham, south-western Queensland].

Status: Not uncommon following good seasons, in a variety of country,

including sandhills, mulga and gidgea woodland, mitchell grass plains and

gibbers,

Taxonomy: See Lidicker & Marlow (1970).

Dasyurus geoffroii Gould, 1841. Western Native-Cat

Map 7

Range: |Well 46, Canning Stock Route, WA, vide Glauert 1933 : 18]

PARKER—NORTHERN TERRITORY MAMMALS 9

Barrow Creek (NMV 1901) and Alice Springs (NMV, donated by Spencer

in 1916).

Status: Finlayson (1961) considered this species to have been formerly

widespread in Central Australia, but now a rare and vanishing form. Spencer

(1897; 5) recorded an incomplete skin probably of this species from Crown

Point,

Taxonomy: The NT populations are generally referred to the nominate

race. Ride (1970) relegated Dasyurinus to the synonymy of Dasyurus,

Dasyurus hallucatus Gould, 1842. Northern Native-Cat

Map 7

Range: Mainly coastal and subcoastal, from the Daly River east to

Cape Arnhem Peninsula, inland to the Pine Creek district; Groote Eylandt.

There is an isolated occurrence at Alexandria*.

Status: Inhabits rocky areas, in which it is sometimes abundant, and

eucalypt woodland.

Taxonomy; The race D, h. nesaeus (Thomas, 1926) has been described

from Groote Eylandt; populations from the NT mainland are generally

referred to the nominate race. Ride (1970) relegated Satanellus to the

synonymy of Dasyurus.

Dasycercus cristicauda (Krefft, 1867). Mulgara

Map 2

Range: North to [Hall’s Creek, WA, vide Glauert 1933 : 20] 20° 09’,

130° 15’ in the Tanami Desert (AM 1965) and Tennant Creek, east to

Tarlton Downs (SAM 1956).

Status; Not uncommon following good seasons, mainly in sandhill

Triodia country, though the specimens from the Tanami Desert were trapped

in a belt of Melaleuca glomerata around a claypan,

Dasyuroides byrnei Spencer, 1896. Kowari

Map 7

Range and Status: Inhabits gibber-sandhill associations of the Lake

Eyre basin, occurring marginally in the NT at Charlotte Waters (NMV 1895,

AM 1895). Four specimens from “Ilamurta” (NMV 1895) and one from

“Barrow Creek” (NMYV 1901) have almost certainly been abusively labelled,

for neither of these localities lies near gibber-sandhill country.

Although the species has not been recorded in the NT since 1901, it

has for the last three or four years been tolerably common in areas of south-

western Queensland and north-eastern SA: Sandringham (NTM_ 1970, a

pair, retained in captivity) and Cordillo Downs (Aitken 1970 ; 44),

* Five specimens from this locality in the BMNH, collected by W. Stalker in 1905-06, were

kindly checked as Lo species by Miss P, D, Jenkins, i /irr.

10) REC, S. AUST, MUS,, 16 (11): 1-57 January, 1973

[Myrmecobius fasciatus Waterhouse, 1836. Numbat

As this species occurred formerly in WA west of the Petermann Ranges,

and in the extreme north-western part of SA, it almost certainly occurred

also in the adjoining south-western NT, although no record exists from this

area. |

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e y z

Z sina op

Poi Be Stay

Barntn, pb arwin, ;

i ” ° ‘ . wo ee° Se ge

Oe z id re,

“Cr Ls ae ow if

rie oon oA AC *y

R ia et 3 A T Sate

| x . | ¥ N\ ay

— ~

| hr :

| o Newcastle Waters | | o Newcastle Waters r

: _ |

| Ak | |

| |

| | | v |

| : | 7 |

| | \" |

| | | |

| | l |

| w@ Alice Springs | | 3 Alice Springs |

| | a | a |

| | | |

| ’ | | |

ae — ee ee See Tene Ca neh te cs , a eve

| |

Map 7 Map 8

Notoryctes typhlops (Stirling, 1889). Marsupial Mole

Map 17

Range: North to [Sturt Creek, WA] Napperby (R. Darken, pers.

comm. ), east to Charlotte Waters. The reports from Wauchope and Arltunga

may be doubted in view of the absence of sandhills at these localities.

Status: Inhabits sandhills; widespread but rarely encountered.

Isoodon macrourus (Gould, 1842). Brindled Bandicoot

Map 8

Range: Mainly coastal and subcoastal, from the Daly River east to

Cape Arnhem Peninsula, inland to Mount Shoobridge and Mount McMinn

(NMV 1912); Elcho Island; Groote Eylandt; Sir Edward Pellew Group

(Keith 1968). The specimen in the MCZ from The Granites, listed by

Tate (1948) as of this species, represents such an isolated record that it

should be re-examined for possible confusion with J. auratus.

PARKER—NORTHERN TERRITORY MAMMALS 1

Status: Not uncommon, frequenting thick grass and other dense under-

growth, with or without tree cover.

Isoodon auratus (Ramsay, 1887). Golden Bandicoot

Map &

Range: |Northern Canning Stock Route; Hall’s Creek, WA] Victoria

River (Depot No, 1?; collected by Elsey, vide Thomas 1904 : 228); South

Alligator River; Mount McMinn (NMV 1912): The Granites (MCZ 1932,

SAM 1936, AM 1952); Lake Mackay and Ethel Creek (SAM 1932):

Tennant Creek and Barrow Creek (NMV J901); Alice Springs (Burt Plain),

Henbury and Charlotte Waters (NMV, donated by Spencer in 1916).

Status; According to Finlayson (1961) this species in Central

Australia “survives in considerable numbers” within an area north of Lake

Amadeus north ta about Winnecke Creek and east to the lower Sandover

River. This is no longer so. The last authentic record appears to be of the

specimen collected at The Granites in May 1952 by the Australian Museum,

This species was most frequent in the extensive sandplain-spinifex areas.

Little is known of its status in the northern part of its range, where it

appears to be marginally sympatric with /. macrourus.

Perameles eremiana Spencer, 1897. Desert Bandicoot

Map 10

Range: Burt Plain, and sandhills about 40 miles north-east of Charlotte

Waters (Spencer 1897).

Status: This species, perhaps never common, has declined to vanishing-

point since the 1930's, Finlayson (1961 : 160) stated that it still persisted

in “sectors 5 and 7°, i.e., the Tanami Desert. There appears to be no

authentic record of its occurrence in this area; the vague records from The

Granites mentioned by Finlayson and Troughton (1965 ; 67) are probably

referable to Isoodon auratus.

Macrotis lagotis (Reid, 1837). Bilby

Map 9

Range: North to [Sturt Creek, WA] Lake Buck (NTM files, 1965),

Attack Creek (NMV 1966) and Dalmore Downs (NTM 1969).

Status: Once common, now generally rare, though still not uncommon

in certain areas; mainly on sandy soils. In the Tanami Desert Wildlife

Sanctuary it frequents the Melaleuca glomerata thickets associated with the

Old Marsh Bed, where it burrows both in the level ground and into the bases

of the giant mounds of Nasutitermes triodiae. See Newsome (1962), Smyth

& Philpott (1968) and Watts (1969).

12 REC. S. AUST. MUS., 16 (11): 1-57 January, 1973

Taxonomy: The NT populations are generally referred to M. /. sagitta

Thomas, 1905.

Macrotis leucura (Thomas, 1887). Lesser Bilby

Map 9

Range and Status: Sandhills about 40 miles north-east of Charlotte

Waters (NMYV 1895); “Barrow Creek” (NMV 1901). A skull with lower

jaw was recently found in a nest of the Wedge-tailed Eagle Aquila audax

amongst sandhills 14 miles south-south-east of Steele Gap, in January 1967,

by P. Hanisch (NTM). The nest was not in use and there is no simple way

of dating the skull. This species appears to be restricted to sandhills;

Finlayson (1932, 1935b) who collected 12 specimens near Cooncherie,

north-eastern SA in December 1931, found its burrows only in sandhills,

while in the same area M. lagotis burrowed only in the flat ground. The

three specimens in the NMV labelled “Barrow Creek” probably came from

further afield, as there are no sandhills close to this locality (see Gazetteer).

Taxonomy: Troughton (1965) and Ride (1970) placed Macrotis

minor (Spencer, 1897) in the synonymy of M. leucura.

pe MA, ng oo HA,

a aed aFty ZF" &

ate ee os ety

2 » q e e

t re, ane Sie

Lf ot ke Cy . 4

TY am wv _

ie Ne D ; Dh nein 8 Fo

x | ia | ba

| Ro | h

| co Newcastle Waters | | o Newcastle Waters i

| | | |

| ; | | |

- Tennant Geek oT nt Creek |

7. | |

Les incl !

| ‘ " | |

"| = | |

| . | | ) |

| , os | |

| ee |

eg mia tea te os |e [aces eee ee if ys

Map 9 Map 10

PARKER—NORTHERN TERRITORY MAMMALS 13

Chaeropus ecaudatus (Ogilby, 1838). Pig-footed Bandicoot

Map 10

Range: Ryan Well (SAM 1891); 40 miles north-east of Charlotte

Waters (NMV 1895); Alice Springs (NMV, donated by Spencer in 1916).

Status: No recent records. T can find no basis for the Barrow Creek

record given by Finlayson (1961).

Petaurus breviceps Waterhouse, 1839. Sugar Glider

Map 10

Range: Mary River east to Cape Arnhem Peninsula and inland to the

Katherine River; Bathurst Island (AM); Melville Island; Elcho Island:

Groote Eylandt.

Status: Generally uncommon, though Johnson (1964 : 452) found it

comparatively abundant in the vicinity of Port Langdon. Groote Eylandt, in

1948. Woodland.

Taxonomy: The NT populations are generally referred to P. b. ariel

(Gould, 1842).

Petropseudes dahli (Collett, 1895). Rock-haunting Ringtail

Map 11

Range: Mary River; Oenpelli: Union Town; Nellie Creek; South

Alligator River; Red Bank Mine near Wollogorang (AMNH 1959); Groote

Eylandt (NTM, specimen in captivity obtained by Miss D. Levitt, 1970),

Status: A rock-frequenting species, widespread and locally common.

Dahl recorded it as fairly numerous in Arnhem Land; Tunney collected eight

specimens from Nellie Creek and 16 from the South Alligator; the Spalding-

Peterson Expedition found it “abundant” at Red Bank Mine in 1959 (Hosmer,

in litt.).

Trichosurus vulpecula (Kerr, 1792), Brush-tailed Possum

Map 11

Range and Status: Spencer (1896b : 16-17) mentioned this species as

“occurring everywhere amongst the eucalypts which border the river-beds”

in Central Australia. Davidson (19054: 8) recorded that while he was in

the Murchison and Davenport Ranges (November 1897-February 1898)

“small opossums [were] found by the natives in the gums along the creck

banks”. Terry (1937 : 100) found one in a hollow log west of the McEwin

Hills in 1932,

Finlayson (1961) considered that an inexplicable collapse in its numbers

in Central Australia had occurred since the 1930's. At present it is a rare

but persistent and possibly increasing species in this area, recent records

14 REC. §. AUST. MUS., 16 (11): 1-57 January, 1973

being: Arthur Creek and Plenty River (Finlayson 1961); Glen Helen Station,

1959 (NTM files); Hatches Creek, two captured on trunks of river redgum,

Eucalyptus camaldulensis, and released, 1968 (R. Berry, pers. comm.);

Charley Creek (NTM 1968); Bushy Park (CSIRO 1966, 1969). Other,

earlier, records given by Finlayson are from Wollara and Lake Mackay, the

latter possibly referring to Terry’s 1932 record.

Taxonomy: Spencer (loc. cit.) referred to the Central Australian

populations as “var. fypicus”. Although it seems likely that they do belong

to the nominate race, this has yet to be demonstrated.

Trichosurus arnhemensis Collett, 1897. Northern Brush-tailed Possum

Map 11

Range: South to [Hall’s Creek, WA] Katherine River and Alexandria*;

Bathurst Island; Melville Island (AM); Croker Island: Elcho Island.

Status: Not uncommon, in woodland with hollow roosting-trees

available. In Darwin it is sometimes found in gardens and buildings.

Taxonomy: Once regarded as a well-marked race of T. vulpecula, this

form has now been accorded the rank of species (Ride 1970).

4 / Wear RR, /

SEP EN gt” SNC? va 2

v yon 2? x on ce

wh Darwin G » pie Darwin Ap>

na a a a

| = | .~ ~

~~ | Pc.

| o Newcastle Walers , | o Newcastle Waters .

| | | |

| a |

| |

| 4 | 4 a |

| |

la loa

| a a | | é& |

| a A 4a | | |

| 4 | ‘ |

| ~ | |

| | | ry |

pee ey Lite an SSeS mesa an

Map 11 Map 12

PARKER—NORTHERN TERRITORY MAMMALS 1S

Bettongia lesueur (Quoy & Gaimard, 1824), Burrowing Rat-Kangaroo

Map 12

Range: North to northern Lake Mackay, a point between the western

Macdonnell Ranges and the Stuart Bluff Range, the Elkedra district and the

Sundover drainage, east to between the Tarlton and Toko Ranges; possibly

north to the Tanami Desert (Finlayson 1958b, based mainly on native

reports and records of abandoned warrens).

Status; No authenticated records since the 1930's. If the species

survives at all in the area, which is doubtful, it may do so in the Lake

Amadeus-Lake Mackay regions and in the Elkedra-Sandover River-Plenty

River area,

Once locally common, with colonies living in large warrens; in grassy

and herbaceous loam flats within the major ranges, open mulga and ironwood

parks skirting the ranges, and in sandridge areas, where, as elsewhere, its

warrens were usually made in firm loam at slight elevations on undulating

swales and not in the sandridges proper. Giles (1889: 280) found great

numbers of this species honeycombing low limestone ridges in the dry bed

of Lake Christopher, eastern WA, in late January 1874, and Finlayson

(1958b : 243) noted a similar occurrence between the Stuart Bluff Range

and the western Macdonnell Ranges in 1950, though the animal had long

been extinct there,

the lirge ‘rabbit’ warrens recorded by M. Terry in 1929 in the

Tanami district in approximately 28° South and 129° 51’ East, would

almost certainly be warrens of Bettongia lesucuri doubtfully parasitized by

rabbits” (Finlayson 1958b ; 241). This rendering is peculiarly Finlayson’s,

Terry's notes apply to the year 1928, not 1929, 28°S does not run through

the NT, Furthermore, Terry gave details of to areas in the Tanami district

where he observed large warrens:

(1) [Between Larrangannie Bluff and the “westward mate of

Killi-Killi”, e., at ea. 19° 40’, 128° 58°, WA near the NT border] “ . . . low

cust-west sandhills and rabbit warrens amongst travertine [a kind of lime-

stone] mounds on the plain in between. The presence of either so far north

was news,

“From this time onwards, increasing as we worked away south, indica-

tions of the northward migration of the rabbit were continually found. But

not one solitary bunny did we see till long after, close to Alice Springs.

Despite burrows dug out where droppings seemed fresh, we got no nearer to

solving the problem as to whether they were hibernating, unusually shy, or

killed off by the drought” (Terry 1930: 133).

1h REC, S, AUST. MUS., 16 (11): 1-57 January, \973

(2) [Between hill 47 (Johnson’s Hill) and hill 49 (Murdoch Cliffs),

ie., at ca, 20° 44’, 129° 59’) “. . . we came to 4 huge rabbit warren hacked

about and gouged out like a battlefield, eloquent testimony to many a busy

black fellow getting a cheap teed, This great sandmound, intermingled with

travertine, was distinctive for numerous native orange trees, down by whose

roots the bunnies had burrowed” (Terry 1930: 178).

Terry recorded seeing rabbits only once in the Tanami Desert, a pair

between Mount Solitaire and the Lander River.

Taxonomy: Wakefield (1967) synonymized Bettongia penicillata

anhydra Finlayson, 1957 with B. lesueur, The specimen upon which anhydra

was bused came from the Luke Mackay area at ca 22” 02’, 129° 47’,

[Bettongia penicillata Gray, 1837. Brush-tailed Rat-Kangaroo

As B, p. anhydra Finlayson, 1957 has been placed in the synonymy of

B. lesneur, no specimens of this species are known from the NT. In view

of the similarity, in form and habit of nesting, between B. penicillata and

Lagorchestes spp., | cannot accept as unequivocal the evidence adduced by

Finlayson (1958b) for the existence of B. pericillata in the NT, past or

present. Itas probable that the northern records from the Barkly Tableland

and Gulf drainage listed by Finlayson are referable to the Spectacled

Hare-Wallaby, Lagerchestes conspicillatus, which is there widespread and

not uncammon. |

Lagorchestes conspicillatus Gould, 1842. Spectacled Hare-Wallaby

Map 13

Range: Daly Waters, Borroloola and upper Creswell Creek districts

south to the Macdonnell Ranges, Huckitta and Tarlton Downs (see below).

A neglected record is of a specimen collected by G. F. Hill “SO miles N of

Chewings’ Well” (j.e,, about 105 miles west of Banka Banka) in July

1911 (NMY).

Stains; Formerly common in dense Acacia and other shrub thickets

amongst the central ranges, but now apparently absent from this urea. — Still

fairly common, however, in Acacia thickets in the northern parts of its range,

notably in low thickets amidst taller woodland such as Acacia shirleyi,

lancewood: 6 miles south of Dunmarra (NTM 1961); upper Creswell Creek,

80 miles north of Alexandria (captured and released, 1964); 33 miles north

of Newcastle Waters (NTM 1968); 18 miles west of Dunmarra (NTM

1968), Daly Waters urea (NTM 1968): 42 miles west-south-west of

Botroloola (NTM 1969); all the museum specimens were road-casualties.

Finlayson’s records of Bettongia penicillata from Tanumbirini, Nutwood

and other northern localities are probably referable to this species.

PARKER—NORTHERN TERRITORY MAMMALS 17

Taxonomy: The NT populations are generally referred to L. c.

leichhardti Gould, 1853.

at pe Ar, SAH

BO SING, ar SO ws. ai

lias — ti, PO ; YJ V8 PGs

at oan v Os a . pt tomn ? wt v

> /

ok se we. we * ie

s re, ater "te

Co at a ?

ae - A -

pal ag 4 OA AC fy,

\ ‘i ' Nera K y Ne, "4

| a aaa | NS

| Aa nen | y~

| | | |

| jen ak | | . |

| oO Tennant Crack | | a Tennant Creek |

| | |

| A | | a |

| | | |

| * i 4 | |". : a “ |

ie aa |

| |

| | | s * |

| | ws |

| | lg : |

ee es ett Le i ee ee ie,

| |

Map 13 Map 14

Lagorchestes hirsutus (Gould, 1844). Western Hare-Wallaby

Map 14

Range: |Canning Stock Route between wells 28 and 43, and north of

Sladen Waters, WA; south of Tomkinson, Mann and Musgrave Ranges,

northern SA] Docker Creek; Mount Jenkins; Lake Amadeus area; between

Mount Conner and Murrachurra, 20 miles to the east; Dare’s Plain, George

Gill Range; Wytookarri; northern Lake Mackay; McEwin Hills; Mount

Doreen; west of Warburton Creek; north of the Sandover about 40 miles

west of the Bundy junction; west of Banka Banka; Tanami Desert (NTM,

see below).

Status: Formerly locally common in spinifex-sandridge country,

occurring in often widely separated colonies, this species has declined

drastically since the 1930's. The only recent records are of two small

colonies on isolated sandhills in the Tanami Desert Wildlife Sanctuary

(observations 1958, 1966 (dead animal found), 1967 and 1970). Little of

the extensive sandridge country north of Lake Amadeus and east of Lake

Mackay has been zoologically surveyed recently, however, and it is there that

this species may still be found in some numbers.

18 REC. 8S. AUST. MUS., 16 (11); 1-57 January, 1973

Taxonomy: The NT populations are generally referred to the nominate

race.

Lagorchestes asomatus Finlayson, 1943. Least Hare-Wallaby

Map 13

Range and Status: Known only from the holotype (skull) in the SAM;

from an animal taken in the flesh by Michael Terry a few miles east of

northern Lake Mackay in 1932. Finlayson (1943) gave the position as 22”

15’, 129° 30’, but Terry’s route strikes 129° 30’ at 22° O04’.

Onychogalea lunata (Gould, 1840). Crescent Nail-tailed Wallaby

Map 15

Range: [South of Tomkinson, Mann and Musgrave Ranges, northern

SA] south of Basedow Range; north of Ehrenburg Range: Red Bank, near

Mount Sonder:; Bond Springs; Alice Springs; Huckitta; between Jervois Range

and Tarlton Range; [near the Warburton Range, WA, 1964, vide Ride 1970],

Status: Extremely rare and little-known; few recent records.

Taxonomy: Frith & Calaby (1969) suggested that this species may be

conspecific with the next.

Onychogalea unguifera (Gould, 1841). Northern Nail-tailed Wallaby

Map 15

Range: [Sturt Creek, northern WA] Tanami (NTM_ 1959, 1970);

Katherine (1967); 15 miles N of Katherine (SAM 1966); Marrakat, Adelaide

River plains (1968); Brock’s Creek; Roper River Mission (1966); Banka

Banka (NTM 1965); Elliott area (NTM 1961, 1962, 1968. SAM 1966);

Beetaloo (NTM 1959); Borroloola (NMV 1911); near Cape Crawford

(NTM 1969); “north-eastern corner of Arnhem Land” (Thomson

1949 : 65).

Status: Widespread and not uncommon: lightly-wooded floodplains

in the north, and open long-grass woodland and shrub-savannah in the south;

occasionally on rocky slopes, as in the Tanami Range,

Taxonomy: The NT populations are generally referred to the nominate

race.

Petrogale penicillata (Griffith, 1827) sensu Ride 1970. Brush-tailed

Rock-Wallaby

Map 16

Range: Central Australia: Petermann Ranges, Mount Olga, Ayers

Rock, Mount Conner, George Gill Range, Macdonnell Ranges, east to the

Sandover and Bundey River drainages, north to the Kintore Range, Mount

PARKER—NORTHERN TERRITORY MAMMALS 19

Doreen (25 miles west of), Cockatoo Creek and the Davenport Range

(Finlayson 1961 and NTM records).

Arnhem Land: north-western escarpment: King River Range; Mount

Borradaile (NTM 1968); Oenpelli; Deaf Adder Creek (NTM 1969).

Status: In Central Australia, widespread and locally fairly common, in

rocky hills and ranges, especially in the vicinity of shady gaps and gorges

with an abundance of rock ledges and scree slopes. In Arnhem Land locally

common on sandstone cliffs, outcrops and scree (talus) slopes.

Taxonomy: The Central Australian populations have yet to be sub-

specifically assessed (Finlayson 1930: 179) but are generally referred to

P. p. lateralis Gould, 1842. The Arnhem Land form venustula Thomas,

1926, resembles the Central Australian one in colour and patterning but is

smaller.

Ride (1970) included two other nominal taxa within P. penicillata:

P. wilkinsi Thomas, 1926 and P. longmani Thomas, 1926. Each is known

only by two specimens collected in 1925, wilkinsi from the Roper River

Mission and /ongmani from Groote Eylandt. More material is needed before

a clear picture can be obtained of the taxonomy and distribution of these

northern Petrogale.

sey Wh. a

AX pe? oe, yer

s Pon 2H oS

nd paw: ° We 137

Ti Darwin “a a

A , ia

Wo Ry

A J %&

, Pate Ls

7 <e ~. ce ef fe

\ or lx Neg

| NS | anne

| es ] Sek | y a

| gg Reweastte Waters | | o Newcastle Waters l

| ° | | |

. | bo |

| 9 Tennant Creek | | 9 Tennant Creek |

— | | |

| | | 2 |

| | | ¢ |

| | "8 .

| A aA | ae s !

| =

= 4 B alice Springs f . _ Hes 4 Bee Spriiies |

| | lwa |

| ray | | "=. . |

pert eee eat Mee eee

20 REC, S. AUST. MUS., 16 (11): 1-57 January, 1973

Petrogale brachyotis Gould, 1841, Short-eared Rock-Wallaby

Map 12

Range: [Forrest River, WA (AM 1952)] Daly, South Alligator und

Mary River drainages (Thomas 1926); Gimbat, headwaters of the South

Alligator (AM 1969); Macarthur River and the Sir Edward Pellew Group

(Keith 1968); Red Bank Mine (AMNH 1959).

Status: Apparently local but fairly common where found. Inhabits low

rockfaces and low rocky hills with large tumbled boulders rather than the

more rugged escarpment country of P. P. venustula. (For a photograph of

its habitat in the Pellews see that given for the habitat of Zyzomys argurus in

Keith 1968, fide Keith, in litt.).

Taxonomy; The populations of the Daly, Mary and South Alligator

rivers were described as P. h. signata by Thomas (1926).

Peradorcas concinna (Gould, 1842), Little Rock-Wallaby

Map 14

Range: Daly River; Brock’s Creek; Nellie Creek; Mary River; King

River, north-east of Oenpelli; Milingimbi, Crocodile Islands; Mount

Borradaile-Cooper Creek (NTM 1968); Deaf Adder Creek (NTM 1969);

East Alligator River (NMV, purchased 1922).

Status: Locally common, in rocky areas. The NTM specimens were

taken on open rock flats and tables with scattered boulders (P. p. venustula,

when it occurred in the same areas, was usually found on scree and boulder-

strewn slopes and cliffs),

Taxonomy: The NT populations are generally referred to P. c.

canescens Thomas, 1909,

Unidentified rock-wallabies have been reported from Mount Alexander,

north-eastern Arnhem Land (Wilkins 1928 : 260) and in the high sandstone

ranges of the Macarthur River between Clyde and Western creeks (Barnard

19142211).

Macropus agilis (Gould, 1842). Agile Wallaby

Map 17

Range: [Forrest River Mission (AM 1952)] Victoria River east to the

Macarthur River, inland to the Katherine River; Bathurst Island (NMV

1912); Crocodile Islands; Elcho Island; Sir Edward Pellew Group (Keith

1968); straggling to Elliott area (NTM 1961).

Status: Locally abundant on subcoastal plains and flood-plains of the

larger rivers; grass-pandanus association, grassy forest; recorded also in

mangrove swamps and on beaches.

PARKER—NORTHERN TERRITORY MAMMALS

Taxonomy:

Wallabia to Macropus.

nominate race.

Macropus robustus (Gould, 1841).

ranges and isolated hills.

See Ride (1970) for the transfer of this species from

The NT populations are generally referred to the

Euro

Map 18

Range and Status: Throughout the NT; fairly common, on slopes of

Taxonomy: B. Richardson, in litt., considers the northern populations

referable to M. r. woodwardi Thomas, 1901 (syn. M. r. alligatoris Thomas,

1904), and the central populations to M. r. erubescens Sclater, 1870.

bof , Fs Re /

5 aN "\ . oe RO a ae nt

tees dy ee AAs

poten arwin a

L . % e i f a oe vA roe

i e e ad v

I . e, E ay re

ae ° fos

G Ly Ls

fT NC * a. Ler.n? *

ers \ gw AY ar

| ° ae \o'# I oan

| ‘ ~ | Shs

| oNewenstle Waters. | | co Newcastle Waters |

| | ‘ |

| a

y | | A |

| | A A

| vy? | | |

| | | i” a a |

| | | a ~ * |

pos a

| 3 |

| Te |

| ey | la |

| a7 ’ | | aa 4 |

[po ee at A oe Le, ee a ee ee Lis

| |

Map 17 Map 18

Macropus bernardus (Rothschild, 1904).

Small Black Wallaroo

Map 18

Range: Western escarpment of Arnhem Land: Mary River (SAM

1913): King River Range (Wilkins 1928: 168); Oenpelli (NMV_ 1912-

1914); Deaf Adder Creek (NTM 1969); head of the South Alligator River.

Status: Locally not uncommon, in rugged sandstone and granite ranges,

REC. §. AUST. MUS., 16 (11): 1-57 January, 1973

Taxonomy: Previously considered by some workers to be a race of

M. robustus, with which, however, it is now known to be partly sympatric

(Parker 1971a).

Macropus antilopinus (Gould, 1842). Antilopine Wallaroo

Map 19

Range: South to [the Negri River, WA] Victoria River, Edith River

and Wimul Swamp (SAM 1966), and east to Cape Arnhem Peninsula,

Status; Less common than M. robustus; on grassy and wooded plains,

often in the vicinity of hilly country into which it retreats when disturbed.

Taxonomy: See Frith & Calaby (1969).

Megaleia rufa (Desmarest, 1822). Red Kangaroo

Map 19

Range: North to the Pedestal Hills (Davidson 1905b : 42), 50 miles

north-west of Lake Surprise (NTM files, 1964), Daly Waters (S, A. Parker,

mys., 1969) and Alexandria.

Status: Widespread and locally common; mainly associated with mulga

(Acacia aneura) and lancewood (Acacia shirleyi) on plains, avoiding the

extensive spinifex-sandridge areas.

RODENTS

Hydromys chrysogaster Geoffroy, 1804. Water Rat

Map 23

Range: Daly River east to Cape Arnhem Peninsula, inland *o Brock’s

Creek; Melville Island; Groote Eylandt; Sir Edward Pellew Group (Keith

1968).

Status: Not uncommon, in lagoons and creeks ( mainly freshwater).

Taxonomy: Tate (1951) assigned the NT populations to the small-

molared race H, c. beccarii Peters, 1874. The large-molared race H. c.

reginae Thomas & Dollman, 1909 may possibly enter the NT from western

Queensland.

Xeromys myoides Thomas, 1889. False Swamp-Rat

Map 23

Range and Status: One specimen collected on the South Alligator River

by Tunney in 1903; otherwise known only from six specimens taken in the

Mackay area of Queensland, five from a permanent swamp thickly covered

with tall grass, shrubs and pandanus (Ride 1970).

PARKER—NORTHERN TERRITORY MAMMALS 23

Tunney’s specimen is dated 5 September (J. Mahoney, in litt.). For

most, if not all, of the period 24 August-19 October 1903 Tunney worked

the coastal plain and tidal section of the South Alligator (Storr 1966 : 64).

he wR Ne “0 f pees AN 4 ra

S 5 Lo, g Ask 2 NS ue

Cs " ev, oh ay Za

Doom “ar> Fa Bom PP

ae) . ae f >

— 'e e » 2 e oy

oe ® Vipers “yas

"oat oe De J frre.

. “%, Tonal ¢ ~ a

TPA rt mo aT ae ,

aN r é eta h ‘ erg

| v — | ° me

| | | ‘

| | | |

| v | 2 |

| o Tennant Creek | | cn rennbortredk |

| ; ! |

|v | | ry |

| | ce ‘ |

| | | A |

| | |

| |

| | |

| | | |

| | |

L ee ee L_ Pe ee oth ee [es

Map 19 Map 20

Pseudomys desertor Troughton, 1932. Brown Desert-Mouse

Map 20

Range and Status: Rare and little-known. Waite (1898 : 128) listed

specimens from “Porcupine Sandhills, Wycliffe Creek”, “Barrow Creek” and

“Untaiinua, Porcupine Grass, Alice Springs”*. The only subsequent records

are: specimen trapped on claypan amongst sandhills east of the Bonython

Range at 23° 42’, 129° 02’ (BMNH 1967); specimen dug from a Netomys

burrow at Yuendumu (SAM 1968): remains from recent owl pellets found

in cave 64 miles south-south-west of The Granites in the Highland Rocks,

low lateritic rises and outcrops in sandhill country (NTM 1970).

Waite’s remarks indicate that some of the specimens were taken in

spinifex, at Wycliffe Creek on sandhills. Finlayson (1941) recorded the

habitat of specimens from northern SA as “sandy loam covered by giant

spinifex”, P, F. Aitken, in Jift., considers this species to be normally an

inhabitant of sandhill country, but mentions finding a thriving colony in

tussocks of the perennial sedge Cyperus gymnocaulos growing on the margins

+ The Alice Springs record was inadvertently omitted from map 20.

24 REC. $8. AUST. MUS., 16 (11); 1-57 January, VTS

of a bore-drain near The Neales, western Lake Eyre: the poputation was

possibly living on a nearby sandhill when the bore was put down.

The term “Untaiinua” relating to the Alice Springs specimen is probabiy

the Aranda name for the species rather than a locality (see note under

Notomys longicaudatts).

Pseudomys gracilicaudatus (Gould, 1845). Chestnut Native-Mouse

Map 20

Range: Humbert River (CSIRO 1968); Nourlangie area (CSIRO

1962); 50 miles north-east of Pine Creek on the South Alligator River

(MVZ 1964); upper Nicholson River and Tin Creek (NTM-CSIRO 1967);

South West Island, Sir Edward Pellew Group (Keith 1968, as P. nanus).

Status: Widespread and locally not uncommon; rocky wooded areas

usually near or along watercourses.

Pseudomys fieldi (Waite, 1896). Alice Springs Mouse

Map 20

Range and Status: Known only from the damaged holotype collected

at “Alice Springs” in June 1895,

Taxonomy: See Troughton (1937).

Pseudomys forresti (Thomas, 1906). Forrest's Mouse

Map 22

Range: Springvale and George Creek (NTM-CSIRO 1967);

Alexandria; Brunette Downs (NTM 1966, 1970); Alroy Downs (NTM

1970); Georgina Downs (NTM 1970); Frazer River; Harts Range; Hamilton

Downs (NTM 1961, 1962); Alice Springs: Burt Creek (NTM 1961);

Wollara,

Status: Seasonally fairly common on the open blacksoil plains of the

Barkly Tableland. Elsewhere uncommon or rare: caught on an open grassy

plain on Hamilton Downs, in the bases of Triodia clumps at Wollara, and in

the vicinity of soakages with stands of tall dense paperbark and eucalypt at

Springvale and George Creek.

Taxonomy: P. waitei (Troughton, 1932) has been relegated to the

synonymy of P. forresti by Ride (1970).

Pseudomys delicatulus (Gould, 1842). Little Native-Mouse

Map 21

Range: South to Delamere (SAM 1966) and Elliott (NTM 1968);

West Island, Sir Edward Pellew Group (Keith 1968); Groote Eylandt.

Status: Widespread and fairly common, on sandy soils with cover

varying from sparse herbage to shrubby woodland,

Taxonomy: The nominate race inhabits the mainland while P. d.

mimulus Thomas, 1926 occurs on Groote Eylandt (Johnson 1964).

PARKER—NORTHERN TERRITORY MAMMALS 25

oe BOM. LY

«tl tll elafy & Wires

po . Bdoonn”

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ie ho |

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Map 21 Map 22

Pseudomys hermannsburgensis (Waite, 1896). Sandy Inland Mouse

Map 21

Range; North to Tanami (AM 1952), Banka Banka and Alexandria,

east to Manners Creek (NTM 1969).

Status: Widespread and seasonally common, generally in the same areas

as the Brown Hopping-Mouse, Notomys alexis (q.v.).

Leporillus apicalis (Gould, 1853). White-tipped Stick-nest Rat

Map 25

Range and Status: Stick-nest rats were first reported in the Centre by

Giles: nests found in dense mulga-dominated scrubs between Mount Peculiar

and Mount Udor in September 1872 (Giles 1889 : 57); nest noted in dense

scrub near Mount Squires, eastern WA, in October 1873, “not the first we

have seen by many on this expedition” (Giles 1889 : 209), a remark that

Finlayson (1941 : 228) rendered as “along the 26° parallel between Ayers

Range and the Cavanagh Range”; nest in dense scrub in the vicinity of

Lightning Rock, eastern WA, in late November 1873 (Giles 1889 ; 232).

The Horn Expedition of 1894 apparently did not encounter this species

at first hand; Spencer, who received two specimens from Field of Alice

Springs, wrote (1896b: 11) that it was evidently a rare form. Probably the

26 REC. S, AUST. MUS,, 16 (11): 1-57 January, 1973

last record from the Centre is of a pair captured eight miles west of Mount

Crombie, north-western SA, in 1933. Finlayson (1941 : 228) attributed

the species’ decline mainly to oyerhunting by aborigines.

Notomys aquilo Thomas, 1921. Northern Hopping-Mouse

Map 23

Range and Status: Groote Eylandt, where it is not uncommon in sandy

areas: Umbakumba (Johnson); ‘open forest between Angurugu River and

Umbakumba’ (D, Levitt, in litt.).

Taxonomy: Ride (1970) considered N. carpentarius Johnson, 1959 to

be conspecific with N. aquilo, which was described from the Cape York

Peninsula.

Notomys alexis Thomas, 1922. Brown Hopping-Mouse

Map 23

Range: North to the Tanami Desert, Banka Banka, Alroy and

Alexandria.

Status: The most widespread member of the genus in the NT, seasonally

very common, Occurs on loamy to sandy soil with various plant associa-

tions, usually Triedia and light shrub cover, but also mulya and dense

melaleuca (Melaleuca glomerata).

Taxonomy: See Finlayson (1940); Aitken (1968). WN. a. alexis

occupies most of the range described above. N. a. everardensis Finlayson,

1940 has been collected in the south at Henbury and N. a. alexis > <

everardensiy in the Basedow Range.

Notomys cervinus (Gould, 1853). Fawn-coloured Hopping-Mouse

Map 22

Range and Status: Charlotte Waters (SAM no date, from P. Byrne).

See note below.

Taxonomy: See Aitken (1968).

Notomys fuseus (Wood Jones, 1925). Dusky Hopping-Mouse

Map 22

Range and Status; Charlotte Waters (NMV 1895). See note below.

Taxonomy: See Aitken (1968).

Notomys longicaudatus (Gould, 1844). Long-tailed Hopping-Mouse

Map 22

Range: “Urwaitcha burrows”, Burt Plain; Mount Burrell: Barrow

Creek",

Status: No records since 1901-2, when Spencer and Gillen obtained a

large series from Barrow Creek.

* Amongst recent owl pellets from a cave 64 miles SSW of The Granites in the Highland Rocks

area is a premaxillary-manillary fragment of a large Nofomys considered by J. Mahoney,

in litr,, to belong to either N, lungicaudauinus or N. amplas.

PARKER—NORTHERN TERRITORY MAMMALS 27

Spencer (in Waite 1898: 121) considered that the species he had

referred to as Hapalotis mitchelli in the Horn Report was in fact Nofonrys

longicandatnuy, This would not be wholly true, however, as he collected 13

specimens of N. alexis on the expedition at Reedy Rockhole and Bayot's

(reck (NMYV), which he would have listed as mitchelli in the Horn Report.

In this report Spencer (1896b: 10) wrote under Hapalotis mitchelli “The

mitive name at Charlotte Waters is Ulabaiya’, which Troughton (1965 ; 320)

misquoted as “the aboriginal name about Burt Plains was given us “Ulabaiya’

by Baldwin Spencer”. Unfortunately there ure no specimens of N. alexis

from Charlotte Waters collected by Spencer himsel!, and no extant specimens

of N. longieandatus trom this locality at all, so the association by Finlayson

(1961) of Spencer’s Ulabaiya (which Finlayson misspelt Ulubsiya) with

N. longicaudatus is unsatisfactory. Finlayson (lec. cit.) further gave

Allabuiya as the East Aranda name for “Neromyy spp, close to alexis but not

specifically identified”.

The significance of “Urwaitcha burrows” in connexion with the Burt

Plain specimens (Waite 1898) has not been appreciated. Aranda of the

Alice Springs district gave me a similar name “Urariteha” for a large hopping

mouse, This upart, it is reasonable to assume that in writing “Urwaiteha”

Gillen was recording the aboriginal name of the species. Cf. “Oorarrie”,

the Wonkanooroo name given by Finlayson (1961) for N. cervinuy.

I cunnot trace the Mount Burrell specimen listed by Finlayson, Possibly

it was collected by T. W. Cornock in 1891 (see Appendix 1), and is the

spiril specimen without data in the SAM,

Notomys amplus Brazenor. 1936. Short-tailed Hopping-Mouse

Map 22

Range and Status: Known only from the two specimens collected ut

Charlowe Witers in 1896 (NMV). See note below,

Norn; Netomys cervinus and N. fuscus apparently have their head-

quarters in the sandhill-claypan-gibber associations of the southern Lake

Eyre Basin, the former burrowing mainly in claypans and the latter in sand-

hills (an ecological separation found also between Macrotis lagotis and

M. leucura) (Finlayson 1939; Aitken 1968 and in /ift.). Both species

occur marginally and sporadically in the NT (Charlotte Waters 1895, 1896)

during periods of increase. Byrne (in Spencer 1896b: 11) wrote in July

1895, after good rains, “The jerboa-like rodents are coming from the eastwards

and (they almost amount to a plague here”, and in September 1895 “they have

again become so searce that the blacks have difliculty in securing a specimen”.

Notomys amplus may similarly have its centre of distribution outside the NT,

Mitchell's Hopping-Mouse Nefomys mitchellii (Ogilby, 1838) has been taken

at Birdsville in south-western Queensland (Tate 1951: 259) and nay, with

the other species discussed here, appear in the southernmost NT during good

seasons.

28 REC. 8. AUST, MUS., 16 (11): 1-57 January, 1973

a \ ee

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Map 23 Map 24

Zyzomys argurus (Thomas, 1889). Common Rock-Rat

Map 24

Range: [Parry’s Creek, near Wyndham, WA] Cooper Creek (NTM

1968); Deaf Adder Creek (NTM 1969); Oenpelli; Boroloola (NMV

1901-2); upper Nicholson River and Springvale (NTM-CSIRO 1967);

Groote Eylandt; Sir Edward Pellew Group (Keith 1968).

Status: Common, amongst boulders and on rocky substrate areas at

the bases of rocky hills.

Zyzomys pedunculatus (Waite, 1896). Waite’s Rock-Rat

Map 24

Range: Alice Springs and Illamurta (1890's); Hugh Creek (1935);

Napperby Hills (1950); The Granites (AM 1952); Davenport Range

(1953); Haast’s Bluff Settlement, Derwent River (NTM 1960).

Status: Rare and little-known; range country and isolated rocky

outcrops. The Haast’s Bluff specimen was caught raiding a stock-camp.

See Finlayson (1961).

The vernacular name employed by Ride (1970), Macdonnell Range

Rock-Rat, is considered no longer appropriate.

PARKER—NORTHERN TERRITORY MAMMALS 29

Zyzomys woodwardi (Thomas, 1909). Woodward’s Rock-Rat

Map 24

Range and Status: Rare and little-known; rocky sandstone outcrops of

the north-western escarpment of Arnhem Land: Oenpelli; Mount Borradaile

(NTM 1968). [Parry’s Creek, near Wyndham, W.A.]

Mesembriomys gouldii (Gray, 1843). Black-footed Tree-Rat

Map 25

Range: Daly River area north-east and east to Cape Arnhem Peninsula,

inland to Katherine River; 5-mile bar, Macarthur River (NMV_ 1911);

Melville Island.

Status: Not uncommon in tall woodland with hollow trees, especially

along watercourses and near the coast.

Taxonomy: The race M. g. melvillensis Hayman, 1936 was described

from Melville Island; see Johnson (1964).

Mesembriomys macrurus (Peters, 1876). Golden-backed Tree-Rat

Map 25

Range: Nellie Creek (1903); Deaf Adder Creek (NTM 1969);

Balanbrinni (NMV 1901).

Status: Much less frequently collected than M. gouldii; wooded water-

courses.

Se? Eo A BE ‘oa ek

aoa “ wep >

a v ats f "jee,

— ° “et Ge ‘ 3

J A e A 7 7 9 7"

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Map 25 Map 26

30) REC. S. AUST, MUS, La (11); 1-57 January, V973

Conilurus penicillatus (Gould, 1842). Brush-tailed Tree-Rat

Map 26

Range: Daly River area north-east and east to the South Alligator River

und Coburg Peninsula; Roper River Mission (AMNH 1959); Bathurst

Island (NMV 1916); Melville Island; Groote Eylandt; Centre Island, Sir

Edward Pellew Group (Keith 1968).

Status; Not uncommon, along the coast and on the subcoastal plains,

with hollow trees; recorded on Coburg Peninsula actually on the tideline

and amongst hollow beach-casuarinas (Casuarina equisetifolia) (Gilbert, in

Troughton 1965 : 311).

Taxonomy: The race C. p. melibius Thomas, 1921 has been described

from Melville Island. The provenance of the Elsey specimen upon which the

name Conilarus hemileucurus Gray, 1858 was based has yet to be precisely

determined; most of Elsey’s zoological material was collected at the Victoria

River Depot No, 1.

Melomys Thomas, 1922. Mosaic-tailed Rats

Map 22

Range: Port Keats, north-east and east to Cape Arnhem Peninsula,

inland to Douglas River and Brock’s Creek; Bathurst Island (NMV 1916);

Melville Island; Elcho Island; Groote Eylandt; Sir Edward Pellew Group

(Keith 1968).

Status; Not uncommon, especially near the coast; grassy pandanus

fringes of fresh- and salt-water streams and lagoons, and the edges of

monsoon forest, in the vicinity of damp ground: mangroves. On Groote

Eylandt, reported entering occupied houses (D. Levitt, im Jitt.).

Taxonomy: See Tate (1951): Johnson (1964). Tate considered most

of the above range to belong to forms of Melomys cervinipes (Gould, 1852),

with the exception of Groote Eylandt, occupied by M]. Jutillus mixtus

Troughton, 1935. Ride (1970) and Calaby, in Jirt., are of the opinion that

the NT populations of the genus are in need of a thorough revision before a

clear picture of their taxonomy and nomenclature can be obtained.

Rattus tunneyi (Thomas, 1904). Tunney’s Rat

Map 27

Range: Douglas Rivet: Mary River; Brock’s Creek; Oenpelli; Coburg

Peninsula (CSIRO 1965); Melville Island; Sir Edward Pellew Group (Keith

1968); Calvert River (OM 1969); Tennant Creek; Alice Springs.

Status: Widespread but apparently local. In the Pellews this species is

the most numerous and widespread rodent, burrowing extensively in the

PARKER—NORTHERN TERRITORY MAMMALS 3I

well-drained sandy ridges (Keith 1968). Elsewhere it has been trapped

along creeks, There have been no records of the Central Australian popula-

tions subsequent to Spencer's donation to the NMV in 1916; the dates of

collection of Spencer's specimens are not precisely known but some at least

were obtained during the period 1894-1897. Of the female from Tennant

Creek listed by Waite (1898 : 124), which could not be traced by Brazenor

(1956), the skull without skin is in the SAM reg. no. M. 2411).

Taxonomy, The race R. ¢. melvilleus Thomas. 1921 has been described

from Melville Island, and R, ¢, dispar Brazenor, 1936 from Alice Springs.

Rattus colletti Thomas, 1904. Northern Territory Dusky Field-Rat

Map 27

Range: King River, north-east of Oenpelli (BMNH, coll. Wilkins,

1924); South Alligator River; Beatrice Hill (NTM 1961, 1967).

Status: Very local but not uncommon where found; clay flats and

moarshy areas,

Tevonomy: Ride (1970) synonymized R. colletti with R, sordidus

(Gould, 1858), but subsequent chromosome studies have suggested that the

former should for the present be maintained as a separate species (J, Mahoney,

in litt.),

Rattus villosissimus (Waite, 1898)*. Long-haired Rat

Map 26

Range and Status: Vhis species appears, fram its distribution during the

less favourable seasons, to have its headquarters in the Lake Eyre basin-

Georgina-Diamantina drainage, where it occurs in highly-localized colonies

around bore-drains, reedy springs and other wet, densely-vegetated spots

(P. F. Aitken, in litt, P. K, Latz and D. Howe, pers. comm.). At intervals

(of five to seven years according to Finlayson 1939, 1941) that may be

correlated with the amelioration of conditions by one or successive seasons

of good rains, the species undergoes a spectacular increase, reaching plague

proportions in the Georgina drainage and on the Barkly Tableland, generally

on open plains in the vicinily of creeks and bores. It may swarm for several

yeurs before declining in numbers as spectacularly as it increased. Such

plagues are accompanied by equally notable build-ups of predators, especially

the Letter-winged Kite, Elanus scriptus, whose movements appear to be

closely-ticd to those of the rat (Parker 1971b), and the Barn Owl.

Tyre alba.

In the NT. peripheral localities reached by the rat are: South-West

Islund, Sir Edward Pellew Group (Keith 1968); Katherine (CSIRO 1968):

Humbert River (CSIRO 1968); Mongrel Downs (196%, J. Mahood, pers.

* Date of publication given incorrectly as 1897 by Iredale & Troughton (1934 ; 73) and Ride

(1970 5 227),

cal

32 REC. S. AUST. MUS., 16 (11): 1-57 Janitary, 1973

comm.); Napperby Creek; Delny (NTM 1968); Tarlton Range; Marqua

(NTM 1968); Pituri Creek; New Crown (possibly from the Dalhousie

Springs colony, fide P. F. Aitken, in /itt.).

CARNIVORES

Canis familiaris Linnacus, 1758. Dingo

Range and Status: Widespread and locally common,

Taxonomy: Australian populations are generally referred to C. f. dingo

Meyer, 1793. The earlier name Canis antarticus |sic| Kerr, 1792 was

suppressed in Opinion 451 of the International Commission on Zoological

Nomenclature, 1956.

vf / ‘ r

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eae % \

| a | ) \

| | : |

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Map 27 Map 28

BATS

Pteropus scapulatus Peters, 1862. Little Red Flying Fox

Map 28

Range: Mainly coastal districts, inland to Deaf Adder Creek (NTM

1969), Wimul Swamp (SAM 1966) and Borroloola (NMV 1912); Elcho

Island. With the flowering of eucalypts it may move much further south,

and has been reported from Buchanan Creek and Playford Creek on the

Barkly Tableland (regularly), Banka Banka, Frew River and Davenport and

PARKER—NORTHERN TERRITORY MAMMALS 33

Murchison Ranges (frequently; specimen from McLaren Creek, NTM

December 1965) and Arthur Creek, Pituri Creek and Sandover River

(occasionally). In adjoining western Queensland it moves seasonally down

the Georgina (D. Howe, pers. comm, ).

Status: A blossom-feeder, common in the northern part of its range,

usually in eucalypt and eucalypt-pandanus woodland, in a variety of country

from coastal flats to sandstone gorges. Observed roosting beneath the dry

skirts of pandanus (S. A. Parker, ms.) and in paperbarks (Melaleuca) along

watercourses (J. L. McKean, in Jitt.).

Pteropus alecto Temminck, 1837. Black Flying Fox

Map 28

Range: Mainly coastal and subcoastal: Victoria and Daly Rivers east

to the South Alligator River and Cape Arnhem Peninsula; sometimes further

inland, as at MeColl’s Bore, Armstrong River (NTM 1958).

Status: More of a fruit-eater than the preceding; recorded in large

numbers from monsoon forest, riverine thickets and mangroves.

Taxonomy: The Australian populations belong to the race P. a. gouldii

Peters, 1867 (Johnson 1964).

Macroglossus lagochilus Matschie, 1899, Northern Blossom Bat

Map 29

Range: Darwin area (W. P. Walsh, in /itt.); Melville Island (SAM

1913); Coburg Peninsula (CSIRO 1965); Tortilla Flats, Adelaide River

(NTM 1967).

Status; “Sometimes locally abundant. Present throughout the year in

Darwin Botanical Gardens, with apparent peak in spring and early summer

(up to 20 seen feeding in same tree). At Rapid Creek, species taken in

Tristania lactiflua forest fringing stream margin” (W. P. Walsh, in litt.).

The Tortilla Flats examples, three, were mistnetted in an arcade in dense

riverine bamboo.

Taxonomy: The Australian populations belong to the race M. lL. nanus

Matschie, 1913 (McKean, in itt.).

Taphozous georgianus Thomas, 1915. Common Sheath-tailed Bat

Map 30

Range and Status: Throughout the NT; common, roosting in caves and

mineshafts.

Taxonomy: McKean & Price (1967) regarded the northern populations

as T. g. troughtoni Tate, 1952, and suggested that the central populations

may represent the nominate race.

34 REC. 8. AUST. MUS., 16 (11): 1-57 January, \973

aus es / chats aug A ?

Se * Ni, oe iA ee StS 1

eee “yoy . Wi PF 6

ta “hfs pies OAT

hje ° i s ae mS A as oe

A . fe Pe fe

a ‘ A bey * at -

Hy A’ S Tr AG A

j q ‘en ; P Nee

= a

| | a

| o Newcastle Waters a I | #Neweastle Waters al

| l a |

| : | |

| { . |

| oT nt Creek | | Tennant Creek |

| | | * A

| | ° |

| | | |

| ! | .

| | | |

| x & Alice Springs ! ea A segetice Spings |

| | | a |

| | Lee |

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Map 29 Map 30

Taphozous flayiventris Peters, 1867. Yellow-bellied Sheath-tailed Bat

Map 30

Range: Point Charles (NMV 1938); Darwin; Lake Deane (W. P. Walsh,

in litt.); Coburg Peninsula (CSIRO 1965); Deaf Adder Creek (NTM 1969);

Cape Arnhem Peninsula; Groote Eylandt; Block Waterhole, upper Nicholson

River (NTM-CSIRO 1967); Tanami Desert at 20° 09’, 130° 15’ (NTM,

AM 1965); Newcastle Waters (NTM 1959); Banka Banka (CSIRO 1968);

Warburton Creek-Tower Creek junction; Temple Bar Creek (CSIRO 1968);

Petermann Ranges and Bonython Range (BMNH 1967).

Status: Widespread but possibly less common than T. georgianus.

Roosts in hollow trees; several of the above specimens were shot over water.

Macroderma gigas (Dobson, 1880). Ghost Bat

Map 29

Range: Macallum Creek area (Finlayson 1958a); Darwin; Mount

Wells, Pine Creek, Cutta Cutta and Owenia Rockshelter, colonies (W. P.

Walsh, in litt.); Oenpelli (MCZ, NMV 1918); King River, north-east of

Oenpelli (NMV 1915); Elcho Island (J. McKean, in Jitt.); Deaf Adder

Creek (NTM 1969); Ronans Cave (SAM 1966); Groote Eylandt (AM);

Red Bank Mine (AMNH 1959); Frazer River, Field River, Alice Springs,

Mount Conway, Ellery Creek (Finlayson 1958a : 923).

PARKER—NORTHERN TERRITORY MAMMALS 35

Status; Rare and vanishing in Central Australia, but locally abundant

in the north. A cave-dweller,

Taxonomy: M. g. saturata Douglas, 1962 is available for the darker

northern populations,

Hipposideros ater Templeton, 1848. Dusky Horseshoe Bat

Map 32

Range: Owenia Rockshelter (W. P. Walsh, in Jitt.); Douglas River;

Ocnpelli; King River, north-east of Oenpelli (NMY 1915); Deaf Adder

Creek (NTM 1969); Cutta Cutta (SAM 1966, CSIRO 1969).

Status; Uncommon, apparently needing pitch-dark caves with humidity

near saturation-point (W. P, Walsh, in Jitt.). Mainly in sandstone areas.

Taxonomy: See McKean & Price (1967). The Australian populations

belong to H, a, aruensis Gray, 1858.

Hipposideros diadema (Geoffroy, 1813). Diadem Horseshoe Bat

Map 32

Range and Status: Known from the NT by 13 specimens mistnetted

amongst tall paperbarks (Melaleuca) in a sandstone gorge on Deaf Adder

Creek in September 1969 (NTM, CSIRO). Doubtless a cave-dweller, as it

is in Queensland,

Yaxonemy: This isolated population has been described as a new sub-

species, HW. d. inornatuy McKean, 1970.

Hipposideros stenotis Thomas, 1913. Lesser Warty-nosed Horseshoe Bat

Map 32

Range and Status: Three of the four NT records are from the western

escarpment of Arnhem Land: Mary River, two collected from caves in May

1895 by Dahl, who found the species not uncommon: King River north-east

of Oenpelli, two collected from “sandstone cliffs” in 1915 by McLennan

(NMYV); Deal Adder Creek, one taken by hand from crevice high in sand-

stone cliff in September 1969 by D. Howe (NTM). The fourth record is

from the Gulf drainage: specimen from Red Bank Mine area near

Wollogorang in 1959 (AMNH).

Rhinonicteris aurantius (Gray, 1845). Orange Horseshoe Bat

Map 33

Range: Darwin (AM), Port Essington; Adelaide River (NMV 1912):

Oenpelli; Maranboy (AM); Cutta Cutta and Smokey Creek (W. P. Walsh,

in litt.).

Status: “Widespread and common near the coast: distribution may

change seasonally, The Cutta Cutta population is c. 50,000 at present unless

| have seriously miscalculated the ratio of Rhinonicteris to Hipposideros ater

there” (W. P. Walsh, in /itt., 1970).

36 REC. §. AUST. MUS., 16 (11): 1-57 January, 1973

Gould quoted the following observation (requoted in Johnson 1964 ;

474), that on the Coburg Peninsula this bat “retires during the daytime to

the hollow spouts and holes of the various species of Eucalypti’. All

subsequent observations of the roosting of this species, however, indicate that

it is a cave-dweller,

Tadarida australis (Gray, 1839). White-striped Bat

Map 31

Range: Scattered localities from the Petermann Ranges (BMNH 1967)

north-east to the Stuart Bluff Range (NTM 1961) and Allambi (NTM

1968).

Status: Locally common,

Tadarida loriae (Thomas, 1897). Little Northern Scurrying Bat

Map 31

Range: Coburg Peninsula; Mount Borradaile (NTM 1968); Seven-Ten

Waterhole, upper Nicholson River (NTM-CSIRO 1967).

Status: Uncommon; the NTM specimens were shot or netted over

waterholes at dusk.

Taxonomy: T. 1, cobourgiana Johnson, 1959 was based on specimens

from the Coburg Peninsula.

SO bAnee RO “ ae Z a

2 vs rom, :

DaPh Darwin . POG

¢ eo een ae

a ies —_— es

eye “ha, ~ R pat ¥ —s e

Nota Nea

oe | ZS.

A |

| co Newcastle Waters 1 | o Newcastle Waters -

| lie |

| | | |

| « | | |

| |

| | | |

| ae ! | |

! Yet | | |

loa * | | |

| | | |

few

Map 31 Map 32

PARKER—NORTHERN TERRITORY MAMMALS 37

Tadarida planiceps (Peters, 1866). Little Flat Bat

Map 31

Range: Lasseter’s Cave, Petermann Ranges (NTM 1969); Alice Springs

area (NTM 1955, 1960, 1969); 14-mile Bore, Alcoota (CSIRO 1968);

Maryvale (NTM 1970); Ross River (NTM 1970); Tea Tree Well (NTM

1963).

Status: Widespread but apparently uncommon; roosting in hollow logs

and branches,

Tadarida jobensis (Miller, 1902). Northern Mastiff Bat

Map 31

Range: Darwin (AM); near White Stone Creek, Marrakai (NTM

1969); Alexandria; Seven-Ten Waterhole, upper Nicholson River (NTM-

CSIRO 1967); [Doomadgee, north-western Queensland (SAM 1963)].

Status: Apparently uncommon; roosting in tree-hollows.

Taxonomy: The Australian populations belong to the race T. j. colonica

(Thomas, 1906).

Myotis adversus (Horsfield, 1824). Large-footed Myotis

Map 35

Range: Coburg Peninsula (CSIRO 1965); Seven-Ten Waterhole,

upper Nicholson River (NTM-CSIRO 1967); Melville Island (NMV 1938).

Status; Uncommon and local. Recorded roosting close to water, in

caves und culverts (within smaller crevices rather than on the bare walls)

and. in hollow trees,

Taxonomy: The Australian populations belong to the race M. a

macropus (Gould, 1855); see McKean & Hall (1965).

Chalinolobus gouldii (Gray, 1841). Gould’s Wattled Bat

Map 34

Range and Status: Widespread and common in the south, north to the

upper Nicholson River (NTM-CSIRO 1967) and Banka Banka; further

north only two isolated records: Douglas River (Johnson 1964) and Coburg

Peninsula (NTM 1961). Roosts in tree hollows and amongst leaves.

Taxonomy: The name C. g. venatoris Thomas, 1908 (type-locality

Alexandria) is employed for the northern populations. Central Australian

populations are variable and represent a cline between C. g. venatoris and

the nominate race (J. L. McKean, in Iitt.).

38 REC. S. AUST. MUS., 16 (11): 1-57 January, 1973

SO we

ei AN

Darwii PAZ

a 8 . Pye

a Ie

% e a

a R ?. Nes

e a | e Sete

| ~ | ye

| é | | ws

| | :

| | | ; fe |

| |

| | |

| | | fy fy

| v Alice Springs | a t Alice Springs 4

! 29 | la 4 |

l | | a” |

be eg ett L_ bi et ee I

| |

Map 33 Map 34

Chalinolobus morio (Gray, 1841). Chocolate Bat

Map 33

Range and Status: In the NT known with certainty only by six specimens

netted over Quarantine Bore, 12 miles south-south-west of Alice Springs, in

1969 (NTM). Finlayson (1961) listed “Chalinolobus cf. morio”™ from

Wollara.

Chalinolobus nigrogriseus (Gould, 1856). Hoary Bat

Map 33

Range: Brock’s Creek; Coburg Peninsula (CSIRO 1965); Cape

Arnhem Peninsula; Groote Eylandt; Sir Edward Pellew Group (Keith 1968);

Red Bank Mine (AMNH 1959); China Wall and Block Waterhole, upper

Nicholson River (NTM-CSIRO 1967); 47 miles north of Daly Waters

(NTM 1961).

Status: A northern species. Widespread but uncommon. Recorded

roosting in rock crevices.

Taxonomy: The NT populations belong to the race C. n. rogersi

Thomas, 1909 (H. M. Van Deusen, in litt.).

Eptesicus Rafinesque, 1820. Little Bats

Map 35

Range and Status: Throughout the NT, common; roosting in caves and

mineshafts.

PARKER—NORTHERN TERRITORY MAMMALS 39

Taxonomy: The name FE. pumilus caurinus Thomas, 1914 is generally

used for the NT populations of this genus. In view of the remarks of McKean

& Price (1967) and McKean, in litt., concerning the need for a revision of

Australian Eptesicus, no specific name is here used.

Nycticeius balstoni (Thomas, 1906). Broad-nosed Bat

Map 36

Range: [Forrest River Mission (AM 1952); Sturt Creek, WA] Darwin;

Fogg Dam (CSIRO 1964); Marrakai HS (NTM 1963); 46 miles south-east

of Adelaide River town (NTM 1958); Red Bank Mine (AMNH_ 1959);

Border Waterhole (NTM-CSIRO 1967); Alexandria; Tennant Creek;

Napperby Creek (NTM 1969); Karanji Bore (NTM 1963); Alice Springs

area (NTM 1960, 1965, 1969); Horseshoe Bend.

Status: Widespread and locally common. A specimen from Wire Creek

was one of five roosting in a hollow tree-limb with two Tadarida planiceps.

Taxonomy: The northern populations are generally referred to N. b.

caprenus (Troughton, 1937) and those of Central Australia to nominate

N. b. balstoni. J. L. McKean, in litt., considers that the type of Scotophilus

greyii Gould, 1858 (type-locality Port Essington) may prove to belong to

the taxon at present known as N. b. caprenus.

ete We! SOM

“ie cis Te he neigey co Wir asy

AP Dain ae f Oy

L, ¥ f JZ Hf

ca re (ore ie

aa fhe t f

“% v Ay j Pt 4

eA AS Fins alt yy Ag io.

nN - \\batq \ 44

wv NE

| v i % | ra

<i | i

| | N vat s

| o Newcastle Waters | ewcastle Waters |

| a | |

| 3 ah

| B Jennant Creek | | @ Tennant Creek

| | |

| |

| | |

l ¥ | | |

’

| | .

| | | . |!

le ¥ Nico spring | | gi dlice Springs

| ’ | | |

| Lome |

Sin ates Le Pe Ics a Rg

Map 35 Map 36

40) REC. §. AUST. MUS., 16 (11): 1-57 January, 1973

Miniopterus schriebersii (Kuhl, 1819). Bent-wing Bat

Map 34

Range: Darwin area; Tortilla Flats (NTM 1967); Virginia Mine

(W. P. Walsh, in Jitt.); nine miles west of Katherine (SAM 1966); Deaf

Adder Creek (NTM 1969); Timber Creek (NMV 1963).

Status: Locally common, in deep caves, mineshafts and abandoned

buildings. W. P. Walsh, in Jitt., estimated the Virginia Mine colony to

number more than 50,000 bats. The nine Deaf Adder Creek specimens were

shot in sandstone gorges, six over a waterhole.

Taxonomy: The NT populations are generally referred to M. s. orianae

Thomas, 1922.

Nyetophilus geoffroyi Leach, 1821. Lesser Long-eared Bat

Map 37

Range: North to False Smoke Hills (AM 1965), Tennant Creek,

Alexandria and Springvale (NTM-CSIRO), with an isolated record from the

Katherine area (NTM).

Status: Scattered but locally not uncommon. Roosts in trees, occa-

sionally in caves and rock-crevices.

Taxonomy: The NT populations are generally referred to N. g.

pallescens Thomas, 1913.

Nyctophilus bifax Thomas, 1915. North Queensland Long-eared Bat

Map 37

Range: Daly River; Brock’s Creek; Melville Island; Coburg Peninsula

(CSIRO 1965).

Status: Unknown. Dahl found it abundant around the Uniya Mission

on the Daly River in 1894, Of the Brock’s Creek specimens, one was flying

over a waterhole, the other clinging to the bark of a tree in daylight (Johnson

1964 : 479),

Taxonomy: The NT populations belong to the race N. b. daedalus

Thomas, 1915 (Johnson 1964).

PARKER—NORTHERN TERRITORY MAMMALS 4)

ERO —_ ae

eral ON oF

cane sd: el

“ fy iy A

’ / i ‘*

Ny 4 “L

fram “ Td ne

AY \ are

any

o Newcastle Waters !

a |

@\ Tennant Creek

ae Alice Springs

Map 37

Nyctophilus arnhemensis Johnson, 1959. Arnhem Land Long-eared Bat

Map 37

Range: King River, north-east of Oenpelli (NMV 1915); Tortilla

Flats, Adelaide River (NTM 1967); Brock’s Creek; Cape Arnhem Peninsula;

Groote Eylandt; Sir Edward Pellew Group (Keith 1968); Springvale

(NTM-CSIRO),

Status: Little-known but probably not uncommon; in dense tall paper-

bark woodland, passages in monsoon forest and the gloomy arcades of

dense riverine bamboo. Found roosting beneath the bark flaps of large

paperbarks,

Taxonomy: There is a possibility that Nyctophilus arnhemensis is a

junior synonym of Nyctophilus walkeri Thomas, 1892, which name was based

on a single specimen from the Adelaide River (J. L. McKean, in Jitt.).

ACKNOWLEDGMENTS

To the following mammalogists my most grateful thanks are due: Mr. P. F. Aitken,

South Australian Museum, for the loan of material, for details of specimens in his care and

for some most fruitful discussions on vernacular names; Mr. M. Archer, Western Australian

Museum, for identifications in the Dasyuridae* and for allowing me to include some of his

unpublished conclusions concerning this family; Dr. J. L. Bannister, also of the Western

Australian Museum, for details of material collected by Mr. H. Butler for the British Museum

(Natural History) in 1967; Mr. J. H. Calaby, C.S.1-R.O. Wildlife Research, Canberra, for

42 REC, S, AUST, MUS. 16 (tl); 1-57 January, 1973

venerously allowing me to include details of material collected on the Coburg Peninsula

(1965) and for his valuable comments on numerous points of taxonomy and nomenclature;

Miss J, Covacevich, Queenslind Museum, for the loan of material and for details af other

specimens in her care; Miss J, M. Dixon, National Museunr of Victoria. for the Joan of

mitterial anc for details of hee entire holdings of N2TL mamoals: Mr J, Edwards Hill and

Miss P, D. Jenkins, British Museum (Nature! History), for their critical comments on

specimens collected by Stalker and Wilkins; Mr. C. W. Mack, Museum of Compurative

Zoology, Harvard University, for supplying details of material; Mr. J. L. Mekean, € SALR.O.

Wildlife Research, Canberra, for identifications in the Chiroptera and for his guidance in the

taxonomy und nomenclature of this order; Mr. J, A, Mahoney, University of Sydney, for

identifications in the Muridue* and for many helpful comments; Mr, Basil Marlow, Australian

Museum. for details of material in his care: Mr, B. Richardson, University of New South Wales,

for his comments on the Osphrafrer-group in the genus Maeropis; Dr A. M. Van Deusen,

American Moseum of Natiral History, foy details Of material in his care.

Mr. B. Hart. Director, Primary Industries Braneh, N/T. kindly allowed ime to tise the

records and files of the Northern ‘Territory Museum, including the results of the numerous

faunal surveys undertaken by the Branch.

For additional records and comments |oam indebted to: Mr, R. Berry, Darwin; Mr.

R. Darken, Alice Springs; my colleagues Messrs. D, Egan, D. Howe and BP. K. Late:

Miss D. Levitt, Groole Bylandts and Mr, W. P. Walsh, North Australian Biological Records

Centre, Darwin, who Allowed me to quote from his extensive fieldnotes on the Chiroptera of

the northern NT.

_ For help with biographical details of collectors 1 thank: Mr. and Mrs. D. Boerner,

Mice Springs; Mr. A. H. Chisholm, Sydney; Mr. Wayne Dodd, Darwin; Dr W) Grant Inglis,

Direetor, South Australian Museum; Mrs. 1. Nluot, Melbourne: Mr, ¥.'T, O'Brien, Director,

Lands and Survey Branch, NTA, Daywin, wid Dr. H. M. Van Deusen

My thanks also to colleagues past and present whose efforts hive helped to build up the

zoological collections of the Northern Territory Museum: Mr. B. E,, Ballon, Mr, b Corbeit,

Mr. P, Hanisch, Mr. W, B. Hitchcock, Mr. D. Howe, Mr, D. Lindner. Dr. A, RB. Newsome,

Mr. A. O. Nicholls, Mr. T, Ryan, Mr, A, K. Shipway, Mr. K. R. Slater, Mr, D. R. Stephens,

Mr. P. Sullivan and Mr, H. Wakefield, Mr. Howe made many valuable contributions to

the draft.

L owe special thanks to Mrs. PL Newnnin und Mes, K) Clark, whose excellent typing was

matched only by their forbearance,

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Aitken, P, F., 1971, Rediscovery of the Large Deserl Sminthopsis (Sriiitlepaty psarinophilus

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Appendix I

NOTES ON COLLECTORS AND COLLECTIONS

_ These notes are fol exhaustive, In the main they treat new information and obscure

points, encountered during the checklists preparation, that would be of yalue in the compila-

hon of & comprehensive bibliography. See also Whittell (1954) and M. Specht (1964).

Cuhill, P. ( - ). Between 1912 and 1920 Cahill collected birds in the Oenpelli-Eust

Alligator River area; most of these are now in the NMV. In 1913 he donated a specimen

of Mesembriomys gouldié, In 1922 the NMV purchased from Mrs. Marie Cahill five

specimens of Mucropus bernardus and one of Periloreas concinna collected by Cahill.

Cornock, Thomas W, ( - ). Went as an assistant with Edward (later Sir) Charles

Stirling (then Hon. Director, South Australian Museum) in (490-91 when Stirling

accompanied the Governor of South Australia, the Earl of Kintore, to the NT. In the

SAM there wire a few bird specimens of Cornock’s, labelled “Mount Burrell, 1891" and

“Port Darwin, March 1891", Possibly he obtained the Mount Burrell specimen of

Notomys longicandatus Cav).

Dahl kK. CUS7I-1951). Victoria River—Daly River—Mary River—Souih Alligator River,

1894-1896 (Dahl 1926), Dahl's notebooks and the great majority of his collections are

iv the University of Oslo, Norway. A comprehensive study of this material has yet to

supersede the piecemeal accounts by Collett and Thomas.

db REC. S. AUST. MUS., 16 (lh): 17 January, VTS

De Teliga, G, W. ( - ). See Schevill, W. B.

Dodd, Walter Dempsey (1891-1969). Born 27 March T1891 al Toowong, Queenstiand, in

1913 Dodd collected mammals and birds (now in SAM) on Melville Island, Bathurst

Ishind and the Oppec Mary Rivet, His Melville Island birds were written op by Zielz

(19l4da, 19}4b); his mammals have never formed the subject of a report, Dodd diced

17 May 1969 at Brisbane.

Elsey, J, Ry (1834-1858). Accompanied A. C. Ciregory’s 1855-1856 expedition weross northern

Australia. Most of bis zoolegical material (now in the BMNH) was obigined at the

Victoria River Depot No. 1. Although Elsey conventiated on birds, he obfained a few

oMmals, Most of which are now without their data. See Chisholn (1964, 1966) und

Macdonald & Colsian (1966),

Finlayson, tl. WW. (1895- ) Ag uuthority on the mammals of Central Austratia,

Honorary Curator of Mammals at the South Australian Musetim from 1940 10 1966, he

amassed a large private collection, the inaccessibility of which is a source of regret

amongst mammilogists. Finlayson's crowning pauper on Ceutrl Australian mumeals

uppearcd in 1961.

villen, Fo J. (1855-1912), Born 28 October 1455. Accompanied W, Baldwin Spencer on

several expeditions, Some, if not all, of the early material from the Burt Plain north of

Alice Spans was Obtained by Gillen, Gillen died of amyotrophic lateral selerosis on

S June 1912,

Toll, G. FB. (1880-1954). Naturalist on the Barclay Exploring Expedition to Central Australia

and the Northern Territory, [O11-1912; Government Entomologist. NT. 1912-1917. No

coniplete report was ever poblished concerning the results of the Barclay Expedition, on

which Hill made collections (now in (he NMY) of birds, nauimals, reptiles, insects and

plants. For a dated itinerary of the outward stage of this expedition (Oodnadatta to the

Micarthur River) see Ewart & Davies (1917. mip). Orher notes may be found i Hill

(1913) and Campbell & Kershaw (1913). Hill's notebooks of 1901-1912. mostly in

shorthand, together with those of H. Vere Barclay, are in the Commonwealth Archives,

Canberta; the journal of the expedilion’s suryeyor, R. McPherson, is in the oflice of the

Lands und Survey Branch, Northern Territory Administration. Darwin, The study and

publication of material relating to this expedition would he a most valuable undertaking,

Hill died wl Sydney on U8 January 14a.

Hosmer. William T. (ra. 1930- ). Born in Ireland. An Australian-based herpetologist,

leader of the [960 Spalding-Hosmer Expedition, whieh visited the NT (specimens,

including w@ few mammals, in AMNH), Also on the 1959 Spalding-Peterson Expedition,

Tluy, C. M, [ - ). Dotelns River and Brock's Creck, 1920, Noy's specimens are in

the USNM and were detuiled by Johnson (1964).

Juhnson, David Horn (1912- ). Born Albany, Oregon, 9 Seplember 1912. Curator of

Mammals. United States National Museum, (941-68, Mammatogist on the 1948

American-Australian Expedition to Arnhem Land (Johnson Lo6d).

Lestung, Albert de ( - ), A Frenchman, an atateue naturalist whe lived on a small

property nmumed Adel's Grove some 90 miles south-west of Rurkelowo on Lawn Hill

Creck, north-western Quecasland, His main interest appeurs to have heen botany, and

he sent plant collechions to the Queensland Merhartum, Brisbane, Troughtaun (1965 5

29-30) mentioned a series of Manigale iigramié (gv.) in the AM collected hy de Lestang.

Melennan, W, R. (U8R2-1995). Coastal NT west to the King River (north-east of Oenpelli),

1915 (McLennan 1917). Although McLennan conceatrated on birds, he obtained a lew

mammals (now in the NMYV) from the siundstone ranges of The Kine River, which Wilkins

(g-v-) visited in 1924, ti omithological Hlerature McLennam's name is Trequently spell

Maclennan (ew. Whittell 1954) and Milennan. However, Melennan is the form

uppearing on his birth-certilicate und) used by him in his letters (Mrs. ‘IV’. Kloot, Archivist,

Royal Australasian Ornithologist Union, bp fr).

Melerson, Russell Francis (1922- ). Born FT) June 1922 af Montelair, New Jersey,

Mammalogist associated wilh (he AMNH 1952-59 Led the 1959 Spulding-Pelerson

Expedition to Queensland and the N'T (specimens in AMNH), Now a free-lanee editor,

author and artist, living in Jamestown. Rhode Iyhind. LLS.A. His several books include

® popular work on bats (Peterson 1964).

Schevill, W. BE, ( - ). Member of the 1931-32 Hurvord Australian Expedition. tA

the MCZ there is a collection of 48 mammal specirhens collected in the tlermannsbure,

Barrow Creek und Teatree Well wreas in 1932 by Sehevill & G. W. De Teliga. Although

‘Tate mentioned 4 few of these specimens in his revisions, the collection wis never reported

in its entirety. See Loveridge (1934 5 244-5),

PARKER—NORTHERN TERRITORY MAMMALS 47

Spulding, Philip Cea, 1903- ), Financed the (959 Spalding-Peterson Expedition and the

1960 Spulding-Hosmer Expedition,

Spencer, W. B. (1860-1929), Zoologist on the Hora Expedition to Central Australia in 1894.

He made several subsequent trips to the NT, some with F. J. Gillen (see Spencer &

Gillen 1912, Spencer 192%), Spencer's contacis of 1894 (P. Byrne of Charlotte Waters,

b. Cowle of tlamurta, e¢ al.) continued sending him muainniuls for some years, “The

Spencer manual collection is a vithuable one of both historical and scientific significance,

However, the pussuge of tine has resulted in considerable dilliculty yo identification of

much of the type material Some specimens baye vanished, possibly destroyed, while

others Have lost most of their date” (Dixon 1970: 105). One might add that in other

cases, us wilh the “Barrow Creek” material (see Gazetteer), some of the specimens seem

to have heen incorrectly localized, Details of the Horn Expedition itinerary more precise

thun those found ip the Horn Report appear in Winnecke (1897).

Stalker, W. (1879-1910), Alroy Downs, Alexandria Station, 1905-1906. At that time

Alexundrin Covered a far greater area than now (see Ingram 1907). Restriction of the

provenance Of Stilker’s material to present-day Alexandria is therefore Unsatisfactory, and

work needs to be carfied out on a dated itinerary. Stalkers material is in the BMNEA,

leriy, M. (1899 ), In the 1920's ant 1930's Terry explored large tracts of Central

Austria, notably the Tanami Desert and the Lake Mackay Lake Amadeus revions. He

obtained mammals for HM, H, Finlayson, amongst them the holotypes of Lagerchestes

asomatus and Betoneia penicillata anhydra, Verry’s books on these surveys (1925, 1927,

1930, 1931, 1937) are full of the most interesting detail,

‘Thomson, D. F, (1901-1970). “Donald ‘Thomson went in 1935 to Arnhem Land

He made an extensive zoological collection, now at Melbourne University, primarily as a

backeround (6 the study of the ecology and food-gathering habits of the aborigines. The

material has been worked Upon, but no systematic publication has been made of the

collections, mainly manimals, reptiles and amphibians” (M, Specht 1964: 5). During

Thomson's liter years at least, these collections were not available to workers, “Their

critical examination and publication is a matter of the first importance.

Thomson dicd on 13 May 1970 at Flthum, Victoria, His ashes were scattered from

a plane over Caledon Bay in north-eastern Arnhem Land.

Vindale, N, B. (P900- ), Roper River 1921: Groole Eylandt 1921-1922, Tindale

collected a few mammals (now in the SAM), Which have not been reported. See Tindale

(1925),

funticy, J. PF) (1871-1929). Arobem Land, 1902-1903. Tunney'’s large bird ind mammal

collections are dispersed amongst the BMNH, WAM, USNM and AMNH.— Storr (1966),

using information from Tunney’s bird specimens, elucidated much of Tunney’s itinerary

in the NI. A study of the mammal labels, in conjunction with Storrs findings. would

prove aw Most profitable exercise,

Wilkins, Sir Cieorge Hubert (1888-1958). For Wilkins’ activities in the northern NT during

1924 and 1925, consult his bowk (1928). His collections, whieh include a good number

of mammal specimens, were deposited in the BMNE, although paratypes of several new

forms were presented to the Queenslund Museum. Wilkins’ “King River” is not the

ivibutary of the Katherine River as Thomas (1926) thought. but the King River north-east

of Oenpelli and south of the Goulburn Ishinds (Wilkins 1928 : 200; Johnson 1964: 455),

Wilkins’ collections have not yet been fully reported,

Wilkins died overnight 30 November-! December 1958. Some months later Com-

mander James Calvert, U.S, Navy, carried his ashes north beneath the Polar lee where

they were scallered on 17 March 1959,

Appendix Il

GAZETTEER OF PLACE-NAMES

_ Lititudes are south, longitudes east, jo the nearest mine. Positions given for cytile

slutions wre these of the present homesteads, See alsa Story (Y67 = 71-83

Adelaide River, rises ot 13° 28% 131° 00% flows N into Van Diemen Gulf at 12° 134 191° Tea

Adelaide River, town on the Adelaide River where it is crossed by Stuart Highway 72 uviles §

of Darwin, at (4° 15", 441° 07°

48 REC. S. AUST. MUS, [6 (11): 1-57 January, 1973

Alexandria, cattle station on Barkly Tableland, 19° 03', 136° 42’, See Appendix Lb under

W. Stalker.

Alice Springs, lOWA On the Todd River, 23° 42', |

Springs” sometimes came from further afield:

were actually from the Burt Plain.

Allambi, cattle station, 24° 13', 134° 25".

Alroy (Downs). cattle station on Barkly Tableland, 19° 17, 136" 03"

Angurugu River, Groote Bylandt, estuary at 13° 58’, 136° 24,

Arllunga, farmer gold town, 23" 25%, 134° 43,

Arnhem Land, formerly denoted the: far north of the NT between the Timor Sea and the

Gulf of Carpentaria; now generally restricted to the Aboriginal Reserve in the eastern

two-thirds of this area. The greater part is occupied by the predominantly sandstone

Arnhen) Land Plateau, the flanks of which are koown as the Arnhem Lund Escarpment.

Arthur Creek, crossed by Jervois Stock Route at 22° 41’, 136° 41’.

Attack Creek, crossed by Stuart Highway at 19° 02’, 134° 08.

Ayers Range. 25° 55’, 133° 09%,

Ayers Rock, 25° 21’, 131" 02’.

Bagot Creek (Camp 23 of the Horn Expedition, 1894), leaves George Gill Range al 24° 24%

131° 48. Not to be confused with the Bagot Creek 9 miles W of Herminnsburg,

Balunbrinni (on label of Mesembriomys macruris, NMV): Spencer (1928 : 552) referred to

“a small waterhole on the Leila Creek, called Ballanbrini”, Leila Creek is a tributary of

the Upper Macarthur River. I have failed to find a waterhole of this name. Lying on

Spencer & Gillen’s route between O.T, Downs and Leila Creek there ts a Balanbrini

Creek (a tributary of the Limmen Bight River) crossed at 16° 35’, 135" 20°. Pending

further research, one must leave the Ballanbrini Walerhole of Spencer & Gillen on the

track between O.T. Downs and the Leila-Macarthur junction,

In the NMY there are several bird specimens from this expedition labelled

“Ballanginie”’, possibly a variant spelling of the ubove.

Bunka Banka, cattle station ou Stuart Highway, 18° 47’. 134° 03".

Barkly Tableland, north-eastern region of plains of great pastoral value, about 300 miles long

by 100 miles wide, orientated NW-SE between 17° and 20°S; bounded in the N and

NE by the Gulf of Carpentaria drainage (Carpentaria full), in the W and S by semi-

desert, and in the SE by the Georgina basin,

Barrow Creek, small town on Stuart Highway, 21° 31, 133° 53 Spencer & Gillen called

here in 1901 and 1902; some of their specimens labelled “Burrow Creck”, however, may

have come from elsewhere (see remarks under Dasyuroides byrnei anc Macrotisy leucura).

Indeed, it is possible that material collected further south and further north was despatched

from Barrow Creck and so labelled upon ils receipt in Melbourne. Similarly, “Alice

Springs” and “Tennant Creek” may often have denoted points of despatch rather than of

provenance. (Dr. G. M. Storr, in fit, now considers that the seven specimens of

Ctenoms robustus Storr (Scincidue) obtained by Spencer and labelled “Barrow Creek

were actually collected considerably further north.)

Busedow Range, 25° 05’, 132° 05".

Rathurst Island, immediately W of Melville Island, 11° 40’, 130° 20°

Bealmée Hill, 35 miles ESR of Darwin, 12° 39, 131° 19’.

Beetaloo, cattle station, 17° 14’, 133° 47°,

Bentinck Islund, Qld, in Gulf of Carpentaria ul 17° 03’, 139° 30".

Birdsville, Qld, town at 25° S54’, 139" 21.

Block Waterhole, upper Nicholson River, 17° 356’, 137° 09’,

Bobby's Well — Kurtitina Well, qv.

Bond Springs, cattle station, 23° 33’, 133° 56’.

Honython Range, WA near the NT border, 23° 38’. 128° 59’.

Border Waterhole, N'T/Old border, 18° 36’, 138° 00%,

Borroloola, small town on the Macarthur River, 16° O04’, 1367 18"

33° 51. Barly material labelled “Alice

some of Gillen’s specimens so labelled

PARKER—NORTHERN TERRITORY MAMMALS 49

Bronce Crees station on North Australian Railway, 27 miles SE of Adelaide River, 13° 28’,

31° 25"

Brunette Downs, cattle station on Barkly Tableland, 18° 38, 135° 37°.

Buchanan Creek, left-bank tributary of the Playford 12 miles SE of Alexandria.

RBundey River. begins at 22° 50°, 134° 35’, flows NE into the Sundover al 21° 45’, 135° 37’.

Burt Creek, crossed by Stuart Highway 37 miles N of Alice Springs, 23° 12’, 133° 45".

Burt Plain, in broadest sense the entire plain immediately N of the western and central

Macdonnell Ranges, but usually restricted to sector N of Alice Springs. Some of Gillen’s

“Alice Springs” material carne from the Burt Plain.

Bushy Park, cattle station, 22° 54’, 133° 56%,

Calvert River, flowing NE into Gulf of Carpentaria at 16° 16’, 137" 45’.

Camooweal, Qld, town, 19° 55’, 138" O08".

Cantyteina Well = Kurtitina Well, g.v.

Cape Arnhem Peninsula, 12° 20’, 136" 50’.

Cape Crawford, northern point of an inland sandstone range, 16° 39", 135° 48".

Carmichael(’s) Crag, western George Gill Range, 24° 13’, 131° 33%.

Cavanagh Range. WA, 26° 13, 127° 56’.

Cavenagh Range — Cavanagh Range.

Cedar Bay, Qld. 15° 49", 145° 22°

Central Australia, officially (formerly) the NT south of 20°,

Centre Iskind, Sir Edward Pellew Group, 15° 41’, 136" 46’,

Charleville, Qld. town, 26° 24’, 146° 15",

Charley Creek, begins S of Mount Hay at 23° 34’, 133° 06’, flowing N to floodout at 23° 07’,

132° 59°,

Charlotte Waters, walerhole on Coglin Creek at 25° 55’, 134° 54’, with the former telegraph

station half a mile io the S at Station Point. Coglin Creek divides sandhill country to

the N from gibber plains to the S. Only here do the Lake Eyre basin gibber associations

intrude into the NI, which may explain the occurrence here and nowhere else in the

NT of Dasvuroides byrnei, Notamys fuseus, N. cervinus and the bird Ashbyia lavensis

(Gibber Bird).

Cheepie, Qld, small town, 26° 38, 145° OV.

China Wall, escarpment on Jeft bank of upper Nicholson River, 17° 46’, 137° 1%.

Clyde Creek (marked Glyde River on current 1 : 250,000 sheet), right-bank tributary of the

Macarthur River which it joins at 16° 25’, 136° O08”.

Cobure (Cobourg) Peninsula, north-western extremity of mainland at 11° 20’, 132° 15%

Cockatoo Creek, 22" 06’, 132° O8”

Cooncherie, SA, waterhole at ca. 26° 48’, 139° 29", ca. 18 miles S of Old Clifton Downs HS.

Cooper Creek, mses in the NW escarpment of Arnhem Land. flows into estuary of East

Alligator River at 12° O07’, 132° 41". NTM collections were formed within 2-3 miles of

Mount Borradaile (qg.v.).

Cooper Creek, drainage channels from the junction of the Barcoo and Thomson rivers in SW

Old at 25° 10’, 142° 55’, flowing SW into Lake Kopperamanna, NE SA, at 28° 35,

138" 45",

Coulthard’s Well = Kurtitina Well, g.v.

Creswell Creek, begins 17° 44’, 136° 40°, floods out at 18° [8’, 135° 035°,

Crocodile Islands, off N coast, centred on 12" 00°, 135° 00’.

Croker Island, off N coast, 11° 10°, 132° 35”.

Crown Point, table-topped hill on the Finke River at 25° 30’, 134° 23’. The site of Old

Crown Station is 2 miles SSW at 25° 32%, 134° 23".

Curtin Springs, catlle station, 25° 19’, 131° 495°,

Cuttu Cutta (Katherine Caves), 16 miles S of Katherine.

Dalhousie Springs, SA, hot springs and permanent swamps, 26" 31', 135° 29”

st REC, S. AUST. MUS., 16 (11): 1-57 January, (973

Dalmore Downs, cattle station on Barkly Tableland, 19° 45’, 135° 59’, Not to be confused

with Delmore Downs on the Bundey River.

Daly River, flowing NW from the confluence of the Katherine and the Flora at 4" 25%,

131° 39° into the Timor Sea at 13° 23’, 130° 18’.

Daly River Mission (formerly Uniya), on the lower Daly at 13° 45", 130° 41%

Daly Waters, town near Stuart Highway, 16° 15 133° 20°

Dare’s Plain, 17 miles NW of King's Canyon, 24° O01, 131° 35°.

Darwin, city on NW coast, 12° 287, 130° 51%

Davenport Range, centred on 20° 50’, 135" O00’.

Deaf Adder Creek. rises on Arnhem Land Plateau, flows W into Nourlangie Creek at 12° 59,

132° 48’. NTM collections were mainly from the sandstone escarpment at ca. 13° 06’,

132° 456’.

Delamere, cattle station in Victoria River basin, 15° 44’, 131° 32’,

Delny, cattle station, 22° 33%. 134° 49"

Desert Block (formerly part af Napperby), part of Amburla station, 23° 04’, 133" 01

Diamantina River, flows SW through SW Qld, into the Lake Eyre basin,

Docker Creek —= Docker River.

Docks ger aboriginal settlement on watercourse of same name, Petermann Ranges, 24” 48",

Douglas River, rises NW of Pine Creek, flows W into the Daly River al 13° 41% 131° 09°

Dunmarra, cattle station on Stuart Highway, 16° 40’, 133° 23°,

Eust Alligator River, rises on Arnhem Land Plateau, flows NW to the coast al 12° 10’,

132° 35’,

Edith River, siding on North Australian Railway where if Crosses the Edith River at 14° 12%

132° 02’,

Ehrenburg Range, 23° 17%, 130° 21’.

Elcho Island, off NV coast, 11" 57’, 135° 44’.

Elkedra, cattle station, 21° 10’, 135° 28",

Ellery Creek; Finlayson (1958a) gave a locality for Macroderma gigas as “the Ellery Creek

gorge in the James Range at about lat. 24° 5’ S. and long. 132° 49" E.”.) This places us

on the Finke River near its junction with Ellery Creek: 24° QS’ strikes Ellery Creek at

132° 50’.

Elliott, town on Stuart Highway, 17° 34’, 133° 31".

Ethel Creek, begins at western end of Treuer Range, flows W to floodout at 22° 124, 130° 04".

False Smoke Hills: 1 have submitted this name to the NT Geographical Nomenclature

Committee, to be used for the feature at 20° 04°, 130° 10° in the Tanami Desert incorrectly

named Smoke Hills on a recent 1 : 250,000 The Granites sheet. The true Smoke Hills,

named by Davidson (1905b:55) and referred to by Terry (1930), are at 20" 15%,

129° 55’. and are marked as Cave Hill on some recent maps. I overlooked this error

when drawing up a previous gazetleer (Parker 1969; 71), Material affected is hal

collected in the vicinity of “Smoke Hills* (—= False Smoke Hills) by the Australian

Museum in 1965.

Field River: Finlayson (1958a) under Macroderma gigas, gave the position of “eaves on the

Field River” as about 23° 34. 137" 53’. This places us amongst sand dunes, most

unlikely country for caves. L. Corbett, who visited the Field River in 1968, was told by

stalion hands at Tobermorey that bat caves existed in hills to the westward of Smith's

Re anne hills in question are almost certainly the Mount Knuckey group at 23° 32’,

Foge Daum, 12° 34’, 131" 18%

Forrest River Mission, WA, 26 miles NW of Wyndham, at 15° [l’. 127° 51"

Four-mile Hole, Wildman River, 12° 36°, 132" 13%,

Fourteen-mile Bore, Alcoota Station, 23° 01, 134° 29/,

Sr rE (Frazer Creek), right-bank tributary of the Bundey, which it joins at 22° 22%

PARKER—NORTHERN TERRITORY MAMMALS 51

Frew River, begins in Davenport Range at 21° 00', 135° 00°, flows NE towards Barkly Table-

land to flaodout at 19" 58’, 135° 40%

George Creek, south of upper Nicholson River, crossed by track at 18° 15%, 137" 16’. Not

fo be confused with creeks of the same name in the Victoria and Mary River drainages-

George Gill Range, 24° 205 131° 45’.

Georgina Downs, cattle station, 21° 07’, 137° 39”.

Georgina River, begins SE of Barkly Tableland, flows SSE through W Queensland.

Gimbal, homestead on the South Alligator at 13° 34’, 132° 36°.

Glen Helen, cattle station, 23° 25’, 132° 15”.

Groote Eylandl, large island in the Gulf of Carpentaria, 14° 00’, 136° 40°.

Haast’s Blufl, aboriginal settlement in western Macdonnell Ranges, 23° 27, 131° 53’, 10

miles SW of mountain of same name.

Hall's Creek, WA, old township at 18° 15’, 127° 47’, 8 miles E of new Hall's Creek,

Hamilton Downs, cattle station, 23° 32% 133° 16.

Harts Range (Depdt), former mining camp at 23° 00°, 134° 55% The present police station

is ca, 2 miles to the N.

Hatches Creck, mining township, 20° 56’, 135° 12’,

Henbury, cattle station, 24" 33’, 133° 15’.

Hermannsburg, mission station on the Finke River, 23° 57’, 132" 47’.

Horseshoe Bend, cattle station, 25° 13’, 134° 13%.

Huckitta, cattle station, 22" 54’, 135° 27’.

Hugh Creek = Hugh River.

Hugh Ee begins in western Macdonnell Ranges, flows SE into the Finke River at 25° 02’,

34° 10".

Humbert River. cattle station, 16° 29°, 130° 39°

Ilamurta, former police camp 26 miles S of Hermannsburg ut 24" 21, 13° 41%

James Ranse, 24° 15’, 133° 05" In broader sense includes the KrichaulT Range.

Jervois Range, 22° 37, 136° 15’.

Karanji Bore, 23° 17% 133° O08’,

Katherine, town on Stuart Highway and Katherine River, 14° 28’, 132° 16’.

Katherine Caves — Cutta Cutta (q.v.).

Kateane da nses on Arnhem Land Plateau, flows W into the Daly River at 14° 25’,

* 39!,

King River, rises on Arnhem Land Plateau, flows NE to the coast at 11° 50’, 133° 32%,

This is the river worked by McLennan in 1915 and Wilkins in (924; it has been

confused by Thomas and others with the King River that flows into the upper

Katherine.

King River Range, a name used by Wilkins (1928) for rocky hills 15 miles up the river

from its mouth. Johnson (1964 : 455) reckoned this camp to be at 11° 56’, 133° 25”

According to MeLennan (1917) the ranges along the King River are of sandstone.

King’s Canyon, George Gill Range, 24° 15%, J31L° 34. A “tourist” name, which some

diehard. Central Australians refuse to use. The native name for this remarkable canyon

and the creek that issues from it is Wudalka.

King’s Creek. strictly speaking, the creek that flows SW out of King’s Canyon to floodout at

24° 25" 131° 25" Formerly used also for the feature now known as King’s Canyon,

Kintore Range, 23° 21°, 129° 23°,

Kurtitina Well, a native well (now probably quite obliterated) between Lake Amadeus and

Ayers Rack at 25° 02% 131° 14’,

Kyancutta, SA. township, 33° 08’, 135° 34’.

Lake Amadeus. large salt-lake. 24° 50’, 131° 00%

Lake Buck. Tanami Desert, 19° 38’, 130° 20°.

52 REC. S. AUST. MUS., 16 (11): 1-57 January, 1973

Lake Christopher, WA, salt-lake, 24° 44’, 127° 34”

Lake Deane, 12° 44°, 131° OV.

Lake Eyre, SA, huge salt-lake. 28° 30’, 137° 20’.

Lake Mackay, large salt-lake on WA/NT border, 22° 20’, 129° O00’.

Lake Surprise. Tanami Desert, on the Lander River at 20° 12%, 131° 49°

Lasseter’s Cave (incorrectly Lassiter’s), Hull River, Petermann Ranges, 25° OJ’, 129° 24’.

Lightning Rock, WA, 26" 04’, 127° 45’.

Lock, SA, township, 33° 34’, 135° 46’.

Macallum Creek: Finlayson (1958) under Macroderma gigas, gave the locality “the table-

land scarp in the Macallum Creek area at about lat. 13° L6’S. and long. 130° 44'E."

This places us a littlke SE of @ Macallum Creek in the Reynolds River area. As the

“tableland scarp” referred to is probably meant to be that of the Arnhem Land Plateau,

the Macallum Creek in question may be the one further east (a tributary of the

Margaret River), crossed by a track at 13° 16°, 131° 27’.

Macarthur River (incorrectly MeArthur), begins on the Barkly-Carpentaria divide, flows

NE into the Gulf of Carpentaria at 15° 50°, 136° 40°.

McColl’s Bore, Armstrong River at ca, 16° 39’, 131° SI’.

Macdonnell Ranges, collective name for ranges along 23° 40’, between 131° 00! and 135° 00’.

McEwin Hills, 22" 00’, 129° 35’.

Mackay, Qld, town on E coast, 21° 09%, 149° LL’.

McLaren Creek, crossed by Stuart Highway at 20° 20’, 134° 14.

Malbon, Qld, small town, 21° 045 140° 18”.

Mangajera Billabong, Roper River (Tindale), probably Mungejirri Yaalput Waterhole, 14° 32’,

135° 15‘, on the Wukointyarra Plains, a salt-marsh area near the mouth of the Roper,

Mann Range, NT/SA border, centred on 24° 05’, 130° 00’.

Manners Creek, cattle station, 22° 07’, 137° 59”.

Maranboy, siding on North Australian Railway 30 miles ESE of Katherine at 14° 40’,

132° 39". Maranboy police station is 14 miles NE.

Marqua, cattle station, 22° 495, 137° 21".

Marrakat, cattle station, 12” 48’, 131° 22”

Mary River, rises on Arnhem Land Plateau E of Pine Creek, flowing NNW to the coast at

12° 17’, 131° 40°.) The species collected on this river by Dahl and Dodd indicate that

the two collectors worked the rugged country of the upper reaches.

Melville Island, large island W of Coburg Peninsula, 11° 35% 131° 00’.

Milingimb), mission station on island of same name off N coast at 12° O05‘, 134° 55’: one of

the Crocodile Islands.

Mongrel Downs (Tallalah Downs), cattle station, 20° 34’, 129° 44°

Mount Alexander, 12° 11’, 136° 40’.

Mount Borradaile, 12° 04’, 132° 51%

Mount Burrell, former overland telegraph depdt on the Hugh River at 24° 36°, 133° 55%

Later became Old Maryvale HS. A hill of the same name lies 4 miles SF. See

Appendix I, under Cornock,

Mount Conner, 25° 30’, 131° 53%.

Mount Conway, 23° 45’, 133° 26”,

Mount Crombie, SA, 26° 39°, 130° 50’.

Mount Doreen, cattle station, see Vaughan Springs.

Mount Heughlin, 23° 21, 132° 14°

Mount Jenkins, 25° 397, 129° 40’,

Mount McMinn (Spencer, 1911), right bank of the Roper River at 14° 43’, 134° 22’,

Mount Olga, 25° 16’, 130° 44’.

Mount Peculiar, 23° 26’, 131° 16’.

Mount Razorback, 23° 32", 132° 27’,

PARKER—NORTHERN TERRITORY MAMMALS 53

Mount Riddoch, cattle station, 23° 02’, 134° 40°.

Mount Riddock — Mount Riddoch.

Mount Shoebridge Mount Shoobridge.

Mount Shoobridge, 13° 32’, 131° 19% Tinfields of same name 2 miles W.

Mount Solitaire, Tanami Desert, 20° 32’, 131° 14%

Mount Sonder. 23° 35’, 132° 34%

Mount Squires, WA, 26” 13%, 127° 29’. Not to be confused with Mount Squire(s), NT,

25° 195 134° 18".

Mount Udor, 23° 30’, 131° O17.

Mount Wells. 13° 30°. 131° 43’,

Murchison Runge, centred on 20° 15’, 134° 25"

Murrachurra (Murrathurra) Springs, 25° 27", 132° 14”

Musgrave Runges, SA, 26° 15’, 131° 30’.

Napperby, cattle station, 22° 31’, 132° 45”,

Nupperby Creek, crossed by North-West Stock Route at 22° 49°, 132° 35’,

Napperby Hills, thase running W-E past Napperby HS.

Negri River, rises at 17° 40, 129° 25’, flows NW into the Ord River. WA,

Nellie Creek, 17 miles NE of Pine Creek.

Neweastle Waters, small town 17 miles NW of Elliott at 17° 22", 133° 22".

New Crown (Point), cattle station on the lower Finke River at 25° 42’, 134° 50’.

Nicholson River, begins at 17° 35’, 137° 00°, flows E into Qld.

Nourlangie, safari camp, 12° 46’, 132° 39%

Nutwood (Downs), cattle station, 15° 49’, 134° 09”,

Qenpelli, Mission station on western escarpment of Arnhem Land 6 miles E of the lower East

Alligator River at 12° 20’, 133° 03’,

Old Marsh Bed, a great depression in the Tanami Desert containing a chain of salt-lakes and

claypuns, churacterized by dense stands of Melaleuca glomerata and the giant mounds of

Nasutitermes iriediae, Most pronounced from ca, 20° 09’, 130° 15’ (swinging SW, S,

SE and E) to ea. 20° 50’. 130° 37’.

Ogualpi Plain, 35 miles NW of Hermanonsburg, a once-famous haunt of the Oqualpi,

Lagorchestes conspicillatus (Finlayson 1961 : 167).

Owenia Rockshelter, 13" 30’, 131° 12’.

Painter Spring, 14 miles NW of Alice Springs, 23° 35’, 133° 42°,

Palmer Creek -= Palmer River.

Palmer River, right-bank tributary of the Finke, which it joins at 24° 46%, 133° 23’.

Purry’s Creek, WA, small watercourse 17 miles SE of Wyndham.

Patonga, 12° 55°, 132° 41",

Pedestal Hills, Tanami Desert, 20° 34, 129° 17°.

Pelermann Ranges, 25° 00’, 129° 30’,

Pine Creek, town on Stuart Highway, 13° S50" 131° 50”.

Pituri Creek, flows past Tobermorey HS, crossing the NT/Qld border 2 miles SSW.

Playford Creek == Playford River.

Playford River, begins on Carpentaria-Barkly divide, flowing SW and W past Alexandria,

Plenty River, begins in Mount Riddoch area, flows E and SE to floodout in northern Simpson

Desert at 24" 20’, 137° 00%.

Point Charles, on coast 15 miles WNW of Darwin.

Port Essington, inlet on N coast of Coburg Peninsula, 11° 15’, 132° LO’. Also used for former

settlement on its shores, officially known as “Victoria”.

Port Keats, mission station on W coast at 14° 144, 129° 32",

Port Langdon, inlet on NE coast of Groote Fylandt.

Quadjinta (Quajinta), see Yaringa.

34 REC. §, AUST. MUS,. 16 (11): 1-57 January, 1973

Quarantine Bore, 12 miles SSW of Alice Springs.

Rapid Creek, 7 miles NNE of Darwin.

Red Bank (Redbank), gorge in the western Macdonnell Ranges halfway between Mount

Razorback and Mount Sonder.

Red Bank (Redbank) Mine, ca. 18 miles W of Wollogorang at 17° [1', 137° 45%,

Reedy Rockhole, George Gill Range, 24" 18’, 131° 36’.

Ringwood, cattle station, 23° 50’, 134° 56’.

Ronans Caye, near Katherine, 14° 28’, 132° 13”.

Roper River, rises near Maranboy, flows E into Gulf of Carpentaria at 14° 43’, 135° 23),

Roper River Mission, on the lower Roper at 14° 44’, 134° 44°.

Ross River, tourist camp, 23° 36’, 134° 29°,

Ryan(s) Well, on Stuart Highway, 22° 43°, 133° 23".

Sandover River, formed by junction of Muller Creek and Waite Creek at 22° 30’, 134° 31’,

flows NE to floodaout at ca. 21° 30’, 137° 00

Sandringham, Qld, cattle station, 24° 03’, 139° 04".

Sevyen-Ten Waterhole, on the upper Nicholson River, 18° QO 137° 17%

Sir Edward Pellew Group, islands in Gulf of Carpentaria opposite the Macarthur River

estuary, centred on 15° 35", 134° 50°,

Sladen Waters (incorrectly Sladden), WA, a short watercourse flowing through the Pass of

the Abencerrages, Rawlinson Ranges, at 24° 5&7 128° 17’, about 5 miles N of Giles

meteorological station.

Smokey Creek, 13° 10’, 131° 06%,

South Alligator River, rises on Arnhem Land Plateau, flows N to the coast at 12° 12’, 132°

23°.

South West Island, Sir Edward Pellew Group, 15° 35°, 136° 52’.

Springvale, south of the upper Nicholson River, 18° 30’, 137° 32’. Not to be confused with

the Springvale 4 miles SW of Katherine.

Station Point, see Charlotte Waters,

Steele Gap. in eastern Rodinga Range, 24° 24", 135° 05%, leading south into the Simpson

Desert.

Stuart Bluff Range, centred on 22” 46’, 132° 34.

Sturt Creek, WA, cattle station on watercourse of same name, 19° 09’, 128° 09",

Tanami, a gorge containing two long-lasting rockholes (the Camel Waterholes of A. A.

Davidson), in the range of the same name, 19° 58’, 129° 40’. The main Tanami gold-

fields are three miles E,

The correct (aboriginal) pronunciation is Tanami (Gee 1910:;5). Nowadays. the

name is frequently pronounced Tanamai, however (ltalian transeription ),

Tanami Desert (Tanami semi-desert), a shrub-sayannah area lying roughly between 18° O0'-

22° OO and 129° O0'-134° 00" First explored by A. A. Dayidson, to whose account

(Davidson 1905b) readers are referred.

Vanami Desert Wildlife Sanctuary, an area of 14,490 square miles in the Tanami Desert.

Holds good numbers of Macrotiy logotis and contains the only extant colonies of

Lagorchestes hirsutus known in Central Australiq.

Tunumbirini, cattle station, 16° 09", 134° SI’.

Tarlton Downs, cattle station, 22° 39’, 136° 54”

Tarlton Range, hills to the S of Tarlton Downs HS.

Teatree Well, small town on Stuart Highway, 22° 08’, 133° 25%

Temple Bar Creek, crossed by Adelaide Road 12 miles SSW of Alice Springs.

Tennant Creek, town on Stuart Highway, 19° 395 134° 114 7 miles S of watercourse and

former telegraph station of same name.

The Granites, former goldfield, 20° 34°, 130° 21’, Tanumi Desert.

The Neales, SA, drainage channels entering NW Lake Eyre.

Timber Creek, police station, 15° 39’, 130° 28, near A.C. Gregory's Victoria River Depot

No. 1.

PARKER—NORTHERN TERRITORY MAMMALS os

lin Creck, south of Upper Nicholson River, crossed by track al 18° 38’, 137° 47%.

Tobermorey,, cattle station, 22° 17’. 137° 56’,

‘Toko Range, lies across NT/Qld border at ca. 22° 45°. 138" 00,

Tombinson Ranges, extreme NW corner of SA, 26° 10%, 129" 20°,

Portilla Flats (Upper Adelaide River Experimental Station), 13° 05‘, [31° 13’.

Tower Creek, begins N of Napperby, flows into Warburton Creek al 22° 12°, 132° 33°.

Umbakumba, mission station on NE part of Groote Hylandt.

Union (Town), former gold town & miles NNW of Pine Creek at 13° 43’, 131° 49"

Uniya, former name of Daly River Mission (q.v.).

Vaughan Springs, site of the new Mount Doreen HS, at 22" 18%, 130° 52’. The old HS

is at 22" 03° 131° 19,

Victoria River, rising at ca. 18" 00%, 130" 00’, flowing NW into the Joseph Bonaparte Gulf

at 15° OO’, 129° 357,

Victoria River Depot No, 1, on the lower Victoria River at 15" 37’, 130° 27’. The provenance

of most of Elsey’s zoological collections.

Virginia Mine. 13" 10’, 131° 03%

Wajelai Billabong (Tindale), not found but prebably on the Wukointyarra Plains near the

mouth of the Roper River; near Mangajera Billabong (see Anrechinonis laniger),

Warburton Creek, begins NW of Napperby., flows into Crown Creek at 22" 03’, 132° 30’.

Warburton Mission, WA, 26° 09', 126° 32’,

Wauchope, small town on Stuart Highway at 20° 39%, 134° 13".

Well 28, Canning Stock Route, WA, 22° 39", 123° 45%,

Well 43 (Billowaggi Well), Canning Stock Route, WA, 21° 12’, 125° 58’.

Well 46 (Kuduarra Well), Canning Stock Route, WA, 20° 08". 126° 17’.

West Island, Sir Edward Pellew Group, 15° 35%, 136° 34’

Western Creek, le(t-bank tributary of the Macarthur River, which it joins at 16° 16’, 136" 12’,

White Stone Creek, Marrakai plains, isolated watercourse crossed by the Woolner track at

12° 33’, 131° 29", Not to be confused with creek of same name in Tortilla Flats area.

Willowra, cittle station on the Lander River at 21° 14’, 132° 38’,

Winmul Swamp, on Wilton River at ce, 13° 36", 134° 18’,

Winnecke Creek, northern Tanami Desert, begins ca. 18° 56°, 130° 02’, flaods out 18° 32’,

131° 39",

Wire Creck, crossed by Adelaide Road 14 miles SSW of Alice Springs at 23° 50’, 133° 49°.

Wollara (Wallara), now a tourist ranch, 24° 38, 132° 17’, Not located by Lidicker &

Marlow (1970 = 222).

Wollogorung, cattle station, 17° 13’, 137° 57’.

Wyclile Creek, crossed by Stuart Highway 11 miles S of Wauchope.

Wyndham, WA, town, 15° 28°, 128° 06’.

Wytookurri (Wylookarri), rockhole at 23° 36’, 129° 46’. Also called Marnpie, which,

according Lo ‘Terry (1937: 139), is a native name for the Common Bronzewing, Phapy

chaleaptera.

Yaringa (specimen of Berrongia lesweur obtained by Finlayson): variously cited as: ‘“Yaringa,

12 miles S.W. of King’s Creek in the George Gill Range” (Finlayson 1958b : 245),

“Yourttga Creek, at the western end of the George Gill Range” (Finlayson 1958b : 249)

and “a water called Yaringa a few miles south of Quadjinta”, the latter being “15 miles

west of Carmichael’s Crag, on the eastern border of the Central Australian Reserve”

(Finlayson 1935b: 91). On the map in Finlayson (1935¢). Quadjinta (Quajinta) and

Yaringa are marked on what is probably Laurie's Creek, the next watercourse W from

King’s Creek. Hf it is Laurie’s Creek then it has been extended much too fur SE.

Yaringa would therefore be W to NW of King’s Creek, not SW.

Yirrkata, mission station on NE coast at 12° 15’, 136° 53’.

Yuendumu, aboriginal settlement, 22° 15’, 131° 48%

56 REC. §. AUST.

Map 1

@ Antechinus bellus

A A. macdonnellensis

VA. bilarni

Map 2

@ Antechinus maculatus

A Planigale ingrami

A Dasycercus cristicauda

Map 3

@ Sminthopsis rufigenis

BS. nifela

A S. froeggatti

S. psammophila

Map 4

A pis crassicaudata

VS. murina ooldea

Map 5

Sminthopsis hirtipes

* Phascogaie calura

@ P. tapoatafa

Map 6

Bi Antechinomys spenceri

A A, laniger (sensu lato)

Map 7

@® PDasyurus hallucatus

W Dz. geoffroii

A Dasyuroides byrnei

Map 8

@ [soodon macrourus

W iJ. auratus

Map 9

BE Macrotis lagotis

* M, leucura

Map 10

A Chaeropus ecaudatus

V Perameles eremiana

@® Petaurus breviceps

Map 11

@ TVrichosurus arnhemensis

AT. vulpecula

W Petropseudes dajili

Map 12

Bettongia lesueur

A specimens

A peripheral localities of

range

® Petrogale brachyotis

Map 13

* Lagorchestes asomatus

L. conspicillatus

MUS.,

16 (11): 1-57

KEY TO MAPS

reported

A Finlayson (1961) and earlier

A 1961 onwards

Map 14

Bi Lagorchestes hirsutus

@ Peradorcas concinna

Map 15

® Onychogalea unguifera

A O. lunata

Map 16

BB Petroeale penicillata lateralis

@ P. p. venustula

A P. longmani

* P. wilkinsi

Map 17

W Notoryctes typhlops

Map 18

V Macropus bernardus

A M. robustus

Map 19

® Macropus antilopinus

WV Megaleia rufa

northernmost records

Map 20

% Pseudomys fieldi

@ P. gracilicaudatus

AP. desertor

Map 21

@ Pseudomys delicaiulus

A P.. hermannsburgensis

Map 22

@ Pseudomys forresti

@ Melomys sp(p)

Notomys amplus

N. cervinus

fuscus

VN. longicaudatus

Map 23

BH Notenys alexis

@ N. aquilo

A Hydromys chrysogaster

* Xeromys myoides

Map 24

Zyzonys

Z. argurus

pedunculatus

woodwardi

4. macrurus

seporillus apicalis

@ %.

Map 25

@ Mesembriomys gouldii

yr Le

Map 26

@ Conilurus penicillatus

Bi Ratius villosissinius

Map 27

WV Rattus colletti

AR. tunneyi

Map 28

V *Pteropus scapulatus

@ P. alecto

Map 29

W Macroderma gigas

@ Macroglossus lagochilus

Map 30

Lf. Taphozous georgianus

BE 2. flaviventris

January,

PARKER—NORTHERN TERRITORY MAMMALS 57

Map 31

@ Tadarida loriae

VT. australis

BB 7. jobensis

A T. planiceps

Map 32

A\ Hipposideros stenotis

W HA. diadema

@ H. ater

Map 33

@ Rhinonicteris aurantius

VY Chalinolobus morio

A C. nigrogriseus

Map 34

A Chalinolobus gouldii

@ Miniopterus schriebersii

Map 35

V *Eptesicus sp(p)

@ Myotis adversus

Map 36

Wi Nycticeius balstoni

Map 37

WV Nyctophilus bifax

@ N. arnhemensis

A N. geoffroyi

RECORDS oF THE x

SOUTH AUSTRALIAN Se

MUSEUM

MORETHIA (LACERTILIA, SCINCIDAE,)

SOUTH AUSTRALIA

By MICHAEL SMYTH

SOUTH AUSTRALIAN MUSEUM

laide

North Terrace, Ade

South Peanalta 5000

VOLUME 16

NUMBER 12

31 August, 1972

MORETHIA (LACERTILIA, SCINCIDAE), SOUTH AUSTRALIA

BY MICHAEL SMYTH

Summary

The genus Morethia Gray, 1845 was formerly part of Ablepharus Lichtenstein, 1823 (Fuhn, 1969).

As more knowledge has accumulated about our local skinks, it has become obvious that the genus

needs revision; the collection of the South Australian Museum contains many specimens under the

name M. lineoocellata (Dumeril and Bibron, 1839) which can in fact be easily distinguished from it.

This paper gives diagnostic characters and, where necessary, descriptions for 4 adelaidensis

(Peters), MZ. boulengeri (Ogilby), M. butleri (Storr), M. lineoocellata and M. taeniopleura (Peters).

I dedicate this paper to the memory of John Mitchell, late Curator of Herpetology in the South

Australian Museum, who was for several years my mentor as I learned herpetology.

THE GENUS MORETHIA (LACERTILIA, SCINCIDAE) IN

SOUTH AUSTRALIA

By MICHAEL SMYTH

Department of Zoology, University of Adelaide, S. Australia 5001

INTRODUCTION

The genus Morethia Gray, 1845 was formerly part of Ablepharus

Lichtenstein, 1823 (Fuhn, 1969). As more knowledge has accumulated

about our local skinks, it has become obvious that the genus needs revision;

the collection of the South Australian Museum contains many specimens

under the name M. lineoocellata (Duméril and Bibron, 1839) which can in

fact be easily distinguished from it. This paper gives diagnostic characters

and, where necessary, descriptions for M. adelaidensis (Peters), M. boulengeri

(Ogilby), M. butleri (Storr), M. lineoocellata and M, taeniopleura (Peters).

I dedicate this paper to the memory of John Mitchell, late Curator of

Herpetology in the South Australian Museum, who was for several years my

mentor as [ learned herpetology.

Genus MORETHIA Gray

Morethia Gray, 1845, Catalogue of the specimens of lizards in the collection

of the British Museum, p. 65.

Type-species: Ablepharus lineoocellatus Duméril and Bibron, 1839.

I have accepted Fuhn’s (1969) distinction of this genus from 4 blepharus

on the basis of skull structure. All members are without moveable eyelids,

the eye being covered by a transparent disc. All are pentadactyl. The two

frontoparictals and the interparietal are fused into a single scale. The

parietals meet in the mid-line.

Morethia adelaidensis (Peters)

Figs. 1, 6

Ablepharus (Morethia) anomalus (adelaidensis) Peters, 1874, Sber. Dt.

Akad. Wiss., Phys.—Math. Klasse, Juni 1874, 376 (formerly Monatsb.

K. Preuss, Akad, Wiss. Berlin).

Ablepharus lineo-ocellatus var. adelaidensis Boulenger, 1887, Catalogue of

the lizards in the British Museum, 3: 349. (Part). “S. Australia.”

Lectotype: Kat. Nr. 4733 in the Museum fiir Naturkunde, Berlin. Adelaide.

Schomburgk. Snout-vent length 5.05 cm. (This is the largest of the

three specimens under this catalogue number. )

Issued 31 August; 1972,

REC. S. AUST. MUS., 16 (12): 1-14 August, 1972

Diagnosis: Five supraciliaries; the third, fourth and fifth all penetrate

between the supraoculars.

Subdigital lamellae acutely unicarinate or tricarinate. Palmar tubercles

elongate, apically acute.

Description: Snout-yent length up to 55 mm. Intact tail 120-165 %

of snout-vent length.

Supranasals present, widely separated, separate from or fused to a

small postnasal. Prefrontals narrowly separated. Frontonasal wider than

long. Frontal longer than wide, in contact with first and second supra-

oculars. Four supraoculars, the second the largest. Frontoparietals and

interparietal fused into a single large scale as wide as Jong, in contact with

the second, third and fourth supraoculars. One pair of nuchals. Seven

Fig. 1. Head of Morerhie acdelaidensis, Wingfield, S. Aust.

labials, the fifth the largest, entirely subocular. Eye entirely surrounded by

granules, Five supraciliaries, the second the smallest, the third, fourth and

fifth about equal in size and all penetrating between the supraoculars

(Fig. 1).

Subdigital lamellae acutely unicarinate or, in some specimens, tri-

carinate. There are 18-24 under the fourth toe, mean 19.5, mode 19.

Palmar tubercles elongate and apically acute. The forelimbs when adpressed

barely reach the eye.

SMYTH—THE SCINCID GENUS MORETIINA IN SAA, 3

Midbody scale rows 26-34, usually 28 or 30; mean 29.4,

Colour in life, grey or olive-grey above; some specimens are browner,

and specimens from Ooldea are distinctly ferrugineous, There are usually

two broad black interrupted dorso-lateral lines from the back of the head

onto the anterior parts of the tail, sometimes continued down the tail as a

single mid-dorsal line. Along cach side is an interrupted and irregular white

line extending from the upper labials, through the ear, above the forelimb,

and along the side to the hindlimb, Above this is a darker band, often

strongly speckled with lighter markings; below is a weaker, speckled dark

band. Ventral surface white. The males in breeding condition develop an

orange colour all around the edges of the ventral surfaces, extending onto

the inside surfages of both fore and hind limbs and being particularly

prominent around the yent and under the anterior part of the tail.

Specimens from the Lake Eyre Basin are distinctly paler, and in some

the dorsal and lateral black bands are barely visible,

Distribution, In South Australia, so far recorded from the arid north-

east, extending south into the coastal habitat along the cdges of the Gults

and west across the Nullarbor Plain into Western Australia (Fig. 6).

Remarks; Peters (1874), in describing M, faeniopleura, compared

it with specimens of what he called Ablepharus (Morethia) anomalis

(adelaidensis). The specimens available to Peters al the time included three

from Adelaide which are clearly M, adelaidensiy as described aboye, It is

from among these that | have designated a lectotype.

Boulenger (1887) distinguished «delaidensis us a variety of M,

linedocellata, and his brief description fits Peters’s specimens. But the

specimens Boulenger lists under this name include, as well as M, adelaidensis,

several bonlengeri as well, The inadequacy of both Peters’s and Boulenger’s

descriptions have compelled me to redescribe M. adelaidensis in detail,

Specimens examined; In the South Australian Museum: RIOIS

Hughes, S.A. (30° 42° S, 129° 31’ BE) (2 specimens); R2585 Price, S.A,

(34° 18’ S, 130° 00° B); R3076 North Tent Hill, S.A, (32° 20” S,

137° 27’ BE), R3186 Yudna Swamp, Moralana Stn. S.A. (31° 32% S,

138° 21’ B) (2 specimens); R3323 Lake Eyre North, Luke Eyre South

(2 specimiens); R3431 Lake Callabonna, S.A, (29° 45’ §, 140° O04 EB);

R3590 Coward Springs, S.A, (29° 24’ §, 136° 49 E); R3836 south end of

Lake Torrens, $.A. (31° 00° 8. 137° 50’ B) (2 specimens); R3842 Kokatha

Hills, SA. (31) 16° S, 135° 15’ BE); R4014 Accalana Crossing, Strzelecki

Ck,, S.A. (29° 14S, 139" 58’ BE); R4990 Lake Coongie. $.A. (27° 11'S,

140° 10’ By: R528) Abracurrie, 30 mi. (54 km) W. Eucla, W.A. (31° 4878S,

128 23° Bit R5283 Ceduna, S.A. (32° 07 S, 133" 40’ EB); R5347

4 REC. $8. AUST. MUS... LG (12): L-l4 August, 1972

Nullarbor HS, S.A. (31° 26’ S, 130° 55’ E); R5864 St. Kilda, S.A.

(34° 45’ S, 138° 31’ E); R5946 “Palmerston, N.T.” (now Darwin, this

locality is probably in error) (16 specimens); R1O882, R1O883 Ooldea,

S.A. (30° 30’ S, 131° 50° E); R11945 Pt. Germein, S.A. (33° OI S,

138° OL’ E); R12013 Edithburgh, $.A. (35° 06’ S, 137° 44’ E); R12689-91,

6 mi. (10 km) N. Pt. Pirie, S.A. (33° 05’ S, 138° OO’ E); R12692-3

Wingfield, S.A. (34° 51’ S, 138" 32’ E); unregistered, “Reuther Collection”

(10 specimens) no locality (probably from Killalpaninna or Kopperamana

Missions, S.A., respectively 28° 36S, 138° 33’ E and 28° 34S, 138° 40’ E).

In the Biitish Museum (Natural History): 64.10.27.9 and 10 “S,

Australia”; 1905.10.31.35 and 36, 100 mi. (161 km) S. Lake Eyre, S.A,

In the Museum fiir Naturkunde, Berlin: Kat. Nr. 4733 Adelaide,

I have also seen specimens from Pt, Gibbon §,A, (33° 47’ S,

136° 47’ B).

Morcethia boulengeri (Ogilby)

Figs. 2. 6

Ablepharus boulengeri Ogilby, 1890, Rec. Aust, Mus., 1: 10-11,

Lectotype: R690 in The Australian Museum, Sydney. Presumably collected

by McCooey at Brawlin, New South Wales (34° 44’ S, 148° 02’ B),

Diagnosis: Six supraciliaries; the first and third are the largest, and the

third, fourth, fifth and sixth are successively smaller.

Subdigital lamellae obtusely unicarinate, palmar tubercles rounded.

This species closely resembles M, butleri, but is distinguishable by its

larger third supraciliary, its much less acutely keeled subdigital lamellae, and

its rounded palmar tubercules,

Description; Snout-vent length in adult males 35-55 mm, in adult

females 40-57 mm. Intact tail 125-165% of snout-vent length.

Supranasals present, widely separated, often fused with the small post-

nasal. Prefrontals usually separated. Frontonasal wider than long. Frontal

longer than broad, contacting the first and second supraoculurs. Four supra-

oculars, the second the largest. Frontopurietals and interparietal fused into

a single large scale about as long as broad, contacting the second, third and

fourth supraoculars. (Ogilby describes a separate interparietal, but all

three syntypes, and all other specimens I have examined, have the inter-

parietal fused with the frontoparietals.) One pair of nuchals.

_ Seven labials (sometimes eight), the fitth the largest, entirely subocular.

Eye surrounded by granules, Supraciliaries six, the first and third the largest,

the first penetrating between the prefrontal and first supraocular, the third

SMYTH—THE SCINCID GENUS MORETINA IN S.A. 5

penetrating between the first and second supraoculars. The fourth, fifth and

sixth supraciliaries do not penetrate between the supraoculars, and decline

regularly in size so that their medial margins form a regular slightly curving

line (Fig. 2), Ear suboval, usually with smaller granules more or less

projecting from the anterior edge.

Five fingers and toes. Lamellae under the fourth toe 15-23, mean 19.5,

mode (9, obtusely unicarinate. Palmar tubercules rounded. The forelimbs

When adpressed reach just beyond the eye.

Midbody scale rows 26-32, mean 29.7, mode 30.

2mm

Rig. 2. Head of Meredida boilengert, Salter Springs, S. Aust.

Colour almost identical to M. butleri, A uniform brown above; most

or all of the dorsal scales have 2-5 (usually 3) fine black lines running along

them, diverging posteriorly; often these lines are expanded and merge into a

black spot or streak, sometimes giving the appearance of interrupted lines of

spots or streaks down the back, but more often distributed irregularly.

A pure white stripe on each side begins on the upper labials and runs

under the eye, through the ear, above the forelimb and along the flunk to

the hindlimb. This white stripe is very prominent, being edged above and

usually below by black stripes. The upper of these black stripes, usually

three scale rows wide, is usually very regular and clearcut on its lower margin,

6 REC. 8. AUST, MUS, 16 (12): 1.14 Aneusi, L972

and runs from the eye to the hindlimb or a little beyond, frequently becoming

less prominent posteriorly, The lower black side-stripe is harrow, irregular

and often interrupted,

Ventral surfaces silver-white. In adult males in breeding condition the

throat becomes bright orange.

The tail of juveniles is pale fawn.

Distribution: Throughout the north of the State and as far south as a

little north of Adelaide and the Murray Mallce south of the River Murray

(Fig. 6). Occurs in the interior of all mainland States.

Remarks: This species is remarkably uniform in colour and morphology

over its range in South Australia, It is sympatric with M. adelaidensiy over

much of its range, with M, taenioplewra in the Northern Territory, and with

M. lineoocellata in several parts of South Australia. It is not yet known to

be sympatric with M, budleri, which it closely resembles,

Specimens examined: In the South Australian Museum: R524

Moolooloo Stn,, S.A. (30° 59° §, 138° 35’ E); R721 “Victorin” (3

specimens); R870 Wynbring, S.A. (30° 33’ S, 133° 32’ BE), RI573

Hermannsburg. N.T. (23° 57’ S, 132° 45’ E) (2 specimens); R2602 (2

specimens), R2648 Mernmerna, S.A. (31° 36’ S, 138° 23’ EB); R3159

“Black Swamp, S.A.” (an unidentifiable locality); R3186 Yudna Swamp,

Moralana Stn., S.A. (31° 32/8, 138° 21’ BE); R3304 Wilpena Pound, S.A.

(31° 30’ S, 138° 37° E) (5 specimens); R3313 > Wonoku Ck. S.A.

(31° 50'S, 138° 24° B); R320 Mt. Aroona, S.A. (30° 34’ 8, 138) 21’ BE);

R3426 6 mi, (10 km) S.W. Wooltana HS., S.A, (30° 25’ §, 139° 25’ E),

R3683 no locality; R3767 Lake Eyre, S.A, (28° 40° 8, 137° 10’ E); R3849

Kondoolka HS., S.A. (32° OL §, 134- 53’ E) (2 specimens); R3938 S.

branch Baleanoona Ck., S.A, (30° 37’ S, 139° 38’ E) (3 specimens);

R401L1 Cordillo Downs HS., S.A. (26° 43’ 8, 140° 38’ E) (5 specimens);

R4012 Innamincka HS,, 8,A. (27° 43’ 8, 140° 45° B)); R4352 Wangoroh

HS., Balranald N.S.W, (34° 38° S, 143° 34’ EB); R4990 Lake Coongie, S.A.

(27° 11° S, 140° 10’ E) (2 specimens); R5484 “Murray Scrub, S.A.”

(5 specimens); R5526-7 Loxton, S.A. (34° 27° S, 140° 34° EB): R5946

“Palmerston, N,T.” (now Darwin; this locality is probably in error); R6008

St. Mary Peak, S.A. (31° 30’ S, 138 33’ E); R10359 Goyders Lagoon,

S.A, (26° 52°’ 8, 139° O00’ E) (2 specimens); R10360 (3 specimens);

R10361 Glengyle HS., Queensland (24° 48’ §, 139° 31’ E); R10936

Paralana Hot Springs, S.A, (30° 12’ 8, 139° 27° BE); RLO962 Yudnamutana

Gorge, S.A, (30° 12’ 8, 139° 17’ EB); R11745 Cradock, S.A. (32° 05% S,

138° 30’ BE); R11936@ North Mulga HS., S.A. (30° 17’ 8S, 139° 32’ BE);

R11942-4 Wertaloona HS., S.A. (30° 38’ S, 139° 21’ E): R11947-51

Bibliando HS,, S.A. (31° 51% S, 139° 07’ B); R12477 Lincoln Gap Stn.,

SM YTH—THE SCINCID GENUS MORETHTA IN S.A. ?

S.A. (32° 37° S, 137° 35’ E); R12677, 8 mi. (13 km) W. of Purnong, S.A.

(34° 52’ S, 139° 32’ E); R12678-9, 4 mi. (7 km) E. of Mantung, S.A.

(34° 36’ S, 140° 03’ E); R12680 Eba, S.A. (34° 04’ S, 139° 36’ E):

R12681-2, 1 mi. (2 km) W. of Telowic, S.A. (33° 02° S, 138° 04° EB);

unregistered, Moorilyanna Well, S.A. (26° 51’ S, 132° 59’ B),

In the Australian Museum, Sydney: R687-9 presumably from Brawlin,

N.S.W. (34° 44’ S, 148° 02’ E).

In the British Museum (Natural History): 74.4.29.1286-8 “Sandhurst,

Victoria”: 90.9,1.4-6 “Brawlin, N.S.W.”.

Fig. 3. Head of Morethia butleri, Western Australia. Drawn in profile; the others

are drawn from slightly above.

Morethia butleri (Storr)

Fig. 3

Ablepharus butleri Storr, 1963, West Aust. Nat., 9: 46-7.

Diagnosis: Supraciliaries form “a straight-sided series of 6 scales, the

first of which is largest” (Storr, 1963) (Fig. 3). Subdigital lamellae

acutely unicarinate, palmar tubercles apically acute.

Remarks: This species was deseribed from the arid eastern part of

Western Australia. Its range probably extends into western South Australia,

and [ have tentatively ascribed a single specimen from Ooldea (S.A. Museum

8 REC. S& AUST. MUS. 16 (12): J-14 August, 1972

R687) to this species. In this specimen the supraciliaries are not as linear

as in the Western Australian specimens, and the line separating them from

the supraoculars is curved rather than straight. As a result the supraciliary

ridge, so prominent in the Western Australian specimens, is absent, and the

supraorbital area is shghtly convex rather than flat,

Morethia lineoocellata (Duméril and Bibron)

Figs. 4, 6

Ablepharus lineo-ocellatus Duméril and Bibron, 1839. Erpétologie général,

5: 817. “Nouvelle-Hollande.”

Cryptoblepharus lineo-ocellatus Gray, 1845, Catalogue of the specimens of

lizards in the collectian of the British Museum, p. 65. “Swan River.”

Morethia anomalus Gray, 1845, Ibid, p. 65. “W. Australia.”

Diagnosis: Six supraciliaries; the fourth is the largest. and the third,

fourth, and sometimes the fifth penetrate between the supraoculars,

Subdigital lamellae obtusely keeled; palmar tubercles apically rounded.

Description: Snout-vent length up to 53 mm. Intact tail 120-160%

of snout-vent length.

The head shields are very similar to those of M7. boulengeri, except for

the supraciliaries. In nearly all South Australian specimens only the third

and fourth penetrate between the supraoculars: the fourth, fifth and sixth are

successively smaller (Fig. 4). But in two specimens from west of Spencer

Gulf, namely from Wynbring and from the Hundred of Nicholls, the fifth

supraciliary is very nearly as large as the fourth and penetrates between the

third and fourth supraoculars; and in a juvenile from St. Francis [sland the

fifth supraciliary, though smaller than the fourth, does penetrate.

Supranasal shields are present in all South Australian specimens,

Subdigital lamellae obtusely keeled; there are 14-22 under the fourth

toe, mean 18.4, mode 19. Palmar tubercles apically rounded.

Midbody scale rows 24-31, mean 27.1, mode 28.

Colour: grey above, The dorsal ocellations which give this species its

name consist each of a single scale, the middle third of which is white and

the two outer thirds black. The ocellation is very variable. On one specimen

from central Eyre Peninsula (R10143) ocellations are quite absent, but on

specimens from the islands off the west coast of Eyre Peninsula they are bold

and numerous, extending from the neck to the end of the tail, and onto the

fore and hindlegs.

SMYTH—THE SCINCID GENUS MORETHIA IN S.A, 9

There is an irregular black stripe along each side, above an irregular

white stripe, sometimes faint, running through the ear, over the forelimb and

back as far as the hindlimb. These lateral stripes are also very variable, but

are never as even or as bold as in M. boulengeri. In some specimens they

are scarcely visible beyond the foreleg. They are best expressed in the most

ocellated individuals.

The male in the breeding season develops a bright orange or orange-

pink throat.

Fig. 4. Head of Morethia lineoocellata, Mallala, S. Aust.

Distribution: This is a southern species, confined to the cooler parts of

the State (Fig. 6) including many of the offshore islands. It also occurs in

Western Australia and Victoria.

Remarks: Gray (1845) described Morethia anomala from “W.

Australia”, distinguishing it from M. lineoocellata because the former, but

not the latter, had supranasal scales. All South Australian specimens have

supranasals, but I have not used the name anomala for them because the

the presence or absence of supranasals is unlikely, by itself, to be a good

specific distinction.

(0 REC, S, AUST. MUS,, 16 (12), 1-14 August, 1972

Most South Australian specimens of M. lineoocellata apparently differ

from many Western Australian specimens in that m the former the filth

supraciliary is smaller than the fourth and does not penetrate between the

supraoculars. Whether this is a good specific distinction, or merely a variant,

I cannot judge on the South Australian material; a careful study of the

material from Western Australia, where both forms are apparently common,

is required,

Meanwhile it should be noted that Loyeridge’s (1934) use of the name

Movrethia lineoocellata anomala tor “the eastern skinks Which are charac-

terized by the almost invariable presence of supranasals and a higher than

average number of mid-body scale rows”, is invalid; he was doubtless

reterring to M. adelaidensis or M, boulengeri, more likely the latter,

Specimens examined: pn the South Australian Museum: R4, R5

between Tanunda and Murray flats, S.A.; R558 Purnong, S.A, (34° 52’ S,

139° 37° E): R870 Wynbring, SA, (30° 33’ S, 133° 32’ BE); R968

Wilkawatt, S.A. (35° 23° 8, 140° 22’ BE): R1i699 Encounter Bay, S.A,

(35° 35° S, 138° 36° E): R2456 Davenport Ck., S.A, (32" 10° S,

(33° 26° BE); R2457 “Palmerston N.T.” (now Darwin; this locality is

probably in error) (35 specimens); R2458 Kangaroo Is., S.A. (5 specimens);

R2473 Flinders Is., S.A. (33° 44S, 134° 31° BE): R2482 St. Francis Is,, $.A.

(32° 30'S, 133° 18 B); R2585 Price, S.A. 34° 18’ S, 138° 00’ E); R3060

North of Buckleboo, S.A. (32° 55° S, 136° 12° EF); R3271 Naracoorte, S.A,

(36° 38’ S, 140° 44’ EB); R8396 Hambidge National Park, S.A, (33° 22’ S,

135° 56’ E); R9005 the southeast of County Chandos, S.A. (35° 16’ S,

140° 47° E); R9262-5 Bascombe Well National Park, S.A. (33° 37° 8,

135° 21’ EB); R9509 Flinders Is,, S.A.; R1O143, R10159 Hd, of Nicholls,

Hincks National Park, S.A. (33° S51’ S, 135° 51’ EB); R10155, RLO167

Hincks National Park, S.A.; R10196, RLO218 Flinders Is., S.A.; R10216,

R10217 Pearson Is., S.A. (33° 57’ S, 134° 16’ EB); R10299, R1O300

Franklin Is.. S.A. (32° 27° S, 133° 39’ E); RIO8&81 Moonlight ‘Tank,

Victoria (35° 45’ S, 141° 23° B): R12444 Streaky Bay, S.A. (32° 48° S,

134° 13’ EB); R1I2610 Big Heath National Park, S.A. (37" 037 §,

140° 33’ S) (5 specimens): R12683 Aldinga Scrub, S.A. (35° 19° 8,

138° 27’ BE); R12684 Spalding Cove, Port Lincoln, S.A, (34° 47° §,

135° 58’ EB) (2 specimens); R12685 Mallala, S.A. (34° 26'S. 138° 31’ B):

R12686 6 mi. (10 km) N.B. Renmark, S.A. (34 09° S, 140° 48’ B);

unregistered, Eyre’s Sand Patch, W.A. (32° 16’ S, 126° 18 B) (3

specimens ) .

In the British Museum (Natural History): 1946,8.15,75 “W Australia”

(syntype of M, anomala Gray),

In the Museum fiir Naturkunde, Berlin: Kat.-Nr. 1355-7. Australia.

SMYTH—THE SCINCID GENUS MORETHIA IN S,A, iN

Morethia taeniopleura (Peters)

Figs. 5, 6

Ablephariis (Morethia) taeniopleurus Peters, 1874, Sber, Dt. Akad. Wiss.,

Phys.-Math. Klasse, Juni 1874, 375-6. (Formerly Monatsb, K. Preuss.

Akad. Wiss. Berlin.) Port Bowen, N.E, Australia.

Diagnosis: Four supraciliaries, the third the largest.

A bold white dorsolateral stripe on each side, from rostral shield to the

fail; suecessively below it a dark band, another white band, and usually a

narrow dark line,

Description: A relatively small member of the genus, obviously flattened

dorsoventrally. Snout-vent length up to 42 mm. Tail 130-160% of snout-

vent length (3 specimens only).

Supranasals present, widely separated, A small postnasal. Frontonasal

wider than lone. Prefrontals well separated. Frontal longer than wide,

contacting the first and second supraoculars, Four supraoculars, the second

2mm

12 REC. S. AUST. MUS, 16 (12): 1-14 Aagast, 1972

the largest, Frontoparietals and parietal fused into a single scale about as

broad as long, with rather concave borders, contacting the second, third and

fourth supruoculars. Parietals meeting in the midline. One pair of nuchals,

almost as large as the parietals.

Seven labials, the fifth the Jargest. Eye surrounded by granules. Four

supraciliaries, the first penetrating between the prefrontal and the first

supraocular, the second penetrating between the first and second supraoculiurs,

the third, which is the largest, penetrating between the second and third

supraoculars, The fourth, which is the smallest, does not penetrate (Fig. 5).

Five fingers and toes, Lamellae under the fourth toe 17-21, mean

19.2. mode 19, acutely unicarinate, Palmar tubercles somewhat clongated.

Midbody scale rows 26-28, mean 26.5.

This species is very distinctively patterned, In old specimens in alcohol

the back is dark brown-black; Peters, however, described it as olive-green,

and Lucas and Frost (1895) give it as greenish-black. In Northern Territory

specimens there Is a faint pale vertebral stripe. In specimen from the Pilbara

in Western Australia, however, this stripe is silvery white, extending from the

middle of the frontal shield onto the tail, Two prominent silvery-white bands

one scale row wide meet across the rostral shield and extend back through

the supraciliaries and dorsolaterally down the body onto the tail. Below

each of these is a black lateral band two scale rows wide, running back from

the rostral to a little behind the hindlimb; it includes the eye and passes above

the ear and the limbs, getting paler as it goes. Below it is another white bund

extending along the upper lip, taking in the ear, running over the forelimb

and along the side. Its lower murgin is bounded by a narrow, irregular,

spotted black line which often does not extend far behind the forelimb, Legs

pale, spotted with black. Tail pale fawn in spirit specimens, but might be

red in life (Lucas and Frost, 1895). Undersurface white,

Distribution: So far not recorded from South Australia, but it

undoubtedly occurs in the Par North of the State. Apparently widespread

in the Northern Territory; Mitchell (1955) records it from Arnhem Land.

Also in northern Western Australia and Queensland.

Remarks: Lucas and Frost (1895) described A. lineoocellatis var.

ruficaudus from Goyder River (25° 45’ S, 134° 30’ E) and Bagot Creek

(24° 40° S, 131° 45’ E), both in the Northern Territory, Later they gave

a coloured illustration of it (Lucas and Frost 1896), These specimens were

clearly M. taeniopleura, On the other hand, Werner's (1910) description

of A. faeniopleurus from around Kalgoorlie in Western Australia is a good

description of M. bufleri. :

SMYTH—THE SCINCID GENUS MORETHIA IN S.A.

Fig. 6.

The known distributions of M. adelaidensis (@), M. boulengeri (Q), and

Specimens examined: In the South

Hermannsburg, N.T. (23° 57’ S,

Australian Museum: R1573

Cockatoo Creek, N.T. (15° 46’ S,

133° 32’ BE) (3 specimens); R1684

129° 08’ BE); R2181 Darwin, N.T.;

R3463 Pilgangoora Well, W.A. (20° 33’ S, 119° 00’ E); R4658 Tambrey

HS., W.A. (21° 38’ S, 117° 36’ E); R4659 Mt. Herbert, W.A. (21° 19’ 8S,

117° 12’ E); R10321 6 mi. (10 km) N. Yuendumu, N.T. (22° 16’ S

131° 49’ B).

14 REC, 8S, AUST. MUS., 16 (12): 1-14 August, 1972

ACKNOWLEDGMENTS

I wish to thank Miss A. G. C. Grandison of the British Museum, Dr. Harold Cogger of

the Australian Museum, Dr. Terry Houston of the South Australian Museum, Dr. Giinther

Peters of the Berlin Museum. and Dr. Glen Storr of the Western Australian Museum, for the

loan of specimens. Keith Newgrain and Beverley Jones drew the profiles and the map.

Glen Storr and Dr, Derek Duckhouse patiently advised me on nomenclature.

REFERENCES

Boulenger, G. A., 1887: Catalogue of the lizards in the British Museum. Second edition.

Vol. 3. British Museum. London.

Fuhn, I. E., 1969: The “polyphyletic” origin of the genus Ablepharus (Reptilia. Scincidae):

a case of parallel evolution, Z. Zool. Syst. Evolutions{., 7: 67-76.

Loveridge, A., 1934: Australian reptiles in the Museum of Comparative Zoology, Cambridge,

Massachusetts. Bull, Mus. Comp. Zool., 77: 243-383.

Lucas, A. H. S. and Frost, C., 1895: Preliminary note of certain new species of lizards from

Central Australia. Proc. R. Soc. Vict,, 7: 264-269.

Lucas, A. H. S. and Frost, C., 1896: Reptilia. In Report on the work of the Horn Scientific

Expedition to Central Australia. Part Il. Zoology. Pp. 112-151. Ed. by

B. Spencer. Melville, Mullen and Slade, Melbourne.

Mitchell, F. J., 1955: Preliminary account of the reptilia and amphibia collected by the

National Geographical Society-Commonwealth Government—Smithsonian Insti-

tution expedition to Arnhem Land (April to November, 1948), Rec. S$. Aust.

Mus., 11: 375-408,

Ogilby, J. D., 1890: Redescription of an Ablepharus from Australia. Rec, Aust. Mus., 1: 1-10.

Peters, W., 1874: Uber einige neue Reptilien (Lacerta, Eremias, Diploglossus, Euprepes,

Lygosoma, Sepsina, Ablepharus, Simotes, Onychocephalus). Sber. Dt. Akad.

Wiss, Phys.—Math, Klasse. Juni 1874.

Storr, G., 1963: Ablepharus butleri, a new scincid lizard from Western Australia. HW. Aust.

Nat., 9: 46-47.

Werner, F., 1910: Reptilia (Geckonidae und Scincidae). Die Fauna Siidwest-Australiens,

“ie 2, part 25. Ed. by W. Michaelsen and R, Hartmeyer. Gustav Fischer,

ena.

REGORDS OF THE

SOUTH AUSTRALIAN

MUSEUM

A NEW SPECIES OF

TROPOSODON BARTHOLOMAI; FROM

THE EARLY PLEISTOCENE KANUNKA

FAUNA, SOUTH AUSTRALIA

(Macropodinae; Marsupialia)

By COLIN R. CAMPBELL

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

VOLUME 16

NUMBER 13

1 February, 1973

A NEW SPECIES OF TROPOSODON BARTHOLOMAIT; FROM THE

EARLY PLEISTOCENE KANUNKA FAUNA, SOUTH AUSTRALIA

(Macropodinae; Marsupialia)

BY COLIN R. CAMPBELL

Summary

The genus 7roposodon was proposed by Bartholomai (1967) for Sthenurus minor Owen, 1877 and

its synonym Halmatrurus vinceus De Vis, 1895 (lectotype QM F3577), the designated holotype

(Sthenurus minor, BM (NH) 48409) being from late Cenozoic deposits in Talbragar Country,

County Bligh, New South Wales. Bartholomai (ibid.) has adequately discussed the taxonomic

history of the genus and it will not be repeated here.

A NEW SPECIES OF TROPOSODON BARTHOLOMAIL, FROM THE

EARLY PLEISTOCENE KANUNKA FAUNA, SOUTH AUSTRALIA

(MACROPODINAE; MARSUPIALIA)

By COLIN R. CAMPBELL

Department of Paleontology, University of California, Berkeley

INTRODUCTION

The genus Treposodon was proposed by Bartholomai (1967) for

Sthenurus minor Owen, 1877 and its synonym Halmaturus vinceus De Vis,

1895 (lectotype QM F3577), the designated holotype (Sthenurus minor,

BM(NH) 48409) being from late Cenozoic deposits in Talbragar Country,

County Bligh, New South Wales. Bartholomai (/hid.) has adequately

discussed the taxonomic history of the genus and it will not be repeated here.

Material from Lake Kanunka (Katipiri Sands, Kanunka Fauna), South

Australia in the collections of the Museum of Paleontology, University of

California, Berkeley includes a complete mandible and some isolated upper

teeth of a new species of Troposodon, herein designated as T. kenti. This

form reveals the previously unknown complete morphology of the lower

incisor and details of the coronoid area.

Specimens of 7. minor available for comparative study included a single

P, (AM F4069), a cast of the holotype (UCMP 55822), and a cast of an

incomplete mandible of BM(NH) 50063a (UCMP 55829). In May 1971 1

was able to examine the extensive collections of T. minor in the Queensland

Museum, and I thank Mr. Alan Bartholomai for his courtesy at that time,

and for his valuable comments on the manuscript of this study.

Thanks are also due to Drs. W. A. Clemens and R. H. Tedford for their

comments on the manuscript. and to Mrs. Pat Lufkin and Mr. Owen Poe for

the figures.

All measurements are in millimetres unless otherwise stated,

Family MACROPODIDAE Owen, 1839

Subfamily MACROPODINAR Thomas, 1888

Genus TROPOSODON Bartholomai, 1967

Type Species Sthenurus minor Owen, 1877

REVISED GENERIC DIAGNOSIS

Moderately large to large: palate entire; upper incisors unknown. P?

subrectangular to subtriangular in basal outline; longer anteroposteriorly

relative to P® than in known species of Protemnodon, longitudinal crest

| Issued 1 February, 1973

2 REC, § AUST, MUS.. 16 (13)> 1-18 Febriary, 1973

slightly labtad of midline, with cusps anteriorly, posteriorly, and postero-

lingually; latter two cusps joined by transverse ridge; lingual basin and

lingual cingulum well developed; broadest posteriorly. dP* molariform, low

crowned, subrectangular in basal outline; swollen laterally at lingual

extremity; forclink and midlink reduced; shelf-like areas behind lophs more

completely developed than in Macropus, Megaleia and Sthenurus. P* with

main crest lubiad of midline, subtriangular approaching subrectangular in

basal outline: well defined lingual basin and cingulum; posterolingual cusp

connected to posterior cusp of main crest by transverse ridge. Eruption of

P* coincides with that of M' in 7. minor, Upper molars with essentially

constant pattern; forelink weak or absent, moderately low to high midlink.

Mandible with moderately deep, elongate syimphysis; prominant genial crest

to below Ps; geniohyal pit shallow to moderately deep: ramus with definitive

groove laterally, from acute diastemal crest to below anterior molars:

pterygoid fossa deep; inferior dental canal extending forward to below Ps;

temporal crest at right angles to horizontal ramus.) small, oval in section,

encircled by enaniel; large, flat, horizontal wear facet dorsally; known

completely only in /. enti, Ps slender, short, subtriangular approaching sub-

rectangular in basal ouvine: weak labial and lingual vertical ridges; main

crest L-shaped in 7, minor; tooth unknown in 7. kenti. dP molariform,

subreetangular, slightly constricted across median valley; shelf-like areas

anterior to lophid crests in 7. minor: tooth unknown in 7. kenti. Ps clongate,

robust, subtriangulur in basal outline: main crest a little lubiad of midline,

L-shaped; lingual base of crown variably swollen: vertical lingual and labial

ridges produce cuspiiles along crest. Lower molars relatively low-crowned,

Subrectangular; anterior cingulum well developed: posterior cingulum well

developed in T. miner, poorly so in T, kenti. Ornamented shelf-like areas

present anterior to lophid crests in| 7. mriner, minimally ornamented tn

T. kemti, Forelink and midlink strong; antero-lubial fossette present in

trigonid basin of 7. minor; weak ridge descends from entoconid to posterior

emegulun in 7. kent, but not in 7. minor,

DISCUSSION

The reason why Owen (1877) failed to recognize Troposedon as a

genus distinct trom Sthenurys was due to his long standing confusion of the

upper dentition of Proftemnedon with that of Sthenurus (Stirton, 1963).

fropesodon appears more closely allied to Protemnoden and those species of

Macropus bearing large premolars than to any other macropodid group

(Tedford, 1966).

[tis here considered probuble that the ancestry of Treposodon was close

to the common mid-late Tertiary origin of Pretemnoden, Sthenurus, and

Wallabia as envisaged by Stirton (1963) and Woodburne (1967), and

possibly represented by the late Miocene Hadronomas Woodburne. 1967, The

CAMPBELIL—A SOUTH AUSTRALIAN FOSSIL MACROPODINE 3

evidence is largely derived from comparison of the premolars, and to some

extent the molars. li P? of Protemnodon there are strong vertical labial and

lingual ridges, a variably developed lingual cingulum terminating opposite

the high anterior cusp and enclosing a lingual basin; longer than any upper

molar and usually wider posteriorly, All these features are found i

Troposodon, and the major structural change required to produce a

Troposodon structure is development of a crest connecting the posterior and

postero-lingual cusps, at right angles to the main crest. Such a change is

already intimated or present in some species of Protemnodon where in Pe

the main crest is slightly inturned postero-lingually or complete.

The upper molars of Protemnodon (and Megaleia) have suggestions of

apical basins above the loph crests, a feature quite characteristic of

Troposodon, The midlink in these two genera is lower and does not turn

so abruptly posteriorly as in Troposedon, and the paracone spur does not

turn sharply lingually but runs up into the median valley directly posteriorly

from the paracone and only near the base of the protoloph does it turn

inward to the bottom of the valley. Thus a basin is not formed on the

posterior face of the protoloph in these genera, but its structural clements are

present. A comparable condition prevails on the metaloph.

Troposodon, with its ornamented brachy-hypsodont molars and broad,

crested premolars, plus the vertical ascending ramus modified basally for the

attachment of heavy jaw musculature, probably occupied a grazing or

grazing-browsing niche. The extension of the inferior dental canal forward

to a point below Py indicates that the mandibles could be precisely manipu-

lated in actions related to cropping with the incisors and cutting with the

premolars (Ride, 1959). The relatively short diastema probably reflects

feeding on heavier vegetation which requires less manipulation in the mouth

to form an ingestible bolus than is the case for soft grasses,

It will be most instructive to discover the nature of the postcranial

skeleton, and that of the cranium and upper incisor dentition in Tropoesodon,

Seemingly, the Pleistocene Katipiri Sands in South Australia offer the best

hopes for such a find at present.

Troposodon kenti sp. nov,’

Hypodigm: The entire known sample.

Holotype; SAM P14507: complete adult left mandible with Li, Ps-Mi.

collected by the late Ruben A, Stirton in 1961. Teeth fully erupted and

well worn. Figs. 1-3,

‘in memory of the four students shot at Kent State University, Ohio, U.S.A, on May 4,

1970, This paper was in preparation at that time and the event served Lo remind that the

freedom of thought so necessary lo human progress, including science, is not yet guaranteed,

and that scientists along with all people should strive to make irrevocable that guarantee,

4 REC. S. AUST. MUS., 16 (13): 1-18 February, 1973

Fig. 1. Troposodon kenti Campbell, n. sp., holotype (SAM P14507), UCMP locality

V5772, Stirton Quarry (Lake Kanunka); Kanunka Fauna, Katipiri Sands. Left mandible

with I); Pg-M4, complete but for tip of coronoid process. Labial view. Scale in cms.

Fig. 2. Tropasodon kenti Campbell, n. sp., holotype (SAM P14507), UCMP locality

V5772, Stirton Quarry (Lake Kanunka); Kanunka Fauna, Katipiri Sands. Left mandible

with T,; Ps-M4, complete but for tip of coronoid process. Lingual view. Scale in cms.

CAMPBELL—A SOUTH AUSTRALIAN FOSSIL MACROPODINE 3

Scns

| a ni,

“

Fig. 3. Troposodon kenti Campbell, n. sp., holotype (SAM P14507), UCMP locality

V5772, Stirlon Quarry (Lake Kanunka); Kanunka Fauna, Katipiri Sands. Left mandible

with [j; Py-My, complete but for tip of coronoid process. Occlusal view. Scale in cms.

Paratypes: UCMP 56898, LdP*; UCMP 56907, RM: or M3, twa LM?

or M*, RM'; UCMP 56920, RP*; UCMP 56921, RM*; UCMP 60805, RM®,

LM"; UCMP 60825, LP*, RM* or M*. Figs. 4 and 5.

AUN

} mee. m

OO aygts

en iT

Fig. 4. Troposodon kenti Campbell, n, sp., paratypes. Top row: RP?,

UCMP locality V5772, No. 56920, Centre row: LdP*, UCMP locality

V5772, No. 56898. Lower row: LP®, UCMP locality V5773. No. 60825,

Labial, lingual, and occlusal views (left to right). Anterior to right.

Seale * 1.5 natural size.

6 REC. S. AUST. MUS., 16 (13); 1-18 February, 1973

Ny See -

2 Weal!”

hy Wf}

[ht .\

Fig. 5, Tiroposodon kenti Campbell, n. sp., paratypes. Top row: RM,

labial view. Centre row: RMt-3, lingual view. Lower row: RM!-*,

occlusal view. RM! (left column) UCMP 56907. LM® (middle column)

UCMP 56907, drawn reversed. RM" (right column) UCMP 56921,

All from UCMP locality V5772. Seale X 1.5 natural size.

Type locality: UCMP locality V5772, Lake Kanunka site | or Stirton

Quarry. A quarry in the basal Katipiri Sands (Stirton, Tedford, and Miller,

1961) at foot of the northern bluffs forming the western side of Lake

Kanunka, eastern Lake Eyre Basin, South Australia.

Age: ?Early Pleistocene, Kanunka Fauna (Stirton, et al., ibid.).

Other localities: UCMP locality V5773 only. This includes a number

of local accumulations of fossil material at base of Katipiri Sands along its

contact with the Etadunna Formation around an isolated hill immediately

east of the type locality.

CAMPBELL—A SOUTH AUSTRALIAN FOSSIL MACROPODINE

Specific diagnosis: Troposodon kenti can be distinguished trom 7. minor

via the following diagnostic characters. Larger dental dimensions; L-shaped

longitudinal crest of Py swollen postero-lingually, conspicuous paraconid

bearing lingually descending ridge. My hypolophid approximately equal in

width to protolophid; accessory ridge from metaconid to forelink weaker than

in 7. minor. Two or more weak vertical ridges between paracone and

metacone on P* ef. two which are low and broad on T. minor, shallow lingual

hasin traversed by anterior low, broad ridge; labial base of crown swollen.

On dP" the anterior cingulum is narrow, midlink weak, transverse valley

V-shaped, lingual stylar cusp absent, apical shelves above loph crests poorly

ornamented, all these in contradistinction to 7. minor. P* broadest medially.

paracone apex close to anterior extremity of longitudinal crest, metacone

higher than hypocone, hypocone rounded, posterior cingulum absent,

posterior fossette absent, lingual cingulum well developed, anterolingual pit

in lingual basin, lingual basin lacking transverse ridges, anterior fossette

present. Accessory links in upper molars paralleling forelink absent, apical

shelves above loph crests minimally ornamented, See Tables 4 and 5.

The most useful of these characters diagnostically are the greater dental

dimensions than 7. fiinor and the apical shelves on the molars being

minimally ornamented. The broader diagnosis offered above should aid in

identification of limited dental samples of 7. kenti.

Description of holotype:

I): exposed portion short, vertically oriented flattened oval in section,

elongate ellipsoid wear facet occupies almost full exposed length dorsally;

enamel extends around circumference except for dentine exposed dorsally on

horizontally flattened wear facet. Figs. 1-3.

P. and dP) unknown,

Py: elongate, robust, longitudinal crest curves sharply lingually at

posterior end of tooth; prominent vertical lingual ridge descends from a

conspicuous cusp at the anterior apex of the crest; behind this three or four

less prominent vertical lingual and labial ridges ure present, each originating

at minor cuspules on crest; postero-lingual expansion reasonably prominent.

Major wear facet an elongate strip labial to and dorsally common with

longitudinal crest, indicating Px occluded with lingual face of P', Tooth

heavily worn occlusally, Fig. 3.

Mi < Mz <M, < Mi: molars subrectangular in basal outline; hypo-

lophid as broad as protolophid in My and My, almost so in Ma, but narrower

in My, Width of trigonid basin (area between protolophid crest and anterior

cingulum) a little less than distance between lophid crests. Forelink

moderately high and strong, descending antero-lingually from the protoconid

8 REC. S. AUST. MUS., Lo Cia): 1-I8 February, V973

to a point a little labial of the longitudinal midline where it turns anteriorly

and runs to the reasonably well developed unterior cingulum, contacting its

approximate midpoint. Forelink occasionally with subsidiary ridging similar

to 7. minor. No antero-labial fossctte in trigonid basin; subsidiary ridge

from forelink near point of anterior flexion runs lingually across anterior

face of protolophid, fading out before reaching metaconid on Ma, probably

reaches metaconid on My where it creates an unornamented shelf-like area

on foreslope of protolophid. Moderately strong midlink descends antero-

lingually from hypoconid to point midway down hypolophid foreslope, there

turning anteriorly to meet very short ridge which vertically descends the mid-

line of protolophid hindslope: accessory ridge from point of midlink flexion

extends lingually in shallow downcurve, fades out before meeting entoconid

on Ms, shorter and abrupty downeurved in My, but creates a small shelf-like

arca on hypolophid foreslope in My and My; My and Ms too worn to reveal

nature of this ridge. Talonid basin broadly U-shaped labially, somewhat

less broadly lingually; small deep pit on basin floor immediately lingual to

midlink on My and My, again ot visible on My and Ms due to wear, No

ornamentation on lophid faces. Posterior cingulum present but less developed

than anterior cingulum, Short, poorly developed ridge descends vertically

fram entoconid to posterior cingulum. Figs. 1-3

TABLE 1 Standard Mandibular Measurements of 7 ynposedon kenti

N.B.—Puarentheses indicate estimated measurements. After Plane (1967) Fig. 5

1, Length of horizontal ramus from incisor alveolus to postalycolar process sper ass 124.0

2. Length of incisor —P, diastem, ... 2... .0.0..0000..-.-. . oo Wer Ee oct. Eon 45.0

3. Depth of ramus below Mz, ttken between roots .... 5s 1 Ta -.. 340

4, Depth of ramus below My, taken between rool& .....0.0., 0-2, OE poms Cees 33.0

5, Length of cheek tooth row, Py—M,. taken at base of crown... 2-0 .. iy ee)

6. Distance from posterior edge of mental foramen ta Py —.--. am ' Otte Ree

7. Height of hypoconid on unworn My, measured from base of enamel .............-.... 5.0

8. Height of condyle aboye alveolar border .....,......0--.-..----.... 1 oe 61.0

9, Meight of ascending ramus above alveolar border... 2. __. SESE INP ESOS EEOC E oT {88.5)

10, Depth of mental foramen below dorsal border of ramus... -.0 0. oot ete! Sa pe 2.5

t1. Width of jaw below My, taken between roots .o 082 8 ce cece ccc cu ueeuues 17.5

Mandible: Figs. 1-3. Table 1. Horizontal ramus deep, relatively

thick; symphysis large, not ankylosed, extremely rugose posteriorly, extending

from anterior extremity of mandible at base of 1, and expanding posteriorly

as u skewed ellipsoid to terminate below the posterior end of Ps; flanged

postero-ventrally into a distinct genial crest below Ps, basal margin acute,

Geniohyal pit slightly indents posterior margin of symphysis: very shallow,

indistinct, Diastema moderately short, about three fourths lengths of cheek

tooth row, slightly convex upward in lateral view, crest sharply rounded;

CAMPBELL—A SOUTH AUSTRALIAN FOSSIL, MACROPODINE 9

mental foramen very close to crest, almost circular, in posterior third of

diastema length, Short eminence present labially at level of mental foramen,

not reaching foramen or anterior extremity of symphyseal region; underlying

shallow trough fades posteriorly into mental foramen. Shallow labial

depression extends from between mental foramen and P. postero-ventrally to

fade out below posterior end of Mz about midway down horizontal ramus.

Digastric Fossa well developed, extending from below transverse yalley of My

near ventral border of horizontal ramus to below posterior moiety of My

where it fades into a very shallow post-digastric sulcus bounded anteriorly by

a very weak digastri¢ process, Shallow depression opens posteriorly into

pterygoid fossa but is not confluent with digastric fossa antero-ventrally, there

being a low, rounded, intervening region of the lingual ramus wall, Post

alveolar shelf short, well developed postalveolar process postero-lingually,

and low postalveolar ridge medially which turns upward to traverse the mestal

wall of the ascending ranius and meet the temporal crest about midway along

its length, above large mandibular foramen. Pterygoid fossa well developed,

deeply excavated labially for insertion of superficial portion of medial

pterygoid muscle; medial border greatly expanded for insertion of deep

portion of medial pterygoid muscle, forming flanged edge of fossa, especially

hear angular process. Condyle well developed, articular surface roughly

elliptical with long axis oriented antero-lingually to postero-labially; extending

labially to level of posterior masseteric eminence. but slightly damaged

lingually. Fovea pterygoidea for lateral pterygoid-temporalis muscle insertion

a triangular pit beneath condyle on mesial face of condyle neck; bounded

untero-dorsally by short sharp crest originating on condyle in region of

mandibular notch, Perpendicular to alveolar margin the temporal crest

ascends to top of slightly damaged coronoid process, posterior slope of

coronoid process a slightly concave curve descending into mandibular notch.

Masseteric crest raised to about level of alveolar margin; masseteric fossa

moderately large, yertically elongated flattened oval in shape with large

masseteric foramen on ventro-mesial wall; leading into short masseteric canal

where deep anterior fibres of the medial masseter and a slip of the deep

masseter insert. Nature and extent of inferior dental canal not visible, but

it probably reaches to beneath the sectorial P; (Ride, 1959; Abbic, 1939),

Angular portion of ascending ramus expands dorsally in posterior view into

u wittt-gluss shape al region of angle.

Description of paratypes

UCMP 56920, P*: Fig, 4. Subtriangular approaching subrectangular

in basal wutline; broader posteriorly than anteriorly; longitudinal crest

concave in side view, straight, Apex of paracone about one quarter distance

ulong crest from anteriormost point: labial and lingual vertical ridges

extremely weak, transect crest, two very small cuspules present but probably

10 REC. S. AUST. MUS., 16 (13): L18 Febraary, (973

more visible in less worn state. Metacone high, at posterior end of crest;

hypecone low, worn, connected to inetacone by low transverse crest which

descends lingual face of metacone, passes through hypocone and turns

anteriorly as a low sinuous lingual cingulum being highest a little anterior

ty the midpoint of the longitudinal crest; cingulum and crest converge

anteriorly but with cingulum expanding lingually and paralleling crest for a

short distance before terminating at u vertical lingual ridge descending from

parucone apex, and matched by a similar labial ridge. Lingual basin

shallow, subtriangular tending subrectangular in shape, traversed anteriorly

by a Jow broad ridge emanating from the highest point of the lingual

cingulum. Lubial base of crown moderately swollen, not quite to extent

that would justify description as a labial cingulum. Posterior ridge from

metacone descends vertically and turns lingually to fade out on posterior

wall of hypocone, No posterior cingulum, but attrition with M! has

produced a wear facet in this location, No posterior fossette.

UCMP 56898, UP: Fig. 4, Molariform, subrectangular in’ basal

outline, unconstricted across median valley, swollen at lingual extremity;

Joplis low, bowed anteriorly, metaloph slightly wider than protoloph,

Anterior cingulum Jow, narrow, short, extending from ridge descending

vertically [rom paracone to anterior base of protocone with slightly developed

forelink meeting it al longitudinal axis of crown. Midlink weak, descending

postero-labially fram apex of paracone then turning posteriorly to meet

metuloph low down and lingual to midline, a little labiad of hypocone apex,

Small shelf-like area with minor ornamentation present posterior to protoloph

crest; short ridge passing posteriorly trom paracone apex descends almost

vertically to floor of V-shaped median valley to meet a similar ridge descend-

ing anteriorly from apex of metacone. Relatively strong ridge descends

postero-labially from apex of hypocone to terminate below apex of metacone.

forming a small posterior cingulum; met by ridge descending almost vertically

from metacone, and by secondary ridge descending from crest of metaloph

labiad of midline, thus delimiting shelf-like area below metaloph crest. No

lingual stylar cusp below protocone in single known specimen,

UCMP 60825, P*; Fig. 4. Subrectangular in basal outline, broadest

point midway ulong length; longitudinal crest almost straight, lower than

apices of paracone and metacone, slightly concave labially; apex of paracone

close to anterior extremity of crown on longitudinal crest: rhree sets of very

weak vertical labjal-lingual ridges transect crest, producing small cuspules.

Metacone highest cusp, near posterior extremity of crown on longitudinal

terest, united to lower rounded hypocone by low posterior transverse ridge:

no definitive posterior cingulum, this area bearing a wear facet produced by

allrition with M': no posterior fossette, Anterior ridge from hypocone

descends sharply and continues anteriorly to form a well developed lingual

CAMPBELL—A SOUTH AUSTRALIAN FOSSLL MACROPODINE MW

cingulum, highest point just posterior to paracone; converges slightly with

longitudinal crest anteriorly to enclose subrectangular, shallow lingual basin

with somewhat sinuous floor profile and bearing small pitlike indentation al

unterd-lingual corner: small antero-lingual fossette anterior to crest connecting

paracone and lingual cingulum. Labial cingulum weakly developed, bearing

two prominent swellings midway between puracone and metacone at base of

crown, poorly defined stylar cusp at anterior extremity of crown on base of

anterior paracone slope. Major wear facet along full length of lingual slope

of longitudinal erest.

Upper rolars, UCMP 60805, LM", RM*, UCMP 56907, two LM® or

M", RM’; UCMP 56921, RM*, Fig, 5, Closely resemble those of T_ minor.

Subrectangular in basal outline, very slightly constricted across median valley;

lophs low, bowed anteriorly; metaloph slightly broader than protoloph i™

M', equal or slightly narrower in M* and M*, M? unknown. Anterior

cingulum low, broad, ascending labially; forelink absent in six teeth known;

labial extremity of cingulum united to paracone by slight vertical ridge

ascending antero-[ubial face of protoloph; two or three faint parallel ridges

occasionally present in labial moiety of anterior cingulum shelf and protoloph

face. Lingual margin of median valley sometimes variably swollen laterally.

Moderately strong ridge descends postero-labially from protocone, turning

posteriorly ucross broadly U-shaped median valley as a moderately low

midimk to unite with extremely short ridge from near midpoint of metaloph,

weaker ridge from paracone descends postero-lingually to turn posteriorly

and closely parallel midlink into floor of median valley, delimiting a shelf-

like area below protoloph crest which is variably but minimally ornamented

by subsidiary ridges and tubercles in M' to M", M' unknown. Strong tidge

curves postero-labially from hypocone to near postero-labial base of crown

where it terminates in a minor swelling of weak posterior cingulum in M’",

or in that position in M' and M®* where posterior cingulum is absent; weaker

ndge descends postero-lingually from metacone towards ridge from hypovone,

delimiting a shelf-like area below metaloph crest which is variably but

minimally ornamented by subsidiary ridges and tubercles in M' to M". Base

of protoloph below protocone usually with variable stylar cusp,

DISCUSSION

The occurrence of Troposoden at Lake Kanunka in the “early

Pleistocene constitutes an important extension of the known geographic

range of the genus. The Katipiri Sands in which the specimens described

were found is composed of channel and floodplain deposits resting dis-

conformably upon the unfossiliferous Tirari Formation at the type locality,

Kalipiri Waterhole on Cooper's Creck, and also at Lake Palankarinna and

Lake Pilikanta where the channels occasionally cut into the top of the

12 REC. S. AUST. MUS., 16 (13): 1-18 February, 1973

Etadunna Formation (?late Oligocene), The deposits at Lake Kanunka

produced Tropasodon, and are of further interest because the fauna contained

is apparently somewhat older than the fauna at the type locality of the

formation, called the Malkuni Fauna. and considered to be late Pleistocene

in age.

The Kanunka fauna contains a diverse assemblage of invertebrates and

vertebrates, including crayfish, lungfish, teleosts, turtles, lizards, crocodiles,

cormorants, ducks, swans, a murid, dasyurids, a thylacoleonid, two vombatids,

potoroine, sthenurine, and macropodine macropodids, and a diprotodontid

smaller than Diprotodon, possibly Euvowenia (Stirton, Tedford and Miller,

1961),

This fauna, yet to be fully studied and described, contains a diversity of

carnivores and herbivores indicative of a situation characterized by areas of

forest and grassland, possibly in the form of savannah, and probably riparian

areas since the existence of watercourses is amply evidenced geologically,

Potoroines, macropodines, and sthenurines are browsing-grazing forms

favouring savannah locales; the murids, dasyurids, and possibly vombutids

indicate the presence of low cover in the form of shrubs or grasses, and if

their present distribution is a faithful guide, a rainfall of 20-30 inches per

annum in a seasonal climate such as prevails over much of the east coast of

Australia today.

As discussed earlier, Troposodon probably occupied a Sthenurus-like

niche, that of a large grazing or grazing-browsing herbivore probably

fuvouring the savannah areas and travelling to water to drink.

STATISTICS

The statistics performed require some discussion. In all cases the

samples involved are too small (7.e., < 10) for reliable conclusions derived

from statistical comparison to be made. Bartholomai (1967) showed that

populations of 7, miner from eastern and western Darling Downs were not

separable on the basis of dental dimensions. His samples were adequate for

Student’s t Test, and the data for his eastern population were chosen for

comparison with the samples here studied (Table 2) because the dental

representation was more complete than in the western population, In

numerous cases in the present study only one specimen was available for

comparison, and although the test applied is perfectly valid (Simpson, Roe

and Lewontin, 1960, pp. 182-3), its limitations due to the lack of variation

considered are self-evident, as are the limitations of Student’s t Test on

samples of Jess than ten.

CAMPBELL—A SOUTH AUSTRALIAN FOSSIL MACROPODINE 13

Thus the statistics are not intended for use as definitive taxonomic tests,

but rather as indications of trends which may be better revealed when larger

samples become available for statistical comparison,

In the case of T. kenti, of twenty tests performed, sixteen involved one

specimen, two involved two specimens, and two involved four specimens. In

twelve of these the values of P (> 0.05 at 5% level of significance) indicate

that the null hypothesis of no difference must be accepted, and that the teeth

tested fall into the dimensional range of teeth from the eastern Darling Downs

sample of 7. minor, In eight cases the values of P (< 0.05 at 5% level)

indicate the null hypothesis must be rejected, and the teeth concerned be

considered as members of a different population (Table 3). The ratio of

acceptances to rejections is taken to indicate that given a larger sample, the

probability for consistently testable significant differences is reasonably high.

This, plus the morphologic differences best represented in Tables 4 and 5

are considered adequate grounds for the designation of new specific rank, in

the purely morphologic sense available to palaeontology. The facts of

geographic and possible temporal isolation of the populations compared are

supporting grounds which cannot be quantified, but are real none-the-less,

TABLE 2 Dental Measurements of Truposedon spp.

Specimen and Species Length Width Width Width

Posteriorly Protoloph Protolophid

Troposodon kenti—

pS UCMP 56920.........

dP? UCMP 56898

P* UCMP 60825,........

UCMP 56907

UCMP 60805

UCMP 56921

UCMP 60825.........

UCMP 56897

UCMP 56897.,....... |

UCMP 56897

UCMP 56907

UCMP 56897

DAB AGRA” REGIS

SoSCeS Vass Ewconua

8.7

10.9

11.0

10.1

10.7, 10.0

11.2

11.4

at es

ROnSH

| AowrwS

cf. Trapesodon minar—

o UCMP 45192

dPs UCMP 45192... _.

P, UCMP 45149

M,; UCMP45192.........

UCMP 45149. ,.....,

M, UCMP 45149

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CAMPBELL—A SOUTH AUSTRALIAN FOSSIL MACROPODINE

TABLE 4 Summary and Comparison of Diagnostic Dental Characters in the lower

Teeth of the Species of Troposodon

Character

Troposodon minor

Troposodon kenti

Longitudinal crest turns abruptly

postero-lingually Crest —L-

shaped i.e. no postero-labial

ridge

Tooth unknown

| Hypolophid much broader than

| protolophid

Tooth unknown

Longitudinal crest turns abruptly

postero-lingually; thus crest

L-shaped. Variably swollen

lingually

Longitudinal crest L-shaped,

Swollen postero-lingually.

Conspicuous paracone with

lingually descending ridge

Lower Molars

Relative width of

lophids

Hypolophid>protolophid M,;

Mo-s;< My

Hypolophid= protolophid M,-s;

almost equal M3;< My,

Forelink ornamen-

tation

Occasionally with subsidiary

ridging

Without subsidiary ridging

Accessory ridge from

metaconid to fore-

Present, moderately strong

Present, weaker than in T.

minor

Apical shelves below

lophid crests

Present, ornamented M,-» with

ridges and tubercles

Present, unornamented

REC. S. AUST. MUS., 16 (13):

1-18 February, 1973

TABLE 5 Summary and Comparison of Diagnostic Dental Characters in the Upper

Teeth of the Species of Troposodon

Character

Treposodon minor

Treposadon kenti

Vertical ridges

Two, low and broad, between

Pparacone and metacone

Two or more, weak

Lingual cingulum . .

Unites with base of crown below

paracone

Low, sinuous, terminates at

vertical lingual ridge des-

cending paracone

Lingual basin

Shallow, unornamented

Shallow, traversed by anterior

low broad ridge

Labial cingulum... .

Absent

Labial base of crown swollen

_ Anterior cingulum. .

Broad

| Midlink

Strong

Narrow

Weak

Transverse valley...

U-shaped

V-shaped

| Lingual stylar cusp..

| Present below protocone

Absent in single known tooth

Apical shelves above

loph crests

Present, variably ornamented

Present, Jess ornamented than

T. minor

Broadest posteriorly

Broadest medially

Paracone apex

4 length along longitudinal crest

from anterior point

Close to anterior extremity of

longitudinal crest

Vertical ridges tran-

secling crest

Up to five sets

Three or four sets

Cusp heights

| Hypocone approximately same

| height as metacone

Metacone highest, hypocone low

and rounded

Posterior cingulum. |

Narrow, short

Absent

Posterior fossette. . .

Present, between posterior

cingulum and ridge connect-

ing hypocone and metacone

Absent

Lingual cingulum, ..

Low

Well developed

Lingual basin

Shallow, traversed by low broad

ridges

Shallow, with antero-lingual pit

Anterior fossette

Absent

Present, antero-lingual to crest

connecting paracone and

lingual cingulum

Labial cingulum ...

“Labial base of crown slightly

swollen

Weak, with two prominent

swellings midway along length

Upper Molars

Representation ... -

M!-* known

M'-* known

Forelink

Reduced or absent, labiad to

crown axis

Absent

Accessory links ....

Slight, parallel forelink in labial

moiety of anicrior cingular

shelf

Absent

CAMPBELL—A SOUTH AUSTRALIAN FOSSIL MACROPODINE 17

SUMMARY AND CONCLUSIONS

A new species of Troposodon Bartholomai, 1967 is described and

named 7. kenti. It is from the ?early Pleistocene Kanunka Fauna at Lake

Kanunka (UCMP localities V5772 and V5773) in the basal Katipiri Sands

of the Lake Eyre Basin, South Australia (Stirton, Tedford and Miller, 1961).

It can be consistently differentiated on morphologic grounds from T. minor

(Owen), 1877, but is represented too poorly numerically for definitive

Statistical differentiation, A large left mandible with Mz-1 (QM F4378)

from the ?late Pliocene Chinchilla Formation and Fauna on Darling Downs,

Queensland, falls into the size range of T. kenti and is morphologically very

similar, It may represent the presence of T. kenfi sympatrically and approxi-

mately contemporaneously with T. minor, but the possibility of its being a

very large specimen of 7. minor cannot be ruled out with the particular

morphology available.

Stratigraphic representation of this genus is too poor to allow a phylo-

genetic sequence to be established. J. minor is present in both the ?late

Pliocene Chinchilla Fauna and the ?early Pleistocene Darling Downs Fauna

on Darling Downs in south-eastern Queensland, a time range overlapping

the ?early Pleistocene provenance of the Kanunka Fauna in South Australia.

The ancestry of Troposodon is considered to have been close to the mid

to late Tertiary divergence of Protemnodon, Sthenurus, and Macropus from

a common ancestor, possibly similar to the ?late Miocene macropodid

Hadronomas Woodburne, 1967 from the Alcoota Fauna (Northern

Territory).

18 REC. S. AUST. MUS., 16 (13): 1-18 February, 1973

LITERATURE CITED

Abbie, A, A., 1939: A masticatory adaptation peculiar to some diprotodont marsupials.

Proc. Zool. Soc. Lond., ser. B, no. 109, pp. 261-279.

Bartholomai, A., 1967: Troposodon, a new genus of fossil Macropodinae (Marsupialia).

Mem. Qd. Mus., 15, pt. 1, pp. 21-33, 3 figs.

De Vis, C. W., 1895: A review of the fossil jaws of the Macropodidae in the Queensland

Museum, Proc. Linn. Soc. N.S.W., 10, pp. 75-133, 5 pls.

Owen, R., 1877: On a new species of Sthenurus, with remarks on the genus Dorcopsis Muller.

Proc. Zool. Soc. Lond. 1877, pp. 352-361, 2 figs.

Plane, M. D., 1967: Stratigraphy and Vertebrate Fauna of the Otibanda Formation, New

Guinea. Bur. Min, Resour. Aust. Bull., 86, 15 figs,, 5 pls.

Ride, W. D. L., 1959: Mastication and taxonomy in the Macropodine skull. in “Function

and Taxonomic Importance”, Ed. A. J, Cain, pp. 33-59, Systematics Associa-

tion; London,

Simpson, G. G., Roe, A., and Lewontin, R. C., 1960: Quantitative Zoology. Rev, Ed.

(New York: Harcourt, Brace and Co.).

Stirton, R. A., 1963: A review of the macropodid genus Protemnodon. Uniy, Calif. Publ.

geol. Sci., 44, pp. 97-162, 15 figs.

Stirton, R. A., Tedford, R. H., and Miller, A. H., 1961: Cenozoic stratigraphy and vertebrate

paleontology of the Tirari Desert, South Australia, Rec. S. Aust, Mus., 14,

No. 1, pp. 19-61, 4 figs.

Tedford, R. H., 1966: A review of the macropodid genus Sthenurus. Univ. Calif, Publ.

geol. Sci., 57, pp. 1-72, 20 figs.

Woodburne, M. O., 1967: The Alcoota fauna, Central Australia. Bur. Min, Resour. Aust.

Bull., 87, pp. 1-187, 34 figs. 7 pls.

RECORDS oF THE

SOUTH AUSTRALIAN (es

MUSEUM ey

RHODACARIDAE (Acari: Mesostigmata)

FROM NEAR ADELAIDE, AUSTRALIA

|. SYSTEMATICS

By DAVID C. LEE

SOUTH AUSTRALIAN Waa

North Terrace, Adelai

South Australia 5000

VOLUME 16

NUMBER 1|4

1 March, 1973

RHODACARIDAE (Acari: Mesostigmata) FROM NEAR ADELAIDE,

AUSTRALIA

BY DAVID C. LEE

Summary

Twenty-five species of Rhodacaridae belonging to 14 genera are listed as collected from the

environs of Adelaide, South Australia. Two new genera (Athiasella and Solugamasus) and 12 new

species (Gamasiphis lenifornicatus, G. saccus, Geogamasus minimus, Gamasellus cophinus, G.

grossi, Acugamasus elachyaspis, Hiniphis bipala, Rhodacaroides minyaspis, Solugamasus mustela,

Antennolaelaps aremenae, A. celox, Onchogamasus virguncula) are described. The description of

adult Gamasellus concinnus (Womersley, 1942) is extended and measurements are given for adults

of all listed species. The larvae of 4 species (Gamasiphoides propinquus, Gamasellus concinnus, G.

cophinus, Acugamasus semipunctatus) are described.

RHODACARIDAE (ACARI: MESOSTIGMATA) FROM NEAR

ADELAIDE, AUSTRALIA

I. SYSTEMATICS

By DAVID C. LEE.

South Australian Museum, Adelaide

ABSTRACT

Twenty-five species of Rhodacaridae belonging to 14 genera are listed

as collected from the environs of Adelaide, South Australia. Two new

genera (Athiasella and Solugamasus) and 12 new species (Gamasiphis

lenifornicatus, G. saccus, Geogamasus minimus, Gamasellus cophinus, G.

grossi, Acugamasus elachyaspis, Hiniphis bipala, Rhodacarvides minyaspis,

Solugamasus mustela, Antennolaelaps aremenae, A. celox, Onchogamasus

virguncula) are described. The description of adult Gamasellus concinnus

(Womersley, 1942) is extended and measurements are given for adults of

all listed species. The larvae of 4 species (Gamasiphoides propinguus,

Gamasellus concinnus, G. cophinus, Acugamasus semipunctatus) are

described,

INTRODUCTION

The Rhodacaridae is a group of mainly free-living, ground inhabiting,

predatory mites, most common and diverse in form in Southern Temperate

regions.

The present study on Systematics formed part of a thesis for the degree

of M.Sc.. University of Adelaide, on rhodacarids from the environs of

Adelaide, South Australia. Further papers will deal with the Ecology

(part IT) and Behaviour (part IIL) of these mites.

The rhodacarids studied were from 4 sites between the summit of

Mount Lofty and the coast-line of the Adelaide Plain. The “Sites”, to be

described more fully in part Il, are as follows:—

(lL) Summit. Approximately 18 km from the sea. Near the summit

of Mount Lofty (715 m), the highest hill overlooking the

Adelaide Plain.

(2) Foothills. Approximately 16 km from the sea, Near the First

Waterfall, at the head of the deepest stretch of Waterfall Gully

which has recently eroded back from where First Creek flows

from the foothills below Mount Lofty on to the Adelaide Plain.

1 Issued 1 March, |973

z REC, §. AUST, MUS., 16 (14): 1-36 Murclr, 1973

(3) Plains. Approximately & km from the sea. Heywood Park

Unley; a suburban park on the Adelaide Plain.

(4) Coastal. Approximately 1.5 km from the sea. Grange Golt

Course.

The terminology and diagnoses of taxa are as used by Lee (1970)

except when otherwise indicated. Measurements are in micrometres, to the

nearest 5 for Jengths and 2.5 for breadths. The idiosomal length given is

the average followed in parentheses by the number of specimens measured

and the range of their lengths. Other measurements given for a specimen

near to the average length are the lengths and breadths of the movable

cheliceral digit (ch), the palp (pa) and the legs (/, //, /// or IV).

Specimens selected for description have been registered in the collection

of the South Australian Museum,

Family RHODACARIDAE Oudemans, 1902

Subfamily RHODACARINAE Oudemans, 1902

Genus RHODACARUS Oudemans, 1902

Rhodacarus roseus Oudemans

Rhodacarus roseus Oudemans, 1902, p. 50.

FEMALE. Not figured.

Measurements: idiosomal length—275 (1 from Summit Site) or 490

(3 from Plains Site, 450-510, measured for appendage lengths and genu

breadths); appendage lengths—ch 100, pa 140, J 405, 1] 285, III 245,

/V 360; genu breadths—pa 22.5, J 25, I 32.5, Il 27.5, IV 32.5. The

female from the Summit Site differs from previously described specimens

from the Plains Site in being smaller with very indistinct punctations on the

idiosoma and an opisthonotal seta 23 of similar length to seta 72 rather than

to Z5.

MALE. Not figured.

LOCAL. Summit Site—temale (197061), moss, 26.4.1968. Plains

3 females (N196879-N196881).

Site

REMARKS. Specimens of R. reseus from the Plains Site have already

been described (Lee, 1970, p. 29). One smaller female, differing from

these as indicated above, was collected from the Summit Site and is for the

time being regarded as belonging to the same species. This species has also

been recorded from the Palaearctic region.

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 3

Genus RHODACARELLUS Willmann, 1935

Rhodacarellus silesiacus Willmann

Rhodacarellus silesiacus Willmann, 1936, p. 282.

FEMALE. Not figured.

Measurements: idiosomal length—305 (1); appendage lengths—ch

30, pa 90, I 230, 17 140, 7 115, /V 200; genu breadths—pa 17.5, / 22.5,

11 27.5, I 22.3, 1V 25. The general appearance is as given for this species

by Sheals (1958), but at the posterior end of the idiosoma the setal lengths

and markings on the shields conform to those figured for this species by

Athias-Henriot (1961, figs. 283, 284 and 291).

MALE. Not collected.

LOCAL. Foothills Site—temale (N197062), moss, 9.5.1968.

REMARKS. The only previous record of Rhodacarellus occurring in

Australia is of a female (N1968280) from Millicent, about 400 km south

of Adelaide (Lee, 1970), which I have since identified as R. silesiacus. This

species has also been recorded from the Nearctic and Palaearctic regions.

Subfamily GAMASIPHINAE Lee, 1970

Genus GAMASIPHIS Berlese, 1904

Gamasiphis australicus Womersley

Gamasiphis (Heteroiphis) australicus Womersley, 1956a, p. 521.

FEMALE. Not figured.

Measurements: idiosomal length 390 (3 from Foothills Site, 360-430);

appendage lengths—ch 40, pa 100, J 290, // 250, I/I 190, 1V 265; genu

breadths—pa 17.5, 1 22.5, MW 30, WI 22,5, IV 22.5.

MALE. Not figured,

LOCAL. Foothills Site—3 females (N197063-N197065), moss or

plant litter, 7.1968-2.1969. Plains Site.

REMARKS. G. australicus from the Plains Site is described (Lee,

1970, p. 50) and appears to be identical with specimens from the Foothills

Site. The only other record of this species is of the type which was collected

at Mylor, about 8 km south of the Summit Site.

“

4 REC. S. AUST. MUS,, 16 (14): 1-36 March. 1973

Gamasiphis fornicatus Lee

Gamasiphis fornicatus Lee, 1970, p. 51,

FEMALE. Not figured.

Measurements: idiosomal length—SOO (20, 480-510); appendage

lengths—ch 72, pa 165, 7 360, /7 310, I 305, IV 385; genu breadths—

pa 20, 7 25, 11 42.5, IM 30, IV 30.

MALE. Not figured,

Measurements: idiosomal length—490 (6, 470-490),

LOCAL. Summit Site—10 females (N197066-N197075) and 3 males

(N197076-N197078), moss or plant litter, 1968-1969. Foothills Site—

14 females (N197079-N197092) and 4 males (N197093-N197096), moss

or plant litter, 1968-1969,

REMARKS. G. fornicatus is also recorded from Mount Remarkable,

about 260 km north of the Summit Site. No differences were noticed

between the type specimens and those from near Adelaide.

Gamasiphis lenifornicatus Lee, n.sp.

FEMALE, Fig. 1; A,B,C,D,E.

Measurements: idiosomal length—390 (1); appendage lengths—ch

40. pa 135, I 305, I 225, IM 220, IV 305; genu breadths—pa 17.75,

1 22.25, M 30, IM 25, IV 25. One pair of pre-endopodal shields. Split

between exopodal and peritrematal shields does not extend posterior to

stigma, and the peritrematal shield is broadly fused to the notal shield. No

conspicuous lateral fissure on notal shield. Leg chaetotaxy is abnormal for

rhodacarids (not as Gamasellus) in lacking 3 setae on leg ['V; seta p/ on the

genu (2, 5/2, 0); setae pd 2 and p/ 2 on the tibia (2, 4/2, 1). The tectum

is conspicuously fimbriated. The idiosoma is acutely convex dorsally and

the dorsal setae are unusually long. Sternal setae are nearly in a straight line

so that a line joining setae sf2, sf3 and st4 enclose an angle of more than 95°;

excepting the atypical Hydrogamasus this is a unique attribute within the

Gamasiphinae. On legs II-I'V, tarsal setae adl and pdl are subequal in

length to pretarsus and bear a broad hyaline flap.

MALE. Fig. 1; F,G.H.

Measurements: idiosomal length—350 (6, 340-370). The distribution

of shields is as for the female except on the venter of the podosoma. The

spermadactyl is slightly longer than the movable cheliceral digit, and has a

hyaline, spatulate tip. On leg II, setae av on the femur and genu are modified

to spurs,

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 5

Fig. 1. Gamasiphis lenifornicatus o.sp.

g Pp

A-E, female: A, soma, dorsum; B, idiosoma, venter; C, pretarsi | and IV;

D, gnathosoma, venter; E, leg TV (part), dorsal setae.

F-H, male: F, leg Il (part), antero-lateral; G, chelicera: H, idiosoma, venter.

6 REC. 8. AUST. MUS., 16 (14): 1-36 March, 1973

| /

Fig. 2. Gamasiphis saccus n.sp.

A-E, female: A, soma, dorsum; B, gnathosoma. venter; C, idiasoma, venter; D,

pretarsi | and IV; E, leg IV (part), dorsal setae.

F-H, male: F, chelicera; G, idiosoma, venter; H, leg IL (part), antero-lateral.

LEE—RHODACARIDAE (ACARL: MESOSTIGMATA) IN S.A, 7

LOCAL. Summit Site—holotype female (N197097) and allotype male

(N197098), plant litter, 24.4.1969; and 5 paratype males (N197099-

N1970103), moss or plant litter, 6-8.1968.

REMARKS. The idiosoma of G. lenifornicatus is globular, but to a

lesser degree to that of G. fornicatus. The leg chaetotaxy is unique amongst

rhodacarids in lacking seta pd2 on the adult tibia [V. The male tibia II is

unusual within the genus (G. saccus n.sp. is the only other species of

Gamasiphis with this attribute) in having a setose seta av.

Gamasiphis saccus Lee, n.sp.

Female. Fig. 2; A,B,C.D,E.

Measurements: idiosomal length—350 (3, 350-360); appendage

lengths—ch 40, pa 120, 7 270, JP 210, WU 190, IV 240; genu breadths—

pa 15, / 22.5, UW 32.5, HI 25, 1V 25. One pair of pre-endopodal shields,

Split between exopodal and peritrematal shields extends backward from

stigma to divide exopodal IV, Lateral fissure runs almost parallel to edge

of opisthonotal shield. Leg chaetotaxy is normal for rhodacarids (as

Gamasellus). The idiosoma is unusually flattened dorsally for a species of

Gamasiphis. There is a conspicuous pit on the anterior edge of the ventro-

anal shield bordering acetabulum IV. This pit appears to be lined by a fine

pile. On legs I-IV, tarsal setae ad! and pdl are simple and less than a

quarter of the Jength of the pretarsus.

MALE. Fig. 2; F.G,H.

Measurements: idiosomal length—340 (3, 330-350). The distribution

of shields is as for the female except on the venter of the podosoma. Sperma-

dactyl is slightly longer than the movable cheliceral digit and, although

blunter, similar in shape. On leg II, setae av on the femur and genu are

modified to spurs.

LOCAL. Summit Site. Foothills Site—holotype female (N1970104).

allotype male (N1970105), 2 paratype females (N1970106 and N1970107)

and 2 paratype males (N1970108 and N1970109), moss, 21.6,.1968.

REMARKS. G. saccus is easily recognized by the large pit on the

posterior edge of acetabulum IY. It is also unique amongst species of

Gamasiphis in having the same leg chaetotaxy as Gamasellus, Specimens

from the Summit Site are indistinguishable from those described,

8 REC. S, AUST. MUS., 16 (14): 1-36 March, 1973

Genus EUEPICRIUS Womersley, 1942

Euepicrius filamentosus Womersley

Euepicrius filamentosus Womersley, 1942, p. 170.

FEMALE. Not figured.

Measurements: idiosomal length—S0O0 (10, 470-550); appendage

lengths—ch 60, pa 140, 1 755, I] 445, IM 440, 1V 490; genu breadths—

pa 22.5, ] distal 25, 1] 45, IT 42.5, IV 47.5.

MALE. Not figured.

Measurements: idiosomal length—490 (10, 460-530).

LOCAL, Summit Site. Foothills Site—10 females (N1970110-

N1970119) and 10 males (N1970120-N1970129), moss or plant litter,

1968-1969,

REMARKS, The above specimens of E. filamentosus are indistinguish-

able from the type specimens (N1970130-N1970139) which are from Glen

Osmond and Long Gully (2 localities within 8 km of the Foothills Site).

A female of an undescribed species from Waimamaku, New Zealand, was

incorrectly listed as belonging to E. filamentosus in the original description.

Genus GAMASIPHOIDES Womersley, |956a

Gamasiphoides propinquus Womersley

Gamasiphis (Gamasiphoides) propinqua' Womersley. 1956a, p. 528.

FEMALE, Not figured.

Measurements: idiosomal length—800 (5. 780-840, measured for

appendage lengths and genu breadths) or 600 (2); appendage lengths—

ch 80, pa 220, 1 570, 11 430, [11 420, [TV 550; genu breadths—pa 35, 1 47.5,

H 60, TT 50, 1 55.

MALE. Not figured,

Measurements; idiosomal length-——730 (4, 720-730) or 540 (1).

LARVA. Fig. 3; A.B,C,D,E,F.

Measurements: idiosomal length—320 (4, 270-350). Seta al on palp

genu is spatulate. Tectum anterior margin is basically trispinate with

numerous spinules, and the three spines are of approximately equal size.

tdiosomal shields clearly defined. Idiosomal setae are simple. Opisthonotal

seta Z3 (Z1 and Z2 are absent) is about half as long as seta 24,

(Mr. R. Domrow of Queensland has pointed out (private correspondence: 30,.9.1970) that

according to the International Code of Zoological Nomenclature, Art. 30 (a) (ii), aides

is masculine and therefore the original ending of this species name should be changed.

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S,A. 9

100

rh ft

ae S

bg pa 4 I

pM al . : a

a choo 4 | et”

hy f I 4 i ) |

4 a i YS Y

Fig. 3. Gamasiphoides propinquus (Womersley )

A-F, larva: A, soma, dorsum; B, leg HI (part), dorsal setae; C, idiosoma, venter;

D, pretarsus T; E, chelicera; F, palp femur and genu, venter.

LOCAL. Summit Site—5 normal-sized females (N1970145,

N1970146, N1970150-N1970152), 4 normal-sized males (N1970147,

N1970148, N1970153, N1970154) and 4 larvae (N1970159-N1970162),

plant litter, 5.1968-1.1969; 2 small females (N1970142 and N1970143)

and I small male (N1970144), plant litter, 2-4.1969.

REMARKS. Most of the above specimens are indistinguishable from

the types of G. gamasiphoides, collected from Belair National Park about

8 km south of the Summit Site, and from specimens described by Lee (1970,

10 REC. S. AUST. MUS., 16 (14): 1-36 Mareh, 1973

p. 63), collected higher up Mount Lofty than the Summit Site. Specimens

only distinguishable by their smaller size were collected at the end of the

summer dry season, a period during which normal-sized adults have not, as

yet, been found.

Subfamily OLOGAMASINAE Ryke, 1962

Tribe OLOGAMASINI

Genus ATHIASELLA Lee, n.gn.

Heydeniella Richters, dentata-complex: Lee, 1970, p. LOL.

Type-species: Hydrogamasus dentatus Womersley, 1942.

DIAGNOSIS. Small to large mites. Holonotal shield. Ventro-anal

shield of female discrete, while on male it is usually fused to the notal shield

and always fused to peritrematal and expodal IV shields. Sterno-metasternal

shield of female never fused to endopodal IV shield. Single pair of

pre-endopodal shields. ‘Twenty pairs of podonotal setae. Leg chaetotaxy

may be normal for rhodacarids (as Gamasel/us) but usually there is one less

ventral seta on genu [V (2, 5/1, 1). On palp genu, seta a/! pectinate with

at least 12 lateral prongs and seta a/2 broadly cuneate in distal half. Dorsal

setae all simple, tapering, Spermathecal access duct opens distally on dorsal

surface of trochanter Il]. Spermadactyl is never conspicuously longer than

the movable cheliceral digit and lies parallel to it. The male corniculus is

slightly attenuated and on a raised base. On the male leg II at least setae av

on the femur, genu and tibia are modified to spurs. Pretarsus I is pedunculate

and conspicuously smaller than other pretarsi. Legs I and IV are long

(0.90 or more of idiosomal length) and there is considerable variation in

leg thickness (using breadth of genu: I is 0.65 or less of Il; IIE is 0.85 or

less of TY).

REMARKS. Nominate species of Athiasella were grouped (Lee, 1970)

in the dentata-complex of Heydeniella. These species are however distin-

guishable from species of Heydeniella by the recognizable location of the

orifice to the spermathecal access duct, the discrete female ventro-anal

shield and the modification of seta av on the male tibia [I] to a spur.

Athiasella dentata (Womersley)

Aydrogamasus dentatiis Womersley, 1942, p. 149.

Heydeniella dentata (Womersley): Lee, 1970, p. 105,

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. tl

FEMALE. Not figured.

Measurements: idiosomal length—740 (14, 720-780); appendage

lengths—ch 125, pa 270, | 750, Hf 570, 11 520, IV 790; genu breadthis—

pa 35, 1 42.5, If 82.5, 1/11 47.5, IV 60.

MALE. Not figured.

LOCAL, Summit Site. Foothills Site

N1970176), plant litter, 27.3.1969. Plains Site.

14 females (N19701653-

REMARKS. All stages of 4. dentata from the Foothills Site have been

described (Lee, 1970), and except for slight variations in their size are

indistinguishable from specimens from the other sites. So far, specimens with

a long “tooth” on trochanter ['V (includes types) have only been collected on

the Adelaide Plain or in the Mount Lofty Ranges. A form with a shorter

“tooth” is found in the Flinders Ranges and the Hummock Ranges at

localities 120-280 km north-west of the Foothills Site (Lee, 1970).

Athiasella relata (Womersley)

Hydrogamasus relatus Womersley, 1942, p. 151, not Hydrogamasus relatus:

Womersley, 1956a, p. 530.

FEMALE. Not figured.

Measurements: idiosomal length—540 (8, 510-570); appendage

lengths—ch 95, pa 205, J 540, II 390, IT 345, IV 510; genu breadths—

pa 27.5, 1 30, 1 50, HT 32.5, IV 42.5.

MALE. Not figured.

Measurements: idiosomal length—540 (9, 520-550).

LOCAL. Summit Site and Foothills Site—8 females (N1970181-

N1970188) and 9 males (N1970189-N1970197), moss or plant litter,

4-9,.1968. Plains Site.

REMARKS. 4. relata is similar to A. dentata, but smaller and the

female lacks a conspicuous “tooth” on trochanter [V, while the male has a

straight, needle-like spermadacty] (Womersley, 1942, fig. 6A). A. relata

has only been previously recorded from Glen Osmond which is close to the

Foothills Site. Specimens of an undescribed species from Taringa, Queens-

land, were incorrectly listed as belonging to A. relata by Womersley (1956a).

12 REC, S. AUST. MUS.. 16 (14): 1-36 March, 1973

Genus GEOGAMASUS Lee. 1970

Geogamasus howardi Lee

Geogamasus howardi Lee, 1970, p. 96.

FEMALE. Not figured.

Measurements: idiosomal Jength—330 (10, 310-340); appendage

lengths—ceh 40, pa 125, 1 300, I] 220, IH 180, /¥ 295; genu breadths—

pa 15,17 17.5, 77 33, HT 22.5, 1V 25,

MALE, Not figured.

Measurements: idiosomal length—310 (10, 300-320).

LOCAL. Summit Site—10 females (N1970198-N1970207) and 10

males (N1970208-N1970217), plant litter, 9.1968. Foothills Site.

REMARKS. The above specimens are slightly larger than the types

from Mount Burr, about 400 km south of the Summit Site, which is the only

other locality record of the species.

Geogamasus minimus Lee, n.sp.

FEMALE. Fig. 4; C.

Measurements: idiosomal length—260 (10, 250-270); appendage

lengths—ch 25, pa 95, 1 245, I/ 170, Il 140, 1V 200; genu breadths—

pa 12.5, 17 15, 11 27.5, JH 17.5, 1¥ 22.5. Similar to Geogamasus howardi

but two characters are conspicuously different. There is a broad strip of

striated cuticle between the ventro-anal shield and the exopodal IV shield,

which is subequal to the distance between the genital seta (st5) and the

posterior margin of the genital shield. On genu IV there is no seta py.

MALE. Fig. 4; B,D.

Measurements: idiosomal length—240 (10, 220-250). Similar to

G. howardi but three male characters are conspicuously different. The

spermadactyl has no elaborate hinge, although it can coil up. On trochanter

Il, seta a/ is enlarged, bent into an “L” shape and spatulate, but the distal

limb is less than twice the length of the proximal limb. On tarsus IT, the

ridge that is regarded as homologous to seta ad3 borders a conspicuous

ventro-lateral depression.

LOCAL. Summit Site—holotype female (N1970218), allotype male

(N1970219), 11 paratype females (N1970220-N1970230) and 11 paratype

males (N1970231-N1970241), moss or plant litter, 1968-1969, Foothills

Site.

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 13

REMARKS. G. minimus is smaller, and more like the only other

nominal species of Geogamasus from Australia—G. howardi—than the South

American species. G, minimus is unique in Geogamasus in haying fewer leg

setae than Gamasellus. Specimens from the Foothills Site are indistinguish-

able from those described.

Fig. 4. Gamasellus grossi n.sp. and Geogamasus minimus n.sp.

A, G. grossi, female, soma, dorsum.

B-D, G. minimus: B, male leg I] (part), antero-lateral; C. female idiosoma, ventral;

D, male chelicera.

14 REC. S, AUST. MUS., [6 (14): 1-36 Mareh, 1973

Fig. 5. Gamasellus concinnay (Womersley)

A-EF, female: A, soma, dorsum; B, gnathosoma, yenter; C, idiosoma, venter; D,

tarsus LV, dorsal setae; E, region of acetabulum TV, showing spermathecal access

tube,

F-H, male: F, idiosoma, venter; G. leg I] (part), antero-lateral: H, chelicera.

LEE-—RHODACARIDAE (ACARI, MESOSLIGMATA) IN S.A, 15

Tribe GAMASELLINI Hirschmann, 1962

Genus GAMASELLUS Berlese, 1892

FALCIGER-complex

Gamasellus concinnus (Womcersicy)

"Digamasellus concina Womersley, 1942, p. 159.

Digamasellus concinna Womersley: Womersley, 1956a, p, 537,

Cyrtolaelaps concinnus (Womersley): Womersley, 1961, p. 194.

FEMALE. Fig. 5; A,B,C,D,E.

Measurements: idiosomal length—S00 (10, 490-510): appendage

lengths—ch 105, pa 180, 1 550, 77 400, 1/7 340, 7V 475; genu breadths—

pa 25, 1 27.5, HW 52.5, WI 37.5, IV 47.5, Three pairs of pre-endopodal

shields; the anterior and posterior pairs are slim and inconspicuous, the

former so much so that it was not noted in the original description. The

peritrematal shield is separate from the ventro-anal shield; this agrees with

original description but not with my examination of the holotype which has

a narrow strip of shield connecting the posterior point of the peritrematal

shield to the ventro-anal shield. There is a reduction in the extent of the

Opisthosomal shields so that the dorsal setae in row R and ventral seta Zy3

are on striated cuticle. Podonotal seta zl is setose and inconspicuous, and

not similar to /1 as originally described. Seta a/l on the palp genu has 6

luteral prongs. The dorsal setae on the tarsi are either setose or slightly

lanceolate in contrast to most such setae on other Australian species in the

Gamasellus faleiger-complex which are conspicuously spatulate.

MALE. Fig. 5; F,G,H.

Measurements: idiosomal Jength-—470 (10, 450-480). Posterior pair

of pre-endopodal shields is as large as the central pair. The ventro-anal

shield is fused to exopodal IV shield and is larger than on the female since

it carries 2 more pairs of setae (Zv3 and Syl). On leg I, setae av or av2

on the femur, genu, tibia and tarsus are modified to spurs; seta pv on the

femur and genu are spine-like; and there are non-setous spurs on the genu

and tibia.

LARVA, Fig. 7; A,B,C\D,E.F.

Measureinents: idiosomal length—230 (10, 210-250). Seta al on

palp genu is spine-like with 6 lateral prongs. Tectum anterior margin is

basically trispinate with spinules, and central spine is longer than lateral

spines, Sternal shield not clearly defined. Idiosomal setae are simple or.

if long, faintly pilose. Opisthonotal seta 73 (41 and Z2 are absent) 1s

subequal to seta Z4. There are 12 or more teeth on fixed cheliceral digit.

16 REC, S. AUST. MUS., 16 (14): 1-36 March, 1973

O—p— ;

ul—o—_o-__

100

A,C.G

B.D.EFH .

Fig. 6. Garmasellus cophinus n.sp.

A, soma, dorsum; B, gnathosoma, venter: C, idiosoma, venter: D. leg

; region of

A-F, female:

1V (femur-tibia), antero-lateral; E, tarsus IV, dorsal setae; F.

acetabulum LV, showing spermathecal access tube.

G-I, male: G, idiosoma, venter; H, leg I (part), antero-lateral; I, chelicera.

LEE RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A, \7

LOCAL. Summit Site—10 females (N1970244-N 1970253), 10 males

(NL970254-N[970263), larva containing protonymph (N1970264) und 16

other larvae (N1970265-N1970280), moss or phint litter, 1968 and 1969.

Foothills Site.

REMARKS. The above specimens ure indistinguishable from the

holotype female (N1970242) and allotype male (N1970243) collected from

Relair National Park (the only other locality record of this species) about

§ km south of the Summit Site, except that the peritrematal shield on the

female is separate from the ventro-anal shield, The unequal leg proportions

of G. concinnus resemble those of Athiasella rather than of characteristic

species of Gamasellus (¢.g., G. tragardhi) in which the legs are subequal in

size. The larva is easily distinguished from the rhodacarid larvae collected

near Adelaide by its large number of cheliceral teeth.

CGamasellus cophinus Lee, f,.sp,

FEMALE. Fig. 6; A.B,C,D,E,F,

Meusurements: idiosomal length—310 (10, 290-310); appendage

lengths—eh 45, pa 90, 7 265, IE 180, HH 160, 7 220; genu breadths—

pu iS, 7 20, 1 27.5, HE 20, 1¥ 27.5, The distribution of ventral idiosomal

shields is similar to other members of the faleiger-complex, but less extensive

in that the peritrematal shield is slim, and the ventro-anal shield does not

extend laterally to merge with the metapodal shield nor anteriorly to carry

seta Jvl. The chaetotaxy is as for the faleiger-complex except on the

opisthosoma where the homologies are not clear (possibly the extra seta in

row Sv is homologous with the seta previously labelled UR|—see Lee, 1970,

fiz, 281). Seta a/l on the palp genu has 6 lateral prongs as in Gamasellis

concinays, but the prongs are proportionately larger. The idiosoma is dorso-

ventrally flattened and sub-rectungular in outline. At the posterior end of

the opisthosoma there is a ridge on both the dorsal and ventral shields. The

ventral ridge is the most heavily sclerotized and convoluted. G. cophinus is

unique within the genus in having only setose dorsal setae on the adult with

no recognizable adhesive exudate on the notum, Leg IV ts large and has

a number of stout, spine-like ventral setae.

MALE. Fig. 6; G,H,I.

Measurements: idiosomal length—280 (10, 270-290). The ventro-

anal shield extends further forward than in the female so that it carries seta

Jv1, but there is still a substantial strip of striated cuticle between it and the

ventral shields on the podosoma, The spermadactyl! lies close to the movable

cheliceral digit, but distally to the single tooth they ure separated, On leg II,

setae ay on the femur, genu, tibia, seta av2 on the tarsus, seta pvl on the

femur and seta pv on the genu ure enlarged into spurs or spines.

18 REC. S. AUST. MUS., 16 (14): 1-36 March, 1973

DEF

C.G,H

AB,

1000p

Fig. 7. Gamasellus larvae

A-F, G. concinnus Womersley:

(part), dorsal setae; D, pretarsus I; E, palp femur and genu, venter; F, chelicera.

G-K, G. cophinus n.sp.:

dorsum; J, pretarsus I; K, palp femur and genu venter.

A, soma, dorsum; B, idiosoma, venter; C, leg IV

G, soma, dorsum; H, idiosoma, venter; I, leg IV (part),

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 19

LARVA. Fig. 7; G,H,LJ.K.

Measurements: idiosomal length—210 (1). Seta a/ on palp genu is

spine-like with 6 lateral prongs. Tectum anterior margin is basically

(rispinate with numerous spinules, and the central spine is longer than lateral

spines although the spines are not so well developed as in Gamiayelluy

conemnnus. Idiosomal shields not clearly defined. Idiosomal setae are

simple. QOpisthonotal seta Z3 (71 and Z2 are absent) is about half as long

as seta 44. There are 4 or 5 teeth on fixed cheliceral digit.

LOCAL, Summit Site—holotype female (N1970281), allotype male

(N1970282), 14 paratype females (N1970283-N1970296) and 9 paratype

males (N1970297-N1970305 ), moss, 7.6.1968-12.9.1968. The morphotype

larva. (NL970306) was bred from adults from Summit Site, moss, 9.8.1968.

Foothills Site,

REMARKS. G. cophinus is an atypical species of Gamuasellus: | group

it in the falciger-complex because of the distribution of idiosomal stiields

(especially the 3 pairs of pre-endopodal shields). the chaetotaxy, the location

of a conspicuous spermathecal access duct, the spine-like seta av2 on the

male tarsus If and the pronged seta a/l on the palp genu which is similar to

the homologous seta on G. concinnus. Attributes dissimilar to those of other

species of Gamasellus are regarded as having evolved relatively recently.

being adaptive to living in the narrow pore spaces of the more mineral soil

layers. Such attributes are the dorso-ventral flattening of the idiosoma, with

no fusion between the podosomal and opisthosomal shields in the male as

well as the female. the short, simple idiosomal setae and the large, spiny leg

IV. | regard the resemblance of G. cophinus to the type of species of

Khodacaroides as superficial. Specimens from the Foothills Site are indis-

linguishable from those described.

Gamasellus grossi Lee, n.sp.

FEMALE. Fig. 4; A,

Measurements: idiosomal Jength—550 (2s, 540-560); appendage

lengths—ch 55, pa 130, 1 365, IT 310, 7 295, IV 420; genu breadths—

pa 25, 142.5, 150, 11 40, 1¥ 42.5. On the notum only 3 pairs of podonotal

setae (/1, 25 and r2) and 2 pairs of opisthonotal seta (75 and S4) are pilose

and spatulate, standing out at right angles to the cuticle.

MALE, Not figured.

Meastirements: idiosomal length—520 (3, 510-530),

LOCAL, Coastal Site—holotype female (N1970307), allotype male

(N1970308), paratype female (N1I970309) and 2 paratype males

(N1970310 and N1970311), moss, 10.6.1965,

20 REC. §. AUST. MUS., 16 (14): 1-36 March, 1973

REMARKS: G. grossi is very similar to two other species of Gamasellus

trom South Australia: G. tragardhi and G. cooperi. G. grossi can be

distinguished from these species by the number of pilose and spatulate dorsal

setae. Otherwise it is so similar to G. tragardhi (see Lee, 1970, p. 135) that

a complete description has not been given. G, grossi has also been collected

from Eyre Peninsula, South Australia—2 females (N19715 and N19716)

and 2 males (N19717 and N19718), moss, Mount Wedge, col. G. F. Gross,

26.9.1964,

Gamasellus tragardhi (Womersley)

Digamasellus tragardhi Womersley, 1942, p. 161.

Cyrtolaelaps tragardhi (Womersley): Womersley, 1961, p. 194.

Gamasellus tragardhi (Womersley): Lee, 1970, p. 135.

FEMALE. Not figured.

Measurements: idiosomal length—640 (10, 620-670); appendage

lengths—ch 60, pa 150, / 390, 17 345, I 320, IV 450; genu breadths—

pa 25,17 52.5, 11 60, 11 47.5, IV 50.

MALE, Not figured.

Measurements: idiosomal length—620 (10, 560-660).

LOCAL. Summit Site—10 females (N1970312-N1970321) and 10

males (N1970322-N1970331), plant litter, 16.8,1968. Foothills Site.

REMARKS. G. tragardhi is also recorded from Adelaide, Bridgewater

(about 6 km southeast of the Summit Site) and near Wilmington (about

280 km north of Summit Site). The specimens from the Summit and Foot-

hills Sites are indistinguishable from the types.

Genus ACUGAMASUS Lee, 1970

PUNCTATUS-complex

Acugamasus punctatus (Womersley)

?Digamasellus punctatus Womersley, 1942, p. 160.

Cyrtolaelaps punctatus (Womersley): Womersley, 1961, p. 194.

FEMALE. Not figured.

Measurements: idiosomal length—605 (4, 560-630); appendage

lengths—ch 105, pa 180, 1 550, JJ 425, IM 385, IV 500; genu breadths—

pa 27.5, 1 42.5, 11 55, 11 45, IV 47.5.

!hE—RHODACARIDAE (ACARI! MESOSTIGMATA) DN S.A, 21

MALE. Not figured.

Measurements: idiosomal length—S80 (5, 560-600),

LOCAL. Summit Site—4 temales (N1970332-N1970335) and 5

males (N1970336-N1970340), plant litter, 10.4.1969.

REMARKS. A. punctatus is also recorded from Adelaide und the

Belair National Park about 8 km south of the Summit Site. The specimens

from the Summit Site are indistinguishable from the holotype female

(N1970341) and the allotype male (N1970342).

Acugamasus elachyaspis Lee, n.sp.

FEMALE. Fig. 8; A,B,C,D,E.

Measurements: idiosomal length—470 (3, 460-480); appendage

lengths—ch 85, pa 170, 1 505, 1 350, HT 305, TV 400; genu breadths—

pa 25,1 30, 1 37.5, 1 30, TV 32.5. The idiosomal shields are distributed

as on other females of the punctatus-complex, but they are the least extensive,

for example the opisthonotal shield is so narrow that it only curries setal row

J and 4 and one seta from row §. The chaetotaxy only differs from other

Species in the punctatis-complex in having 4 setae in row W/R. None of the

setae on the idiosoma and legs are spatulate and only a few are pilose, No

spermathecal access duct is visible (the structure on the adaxial edge of

acetabulum [V—fig. 8C—is the genital apodeme and attached muscles ).

MALE. Fig. 8: F,G,H.

Measurements: idiosomal length—430 (2, 420 and 440), ‘The

distribution of idiosomal shields is as on the female except on the venter of

the podosoma. This contrasts with other Australian members of the

punctatus-complex in which the males have a section of the podonotal shield,

carrying at least setae r2 and r4, that is split away posteriorly from the rest

of the shield and fused to the peritremuatal shield. Seta s@5 is on striated

cuticle. The spermadactyl is short and twists under the movable cheliceral

digit so its spatulate tip lies close to the adaxial surface of the digit. On leg

IT, setae ay on the femur, genu and tibia are enlarged mto spurs while setac

pyvl on the femur and py on the genu are spine-like.

LOCAL, Coastal Site—holotype female (N1970343). allotype male

(N1970344), 2 paratype females (N1970345 and N1970346) and | para-

type male (N1970347), moss, col.: M. Fagg, 23.5.1965.

REMARKS. 4, elachyaspis is the smallest species in the punctatis-

complex, has the simplest setae and Is the most sparsely covered by shields,

March, 1973

a ey

Keo =

ee ee i>, Semel

— ~~ } / fet

ae ae el ere Le

EFM

A.CG

100}1

Q— 5 j ~O-

venter:

idiosoma,

1-36

MUS., 16 (14):

REC, §, AUST,

B, pretarsi | and IV: C,

1V (part), dorsal setae,

F, Jeg IL (part), antero-lateral; G. idiosoma, venter; H. chelicera.

Acugamasus elachyaspis n.sp.

E, leg

Fig. 8.

A, soma, dorsum;

gnathosoma, venter:

A-E, female:

F-H, male:

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 23

Acugamasus semipunctatus (Womersley)

?Digamasellus semipunctatus Womersley, 1942, p. 163.

Digamasellus semipunctatus Womersley: Womersley, 1956a, fig. 20.

FEMALE. Not figured.

Measurements: idiosomal length—725 (4, 700-760); appendage

lengths—ch 135, pa 240, J 710, 17 480, 177 450, IV 620; genu breadths—

pa 30, 155, 1 65, 111 52.5, IV 55. Opisthonotal shield is reticulated and

the podonotal shield is rugose, but neither shield has the raised punctations

that are characteristic of the notal shields of A. punctatus.

MALE. Not figured.

Measurements: idiosomal length—670 (4, 650-690). On leg II, setae

ay on the femur, genu and tibia are modified to spurs, while seta py1 on the

femur and seta pv on the genu are spine-like. There is a non-setous spur on

the antero-lateral surface of genu II.

bal

| bd \

| |

| /

7 ae

—

}

a /

—~. /

Pail ~

aon

x P

ye, ¢ 2AM

— \ \

Fig. 9. Acugamasus semipunctatus (Womersley) larva

A. soma, dorsum; B, idiosoma, venter; C, pretarsus I; D, leg IIL (part), dorsal setae:

EF. chelicera: F, palp femur and genu, venter.

24 REC. S, AUST, MUS,, 16 (14): 1-36 March, 1973

LARVA. Fig. 9; A.B,C.D.E.F.

Measurements: idiosomal length—340 (3, 320-360). Seta al on palp

genu is lanceolate with one basal prong as in the adult. Tectum anterior

margin is basically trispinate with numerous spinules, and central spine is

longer than Jateral spines. Idiosomal shields clearly defined. [diosomal

setae simple. Opisthonotal seta 73 (71 and 72 are absent) subequal in

lensth to seta Z4.

LOCAL. Summit Site—larva (N1970397), litter, 24.4.1969 and larva

(N1970398) bred from adults from this Site. Foothills Site—4 females

(N1970350-N1970353) and 4 males (N1970354-N1970357), moss,

24.5.1968, and larva (N1970399) containing protonymph, moss, 5.8.1968.

REMARKS. 4. semipunctatus is the largest species in the punctatus-

complex. Its name is misleading, since it does not bear any notal punctations

similar to those on A. punctatus. Womersley (1942) describes these puncta-

tions on the podonotal shield, but has drawn this shield as being rugose. The

females from the Summit and Foothills Sites are indistinguishable from the

holotype female (N1970348), moss, Bridgewater (about 6 km south-east of

the Summit Site). A male (N1970349) labelled “allotype”, moss, Muston,

Kangaroo Island (about 65 km south-west of the Summit Site and separated

from the mainland by 11 km of sea) is probably the specimen drawn by

Womersley (1956a) with no text description. This male differs from those

from near Adelaide in not having a non-sctous spur on genu II, but is

otherwise indistinguishable.

Genus HINIPHIS Lee, 1970

Hiniphis bipala n.sp.

FEMALE. Fig. 10; A,B.C.D.

Measurements: idiosomal length—300 (3, 300); appendage Jengths—

ch 35, pa 100, | 205, 17 185, H/F 140, 1 190; genu breadths—pa 15, J 20,

127.5, HE A7.5, 1¥ 17,5. The notal shield is divided in two. The meta-

sternal shield is fused to the endopodal IV shield. The dorsal and ventral

shields are widely separated by striated cuticle except for the narrow anterior

fusion of the peritrematal and podonotal shields. Exopodal III shield is not

split. The idiosomal chaetotaxy is as for Hiniphis hinnus except that there

are 3 setae in row Sv, Leg chaetotaxy is normal for rhodacarids (as

LEFE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 25

Syeda

Fig. 10. Hiniphis bipala n.sp..

A-D, female: A, soma, dorsum; B, tarsi | and 1V. dorsal setae; C, idiosoma, venter:

D, gnathosoma, venter,

E-G, male: E, idiosoma, venter; F, leg Il (part), antero-lateral; G, chelicera.

26 REC. S. AUST. MUS., 16 (14): 1-36 March, 1973

Gamasellus). On the palp genu, seta a/1 is spine-like with about 4 pairs of

short, fine lateral prongs. On the palp femur there is a conspicuous non-

setous tubercle near the ventro-distal edge. There is a pair of conspicuous

pits near the anterior margin of the opisthonotal shield. No recognizable

spermathecal access duct.

MALE. Fig. 10; E,F,G.

Measurements; idiosomal length—290 (2, 280-300). Idiosoma is

encased in a single continuous shield with a dorsal split, except for the discrete

pre-endopodal and anterior exopodal shields. The spermadactyl lies close

to the movable cheliceral digit but distally to the single tooth they are

separated. There is a tubercle on the palp femur similar to that of the female.

On femur IT, seta av is enlarged to a lumpy tubercle and seta pvl is modified

to a small, globular tubercle. Seta av on tibia I ts spine-like, but seta av on

genu I] is only very slightly stouter than the other setae.

LOCAL. Summit Site—holotype female (N1970358), allotype mule

(N1970359), 2 paratype females (N1970360 and N1970361) and one

paratype male (N1970362), plant litter, 5 or 12,8.1968.

REMARKS. H. bipala is the only species allotted to Hiniphis other

than the type (H. hinnus). The males of these two species are similar, but

the distribution of idiosomal shields on the females differ in that the dorsal

Shields are extensively fused to the ventral shields of H. hinnus. On the other

hand females of both species have two attributes (fusion of the metasternal

and endopodal IV shields, and separate podonotal and opisthonotal shields)

not found together on other females of Ologamasinae, although they occur

together in Evepicrius (Gamasiphinae) and Onchogamasts virguncula

(Sessiluncinie ).

Genus RHODACAROIDES Willmann, 1959

Rhodacaroides minyaspis Lee, n.sp.

FEMALE. Fig. 11; A.B,C.D,

Measurements: idiosomal length—300 (5, 300-310); appendage

lengths—ceh 80, pa 115, J 310, I 250, I 220, IV 280; genu breadths—

pa 25, 125, 11 27.5, TH 25, 1 32.5. The extent of the idiosomal shields is

unusually reduced; only part of setal row J and Z are on the opisthonotal

shield; the peritrematal shield hardly exists; there 1s no ventral shield so that

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 27

setae in row Jv and Zy are on striated cuticle. The dorsal chaetotaxy of the

idiosoma is reduced, with only 12 pairs of setae on the opisthonotum. Leg

chaetotaxy is normal for rhodacarids (as Gamasellus). On the palp genu,

seta all is spine-like with 4 prongs on one side and 2 prongs on the other

side. On the palp femur there is a small, non-setous tubercle on the mid-

ventral surface. No spermathecal access duct is visible. Pretarsus I is absent.

700. IW

Fig. 11. Rhodacaroides minyaspix n.sp., female

A, soma, dorsum; B, gnathosoma, venter; C, tarsi | and IV, dorsal setae; D, idiosoma,

venter,

MALE. Not known,

LOCAL, Summit Site—holotype female (N1970363) and 4 paratype

females (N1970364-N1970367), plant litter, 12.8.1968 (holotype) or 10

or 24.4.1969.

ae REC. 8, AUST, MUS... 16 (14): 1-36 Mareli, 1973

REMARKS. R. minyaspis is unique amongst rhodacarids in having

only an anal shield on the venter of the opisthosoma. Because I have no

males of this species it is tentatively grouped in Rhedacarvides. There are

3 nominal species in the genus, und R. minyaspis is more similar to R. costet

from South America than to the type (R, aegyptiacus) from Egypt.

Genus SOLUGAMASUS Lee, n.gn,

Type-species: Solugamasus mustela Lee, n.sp.

DIAGNOSIS. Small mites, Separate podonotal afd opisthonotal

shields. Ventro-anal shield discrete in both sexes, Sterno-metasternal shield

of female never fused to endopodal IV shield. Two pairs of pre-endopodal

shields (in series rather than parallel), Twenty-two pairs of podonotal setae.

Leg chaetotaxy is normal for rhodacarids (us Gamasellus), On palp genu,

seta a/l has two pairs of lateral prongs and seta al2 is lanceolate. Dorsal

setae all simple, tapering. Location of spermathecal access duct is unknown,

The spermadactyl is strongly recurved. On the male leg II at least setae aw

on the femur, genu and tibia are modified to spurs. Pretarsus | sometimes

absent. Legs Land IV are long (0.9 or more of idiosomal length) and there

is medium variation in leg thickness) using breadth of genu: [ is approxi-

mately 0.8 of Il: TIT is approximately 0,75 of TV).

REMARKS, Solugamasus is similar to Rhodacaroides, but 1 recognize

it as a distinct genus because of the strongly recurved spermadactyl in the

male. A recurved spermadactyl occurs in widely differing rhodacarid genera

such as Rhodacarus, Pyriphis and Sessiluncus but its occurrence never varies

within a genus. The female of the only nominal species in Solugamasus is

cusily distinguished from the 3 nominal species of Rhodacarnides by, among

other attributes, its unusually short idiosomal setae. On the other hand, I do

do not specify any female attributes as distinguishing these two genera,

Solugamasus mustela Lee, n.sp.

PEMALE. Fig. 12; A.B,C,D,

Measurements: idiosomal length—290 (7, 270-300); appendage

lengths—ch 50, pa 90, 7 280, // 240, HI 200, 7¥ 290; genu breadths—

pa 17.5, 120, 11 25, 11 20, 1 27.5. The extent of the idiosomal shields ts

reduced so that setae 73, r4, RI and Zv1 are on striated cuticle. The opistho-

notal chactotaxy is reduced to 12 puirs of setae. On the palp genu, seta a/]

has two pairs of lateral prongs very near the tip, On the palp trochanter,

seta av is spine-like and set on a tubercle. The idiosomal setae are simple

and very short. Pretarsus | is absent,

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 29

_0———0-—=

o=5 O50

————

Fig. 12. Solugamasus mustela n.sp..

A-D, female: A, soma, dorsum; B, tarsi I and IV, dorsal setae; C, idiosoma, venter:

D, gnathosoma, venter.

E-G, male: E, idiosoma, venter; F, leg Il (part), antero-lateral; G, chelicera.

30 REC, §, AUST, MUS., 16 (14): 1-36 Mareh, 1973

MALE. Fig. 12; E,F,G.

Measurements: idiosomal length—270 (3, 270-280). The distribution

of idiosomal shields is similar to the female, but the anterior shoulders of the

ventro-anal shield extend forward to carry seta Zvl. The spermadactyl is

strongly recurved and there is a small dorsal process on the fixed digit. On

leg I, seta av on the femur, genu and tibia are enlarged into spurs while

some other ventral setae are long and spine-like.

LOCAL. Foothills Site—holotype female (N1970368), allotype male

(N1970369), 6 paratype females (N1970370-N1970375) and 2 paratype

males (N1970376 and N1970377). moss and plant litter, 9.5.1968-

30.1.1969.

REMARKS. See remarks on genus.

Subfamily SESSILUNCINAE Lee, 1970

Genus ANTENNOLAELAPS Womersley, 1956b

Antennolaelaps aremenae Lee, n.sp.

FEMALE. Fig, 13; A,B,C,D.E.

Measurements: idiosomal length—745 (3, 720-760); appendage

lengths—ch 75, pa 210, 1 870, 1 570, HI 525, IV 730; genu breadths—

pa 32.5, 1 55, JI 75, HI 52.5, IV 55. Horizontal outline of idiosoma is

nearly parallel-sided. The pre-endopodal shields lie very close to the sterno-

metasternal shield but are probably separate from it. The opisthonotal

chaetotaxy is reduced to 15 pairs of setae (as Antennolaelaps testudo). Leg

chaetotaxy is normal for rhodacarids (as Gamasellus). On the palp genu,

seta all has 4 pairs of lateral prongs, and seta a/2 is lanceolate. On the

gnathosoma, the fourth hyposternal seta is pilose. The idiosomal setae are

all simple and some podonotal setae are very small. Pretarsus I is

pedunculate.

MALE. Fig. 13; F,G,H.

Measurements: idiosomal length—690 (4, 670-710). Ventro-anal

shield is not fused to the sternito-genital or exopodal [V shield; on the other

hand it is more extensive than in the female so that posteriorly there is hardly

any striated cuticle between it and the notal shield. The tectum bears an

extra pair of spinules on the central spine that are not present on the females,

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A. 44

Fig. 13, Antennolaelaps aremenae n.sp..

A-F, female: A, soma, dorsum; B, pretarsi | and IV: C, leg ITV (part), dorsal setae;

D, idiosoma, venter; E, gnathosoma, yenter.

P-H, male; F, idiosoma, venter; G, chelicera; H, Jeg IL (part). antero-lateral.

32 REC. S. AUST, MUS., 16 (14): 1-36 Mareh, 1973

The spermadactyl is of similar shape to the movable cheliceral digit except

that the tip is almost spatulate. On leg II, setae av on the femur and genu

are enlarged into spurs, while seta ay on the tibia is spine-like.

LOCAL. Summit Site—allotype male (N1970380), 2 paratype

females (N1970381 and N1970382) and 3 paratype males N1970383-

N1970385), moss or plant litter, 9.5.1968-12.8.1968. Foothills Site—

holotype female (N1970379), moss, 24.5.1968.

REMARKS, 4. aremenae is the largest species in the genus and its

idiosoma is nearly-parallel sided, as for A. celox, in contrast to the 3 species

described from Queensland which are suboval or subcircular in horizontal

outline. There are many attributes which distinguish this species from the 3

previously described species (see Lee, 1970), amongst which is the variable

size of the podonotal setae with both very small and averaged sized setae in

rows j and z. A, aremenae is unusual amongst species of Sessiluncinae in

having sexual dimorphism in the size of the ventro-anal shield.

Antennolaelaps celox Lee, n.sp,

FEMALE. Fig. 14; A.

Measurements: idiosomal length—S45 (5, 530-560); appendage

lengths—ch 55, pa 160, I 600, // 460, 1/1 400, JV 580; genu breadths—

pa 25,1 40, 11 50, 111 37.5, 1¥ 40. A. celox is similar to A. aremenae. The

following attributes differ: it is smaller; the fourth hypostomal seta is not

pilose; there is a semicircular ridge around sternal pore 2 and seta st2 (as on

male see fig. 14B); there are 18 pairs of opisthonotal setae; the longer

opisthonotal setae are pilose and more of the dorsal setae are very small,

including some on the opisthonotum.

MALE. Fig. 14; B and D.

Measurements: idiosomal length 505 (3, 500-510). Ventro-anal shield

is of similar size to that of the female so that posteriorly there 1s a conspicuous

strip of striated cuticle between it and the dorsal shield. On leg Il, seta py

on the genu is pilose and of a similar length to seta py on the tibia.

DEUTONYMPH. Fig. 14; C.

Measurements: idiosomal length—415 (2, 410-420). The majority

of dorsal setae are nearly as long as the distance between their setal bases in

contrast to their small size in the adult.

LEE—RHODACARIDAE (ACARI: MESOSTIGMATA) IN S.A.

6, |

vista

\ 4

ae |

o whe |

shoe, |

i Ay

| t i |

(ome) fi yg é

\} LG J

Ne oval H i

5Z5y

Fig. 14.) Antennolaelaps celox n.sp..

A, female, soma, dorsum; B, male, idiosoma, venter; C, deutonymph, soma, dorsum:

D, male, leg Hl, antero-lateral.

LOCAL. Summit Site—holotype female (N1970386), allotype male

(N1970387), 4 paratype females (N1970388-N1970391), 2 paratype males

(N1970392 and N1970393), one morphotype deutonymph (N1970394)

and one paratype deutonymph (N1970395), moss or plant litter, 9.5.1968-

4.7.1968. Foothills Site.

REMARKS. A. celox is similar to A. aremenae, but smaller and more

dorsal setae are very small. Specimens from the Foothills Site are indis-

tinguishable from those described,

34 REC. S. AUST. MUS., 16 (14): 1-36 March, 1973

Genus ONCHOGAMASUS Womersley, 1956b

Onchogamasus virguncula Lee, n.sp.

FEMALE. Fig. 15; A,B,C,D,E,

Measurements: idiosomal length—310 (1): appendage lengths—

ch 40, pa 110, 1 275, W200, I 175, IV 255; genu breadths—pa 15, J 20,

HT 30, HT 17.5, IV 25. Separate podonotal and opisthonotal shield, with

opposing edges touching. Three pairs of pre-endopodal shields. The sterno-

metasternal shield is fused to endopodal IV shield. Ventro-anal shield is

discrete, but its anterior edge lies very close to the ventral podosomal shields.

Exopodal IT and III shields are split. The peritrematal shield is free

posteriorly. Idiosomal chaetotaxy—6j, 62, 5s, Sr: Sst: 3Jv, 3Zv, 2Sy,

3 anal, Leg chaetotaxy is abnormal for rhodacarids (not as Gamasellus) in

lacking seta pd4 on tarsus 1V. Movable cheliceral digit has at least 5 teeth.

On palp genu, seta a/l pilose with 5 pairs of lateral prongs and seta al2 1s

spine-like. Dorsal setae simple and tapering. On the sternum, a line joining

setae sf2, st3 and st4 would enclose an angle of less than 95°. Pretarsus I

present but not pedunculate. Femur IY with 2 non-setous tubercles on

ventral surface.

Fig, 15. Onchegamasus virgunculu o.sp., female

A, soma, dorsum; B, gnathosoma, venter: C, tarsi | and 1V; D, idiosoma, venter:

EB, femur IV.

LER—RHODACARIDAF (ACARI: MESOSTIGMATA) IN SA 35

MALE. Not known.

LOCAL. Summit Site—holotype female (N1970396), plant litter,

5.8, 1908,

REMARKS. Amongst species of Sessiluncinae. O. virgunela is unique

in having 3 pairs of pre-endopodal shields, and the only other species with a

divided dorsal shield belongs to Paragamasellevans. The chaetotaxy is

unique amongst rhodacarids; combining the absence of seta pd4 on tarsus TV

(as Gamasellopsis) with an otherwise normal leg chaetotaxy (as Gamasellus ).

Because of the position of the female sternal setae (which would be diagnostic

of communis-complex) and the absence of fusion between the peritrematal

and exopodal shields (diagnostic of pumilio-complex ), | revoke my previous

concept (Lee, 1970, p. 189) of Onchogamasus including two species-

complexes. Furthermore, the attributes of O. virguneula lessen the gap

between the attributes of Onchogumasus and those of Gamasellopsis and

Gamasitus, so that f males of the types of these 3 genera prove to be similar

the possibly synonomy of these names should be considered.

ACKNOWLEDGMENTS

lan indebted lo Dr. K. EB. Lee for permitting me to use the core sample dessivaling

funnels pl the Soil Zoology Section of the C.S.1.R.O. Soils Division, Glen Osmond, in order

to collect some of the mites described here. | wish to acknowledge the helpful advice and

efilicism received from Dr. D, A, Duckhouse, University of Adelaide. during the preparation

of the M.Se, thesis which formed the basis of this study.

[am most grateful lo Mrs, Curol Aitken for preparing some drawings (Figs. 1, 2, 3, 4,

7,89, 13 and }4) and to Mrs. Brenda Head for preparing the other drawings (Figs. 5, 6

WO, 1h, 12, 03 part, 15), .as well as to Mrs. Joan Murphy for carefully typing: this manuseript,

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Rerlese, A., 1904: Acari nuiovi. Manipulus IL Redia, 2: 257-280.

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Teil 5, Gamasiden, dAearvlogie,