RECORDS
OF THE
SOUTH AUSTRALIAN MUSEUM
VOLUME 19
Published by the Museum Board
Adelaide 1985-1986
iii
CONTENTS
Nos. 1-5. Published May 1985.
No. 1. A bibliography of Aboriginal archaeology in
South Australia, (Philip G, Jones) ......... ]
No. 2. A new and interesting species of Cerhegus
Thorell (Ischnothelinae, Dipluridae) from South
Australia, (Robert J. Raven).............. 15
No. 3. Millipedes from Australia, 5: Australio-
somatini from South Australia, with a note on the
status of Polvdesrynus inmotatus Karsch, and first
record of a second Mediterranean julid in
Australia (Diplopoda: Polydesmida, Paradoxo-
somatidae & Julida, Julidae), (C. A. W. Jeekel)
ease a, ea ee ce tak Saeco’ Aa soem rales 19
No. 4. Sarcoptiformes (Acari) of South Australian
soils. 4. Primitive oribate mites (Cryptostigmata)
with an extensive, unfissured hysteronotal shield
and aptychoid. (David C. Lee) ........... 39
No. §. Mollusc type-specimens in the South
Australian Museum. 2. Gastropoda: Conidae.
(Wolfgang Zeidler)... 212... .4 2226.2. ene 69
Nos. 6-10, Published June 1986,
No. 6. A revision of the troglophilic genus Brises
Pascoe with a discussion of the Cyphaleini
(Coleoptera, Tenebrionidae). (E. G. Matthews)
ee ey Phy) Et a | wih
No. 7. Additions to the collembolan fauna of Heard
Island. (Penelope Greenslade) ...... ht 9]
No. & Molluse type-specimens in the South Aus-
tralian Museum. 3. Polyplacophora. (W. Zeidler
FDL cs PR OPICTE Vises cw dosocnetrs Cxermawrn dda 97
No.9 A revision of the southern Australian starfish
genus Vecrria (Asteroidea: Oreasteridae), with the
description of a new species. (W. Zeidler and F. W.
Rowe) . 5 ae Het eee + ol NS
No. 1. Redescriptian ‘of ‘the mysid crustacean,
Noromvsis australiensis (Tattersall) Comb. Nov.,
representative of a new genus, (Karl J. Wittmann)
139
Nos. Il-l4. Published May 1986.
No. 11. On Trombella alpha n.sp. (Acarina: Trom-
bellidae) from Australia: correlation, description,
developmental abnormalities, systematics and
possible auditory structures. (R. V. Southeott).
No, 12. The genus Odontacarus (Acarina: Trom-
biculidae). Il. Observations on the life history and
morphology of Qdontacarus swani np. and
related forms (R. V. Southcott).......... 169
No. 13. History of the discovery of Speleognathus
australis Womersley (carina: Trombidiformes),
with notes on its natural history and behaviour.
(RO 8) Southcotty .. cee cea ee eee: 201
No. 14. Description of Odontacarus veitchi sp. nov.
(Acarina: Trombiculidae). (R. V. Southeott) 213
No, 15, Published August 1986.
A checklist of helminths from Australian birds
(Patricia M. Mawson, L. Madeline Angel and 8, J.
Podlnaraie)! sue ct 4 eee eee ere
Nos. 16-19. Published November 1986,
No. 16. Anatomical notes on the land snail Borh-
riembryon (Pulmonata: Bulimulidae) from South
Australia and Western Australia. (Ron C. Kershaw)
che a uener | aD.
No. ‘It. The Ephemeraptera ‘{tnayflies) of South
Australia. (P. J. Suter). . aes » a3
No, 18. First representative of the order Macro-
stomida in Australia (Platyhelminthes, Macro-
stomidae). (Ronald Sluys)..............+ 399
No. 19. A reassessment of the Papuan subfamily
Asterophyryinae (Anura: Microhylidae). (Thomas
Ae VTE Na iets, eae» aby wn c:nimiow one a et) ee
CONTRIBUTORS
ANGEL, L. MADELINE
See Patricia M. Mawson...............++:
BURTON, THOMAS C.
A reassessment of the Papuan subfamily Astero-
phyryinae (Anura: Microhylidae) 405
EDMONDS, 5. J.
See. Patricia M. Mawsonm............2.-5.- 219
GOWLETT, K. L.
See Wolfgang Zeidler :
GREENSLADE, PENELOPE
Additions to the collembolan fauna of Heard Island
|
JEEKEL, C, A. W.
Millipedes from Australia, 5: Australiosomatini from
South Australia, with a note on the status of Poly-
desmus innotatus Karsch, and first record of a
second Mediterranean julid in Australia (Diplopoda:
Polydesmida, Paradoxosomatidae & Julida, Julidae)
ee oe Fh eS eee ne
JONES, PHILIP G.
A bibliography of Aboriginal archaeology in South
Australia
KERSHAW, RON C.
Anatomical notes on the land snail Bothriembryon
(Pulmonata: Bulimulidae) from South Australia and
Western Australia. .........-2.56e6-e05 . oot
LEE, DAVID Cc.
Sarcoptiformes (Acari) of South Australian soils, 4,
Primitive oribate mites (Cryptostigmata) with an
extensive, unfissured hysteronotal shield and
BINMONMEL s-z. eile sy, pale ae Paes ats 39
MATHEWS, E. G,
A revision of the troglophilic genus Arises Pascoe
with a discussion of the Cyphaleini (Coleoptera,
CTE Ca its [rrr TT
MAWSON, PATRICLA M., L. MADELINE ANGEL
and 5. J. EDMONDS
A checklist of helminths from Australian birds 219
RAVEN, ROBERT J.
A new and interesting species of Cerhegus Thorell
(Ischnothelinae, Dipluridae) from South Australia
ROWE, F. W.
See Wolfgang Zeidler ................+.+.. INF
SLUYS, RONALD
First representative of the order Macrostomida in
Australia (Platyhelminthes, Macrostomidae). 399
SOUTHCOTT, R. V.
On Trombella alpha n.sp. (Acarina: Trombellidae)
from Australia: correlation, description, develop-
mental abnormalities, systematics and possible
AUGILOCY SIFUCUITES:. ce dee et tegen 145
The genus Odontacarus (Acarina: Trombiculidae).
II. Observations on the life history and morphology
of Odontacarus swani n.sp. and related forms. . 169
History of the discovery of Speleognathus australis
Womersley (Acarina; Trombidiformes), with notes
on its natural history and behaviour ....... 201
Description of Odontacarus veifichi sp. nov.
(Acarina: Trombiculidae)............25..55 213
SUTER, P. J.
The Ephemeroptera (mayflies) of South Australia
339
WITTMANN, KARL J.
Redescription of the mysid crustacean, Nofomysis
australiensis (Tattersall) Comb, Nov., representative
CE CVE POON, = nich lem cake Peay aha eln seth oe
ZEIDLER, WOLFGANG
Molluse tyfie-specimens in the South Australian
Museum. 2. Gastropoda: Conidae ........., 69
ZEIDLER, WOLFGANG and K. L. GOWLETT
Molluse type-specimens in the South Australian
Museum. 3. Polyplacophora .........-..... 97
ZEIDLER, WOLFGANG and F. W. ROWE
A reviston of the southern Australian starfish genus
Nectria (Asteroidea: Oreasteridae), with the
description of a mew species............... 117
Vii
INDEX
PRCT UAL <a osc Sem elaionecint « « a ee Pee (Ui) Ts EBay Re ne A, ee aa es op os ee, 19
(POS ie ea eh TT Ne PeeWee Slee er Lie Us PPM UNLIELE dct asso iok-as area als Reuter 19
PATNA eee ee kt ae 145, 161, 165, 213 f
Gove Cutan, ......2..:08 ee 145 [ethers fea; til Pe aes eee i eee erage, Skee 327
PRUNE Roe hechey uss Mian &hrdnes Peas lames yee ee AGS MIAChOStOMIM ss eck bic eee cae ee ee 399
PN TALNEANUEND Oe a vaturn = os Wes es Cea Re ed ye } OiNTaereatoninadie sucha ae. se eka ee oe es 399
seth Rene tt | cts Up eet ea 117 Mayflies (of South Australia) ............... 339
ASCO PITVINAG 6s eds ce ee Pa ee ADS” Microhylidat so 22. Pac. oP ice Cine i ce rae 405
australiensis, Notomysis .................0.: 139 Millipedes (from South Australia)............. 19
AustraliosOMatinae...........00 cess eee beeen LS Se pei a te oer Re, I 39
australis, Speleagnathus ...,.....:.0.000008: 201 Mollusc (type-specimens)..............25 69, 97
: ; Nip Ses oy CR ee ee 139
Birds (helminths from) .........0.....--0005 219
FOUTS VON oa eae Ah eee eget ee ee BAT UN BCRE DR Pou sist a oa acta ie eee oe eae ota 117
FBR es pe eh tas Sire easel cideakel a al eaek acpn meee ol AP MENLEMTISAIM sche cae ey oes oh old + a ttt sm tee er ae 139
meer ie? sass aoa a tts wep SeammemMartt 22. coia Sova cd ghee 169, 213
CSIP TE jig, lat eae, dea 2 RRR aD Tere Re REE ALA rs 5 ahd nor aan rates ah a ate ee keg 117
Coleoptera... 2.16. eee eee es 2 Sen sire, ante Mat Polio ENDS TIMES 2 Sy co ws thn eceer then ts Pe Se aaa 39
Collembola el Soa ea ata Ra pee Etat Paradonosomatidae .........- cece eee street 19
MOUNTING! 5. '. eens aoa Pe ta ars a alin ae ee ee 69 ;
Plotynetiinthes ceca ek hss Piaie. oh cade. 399
PEDAL ACA a pare ese tra aie as oo ate em Thiptc waste Gales 139 i
‘ Polynit ©. 55.0. is fad tui oe eae ee gee 19
i mycin) Ci Jnr it Oh Eee RNR Pei a ee ere oo 39
Cyphaleini 7 POUWP SINE S nr ba). a Re eee als See 19
ce a ai ys aa Se PolyplaGoplord... essen se eee ee ceaeeentawes OT
Pp pOUR 2.1). ck ate ee Oe as ad ae 19 Pe 2 oe wakes eek ee eee ca ee 327
Belinda ss rca se stein Pleo. s Sate ee eo 15 :
Shel DCC PUP CRUE oe, gee cher garg hk ete ee erate ag
Fphemens (Hert suerss-.ceagu hs le. es S30 RE neleneriaeaus 2) yaa actevayg eet a-un 5 pee 201
fe ey ee ae CO oe ee 69 swani, Odomtacarus . 00.6606 c eee c eee eens 165
Heard Island (Collembola of) .............--- Pe iccicecypett ia iain Reketee bee cote 0 a
[OA RG RN” Gee sl cin iia Mee 219 Trombella a eed eae ce eRe: o ee eee eres y 145
; Thanibellidne: oc Cicer on ei eek Sete es 145
innotatus, Polydesmus........0:0+eeeeeeecees 1D Trombieubidee... . 24... 2.6 ese see ees 165, 213
[schinothelinaes «oa 202. Sx ewe ee oe sc es 1S ‘Thonibidiformes 2.0... ed ae eee ee 201
weibehi, Qdontacarus: .s4 vis vece csc ee eew ens 213
RECORDS oF THE
SOUTH AUSTRALIAN
MUSEUM
VOLUME 19 NUMBERS 1-5 MAY 1985
No. 1 A BIBLIOGRAPHY OF ABORIGINAL ARCHAEOLOGY IN
SOUTH AUSTRALIA
by PHILIP G. JONES
No. 2 A NEW AND INTERESTING SPECIES OF CETHEGUS
THORELL (ISCHNOTHELINAE, DIPLURIDAE) FROM SOUTH
AUSTRALIA |
by ROBERT J. RAVEN
No. 3 MILLIPEDES FROM AUSTRALIA, 5: AUSTRALIOSOMATINI
FROM SOUTH AUSTRALIA, WITH A NOTE ON THE STATUS
OF POLYDESMUS INNOTATUS KARSCH, AND FIRST
RECORD OF A SECOND MEDITERRANEAN JULID IN
AUSTRALIA (DIPLOPODA: POLYDESMIDA, PARADOXO-
SOMATIDAE & JULIDA, JULIDAE)
by C. A, W. JEEKEL
No. 4 SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS.
4. PRIMITIVE ORIBATE MITES (CRYPTOSTIGMATA) WITH AN
EXTENSIVE, UNFISSURED HYSTERONOTAL SHIELD AND
APTYCHOID.
by DAVID C. LEE
No. 5 MOLLUSC TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN
MUSEUM. 2. GASTROPODA: CONIDAE
by WOLFGANG ZEIDLER
SOUTH AUSTRALIAN MUSEUM
North Terrace, Adelaide
South Australia 5000
A BIBLIOGRAPHY OF ABORIGINAL ARCHAEOLOGY IN SOUTH
AUSTRALIA
BY PHILIP G. JONES
Summary
A bibliography of South Australian Aboriginal archaeology, indexed accordingly to subject and
region, and citing 600 references, including published and unpublished material is presented.
A BIBLIOGRAPHY OF ABORIGINAL ARCHAEOLOGY IN SOUTH AUSTRALIA
Compiled by
PHILIP G. JONES
South Australian Museum, North Terrace, Adelaide, South Australia 5000
(Manuscript accepted 30 August 1984)
ABSTRACT
JONES, PGi. 1985, A Bibliography of Aboriginal archeology in South
Australia, Ree S Aust, Mus. Py) 1-14.
A bibliography of South Australian Aboriginal
archaeology, indexed according to subject and region,
and citing 600 references, including published and
unpublished material is presented.
CONTENTS
INTRODUCTION
SERIES ABBREVIATIONS USED
A. GENERAL
(1) Legislation
(2) Origins and General Texts
(3) Trends and Methods
B SUBJECT LISTING
(1) Incised Stones, Cyleatis
(2) Stone Tools and lidustries
(3) Stone Arrangements, Structures
(4) Rock Engravings and Paintings
(5) Caves and Rockshelters
(6) Shell Middens
(7) Bone Points, Wooden Artifacts
(8) Canoe Trees, Scarred Trees
(9) Excavations of Human Remains
(10) Radiocarbon Dates
Cc, REGIONAL LISTING
General
(a) Western, North-Western 9
(b) Eastern, North-Eastern 9
(c) Mid-North, Flinders Ranges 10
(d) Eyre Peninsula, Yorke Peninsula 10
(c) Lower North, Adelaide Region, Fleurieu Peninsula = 11
(f) River Murray Region aI
oo coer nmr anorrsk be VYHnvht w=
(g) Lakes, Coorong, South East i
(h) Kangaroo Island i2
INTRODUCTION
Archaeological knowledge is constructed from the
available evidence; this bibliography represents an
attempt to document this evidence as it relates to South
Australia and to make it accessible to researchers.
The bibliography contains a full listing of references
relating to the theory and practice of Aboriginal
archacology in South Australia, It cites published and
unpublished material extracted from a wide range of
sources. Unpublished material has been chosen
selectively; only notes or documents of direct
Mey, [UN
significance have been included. Recent newspaper and
magazine articles about archaeology are numerous and
vary in quality. Only significant examples have been
included,
The bibliography falls into three main sections:
A. General
Section A is an introductory section covering relevant
legislation and general texts relating to theory, method
and developments in South Australian Aboriginal
archacology.
B. Subject Listing
Section B contains subject-related references (“Rock
Engravings and Paintings’, “Shell Middens’ etc.)
organized under ten subject headings. Individual
entries are cross-referenced to the regional divisions
dealt with in Section C by means of alphabetical key
letters which follow each entry,
C. Regional Listing (see Fig. 1)
Section C includes references which relate to
particular areas in South Australia rather than to
specific archaeological subjects as dealt with in Section
B. These references have been arranged under a general
heading, relating to the State as a whole, and under
cight regional headings (a-h), These broad regional
divisions reflect a convenient organization of the
reference material. They do not represent precise
geographic or topographic boundaries or cultural areas,
The subject and regional listings have been used as
(he most effective means of arranging the quantity and
diversity of reference material. The two listings are not
mutually exclusive; regional listings in Section C should
be supported by relevant material from Section B.
For those conducting archaeological research in
specific localities there are two additional sources which
may be consulted. Both contain restricted information:
(1) The annual Bibliography of Consulting Reports
prepared by the Australian Association of Consulting
Archaeologists Incorporated. This can be obtained
through the Association, PO. Box 214, Holme
Building, University of Sydney, N.S.W, 2006.
(2) South Australian Archaeological Site Reports.
Two different files of these Reports are maintained, one
in the South Australian Museum (Anthropology and
Archacology Section) and the other in the South
Australian Department of Environment and Planning
(Heritage Conservation Branch).
The writer wishes to thank Mrs Joan Murphy for
her cooperation and patience in typing this manuscript.
rs
A.HLR.
A.LA.S.
Am. Antiquity
American Philosoph. Soc. Mem.
Annals Aust. Coll. of Dental Surgeons
Ann. Rev. Anthrop.
Archaeol. in Oceania
Arch. Phys. Anthrop. in Oceania
Artefact
Asian Persp.
Aust. Ab. Stud
Aust. Archaeol,
Aust. Archaeol. Assoc. Newsletter
Aust. J. of Botany
Aust. J. Set.
Aust, Mus. Mag,
Aust. Mus. Mem,
Aust, Nat, Hist,
Aust. NZ. Ass. Ady, Sei,
Aust. Quit. Newsl.
Cam. U. Press
Curr. Anthrop.
ELS,
FOS.AM.
J. Arch, Science
J. Anthrop. Soc. S, Aust.
J. Biogeogr.
J. Roy, Anthrop. Inst.
J. Roy. Soc. W. Aust.
J. Sov. Oveanist.
Nat. Hist.
Nat. Mus. Vie. Mem,
New Quart. Cave
OUP.
Proc. Prehistoric Society
Proc. Roy, Geog. Soc. Aust., 5. Aust. Branch
Proc. Roy. Soc. Tas.
Proc, Roy. Soc. Vic.
Rec, Aust. Mus.
Rec. S. Aust. Mus,
Report Aust, Assoc. Adv. Sci.
Rep, Wash. Geol, Cong.
Revue Ethnogr. Tradit. Pop.
Site Rec, Newsletter
S.A. Mus. A. D.
S. Aust. Nat.
S. Aust Ornithologist
Syd. Univ. Spel. Soe. J.
Trans. Br, Cave, Res, Ass.
Trans. Roy. Soc. S, Aust.
View Nal,
A. GENERAL
(1) Legislation
RL. S. AUST, MUS. 19 (1) 14 \fa, [ONS
SERIES ABBREVIATIONS USED
Aboriginal and Historie Relics Unit relocated from South Australian Museum in 1979
Currently operates as part of the Heritage Conservation Branch, Department of
Environment and Planning, South Australia
Australian Institute of Aboriginal Studies, Canberra
American Antiquity
American Philosophical Society Memoir
Annals of the Australian College of Dental Surgeons
Annual Review of Anthropology
Archaeology in Oceania
Archaeology and Physical Anthropology in Oceania
The Artefact
Asian Perspectives
Austrahan Aboriginal Studies
Australian Archaeology
Australian Archeological Society Newsletter
Australian Journal of Botany
Australian Journal of Science
Australian Museum Magazine
Australian Museum Memoirs
Australian Natural History
Australian and New Zealand Association for the Advancement of Science
Australian Quaternary Newsletter
Cambridge University Press
Current Anthropology
Environmental Impact Statement
Friends of the South Australian Museum Newsletter
Journal of Archaeological Science
Journal of the Anthropological Society of South) Australia
Journal of Biogeography
Journal of the Royal Anthropological Institute
Journal! of the Royal Society of Western Australia
Journal de Ja Socieie Oceanistes
Natural History ;
National Museum of Victoria Memoirs
New Quarterly Cave Research
Oxford University Press
Proceedings of the Prehistoric Society
Proceedings of the Royal Geographical Society of Australia, South Australian Branch
Proceedings of the Royal Sociery of Tasmania
Proceedings of the Royal Society of Victoria
Records of the Australian Museum
Records of the South Australian Museum
Report of the Australian Association for the Advancemem of Science
Report of the Washington Geological Congress
Revue D'Ethnographie et des Traditions Populairs
Site Recording Newsletter, Heritage Conservation Branch, Department of Environment
and Planning, South Australia
South Australian Museum Archival Documentation
South Australian Naturalist
South Australian Ornithologist
Sydney University Speleological Society Journal
Transactions of the British Cave Research Association
fransactions of the Royal Society of South Australia
Victorian Naturalist
(2) Origins and General Texts
BAGLIN, D, and MULLINS, B. 1969 “Aborigines of Australia:
Horwitz, Sydney.
BAILEY, G. N. 1980. Holocene Australia. /n “The Cambridge
ELLIS, R. W. 1972. South Australia: Existing legislation and its
implementation. /n The Preseryation of Australia’s Aboriginal
Heritage, Report of National Seminar on “Aboriginal Anriquines
in Australia” 23rd and 24th May 1972, A.LA.S., Canberra,
EDWARDS, R. 1970. Legislation for the protection of Aboriginal
relics in South Australia, /n “Aboriginal Antiquities in Australi;
Their Nature and Preservation” (F. D. McCarthy, ed,), pp.
159-165. A.1.A.S., Canberra.
SOUTH AUSTRALIA (STATUTES) 1965. Aboriginal and Historic
Relics Preservation Act. Govt. Printer, Adelaide,
SOUTH AUSTRALIA (STATUTES) 1979. Aboriginal Heritage Act.
Govt. Printer, Adelaide.
SOUTH AUSTRALIA (STATUTES). Aboriginal Heritage Bill (Draft
form). Act not yet proclaimed (August 1984),
WARD, G. K. 1983. Archaeology and legislanion in Australia, Jn
“Australian Field Archaeology. A Guide to Techniques?
(G. Connah, ed.) pp. 18-42. A.1.A.S., Canberra.
Encyclopedia of Archaeology” (A. Sherratt, ed.), pp. 333-341.
Cambridge University Press, Cambridge.
BASEDOW, H. 1925. “The Australian Aboriginal’ Preece & Sons,
Adelaide.
BERNDT, R. M., and C. H. (eds.). 1965. “Aboriginal Man in Australia!’
Angus & Robertson, Sydney.
BERNDT, R. M., and C, H. 1981. “The World of the First Australians”
Lansdowne Press, Sydney.
BIRDSELL, J. B. 1953. Same environmental and cultural factors
influencing the structure of Australian Aboriginal populations.
The American Naturalist, 87: 171-207.
BIRDSELL, J. B. 1957. Some population problems ifvolving
Pleistocene man, Cold Spring Harbor Syriposta on Quantitative
Biology, 22: 47-69.
KIRDSELL, J, B. 1967, Preliminary data on the tribybrid origin of
ieee Aborigines. Arch, Phys. Anthrop. in Oceania, 2:
155.
BIBI MOGRAPHY OF ABORIGINAL ARE LIALOLOGY
BIRDSELL, J. B 1977. The reenlibrarion of a paracigin for the thrst
peapling of Greater Australia. Jv “Sunda and Sahl: prehisiane
studies jn Sourneast Asia, Melanesia and Australia!’ (J. Allen,
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BLAINEY, G. N. 1975. “Triumph of the Nomads: A Histury of
Ancient Australia!’ Maemillan, Melbourne
CAMPHELL, A. H. 1967. Aboriginal traditions and he prehstary
of Australia, Marikirid, 6, 476-481
CASEY, D, A. 1940, The present siate of our knowledge of (he
archaeology of Ansitalia 7 “Proceedings af the third Congress
of Prehistorians of the Par East", Sinvapore
COTTON, HC. (ed) 1966. “Aboriginal Man in South and Central
Australia!” Part 1, Giovt, Pronter, Adelaide,
COUTTS, Pd, bh 1886. “Features of prehistoric campsites jn Auginalia’’
Mankind, Gi, 338-346.
DIAMOND, J. M. 1977. Distibunenal strategies. fy “Sunda and
Sahul: prehistoric studies an Southeast Asia, Melanesia and
Australia” (I. Allen, J, Golson, and R- Jones, eds.), pp. 295.316.
Academic Press, Landon,
EDWARDS, R.. (ede). 1975. “The Preservanon of Australia's Abonginal
Heritage. Report of National Seminar on Aboriginal Antiquities
in Australia, Many 1972! ALAS. Canberra.
FLOOD, J, 1983, “Archaeology of the Dreanjime’, Collins, Sydney.
OL, BE, DB, 1963, Australian Aborigines and (he giant extinct
marsupials. dase Nae Hist, be 263-266.
GOULD, R. A. 1973, Ansrralian archaeology in ecological and
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GREENWAY, | 1962 Archaeology in Australia. Southwestern Lare,
28; 25-30,
HAMMOND, G, 19]. Aspects of leva) significance in Archaeology,
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HORTON, BD. R. 1981. Water and woodland: the peopline of Australia,
ALAS. Newsletter We 21-27.
HORTON, D. R. 1981, Eurly thought on early qe i Australia.
Artehier @: 33-69,
HOSSEEL D, BS. 1966. Antiquiry of nan in Australia Jn “Aboriginal
Manin South and Cerrral Ausiclia, Part I” |B OO. Cotten, ed },
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HOWCHIN, W, 1933, A rejoinder to some recent ctlinological papers,
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JENBINGS, JN. 1971. Sea level changes and land links. Jn
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MeCARTHY, | OD 1963, The prehistory of the Australian
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‘\\ KANGAROQ ISLAND " yr
\ eto!
Ne
Ny
oN
© >10,000 years B.P. res!
vw = €10,000 years B.P. XQ
wo 5,000 years B_P, OR 90d
*% Stone structures
( Rock paintings.
@ = Rock engravings
|
FIG. 1. Map of South Australia showing the regions referred to in the text. Symbols denote localities of different subjects listed (see key
for explanation).
A NEW AND INTERESTING SPECIES OF CETHEGUS THORELL
CISCHNOTHELINAE, DIPLURIDAE) FROM SOUTH AUSTRALIA
BY ROBERT J. RAVEN
Summary
A new species, Cethegus ischnotheloides, is described from South Australia. Males possess spine-
like bristles on the palpal tarsi that are attenuate as in Ischnothelini.
A NEW AND INTERESTING SPECIES OF CETHEGUS THORELL (ISCHNOTHELINAE, DIPLURIDAE)
FROM SOUTH AUSTRALIA
by
ROBERT J, RAVEN
Queensland Museum, Gregory Terrace, Fortitude Valley, Queensland, 4006
(Manuseript accepled 22 March 1983)
ABSTRACT
RAVEN, Ro 1985. A new and interesting speeies of Cerrenns
Thorell (ischnorheliiae: Dipluridaed fram South Australia, Rec.
S. lust Mus. 42). 15-17,
A new species, Cethegus ischnotheloides, is described
from South Australia. Males possess spine-like bristles
onthe palpal tarsi (hat are attenuate as in [schnothelini.
INTRODUCTION
Until recently, only three species of Cefhegus were
known trom Australia (Main 1960), In a revision of
Australian Ischnothelinae, Raven (1984) clarified the
diagnostic characters of Cethegus and C. fugax
(Simon) and described eight new specics, Main (1960)
included specimens from eastern South Australia in C.
fugax, However, Raven (1984) included only specimens
from southwestern Australia in that species. When
Raven's study was completed, Mr David Lee, Senior
Curator of Arachnids and Helminths at the South
Australian Museum, presented me with two interesting
males of Cethegus from central South Australia. One
character of these males is unlike that of any other
Australian ischnothelinid and requires amendation of
the diagnosis of the Euagrini.
Abbreviations are standard for the Araneae and with
methods used may be found in Raven (1984), All
measurements are in millimetres except eye measure-
ments (hat are in graticule units, each being 0.025 mm.
SYSTEMATICS
Tribe Euagrini Raven, 1979
Diagnosis
Diplurid spiders with long posterior lateral spin
nerets with long apical segment; a hirsute pigmented
cuticular crescent surrounds the base of the posterior
median spinnerets. No cuspules on labium or maxillae.
Spines present or absent on normal or attenuate male
palpal tarsi. Cheliceral furrow with one row of teeth
on promargin. Trichoborhria with corrugiform collar
around bases. Jarsal organ low.
Remurks
Previously, the presence of spines on male palpal
tarsi was known only for the western European genus
Phyxioschaema in the Euagrini (Raven, 1981). However,
only [schnothelini (in the Dipluridae) have the
characteristic extension of the male palpal tarsi (see
Raven, 1983), In most Euagrini, palpal tarsi of males
are short and truncate; in Cefhegus ischnotheloides,
the male palpal tarsi are ‘spinose’ and are slightly
elongate apically (unlike other known Cethegus),
Tribe Euagrini Raven
Cethegus ischnotheloides 1. sp.
(Figs 1-4) (Table 1)
Diagnosis
Males with elongate embolus reaching proximal
palpal patella; spine-like bristles present on attenuated
palpal tarsi. Sternum with blunt hairs. Females
unknown.
Holotype male SAM NI19&1 394
Carapace 6.56 long, 5.69 wide. Abdomen 5.38 long,
4.13 wide.
Description:
Colour in alcohol: Carapace and legs reddish brawn,
chelicerae maroon, Abdomen entirely brown,
Carapace: Fovea short, semicircular, deep. Lateral
margins with few Weak lateral bristles. About 3 pairs
of foveal bristles. Striae deep, glabrous. Clypeus 0.18
wide. Black hairs on interstrial ridges.
Eyes; On strongly raised tubercle; group is 0.27 times
head-width, and is 1.59 times wider than long, Front
row centres form strongly procurved line; back row
centres form slightly procurved or straight line, Ratio
of AMB:ALE:PME:PLE, 18:14:11;13. Ratio of MOQ
front width:back width:length, 30:33:28. Eye inter
spaces as AME diameters; AME-AME = ALE-PLE,
0.3; AME-ALE = PME-PLE, 0.05,
Chelicerae: With fine transverse dorsal ridges.
Promargin with 4 large and 8 small teeth; basally with
2 fine teeth.
Maxillae: 1.80 long, 1.25 wide; with some short
pointed setae; serrula ridge-like.
Luhium: 0.63 long, 1.13 wide; groove broad and
continuous.
Sternum: 3.52 long, 2.92 wide; all sigilla oval;
posterior and middle sigilla 0.18 long and away from
'f REC, S, AUST. MUS. 19 (2), IST
Vins LVRS
FIGS, 1-4, Cethegus ischnotheloides fesp, mae holotype. |. carapace and cheliverae. 2, sternum, maxillae and labium. 3, dorsal palpal
larsus showing al(cnuation and spine-like brisiles, 4. rewolateral palpal patella, fibia aud tarsus. Seale fine = 2mm for fig. by) mu
for figs 2, 4; 0.5 mm for fig. 3.
margin; anterior sigilla 0,20 long and 0,25 away from
margin, Setae Jong erect or short blunt.
Palp; Bulb pyriform; embolus long, tapering,
extending to proximal patellae; tibia with slight ventral
groove; tarsus slightly pointed with cluster of slender
spine-like bristles,
Legs; Leg formula, 4321. Leg | smooth, cylindrical.
Tarsi 11] and (V erect hairs forming weak scopula
divided by setae. Spines. No true spines on femora or
patellae. Leg 1: tibia, pl v2; metatarsus, p2 v2; tarsus,
v4, Leg 2: tbia, p2 v3; metatarsus, p3 V5; tarsus, v5.
Leg 3: tibia, p3 di r3 v6; metatarsus, pll r5 v8; tarsus
d3 v7. Leg 4: tibia, p4 r4 v7; metatarsus, p9 d3 5 va;
tarsus d4 v9. Palp: 0. Claws: 12 tine teeth on paired
claws; 3 fine teeth on unpaired claw. Trichobothria: Two
rows, each of 10, on tibiae; abour 10, in a line on
metatarsi; 8, of different length on tarsi in irregular row,
Spinnerets: Posterior medians 1.20 long, 0.36 in
diameter and 0.68 apart; crescent of cuticle present as
darkened area forming an anterior fold. Basal, middle,
apical and total articles of posterior laterals, 1.92, 1.64,
2,04, 5.60 long respectively,
Material examined: Holotype male (N1981394),
paratype male (IN1981395), sand dunes, Commonwealth
Hill Station, 29°57'S,, 134°10'E,, South Australia,
April, 1981, P. Bird, R, Sinclair, deposited S.A.M.
Disiributien; Central South Australia.
TABLE I. LEG MEASUREMENTS OF CETHEGUS
ISCHNOTHELOIDES.
VALUES ARE FOR MALE HOLOTYPE.
Leg I oLeg Hl OLeg WE) Leg iV Palp
Femur §.13 5.06 5.00 6.25 3.44
Patella 2.94 2.81 281 3.19 1.94
Tibia 3.19 3.19 3.31 3.56 2.81
Metatarsus 4.69 5,06 5.94 6.44 —
Tarsus 2.25 2.69 3.19 3.75 1.69
Total 18.20 IB8T 20 25 2IA9 9.R8
Remarks: Males of Cethegus ischnotheloides differ
from those of © fugax in the ‘spinose’ pointed palpal
tarsi.
PHYLOGENETIC AND
BIOGEQGRAPHIC SIGNIFICANCE
Inasmuch as males of C. ischnoiheloides possess
‘spinose’ pointed palpal tarsi, they differ from males
of all other Australian Euagrini and bear some
resemblance to male Ischnothelini, However, because
spinose male palpal tarsi are also found in genera of
Masteriinae and Diplurinae, the presence of spines may
be regarded either as. a symplesiomorphy or a
parallelism—a hypothesis involving their
synapomorphy is falsified by the absence of cuspules
and the presence of corrugiform trichobothrial base
collars in Euagrini, | here regard the presence of spinose
palpal tarsi in Ischnothelinae as a plesiomorphic
retention. However, parsimoniously, the attenuated
palpal tarsus of males of C jschnothelaides is
presumably a parallelism otherwise Ce/hegus and
Ischnothele are paraphyletic.
That a Western Australian cuagrine should retain
characters otherwise present only in Indian and
Neotropical (/schnothele) and Ethiopian (Lerhrothele
and Thelechoris) genera is probably predictable. Same
geologists (e.g. Powell ef al., 1981) propose that pre-
drift India was in close contact with Western Australia.
if that were so, some sharing of taxa dating to that
period is to be expected. Spiders of the genus Ce/hegus,
as defined by Raven (1984), are predominantly
xerophilic although a number of species occur in
northeastern Queensland, Throughout its range, to my
knowledge, Cethegus is allopatric with other euagrine
genera. | suggest that Cerhegus represents a group that
was first isolated in Australia as the most plesiomorphic
A NEW SPECIES OF CETHEGUS 17
genus (that proposition will be discussed more
completely elsewhere) and as such has retained
characters of the presumed sister group of the Euagrini,
the Ischnothelini. Therefore, Cethegus ischnotheloides
may be the most plesiomorphic Cethegus species.
ACKNOWLEDGMENTS
I am grateful to Mrs V. E. Davies and R. Molnar
for commenting upon this manuscript, to Mr David
Lee for bringing the spiders to my attention and to Ms
R. Owens for carefully typing the manuscript.
REFERENCES
MAIN, B. Y. 1960. The genus Cethegus Thorell (Mygalomorphae:
Macrothelinae). JR. Soc. West. Austr. 52: 9-11.
POWELL, C. McA., JOHNSON, B. D., and VEEVERS, J. J. 1981.
Gondwanaland, the separation of Australia and India and their
interaction with southeast Asia, 17-29. Jn Keast, A. (ed.).
Ecological biogeography of Australia. Monographie Biologicae
41. London: W. Junk, The Hague.
RAVEN, R. J. 1981. The mygalomorph spider genera Phyxioschaema
Simon and Sfenygrocercus Simon (Ischnothelinae: Dipluridae:
Araneae). Bull. Brit. arach. Soc. 5(5): 225-31.
RAVEN, R. J. 1983. Notes on some mygalomorph spiders in the
Musee Royal de |’Afrique Centrale (Dipluridae, Hexathelidae,
Ctenizidae: Araneae). Rev. zool. Afr. 97: 549-562.
RAVEN, R. J. 1984. Systematics of the Australian Curtain-web
spiders (Ischnothelinae: Dipluridae: Chelicerata). Aust. J. Zool.
Suppl. Ser. 93: 1-102.
MILLIPEDES FROM AUSTRALIA, 5: AUSTRALIOSOMATINI FROM
SOUTH AUSTRALIA, WITH A NOTE ON THE STATUS OF POLYDESMUS
INNOTATUS KARSCH, AND FIRST RECORD OF A SECOND
MEDITERRANEAN JULID IN AUSTRALIA (DIPLOPODA:
POLYDESMIDA, PARADOXOSOMATIDAE & JULEDA, JULIDAE)
BY C. A. W. JEEKEL
Summary
Two new species of the genus Heterocladosoma Jeekel, H. zebratus and H. galaxias, and one of the
genus Somethus Chamberlin, S. grossi, are described. Australiosoma castaneum Attems is
redescribed and made the type-species of a new genus Oncocladosoma, which is probably related to
Somethus, Dicladosomella Jeekel and Phyllocladosoma Jeekel. A new subspecies, O. castaneum
ingens, and two new species, O. conigerum and O. clavigerum, are added to the new genus. The
holotype of Polydesmus innotatus Karsch, a subadult female, has been re-examined; it may be
referable to a genus close to Otoplacosoma Verhoeff (tribe Antichiropodini). A list of the known
South Australian Paradoxosomatidae is given, and some distributional patterns are briefly
discussed. Brachyiulus lusitanus Verhoeff, a west Mediterranean julid, is recorded from a number
of South Australian localities.
MILLIPEDES FROM AUSTRALIA, 5: AUSTRALIOSOMATINI FROM SOUTH AUSTRALIA, WITH A
NOTE ON THE STATUS OF POLYDESMUS INNOTATUS KARSCH, AND FIRST RECORD OF A
SECOND MEDITERRANEAN JULID IN AUSTRALIA (DIPLOPODA: POLYDESMIDA, PARADOXO-
SOMATIDAE & JULIDA, JULIDAE)
by
Cc. A. W. JBEREL
Institure of Taxonomic Zoology (Zoological Museum), University of Amsterdam, P.O. Box 20125, 1000 HC
Amsterdam, The Netherlands
(Manuscript accepted 22 September 1983)
ABSTRACT
JEEREL, C, A, W. 1985, Millipedes from Australia, Ss
Austriliosamatini trom South Australia, with a bete on the seins
of Palvdesiius tinetats Karsel and first recor af a second
Mediterranean julid in Australia (Diplopoda: Polydestmita,
Paradoxosoniiudac ad Dulida, Fulichie), Kee S, dash Ates, (93):
19-37,
Two new species of the genus Heteracladasoma Jeekel,
HY. zebratus and A. galaxias, and one of the genus
Semethus Chamberlin, S. grossi, are described.
Australiosoma castaneum Attems is redescribed and
made the type-species of a new genus Oncecludosome,
which is probably related to Somethus, Dicladosomella
Jeekel and Phyllocladosoma Jeekel. A new subspecies,
O. castaneum ingens, and two new species, Q. conigeruri
and O clavigerum, are added to the new genus. The
holotype of Polydesmus innolatus Karsch, a subadult
female, has been re-examined; it may be referable to a
genus close to Oreplacosoma Verhoeff (tribe Antichiro-
podini). A list of the known South Australian Para-
doxosomatidae is given, and some = distributional
patterns are briefly discussed, Bracltyuylys lusizenus
Verhacff, a west Mediterranean julid, is recorded from
a number of South Australian localities.
INTRODUCTION
This is the second of two papers treating the
Paradoxosomatidae of South Australia, The first paper
dealt with the Antichiropodini (Jeekel 19824) and the
present contribution treats the Australiosomatini of the
State. As pointed out already, our knowledge of the
mullipedes of Souch Australia is extremely scanty, In the
family Paradoxosomatidae only two species had been
described: Paolydesmus (Strongvlasoma) innoratus
Karsch, 1881, an unrecognizable species of uncertain
taxonomic status, and Austreliosome castaneun Attems,
1944, a reasonably well characterized bul as yet not
correctly classified species (cf. Jeekel 1968: 26).
The material reported upon was received on loan from
the South Australian Museum, Adelaide. lt had been
collected mostly in the surroundings of Adelaide, and
only lew samples from elsewhere were available.
Nevertheless, it gives a fair lirst impression of the
composilion of the fauna, and indicates that South
Australia has a relatively rich representation of
Paradoxosomatidae (together with the suborder
Cambalidea of the order Spirostreptida the family
appears to be the dominant millipede group) with still
many new species to be expected.
The type-specimen of Polydesmus innotatus Karsch
has been re-examined, but it is a subadult female, which
cannot be associated with any of the other South
Australian paradoxosomatids at hand. Neither can it be
referred with certainty to either of the two relevant
Australian paradoxosomatid tribes.
It has been pointed out carlier (feekel 1981; 20) that
records of millipedes introduced into Australia from
elsewhere are few. The material in the South Australian
Museum contains, besides some specimens of
Ommatoiulus moreleti (Lucas), a species now known to
have become widespread in a large part of South
Australia, also a number of samples of Brachyiulus
lusitanus Verhoelf, a Mediterranean julid known to have
synanthrope tendencies, bul as yet recorded only a few
times from outside the palearctic region.
The bulk of the material treated, including the types
of the new species, is preserved in the South Australian
Museum, some paratypes and voucher specimens lave
been retained by the author for the Zoological Museum
at Amsterdam.
SOUTH AUSTRALIAN AUSTRALIOSOMATINI
Heterocladosoma Seckel
Heierucladosoma Jeekel,, 1968: 144.
Type-species! Eustrongylosomea bifaleatum Silvestn, 19K,
Remarks
This genus was proposed for three species occurring
in the coastal region of Queensland. The quile
unexpected discovery of two new species in South
Australia extends the range of the genus considerably
and establishes an important faunistic link between the
fauna of South Australia and that of the Australian cast
coast.
2 EL
Heterocladosoma is well characterized by two
libiotarsal branches arising from the base of the
acropodile of the gonapod, one narrow lanceolate, the
other more voluminous, broadly lamimate and apically
more or less recurved. The femoral process is coalesced
with the solenomerite over most af jts length, its Iree
part arising from the distal part of the channel-bearing
branch of the gonopod as in Somethus Chamberlin. But
contrary to Sentethus, which has only one tibiotarsal
branch, the spermal channel does not make a loop, but
runs straight towards the apex of the solenomierite
Heterocladosoma zebratum n. sp.
Material
Coralbignie to Buckleboo, South Australia,
15-16,111.1950, leg. G, F, Gross,¢ holotype, 3 ? paratypes,
North of Kokotha, South Australia, 11.V1.1956, burnt
out of spinifex, leg. G, F Gross, 1 3 paratype.
Nonning, Gawler Ranges, South Australia, 17. VI.
1956, burnt out of spinifex (Veda sp.), leg. G. F Gross,
1 ¢ paratype.
Description
Colour Probably somewhat faded. Head with
clypeus, frons and anterior part of vertex brown;
remainder, including lateral sclerites brownish yellow.
Antennae light brown, infuseate towards apex of fth
antennomere; 7th antennomere black, its tip whitish.
intersegmental membranes of antennae brownish yellow,
Callum with a broad zone along anterior and lateral
margins brown, remaining part brownish
yellow. Subsequent somites with a rather narrow zone
of the prosomites adjacent to waist, the waist itself, and
the part of metatergites adjacent to waist, abaut halfway
towards the transverse furrow, brown, fading below the
level of paranota to brownish yellow. Remaining parts
of pro- and metasomites brownish yellow. Venter and
sternites brownish yellaw. Legs brown, with pale
intersegmental membranes and a pale tarsal np. Anal
somite dorsally brownish yellow, sides brawn; margins
yellowish. Paraprocts brown, with brownish yellow
margins, Hypopract yellowish,
Width: d= 2.$-3.5 mm, 2; 3.23.5 mm,
Head ard antennae: Labrum widely and rather weakly
emarginate Clypeus rather strongly impressed towards
Jabrum and with an impression on each side belaw
antennal sockets. Lateral border widely canvex, weakly
emarginate near labrum. Surface uneven due to the
presence of setiferous pits, Pubescence moderate, setae
of moderate length. Lateral parts of head almost
hairless. Froms not demarcated from clypeus or vertex,
sparsely setiferous, Antennal sockets Separated by 1.5
times diameter of a socket or by 0.7 times the length
of 2ad antennomere. Postantennal groove rather deen
and wide, the wall in front moderately prominent,
Postantennal bean-shaped area indistinct, weakly
Ss. AUST. VEUS. 19 (3); 19:97
Atay, FOSS
demarcated and not inflated. Vertex longitudinally
widely convex, more strongly so near collum;
transversely faintly concave, and laterally rather strongly
convex, but without inflated lateral edges. Vertigial
sulcus moderately impressed, running downward to
upper level of sockets, Vertex hairless, Antennae rather
long and slender, weakly clavate, with Sth and 6th
antennomeres thickest. Antennomeres subcylindrical,
but Sth-and 6th a little more abeonical; 6th antennomere
not inflated. Pubescence moderate in proximal
antennomeres, becomung rather dense in the distal ones,
Relative length of antennomeres 2 to 6; 0.95, 1.00, 0.95,
0.95, 0.90,
Collum: About as wide as head, subtrapezoidal in
dorsal outline. Anterior barder straight in middle, widely
rounded mote laterally and straight again towards lateral
sides. Posterior border widely and weakly concave,
laterally rather weakly convex. Lateral sides almost
evenly and rather narrowly rounded. Marginal rim
jaterally a little incrassate, not brimlike. Premarginal
furrow distinct, vanishing towards middle of anterior
border. Surface smooth, hairless, transversely widely and
evenly convex in middle to become samewhat more
strongly convex laterally; longitudinally widely canvex,
a lithe more strongly so near anterior border.
Sones: Rather weakly constricted. Prosomites dulled
by a fine cellular structure. Waist narrow, rather sharply
demarcated from pro- and metasomile, dorsally
distinctly beaded down. to level of paranota, weakly
siridlate along sides. Metatergites smooth, shiny,
hairless, Transverse furrow present from 5th to 17th
somite, weakly indicated on 4th and 18th somites.
Furrow father well impressed, with a vague sculpture,
disappearing laterally ai a distance fram dorsal furrow
of paranota about equal to dorsoventral diameter of a
poriferous paranotum. Sides smooth in general, but up
ta 4th somite somewhat granulose. Pleural keels up to
3rd somite represented by rather distinct curved erenulate
ridges, concavity upwards, without posterior lappet.
Pleural keels in 4th somite vestigial, in 5th absent. Sixth
somite with a weak posteriar swelling and 7th with a
low conical swelling near posterior margin of somite,
Paranota: 2nd somite a little wider than collum. in
dorsal aspect anterior border rounded and slightly
shouldered at base; latero-anterior edge narrowly
rounded, without distinct lateral tooth. Lateral border
widely and almost evenly rounded. Latero-posteriar edge
subangular, narrowly rounded, a little produced caudad
and projecting weakly behind margin of somite.
Posterior margin very short and a little concaye, In
lateral aspect upper side straight, sloping a Tittle in
anterior direction, Marginal rim rather thick, a little
callous; upper furrow distinct, also along anterior and
posterior margins. Third somite a little wider than 2nd
and as wide as 4th. Paranota of 3rd somite in dorsal
aspect widely and evenly rounded, a little more narrowly
rounded anteriorly. Latero-posterior edge angular and
SULT TPEDES PROM AUSTRALIA S 2l
a little produced caudad, but not projecting behind
margin of somite. Posterior border short, straight, In
lateral aspect upper margin straight, curving abruptly
dorsad anteriorly. Dorso-ventral width not much larger
than that of paranota of 2nd somite; ventral
demareation by a distinct impression, reaching forward
to about whree-fifths length of metasomite. Paranota of
4th somite rather similar to 3rd. In dorsal aspect more
widely curved than in 3rd somite, with posterior edge
subangular, not produced caudad, and posterior barderc
straight. In lateral aspect upper margin curved darsad
more widely, Dorso-ventral width as in 3rd somite, but
ventral demareation reaching cephalad to about halfway,
Paranata of Sih somite rather weakly developed and
rather weakly prominent. Margin in dorsal aspect widely
convex, Posterior edge subangular in porcless somites,
narrowly rounded in poriferous somites, In posterior
somites posicrior edges become minutely angular and
produced a little caudad from 14th somite onwards,
projecting only a Lt behind caudal margin of somite
in 17th and 18th somites. In lateral aspect dorsal
demarcation of paranota weakly concave anteriorly,
weakly convex posteriorly in poriferous somites, about
straight in poreless somites. Darsal furrow anteriorly
rather abruptly curving upwards, but not reaching waist.
Paranota dorsoventrally not wide, rather narrow,
especially in porcless somites. Ventral demarcation by
a depression reaching cephalad to about halfway or two-
fifths length of metasomite, Posterior edges in lateral
aspect acutely angular, especially in poreless somites, [n
poreless somites ventral demarcation a little coneave.
Sternites and legs: Sternites of middle somites longer
ihan wide (ratio 1.3:1.0). Cross impressions weil
developed; longitudinal impression rather wide;
transverse impression also rather wide, but narrow
between coxae. No sternal cones. Pubescence rather
dense, especially near bases of coxae; hairs of moderate
length. Sternite of 4th somite rather broad, rather
densely setiferous, and transversely rather widely
cancaye. Siernite of Sth somite with a broad pracess
arising between and slightly in front of anterior coxae;
at base the process is a little wider than distance between
coxae. Process short, projecting downward, widely
rounded. Anterior side a lille concave with dense brush
ol short setae, Posterior side faintly convex with some
long hairs. ‘Transverse impression deep, Posteriar part
of sternite excavate, but not down (0 level of metasomal
Tillis; Nol taised at base of coxae. A group of long setae
in the middle, Sternite of 6th somite deeply excavate;
posteriorly level with metasomal ring bul anteriorly a
little raised above level as in posterior pari of sternite
of Sth somite. Coxal bases not raised, but coxae of legs
cistinelly elonwate. ‘Transverse impression wide and
shallow. Four areas with long setae arranged mm a square.
Sternite of 7th somite with u low and rather narrow,
finely granular ridge lateroeephalad of gonopod
aperture, Sternite af 8th somite excavate, particularly in
anterior half, and raised only a little above ventral level
of metasomal ring. Transverse impression weakly
developed. Legs rather long and slender, prefemora
rather convex dorsally. Pubescence on ventral side rather
dense in all podomeres, Huirs moderately long, Tarsi
pubescent on all sides, Femora almost straight. Tibial
and tarsal scopulae present on anterior legs but soon
thinning out and absent from legs of 7th somite
onwards. First leg strongly incrassate, Coxa of 2nd leg
with a short medial rounded cone. Relative length of
podomeres 2 to 6 in middle somites: 0.65, 1.00, 0.60,
0.55, 0.75.
Anal somite: Dorsal profile straight or faintly conyex.
Sides of epiproct concavely converging, before apex a
shght indication of an abrupt stepwise narrowing, quite
near apex. Anex of epiproct with lateral edges narrowly
rounded and posterior margin weakly concave. Epiproct
broad at base, distally still rather broad. Dorsoventral
width moderate, length moderate. Setae not on tubercles.
Paraprocts with narrow, moderately high rims. Selac nat
on tubercles. Hypoproct large triangular, parabolically
rounded with sides convex and apex more narrowly
rounded. Setae not an tubercles.
FIG OL. Hererovudosome cebrani on sp., holotyped, rigit gonopod,
medial aspect.
Gonopods: (Fig, |) Coxa somewhat clangate, rather
stout at base, but narrowing distad, Setiferous area
rather large. Prefemur short, ovoid; its distal
demarcation transverse on axis of acropodite. Tibiotarsal
branches both well developed; caudal one lanceolate,
widening a litthe distad; anterior one stoyter, more
irregular in shape, and ending in an acuminate
somewhat uncate apex, Solenomerite well developed,
With permal channel running along medial side, widely
curved and apically narrowing to solenomerite proper,
which curves a little mesad. Femoral process arising
guile near apex of salenomerite, curving caudad, about
as long as solenomerite proper, apex acuminate,
Female: Sternites.as long as wide, Pubescence of legs
and sterna less conspicuous than in male. Legs shorter,
prelemora not incrassate, Relative length of podomeres
2 to 6 in middle somites: 0.80, 1.00, 0.50, 0.50, 0.85.
Head with vertex transversely more evenly convex, not
particularly flattened in middle and without lateral
swellings. Collum with anterior border much more
evenly convex and only a little more strongly rounded
laterally, Pleural keels of 2nd somite produced into a
long process; of 3rd somite in a long posterior cone;
in 4th somite only a minor cone near caudal margin.
Coxa of 2nd leg with a dagger-like process arising from
ecto-caudal side of apex, pointing Jatero-ventrad,
Epigynal structure consisting of two pararnedian wide
emarginations separated medially by a low conical
process painting cephalad.
Remarks
The three previously described species of
Aeteracladosama, viz A, bifaleatum (Silvestri, 1898), A.
lransversetaeniatum (L, Koch, 1867), and &.
hamuligerum (Verhoeff, (924), are all from the coastal
region of Queensland between Cairns and Brisbane.
Compared to H. zebratum and A. galaxias these three
species are differerit in that the largest tibiotarsal branch
projects a little distad of the seminiferous branch, 1,
hamuligerum is distinct in having the seminiferous
branch apically divided into three processes, a
soleriomerite and two, or possibly ane, deeply split
femoral branches. in A. transverselaeniatum and 4.
bifalcatun: the bifurcation of the seminiferous branch
into solenomerite and femoral process is situated closer
to the base of the branch, resulting in a relatively greater
length of the two distal processes. #H,
transversetaeniaiuim is particularly disunct in is
voluminous gonopod telopodite and in the apex of the
larger tibiotarsal branch which is strongly recurved
forming an elongate spinelike production. As yet ii is
not clear to which of the three Queensland species H.
rebratum and J. palaxias are most closely related, but
on account of the total configuration of the gonopads
these new species seem ta approach A. bifalcatum more
than the others.
Heterocladosoma galaxius n. sp.
Material
Gamman Plateau, South Australia, 19,1X.1956, lee.
G. F. Gross, ¢ holotype.
Description
Colours Head dark brown, with labral area, an area
around antennal sockets and sutures of lateral sclerites
22 RECS. AUST, MUS IN (3), 12-47
Mux [08s
yellowish brown. Antennae dark brown, with 6th and
7th antennomeres darkest; intersegmental membranes
yellowish, tip whitish. Collum blackish brown, with a
pair of semilunate yellowish white spots at caudal
margin, almost touching medially and continued
laterally im a yellowish sireak along caudal margin,
disappearing near latetal rounding, Somites blackish
brown, ventrally paler. Sternites and legs rather dark
brown, Dorsum marked by two series of paramedian
oval yellowish white spots, occupying the metatergites
from halfway between waist and transverse furrow, and
prosomites of the next somite to about halfway to its
waist. Lateral sides.a shade paler brown in their posterior
third. Anal samite dark, but ventral side including
hypoproct paler brownish; epiproct entirely yellowish
white,
Width; 3,3 mm.
Alead and antennae: As in the preceding species, but
differing in antennal sockets being a little closer to each
other, and separated by 1,35 times diameter of a socket,
or by 0.6 times length of 2nd antennomere. Antennae
with distal antennomeres a little shorter; relative length
of anternomeres 2 to 6; 0.95, 1.00, 0.95, 0.85, 0.75.
Callum: Differing only in being a litthe wider than
head.
Somites: Waist rather narrow, distinctly demarcated
from both pro- and metasomites, dorsally not headed,
but faintly longitudinally striate down to level of
stigmata. Pleural keels in 4th somite weakly. developed,
Paranota: Posierior edge of paranota of 2nd somite,
not caudally produced, posteriar border obsolete, In
lateral aspect the upper margin curyes a little upward
caudally, Posterior edge of paranata af 3rd somite
produced a little and projecting slightly caudad of
margin. Posterior edges of paranota of Sth and
subsequent somites narrowly rounded in dorsal aspect,
becoming subangular in 14th samite only, not produced
caudad. Vertral demarcation of paranota reaching
cephalad to about three fifths of length of metasomite,
Posterior edges in lateral aspect rather narrowly truncate
in pareless somites, more widely and more obliquely
truncate in poriferous somites.
Sternites and legs: Similar to those of the preceding
specics. Sternal cones are weakly indicated in sternites
of middie part al’ body. Sternite of Sth somite with
process directed downward and a little cephalad,
projecting scarcely in tront of the sternite. Process rather
short, its apex quite widely triangular, medially rounded.
In lateral aspect posterior side of process straight, with
a transverse concavity at its base, Legs rather long and
moderately stout. Relative length of podomeres 2 ta 6
in middle somites: 6,50, 1,00, 0.55, 0,55, 0.70.
Anal somite: Epipract almost parabolically rounded;
apex rather narrowly rounded, medially straight and not
emarginate,
MILLIPEDES FROM AUSTRALIA 5
FIG. 2. Helerovladasoma zaluxtus 1. sp., holotype 3, right gonopod,
medial aspect,
Gonapods; (Fig. 2) Largely similar to those of the
preceding species, but anterior tibiotarsal branch quite
different: narrow at base, gradually widening distad,
sharply crooked, and curving rather abruptly distad
again, terminal part rather voluminous and tapering
towards apex and somewhat uncate, Posterior tibiotarsal
process distally widening, then rather abruptly
narrowing and ending in a narrow spinelike process
(strongly reminiscent of a similar structure in
Streptocladasoma dissimile Jeekel (1980: &, Figs 8-9.)
Female: Unknown,
Remarks
Although this species is obviously closely related to
H, zehraium, it is easily distinguished by its very
distinctive colour pattern, by differences in the structure
of the gonopods, and some other features like the
different sculpture of the waist, the slightly stronger
development of the pleural keels, the less prominent
paranota, and the presence of weak sternal cones.
Somethus Chamberlin
Somethus Chamberlin, 1920; 651; Jeekel, 1968; 27;
Jeekel, 1979: 651.
Type-species: Somethus fuscipes Chamberlin, 1920.
vw
Remarks
This genus was erected by Chamberlin for the
reception of a single species from an unknown
Australian locality. Unfortunately, the original
description was not accompanied by a drawing of the
gonopod of the type-species, On that account the
placement in the tribe Australiosomatini was somewhat
arbitrary (Jeckel 1968), A re-examination of the type
material (Jeekel 1979) confirmed the placement.
However, with regard to the gonopod structure, doubt
still exists on the true identity of the species,
The discovery of some species of the genus in
northern Tasmania and Victoria, to be published
elsewhere, and the new species described hereunder has
proved that Somethus is a well-defined genus
characterized in particular by having a single tibiotarsal
branch, arising from near the base of the acropodite of
the gonopad, a small femoral process emanating from
the channel-bearing branch of the gonopod more or less
near jts apex, and a short solenomerite proper, haying
a triangular additional process, in which the spermal
channel makes a loop before running towards the apex
of the solenomerite.
The record of Somethus grossi n, sp. from South
Australia extends the distribution westward into the
moister parts of that State. Ii seems likely that the type-
species may be found sooner or later somewhere in
southeastern Australia also.
Somethus appears to be closely related on the one
hand to PhAyllocladosoma Jeekel, 1968, and
Dicladosomella Jeckel, 1982, in which the femoral
process is completely lost and the tibiotarsal branch
more broadly expanded, and on the other hand to
Oncocladosama n. gen., in which the femoral process
is vestigial or absent and the tibiotarsus is also more
voluminous and more or less clubshaped, The four
mentioned genera are furthermore each characterised
by a particular size and curvature of the solenomerite
proper.
Somethus grossi n. sp.
Material
Rankin Creek, near Melrose, South Australia,
13.V1,1954, leg. G. F Gross (E.S.1. 1355), & holotype,
1? paratype,
Same locality, date and collector (E.S.1. 1335), 3¢
paratypes.
Descriptian
Colour: Head brown, with labral area, an area around
the antennal sockets, and lateral sclerites of head
yellowish, Antennae yellowish to brown, with 6th and
basal part of 7th antennomere darkest, Collum brown,
with traces of a rather narrow, medially widest,
transverse band along posterior margin. Somiles alsa
brownish, with most of area behind the transverse
furrow, including posterior half of paranota, yellowish.
a RECS. AUST. MOS
Venter, sternites and Jegs yellowish. Anal somite brown,
but most of epiproact, and margins of parapracts, and
hypoproct yellowish. The material appears to have lost
much of its original colour and the brawn ecalour may
have been blackish brawn in living specimens.
Widihid + 3.23.4 mm,?: 3.7 mm,
Head and antennae: Labral emargination of moderate
width and depth. Clypeus very strongly impressed
towards the labrum, the impression erescentic in shape,
and surface above it a little inflated. Clypeus with
svtiferous pits, moderately densely pubescent; on each
side below antennal sockets a rather Weak impression.
Setae rather short. Lateral border of clypeus widely and
weakly convex, a notch near the labrum. Lateral sclerites
of head hairless, Frons not demarcated fram clypeus
or vertex, smooth, hairless. Antennal sockets separated
by 1.4 times diameter of a socket or by 0.7 times length
of the 2nd antennomere. Postantennal groove rather
deep and moderately wide, wall in lront rather
prominent. Bean-shaped area at posterior margin of
antennal sockets rather distinctly demarcated, slightly
inflated. Vertex almost evenly convex longitudinally,
slightly more convex near collum; almost flat or even
@ litle concave transversely, lateral edges rounded and.
somewhat inflated. Vertex smooth, shiny, hairless;
vertizial sulcus rather deeply impressed, not reaching
upper level of the antennal sockets. Antennae rather
long, Slender, hardly clavate. Antenomeres
subeylindrical: 6th slightly more obconical but nor
inflated. Pubescence moderate in proximal
antennomeres becoming dense in distal ones. Relative
length of antennomeres 2 to 6: 1.00, 0.95, 0.90, 0.80,
0.70.
Collum: A little wider than head, subtrapezoidal in
dorsal outline. Anterior border straight or very fairly
voneave, widely rounded more laterally and straight
again towards lateral sides. Posterior border widely
emarginale in middle, widely rourded towards lateral
sides. Lateral border moderately widely anc
symmetrically rounded. Surface transversely weakly
convex, laterally more strongly so, and even slightly
incurved at sides. Longitudinally surface evenly widely
convex. Lateral margin with a narrow and low rim:
premareinal furrow distinet, disappearing at level of the
lateral edge of the vertex. Surface smooth, shiny and
hairless,
Somites: Rather weakly consiricted. Prosomites dulled
by a fine cellular structure and with fine striae. Waist
narrow, rather distinetly demarcated trom pro- and
metasomites, dorsally finely but distinctly longitudinally
ribbed down to the level of the paranota, faantly striolale
below that level. Metatergites smooth or with some fine
wrinkles, shiny and hairless. Transverse furrow present
on Sth to léth somites, vaguely also on 17th samite.
furrow moderately impressed and with some vague
sculpturing, running laterad to a distance trom upper
margin of paranota equal to the dorsa-ventral diameter
paps W837 May, Twas
of a poriferous paranotum. Sides smooth, or somewhat
irregularly and finely wrinkled, shiny. Anterior sanqites
up to 4th with sides. a little Subgranularly uneyen. No
pleural keels.
Paranola; Ind somite a little wider than the collum.
Lateral border in dorsal outline straight anteriorly and
a litile diverging in caudal direction, and widely convex
pasteriorly. Latero-pasteniar edge narrowly rounded,
incurved se as to be slightly produced and projecting
a little behind margin of somite. Posterior border almost
obsolete, Anterior border moderately widely rounded,
latero-anterior edge narrowly rounded, wilh a small,
almost obsolete lateral toath, In lateral aspeet upper
margin widely and weakly concave dorsally, situated on
a low level, and therefore scarcely visible from above;
upper margin slightly sloping in anterior direction,
Paranotum dorsaventrally rather narrow, The upper
furrow distinc! along all margins including the antenor
and posterior. Third somite a little narrower than 2nd
and a little wider than 4th. Paranota in dorsal aspect
evenly rounded, anteriorly and posteriorly more
narrowly. Posterior edges particularly in 3rd somite
angular, in 3rd somile slightly produced caudad, but not
projecting behind margin of the somite; in 41h narrowly
rounded and not produced, Posterior border in 3rd
somite very short, almost obsolete, in 4th obsolete. In
lateral yspeet paranota of 3rd and 4th somites with a
widely concave upper margin, rather abruptly curving
dorsad anteriorly, posteriarly curving upwards and
shortly paralleling posterior margin of somite Darso-
ventral width rather narrow to moderate. Lower
demarcation formed by a depression reaching cephalad
to about two fifths of lenuth of metasorite, Posterior
edges of paranota in lateral aspect acuminate. Paranota
ol Sth and subsequent somites rather weakly prominent.
In dorsal aspect lateral margin of poriferous paranota
widely rounded, with a faint emargination from pore
area to caudal edge. Lateral rounding ol poreless
paranola a little wider. Posterior edge of paranota
narrowly younded, but in poreless somites and in
poriferous somites of caudal half of body posterior
edges become more angular and quite weakly produced
caudad, though nat projecting behind the margin.
Posterior border quite short, convey in porilerous ta a
little emarginate in poreless somites. In lateral aspect
upper margin is faintly concave aiiteriorly and a little
convex posteriorly in poriferous somiles, siraight or even
faintly concave im pareless somites. Dorsal furrow
curving a little upward anteriorly, not reaching waist but
running cephalad to about four-fifths length of
metasamite. Dorsal furrow caudally curving abruptly
dorsad and briefly paralleling posterior margin of
somite Ventral impression demarvating the paranola
visible up to about twa-lifths of length af niciasomite
in both poriferous and porcless somites, Dorsa-ventral
width of paranota rather narrow, the poreless only a
little narrower than the porilerous. Paranola oot callous,
Pores of moderate sive, situated at anterior cod of an
MITE IPE DES FROM AUSTRALIA 4
oval excavation. Posterior edges of paranota in lateral
aspect more or less acuminate.
Sternies and legs: Sternites of middle somites longer
than wide (ratio: 1,6:1.0). Cross-impressions strongly
developed, with deep longitudinal and transverse
impressions. At bases of coxae rather large rounded
conical protuberances projecting downward and a little
caudad, distinet particularly at anterior pair of coxae
and especially in 9th, 10th and th somites, becoming
less distinct in posterior somites. Pubescence of sternites
dense, with setae ol moderate length, Sternite of 4th
samile rather widely and moderately deeply excavate,
moderately setiferous, with long setae. Sternite of 5th
somite with 4 low hump between anterior legs, which
is produced into a large shovel-like process a little in
front of coxal bases. This process is directed cephalad
at base, crooked halfway so as to become directed more
ventrad distally, Process projecting distinctly in front of
sternite, Posterior surface in lateral aspect corvex at base,
more distally concave; anterior surface convex in profile,
apically provided with a dense brush of short setae,
Process a little broader than distance between anterior
coxae, distally quite broadly rounded, subangular in the
middle. Posterior surface rather weakly pubescent with
long setae. Transverse impression distinct and deep.
Pastenior part of sternite raised as a transverse, rounded
wall, Without longiludinal impression, densely set with
long setae. Sternite of 6th somite not raised above ventral
level of metasomal ring, deeply excavate, Coxal bases
scarcely raised, Pubescence toderate, with long hairs.
‘Transverse impression weak. Sternite of 7th somite with
a large gonopod aperture pressing the ambulatory legs
a little laterad. Latero-cephalad of aperture a rather low
callous wall, Sternite of 8th somite anteriorly widely
concave and only slightly raised above ventral level of
metasomal ring. Anterior coxal bases not raised,
distinctly more separated than posterior. Transyerse
impression weak. Pubescence dense, with long setac.
Legs of moderate length, rather stout. Prefemur dorsally
moderately convex, femora faintly arched. Pubescence
dense on ventral side of all podomeres and on all sides
of the tarsi and the anterior tibiae, remaining surlaces
without apparent pubescence. Hairs rather Jong,
Scopulae present on tibiae and tarsi of all legs up to
6th somite, absent on all postgonopodal legs. Relative
length of podomeres 2 to 6 in middle somites: 0.60, 1.00,
0.65, 0.55, 0.75, First leg strongly incrassate, with the
usual ventral femoral tubercle. Coxae of 2nd legs ventro-
apically thickly rounded, bul not projecting.
Anal somite: Upper profile faintly convex, almost
straight. Surface somewhat wrinkled. Epiproct of
moderate length and thickness, slightly concave on
ventral side, broad at base with sides concavely
converging, more distally straight and convex towards
apex. Near apex a slight stepwise narrowing. Apex
truncate and a little emarginate, lateral edges narrowly
rounded, Setae not on tubereles. Valves with rather
tw
ay
narrow and moderately high rims, Setae on minute
tubercles, Hypoproct large and triangular, parabolically
rounded with sides widely and apex more narrowly
rounded, Setae on small tubercles which do not project
outside margin.
Va
AA Veh
SS}
\\
TG, 3, Semeihus grossi io, sp. holotype ?, right gonopod, medial
aKpeel.
Gonopods: (Fig. 3) Coxa long and stout, tapering a
little towards apex and curving a little caudad. At medio-
anterior side of apex a small conical process, Prefemur
rather shart, rounded, its longitudinal axis transverse
to axis of acropodite, Demarcation from acropodite
transverse. Acropodite split into two main branches: a
caudal tibiotarsus, and a combined solenomerite and
femoral process, Tibiotarsus long, narrow at its base but
widening halfway to a laminate apical part. Combined
solenomerite and femoral process split at about three
quarters of length into a solenomerite proper and
separate [emoral process. Latter tapers apically and
curves caudad. Spermal channel runs along anterior side
of acropodite, and finally along medial side of
a REO
solenomerite proper, making a loop in a triangular
preapical process. Apex of salenomerite narrow, point-
ing caudad.
Female: Dilfering from the niale, aside from the usual
sexual characters, in having vertex of head Lransversely
widely and evenly convex. Vertigial sulcus quite deeply’
impressed. Antennae shorter, with the antennomeres.
more obconical, Relative length of antermomeres 2 to
6: 1,00, 0.95, 0.95, 0.95, 0.90. Third somite with a low
transverse pleural keel along posterior margin. Sternites
flattened, impressions distinctly less. deep than in the
male, langitudinal ones very wide, Cones absent, Ratio
of length/width: 1.1:1.0. Pubescence rather dense, hairs
shorter than in male, Legs. with straight femora, Relative
length of pocdomeres 2 to 6 in middle somites: 0.80, 1.00,
0.55, 0.55, 0.70. Coxa of 2nd pair of levs distally
produced into a large caudal transverse, callous process,
giving coxa from behind a broad Iriangular aspeet, as
broad at base as long medially, and basally produced
laterad of prelemoral basis. Epigynal structure
consisting of two paramedian rather narrow
emarginations embracing coxal bases of 2nd pair of legs,
and medially separated by a triangular prominence,
Remarks
The new species is easily distinguished from the type-
species, S. juscipes, by the gonopods having a long,
distally widening tibiotarsus (Chamberlin (1920)
describes the “basal spur"! as a "thin lanceolate blade
narrowed at cach end, a little twisted . . ’). the absence
of pleural keels in the male, the presence of sternal cones
in the male, and the larger size (in S. /uscipes. 2.8 mm).
The species is named after its collector, apparently
one af the very few people in South Australia who took
the trouble to collect millipedes, including most of the
material treated in this paper.
Oncocladosoma 0, gen,
Generic diagnosis
Rather robust to small Australiosomatini with 20
somites and a normal pore formula. Head with vertex
transversely flattened in male, normally convex in
feinale, Antennae of moderate length, slightly to
distinctly clavate, basal antennomeres subcylindrical,
distal anes more obconical in shape,
Somites rather weakly to moderately consiricted,
waist tather narrow, distinetly longitudinally ribbed or
beaded. Metatergites smooth, hairless, Transverse furrow
present from Sth somite onwards, rather weakly to rather
deeply impressed. Pleural keels vestigial or absent in
male, moderately developed up to 4th somite in female.
Paranota rather weakly developed.
Sternites distinctly longer than wide in male, about
as long as wide or 4 little longer than wide in female.
Slernai cones weakly developed to vestigial. Sternite of
Sth samite of male with a process between anferior legs.
h AGE. MILL 19 (3)2 1-87
May, LOSS
Legs rather long; first leg of male incrassate and with
a ventral femoral (ubercle, Tibial and tarsal scapulae
present only in 4 number of legs in (he anterior half
of the body of male.
Gonopods with prefemur ovaid, its longitudinal axis
almost transverse on the axis of the acropodite,
Actopadile deeply split into two main branches, femaral
part almost vestigial. Tibiotarsus undivided, more or less
clubshaped, with a rather narrow “'stem" and a widened,
more or less complicated distal half, Seminiferous
branch exceeding tibiotarsus in length, end typically
uncate and recurved. Spermal channel running along
posterior side ol seminiferous branch, turning distally
towards medial side, aid runaing lo extreme distal part
of the seminiferous branch before abruptly recurving
towards apex, Just proximad of distal edge of
semuniferous branch and cephalad af course of spermal
channel there is a vestige af the femoral process, which
is completely absent, hawever, in the type-species. A little
more proximad on mediocaudal side af serniniferous
branch there is in the type-species a short spinelike
process, vestigial or absent in the other species,
Type-species: Australiosoma castaneum Atierms, 1944,
Remarks
This genus is characterized mainly by the peculiar
shape of the acropodile of the gonopods, with its single
clubshaped tibiotarsus, its typically uncate seminiferous
branch, the absence or almost camplete reductian of
the femoral process, and the course of the spermal
channel, which makes a slight Joop before running.
towards the apex of the solenomerite.
fn haying the ganopod telapadite deeply split into two
main branches, Queocladosoma bears some similarity
to the genera Dicladosoma Brolemann, 1913,
Dicladosomella Jeekel, 1982, Phyllocladosoma Icekel,
1968, and Somerhus Chamberlin, 1920. Dicladosama,
trom Mt. Kosciusko, is distinct in having the base of
the tibiotarsus twisted towards the lalero-anterior side
of the seminiferous. branch, It moreover seems distinct
in) haying apparently no loop in the course of ihe
spermal channel. Prcludosamella, Ph\llocludosomea and
Soniethus have a more erect telopadite, and the spermal
channel makes a loop in a distinctly developed process.
Dicladosomella, from southeastern New South Wales,
hag a more laminate tibiotarsus, and the apex of the
solenomerite is typically directed dijstad
Phyllocladosoma, from northeastern New South Wales
and southeastern Queensland, resembles Dicladasemella
in most of the characters mentioned, but it ts
distinguished by the abbreviare solenomerite, the apex
of which extends searcely beyond the loop-bearing
process, Sovnethus is distinguished by having a distinctly
developed femoral process, Both, Dicladosomella and
Phylloctadosama, lack any indication of a femoral
process.
The pew genus contains, besides its type-species,
which is divided into two subspecies, two new species.
MILLIPEDES FROM AUSTRALIA §&
Geographically it seems to be confined to the Mt. Lofty
Ranges,
Oncocladasama castaneum (Attems)
Australiosoma castaneum Attems, 1944; 249; Jeekel,
1968: 26.
This species is represented by two subspecies, O &.
castaneum and QO. c. ingens n-subsp., characterized by
a significant difference in size
Material
Qc. castaneum (Attems):
Mt. Lofty, 21.PX1883, leg. Tepper, 1 @ , 19.
Belair, 15V.1938, lex, R, V. Southeott, 23 .
Belair, Mt, Lofty Ranges, leg. N. B. Tindale, }o .
Oc. ingens n subsp.
Near Mt. Lofty Station, 1V.1883, Dr. Haacke don., ¢
holotype, 23.2, 112 , 6 juve (19 somites), 2 juv, 2
{19 somites) paratypes,
Bridgewater, 2.111884, leg. Tepper, 1d paratype.
Norton Summit, Mr, Lofty Ranges, 7,1V.1884, leg.
Tepper, 2d paratypes,
Mt, Lofty, Waterfall Gully, 23V 11884, les, Tepper, |
paratype.
Onkaparinga River, near Mylor, 11.1947, les. G, F
Gross, 2¢° paratypes,
Mylor, 20.FV.1957, leg. G. EF. Grass, 1 o° paratype,
Upper Sturt, Soil Tx L04, 5.1962, leg R. V. Southeott,
Lé paratype
Bridgewater, Lor no, 11-72a, IX.J972, leg. G. H, Baker,
1d ,39 paratypes.
Without locality or other data, 9¢ , 52 , 1 juv 2
somites) paratypes.
Without locality or other data, 2v , 2 2 paratypes.
(19
Description
Colour; Head brown, vertex and lateral sclerites
blackish. Area around the antennal sackets and margins
of lateral scleriies pale brownish. Antennae brown, 6th
antennomere and basal part of 7th infuscate; tip whitish;
the intersegmental membranes pale brownish. Collum
blackish, margin behind vertex reddish brown; a rather
narrow zone along posterior margin, broadest medially
and tapering towards sides, brownish yellow, Prosamites
and anterior part of metasomites blackish brown,
posterior pact of metatergites, ic. medially from just in
front of transverse furrow caudad, yellowish brown,
Posterior part of paranota and posterior zone of sides
also yellowish brown. Demarcation between the darker
and lighter colours not sharp. Venter, sterrites and legs
brownish, three distal podomeres infuscate, inter-
segmental membranes and tip of tarsi yellowish brown,
Anal somite dorsally, including epiproct, yellowish, sides
blackish. Paraprocts black, margined with yellowish
brown. Hypoproct yellowish brawn. More heavily
infuscated specimens have pale colour only along
posterior margin of metasomites, Specimens which have
te
vel
been stored in alcohol for a long time are brown in
general, with the lighter colour only vaguely indicated.
Juveniles. dirty whitish, without colour pattern.
Width: O. c, castaneum; ¢ + 2,8-3,2.mm, ¢ +: 3,0 mm,
O. ¢. ingens: &: 3.5-4.3 mm, ¢: 3.4-4.0 mm, juy ¢ U9
§.)) 2.9-3.2 mm, juv 2 (19 sj) 2.7-2,9 mm.
Head and antennae: Labrum moderately widely and
moderately deeply emarginate, Clypeus strongly
impressed towards labrum, surface uneven due to
presence of setiferous pits: on each side below antennal
sockets wide impression. Lateral barder of clypeus
faintly convex, with a notch near labrum. Pubescence
moderate becoming sparse in frontal region and on
lateral sclerites; above labrum a series of hairs, hairs
rather short. Frons not demureated from vertex or
clypeus. Antennal sockets separated by 14 times
diameter of a socket or by 0.7 times length of 2nd
anteonomere. Vertex shiny, hairless, transversely faintly
convex, a little more sa near lateral edges, longiudi-
nally almost evenly and rather widely conyex.
Postantennal groove rather deep and rather wide; wall
moderately prominent. Beanshaped area at posterior
side of antennal sockers well demarcated and inflated.
Vertigial sulcus well impressed, not reaching upper level
of antennal sockets, with short fine transverse wrinkles.
Antennae of moderate length, moderately stout,
slightly clavate, Antennomeres sabcylindrical, bul Sth
more obconical, and 6th still more obconical but not
inflated, Pubescenee moderate in proximal anten-
nomeres, becoming dense in distal ones. Relative length
of 2nd lo 6th antennomeres: 0.95, 1.00, 0.95, 0.85, 0.70,
Collunu Subtrapezoidal in dorsal outline, a little wider
than head. Anterior border straight in middle, widely
rounded more laterally and straight again towards lateral
sides. Posterior border widely and weakly concave in
middle, straight laterally. Lateral sides. asymmetrically
and rather widely rounded, with strongest convexity
caadal. Surface hairless, shiny and almost polished, with
weak irregular wrinkles. Surface transversely widely
conyex, more strongly so towards lateral sides and
laterally even slightly incurved; longitudinally weakly
convex, slightly more so towards anterior and posterior
margins, Marginal rim laterally moderately wide, nat
particularly raised; premarginal furrow distinct,
disappearing at lateral edge of vertex.
Somes: Constriction rather weak, Waist rather
narrow, sharply demarcated from. pro- and metasamites,
distinctly longitudinally ribbed down to upper Jevel of
paranota, finely striate below that level. Prosomites dull,
with a fine cellular structure and fine short. striae.
Metatergites smooth, hairless, shiny, with some irregular
weak wrinkles, Transverse furrow finely and not deeply
impressed, without apparent sculpluring, present from
Sth to L7th somite, vaguely indicated on 18th, Furraw
running laterad ro about a distance from upper
demarcation of paranota of about one and a half times
as EC SL AUST. MUS, 10 ta) 1937
ora little more the dorsoventral width of a poriferous
Paranotum,. Sides smooth or slightly wrinkled, granulate
up to 4th somite, Pleural keels abortive or absent.
Puranota 2nd somite a little wider than collum; 3rd
somite a little wider than 2nd and about as wide as 4th,
Paranota of 2nd samite with anterior margin a little
thrust torward, widely convex. Latero-anterior edge
subangular, narrowly rounded, without distinct tooth.
Lateral margin Widely and almost evenly rounded, but
behind middle a lillie more strangly convex and caudally
straight, hardly diverging. Latero-posterior edge
subangular, slightly produced and projecting slightly
behind posterior margin of somite. Posterior margin
short, widely canvex. Paranota situated ona low level,
rather weakly prominent, although margin visible from
above. Upper margin in lateral aspect widely and weakly
convea, Sloping a little cephalad, Marginal callus rather
wide from the middle onwards, tlarrower ariteriorly, its
lower side siraight. Premarginal furrow distinct on all
sides, briefly parallelling the poslenor margin of the
somite. Paranata of 3rd somite with anterior margiu not
shouldered at base, widely convex, merging via a
stronger rounding into the widely convex lateral margin.
Posterior edge subangulat, slightly produced caudad,
Posterior margin short, a little concave. In lateral aspect
marginal callus moderately wide; ventral demarcation
by a depression is present only in posterior half,
converging dorsad somewhat convexly and meeting
dorsal demarcation in an acute angle. Paranota of 4th
samite similar to those of 3rd, but lateral margin a Jntle
more widely rounded, and posterior edge caudally not
produced. Paranota of Sth and subsequent somites
cather weakly prominent, Lateral margin in dorsal aspect
weakly ta faintly convex, anteriorly slightly more
rounded; poriferous paranota caudally slightly more
convex than the poreless. Latero-posterior edge
subangular, quite narrowly rounded, faintly produced
caudad in most somites, more so in the somites of
posteriar half of the bady, but projecting behind the
caudal margin of the somite only in the 16th, In lateral
aspect upper demarcation of the callus convex
(poriferous somites) or straight (poreless somites),
premarginal furrow turning upward near waist buc not
reaching it. Ventral demarcation by a depression present
only in the caudal half or three-fifths of the paranola,
converging straight (poreless) ar convexly (poriteraus
somites) towards the dorsal demarcation in ari acute
angle, more acute in pareless than in poriferous somites.
Dorsoventral width of paranota moderate, in poreless
somites about two-thirds of poriferous, Posterior margin
finely rimmed, the premarginal furrow briefly
parallelling caudal! margin of metatergite. Pores situated
quite neat the ventral edge of the rnarginal callus, in
a father srriall oval pit.
Sternites and fegs: Sternites of middle somites longer
than wide (ratio §,6:1.0). Cross impressions well
developed, rather wide. Sternal cones. present on all
Nhat, 1485
postganopodial sternites up to [7th somite, rather small
near anterior coxae, and still more weakly developed
near posterior coxae, Pubescence rather dense to dense,
loeated in four quadrants of each stermte. Median and
iransyerse impressions hairless; hairs of moderate lenzth,
Sternile af 4th somite rather broad, with a well-
developed median impression, no sternal canes,
Pubeseence rather dense. Sternite of Sth somite with a
subtrapezoidal process arising between and a liltle in
(ront of anterior coxal sockets; process broader than
long, a little wider than width between coxae, directed
downward and cephalad, bending abruprly downward
in distal hall, projecting distinctly in front of anterior
margin of sternite. Anterior surface of process in profile
convex hallway, apex with a dense brush of short setae.
Pasterior surface widely concaye, moderately densely set
with rather long selae. Apex of process in caudal aspect
broadly rounded, medially fainily angulate, lateral edges
narrowly rounded, Behind process a deep transverse
Impression, No longitudinal impression between
posterior coxae, but posterior coxal sockets connected
by a transverse wall which is densely set with long setae.
Sternite of 6th somite Ssearcely (anteriorly) or not
(posteriorly) raised above ventral level of metasamal
ring. Coxal sockets searcely raised and coxae nor
elongate. Pubescence consisting of two transyerse zones
of densely set long hairs. Sternite of 7th somite
lateroanteriorly with a finely rugulose-granulose
transverse wall. Gonopod aperture large, separating
coxal sockets of posteriar legs, Sternite af Sth somite
anteriorly not raised aboye ventral level of metasomal
ring; anterior coxae sommewlial separated. Transverse
impression weakly developed. Posterior part not
modified, cones abortive. Pubescence moderate, hairs
rather long. Legs in general rather long and stout,
prefemora dorsally convex, femora faintly arched.
Pubescence ventrally dense on all podomeres, hairs
rather long. Dorsal pubescence nor obvious, except on
tibiae and tarsi and partly also on postfemora. Scopulac
of tibiae and tarsi present up to anterior legs of 8th
somile, totally absent from posterior legs of that somite
onwards, Relative length of podameres 2 to 6; 0.80, 1.00),
0.65, 0.60, 0.80. Legs of first pair strongly incrassate,
with a ventral fernoral process. Coxae of 2nd pair
medially rather thickly rounded but only weakly
produced distad.
Anal somite; Upper profile about straight or taintly
convex, Epiproct broad, moderately thick, rather short,
ventrally samewhat concave, Sides concavely converging,
narrowing abruptly just before apex, which is truncate,
caudally weakly emarginate, lateral edges parrowly
rounded. Setae not on tubercles, Paraprocts with narrow
and rather low rims; setae not on tubercles or latrer
abortive. Hypoproct parabolically rounded, setae on
abortive tabercles.
Gonopods: (Pigs 4-5) Characterized mainly by the
shape ef the tibiotarsus, Moreover there is no trace of
MILLIPLDES FROM AUSTRALIA 5 39
ess
FIGS, 4-5. Oncoeladasama castaneuin ingens n. subsp., holotype d. 4: right gonopod, medial aspect. 5: telopodite of left gonopod, anterior aspect.
a femoral process. On the other hand the species has
a spinelike process arising from the medio-caudal side
of the seminiferous branch a little proximad of the base
of the distal hook. The morphological status of this
process is not yet clarified.
Female: Head with the antennal sockets separated by
1.4 times diameter of a socket or by 0.8 times length
of 2nd antennomere. Vertigial sulcus slightly less
impressed, vertex transversely widely and evenly conven.
Relative length of antennomeres 2 to 6: 1.00, 0.95, 0.90,
0.85, 0.80. Somite 2 with pleural keels represented by
a strongly developed rounded ridge. A thick dorsally
sharply demarcated rounded swelling in 3rd somite, in
4th an abortive swelling. Sternites about as wide as long.
The cross-impressions much weaker, without sternal
cones. Setation rather dense, but setae rather short. Legs
with ventral pubescence rather dense, hairs of moderate
length. Relative length of podomeres 2 to 6: 0.70, 1.00,
0.50, 0.45, 0.80. Coxa of 2nd pair of legs with a caudo-
lateral callous process, pointed and directed caudo-
laterad, medially not produced. Epigynal structure
consisting only of two emarginations of ventral side of
3rd somite, embracing coxae of 2nd pair of legs, and
medially separated by a low triangular process pointing
cephalad. Lateral border of emarginations raised.
Remarks
When we compare the present drawings of the
gonopods with the one which Attems (1944) published,
there can be but little doubt that the presently recorded
material belongs to the same species, It will be noted,
however, that there are important discrepancies between
the drawings. In the first place Attems misidentified the
small process at the medio-caudal side of the
seminiferous branch as a side branch of the tibiotarsus
(the latter erroneously called ‘“Femoralfortsatz”).
3 RIC. S. AUST, MUS, 19.43)) 19-37
Furthermore, he illustrated a separate femoral process
(erroneously indicated as “Tibiotarsus”), emanating
distinctly proximad of the distal edge of the seminiferous
branch. Such a process js not observed in the present
material, and we must assume thar Atlems either
misinterpreted the chilitious structures in his preparation
or that the gonopod he examined was damaged.
The subspecies ingens appears. to differ from the
nominate subspecies mainly in its larger size, Differences
in the details of the gortopods must be substantiared
by future research when mote material becomes
available, Whereas the subspecies castaneuii appears 10
be confined to Belair (unfortunately Attems gave only
“Sudaustralien” as type lacality), the subspecies ingens
occurs in a much larger area of the Mt, Lofty Ranges.
Oncocladesoma conigerum n_ sp,
Material
Without locality label, but found in a tube together with
OQ. castaneum ingens: ¢@ holotype, 4 ¢ paratypes.
Near Mt. Lofty Station, (V.1883, Dr Haacke don., | ¢
Paratype.
Without locality label, 6 # paratypes, 2 » paratypes.
Description
Colour: Essentially the same as that ef the preceding
Species, but less infuscate (possibly due to prolonged
preservation in alcohol). Dark colour of head confined
to vertex and frontal region; lateral sclerites of the head
pale brownish. Antennae brawn, joints paler, Collum
and somites as jn castaneym but pale and dark colours
less contrasting. Distal podomeres scarcely infuscate.
Width; # + 2.1-2,4 mm; ? : 2.3-2.6 mm.
Head and antennae Lateral border af clypeus with
notch indistinct, Antennal sockets separated by 1.3 times
diameter of a socket or by 0,55 times length of 2nd
antennomere, Postantennal bean-shaped area a little
inflated, moderately demareated, Antennae rather stout,
Relative length of anternomeres 2 to 6: 1.00, 0.95, 0.80,
0,80, 0.75,
Collum: A little narrower than head, Lateral sides
almost evenly rounded,
Somiles; Waist distinctly beaded dorsally. Transverse
furrow disappearing laterally at a distance fram dorsal
demarcation of paranota equal to dorso-ventral diameter
of a poriferous paranotum. Pleural keels weakly
developed up to 3rd somite, faintly indicated in 4th,
Paranota: 3rd somite about as wide as 2nd; 4th a little
wider chan 3rd. Paranota of 2nd somite with marginal
callus of equal width. Paranota of 3rd samite without
anterior edge, lalero-anterior border rather strongly
rounded, In lateral aspect upper demarcation of
paranota of 3rd and 4th somites slightly concave, turning
upward anteriorly and posteriorly; the upper
demarcation not sloping caudad. Paranota of 5th and
subsequent somites with posterior edge narrowly
Mut, 1985
rounded, scarcely produced except very shghtly so in
posterior somites but not projecting behind posteriar
margin of somites, In lateral aspect veritral demarcation
Of paranota more strongly convex, meeting upper
demarcation in a wider angle than in castaneum, giving
posterior cdge of paranota a more subiruncate
appeararice,
Sternites and legs: Sternites af middle samites longer
than wide (ratio 1,65: 1.00), Process of sternite of Sth
somite with process about as long as wide at base, about
parabolical in posterior aspect. Posterior half of sternite
with a median rather deep furraw; pubescence rather
dense, setae longish. Sternite of 6th somite with a
distinct transverse furrow, but without Jongitudinal
impression, Sternite of 7th somite with a weak callus
latero-cephalad of gonopod aperture Legs with
scapulae up to first pair of 8th somite as in preceding
species. Pubescence on ventral side of legs long. Relative
length of podomeres 2 ta 6 in middle pan of body: 0.63,
1.00, 0.60, 0.55, 0.80,
Anal somite: Upper profile faintly convex, weakly
convex in Front of base of epiproct. Epiproct longish,
broad. Caudal emargination very weak. Hypoproct
rather large, about evenly semicircularly rounded,
Gonopods: (Figs. 6-7) Coxa relatively a little more
robust than in ©. cusfaneum. Prefemur with a weakly
chitinized conical process emanating from
intersegmental membrane connecting coxa arid
prefemur. Acropodite largely similar to that of @
castaneum, differing mainly in shape of tibiotarsus.
Seminiferous branch with a slight vestige of a femoral
process, The larger spinelike process of OQ. casfanetin
is represented here by a tiny toorh
Female: Head with clypeus moderately impressed
towards labrum. Antennal sockets separated by 1,3 times
diameter of a socket or by 0.7 times length of 2nd
antennomere. Antennae relatively shorter, relative length
of antennomeres 2 to é& 1.00, 0.90, 0.85, 0.90, 0.75.
Collum more evenly rounded transversely. Somites with
pleural keels on 3rd somite represerited by. a low cone
near caudal margin. On 4th somite a faint ridge
Sternites of middle samites 1.2 times longer than wide.
Legs less incrassate, femora straight. Relative length of
podomeres 2 to 6 in middle part of body: 6.70, 1.00,
0.55, 0.50, 0.70. Epigynal structure with paramedian
emarginations deeper than in castaneu, surface of the
ventral side of 3rd somite anteriorly raised, and median
conical process directed more downward,
Reniarks
In the characters not mentioried above, the description
of O. castaneun? applies.
This species looks like a diminutive form of O
casfaneum, being even smaller than the subadult
specimens of that species, 11 differs only in small
proportional details of jts external morphology, but it
is well characterized by the structure of the gonopods.
MILLIPE DES FROM AUSTRALIA 5 3
in
\
FIGS, 6-7, Oneoctadosama conigerum . sp., holotyped' -
Although most of the material was lacking a locality
label, the fact that part of it was found in a tube
containing O. castaneum ingens seems to imply that it
occurs sympatrically with that species in the Mt. Lofty
Ranges.
Oncocladosoma clavigerum n. sp.
Material
Belair, V1.1883, leg. J. W. Haacke, o holotype, 15 ¢,
8? , 2 juv.s (19 somites), 1 juv. (19 somites)
paratypes.
Blackwood, V1.1883, leg. J. W. Haacke, 2¢, 1¢
paratypes.
Woodley’s Vineyard (near Glen Osmond, SE Adelaide),
soil, Berlese funnel, 30VII-8.VIII.1950, leg. R. V.
Southcott, 2 .¢ paratypes.
Sellick’s Hill, 23.0X.1954 (E.S.1, 1592), leg. G. F. Gross,
ld.
"
EON
6: oght gonopod, medial aspect, 7: left gonopod, anterior aspect.
Descriplion
Colour: Similar to that of the two preceding species,
but less pronounced possibly due to preservation. Dark
colour of head confined to vertex and frontal regions;
Jateral sclerites of head pale brownish. Antennae dark
brown, distal annuli of antennomeres and
intersegmental membranes pale brownish. Collum
brown, without distinctly paler marginal areas. Somites
brown, darkest in waist area, without paler bands; only
posterior halyes of paranota a somewhat paler brown,
and posterior halves of metatergites also slightly paler.
Anal somite also with a paler epiproct, and margins
vaguely paler. Hypoproct pale brown. Sternites and legs
pale brown; three distal podomeres somewhat infuscate,
but no pale annuli. Tip of tarsi pale.
Width: ¢ + 1.51.8 mm,? : 2.0-2.3 mm, juv. ¢ (19
somites) 1.4 mm, juv.? (19 somites) 1.4 mm. The
43
s- REC
matenal from Belair seems Slightly smaller (4; 1,5-1,8
mum, ? > 2.0-2.2 mm) than the specimens from elsewhere,
Which measure 1.8 mm and 2,3 mm for the lwo sexes
respectively,
flead und anrennaes Labrom rather widely
emarginate. Clypeus rather strongly impressed towards
labrum. Antennal sockeis separated by 1,6 limes
diameter of a socket or by 0.85 times length of 2nd
antenomere, Anlennae rather stout, distinctly clavate
(involving 5th and especially 6th antennomere), 6th
antennomere a little inflated. Relative length of
anrennomeres 2.10 6: 1.00, 1.00, 0.95, 0.95, 0.90.
Collum: Distinctly narrower than bead, subreniform
to subtrapezoidal in dorsal outline. Anterior border
almost evenly rounded, only slightly more convex behind
edges of vertex, Posterior border laterally widely convex,
Lateral sides rather narrowly and almost evenly rounded.
Surface af collum Jongitudinally evenly and widely
convex.
Somiies; Canstriction maderate. Waist dorsally
distinctly beaded, laterally distinctly to finely striate.
Prosomites dull, with rather pronounced fine cellular
structure, Transverse furrow of metatergites sharply and
rather deeply impressed, with fine longitudinal striation,
disappearing laterally at about a distance rom dorsal
demarcation of paranota equal to dorsa-venteal diameter
of a poreless paranotum. Transverse furrow indicated
on 4th, and very vaguely also on 18th somite. Pleural
keels an 3rd somite weakly indicated by a furrow;
abortive on 4th somite.
Paranota: 2nd somite searcely wider than collum.
Paranota of 2nd somite with latero-anterior edge
narrowly rounded, without tooth, Lateral margin only
faintly convex, a lithe more so towards pasterior edge,
searcely visible from above. In lateral aspect margin
almost straight, moderately thick and of equal width,
Paranoia of 3rd samite lalero-anteriorly evenly and
rather widely convex. Posterior edge angular, slightly’
produced caudad bul not projecting behind margin of
the somite. Paranota of 4th somite with posterior edge
faintly produced. In lateral aspect dorsal demarcation
of paranota of 3rd and 4th somites faintly cancave,
turning upward gradually at anterior end. Paranota of
Sth and subsequent socutes with ventral demarcation
rather convexly converging with upper demarcation,
giving paranola, particularly the poriferous. ones, an
obliquely truncate aspect in lateral view. Caudal edges
not projecting behind margin of samites. Pores in a
relatively large and deep pil, nearer ventral margin,
rather near posterior edge.
Sternites and legs; Sternites of middle somites
longer than wide (ratio 1.6: 1,0). Sternal cones not
obvious, quite weakly developed Pubescence moderately
dense, all aver sternite, the setae of moderate lengih.
Sternite of Sth somite with the process between the
anterior legs subtriangular, the apex rather widely
S AUST MES TY pa): 1847
ND ub, PONS
rounded, about as long as wide. Anterior side in profile
weakly convex, the process almost perpendicular,
scarcely projecting in front of sternite, Caudal half of
sterhite widely transversely concave, rather densely: set
with longish setae. Sternite of 6th somite with coxal
sockets conspicuously more widely separated. Sternite
of 7th somite with pregonopadial wall weakly
prominent, Sternite of 8th sornite with anterior coxae
widely separated. Sternile anteriarly flattened, a tittle
raised, Transverse impression and posterior hall of
sternite not modified, Pubescence moderate, with setae
longish. Legs of moderate length or longish, rather
stout. Relative Iength of podomeres 2 ta 6 in middle
somites: 0.75, 1.00, 0.65, 0.70, 0.85, Coxae of 2nd pair
not fnedially produced, Coxae of first pair of legs of
6th samite and of anterior pair af &th somite slightly
produced tnto.a weak nmounded enone. Ventral pubescence
of legs rather dense,
Anal somite; Upper profile straight, slightly convex
anteriorly. Apex of epiproct relatively narrowly truncate,
scarcely emtarginate, Hypoproct silbtrapezoidal, with
median part of apex somewhat triangularly produced;
setae on faint tubercles which are faintly produced.
Gonopvuds: (Figs 8-9) Coxa relatively slightly mare
slender than in the ather two species. Tibiotarsus
typically club-shaped, with a relatively long stem.
Seminiferous branch characterized by (he rather strong
curvature; femoral process reduced io a tiny lobe The
More proximal spine, present in Q castanewn and
vestigial in QO. conigerum, is totally absent here.
Female: Head with anterinal sockets separated hy 1.5
times. diameter o| a socket or by 0.9 umes length af 2nd
antennomere, Relative length of antenniomeres 2 10 &:
1.00, 0.90, 0.85, 0.90, 0.85. Pleural keels of 2nd somite
well developed, those of 3rd somite represented by a low
conical swelling neur posterior margin; in 4th somite this
swelling is very weak and small, Keels are dorsally not
demarcated by a furrow. Sternites of middle somites 1.1
times langer than wide, Setae of moderate length.
Relative length of podomeres 2 to 6 in middle part of
bady: 0.80, 1.00, 0,55, 0.45, 0.95. Coxae af 2nd pair of
legs on caudal side with a low rounded transverse callus.
Epigynal structure with median paint durected obliquely
downward; lateral anterior projection also directed
obliquely downward,
Remarks
In points not mentioned the description of O
castaneunr applies.
This is the smallest representative of the genus
Oncocludosoma, dillering from the other species in
small details of the external morphology and
particularly in the structure of the gonopods.
It may be of imerest to note that the specimens for
Belair are slightly smaller on the average, parallelling
the vonditions in O. eastaneum. In the present case,
however, the differenves seem jusignificant.
MILLIPEDES FROM AUSTRALIA 5 33
FIGS, 8-9. Oncocludosoma clavigerum i. sp., holotyped . 8; right gonopad, medial aspect. 9: left gonopod, anterior aspect.
THE STATUS OF POLYDESMUS
(STRONG YLOSOMA) INNOTATUS KARSCH, 1881
The taxonomic position of this species, allegedly based
on a female specimen from Adelaide, has been uncertain
since the publication of its description. Through the
kindness of Dr M. Moritz of the Museum fir
Naturkunde der Humboldt-Universitat in Berlin, | was
able to re-examine the type-specimen, hoping that it
could be associated with one of the paradoxosomatids
received from the South Australian Museum.
The type-specimen is in bad condition; its antennae
and most of its legs are missing. Moreover, it is not an
adult female, but a juvenile with 19 somites.
Nevertheless, on account of its external characters it has
become clear that the species is different from other
Paradoxosomatidae reported from South Australia. In
case the species is rediscovered in South Australia its
most salient characters are mentioned here,
“Strongylosoma™ innotatum (Karsch)
Polydesmus (Strongvlosoma) innotatus Karsch, 1881: 42.
Strongvlosoma innotaium; Attems, 1898: 307.
Material
Adelaide, Schomburgk leg., 1 juy. % (19 somites),
ZMB Kat.Nr. 560, holotype.
Descriptive notes
Colour; Dull brownish, apparently faded.
Width: 3.5 mm.
Head and antennae: Without particulars.
Collum; Wider than head (Fig. 10). Lateral sides
flaring, rather widely rounded and without posterior
edge; lateral border merging gradually into caudal
border.
Somites; Waist rather narrow, without apparent
sculpture. Metatergites of 4th to penultimate somite with
a generally deeply impressed transverse furrow, running
far laterad and reaching in most cases dorsal
premarginal furrow of paranota.
Paranota: (Figs 10-11) 2nd somite distinctly wider than
collum; 3rd and 4th each slightly narrower than
preceding somite. Paranota of 2nd somite on a low level,
well developed; anterior border shouldered at base and
34 REC. 8. AUST. MUS, 19 (3): 19-37
FIGS. 10-11. “Strongylosoma” innotatum (Karsch), holotype ? juvenile
(19 somites), 10; left side of head, collum and 2nd and 3rd somites,
dorsal aspect. Il: left side of 9th somite, dorsal aspect.
thrust forward a little, widely rounded, merging into
rather faintly convex lateral margin via a somewhat
stronger rounding, Posterior edge rather narrowly
rounded but not angular. Posterior border a little
convex; posterior part of paranota projecting distinctly
caudad of margin of somite. Paranota of 3rd and 4th
somites also prominent, widely rounded anteriorly and
laterally, more narrowly rounded caudally and project-
ing a little behind margin of somites. Paranota of Sth
and subsequent somites at first subangular caudally, but
in second half of body becoming more distinctly
angular, though produced caudally and projecting only
a little from 15th somite onwards.
May, [985
Sternites and legs: Sternites without distinctive
characters; legs mostly absent (in specimen).
Anal somite: Epiproct moderately developed, apex
truncate with a weak caudal emargination. Hypoproct
rather broadly triangular, with sides and apex rounded.
Remarks
Since the specimen is a juvenile, most of its external
characters may be less pronounced than in the adult,
in particular the adult male. The paranota are different
from those of any of the other South Australian
paradoxosomatid genera. In particular the relatively
wide expansion of the paranota of the 2nd somite seems
characteristic. In this respect the species appears to have
a certain similarity with Ofoplacosoma bivittatum
Verhoeff, 1924, from the Kimberley district of Western
Australia, which would mean that eventually it may
prove to belong to the tribe Antichiropodini. It should
be borne in mind though, that it is also possible that
the locality label is wrong, in which case the systematic
position of the species probably will be an enigma for
ever.
SUMMARY OF SOUTH AUSTRALIAN
PARADOXOSOMATIDAE
With the completion of the study of the Para-
doxosomatidae of the collection of the South
Australian Museum, it may be useful to summarize the
results which have been obtained so far.
The number of known species has increased from two
to ten and one subspecies. The list is as follows:
Tribe Antichiropodini
Antichirapus mammillifer Jeekel
Aulacoporus pruvoti (Brolemann)
Tridactylogonus obscurus Jeekel
Incertae sedis
“Strongylosoma” innotatum (Karsch)
Tribe Australiosomatini
Heterocladosoma zebratum Nn. sp.
Heterocladosoma galaxias n. sp.
Somethus grossi n. sp.
Oncocladosoma c, castaneum (Attems)
Oncocladosoma c. ingens n. subsp.
Oncocladosoma conigerum 1. sp.
Oncocladosoma clavigerum n. sp.
Considering the fact that the southeastern part of
South Australia, where these paradoxosomatids were
obtained, is quite remote from the nearest occurrence
of the family elsewhere and is moreover drier than the
areas in eastern Australia where the majority of the
other described species occur, it is remarkable that the
number of endemic genera is so relatively low. To this
category belong only Tridactylogonus and
Oncocladosoma. The other genera, viz. Aniichiropus,
Aulacoporus, Heterocladosoma and Somethus, are
MILLIPEDES |}ROM AUSTRALIA 5 35
shared with other Australian states, where, according to
our present knowledge they all show a greater diversity.
Antichiropus has quite a number of species occurring
along the west coast of Western Australia.
Heterocladosoma has three distinct species in the coastal
area of Queensland from Cairns to Brisbane, and,
according to as yet unpublished data, Somethus has two
species in Tasmania and one in eastern Victoria.
Aulacoporus has a number of species in Queensland and
the northern part of New South Wales, but the South
Australian record of A. pruvolti probably represents an
introduced population. It is evident anyway that the
fauna or South Australia, as far as the non-endemic
genera are concerned, is composed of a mixture of
western (Antichiropus), northeastern (Heterocladosoma),
and southeastern (Somefhus) elements (Fig. 12).
Considering the diversity within each of these genera
“/
2
- a
“or
i
9
O (e)
-O
Pie oT
elsewhere it seems probable that South Australia became
populated along these routes, rather than the reverse.
The relationship of the two endemic genera has not
yet been satisfactorily determined. Tridactylogonus
seems to belong to a group of small antichiropodine
paradoxosomatids which focuses in Victoria, Tasmania
and southern New South Wales. Oncocladosoma
appears to be closely related to Somethus,
Dicladosomella and Phyllocladosoma, indicating a
faunistic connection between South Australia, Tasmania,
eastern New South Wales and southern Queensland.
Oncocladosoma represents, moreover, a particularly
interesting case. The genus has been found only in the
Mt. Lofty Ranges, where it is represented by four partly
sympatric taxa. No other paradoxosomatids have been
reported from this probably best-explored part of South
Australia, and the faunal composition shows the aspects
ag\!
FIG. 12. Map showing the distribution of the non-endemic South Australian australiosomatine genera Heterocladosoma (*), Somethus ( @ ),
and Antichiropus (0), and the endemic genera Oncocladosoma ([~) and Tridactylogonus (_ ).
3A REELS. AWIST, MUS, 10 (4)5 19-97
of that ofa long isolated island. During an early period
this area must have been populated by a single species,
possibly from the Sume/ius-stem, which, in the course
of a long period of isolation, radiated in a number of
closely related taxa,
The presefr picture of the distribution of the South
Australian Paradoxosomatidae, incomplete as it
undoubtedly is, seems to indicate that the fauna is the
result of exchanges during at least two widely separated
periods in history. The lirst of these periods, which may
he more or less arbitrarily dated at the late Mesozoic
or early Tertiary, saw a faunal connection between
Victoria atid southern New South Wales and South
Austraha, and is witnessed by Yridactvlogonus and
Oncocladosoma, The second, possibly occurring during
the Jate Tertiary, brought representatives of
Heterocladosoma, Somethus and Antichiropus to South
Australia,
Of course, this is a very simplified explanation of
What actually may have happened, but it will be
interesting to assemble further data lo verify and work
out the details of the hypothesis.
RECORDS OF SOME MEDITERRANEAN
JULIDAE INTRODUCED INTO SOUTH
AUSTRALIA
Records of millipedes introduced into Australia from
elsewhere are few. Baker (1978) extensively reparted on
the distribution and dispersal of the west Mediterranean
julid Onrmeatoiulus moreleti (Lucas) in South Australia
and Vicloria. | have summarized the known records of
European millipedes in Tasmania (Jeekel 1981) and more
reeently added the oriental paradoxosomatid
Orthomorpha coarcnita (De Saussure) to the Australian
list (Jeekel 1982a)
The material of the South Australian Museum
contained, besides some specimens of Onwndraiilus,
also a number of samples of a Mediterranean species
of Bravhviulus not previously known to occur in
Austraha.
Brachyiulus iusitanus Verhoell
Brachviylus pusillus lisitunus Verhoell, 1898: 153, PL.
6, Fig, 28,
Microbrachyiults calcivagus Verhoef, 1910; 225,
Figs. 33-34.
Brachviulus lustianus; Sawlowski, 1930: 183, big. 4;
Strasser, 1976: 606, Fig. 37-
Pl. 2,
Material
Woodley’s Vineyard (near Glen Osmond, SE
Adelaide), soil, Berlese funnel, 30V1I-8VHL1950, les,
R. V. Southcott, 3¢, 52, 8 juvs,
Woodley’s Vineyard, soil, Berlese funnel, 3.18.1950,
lex, R. V, Southeotl, 4 juvs,
Sellick’s Hill, 23.16.1954 (E.S.J. 1953),
Gross, 12,
lee Ce 1.
Maw, 1985
Burnside, 12.VI1962, leg. Mr Waxman, 2¢, 39.
Burnside, 24.1.1963, leg. Mi’ Wasman, 1d’, 16 2.
Remarks
This species was originally described froin Portugal,
and has since been recorded from numerous localities
in the Mediterranean region, eastward to Greece, Turkey,
Syna and Turkestan. Apparently most of the records
outside the western Mediterranean range concern
synanthropic localities, and the species is obviously
casily distributed by. commerce. Outside the palearetic
region records are still few, and since Jawlowski (1930)
reported B /usttanus from Mexico the species was only
recenuy mentioned as occurring in the southeastern
United States by Filka.& Shelley (1980) where it seems
to have beeh previously misidentilied as Brachyiulus
pusillus (Leach).
B. lusttanus is similar to B pusillus, but differs in its
slightly larger maximum size; it can be easily separated
by the fanlike phylacum ol the posterior gonapods
(spiniform in B pusillus) (see the Wustrations in Verhoeft
(1910), Jawlowski (1930) and Strasser (1976)).
Ommatoiulus moreleti (Lucas)
fulus inoreleti Lucas, 1860: 96,
Archiulus toreleri; Attems, 1928: 291, Pl. 18, Figs.
427-432.
Schizophyviliam morelet, Schubari, 1966: 23, Migs. 12-20.
Ommataiulus moretletii; Baker, \978: 1.
Material
North End Pt. Lincoln, 3.01955, leg, M. Carrick,
2%,
Retiarks
This species originates froin the [berian Peninsula,
where il apparently occupies a rather restricted range
in the northern half of Portugal and adjacent Spain.
It has been reported from a number of Atlantic islands
(Canary Islands, Madeira, the Azores, Bermuda, St.
Helena), the Cameroons and South Africa. In Australia
it is known to become more and more wide-spread in
South Australia and Victoria (Baker 1978) and it has
also been reported from Tasmania, although as yet it
does not seem to be so well established there,
Once it is introduced into a suitable environment ii
spreads rapidly and becomes a dominant clement of the
millipede fauna of the area. On Teneriffe (Canary
Islands) for instanee, it is seen almost anywhere in areas
under human influence, Schubarl (1966) gave an
extended report on its distributional aspects in Sourh
Africa, where it appears to have spread over a large part
of the country in the course of a lew decades, Baker
has deseribed a similar phenomenon jin South Australia.
Ommataiulus and Brachyiulus ave easily distinguished
from) the autochthonous jnliform millipedes oF Sauth
/Nustraia by the longitudinal striation of the dorsal side
MILLIPEDES FROM AUSTRALIA 5 37
of the metasomites and by the freely projecting cauda,
the latter being particularly distinct in Ommatoiulus.
Brachyiulus is further distinguished by having two
yellowish dorsal paramedian bands.
REFERENCES
ATTEMS, C. 1928. The Myriopoda of South Africa. Arn. S. Afr
Mus. 26: 1-431, pls. 1-26,
ATTEMS, C, 1944. Neue Polydesmoidea, Zool, Anz. 144: 223-251.
BAKER, G. H. 1978. The distribution and dispersal of the introduced
millipede, Ommatoiulus moreletii (Diplopoda, lulidae) in
Australia. J Zool, Lond. 185: I-11.
CHAMBERLIN, R. V, 1920. The Myriopoda of the Australian region,
Bull. Mus. comp. Zool, Harv. 64: 1-269.
FILKA, M. E., and SHELLEY, R. M. 1980, The milliped fauna of
the Kings Mountain region of North Carolina (Arthropoda:
Diplopoda). Brimleyana 4: 1-42.
JAWLOWSKI, H. 1930. On European Diplopoda introduced to
America. Frag. Jaun. Mus. zool. polon. 1: 183-185, pl. 5.
JEEKEL, C. A. W, 1968. On the classification and geographical
distribution of the family Paradoxosomatidae (Diplopoda,
Polydesmida): i-vii, 1-162, Amsterdam (privately printed).
JEEKEL, C. A. W. 1979. Notes on the classification of some little-
known Australian paradoxosomatid genera (Diplopoda,
Polydesmida) J nat. Hist, 13: 649-658.
JEEKEL, C. A. W. 1980. A new genus and two, new species of the
family Paradoxosomatidae from Australia (Diplopoda,
Polydesmida) Bu/l zool Mus. Univ, Anist, 7: 1-10.
JEEKEL, C. A. W. 1981. Australia Expedition 1980; legit C. A. W
Jeekel and A, M. Jeekel-Rijvers. List of collecting stations,
together with general notes on the distribution of millipedes in
eastern Australia and Tasmania. Vers/. techn. Gegey. Inst. taxon.
Zool. Amst, 30; 1-59,
JEEKEL, C. A. W. 1982a. Millipedes from Australia, 1;
Antichiropodini trom South Australia (Diplopoda, Polydesmida,
Paradoxosomatidae) Bull. zodl, Mus. Univ, Amst, 8: 121-132.
JEEKEL, C. A. W. 1982b. Four new or little-known Australiosomatini
from New South Wales (Diplopoda, Polydesmida,
Paradoxosomatidae) Bijdr. Dierk, 52; H 137-154.
KARSCH, F. 1881. Zum Studium der Myriopoda Polydesmia. Arch.
Naturgesch. 47(1): 36-49, pl, 3.
LUCAS, H. 1860. Myriapodes, /a “Notice sur histoire naturelle des
Acores” (Morelet, A.), pp. 96-97. Bailliere, Paris.
SCHUBART, O. 1966. Diplopoda IIl, Pselaphognatha,
Opisthospermophora, Colobognatha. 8. Afr anim. Life. 12: 9-227.
STRASSER, K. 1976. Uber Diplopoda-Chilognatha Griechenlands
II. Rev. suisse Zool. 83; 579-645,
VERHOEFF, K, W, 1898. Ueber Diplopoden aus Bosnien,
Herzogowina und Dalmatien, [V. Theil: Julidae. Arch. Naturgesch.
64(1): 119-160, pls. 5-6..
VERHOEFF, K. W. 1910. Uber Diplopoden. 11-15, Aufsatz (31-35):
Beitrage zur Kenntnis der Glomeriden, Juliden,
Ascospermophoren und Lysiopetaliden, sowie zur Fauna Siziliens,
eic. Nova Acta Acad. Caesar. Leop, Carol, 92: 139-448, pls. 1-9.
SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS.
4. PRIMITIVE ORIBATE MITES (CRYPTOSTIGMATA) WITH AN
EXTENSIVE, UNFISSURED HYSTERONOTAL SHIELD AND APTYCHOID
BY DAVID C. LEE
Summary
A study of sarcoptiform mites from surface soil (greatest depth usually = 4cm) at 9 florally diverse
sites in South Australia is continued. Opisthosternal shields, supernumerary leg setae, gnathosternal
fissures and cheliceral spatulae are considered separately. The Cryptostigmata is treated as
including 5 suborders: Palaeosomatida, Retrofissurida, Afissurida, Profissurida and Comalida. The
Afissurida and primitive Comalida (i.e. Mixosomatina and Clinofissurae) are considered further in
order to complete the lower oribate mites within this study.
SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS, 4. PRIMITIVE ORIBATE MITES
(CRYPTOSTIGMATA) WITH AN EXTENSIVE, UNFISSURED HYSTERONOTAL SHIELD
AND APTYCHOID,
by
DAVID C, LEE
South Australian Museum, North Terrace, Adelaide, South Australia SOQ).
(Manuscript accepted 20 December 1983)
ABSTRACT
LEE, D.C, 1985. Sarcoptitormes (Acuri) of South Australian soils.
4. Primitive oribate mites (Cryptostizgmata) with an extensive,
untissured hysteronotal shield and aptychoid. Rec, S. Aust. Mus,
14): 39-67.
A study of sarcoptiform mites from surface soil
(greatest depth usually 4 cm) at nine florally diverse sites
in South Austraha is continued. Opisthasternal shields,
supernumerary Ieg sctac, gnathosternal fissures and
cheliceral spatulae are considered separately. The
Cryptostigmata is treated as including five suborders:
Palacosomatida, Retrofissurida, Alissurida, Profissurida
and Comalida. The Afissurida and primitive Comalida
(ic. Mixosomatina and Clinofissurae) are considered
further in order to camplete the lower oribate mites
within this study. Five families are discussed although
not tepresented by specimens in this study: the
Eulohmanniidae to confirm its inclusion in the Mixo-
somatinay the Malaconothridae, Trhypochthonidae,
Trhypochthoniellidae and a new family, Allonothridae,
to delineate the two infraorders (Mixosomatina and
Holosomatina) of the Comalida. Seven species of
Afissurida, Mixosormiatina and Clinofissurae were
collected. Four species are new: Plafynethrus
hreviserosus, Croronia jethurmerae, Nanhermannia
grundjeuni, Phyllhermannia eusetosa. Two subspecies are
new: Epilohmannia cylindrica media, Epilohmannia
pallida australica. Papillacarus pseudoaciculatus 1s newly
recorded from Australia, A new synonym is kepracerus
under Papillacarus. New combinations are egypfica and
imdica (ex Papillacarus) with Lohimannia, and kereanus,
ogawai and ramirezae (ex Vepracarus) with Papillacarus.
INTRODUCTION
This publication is a further part of an ongoing study
(Lee 1981, 1982). Its intent is to complete the systematics
of what are generally regarded as the primitive, lower
or macropyline aribate miles. Because the higher
classification has changed sa much since this study
began, the oribate mites considered here are not all
closely allied. They appear to be similar because they
are primitive but have an extensive unbroken
hysteronotal shield (a derived character state convergent
in Wwo suborders) and are aplychoid (a character state
ancestral to the derived ability to fold the proterosoma
back onto the hysterosoma which has arisen in three
suborders). The mites ta be considered include the
Lohmannioidea (Afissurida), the Mixosomatina
(Comalida) and the Clinofissurae (Comatlida-
Holosomatina). The following families from the
Mixosomatina have already been dealt with:
Phthiracaridae and Euphthiracaridae under Euptyetima
Lee (981) as well as Gehypochthoniidae under
Monofissurae (Lee 1982).
The numerous higher categories of oribate mites may
be reduced, some becoming superfamilies, if there is
compensation for the disproportionate tendency to
“upgrade” the classification (Krantz 1978), However,
these categories are retained here, | follow the recent
discarding by O’Connor (1984) of the abnormal use of
“cohort” in acarology, Instead, between infraorder and
superfamily, the rank of section is used as for
crustaceans (Kaestner 1970).
The mounting procedures used have sometimes
resulted in squashed specimens. The possibility of such
distortion, always represented in illustrations of
pnathosterna where tt facilitates description, should be
considered,
All mites described in this study are deposited in the
South Australian Museum, Adelaide.
MORPHOLOGY
Opisthosternal Shields and Setae
The opisthosternal shields may be defined by the setae
they bear. Unfortunately, in the reduction from the
primitive three pairs of setal files, it is difficult to assess
which files are lost or merged. Therefore, the geni/a/ and
anal shields are regarded as the movable shields covering
the relevant orifices and bearing setal files Jg and Zz
or Ja and Za; and if these shields bear only one setal
file it is referred to as either J/Zg or JZa whether or not
it appears likely that a specific file (eg. Ja in some cases)
is missing, The more lateral shields are the aggenital
shield bearing setal file Sz and the edanal shield beanne
setal file Sa, which may be merged to form the
opisthoventral shield. \n the Retrofissurida and
Afissurida, the anal shield may be split into a peronal
shield (bearing one or two setae Ja) and a paranal shield
ai RECS. ACIST. MUS) JO (lps 3U.67
(bearing two, three or four setae Za), In such cases, |
have in the past (Lee 1982, Figs. 7 and 16) referred to
these setal files with a different notation. In exceptional
cases, as Hoplophihinicarus shealsi Lee, 1981 (see Pix.
25), the movable shield over the genital orifice bears
three setal files indicating it is a merged genital and
agpenital shield.
Chaetolaxy of Legs and Coxites
The Ieg tarsi of some species described below are
hypertrichous. Lying proximally to the normal
coniplement setae, the exira supernumerary setae are
restricted to the four following positions: two
dorsolateral—da, dp and two ventral—u\, pu The
suffix “xv? is added to the Sienature lor each
supernumerary sctac, followed by a number indicating
the position of the including whorl or rank, with "1"
the most proximal.
The ventral conite setae are in approximately single
file and are numbered as such [rom the adaxial end
without any attempt to hamologize them with the files
(anterior, median, posterior) from which they might
be derived.
Gnrathosternal Fissures and Cheliceral Spatula
Gnathosternal fissures are regarded as derived and
may be functionally correlated with greater rigidity af
the idiosoma requiring greater mobility of the
mouthparts. The two types of fissure have been detined
(Lee 1984). The species described below have wento-
coxal fissures, whilst what was referred to as “diarthrie”
(Grandjean 1957) or a “quadrangulate mentum" (Lee
1982) occurs in more advanced taxa with dicoxal
Jissures, These (wo types of Tissures split different parrs
of the gnathosternum (although the homologies of
areas regarded as derived from either Lhe palp coxile
or mentum are not confidently held), whilst before the
differences were regarded as due to a change in shape
of the mentum, Apparently in rare cases both types
of fissures occur on the same animal (see Nesfris
species, Tragardh 193la; Fig, 6),
As previously (Lee 1984), the term eheliceral spatula
is used for 'Tragdrdh's argan'. The presence of a
cheliceral spatula was regarded as a possible
synapomorphy for the Holosomatina (Comalida) (Lee
1984), Norton (personal communication 1982) has
since pointed out that the cheliceral spatula is present
in a number of families of Mixosomatina (Comalida)
and | discuss this further below under Remarks” en
that intraorcter.
SYSTEMATICS
| acvepl the concept that the Cryptostizmata is a
paraphyletic taxon as suggested by O'Connor (1984),
with the Astigmata regarded as a sister-group to part
of the Comalida (in the restricted sense used below).
Muy, [UNS
On the other hand, t-do not in this instance follow the
principle of Wiley (1981) that paraphyletic groups be
rejected as Unnatural, arbitrary, hurrian constructs, |
consider it useful to retain the Cryptostigmata as a
grade, sharing similar primitive character states, and
“left behind” by the considerable differentiation of the
Astigmala.
The higher classification of the Cryptostigmata is
modified so that it is regarded as including live
suborders, sequenced to reflect their supposed level of
advancement as follows: Palacosomatida (see
Bifemorata Lee, 1981), Retrofissurida (see Retrofissurae
Lee 1982), Afissurida (see Afissurina Lee, 1984 and
below), Protissurida (See Profissurae Lee, 1982 and
Pediculochelidae) and Comalida (see Comalida, in
part, Lee, 1984 and below), Changes. from the
classification presented at the 6th International
Coneress of Acarology in 1982 (Lee 1984) are that the
Dismatida is disbanded and the included three distiner
primilive lineages upranked to suborders, whilst the
Profissurina is excluded from the Comalida and also
upranked to a suborder. Four of these suborders may
be regarded as clades, bul the Comalida is not a
complete clade since the Astigmata is excluded from
it (although Wt may prove arguable that the entire
Comalida is a sister-group to the Astigmata).
Upranking of the Profissurida is a response to {he
proposition (Norton, O’Connor and Johnston 1983)
that the Pediculochelidae and Haplochthoniidae
constitute the latest derivative lineage within the
Protoplophoroidea (=Prolissurida). Eyen if this
phylogeny was Well Supported, [| would maintain the
Pediculochelidae as a separate superlamily on the basis
of morphological divergence (widely separated coxites
WW-LV from J-LL, terminal anus, pretarsal stalked pad),
although I do tentatively accept that with the
Protoplophoroidea it constitutes a monophyletic group,
On the other hand the /ate derivative phylogeny of
Pedieulochelidae is nat acceptable since there is no
evidence suggesting an ancestor with hysteronotal inter
calary sclerites and erectile setae, or that the absenve
of sclerotization and reduced setation (often a
convergent sinilarity) are synapomorphies fora lineage
composed of it and the Haplochthoniidae. | propose
that the Pediculochelidae is regarded as an early
derivative group as indicared by the primitively disjunct
external malae. This suggests that fhe conjunet external
malae of other Profissurida have arisen separately as
an apomorphy convergent lo a similar charaeter state
in the majority of the Comalida and (herefore supports
the upranking and exclusion of the Profissurida fram
the lanier sitborder,
Suborder AFISSURIDA
Diagnosis: Cryptostigmata. ELysteronotal shield lacks
transverse Fjssures, bul iransverse furrows olten present.
Hysieronotal gland absent. Hystcronotal chactotaxy
SARC OPITIPORMES (AC ART) OF SOLUTE AUSTRALIAN SOIDS 4 4)
holotrichous or hypertrichous. Setal files wilh lwo setae,
both similar in size and shape to other proleronotal
setae. Coxites merged, forming two podosternal shiclds
(conites 1, 1 or Ith, WV), Cheliceral spatula ubsent,
External malae disjunct. Gnathosternal fissures absent,
but mentocoxal furraw sometimes present, Adoral setae
in transverse row, at least adaxial two pairs large and
flattened. Pretarsi with one claw. Tarsus | and genu J
cach with two solenidia. Tibia | solenidia flagelliform.,
Femora undivided, Nymphs with similar facies to adult.
Remarks: The Alissurida, alsa referred to as
Afissurina (Lee L984), Includes only one superfamily,
the Lohmannioidea; Grandjean, 1969, The
Lohmannioidea was then included in the Mixonomata
which approximates to the Mixosomatina (Comalida)
considered below. Despite this, Grandjean (1950), in a
thorough work on the Lohmannidae, has. indicated that
this family is very distinct from other oribate rites, but
with distant affinities to certain “Enarthronota”. I agree
that it Shares.a number of primitive charaeter states with
the Retrofissurida (part of old “Enarthronota’) such as
disyunel external malae and a complete notal chaelotaxy;
including two large, well-separated setae in file s» and
siateen pairs o! hysteronotal setae. On the basis af this,
it is reasonable to regard the absence of a hysteronotal
gland as primitive and nat secondary as in some
Mixosomatina (Comalida). Therefore, my inclusion of
the Lohmannioidea in a separate, primitive suborder,
with its extensive hysteronotal shield considered as
convergent with thal slate amongst the Comalida,
reflects the earlier (950) conclusions of Grandjean
rather than his later (Grandjean 1969) work.
Besides the Lohmanniidae, two other families have
been recognised in the superfamily, each including only
a single species. The Nothrolohmannidae Balogh, L968
is recognisable by bifurcate seta /l, humeral processes
bearing seta Sl and large triangular aggeniral shield
fused to posterior margin of coxile IV. The
Xenolohmanniidac Balogh and Mahunka, 1969 has a
“menton divided into two parts, not meeting medially”,
As illustrated, this structure appears as if it might
represent a pair of broad Ilat pracesses extending
forward from coxites | rather than an unusual
gnathosternum.
Family LOHMANNSIIDAE Berlese
Lohmannini Berlese, 1917b:; 176
Lohmannidae: Grandjean, 1950; 100
Lohmanniidae: Wallwork, 1962a; 457
Lohimanniidae: Corpuz-Raros, 1979: 315
Type-genus: Lohimannia Michael, 1898: 75
Diagnosis Afissurida, Preanal shield present: either
bar-shaped, “Eshaped, ar reduced fo central knob,
Aggerital shield sometimes present, but not fused ta
coxite IV or extending posteriorad to halfway along
genital shield, No broad flat process extending forward
from coxite £ under gnathosternum. Hysteronotal seta
St not on conspicuous humeral proeess, Proteranotal
stta JL not bifureate,
Morphology: Small to large (480-900) dull yellow or
brown mites, Proteronotum long (0.4% length of
hysteronotum) and broad (as wide as hysteronotum).
Opisthosternum usually shorter (han podosternum, Legs
short (leg ] longest, less than 0.4x idiosomal length).
Palp femur and genu fused or partially fused. Genital
shield bears LO or more setae in two files (usually 6/e,
4Ze), may be divided into subequal anterior and
posterior paris. Setal files. Sz and Sa absent. Anal shield
may be divided into petanal and paranal parts.
Solenidiotaxy [ (2-1-2), 11 (1-1-1 or 2), U1 (1-1-0), TV
(1-0-0),
Distribution, Widespread in tropics, where greatest
poneric diversity Occurs. Particular species may
themselves be widespread, Achieved “a greater deyree
of evolutionary radiation in Africa than in Sauth
America’ with “a wide distribution in those parts of
the world which belonged, formerly, to Gondwanaland”
(Hammer and Wallwork 1979). [nm temperate regions,
but generic diversity decreases away from tropics,
Lohmannia beng most widespread. In northern
hemisphere, genera other than Lofimannia only found
south of 45°N.
Some species known to burrow in roots or bark,
Represented. both in deserts and moist habitats such as
coastal bogs, Ecological notes on species collected in
Philippines (Corpuz-Raros 1979) indicate wide variety
of niches which family can oecupy in limited
peographical area.
Remarks; The above diagnosis distinguishes the
Lohmanniidae from the two very similar monospecific
families alsa in the Afissurida and is, therefore, relatively
trivial. A number of further character states are listed
under “Morphology” which may apply to all three
families.
Balogh (1972) included 20 genera in the
Lohmanniidae. Amongst these, authors have regarded
the following four genera as similar to Papillacerus
(Which 15 represented in this study): Crvpracarus
Grandjean, 1950; Dendracarus Balogh, 1960;
Lepidacarus Csiszar, 196); Kepracerus Aoki, 19656. On
the basis of the disposition of ihe opisthosternal shields
Cryptacarus and Dendracarus can be excluded. Whilst,
as elaborated on below, Lepidacarns can also be
distinguished, béepracarus is regarded as synonymous
with Papillacariss,
PAPILLACARUS Kunst
Papillacarus Kunst, 1959: 70, Type designation (original):
“Lohmannia nurcioides Berlese v. aciculuta Berlese
1905",
42 REC. S. AUST. MUS 19 taj: 40-67
vepracurus Aoki, 1965b: 142. Type designation
(original): “* Fepracarus ogawai Aoki,spec.wov.—n.sy i.
Type-species: Pupillucarus aciculams (Berlese, 1905; 24).
Diagnosis: Lohmanniidae. Genital shield divided by
transverse fissure into two subequal shields, Anal shield
divided by longitudinal fissure into peranal and paranal
shields, Preanal shield reduced to central, bicornate
knob. Rostral tectum blunt, anterior margin cither
smooth arc or sinuous with four slight tubercles or with
three conspicuous tubercles (lateral tubercles where
lamellae meet margin), Proieronotal plasmic seta (32
with long cilia on only one side of slim or lanceolate
main stem. Notal setae with cilia which may be
IMconspicuaus or so long that setae bush-like, Posterior
notal hypertrichy, never anterior to seta J2. Notal
integument papillate. Yentral ridges on femora | and I.
Analchaerotaxy: 2/a, 42a, Tarsi UT and IV each with
10 setae. Tarsus UL with two solenidia.
Distribution. Possibly almost limited to Old World,
Virgin Islands (NTa); Anuobon Island, Ghana (Ew);
Crimea, Moscow, Ukrainia (Pe); Bulgaria, Greece, [aly,
Spain, Tunisia (Pm); Korea, Japan (Pe); India (Oi);
Thailand (Os), Java, Philippines (Om); South Australia
(Aa). Known Oriental fauna Shows greatest
morphologival cliversity,
Found in grassland, feeding on grass roots, also under
trees, where grass may or may not have been present.
Remarks: Three genera (Lepidacarus, Pupillacarus;
and beprecarus) have the disposition of opisthosternal
shields given in the above diagnosis, Lepidacaris can
be distinguished from the other two genera by the
preanal shield betng an undivided central knob, as well
as by the hysteronotal setac being Ical-like.
The only species af Papillacarus and bepracarus that
has beer described in enough detail to melude characters
of the gnathosoma and legs ts P angulatus. Therefore,
characters used to delineate these genera and their
included species have to be limited to those of the
idiosoma and in Some cases to the notum. On the basis
of such characters | have not been confident about
distinguishing the two genera. Instead | have merged
them and regarded Papi/lacarus as including the four
following species-complexes based on the form and
chaetotaxy of the hysteronotal setae. This is intended
as a temporary measure until more characters are
considered. Although the aciew/arus-complex (includes
type of Pupillucarus ) and ogawal-complex (includes type
of Fepracarus) are easily delineated, this is undermined
by the other two complexes. The fact that the Airsuriys-
complex ificludes species from both genera illustrates
this confusion. The type complex is considered last since
it includes the species found in South Australia, P
egypticus Elbadry and Nasr, 1977 and P indicus Haterz
Kardar, 1972 are grouped in Lokmannia, neomb,
Mul, [as
Aursurs-camplex
Diagnosis: Papillacarus, Hypertrichy on hysteronolurmn
posterior to seta /4 ar JS, area bearing 50-60 setae
Supernumerary setae shore (length subequal to or less
than distance between posterior setal bases), bush-like
(because long cilia), Complement setae similar,
indistinguishable. If any Jonger seta (uncertain if super-
numerary or complement), then around posterior
margin,
Remarks: Three species included in the complex, P
hirsutus (Aoki, 1961), ex Crypracurlis —Po, Om, Ap. P
vhanarrinesis Perez-Imgo, 1967—Pm. PF koreanus
(Mahunka, 1973), ex Feprocerus, nxomb.—Pec.
ogawai-complex
Diagnosis: Papillacarus. Hypertrichy on hysteronotum
posterior to seta /4, area bearing 50-80 setae. Super-
numerary setae short, bush-like. Complement setae at
least twice as Jong, with inconspicuous cilia, Single rank
of tubercles between setal ranks 5 and 6.
Remarks: Two species included in complex. P ogawal
(Aoki, 1965b), ex type of bepracarus, ncomb—Os. P
ramirezae (Corpus-Raros, 1979), ex bepracaris;
n.comb.—Om.
rantasus-complex
Didgnosis: Pallacarus. Hypertrichy on hysteronoturn
posterior Lo seta J2, area bearing 120-170 setae. Super
numerary setae shart, bush-like, Complement setae /4,
and those posterior to it, more than five times as long,
with imconspicuous cilia.
Remarks: Two species included in complex, P nameasts
Balogh, 1961—Om. P crugae Corpuz-Raros, 1979—Om.
aciculatus-complex
Diagnosis! Papillacarus, Hypertrichy on hysteronotum
posterior to seta J/4, area bearing 20-45 setae, Super-
numerary setae short but similar in form to complement
setae, 60 both with inconspicuous cilia, not bush-like
Remarks: Six species included in complex. P
ucieulatus Berlese, 1905—Pe, Pm. P angulatus
Wallwork, 1962a—Ew, P undiras(ralus Aoki, 1965b-Os.
FP. ondriasit Mahunka, 1974—Pm. P pseucdeaciculaius
Mahunka, 1980a—Pm, Aa. P vwitis Elbadry and Nasr,
1977-——-Pin,
Papillacarus pseudoacicalatus Mahunka
(Figs 1, 2)
Papillecarus pseudoacioulaius Mahunka, 1980a: 126
Triranymph
Dull, ochre-coloured with yellowish brown cheliceral
extremities, external malae and legs. Cuticle of shields
mainly vranulale except in transverse hysleranotal
furrows and in small, usually marginal patches, Similar
SAKCOPTIFORMES (AC ART) OF
but less extensive arcas papillate, protuberances being
aciculate (as few on gnathosternum, illustrated Fig, 1),
Cuticle of prehysteronotal fissure strongly strated
suggesting, propodosoma can flex downward. [diosomal
length 510 (1); appendage lengths—ch 42.5, pa 45, / 180,
{1 160, 177135, 7175; fetnur breadths—pa 12,5, 750,
HE 45, HF 37.5, (42.5.
Cheliceral seta chl much shorter than ch,
Mentocoxal furrow present, clearly not fissure, Rostral
teclum hyaline, indistinct, possibly smooth are:
Dorsolateral longitudinal fissure from. anterior margin
of hysteronotal shield back (o level of preanal shield.
This delineates dorsal margin of pleural shield which
bears no setae bul slit-like pore (#f1) at anterodorsal
corner. Few supernumerary hysteronotal setae (?six} so
area behind /4 bearing about 18 setae. Cilia on
hysteronotal setae Jonger than depicted on type
specimen, more like & endriasi (Mahunka 1974: 575,
Fiz. &), but supernumerary setae relatively longer. Scta
Jé conspicously longer than surrounding setae, Six
Iransverse hysteronotal furrows, only anterior furrow
just behind scta /1 complete, posterior lurrow Jevel with
seta J/4 reduved to two iudistiner pits. [ndistiner
structures may represent two pairs of genital papillae,
Five pairs of setae on anterior genital shield and four
pairs on posterior genital shield (7./z, 2Z2), possibly Ze3
missing from adult complement.
Material examined: Ong tritanymph (N19831), bases
of grasses and plantains, Glenthorne, 12.6.1974, D.C.
Lee.
Lysiribution: Tunisia (Pm); South Australia (Aa),
South Australia: Glenthorne, cuitivated pasture, | trito-
nymph (« /8).
Remarks: Vhe referral of this single specumen to be
a tritonymph of A pseidouciculatus is not done with
confidence. The identilicatian is based mainly on the
low number of supernumerary hysteronotal setae,, but
this may be related (o at being an immature stage. The
extensive somal coverage with aciculate protuberances
in dense patches excludes some other species. Since the
species are So similar in the aciewlatus-complex, the
above iyitonymph could represent a new species.
Suborder COMALIDA
Diagnosis: Cryplosigzmala. Hysteronotal shield
usually lucks transverse fissures (exceptions: 782 present
in most Lumixosomatae, 7781 present in some
Platynothrus and Crotonia species). Hysteronotal gland
usually present (exceptions: abseat in Eulohmanniicae,
Nanhermanniidae, Phthirsearidac, Synichotriliinae).
Hysterorotal chaetotaxy usually hypotrichaus, with at
least seta /4 and/or 44 vesugial or absent (holotrichous
exeeptions: some Eumixosamatac, Eulohmanniidae,
some Clinolissurae), Setal file s with one or iwo setae,
if two, then at least one 0.5x length or less, of
proteronatal seta /2. Coxites rarely discrete, varying
degrees of lusion fay merge into single podosternal
“SOUTH AUSTRALIAN SOILS 4
shield, Cheliceral spatula present or absent. External
malae ¢onjunct or eoarctate. Gnathosternal fissures
absent or mentocoxal and/or dicoxal fissures present.
Adoral setac positioned so that at least gel antenor ta
qo2. Pretarsus with one, three or rarely two vlaws. Genu
1 with one (true for all species with two solenidia on
tarsus |), two or three solenidia, Tibia T solenicdia
baculiform, piliform or flagelliform. Kemora undivided.
Nymphs with similar or dissimilar facies to adult.
Remarks: Vhe Comalida (Lee 1984) is diminished to
exclude the Profissurina as cammented on under
“Systemalics” above, This leaves it restricted to two
infraorders: Mixosomatina and Holosomatiia, The
Mixosomatina and amongst the Holosomatina, the
Clinofissurac, are dealt with below. Thus, within my
study of South Australian oribate mites, three sections
(Pherenotae, Gymnonolae and Paronotae) of
Holosomatina will nor have been considered.
The synapomorphy of the Comalida is still regarded
as the canpinct or coarclate position of the external
malae, but it is now treated as convergent with thal
character state in (he upgraded Profissurina. A general
trend can be recognised in the adults to derived character
states such as a strongly sclerotized integument, fused
somal shields, fissured gnathoslernum, conspicuous
somal ridges (lamellae, pteromorphs, teeta), fewer hairs
(setae, solenidia), heteromorphie leg segments and a
more complex respiratory system (pores, Wachee). Since
a similar Lrend does not occur in the immature Sages,
they tend to be dissimilar to the adults in all bur the
more primitive taxa.
Relationships within the Comalida, which inechides
the preat majority of extant oribate species, are nol well
understood. | have assigned the advanced taxa to the
Holosomatina, which is regarded as monophyletic. The
remaining distanily related taxa are provisionally referred
to the Mixosomatina, which is regarded a5 a primitive
grade.
The Comatida is regarded as paraphyletic beeause the
Astiginata is not inevluded (see “Systematics” above).
This is preferred since it provokes the search amongst
the Comalida for the siste-group to the Asrigmalta, &
more acceptable phylogeny, since the primitive nature
of Parhypochthonius suggests that it is angestral within
a lineage including both of these taxa. If this seareh ts
not fruitful, however, iL may prove desirable to regard
the entire Comalida as a sister-group to the Astigmata
and so monophyletic.
Intraorder MIXOSOMATINA
Diuznosis: Comalida. Transverse midpadasrernal
fissure between coxifes Hand U1. Qhien twa salenidia
on gen I, if only one solenidium then either crinsverse
hysteronotal fissure TB2 present (some
Gehypochthoniidae) or omsihosterial setation reduced
(at least setae Se absent and usually only one seta /Ze—
44 i
Neomixosomatae). Adanal shrelds never merge behind
anal shields. Cheliceral spatula present ov absent-
Remarks: The Mixosomatina includes. three sections
(ELumixosamatae, Mesomixosomalae and
Neomixosomatae) much as before (Lee 1984) except that
the latter section has been extended to include (he
Trhypochthoniidae and Trhypochthoniellidae. Within
my study of South Australian oribate mites, members
of the Eumixosomatae have already been considered
under “Monofissurae” within the “Arthronotina™ (Lee
1982), but it should be noted that the recently included
(Lee 1984) Narth American Nehypachthontidae Norton
and Metz, 1980 Jacks a transverse hysteronotal fissure,
which requires a change in the diagnosis. The other two
sections are considered below, although no members of
the Neomixosomatae were found in this sticdy.
The Mixosomatina has no synaponiorphy, but is
recognisable by the primitive character state of a
podosternal lssure which allows che propodosoma to
move in relation Lo the metapodosoma. In some taxa
with a strongly sclerotized cuticle, this Mlexibility is
accentuated by a number of derived states such as in
prychaidy, when the legs are pulled up into the soma
and the proteronotum flexed downward to close on a
forward facing section of the genital shield, or when the
propodasoma can partially telescope into ihe meta-
podosama,
Relationships within the Mixosomatina are uncer Lain,
partly because some primitive character states may or
may not be secandarily reverted to in advanced taxa.
The weakly sclerotized cuticle of the Eumixosomatae
is primitive in the Parhypochthioidea and the
Nehypochthoniidae but in the Neomixosomutae it may
be eilber reverled to or there may not have been any
strangly sclerotized ancestors. Furthermore, whilst £ (Lee
1984) considered that the absence of cheliceral spatulac
was primitive but diagnostic of the Mixosomatina,
Norion (personal coramunicanon, 1982) has painted out
that cheliceral spatulae do occur in some Mixosomatina.
He has observed cheliceral spatulae in Collofinannia
gigantea and a new North American species of
Collohmanniidae. Also, their presence in
Perlohmanniidae 1s suggested in an illustration by
Grandjean (1958: Fig. SA), and [ have now included the
Trhypochthoniidae and Trhypochthoniellidac in
Mixosomatina, both of which have cheliceral spatulae.
Tragdrdh (1931b) recorded cheliceral spatulae on
Phihiracarus maculatus, but this was refuted by
Grandjean (1959) who considered the structure to be an
oncophysis. Cheliceral spatulac are considered to be
absent in all Eumixosomatae, some Mesomixosomatae
(Epilohmanniidae, Eulohmanniidac, Phthiracaridac,
Eupthiracaridac) and some Neomixosomatae
(Malaconothridae). Therefore, although primitively
absent, some taxa without cheliceral spatulac may have
ancestors that possessed them.
S OAUIS TD MLIS
(4) WRG? \iay 1488
The three sections within the Mixosomatina are
grades. The Mesomixosomatae include a number of
specialized lineages, whilst the Eumixosomatae are
apparently primitive (cettainly so in the case of
Parhypochthonius, bucpossibly EViplochihonius reflects
specialization to living in the deeper soil layers) and ihe
Neomixosomatae include advaneed lineages which may
be ancestral to (he Holosomatina, Regarding the
relationships of the Astigmata, I hald to a conservative
belief that weak selevotization in the Astigmata is an
ancestral character state and nota reflection of neoteny
as proposed by O'Connor (L984), Therefore, a sister
group to the Astigmata would have a weakly selerolized
adult, as well as lacking a rostral tectum and cheliceral
spatulac. Possibly, reduction in the fourth hysteranotal
setal rank would be the synapomorphy, placing this
sister-group within the Eumixosamatae, However, a
stringent analysis will be a necessary prelude to
developing a durable model for the areas only speculated
on here.
Section MESOMIXOSOMATAE
Diagnosis’ Mixosomatina. Hysteronotal shield
without transverse fissures. Rostral tectum present.
Gnathoslermum with A-shaped mentocoxal fissure
Adoral setae in three conspicuous pairs. Opisthosternal
setal files Se with at least one seja, #Za with at least
two setae. Protcronotal setal file « with one or two setae.
Gent | with two or rarely three solenidia.
Remarks: The Mesomixosomatae is a diverse group
of small farnilies which, allhough having a strongly
sclerotized adult culicle, have maintained flexibility by
developing various specialized ways of moving the
propodosoma in relation to the hysterosama. | have not
attempted any superfamily groupings, Even grouping the
six. families into aptychoid (Callohmanniidae
Grandjean, 1969; Epilohmanniidae Oudemans, 1923;
Eulohmanniidae Grandjean, 1931; Perlohmanniidae
Grandjean, 1954a) and ptychoid (Euphthiracaridae
Jacol, 1930; Phthiracaridae Perty, 1841) taxa ts
questionable. The piychoid families have already been
considered in this study (Lee 1981) as the Buptyctima,
The Epilohmanniidae is represented in collectians for
this study and is considered below, Although nor
represented, the Eulohmanniidae is also considered in
order Lo confirm ils grouping in this section despite
possessing same primitive character stales sugwesting i
might even be grouped with the Afissurida.
Family EPILOMMANNITDAE Oudemans
Lesseriidaec Oudemans, 1917: 78
LEpilohmanniidae Oudemans, 1923: 79
Epilohmanniidag: Grandjean, |954a: 430,
Epilohmanniidac: Grandjean, 1969: 144.
Epilolimannidae: Norton, Metz & Sharma. LOTS: 15.
‘Type-genus: Epilohimannia Berlese, 19/7b,
SAKCOPPTIFORMES (ACART) Ol
Diagnosis; Mesomixosomatae. Minute lo medium-
sized (320-800) yellow ta brown miles. Extensive non-
sclerotized cuticle al transverse midpodosternal fissure
between conites I} and lil allows prapodosoma to
partially telescope in and oul of metapodosama.
Otherwise, extensive sclerotized somal shields, with
coxifes fused to cach other, aggenital shield and usually
mentum (exception Lpilohmarnnoides yacoli). Posterior
end of avgenital and anierio: end of adanal shields
truncated or fused together so that ventrolateral
longitudinal fissure Straight. Adanal shield) merges
medially as broacl band in front of anal shields.
Cheliceral spatula absent. Palpal segments fused.so that
only 1Wo separate. Adoral seta vol bilurcate.
Hysteronotal gland present, pore apens into depression
also containing alyeolus ol relic seta 44, whilst seta /4
similarly absent, Opisthosternal setal file Sg ineludes 3-7
pairs, Opisthasternal pores Z4af and Se/ present, On
larsus | sela pd/3 reduced to scale closely associared with
distal lace of solenidium so4. On tibia 1 solenidium
Nagelliform, longer than segment.
Distribution: Possibly cosmopolitan. Wallwork
(1942b) states that Epilohmanniidae “appears to be
widely distributed through the warmer regions af the
world", Records from southern Canada (Nn) and
Moscow (Pe) could be most northerly, with records from
South Australia (Aa) being most southerly, suggesting
predominantly pantropical distribution becoming sparse
in temperate regions up to 57°N and 35°S. Outside Ant-
arctica and Subantaretic, South Ethiopian and New
Zealand-Australian may be only minor regions in which
family not represented.
Microhabitats rane widely from = grass Tools,
vineyards and forests, but possibly do not include and
or semi-arid environments.
Remarks: The Epilohmanniidae includes the
lypeeenus and Epilohmannides. Norton, Metz and
Sharma (1978) in describing lwo speeies of
Epilohimannoides also give u diagnosis lor the family
on Which the above diagnosis is based.
EPILOHMANNIA Berlese
Epilohmannia Berlese, 1917b; 176. Ivpe designation
(arwinal: “Lohmainia eylindrica Berl”.
Lesseria Oudemans, 1917: 78. Type designation
(original): by monatypy (“Lessert ssanisloi Oudms,
1915". synonym of Epilohmeannia evilindrica by van
der Hammen 1959; 34).
‘Wvpe-species: Epiohmannia cylindrica (Berlese, 1905)
23).
Diugnosis: Epilohmanniidac. Opisthoventral shield
divided into 1wo parts by Lransyerse fissure jist posterior
1a genital Shield. Genital and anal stields oblong rather
than subovals posteriar margin of genital shield and
anterior (nargin of anal shield beg broad-shouldered.
Trochanter Hd) and LY with distal axis at right angles
SOUTH AUSTRALIAN SOILS | 4°
to proximal axis, Acetabulum LV on same longitudinal
line as LIL, both divided by spurs into double apening.
Distribution: As for Epilohmanniidae.
Remarks; The more complex form of trochanter 1]
and LV and their acebatula with double openings,
Suggest that these legs can lock into either of twa basic
positions and, further, indicates that Epilohimannia may
be derived from the much less diverse (four compared
with 26 species) Epilohmunnoides with simpler posteriar
trochanters and acetabula.
Balogh and Mahunka (1979) have distinguished
within Epilohmiennia a Subgenus, Sinolohmuannia, by the
presence of a spine-like seta d associated with the
solenidium on tibia VV. The eviindrica-complex of
Schuster (1960) can be regarded as equivalent to the
nominate subgenus, A more conspicuous character state,
the position of the acetabulum on coxite TV, has nat
been referred 10 in grouping species, Both the species
considered below have the acetabulum IY on the
posterior margin of conite 1V, bul it may be near the
anterior margin or in between, However, the included
species are similar and, untortunately, the specimens
collected in this study belong to species with a number
of even more similar subspecies. Since | am not
confident about grouping them in any of the subspecies,
new subspecies have had to be established. Because of
the similarity of the established subspecies, relerences
to them are included under the new subspecies.
The illustrations of one specics in this paper are
intended to show character states delineating the family
and genus rather than the subspecies.
Epilohinannia cylindrica (Berlese) media n.ssp.
(Figs 3-5)
Lohomannia evlindrica Berlese, L905; 23.
Lesseria szanisloi Qudemans, 19)7: 78.
Eptlohmanma ssunislor Schuster, 1960) 202 (inchiding
mininid, fy. 205).
Epilofinannia cylindricu Aoki, 19635e: 309.
Epilohinannia evlindrica Bayoumi aud Mahunka, 1976;
6.
female
Dull (slight greasy shine) chestnut brown, slightly
darker cheliceral extremities and extermal malag,
oxtensive pule straw-coloured cuticle hetween
proterosoma and hvsterosoma when extended. Most of
hysterosternum with sparse punela, mited in other
parts of venter to areas illustrated (Fig. 3). Idiosamal
length (abutting margiis to midpodosomal fissure,
possible extension SQ, retraction 30) 465 (10, 415-525);
appendage lengths (for 475)—ch 44, pa 50,7 195, 21175,
11 N95, 7 275: femur breackths—pa 7.5, £20, 1/20, LIT
15, 7b 205 broadest seement breadth—/ eenu 32.5, 7
venu 30, //f trochanter 25, JW trochanter 37.5.
Appendage sclae: clr (2). pu (3-8), 7 (1-4-5-5-17), IT
(14-5-5-12), 7277 (2-3-4-4-10), 71 (2-3-4-4-9). Solentchia:
7 (2-1-3), 22 1-1-2), (7 1-1-0), 2 O-L-O), Solenidia of
46 REC. 8. AUST. MUS. 19 (4); 39-67 May, 1985
FIGS, 1-5. Papillacarus and Epilohmannia: 1-2, Papillacarus pseudoaciculatus Mahunka, tritonymph; 1, gnathosternum; 2, right chelicera,
anterior surface; 3-5, Epilohmannia cylindrica (Berlese) media n.ssp., female; 3, idiosternum; 4, gnathosternum; 5, right chelicera,
anterior surface.
SARCOPTIFLORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 4 Ay
genua and tibiae flagelliform with minute, closely
associated seta d (possibly partially coupled—see
Epilohmannoides terre: Norton, Metz and Sharma,
(978), on legs HL and LV seia d only recognised when
separated from solendium by preparation of specimen,
Solenidium so4 (so3 regarded ay absent) on tarsus [also
flagelliform with minute seta pd3 clasely associated with
distal face. Other solenidia on tarsi | and 11 ceratiform.
Chelicera without spatula, small process (Fig, 5)
posterior to movable digit probably chitinous plate
representing reduced trochanter. Adoral setae gol and
ao2 bifureate. Genital shield setaer 5/g, 3Ze.
Tarsus IY with length 5,5x breadth, and setae vl and
v2 (see Schuster 1960; Fig. 5 a- K) and £2) spine-like,
but less robust than for & evlindrica minima (see
Schuster 1960; Fig. $b), Seta v2 nearer to vl than to v3.
No eggs observed, specimens assumed female since,
amongst eight pairs of setae on posilor, three pg pairs
well apart [rom rest, two dg pairs longer, One to three
boli in cach specimen, light to dark brown
heterogencously granular contents without recognisable
cellular structures,
Material examined: Seuth Australia (Aa). Holotype
female (N19832), nine paratype females
(N19833-N19831]), bases af grasses and plantains,
Glentharne, 12.6,1974, D.C. Lee.
Distribution: E. cylindrica (subspecies media only
known from South Australia), Termmessee (Na); Tehad
(Ee); Austoia, Egypt, Hungary, Italy, Moscow, Sicily,
Spain (Pe); Turkestan (Ps); Philippines (Qm); South
Australia (Aa); Hawaii (Ap), South Australia:
Glenthorne, cultivated pasture, 10 (2/8).
Remarks: ©. cylindrica media \ies between Ec.
tviinadriva and E. o minimain a gradation of character
states, As indicated by Schuster (1960), B. ¢. cylindrica
is 435-573 long, has tarsus LY length 5.6x breadth, with
selae vl and v2 setose, whilst 2. c. minima is 370-440
long, lias tarsus LY length 3,6x breadth with setae vl
and v2 robust and spine-like (more so than E. &. medica).
Epilohinannia pallida Wallwork australica n.ssp.
(Fig. none)
Epilohinanna pallida Wallwork, 1962b: 689.
Spilahindanta pallida pacifica Aoki, 16Sc: 312.
&piehmanma pallida cegvplica Bayoumi & Mahunka,
L976; 8.
(pilohmannia pallida indica Bhattacharya & Banerjee,
F980; 19.
Epilohmannia pallida americana Balogh & Mahunka,
I9BL: 59,
Female
Dull brown, paler but similar to &. evlindrice.
Idiosomal length (abutting margins to midpodosomial
fissure, possible extension 42.5, retraction 25) 400 (2 cx
Piccaninnie Ponds, 420-425; 3 ex Chambers Gully,
350-405); appendage lengths (for 405)—ch 40, pa 45,
#155, JT 130, 177 140, 7¥ 190; femur breadths—pa 10,
722.5, 1720, 7/1715, JV 15; broadest segment breadths—/
genu 27,5, // genu 25, /// trochanter 17.5, /¥ trochanter
25,
Appendage setae: ci 2, pa (3-8), J (1-3-4-5-15), i
(1-4-4-4-12), /// (2 or 3-3-3-3 or 4-10), [W (2-3-3-4-9),
Compared with £. ¢. media seta ve absent on femur |,
genua | and II, whilst seta vp absent on genua III and
IV. On genu | seta v small, Solenidia similar in number,
shape and size ta E. ¢. media, except on genua II], 1V
and tibiae JJ, 1V, solenidia relatively longer.
Chelicera without spatula. Adoral setae go] and #o2
bifurcate. Genital shield setae: S/z, 3Zg,
Characters previously used to delineate subspecies as
follows, Hysteronotal setae sparsely ciliate and tapering,
scia ZI posterior to /I, distance J/I-Z7) subequal to 14/1,
Anterior apodeme to coxite | curves back to single
apodeme separating coxites | and I] without connecting
ridge Lo partner, conil¢ seta HY 2 subequal in Jength to
fi3, lateral margin of coxite 1V straight. Opisthasternal
seta Se? level with Jel, setal file Ze separate trom file
Je by distance subequal to length af Zel, slit pore Sa/’
inclined at angle less than 25° from transverse axis.
Distal end of trochanter LI without dorsal spur on
tarsus LV seta v2 nearer vl than v3. —
No eggs observed, specimens assumed female sirice
amongst eight pairs of setae on positor, three pg pairs
well apart from rest, two dg pairs longer, One or twa
boli in each specimen, light to dark brown
heterogeneously granular contents without recognisable
cellular structures.
Material examined: South Australia (Aa). Holotype
female (N198312) and two paratype fenyales (N198313,
N198314), grass and mass or litter under Lycalyptus
viininalis, Chambers Gully, 12.6,1974, D, C, Lee, Twa
paratype females (NI98315, N1983J6), litter and sparse
grass under Acacia sophorae, Piccaninnie Ponds,
20.8.1975, D. C. Lee.
Disiritution: &£: pallida (subspecies australica only
known fram South Australia). Texas, Minnesota, North
Dakota, South Dakota (Na); Paraguay (NTb); Ghana
(Ew); Egypt (Pe); West Bengal (01); Hong Kong (Qs);
South Australia (Aa); Hawaii (Ap), South Australia:
Chambers Gully, savannah woodland, 3 (2/8);
Piccaninnie Ponds, coastal cloged-scrubland 2 (2/8),
Remarks: E. pallida is not very different from E
cvlinedrice (especially if the comparison is made to &
c ining) except that seta d2 on tarsus IV is
conspicuously spine-like.
The other five subspecies of £ pallida are very similar.
& p, australica can be distinguished fram these by the
character states listed above On the basis of these il is
most similar tod. 7 indica and & op, pacifica, bur ditters
in [wo states regarded as important: the shape ol the
anterior coxjte apodemes and. the position of seta v2
on tarsus JV. One character, the chaetotaxy of the genilal
4h Rie. S: AUST, MUS. J9 (4); 29-67
shield sometimes used to distinguish subspecies, has not
been listed as such because of its intraspecilic variation
(see MeDaniel and Bolen 1983).
Family EULOHMANNIIDAE Grandjean
Ewlohmanniidae Grandjean, L931: 144
Eulohmanniidae: Grandjean, 1954a; 429
Woe-genus: £iv/ehmuannia Berlese, 1910.
Diuenosis: Mesomixosomatae Medium-sized
(650-700) siraw-coloured mites. Extensive nonsclerotized
cuticle at transverse midpadosternal fissure between
coxites It and [ff allows propodosoma. to partially
telescope in and out of metapadosoma. Otherwise,
extensive sclerotized somal shiclds, with coxites fused
to each other and aggenital shield. Posterior end of
aggenital and anterior end of adanal shields taper so
that ventrolateral longitudinal fissure V-shaped. Adanal
shield not merged medially in front of anal shields,
Cheliceral spatula absent. Palpal segments fused so four
separate. Adoral seta aol not bilurcate. Hysteronotal
gland absent. Hysteranotal chaetotaxy holotrichous,
fourth setal rank not reduced, Opisthosternal setal file
Se bypertrichous and dispersed, 15 or more setae,
Opisthosternal pore Suf present, Zaf absent, On tarsus
[, seta pa3 setose and well separated from solenidium
so4, On tibia 1, solenidium piliform, shorter than
seement.
Distribution, Holareue.
Remarks: The Eulohmariniidae includes one species,
FEulohmunnia ribagai Berlese, which has been described
a number of times, perhaps the most accurate ancl
comprehensive illustration being that by Lebrun and
Wauthy (1981: Fig, 2). But there appears to be no
detailed description of the gnathosternum, Since the
species has a number of primitive character states such
as a holotrichous hysteronotal chactotaxy and no
hysteronotal gland, it was necessary to check the
enathosternum to evaluate whether ar nat the species
was correctly grouped in the Comalida. This was made
possible by a gift of two specimens from Dr, Georges
Wauthy.
Bulohmannia ribagai Berlese
(Fig. none)
Lohmannia (Eulohmannia) ribagai Berlese, 1910; 223.
Adult
Idiosomal length 655 and 695. Cheliceral spatula
absent. Mentocoxal fissure present, A-shaped. External
malae conjunet, distally with two lateral robust refractile
teeth and median hyaline flap. Gmathosternal chaetoaxy
reduced: 3zo, 2cy, Jpo. Gnathosterniim elongate much
Via, LYS
as Perlohmannia dissimilis see Woolley 1969; Fig, 10).
Yarsus | with 23 setae and three soleridia, The shape
and positioning of plasmic seta dl and three solenidia
much as Perlohmannia dissimilis (see Grandjean 1958:
Fig. 6C}; seta dl short with distal knob, solenidia well
separated from setae.
Material examined: Fwo adults (N1983103, N1983104),
litter and humus, Lauzella Wood, Belgium, 4.1980, G,
Wauthy.
Remarks: Having seen details of the gnathosternum,
there is no doubt that Eulohimannia is well placed in the
Comalida, and (heréfore that some apparently primitive
character states, such as the absence of hysteronotal
glands and holotrichous hysteronotal chactotaxy, are
derived reversals. Although Au/ohmannia in general
form and extent of selerotization resembles
Epilohmannia, it exhibits similarities to Perlohimannia
of the gnathosternum and tarsus | which suggests they
may be regarded a Sister-zroups.
Section NEOMIXOSOMATAE
Diagnosis: Mixosomatina, Hysteronotal shield
without transverse fissures. Rostral leclum present.
Gnathosternum either without fissure or A-shaped or
what may be transverse linear mentacoxal fissure
present. Acoral setae present or absent, Opisthosternal
setal file Se absent, JZa usually with only one seta
(exception: Mucronothrus—2JZa). Proteronotal setal tile
s absent, plasmic seta z2 may be reduced and setiform,
On genu J, one solendiurn,
Remarks: The Neomixosomatae was established (Lee,
1984) to include the Malaconothridae which has a
conspicuous fissure between coxites H-HI and Jacks a
vheliceral spatula, but is in seme ways similar Lo the
Nothraidea, the most primitive of the Holosomatina,
As indicated in the above “Remarks” under
Mixosomatina, it has now been established that
cheliceral spatulae occur on some of its members. This
means that the Trhypochthontidac: Balogh, 1972 is nol
excluded from the Mixosomatina by possessing
cheliceral spatulae and so the relevant literature has been
examined, even though no representatives were collected
in this study, The result is that a number of changes
are made to the classification, most genera of
Trhypochthoniidae including the type-genus being
included in the Mixosomatina, On the other hand, two
genera have been grouped in a new family retained
within the Holosomatina. The three tamilies
(Malaconothridae, Trhypochthoniidae and
Trhypochthoniellidae) now in the Neomixosomatae are
briefly commented on below,
1 wish to emphasize the provisional nature af this
classification and that | have ignored some character
states considered in the more comprehensive studies of
relevant families by Knulte (1957) and van der Hammen
(listed by van der Hammen, 1959) because they are only
SAKEOPLIPORAMIES (AC ARE) OF SGUTID AUISTISAL DAN SCHLS 4 ay
known lor a lew species. Since those sludies, lour genera
have been added to the eight genera (included by
Balogh, 1972 within the ‘Irhypochthoniidae and
Malaconothridae) and, both before and after the studies,
the majority of species deseriptions have not been
extensive enough, in same cases with debatable generic
combinations, Therelore, g thorough reinvestigation of
established taxa is needed before reliable diagnoses can
be given lo them, and their considerable relevance to
the classification of Comalida understood.
amily MALACONOTHRIDAE Berlese, 1917b
Diagnosis: Neomixosomatrae, Cheliceral spatula
absent, Adoral setae reduced to two pairs or all absent.
Gnathosternal fissure absent or, if part ot mentocaxal
fissure present, lateral parts never meet at mid-point.
Proteronotum without setal file s and seta 22 setiform
and less (han 0,5 leneth /2, Hysteronoral seta 22 nearer
73 than J/2. Coxite LL posterolateral corner extended as
carina with backward facing socket. Opisthosiernal setal
file Sa with three setae, On tarsus L, three solenidia
chistered between level of setae pd2-pd3. On tibia l, one
solenidium.
Remarks: The posterolateral corner of coxite IT may
act as a socket jnto which (rochanter IIE fits, possibly
functioning as a pivot when the propodosoma flexes to
one side. Such a movement could be limited in some
species by a large rectum, projecting laterally from just
pasterodorsally to acetabulum UH, hitting a similar
smaller tectum anterior fo adetabulum ITI.
The Malaconothridae, as by Balogh (1972), includes
the following four genera: Fossonatirus Hammer, 1962;
Mataconothrus Berlese, 1905; Trjmalaconothryus Berlese,
I9{7e, Zeanothrus Hammer, 1966,
Family TRHYPOCHTHONTIDAE Willmann, 1931
Diaenesis; Neamixosomatae, Clreliceral spatula
present. Adoral selae with three pairs present,
Mentocosal /issure present, A-Shaped, Proteronotim
with seta 22 club-like os, if setiform, either subequal in
length to /2 or, if less than OS» Jength 2, seta.s present,
Hysteronotal seta 42 nearer /2 than 23. Coxite Ll
posterolateral corner sometimes extended as carina with
backward facing socket, Opisthosternal seral file Se with
two or three setae, On tarsus |, rhree solenidia usually
widely spaced between level of setae pd2-pdl4 (exception:
Hydronothrus), On tibia 1, usually two solenicia
(exceptions: Ardronotiirus and Muacronothyius with one
solenidium).
Renuarks: The eight genera included by Balogh (1972)
in the Trhypochtheniidae are decreased in number by
subdivision either into the Irhypechthoniellidae or the
Allonothridae (Clinofissurae). The following four
genera are scill included Archegozeles Grandjean, 1931;
Hlydronoihryus: Aoki, 1964; Miteronuthirus Tragdrdh,
W3le; Trhypochthonius Berlese, 1905, The conservative
eniphasis on notal setae results in J¥ydrenothrus being
grouped in this family, although the leg [ setation is
similar to that al’ the Malaconothridae, Archegazetes
differs considerably from the other three genera,
especially with regard to its lang leg tarsi-and seta 72.
But some recarded differences, the absenee of both a
cheliceral spatula and a fissure between coxites LH and
Hl (Beck, 1967), are regarded as errors. The presence
of a cheliceral spatula and a coxite H/11) fissure was
recorded by Grandjean (1959) and van der Hatmmmen
(1955) and has been confirmed by Dr. R. A. Norton
{personal communivalion, 1983) of specirnens from
Mexico, Panama (N'Tm), Brazil (NTb) and Malaysia
(Om),
Family TRHYPOCHTHONIELLIDAE Knulle, 1957
Diagnosis: Neamixosomatae. Cheliceral spatula
presen|. Adoral setae with three pairs present.
Gnathosternal line (7fissune) present, transverse, linear.
Proteronotum with seta 52 club-like or setiform and less
than 0.8 length j2, Hysteronotal seta 72 neurer J2 than
73, Coxite 1] posterolateral corner not extended as
caring. Opisthosternal setal tle Sw with two setae On
tarsus 1, three solenidia widely spaced between level af
setae pd2-pd4, On tibia [, one or two solenidia.
Remarks: Trhypochthoniellidae includes two genera:
Trhypochthoniellus Willmann, 1928; Afronuthrus
Wallwork, 1961, The grouping together of these genera
is mainly based on an assumption about ynathosterrial
structure, They can be regarded as having a quadrate
mentum; & “hypostome rechteckig’’ Tor
Trhvpochthoniellus (Knulle 1957: 151) or as illustrated
for Afronothrus (Wallwork 196]; Fig. 7, Hammer 1972:
Fig. 19a). The hne illustrated on Afronvihrus could not
be the dicoxal fissure which delineates (he atucrior
margin of the “quadrate mentum” of advanced
Holosomatina. It is therefore assumed that it is a
mentocoxal fissure and there may be a valid quadrate
mentum, but carelul evaluation of what the drawn line
represents 1s sull needed.
Infraorder HOLOSOMATINA
Dyaenosis: Comalida, Coxites fused together into a
Single shield. On genu 1, one solenidium. Tibia 1
salenidia baculiform, piliform or, if Magelliform, closely
associated with similar seta. No transverse hysteronotal
fissures (exceptian;: ?7'BL present in some Crofonia and
Plarynothrus species). Usually two setae in file /Za,
always when adanal shields not merged behind anal
shields. Cheliceral spatula present.
Remarks: Vhe Hoalosomatina is regarded as
monophyletic, whilst its tour subordinate sections
(Clinolissurae, Pherenolac, Gymnonorae and
Poronotae) are likely to be grades. The lineayzes have not
au Ret:
been identified, although the Poronotae (being the most
derived) could be a clade. Only the mast primitive
section, the Clinofissurae, is considered in detail below.
A major taxon, Lhe Circurndeliscentiae Grandjean,
19§4a (= either the Euoribatida Balogh and Mahunka,
1979 or a major part of the Brachypylina: Balogh, 1972
or the Pherenotae, Gymnonotae plus Poronotae), 15
regarded as monophyletic bul remains unnamed in this
classification as before (Lee 1984). This is because jt
approximates to the Holosomatina (excluding only the
six families of the Clinofissurae) arid because the fusion
of coxites inta one shield (which is the synapomorphy
of the Holosomatina) is regarded as a predominant step
that precludes conditions such as pty¢hoidy and is a
prelude ro other states correlared with a thick, rigid
idiosomal integument, such as a circular hysterosomal
dchiscence line, a dicoxal enathosternal fissure and a
tracheal system opening ventrolarerally between legs 1I-
Lil or in the acetabular cavities of legs | or Ill, The
presence of a dicoxal gnarhosternal fissure may be
correlated with the circular hysterosomal dehiscence line
of the “Circumdehiscentiae’, but a Notrrus species is
clearly described by Tragardh (193la) as having both
a dicoxal and a mentocoxal gnathosternal fissure,
although this needs confirmation since it is difficult to
conceive the function of such a double-jointed system,
The presence of a tracheal system appears to be the
synaparmorphy of a slightly smaller taxon than the
“Circumdechiseentiae’ since it is absent from the
Hermannicllidac:
Section CLINOFISSURAE
Diagnosis: Holosomatina. Gnathosternum usually
with A -shaped mentocoxal fissure present (exceptions:
no fissure in Allonothridae, possibly both mentocoxal
and diconal fissure present in one Notfirus species). If
adunal shields separate from hysteronotal shield (i,¢,
exclude Nanhermanniidae) they do not fuse behind anal
Shields. Genua subquadrangulate in outline and similar
in Size Lo Libiae, Hysterosomal dehiscence line T-shaped,
midnotal.
Remarks: The Clinafissurae is diagnosed by character
states primitive to the Holosarpatina. It is equivalent
ia the majority of the Nothroidea plus the
Nanhermannioidea and Hermannioidea in the previous
classification of Balogh (1972), when the latter two
sUperfamilies were included in the higher oribate mites
(as “Brachypylina"), although they are excluded from
the smmilar Circumdehiscentiae Grandjean, 1954a. On
the other hand, this js the Jirst time thal members of
the Nothroidea have been included amongst the higher
Gribare mites (if considered as referring to the
Holosomatina),
The following six families are included in this Section:
Allonottividae mf. Carnisiidae Oudemans, 1900;
Crotoniidae Thorell, 1876; Hermanniidae Selinick, 1928;
Nanhermanniidae Sellnick, 1928; Nothridae Berlese,
SAMS E MUS TY (dy: d9-87
Vein, 1985
1885. AU but the Allonothridac and Nothridac are
represented in this study and the Allonothridae is
considered further as if has to be defined. The
Allonothridae, Camisiidae, Crotoniidae and Nothridae
can be grouped in the Nothroidea Grandjean, 19544,
hut superfamilies will not be considered here,
Family ALLONOTHRIDAE nf.
Type-genus; A/lonothrus van der Hammen, 1953
Diagnosis: Clinofissurae. Gnathosternal fissures
absent, or, if part of mentoenxal fissure present, lateral
parts tever meet at mid-point. Three pairs of
adoral setae, Rostral tectum without miediat mcision.
Proteronotal plasmic setae 72 at least 2x as long as
distance /2-22, and setiform or slightly swollen distally.
Hysteronotal seta J4 absent. Hysteronotal gland present.
Natal setae zl and /5 not on apophyses. Coxtte selae
not hypertrichous (3-1-3-3). Setal file Sg absent,
combined setal file /Z%2 on median margin of genital
shield. Idiosoma not almost covered in continuous
shield, No separate preanal shield. Palp tarsus with nine
setae, No dorsolateral supernumerary setae on tarsus |,
Nymphs without small shields around hysteronotal setal
bases.
Remarks: The Allonothridae includes the following
twa genera: Allonoihrus yan der Hammen, 1953;
Pseudonothrus Balogh, 1958, The genera are similar to
each other and were included in the Trhypochthonidae
(Neomixosomatae, see above), but because of the
apparent fusion of all the coxites into one shield they
are now grouped in the Holosomatina as a Hew family,
The Allonothricdae exhibits similarities to both the
Neomixosomatae and Clinofissurae. suggesting that they
belong to the same lineage, but much more data is
needed before a cladistic classification can be proposed
within the Comalida.
Family CAMISIIDAE OQudemans
Camisiidae (part) Oudemans, 1900: 142.
Camisiidae (part): Sellnick, 1928: 18.
Camisiidae: Grandjean, 1954a: 431.
Camisiidae (part): Sellnick and Forsslund, [9S4: 473.
Camisidac: van der Hammen, 1959: 65.
Type-gerus: Camnisia von Heyden, 1826
Diagnosis: Clinofissurae. Gnathosternal A-shaped
mentocoxal fissure present. Three pairs of adoral setae.
Rostral tectum without median incision, Proteronotal
plasmic seta z2 may be reduced and globular but never
enclosed in bothridium, if filamentous length 0.75x-1.5x
distance 2-22, Hysteronotal sela /4 present or absent,
Hysteronotal gland present. Notal setae sl and J5
sometimes on apophyses. Coxite setae not hypertrichous
(3-1-3-3 or 4). Coxite shields usually not merged with
SARCOUTIORMES (ACARD Ol
apgenital shields. Setal file Sg includes two setae,
combined setal file /Z¢ on median margin of genital
shield. Idiosoma not almost covered in continuous
shield, Discrete preanal shield usually as wide as anal
shield, Palp tarsus with seven setac. Tarsus | with four
or fewer dorsolateral supernumerary setae, Nymphs
without small shields around hysteronotal setal bases.
Distribution: Possibly cosmopolitan, greatest diversity
in temperate regions, represerited in tropical montane
ar Oceame regions,
Remarks, Camisiidae are smail to gigantic (530-1225)
dull brown mites, usually covered in cerotegument, thick
in parts and with adhering detritas and fungus.
Camisiidae has been considered as synonymous with
Nothridae, but both names have been in use since
Grandjean (1954a) separated them. The family is now
still as regarded by Balogh (1972) except that
Austronathrus is grouped ini Crotoniidae (Ramsay and
Luxton 1967), Comprehensive descriptions have been
made for Cantisia (Behan 1978; André L980) and
Platynothrus (Fujikawa 1982),
The following four genera are included in Camisiidae;
Camisia von Eleyden, 1826; Heminoihrus Berlese, 1914;
Neonothrus Forsslund (in Sellick and Forsslund 1955);
Pluiynothrus Berlese, 1914. Heminothrus and
Neonothrus are very similar to Plat\nothrus,
Heminothrus being established earlier (p. 38) in the same
reference.
PLATYNOTHRUS Berlese
Platynethrus Berlese, 1914; 99. Type designation
(original): “Nathrus palliatus K.(=N. bistriatis K.)".
Type-species: Plawvnathrus peltifer (Koch, 1839: 29/9),
Diagnasis! Camisiidae. Proteronotal plasmic seta 22
vermiculate, usually slightly dilated and ciliate distally,
One seta in file s, Seta 1 without conspicuous apophysis
(at most, shorter than 0.5x distance zl-zl), Bothridial
cavity with only one side pocket situated near base of
seta z2.. Hysteronotal seta /4 absent or minute, Seta J5
without conspicuous apophysis (at most, shorter than
its own diameter). Two setae in file Sg on inner margin
of aggeriital shield, Two setae in file JZa. Coxites |, UI,
{]] merge across midsternal line, coxites |V may be
partially or completely separated from each other by
fissure. Leg [| with six solemdia (1-2-3),
Distribution; Widespread within temperate regions,
usually montane or oceanic in tropics. Canada,
Greenland (Na): Argentina, Bolivia, Chile, Peru (N'Te);
St. Helena (Es); Finland, Norway, Sweden, U.S.S,R.,
other parts of Burope (Pe); Japan (Ps); Himalayas (Oi);
South Australia (Aa); New Zealand (An); South Georgia
(ACS).
SOUTH AUSTRALIAN SOUS 4 31
Remarks: Major works on Platynothrus are included
in studies on the Swedish fauna (Sellick and Forsslund
1955: 513), Berlese’s collection (van der Hammen 1959:
71), Himalayan collections with a key to species (Aok|
1965a; 290) and the northern Japanese fauna (Fujikawa,
1982; 279), Balogh (1972) regards Platvnothrus as having
one pretarsal claw so his keys do not work for species,
Such as the one deseribed below, which have three
pretarsal claws. Heminothrus and Neonothrus are very
similar to Platvrothrus, and might be grouped in this
genus, Platynothrus includes seventeen species and two
of these have a subspecies.
Platpnatheus breviserosus n.sp,
(Figs 6-12)
Female
General appearance und measurements: Red-brown,
covered in cerotegument, thick with adhering detritus
and fungus laterally anc posteriorly on hysterosoma and
proximally on legs. Notal minute pits and lateral low
bumps distributed as illustrated (Fig. 6), whilst all shields
coyered with fine punctuation except on rostral tectum.
Legs similar sculpturing, mainly proximal, ventral and
sometimes anterior on individual sezments. Setae; claws,
external malae, cheliceral digits clear or light brown and
refractile, Idiosomal length IW40 (1); appendage
leneths—ch 65, pa 100, 1 580, 11 490, TH 470, FV 615;
femur breadths—pou 25, 7 75, 7775, fi 65, IV 62.5.
Prosternum: Lateral refractile half of external malae
in vertical plane when unsquashed (ic. not fattened as
illustrated, Fig. 8) and bears two rows of ciha on dorsal
surface, Adoral setae aol blunt-ended- Setae pel and c2
in shallow, punctated depressions. Coxites all fused inta
ane shield, but clearly delineated by grooves, and
separate from aggenital shields,
Proferonotum: Seta 22 vermiculate, but slightly dilated
and ciliate distally, Bothridial cavity with only one side
pocket just median to base of seta 72, so posterior
median wall forms unbroken arc. Surface has central
flat-topped mound with two mounds behind ti, and
mounds above acetabula for legs,
Opisihasternum: Qpisthaveniral shield separate from
coxites, without protrusions [ram inner margin to bear
Sg setae. Preanal shield abuts mare closely on to anal
shield than illustrated (Fig. 7) and appears foreshortened
Sifice extends upward in vertical plane. Opisthosternal
selae in file Sa blade-like, with hyaline flaps on setal core,
Genital shield with anteriorly twisted downward median
margin bearing setae (Fig. 7) rather (han median ridge,
also transverse central unpigmented zone. Twenty-three
marginal /Ze setae Mattened with slightly ragged, blunt
ends. Pores Zaj and Saf present.
a
t+
REC. S, AUST, MUS, 19 (4): 39-67 Mery, 1985
i
teri”, jf
.¢
ove
cio the
Oat tae
> Ee
eo felnsted foe."
. aw!
FIGS. 6-9. Platvnothrus brevisetosus nsp., female; 6, notum; 7, idiosternum; & gnathosternumy; 9, right chelicera, anterior surface,
SARCOPEIORMES FAC ART Ob
Aysteronoium: Fissure runs transversely along first
setal rank and backward along setal files Z and 5 (SI,
§2, Z3, Z4) possibly representing longitudinal pleural
fissure, Setae 72, 22, J3 lic between median ridge and
lateral furrow. Many hysteronotal setae appear bladelike
with hyaline flaps to setal core and marginal cilia, not
always illustrated (Fig. 6). Seta /4 apparently present
but minute,
Appendages: Chelicerae relatively small, On fixed digit
five teeth including distal point; anterior and posterior
teeth in twa pairs. Seta cA) originally present, but both
lost during dissection. Setae: ch (2), pa (1-0-3-7), 1
(0-10-5-6-28 or 29), /7 (1-10-5-6-23), TM (4 or 5-6-5-6-23),
fP (0-S-4-6-23). Solenidia: pa (0-0-1), J (1-2-3), {7 (1-1-2),
Hi (1-1-0), 4" (1-1-0), Pretarsus with three subequal claws,
lateral claws with inconspicuous dorsal cilia file,
Terminal plasmic setae on palp tarsus spine-like, in
recess. Solenidia baculiform, relatively short never 2x
length of associated seta, longest on genu J. Tarsi all
with supernumerary setae. Setae colourless, on tarsi have
strongly refractile bases bur hyaline tips.
Somal inclusions’ No eggs or boli, Ovipositor
involuted, if extended would be about 3x length of
genital shield, Breadth slightly less than that of one
genital shield. Bears cight pairs of setae, two dg pairs
longer than seiae /Zg, three pg pairs more than length
of genital shield away trom three /g pairs.
Mule
Unknown,
Material examined: Holotype female (IN198328), litter
under Pinus pinea, Knott Hill Forest, 22.35.1974, D. C.
Lee.
Distribution: Souwh Australia—Aa: Knott Hill,
cultivated pine forest, 1 (1/2),
Remarks: P. brevisetosus, with three pretatsal claws,
short hysteronotal sctae and well-spaced setae J2, keys
to 2 altimonianus Hammer, 1958 [rom Bolivia in Aoki’s
(1965a) work as would P ferniclava Hammer, 1966 from
New Zealand which is possibly even more similar. P
brevisetosus is distinguishable from these two species by
the presence of a rudimentary /4, hysteronotal fissures,
large size, blade-like hysterosomal setae and stout setae
fl and zi.
Family CROTONIIDAE Thorell
Crotonoides Thorell, 1876; 452-508.
Holonothridae Wallwork, 1963; 727.
Crotoniidae: Ramsay and Luxton, 1967: 479
Type-genus: Crotonia Thorell, 1876,
Diagnosis: Clnofissurae, Gnhathosternal A-shaped
mentocaxal fissure present, Three pairs of adoral setae:
SOULE AUSTRALIAN SULS 4 52
Ros(ral tectum without median incision. Proteronotal
plasmic seta 22 reduced, globular and enclosed within
bothridium. Hysteronolal seta J/4 present,
Hlysteronotal land absent. Notal serac z) and J5 usually
on conspicuous apophyses, Coxife / or // setae
sometimes hypertrichous but not /277 and /¥ (3 or 4 -
| or 2-3-2 or 3). Coxite shields merged with aggenital
shields. Setal file Se includes two or three setae,
combined setal file /Zzg on median margin of genital
shield, Idiosoma not almost covered in continuous
shield. Diserete preanal shield as wide as anal shield.
Palp tarsus with nine setae, Tarsus / with five pairs of
dorsolateral supernumerary setae. Nymphs with small
shields around hysteronotal setal bases.
Distribution: Southern temperate regions as for
Crotonia. Austronathrus from New Zealand,
Holonorthrus trom New Zealand and Macquarie [sland
(An, Sa).
Found in plant litter, and on lichen, liverworts, moss,
ferns and above-ground parts of shrubby plants.
Remarks: Crotonudae are large to gigantic (801-1500),
dull, dark brown mites, extensively covered in
cerolegument and detritus which may be accompanied
by nymphal skins and fungi. The unique structure of
the bothridium enclosing a globular 72 is regarded as
the family synapomorphy.
The following three genera are included in
Crotoniidae: Austronothrus Hammer, 1966;
Holonothrus Wallwork, 1963; Cratonia Thorell, 1876,
CROTONIA Tharell
Westwoodia Pickard-Cambridge, 1875; 383-390. Type
designation (original): “Westwoodia obtecta sp.nov.’
Crotania Thorell, 1876; 452-508. Type designation
(original): “Crotania obtecta (Pickard-Cambridge,
1875)”.
Acronothrus Berlese, 1917a: 65. Type designation
(original); “Nothrus (Acrenothrus) cophinarius
Mich. 1908”,
‘Type-species: Crosonia ohtecta (O. Pickard-Cambridge;
1875; 386).
Diaznosis: Crotoniicae, Hysteronotal setation reduced
to thirteen pairs (/2, J3, 241 missing). Bases of setae J]
and Z2 without connecting ridge. Rostral tectum with
single prominence bearing both setae /l. Setae 7] on
separate conspicuous apophyses, Selae /5 (and other
posterior setae) on separate apaphyses or on single
median branched apophysis. Posterior margin of coxites
fk in smooth are not broken by median notch.
Opisthosternal file JZu includes three setae.
Distribution, Widespread in southern temperate
regions (NTe; Be, Es; Aa, Ap, An; Saj—see Hammer
and Wallwork (1979) for a teview, Nate that South
34 REC, S. AUST. MUS. 19 (4): 39-67 May, 1985
FIGS. 10-12. Platynothrus brevisetosus n.sp., female; 10, leg I, dorsal surface of genu, tibia and tarsus; 11, leg tarsi, posterior surface of distal
ends; 12, leg III, posterior surface of trochanter and femur.
SARCOPTIFORMES (ACARI) OF
Ethiopian record is St. Helena. More recent records from
Tierra del Fuego (Mahunka 1980b) and New Zealand
(Luxton 1982).
Usually found in moist forest or heath in the plant
litter, moss or on the above-ground parts of shrubby
plants (Luxton 1982).
Remarks: The thick layer of cerotegument, detritus
and nymphal skins obscures the dorsal features of
Crotonia mites as illustrated by photograph of C. obtecta
(O. Pickard-Cambridge) (Ramsay and Luxton 1967: Fig.
8). Crotonia may currently include twenty-four species,
and Luxton (1982) provides a key for fifteen adequately
described species.
Crotonia jethurmerae n.sp.
(Figs 13-19)
Female
General appearance: Dark brown, thick cerotegument
with adhering detritus (including charcoal, left by forest
fires), fungal hyphae and tritonymphal integument.
Encrustation forms anterior protrusion encompassing
setae zl and posterior protrusion encompassing setae
J4, J5, Z5, S5 (small on some specimens, then bilobed).
Notal minute pits and low bumps distributed as
illustrated (Fig. 13). Setae, claws, external malae,
cheliceral digits clear or light brown and refractile.
Idiosomal length 1400 (3, 1390-1405). Proportions of
appendages similar to those for male holotype (see
below).
Prosternum: Lateral refractile half of external malae
in vertical plane when unsquashed (i.e. not flattened as
illustrated, Fig. 15) and bears two rows of cilia on dorsal
surface. Three adoral setae, aol bifurcate, flattened, one
branch forming denticulate hyaline flap. Coxites all
fused into one shield, partly delineated by grooves but
leaving broad, flat mid-sternal zone, fused to aggenital
shields. Hyaline cap on coxite seta may be broken off,
leaving slim setal core. Coxite setae ///3 and /V1 on
apophyses.
Proteronotum: Seta 22 globular, enclosed in bothridial
cavity with faint reticulate markings on lining, and
appears as if slit-like opening to exterior between pair
of ear-like folds. Cuticular pits confined to concavities
at sides of high, flat median zone, which falls steeply
to rostral prominence. No setae observed in file s, but
if small would be difficult to distinguish amongst
adhering detritus. Apophysis to seta z1 medium length
(about equal to distance zl-zl) with curved lateral flap.
Opisthosternum: Aggenital shield fused to coxites and
only narrowly joined to adanal shield, with raised
median rim near preanal shield and small notch anterior
to seta Sg2 (one female has 3Sg on one side). Preanal
shield well separated from other shields, foreshortened
as illustrated (Fig. 14) since extends upward in vertical
SOUTH AUSTRALIAN SOILS 4
plane. Number of opisthosternal setae blade-like, with
hyaline flap on setal core. Genital shield has central
transverse unpigmented zone embracing setae JZg5,
JZg6 and notch between them (structure suggests shield
may fold along this line). No pore Zaf located on anal
shield although Saf and 4/3 conspicuous.
Hysteronotum: Fissure runs transversely along beside
first setal rank and backward along setal files Z and S$
(S1, $2, Z3, Z4) possibly representing dorsolateral
longitudinal fissure. Ventrolateral longitudinal fissure
terminates anteriorly, dorsal to posterior margin of
acetabulum /V, so that anterior part of opisthosomal
pleural shield merges with podosomal shield.
Comparison with Camisiidae, adult and nymphs,
suggests seta “dl” is Z2 not J2, so hysteronotal
chaetotaxy 4/, 5Z, 45, but concept of centralward
migration of Z2 and lost J2 debatable.
Appendages: Chelicerae relatively small, both digits
terminating in paired, nearly parallel teeth (including
distal points), with single large tooth proximally. Setae:
ch (2), pa (1-1-3-9), I (0-12-5-6-38), I (0-12-5-6-30 to 32),
IIT (5-7-5-5-28 or 29), IV (1-8-5-5-29 to 31). Solenidia:
pa (0-0-1), 7 (1-1-2), I (1-1-2), I/7 (1-1-0), TV (1-1-0).
Pretarsus with three subequal claws, lateral claws with
dorsal cilia file. Four terminal plasmic setae on palp with
minute distal knob. Solenidia piliform, never more than
1.25x length of associated setae. Solenidia on genua and
tibiae ZZ, J/Z, IV similar to one on genu I (Fig. 17), but
similar-sized or smaller than associated setae. Tarsi all
with supernumerary setae. Setae pale brown, or colour-
less on tarsi with strongly refractile base but hyaline
distally.
Somal inclusions; Three paratype females each with
four or five eggs. Eggs smooth, suboval, 305-310 long,
170-175 broad. Ovipositors involuted, extended would
be about 2x length of genital shield. Breadth slightly
less than that of one genital shield. Difficult to make
out setae, setae dg longer than J/Zg. Two clearly
delineated boli present per specimen, mainly cellular
material, spores, hyphae, cellular sheets of plant tissue.
Male
Measurements and spermapositor (otherwise as
female): Idiosomal length 1185 (8, 1125-1290);
appendage lengths (for 1185, holotype)—ch 77.5, pa 110,
T 1220, 17 747.5, [17 762.5, 1V 905; femur breadths—pa
22.5, J 110, ZT 100, 1 92.5, IV 87.5. Opisthosoma
contains large, granular horseshoe-shaped organ (open
end at posterior), possibly gonad. Spermapositor short,
breadth about half that of one genital shield, setae dg
about half length of JZg.
Material examined: Holotype male (N198317), seven
paratype males (N198318-N198324), three paratype
females (N198325-N198327), litter and sparse moss,
56 REC. S. AUST. MUS. 19 (4): 39-67 May, 1985
100,um
15,16
13,14
FIGS. 13-16. Crotonia jerhurmerae p.sp., male except when otherwise indicated, 13, nolumy 14, female idiosternum; 15, gnathosternum; 16,
right chelicera, anterior surface.
SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 4 37
d2
18 al2 f Li ae
dpx5 iy /
100mm
W719
FIGS. 17-19. Crotonia jethurmerae n.sp., male; 17, leg 1, dorsal surface of genu, tibia and tarsus; 18, leg tarsi, posterior surface of distal
ends; 19, leg III, posterior surface of trochanter and femur,
Lav
ed
under Eucalypius obliqua, Mt. Lofty, 9.5.1974, DL ©.
Lee.
Mt,
Distributions South Austealia—Aa: Lofty,
selerophyll open-forest, LL (5/8).
Remarks: Because one male has little pigment or
cerategument and therefore ig easy to examine, it is
designated holotype in cantrast to the usual procedure
of so designating a female.
Luxton (1982) divides Crofonia inta five spevies-
complexes, amongst which C jetiurmerue would be
grouped in the cophinaria-comptex. The inclusion of &
Jethurmerae requires modification of the characteristics
of that complex in that hysteronotal setae J] and SI are
approximately equal in length, also setae /l 15 Lhorn-
like and straight. The curved lateral Map on the
apaphysis to seta z] appears unique within the genus.
Family NANHERMANNIIDAE Sellnick
Nanhermanniidae Sellnick 1928: 17
Nanhermanniidaec: Woolley and Higgins, 1956: 913.
Type-genus: Nanhermannia Berlese, 1914! 100.
Diagnosis: Clinofissurac. Gnathosternal A-shaped
mentocoxal Fissure present, Three pairs of adaral setae.
Rostral tectum without median incision. Proteronotal
plasmic seta <2 baculiform or dilated and ciliate distally,
length lx-3x distance /2-32. Hysteronotal seta ./4 present,
Hysteronotal gland absent. Notal setae sl and /5 not
on apophyses, Coxites 7, //7 and /k may he
hypertrichous. Setal file S¢ includes two setae, combined
setal file JZg on median margin of genital shield,
Idiosoma almost covered in continuous shield except for
relatively well separated genital and anal orilices, and
prehysteronotal fissure extending ventrally (? anterior
part of ventrolateral longitudinal fissure) as crescentric
split vearly meeting midway between genital and anal
orifices. Discrete preanal shield but internal under
anterior margin of anal shields. Palp trochanter Fused
to femur, tarsus with seven setae. Tarsus / with one or
two pairs of dorsolateral supernumerary setae
(solenidium sol level with them). Nymphs without small
shields around hysteronotal setal bases.
Distribution: Probably cosmopolitan. Beside
Nanhermannia, Masthermannia may be cosmopolitan.
Cyrthermannia is tropical (Cuba, N'Ta; Thailand, Os)
as well as oecurring in Japan (Pe). Whilst
Cosmohermannia is known from southern Japan (Pe)
ahd New Guinea (Am).
Found if woodland and
fermentation and humus layers).
forest litter (both
Remarks; Vhe only thoroughly described species in
Nanhermanniidae is Masthermennia (as Posthermannia)
REELS. AUST. MLS. 19 (4p) ON?
May, 19K5
nematophora (Grandjean, 1954b), The family was
revised by Woolley and Higgins (1956), Although two
genera and a number of species have been deseribed
since then, the form of Nanhermannidae js without
great variations from that of the original species.
Nanhermanniidae includes the following four genera:
Cosmohermannia Aoki and Yoshida, 1970;
Cyrthermannia Balogh, 1958; Masthermannia Berlese,
1914) Nanherninnia Berlese, 1914.
NANHERMANNIA Berlese
Nanhermannia Berlese, 1914: 100. Type designation
(original); ““lermannia nana Nic’.
Type-species: Nanhermannia nana (Nicolet, 1855: 458).
Diagnosis: Nanhermanniidac. Hysteronotal setae not
on swollen tubercles, simple, either sefose or lanceolate,
sometimes minute forwardly directed proximal spur.
Posterior marvin of hysteronotum evenly convex without
prominent protuberances. Coxires 7, 7k with two to
four and three or four setae respectively. Trochanter /
with three dorsal setae,
Distribution: Probably cosmopolitan. Canada (Nn);
Washineton (Ne); Colorado, Idaho (Nr); Maryland,
North Carolina (Na); Chile, Patagoma, Peru (NTt);
Angola, Rhodesia (Ee); St. Helena (Es); Europe—most
northern records leeland and Kola Peninsula (Pe); Italy
(Pm); Altay Mountains, Kunashir Island, Sakhalin
Island, Samarkand (Ps); Japan (Pe); Thailand (Os);
Philippines (Om); New Guinea (Am); Queensland,
South Australia (Aa); New Zealand (An).
Remarks: Nanhermannia is the most widely recorded
venus in the family and currently includes at least
nineteen species, Balogh and Mahiinka (1978), in
describing N. donirowi from Queensland, refer to a
thaiensis-complex based on N, thaiensis Aoki, 19656
characterized by “medially confluent posterior
protuberances of rhe prodorsum and (hat they do not
have longitudinal furrows among interlamellar setae’.
For convenience, J will redefine the (Aveinsis-complex
which ineludes the species from this study.
thaiensis-complex
Diagnosis: Nanhermannia. Pair of posterior
proteronatal protuberances basically semicircular each
with four to sever) minor protuberances. Furrow between
seral pair 2 absent or shallow, not breaking connection
between protuberances, Genua and tibiae / and #7 seta
Hot bifureate.
Remarks Unfortunately the diagnosis has to be based
on the pasterior sculpturing of the proteronotum, which
SARCOPTIPFORMES (ACARI) O1
is difficult to use for some intermediate species. The
lhaiensis-complex is regarded as including the following
six species: N. domrowi Balogh and Mahunka,
1978—Queensland (Aa); N. forsslundi Karppinen,
1958—Finland (Pe); NM. goredkov{ Sinikova,
1975—Aliay Mountains and Kunashir Island (Ps); WN.
gtandjeani n.sp—South Austraha (Aa), M. pectinara
Strenzke, 1953—Germany (Pe); N, thaiensis Aoki,
1965b—Thailand (Os).
Nanhermannia grandjeani n.sp.
(Figs. 20-26)
female
General appearance and measurements: Light brown
generally, darker around leg acetabula and posterior
proteronotal protuberances. Shallow, clear
cero(cgument, some adhering detritus around Jeg bases,
Setae, claws, external malae, cheliceral digits clear and
refractile, Coarse puficta over much of soma excluding
lateral regions of proteronotum, coxites, genital and anal
shields, and crescent shape on both sides of setal file
Sa (see Fig, 21), Much smaller but deeper puncta located
dorsally on proteronatum and central part of each
caxite, Idiosomal length 637.5 (1); appendage lengihs—
ch 42.5, pe 55, f 265, 17 250, 777 220, 280; femur
breadths—pa 12.5, / 52,5, U 55, 1/7 40, 7 40,
Prostermmne External malae without adaxial hyaline
flap, but midanterior flap ventral to discrete tubercle,
twa rows of long cilia on dorsal surface. Three adoral
setae, aal very fine. Coxites merged with cach other and
sucrounding shields, although delineated by grooves.
Proteronotunt Seta 22 club-like, ciliate on dilated area,
Seta j2 has inconspicuous posterior spur at base.
Middorsal T-shaped [lat-lopped mound with setae zl
at anterior end and setae 22 al end of Jateral arms.
Region often used in species diagnosis illustrated in
detail (Fig, 24), Furrow between sctae /2 shallow,
aecentuated by absence of small puncta that cover ‘T-
shaped mound, Posterior proteronotal protuberances
obscure matching but smaller ventral protuberances (Fig.
20),
Opisthosternum: Shields distributed in manner unique
to Nanhermannia. File /Zg with nine setac and Sa with
three setae, Pores Zuf and Sef present, other two pores
illustrated (Fig. 22) regarded as hysteronotal (/1/).
fivsteronolum: Some setae (at least J2, 242, 23, Sl,
S2, SS) with inconspicuous anterior spur at base. All
setae blade-like, with hyaline dorsal and ventral flaps
along entire length, nor long enough in firsr three ranks
to reach base of following seta.
Appendages: Chelicerae relatively small. On fixed digit
five tecth including distal point, two small adaxial teeth
parallel to main two proximal teeth. Movable digit with
SOUTLE AUSTRALIAN SOLES 4 SY
(three tecth, including paired distal points and large
proximal tooth. Setae: ch (2), pa (1-0-2-7), 1 (I-5-5-6-23),
UL (1-7-5-5-22), #1 (5-2-2-3-18), J (1-3-2-2-16). Solenidia:
pa (O0-O-1), 7 (i-1-2), (7 (1-1-2), 277 (1-1-0), 7 (1-1-0),
Pretarsus with one claw, Terminal pair of plasmic setae
on palp tarsus spine-like, in recess. Anterodorsal edge
of tarsal tibia with refractile spur. Solenidia baculiform,
relatively. short, never as long as associated setae: Tarsi
Fand ffeach with two superhumerary setae; possibly
both dp. On tibiae 7 and 7 (only illustrated on 7F—
Fig. 25) seta wv conspicuously longer than segmen,
Somal inclusions: No eggs. Ovipositor involuied,
extended would be abow! 2% length of genital shield.
Breadth slightly less than that of one genital shicld
Bears eight pairs of setae, two dg pairs about 0.6% length
of setae J 42 and 4x length of setae pg. One simall bolus,
granular, particles many shapes, no complete cell walls
present,
Male
Unknown.
Material examined: Holotype female (NL98329), litter
and sparse inoss, under Eucalyptus ebliqua, Mt, Lofty,
95,1974, D.C. Lee.
Distribution: South Australia—Aa: Mi. Lofty,
scleraphyll apen-forest, | (1/8).
Remarks: Amongst the thaiensis-complex, differences
between species in the description of proteronotal
protuberances. and the intermediate furrow may be
actual or represent different interpretations by authors.
Within such variations N, grandjeani lies between N.
pectinaia and N. domrawi, The shortness of the
hysteronotal setae distinguishes N, grandjeani from N.
damrowi, N. forsslundi or N. zorodkovi. N. pectinata
has no large puncta on the proteronotal T-shaped
mound and seta s2 is not dilated distally. N. /halensis
has shorter, laterally biased protuberances, hysteranatal
puncta are larger and uneven i size and shape whilst
opisthosternal file Sa includes only two setae arid the
coxites bear only nine setae ventrally (3-1-2-3).
Family HERMANNITDAE Sellinick
Hermanniidae Sellnick, 1928: 18.
Hermannidae: Woas, 198k: 7.
Type-genus: Hermannia Nicolet, 1855.
Diagnosis: Clinolissurae. Gnathosternal A-shaped
mentocoxal fissure present, at least median part. Three
pairs Of adoral setae. Rostral tectum without median
incision. Proteronotal plasmic seta (32) filamentous or
club-like; length 0.75x-1.5x distance 2-72, Hysteronotal
Seta /4 present, Mysteronotal gland present. Notal setac
zl and J5 not on apophyses, Coxites /// and /¥ setac
60) REC. S. AUST, MUS. 19 (4): 39-67 Miav, 1985
100.Lm
22,23,24
FIGS. 20-24. Nanhermannia grandjeani n.sp., female; 20, notum and right pleura; 21, idiosternum; 22, gnathosternum; 23, right chelicera,
anterior surface; 24, proteronotum, posterior protuberances.
SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 4 4)
hypertrichous (3-1-4 or 5-5 or 7). Setal file Sg includes
two to five setae, setal files Jg, Zg separate but may be
close together. Idiosternum almost covered in
continuous shield but well separated from notal shield.
Discrete preanal shield, narrow (width about distance
JZal-JZa2). Palp tarsus with nine setae. Tarsus / without
dorsolateral supernumerary setae. Nymphs without
small shields around hysteronotal setal bases.
Distribution: Possibly cosmopolitan. Phyllhermannia
found mainly in southern hemisphere (see below) whilst
Hermannia confined to Palaearctic and Nearctic regions.
Species of Hermannia occur on bark of living trees,
regularly in forest litter, often at high altitudes, also in
littoral habitats and salt marsh (Krantz, 1978).
Remarks: A recent study of the Hermanniidae by
Waos (1981) makes Phyl/hermannia a junior synonym
of Hermannia, so that all species are in a single genus,
with both types in the gibbia/conyexa-complex. His
study is disadvantaged by the brief descriptions of
southern hemisphere species, i.c. those previously
grouped in Phyllhermannia. The below description 1s the
first comprehensive study of a Phyllhermannia species,
FIGS. 25-26. Nanhermannia grandjeani n.sp., female; 25, legs, dorsal and posterior surfaces of genua, ibiae and tarsi; 26, leg II] posterior
surface of trochanter and femur,
h2 RFC. S, AUST, MUS, 19 f4); 30.47
and character states such as the reduced setation of the
palp coxite, palp femur and legs, as well as the shape
of adoral seta ao2, suggest that Phyllhermannia may still
be a valid taxon, Therefore, 1 have chosen what may
appear a weak character (the position and shape of seta
zl) ta diagnose (he two genera, because it has always
been described and it makes a conservative grouping,
requiring only one species to be excluded from
Phyllhermannia, This i8 a temporary measure until
Phyllhermannia phyllophora is properly described.
The following two genera are included in
Hermanniidae: AHermannia Nicolet, 1854;
Phyllhermannia Berlese 1917a.
PHYLLHERMANNIA Berlese
Phyllhermannia Berlese, 1917a: 65, Type designation
(original): ““Hermannia phyllophara Mich?
Phyllhermannia: Tragirdh, 1931b: 576.
Hermannia (in part): Woas, 198): 36.
Type-species: Phyllhermannia phyllaphora (Michael.
1908: 140),
Diagnosis: Hermanniidae. Proteronotal seta zl with
distal half tapered off to a point, and marginal, lateral
to line j1-/2. Opisthoventral shield with transverse strip
between genital and anal shields. Mentocoxal fissure
usually complete, reaching edge of gnathosternum
(exception: PhyllAerinannia tuberculata).
Disiribuiion: Widespread in southern hemisphere,
known range extending into northern hemisphere along
western border of Pacific ovean. Chile, Juan Fernandez
Islands (NTc); Cape Province, Natal (Es); Madagascar,
Mauritius (Em); Tanganyika (Ee); southern Japan (Pe);
Thailand, Vietnam (Os); Java, Philippines (Om); South
Australia (Aa); New Zealand (An); Puntas Arenas (Sm),
Remarks: Phylihermannia was established without any
diagnosis, probably on the basis of the leaf-like leg setae.
The first detailed consideration of the genus was by
Trégdrdh (1931b) and keys to some species are given by
Aoki (1965b) and Balogh and Mahunka (1966), Woas
(1981) regards this genus asa synonym of Hermannia
and includes most species in his gibba/convexa-camplex.
The diagnosis of the genus used here is weak since
the character states of the opisthoventral shield and
mentocoxal fissure also occur in the small (afier the
exclusion of Phyllhermannia species) vibba/
convexa-complex within Aerniannia. My approach has
been to maintain PAyl/hermannia until more extensive
descriptions of included species, especially the type. are
available. Only one species has to be excluded from this
genus: Hermannia aeralata (Aoki, 1970) from Japan,
This is still grouped in the gibbia/convexd-complex as
by Woas (1981; 34),
MTak, TYRs
Twenty-four species. and one subspecies are included
in Phyl/hermannia: One species (P africana Balogh,
1958) has nol been considered because of the insufficient
description, Of the remainder, ten species are considered
similar to the new species described below and these are
grouped in a species-complex.
elisetosa-com plex
Diagnosis: Phyllhermannia, Hysteronotal seta Z1
central, not migrated laterally in front of Sl, distance
Z1./2 subequal to or less than J1V2. Hysteronotal setae
shorl, J] mot reaching J2 base. Apodemes between
coxites /J/ and J/-//7 oblique, at least at 22.5° angle ta
transverse axis. Genital setal file Ze with at least one
seta twice length of Sg setae, ar more. Leg sctae never
leaf-like or spatulate and number reduced, tarsus | with
24 or fewer setae and one solenidium.
Remarks: Members of the eusefosa-complex are
usually only known by idiosomal character states, The
three species from Mauritius, with only one setae in
opisthosternal file Sg, may form a separate complex, P
luberculata from Chile is included although unique
within the genus in having a restricted mentocoxal
fissure as in many Hermannia species.
Elevert species are included m the complex. P
bimaculata Hammer, 1979—Java (Om); PB eusetosa
tisp.—South Australia (Aa) P Joliata Hammer,
1966—New Zealand (An); AP mauritii Mahunka,
1978—Mauritius (Em); PRP medesfa Mahunka,
1978—Mauritius (Em); P’ mollis Hammer, 1966—New
Zealand (An); PB pacifica Hammer, 1972—Tahiti (Ap);
P paulieni Balogh, 1962—Madagasear (Em); P rubra
Hammer, 1966—New Zealand (An) A tremicta
Mahunka, 1978—Mauritius (Em); FR tuberculata
Covarrubias, 1967—Chile (NTe).
Piyllhermarnia eusetosa n.sp.
(Figs 27-32)
Female
General appearance and measurements: Red-brown,
covered in cerotegument, thin with sparse adhering
detritus. Beside small tubercles laterally and around
coxites, acetabula and genital and anal orifices, soma
(excepting proteranotum) covered in low, minute,
superficial bumps, with pale strips between forming a
reticulate pattern. All shields covered with fine puncta.
Similar puncta and pattern on legs. Idiosomal length
795 (25, 675-917,5); appendage lengths (for holatype,
915)— ch 57.5, pa 90, I 380, IT 430, 77 440, TF 585;
femur breadths— pe 27.5, F 117.5, 17105, 11790, 1¥ 95_
Prosternuin: On external malae median hyaline lap
hanvontal with posterior notch through which adoral
seta. wo2 protrudes. Internal mala with distal spike
leading back to minute dorsal furrow, Lateral réfractile
SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 4 03
part of external malae bears two rows of cilia on dorsal
surface. Lateral and central gnathosternum with finely
punctate patches as illustrated (Fig. 29). Coxites all fused
into one shield but clearly delineated by grooves, deep
along midsternal line.
Proteronotum: Seta z2 vermiculate, but slightly dilated
and ciliate distally. Bothridial cavity for seta 22 with
number of short pockets and one long pocket. Two pairs
27 ‘i
|
on
of sharp tubercles near posterior margin face backwards,
dark and conspicuous although small: one lies at
posterior end of ridge running backwards from
acetabulum /, other lies posterior to setae /2 and z2 and
equidistant from both.
Opisthosternum: Continuous opisthoyentral shield
with strip between genital and anal shields, broadly
fused to coxites, but not fused together posterior to anal
10Q.um
29,30
27, 28
FIGS. 27-30. Phyllhermannia eusetosa n.sp., female; 27, notum; 28, idiosternum; 29, gnathosternum; 30, left chelicera, anterior surface.
64 REC, S. AUST. MUS. 19 (4): 39-67 Muy, 1985
FIGS, 31-32. Phyllhermannia eusetosa n.sp., female; 31, legs, dorsal and posterior surfaces of genua, tibiae and tarsi; 32, leg IL] posterior
surface of trochanter and femur.
SARCOPULIPORMES (AC ART) GF SOL TH AUSERALLAN SOILS 4 As
shiclds. Groove behind coxite Ji Cresent-shaped
thickening of integument behind acetabulum /F Anal
shield with longitudinal ridge endimg anteriorly in
tubercle that fits around preanal shield. Both this
tubercle and crescent-shaped thickening behind
acetabulum /¥ conspicuously dark-coloured, although
with pale shading (Fig. 28) because they stand proud
from surrounding shields, Pare Sa/ present bul not Za
Hysteronoium: Two pairs of tubercles near anterior
margin, ventral to cireumhysteronotal fissure and
opposing proteronotal tubercles, central pair small and
inconspicuous. Pair of dark semicircular integument
thickenings behind first rank of hysteronotal setae (/1,
Z\, Sl), from which central arm borders on
inconspicuaus furrow running back to bend as faint
ridge around seta ./4. All hysteronotal setae blade-like.
Appendages: Chelicerae relatively small, Both digits
with four teeth, terminating in paired, nearly parallel
leeth (including distal points). Palp femur with
anteroventral Mange, Setae: ch (2), pa (O-l-1-3-9), T
(0-6-5-5-22), FF (1-7-5-5-17), MP (2-3-2-4-15), 7
(2-4-3-4-14), Soleridia: pa (0-0-1), (1-241), 4-1-1), 0
(1-1-0), /¥’ (0-1-0). Pretarsus with single claw bearing pair
of inconspicuous dorsal cilia files, Terminal pair of
plasmic¢ setae on palp tarsus spine-tike, with anteradarsal
flange around their bases. All but one solendium
baculiform, relatively short, less than 0.5x length of
segment bearing them, Solendium so2 on tibia |
flagelliform, very lony, about equal ta length of genu,
libia and tarsus J together. On genua /, 77, J, and on
all tibia (excluding so) on tibia 1) solenidia coupled with
dorsal setac, whilst solenidia on tarsi 4, /7 and sol on
tibia / at most only associated with dorsal setae.
Reticulate pattern on dorsal and lateral surfaces of most
ley segments (Fig, 32—part drawn on femur), but not
on dorsal surface of trachanter or any part of tarsi. This
pattern consists of darker raised ridges unlike similar
hysteronotal pattern.
Samal inclusions: Amongst twenty-five registered
specimens, three contain two eggs, one contains ciree
eges, five contain four eggs, remainder without eggs.
Eges about 240 (225-265) long, ellipsoid, with uniform
smooth surface, Ovipositor 2x length of genital shield.
Breadth about 15x that of one genital shield. Bears eight
pairs of setae, two dg pairs subequal in length ro setac
Ze2 but thom-like (x2 breadth) and more refractile, three
pe pairs about length of genital shield away from three
me pairs. Only one bolus seen with mainly
unrecognisable (razments, but some spherical spores and
one multicellular strip of tissue,
Male
Unknown.
Miuaterial examined: Holotype female (NL98330), and
twenty-four paratype females (N1I983417-N1983440),
under uvalyplus oblique, sclerophyll forest, Mt Lofty,
95.1974,
Distribution: South Australia—Aa: M1.
sclerophyll forest, 118 (6/8),
Lofty,
Remarks: P. eusetusa@ is distinguishable from other
members of the eusefesa-complex in possessing the
fallowing combination of character states; hysteronotal
setae blade-like Withaul cilia, no proteronotal ridge runs
cither between setae j2-/2 or seta j2 and the median
tubercle near posterior margin; coxite / bears five setae.
ACKNOWLEDGMENTS
J am indebted ta the Interim Council of the Australian
Biological Resources Study for funds for equipment,
and. to the Science and Industry Endowment Fund for
funds for travel in conneclian with this project,
Special thanks are due to Mr D. MacFarlane,
Commonwealth Institute of Entomology, London, for
an English translation of a paper written ju French
(Grandjean, 1954a) and to Dr G. Wauthy, Carholic
University of Louvain, Belgium, for a gift of specimens
of Eulohmannia ribagai. Thanks are also due to Dr
8, G. M. Jamieson, University of Queensland, Brisbane
and Dr R, A. Norton, SUNY. College of Environ-
mental Science and Forestry, Syracuse, for commenting
on the manuseript.
My greatest debt of gratitude is to Ms Jenni Thurmer
for her excellent illustrations and to Mrs Debbie Metloy
for typing this manuseript-
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WALLWORK, J. A. 1963. The oribatei (Acari) of Macquarie Island.
Pacif. Ins. 5: 721-769.
WILEY, E. O. 1981. Phylogenetics, the theory and practice of
Phylogenetic Systematics. New York, John Wiley and Sons, 439 pp.
WILLMANN, C. 1928. Neue Oribatiden. I. Zool. Anz. 76: 1-5,
WOAS, S. 1981. Zur Taxonomie und Phylogenie der Hermanniidae
Sellnick 1928 (Acari, Oribatei), Andrias 1: 7-88.
WOOLLEY, T. A. 1969. The infracapitulum—a possible index of
oribatid relationship. Jnt. Congr Acarology: 2. Budapest: 209-221.
WOOLLEY, T. A. and HIGGINS, H. 1956. A revision of the family
Nanhermanniidae (Acari: Oribatei), Proc, Tenth Int. Congress
of Ent. I: 913-923.
MOLLUSC TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM
2. GASTROPODA: CONIDAE
BY WOLFGANG ZEIDLER
Summary
Type-specimens of 21 species of Conus in the South Australian Museum are catalogued and
illustrated. All are holotypes or syntypes. Of these at least three, C. cumingii Reeve, 1848, C.
metcalfei Angas, 1877 and C. sydneyensis Sowerby, 1887 are almost certainly not types while the
status of another two, C. kermadecensis Iredale, 1912 and C. rossiteri Brazier, 1870 is in doubt.
Only 10 of the specimens are from Australian waters, another 10 are from various South Pacific
islands and one is from Mauritius. Species are listed alphabetically according to the original name
of the species or variety. In addition, four species of cones recorded as types for which no reference
can be found are listed in an Appendix.
MOLLUSC TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM
2, GASTROPODA: CONIDAE
by
WOLFGANG ZEIDLER
South Australian Museum, North Terrace, Adelaide, South Ausiralia 5000
(Manuscript accepted 29 August 1983)
ABSTRACT
ZEIDLER, W, 1985. Mulluse type-specimens in the South Australian
Museum, 2. Gastropoda: Conidae. Rec. 8. Aust, Mus. 15). 69-75
Type-specimenis of 21 species of Conus in the South
Australian Museum are catalogued and illustrated. All
are holotypes or syntypes. Of these at least three, C
cumingii Reeve, 1848, C. metcalfei Angas, 1877 and C.
sydneyensis Sowerby, 1887 are almost certainly not types
while the status of another two, C kermadecensis
Iredale, 1912 and C. rossiferi Brazier, 1870 is in doubt.
Only 10 of the specimens are from Australian waters,
another 10 are from Various South Pacific islands and
one is from Mauritius. Species are listed alphabetically
according to the original name of the species or variety,
In addition, four species of cones recorded as types for
which no reference can be found are listed in an
Appendix.
INTRODUCTION
Most of the indigenous Conidae collection in the
South Australian Museum (SAM) was collected, or
otherwise obtained by Sir Joseph C. Verco around the
turn of the century. However, most of the exotic species
were acquired by purchasing other people’s collections.
One of the most significant of these was the A, F
Kenyon collection which the Museum purchased early
this century. Mrs Kenyon accumulated a considerable
collection and acquired many types, particularly of
cowries and cones, She also acquired much of the J.
W. Brazier collection which accounts for about half of
ihe cone types listed here and only three of Brazier’s
species are not represented: C. cooki Brazier, 1870 (two
syntypes in Australian Museum, Sydney), C coxeni
Brazier, 1875 (type in Coxen collection—lost?) and C.
sophiae Brazier, 1875 (type in Australian Museum,
Sydney). Unfortunately, several of the original labels are
missing and the type status of some specimens is in
doubt, There are also a number of specimens which have
been labelled type” but for which no reference can be
found and these are listed in the Appendix and are only
included because fiituire workers may stumble upon a
reference to them.
The South Australian Museum holds 21 Conus
“types” of which three, C. cumingii Reeve, 1848, C.
metcalfei Angas, 1877 and C sydneyensis Sowerby, 1887
are almost certainly not types while the status of another
two, C. kermadecensis Vredale, 1912 and C. rossiferi
Brazier, 1870 is in doubt.
The species are listed under the original name at the
time of description and in alphabetical order of species
or variety. All of the types are represented by shells only,
and only two have the operculum with them indicating
that they were alive when collected. Measurements for
height and maximum width are given for each specimen,
as these were often inaccurate or not given in the onginal
description. Each specimen has also been photographed
(Figs, 1-3) as the types were rarely figured and
photographs of only a few have appeared in recent
publications.
The present status of each species is, in most cases,
according to Walls (1978).
LIST OF TYPES
Conus kenyvonae var. arrowsmithensis Brazier, 1896
(Fig. la)
Prac, Linn. Soe. New South Wales 21; 346,
Holotype: SAM Reg, No. D5459. Kenyon No, 337,
Dimensions, 35.5 ™ 20.5 mm (Brazier gives 36 x 2]
mm).
‘Type-locality: Arrowsmith Island, Marshall Islands;
collected by J, Brazier, 22-ix.1872.
Present Status: Synonym of C. distans Hwass, 1792,
Remarks: A very worn specimen with Brazier’s original
label. Considered a juvenile of C disrans, Type figured
by Cotton (1945), pl. 4, Fig. 4.
Cunus barbara Brazier, 1898
(Fig. Ib)
Proc. Linn. Sac, New South Wales 22; 784,
Holotype: SAM Reg. No. D6176. Kenyon No, &3,
Dimensions, 40.0 « 20.3 mm (Brazier gives 40
20 mm).
‘Type-locality: Solomon Islands; collected by Mrs A. F-
Kenyon, date of collection unknown,
Present Status: Synonym of C. monachus Linnaeus,
1758,
Remarks: A worn specimen with Brazier's original label.
Type figured by Cotton (1945), pL 4, Fig. 2.
7 REOLS. AUST MUS, 19 (5) 69.78
Conus cumingii Reeve, 1848
(Fig, le)
Conchologia Ieonica | (Conus Suppl.): Pl. 3. species 282.
Holotype: SAM Reg. No. D6208. Kenyon No. 372.
Dimensions, 48.5. x 24.5 mm (Reeve gives none).
Type-locality: None given on any labels but Island of
Mindanao, Philippines; collected by Mr Cuming,
according to Reeve.
Present Status: Valid species.
Remarks: This specimen cannot be Reeve's type of
cumingii despite the label which says “Reeve’s type’. It
does not agree with Reeve’s description and is too large
for the species. It seems to be C. anemone Lamarck,
18L0 and was not listed by Cotton (1945),
Conus flindersi Brazier, 1898
(Fig. Id)
Proc. Linn. Soc. New South Wales 22; 780.
Holotype: SAM Reg. No, D14129. Kenyon No. 139.
Dimensions, 28.0 x 15.0 mm (Brazier gives 29 x
19 mm).
Type-locality: Flinders, Western Port, Victoria, wider
stone at low water. (Not Flinders [sland, Bass Strait).
Probably collected by Mrs A. F, Kenyon, date of
collection unknown.
Present Status: Synonym of C. anemone Lamarck, 1810.
Remarks: This specimen does not match up with the
dimensions given by Brazier, but as the original label
by Brazier is present and in the absence of any other
possible types, We must accept it as Brazier's Lype and
conclude that his measurements are inaccurate, Cotton
(1945) maintained that he had not seen the type.
Conus frostiana Brazier, 1898
(Fig. le)
Proc. Linn. Soc. New South Wales 22: 781.
Holotype: SAM Reg. No. D6170, Kenyon No. 138,
Dimensions, 18.2 * 9.5 mm (Brazier gives 18 *
10 mm).
Type-locality: Solomon Islands; collected by Mrs A. F.
Kenyon, date of collection unknown.
Present Status: Uncertain, Walls (1978) thinks this
species could be synonymous with C. aigropunctatus
Sowerby 1858 or C. wtonachus Liane, 1758 or even C.
magus Linne, 1758. 1 am inclined to believe that the type
may be a small CG monachus.
Remarks: Specimen with Brazier’s original label.
Conus kenyonae Brazier, 1896
(Fig. 1f)
Proc, Linn. Soe. New South Wales 21; 346,
Holotype: SAM Reg. No. D14194. Kenyon No. 336.
Dimensions, 42,0 * 23,5 mm (Brazier vives 43 x
24 mm).
TVpe-locality: Shark’s Bay, Western Australia; collected
by Mr Podesta, date of collection unknown,
Present Status: Synonym of C. distans Hwass, 1792.
Remarks: A very worn specimen with Brazier’s origmal
Nay TAs
label, Considered a juvenile of C distans, Type figured
by Cotton (1945), pl. 4, Fig. 6.
Conus kermadecensis {redale, 1912
(Fig, 1p)
Proc, Mal. Sac, Land, WW(3): 227, pl. 9, Figs. 13 and 16,
Syntype: SAM Reg. No. D6I69. Kenyon Na. 420,
Dimensions, 43.0 * 23.4 mm,
Type-locality: Original label lost? Iredale gives Sunday
(=Raoul) Island, Kermadee Group.
Present Status: Valid species,
Remarks; Specimen with operculum, corresponding
venerally 10 lredale’s description, The type status of the
Specimen, however, isin doubt as Iredale stated that the
type would go to the Canterbury Museum, New Zealand
(which apparently has two) and the paratypes would ga
to the Australian Museum, Sydney. The Australian
Museum has three lots of C. kermadecensis collected by
Iredale and Oliver from the type-locality, but nane have
been designated type or paratypes. Since Iredale
mentions only three specimens, and two are accounted
for, the above could be a syntype corresponding to the
one lredale measured as 42 % 25 mm. Cotton's (1945)
measurement Of 45 mm for this specimen is obviously
an error.
Rhizoconus klemae Cotton, 1953
(Fig, Ih)
Trans. Roy. Soe. South Aust, 76: 24, pl, 3, Figs, | and 3.
Holotype; SAM Reg. No. D14465. Dimensions:. 46.5 *
26.0 mm (Cotton gives 47 *% 26 mm).
‘Type-locality:; Corny Point, Yorke Peninsula, South
Australia; collected by Miss M. Klem, date of collection
unknown.
Present Status: Conus klemae (Cotton, 1953), valid
species.
Remarks: Most of the specimens mentioned by Cotton
(1953) are in the SAM collections but none were
designated paratypes, A juvenile specimen (SAM Reg.
Nou. 1916233), callected with the type, Was also figured
by Cotton (pl, 3, Fig. 2).
Conus mietcalfei Angas, 1877
(Fig. 2a)
Proce. Zool. Soc. (London): 173, pl. 26, Fig. 13,
Holatyper SAM Reg, No. D6207, Kenyon Na. 200,
Dimensions, 17.6 ~ 95 mm (Angas gives 21.2 x
12,7 min),
Type-locality: Dredged at “Sow and Pigs” reel, Port
Jackson, New South Wales; probably colleered by J.
Brazier, date of collection unknown,
Present Status: Synonym of C. atgasi Tyron, 1883.
Remarks: This specimen is almost cerlainly not the type.
The original label is missing and the measurements do
not correspond to those given by Angas. The type is
mast likely in the Australian Museum, Sydney who have
a specimen (C1LO3598) with an old label marked “Type”
measuring 20.5 « ibd mm which is a closer
MOLLUSC TYPE-SPECIMENS 2. GASTROPODA: CONIDAE 71
FIG. 1. a. Conus kenyonae var. arrowsmithensis, holotype 35.5 x 20.5 mm (x1): b. Conus barbara, holotype 40.0 x 20.3 mm (x1): c. Conus
cumingii, 48.5 x 24.5 mm (x1): d. Conus flindersi, holotype 28.0 x 15.0 mm (xX1.5): e. Conus frostiana, holotype 18.2 x 9.5 mm (x2):
f. Conus kenyonae, holotype 42.0 x 23.5 mm (x1): g. Conus kermadecensis, ? syntype 43.0 x 23.4 mm (x1): h. Rhizoconus klemae,
46.5 x 26.0 mm (x1).
approximation to Angas’s description. Not to be acone. It is most likely a turrid similar to Conopleura
confused with C. metcalfii Reeve, 1843. Hinds, 1844 (Cernohorsky, 1974).
Remarks: A worn specimen with Brazier’s original label.
Kenyonia pulcherrima Brazier, 1896 Not to be confused with Conus pulcherrimus Brazier,
(Fig. 2b) 1894 or Conus pulcherrimus Heilprin, 1879—a fossil.
Proc. Linn. Soc. New South Wales 21: 347. . ;
Holotype: SAM Reg. No. D6181. Kenyon No. 175. Conus pulcherrimus Brazier, 1894
Dimensions, 28.0 x 9.4 mm (Brazier gives 28 x (Fig. 2c)
10 mm). Proc. Linn. Soc. New South Wales 9(1): 187.
Type-locality: New Hebrides; collected by Mrs. A. F. Holotype: SAM Reg. No. D6172. Kenyon No. absent.
Kenyon, date of collection unknown. Dimensions, 80.0 x 27.2 mm (Brazier gives none)
Present Status: Uncertain. This is almost certainly not Type-locality: Tanna, New Hebrides, on beach after
we REC, S. AUST. MUS. 19 (5): 69-75
May, 1985
FIG. 2, a. Conus metcalfei, 17.6 % 9.5 mm (* 2): b. Kenyonia pulcherrima, holotype 28.0 x 9.4 mm (x 1.5): &. Conus pulcherrimus, holotype
80.0 x 27.2 mm (x 0.5): d. Conus remo, lectotype 35,0 « 17.5 mm (1): &. Conus rossiteri, Tholotype 10,6 x 6.4 mm ( *3): f. Floraconus
sautdersi, holotype 57.0 x 32.0 mm (x1): g. Floraconus singletoni, holotype 43.5 * 22.0 mm (x1): h. Conus superstes, syatype 3.3
x 1.7 mm (x10).
submarine voleanic eruption; collected in 1878; collector
unknown.
Present Status: Synonym of C. excelsus Sowerby, 1908.
Remarks: A worn specimen with Brazier’s original label
indicating the species and locality but not the fact that
it was Brazier’s type. Cotton (1945) apparently ignored
the original label and did not realize that Brazier had
given a short description under “Notes and exhibits”
and thus described this specimen as the unique type of
a new species, Asprella tannaensis.
Not to be confused with Kenyonia pulcherrima
Brazier, 1896 which is now considered a turrid or C.
pulcherrimus Heilprin, 1879—a fossil.
Conus remo Brazier, 1898
(Fig. 2d)
Proc. Linn. Soc. New South Wales 23: 27).
Lectotype: SAM Reg. No. D14128. Kenyon No. 186.
Selected by Cotton (1945), p. 265, pl. 4, Fig. 9,
Dimensions, 35.0 * 17.5 mm (Cotton gives none but
Brazier gives 35 x 154-17 mm for the type).
Type-locality: San Remo, Port Phillip Bay, Victoria;
collected by Mrs A. F. Kenyon, date of collection
unknown.
Paralectotypes: SAM Reg. No. D15948, 8 specimens with
same collection data as type.
MOLLUSC TYPE-SPECINMIENS 2, GASTROPODA: CORIDAL TS
Present Status; Synonym of C. anentone Lamarck, 1810,
Remarks; Specimens with Brazier’s original label but the
type had not been indicated.
Conus rossiteri Brazier, 1870
(Fig. 2¢)
Prac. Zool, Soe. (London) (1870); 109,
Holotype: SAM Reg. No. D5975. Kenyon No. 312.
Dimensions, 10.6 x 6.4 mm (Brazier gives approx. 16
x 8 mm).
Type-locality: Cape Solander, Botany Bay, New South
Wales; collected by J. Brazier, date of collection
unknown,
Present Status: Synonym of C anemone Lainarck 1810,
Remarks: This is a doubtful type-specimen as it is much
smaller than the measurements given by Brazier, Also
the original label is missing and there is no indication
that this is the type apart from the museum register
referring lo il as “Llype”’ and Cotton's (1945, 1958)
reference to it as the type. The Austalian Museum,
Sydney, has a specimen registered as type in 1902, from
material purchased from Brazier in 1&89, measuring
14.5 x 9mm.
Floraconus saundersi Cotton, 1945
(Fig, 2f)
Ree, §. Aust. Mus. 8(2): 264, pl. 4, Fig. 8.
Holotype: SAM Reg, No, D14198, Dimensions, 57.0 x
32.0 mm (Cotton gives none).
Type-locality; Levens Beach, Edithburgh, Yorke
Peninsula, South Australia; collector and date of
collection unknown,
Present Status: Synonym of C. avremone Lamarck, 1810,
Remarks: Two specimens, SAM Reg. No. D16232, were
with the holotype and may have been used in the
original description but were not designated paratypes.
Floraconus singletoni Cotton, 1945
(Fig. 2g)
Ree, S, Aust, Mus, 8(2): 263, pl. 4, Piz, 10,
Holotype: SAM Reg. No. D14195. Dimensions, 43.5 x
22.0 mm (Cotton gives 45 x» 22 mm),
Type-locality: Western Port, Victoria; collected by C. J.
Gabriel, date of collection unknown,
Present Status: Synonym of C anemone Lamarck, 1810,
Remarks: Considered an albino variant of C. anemone,
TWo specimens, SAM Reg, No, D16236, were wilh the
type but were not designated paratypes.
Conus superstes Hedley, 1911
(Fig. 2h)
Zoo), Results ELS. Endeavour 1904-10, part 1; 111, pl. 20,
Figs, 345, 36,
Syntypes: SAM Reg. No. D15904, 11 specimens, the
specimen figured here measures 3.3 % 1.7 mm.
Type-locality: Forty miles south of Cape Wiles, South
Australia in 100 fms; collected by FILS. Endeavour
28,viil, 1909,
Present Status: The species description is based on small
juveniles, so (hat it would be difficult to determine the
specific status until the life histories of South Australiati
cones are better known,
Remarks: Specimens with the original label with the
ward “Co-type” and registration number “E.3800".
Cantus sydneyensis Sowerby, 1887
(Fig. 3a)
Thesaurus Conchyliorum, 5 (Conus Suppl.); 260, pl. 32
(510), Fig. 694,
Holotype: SAM Reg, No, D6I83. Kenyon No. 192.
Dimensions, 17.0 * 9.0 mm (Sowerby gives 24
x 12 mm),
Type-locality: Port Jackson, New South Wales, collected
by Brazier; date of collection unknown.
Present Status: Synonym of C. aplusire Reeve, 1843
according to Walls (1978) or of C angasi Tyron, 1883
according to Cotton (1945). The specimen at hand looks
more like C angasi than C. aplustre,
Remarks: This specimen cannot be Sowerby’s (ype, even
though the original label with the specimen, designating
it as a type, appears to be in Brazier’s handwriting. The
locality data is given as Port Stephens and the
measurements do not agree with the original description
given by Sowerby. According to Moolenbeek (pers.
comm.) the type appears to be in the Institut Royal des
Sciences Naturelles de Belgique (Brussels).
Asprella tatinaensis Cotton, 1945
(Fig. 2c)
Rec. S. Aust. Mus, 8(2): 270, pl. 4, Fig, 3,
Holotype: SAM Reg. No. 124172.
Remarks: Based on the same specimen as the type of
C. pulcherrimus Brazicr, 1894 also listed in this paper,
The operculum figured with the type by Cotton (1945)
probably belongs to another species as the type is worn
and faded and was obviously collected devoid of the
animal.
Alermes trigei Cotton, 1945
(Fig. 3b)
Rec, S. Aust, Mis, 8(2): 267, pl, 4, Fig. WU,
Holotype: SAM Reg. No. 914324. Kenyon No,—absent.
Dimensions, 61.5 * 27,4 mm (Cotton gives 60
x 27 mm).
Type-locality: New Hebrides; collector and date of
collection unknown.
Present Status: Synonym of C wximenes Gray, 1839.
Remarks: Described froma unique specimen from the
Kenyon collection,
74 REL
S AUST. MUS, 19 (3S): 64-75
Wav, 1985
FIG, 3, a, Conus sydneyensis, 17.0 * 9.0 mm (2); b. Hermes rriggi, holotype 61.5
~ 24.5 mm (x1).
<~ 16.0 mm (“L.5): d. Conus worcesteri, holotype 47.6
Conus waterhouseae Brazier, 1896
(Fig. 3c)
Proc. Linn. Soc. New South Wales 21: 471.
Holotype: SAM Reg. No. D5786. Kenyon No,—absent,
Dimensions, 28.5 »* 16.0 min (Brazier gives 30
x 15 mm),
Type-locality: Solomon Islands; collected by Mrs. G. J.
Waterhouse, date of collection unknown.
Present Status: Synonym of C. disfans Hwass, 1792.
Remarks: Specimen with Brazier’s original label, Type
figured by Cotton (1945) pl. 4, Fig. 2. Another specimen
(SAM Reg. No. DS787) with the type and marked “type
of variety” was also figured by Cotton (pl. 4, Fig. 7).
This specimen, from Mauritius, was introduced as a new
variety by Kenyon (1906) who gave a brief description
but it was not given a name as it was considered to be
a juvenile of C distans.
Conus (Chelyconus) worcesteri Brazier, 1891
(Fig. 3d)
Proc. Linn. Soc, New South Wales 6: 276, pl. 19, Fig. 4.
Holotype: SAM Reg. No. D6178. Kenyon No, 155.
Dimensions, 47.6 x 24.5 mm (Brazier gives 48 x 25
mm).
Type-locality: Island of Mauritius; collected by Mr
Robillard, date of collection unknown.
Present Status: Synonym of C magus Linnaeus, 1758.
Remarks: The original label by Brazier is missing but
there is little doubt that this is the type. The specimen
was collected live as the operculum is present. Type
figured by Cotton (1945), pl. 4, Pig. 1.
» 274 mm (XI1)oc. Conus waterhouseue, holotype 28.5
ACKNOWLEDGMENTS
1 wish to thank Mr I. Loch of the Australian
Museum, Sydney for supplying information on the cone
types in that institution and Dr R. G. Moolenbeek,
Institut Voor Taxonomische Zoologie, Zoologisch
Museum, Universiteit van Amsterdam, The Netherlands,
for his comments on the status of C. kermadecensis and
C. sydneyensis. Thanks are also due to Mrs J.. Forrest,
South Australian Museum, who took, and also
prepared, the photographs.
REFERENCES
ANGAS, G. 1}. 1877. Descriptions of one genus and twenty-five species
of marine shells from New South Wales. Proc. Zool. Soe.
(London): 1714177, pl. 26.
BRAZIER, J. 1870. Descriptions of three new species of murine shells
from the Australian coast. Proc. Zoal. Soe, (London): 109,
BRAZIER, J, 1875a. Description of fourteen new species of shells
from Australia and the Solomon Islands. Proc, Linn, Soe, New
South Wales be 1-9.
BRAZIER, J. 1875b. Descriptions of ten new species of shells, from
the collection of Mr. Charles Coxen, of Brisbane, Queensland.
Proc, Zool Soe. (London): 3-34, pl. 4.
BRAZIER, J, 1891. Description of a new cone from Matiritius. Pree.
Linn Soe. New South Wales 6: 276, pl. 19.
BRAZIER, J, 1894, Notes and exhibits, Proc. Linn. Sac. New South
Hirles 91): 187,
RRAZIER, J. 1896a. A new genus and three new species of mollusca
from New South Wales, New Hebrides and Western Australia.
Proc. Linn. Sov. New South Wiles 21: 345-347,
BRAZIER, J. 1896b, New species of Cone trom the Solomon Islands,
Proc. Linn. Soc, New South Wales 2h; 471.
BRAZIER, J. 1898a. New marine shells from (he Solomon Islands
and Australia. Proc. Linn, Soe, New South Wales 22: 779-782,
BRAZIER, J. 1898b, Four new species of Mollusca from Victoria.
Proc. Linn. See. New South Wales 23, 271-272,
CERNOHORSKY, W.. 1874. The taxonomy of some Indo-Pacific
Mollusca with description of a new species. Part 2. Ree. Aucklarul
Inst. Mus, Vy 121-142.
MOLLUSC TYPE-SPECIMENS 2. GASTROPODA: CONIDAE 75
COTTON, B. C. 1945. A catalogue of the cone shells (Conidae) in
the South Australian Museum. Rec. S. Aust. Mus. 8(2): 229-280,
pl. 1-5.
COTTON, B. C. 1953. New species and records of Mollusca from
South Australia. Trans. Roy. Soc. South Aust. 76: 21-26, pls 2, 3.
COTTON, B. C. 1958. Australian recent and tertiary species of the
molluscan family Conidae. Publ. B. C. Cotton. 4 pp. Sept 1.
HEDLEY, C. 1911. Report on the Mollusca obtained by the ELS.
Endeavour chiefly off Cape Wiles, South Australia. Zool. Results
FILS. Endeavour 1904-10, pt. 1: 90-114, pls 17-20.
IREDALE, T. 1912. New generic names and new species of marine
Mollusca. Proc. Mal. Soc. (London): 10(3): 217-228, pl. 9.
KENYON, A. F. 1906. Notes. Proc. Mal. Soc. (London): 7(1): 5.
REEVE, L. A. 1848. Monograph of the genus Conus. Conchologia
Iconica | (Conus Suppl.).
SOWERBY, G. 1887. Thesaurus Conchyliorum 5 (Conus Suppl.).
WALLS, J. G. 1978. Cone shells: a synopsis of the living Conidae, 1011
pp. T.F.H. Publications.
APPENDIX
Specimens labelled or registered as “type” for which
no reference can be found.
Conus noumeensis Brazier, SAM Reg. No. D5729.
Kenyon No. 69. (68.5 x 39.5 mm).
Locality: Anse Vata, Noumea, New Caldonia.
Remarks: Specimen with Brazier’s label marked “Type
specimen”. Cotton refers to this specimen and says that
it is C. chenui Crosse, 1857 and has nothing to do with
C. suffusus var. Noumeensis Crosse, 1872 which is a
synonym of C. marmoreus Linnaeus, 1758.
Conus pilcheri Brazier, SAM Reg. No. D6171. Kenyon
No. 247 (22.3 x 14.0 mm).
Remarks: Specimen with Brazier’s label marked “Type
specimen” but no locality data. It is very similar to C
cyanostoma A. Adams, 1854.
Conus sydneyensis var. sowerbyi Kenyon, SAM Reg. No.
D6184. Kenyon No. 192 (27.0 x 15.5 mm).
Locality: Port Stephens, New South Wales.
Remarks: Specimen with Kenyon’s label marked “type”.
Another specimen with a similar label but not marked
“type” is also in the collection. The specimens seem to
be juvenile C. purpurascens Sowerby, 1933.
Conus wisemani Brazier, SAM Reg. No. D6179. Kenyon
No. 199. (28.2 x 17.3 mm).
Locality: Low Island, Trinity Bay, north Queensland.
Remarks: Specimen with Brazier’s label but not marked
“type” yet registered as a type in Museum register. It
seems to be a white form of C. suturatus Reeve, 1844.
RECORDS oF THE
SOUTH AUSTRALIAN
MUSEUM
VOLUME 19 NUMBERS 6-10 JUNE 1986
No.
No.
No.
A REVISION OF THE TROGLOPHILIC GENUS BRISES PASCOE,
WITH A DISCUSSION OF THE CYPHALEINI (COLEOPTERA,
TENEBRIONIDAE)
_ by E. G. MATTHEWS
ADDITIONS TO THE COLLEMBOLAN FAUNA OF HEARD
ISLAND
by PENELOPE GREENSLADE
MOLLUSC TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN
MUSEUM. 3. POLYPLACOPHORA
by W. ZEIDLER and K. GOWLETT
A REVISION OF THE SOUTHERN AUSTRALIAN STARFISH
GENUS NECTRIA (ASTEROIDEA: OREASTERIDAE), WITH THE
DESCRIPTION OF A NEW SPECIES
by W. ZEIDLER and F. W. ROWE
REDESCRIPTION OF THE MYSID CRUSTACEAN, NOTOMYSIS
AUSTRALIENSIS (TATTERSALL) COMB. NOV., REPRESENTATIVE
OF A NEW GENUS
by KARL J. WITTMANN
SOUTH AUSTRALIAN MUSEUM
North Terrace, Adelaide
South Australia 5000
A REVISION OF THE TROGLOPHILIC GENUS BRISES PASCOE, WITH A
DISCUSSION OF THE CYPHALEINI (COLEOPTERA, TENEBRIONIDAE)
BY E. G. MATTHEWS
Summary
Eight species and one subspecies of Brises Pascoe are recognized, keyed, and briefly described.
Four of the species are new and named caraboides, katherinae, occidentalis and nullarboricus. The
specific name duboulayi Bates becomes a subspecies of acuticornis Pascoe, and granulatus Carter
becomes a synonym of the latter. There is a brief discussion of the cave and burrow frequenting
habits of Brises.
A number of former subfamilies and tribes of Australian Tenebrionidae are combined into the
single tribe Cyphaleini and adult diagnostic characters for the latter are proposed.
A REVISION OF THE TROGLOPHILIC GENUS ARISES PASCOE, WITH A DISCUSSION OF THE
CYPHALEINI (COLEOPTERA, TENEBRIONIDAE)
&. G. MATTHEWS
South Australian Museum, North Terrace, Adelaide, S.A. 5000
(Manuseript accepted 5 Noveinber 1984)
ABSTRACT
MATTHEWS, Bb. G. 1984. A revision of the troglophilic genus Brives
Pascoe, will) a discussion of the Cyphaleiii (Coleoptera). Kee
S Austr. Mus. V9(6): 77-90,
Eight species and one subspecies of Brises Pascoe are
recognized, keyed, and briefly described, Four of the
species are new and named caraboides, katherinae,
accidentalis, and aullarboricus. The specifie name
duboulayi Bates becomes a subspecies of aeuticprnis
Pascoe, and granulatus Carter becomes a synonym of
the latter. There is a brief discussion of the cave and
burrow frequenting habits of Brises,
A number of former subfamilies and tribes of
Australian Tenebrionidae are combined into the single
tribe Cyphaleini and adult diagnostic characters for the
jaller are proposed.
INTRODUCTION
The genus Brises Pascoe, 1869, comprises cight known
Species whose distributions are practically confined to
the arid zone of Australia, with most known records
being within Sawh Australia (Fig. 23). lt is unique
among Australian ‘Tenebrionidae in that the majority
of species have been recorded from caves and mammal
burrows,
Brises belongs to the very large and diverse
Australasian endemic (mbe Cyphaleini (subfamily
Tenebrioninae) Which represents the most important of
the three dominant elements of the Australian
fenebrionid fauna, the others being the Adeliini
(subfamily Lagriinae) and Amarygmini (subfamily
Tenebrioninae) (Kaszab 1982), Having probably
originated as forest-inhabiling, rotten-wood-feeding
forms (as many still are) cerlain elements of the
Cyphaleini invaded the arid zone lo become soil
inhabiting as larvae and ground foraging as adults, but
they do not remain active under daytime conditions as
do many African desert tenebrionids. The concept of
the tribe Cyphalcini adopted here largely conforms Lo
that implied by Doyen and Tschinkel (1982) and includes
all the members of the “subfamilies” Helaeinac,
Nyetozoilinae, Briseinac, and Cyphaleinae of Carter
(1926), 4 lotal of 46 genera and some 450 species. in
Australia and New Guinea, plus the 14 species of
Mimopeus Pascoe in New Zealand (Watt 1968),
June, 1986
The following abbreviations for institutional names
are used in the (ext and follow the four-letter system
proposed by Watt (1979):
AMSA —Australian Museum, Syciey.
ANIC —Australian National Insect
Canberra,
BMNH —British Museum (Natural History),
MYMA— Museum of Victoria (Natural History and
Anthropology), Melbourne.
QMBA —Queensland Museum, Brisbane.
SAMA —South Australian Museum, Adelaide
UOBA —University of Queensland Entomology
Department, St Lucia,
WAMA —Western Australian Museum, Perth.
The primary types of all ihe taxa of Brises have been
seen by the author,
Collection,
The Limits of the Cyphaleini and Tenebrionini
As discussed by Doyen and Tschinkel (1982), these two
tribes belong to the “tenebrionine lineage’ which is
characterised in part by having the spermatheca derived
from the original bursa copulatrix (which |ras
disappeared as a recognizable entity), by the orientation
of the coxite and paraproct baculi of the ovipositor
being transverse and oblique respectively (Fig, 1), and
by the defensive gland reservoirs opening between
sternites 7 and 8 and being small and not reinforced by
helical thickenings (Tschinkel aid Doyen 1980). These
three features are shared with the closely related tonicine
and opatrine lineages (best treated as further tribes of
the Tenebrioninae) and sharply distinguish these lineages
collectively (in effect the subfamily Tenebrioninae) from
other subfamilies. The above authors do not propose
any formal classification, but | equate Lhe subfamily
Tenebrioninae with their tenebrionine, toxicine, and
Opatrine lineages,
The Tenebrioninae (as understood here) are further
characterised by a number of primitive features,
including the almost complete absence of the clustered
antennal sensoria termed “tenebrioid sensory organs”
by Medvedev (1977) and, inconsistently in apparently
primitive elements, internally open fore coxal cavities
and a subcubital fleck on the wings (Fig, 2).
The two tribes Tenebrionini and Cyphaleini share all
of the above features, at least in part, and in a natural
system could not be satisfactorily separated. However,
78 REC. S. AUST, MUS, 19(6): 77-90
the resulting group would be rather unwieldy on a
world-wide basis and it is desirable for purely practical
purposes to try to find an arbitrary line of separation.
‘att (1974) suggests a larval character—a crenulate
spiracular peritreme, combined with a primitive adult
character—the subcubital fleck, to characterise
Cyphaleini. However, too few larvae are known for us
to be able to understand the significance of the
spiracular character, and the adult one is not only
plesiomorphic but also by no means universal in the
Cyphaleini. For practical reasons | prefer to draw the
line at an adult apomorphic character—the absence of
original elytral striae—to recognise Cyphaleini, when
combined with certain other features.
paraproct
accessory gland
June, 1986
Specifically, Cyphaleini may be considered to be
members of the Tenebrioninae which share the
following adult characters (A—apomorphic; P—plesio-
morphic): (1) absence of tenebrioid sensoria (P),(2)
internally open fore coxal cavities (P), (3) the third
antennal segment longer than the preceding or
following ones (A), (4) the original nine elytral striae
entirely absent, the elytra either non-striate or with
secondary, usually supernumerary striae (A), (5) no
sexual dimorphism in the shape of the fore legs (P),
and (6) the parameres of the aedeagus with a vestiture
of minute spines (A?). As far as I know, all Cyphaleini
share the above features, but conversely none of the
features are exclusive to the group. 7eneristes Pascoe,
new crossvein
Ss, i =
1A interrupted
CuP and 1A
partly merged
FIGS, 1-4. 1. Ovipositor and iniernal female genital apparatus of Brises a. acuticornis Pascoe, 2, The renebrionine wing venation (Meneristes
australis Blessig). Venation nomenclature after Medvedev (1968). 3. The cyphaleine wing venation (Brises uw. aeuticarnis Pascoe). 4.
The coelometopine wing venation (Enevalesthus utraviridis Macleay).
REVISION OF THE GENUS BRISES a4
int particular, is a transitional genus be ween Cyphalein
and Tenebrionini, since it displays characters |, 2, 3,
und 6, but not 4 and 5, and it was placed in the
Cyphalemi by Watl (974), a position supported by
Doyen and Tschinkel (1982), | preter to cansider the
clytral character (4) to be decisive, because of the case
with which it can be seen, and so place Weneristes in
the Tenebrionini.
Additional characters which are frequent bul nat
universal in the Cyphaleini are the presence of a
subcubital fleck on the hind wings (P) (in ten of the
20 winged gener examined, but not in Brises), the last
maxillary palpal segment triangular or securiform (A)
(29 of 33 penera), a Shortening Of the parameres in
relation to the tegmen of Lhe aedeagus (A) (14 of 33
genera examined) and a sclerotisation of the median
lobe, olen in a characteristic form (arrow-shaped in
Brises) (A), In addition, eight of the wiliged genera
examined (B8risey Pascoe, Prerohelteus Breme,
Emeephalus Wivby, Paraphines Nacleay, Ospidus
Pascoe, Burvliphia Pascoe, Anausis Bates and
Chartoptervxy Westwood) display a venauional
peculiarity illustrated tn Fig. 3, whereby }A is broken
at one spor and a new cross vein appears, ustially only
asa stub, further proximad between CuP and 1A
(compare the normal 1enebrionine venation seen in
Meneristes, Fig. 2), The later configuration also
appears in at least 12 geaera of Cyphaleini, so wing
venation cannot be used reliably as a tribal character.
The genus 7irwena Erichson clearly belongs in the
Cyphaleini as here conceived since 1 shaws all the
diugnostic features mentianed above except (3) and
some of the additional characters (a subcubital Meck,
securifoun maxillary palpal segment, and selerotisation
of the median lobe), However, Titaene is apparently
also close to Ariys/ona Baies of New Zealand and
Cullismilax Bates of New Caledonia, all three placed
in a tribe Titaenini by Kaszab (1982). The latter lwo
genera display the full complement of original elytral
striae (al lease in some species) and so do not fall within
the Cyphaleini as here arbitrarily defined. | have not
closely examined Artysrend and Callismilay and prefer
to leave the question of their relationship (and therefore
that of Ti@era) in abeyanee,
Ir shouldalso be mentioned here that the subfamily
Coelometopinae, which is sharply distinclive on female
vonitalic and gland reservoir characters (Doyen and
Tschinkel 1982), can often alsa be recognized hy a
characteristic Wing venation in which the veins CuP
and tA have completely merged for a short distance,
swallowing crossvein cu-n (Pig. 4). This modifigation
is aecampanied by a redaction of the vannal atea,
However. a few eoelomeropines have a normal
lenebrionine venation (ag. Teragonomenes Chevrolat,
Charivihecu Pascoe), Coclomeopinae strongly
resemble Tenebrionint in exterial form but can always
be recoenized without disseclion by the presence of
tenebnioid sensoria on the antennae, universal i the
group. The following coelamelopine genera were
erroneously placed in the Tenebrioninae by Carter
(1926); //ypuulux Bates, Hyedissus Pascoe, Encvalesthus
Motschulsky, Serenis Motschulshy, Proiefhis Pascoe,
Tetrazonomenes Chevrolat, Gectosis Pascoe and
Zophophilus Fairmaire (the correct placement of some
of these genera has already been proposed by Doyen
and Ischinkel (1982)).
The Australian genera which L consider to be true
Tenebrionini are -Asphales Pascoe, Sladnea Carter,
Menerisres Pascoc, an unnamed genus comprising Lhe
species Colydioides Enichson, rectibusis Carter and
srdnevunus Blackburn, all three erroneously put in
Menephilus, and Paratoxicum Champion,
The 46 genera of Cyphaleini will not be listed here
but can be obtained from Carter's (1926) checklist
under the “subfamilies’’ Helaeinae, Nyetozailinac,
Briscinae, and Cyphaleimae.
Subgroups of Cyphaleini and Recognilion of Brises
The placing of the eyphaleine genera in four separate
subfamilies by Carter (1926) suggests Chat some natural
groupings may exist within the tribe, perhaps at
subtribal level. In the course of the present study |
dissected representatives of 33 cyphaleine genera but
was tunable to arrive at any arrangement, and believe
that the former categories cantar be maintained at any
level. There is no concordance in the pattern of
variation of characters, whose States consequently
follow a mozaic distribution, This is a characteristic
feature of the Tenebrionidae as a whole and one whieh
has so far defeatedall attempts to arrive at a convincing
natural classification of the family (Doyen ana
Lawrence 1979; Doyen and Tschimkel 1982).
Brises \iself was singled out by Carter in 19]4, and
placed in its own subfamily in 1924, because of a
number of ostensibly unique features, specifically the
narrowly (triangular shape of the last maxillary palpal
segment (normally strongly secuntonn in Cyphaleini),
(he Narrow posterior intercoxal process, the elongate
lugs, and the long tibial spurs and tarsal claws, After
a detailed examination of tepresentative Cyphaleini 1
find jt difficult to understand just what is so unusual
about Brises, Allo rhe above characters recur in other
genera (but not in the same combination). Like all
eyphaleine genera Arises may be recognized anty
through a unique cembinayen of a number af
characters, enumerated for Brises below under
“Diagnosis” ff also has a distinctive habitts fesembling
that ol cursorial carabid beetles, due mainty la its
relatively narrow prothorax and long legs. This charac
teristic build ts the principal feature distinguishing i
from same species of Plerohelaeus Breme, in which the
outlines of rhe promotum and elyira are confluent, but
which in other respeers resemble Brises. Plerohelacus
itself appears (o be a Compostie taxon.
RO REC, 8, AUST, MUS. 19(6): 77-90
Adaptations of Brises
The Cyphaleini are a relatively primitive group of
Tenebrioninae which has radiated extensively in the
Australian region in the absence of the subfamily
Pimehinae (Tentyriinae), which elsewhere tends ta
became dominant in similar xeric situations. Brises
itself has evolved a strategy of utilising caves and
burrows lor daytime shelter at Jeast, Six of the nine
taxa have been recarded fram caves (caraboides, a.
acuticornis, a. duboulayi, katherinae, occidentalis, and
trachynotoides), and three of these {the two subspecies
of aculicernis, and frachynatoides) fram rabbit
burrows as well, Two of the remainder are recorded
from rabbit burrows only (b/airi and parvicollis), only
nullarboricus being without any data on shelters used.
G. B. Monteith (in ditt.) writes that south of Birdsville
Hlairi once came out at night in August in enormous
numbers foraging on the crests of sandhills, even
though it was freezing cold. B a. acuticornis feeds
inside caves and so, presumably, does kutherinue (sec
avcounts uiider species), Feeding habits of the other
species are not known, Larvae which have been found
in caves and borrows in association with Arises adults
are assumed 1o belong to the latter, but some Helaeus
species also use burrows (Matthews 1985) and it 1s
necessary to rear larvae through to confirm their
identity.
By examination of gut contents and through
observation of captive specimens of acuticornis | infer
that the adults are general scavengers like mast tene-
brionids. In burrows ihey (and presumably all other
species of the genus) are probably coprophagous on
the droppings of the mammal inhabitants, and in caves
they appear to feed on all faecal matter available and
an carrion.
Adaptation of &rises ta hfe in caves and burrows is
barely reflected in theic structure. Long appendages,
small eyes and a large hind body are features often seen
in cavernicolous beetles. as is wing reduction. [n all
these respects Brises is only at the earliest stage af
modification, caraboides being the most modified.
Only two species appear to be flightless and only five
haye eyes measurably smaller than those of winged
epigean genera. Intensity of pigmentation js al normal
levels, but paler specimens are common in duboulayi.
Altogether, the morphology of the specics suggests that
there is frequent migration from one burrow or cave
to another.
SYSTEMATICS
Brises Pascoe
Brises Pascoe, 1869, p. 145; Carter, 1914, pp. 45, 46;
Carter 1926, pp. 127, 145.
Ephidonius Pascoe, 1869, p. 15]; Carter, 1914, p. 45
(syn.).
June, 1986
Type Species: OF Brises: Brises trachynotoides Pascoe,
1869, by monotypy. OF Ephidenins: Ephidenius
acuticornis Pascac, 1869, by monotypy.
Descripiien: Entirely black.
Head; Anterior margin of clypeus concave. Clypeo-
frontal suture complete or not. Basal membrane af
labrum visible from above, Eyes entire bul constricted
by both canthus and edge of occiput, the dorsal
interocular distance 23-5 tires width of one eye, Gular
Sutures meet near middle, no gular pits, anterior edge
of gula unmodified. Bridge of tentarium straight. Third
antennal segment longer than 2nd or 4th. Tormae of
labrum transverse, wilhoul prominent anterior
extensions. Mandibles bidentate, mola not striate.
Lacinia unarmed. Terminal segment ol maxillary
palpus narrowly triangular to sublinear (Fig. 5), of
labial palpus sublinear. Angles of mentum anterior,
subacute,
Thorax: Margins of pronoium prominent but rarely
strongly explanate, anterior edge evenly arcuate.
Outline of pronotum not confluent with thar af elytra,
narrower, Pronotum glabrous or wilh minute, very fine
setae. No sharp prosternal keel. Fore coxal cavities open
internally, Mesasternum strongly excavate to receive
prosternal process, Metendostermite Y-shaped, with
laminae. Scutellum visible, small, in form of
subequilateral triangle, Elytra glabrous or with minute
setae, true striae absent, cannae present or nol.
Epipleura moderately wide, complete. Wings withour
subcubital fleck or with famine trace of one, with base
of first branch of 1A broken and partial supplementary
cross vein cu-a present (Fig. 3). Tarsi slender, claws
long, equal in length Lo about 24 of last tarsal segment.
Tarsal vestiture in form of long bristles. Tibial spurs
equal in length to 4-4 length of metatarsus. Legs
slender, withoul carinae.
Abdomen: \ntercoxal process of first visible
abdominal sternile narrowly triangular, Reservoirs of
defensive glands small, simple. Ovipositor and internal
female genital tract of tenebrionine type (Fig, 1).
Parameres. of aedeagus spinose, comprising |4-'4 of
tolal aedeagal length, with backward prolongations,
Median lobe arrow-shaped or simply expanded
Sulbapically (Figs. 11-14).
Tota! length J0-25 mm,
Dragnosis: Eyes in dorsal view ovoid or subquadrate,
not strongly (ransverse, interocular distance equal to
2¥2-5 eye widths. Membrane at base of labrum exposed
by concavity of elypeal margin. Lasl segment of
maxillary palpus narrowly triangular. Pronotum
glabrous, markedly narrower than elytral bases and not
confluent in outline with elytral edges, Elytra glabrous
or with minure setae, the lateral margins not expanded.
Wings (when present) without distinct subcubital tleck.
Legs slender, femora extending Jor about 4-7 of their
length beyond bady sides. Tibial spurs equal 10 ‘4 to
REVISION OF THE GENUS BRISES
6
ae ip
oe
f
N
hi |
a nm 1
10
—
ah LAR
ye “
5 if th ADs h
te \ I i
| hp 9s a
; ‘:
\
11
14
FIGS. 5-14. 5. Outline of terminal maxillary palpal segment, Brises trachynotoides Pascoe. 6. Antenna, Brises nullarboricus n. sp. 7.
Antenna, Brises a. duboulayi (Bates). 8. Hind tarsus, Brises caraboides n. sp., setae omitted. 9. Hind tarsus, Brises occidentalis n. sp.
10. Hind tarsus, Brises trachynotoides Pascoe. 11. Aedeagus of Brises a. duboulayi (Bates) in side view. 12. Apex of aedeagus of Brises
a, duboulayi (Bates) in ventral view, showing arrow-shaped end of median lobe. 13, Aedeagus of Brises trachynotoides Pascoe in side
view, 14, Aedeagus of Brises trachynotoides Pascoe in ventral view, median lobe also shown partly extracted,
#2 REC, S. AUST,
—
© length of metatarsus. Claws long, equal in length
to about *: of last tarsal segment. ‘larsi bristled
beneath, nor tomentose. |ntercoxal process of first
visible abdominal sternite narrowly triangular,
Distribution (Pig. 23): All of South Australia exeept
the southeastern portion; New South Wales west of the
Great Dividing Range; southwestern Queensland; the
Northern ‘Territory trom Katherine southward; the
Nullarbor, Central, and North West districts of Western
Australia, Probably occurring in all parts of Australia
receiving less (han 300 mm ol annual rainfall, with
populations substantially outside this area possibly
isolated and restricted to caves.
KEY TO THE SPECIES AND SUBSPECIFS OF
BRISES
1 (2) Homeralanules stronely explanule, Llyiral length about 13
times basal elyrral width, Wingless (Pin 24)
hire Aer trie | eee eee F Adtire Carrer
Huinnaeat unules rounded. Elytral length 16-2.1 cimnes hasal
culyiral width. Wings present bul may be reduced .., 2
21) Elyrral surlace with punctires arranged in rows, sametinies
integular, Granules or spines if present, also in rows. Head
und pronotum usually shayieerned and punctate, Terminal
antennal seument more or less acuntinale (Fiz, 7). No sexual
dimorphisny .0 . 2.00 2-2 2222 eee eee -Aota
Elytral surfaces coursely reticula: pimeiate arid vranulate, the
punetures and granules never in rows on dise at least. Head
and pronotum densely vermiculate or eranulare. ferminal
antennal segment nor acuminate, at mast somewhat
nurrowed apically (hig. 6), Male with middle and hind
POC EMT 8
femora modificd (Kies, 19-22
4(2) Pronotal surface shiny, finely Shavreened and purictate only,
without granules Se 62 5 Seg tuuncoct® 4
Pronou suriaee malt, with at Thad a few ‘small aranules
mai)
4.(3) Hind beds oval (hig. 25), Mlighiless. Livtea with the mirervals
between low, rounded carinaé enarsely and Uensely pune
fale, without eranmules 6). 02.0, 2) carahoides in. sp,
Hind body oblong (lip, 2A), wines normal blytral surface
finely punckile, Wily Ur Without granules and carinac
5
S (4) Elytra withoue trace of spines, granules or sharp cariae
3 acullcormis acuticarnis (Pascoe)
L lyirs with au lease a few small oranules, niay We distenenly
eranulate and caripate
. 8 deulleorais
+ idehe naleyl (Bates)
(3) Head and pronoun coursely bupose und yranulate. Prosternal
process with the margins sharply raised between fore corue.
Liviral surface With short setae, more prominent laterally,
ely tral edges nearly straight in middle (Pig. 29)
A S. katherinae n. sp.
Head and provonin witht small erannles, the surtace berween
them shagreened and finely ruigose. Prostermal pracess With
margin not raised, Elyira Wwithoue serae, elytral cdyes
distinedly comes throughout (Figs, JO and 31} 7
TH) byes wider than lone, separated by about 4 eye widilrys
(Fig, JT), Elytpal carinae very prominenr the suituee
herween rhem concave and distineily eriqulace
6 vecidentals mh. 4p.
MUS. JOG): 77-90
June, 1986
Fyes about as long as wide and separated by ahout 3 eve
widths (Pip. (8), Elyiral carinae ting, rhe surface berween
thom Mal, minutely eranulate, with fine supplementary
fongiiudinal ridves .., 7 pervicollis (Blackburn)
8 (2) Clypeusin lower pline than trons, clearly demarcated ar hinse
Front cdges of eves strongly oblique (Fig. 14), Elyirawith a
shor) furmeral carina (hig, 32) 8 auad/arhorteas ne sp,
Clypeus in same plane as frons. not sharply delimited at base.
Dron edges of eyes more or less uransverse (Fav. 16). Ebytey
withoutahumend carina ., 9 dwehvaotoides Paseo
lL. Brises blairi Carter
(Fig. 24)
Brises blairi Carter, 1914, p. 58; Carter, 1926, p. 145.
Descriptions Mentum with two pits and broad
anterior membranous area. Terminal antennal segment
apically narrowed bul obtuse. Clypeus in lower plane
than frons. Eyes with length subequal to width in dorsal
view, separated by a distance equal to about 2'4 eye
widths. Edges of prosternal process not raised.
Pronotum 1.7-1.9 times as wide as long with margins
stronyzly explanate. Pronotal surface alutaceous, with
scattered very fine punctures. Elytral surfaces with yery
small granules, denser anteriorly, and very small
punctures more evident anteriorly and along sides, the
surface between them alutaceous, without setae. Two
incomplete raised carinae or rows of granules on each
elytron. Ratio of elytral length to width across bases
about (3:1. Hind wings almost entirely atrophied.
Longest hind spur equal to more than '% length of
metatarsus, Parameres making up about 4 of total
aedeagal length. No sexual dimorphism. Total length
15-20 mm.
Distribution and habitat The north-east quarter of
South Australia and adjacent arcas of Queensland and
the Northern Territory. Not recorded from caves but
found in rabbit burrows and in the open on sand ridges
al night, sometimes in large numbers. Collected from
July to May.
Type: Killalpanima, 5.A., 100 miles east of Lake Eyre
(H. J. Hillier), o. BMNH.
Specimens examined (119); SOUTH AUSTRALIA,
Alton Downs (old) H.S., 48 km SW by W of Birdsville.
Birdsville, 25 mi S of Clayton R., near bore drain.
Cooper's Creek, Diamantina R., 25 mi S of Birdsville.
Hay R., 24 mi W of camp J3. Innamincka, Brodic’s
Water Hole, Lake Byre, Madigan Gulf, Sulphur
Peninsula, Lake Eyre, Prescott) Peninsula. Lake
Kittakittaooloo, S shore. Lake Palankarinna. Minnic
Downs, NE corner. Mt. Gason, 4] km SSW of Clifton
Hills. Mungeranic Sta., water hole, Mungeranie, 20 mi
S of. Purni Bore, Simpson Desert, 6 km WSW of, pit
traps in sand ridge. Simpson Desert. Warburton R.,,
2km NE of Kalamurina H.S. Warburton R,, New
Kalamurina H.S. NORTHERN TERRITORY. Andado
H.s., 15 km ENE of. Vinke R., MeDonnell Ranges.
Hermannsburg. Indracowra, 5 mi N of, ex pit traps in
rabbit burrows. QUEENSLAND, Arrabury, 20 mi N
REVISION OF THE GENUS BRISES 83
19
21
16
20
22
FIGS, 15-22. 15. Head, Brises nullarboricus n. sp. 16. Head, Brises trachynotoides Pascoe. 17. Head, Brises occidentalis n. sp. 18. Head,
Brises parvicollis (Blackburn). 19. Hind femur of ¢, Brises trachynotoides Pascoe. 20. Middle femur of ¢ , Brises trachynoloides Pascoe.
21. Hind femur of ¢, Brises nullarboricus n. sp. 22, Middle femur of 3, Brises nullarboricus 0. Sp.
of, on road to Planet. Birdsville. Birdsville, 30 km W
of, Bluff Sta. Diamantina R. Kaliduwarry Sta., Camp
20. Specimens are located in AMSA, ANIC, QMBA,
SAMA, UQBA, and WAMA,
2. Brises caraboides, Nn. sp.
(Figs 8, 25)
Description: Mentum without pits, with irregular
surface and narrow anterior membrane. Terminal
antennal segment subacuminate. Clypeus in lower
plane than frons. Eyes somewhat wider than long in
dorsal view, separated by a distance equal to about 3%
eye widths. Edges of prosternal process not raised.
Pronotum about 1.3 times as wide as long. Pronotal
surface finely shagreened, with scattered fine
punctures. Humeri rounded. Elytral surfaces with
about 15 rows of coarse punctures in straight lines,
closely spaced between four low, more or less evenly
spaced rounded longitudinal ridges on each elytron,
without carinae, with short fine setae. Ratio of elytral
length to width across bases about 1.9:1. Hind wings
reduced to about half of elytral length. Longest hind
spur equal to about 4 of metatarsus. Parameres
Keb REC, S, AUST. MUS. 19(6): 77-90
making up about '4 of total aedeagal length. Total
length 18-19 mm.
Remarks: The species is closely related to acuticornis,
within whose range it occurs and from which it differs
mainly in pronotal and elytral shape and puncturation,
by being flightless, and by having longer legs. The
narrow bumeri indicate that wing atrophy js of long
standing and no intermediate forms are known to exis
between caraboides atid deuticornis.
Distribution and habitat; The type locality only.
Collected im an “alcove al base of cliffs”.
Tipe; Wwilight Cove, Eucla Basin, WA., 5.X1.1966,
J, Lowry, or, ANLC, Paratype: One or, with same data
as holotype, ANIC.
Brises acuticornis (Pascoe)
Description: Mentum without pits, sparsely punctate,
wiih posterior V-shaped grooves enclosing raised area.
Terminal antennal segment acuminate. Clypeus in lower
plane than trons. Eyes about as wide as long, separated
by a distance equal to about 4 eye widths. Edges of
prosternal process not raised. Pronotum |,4-1,.5 times
as Wide as long, Pronotal surface alutaceous, finely and
sparsely punctate. Elytral surfaces variable (see under
subspecies), with short setae. Ratio of elytral length to
width across bases 1.6-1.7:1, Hind wings fully
developed, Longest hind spur equal to about ' length
of metatarsus. Parareres making up about of total
aedeagal length. No sexual dimorphistn. Total length
15-25 mm.
I recognize the following two subspecies (taxa 3
and 4),
3. Brises actiticornis acuticornis (Pascoe)
(Figs I, 3, 26)
Ephidonius ucuticarnis Pascoe, (869, p. 151.
&rises weuticornis, Carter, 1914, pp. 45-46; Carter,
1926, p. 145; Hamilton-Smith, 1967a, pp, 37-39, 41;
Hamilton-Smith, 1967b, pp. 115-116; Richards, 1971,
pp. 17-45 passim.
Description: Elytral surfaces with small punctures
set in longitudinal rows between three low ridges on
vach elytron, without granules or spines,
Distribution and habtiat; This subspecies generally
occurs in the coastal areas of South Australia and the
Nullarbor Plain, There are numerous records fram
caves, from both the light and dark zane, often
associated with guano and carrian. Probable larvae
have been colleeted inside caves and one collector
(M, Gray) reports that adults and larvae feed on weta
(raphidophorid cricket) excreta in Weebubbie Cave.
Collected from July to April.
In her study of the cavernicolous fauna of the
Nullarbor Plain, Richards (1971) reports B acuricornis
from 25 caves throughout the region, mostly in the dark
June, LYKA
zone up to 4 km from the entrance, Dry, powdery bat
guano supports laree populations of adults and larvae,
anc this species is the only coprophage to oceur in both
bird (swallow) and mammal faeces, including humati
excrement and fox droppings, It also occurs im rabbit
burrows on the plain.
Type: Lectotype (sex undetermined) [rom Gawler,
S.A., herby designated from two syntypes in the Pascoe
Collection (BMNH).
Specimens examined (174): SOUTH AUSTRALIA,
Ardrossan. Ceduna, Coak, 9 mi E of. Denial Bay, | mi
8 of, Diprose Cave No. 3. Eucla Basin, eave N 149,
Fisher, E-W Railway. Fowler's Bay. Crawler. Koonalda
Cave sinkhole. Koonalda H.S., 10 km SE by E of.
Koonalda Sta. blowhole entrance. Koonalda, N 33
sinkhole (amongst dead birds). Koonibba.
Kooringabie Sta. Murrawijinee Cave N 7, Nullarbor.
Murray. R, Ooldes, Pt Pierce Mts, Yorke Peninsula,
Swan Reach, Punyelroo Cave. Thylacine Hole, Eucla
Basin. Wardang 1. White Wells Cave, Winbirra Cave,
Eucla Basin. Yorke Peninsula. WESTERN AUST-
RALIA. Abrakurrie Cave N 3, Nullarbor, Caiguna and
Cocklebiddy, halfway between, ex rabbit burrow.
Cocklebiddy Cave, doline. Dingo Cave N_ 160,
Nullarbor Plain, on guano. Eucla. Eucla, 32 mi SE of
Coongana, Nullarbor. Eyre’s Sand Patch. Madura,
6 miS of, cave. Madura, 8 mi Cave, Madura, main cave
N 62. Moonera Tank Cave, Nullarbar. Mullarmullang
Cave, Nullarbor, dark zane, feeding on dead bat,
Murra-cl-elvyn Cave, Nullarbor, light and dark zones,
an dry guano, Nullarbor, 50 km W of; 64 km E of;
7 km WSW of. Nullarbor H.S. Pannikin Plain Cave.
dark zone. Petrogale Cave, 14 mi S$ of, N 79, E of
Madura. Swallow Cave, Cocklebiddy, near dead bat-
Tommy Graham's Cave N 56, Nullarbor. Weebubbie
Cave N 2, Nullarbor. Specimens are located in AMSA,
ANIC, BMNH, OMBA, SAMA, UQBA, and WAMA.
4, Brises acuticornis duboulayi (Bates), new siatus
(Figs 7, 11, 12, 27, 28)
Ephidonius duboulayi Bates, 1872, p. 279.
Brises duboulayi, Carter, 1914, p. 45; Carter, 1926,
p. 151.
Brises granulatus Carter,
synonymy),
1921, p. 314 (new
Description: Distinguished from a, aculicernis only
by the sculpturing of the elyira, which shaw at lease
a trace of small granules or spines and an aceentuation
of the ridges. In the extreme form, the elytra are
distinctly spinose and strongly carinate. There is a
gradual intergradation between typical a, wcntcarnrs
and a, dubouluyi along geographical gradients (sec
below).
Distribution and habitat; This widespread subspecies
is essentially the inland race ot acuricornis, bur it
reaches the coast on Eyre Peninsula, where jt 35 also
REVISION OF THE GENUS BRISES 85
in its least granulate form. This form could have been
ascribed to a. acuticornis, but it is more convenient for
nomenclatorial purposes if the latter name is arbitrarily
restricted to specimens without any trace of granules,
spines, or carinae. Moderately granulate forms without
carinae (the former granulatus, Fig. 27) are to be found
in eastern South Australia from the Adelaide Hills
northward and in New South Wales as far east as the
foothills of the Great Dividing Range, whilst the most
strongly carinate and spinose form (@. duboulayi
proper, Fig. 28) occurs in northern South Australia
from Port Augusta northwards to adjacent states and
westwards through the driest parts of Western Australia
to the west coast. In parts of South Australia it is
difficult to assign specimens to one form or the other.
The Ashford Cave (N.SW.) population of this
subspecies seems to be isolated by many hundreds of
kilometres from the nearest other records in western
New South Wales, but the intervening area is too poorly
collected for us to be sure of this, The few Ashford
Cave specimens seen are smaller and paler than other
duboulayi. The other extreme eastern record, Yiddah,
N.SW., is based on a single old specimen and needs
investigation.
250
There are several other records of the occurrence of
this subspecies in caves, and in rabbit burrows (see
below). Records are from October to July.
Types: Of duboulayi: Champion Bay, W.A., Duboulay,
3, BMNH. Of granulatus: Broken Hill, N.SW., R. J.
Burton, 1/21, 9, MVMA.
Specimens examined (108); SOUTH AUSTRALIA,
Adelaide. Buckalowie Creek, from small cave a little
beyond light. Clara St Dora Cave. Cleve. Coober Pedy.
Evelyn Downs Sta., Qodnadatta. Everard Ranges to
Warburton Ranges. Franklin I. Kingoonya. Kokatha
H.S., 17 km SE of, ex pit traps in rabbit burrows. Leigh
Creek. Minnipa, Mt Finke, 40 km S Malbooma H.S.,
ex rabbit warren. Mt Lofty Ranges. Moralana H.S., 22
km WNW of. Murray R. Pt Augusta. Pt Lincoln.
Purple Downs, 160 mi NW of Pt Augusta. Streaky Bay.
Stuart Ranges. Tarcoola. Whyalla. Wilgi near Loveday,
Gilbert's Well. Wilpena Pound, in bat cave in dung.
Wooltana Cave, 200 ft underground in bat guano,
NEW SOUTH WALES. Ashford Caves via Inverell.
Broken Hill. Culpaulin, Darling R. Yiddah,
QUEENSLAND. Diamantina. NORTHERN
TERRITORY. Alice Springs, 15 km S of, bat caves.
FIC), 23. Distribution of the species of the genus Brises. 1—B. blair Carter. 2—B. caraboides vn. sp. 3—B. a. acuticornis (Pascoe). 4—B
a. duboulay! (Bates), 5—A, katherinae 0. sp. 6—B. occidentalis o, sp. 7—B, purvicollis (Blackburn). 8—B. nullurboricus 0, sp. Y—B.
trachynotuides Pascoe, 250 mm annual isohyet shown,
86 REC &. AUST, MUS, 19(G): 77-90
WESTERN AUSTRALIA. Coalgardie. Cue, Onslow.
Specimens are located in AMSA, ANIC, SAMA, and
UQBA.
5, Brises katherinae, n. sp,
(Fig. 29)
Description: Mentum variable, without grooye but
with cither large punetures ar oblique depression
posterolaferally. Terminal antennal seement acuminate.
Clypeus in same or higher plane than trons, not
delimited by a sharp depression, Byes wider than long,
Ihe anterior edge strongly oblique, separated by a
distance equal to about 2 eye widths, Prosternum feebly
ridged before coxae, edges of prosternal process
strongly raised between coxac. Pronotum [.3-1.5 times
as wide as long. Pronotal surface irregularly rugose and
shagreened, with numerous large shallow punctures
and some slightly raised shiny spots on dise. Elytra each
with three nearly Complete carinae and part ofa fourth,
and a sutural row of granules, surface between Carinae
flat, shagreened, with rews of fine but deep punctures
and sparse short, curved setae. Ratio of elyirval length
to Width across bases 1.8-2.11, Hind wings fully
developed. Longest hind spur equal to abour |i of
length of metatarsus. Parameres making up about '4
of total aedeagal length, No sexual dimorphism. Total
length 15-25 mm.
Distribution and habitats ’nown anty tram the
complex of caves near Katherine, NT. It ovcurs well
inside the dark zone, and has not been collected from
outside caves. More collecting in inland northero
Australia is needed to establish the distribution limits
of this and any other species in the region, in ordet
to determine Whether katherine is really as isolated
us iL appears to be.
Type: Kintore Cave, Katherine, N_T., BS 2134,
12v.1974, @ SAMA, | 21, 155. Paratvpes: Same data
as holotype, 5, SAMA, Cutta Cutta Cave, N.T,, 200 m
from entrance, 5,x11.1983, W. D. Williams, 2, SAMA.
Cutta Cutta Cave, Katherine, NT. 13.i1x.1973, A. Goede,
1, SAMA. Three Mile Cave, Katherine, NLT. 24.yi,1962,
R, V. Southeott and NT, Cave Exploration Group, 2,
SAMA. Katherine, N-T., !¢ mile from entrance to 16
Mile Cave, June 1962, 1, ANIC. Katherine, N.T, 16
Mile Cave, 12.iv.1963, W. Penman, 2, WAMA.
6. Brises avefdentalis, n. sp,
(Pigs 9, 18, 30)
Description: Mentum with postero-lareral grooves,
not joining. Terminal antennal segment acuminate.
Clypeus in somewhat lower plane than trons bul not
sharply delimited. Byes aboul as wide as long,
separated by a distanee about equal to 4 eye widths.
Head somewhat elongated behind eves, Prosternum
distinctly carinate belare voxae, edges ol prosternal
process not raised. Pronotum 1.5-1.6 times as wicle ag
long. Pronoral surface Shavreened, with indistinel
punctures and very small granules regularly spaced.
Aine, 1986
Elytra each with two strong, nearly complete carinac
and two indistinct ridges externally, intervals between
them with sparse granules and punctures in irregular
rows, Without setae. Lateral edges of elytra markedly
convex. Ratio of elyoral length to width across bases
1,8-1,9:1, Hind wings fully developed, Largest hind spur
equal to about ' of length of metatarsus. Parameres
making up about '4 of total aedeagal length. No sexual
dimarphism, Total length 19-22 mm.
Distribution and hubitaty Known only fron three
localities in Western Australia situated between
Meckatharra and the coast. Ore of the Wilgie Mia
specimens bears the label “Found in cave”,
Tipe: Weld Range, W.A., 2.111.1963, A. Douglas, o,
WAMA. Parauivpes: Same data as holotype, 4, SAMA,
WAMA. Mileura H-S. 4 mi W of, 8.xii.1966, W. H-
Butler, 1, WAMA, Wilgie Mia W.A. (27 285 114 21
E), ix, 1961, D, Merilees, 2, WAMA, Ditto, 1963, A.
Douglas, 3, WAMA. Ditto, 16.v.1973, M. Thomas, 1,
SAMA.
7. Brives parvicollis (Blackburn)
(Figs 17, 31)
Ephidonius parvicallis Blackburn, 1895, p, 52.
Brises parvicellis, Carter, 1926, p. LSI,
Deseripijon: Mentum with postero-lateral
depressions, not meeting. Terminal antennal segment
narrowed distally bul bluntly rounded. Clypeus in lower
plane than Frans but not sharply delimited. Eves laree,
Jonger than wide, separated by a distance equal to
about 3 eye widths, Head surface finely, densely
granulate and shagreened. Prosternum moderately
nidged before coxae. Pronotum 1.5-1.7 times as wide
as long, Pronotal surface shagreened with numerous
very small granules, Elytra each with two sharp but
fine, nearly complete carinag, and one Gr (wo indistinet
ones laterally. Intervals between them shagreened, with
distinct straight rows of small punctures and minute
granules, without setae, Lateral edges al elytra
markedly convex. Ratio of elytral length to width across
bases 1.8-2.0:1. Hind wings fully develaped. Lonvest
hind spur equal to abour ' length of metatarsus,
Parameres making up about 4 of toral aedeagal length.
No sexual dimorphism, Total length 17-22 min.
Distribution and habiiau The few avaiable records
suggest One focus of distribution in the cast central part
of South Australia in low-lying aceas and another in
the mountains of the north-west corner and adjacent
ranges of neighbouring states. This species has been
collected from rabbit burrows on several oecasions but
is not known Irom caves and may be an open-ground
forager like bloiri, September to March.
Teper Lake Callabonna, SA. A, Zietz, 2, SAMA,
There is Ho specimen by this name in BMNH, The one
in SAMA is not marked type bul if bears the name
Ephidonius parvicallis in Blackburn’s writing and the
REVISION OF THE GENUS BRISES ‘BT
correct locality and collector, and has been designated
as lectalype. Two ather specimens with the same data
are labelled co-types (that is, paratypes) in Blackburn’s
hand, and (here are five others with the Zietz label,
determined by A. M. Lea as co-types, all designated
paralectotypes and all in SAMA.
Specimens examined (36) SOUTH AUSTRALIA.
Agnes Creek Sta. ex pitfalls in rabbit burrows.
Eradunna H.S., 153 km W oof Everard Ranges to
Warburton Ranges. Glenmanyie Bore E of Lake Frome,
ex rabbil burrows. Lake Callabonna. Lake Mulligan
(possibly Mulligan Springs near Lake Callabonna).
Lake Palankarinna (28 46§ 138 25 EB), rabbit burrows.
Moralana, 22 km NNW. of, rabbit burrows.
NORTHERN TERRITORY, Petermann Ranges, Hull
R. 33 km ESE of Dorker R. Specimens are located in
AMSA, ANIC, and SAMA.
8, Brises nullarboricus, n. sp.
(Figs 6, 16, 21, 22, 32)
Descriptions Mentum with transverse ridges and
lateral concavilies. Verminal antennal segment broadly
rounded. Clypeus in lower plane than frons, sharply
set off by a step, Eyes with anterior edges. oblique, a
little longer (han wide, separated by a distance equal
to abou 3 eye widths. Head surface densely granulate.
Prosternum not ridged, Pronotum 1.6-1.7 times as wide
as. long. Pronotal surfaces densely granulate, Elytra
each with two nearly complele carinae and one short
humeral varina, rest of surface coarsely reticulo-
punctate and granulate, the punctures and granules not
in foWs, Without setae, Ratio of elytral length to width
across bases 1.8-1.9:1. Hind wings fully developed.
Longest hind spur cqual to about 74 length of
metatarsus. Parameres making up about '% of total
avdeagal length. Sexual dimorphism evident in shape
of middle and tind femur, the hind margin of which.
is Slightly expanded at base in the male. Total length
13-17 mm.
Distribution and habitat: Known only from the
Nullarbor Plain, without habitat data.
Type: Nullarbor H.8., 7 km WSW of, 5.A,, 31 28S
13050 E, 11.41.1978, D. C.F Rentz and M. 1 BD. White,
Stop 25, 9, ANIC. Paratypes: Same data as holotype,
3, ANIC. Nullarbor H.S., S.A. 101.1960, PB Aitken,
1, SAMA. Kooringibbie (Well), S.A. 3, SAMA, 744
Mile Camp, East-West Rallway, W.A,, Fo Mack, J,
SAMA. Fisher, East-West Railway, S.A., Troughton and
Wright, 1, AMSA. Reid, 40 mi N of, W.A,, 41.1968,
59/3 A. M- Richards, 1, ANIC,
9, Brises trachynotoides Pascoe
(Figs 5. 10, 13. 14, 15, 19, 20, 33)
Brises trachynoteides Pascoe, 1869, p. 146, pl. xi,
fig, 5; Carter, 1914, p. 46, Carter, 1926, p. 151,
Description: Mentum with indistinet postero-lateral
depressions, Terminal antennal segment broadly
rounded, Clypeus in same plane as frons. Eyes with
anterior edges nearly transverse, wider than long,
separated by a distance equal ro abour 2! eye widths.
Head surface vermiculate. Prosternum shghtly ridged
before coxae, Pronotum |,6-1.9 times as wide as long.
Fronotal surface vermiculate-granulate. Elyira each
with two sharp, nearly complete carinae, no humeral
carina, rest of surface reticulo-punctate and granulate,
the punctures and granules not in rows, without setae.
Ratio of elyctral length to width across bases 1.6-1.8:1.
Hind wings fully developed. Longest hind spur equal
ta about 24 of length of metatarsus, Pararmeres making
up about % of tofal aedeagal length. Sexual
dimorphism evident in shape of legs, the males having
the hind margin of the middle and hind fernera
angularly expanded and all tibiae distinctly arcuate
(straight in the female). Total length 10-17 mm.
Remarks; There is considerable vanability both in
size and shape in this species, the larger individuals
having more explanate and sinuate pronotal margins.
Distributien and habitat; A widespread species
occurring in the northern half of South Australia,
south-western Queensland, the south of the Northern
Territory and all of Western Australia except the South
West. It is not frequently encountered, however, Found
iw rabbit burrows, and there is one series from a cave
on Barrow Island, W,A, August to May,
Tipe: Charnpion Bay, W.A,, 2, BMNH.,
Specimens examined (52); SOUTH AUSTRALIA.
Agnes Creek, ex pitfalls in rabbit burrows, Ediacara,
Emu, 300 mi NW of Woomera. Everard Ranges. to
Warburton Ranges, Mt Irinke, 40 km S of Malbooma
H,S,, ex rabbit warren, Purai Bore, Simpson Desert,
6 km WSW of, pit traps on sand ridge, Stuart Creek
H.S., 10 km E of. Wynbring. QUEENSLAND.
Cunnamulla. NORTHERN TERRITORY,
Hermannsburg. Idracowra H.S,, 5 mi N of, ex pil raps
in rabbit hurrows. MacDonnell Ranges. Petermann
Ranges. WESTERN AUSTRALIA, Barrow [,, cave N
of Flacourt Bay, Forrest River District, Gill Pinnacle,
Mural Crescent. King Sound. Kookynie. Mt Linden,
7.5 km NNW of, Specimens are located in ANIC,
BMNH, QMBA, SAMA, and WAMA,
ACKNOWLEDGMENTS
| wish to thank the curators of the collectians
consulled for arranging loans of specimens, in
particular Mr M, J. D. Brendell (British Museum), Mr
G. Holloway (Australian Museum), Dr T. F. Houston
(Western Australian Museum), Dr J. FO Lawrence
(Australian National Insect Collection), Dr G, B.
Monteith (Queensland Museum), and Dr A, Neboiss
(Museum ol Victoria, Division of Natural History and
Anthropology).
88 REC. S. AUST. MUS. 19(6): 77-90 June, 1986
{ ‘a
memos
NO
nn
FIGS. 24-27. 24. Brises blairi Carter,d. 25. Brises caraboides n. sp., ¢ . 26. Brises a. acuticornis (Pascoe), o . 27. Brises a. duboulayi
(Bates), @ , granulate (eastern) form. Photos: J. Forrest.
REVISION OF THE GENUS BRISES 89
FIGS. 28-31. 28. Brises a. duboulayi (Bates), ¢ , carinate (western) form, 29, Brises katherinae n. sp., o . 30. Brises occidentalis n, sp.,
3. 31. Brises parvicollis (Blackburn), o . Photos: J. Forrest.
90 REC. S. AUST. MUS, 19(6): 77-90
June, 1986
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Tenebrionidac. Proc. Linn Soc. N.S.W. 39: 44-86,
CARTER, H. J. 1926, A check list of the Australian Tenebrionidae,
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DOYEN, J. T., and LAWRENCE, J. F. 1979. Relationships and
higher classification of some Tenebrionidae and Zopheridae
(Coleoptera). Syst. Ent. 4: 333-377.
DOYEN, J. T., and TSCHINKEL, W, R. 1982. Phenctie and cladistic
relationships among ténebrionid beetles (Coleoptera). Syst. nr.
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HAMILTON-SMITH, E. 1967a. Fauna of the Nullarbor Caves. /n
“Caves of the Nullarbor” (J. R. Dunkley and T. M. L. Wigley,
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Sydney. 61 pp.
HAMILTON-SMITH, E. 1967b, The Arthropoda of Australian caves,
J. Aust. ent. Soc. @: 103-118.
KASZAB, Z. 1982. Die Tenebrioniden Neukaledoniens und der
Loyauté-Inseln (Coleoptera). Folia ent. Hungarica 43: 1-294.
MATTHEWS, E. G, 1985. Foraging activity of some tenebrionid
beetles in a South Australian mallee area. Jn “Soil and Litter
Invertebrates of Australian Mediterranean-type Ecosystems” (P,
Greenslade and J. D. Majer, eds.). WA. Inst. Tech., Sch, Biol.
Bull. 12: 63-64.
MEDVEDEYV, G, S, 1968, “Zhuki chernotelki (Tenebrionidae),
Podsemeistvo Opatrinae!” Fauna SSSR Vol. 19, part 2.
Leningrad.
MEDVEDEV, G. S, 1977, Taksonomicheskoe znachenie
antennal‘nykh sensill zhukov-chernotelok (Coleoptera, Tene-
brionidae). Tr Vsesovusn. ent, Obshch, AN SSSR 58: 61-86,
PASCOE, F. P. 1869. Descriptions of new genera and species of
Tenebrionidae from Australia and Tasmania. Ann, Mag, Nat.
Hist. 3: 29-45, 132-53, 277-96, 344-351.
RICHARDS, A. M. 1971, An ecological study of the cavernicolous
fauna of the Nullarbor Plain, Southern Australia. J, Zool, Lone.
164: 1-60.
TSCHINKEL, W. R., and DOYEN, J. T, 1980. Comparative anatomy
of the defensive glands, ovipositors and female genital tubes
of tenebrionid beetles (Coleoptera), Jat. J Insect Morphol.
Embryol, 9: 321-368.
WATT, J. C. 1968: Specific synonymy in Miniopeus Pascoe (Cilibe
auctorum), and the nomenclatoral status of some related genera
(Coleoplera, Tenebrionidae), NZ. Ent, 4; 35-39,
WATT, J.C, 1974. A revised subfamily classification of Tenebrionidae
(Coleoptera). NZ. J. Zool 1: 381-452.
WATT, J.C. 1979, Abbreviations for entomological callections, NZ.
J. Zool. 6: 516-520.
FIGS, 32-33. 32. Brises nullarboricus n. sp., 2. 33. Brises trachynojvides Pascoe, & . Photos: J, Forrest:
ADDITIONS TO THE COLLEMBOLAN FAUNA OF HEARD ISLAND
BY PENELOPE GREENSLADE
Summary
The fauna now consists of 8 species and 2 doubtful records. Friesea tilbrooki, Cryptopygus caecus,
Cryptopygus tricuspis, Isotoma (Sorensia) punctata and Isotoma sp. indet. Are new records for the
island, and the presence of Tullbergia bisetosa and Cryptopygus antarcticus antarcticus is
confirmed. Tullbergia templei was described from Heard I. As was Friesea viennei which is here
synonymised with Friesea tilbrooki, and Cryptopygus quadrioculatus is synonymised with C.
tricuspis after examination of types. Records of Isotoma (Sorensia) subflava and Parisotoma
octooculata from Heard I. Are no longer valid since they depend on incorrect identifications and
Tullbergia antarctica and Cryptopygus antarcticus reagens are considered doubtful records. The
fauna consists mainly of widely distributed species and there is no sign of endemism.
ADDITIONS TO THE COLLEMBOLAN FAUNA OF HEARD ISLAND
by
PENELOPE GREENSLADE
Honorary Research Associate, South Australian Museum, North Terrace, Adelaide, South Australia S000
(Manuscript accepted 30 April 1985)
ABSTRACT
GRELNSLADE, PENFLOPE 1986. Additions to the collembolan
fauna of Heard Island, Ree. 8. otuer Mus. F907): 91-96,
The fauna now consists of 8 species and 2 doubtful
records. Friesea tilbrooki, Cryptopygus caecus,
Cryplopyeus tricuspis, satoma (Sorensia) punciata
and /soroma sp. indet. are new records for the island,
and the presence of Tullbergia bisetosa and
Crvptopyeus aniurcticus antarclicus is confirmed,
Tullberzia templei was described from Heard J, as was
Kriesea viennei which is here synonymised with Friesea
tilbrooki, and Cryptopyeus quadrioculatus is
synonymised with C. tricuspis after examination of
types. Records of /sotoma (Sorensia) subflava and
Parisotoma oelocculata from Pleard Ll. are no longer
valid since they depend on incorrect identifications and
Tullbergia antarclica and Cryptopygus antarclicus
reazens are considered doubtful records. The fauna
consists mainly of widely distributed species and there
is nO sign of endemism,
INTRODUCTION
Heard [. is a small (43 km 19 km) ice-capped
Antarctic island lying in the South Indian Ocean at
53°06'S, 77°30'E about 440 km SE of the Kerguelen
Is and 4100 km WSW of Perth. Parts of the island are
permanently glaciated and it is dominated by a
quiescent voleano, Big Ben, which rises to 2745 m (Law
and Burstall 1953; Horne 1984), Other areas near the
coast and on headlands are free of ice and snow for
sore months of the year and Support vegetation. Seven
vascular plants have been recorded, Several collections
of Collembola have been made from the island
(Table 1). Wise (1970b) dealt with the fauna in detail
and listed seven species while the most recent review
is that of Deharveng (1981) who did not examine
previous collections and gave three definite records and
five doubtful identifications. A new collection of
Collembola from Heard |, was found to contain two
species not recorded before and this prompted an
examination of material collected previously and a
review ol the faunal list, Identifications were made
using Deharvenp's (1981) keys, descriptions, figures and
specimens identified by him unless indicated otherwise.
Subspecies have been retained although this should not
be taken as support by the author for the concept
generally.
Aine, Laser
TABLE IL. SUMMARY OF COLLECTIONS OF COLLEMBOLA
FROM HEARD ISLAND
Records Institution
published of
Collection Year by Deposition
Deutsehe Sud- Vanhoffen 1908
Polar Expedi- \901-3) Enderlein 1909 Berlin?
tion
BANZARE 1929 Womersley 1937 SAMA
(T. Hy Johnston)
ANARE 1951-2 Brown 1964 USPHTM
(kK, G. Brown)
P. Temple 1965 ‘Wise 1970b ANIC (holorypes)
BPBM
5. Tremont 1983 Tremone 1983 SAMA
inecnslade
this work
The following abbreviations are used: AMNZ,
Auckland Institute and Museum, New Zealand; ANIC,
Australian National Insect Collection, Canberra;
BANZARE, British, Australian and New Zealand
Antarctic Research Expedition; BPBM, Bernice P,
Bishop Museum, Honolulu, Hawaii; LDColl, Louis
Deharveng Collection, University Paul Sabatier,
Toulouse, France; NMNZ, National Museum of New
Zealand, Wellington; SAMA, South Australian
Museum, Adelaide; USPHTM, Department of Public
Health and Tropical Medicine, Sydney University.
SYSTEMATICS
KEY TO HEARD ISLAND SPECIES
| Thorax | carrying setae, mot reduced, abdomen V and
VI separate cncien cern cers cutter recresreere 2
Thorax | wilhout setae, reduced, abdomen V and V1
TE): s ee an a, aS 5
2 White, ocelli absent, two spines posterodorsally. on
abdomen VI, grinding mandibular plate and pscudo-
celli present, furca absent oo... csc cre py eee eee 3
Grey, ocelli 3+3, 7-1) spines posterodorsally on
abdomen VI, no grinding mandibular plate or pseudo-
celli, furca present Frieseu tilbrooki Wise
3 Empodial appendage present with seta at least half
as long as claw Tullbergia bisetosa Barner
Empodial appendage absent or rudimentary, lacking
seta
4 Postantennal organ with 50 tubercles ar less, small
species, less than 1.5 mm long .-..-
Tullhergia templei Wise
Postantennal organ with 80 tubercles or more, large
species, More than 2 mm long
Tullbergir antarctica Lubbock
tee eho teens
5 White, acelli ‘and pigmented eye palch absent .... 6
Greyish or black, acelli and pigmented eye patch present
seco ales SlenosMled ee! seven anes Mines Siocon Sasneb eee xb sneeL | sll | oon] jane Be 7
92, REC. 5, AUST. MUS. 19(7): 91-96
6 Dens about 3 times longer than manubrium, with more
than 18 anterior setae anid more than 6 posterior setae,
mucro with three teeth Isotoma sp, indet.
Dens aboul the same length as manubrium, with 9-13
anterior and 4-5 posterior seve, mucro with 3 teerh
i Ae RP wa ta “ryplopveus vaecus Wahlgren
7 Occelli 242 or less, no clayate tenent hairs, mueng with
H deethens reer et ect ag mind amine eels &
Ocelli 646, 2 clavate tenent hairs present on all leys,
mucro with 2 teeth -.. 222.2. 222022 Lee. 9
8 While with sparse black suecijes, ocelli | A, dens nearly
3™ longer than manubrium with about & posterior
and more than 50 anterior Setae, body with long ciliated
macrochaetae distally abundant about 4« Jonger than
ordinary setae On abd V/VI, distal inner anterior
margin of the manubrium with } +1 spines with swollen
hases lsotama (Sorensia) punciata Wahlgren
Grey, ocelli 24+ 2, dens with 5-6 posterior setae and 13-15
anterior setae, dorsal macrochaelae smoath and about
2™ as long as ordinary setae on abdomen V/ V1, distal
inner anterior margin of the manubrium without spines
Lh +) kletm 9 0 Cryptopyeus trieyspis Enderlein
9 Dens shorter Than manubrium with 5 anterior setae and
four posterior setae, mucro with two teeth, macro-
chaetae smooth and only about a third as long again as
ordinary setae on abdomen V/VIE .,..,......0005
eet Bk ERM mene AGE sce Cryptopyeus antarcticus Willem
i 5 (rarely 4) posterior subeoxal setae on fureal segment
and 15-18 anterior sub-coxal setae, 11-18 posterior
manubrial selae ......,.
vita i. Cryptopygus unitarctic: us redgens Enderlein
ii 4 (rarely 5) pasterior subcoxal setae on fureal segment
and 8-14 anterior sub-coxal setae, 9+9 (range 8-10)
posterior manubrial setae
Family NEANURIDAE
|. Friesea tilbrooki Wise 1970
=Friesea viennei Deharveng 1981 syn. nov.
(Figs 1, 2)
Type Locality: South Georgia.
Material Examined: Holotype: South Georgia,
Busen Peninsula, Enten Bay, under rocks on beach,
7,xi,1963, H, D, Clagg, SG 28A, KWSG 2i6c (BPBM),;
Heard 1., Mt Aubert de la Rue, grassy lawn, 26.11.1941.
Dr Vienne, KWHI 39, (Holotype of Frvesew wiennei)
?_LDColl.; Macquarie [., various localities, 32 exs,
Wise (1970a) described F tilbrooki from three
specimens apparently with 10 anal spines collected on
South Georgia. [n 198] Deharveny described F viennei
from Heard J. from a single specimen with seven anal
spines. He mentioned that F) yienner was close ta F
ri/brooki differing only in the number of anal spines
and in having at least four sensory setae (soies 5s’) on
antenna IV compared to F /ilbrooki’s three. Re-
examination of the type of F rilhrooki shows that it
possesses live sensory setae (Fig. 2) in this pasition and
that in all other respects apart from the anal spines it
is identical to Deharveng's description of F) werinet.
The two paratypes of /) ri/breoki have recenily been
examined ancl possess 7 and 8 spines respectively on
abdomen VI (K. Wise pers. comm.). Friesea viennel
June, 1986
was described with a small tooth to the claw and
although F. filhroak lacks this tooth according to Wise,
a small tooth was observed by the present author on
claws | and ff of the F albrooks holotype.
U2 4pacimena
~~ "10 4pecimona trom onw population
Number of spacimens
Number of spinka
FIG. |. Number of spines on abdomen VI found on Friesea Nlhrookt
Wise specimens from a number of localities on Macquarie !.
Morphological variation) Examination of a long
series of specimens of F (//bravki froma single locality
on Macquarie I. showed that the number of anal spines
ranged rom 7-1] in adulls (Fig. 1) and that this
variation had a similar distribution within a single
population to that found for all specimens examined.
A third of the specimens were asymmetric and seven
was the most common number of spines with aynajm,
and p, always spinose. The holotype of F Lilbrooki
from South Georgia had my and a, spinose also (Wise
1970a, Fig, 2B); one of the paratypes was asymmetric.
A single specimen from Macquarie [. had 1] spines
(Fig. 2E). No correlation was found between age, size
or sex of specimens and the number of anal spines.
Bifurcate and double spines inserted in a single posiuon
were seen occasionally.
The sensory setae on antennae JV varied in size in
adults. Occasionally individuals with 4 or 6 well
developed sensory setae were found. The tooth on the
claw was small and hard to distinguish. It appeared to
be absent in some specimens and on some legs,
Other variation found in the Macquarie 1,
specimens was in the tenent hairs from fairly shart to
long and slightly clavate or bent al the tip. The length
of the abdominal macrochaetae varied also and when
longest they were slightly serrated and bent at the tip.
Deharveng mentions that & Wernne/ has a tendency
to plurichaetocity and was asymmetric in ifs setal
arrangement particularly on thorax I and abdomen
IV. He compared its plurichaetocity to Friesea Jara
which was studied by Grow and Christiansen (1974).
COLLEMBOLA OF HEARD ISLAND 93
KP WIOVVII ¥
een WOW Yoo uy
OF
Se)
FIG. 2. Friesea tilbrooki Holotype, A. dorsal view of antenna IV (sensory setae stippled), B. mandible, C. maxilla, D. genital opening
of @ specimen from Macquarie I., E. dorsal view of abdomen V1, (cuticle only partly drawn). F. genital opening of ¢ . Scale line=0.] mm.
54 REC. S. AUST, MUS. 19(7): 91-96
These authors noted that another species, Friesea
grandis, had a “remarkable amount of setal variation”
and that specimens were rarely symmictrical. Series of
specimens of /) ti/brooki from Macquarie |. also
showed much yariation in setal number and
arrangement,
TABLE 2, MORPHOMETRY OF FRIESEA SPECIMENS FROM
HEARD ISLAND AND MACQUARIE ISLAND
| Macquarie 1. Specimens
Number of specimens = 17
Ratio
A dorsal tenent hair Ht tol. claw Wi 1,062 mean @ —.255
95% CL Wl, 8S range .7
y= 1153
B Int, claw Uh: diam. oe A» 3.2 mean.e —9481
95% Cl. 4.6, 2.8
+ 1484
C anal spine 9, Int, claw lll=.811 mean ¢ =.19
95% Cl, 89, 73
3-145
range 21-57
range -36-)-4
¥=.2950
If Comparison of Ratios
A B Cc
Holotype & Wenner (Heard |.)
faccording to Dehurveng (1981)) 1.25 3 TY (0.7-0.9)
Holotype F-ti/brooki (8, Georgia) ,1460 0 2.5 (R66
Macquarie lL, specimens L.06 3.2 O.811
Measurements of the dorsal tenent hair and internal
claw length of leg HI, diameter of ocellus A and anal
spine p, were made and ratios of their lengths
compared with those given by Deharveng (1981) for /;
viernei (Table 2), Deharveng's value of 1.25 for ratio
A (dorsal tenent hair: internal length of claw IIL), is
higher than the mean found here but within the range
of these observations. Ratio B (internal length of claw
Iff: diameter of ocellus A) he gives as 3 and ratio C
(anal spine: internal length of claw II) as 7-9. The
latter figure is certainly a misprint for 0.7-0.9 which
falls well within limits found here. This is good
agreement considering the variation im size of different
spines on a single animal and that Deharveng does not
state which spine he measured,
In view of the measured variation within the
Macquarie 1, populations and known variation in other
+riesea species (Grow and Christiansen 1974) J consider
the specimens from Macquarie |,, Heard |, and South
Georgia to be conspecific.
No other southern hemisphere species of Friesea has
the same combination of eye number (3+ 3), anal
spines (7-11) and well developed furca. Friesea
multispinosa Denis 1947 is nearest, described as having
14-15 spines (13 in Denis’ figure), 343 ocelli and with
a similar furca, Denis (1947) based his description on
a single specimen from Kerguelen Island found in a
similar littoral habitat in which F /i/brooki is found.
In view of the considerable variation in the number of
abdominal spines of F) (//brooki and its similarity to
F, multtspinosa in habitat, ocelli number and furea, it
seems possible that it is a synonym of F mu/tispinosa:
However until the holotype af A niallispinasa is
examined no change of status of these two species is
advisable.
June, 1986
Soine additions to the description of & ¢i/breoki are
given. below, although details piven by Deharveng and
by Wise, apart (rom those already corrected above, still
stand,
Antenna IV. § well developed sensory setae (acies s),
small organite and accessory sensory setae present,
single apical bulb with slight suggestion of division.
Mouthparts: mandible with 7 teeth; maxilla with
dentate lamella carryme about 12 Leeth.
Genital Aperture: 2 with up to 30 short setae and
one pair on anterior lip of genital opening; o with 4+4
broad round sensillae internally and abouc 50 short
setac.
Chaeloiaxy: abdomen VI with tendency towards
plurichactocity and asymmetry, a,m,m,p, (practically)
always Spiftase, m, usually absent,
Comments on the biology of this species and details
of Macquarie J. collections will be given in a later paper
(Greenslade and Wise unpubl. results),
Distribution; Heard |,, Macquarie |, South Georgia,
Family ONYCHIURIDAE
2. Tullbergia antarctica Lubbock 1876
Type Locality: Kerguelen 1,
First recorded for Heard |, by Vanhotfen (1908) and
Enderlein (1909). This record was noted by Salmon
(1949), Brown (1964) and Wise (1970b) but ii has nar
been recollected,
The specimens have not been found. This is a
doubtful identification according to Deharyeng (1981).
Distribution; Antarctica, Kerguelen L.
3. Tullhergia bisetosa Borner 1903
Type Locality: Kerguelen Is,
First recorded for Heard 1. by Womerslcy (1937). This
record was noted by Denis (1947), Salmon (1949), and
Deharveng (1981). The species was recollected by Brown
and by Tremont.
Materio! Examined: Heard 1., Skua Beach, detritus
under rocks, in Azorella and in rack crevices, 18,i),1983,
5. Tremont, 26 exs (SAMA): Atlas Cove, beneath Poa
near elephant seal wallows, 20.11.1983, S. Tremont, 50
exs (SAMA): Atlas Cove, beneath rocks, in Azorella
near elephant seal wallows, 20.11.1983, S. Tremont, 31
exs (SAMA); Heard I, BANZARE coll. 356, det. H.
Womersley, 3 exs (SAMA); 31 slides (about 100 exs)
K. G. Brown, no further data, (USPHTM),
Distribution: Kerguelen 1s, Macquarie J., Heard I,
4. Tullbergia templei Wise 1970
Type Laculity:; Heard I.
Nor recollected but record noted by Deharveng
(1981),
Material Examined: Holotype: Heard |., S. Barrier,
Cairn 5, 457 m, 81.1965, P. Terriple, (ANIC),
COLLEMBOLA OF HEARD ISLAND 95
Distribution: at present only known ftom Heard I.
and Macquarie |, (Greenslade & Wise unpubl. results).
Deharveng (1981) recorded a species from (he Kerguelen
Is which he identified tentatively as Tul/bergiu cf.
templei.
Family ISOTOMIDAL
5. Crypropygus antarcticus antareticus Willem \901
Tpe Lovality: Antarctica,
First recorded for Heard |. as Cryplopyeus
antarcticus by Womersley (1937) and recollected by
Brown, Temple and Tremont,
Material Examined: Ueard 1., Atlas Cove, beneath
Pou near elephant seal wallows, 20.11,1983, 5, Tremont,
2 exs, (SAMA); beneath Azore/la on hillside, 112.1929,
BANZARE call. 349, det. H. Womersley, 3 exs
(SAMA); |2 slides (about 20 specimens) K. G. Brown,
no further data (USPHTM).
Distribution: Widespread in Antarcuca and
Subantarctic islands, ?New Zealand, ?Australia.
6. Cryplopygus antarcticus reagens Enderlein 1909
Type Locality. Crozet 1.
First recorded for Heard |, by Vanhoffen (1908) and
Enderlein (1909) as Crypropygus reagens Enderlein.
Not recollected although the record was noted by
Brown (1964) and Wise (1970b). Deharveng (1981)
considers this to be a probable misidentification for
Ca. anlarcticus Willem, The specimens have not been
found.
Possession 1., Tle des
Disiribuliton: Crozet 4.
Cochans,
7. Cryptopygus caecus Wahlgren 1906
Type Locality: South Georgia.
First collected [rom Heard |, by Tremont.
Material Examined: Heard |., Skua Beach, detritus
under rocks near Azorella and in rock crevices,
18.11.1983, S. Tremont, 38 exs (SAMA); Atlas Cove,
beneath Pou grass near elephant seal wallows,
20.47.1983, S. Tremont, § exs (SAMA).
Distributton: widespread in 8. America, Australia,
New Zealand, Subantarclic islands and Antarctica.
8. Cryptoprgus tricuspis Enderlein 1909
= Purafolsemia quadrioculata Wise 1970
-Cryplopygus qhadriocularus Wise
(Rapoport 1963), nec (Martynova 1967)*
* The two different species described by Rapoport and
Martynova and named /sofomina guadrioculuta were
automatically transferred to the genus Cryplopvets
when Jsofoniela was synonymised with Cryprapyeus
by Massoud and Rapoport (1968).
1974 nec
Type Locality, Kerguelen Is.
First collected fram Heard |, by Tremont.
Material Examined: Heard J,, Atlas Cove, beneath
Pow grass near elephant scal wallows, 20,i1,1983, S.
Tremont, 2 exs (SAMA); Atlas Cove, beneath rocks
near Azorella, 20.11.1983, S. Tremont, 1 ex (SAMA),
Deharveng (1981) notes that C. trieuspis from Kerguelen
I. and Wise's C guddrioculatus from South Georgia
are very close except that C. guadrioculatus has only
9 ‘ventral’ (anterior) setae on the dens and that
according to Wise’s text the species seemed not to have
dilferentiated macrochaetae on the thorax or abdomen
except al the extreme posterior end of the body,
Examination of the holotype of Parafa/somia
quadrioculala showed that both dentes carried 13
anterior and 6 posterior setae. Macrochaetae were
present as follows; on thorax Il 1, thorax 11 1,
abdomen | 3, I] 3, 11 24empty sockets. and IV
3+empty sockets, This pattern conforms with that
given) by Deharveng. The macrochaetae seem to be
easily detached. A paratype carried 12+5 and 13+5
setae on the dentes, Other details of chactotaxy were
as follows:
Numbers of setue
C, tricuspis P. P.
cet quedrioculata quadrivculata
Deharveny Holotype Paratype
Manubrium 13-15 7 13
Furcal subeoxa
Anterior 13-20 i4, 17 17, 21
Posterior 4-6 5,6 6, 6
In all other respeets the holotype and a paratype of
Pardfol-somia quadriveulata agree with the
description given by Deharveng (1981) for C. (ricuspis,
and 1 therefore consider them conspecific.
Distribution; Kerguelen [s., Crozet 1., Marion. L,
South Georgia, Heard |.
9. Isatoma (Sorensia) punctata Wahlgren
Type Locality. Tierra del Fuego.
Material Examined: Heard 1., “from damp
situation”, 1.x11.1929, BANZARE Coll 356, | example,
labelled /sotema octo-oculata Willem, det. H.
Womersley; Heard |., Poly Gully, from dove prion’s
(Pachyptila desolata) nesting material, 811.1965, P.
Temple KWHI21, about 19 exs (BPBM), 2 slides (3 exs)
kK. G. Brown, (labelled Parisoroma octo-oculata), no
further data, (USPHTM),
This species was recorded by Womersley (1937) and
Brown (1964) as /solonia octooculata Willem 1901.
Wise (1970b) recorded Temple's specimens as Sorensia
subflava Salmon. (now Jsotoma (Sorensia) subflava
according to Deharveng (1981)). Material has been
compared with the holotype of /, (S) subflava and
found to difter from it in a number of characters. of
%6 REC. 5, AUST. MUS. 1[9(7): 91-96
specific importance; Le, Sensory setae on abdomen V
and V1, and chaetotaxy of the manubrium. Specimens
are similar to /, (S) punctata in these characters and
are identified as this species, A revision of the genus
is in preparation.
Distribution: Tierra del Fuego, Crozet [., Possession
L, Argentina, Heard [. (all other records of this species
require verification).
10. Jsotoma (Parisotoma) octooculata Willem 190)
Type Locality: Antarctica.
This species was determined by Wornersley (1937)
and Brown (1964), Both Womersley’s and Brown's
specimens are now found to belong to /. (S) punclata
(see above). Wise (1970b) noted the species record but
Deharveng (1981) considers the record of this species
doubtful. There is clearly now no evidence to suggest
it is present on Heard I
Distribution: Antarctica only (all records of this
species from Subantarctic islands, such as Sauth
Georgia (in Wise 1970a), require re-examination),
ll. Jsetoma sp. indet.
Material Examined: Heard [., Poly Gully, Winston
Lagoon, P. Temple, 2.ii.1965, KWHI2, 3 exs (BPBM).
These three specimens were identified by Wise
(1970b) as Sorensia subjlava, They are in poor
condition and it is not possible to determine them
further than to genus. They are unlikely to be /. (P)
oclootulata since the postantennal organ and
chaetotaxy of the manubrium differ from those of that
species. Similarly the structure and chaetotaxy of the
manubrium differ from /. (S) punctata.
DISCUSSION
Of the cight species definitely recorded, six occur
widely on Subantarctic islands and the seventh, T
June, 1986
templet oecurs on Macquarie |, as well as Heard L.
(Greenslade & Wise unpubl. results). One species is of
unknown distribution, No island endemism was detected-
Deharveng (1981) noted some island endemism lor the
Kerguelen Is, but the fauna af these islands is richer
in species, and the islands are larger in area and further
north. However the fauna ol Macquarie 1. with 22
species also shows little island endenvism (Greenslade
and Wise unpubl, results) and it is al about the same
latitude and of the same size as the Kerguelens and
hear the rich source area of the Auckland and
Campbell [slands.
A large amount of morphological variation,
particularly in chactotaxy, was found in these Sub-
antarctic species, Other invertebrates from polar regions
have been found to exhibit a high degree of variability
(Downes. 1965), It has been suggested by Greenslade
(1983) that this is because these faunas are subject
to A selection where relaxation of constraints
on conservative morphology occurs. Whether the
variation documented here js due to a genuine
divergence of lificages on different islands, or
to relaxation of selection pressures or even to the
founder effect can probably only be determined by
breeding, cytological or biochemical techniques.
ACKNOWLEDGMENTS
J would particularly like to thank K, A. J. Wise of
the Auckland Institute and Museum for much help of
various kinds during the preparation of this paper and
for critically reviewing the manuscript and also for the
loan of apecimens. Thanks are also due to Dr R. G,
Ordish, National Museum, Wellington, Professor J,
Brian of the University of Syducy’s Department of
Public Health and Tropical Medicine, L. Deharveng,
University Paul Sabatier, Toulouse, and the Bernice P,
Bishop Museum for the loan of specimens and to the
British Trans-Antaretic Fund for Financial support,
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isolomides d'Amerique du Sud ef de VAntarcliquce. Rial Miner
aust, INO 307-337.
SALMON, J, T. 1949, New sub-anturetic Collembali. Cupe Exped.
Ser Bult. 4: 1-56.
[REMONT,S, 1983, Collembata. In Heard Island Expedition 1984
Seientific Repo. Ed, R, Vining, p, 31,
VANHOPFEN, G. 1908. Tiere und Pflanzen der Hearct-Insel.
Deutsche Sudpolar-Exped 1901-1903, 2: 265-271, Berlin
WISE. KA... 19708, Collembola of South Georgia Pucific liseers
Monoagniph 23. 143-208,
WISE, RK. ALJ, 1970b. Collembola of Heard Islind, Puetfic Insevis
Monngraph 23: 209-215,
WOMERSLEY, H. 1947. Collembola (Springtails), BANZARL Rep,
Ser 8, Vola: 1-7.
MOLLUSC TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM.
3. POLYPLACOPHORA
BY W. ZEIDLER AND K. L. GOWLETT
Summary
The South Australian Museum collection of chiton types is the largest in the Southern hemisphere.
It contains primary type material, and some secondary types, of 123 species, subspecies or varieties.
A further 15 species are represented only by secondary types. The type status of at least 11 of the
primary types 1s considered very doubtful or, in the case of neotypes, invalid. Listed in an appendix
are an additional 21 specise represented by types of dubious status from the Dupuis collection.
Species are listed alphabetically according to the original name of the genus or species.
MOLLUSC TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM. 3. POLYPLACOPHOKA
by
W. ZEIDLER and K, |. GOWLETT
South Australian Museum, North Terrace, Adelaide, South Australia S000
(Manuscript accepted 6 February 1985)
AUSTRACT
ZEIDLER, Wy and GOWLETT, KROL, 1986. Molliise type-specimens
in the South Australian Museum. 3, Polyplueephora Ree S.
Aste Mus, (8) 97-115,
The South Australian Museum collection of chiton
types is the larvest in the Southern hemisphere. It
contains primary type material, and some secondary
iypes, of 123 species, subspecies or varieties. A further
15 species are represented only by secondary types. The
type status of at least 11 of the primary types is
considered very doubtful of, in the case of neotypes,
invalid, Listed in an appendix are an additional 21
species represented by types of dubious status from the
Dupuis collection. Species are listed alphabetically
according to the original name ol the genus or species,
INTRODUCTION
Mast of the chiton types in the South Ausrratian
Museum are due to the early work of W. G. Torr and
b. Ashby al the turn of the century, particularly Ashby
who continued his active interest ia chitons well into
the 1930s,
Ashby donated his collection of chitons, including
most of the types of Torr, to the South Australian
Museum in 1932, but he continued to borrow material
freely for his research, In 1934 Ashby’s home at
Blackwood in (le Adelaide Hills was destroyed by fire
and some type material af chitons and birds wats lost.
As there are no accurate records of the material on loan
to Ashby at the time of the tire, one must assume that
any Ashby or Torr types, listed in the literature as being
in the Ashby or Torr collection, and which cannol now
be located in.an Australian museum were destroyed at
that lime.
Ashby also acquired a small vollection al chittons
from Commandant Paul Dupuis, once conchologist at
the Museum Royal d'Histoire Naturelle de Belgique,
Brussels, including some important historical and
probable type naterial of Blainville, D’Orbigny, Quay
and Gaimard, Rochebrune, Jousseaume and Dupuis. The
Dupuis collection is wot well curated and original labels
are often missing. Types have been indicated by a small
label “TYPE” in red print glued to the label with the
specimen and appears to be a later addition, As the
type stalus of most of these specimens is very doubtful
they have not been included in the main part of the
following list bur are listed separately in on appendix,
Fine, 1986
Other chitan types in the South Australian Museum
came from the May collection, acquired by J. C. Verco
and presented to the musetim in 1929, and as a result
of studies by Cotton, mostly in conjunction with
Ashby, Godfrey and Weeding. Since Cotton and
Godtrey (1940) little of consequence has been published
on Australian chitons and type material of only one
species, Lucilina lilbrooki Milne, 1958 (a paratype), has
been added to the collection.
fn their monograph on South Australian chitons,
Catton and Godfrey (1940) and also Cotton (1964)
selected neotypes for South Australian Bednall and
Matthews species an the basis of Ashby (1918) who
recourits the loss of Bednall and Marthews types at sea,
However they failed to verify which types were lost as
al least some of the original type material (syntypes)
of Bednall is known to exist (Davis ef al 1979).
Similarly, neolypes were also designated for other
species where The type was presumed lost. We believe
that in most cases these ncotypes were designated in
a somewhal arbitrary manner, the possible existence
of type material was not sufficiently investigated and
there were no further statements regarding the
specimens, other than the locality and that they had
been designated as neotypes and given muscum
rezistration numbers. We therefore regard all of the
neatypes selected by Cotton and Godfrey (1940) and
Cotton (1964) and listed here, with the possible
exception of Acanthochites tatei Torr & Ashby, 1898,
as invalid.
We further believe that the South Australian
Museum collection of chiton types is the largest in Lhe
southern hemisphere and one of the more significant
collections in the world. It includes type material for
138 species or subspecies (excluding the Dupuis
collection) and the type status of only 1118s in doubt
or, in the case of neotypes, considered invalid, the
yemainder have been verified by us according ta
available material and information. A further 21
species are represented by types of dubious status in
the Dupuis collection.
In the following list species are arranged
alphabetically in families under the original name at
the time ol description, Changes in familial status have
been cross-referenced. The present status of each
species is, unless indicaied otherwise, according to Kaas
and van Belle (1980), The specimens are all dry except
for some of the paratypes of Isehnochitan johnstant
58 REG, S$. AUST. MUS. 18(Rp OT-ATS
and /. mawson’ and are listed as “entire” when the
articulated valves and girdle are present or as “entire
With animal” when the dried aninial is alsa present.
The following abbreviations are used in the text.
AItM=Auckland Institute and Museum, New Zealand:
AM =The Australian Museum, Sydney;
ANSP= Academy of Natural Sciences of Philadelphia,
US.A; BANZARE=British, Australian and New
Zealand Aniaretic Research Expedition, 1929-31;
MCZ=Museum of Comparative Zoology, Harvard
University, U.S.A MNHN=Museum National
D'histoire Naturelle, Paris; NMN#Z=National Museum
of New Zealand; MV=Museum of Victoria;
NUSW.=New South Wales; N.Z.=New Zealand;
Qld.=Queensland; QM=Quceensland Museum;
S.A.=Soauth Australias SAM=South Australian
Museum; Tas.=Tasmania; TM=Tasmanian Museum
and Art Gallery; W.A.=Western Australia;
WAM=Western Australian Museum,
Family ACANTHOCHITONIDAE
Genus Acanthochites Risso, 1826
Acanthochites cornutus Torr & Ashby, 1898
Trans. R. Soc. S. Aust. 22: 217, pl. 6, fig. 3a-P.
=Craspedochiton cornutus (Torr & Ashby, 1898),
Holotype: D12188, 5 articulated median valyes and
associated girdle, from Marino, near Adelaide, S.A.,
at low tide, collected by E. Ashby, date of collection.
unknown,
Note: Rest of type presumed lost. Type unique.
Acanthochites crocodilus Torr & Ashby, 1898
Trans. R, Soc, S, Aust, 22: 216, pl. 6, fig. 25-f.
=Notoplax crocodilus (Torr & Ashby, 1898)
Holotype: 012137, posterior valve, one median valve
and severs! fragments of valves and girdle, from
Marino, near Adelaide, S.A., collected by W, G, Torr,
date of collection unknown.
Paratype: D12195, entire specimen with animal with
same collection data as holotype, labelled “cotype”.
Acanthochires exilis Torr & Ashby, 1898
Trans, R. Soe, §. Aust. 22: 218, pl. 7, fig. 6a-f.
=Craspedochiton cornutus (Torr & Ashby, 1898)
Holotype: D1225], posterior valve and 3 disarticulated
median valves, from Spencer Gulf or Investigator
Strait, S.A., dredged by J. C. Verco in 10-15 fathoms,
date of collection unknown.
Note: Rest of type presumed lost,
Acanthochites jucundus Rochebrune, 1882
Bull. Soc, Philom, Paris Ser, 7, 6: 194,
=Acanthochitona jucunda (Rochebrune, 1882)
Holotype: 212267, 1 median valve only, from New
Holland, collected by Belligny, date of collection
unknown,
June, 1986.
Note: According to Ashby (1926) this valve came from
the holotype and was given to him by Dr Lamy. Recent
enquiries indicate that the rest of the type cannot be
located in MNHN although Ashby (1922) apparently
saw the type and says “there are a number of specimens
in spirit —-- all much worn!"
Acanthochites kimberi Torr, 1912
Trans. R, Soc. 8. Aust. 4: 67, pl. 6, fig, Sa-f.
=Acanthochitona Kimberi (Torr, 1912).
Neotype: D13758, entire specimen with animal, from
Aldinga, near Adelaide, S.A., collector and date of
collection unknown. Selected by Cotton and Godfrey
(1940; 522, fig, 515).
Syntypes: D12220, 3 entire specimens with animal,
from Corny Point, Yorke Peninsula, S,A., collector and
date of collection unknown, DL2221, 2 entire specimens
with animal and a median, posterior and anterior valve,
from Kangaroo Island, S.A., callector and date of
collection unknown. DI2227, entire specimen with
animal, stuck on card with other specimens from
various localities—siace remaved, from Por
Noarlunga, S,A., collector and date of collection
unknown.
Nate: In the presence of syntypes a neotype should not
have been erected, The type status of 112220 and
particularly D12227 is very doubtful as the locality data
do not match the original description which was
based on 4 specimens from Aldinga and Kangaroo
Island, However, D12220 and D1222) has the label
“Type CO-59” which corresponds to the number of the
Species in the publication, indicating that these
specimens may have been part of the original material,
D12221 also has a label which says “cotypes. type to
be selected", Therefore these are almost certainly
syntypes.. The loose valves with D1222! could have
come from the specimen illustrated by Torr and thus
the intended type, although Torr apparently intended
the specimen collected by Kimber from Aldinga to be
the type. None of the above specimens are as large as
10*4 mm which is a measurement given by Torr and
it must be presumed that this specimen has been lost.
Acanthochites (Loboplax) mariae Webster, 1908
Trans, Proc. N.Z. Insi. 40; 254, pl. 20, figs 1-11.
=Notoplax mariae (Webster, 1908),
Paratype: DIII08, median valve 4, girdle fragment, and
radula, without collection data.
Note: With label ‘valve from Webster's paratype”.
Webster apparently did not designate paratypes
although he mentions at least 7 specimens in addition
to the type. Type in AIM (TM-1).
Acanthochites matthews! Bednall & Pilsbry, 1894
Nautilus 7 (10): 120.
=Notoplax (Bassethullia) matrhewsi (Bednall &
Pilsbry, 1894),
MOLLUSC TYPE SPECIMENS 4. POLYPLACOPHORA Sy
Neotype: D13741, entire specimen with animal, from
Sultana Bay, Yorke Peninsula, S.A., collectorand date
of collection unknown. Selected by Cotton and
Godfrey (1940: 536, fig. 534).
Note: tredale and Hull (L925b) selected a neotype lar
this species presuming the type to be lost (probably
AM, C10411), Cotton and Godfrey (1940) unaware of
this selected yet anather nealype. However, according
to Davis et af. (1979) the holotype is in ANSP (Sheil
Car, No, 64916), We therefore consider both neatypes
invalid.
Acanthochites maughani Yorr & Ashby, 1898
Trans. R. Soc. S. Aust, 22; 218, pl. 7, fig, Sa-f.
=Acanthachitona pilsbryi (Sykes, 1896).
Holotype: 912264, 5 articulated median valves with
remains of girdle and animal, plus one loose median
valve, from Victor Harbor (Port Victor), S.A,, collected
by M. M, Maughan, date of collection unknown.
Note: Rest of type presumed lost,
Acanthechites rubrostratus Torr, 19\2
Trans, R, Soc, S, Aust, 36: 169, pl. 7, fig. Ta-f.
= Notoplax rubrostrata (Torr, 1912).
Holotype: D137L7, entire specimen with animal, from
St Francis Island, Nuyts Archipelago, §.A., callector
and date of collection unknown,
Paratype: D16020, entire specimen with animal, from
Corny Point, Yorke Peninsula, S.A,, collector and date
ol collection unknown,
Note: A label with the holotype says "valves figured
plvii have been lost", The type status of D16020 is in
doubt as Torr (1912) only lists specimens from St
Francis Island and Henley Beach; however a label with
the specimen says “CO-TYPES 41” which corresponds
to the number of the species in the publication,
Acanthoachites rufus Torr, 1912
Trans. R. Soc, 8. Aust, 36: 167, pl. 6, fig. 4a-t.
=Craspedochiton variabilis (Adams & Angas, 1864).
Holotype: DJ2208, entire specimen with part of animal,
from Kanyaroo Island, SA., collector and dale of
collection unknown,
Note: Type unique,
Aeanthochites subviridis Vort, 1911
Trans. R. Soc, 8S, Aust. 35: 104, pl. 25, fig. Sa-f.
=Notoplax subviridis (Torr, 1911).
Holotype: 112872, 4 articulated median valves with
girdle and animal, plus the 4 remaining valves
disarticulated, from Rabbit Island, Albany, W.A,,
presumably collected by Torr, Christmas, 1910-11,
Paratype: D14490, entire specimen with animal, with
same collection data as holotype and labelled
“co-type”.
Acanthochites tatei Torr & Ashby, 1898
Trans. R. Soc. S. Aust. 222 219, pl. 7, fig, 7a-t
=Acanthochitona granastriata (Pilsbry, 1894).
Neotype: D13732, entire specimen with animal, from
Middleton, Encounter Bay, S.A., collector and date of
coflection unknown, Selected by Cottan and Godfrey
(1940: 527, fig. 523).
Note: The neotype was selected from a lot from the
Torr collection (D12260) with the label “our type of
Acan, tatei was very diminutive and was ruined in
dissecting. These are from the same spat, , 2.
Acanthochites tristis Rochebrunc, 1882"
Bull. Soe. Philom. Paris Ser 7, 6; 194,
=Nom. inquir.
Holotype: D12269, | median valve only, from New
Holland collected by Dussumier, date ol collection
unknown.
Note: According to Ashby (1926) this valve came from
the holotype and was given to him by Dr Lamy. Rest
of type in MNHN,
Acanthachites verconis Torr & Ashby, 1898
Trans. R. Soc. 8S. Aust. 22: 217, pl. 6, fig. 4a-f.
= Notoplax verconis (Torr & Ashby, 1898).
Holotype: D12201, 3 articulated median valves with
fragments of girdle and animal, plus fragments of the
posterior valve and median valves, from Gulf St
Vincent, 5.A., dredged by J. C. Verco, date of collection
unknown,
Note: Anterior valye presumed lost. Kaas and van Belle
(1980) regard this species as a synonym af Notoplax
wilsoni (Sykes, 1896) however. one of us (K.L.G.) has
examined this species in detail and regards it a valid
species.
Genus Acanthochiton Gray, 182! em, Iredale, 1915.
Avanthochiton bednalli var johnstani Ashby, 1922
Trans, R, Soe. §. Aust 47: 231.
=Acanthochitona bednalli (Pilsbry, 1894),
Holotype: D12185, entire specimen, from Carnarvon,
Shark Bay, W.A., collected by W, C, Johnston, date
of collection unknown,
Note: Kaas and van Belle (1980) follow Cotton and
Weeding (1939) who elevate this variety ta specific rank,
however, one of us (K.L,.G.) has examined this species
in detail and regards il as a deep-water form af A,
bednalli.
Acanthachitan broakesi Ashby, 1926
Proc. Malac. Soc. Lond, V1(1): 14, pl. L, fig. 3a-c, pl. 2,
fig, 7
=Acanthochitana brookesi Ashby, 1926.
Holotype: D11025, 5 disarticulated median valves and
girdle fragments from Auckland Harbour, N.Z.,
collected by H. Suter, date of collection unknown.
Paratype: D11031, entire specimen without girdle, with
same collection data as holotype.
Note; Remainder of holotype in AIM (TM-2).,
100
Acanthachiton gacliffi Ashby, 1919
Trans. R. Soc. 3, Aust, 43: 398, pl, 42, figs 2-3,
=Aecanthochitona gurliffi Ashby, 1919,
Holotype: D12189, 2 median valves, a valve fragment,
remains of girdle and animal, from Port Lincoln, S.A.,
collected by E, Ashby, Jan, 1917,
Note: Rest of type presumed lost.
Acanthochiton heterochaetus Bergenhayn, 193)
Arkiv. Zool. 234 (13): 20, pl. 1, figs 38-42, pl. 3, figs
67-74,
=Acanthochitona gracilis (Jeffreys, 1859).
Syntype: DIGl85, entire specimen with animal but with
anterior valve missing, from off La Luz, Gran Canaria,
Canary Islands, dredged in 100 m on red algae by N,
Odhner, 1930.
Note: Bergenhayn (1931) records 17 specimens but
apparently did nor designate a holotype, Our specimen
is labelled “cotyp”.
Acgnthochiton kimberi vallingupensis Ashby, 1925
Rep. Aust. Assoc. Adv. Sci. V7; 382.
=Acanthochitona kiniberi (Torr, 1912),
Holotype: D13718, entire specimen with animal, from
Yallingup, W.A,, collected by E. Ashby, 23-x.1920.
Acanthochiton macrocystialis Ashby, 1924
Trans. R. Soc, § Aust. 48: 324, pl. 31, fies 3, 3a.
=Acanthochitona macrocystialis Ashby, 1924.
Holotype: D12248, 5 articulated median valves with
remains of girdle, plus.disarticulated anterior, posterior
and tnedian valve, from Point Puer, near Port Arihur,
Tas., amongst holdfasis of Macrocysris pyrifera,
collected by E. W. Mawle, date of collection unknown,
Paratypes; D10704, one entire specimen with same
collection data as holotype. DI2563, one entire
specimen, one lot of 6 disarticulated median valves and
one lot of 3 disarticulated median valves together with
2 anterior and one posterior valves, all with the same
collection data as holotype.
Note: The label with 912563 indicates that the paratype
measuring 20%9 mm was sent co Thackway,
Acanthechiton manxillaris Ashby, 1919
Trans. R. Sac. 8S. Aust. 43: 397, pl. 41, figs 5 & 6, pl.
42. fig. |.
=Aeonthachitona kimberi (Vorr, 1912).
Holotype: 012253, 4 median valves and anterior valve
articulated with girdle, plus ane loose median valve,
trom Marino, near Adelaide, S.A., on rocks at low tide,
collected by E. Ashby, date of collection unknown,
Note: Rest of type presumed lost. Type unique.
Acanthochiten pilsbryi maughaneanus Ashby, 1919
Traps, R, Soc, 8, Aust, 43; 395, pl. 41, fig. 4.
=Acanthochiiona pilshrvi (Sykes, 1896).
Holotype: D15536, 3 disarticulated median valves,
anterior valve and valve fragments and remains of
REC 5, ALIST. MUS. 198): 97-118
dune, [986
girdle, from Middle Harbour, Sydney, N.S.W., collected
by BE. Ashby, 625.1903,
Note: Rest of type presumed lost.
Acanthochiton (Notoplax) porcing Ashby, 1919
Trans. R. Soc, S. Aust. 43: 395, pl. 41, figs 7-10.
=Noatoplax (Bassethyllia) matihews) (Bednall &
Pilsbry, 1894).
Holotype: 112250, specimen in two pieces wilh
articulated median valves and remains of girdle and
animal, plus disarticulated anterior, posterior and one
median valve, from Gulf St Vineent, $,A., dredged by
J.C. Vereo, date of collection unknown.
Note: Type unique.
Acunthachiton retrojectus var. pustulosus Ashby, 1922
Trans. R, Sac. §. Aust. 46: 15,
=Acanthochitona retrojecta (Pilsbry, 1894),
Holotype: D12205, entire specimen with animal, fram
Quarantine Station, Sydney Harbour, N.SW.,, collected
hy E. Ashby, Nay, 1918.
Acanthechiton shirleyi Ashby, 1922
Trans, R. Soc. S, Aust. 46; 13, pl. 3, fig, Ja-c
= Acanthochitona shirlevi Ashby, 1922,
Paratype; D10715, disarticulated valves stuck on card,
from Northwest Reet, Capricorn Group, Old., collector
and date of collection unknown,
Note: Type in QM (MO 4043),
Acanthochiten ¢{Natoplax) spongialis Ashby, 1923
J. Roy, Soc. Wo Austr. 10(4); 13, pl. 1, fig. 1.
=Neloplax spenzialis (Ashby, 1923).
Holotype: 013733, entire specimen from D’Enrre-
casteaux Channel, southern Tas., dredged in 9-10
fathoms by W. L. May, date of collection unknown.
Note: Cotton (1964) indicated that this specimen may
not be the holotype presumably because it does nat
match the measurements given (38 X16 min for D13733;
35% 18 mm given lor the type) and does not appear Lo
match the rather poar photo of the type. However,
SAM seems to have the other 3 specimens mentioned
by Ashby (D12803 and 014471) and as none approach
the dimensions of [he type, and in the absence of
evidence Lo (he contrary, we must assume (hat D13733
is the holotype.
Acanthochiton thackwayi Ashby, 1924
Trans, &. Soc. S. Aust. 48: 318, pl. 31, figs | & 2,
=Acanthochiiona thackwayi Ashby, 1924.
Holotype: D10716, entire specimen with animal, from
Fly Point, Port Stephens, N-SW.,, collected by EB, Ashhy,
October, 1923,
Paratype: DI6548, entire specimen with animal, fram
satne locality as holotype, collected by A. E, J
Thackway, dale of collection unknown.
Nate: Smith and Robertson (1970) list 2 paratypes in
the MV (F16376) with the locality as Shell Harbour.
MOLLUSC TYPE SPECIMENS 3. POLYPLACOPHORA Ol
We regard (hese specimens as very doubtiul paratypes
as Ashby only mentioned one other specimen which
is the paratype in SAM,
Acanthochiton turton Ashby, 1928
Proc, Malac, Soc. Lond. 18(2); 79, pl. 6, figs 1-4.
= Acunthoehitona garnou (Blainville, 1825).
Holotype: 210985, 5 disarticulated median valves and
girdle fragment, from Port Alfred, South Africa,
collected by W. H, Turton, date of collection unknown,
Paratypes: DIUY84, one entire speciinen and two broken
median valves, with same collection data as holotype,
Note: The type status of some ol the valves labelled
“holotype” is in doubt. There are two lots, One lot of
4 yalves does not appear to belong to the type. The
valves are from two different specimens and measure
§.3%3.2, 5.03.2, 4.1%2.5 and 2.118 mm respect-
ively, The first two valves are too large for the type and
the last one too small (Ashby gives a width of 4.5 mm
for the type) and the third valve, and also (he last one,
are green in colour and not pink as stated for the type.
The other lat, consisting of only one valve, with the
label “valves 6 & 7 not photo'd” would appear to
belong to Lhe type.
Giles and Gosliner (1983) record another valve from
the holotype in the South African Museum (432649),
Acanthachiton zealandicus doubtlessensis Ashby, 1926
Proc, Matac. Sac, Lond. 11(1); 12, pl. 1, fig. 2a-c; pl.
2, lig. 6,
= Acanthochitona sealandica (Quoy & Gaimard, 1835).
Holotype: DIN09, disarticulated specimen, from
Doubtless Bay, N.Z., collected by A, E. Brookes, date
of collection unknewn,
Paratypes: DI1118, 3 entire specimens, with same
collection data as holotype.
Nate; Paratypes correspond to Ashby’s paratypes
No. 1, 3 and 4.
Genus Craspedachiton Shuttleworth, 1853
Craspedochiton jaubertensis Ashby, 1924
Trans. R, Sac, S. Aust. 48: 326, pl. 31, lig. Sac.
Holotype: D11234, 2 articulated median valves with
fragments of one valve and remains of girdle and
animal, plus disarticulated anterior, posterior and
remaining 3 median valves, dredged in 70 [t, 42 miles
W,.SW. of Cape Jaubert, northern W.A., by Dr E,
Mjoberg, 26.1911. (Swedish Scientific Expeditions
1910-19]3).
Genus Lophoplax Ashby, 1926
Lophoplax finlayl Ashby, 1926
Proce. Malac. Soe. Land, V1(V): 30, pl. 3, fig. 4) pl 4,
fies 1-4,
= Craspedachiron rubiginosus (Hulton, 1872).
Holotype: 914318, 5 disarticulaled median valves,
dredged in 60 fathoms off Otago Heads, N.Z., by H. J.
Finlay, date of collecuon unknown,
Note: Rest of type presumed lost. Type unique.
Genus Notoplay Adams, 1861
Natoplax (Ainblyplax) braokes/ Ashby, 1929
Trans. Proc. N.Z. Inst. 60: 370, pl, 32, figs 1-4,
= Notoplax brookesi Ashby, 1929.
Holotype: 011030, 2 disarticulated median valves, from
Tauranga Harbour, N.Z., dredged in 3 fathoms,
collector and date of collection unknown,
Paratype: D11113, entire specimen with same collection
data as holotype.
Note: Rest al lype presumicd lost.
Notoplax (Amblyplax) foveauxensis Ashby, 1926
Prac, Malac. Soc. Lond. 1701): 20 pl. |, fig, Sa-c
~ Craspedochiton rubizinosus (Wutton, 1872).
Holotype: D16021, | median valve, from Foveaux
Strait, N.Z., dredged in 15 fathoms on oyster shell,
collected by W, R, B. Oliver, date of collection
unknown.
Paratype: D16022, 4 articulated median valves with
remains of girdle and animal, plus disarticulated
anterior, posterior and 2 median valves, with same
collection data as holotype.
Nore: Rest of type in NMNZ (M1584),
Notoplax (Amblyplax) mariae haurakiensis Ashby,
1926
Proc. Malac, Soc. Lond, 17(1): 26, pl. 2, fig. Aa-c.
=Notoplax mariae (Webster, 1908).
Holotype: 11043, 4 articulaled median valves aiid
remains of girdle, plus one disarticulated median valve,
from Hauraki Gulf, N.Z., dredged in 20 fathoms, off
Atrina shell, collected by A, E. Brookes, date of
callection unknown.
Paratype: D16550, remains of girdle, stuck on card,
wilh same collection data as holotype.
Note: Rest of type in ALM (TM 535),
Notoplax (Aniblyplax) oliveri Ashby, 1926
Proc. Mualae. Soc. Lond. W711): 18, pl. 1, fig. 4a-e.
=Craspedochitan rubizinosus (Hutton, 1872).
Holotype; D11044, 1 median valve, from between
Kawau and Tiritirr islands, Hauraki Gull, near
Auckland N.Z., dredged in 20 fathoms in dead fring
shell, collecied by A. E. Brookes, date of collection
unknown,
Paratype: DI6551, 5 articulated median valves with
remains of girdle, plus disarticulated anterior, posterior
and median valve, with same collection data .as
holotype.
Note: Rest of type in NMNZ (M1585). Paratype has
label “Ashby's No, 2"
Notoplax rotinestensis Ashby, 1929
J. Roy. Soc. Wo Ausr. 15: 47, figs 10-13.
12
Holotype: D12565, 3 disarticulated median valves (2,
3, & 7), fragments of girdle and radula mounted on
slide, [rom Bathurst Point, Rottnest Island, W.A.,
collected by L. Glauert, date of collection unknown.
Nate: Rest al type in WAM (12885).
Family CALLISTOPLACIDAE
Genus Callistochiton Dall, 1882
Callistochiton antiquus miayi Ashby, 1919
Trans. R. Soc. 8, Aust, 43: 401, pl. 42, figs 8 & 9.
=Callistochiton antiquus meridionalis Ashby, 1919.
Holotype: D12550, | median valve from Penguin rocks,
north-western Tas., collected by E, Ashby, 11.x.1916.
Note: Rest of type presumed lost.
Callistochtton antiquus meridionalis Ashby, 1919
Trans. R. Soc. §. Aust, 43; 400, pl. 42. fig. 7.
Holotype: D13716, disarticulated valves and piece of
girdle and radula, from Marino, near Adelaide, 5.A.,
collected by E. Ashby, date of collection unknown,
Ischnochiton (Lepidozona) asthenes Berry, 1919
=Callistochiton asthenes (Berry, 1919),
see ISCHNOCHITONIDAE
Callistochiten auzustensis Ashby & Cotton, 1937
Trans. R. Soe. S. Aust. 612 145, pl. 8, figs 2-4,
Holotype: DI2952, anterior valve, 6 disarticulated
median valves with one broken and remains of animal,
trom Port George IV, Augustus Island, northern 'W.A.,
collected by B. Bardwell, October, 1933.
Note: According to Ashby and Cotton (1937) there were
only six detached valves so presumably the posterior
valve was lost betore the description of the species. Type
unique.
Callistochitan broomensis Ashby & Cotton, 1934
J Roy, Soc. W. Aust. 20: 213, pl. 13, fig. 3,
Holotype: D10723, entire specimen with animal, from
Gantheaume Point, Broome, W.A., collected by MCZ
expedition to Australia, September, 1929.
Note: Type unique.
Callistochiion clenchi Ashby & Cotton, 1934
J. Roy. Soc. Wo Aust. 20; 214, pl. 13, fig. 1.
Holotype: D10724, entire specimen with animal, from
Gantheaume Point, Broome, W.A,, collected by MCZ
expedition to Australia, September, 1929.
Note: Type unique,
Callistochiton mawlei Iredale & May, 1916
Proc, Malac, Soc. Lond. 12(2/3): 113, pl. 4, fig. §.
Paratypes: D12019, one disarticulated specimen with
dried radula, locality not indicated but probably Tas.,
collector and date of collection unknown; D12026,
disarticulated valves of one specimen with same
collection data as 12019,
REC. S AUST. MUS. 19(8): 97-115
June, 1986
Note: An old label with the specimens says. that they
were disarticulated by Iredale and used in the original
type description but there is no indication that they
were designated paratypes. Types in TM (E 201/7542).
Cullistochiton oveidus Ashby & Cotton, 1934
J. Roy. Soc, Wo Aust. 20: 215, pl. 13, tig. 4.
Holotype: DLO725, entire specimen with animal, {rom
Gantheaume Point, Broome, W.A,, collected by MCZ
expedition to Australia, September, 1929.
Note: Type unique.
Genus Lophockhiton Ashby, 1923
Lophoehiton johnstoni Ashby, 1923
Trans. R. Soc, 8. Aust, 47; 234, pl. 16, fig. 7a-cy pl. 17,
fig. la-d,
=Callistovhiion coceus (Menke, 1844).
Holotype: D12152, disarticulated specimen with one
median valve fragmented, dried radula and piece of
girdle, from Carnarvon, in extreme north of Shark Bay,
W.A., collected by W, C, Johnston, date of collection
unknawn.
Note: Type unique.
Family CALLOCHITONIDAE
Genus Acuteplax Cotton & Weeding, 1939
Acutoplax collont Weeding, 1940
Trans. R. Soe. S. Aust. 64(1): 48, pl, 4, fiz, 1, la,
=Cuallachiton cottoni (Weeding, 1940).
Holotype: 113766, entire specirnen with animal,
dredged in shallow water, Spencer Gulf, S.A., ‘by
Fisheries launch “Whyalla”, K, Sheard, March, 1938.
Note: P, Kaas, Rijksmuseum, Leiden, has examined the
type and considers it a synonym of Callochiton mayi
Torr; 1912.
Genus Callochiton Gray, 1847
Callochiton elongatus May, 1919
Pap. Proc, R. Soc. Tas. 1919: $5, pl. 14, fig, la-b.
Paratypes: D12696, 7 entire specimens (3 with animal),
from Norfolk Bay, Tas., collected by E. Mawle, 1915,
D10681, 6 entire specimens stuck on card, 2 from Port
Arthur, Tas., collector and date of collection unknown,
and 4 from Woodbridge, Tas., collected by E. Ashby,
26.111,1920.
Note: D12696 are from the May collection (Na, 230)
and are labelled '‘cotypes’. DIO681 are from the Ashby
collection and are also labelled “cotype”’, probably
referring to the two specimens from Port Arthur,
however May notes only 7-8 specimens so the type
status of hese specimens 1s in doubt. ‘The 4 specimens
from Woodbridge cannot be types. Fype in TM (E177/
7518),
MOLLUSC TYPE SPECIMENS 3, POLYPLACOPHORA 103
Callochiton klemi Ashby, 1926
Trans. R, Soc, 8. Ausr. 50: 243, tig. 4.
Holatype: DI1703, | median valve, from amongst shell
grit, Daly Head, Yorke Peninsula, S.A., collected by
W. Klem, date of collection unknown,
Note; Type description was based on this single valve.
Callochiton platessa var. Jossa Ashby, 1922
Trans, R. Soc, 8, Aust, 46: 19, pl. 3, fig. 4.
=Callochiton crocinus (Reeve, 1847)
Syntypes: D11699, 3 entire specimens (2 with animal
remiains), from Watson's Bay, Port Jackson, N.S.W.,
collector and date of collection unknown.
Nate- The above three specimens were sent to Ashby
by May and Ashby refers to them in his description
and jllustrated the larger specimen. The other
specimens mentioned by Ashby do not appear to be
in SAM although one specimen, D11694, with he label
“variety with pits on valve 7" could be the SAM
specimen mentioned by Ashby. Another specimen in
SAM, D1067), is labelled “Holotype” but it does not
matvh Ashby’s description as the pits on valve 7 are
pworly defined and only 6-7 in number. Lt is stuck on
a card with 3 other specimens and is the only one
labelled “var fossa’ and we suspect that this
information was added alter Lhe publication of the
original description.
Callochiton rufus Ashby, 00
Trans. R. Soe, §. Aust, 24: 87, pl. 1, tig, 2a-g.
Holotype: D11700, specimen with first median valve
removed and anterior valve missing, presumed lost,
with animal, from Gull St Vincent, $.A., dredged by
J.C. Verco, date of collection unknown.
Nate: Type unique
Family CHITONTIDAE
Genus Acanthoplenra Guilding, 1829
Acunthopleura gemmata var. queenslandica Ashby,
192]
J. Roy. Soc, Wo Aust, & 30.
=Acanthopleura gemmata (Blainville, 1825).
Holotype; 912459, disarticulated specimen with girdle
fragments, from Dunk Island, Qld., collected by J,
Shirley, date of collection unknown.
Note: There is also a radula, mounted on a slide,
labelled “Acanthupleure gemmata Bl. var 1, Dunk Id.
Q", which could have come from the above specimen.
This same specimen was selected by Ashby (1928) as
the neatype of Chiton gemmatus Blainville, 1825 (also
listed here).
Acanthopleurd gemmatus maudensis Ashby, 1928
Trans. R. Soc. 8: Aust 52; 172, pl. 12, figs 8 & 9.
=Avcanthopleura gemmata (Blainyille, 1825),
Holotype: 10775, disarticulated anterior, posterior
and 4 median valves, from Maud's Landing, north of
Shark Bay, northern W.A., collected by T. Curton, date
of collection unknown,
Note: Median valves 2 and 6 presumed lost.
Genus Chiton Linnaeus, 1738
Chiton aureomaculata Bednall & Matthews, 1906
Proc. Malac. Soc. Lun. 7(2); 91, pl. 9, figs 3 & 3a-h
=Chiton (Rhyssoplax) tricostalis Pilsbry, 1894.
Neotype: D14135, entire specimen with animal, from
Cape Banks, S,A., collector and date of collection
unknown, Selected by Cotton (1964: 94).
Note: From Torr collection. Listed by Cotton (1964)
as neolype without discussion and considered, by us,
as invalid.
Chiton bednalli Pilsbry, 1895
Nautilus 9: 90.
=Chiton (Rhyssoplax) bednalli Pilsbry, 1895.
Holotype: 113742, disarticulated specimen with
remains of animal, from Sultana Bay, Yorke Peninsula,
S.A., collected by W. T. Bednall, date of collection
unknown,
Note: From Bednall collection.
Chifon exaptandus Bednall, 1897
Prov. Malac. Soc. Lond, 24); 152.
=Chiton (Rhyssoplax) exoptandus Bednall, 1897.
Neotype: D13744, entire specimen, from Sultana Bay,
Yorke Peninsula, 5.A,, collector and date of collection
unknown. Selected by Cotton and Godfrey (1940: 559),
Note: Listed by Cotton and God!rey (1940) as neotype
without discussion and considered, by us, as invalid.
Chiton gemmuatus Blainville, 1825
Dict. Sci. Nat, 36: 544,
=Acanthopleura gemmata (Blainville, 1825)..
Neatype: D12459, the same specimen as the holotype
of Acanthopleura gemmuata var queens/andica Ashby,
192], also listed here, Selected and figured by Ashby
(1928; 172, fig. @ & 7).
Chiton (Rhyssoplax) kimberi Ashby, 1928
Trans. R. Soe. 8. Aust, 52; 170, pl. 12, figs 10-12.
Holotype: D12393, disarticulated anterior, posterior
and 3 median valves plus fragments of girdle and radula
mounted on slide, fram Capricorn Group, Qld.,,
collected by W. J. Kimber, date of collection unknown,
Note: Rest of [ype presumed lost. Type unique.
Chiton marmoreus var. coeruleus Winkley, 1894
Nautilus (4): 78.
=Tonicella marmorea (Farbicus, 1780) (ISCHNO-
CHITONIDAE)
Paratype: D10467, enlire specimen, from Eastport,
Maine, U.S.A,, collected by H. W, Winkley, date of
collection unknown,
104 REC. S. AUST. MUS. 19(8): 97-115
Note: Specimen with original MCZ label and with the
number “1707" written on the shell. From Ashby
collection. Type in MCZ (MCZ 32856).
Chiton oruktus Maughan, 1900
Trans. R. Soc. S. Aust. 24: 89, pl. 1, fig. 3a-g.
=Chiton (Rhyssoplax) oruktus Maughan, 1900.
Holotype: D1489, disarticulated specimen, plus girdle
fragment, from MacDonnell Bay, S.A., collected by
W. G. Torr, date of collection unknown.
Paratype: D12383, entire specimen, from Robe, S.A.,
collector and date of collection unknown.
Note: The type status of D12383 is doubtful as the only
locality listed by Maughan (1900) was MacDonnell Bay.
Chiton scaber Blainville, 1825
Dict. Sci. Nat, 36: 553.
=Nom. inquir,
Holotype: D12271, 1 median valye only, from New
Holland, collector and date of collection not indicated,
Note: From Ashby collection with label “valve of type”
presumably obtained by Ashby during his visit to
Europe in 1922. However, the species is not mentioned
by Ashby (1922).
Chiton tulipa alfredensis Ashby, 1928
Proc. Malac. Soc. 18(2); 87, pl. 8, figs 19-21.
=Chiton (Rhyssoplax) tulipa Quoy & Gaimard, 1835.
Holotype: D10983, 3 disarticulated median valves,
girdle fragments and loose girdle scales, from Port
FIG, 1. Chien verconts Torr-& Ashby, (898. AL Lectorype, 17> UL mim (curled)
June, 1986
Alfred, South Africa, collected by W. H. Turton, date
of collection unknown.
Paratypes: DI1157, 3 entire specimens with same
collection data as holotype.
Note: Holotype with label “these valves not photo’d”,
Rest of type presumed lost.
Chiton verconis Torr & Ashby, [898
Trans. R. Soc. §. Aust, 22: 215, pl. 6, fig. la-f,
= Chiton (Mucrosquama) verconis Torr & Ashby, 1898.
Lectotype: D16546, entire specimen with animal, from
Gulf St Vincent, S.A,, dredged by W. J. Kimber, date
of collection unknown. (Lectotype chosen here).
Paralectotype: D16547, entire specimen with animal,
with same collection data as lectotype.
Note: The SAM register and Cotton (1964) list D12380
as the holotype but this lot consists of one complete
specimen, one median valve and the dried remains of
a specimen from which the valves have been removed,
none of which corresponds to the original description
and is not even C. verconis but Callochiton cottoni
(Weeding, 1940). In view of this confusion we have
selected the above lectotype and paralectotype from a
lot which we believe constituted syntype material. The
lectotype (Fig. 1A) matches the original description and
measures 1711 mm (curled) and 31 mm along the
curled dorsal surface, which approximates to the
measurement of 24x12 mm given by Torr and Ashby,
and may have been the specimen for which this
measurement was given. The paralectotype (Fig. 1B)
B. Paralectotype, 1S *6.6 mim,
MOLLUSC TYPE SPECIMENS 3, POLYPLACOPHORA
which was collected with the lectotype is less curled and
measures 11.5%6.6 mm.
Another 3 specimens (D10739, 012379 and D12674) all
dredged in Gulf St Vingent by Vereo may also have
belonged to the original type series bul they are not
given ype status by us.
The valves figured by Torr and Ashby are presumed
lost
Genus Liolophura Pilsbry, 1893
Plaxiphora pustulosa Torr, 1911
= Liolophura (Clavarizona) hirtosa (Blainville, 1825).
see MOPALIIDAE,
Genus Lucilina Dall, 1882
Lucilma rainfordiana Hull, 1924
Proc. R. Soc. Old. 36: 118, pl. 21, fig. 3.
=Tonicia (Lucilina) fortilirata (Reeve, 1847).
Paratype: D14496, entire specimen with animal, from
North Head, Port Denison, Qld, collected by E, H.
Rainford and A. F. B. Hull, 18.ix.1923.
Note: From Hull collection, Type in QM (MO 1139).
Lucilina lilbrooki Milne, 1958
Proc, R, Zool. Soe. N.S.Meo 1956-57: 152, fig. 1-4.
=Tonicia (Lucilina) tilbrooki (Milne, 1958),
Paratype: D14591, entire specimen with animal remains,
fram Heron Island, Qld., collected by K. L. Milne,
1952.
Note; Type in MV (F18471).
Genus Mucrosquama tredale & Hull, 1926
Mucrosquavia nielsent Cotton & Weeding, 1939
Trans. R. Soc. S. Aust. 63(2): 190, pl 7, fig. 8.
=Chiton (Mucrasquama) carnosus Angas, 1867,
Holatype: D13720, entire specimen, from Hardwicke
Bay, Yorke Peninsula, S.A., dredged by J. C, Verco, date
of collection unknown.
Note: Type unique.
Mucrosquama sheardi Cotton & Weeding, 1939
Trans. BR. Sac, S. Aust, 63(2); 190, pl. 7, fig. 3.
=Chiton (Mucrosquama) verconis Torr & Ashby, 1898.
Holotype: 013721, entire specimen with animal, from
Spencer Gulf, S.A., dredged by Fisheries Launch
“Whyalla, K, Sheard, March, 1938.
Genus Onithochiton Gray, 1847
Onithochiton ashbyi Bednall & Matthews, 1906
Proc. Malac, Soc. Lon. 7(2): 92, pl. 9, figs 2; 2a-e.,
Neotype: D13745, entire specimen with animal, from
south of Port Willunga jetty, near Adelaide, S.A.,
collected by E. Ashby, 2.iv,1926. Selected by Cotton and
Godfrey (1940: 566).
105
Note: From Ashby collection. Listed by Cotton and
Godfrey (1940) as heotype without discussion and
considered, by us, as invalid.
Onithochiton quercinus occidentalis Ashby, 1929
Trans, R. Soc, 8, Aust, 53: 65,
Holotype: D12527, entire specimen, from Dongarra,
W.A., from exposed outer reef, collected by E, Ashby,
10,xi.1920.
Genus Rhyssoplax Thiele, 1893
Rhyssoplax jacksonensis Ashby, 1921
Proc. R. Soc. Vie. (NS) 33: 153, pl. 8, fig. la-b.
=Chiton (Mucrosquama) carnosus Angas, 1867,
Holotype: 10720, entire specimen, from Quarantine
Station, Port Jackson, N.SW,, collected by E. Ashby,
23.xi.1918.
Rhiyssoplay surrecta Hull, 1922
Aust, Zool, 2(3); 84, pl, 24B, figs 1-4,
=Chiton (Rhyssoplax) bednalli Pilsbry, 1895.
Holotype: D12671, entire specimen, [from Port
Willunga, near Adelaide, S.A., collected by W. J.
Kimber, date of collection unknown.
Note: Type unique.
Genus Sypharochiton Thicle, 1893
Sypharochiton pellisserpentis septentriones Ashby,
1924
Trans. R, Sac. 8. Aust. 48; 321.
=Chiton pellisserpentis Quoy & Gaimard, 1835.
Holotype; D17238, entire specimen with animal, from
Nelson Bay, Port Stephens, N.SW., collected by H. W.
Thackway, October, 1923.
Genus Tonicia Gray, 1847
Tonicia hullianus Torr, 1911
Trans. R. Soc. S. Aust, 35: 104, pl. 25, figs 4a-l.
=Tonicia (Lucilind) hilliana Torr, 1911,
Holotype: D12873, entire specimen trom Ellenbrook,
south of Cape Naturaliste, W.A,, collected by W, G,
Torr, Christmas, 1910-11.
Note: Type unique.
Family CHORIPLACIDAE
Genus Choriplax Pilsbry, 1894
Choriplax grayi pattisoni Ashby, 1921
Trans. R. Soe, 8 Aust. 45: 137, pl. 9, fig. la-e.
Holotype: D15019, entire specimen, found with giant
kelp (Laminaria) washed ashore near Cape Banks
Lighthouse, S.A., collected by G. Pattison, 1921.
Note: One of us (K.L,G.) has examined this species in
detail and regards it a synonym oat Cheriplan gravi
(Adams & Angas, 1864). Type unique.
106 REC. S. AUST. MUS. 19(8): 97-115
Family CRYPTOPLACIDAE
Genus Cryptoplax Blainville, 1825
Cryptoplax iredalei Ashby, 1923
Trans, R, Soc, 8S, Aust, 47; 238, pl. 19, fig. 4,
Holotype: D12306, entire specimen, from Port Lincoln,
collected by E. Ashby, January, 1917,
Cryploplax striatus var. weslernensis Ashby, 1923
Trans, R, Soc, S, Aust, 47; 238,
=Cryptoplax striata westernensis Ashby, 1923.
Holotype; D1I0717, entire specimen with amimal
remains, from Rottnest Island, W.A., collected by E.
Ashby, 2,x,1920,
Note: Type unique.
Family ISCHNOCHITONIDAE
Genus Anisoradsia Iredale & May, 1916
Anisoradsia mawlei saundersi Ashby, 1918
Trans. R. Sac, 8. Aust. 42: 82.
=Ischnochiton (Heterazona) cariosus Pilsbry, 1892,
Holotype: D11961, specimen with anterior, posterior
and | median valve disarticulated, from Port Lincoln,
S.A., collected by E, Ashby, January, 1917,
Genus Dinoplax Dall, 1882
Dinoplax gigas var. validfossus Ashby, 1934
Ann, Durban Mus. 3(4): 79, pl. 9, fig. 3.
= Dinoplax validfassus Ashby, 1934.
Holotype: D10971, entire specimen, from Durban,
Natal, South Africa, collector and date of collection
unknown,
Note: From J. D. Casey collection.
Genus /sehnochiton Gray, 1847
Ischnochiton (Lepidezona) asthenes Berry, 1919
Lorquinia 2(6); 47,
=Callistochiton asthenes (Berry, 1919) (CALLISTO-
PLACIDAB),
Paratype: DLO404, entire specimen, from While's Point,
Los Angeles County, California, U.S.A., collected by
A.G, Smith, 14-18.vii.1916.
Note: Label with specimen has the number “SSB 1196”.
Type in Berry Collection,
fschnochiton atkinsoni Iredale & May, 1916
Proc, Malac, Soc, Lon, 12(2/3): 110, pl. 4, fig. 3.
Paratypes: D15678, 8 entire specimens with animals,
from Stanley to Devonport, Tas., collected by W. G.
Torr, date of collection unknown.
Note; From May collection (No. 233), Type in TM
(E179/7520).
Ischnochiton atkinsoni bruntensis Ashby, 1927
Pap, Proc, R, Soc, Tus, 1926: 111.
Sune, 198:
Holotype: D11966, entire specimen, from Luyawanna,
South Bruny Island, D’Entrecasteaux Channel, Tas.,
collector and date of collection unknown.
Ischnachiton atkinseni lincolnensis Ashby, 1920
Trans. R, Soc. &: Aust. 42275, pl. 12, fig. Sa-b.
=Jschnochiton variegatus (Adams & Angas, 1864),
Holotype: D11763, entire specimen, from Port Lincoln,
S.A,, collected by EB. Ashby, January, 1917.
Ischnochiton auratus Ashby, 1920
Trans. R, Soc. 8S. Aust, 44: 277, pl 12, fig, 6a-b,
=Ischnochilon variegatus (Adams & Angas, 1864).
Holotype; D118450, entire spegimen with animal, from
Marino, near Adelaide, 5.A., collected by E. Ashby,
date of collection unknown,
Paratype: D10738, entire specimen with animal, with
same collection data as holotype,
Ischnochiton bakeri Torr, 1912
Trans. R, Soc, §. Aust. 36; 169, pl, 7, fig. Sa-v, f.
=Nom, inquir,
Holotype: 012145, entire specimen with animal but
posterior valve missing, from Henley Beach, Adelaide,
S.A,, collected by W, H, Baker, date of collecnon
unknown.
Note; The specimen is very eroded and the posterior
valve was missing at the time of collection, Type unique.
Ischnachiton (Isochitan) burdwelli Ashby & Cotton,
1934
JR, Soe. Wo Aust, 20: 217, pl. 13, lig. 4.
=Isehnochiton bardwelli Ashby & Cotton, 1934,
Holotype: D11978, disarticulated specimen and virdle
scales mounted on slide, from off Broome, W-.A,,
dredged in 7 fathoms by B. Bardwell, date of collection
unknown,
Paralypes: DI1979, 2 entire specimens wilh same
collection data as holotype,
Note: Smith and Robertson (1970) list the holotype as
being in the MV (F17977), However, our specimens
match the original deseription and are elearly marked
“holotype” and “paratypes”, Also we have the original
label “‘Solvaga recens Thiele” as mentioned by Ashby
and Cotton, so that the type status of the MV specimen
must be very doubtful,
Ischnochiton bednalli Torr, 1912
Trans. R. S. Aust. 36: 166, pl. 5, fig. 3a-f.
=Subterenochiton bednalli (Torr, 1912)
(SUBTERENOCHITONIDAB)
Holotype: D11792, entire specimen and radula mounted
on slide, from Sct Francis Island, Nuyts Archipelago,
S.A., collector and date of collection unknown,
Ischnochiton (Haploplax) broomensis Ashby &
Cotton, 1934
JR. Soc, Wo Aust, 20: 216, pl. 13, fig. 2.
MOLLUSC TYPE SPECIMENS 3. POLYPLACOPHORA
Holotype: D10729, disarticulated specimen with radula
and girdle rermains mounted on slides, from
Gantheaume Point, Broome, W\A,, collected by MCZ
expedition to Australia, September, 1929.
Paratype: 1915537, entire specimen with animal, with
same collection data as holotype.
Isehnochiton (Hetervzona) cariasus var occidentalis
Ashby, 1921
Trans. R. Soc. S. Aust. 45; 42.
=Isehnochiton (Heterozona) cariosus Pilsbry, 1892.
Holotype: DIL884, entire specimen with animal, from
Point Peron, near Rockingham, W.A,, collected by B.
Ashby, 1920.
Ischnochiton (Radsiella) délagouensis Ashby, 1931
Ann S, Afr, Mus. 3001): 40, pl. 6, figs 63-66.
=Isc¢hnochiton delagoaensts Ashby, 1931,
Holorype; D16019, girdle fragments mounted on slide
only, from Delagoa Bay, South Africa, collected by
K. H. Barnard, date of collection unknown.
Note; Rest of type in South African Museum (A 6589),
Type unique.
Ischnochiton hewitt Ashby, 1931
Ann. 8. Afr. Mus. 3001): 33, pl. 5, figs 50-53,
=Isehnochiton bergoti (Velain, 1877).
Holotype: DI0997, entire specimen and radula
mounted on slide from Table Bay, South Africa,
collector and date of collechon unknown,
Note: The above specimen is labelled type’ and with
the South African Museum No, ‘6757", However,
Ashby (1931) figured the specimen from the Albany
Museum (No. 8085) as the type. The above specimen
therefore cannot be the type but is probably a paratype.
Giles and Gosliner (1983) are similarly in error as they
list a valve from the type, also numbered A 6757,
Ischnockiton jredalei Dupuis, 1918
Bull, Mus. Nat. Hist. Nai. 24(7): 526.
=Ischnochiton lineolatus (Blainville, 1825).
Syntypes: 012718, 7 entire specimens, 6 from Penguin
and one from Stanley, north-westerti Tas,, collector and
date of collection unknown.
Note: From May collection (No, 236) labelled “para-
types", The type status of these specimens is very
doubtful as Dupuis (1918) did not give specific details
of Specimens and says that he did not sce any local
examples.
{schnochiton tredalel kingensis Ashby & Hull, 1923
Aust, Zoal, 3(2); 81, pl. 8, figs 1-4.
=Ischnochiton lineolatus (Blainville, 1825),
Holotype: D13731, median valves 3-5 and parts of
girdle stuck on card, remaining valves disarticulated,
trom Fraser Bay, King Island, Bass Strait, collected by
A, F, B. Hull, December, 1922.
107
Ishnochiton (sic.) jervisensis Ashby & Cotton, 1937
Trans. R, Soc. S, Aust, 6101): 147, pl 8, fig. 1.
=Ischnochiten pilsbryi Bednall, 1897,
Holotype: 513282, entire specimen, from Cape Jervis,
S.A., in a sheltered pool at low tide, collected by F,
Ashby, 28,1,1937.
Note: Type unique?
Ischnachiton johnstoni Cotton 1937
BANZARE Rep. Ser, B. 4(1); 11, figs 10-18.
Holotype; 014456, disarticulated valves stuck on glass
and radula mounted on slide, trom BANZARE Stn.
47 (49°S0'S, 63°33’E), off Kerguelen Island, depth
150 m, collected by BANZARE, 7.i1.1930.
Paratypes: D14456, disarticulaied valves of one
specimen stuck on glass with holotype and radula
mounted on slide, with same collection data as
holotype, D15162, 2 entire specimens with animal, with
same collection data as holotype, DI6015, 3 entire
specimens with animals, in spirit, with same collectior
data as holotype.
Ischnochiron levis Torr, 19t2
Trans. R, Soc, 8. Aust, 36: 168, pl, 6, fig. Ga-f.
=ts¢hnochiton (Autochiton) levis Torr, 1912,
Holotype: D1I1976, 5 articulated median valves with
girdle and animal and disarticulated posterior valve;
from Edithburgh, Yorke Peninsula, $,A., collected by
E. H. Matthews, date of collection unknown,
Note: Rest of type presumed lost. Type unique,
Ischnochiton (Anisoradsia) mawlei lredale & May, 1916
Proc. Matuc. Sec. Lond, 12(2/3) 108, pl. 4, fig. 4.
=lschnochiton mawlei Iredale & May, 1916.
Paratype: D12546, disarticulated specimen and remains
of girdle, trom South Tas,, collector and date of
collection unknown.
Note: A label with the specimen says “dissected by
Iredale’, a later SAM label says “Holotype (?)'. The
type does not appear to have been disarticulated,
according to the original description, and is located in
TM (E 196/7537) thus the SAM specimen is most likely
a paratype,
Ischnochiton mawsoni Cotton, 1937
BANZARE Rep. Ser. B, 4(1): 9, figs 1-9.
Holotype: Di4457, disarticulated valves stuck on glass
and radula and girdle mounted on slide, from
BANZARE Stn, 83 (54°42'30"S, 158°54'30"E), off
Lusitania Bay, Macquarie Island, depth 69 m, collected
by BANZARE, 5.xi1,1930.
Paratypes: D15160, abour 300 specimens with animals,
with same collection data as holotype. DI5J4l
disarticulated valves of one specimen stuck on glass
with holotype and radula and girdle mounted on slide,
with same collection data as holorype, DI6014, 55 entire
specimens with animals, in spint, with same collection
data as holotype.
108 REC
Ischnochiton (Haploplax) metyi var, viridis Ashby, 1920
Trans. R. Soc. S. Aust. 44: 264.
=TIschnochiton (Haploplax) mayi Pilsbry, 1895,
Holotype: 511972, entire specimen with animal, from
Lunawatina, South Bruny Island, D'Entrecasteaux
Channel, Tas., collected by E. Ashby, 22.iii.1920,
Ischnochiton (Haploplax) misimdensis Ashby, 1923
Trans. &. Soc, S$. Ausi. 47: 228, pl. 16, figs 6, 6a-c.
=Ischnochiton (Haploplax) adelaidensis (Reeve, 1847).
Holotype: D12498, disarticulated specitnen and
fragments of girdle, from Misima, Papua New Guinea,
collected by R. Andrew, date of collection unknown,
Paratype: D16552, entire specimen with animal, with
same collection data as holotype.
Ischnochiton (Stenachitan) pallens Ashby, 1900
Trans. R. Soe. 8. Aust. 24: 86, pl. 1, fig. la-e, p.
=Srenochiton pallens (Ashby, 1900).
Holotype: D978, 5 disarticulated median valves and the
anterior valve, from Gull St Vincent, S.A., dredged by
J, C, Verco, date of collection unknown.
Paratype: DI1728, entire specimen with animal, with
same collection data as holotype (labelled “cotype’).
Note: Rest of type presumed lost,
fsehnachiton (Stenachiton) pilshrvanus Bednall, |897
Proe. Malae. Soe. Lonel, 24): 142.
= Stenachiton pilsbryanus (Bednall, 1897).
Neotype: D11729, posterior and 3 median valves
articulated with remains of girdle and disarticulated
anterior and median valve, from Tapley Shoal, Gulf St
Vincent, S.A., on Zostera bed, collected by R. Tate, date
of collection unknown. Selected by Ashby (1919: 67,
pl. 1, figs 2, 2a-c).
Paraneatype: D11727, entire specimen from Marino,
near Adelaide, S.A., collected by E. Ashby, 19.11.1910.
Note: The above specimens are the “holotype” and
“paralype” respectively of Ashby’s (1919) description
of Stenochiton (Zostericola) pilsbryanus (Bednall,
1897), According to Davis ef a/, (1979) there are at least
3 syntypes of this species in ANSP (Shell Cat. No.
69142) however, Ashby (1919) refers to this material and
says “I easily identified in the material shown to me
the three species S. juloides, Ad. and Ang.; SS
cymodocealis, Ashby; and S. posidenialis, Ashby, all
very small and juvenile’. Later, Ashby (1927) doubted
the validity of his identification of the neorype and
renamed it us a new species, Sfenovhiton tatei, also
listed here,
Rest of neatype presumed lost.
fschnochiton pilshryi Bednall, 1897
Proce. Malae. See. Lond. 2(4): 143, pl. 12, figs 2, 2a-e
Neorype: D11766, entire specimen, frotn Sullana Bay,
Yorke Peninsula, S.A., collector and date of collection
unknown. Selected by Cotton and Godfrey (1940: 491,
fiz. 477),
S.ALUST, MLS. 19(K): 97-115
June, L986
Note; Fram Matthews and Bednall collection. Accord-
ing to Davis ef al. (1979) there are 3 syntypes in ANSP
(Shell Cay. No, 67369); we therefore consider the
selection of a neotype invalid. Davis er w/, (1979) also
erroneously list the type locality as Cape Yorke
Peninsula, Qld.
Since the above specimen came from the Matthews and
Bednall collection and is from the type locality we
consider it likely to be a syntype.
Ischnochiton (Heterozona) properensis Ashby, 1920
Trans. R. Soc. S. Aust. 44: 278, pl. 12, fig. Ta-b.
Holotype; 11896, entire specimen, from Praper Bay,
Port Lincoln, 8.A,, collected by E, Ashby, Tanuary,
1917
Isechnuchiion resplendens Bednall & Matihews, 1906
Proe. Malac. Soc. Lond, 1(2); 91, pl, 9, figs 4, da-f,
Ischnochiton (Haploplax) smaragdinus resplendens
Bednall & Malthews, 1906,
Neotype: 113739, entire specimen with animal, from
Marino, near Adelaide, S.A., collector and date ol
collection unknown, Selected by Cotton and Godtrey
(1940: 503, lig. 491).
Note: Listed and figured by Cottan and Godfrey (1940)
as neatype without discussion and considered, by us,
as invalid.
ischnochiton (Haploplax) smaragdinus resplendens
var. wesfernensis Ashby, 1923
Trans. R. Soe. 8. Aust. 47: 226.
Holotype: D13722, entire specimen, from Yallingup,
W.A., collected by BE. Ashby, date of collection
woknown.
Ischnochiten (Haploplax) smaragdinus var. funereus
Ashby, 1924
Trans. R. Soc, S. Aust. 48: 315.
Holotype: 011237, entire specimen, wilh animal, from
Long Reef, N-SW., collected by W. H. Hatcher, date
of collection unknown,
Ischnochion siramfeln Bergenhayn, 1931
Arkiv, Zool, JBA(3); VW, pl. |, figs 14-165 pl 2,
figs 52-56.
~Lepidochitona stroemfelti (Bergenhayn, 1931).
Syntype: DI6186, entire specimen with animal, fram
Fuertaventura, Puerto Cabras, Canary Islands, on
rocks al low tide, collected by N. Odhner, 1930.
Note: From Ashby collection.
Ischnochitan tateanus Bednall, 1897,
Prac, Malye, Sec. Lond. 2(4); 147, pl, 12, lig, Jac.
Neotype: 913738) entire specimen with animal, from
Sultana Bay, Yorke Peninsula, S.A., colleeror atid date
of collection unknown, Selected by Cotton and
Godfrey (1940: 493).
Note: Listed by Cotton and Godfrey (1940) without
MOLLUSC TYPE SPECIMENS 3. POLYPLACOPHORA
discussion, also Davis ef af. (1979) list 7 syntypes in
ANSP (Shell Cat. No. 69143) and we therefore consider
the selection of @ neatype invalid.
Ischnachtion thamuast Bednall, 1897,
Proc. Malac. Soe, Lond, 2(4): 149, pl. 12, figs 4a-d,
Sa-d,
=lsehnachiton (Haploplax) rhamasi Bednall, 1897,
Neotype: D13737, entire specimen with animal, [rom
Marino, near Adelaide, S,A,, collector and date of
collection unknown, Selected by Cotton and Godfrey
(1940; 504),
Note: Listed by Cotton and Godfrey (1940) without
discussion, also Davis ef a/. (1979) list 8 syntypes in
ANSP (Shell Cat. No. 69144) and we therefore consider
the selection of a neotype invalid.
Isehnoehiton tindalei Ashby, 1924.
Trans. R. Soc. 8. Aust, 48: 323, pl. 31, fig. 4a-c.
Holotype: D4656, disarticulated specimen with pieces
of girdle, from Groote Eylandt, Gulf of Carpentaria,
N.T., on a block of dead coral in deep water, collected
by N. B. Tindale, date of collection unknown.
Note: Type unique.
Lepidopleurus variegaius Adams & Angas, 1864.
~Isehinochiton variewatus (Adams & Angas, 1864),
sce LEPIDOPLEURIDAE.
Ischnochiton vercanis Tory, V1.
Trans. R. Soe, 8. Aust. 35; 102, pl. 24, fig. la-t.
Holotype: DI2868, entire specimen, from Ellenbrook,
south of Cape Naturaliste, W.A., in rockpool on inside
reef, collected by W. G. ‘Torr, Christmas, 1910-1).
Note: Type unique,
Genus Ischnoradsia Shuttleworth, 1853
Ischnoradsia papuaensis Ashby, (923.
Trans. R. Soe, S. Aust, 47; 227, pl. U7, fig. 2a-e.
=Isehnachiion (Ischnoradsia) pupuuensis (Ashby,
1923).
Hololype: D14316, posterior and 2 median valves
articulated with girdle, rest of specimen disarticulated,
from Normanby Island, Papua New Guinea, collected
by R, Andrew, date of collection unknown,
Note: Type unique.
Genus Stenochiton Adams & Angas, 1864
Stenochiton cvmodacealiy Astiby, 1918.
Tiuns. R. Soe. S. Aust. 42: 70, pl. 13, figs 1.4, 5; ph. 14,
figs Jl, J2a-e.
Holotype; D981, entire specimen with animal, from
Marino, near Adelaide, $.A., on Cymodoced untaretica
stems, collected by EB, Ashby, date of collection
uoknown,
109
Pararypes: D980, 2 entire specimens with animals, with
same collection data as holotype, D16545, disarticu-
lated specimen with posterior valve presumed lost, with
same collection data as holotype.
Note; D16545 is labelled “type” but was obviously
disarticulated for fig. 12. According to Ashby (1918)
the type is the specimen illustrated in liz. 5 which
corresponds to the above,
Srenochiton posidonialis Ashby, 1918.
Trans. R. Soe. S. Aust, 42: 72, pl. 13, figs 2, 6, pl. 14,
fig. W3a-d.
=Slenochiton pilsbryanus (Bednall, (897).
Holotype: D11714, entire specimen with animal, from
Marino, tear Adelaide, S,A,, collected by E, Ashby,
date of collection unknown.
Paratypes: D11718, 5 entire specimens stuck on one
eard; 2 from Cape Jervis, S.A., collected by E. Ashby
14.11.1918: 1 From Outer Harbour, S.A., and one from
Marino collected by E, Ashby, date of collection
unknown and | from Largs Bay, $.A., collected by
E, H, Matthews, date of collection unknown, D16544,
disarticulated specimen with same collection data as
holotype,
Note: 916544 is labelled “type” but was obviously
disarticulated tor fig, 13, According to Ashby (1918)
the type is the specimen illustrated in fig. 6 which
corresponds 10 the above.
Stenochiton tatei Ashby, 1927,
Pap. Proc. R. Soc. Tas. 1926: 113.
=Stenochiton pilsbryanus (Bednall, 1897).
Holotype: D1L1729, based on the same specimen as the
neotype of /schnochiton pilsbryanus Bednall, 1897,
selected by Ashby, (1919) and also fisted here,
Note: A new name for the shell described as /.
pilsbryanus (Ashby, 1919) as the identification of the
specimen selected as neotype was in doubt.
Genus Tonicella Carpenter, 1873
Chiton marmeoreus var., caeruleus Winkley, 1894,
=Tonicella prarinoreus (Pabricus, 1780).
see CHITONIDAE
Genus Trachydermon Carpenter, 1864
Trachyderinon lowei Pilsbry, 1918.
Nautitus 31(4): 127.
=Lepidochitona lowei (Pisbry, 1918).
Paratype: 011275, entire specimen, from San Pedro,
California, U.S.A., collected by H, N. Lowe, date of
collection unknown.
Note; From Ashby collection, Type in ANSP (Shell
Cal. No, 117958).
Truchivdernian (Craspedochilus) nirtoni Ashby, 1928,
Proc. Malac. Soc. Lond. V8(2); 80, pl. 6, figs 5-8,
Ho REC. 5. AUST. MUS. 19(8); 97-Lis
= Lepidochitona turtony (Ashby, 1928)
Holotype: D10982, 2 median valves, valve fragments
and lragment of girdle, from Part Alfred, South Africa,
collected by W. H. Turton, date of collection unknown.
Note: A label with the valves Says “valves not
photographed", Rest of type presumed lost, Type
unique.
Family LEPIDOPLEURIDAE
Genus Lepidopleurus Risso, 1826
Lepidopleurus badius Hedley & Hull 1909,
Rec, Aust. Mus. 7: 260, pl, 73. figs 1 & 2,
= Leptochiton badius (Hedley & Hull, 1909),
Paratypes: DI0668, 5 entire specimens with animals,
from Lony Reef, near Narrabeen, N.S.W,, collector and
date of collection unknown. 112532, one entire
specimen with animal, with same collection data as
D10668.
Note: The localities “Port Stephens” and “Kangaroo
Island S.A? have been added to DI0668 (in pencil) and
there 1s therefore some doubt regarding (he lype status
of these specimens, Type in AM (C30459),
Lepidopleurus columnarius Hedley & May, 1908,
Rec. Aust. Mus. 7(2): 123, pl. 24, Figs 27 & 28.
=Leptochiton coluennarius (Hedley & May, 1908).
Paratypes: D10667, 2 median valves, from 7 miles cast
of Cape Pillar, Tas,, dredged with holotype in 100
fathoms, collected by W. L. May, 18-xii.1907. D15679,
posterior and 4 median valves, with same collection
data as D10667,
Note: D15679 from May collection (No. 226). Type in
AM (C29060),
Lepidopleurus fintayi Ashby, 1929.
Trans, Proc, NZ, Inst, 60; 372, pl. 32, figs 5-7.
= Leptochiton finlayi (Ashby, 1909).
Holotype: D11061, 5 disarticulated median valves and
dried radula, from off Otago Heads, N-Z., dredged in
60 fathoms by H. J. Finlay, date of collection unknown.
Paratype: D16188, entire specimen with animal but
posterior valve absent, with same collection data as
holotype,
Note: Rest of type m AM (C95165),
Lepidopleurus glauerti Ashby, 1929,
JR. Soc, Wo Aust, 15: 50, fig. 16.
=Leptochiton glauerti (Ashby, 1929),
Paratype: D16152, 5 median valves articulated with
animal and remains of girdle, anterior valve and other
median valve disarticulated, from Barhurst Point,
Rottnest Island, W.A., collected by L.. Glauert, date of
collection unknown,
Note: Posterior valve presumed lost. Type in WAM
(12876).
June, 1986
Lepidopleurus tredalei Ashby, 1921.
Proe, R, Sac. Vic. (NS) 33: 187, pl. 8, fig. 3a-b,
=Leptochiton tnquinatus (Reeve, 1847).
Holotype: D11240, entire specimen with animal, from
Doubtless Bay, N.Z., collected by A. E. Brookes, date
of collection unknown,
Lepidopleurus liratus Adams & Angas, 1864.
Proe, Zool, Sac. Lond. 13: 192.
=Leptochiton liratus (Adams & Angas, 1864).
Neotype: D13735, entire specimen with animal, from
Sultana Bay, Yorke Peninsula, S.A., collected by
Matthews and Bednall, date of collection unknown.
Note: A neotype for L, /irafus was selected by Lredale
and Hull (1925a; 343) and this specimen is in the AM
(C10410), The above specimen was presumably selected
by Cotton and Godfrey (1940: 477, lig. 458) as yet
another neotype for L.. liratus; however, the specimen
does not match fig. 458. The type status of the above
specimen is therefore extremely doubtful and is invalid
in any case as Iredale and Hull (1925a) had already
selected a neotype.
Lepidopleurus matthewsianus Bednall, 1906.
Prac, Malae, Sac. Lond, 7: 92, pl, 9, fig. Ja-f
=Leptochiton matthewsianus (Bednall, 1906),
Neotype; D13734, entire specimen with remains of
animal, from Marino, near Adelaide, 5.A,, collected
by E. Ashby, 1919. Selected by Cotton and Godfrey
(1940; 476, fig. 455),
Note: Listed and figured by Cotton and Godfrey (1940)
as neotype without discussion and considered, by us,
ag invalid,
Lepidopleurus niger Torr, 1911.
Trans. R. Soc, S, Aust, 35; 105, pl, 25, fig. Sa-f.
=Leptochiton niger (Torr, 1931),
Holotype: D11686, entire specimen with animal but
with anterior valve missing, from Hopetoun, W-A.,
uncer stones in shallow pools, collected by W. G, Torr,
Christmas, 1910-11,
Note: Anterior valve presumed lost. Type unique.
Lepidopleurus pelagicys Torr, 1912,
Trans. R. Soe. 8. Aust. 36: 165, pl. 5, Vig, 2a-f,
=Leplochiton columnaris (Hedley & May, 1908).
Holotype; DI1688, anterior, posterior and one median
valve, dredged in 130 fathoms, off Cape Jaffa, S.A.,
collected by J. C, Verco, 25.xii.1905.
Note: Rest of type presumed lost.
Lepidopleurus profundus May, 1923.
lust, Index Tas, Shells, pl. 14, fig, 2, Appencix
—Leptochiton prafundus (May, 1923),
Holotype: D12533, entire specimen, dredged in 10
fathoms, off Pilot Station, Derwent River, Tas,
collected by W. L. May, date of collection unknown,
Paratype: DI6189, entire specimen with animal, with
MOLLUSC TYPE SPECIMENS 3. POLYPLACOPHORA uy
anterior valve disarliculaled, with same collection data
as holotype.
Note: May (1923) illustrated L. inquinatus Sykes, 1896
but in an appendix says “An examination of the type
in the British Museum shows that this is not (ageuinatus,
which is the N.Z. species. Our shell is now to be known
as profundus Ashby". Unfortunately Ashby’s
description of prafundus was published after May's
and in any case is a different species.
Lepidopleurus prafundus Ashby, 1923.
Trans, &, Soc, S, Aust, 47: 221, pl. 16, figs 2, 2a.
=Leptochiton collusor (Iredale & Hull, 1925).
Holotype: D11288, entire specimen with animal, from
Gulf St Vineent, $.A., dredged by J. C. Verco, date of
collection unknown,
Paratypes: D11691, 2 specimens, one entire, the other
with animal but with anterior, posterior and one
median valve disarticulated, frorn Port Phillip Bay, Vic.,
dredged by Bracebridge Wilson (No. 881), date of
collection unknown.
Note: Renamed Parachiton collusor by Iredale and
Hull (1925a), also listed here, as 1. profundus was
preoccupied,
Lepidopleurus variegaius Adams & Angas, 1864,
Proc. Zool. Soe. Lond. 13: 192.
=Ischnochiton variegurus (Adams & Angas, 1864)
(ISCHNOCHITONIDAE).
Neotype: D13736, entire specimen with animal, from
Minlacowie, Hardwicke Bay, Yorke Peninsula, S.A.,
eollected by E. H. Matthews, date of collection
unknown, Selected by Jredale and Hull (1927: 13).
Genus Parachiton Thiele, 1909
Parachiton colluser \redale & Hull, 1925,
Aust, Zool, 3(8): 346, pl. 39, fig, 22.
=Leptochiton collusor (lredale & Hull, 1925),
Holotype: D11288, based on the same specimen as the
holotype of Lepidopleurus profundus Ashby, 1923, also
listed here,
Note: There is some doubt that this specimen was the
one selected as type by [Iredale and Hull (1925a) as
Iredale and Hull (1927) list the type of P cullusor ir
SAM and the type of L, profundus in the Ashby
collection. However, later labels with the specimen say
that it is the type of P collusor and Cotton and
Godfrey (1940) also list and figure it as the type,
Parachiton verconis Cotton & Weeding, 1939,
Trans. R, Soc. S. Aust. 63(2); 183, pl. 7, fig. 2.
= Leptochitan verconis (Cotton & Weeding, 1939),
Holotype: D11689, posterior valve only, from near St
Francis Island, Nuyts Archipelago, S.A., dredged in
15-20 fathoms by J. C. Verco, date of collection
unknown,
Note: Type description was based on this single valve.
Genus Terefiochiton Iredale, 1914
Terenochiton iscus Cotton & Weeding, 1939.
Trans. R. Soc. S. Aust. 63(2); 182, pl. 7, fig. 1.
=Leptochiton iscus (Cotton & Weeding, 1939).
Holotype: D1232, entire specimen with animal, from
Cape Jervis, S.A,, collected by F, L. Saunders, 1917.
Family MOPALIIDAE
Genus Kepionella Ashby, 1919
Kopionella matthewsi var. tnfermedia Ashby, 1927,
Pap. Proc, R. Soc, Tas, 1926; 101.
=Plaxiphora (Fremblya) matihewsi (iredale, 1910).
Syntypes: D12115, 3 entire specimens with animal
remains, from Penguin, north-western Tas., collected
by E. Ashby, November, 1924,
Kopionella tasmanica Ashby, 1920.
Trans. R. Soc. 8, Aust. 44; 268, pl. 11, fig. la-d,
= Plaxiphora (Fremblya) matthewsi (Iredale, 1910),
Holotype: 112156, disarticulated specimen, from
Lunawanna, South Bruny Island, D’Entrecasteaux
Channel, Tas., collected by E. Ashby, March, 1920,
Note: Smith and Robertson (1970) list the holotype as
lost as it could not be located at the time.
Genus Plaxiphora Gray, 1847
Plaxiphora hedleyi Torr, 1911.
Trans, R, Soc, 8, Aust, 35; 103, pl. 24, fig, 2a-t
=Plaxiphora (Fremblya) matthewsi (Iredale, 1910),
Holotype! D12871, entire specimen with anterior and
posterior valve disarticulated, from Rabbit Island, near
Albany, W.A,, collected by W. G. Torr, Christmas,
1910-11.
Plaxiphora pustulosa Torr, (911,
Trans. R. Soc, 8S. Aust. 35: 107, pl. 25, fig. 7.
=Liolophura (Clavarizona) hirtosa (Blainville, 1825)
(CHITONIDAE).
Holotype; 113719, one median valve only, from
Albany, W.A,, collected by W, G, Torr, Christmas,
1910-11.
Note: Type description was based on this single valve.
Plaxiphora zebra Torr, 1911
Trans, R, Soc, S, Aust, 35: 106, pl. 25, fig. 6.
=Plaxiphora (Fremblya) matihewsi (redale, 1910),
Holotype: D12869, one median valve only, from Port
Esperance, W.A., collected by W. G. Torr; Christmas,
L910-J 1.
Note: Type deseription was. based on this single valve,
2 REC. S. AUST. MUS. 19(8E YT ATS
Family SCHIZOCHITONIDAER
Genus Lerica H & A Adams, 1852
Lorica elliottae Cotton & Weeding, 1939,
Trans. R. Soe. S. Aust, 63(2): 189, pl. 7, fig. 9.
Holotype: D11658, entire specimen, from Rottnest
Island, W.A., collected by L, A. Elliott, January, 1933.
Note: Type unique.
Lorica haurakiensis Mestayer, 1921.
Trans, Prac. N.Z, last. 53; 177, pl. 38, figs 1-3.
Paratype: D1J089, entire specimen, dredged in 20
fathoms, off Kauwau and Tiritir islands, Hauraki
Gulf, near Auckland, N,Z., collected by A, E. Brookes,
date of collection unknown.
Note; Type in NMNZ (M1121),
ine, 1986
Genus Lori¢vella Pilsbry, 1893
Loricella terri Ashby, 1919,
Trans. R. Sac. S. Aust, 43: 62, pl. 10, fig. 16.
= Loricella angasi (H, Adams in H. Adams & Angas,
1864).
Holotype: D12440, entire specimen, from Quaranline
Station, Port Jackson, N.SAW., in shallow water at low
tide, callected by E. Ashby, November, 1918.
Family SUBTERENOCHITONIDAE
Genus Subterenochiton |redale & Hull, 1924
ischnochiton bednalli Torr, 1912.
=Sublerenochiton bednalli (Tarr, 1912),
see ISCHNOCHITONIDAE.
REFERENCES
ASHBY, FE. 1918. Monorrph of the genus Srevociifan (Order
Polyplacophora), with descriptions of (wonew species, Trams.
K, Soe, S Aust 42: 63-78, pls 13 & 14.
ASHBY, E. 1919. Noics on Austrian Palyplucophors. Including
descriptions of two few venera, & aew variety, and the
description and propesed recognition of Mr, Bednall’s
Stenachitan pilsbryanus, Trans. Ro Sov, S. Aust, 43: 66-73, pl. 1),
ASHBY, F. 1922, Types of species of Austration Polyplacophora
described by De'Blainville, Lamarek, De’ Rachbrune, and others,
now in the Museum d'Histoire Naturelle, in Paris. Jravts. R. Soe.
S. Aust 4: 572-542.
ASHBY, LE, 1926, The Acanthoid Chitons of New Zealand, with
deseriprions and figures, incliding several new species. Pree,
Malue. Soe. Lond. 17h 5-34, pls 1-4.
ASHBY, E. 1927. Notes on, and additions to, the Chiton fauna of
North-West Tasmania, together with a brief review af the genus
Srennchiton. Pap. Proe. RB. Soe, Tasin, W26: 92-17.
ASHBY, E, 1928. Notes ona collection of chitons (Polyplacaphora)
trom the Capricorn Group, Queenslancl. Tryas. R, Sac AS Ausy
52: 167-173, pl. 12.
ASUIBY, B. 1931. Monograph of the South African Polyphicophorit
(Chitonsy) Avi S. Ay Mus, SQL): 1-59, Oly 1-7.
ASHBY, E,, and COTTON, B.C. 1937, Deseription of two new
species of Australian Cliitoims with additonal notes and records.
Trans, R. Sov. 8. Aust, Als AS-148, pl 8.
BERGENHAYN, J. R. M. 1931. Beitrage cur Malakovoologie der
Kanarisehen Inseln. Die Loricaten. “Arkiv. Z00/. 23A(13): 138,
pis, 13,
COTTON, 8. C. 1964, Seuil Ausiralian Moallusea: Chitans, Govt.
Printer Adelaide, 151 pp,
COTTON, B.C. and GODEREY, FF K, (940. Phe Mollisey of South
Australia: Part tl Scahapode, Cephalapoda, Aplacepohora
und Crepipoda. Cove, Printer Adelaide, 284 pp.
COTTON, B. C., and WEEDING, B. J, 1939. Flindersian Lorjcales,
Trans, Ry Soe S. Aust, 63; (80-199, pl 7
DAVIS, G. M., ROBERTSON, R., and MILLER, M. 1979, Catalogue
al the Chiton types of the Academy of Warural Setences uf
Philadelphia. Teveria Po 1-60.
DUPUIS, P1917. Notes prises au conrs de Vexumien de la Collecion
de Polyplacophres du Mustum de Paris. Bull, Mts. Neti, Hist.
nal. Paris 23. 553-8.
DUPUTS, P1918, Notes convernanr les Pelyplavephores, Bull, Mrs.
Nat. Hist. Nat. 24; 525-533.
GILES, E,, and GOSLINER, T, (983. Primary rype specimens of
marine Mollusca (excluding Cephalopoda) inthe Sour Africa
Museum, Ann Si d/h Mus. 925 1-52,
IREDALL, T, and HULL, A, OB. 1925a, A Monograph ot
Abstraliag Loricates, Part IV. lst Zool 3: 339-362, pls
39-40),
IREDALE, 1., and HULL, AL FB. 19256, A Monograph ot
Australian Loricates, Part V, dust Zool 4: 75-11, pls 9-12.
IREDALE, T.. and HULL, AL FB. 19270 4) Movtoeruph uf
Austratian Larteates (Phvliimn Malhiscd Order Liricatiu. R-
“001. Soc. NUSAV, publ. 168 pp,, 2) plates,
KAAS, P., und VAN BELLE, R. A. 1980. Cotulogie of Living
Chitoms. Dr W. Backhitys, Publ, Rerlerdan, 144 pp
MACPHAIL, M. K., and ZFIDLER, W. 1978. History In a
Tasmanian Chiton, Tes, Net. No, 53: 1-7,
MAUGHAN, M. M.. 1900, Definition of a new species of South
Australian Polyplacophora. Tras, Ro Soe, 8. Aust, 24: 89, pl |
VIAY, WoL. 1923. 44 Sihistrated Index of Tesiianian Stiedls, Gav.
Printer Flobarl, 54 pp., 50 pls., Appendix.
SMITH, B. J, and ROBLRTSON, R.C. 1970. Catalogue af Chiton
(A phineura, Matlusea) Iypes fa ric National Muse ol
Victoria, Australia, Wer, Nat Mates, Pie. 3p 8190,
APPENDIX
List of Chiton “types” from the Dupuis collection,
given to Ashby by Dupuis, All are labelled “TYPE”
or “CO-TYPE” usually with a Dupuis label with the
word "TYPE" in red print, glued to the label and the
prefix “co?’ added by hand. The specimens are listed
in alphabetical order according to the original name
at the time of description,
Acanthopleura balansae Rochebrune, 1882.
Bull. Soe. Philam. Paris, Ser. 7, 6: 197.
=Acanthaplenra heddoni Winkworth,
(CHITONIDAE).
Syntype: D10231, entire specimen with animal, from
Timor, collector and date of collection unknown.
1927
Note: Speeimen with label “co TYPE de. baluasae
Roch!" and reidentified as A. spinigera Sowerby, The
locality given by Rocheburne (1882) is “Australie (Peron
et Lesucur), Nouvelle Caledonie (Balansa; Ciermain)"”.
The type status of the above specimen is therefore very
doubtful.
Chiton elongatus Blainville, 1825,
Dict. Sei. Nat. 36: 352,
=Ischnochiton elongaius (Blainville, 1825)
(SCHNOCHITONIDAE).
Syntype: D10237, entire specimen, from King Island,
Bass Strait, collected by Péron and Lesueur, 1802.
Nate: The shell is marked on the inside “ile King”
MOLLUSC TYPE SPECIMENS 3, POLYPLACOPIIORA tls
presumably in the handwriting of Péron or Lesueur and
is a historically significant shell (Macphail & Zeidler,
1978). Jt was sent to Ashby by Dupuis as "“'co-types”
of f. lineolatus (Ashby, 1922) and a Dupuis label says
the same, Ashby (1922) determined it as J. crispiis
(Reeve, 1847) which is now considered a synonym ot
f, elongatus, The above specimen matches the original
deseription of C. elongarus and considering that it was
collected by Péron and Lesueur it was most likely part
of (he series of specimens originally seen by Blainville.
Syntypes: DLO238, 3 entire specimens with same
collection data as DI0237.
Note: =f subviridus (tredale & May, 1916) but would
have been considered /, e/ongarys in the past. There
is no original label with the specimen but a label
presumably written by Ashby says “col. by Péran and
Lesueur, [s, King (802 (with Blainville’s type of
lineolatusy”. If the information with the specimens is
correct then they could also have been part of the
original material seen by Blainville.
Chiton lamsyi Dupuis, 1917.
Bull. Mus. Nain, Alist. nat. Paris 23: 538,
=Chiton peregrinus Thiele, 1910 (CHITONIDAEB),
Syntypes: D102), 10 entire specimens, only one with
animal, from Aden, Red Sea, collected by Dr
Jousseume, clate of collection unknown. D10252, one
disarticulated specimen with piece of dried girdle, fram
the Red Sea, collected by Dr Jousseume, date of
collection unknown. DLO0255, one disarticulated
specimen with piece of dried girdle with same collection
data as D10252, D11200, one entire specimen from the
Red Sea, collector and date al collection unknown,
Note: All of the above are labelled “TYPE” except for
011200 which is labelled “vo-type’. As Dupuis did nat
clearly designate.a type and since the aboye specimens
were given ta Ashby by him, it is likely thar they are
all syntypes.
Chitan tehuelehus YOrbigny, 1841.
ov. Amer. merid., Mall, 3(3): 488, pl. 65, figs 7-13,
=Chuetopleura angulata (Spengler, 1797)
USCHNOCHITONIDAEB).
Syntype: 10293, entire specimen, from Bay of San
Blas, Patagonia, colleclor and date of collection
unknown.
Note: The above specimen is from the type locality but
in the absence of orivinal labels the type status must
be very doubtful,
Chiton undulatus Quoy & Gaimard, 1835.
Voy. de l'Astrolabe, Zool. 3: 393, pl. 75, figs 19-24.
=Onithochiton nezlectus Rochebrune, 1881
(CHITONIDAR).
Syntypes: D10223, 2 entire specimens, from N.Z.,
collector and date of collection unknowh,
Note: Specimens of very doubtful type stabus. The type
is in MNEIN and was seen by Ashby (1922). However,
Quoy and Gaimard recorded several specimens but
apparently did not designate a type so the above
specimens could have been part of Lhe Lyne Series,
Chiton violaceus Quoy & Gaimard, 1835,
Voy. de ‘Astrolabe, Zool, 3: 403, pl, 73, figs 15-20,
=Notophix violaceus (Quay & Gainard, 1835)
(ACANTHOCHITONIDAEB),
Syntype: DIL0S4, disarticulared specimen, from N.7.,
collector and date of collection unknown.
Note: A specimen af doubtful type status, The Dupuis
label with the specimen has “Tasman Bay nr. Nelson”
added in pencil, which is the type locality.
Cryptoplax caledonicus Rochebrune, 1882.
Bull, Soc. Philom. Paris, Ser. 7, 6: 196,
=Criptoplux larvaeforinis (Burrows,
(CRY PTOPLACIDAE).
Paratypes: DLO330, 3 entire specimens with animals,
fram New Caledania, collector and date of collection
unknown.
Nore; A Dupuis label with the specimens designates
them as “‘co-types” but a later Ashby collection label
has the wards “with TYPE" on it thus indicating that
the above specimens could be paratypes.
1815)
Cryptoconchus stewartianus Rochebrune, 1882.
Bull. Soc. Philom. Paris, Ser. 7, 6: 194.
=Cryptocanchus porasus (Burrows,
(ACANTHOCHITONIDAE).
Paratype: D11053, disarticulated specimen, from Cock
Strait, N.Z., collector and date of collection unknawn.
Note: The above specimen is most likely a paratype as
Ashby (1926) deseribed and illustrated it and referred
fo it as “ane of Rochebrune’s cotypes”. The type, a
spirit specimen, is in MNHWN (Ashby 1922).
1815)
Gymnoplax spiciierus Rochebrune, 1884.
Bull. Sac. Philom. Paris, Ser. 7, 8: 36.
=Isehnochiton (Ischnoplax) pectinatus (Sawerby,
1840) (SCHNOCHITONIDAEB).
Syntype: DI0297, 1 median valves, from Cochino
Island, Guadeloupe, callector and date of callection
unknown.
Note: The above material is from Lhe type localily and
was probably part of the type series.
Lenidopleurus campbelli Pilhal, 1880,
Comptes Rendus hebd. Séanc. Acéd. Sei. Paris, tl
sags,
= Ischnochiton cireumvallatus (Reeve, 1847)
GSCHNOCHITONIDAE).
Syntype: D11022, animal and disarticulated valves,
from Campbell Island, N.Z., collector and date of
collection unknown.
Note: There is some doubt that this is one of Filhol’s
types; however, it matches the description given by
Dupuis (1917) for specimen “B” of two specimens that
hi
he regarded as type material of L, cumpbelli, There is
also a radiila slide with D11022 but it does not belong
to the above specimen; however, [he SAM register refers
to lwo specimens one of which may have been used
for the radula preparation.
Lepidopleurus cessaci Rochebrune, 188).
Bull, Soc. Philom, Paris, Ser. 7, 5: 118.
=Ischnoachiion cessaci (Rachebrune, 1881) ISCHNO-
CHITONIDAE).
Syntype: D10324, disarticulated specimen, from Cape
Verde Island, collector-and date of collection unknown,
Nore: SAM records indicate that there were another
§ specimens which Ashby kept and were presumably
lost in the fire at Ashby’s house. The above specimen
is from the type locality arid was probably part of the
Type series,
Lepidopleurus fodiatus Rochebrune, 1881.
Bull. Soe. Philom. Paris, Ser. 7, 5: 119.
=Isehnochiton textilis (Gray, 1828)
(SCHNOCHITONIDAE),
Syntype: D10286, entire specimen, from Natal, South
Africa, colleered by M. Verreaux, date of collection
unknown.
Note: The type status of this specimen is in doubt. The
label with the specimen reads “Jschnochiton tigrinus
Krauss (co-TYPE de Lepidopleurus fodiatus Rocheb.
Australia!!) Verreaux Nazal?”, Obviously there is some
confusion regarding the localily data.
Ashby (1922) says that he has never seen this species
in Australia and is confident that New Holland, the
type locality, is erroneous.
Lepidopleurus melanterus Rochebrune, 1884,
Bull. Soc. Philom. Paris, Ser. 7, 8: 37.
=Ischnochiton circumvallatus (Reeve, 1847)
(ISCHNOCHITONIDAE),.
Syntypes: D10292, 7 ertire specimens with animals,
from Campbell Island, N.Z., collector and dale of
collection unknown, D10319, one entire specimen with
animal, with same collection data as D10292. D13061,
one entire specimen with animal, with same collection
data as DI0292,
Note: The above material 1s from the type locality and
al least D10292 could have been part of the type series.
D10319 and D13061 are without the usual Dupuis label
and may have been extracted from D10292.
Lepidopleurus rochebruni Jousseaume, 1893
Bull. Soc. Phila. Paris, Ser. 8, 6: 02.
=Callistochiton adenensis (E. A. Smith, 1891)
(CALLISTOPLACIDAE}),
Syntype! 010326, one entire specimen, from Djibouti,
Gulf of Aden, collector and date of collection
unknown. DI0342, 2 disarticulated specimens, one
from Djibouti, the otber from Aden, Gult of Aden,
collector and date of collection unknown,
KEC, 5, AUST. MUS. 1908); 97-115
Sune. (986
Note: D10326 is clearly marked “cotype du Dr
Jousseaume” and the specimen matches the description
and the largest measurement given by Jousseaume. It
is therefore most probably a syntype, The type status
of D10342 is rather more doubtful.
Notachiton mirandus Thiele, 1906
Wiss. Ergebn. d. Tiefsee Exp. 9(2): 332, pl. 29, figs
11-16.
=Nuttallochiton mitandus (Thiele,
NOCHITONIDAE).
Syntype: D10213, 5 median valves articulated with
airdle and disarticulated anterior, posterior and median
valve 2, [rom Antarctica, collector and dale of
collection unknown,
Note: Despite the fact that D10213 is clearly labelled
“cotype” it cannot be Thiele’s type of N, wirandus as
Thiele mentions only one specimen and an anterior
valve from east of Bouvet Island (Valdivia Sti. 127)
and this specimen was several times larger than D10213.
1906) (ISCH-
Onithochiton filhali Rochebrune, 1881
Bull. Soc. Philom., Paris, Ser. 7, 5: 120.
=Onithochiton neglects Rochebrune,
(CHITONIDAE)
Syntype: D10215, disarticulated specimen, from Cook
Strait, N.Z., collector and date of collection unknown,
Note: Clearly labelled “coTYPE de O. filholi Roch”,
1881)
Onithochiton Rochebrun¢, 1881
(CHITONIDAE)
Bull, Soc, Philom, Paris, Ser. 7, 5: 120.
Syntypes: D10221, two entire specimens with animals,
without collection data.
Note: [hn the absence of original Jabels and pood
locality data it is difficult to ascertain the type status
of the above specimens.
neglectus
Schizachiton hyadesi Rochebrune, 1889
Miss, Sei, Cap, Horn 4 Zoal,, : 132, pl. 9, fig. 1.
=Nuttallochiton hyadest (Rocheburne, 1889) (ISCH-
NOCHITONIDAE)
Syntype: D10189, entire specimen with animal, from
Terra del Puego, collector and date of collection
unknown.
Note: The type status of this specimen is very doubtful.
The original label does not indicate the type status but
the words “cotype-paratype” have been added in pencil
on the back of the label.
Sehizochiton jousseaumei Dupuis, [917 (SCHIZO-
CHITONIDAE)
Bull. Mus, Natn. Hist. nate: Paris, 23: 536, figs 1-4.
Syntypes: D10233, 2 entire specimens with animals plus
one anterior valve, from the Red Sea, collected by
Jousseaurne, date of collection unknown. D10235, one
entire specimen with animal wilh same coilection data
as DI0233
MOLLUSC TYPE SPECIMENS 3. POLYPLACOPHORA 15
Note: Although the above specimens are labelled
“TYPE” it is unlikely that any of them are types as
the species was based on a unique specimen which Is
in MNHN. The single anterior valve of D10233 is
labelled ‘‘Perim’”’, the type locality.
Tonica fontainei Rochebrune, 1882
Bull. Soc. Philom. Paris, Ser. 7, 6: 193.
=Ischnochiton punctulatissimus (Sowerby,
(ISCHNOCHITONIDAE)
Syntypes: D10291, 4 entire specimens with animals, 2
disarticulated specimens plus 3 end valves and 4 median
valves from at least 2 specimens, from Chile, collector
and date of collection unknown.
Note: The specimens are clearly marked “co-TYPE”
and are probably types.
1832)
Tonicia lebruni Rochebrune, 1884
Bull. Soc. Philom. Paris, Ser. 7, 8: 35.
Syntype: D10196, entire specimen with animal, from
Patagonia, collector and date of collection unknown.
Note: The specimen is labelled ‘““TYPE” and in the
absence of evidence to the contrary it must be
considered possible type material.
ACKNOWLEDGMENTS
We wish to thank the following for supplying us with
information regarding types held by their respective
institutions: Dr W. O. Cernohorsky, AIM; Mr I. Loch,
AM; Dr B. A. Marshall, NMNZ; Mr J. Stanisic, QM;
Dr A. M. Testud, MNHN and Dr F. E. Wells, WAM.
We are especially grateful to Dr M. K. Macphail who
initially curated the mollusc type collection under a
State Unemployment Relief Scheme. We also wish to
acknowledge Mrs Jan Forrest, SAM, who took the
photographs and prepared Figure | and Miss J. Connor
who typed the manuscript.
A REVISION OF THE SOUTHERN AUSTRALIAN STARFISH GENUS
NECTRIA (ASTEROIDEA: OREASTERIDAE) WITH THE DESCRIPTION
OF A NEW SPECIES
BY W. ZEIDLER AND F. W. E-. ROWE
Summary
The status of the genus Nectria is discussed and the genus transferred from the family Goniasteridae
to the family Oreasteridae. A previously closely allied genus, Nectriaster, is also transferred to the
Oreasteridae. The status of the subfamily Nectriinae is discussed and maintained for the unique
genus Nectria. A new subfamily, Oreasterinae, is erected for the remaining genera of the family
Oreasteridae including Nectriaster. Problems relating to Gray’s (1840, 1847a, b) misunderstanding
of N. ocellifera which he based on Oudart’s (1827) figure of Lamarck’s species are discussed and
Oudart’s figure is reproduced for the first time. Eight species are currently recognised including a
new species from Tasmania. Each of the eight species is diagnosed and geographic ranges detailed.
A lectotype is selected for N. ocellata Perrier. N. pedicelligera Mortensen is recognised as a valid
species restricted to the southern Australian coast. The species potential of the genus is considered
not yet to have been reached. Nectria is recognised as an endemic southern Australian genus.
A REVISION OF THE SOUTHERN AUSTRALIAN STARFISH GENUS NECTRIA (ASTEROIDEA;
OREASTERIDAE), WITH THE DESCRIPTION OF A NEW SPECIES
W, ZEIDLER! and F, W. E, ROWE?
‘South Australian Museum, Narth Terrace, Adelaide, South Australia 5000
* The Australian Museum, 6-8 College Street, Sydney, New South Wales 2000
(Manuscript accepted 3 Murch 1985)
ABSTRACT
ZLIDLI-R, W., and ROWE, F. WE. 1986. A revisiorcof the southern
Australian sturlish genus Neerria (Asteroidea; Oreastenidae),
with the deseription of a new species. Ree. S, Ada, Mus, 19):
117-138.
The status of the genus Neertria is discussed and the
venus lransferred (rom the family Goniasteridae to the
family Oreasteridae. A previously closely allied genus,
Nectriaster, is also transferred to the Oreasteridae. The
sratus of the subfamily Nectriinae is discussed and
maintained for the unique genus Nectria. A new
subfamily, Oreasterinac, is erected for the remaining
genera of the family Oreasteridae including Nectriaster,
Problems relating to Gray's (1840, 1847a, b) misunder-
standing of N, ocellifera which he based on Oudart’s
(1827) figure of Lamarck’s species are discussed and
Oudart’s figure is reproduced for the first time, Eight
Species are currently recognised including a new species
from Tasmania. Each of the cight species is diagnosed
and geographic ranges detailed. A lectotype is selected
for N. ovellate Perrier. N. pedicelligera Mortensen 15
recognised as a valid species restricted to the southern
Australian coast. The species potential of the genus is
considered not yet to have been reached, Necsria is
recognised as an endemic southern Australian genus.
INTRODUCTION
Although Necrria is a well established southern
Australian genus andl is easily recognisable, there has
been some confusion regarding the type species and
the identity of a number of the species despite the
reviews of Clark (1966) and Shepherd (1967),
We have examined all available type material for cach
of the species together with the collections held in the
Australian Museum Sydney (AM); Museum of
Victoria, Melbourne (MV) (specimens not measured);
South Australian Museum, Adelaide (SAM); Western
Australian Museum, Perth (WAM): Tasmanian
Museum and Art Gallery, Hobart (TM); Queen
Victoria Museum and Art Gallery, Launceston (QVM)
(specimens not measured) and Museum of
Comparative Zoology, Harvard, U.S.A. (MCZ).
Following our examination of this material it is
apparent that Clark (1966) and Shepherd (1967) did not
appreciate (he limits of (he species,
We have also obtained a copy of Oudart's (1827)
figure of Asterias ocellifera Lamarck 1816 (there is no
accompanying text) which was used by Gray to identify
his specimens when he established the genus Nec/ria.
This figure has apparently not been sighted since
Perrier (1875) and we reproduce it here (Fig. 1) in order
to clarily the confused situation created by Gray's
(1840, 1847a, b) use of this figure.
We are now able to recognise cight species of Nectria
one of which is new to science. We are further able to
detail the distribution of these eight species.
Neciria is transferred to the family Oreasteridae but
the status of the subfamily Nectriinae is maintained.
The following additional abbreviations are used in
the text.
BMNH =British Museum (Natural History)
MNHN=Museum National D’Histoire Naturelle, Paris
NSW = =New South Wales
SA =South Australia
Tas =TJasmania
Vie =Victoria
WA =Western Australia
SYSTEMATICS
Family OREASTERIDAE Fisher, 1911
Subfamily Nectriimae Perrier, 1894
Genus Nectria Gray, 1840
Nectria Gray, 1840 (1841): 287; 1866: 15; Dujardin
& Hupe, 1862: 406; Perrier, 1875: 185; 1876: 1; 1894:
333; Viguier, 1878: 187; H. L. Clark, 1946: 85; A. M.
Clark, 1966; 309,
Diagnosis: An oreasterid genus with § tapering arms
(R up to 125 mm) and relatively large disc (r up to
45 mm), Abactinal surface convex, actinal surface fat.
Abuactinal skeleton comprises distinctive raised tabulae
cach supported by 6 radialing ossicles at the base
forming a network. Disc tabulae crowned with
peripheral ring of usually prominant granules encircling
central granules, remainder with granules of varying
shape and size. Tabulae distinguished on dise,
decreasing in size along arm to about '2 R after Which
they become unrecognisable as tabulae, Marginal plates
obvious, flat, squarish, infera- and supero-marginals
of similar size and number decreasing regularly in size
118 REC. S. AUST. MUS. 19(9): 127-138
16 arm Lip, covered with even-sized granules. Actinal
plates up to 6-8 rows at base of rays, decreasing distally,
covered with coarse granules. Adambulacral plates with
furrow spines being lice larger than adjacen| granules
on actinal plates, Papulae in discrete groups between
radiating ossicles of dorsal skeleton sometimes
occurring between marginal plates but rarely on actinal
surface immediately below inferomarginal plates.
interradial septae partially calcified. Pedicellariae alten
present (Fig. 2). Superambuluacral plates present-
Nectria is an endemic Australian genus occurring
from the mid-west coast of Western Australia
(Houtman Abrolhos) along the southern coast to
Broughton Loff Port Stephens, N.S.W. in the east (Fiz.
3). Records of specimens from Fiji (Sladen 1889), New
Zealand (Mortensen 1925) and Mauritius (MV
specimen) are erroneous, Species oceur from the
shoreline down to 550 m and are recorded fram habitats
associaled with rocky reels.
Remarks; The genus was established by Gray (1840)
based on a specimen or specimens without locality data
which he identified as Asterias ocellifera Lamarck, 1816
(as “oculifera” \apsus cal.) using Oudart's (1827) figure
(Fig, 1) of Lamarck's species. However we have
determined that of the few dried specimens of Necttia
in ihe BMNH from the mid-l9th century likely to have
been seen by Gray (1840), none are N. ocellifera.One
specinien in particular (BMNH; 1953; 4.27.24) (Fig. 6b)
which was cerlainly seen by Gray “as its oldest label
has ‘Neerria ovulifera’ stuck on the back of the board
with ‘=A sterias acellifera Lamarck’ belaw and “Gray"
pencilled after ocilifera’’ (A. M. Clark, pers. comm.)
is clearly N. ocellata.
Thus. like Perrier (1875) we canelude that Gray (L840)
mistook one species for the other. Clearly Gray never
saw a specimen of N, ecellifera for later (1847a, b) he
identified a specimen as Poerria ovellifera, nating that
he may have mis-identified the BMNH Nectria since
he says “ihis species [A ocellifera| more nearly
resembles Oudart’s figure than the species | have
described under the name Nectria oculifera’”’. In shape,
but not structure, NV. ocel/ifera resembles P ocellifera
more sa than daes N. ocellate, We regard Gray’s (1847)
Poariria ocellifera another mis-identification based on
Oudart’s figure and not a description of anew species,
However, Clark (1963) has redescribed and figured
Gray's specimen of P ocellifera as the holotype of that
Species since she does not consider Gray mis-identified
the specimen.
The above confusion raises the problem of whether
acellifera or ocellata Should now be the type species
of the genus. Although H. L. Clark (1946) has stated
“sd sterias ocellifera is the type species af Nectria we
concur with A. M. Clark (1966) in that ‘strictly
speaking, N. oce/fara Perrier tnight be considered as
the type species since that 1s the one thal Gray had
before him when he diagnosed the genus Neetria,
though he mistakenly identified it with Lamarck's.
June, 1956
species” According ta Article 70a of the Internatianal
Code of Zoological Nomenclature (LCZN) a decision
on such a matter should be lelt ta the Commission.
Neetria was first placed in the family Pentacerotidae
by Gray (1840); subsequently Perrier U875) included
the genus in the Goniasteridae, dropping all reference
to the Pentacerovidae. Later (1894) Perrier plaved the
genus in its own subfamily the Neetriinge on the basis
of the distinctive tabulae, Following the exammation
af two specimens Fisher (1911) maintained the sub-
familial status in the family Goniasteridae on the basis
of the presence of intermarginal papulae which he
considered a most important character; he also
deseribed the occurrences of superarmbulacral plates in
a number of goniasterid genera including Neciria.
Spencer and Wright (1966) uphald Nectriinae on the
same basis as Fisher (1911). They include wirhin the
subfamily one other genus, Nectriasier H, L. Clark,
1946. However, we transfer both of these genera to the
tamily Oreastertdae, since, like such genera as Oreaster
and Penraceraster, Nectria has a substantially complete
caleartous. interbrachial septum, and Neerriester
possesses a complete, calcareous interbrachial septum.
The inferbrachial septum of zoniasterids is cypically
membranous. Also, in both Neetria and Neetriaster,
the alignment of marginal plates, their granulose
covering, and the arrangement of abactinal plates in
particular, are similar to other oreasterids.
The relationship between Neeiriag and Necfriaster
does not appear as close as that Suggested by H, L,
Clark (1946) and subsequent authors. Indeed the dorsal
armament of Nectriaster is not unlike that af Oreaster
and Pentaceraster, to which it may be considered more
closely aligned. However, the distinctive abactinal
tabulae af Nectria stands it apart from all other
oreasterids and for this reason we prefer to retain the
subfamily Nectriinae, as did Perrier, but within the
family Oreasteridae. With the recognition of this
subfamily for the type-genus Neetrig, the remaining
genera within the Oreasteridae should be referred (until
otherwise revised) to the oominative subfamily
Oreasterinae sub.fam.nev, which is diagnosed herein
as a subfamily of Oreasteridae which lacks tabulac and
for which Qreuster is nominated as the type-genus.
Clearly, however, the family Oreasteridae is in need
of entical reappraisal.
KEY TO THE SPECIES OF NECTRIA
| Inrermarginal papulae present, when absent many distal
ur plates enlarged .2.. 02... ee ee ee eee 2
—Intermarginal papulae absent, distal arm plates never
pyle ey ca eg oh ees tet opal oy ot ge 5
2 Tabulae with concave aspect, with central granules lower
in. protile and smatier than periphals
REVISION OF THE GENUS NECFRIA Wy
3 Dorsal arm plates decrease rezularly in size towards arm
tip; pediccllariag with slender valves oo) ec... eee
Why ebb sel pede reo N. pedicelligera
— Dorsal arm plates of mixed sizes towards arnt Up, some
almost as larve as.dise plates: pedicellariae with slender
or broad valves
4 Purrow spines 4-6 (rarely 3 or 7), pedivellariae common
with broad valves; imlenmarginal papulae always present
eens el Fes feet cs cer N. mmultispiita
— Furrow spines 3-4 (rarely 2) pedicellariae rare with slender
valves; intermarainal papulae sometimes absent —-.
ics oy bet fn fs wees as ey, Oh ye pte nne hale <A?
5 ‘tabulae low abou! | mm high 2... A. /emilis: sp.nov.
— Tabulae well developed more than | omm high .... 6
6 Marginal plates with granules discinetly smaller (han on
actinal surface; tabulae with very low, at central
granules and radiating scale-like peripheral granules
R/r= 2.65 =0.06 (495% cl.) N. ecellifera
— Marginal plates with coarse granules or granules similar
Lo aclinal surface; tabulae witlt central granules rarely
flat, peripheral granules wedge-shaped sometimes
radiating. LU marginal plares wilh fine granulation and
tabuluc with law Plat central granules, chen peripherals
not radiating and R/r «3114002 (495% eb) 2. 7
7 Tabulae with peripheral abd central granules few in
number (usually each < 20), irrewular in size and shape,
round in cross-section, peripherals radiating; dorsal arm
plates always indistinct distally with enlarged central
GIUATTHOS. oc eines Detter reads N. wilsant
— Tabulae with numerous peripheral and central geaiules
(usually >20, often >30), usually regular in size and
shape, peripherals forming compact ring around
centrals, rarely radiating; central granules convex,
usually crowded, larger than peripherals, dorsal arm
plates will more or less clistinct limits distally and with
celal granules not prominent (FORM 1); central
granules irregular in size, dorsal arm plates indistunee
distally with central granule or granules prominent (as
in wilyoni) (FORM 2); central granules lower than
peripherals, becoming flat in extrenme cases (as in
ocellifera), dorsal.arm plates with distinct Imits distally
anc wilh even granulation (FORM 3). N. veellata
Nectria ocellifera (Lamarck)
(Figs 1 & 4a)
Asterias ocellifera Lamarck, 1816: 553; 1840: 239 (part);
Oudart, 1827: plate 1.
Goniodiscus ocelliferus: Miller & Troschel, 1842: 60.
Neetria ocellifera: Dujardin & Hupe, 1862; 406; Perrier,
1869: 283; 1875: 147; 18746: 3; A. L. Clark, 1914; 139;
1916: 35; 1928; 379 (key); 1946: 86; Shepherd, 1967:
464, fig. 3 (part); Marsh, 1976: 217 (table).
Nectria ocellata: Shepherd, 1967: 467 (parr).
Muareriul examined: Holotype, MNHN; ECAS 670
(R/r=55/20 trun), mers australes?, Péron et Lesueuw,
1803 (Shepherd (1967) has suggested ihat Ihe type
probably came from Geographe Bay, W.A.).
In addition 65 specimens fram WAM, 3 specimens
from AM and one specimen from MCZ.
Diugnosiss R up to 75 mm, ¥ up ta 30 mm.
R/r=2.6540,06 (£95% el.), Tabulae with ventral
vranules irregular in size and. shape, flat, very low in
profile (approx, 0.2 mm above tabula), slighuy spaced,
angular in cross-section. Peripheral granules radiating,
about twice as long as wide, up fo | min high, thin,
scale-like, lapering to rounded tip, Tabulae more or less
confined to disc. Dorsal arm plates slightly convex,
decreasing in size towards arm tip, with low granules
giving smooth covering, peripheral granules reduced
not distinetive, centrals larger than peripherals, limits
of plates distinct to naked eye. Marginal plates distinct,
granules of similar size to each other, close but not
compact, distinctly smaller than those on uctinal or
abactinal surfaces. Aetinal plates up to 6 rows
proximally, Furrow spines 2-4. Subambulacral spines
3-4 an lirst row and 2-3 on second row. Papulue
numerous dorsally extending almost to arm tips, absent
intermarginally and actinally, Pedicellariae absent.
Colour red or orange according (o Clark (1946), or pale
pink to pale orange-pink (L. M. Marsh, pers. comm,).
Remarks: The form and granulation of the tabulae
(Fig. 4a), the distribution of the papulae, the fine
pranulation of the marginal plates and the relationship
of R/r gives this species a particularly characteristie
appearance, The relationship of this species with
extreme forms of N, oce/lata from the Great Australian
Bight are discussed under that species.
On the basis of a poorly preserved specimen (WAM;
9676, R/r=35/14 mm) from Bald L,, wear Albany, W.A.
in 51.2-64.m, which we recognise as ocellifera, we extend
the range of the species to the south coast of W.A.,
otherwise all known specimens have been collected
berween Geraldton and Cape Naturaliste, W,A., in
depths of 45-180 in. The specimen from Bald |. was
identified by Shepherd (1967) as N. ace//ata,
Distribution: Geraldton ta Bald L., near Albany,
W.AS 45-180 m (Fig. 3a),
Nectria dcellata Perrier
(Figs 2a & 4h-8a)
Asterius ocellifera Lamarck, 1816: 553 (part); 1840: 239
(part).
Nectria oculi/era (lapsus cal. for ocellifera): Gray, 1840:
287.
Nectria ocellifera, Gray, 1866: 15; Sladen, 1889: 319,
pl, 55, figs 1-7; H. L. Clark, 1909; 529.
Nectria ocellata: Perrier, (875: 188; 1876: 4; FL.
Clark, 1916: 34: 1928: 378 (part), 379 (kev); 1938; 78;
1946: 85; Cotion & Godtrey, 1942: 197 (part); A. M
Clark, 1966; 313, 315 (table-part), pl. 2, fig, 3;
Shepherd, 1967: 468 (part), fig, 45 1968: 738 (part);
Marsh, 1976; 217 (table); Zeidler & Shepherd, 1982:
406 (part), fig, 1O4a,
Nectria multispina: Shepherd 1967: 468 (part).
Material Examined: Lectotype (designated herein),
BMNH; 1958: 7.30.20. (R/r—68/23 mm), Tasmania,
purchased E. Gerrard jun. One specimen BMNH, 1953)
4.27.24 (R/r—53-56/16 mm), without data.
In addition 98 specimens tram AM, 38 specimens
from MV, 34 speciiners from TM, 18 specimens from
1 REC, S. AUST. MUS. 19(9): 117-138
SAM, 15 specimens from MCY, 7 specimens from
QVM and 4 specimens. from WAM,
Diagnosis: Ro up to 125 mm, r up to 4§ min,
R/r=2,9740,05 (495% lj, Tabulae with central
granules of similar size and shape, close but usually
Nol campaer, rounded (o angular in cross-section,
rarely lower than peripherals, Peripheral granules of
similar size to each other, usually smaller than centrals,
wedge-shaped, forming more or less compact ring
around centrals, tending to radiate In some specimens.
Tabulae varying fram flat to convex, extending beyond
dise 1a about 24 R. Dorsal arm plates usually convex
sometimes flat, decreasing regularly in size towards arm
lip, peripheral granules becoming indistinct, central
granules often enlarged and prominent giving plates
rough appearance, limits of plates varying from distinct
to indistinct to naked cye. Marginal plates distinct,
granules of similar size ta each other, convex, clase bul
not compact, similar to thase on actinal surface,
Actinal plates up to 6 raws proximally, Furrow spines
2-4 (rarely 5). Subambulacral spines 1-3 on first and
second row. Papulae numerous dorsally extendiag
almost lo arm tips, absent intermarginally and
attinally, Pedicellariae with 3-4 (rarely 2 or 5) usually
clorigate valves (Fig, 2a), common on all surfaces,
frequently replacing subambulacral spines. Colour
uniformly brick red to orange, sometimes. mottled with
lighter and darker coloured tabulae.
Remarks This is the most variable and frequenily
mis-identified species of Necirio. This has been so since
Gray (1840) originally mis-identified the species as
ocellifera Lamarck. Perrier (1875) examined a number
of specimens in the BMNH, all apparently from
northern Tas. and concluded that Gray mis-identified
ocellifera of Lamarck and praposed the name ocellata.
We have examined 3 historical specimens each of which
was possibly examined by Gray and subsequenily
Perrier (A. M. Clark, pers, comm.), of these one
specimen (BMNH; 1846.8.3.14) we identify as WN.
pedicelligera (Fie, Ya) (see p. 000), of the ather twa,
one (BMNH; 1958.7.30.20) has tabulae and tabular
granulation conforming closely to Perrier’s description
and this specitnen we mominale as a lectotype for the
species ocellara (Fig. 4b). We have rejected the second
specimen (BMNH;: 1953.4.27.24) (Fig. 6b) as a possible
lectotype or paralectotype, even though Perrier
probably saw it, as the labujae and tabular granulation
does not conform to Perrier’s description of WN.
oceillaia, and we are therefore uncertain (hal it qualifies
usa Lype specimen. Perrier (1875) also refers to a poor
specimen in the MNHN which he was unable to
determine, This specimen (ECAS 668) which may have
been seen by Lamarck and identified as ove/lifera has
been labelled as a type of ocellata which it clearly is
not as the species description af ocellata was based on
the material in the BMNH.
Following our examination af 218 specimens from
southern and south-eastern Australia we have recog-
June, 1986
mised 3 more or less intergrading forms within the
species ocellata.
The first form conforms most closely with the
lectotype in that the tabulae are slightly convex and
covered with even sized granules (Figs Sa, 6), This form
has its centve of distribution at the eastern and north
coast of Tas. extending north to Port Jackson, N.S.W.
and west to Phillip L. Vie. and is found in very shallow
water [ror the shoreline down ta 20 m (Fig. 3b).
Along the east coast af Australia we recognise a
second form with tabulae with central granules which
become shorter than the peripheral granules, the later
radiating giving the tabula a [aller appearance and
with dorsal arm plates which Frequenily possess
ertlareed central granules giving the arms a rough
texture (Figs Sc, 6a) similar to thal of wilson; from
W.A. This form is disiribuied between Broughton [.,
off Part Stephens, N.SW. and Erith |, Bass Straii, but
has a greater depth range than form | oceurring in 0-90
m (Fig. 3b).
The third form (Figs 6b-d; 7a) has tabulae with
central granules. distinctly shorter than the radiating
peripherals, becoming flattened and gcellifera-like at
the extreme western end of their range (Figs 7b; 8a),
The dorsal arm plates have an even covering of granules
and the limits of the plates are relatively distinct, We
de not identify the specimens from the Great
Australian Bight with oce/lifera because of the elongate
arms and the smaller size and distribution of the
tabulae despite the fact that the marginal granulation
is finer than the actinal granulation. These extreme
variants may paint to a relationship between ocellaia
and acellifere which is not obvious when comparing
ocellifera from W.A, (Fig. 4a) with typical ecellata trom
Tas, (Fig. 5a), The relationship between ecellate atid
ocellifera can only be resolved satisfactorily when more
speeimens fram the Great Australian Bight become
available,
Form 3 extends fram south-eastern Tas. west to the
Great Australian Bight (125 °30'E) occurring in depths
of 5-230 m (Fig, 3b),
Although form 1 clearly intergrades with form 2
along the east coast and with tarm 3 along the south
coast, forms 2 and 3 are easily separable and may have
originated as a result of the Pleistacene land bridge
between Vic, and Tas, separating the twa populations
which then developed their different characteristics.
Along the NSW coast. at least as far South as Eden,
NN. ocellata is the only known species of Nectria so thal
its identity has not been confused, However, along the
south coast the species most often confused with
ocellata is pedicelligeru (Figs 9b; (0a) which most
resembles our form 3 and can most easily be
distinguished trom it by the presence of intermarginal
papulac.
OF the first L3 records of ocellara listed by Clark
(1966, table 3) al least two (BMNH; 1862.7.9.51 and
-52) from W.A. and Dirk Hartog lL. (Shark Bay, W.A.)
REVISION OF THE GENUS NECTRIA ea
respectively are unltkely to be ocel/ata. Further we
doubr the likelihood of Dirk Hartog lL. as a locality
from which Neciria would have been collected,
Shepherd (1967) records a juvenile specimen from
Caloundra, Queensland; we have examined this
specimen and confirm its identity but consider the
locality data to be erroneous.
Distribution, Western end of the Great Australian
Bight (125°30'E) through Bass Strait south slong ihe
eastern coast of Tas, and north along the NSW, vcoast
to Broughton 1; intertidal to 230 m (Fig, 3b).
Necwia macrobrachia H. L, Clark
(Figs 2b & Bb, c)
Nectria mactobrachia WH. L, Clark, 1923: 236, pl, 13,
ligs 5-6; 1928: 379 (key); 1946: 86; A. M. Clark, 1966:
311, pl 3, figs 2-3, text fi, 2; Shepherd, 1967: 474,
fig, 2 (part); 1968: 738; Marsh, 1976: 217 (table);
Zeidler & Shepherd, 1982; 406, fig. 10.4e.
Material Examined: Holotype, BMNH, 1929: 6,121
(R/r=60/17 mm), Pelsart Group, Houtman Abrolhas,
shore, Prof. Dakin-
In addition 32 specimens tram WAM, 23 specimens
from SAM and 20 specimens frant A.M,
Diugnosiss R up to 75 mm, r up to 2] mm,
R/r=3,47+0,08 (495% cl), Tabulae with ceniral
granules irregular in size aud shape, flat, low in profile,
close but not compact, angular in cross-section.
Peripheral granules of similar size to each other,
conspicuously larger than centrals, sometimes radiating
particularly in deep-water specimens (Fig, 8c), generally
wedge-shaped, raised well aboye centrals to form
coneave tabulae. Tabulae low, crowded extending
beyond dise to abuut 44 R. Dorsal arm plates flat,
deereasing regularly in size towards arm tips, peripheral
granules becoming indistinct from centrals, limits of
plaies indistinct to naked eye. Marginal plates relatively
indistinct due to close, ever’ covering of granules,
similar to those on actinal surface. Actinal plates up
to 3 (rarely 4) rows proximally, Furrow spines 2-3,
Subambulacral spines 2-3. in first and second row.
Papulae numerous dorsally extending 1a. arm lips also
present intermarginally and actinally. Pedicellariae with
3-4 broad valves (Fig. 2b) resembling those of NV.
multispina, rarely present, Occurring at least on actinal
and abactinal surface, Colour light yellow to orange,
brown of even pink with peripheral granules on tabulae
lighter and arm tips. darker than rest of body (Shepherd
1967),
Remarks: The torm and granulation of the tabulae
(Figs 8b, c) and the distribution of the papulae easily
separates this species from others in the genus.
Distribution: Houtman Abrolhos, W.A. to Wilson’s
Promontory, Vic. and Brith lL, Bass Strait, but has nor
been recorded from Tas.; intertial lo 180 m (Fig. 3c).
We have been unable to substantiate the depth recard
of 350 m given by Shepherd (1967) which we consider
to be a misprint,
Neetria pecicelligera Mortensen
(Figs 24. 9 & 10a, b)
?Chaetasier munitus, Mobius, 1859: 3, pl, 1, figs 1-2.
Nectria sp. Fisher, 1911: 163; H. L. Clark, 1914: Jf).
Neciria pedicelligera Mortensen, 1925: 291, pl, 13, figs
5-6, text fig, 9H, L; Clark, 1928: 370 (key) A, M.
Clark, 1966: 315 (table); Shepherd, 1967: 478.
Nectria ocellatu: A. L. Clark, 1928: 378, 379 (key); A.
M. Clark, 1966; 315 (table-part), pl. 2, fig. 4;
Shepherd, 1967; 465 (part); Zeidler & Shepherd;
(982: 406 (part); fig. LO.4b,
Nectria multispina: Catton & Godfrey, 1942: 197;
A. M. Clark, 1966: 314 (part); Shepherd, 1967: 468
(part), fig, 3 (part).
Material Examined: Holotype, Otago Museum; A
§3-47 (Rér =53/18 mam), Gisborne, east coast North L.,
New Zealand, H. Suter? (see remarks), One specimen,
BMNH; 1846; 8.3.14 (R/r=55/18 mim), Flinders L,
Tasmanid, Lord Stanley.
In addition 146 specimens irom SAM, 26 specimens
from MV, 25 specimens fram AM, It specimens fram
WAM, 4 specimens from MCZ and 2 specimens fram
TM and QYM.
Diagnosis: R up to 120 mm, r up to 36 mm.
R/r=3.1640.04 (+95% cl). Tabulae with central
granules of similar size and shape, close or compact,
angular in cross-section, larger ihan peripherals.
Peripheral granules of similar size to each other, wedge-
shaped, forming a more or less compact ring around
centrals, rarely radiating. Tabulae slightly convex, rarely
flar, extending beyond disc for up to 4% R, Dorsal arm
plates more or less flat, decreasing regularly in size
towards arm lip, peripheral pranules becoming
indistinct, central granules similar ta peripherals or
enlarged and prominent giving plates rough
appearance, limits of plates indistinct to naked eve.
Marginal plates distinct, granules of similar size ta each
other, convex, close but not compact, coarse, similar
to those on actinal surface. Actinal plates up to 7 rows
proximally, coarsely granulated, Furrow spines 3-4
(sometimes 5, rarely 6). Subambulacral spines 2-3 on
first row and 3-4 (rarely up to 6) on second sow,
Papulae numerous dorsally extending almost to arm
lips, also present intermarginally but rarely actinally.
Pedicellariae with 3-5 (rarely 6) very slender valves
(Fig. 2c), valves particularly slender on acunal surface,
usually common on all surfaces and often replacing
a subambulacral spine on the adambulacral plates.
Colour uniformly brick red to orange, often mottled
with lighter coloured tabulae, especially at the base of
the arms.
Remarks; This is the first time that this species has
been fully recognised since Mortensen (1925) described
it, from Gisborne, New Zealand. However, according
to A. N, Baker (pers. comm,.), despite extensive
collecting eftorts in New Zealand particularly off
Gisborne, no species of Nectria have ever been
{22 REC. 5. AUST. MUS. 19(9): 117-138
recovered. We therefore believe the New Zealand record
to be an error and we recognise it as an endemic
southern Australian species. This species 1s now known
(io range fram) Denmark, WA. ta Eden, N.SW.
including the north coast of Tas, and is a shallow-water
species occurring in 0-20 m,
This species has been contused mast frequently with
ocellata and iiultispina. This.confusion has probably
arisen because previous authors have fot examined the
type of pedicelligera thus oot appreciating it as a
southern Australian species believing it to be endemic
to New Zealand, N, pedicelligera is distinguished from
ocellata by the intermarginal papulae, offen the
presence af a rosette of 6 prominent tabulae at the base
of the arms and the more regular form of the
granulation (Figs 9a, c: 10a), Superficially it resembles
some specimens of mudfispina (Fig. 10b) hawever,
multispina differs in the farm of its granulation, the
enlarged plates extending along the arm, the
consistently higher number af furrow spines and the
much broader valved pedicellariae.
We have examined the specimen from Westernport,
Vic. (MCZ, 1932), described by Fisher (1911) and
examined by Clark (1914) and have identified it as
pedicellizera, The holotype of Chaetaster munitus
Mobius 1859 is presumed lost as 1 1s not in the Kiel
muscum (Shepherd 1967) and was not found by one
of us (FW.E,R.) in the Hamburg museum. However,
the figure given by M@bius (1859) is mast like
pedicelligera, particularly in the arrangement of the
tabulae and their granulation, The apparent absence
of pedicellariae with large valves on the actinal surface
also excludes il from mru/tispina with which Clark
(1966).and Shepherd (1967) thought it was conspecific.
In order to maintain stability and due to the inadequate
description given by MGbius and the loss of type
material we agree with Clark (1966) that the name
Chaetasier mynitus be treated as a nomen oblitum,
such action however requires & case fo be put to the
ICZN (article 23b).
The three specimens (BMNH; (862.1-810 & LH;
1846,8,3,14) recorded by A. M. Clark from Tas, (1966,
table 3) as “iolermediate between ocel/ara and
mnuluispina” we identify as pedicelligera.
Distributions Denmark, WA. to Eden, N.SW,
including Bass Strair and the north-west of Fas.;
intertidal to 20 m (Fig. 3d).
Nectria niultispina H, 1, Clark
(Figs 2d, 1Oe & \la)
Nectria mullispind H, L, Clark, 1928: 375, figs 1a-b;
1938; 77, 1946: 86; Cotton & Godfrey, 1942: 197 (part);
A.M. Clark, 1966; 314, 315 (table-part), pl. 2, figs
1-2; Shepherd, 1967: 468 (part), 1968: 739; Marsh,
1976; 217 (lable); Zeidler & Shepherd, 1982; 406, figs
10.4¢, d.
June, 1986
Material Examined: Holotype, SAM; KS50
(R/r=80-85/30-32 mm)? $.A., probably Spencer Gulf
or Gulf St Vincent, H, L, Clark (1928); 2 paratypes,
SAM; K52 (R/r=56/19, 51/17 mm) without locality
data, J, C. Verco Feb 189); paratype, MCZ 2904
(R/r=60/19 mm) with same data as K52.
fn addition 22 specimens fram SAM, 7 specimens
from AM, 4 specimens from WAM and cme specimen
from MV.
Diagnosis) R up to 102 mm, r up to 34 mm.
R/r=3.01 40.08 (+95% cL). Tabulae with central
granules irregular in size with central ones being larger
thar hase at periphery, Very compact giving central
tabulae smooth convex appearance, angular in cross-
section, all larger than peripherals. Peripheral granules
of similar Size to cach other forming distineL compact
Timg around centrals, Tabulae appear to extend to arm
lips bur from 24 R become indistinguishable tram
convex plates, interspersed with smaller, Matter plates.
Dorsal arm plates with granules. maintaining their
relative sizes but peripherals becoming less distinct than
on disc, limits of plates distinct to naked eye. Marginal
plates distinct, granules often similar in size, sometimes
the peripherals are noticeably smaller, clase, sometimes
compact, similar in size to those on actinal surface.
Actinal plates up to 6 rows proximally, Furrow spines
4-6 (rarely 3 or 7). Subambulacral spines 3 (rarely 44
in first row, 3-8 im second row. Papulae numerous
dorsally, extending to arm Ups, also’ present
infermarginally, absent actinally, Pedicellaraic with 3-4
(rarely 2) distinctive broad valves (Fig. 2d), common
an all surfaces but particularly characteristic of lirst
actinal row of plates. Colour orange to briek red,
matted with darker tabulae, often only some of the
central granules of the tabulae are darker,
Remarks: This is a clearly defined species easily
distinguished by the form of the tabulae, the distinctive
pedicellariae and the consistently high number of
furrow. spines (Figs lOc; ba), Although miuiiisping has
not been confused with orber species it ts clear that
other species have been confused with rmuitispina die
toa lack of understanding of rhe limits ef each of the
species of Nectrid, N. ocellara and N. pedicelligera have
been the species most Irequently confused with
muttispina and this is discussed under thase species,
Of the 9 lots of specimens referred to pultispina by
Clark [1966, table 3) only the type specimens and
specimens under the number BMNH; 1962,4,97 are
multispina.
The species which most resembles and appears most
closely related ta mulrispina is. saoria, particularly in
the arrangement of the tabulae and dorsal arm plates.
However it is clearly distinguished by its smaller dise,
the vansistently fewer furraw spines and the absence
of the charaeteristically large pedivellariac found in
mullispni,
Distribution: Shoalwater Bay, near Cape Peron,
south of Fremantle, W.A. to Wilsan’s Promantory, Vie.
REVISION QF THE GENUS N&CTREA 123
but mot from Bass Strait or Tas., intertidal to 20 m
(Fig, Ja).
Necirid wilsoni Shepherd & Hodgkin
(Figs 2e, 11b & 12)
Neetria wilsant Shepherd & Hadekin, 1965; 119, fig. 1;
Shepherd, 1967: 474; Marsh, 1976: 217 (table);
Zeidler & Shepherd, 1982: 408, fig. 10.4e,
Neerrid ocellata: A, M, Clark, 1966; 315 (table-part),
pl. 1, ligs 4-6; Shepherd, 1967: 465 (part).
Muarerial Examined: Holatype, WAM, 3-65
(R/;=77/26 mm), Sorrento Beach, W.A. 1.8 m, B. R-
Wilson, 27. L1963; paratype, WAM; 18-59 (R/r=63/
20 mm), Eagle Bay, Cape Naturaliste, W.A., 9.2 m, B.
R. Wilson & L, Marsh, 27.X11,1958; paratype, WAM;
19-59 (R/r=75/25 mm), Dunsborough, WLA., 9.2 m,
B. R. Wilson & L. Marsh, 25.X1LI958; paratype,
WAMS 2-62 (R/r=88/31 mm), Hamelin Bay near Cape
Lecuwin, W,A., on jetty piles, B, Ro Wilson & R. Slack-
Smith, 30.X11196); paratype. WAM, 1-65
(R/r=56/19 mm), BDunsboraugh, W.A., 16.9 m on
rock, B. R, Wilson, (5.1V.1963; pararype, WAM; 2-65
(R/r=90/34 mm) Sorrento Reef near Fremantle, W.A.,
On sea grass, B. R. Wilson; paratype, WAM; 4-65
(R/r=90/30 mm) Sorrento Reet, W.A., BR. Wilson,
2LXIL 1963; paratype, WAM, 5-65 (R/t =83/27 mm)
Sorento beach, W.A., 5.5.m on sea grass near
limestone, BR. Wilson, 23.X1L. 1963; paratype, WAM;
3-62 (R/r=120/45 mm), off Beagle Island (29°50'S),
W,A.,, Poole Brothers, Mar. 1961; paratype, SAM; K613
(R/r=—69/22 mm) Hall Bank near Fremantle, W.A.,
B. R. Wilson, 111.1963.
In addition 40 specimens from WAM, 9 specimens
from SAM and 7 specimens. from AM.
Diagnosis’ R up to 120 mm, rc up to 40 mm,
R/r=3,.08+0,06 (+95% cl). Tabulae wirh central
granules irregular in size and shape, a mixture of
granules larger and smaller than peripherals, markedly
convex, raised well above tabula, well spaced (rarely
crowded), round in cross-section, Peripheral granules
rachialing, irregular in size and shape, but forming
distinet peripheral ring. Tabulae more or less confined
to disc, Dorsal arm plates, flat decreasing in size
towards arm tip, with very coarse granulation, | or 2
central granules on each plate often enlarged and
prominent, limits of plates indistinct to naked eye.
Marginal plates distinct, granules coarse, spaced, some
central ones very large similar to darsal arm plates.
Actinal plates up to 7-8 rows proximally. Furrow spines
3-4 (rarely 2). Subambulacral spines 2-3 on first and
second row. Papulae numerous dorsally extending
rarely beyond 3 R, absent inlermarginally and
actinally. Pedicellariae with 4-5 (rarely 3, 6 or 7) slender
valves (Fig 2e), usually present an all surfaces, some
specimens without.
Colour deep orange to magenta, papular area lighter
(Shepherd and Hodgkin 1945).
Remarks: Despite some confusion between this
species and oce//ate the large and irregular granules of
the tabulae, marginals and arm plates particularly make
wilsont a very distinctive species (Fig. Jb). The
granulation of this species is very variable (Pig, 12).
It is interesting to note that some N.SW, specimens of
ocelluta (Fig, 6a) resemble wilson/ in that the granules
become irregular, However, the tabular granulation is
much finer and the granules.are more numerous in the
N.S.W. Specimens so thal these two widely
geographically separated species are not confused. We
agree with Shepherd (1967) that the three specimens
(WAM: 18,59; 19,59 & 2.62) recorded by Clark (1966,
lable 3) as being “alfiliated to ecellafa” are wilsoni,
The species is now extended in range casiward across
the Great Australian Bight to Cape Jervis, S.A,
Distribution Beagle 1, WA, to Cape Jervis, S.A.
intertidal to 44 m (Pig. 31).
Nectria saoria Shepherd
(Figs. 2F & 13a)
Necrria suoria Shepherd, 1967: 475, fig. 2 (part); 1968:
739; Marsh, 1976: 217 (table); Zeidler & Shepherd,
1982; 408, fig, 10,41,
Nectria mullispina: A. M. Clark, 1966: 315 (lable-part).
Material Examined: Holotype, SAM; K670
(R/r=56/15 mm), submerged lirnestone reef between
Wright I. and The Bluff, Encounter Bay, S.A., [0 m,
8, A. Shepherd, 81,1963; 4 pararypes, SAM; K628
(R/r=56/15, 53/16, 50/13, 47/14 mm), collected with
type; 3 paratypes, SAM; K627 (R/r=41/16, 52/15,
40/13 mm), West |., Encounter Bay, S.A., 10m, S. A.
Shepherd, 23.VI11.1964; paratype, SAM; K656
(R/r=65/17 mm) collected with type; paratype, SAM;
K658 (R/r=64/20 mm), limestone reef between Thistle
and Hopkins |, Spencer Gulf, S.A., 10 m., 5S. A.
Shepherd, Jan. 1964; paratype, WAM 8-64 (R/r=60/17
mm), Hamelin Bay near Cape Leeuwin, W.A,, B. R-
Wilson and. R. Slack-Smith, 30_XI1L.61.
In addition 24 specimens from SAM, 23 specimens
trom WAM, 7 specimens from MY and 6 specimens
from AM-_
Diagnosis; R up to 83 mm, r up to 28 mm.
R/r=3.49+0.09 (+95% el.) Tabulae with central
granules similar in Size, Sometimes 1-2 very latge
granules bul these usually mixed with smaller ones,
compact giving tabulae canvex appearance, angular in
cross-section, all larger than peripherals. Peripheral
granules slightly irregular in size forming an irregulat
peripheral ring. Tabulae appear to extend to arm tips
but from about %4 R become indistinguishable from
convex plates; interspersed with smaller plates, Dorsal
arm plates with granulation similar to disc bul more
compact giving the larger distal plates a smoother
aspect, limits of plates distinet to naked eye, Marginal
plates distinct, granules often similar in size to each
other, close, sometimes compact, similar in size to those
124 REC, 5, AUST. MUS. 19(9); 117-138
ou actinal surface. Actinal plates up to 2-3 rows
proximally, Furraw spines 3-4 (rarely 2), Subambulacral
spines 2-3 in first row, 2-4 in second row. Papulae
numerous dorsally, extending to arm lips, less commot
and sometimes absent inlermarginally, absent actinally.
Pedicellariae with 3-5 (rarely 2 or 6) clongate valves
(Fig. 21), common abactinally, less conimnon er absent
iatginally and actinally. Colour bright red, papulac
black (Shepherd 1967),
Remarks; This distinctive species (Pig. 13a) has been
adequately described by Shepherd (1967) and its
relationship to smu/tisping has been diseussed under
that species. Prior to Shepherd’s deseription of this
species it has been identified with zultspinu.
Of the 9 lots of specimens referred to nultispine by
Clark (1966, table 3) (hose fram Rottnest |, Albany and
Recherche Archipelago are sworin as suggested by
Shepherd (1967).
Distribytion; OFT Perth and Rottnest 1, W.A. to
Encounter Bay, S.A. intertidal to 25 mi (Fig. 3g),
Neciria humilis sp.riov.
(Figs 28; I3b & 14)
Material Examined: Holotype, TM; H 1476, 80 km
west of Woolnorth, north-west Tas., 550 m, A.
MeGilford (“Sea Fisheries’’), 28.11,1979; paratype, TM;
H 1844, collected with holotype, 7 paratypes, AM)
J 18467, labelled “80/130 off Storm Bay near Tanvar
River 25-30 fms" (approx, 45-55 m), no other data,
Descriptions The holotype (Fig. 13b) measures
R/t=53-58/17.5 min br =22 mm. Arms taper regularly
to narrow tip. Abactinal tabulae low not more than
J mm high, very slightly convex, longitudinally ovate
ip to 3.52.9 mm, spaced, arranged in a carinal row
wilh two dorsal laleral rows to about 2 R, a third
dorsal row of plates also exiends to abour 2 R, beyond
¥a R the tabulae become indistinguishable from arni
plates, tabulae/arm plates decrease regularly in size
distally. Tabulae with central granules of even size,
convex, low, angular in cross-section, peripheral
eranules marginally Sinaller than centrals forming
compact ring around centrals. Marginal plates distinct,
22-23 supero- and infero-marginal plates with an even
covering of small granules (approx. 7 granules/2 mm),
finer than those on actinal plates. Actinal plates of 6
rows. proximally, the first row extends almost to ann
tip (lo 17th intero-marginal plate) the second, third and
fourth row extend to the Sth, 3rd and 2nd infero-
marginal plares respeerively, the fifth and sixth row do
not extend beyond the Ist inferamargina! plate, Actinal
plates covered with coarse, spaced granules (approx.
§ eranules/2 mm), Adambulacral plates with 3-4
furrow spines, where there are 4 the first one is smallest
otherwise the other 3 are similar in size to each other.
Subambulacral spines, 3 on first row, about half-length
al’ furraw spines; 3 on second raw, barely indistin-
euishable in size from adjacent dctinal granulation,
dune, LON6
Oral plates with 8 furrow-spines, 4 subambulacral
spines. and 7-8 law spines/granules on actinal surlace
of plate. Papulae abundant dorsally in discrete groups
of 810, confined by the skeletal reticulum, absent
intermarginally and actinally, Pedieellariac absent.
Colour unknown (white in alvohol).
The paratype (Fig. 14) (TM: H 1844) measures
R/r=46-51/1719 mn, br=23 mm. Essentially similar
to holotype differing only in having stouter arms,
The seve paratypes (AM; J 18467) range in size
from R/r=66/18 mm to R/r=48/25.5 mm and dilter
from the type only in the presence. of pedicellariae
which occur on the dorsal and actinal surfaces, but are
most common on the adambulacral plates,
PedivcHariae with 3-4 valves, triangular in shape, about
twice as lang as wide (Fig- 2g).
Enunology
From the Latin Givilis=low, referring to the very low
tabtlac,
Renarks: This species stands apart from other
species of Neciria by the granulation and form and
arrangement of jhe dorsal tabulac. The only other
species with such low fabulae is macrobrachia, but in
this species the tabulae are close-packed and
transversely hexayonal or ovate, the granulation is quile
distinctive and the distribution of papulae immediarely
separates the two species. Amongst the species in which
the papulae are restricted to the dorsal surface, /iaiilis
15 most closely related to acel/lafe. However, the tabulae
of eeellata never become as low, or the granulation
cavering the tabulae as. even, as in /rumiliy.
Tt is unfortunate that the data with the 7 paratypes
48 ambiguous as the Tamar River is in the mid-north
of Tas. and the only known Storm Bay is that near the
mouth of the Derwent in south-eastern Tasmania
(Fig. 3h),
CONCLUDING REMARKS
In the most recent revision of the Asteroidea, Neorria
has been allied with Necrriaster in the subfamily
Nectriinge of the family Gonasteridae, However, we
have concluded that skeletal lealures, including
particularly the calcified form of Lhe interbrachial
septum, show that these genera are better placed in the
family Oreasteridae, We have further concluded that
Neciria, by virtue of its abactinal tabulae, stands apart
from all other genera in that faintly, We prefer to
recognise this isolation by maintaining the status af the
subfumily Nectriinge from which Neerrinster is
excluded. All other genera within the tamily
Greasteridac are referred to the new subfamily
Oreasterinae
Within the genus Neciria we have found the presence
or absence of intermarvinal papulae to bea very useful
and consistent character Jor all of the species except
seoria, This character in combination with the form
of abactinal and marginal granulation and the form
REVISION OF THE GENUS NECTRIA 125
of the pedicellariae have enabled us to determine the
limits of the species.
We have determined that Necfria is an endemic
southern Australian genus and that records from Fiji
(Sladen 1889), New Zealand (Mortensen 1925) and
Mauritius (MV specimen) are erroneous.
Regarding the origin of the genus Necfria in
Australia, lack of fossil material prevents us from
determining its evolutionary or temporal origin. From
our knowledge of the species distributions we would
infer that the genus arose in south-western Australia
and distributed along the south coast; this is consistent
with our present knowledge of major ocean currents
in southern Australia. However, only one species, N.
ocellata, has been successful in reaching the N.S.W.
coast. Considering our interpretation of the complexity
of this species we conclude that the full potential for
speciation in the genus Nectria has not yet been
realised.
ACKNOWLEDGMENTS
We would like to thank Miss A. M. Clark, British
Museum (Natural History), London; Miss A, Green
and Mrs E. Turner, the Tasmanian Museum and Art
Gallery, Hobart; Dr A. Guille, Museum National
d’Histoire Naturelle, Paris; Dr A, C. Harris, Otago
Museum, Dunedin, New Zealand; Mrs L. M. Marsh,
Western Australian Museum, Perth and Dr R.
Woollacott, Museum of Comparative Zoology,
Harvard, U.S.A. for the loan of material. Collections
have also been examined in the Museum of Victoria
and the Queen Victoria Museum and Art Gallery,
Launceston by the authors access to which was given
by Sue Boyd and Bob Green who are respectively
thanked.
Miss A. M. Clark and Mrs L. M. Marsh also kindly
reviewed the manuscript and made helpful suggestions.
The photographs were taken by Miss H. McClelland
of The Australian Museum, who is duly acknowledged.
Ms J. M. Thurmer of the South Australian Museum
prepared Fig. 3. We are especially grateful to Dr D.
Kuhlmann, Museum fur Naturkunde der Humboldt—
Universitat, Berlin, DDR, for supplying us with copies
of Oudart’s figure.
The Mark Mitchell Research Foundation is acknow-
ledged for granting financial assistance to one of us
(W.Z.) for travel to Sydney to complete the study.
REFERENCES
CLARK, A. M. 1963. A note on Patiria ocellifera Gray, 1847
(Asteroidea), Doriana 3: 1-9, 4 pls.
CLARK, A. M. 1966, Port Phillip Survey 1957-1963. Echinodermata.
Mem. Nat. Mus. Vic. 27; 289-355, pls i-iv.
CLARK, H. L. 1906. Echinodermata. Sci. Res. “Thetis”. Mem. Aust.
Mus. 4: 519-564, pls 47-58.
CLARK, H. L, 1914, The echinoderms of the Western Australian
Museum. Rec. W. Aust. Mus. 1: 132-173, 1 fig., pls xvii-xxvi.
CLARK, H. L. 1916. Report on the sea-lilies, starfishes, brittlestars,
sea-urchins obtained by the F.1.S, ‘Endeavour’ on the coasts of
Queensland, N.SW., Tasmania, Victoria, S. Australia, W.
Australia, Endeavour Res. 4: 1-123, 11 figs, 44 pls.
CLARK, H. L. 1923. Some echinoderms from West Australia. J.
Finn. Soc., Zool. 35: 229-251, pl. 13.
CLARK, H. L. 1928. The sea-lilies, sea-stars, brittle-stars and sea-
urchins of the South Australian Museum. Rec. S. Aust. Mus.
3: 361-382, figs 108-142.
CLARK, H. L. 1938. Echinoderms from Australia, Mem. Mus. comp.
Zool. Harvard, 55: 1-596, 63 figs, 28 pls.
CLARK, H. L. 1946, The echinoderm fauna of Australia, Publ.
Carneg. Instn, 566: 1-567.
COTTON, B. C. and GODFREY, F. K. 1942. Echinodermata of the
Flindersian Region, southern Australia. Rec. S. Aust. Mus. 7:
193-234, pl. 12.
DUJARDIN, M, F. and HUPE, M. H. 1862. Histoire Naturelle des
Zoophytes. Echinodermes. Paris. 628 pp., 10 pls.
FISHER, W. kK. 1911. Asteroidea of the North Pacific and adjacent
waters. Part 1. Phanerozonia and Spinulosa. Bu/l. US. Nat. Mus.
76: 1-419, 122 pls.
GRAY, J. E. 1840 (1841). A synopsis of the genera and species of
the class Hypostoma. Ann. Mag. nat. Hist, 6: 275-290.
GRAY, J. E. 1847a. Descriptions of some new genera and species
of Asteriadae. Proc, Zool. Soc. Lond. 1847; 72-83.
GRAY, J. E. 1847b. The description of some new species of starfishes,
Asteriadae. /n Jukes J. B. Narrative of the surveying voyage of
H.M.S. ‘Fly’; commanded by Captain F. R. Blackwood, R.N.
in Torres Strait, New Guinea, and other islands of the Eastern
Archipelago during the years 1842-1846, together with an
excursion into the interior of the eastern part of Java. Vol. II.
London, pp. 339-354,
GRAY, J. E. 1866. Synopsis of the species of starfish in the British
Museum (with figures of some of the new species), London.
iv+17 pp., 16 pls.
LAMARCK, J. B. P. 1816. Histoire naturelle des animaux sans
vertebres. Ed 1, Vol. 2. Paris. 568 pp.
LAMARCK, J. B. P. 1840. Histoire naturelle des animaux sans
vertebres. Ed. 2, Vol. 3 (reviewed and augmented by G. P.
Deshayes and H. Milne-Edwards) Paris. 770 pp.
MARSH, L. M. 1976. Western Australian Asteroidea since H. L.
, Clark. Thalassica Jugoslavica 12: 213-225, 2 figs, | table.
MOBIUS, K. 1859. Neue seesterne des Hamburger und Kieler
Museums. Hamburg. 14 pp. 4 pls.
MORTENSEN, T. 1925. Echinoderms from New Zealand and the
Auckland-Campbell Islands, II]. Asteroidea, Holothurioidea,
and Crinoidea. Vidensk. Medd. naturh. Foren. Kbh, 79: 263-420,
, 70 figs, 3 pls.
MULLER, J. and TROSCHEL, F. H. 1842. System der Asteriden.
Braunschweig. xx+135 pp., 12 pls.
OUDART, M. P. 1827. Cours d'Histoire Naturelle contenant les
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PERRIER, E. 1869. Reserches sur les pedicellaires et les ambulacres
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PERRIER, E. 1875. Revision de la collection de stellerides du
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PERRIER, E. 1876. Revision de la collection de stellerides du
Museum d'Histoire naturelle de Paris. Arch, Zool. exp. gen. 5:
1-104, 209-304.
PERRIER, E. 1894. Stellerides. Exped scient. Travailleur-Talisman,
1-431, 26 pls,
SHEPHERD, S. A. 1967. A review of the starfish genus Nectria
(Asteroidea: Goniasteridae). Rec. S. Aust. Mus. 15: 463-482,
4 figs, 1 table.
SHEPHERD, S. A. 1968. The shallow-water echinoderm fauna of
South Australia. 1. The asteroids. Rec. S. Aust. Mus. 15: 729-756,
1 fig., 5 tables.
SHEPHERD, S. A. and HODGKIN, E, P, 1965. A new species of
Nectria (Asteroidea: Goniasteridae) from Western Australia. J.
Roy, Soc. West. Aust. 48: 119-121, 1 fig.
SLADEN, W. P, 1889, Asteroidea, Rep, Sci. Res. “Challenger’} (Zool)
30: 1-935, 118 pls.
SPENCER, W. K. and WRIGHT, C. W. 1966. Asterozoans. /n
Moore, R. C. (ed.). Treatise on Palaeontology, Part Il.
Echinodermata 3. Vol. 1. Geological Soc. America Inc. and
University of Kansas Press: pp. U4-U107, figs 1-89.
VIGUIER, C. 1878. Anatomie compare du squellete des stellerides.
Arch, Zool. exp. gen. 6: 33-250, pls. 5-16.
ZEIDLER, W. and SHEPHERD, S. A. 1982. Sea stars (Class
Asteroidea), /a (Shepherd, S. A. and Thomas, I, M,, eds).
“Marine Invertebrates of Southern Australia? Part 1, Adelaide,
S.A. pp. 400-418, figs 10-3-10-9, pl. 30-2.
126 REC, 8, AUST. MUS. 19(9): 117-138 June, 1986
e bh ra be f On ellejer €
’ Usterva cvelli7 Pra
Cry a yaw On r/ tu ne
FIG. 1. Oudart’s (1827) figure of N. ocellifera Lamarck.
FIG. 2. Pedicellariae from (a) N. ocellata, R/r= 67/22 mm, West L., Encounter Bay, S.A., (SAM; K 1754); (b) N. macrobrachia, R/r= 69/19 mim,
Fisherman |., Green Head, W.A. (WAM; 493-79); (c) N, pedicelligera, R/r=69/21 mm, Wright I., Encounter Bay, S.A, (SAM; K 663);
(d) N. multispina, R/r=70/25 mm, Smooth L., Nuyts Archipelago, S.A, (SAM; K 1755); (e¢) N. wilsoni, R/t=65/18 mm, St Francis L.,
Nuyts Archipelago, S.A. (SAM; K 1756); (f) N saoria, R/r=68/19 mm, St Francis 1., Nuyts Archipelago, S.A. (SAM; K 1757); (g) N.
Aumilis, R/r=64/18 mm, off Storm Bay/Tamar R., Tas,, (Paratype, AM; J 18467).
REVISION OF THE GENUS NECTRIA 127
Dirk Hartog Ig.
Caloundra
od
Port Stephens
Naturaliste
Adelaide
f ——
Melbourne |.
Basse Strait
N. ocellifera
TAS, \H Hobart
Houtman
Abrolthos
Denmark
Wilsons Prom.
NM. macrobrachia JY Erith Is N. pedicelligera
Shoalwater|Bay Beagle ls.
my
Cape Jervis
Wilsons Prom,
N. multispina N. witsoni
Rottnest Is.
Fremantle
Encounter Bay
N. saorla N. humilis Wootnorth- Tamar River
‘Storm Bay
FIG. 3. Maps showing distribution of Necrria species.
REC. S. AUST. MUS. 19(9): 117-138 June, 1986
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FIG. 4. (a) N. ocellifera, R/r=40/15 mm, “Diamantina” Stn. 68 (30°37'S; 114°44’E), 139-146 m. (WAM; 1375-74); (b) N. ocellata,
R/r=68/23 mm, Tas. (Lectotype, BMNH; 1958: 7.30.
REVISION OF THE GENUS NECTRIA
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FIG. 5. N. ocellata (a) Form 1, R/r=123/40 mm, off Maria I., Tas. (TM; H 1293); (b) Form 1, R/r
Vic.,
REC, S. AUST. MUS. 19(9): 117-138 June, 1986
FIG. 6. N. ocellata (a) Form 2, R/r=97/31 mm, off Norah Head, N.SW_, 50-70 m (AM; J 3466); (b) Form 3, R/r=56/16 mm, without
data (B\INH; 1953: 4.27.24); (c) Form 3, R/r=87/29 mm, Westernport, North Arm Channel, Vic. (AM; J 9920); (d) Form 3,
R/r=80/27 mm, Great Taylors Bay, Bruny I. Tas. (TM; H 450).
REVISION OF THE GENUS NECTRIA
FIG, 7, N. ocellata, Form 3 (a) R/r= 78/25 mm, Cape Jervis,
146-230
REC. S. AUST. MUS. 19(9): 117-138 June, 1986
FIG. 8. (a)N. ocellata, Form 3, R/r =85/28 mm, Great Australian Bight, 146-230 m (AM; E 3645); (b) N. macrobrachia, R/r=69/18 mm,
Fisherman I., Green Head, W.A., 21 m (WAM; 493-79); (c) N. macrobrachia, R/r=57/16, off Port Gregory, W.A., 110 m (WAM; 27-71).
REVISION OF THE GENUS NECTRIA
FIG. 9. N. pedicelligera (a) R/r=55/18 mm, Flinders I., Tas. (BMNH; 1846: 8.3.14); (b) R/r=85/28 mm, Cape Jervis, S.A., 10 m (SAM;
K 596); (c) R/r=85/25 mm, Doubtful Islands Bay, W.A., 1-2 m (WAM; 367-75).
REC. S. AUST. MUS. 19(9): 117-138 June, 1986
FIG. 10. (a) N. pedicelligera, R/r=49/15, West I., Encounter Bay, S.A., 4-5 m ( J 7554); (b) N. pedicelligera, R/r=72/22 mm, Stanley
1., Denmark, W.A., 12-23 m (AM; J 12709); (c) NM. mudltispina, R/r=80/28 m Shoreham near Melbourne, Vic. (WAM; 1399-74).
REVISION OF THE GENUS NECTRIA
/33 mm, Sorrento Reef, Perth, W.A., 4-5 m (AM; J 7834).
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94/32 mm, Sorrento Reef, Perth, W.A. 4:5 m
; 2-65); (e) R/r=64/18, off Dunsborough, Geographe Bay, W.A., 20-30 m (WAM; 746-76).
-62); (d) R
117-138
REC. S. AUST. MUS. 19(9)
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REVISION OF E GENUS NECTRIA
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——
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7-5 mm, 80 km W, of
=74/21 mm, off Rottnest [., W.A., 25 m (WAM; 33-71) (b) is, R/r=53-58/1
as., 550 m (Holotype, TM; H 1476).
_ la) N, saoria, R
Woolnorth,
138 REC. S, AUST. MUS. 19(9): 117-138 June, 1986
FIG. 14. N. humilis, R/r=46-51/17-19 mm, collected with holotype (Paratype, TM; 1844).
REDESCRIPTION OF THE MYSID CRUSTACEAN, NOTOMYSIS
AUSTRALIENSIS (TATTERSALL) COMB. NOV.: REPRESENTATIVE OF A
NEW GENUS
BY KARL J. WITTMANN
Summary
Distinctive features of the mouth-parts and ventral setae on the telson necessitate the establishment
of a new genus, Notomysis, for Leptomysis australiensis Tattersall.
REDESCRIPTION OF THE MYSID CRUSTACEAN, NOTOM YSIS AUSTRALIENSIS (TATTERSALL}
COMB, NOV.;: REPRESENTATIVE OF A NEW GENUS
by
KARL, J. WITTMANN
Institut fdr allgemeine Biologie der Universitat Wien, SchwarzSpanierstr, 17, A-1090 Wien, Austria
(Manuscript accepted 30 April 1985)
ABSTRACT
WITTMANN, K J, 1986. Redescription of the mysid crustacean,
Notomysis ausiraliensis (Tattersall) comb, nav: representative
of a mew genus. Ree. S. Aust. Mus. 19010): 139.143,
Distinctive features of the mouth-parts and ventral
selae on the telson necessitate the establishment of a
new genus, Notomysis, for Lepromysis australiensis
‘Tattersall,
INTRODUCTION
While preparing a revision of the genus Lepronmiysis
it became obvious that L. australiensis Tattersall has
several characters which are not Found in any other
known species. Tattersall's (1927) description was based
on material collected by H. M. Hale in late 1926 in Gulf
St Vincent, South Australia, No other Leptomypsis
Species has been found in Australian waters since then,
In his description Tattersall states; “This is a character-
istic species of the genus .. 7’; however, he gives no
description of the mandibles and maxillag which are
crucial taxonomic characters al the species and genus
level,
The doubtful status of the species made it necessary
to examine the type material which ts kept at the South
Australian Museum (Reg, No, C, 1617), The material,
labelled as syntypes, comprises five adult and three
subadult females, seven adult males, and several parts
of animals. A male with a body length of 9 mm was
chosen as lectotype, dissected completely, and mounted
in Swan medium on slides. Seven further specimens
collected by H, M. Hale in 1925 at the same location,
not labelled as synlypes, are in the Tattersall collection,
Briiish Museum (Natural History) (Reg. Nos 1964,1.21;
2257-2275). The examination revealed that the
description given by Tattersall (1927) is somewhat
incomplete; also there are Several conflicting points
cancerning features common to all specimens studied:
the eyes are much smaller; the exapod of the fourth
male pleopod has four instead of five large modified
setac; the dactylus. of the third to cighth thoracic
endopods is much smaller than described; and the
telson bears setae and has a minute apical incision,
These findings show that a redeseription is necessary.
From the following deseription of mouth-parts and
telson it becomes clear that the species cannot remain
in the genus Lepfomysis but should be placed in a new
genus,
Notomysis noy. gen.
Diagnosis: Mysidae, Leptomysini with eyes normal.
In addition to the usual sexual dimorphism of the
antennula the males are characterized by a hairy
rounded organ located dorsally at the distal segment
of the syrmpod. Antennal scale sétase all around, with
small terminal joint. Labrum normal, without
spiniform process. Mandible with processus molaris
reduced, palpus normal. Maxillula with endite small,
distal joint slender, Maxilla with exopad normal,
endopod without spines, proximal endite as broad as
other 2 endites combined. First thoracic endopod
without endites, ischium distinet but very short. Third
to eighth thoracic endopod with carpopropodus
3-segmented by transverse articulations. Female with 3
pairs of oostegites. Pleopods of female represented by
simple setose plates. Pleopods of male all biramous,
well segmented except first endopod; fourth exopod
with a total of 4 large modified setae on distal 3
segments. Endopod of uropod with spines on inner
margin. Exopod without distal joint and without
spines. Telson with numerous setae ventrally, minute
apical incision, and spines on lateral margins,
Tvpe-species: Lepromysis australiensis Tattersall.
Etymology: From the Greek term ‘ndtos' (= south),
Relationships; The genus Noromysis is closely allied
to the genus Promysis, sensu Ti (1964), with which it
shares features of the antennae, mouth-parts,
thoracopods, and pleopods. Maxilla, maxillula,
mandible, and first maxilliped are unlike Leptomysis
but clearly belong to the type represented by the genera
Promysis and Prionamysis, The new genus is
distinguished from these and all other genera of the
family Mysidae by the unique structure of the telson,
Notomysis australiensis (Tattersall) comb. nov.
(Figs 1-26)
Diagnosis: Middle segment of antennular peduncle
with strong modified seta directed laterad. Inner
margin of endopod of uropod with [6-28 spines
increasing in length distally; 11-22 of these spines
irregularly arranged on statocyst; 5-7 spines linearly
arranged distal to statocyst. Telson with about 45-60
closely set spines on each lateral margin; apex with
minute narrow incision about i, length of telson.
Ventrally telson bears 8-12 plumose setae on
140
longitudinal ridge running half-way between centre and
tip.
Description: General body proportions slender,
closely similar to Prionamysis aspera li, A farther
coincidence with this species is that body and
appendages are densely covered with small cuticle
structures (Fig, 5). These are of about cylindrical shape
on the carapace, body trunk, eye-stalks, and bases of
appendages. Towards tips of appendages they tend to
become acute and scale-like (Figs 3, 14, 19, 21, 23).
Carapace with a transverse straight row of ca 13 pores
in median position a short distance anterior to cervical
sulcus; a further row of ca 27 pores at the cardial sulcus
and a larger pore surrounded by 7 smaller ones a short
distance anterior to the rounded posterior margin of
carapace, Rostrum large, longer than distal segment of
the antennular peduncle. Eyes as in Fig, }. Last
abdominal somite terminally produced into 2 strong
Spiniform projections on each Jateral margin.
Mouth-parts: Frontal border of labrum bluntly
rounded. Mandibles with processus molaris reduced to
two lobes, masticatory lamellae completely lacking.
Teeth of pars incisivus and lacinia mobilis larger on
left than on right mandible. Left pars incisivus with
3 large and 3-4 smaller teeth, with the size decreasing
proximally, Right pars incisivus distally with one large
tooth flanked on each side, rostrally and caudally, by
one masticatory plate carrying several small teeth, Left
lacinia mobilis with 5-6, right one with about 4 teeth
of varying size, Each pars centralis (=spine row) with
4-5 teeth of about equal size. Left mandible only
densely covered with bristles on caudal face in region
of lacinia mobilis and pars centralis, Palpus with
unusually dense setation, Maxillula as is typical of
Mysidopsis and Promysis, distal joint without array of
pores found in Leptoniysis. Maxilla essentially as
figured by Ii (1964) for Promysis orientalis Dana,
Thoracopods: First exopod with 9 segments, second
to seventh with LO, eighth with 9-10, Proximal segment
flanked by large intersegmental joints; acutely pointed
at laterodistal corner; small plumose seta usually
present close to this corner. Epipodite | linguiform,
without seta, First and second endopod like those in
Pramysis. Merus of first endopod without array of
pores that occurs in Lepromysis; propodus and dactylus
without spines except a strong nail al apex of dactylus.
REC. 3, AUST. MUS. 19(10): 139-143
June, 1986
Carpopropodus of third to eighth endopod with 3 short
joints; dactylus minute with slender nail. First pair of
oostegites reduced, but larger than usual in Mysidac,
with same basic setation patiern as in second pair (Figs
17, 18). Penis short, with about 11 smooth, curved setae
around ejaculatory opening, and a series of plumose
setae along outer margin,
Pleapods: In females all 5 pairs represented by simple
setose rods increasing in length caudally, lm sub-basal
position each bears a more or Jess indistinct apophysis
directed laterad. All features of male pleopods
essentially as in Promysis orientalis, sensu Vi (1964).
First to fifth endopod with 1, 9, 9, 9, and 8-9 joints,
respectively, Basal joints each with a well-developed
apophysis which is slender in first endopod, but
rounded, plate-like in second to fifth, Basal segment
of fifth endopod with small additional apophysis. First
to fifth exopod with 8, 9, 10, 11-12, and 9-10 joints,
respectively. Fourth exopod with large modified setae
on last 3 joints. Antepenultimate and penultimate
segment each with one large’ modified seta and an
additional small smooth seta. Distal segment minute,
with 2 less powerful modified setae.
Uropods: Bxopod setose all around, |.6 times as long
as telson, or 1,5 limes endopod. Endopod with
spiniform projection dorsally at statocyst. Distance
between apical spine al inner margin and tip is 25-30%
length of endopod.
Nauplioid stage: Antennula distally with acute scales
arranged in comb-like units, Antenna and mandible
smooth. Abdomen covered with small hairs.
Length: Measured from rostrum to tip of telson
excluding spines. Length is 8-13 mm in adult females
(n=8) and 7-11 mm in adult males (n=9), Mean egg
diameter is 0.49 mm (n=4),
ACKNOWLEDGMENTS
lam greatly indebted to Dr Joan Ellis (London) and
Mr Peter Aerteldi (Adelaide) far the trouble they have
taken to send the material to Vienna.
REFERENCES
Hl, N,, 1964, Fauna Japonica, Mysidae. 610 pp, Biogeographical
Sociery of Japan, Tokyo,
TATTERSALL, W. M,, 1927. Australian apossanmr shirinipis
(Mysidacea).. Ree. S. Aust Mus. 3: 235-257.
THE MYSID CRUSTACEAN, NOTOM YSIS AUSTRALIENSIS 14
FIGS J-9. Nofomysis australiensis (Vattersall), Lectotype, ¢ 9 mm. 1, Anterior body region, dorsal view. 2. Antenna, ventral view. 3,
Antennula, dorsal view. 4. Right mandible, caudal view. 5. Examples of cuticle structures, schematically. 6. Labrum, ventral view. 7.
Manxillula, caudal view. 8, Maxilla, caudal view, 9. Masticalory margins of mandibles, caudal view; details show dentation of pars incisivits,
lacinia mobilis, and pars centralis of left mandible.
142 REC. S. AUST. MUS, 19(10); 139-143 June, 1986
a
FIGS 10-16. Netoutysis ausiratiensiy (Tattersall), Lectotype, ¢
thoracic appendage, rostral view, 12. Distal portion af second thoracic appendage, setae omitted, 13-15, First, fourth, and lifth pleopod,
caudal view. 16. Penis.
Qmn, 1) First Tharacie sternite with appendage, caudal view, If, Second
THE MYSID CRUSTACEAN, NOTOMYSIS AUSTRALIENSIS 143
O.2 mm 17-22
FIGS 17-26. Notompsis australiensis (Tattersall). 17-20. ¥ 9 mm. 17. Second oostegite, inner face. 18. First oostegite, inner face. 19. Filth
pleopod, rostral view. 20. First pleopod, rostral view. 21, 22. Lectotype, ¢ 9 mm. 21. Uropods, ventral View. 22. Telson, ventral view,
23. d¢ & mm, ‘tarsus’ of third thoracic endopod, detail shows dactylus with nail, 24-26, Nauplioid larva, lateral views. 24. Distal portion
of abdomen. 25. Nauplius appendages. 26. Distal portion of antennula.
RECORDS oF THE
SOUTH AUSTRALIAN
MUSEUM
VOLUME 19 NUMBERS 11-14 MAY 1986
No
No.
No.
No.
1
12
13
14
ON TROMBELLA ALPHA N.SP. (ACARINA:
TROMBELLIDAE) FROM AUSTRALIA: CORRELATION,
DESCRIPTION, DEVELOPMENTAL ABNORMALITIES,
SYSTEMATICS AND POSSIBLE AUDITORY STRUCTURES
by R. V. SOUTHCOTT
THE GENUS ODONTACARUS (ACARINA:
TROMBICULIDAE). Il. OBSERVATIONS ON THE LIFE
HISTORY AND MORPHOLOGY OF ODONTACARUS
SWANI N. SP, AND RELATED FORMS
by R. V. SOUTHCOTT
HISTORY OF THE DISCOVERY OF SPELEOGNATHUS
AUSTRALIS WOMERSLEY (ACARINA: TROMBIDIFORMES),
WITH NOTES ON ITS NATURAL HISTORY AND BEHAVIOUR
by R. V. SOUTHCOTT
DESCRIPTION OF ODONTACARUS VEITCHI SP. NOV.
(ACARINA: TROMBICULIDAE)
by R. V. SOUTHCOTT
SOUTH AUSTRALIAN MUSEUM
North Terrace, Adelaide
South Australia 5000
ON TROMBELLA ALPHA N.SP. (ACARINA: TROMBELLIDAE)
FROM AUSTRALIA: CORRELATION, DESCRIPTION,
DEVELOPMENTAL ABNORMALITIES, SYSTEMATICS
AND POSSIBLE AUDITORY STRUCTURES
BY R. V. SOUTHCOTT
Summary
A trombidoid mite (Acarina: Trombellidae) larva, parasitic upon field crickets Teleogryllus
commodus (Walker) in northern New South Wales, has been reared experimentally to the active
deutonymph. The larva, protonymph and deutonymph are described as Trombella alpha n.sp., and
the deutonymph compared with previously known species of the genus, particularly T. warregensis
Hirst and T. adelaideae Womersley from Australia.
ON TROMBELLA ALPHA NSP. (ACARINA: TROMBELLIDAE) FROM AUSTRALIA;
CORRELATION, DESCRIPTION, DEVELOPMENTAL ABNORMALITIES, SYSTEMATICS ANIt
POSSIBLE AUDITORY STRUCTURES
by
R. V. SOUTHCOTT
Honorary Researeli Associate, South Australian Museum, North Terrace, Adelarde, South Australia SUUU
(Manuscript accepted 23 January 1985)
ABSTRACT
SOQUTHCOTT, KR. VY. 1986. On Trompbetlu uiphu osp. (Acarina:
Trambellidae): Gorrelation, description, developmental
abnormalities, systematics anid possible auditory structures, Ree,
S. Aust. Mus, W(11); 145-168.
A trombidoid mite (Acarina: Tronmbellidac) larva,
parasitic upon field crivkets 7e/eogrvilus commodus
(Walker) in northern New South Wales, has been reared
experimentally to the active deulonymph. The larva,
protonymph and deutonymph are described as
Trombella alpha nsp., and the deutonymph compared
With previously kiown species of the gents,
particularly 7. warregensis Hirst and 7. adelaideae
Womersley from Australia,
One reared nymph of T alpha had gross
developmental abnormalities of the legs, with
shortening or even loss of segments, and non-
development of setae, here termed brachymely. Similar
oecurrenees in this and other mites are instanced,
A correlation ard a principal component analysis
was made of The larval dimensional variates, using
various clorsal scutum, leg segmental, and body sctac
measurements. Three principal components were
delined, each including one or more idiosomal variates
and one or more leg variates.
A key is presented for the known adults and
deulonymphs of Trombella of the world,
Leg chaetotaxy of Trombella alpha, the only
deseribed larva of Trombella, is discussed.
The family Trombellidae and its subfamilies
Chyzeriinae and Trombellinae are redefined, Generic
classification of the Trombelhdae is revised, A new
genus jis erected for adults and nymphs:
Maiputrombella yn, gen. with type species Mf.
americanum (Robaux, 1968) its only member, from
South America, Keys are provided to the genera of both
larvae and of adults and deutonymphs, HWomerslevia
Radford, 1946 is restored to full generic status,
The adult genus Perachyzeria Hirst is cemoved from
the family and placed in the Johnstonianidae, as is also
the larval genus Ru/phandyng Vercammen-Crandjean
et al. (1974).
The possible functions of some of the structural
peculiarities of 7rombella alpha \arva are discussed in
relation tO existing knowledge and suggestions that
have been made for some other prastigmatic mites, The
Hine, 1986
attenuation of the larval tarsi is believed to be related
toils function as a sound receptor, serving as a sunar
device for the location of the sound-emitting cricket
hosts, [his suggestion is supported by a significant bias
(over 3:1) in the numbers of mites obtained from
vocalizing male crickets as against the suent females
and because this bias was nol found in the sub-adult
(ic. non-stridulating) male instars.
INTRODUCTION
Berlese (1887) proposed the genus Trombella for F
glandulosa Berlese, 1887 lrora northern Ltaly; a species
since recorded from Austria (Schuster, 1960). Originally
Berlese gave a definition and description only of the
type species, ie, by referring to the sixteen dorsal
depressions. In 1888 he added the South American 7;
nothroides Berl, 1888, which lacks the rounded dorsal
opisthosomal depressions.
Woinersley (1954b) reviewed the subfamily
Trombellinae Thor, 1935 and erected the genus
Nothrotrambidium lor Trombella otiorum Berlese,
1902, from Europe (see Feider 1955, p. 68; 1958, p. 265),
thereby restricting Berlese’s genus; he placed if) it also
T. nothroides Berlese, and T. lundbladi Willmann, 1939
from Madeira (North Atlantic region). Additional
species include Nothrotrombidium bulbiferum
(Willmann, 1940) (nom, enend.), from Europe and
Nothrotrombidium brevilarsumn André, 1960 (vom,
emend.) (1960a) [rom Tonkin, Indochina, Tronrbella-s, |
sth, has currently five described species: T glandulosa
Berlese (ype), T warregensis Hirst, 1929 (Australia),
T. adelaideae Womersicy, 1939 (Australia), 7. /avosa
André, 1936 (1936a) (Africa) and 7) /usianica Andre,
1944 (Europe),
Genera now placed in the family Trombellidac
include Trombella Berlese, Puruchyzeria Hirst, 1926
(however, see lurther below), Para/hrambella Andre,
1958, Neonothrothromiidium Robaux, 1968, based
purely on the adult forms, CAyzeria Canestrini, 1897
and Nethrotrombidiinm Womersley, 1954, known from
the adult and the larva, dudyvana Womersiey, 1954
(195da) und Durenia Vercammen-Grandjean, 1955,
known from the larva and deutonymph, and
Womerslevia Radlord, 1946, Ra/phutudyna Vereammen-
Grandjean et wl, 974, and Nothrotrombicula
Dumbleton, 1947, known only from the larva.
146
I reported that larval trombidioid mites, parasitic
upon field crickets, 7é/eogryllus commodus (Walker),
in northern New South Wales, and classifiable to
Womersleyia Radford, had been reared experimentally
to the nymphal stage, and that these nymphs were
classifiable as Trombella (Southcott, 1982), It was thus
proposed that Womersleyia Radford is a junior
synonym of 7rombella Berlese. Previously it was known
only from its type species, W. minuta Radford, 1946,
a larva taken either free-living from mud, or parasitic
under the wings of grasshoppers (Acridoidea:
Tetrigidae and Tettigoniidae) on the Island of Gan,
Addu Attoll, Maldive Islands; original specimens taken
on 20.xii.1944. (A paratype slide in the South Australian
Museum collection is labelled (in the writing of one
of Womersley’s technical assistants) “ex Grasshopper/
Maldive Is/13. Jan 1945 C.D.R.” This slide bears also
a label in pencil showing that it was used by
Vercammen-Grandjean in 1970 in his revision (1972)
of the species. | have added an identifying number
ACB731 to each label.)
This paper describes the larva, protonymph and
deutonymph instars of the New South Wales species
of Trombella, details of the experimental correlation
and its taxonomic significance.
The reared deutonymphs appeared to be
morphologically similar to the adult Trombella
adelaideae Womersley, 1939, known from a single
specimen collected. “from under a stone at Burnside”,
in the Adelaide district, South Australia, in August.
However, there are differences. One other Australian
species is known, based also on a single specimen; this
is T. warregensis Hirst, 1929, found “under a log on
the bank” of the River Warrego, 4 miles west of
Barringun, New South Wales, in August 1928 by Hirst.
The total number of species of Trombella in Australia
is conjectural, as these mites have been collected and
surveyed very inadequately. I have seen several as yet
undescribed species of larvae referable to Trombella s.
str. parasitic upon grasshoppers, whilst surveying the
mite ectoparasites of grasshoppers in the Australian
National Insect Collection, CSIRO Division of
Entomology, Canberra. In view of the uncertainties of
attempting to correlate deutonymphal and adult
trombidioid (and other prostigmatic) mites on
morphological features, it is considered wisest to erect
a new taxon, 7rombella alpha n. sp., for this species.
After again studying the characters of 7rombella
alpha and other larvae known in the family,
Womersleyia is restored to full generic status (see
further below).
All measurements are given in micrometres (um)
unless otherwise stated.
Genus Trombella Berlese
Synonymy
Note: I have not attempted in the synonymic list for
Trombella below to define where it covered also
REC, S. AUST. MUS. 19(11): 145-168
June, 1986
Nothrotrombidium Womersley, 1954. All authors used
the genus name of Trombella for all species here
considered until Womersley’s action. It may therefore
be taken that from 1887 to 28 May 1954 (the date of
publication of Womersley, 1954b) all usages of
Trombella included, or implied the inclusion of,
Nothrotrombidium. Subsequently, the great majority
of authors used this separation, exceptions being Feider
(1955, 1959a), Daniel (1959) and Krantz (1978), who
still included T. otiorum Berlese, 1902. These remarks
apply also to the incorrect spelling as Thrombella by
some authors. Feider subsequently (1958) became aware
of, and accepted, the use of Nothrotrombidium.
Trombella Berlese 1887, fasc. 40, no. 2; 1888, p. 180;
1893, pp. 91-92, 96, 138, 149; 1894, fasc. 72, no. 6; 1902,
p. 127; 1912, pp. 2, 4, 8, 9, 11, 14-16, 18, 22-30. Hirst,
1929, p. 168; Vitzthum, 1929, pp. vii, 63; 1931, 3(I1),
p. 146; 1940, pp. 144, 145; 1941, pp. 506, 826.
Womersley 1934, pp. 181, 185; 1937, pp. 75, 76; 1939,
p. 149; 1954b, p. 125. Thor 1935, p. 108. Willmann
1939, p. 15; 1940, p. 215; 1941, p. 59. Thor and
Willmann 1947, p. 199. Baker and Wharton 1952, p.
250. Feider, 1950, p. 4; 1955, pp. 26, 41, 68; 1958, p.
265; 1959b, p. 541. Vercammen-Grandjean 1955, pp.
253, 260; 1973, p. 109. Vercammen-Grandjean and
Kolebinova 1968, p. 250. Schuster 1960, p. 5. Krantz
1978, pp. 278, 351. Southcott 1982, pp. 286, 290.
Thrombella André 1934, p. 472; 1936a, p. 9; 1936b,
p. 325; 1938a, p. 215; 1944, p. 230; 1958, pp. 14, 15;
1962, p. 63. Robaux 1967, pp. 3, 4, 7, 109; 1968, p. 453
(incorrect spelling).
Trombella alpha n. sp.
Description of holotype larva (specimen ACB713A;
also supplemented from other specimens) (Figs. 1A,
B, 2, 3, 4A, C, SA, B).
Colour in life orange. Length of idiosoma (partially
engorged specimen, slide-mounted) 355, width 215;
total length of animal from tip of chelicerae to
posterior pole of idiosoma 445.
Dorsal scutum trapezoidal, with a broad projecting
anterior “nasus”; anterolateral angles (“shoulders”)
rounded, obtuse-angled; posterolateral angles evenly
rounded. Anterolateral margins indented, lateral and
posterior margins concave. Scutal sensilla placed
towards rear of scutum, well separated, and closer to
level of PLs than ALs with only a faint indication of
setules (branches, see Goff et a/. 1982) with phase-
contrast oil immersion microscopy.
Scutal scobalae (non-sensillary setae) six in number,
with two AM setae at about middle of “nasus”, the ALs
and PLs towards their respective angles; these setae
ciliate, slender, tapering, a little blunted, with a single
large terminal setule in many instances.
Standard data of the holotype and a series of other
larvae are summarized in Table 1. Raw data are held
on file at the South Australian Museum.
TROMBELLA ALPHA N.SP. (47°
PIG. 1. Trontbella alpha sp. noy, Holotype larva. A Dorsal view, to scale on left, (Legs on left side omitted beyond trachanters,) B, Dorsal
idiosomal sela, further enlarged (not to scale). In this and in Subsequent figures the A sign indicates the seta is shown in both the
dorsal and ventral fiers
148
TABLE 1, STANDARD DATA AND OTHER MORPHOMETRIC
DATA OF A SERIES-Ob LARVAE OF TROMBELLA ALPTIA
N.SP,
(ALDI ASUrerBeN Ly Hi eLONEtes. ar
Character olplype Range Meat Su. n
AW 33 55-04 S607 234 12
BPW R4 TAY 84,50 419 \2
SK 25 22-24 24,25 y.6ty 12
ASB RI SSNI 7 9.02 u
PSB uw 25-40 3542 4 Atl {2
| 120 §7-|20 108,33 [4.27 4
W 109 100-108 [D567 a.t4 12
ar 43 34-14 RU AUY 44h 011
AM fy Joan 44.75 49 9
AL Lr 206 34 24.55 346 WN
" ay cab s-27 Tl. Mt 255 10
AMH a2 is.22 10 1.66 Ww
Sens — 4h3 | ASAT 3
Ds W-3h GLAD (35-49) 4 DAT
us 40-46 (33-42)-(44-49) _— i
PDS 34-4) (91-37)-( 36-48) - iv
Gel 5S 51-60 34,83 298 32
til 73 62-73 60.34 1470 2
Gell al 47-55 Sy.00 2.13 42
i) 7 S709 62,50 4.37 612
Cell 60) 31-60 5542 s-< a es
Tl 87 73-87 749,83 4 12
* For maximum values of DS.
At times setae are broken in the specimens. As Table
| indicates, It is unusual to find the scutal sensillary
setae intact in a mounted spe¢imen, During the
moulfing process, scutal scobalae commonly fracture.
A number of specimens measured here have been
allowed to moult. This often results in disruption of
the skin, or its crumpling and folding and usually
increases the difficulties of measuring. Im some cases,
however, 4g. in the cast dorsal scurum, it may result
in-a flattening and better display of the more anterior
parts of the sculum, aS Well as better estimates of the
rectilinear length. In fully or partially fed specimens
of the larvae the shield is often, in its anterior part,
too curved and obscured for accurate measurements
on the slide, and in such cases estimates of Land ASB
tend to be unreliable,
yes 2+2, sessile, cach lateral pair on a distinct
ocular plate near PL angle of scutum. Corneae oval,,
anterior with longest diameter 13, posterior with
longest diameter 17,
Dorsal idiosomalae slender, ciliate, blunt-ended,
arising from che usual small plates or annuli set in the
epicuricle, in rows arranged 6, 6, 6, 6, 4, J; Cotal 29.
Venter of idiosoma with a pair of scobalae in area
between coxae [f and I, slender, tapering, pointed,
ciliate, 41 long. Behind the level of coxae I and about
38 setae, tapering, ciliate, pointed or slightly blunted,
in irregular transverse rows across the opisthosoma,
25-35 Jong. Anus (uroporus auci,), of (wo longitudinal
valves (obscured in holotype; 36 long mi ACB71I2E).
Coxalae 2, 1, 1, normal, pointed, with long setules.
Medial coxala | placed over about the mid-point of the
anterior coxal border, or a litle medial to it, 54 long;
lateral coxala | placed well laterally, over the beavily
chitinized anterolateral rim of the coxa, 5] long, Coxala
Ef placed towards ouier part of coxa, about 1/3 back
from the anterior horder, 47 long. Coxala LLL placed
REC S AUST, MUS f9(11): 145.468
June, 1986
over anterior border, about 1/5 back from its heavily
chitinized and projecting lateral part; 46 long.
Lee seemental formula 6, 6, 6. Legs slender, seemenirs
from femur to tibia more or less evlindrival; tarsi
tapering, attenuated, Leg | 430 long, Li 415, U1 455
(each including coxa and claw), Pedocoxal supracoxalae
nol identified, presumably absent,
Tarsus | 116 long, by 18 high at its thickest part, near
its origin, Til/Gel ~ 1.33. Tarsus IL U8 by 18;
Till/Gell = 1.3L. Tarsus Hl 124 by 18 THY Gell =
1.45. (Tarsal nveasurements exclude claw and pedicle.)
For other leg metric data, see Table 1, One faleiform,
slender claw to ¢aech tarsus.
Leg scobalae (ie. branched, barbed setae) slender,
tapering, pointed, the setules (barbs) moaderarely
outstanding, Leg scobalar formula: trochanters |, I,
1H 101), 101), 101), fernora 6-7(5-7), 7(6-7), 4(6), getiua
4(4), 4(4-5), 4(4), tibiae 7(6-9), 6(6-7), 7(7) (figures given
for holotype, followed by the range observed in at least
six Specimens, in brackets),
Fernora, genua, libiae and tarsi with specialized setae
isee Figs 1A, 2). Large striate solenoidalae are present
on leg segments as figured, Ln addition to these easily
identifiable setae, the femora, genua and tibiae carry
small, slender, pornted, smooth i.e unbranched setae,
not optically active, In previous papers | have identified
these setae by the name of “spinalac”. These setae will
be referred to here as spinofemoralae, spinogenualac
ete, according to my previous system of nomenclature
(Southcott 1961a, 196lb, 1963; see also the further
comments below), Solenotarsalae 1, 1, 0. Solenotibialae
2(71), 2070, (20) (there is some difficulty in
differenhating the more slender of these setae from
spinalae). (A previous statement (Southcou 1982, p.
317) that the formula for the solenotibialae in Chyseria
is 0, 0, 0 was wrong, it should be 2, 2, 1 as shown in
the figures to that article.) Vestigiotibialae T, 0, th
Vesliziogenualae |, I, 0.
Grathosoma small, compact, the combined chelac
bases from above almost hemispherical; gnathosoma
9§ long to front of cheliceral blades, by 82 wide.
Cheliceral blades large, transverse, each with three
(range 0-3) retrorse teeth along posierior edge, Anterior
hypostomala pointed, ciliate, 22 long, Palpal coxala
present, lapering, pointed, ciliate, 34 long, Palpul
formula 1, 0, 2, 1, 3, 8 of 9, Dorsal palpfemorala
lapering, ciliate, 35 long, ventral similar bul more
slender, 35 long. Palpgenuala tapering, ciliate, placed
dorsolaterally, 31 long (from AC'B7UC; broken in
Holotype), Palpal tibialae and tarsalae as figured,
Palpal tibial claw bifid, with the tines short, curved,
pointed, a lithe separated, shghily unequal, Palpal
supracoxala not identilied. Galeala absent,
Analysis of the Larval variates
In an analysis of the data in Table |, | have
eliminated the Sens figures, since only three are
recorded, Also, as the data for the various estimates
TROMBELLA ALPHA W.SP. 149
=)
FIG, 2. Trombella alpha sp. noy. Holotype larva. Ventral view (legs on left side of drawing omitted beyond trochanters).
150 REC, 8. AUST, MUS, 19(1}): 145.168
for DS are to Some extent redundant, these figures have
been restricted to the maximum of the DS. The data
are thus reduced to a 19 « (2 table of variates.
A correlation matrix was first calculated from these
figures, followed by a principal component analysis
(Hotelling 1933), In view of the small number of mites
only the first three (independent) patterns are
considered.
Patlern 1; The variates ASB, L, A-P and Ti] (and
to a less extent Til] and GelLL) varied in the same way,
and accounted for about 40% of the patterning, These
variates refer mainly to estimates of shield length,
together with that for Tibia 1,
Paitern 2: The variates AW, PW, PSB, AM and Gel
vary together and account for about 30% of the
patterning. These refer mainly to shield width
estimates, plus Gel,
Pattern 3: The variates DS and Til vary together and
make up about 20% of the patterning. This pattern
refers only to estimates of the maximum length of the
dorsal idiosomal setae and of Tibia 1, and has no shield
variate component.
I. is interesting that there is in each component a
representation of a leg variate. [t may be commented
that these findings appear to differ from results
commonly obtained from morphometric studies upon,
for example, vertebrates, where the first pattern is
usually of a size variable, 1! such were the case here
one would expect that all shield measurements would
tend to bé represented in Paitern 1, The results could
be, to some extent, an indication of the uniformity of
the parasite samples available for study.
Description of Protonymph
(see pp, 160-161)
Description of Deutonymph
(Figs 6, 7A, B, 8A, B, see also Figs 4C, D, 7, B)
Description based mainly on reared specimen
ACB712B,
Colour in life orange. Idiosoma slender; cordate,
flattened dorsally, with prominent division between
propodosoma and metapodosoma. Propodosoma
more or less conical, with base considerably narrower
at junction with metapodosoma, Anterior edge of
metapodosoma more of less straight, terminating
laterally in slightly obtuse but rounded shoulders,
continuing into the posterolateral and posterior mareins.
(In the unfed and unmounted newly emerged nymph
the posterolateral borders are somewhat concave; see
Fig. 4D). Length of idiosoma from tip of “nasus" to
posterior pole 840, greatest width 425, length of
propodosoma 200, of metapodosoma 640,
Propodosoma with a pair of dorsal sensillary areas,
each sensillum being mounted in a small boss carrying
chitinous projections with pointed tips, the whole
appearance burr-like. Each boss laterally with a pair
June, 1986
BG, 3, Tiombellawulpha sp. nov, Larva in situ, parasitic on vw ericket
(SEM by courtesy of Mt 8. J, Davidson.)
of short, blunted, ciliate setae, 12, 15 long; each of these
setae projecting from individual papillae. Anteriorly,
propodosoma produced into a short blunted point
overlying cheliceral fangs; from this point the border
runs back posterolaterally and almost transversely to
an obtuse shoulder, and then more posteriorad,
Anterior and peripheral part of dorsal surface of
propodosoma with almost smooth “tear drop” setae,
each mounted on an individual papilla, an enlarged
and heightened annulus of the seta. Long axis of
scobillum of each seta lies more or less transversely
upon seta shaft, pointing slightly upwards, The more
antenor of these on propodosoma point more or less
medially; those on anterolateral parts of dorsum of
propodosoma point more Or less anteriorly (Pig. 7A).
The more central and posterior part of propodosoma
rugose and devoid of setae, the anterior portion, lying
between (he sensilla, being level, (he posterior partion
formed into two large projecting bosses, with burt-like
projections of epicuticle; propodosoma bordered by
“tear drop” scobalae, and, more laterally, more
attenuated setac,
Eyes 2+ 2, sessile, lateral upon propodosoma, lateral
and somewhat posterior to sensilla, Each pair consists
of an anterolateral and a posterolateral eye, with a thick
cornea, each about 16 across.
A single filiform sensillary seta, 96 long, emerges
from each sensillary boss.
Hysterosoma dorsally with 146 oval or circular
shallow pits i two lateral lines, each of six depressions,
and a medial longitudinal row of four. Depressions
fairly close to each other, and collectively occupy a
considerable proportion of dorsum of hysterosoma,
Anleromost pit of median row elliptical, somewhat
posterior to two anteromost pits of lateral rows; behind
it are two circular pits, then an elliptical pit, sel among
the six most posterior of the lateral pits of the dorsum.
Each pit of lateral row somewhat elliptical or ovoid,
second the largest, posteriorad they become
progressively smaller.
TROMBELLA ALPHA NSP. 151
—
Suu ow
% eT ‘dv
FIG. 4, Trombella alpha sp. noy. A Larva in oblique lateral view, undergoing transformation to nymph, specimen AC B712C on 9.iv.1980.
e eyespot. B Later stage in the larva to nymph transformation, from below, from specimen ACB712B on 9.iv.1980, The nymph is
developing within the uneast larval skin, two of the larval legs being shown; UII indicates larval leg HI. In the nymph c¢ indicates
the developing chelicerae, p the palpi, and 1, 2, 3, 4 indicate the developing nymphal legs. C Cast larval skin and some of the protonymphal
skin (latter drawn in heavier line) (specimen ACB712D). The palpi and basis of the gnathosoma are laid back and these parts of
the mouthparts are seen in ventral view. a the “Y”- or “T”-structure of the deutonymphal skin. 6 another part of the deutonymphal
skin. c the “mateola” or studded boss of the deutonymphal skin. D Deutonymph (ACB712C) seen in transparency. (All to nearest scale).
152 REC. 8. AUST. MUS. 19(11); 145-168 June, 1986
FIG. 5. Trambella alpha sp. nov., protonymph. A An almost intact protonymphal skin, from specimen ACB712A. @ anal valves from
the larva, as area setosa, # mateola or studded boss, 7 one of the nodular bodies inside the skin (shown cross-hatched), /s attached
piece of idiosomal skin of larva, ds dorsal scutum of larva in a fragment of larval skin (to scale on lett). B Anal valves and area
setosa of another specimen (ACB714B) (to scale on right).
TROMBELLA ALPHA N.SP.
100
500
um
i a
ae
aetheh
i oN ii
ft
)
Y
° ae noe Vy
Sat.
—!
—
rc
rs
ar
>
Mh
e |
"03
FIG, 6. Trombella alpha sp. noy, Active nymph (deutonymph), specimen ACB712B, dorsal view, entire.
Most of dorsum of hysterosoma densely covered
with robust setae, with tear-drop shaped scobillum
perched transversely on chitinized “papilla” (seta
annulus). Surface of scobillum with faint more or less
oblique pattern, visible at high magnification, but
scobillar surface smooth in lateral view. These setae
occur over most of dorsum, including pits, but are
sparser in inner parts of pits. Setae smaller in anterior
part of dorsum of metapodasoma, and surfaces tend
to project, thus resembling small burrs; laterally setae
tend to elongate and be ciliate.
The second, fourth and fifth of lateral row of pits
contain, more or Jess centrally, ring of chitin (appears
as “C” in pit 5) set lower than bases of setae; it possibly
functions as a respiratory aperture, but could function
as muscle insertion; presumably derived from seta base
(annulus).
Ventral surface: not clearly seen in ACB712B, but
appears to be normal for 7rombel/a. Genital aperture
and anus appear to be normal for deutonymphs
(somewhat obscured in mounts).
Legs (Figs 6, 8A, B) short, fairly robust for a
urombidioid mite, segments beyond trochanters more
or less cylindrical, Leg 1 620 long, II 415, III 445, 1V
520 (including trochanters to tips of tarsi, without
claws). Each tarsus bearing two claws. All segments of
legs with irregular surface due to leg setae (scobalae)
originating from small papillae. These leg setae
generally robust, and tend to resemble body setae but
are much more slender, tapering and blunt-pointed. In
more proximal and more distal parts of legs these
scobalae tapering, more slender and flexible, more like
usual setation of trombidioid mites. Leg segments
carrying small spiniform sensory hairs (spinalae)
interspersed among scobalae, from telofemora to tibiae,
In leg | (Figs 6, 8A) tarsus from above more or less
parallel-sided, 198 long by 67 wide. In leg 1V (Fig. 6)
REC. 8S. AUST. MUS, 19(11): 145-168 June, 1986
154
TROMBELLA ALPHA N.SP. 155
FIG. 8. Trombella alpha sp. nov, Deutonymph. A, B Normal specimen ACB712B, A left leg I, 8 left leg 11. C-F Abnormal legs of specimen
ACB7I2A (see text), € left leg 1, showing extensive deformity, with almost complete loss of all segments beyond the telofemur, but
al] sezements somewhat abnormal. D Right leg I, showing a generalized deformity with progressive shortening of segments; note also
loss of setae. E Left leg Il, similarly affected, F Right leg 1, with considerable deformity, including shortening of all segments and
gross reduction of setation, and reduction of the paired claws to a single short peg. (All to scale shown.)
FIG, 7, Trombella alpha sp. nov. Deutonymph. A Dorsal view of anterior part of idiosoma and tips of chelicerae, specimen ACB7I2B.
B Further enlargement of the setae of the anterior of the median column of depressions, The surface patterning of three of these
setae is shown at the lower right, C Chelicera of specimen ACB712C, D Medial aspect of R palp of ACB7I2C. (All to nearest scale.)
156 RECS. AUST. MUS, 19( LI: 145-168 Aine, 1986
TABLE 2. MEASUREMENTS OF LENGTHS (um) OF LEG SEGMENTS OF A SERIES OF NYMPHS OF (ROMBELL.A ALPHA
N.SP; AND SIMILAR DATA FROM THE HOLOTYPES OF TROMBELL Al ADELATDEAE WOM, AND T. WARREGENSIS HURST,
TOGETHER WITH RATIOS OF LEG SEGMENTS AS A PROPORTION OF TARSAL. LENGTH
: ap Acuile ot Adult of
Nyniplis at Tromhella alplia T. adlelaideae TS wareepensis
Specimen number ACBTIZR ACBIII4* ACRB712€ ACB7]2D AC B7I4R' ACBT29 ACHT30
Segment wm palia pm = ratio wm ratio wm ralia wm ratio pm rati wm rho
Tarsus | 198 1,00 42 1.00 R30 1.00 1 Loo WO 1.00 20 1K) 30).
Tibia 1 120 60 fil 98 106 58 120 63 98 58 180 64 320 RD.
Genus [ RO au a9 19 —! — 90 Al —f — 148 A3 230 JY
Tarsus IV 126 1.00 122 LOO 35 (1.00 133 100 125 1,00 Za naw nib 1.00
Tibia 1V 4 4O 113 93 117 BT 1] BI 105 (84 192 30 328 1,05
Genu TV 72 st 73 60 75 56 78 59 72 58 128 60) 197 63
* Teratological specimen. The treasurements are daken from the less deformed right side (see text), Legs Li and UL also show the same
deformity (brachymely).
| This specimen has a brachymelic right leg WI. However legs | and TV appear nermal {see text),
+ Specimen flexed, nol measurable in mount.
farsus tapering, 126 long by 36 wide. For other leg
dimensions see Table 2,
Palpi fairly stout, provided with robust scobalae over
TABLE 3. PERCENTAGE OF 7 COMMODUS PARASITISED
(BY 7, ALPHA) AND THE MEAN PARASITE BURDEN PER
CRICKET, OVER TWO SUCCESSIVE SEASONS OF PLT FALL
IRAPPING AT KIRBY STATION.
femur and genu, more distally setae more slender, aod Number of Percentage of = Meati number
: ad (& 7 ~ dias Collection crekers crickets with of mites
some are barbed (see Fig, 7D, from ACB712C), Palpal date examined mite parasites per cricket
tibia with Strong curved claw, and accessory claw-like
Sain " ; ey - 3.1.79 3i 42 07
seta on medial side, Palpal tarsus with sparse setation i179 55 9] 793
of slender pointed hairs, some barbed. Pit 1s +4 soa
Cheliceral Fang robust, curved, pointed, 71 long, “Oo iv.79 46 50 0.80
ansing [rom a basis chelicerac about twice as long (Pig. 24.iv,79 43 18 O.18
ka a Ae aTIe : BOP A pean (1979) 44 61 zd
ry Ato aC). 11.80 ny 4 0.34
a , 811-80 M4 21 0.36
Origin of the Specimens of Trombella alpha 711,80 39 18 0.23
I 4.iy.80 75 15 0.19
This study of Trombella alpha is based upon larvae = Mean (1980) #2 2) 0.3
Which have been obtained parasitic upon field crickets,
Teleogryllus commmodus (Walker), in the New England
District of New South Wales, by Mr 8, J. Davidson,
who commented (pers, comim,, 5.86.1979) on“... a red-
orange coloured mite which appears to be parasitic on
gryllids. Up to 50 per cricket have been recorded, with
mouthparts attached to the hosts’ pleural regions,
especially on the metathorax” (see Fig. 3). Ina further
note (pers, comm., 1982) Mr Davidson has summarized
his observations On the parasilisation of the crickets
by this mite species as follows: The mite “attaches
principally to the soft pleural regions of the
hosr.., Numbers of this mite were monitored by
examination of most of the T commodyus trapped in
pillalls from 1979 to 1980. A higher percentage of
vrickets was parasitised in 1979 than in 1980, with
respective means of 61% and 20% . . . Similarly, there
was a higher mean parasite burden per ericket in 1979
than in 1980, The mite was most prevalent on crickets
in about February, and both percentage parasitism and
parasile burdens declined in late autumn..." [Table 3]
Material examined
ACBT7I1; 4 specimens, collected from paddock
number K2, Kirhy Rural Research Station, near
Armidale, N.SW., 12.44.1979, Preserved in alcohol,
decolorized.
ACB712; 5 specimens, live, eriekets colleeted
12.j11, 1980; mites removed 19.11.1980, (Transformed to
nymphs in Adelaide.)
Note: These data refer to all instars af che erieket in which the sex
can he distinguished, ie. the last three instars (see also Table 7, and
lext thereto).
ACB7I13: Several
preserved iy) alcohol.
ACB714: 3 specimens, live, collected 19,111,1980,
ACB712, ACB7\3 and ACB7/4 were all collected 10
km N of Armidale, N.S.W.
Type and paratypes to be deposited in the South
Australian Museur,
larvae collected 19.111. 1980;
Rearing Experiments: Larvae lo Nymphs of V. alpha
Deuronymphs were reared from two batches of
larvae forwarded to my laboratory, on strips af wet
blotting paper in sealed Lubes, Experimental details arc
as follows:
(1) Experiment ACB712. The cricket was collected on
12.41.1980 and the mites removed on 19.ii/. L980. The
five larvae sent were received by me on 21.11.1980.
On feceipt the mites were immobile, ic. they were
possibly in a pre-pupal condition, One mite was
rather swollen, on its back in a film of water on (he
site of the tube. The legs looked decolorized. One
larva Was waving its legs abour, stuck to glass. The
three other larvae immobile, not swollen. Later in
the day two mites became immobile and were
considered swallen, while two were waving their legs
about. All live were stuck ina Film of water and the
tube was wet.
23.11.80. All larvae immobile, plump,
TROMBELLA ALPHA NSE is7
24..80. All immobile, not shrivelled, not mouldy, One
or more were considered to be in a pre-pupal stage.
25-24. 10,80. All mites observed daily, and recorded as
immmdabile,
S047. 80. In (wo specimens “frosting” under the skin
ol) the mite was detected. This was interpreted as
representing the development of nymphal setae
within the exuviae,
311.80. 3,00-3.15 pum. One larva appeared to be sitting
upon a pupa larger than tlsell. Skin of Jarva
appeared transversely wrinkled. The other larvae
were smooth, swollen, not mouldy, and were
considered to be in a pupal state,
3,30) pam, The legs af the nymph were clearly visthle.
Anterior part of the larval skin had gradually
become whiter and more opaque, as. though filling
with air. A clear gap was Visible, most noticeable in
the right rear leg,
The legs of the nymph gradually exiended away
from the body of the animal.
Other larvae also had air under the skin.
Liv.80. Nymphs had not as yet emerged. Two larvae
in the tube had promincut protuberances,
8.iv.80. Nymphs had still not emerged, despite nymphal
palpi and legs being clearly visible within the larval
skin,
9.jv.80. Camera lucida sketches were made of the
transtorming larvae (see Figs 4A, B).
15,iv.80. Two nymphs had emerged, these being labelled
ACB7I2C and ACB7IZD. These were not the
specimens mainty described above. The cast larval
skins of rhese two mites, also a dead larva
(ACB7I2ZE), were mounted.
164¥,80. One more nymph (ACB712A) emerged in the
morning, and one (ACB712B) at night. The cast
skins of these two specimens were mounted,
Wiv.80. One nymph (ACB712D) was immobile,
possibly dead; mounted. I attempted to feed the
nymphs with pieces of apple, culicine mosquilo eges,
and squashed adults of Aedes notoscripius, also a
sainple of bird dung, and the juice of a squashed
grasshopper (Phaulacridium vittatum), ete. One
nymph appeared to feed upon Lhe piece of apple, and
possibly feeding could have occurred with the other
materials. One nymph (ACB712D) was seen moving
slowly as late as 17.v.80; the others had died earlier.
Eventually all nymphs were mounted in gum chloral
mountant for study; none had transtormed to any
later instar,
(2) Experiment ACB714, Three plump larvae were
received in wet preparation on 21,111.80, having been
collected on 19,i1,80, All remained immobile, and
unattected by mould filaments spreading slowly
across the inner surface of the glass tube, which
began lo produce spores on 27.iii.80.
On 281.80 one mite was observed to have
decolorized legs and mouthparts.
On 31.11.80 nymphal legs developing in one larva,
In ats unchanged larval skin, were detected.
On 8.iv.80 one specimen (ACB7I4A) became
mouldy. Another specimen (ACB714C) eventually
shrivelled to a more or less spherical object without
recognizable features. One larva (ACB7IL4B) was seen
to have protruding limb masses by 8.iv.80, and a
nymph emerged on 20.iv.80, Food was offered ta it
in the form of a mass of culicine eggs, pieces of grass,
a squashed dipteran, a squashed small weevil, a
squashed Phaulacridium vittatum etc, and the mite
possibly fed, It Was not seen moving after 13.v.1980.
The mile and its cast skin, and the other larvae used
in the experiments, were eventually mounted on
slides for study.
From these observations a confiden! carrelation of
the larva with the nymph of this species can be made.
Table 4 summarizes the details of the successful
larva-nymph transformations.
Remarks on the Larva to Nymph Transformation
The active nymphal stage of the Parasitengona is
currently recognized as the deutonymph, Between the
larva and the deutonymph is the protonymph, which,
as with most prostigmatic mites in the Parasilengana,
is calyptostatic and takes place within the larval skin.
In the Erythraeoidea the skin tends to retain its shape
when shed by the protonymph, and can commonly be
retrieved in one piece after such eedysis (Frauenfeld 1848;
Womersley and Southcott 1941; Southcott 1946a, 19460,
1961a), In the case of the smaller Trombidioidea the
protonymphal skin is thinner and more fragile, so that
although there have been many studies on larva fo
nymphal transformations in the Trombiculidae and
ather families, rarely is there an attempt to describe
the protonymphal stage between the larva and the
deutonymph. However, modern exceptions to this
statement can be found in the works of Jones (1951,
(ABLE 4.. DETAILS OF SUCCESSFUL TRANSFORMATIONS OF LARVA TO ACTIVE NYMPH IN TROMBELLA ALPHA NASP.
Date
bxpenment and Date Date Date became Achve nymph Duration of the
specimen mumber collected received immobile emerged immobile phase*
ACHMIZA 12.111.80 21 .i11.80 19-23.41.80 1h.iy.80 24-28 days
ACB?I2B 12.in.80 21.11.80 19-223.471.80 16.iv.80 24-24
ACKRTIIC 12,111.80 21 ii. 80 19-23.111.80 15,iy.80 3-9
ACBTIZD 12 i. R0 ZI iii, BO 19-72 HiLRO 1S.iy 80 23-27
ACBTI4B 19 411,80 21111, 80 1O7L Hi8O 20.80 30-32
* By subtraction, The figures given express a ninge of possible dates from the data availuble, Since there #4 nO precise indication of the
time of Commencement of the protonymptial stage this nor possible to give ay éstimate of the durauion of that period.
iss
1954), Neal and Barnett (1961), Johnston and Wacker
(1967) and Robaux (1974),
An intermediate membrane seen within the investing
integuiments of the prelarva, or later stages, was named
the “Zwischenhaut” by Claparéde (1868) in tis study
of the development of the water mite Atax bonz
Claparede, the “apoderma" by Henking (1882) in his
study al Trombidium fuliginosum Hermann, ar the
“Fntermediate skin” by Jones (1951, 1954), in his stucty
on Neorrombicula autumnalis (Shaw). Since
deutonymphal development takes place entirely within
the expanding larval skin, there appears no need for
the protonymph of the Trombidioidea to develop the
distinct pupal setation as in the Erythracoidea, and
which presumahly has a deferisive and possibly
substrate-anchoring function.
In the case of the developing Trembella alphu
deutonymph, some of the suecessive stages of ihe
gradual transformation from immobile larva through
the protenymiph to the active deutonymph are shown
in Fig. 4A, B. Ininally the legs and palpi lose their
orange coloration, os the living substance of the aninial
is withdrawn to a more compact mass, The ventral part
of the larva enlarges, so that after several ditys the
appearances are of a larva riding upon a larger rounded
structure, a flattened prolate spheroid. The nymphal
leg prominences gradually appear, and project
increasingly from the main mass. The larval skin
appears to separate from the developing nymph, so that
the darsal surface at least appears as whitish and more
opague. The emerging nymph may or may not leave
behind a more or less intact larval skin. Generally the
“intermediate skin”, i.c. the protonymphal skin, remains
inside the cast larval skin, so that il is not easy Lo
recoginze. Ovvasionally parts of the protonymphal skin
separate from the larval skin, and rarely, the
protonymphal skin may appear as a more or less intact
structure. Even within ihe east larval skin, however,
distinet structures may be seen in the exuviae discarded
by the nymph. Thus in Fiz, 4C there is visible a Y- or
T-shaped (hickening of the integument (“a"), shown
underneath the larval scutum, More posteriorly is a
distinct structure, near the midline, which appears as
a studded boss or “mateola", with folds of thin skin
runing up to it (“ein Pig, 4C), Other less clearly
recognizable structures can also be seen, with vague
outlines suggesting that ihey may have been investments
of the developing limbs, and sometimes with numerous
small elevations (eg., as shown in Fig, 4C, “b"),
One distinet structure is made up of twa elonpale-
oval or kidney-shaped valves, surrounded by a large
oumber of small conical elevations in.an “area sctosa”
(‘as" in Fig. 5A; see also Fig.. $B), These valves are the
same structure as the larval anus or “uroporus”, haying
the same appearance of two apposed valves, but
surrounded by a highly modified area of skin,
Camparison of the cast larval and pratorymphal skins
shows thal between them they possess only ane pair
REC, $8. AUST. MUS. 19(11): 145-168
June, 1986
of valves or valve-like structures, which are ot the same
dimensions as those of the larvae, ic. these are of larval
Oligin, the area setosa, however, is purely af
prolonymphal origin, The more or less concentrically
arranged elevations are short cones with blunted tips.
They are setiform in shape, and either represent
precursers to sttac, or possibly serve as investing
structures for the developing setae of this area of the
deutonymph. These protonymphal setae are similar in
appearance to che projections of the “studded bass"
(“miateola”, or sinall mace) mentioned above
Although i! is not al present possible to allocate
analomical positions or functions to all of the
structures of the protonymphal skin, at least some of
the main structures can be postulated.
In Fig. SA is shown a more or Jess intact
prolonymphal miegument, after slide-mounting
through lactophenol and gum chloral media (specimen
ACB7I2A). This is described briefly in the following
section,
Peseription of Cast Protonymphal Skin
(Fig, 3A, B)
This skin (ACB7IZB) is more or less in one piece,
transparent and colourless. Overall length 875, to tip
of projecting anterior pointed structure; width 445_
General outline oval, with broad rounded posterior end
and vrumpled but narrower anterior end. Whole of skin
with crumpled appearance; only some of the many
minor folds are shown in Fig. 5A. The appearance of
the skin matches roughly the shape of developing
nymph (see Fig. 4B), with widest part and lateral bulges
corresponding to legs 1V ol nymph. The position al
anal valves and surrounds corresponds to posilion of
these structures in larva and deutonymph. The anterior
part of cast skin is more difficult to assign to
corresponding structures, but presumably the various
parts-are investing integuments to chelicerae, palpi, and
the more anterior legs.
The mateola appears in the skin, projecting
anteriorly, about midway between the anal area and
ihe front of the skin with folds of integument leading
to the base. The “Y-structure”’ was not certainly
identified, bur possibly was contained in the folds
drawn around the mateala (“win Fig. SA).
Rounded markings present over various parts of cast
skin, recognizable as founded elevations on
protonymphal skin, There is no evidence of seta
formation in (hem (such skin elevations are a well-
marked feature of the few developing, protonymphs of
(rombidioid mires that have been described; eg.
Johnston and Wacker, 1967, Fig. 5, p. 307, for
Eutrombicula splendens),
Several thickened, almost nodular deposits appear
Wilhin the skin, shown cross-hatched in Fig, 5A, one
being labelled "9?" Whether these are mere accidental
inclusions, or are representative of some consislent
PROMBELLA ALPHA NSP.
process ts mol known, because only one reasonably
intact protonymphal skin was available
Anal valves (irom larva) atid area Setosa as
deseribed above,
Associated with the protonymphal skin in this
preparation are only two other pieces of larval skin,
One (‘ds” in Fig. 5A) consists of the larval dorsal
scurum arid a small amount of its surrounding skin,
detached from the protonymphal skin, The setae of
sculum are severely damaged from the processes of
transtormation, and possibly by the mounting
procedure and no seta is intact. The other small larval
skin piece is at the posterolateral aspect of the
protonymphal skin (Us? in Pig. 3A}.
varlunions ut (he Leg Structure af the Deutonvinph,
and the Developmental Abnormality “Brachymely”
Since dimensions of leg segments are commonly used
as species-dilferentiating characteristics in these and
other mites, i 15 instructive to examine the intraspecitic
variation of a batch of mites which have been collected
from the same area, and handled in the laboratory as
uniformly as possible.
Table 2 lists tarsus, tibia and genu lengths of legs
hand JY of the reared 7rombella alpha nymphs (also
those of holatypes of 7. adelaideae and T. warregensis)).
There is some similarity be(ween tatios of the
segmental lengths in the three nymphs ACB71I2B,
ACR712C and ACB714B. However, ACBR71ZA shaws
#rass shortening of the three measured segments of Leg
[, but no abnormality in leg LY. This occurs in greater
desree distally; and is accompanied by other structural
abnormalities. It occurs in legs other than those shown
in Table 2, and may vary between the two sides of the
animal. This developmental abnormality will be termed
“brachymely”,
Brachvnely ie Vrombella alpha and in some other
Prostizmatic Mites
Fig. 8C-F shows the deformed legs | and UW of
ACKTI2A, compared with the normally developed legs
Tand Woof ACB7IZB (Fig. 8A, B).
JABLE 5, COMPARISONS OF ABSOLUTE LENGTHS (um) OF
LEG SEGMENTS OF LEGS | AND [TV DF REARED
TROMRBELLA ALPHA NYMPHS AND RATIOS
mn
ACBTIRA* ACBTI2ZB ACB712C ACBTISB yaziw X/m
® y ? w 3
Segment
Tal fd 198 83 170 {83,7 35
Til ol 12 D6 OR 108 56
Gel 49 wi) — — 80 61
Tal¥ 122 126 135 125 128.7) 95
hiv 113 Ita WwW 105 112 Or
ciel) 73 72 75 72 73 1.00
* Measuring the loss deformed R leg, for purposes.of Comparison,
Tn the left leg the sepments beyond the femur are worecornizable
from the gross shortening and fusion,
ford, in the cuse of Gel,
459
Table 5 shows normal variation for leg 1V in
specimen ACB712A; deformities in this specimen. are
restricted to legs 1-111, In the right leg J the tarsus, tibia
and genu are considerably shortened, and the setae
much reduced (Fig. 8D), The crochanters, basilemora
and telolemora appear more or less normal, with a
minor degree of shortening and some loss of setae iti
these segments also. The degree of shortening is grealer
distally. In legs Hl (Figs. 8E, PF) and II there ts a
reduction of setae throughout the legs, including the
two parts of cach femur. In right leg 1 (Fig. 8F) and
right leg 111 there is only one tarsal claw to each tarsus.
The most extensive deforrnily occurs in left leg 1, where
there is some thickening of the femur, and beyond the
femur the remainder of the leg is reduced to a single
fused segment (Fig, 8C}, [A this segment there is no
recognizable tarsal claw or claws, with only a
rudimentary indication of setae, without any seta basis
or other part being distinctly recognizable. (In specimen
ACBT7I4B there appears to be a minor degree of
shortening of right leg JH, apparently without other
abnormalities in the body or other legs, The
measurements ol the lengths are: Tarsus Ill 66, tibia
ILL 76, genu HT 57, In tarsus 11 and tibia IL there is
a reduction of setation, and possibly atso in genu FFE,
For left leg Hl of the same specimen: tarsus ILI 93 long,
tibia IT 73, genu III 58.
There was nothing, that occurred in the experimental
handling of specimen ACB712A to Suggest a reason
for the remarkable degree of deformity observed, All
of the Jarvae in experiment ACB7I2 underwent
transformations at about the same time intervals. The
abnormalities cannot be attributed to a failure of the
production of moulting fluid, as such would siniply
fail to allow release, but would not be expeeted to affect
limb development, Abnormalities other than in the
limb segments were not detected. The cast skin of the
larva and protonymph appear normal. The lengths of
the leg segments of the larva, also the shield data (see
Table 1), are normal. The larval tibiae and tarsi, for
example, are normal in structure. The cast
protonymphal skin (see Fig. 5A) appears normal. It
is the most intact of the protonymphal skins available
for study, at least of those that have separated from
the larval skin. It would be inappropriate to attempt
to connect some of the parts of the ACB7IZA
protonymphal skin with structural abnormalities of che
nymphal legs at the present time, because of the lack
of knowledge of the correspondence of nymphal and
protonymphal structures,
It as not uncommon for minor abnormalities in leg
claw structure to be seen in larval erythraeid mites
captured if) the field, This has been referred to as
“twisted claws" or TC abnormality (Southcott, 19612,
p. 481; 1966, p, 732; 1972, p. 29), More severe
deformities are also seen im Lhe legs of larvae caughi
in the field, Thus in one batch of larvae caughi aj
Inman Valley, South Australia in 1952 many larvae of
Lob
Erythrites urrhrae (Wam.) and E. reginae (Hirst)
showed a mild tarsal deformity, and one specimen af
E. resinae showed an extreme deformity of all
pedotarsi, which were grossly contracted and distorted
(Southcott, 1954).
The cause of ihese deformities is unknown. An
attempt to reproduce them in larval erythraeid mites
by chilling hatches of ees resulted in scutal deformities
bui not leg deformities (Southeolt, 1955),
Although it is not unusual for minor abnormalities
of scutal structure to Occur among prostigmiatic mites,
involving. Supernuiricrary setae of loss of setae (sec
Southcotl, 1966, pp. 756-758), 1t appears thal (he limbs
of developing mires are more at risk of structural
abnorinalities (in the form ol’ shortening of segments
and loss of sefarional pattern) than the structures
associated with the idiosama. The claws may suller a
minor detormity, may he grossly distorted, or virtually
absent.
Various ather developmental abnormalities have
heen recorded among the Acarina and in other
arthropods. Among prostigmatic mites there may be
complele abserice of a leg, eg. in the water mite
Sperchon glandulosus thienemanni Koenike (Szalay,
1932), arin Exrrombidium olorheiense Ceider (Feider,
1946), Simple shortening of a leg, but without other
atmormatlity, has been recorded by Feider (1946) in £.
atorheiense, and termed micromely, Sehizoniely
{complete or incomplete duplication of a leg) has been
recorded by André (1949, 1960b) in the trombidioid
miles Microtrombidium sucidum (L. Koch) and
Carporhrombium carduigerum (Berlese). Various lee
und other abnormalines in Hydrachnellae and other
mites were recorded and discussed by Thor (1926),
Exposure ol the developing egys of other arachnids
[oO supraoptimial temperatures has produced limb, eye
and other deformities. Among the limb deformities are
brachymely and schizomely (opilionids: Juberthie,
1968; spiders: Jactinski, 1971, Mikulska and Jacunski,
1971, Mikulska, 1973).
The observations reported here indicate thac
developing limbs of Acarina are sensitive to teratogenic
agents, as yet unknown,
Classification of Post-larval Trombella
Known post-larval forms of Troambella 5. str.
comprise six closely related species, charactor ized by
the lack of a crista metopica and the presence of large
idiosomal depressions, allotted a glandular funetion by
some authors, The genus and species were founded on
uwsingle specimen of 7 glandulosa, found deeply buried
in soil, Adria, narthern Italy (Berlese, 1887). This
specimen, which las not been restudied by any Later
author, had several Unusual morphological features,
nat recorded for any other species since plaved in
Trambetla §& stri (1) the presence of a ehitinized pore
in each of the idiosermal pits, whether dorsal, lateral!
of vertral; (2) the dorsal idiosomal pits not contiguous,
REC, 8. AUST. MUS, 19(11)> 145-108
June, 1986
i fact well separated and occupying only a small
proportion of the surtace; (3) {he posteromost median
dorsal pit being set ina group of four of the lateral
pits, instead of being set in the midst of a group of
six pits; (4) the presence of a Jong sinuous idiasomal
seta arising from a small prominence an the lateral side
of each propodosomal sensillum. These fealures of
Berlese’s original drawing were reproduced in Thar and
Willmann (1947), and the sole further specimen of T
glandiilosa recorded, from Austria, by Schuster (1960),
is Stated fo carrespond avcurately in all morphological
features they give, apart from being somewhat smaller,
One may therefore accept Berlese’s drawings as
accurate.
The African 7) javosa appears to be the species mast
separate from 7. glandulosa, Dorsally it carries large
idiosomal depressions, all Jacking a central pare. bul
surrounded by a prominent ring of chitin, The dorsal
idiosamal setae are also markedly different, being in
the form of large flattened scales, the scobillum ending
in a long ceutral point and being provided with large
projections along its lateral edges (Andre 1936@a, 1958).
In T. glandulosa the dorsal idiosomal setae are
spinelike, simple, and bent near the base of the shalt
so that the main part of the shaft is parallel to the
surface. In 7. /usitanice the general icdiosomal setae are
Spiniform, simple, curved, sharp, on short tubereles
(papulae), only the more peripheral ones of the
propodosoma bearing fine setules. la 72 werregensis
the dorsal idjosomal seta is papillare, clongate, curved
near its base, and ornamented with prominent broad
selules in (he farm of scales or lone barbs; in both 7
udelaideae and TZ. alpha the seta presents asa lear-drap
perched transversely upon a papilla, In 7. adelaidece
the scobillum is ornamented with prominent broad
scales, while in 7 ulpha only a reticular pattern is
visible {see Figs 7B, ¥A-D),
The other hitherto recorded species of adults of
Trombella—T. lusitanica André, T. warregensis Hirst
(which appear to be close to each other) and 7:
udelaideae Wom.—dilfer in having different members
of the idiosoimal pits with central pores, in -setal aud
leg characters, ete. (One feature used by André in
proposing 7 /usitaniea and separating it from 7:
glandulosa and FT warregensis, was stated to be jts
possession of a punctate seta-less area on fhe
propodosoma. However in TD warregensis re-
examination of the holetype under oil immersion
sliows minute puncta in this glabrous area. The same
glibraus area in T edelaideve and T, alpha has small
rounded tubercles.)
Thus in © adelaideae and To alpha the dorsal pits
carry (hese ehirinous pores or rings more or less
centrally in lateral depressions 2, 4 and 5; in 7
warrevensis {hey are present in lateral depressions 2,
3, 4.5 and 6) in & luslraniea (ey ave present in literal
depressions 2, 3, 4 and 5 (Andre, 1944),
TROMBELLA ALPHA N.SP. 16]
Cc
FIG. 9. Dorsal idiosoma setae in adult Trontbella. A, B Trambella adelaideae Wom., Holotype, A setae from the anterior median dorsal
depression; B setae from the posterolateral edge of the dorsum. C, D T. werregensis Hirst, Holotype. C Group of setae from posterolateral
part of dorsal ididsoma; D Two setae in lateral view at the posterolateral edge of the dorsum. (All to scale shown.)
The arrangement of the setae of the dorsal idiosomal
pits has been used by authors (e.g. Womersley, 1954b)
as taxonomic criteria. Thus the central areas of these
pits in 7 favesa bear slender papilla-less spiniform
setae (Andre, 1936a), while in 7: /usitanica the pits bear
spiniform, conical setae arising from a short tubercle
(André, 1944); in 7. glandulosa they are short, conical
and curved (figured by Berlese, 1887). In the three
Australian species recorded here the setae of the pits
do not differ essentially from the general dorsal
idiosomal setae. In both 7? adelaideae and T. alpha the
peripheral setae of the dorsal pits may form a single
vague row, but in 7. warregensis the central part of the
dorsal depressions is nude, with peripherally the setae
in one or two fairly regular concentric rows (in 7;
glandulosa Berlese figures one row of setae near the
centre, and one row at the rim).
Some use has been made of the relative shapes of
leg segments and the rations of various segmental
lengths as taxonomic characters of trombidioid and
other mites, although with soft-bodied mites such as
the trombidioids too much reliance should not be
placed upon the shape characteristics of the tarsi. For
the three Australian species of Trontbella the ratios of
tarsus I length/tibia I length and tarsus I length/tarsus
I width, are given in Table 6.
The data in Table 6 indicate that 7) adelaideae and
T. alpha resemble each other more than either
resembles 7. warregensis, which agrees with the
idiosomal seta characteristics.
The data presented or previously published appear
adequate to separate species of Trombella on a world
basis, even though we do not have any species recorded
ta REC. 5. AUST. MUS. 19(LI): bas. 168
TARLE 6.
LENGTH OF TARSUS I/WIDTH OF TARSUS I.
dune, 1986
COMPARISON OF LEG'SHAPE RATIOS IN TROMBELLA FROM AUSTRALIA, USING TARSUS 1 CIBIA T, AND
IN NYMPHS OF 7. ALPAA AND ADULTS OF T. APELAIDEAE AND
T. WARREGENSIS (See also Table 5 hor leties to lengths)
Trombella alpha nymplis*
T, adelaideae| T. warregensis}
ACB7125 AGB712C ACB7|20 ACHTI4 ACB7249 AC BI30
Tarsus |
Tihia 1 1.65 1.73 159 1.73 1.56 1,22
Carsal width) a7 or 7s As JODO), WZ E bey T7hin(h }, Lode)
(L| {R) iL) (RB)
Length Tal 2h 2.73 2.55 2.2 2st 282 340 cads0
Width Tal Hine 102 Vit (64
ZAG; 2.763 2.224 2.325
* omilting brachymellc specimen ACB7I7A_ See data jn Table Z.
} Re-measured from holotype, See Jala in Table 2, In both tee I is seen in lateral view.
7 Womersley (1954b, pp, 127, 128) gives these ligures as 270 and 90, and the ratio us 3-1
1939, p. 149, he wave these figures as 260 and 90.)
§ Womersley (19546, pp. 126, 128) (who had remounted the spceimen) gives these figures
(1929, p. 170) gave 370 and 740, iu. a matio af 2.64).
in which both adults and deutonymphs have been
described. The following key is therefore presented,
KEY TO THE KNOWN WORLD SPECIES OF POSELARVAI,
STAGES OF TROMBELLA 8. STR,
1, All dorsal pits of idiosoma lack a central chitinized pore-
structure. Dorsal idiosomal setae flattened, with long
median end and 3-5 strong lateral projections, Central
areas of dorsal pits with long spiniform setae, not
mounted on papillae ..,.., . T favosa André
Ai least some of the dorsal pits of the idiosama wirh
a central chitinized pore-structure ........-..--..2
2(1) All dorsal idiosomal pits with a central chitinized pore-
structure. Dorsal idiosomal setae spimiform, bent near
base sa thal the shaft lies parallel to Bed surface, Ratio
of length mrsus I/iibia | about 2.0 . SPA giipe 2f22
r pluncdiedeese Berlese
At least the antcromost of the lateral row of dorsal
idiosomal pits lacking a central chitinized pore-structure
3(2) Dorsal idiosomal setae long and pointed, the seta
scobillum not wider than its basal pupilla. Third lateral
dorsal depression with cesiral pore-structure.,....4
Dorsal idiosomal setae tear-drop shaped, the scobillum
wider than the basal papilla, Third lateral dorsal
depression without central pore-strueture 2.0.0.4 §
4(3) Dorsal idiosemal setae with numerous prominent
setules. Chitinized pores present in lateral row pits of
dorsum numbers 2, 3, 4, Sand 6, Tarsus 1/tibia | about
Diditga tee camer ste ieatmere T. warregensis: Hirst
Dorsal idiosomal setae either spiniform or with slender
setules. Chitinized pores present to lateral row pits of
dorsum Humbers 2, 3, 4 and 5, Tarsus I/tibia | about
Py. ae ce PR Ce ood De Pt T. dusitanica André
5(3) Scobillum of dorsal idigstimel setae with distinct ciliate-
barbed surface... 22,2202, T adelaideae Wamersley
Scobillum of dorsal idiosomal setae with smooth
surface, not presenting as a ciliate-barbed structure -
STTTITTO Ee ee F. alpha n. sp, (deutonymph}
Ley Chaetotaxy of Trombella alpha Larva
Specialized sensory setae upon the legs of tarval
prostigmatic mites have been of interest to taxonomists
for many years, particularly in classification, Although
they function as physical and chemical receptors, there
have been few functional studies upon therm, efforts
having been mainly in description and classification,
with their functions only rarely being postulated.
Classification of these setae has been contentious, with
The specamen has por been remounred. (Ln
is 37S anid (495, and the ratio as 2.75:1- Hirst
two main systems being used in the terrestrial
Parasilengona, which [ have designated the
“trombiculid systern” and the “Grandjean system"
(Southcott, 196la), Despite these differing classificutory
systems, there are no doubts that setal patterns differ,
not only between families, but between genera. Within
the genera allotted Lo the family Trombellidae
considerable differences in leg setational patterns occur
(Southcotr, 1982),
Although the presence of modified “normal setac”
or scobalae—"maslisetae”’—is well-known on the
femora of trombiculid mites (Audy, 1954; Southcott,
1961b), the presence of simple sensory setae upon the
femora is more restricted. In the Trombellidae, in
addition to being present on femur |, Il and HL of
Trombella alpha, they were recorded earlier on the
telofemora | and Ill in Ralphaudyna (as nude
femoralae) by Vereammen-Grandjean er a/. (1974),
However, they occur more widely, being also present
upon the femur of larval Neotromhidium (Southeott
1954; Borland, 1956) and Manunguis (Lindquist and
Vercammen-Grandjean, 1971) (Neotrombidiidae or
Neotrombidiunae), in Burenia (Vercammen-Grandjean
1955; Vereammen-Grandjean and Audy, 1959) (Trom-
bellidae) and Hannemuniu (Hyland, 1956) (Leeuwen-
hoekiidae or Lecuwenhoekiinae) and more widely in
the Trombidioidea, such as in Megophthrombium
(Mullen and Vercammen-Grandjean, 1978) (Micro-
trombidiinae or Microtrombidiidac), and in various
genera of the Johnstonianidae (Newell, 1957; Robaux
1978), It would thus be unwise to place too much stress
on the presence of these setae on the femora of the
larval instars as a taxonomic character.
Similar smooth sensory setae occur also on the
adults and nymphs of Thombella (Pig. 8A-F'). They also
occur more widely in the Prostigmata eg Srrariy
coopert Southcout (Smarididae), referred to as spinalag,
spinolemoralae, ete. (Southeort, 1961b), and on genus
I-lll of the nymph of Micrerrombidium firsutuin
Wom. (Southeott, 1946b),
Another resemblance between Trobella and
Ralphavdyna larvae lies in each possessing two palpal
fermoralae. In Trowbellu, however, there are oo
TROMHELLA ALPHA NSP. 163
vesUizialae; in Ralphaudvng these are present as distinct
‘“mushroom-like microspurs” (Vercammen-Grandjean,
ét al, 1974, pp. 248-9),
Generic Classification of the Trombellidae
1, Adults and nymphs
Thor (1935) defined the Trombellinae as those
Trombidiidae which Jacked a crista, with eyes 2 +
2. sessile, sclae short and sharp, with two sensory setae
on [he propodosoma at the level of the eyes, and some
lesser eharacters, to include only 7romtbella Berlese, In
1937 Womersley added Chyzeria Canestrini 1897 to be
subfamily, as well as Paravhvzeria Hirst 1926 (syn.
Thaumatothrombium André 1938) (1938b)); this was
accepted by ‘Thor and Willmann (1947), However, both
Parachyzeria indica Hirst and FP poecilotrichum
(André) have two pairs of prosomal sensilla, and
therefore should be placed in the family Johnstoman-
idac Thor 1935, subfamily Johnstonianinae as defined
by Newell (1957). Womersley (1954b) described the
reared ¢yeless deutonymphs of Axudyana Wom.,
dividing the subfamily into the tribe Chyzeriini for
Chyzeria and Parachyzeriv, and Vrombellini for
Trambella, Audyvana and Nothrotrom/ridjum Wom.,
1954 (erecled for rhe European 7. ofierwm Berl), but
did not redefine the subfamily. Vercammen-Grandjean
(1955) deseribed the reared nymphs af Durenia, and
provided a revised key for the tribes and genera of the
post-larval Trombellinae. Feider (1955) elevated the
Trombellinae to (he family Trombellidae, a status which
most later authors have accepted (see Southcott 1982),
Within this family (or subfamily) two further genera
have been proposed for adults or adults and nymphs);
Parathrombella André, 1958 and Neonoathrothram-
bidium Robaux, 1968.
In defining these two last-named genera cach author
slated Lhal a crista melopica ts presenl. In
Parathroambella nasata Andre, the type species of tts
genus, the propodosoma bears dorsally | + 1 eyes
laterally, and in the central pan there is a thickened
chitinous plaque, roughly in the shape of a trapezium,
with prominent anterolateral angles, cach bearing a
sensillary seta, which are thus well separated, There is
no anterior linear projection of chitin from this plaque
or Shield. The hysterosomna bears 10 (or 11) contiguous
mares, which resenble (hose of Thombella, there being
four in each lateral row and two or three median. Andre
therefore redefined the Trombellinae, saying that. the
erste] was limited (0 a more or less developed transverse
band, with “rarement un vestige, trés réduit, médian™.
The use of the term crista ete. seems to be largely a
matter of terminology, There is Wo daub that the two
species André classilied in Parathrambella belong ta
the Trombellidae. However the other two species that
he placed in Perarhrombella ditler fram & vasuta in
having the following three characters; (1) eyes 2 + 2,
(2) no large dorsal hysterosomal depressions but instead
a hexagonal network connecting the bases of the seta
papillae, (3) twa small circular pits anterolaterally on
dorsum of bysterosoma. André (1962) recognized that
the (wo later species come within Dyrenia,
Robaux (1968) erected Neonorhrorirambidiani, wilh
type species N. /ranzi Robaux, 1968 from South
America, stating that in this genus the crista is well
developed, but did nat discuss any consequent revision
of the definition of the Trombellinae. As earlier (L966)
he had relerred to @ crista in Nethrorronmibidium
otiorum (Berl,) as consisting of na more than the two
prosomal sensillary bosses, he was clearly using the
term crista i a wider sense than the majority af
authors. Even so, he later (L968) figured in N. /ranzi
@ broad chitinous thickening of the dorsum of the
propodosoima extending from its rear border, atiteriorly
enclosing (he sensillary bosses and with an anterior
putter-like seta-less delimited extension running into the
nasus, In the second species he placed in this gents,
N. antericanum Robaix, 1968, the propodosoma beans
dorsally an oblong plaque, extending only as. far
forwards as to-enclose the sensillary bosses (which are
anterior to the eyes), and without any further anterior
exfeasion.
In my opinion these large thickened areas on the
dorsum of the propodosoma correspond to the dorsal
shields of other proshgmalic miles, and could well be
termiced a scutum rather than a ecrista. The same
situation occurs in Chyzgenia, where there is evidence
of a dorsal propodosomal scutum and also of a
rudimentary crista (Southcott, 1982), The erista may
be considered as absent in N. americanum, and
rudimentary in N. franzi,
The differences enumerated between N, sranzi and
N. dinericoium justify their generi¢ separation, and
for N. americanue the genus Maiputrombella no. gen-
is proposed.
Although (he definition of the Trombellinae giver
by Thor (1935) and Thor and Willmann (1947) included
2) 2 eyes, yet eyes are absent in Audvana, present
as) + lin Paraihrombella, and as2 + 2 in Trombella
and other genera. Far Nathrotrombidium Wom, the
status of the eyes is unclear, as Robaux (1966, Fig. 1D)
shows only] 4 L for NV. orforuim, the type species (bur
does not comment on this in the text), while Andre
(1960a) records thai N, Arewrersum Andrée, (960, has
2 4 2 eyes.
From the foregoing, a revised definition of the
Trombellidae may be offered, with also subfamilies,
and a definition of a new genus.
Family Trombellidae Thor
(Originally as Trombellinae Thor, 1945)
Definition: Trombidiojdca with absent oar
rudimentary crista metopica, Eyes absent, or present
asl + Lor2 + 2; if present, sessile. Propodosoma
dorsally with a pair of sensilla (trichobothria), set in
a fossa which may have an elevated rim, or be present
\64 REC, 8, AUST, MUS, I9(1TE [45-168
as a papilla with a central hole. Dorsum of
propodosoma commonly with a thickened scuyum,
which may be poorly defined at its borders. Ldiosoma
commonly highly modified, with pits or depressions
with a possible glandular function, and bearing
modified setae. Body and leg scobalae borne singly, or
in groups upon small cuticular plates,
Type genus Jrombella Berlese, 1887.
KEY TO THE SUBFAMILIES AND GENERA OF ADULTS AND
DEUPTONYMPHS OF TROMBELILIDAE
1, Dorsum of hysterosoma with prominent sefa-bearing
projections... ..60. veoy say Chyverilnae, new starus
Sole genus 2... .....5-)5 Chyzeria Canestrini, 1897
Dorsum of hysterosoma without prominent sela-bearing
projections, but may bear depressions of possible
glandular function (Trombellinae, $s. str)... -. -2
Dorsum of hysterosoma wilh pits or depressions ., 3
Dorsum of hysterosoma withoul pits or depressianss
3(2) Dorsum of hysterosoma with two small anterolateral
pils. Propodosoma may bear a chitinized plaque
dorsally, adjoining or enclosing the pfetpn geomet
sensilla. Eyes 2 + 2, aboui level with sensilla . ;
. Durenia Vercanimen-Grandjean, 1955
Dorsium of hysterosoma with three bonxitueiaal¢ raws uf
circular, oval or oblong depressions... .. - 4
4(3) Six depressions in lateral dorsal hysterosomal TOW. Eyes
Ja 2 Trombella Berlese. (887
Four depressions in lateral dorsal opisthasemal row,
Eyes 1 + | - Parathrombella Andre, 1958
5{2) Dorsal idiosonal setae on papillae, in groups of 2-(2
in small cuticular plaques, Sersilla well separated, Eyes
(1)
iy Te a Audvana Womersley, 1954
Dorsal ididsormal setae on papillae, arising individually
from cuticle ., 2. - a eer
6(5) Dorsum of propodosonia ‘without ‘significant thickening
to a scutum or plaque-like structure, Propodosomal
sensilla vlose together, somewhat posterior to level oF
eyes (for N. oforum (Ber).), from Roba. 1986)
rt UTA , Nothrotrambidium Wamersicy, 1954
Dorsuim of propodosoma with a well-developed seutum
or chilinized plaque In its poatoripeeeeasl part, Eyes z
i SRT OTT ee eo vitals
7(6) Propedosomal sensilla approximneteck levee “witht eyes,
on an elongate propodosomal scum which extends
forward on to nasus, there forming @ median furrow
Pernt gee ae _ Neonothrathrombidiunr Robaux, 1968
Propodosomal sensilla well separated, anterior (oO level
of eyes. Propodosoma bears dorsally a large transverse
oblong scutum or plaque, enclosing sensillary bosses;
there is no antenior extension of this scutum, Palpal genu
bears distally on its lateral face 4 row of broadened,
spatulate setae, op ane Maipuirombella n. gen.
Definition af a New Genus of Trombellidue
Maiputrombella 7. gen.
Definition: Trombellidae with 2 + 2 eyes, pasterior
to level oF propodosomial sensilla. Propodosama hears
dorsally a large transverse oblong scutum or plaque,
enclosing the sensillary bosses; anterior chitinized
extension of propodosonial scutum absent. Body setae
short, blunt-ended, on Short papillae. Palpal genu bears
distally upon its lateral face a row of broadened,
spalulate setae.
June, 1986
Type species Neonothro(hrombidium americanum
Robausx, 1968.
This genus is known only from its type species. The
generic name is derived lrom Maipu, Chile, the place
of collection, and Tromrbella.
(2) Larvae
Southcott (1982) discussed the characters of larval
Trombellidae, concluding thal it was not possible to
give a formal definition of a larval trombellid, OF the
two subfamilies into which the Trombellidae is here
divided, the reared larvae, except CAyzeria, belong bo
ihe Trombellinae,
A review of the characters of the larval Trombellidae
shows that Momerslevia Radford has a number of
features in common with Durenta, so thal | have
decided to restore the former to full generic status (see
key belaw.)
The following ts a revised key to the larvae at present
allotted to Trombellidace. (Ra/phovdyvne Vercammen-
Grandjean ef a/., 1974, has been included, although a
review of its characters, with two pairs of se\tal sensilla,
shows that its affinities lie with the family Jolinstonia-
nidae, near the subfamily Lasseniinae Newell, 1957,
where if is provisionally placed.
KEY TO THE LARVAE OF TROMBELLIDAE
}. Lee segmental formula 7, 7, 7.-..-.--..-.--.--.-2
Lee seemental tornula 7. 6, 6 or 6, 6, 6. rom 4
2(1) Dorsal seuium Jacking anleromedian projection
(“nasus")...-._- _Chyzeria Canestrini (897
Dorsal scutum with anteroutedian projection (nasus")3
(2) Dorsal seutum with 8 setae — 2.
Ralphaudyna Vercanimen- Grandjean otal, \74
Dorsal scolum wilh @ Sefa4e ..,.,..
Tenet Nothotrambicula Dumbleton, 1947
Leg sepmental formula 7,6, 6. AM and AL (laltler clase
to Sens) setae short, clavate. Two claws ta each
a(t)
pedotarsus. Palpal tibial claw with four prongs
(Vereammen-Grandjean, 1972)...., 2... -
eeet tas becuse) Audvana Womersley, 1954 (1954a)
Leg segmental formula 6, 6, 6. ete eae 4S
§(4) Tarsus ol cach ley with a single claw jn. Sees. 2 8
Tarsus of legs Land U1 with a single apically Lrifureste
claw, tarsus of jeg LIL with two claws , ; 7
6(5) Scutal sensilla well behind level of AL puiiealae. ssbisti
midway between levely of AL and PL. Chelwerae
compact, rhe combined chelae buses not more than twice
as long as Wide,........-,- Trombella Berlese, |887
Sculal sensilla only a little behind level of AL, sculalac
Chelicernae elongate, the combined chelae bases more
than twice as. Jong as wide (lor N. odiorum (Berl), from.
Feider, 1958)...
_ Nothrotrombidium Womersley; 1954 (1954b)
75) Nasus ot sculun) small, largely occupied by the bases
af the AM seutalae, and with a deep constriction behind
Leg tibia 11) with a large solenoidala an
eS RA TY, Monrersleyia Rad ford, 1946
Nasus Gf seutun large, triangular, its lateral borders
continuous with anterolateral borders of seutum, with
al most anly minor constrivian. Leg tibia HL withour
a large solenoidala,..... Au
cowl cies «7 Durenia Vercammen- Grandjean,
tab bot eet Pht br bi fen te
1958,
VTROMBELLA ALPHA NSP. 165
AUDITORY FUNCTION IN ACARINA
(a) Function of Tarsi in Trombella alpha Larva
The attenuated tarsi of larval Trombella alpha
represent an unusual shape among larval prostigmatic
miles, suggesting (hat [hey might serve as sound-
receptor organs. If so, they could possibly be useful
in locating males of (he host cricket. At my request,
Mr Davidson, the collector of the specimens and data,
has provided information on the numbers of mites
parasiiic upon male and females of Teleogryllus
commodus, as well as for sub-adults (A) and A2
instars), As only the adult males stridulate, for the
hypothesis lo be valid there should be a greater
tendéeney for parasitization of the adult males. In Table
7 data are provided on the numbers of mites on the
various instars of the crickets, by date of collection.
Exainination of the data in Table 7 shows that there
is a greater number of larval mites attached to the adult
male crickets (han to adult females, the proportion
being 3,70/1,15 overall, or greater than 3:1, Submitting
the data to a test of the null hypothesis, i.e. that the
proportions of the mites on the crickets are
independent of sex, we find that X¥* = 83.52 on
Id. f., ie, P<OOlt**,
A significantly greater degree of parasitization of the
adult male crickets ovcurs than in the females. It
should be pointed out also thal as the crickets were
captured in pitfall traps (using Vacola jars as traps),
there could have been some transfers of mites between
the crickets, since these traps functioned over several
days and the conditions in the traps the crickets were
extremely crowded When many were caught, Although
only a proportion of the mites were studied, among
the samples submitted there was only this one species
of larva,
‘This finding therefore is consistent with the
hypothesis that the mutes may lind their hosts, at least
in part, by the use of a sound-detection mechanism.
Since only the adult male crickets stridulate, the
hypothesis may be exainined further by comparing the
figures of parasitization for the sub-adull crickets; the
first and second instar sub-adults are the only sub-adult
instars in Which the sex 1s readily determinable. (Mr
Davidson advises (pers, commm,, 1983) that difficulties
in sexing account for some differences between the
totals in the tables.)
Table 7 shows that inthe sub-adult instars the mutes
appear to have a preference for the female crickets, with
a mean of |,57 mites per female against a mean of 1.25
mites per male. If however we compare these figures
by the same null bypothesis as above, we calculate thal
x? = 2,90 on i d. f., not significant, ie. the null
hypothesis is not disproved.
There is thus io evidence of a sexual preference by
the mites in parasitizing the sub-adult crickets.
These results thus are consistent with the hypothesis
that the mites make some use of sound-detection in
their searches for their cricket hosts.
(b) Possible Audifory Oreans in Acarina
It has been suggested above that the larvae of
Trombella alpha use the leg tarsi as sound-receptor
organs for locating the sound-emitting adult males of
the erickets, which is supported by the greater Incidence
of parasitization in adult males than in adult females.
Other instars of the crickets are also utilized by the
miles (compare Tables 3, 7).
Earlier suggestions have been made that there are
structural adaptations in ectoparasitic mites which
serve an auditory function in host location. Newell and
Vercammen-Grandjean (1964) described two species of
mites. from Africa in the family Johnstonianidac
(Trombidioidea) as Pleridopus auditor Newell and
Vercammen-Grandjean, 1964, and PR pseudohanne-
mania Newell and Vercammen-Grandjean, 1964. P
auditor Wad been collected parasitic Upon crickets,
while PR pseudohannemania was described from a
single specimen collected free on the forest floor,
In both of these species there is an unusual
morphological arrangement and modification of setae:
tarsus Ill ane tibia U1 carry a dorsal row of long
feathered setae, each of which has a preformed fracture
line near ils base, corresponding lo an interruption af
the actinochitin. They proposed that these setae
TABLE 7 INCIDENCE OF TROMBELLA ALPHA ON THE CRICKET TELOGRYLLUS COMMODUS
(Only live erickets examined)
Cricket
Cutegory Adult Males Adult Females Sub-Aduli* Males Sub-Adult* Females
Ditte of No of No of No of No. of No af Nou. ol
Collection Crickets Mitws NimeNe Crickets Mites Nm/Ne Crickets Miles Nin’Ne Crickets Miles Nii Ne
N Nis N. Nin N. Nu N. Na _
304,79 f 3 0.5 | 0 On 15 7 (1.47 9 \2 1.35
9.7), 74 W 143 13,3 8 62 7.75 16 87 S.dd 20 Hy 3.45
26.11.79 4 127 O07 4 2A 6.30 0 i} — 0 0 =
26.411,79 47 149 4.43 42 72 1,71 0 it) — " 0 -
QAV.TY is Is OR3 28 A O.K2 13) 0 _ i] u -
24iv.79 5 | 0.2 25 $ 20 | i) oO. | ) OO
Wao i) “ — ih 0 38 9 0,24 22 3 O59
8.80 2 u oo | a Hu 7 1 Old 4 4 0
7.80) It & 0.50 ia Z O18 b ti ind) + l Hae
480) 10 | Ot 37 13 (123 I u a 4 () Ul
Totils 11s 437 3,70 177 m9 115 34 ia i.24 63 99 157
* Ade Pane oubult 2 instars.
166
subserye an auditory function, allowing the larvae to
locate their hosts by a sonar technique. These setae are
venerally broken off in the older tie. fed) larvae. These
aujhors proposed chat this row of setae served no
further useful function after attachment to the host,
(This Would be consistent with the general finding that
such larvae feed Fully ona single host.) Additionally,
the tarsi of P pseudohunnemania (but only tarsus I
in P auditor) are highly attenuated, cach tarsus ending
in @ short pedicellus and three claws,
The arrangement of the setae was considered
suggestive “ol che setae of the antennae of male
Culigidac, which are known auditory organs”,
Ina tater paper Vercammmen-Grandjean er al, (1965)
discussed another form of setal modification in the
North American leeuwenhoekiid mite, Whartonia
glenni Brennan, 1962, which is an ectoparasite of two
species of bats of two families: Phyllostomatidae and
Embailuromdae. In this species of mite chere are greatly
elongated setae (“mastisetae”) on che larval tarsus ILE.
These anisotropic setae, identified az mastitarsalae,
have two conspicuous bends and points of weakness.
Most specimens of setae from mifes taken in the field
are broken (92.1%o of 441 setae), a situation analogous
to that seen in the genus Preridopus, These authors
speculate that such setae assist in locating sound-
emitting vertebrates, such as bats which emit their high-
pitched sounds even when roosting, when almost all
of the opportunities of attachment of these mites can
be conecived to occur,
Although these authors do not mention it, some
other prostigmatic mites have highly modified setae
which might conceivably serve as sound-detecting
organs. Thus in the family Ery{hracidae the genera
Fatoniana Cambridge, 1898 and Piflophus Berlese,
1916 have such upon the tibia IV of the adults (and
possibly also on genu TV), With regard to the possible
function of these setae, there termed “plumalae”, the
author (Southcott, (961a, p. 486) accepted the views
of previous students that these setse were used as sails
4s a means of progression. [t is now apparent that a
more detailed study of their function is warranted-
In the case of Trambella alpha larvae, the setational
pattert; does not suggest that the setae have any
possible specialized auditory function, However, the
alienuation of all tarsi does suggest that they could
function as sound receptor organs in host detection,
Such a funetion might be enhanced by the possession
of a single tarsal claw to each lee. The use by this larva
of a cricket as its host species. is possibly of significance.
Since many animals erriit sounds for a variety of
Purposes, such as territory-establishment, echo-
location of surroundings and prey, sexual attraction
and other functions, it would appear that the
development of host-locating organs by their potential
Parasites is worthy of a wider study,
REC. 8. AUST. MUS. 9Qilp 145-168
June, 1986
ACKNOWLEDGMENTS
Lam indebted to Mr Steve J, Davidson, Zoology
Department, University of New England, Armidale,
New South Wales, lor collecting and forwarding both
preserved and live larval mites and data for the present
Study, The South Australian Museum has made
available the holotypes of Trornbella warregensis Hirst
and VT. uwdelaideae Wom,, and a paratype of
Womerslevia minuta Radford, for study. | thank Mr
L, G. Veitch, Prinvipal Research Scientist, CSIRO
Division of Mathematics and Statistics, Glen Osmond,
South Australia, for aid and advice in siatistical
matters.
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THE GENUS ODONTACARUS (ACARINA: TROMBICULIDAE).
Il. OBSERVATIONS ON THE LIFE HISTORY AND MORPHOLOGY OF
ODONTACARUS SWANIN. SP., AND RELATED FORMS
BY R. V. SOUTHCOTT
Summary
The ovum, larva and adult of Odontacarus swani n. sp. are described, being the first recorded
correlation between adults and larvae in Odontacarus. However, as is customary in this group, a
larva has been selected as the type.
A key is given for all species described as larvae from Australia, New Guinea and South East Asia.
THE GENUS ODONTACARUS (ACARINA: TROMBICULIDAR). Ul, OBSERVATIONS ON THE LIFE
HISTORY AND MORPHOLOGY OF ODONTACARUS SWANI N. SP, AND RELATED FORMS
by
R, V. SOUTHCOTT
Honurary Research Associate, South Australian Musetim, North Terrace, Adelaide, South Australia S000
(Manuscript accepted 23 January 1985)
ABSTRACT
SOUTHCOTT, RK. V. 1986. The genus Odentacarts (Acurimis:
Trombieulidae) UH, Observations on the life history and
morphology af Odortacurus swarl n. sp. and related forms.
Ree. §. Aust. Mus. V(12): 169-200,
The ovum, larva and adult of Qdentacarus swani
n, sp. are described, being the first recorded correlation
between adults and larvae in Odontacarus, However,
as 3s customary in this group, a larva has been selected
as the type.
A key is given for all species described as larvae from
Australia, New Guinea and South East Asia.
A correlation analysis is made of the dimensions ol
the chilinized parts (scutum, setae and leg segments)
of the (ype series of O. swany, A principal component
anulysis defined two components, both of which
inchide variates derived trom the dorsal scutum and
the lez segments.
A similar analysis is made for the larvae of two other
species of Odontacarus: O. atherlonensis (Womersley,
1945) from north Queensland, and QO. mecullochi
(Womersley, 1944) recorded from north Queensland
(and Papua New Guinea), In each of these species two
principal components of correlation are defined which
are nol completely duplicated in the other two species.
All four of these components include variates based
upon scutal dimensions, three of them include yariates
based on leg Segmental lengths, and two of them (each
of these latter being the second principal components)
include variates based on lengths of idiosomal (body)
setac. Some components contain also variates derived
from body hair dimensions,
It is concluded that no standard components can be
defined and in general there is not a high degree of
correlation between the various standard (and other)
variates customarily used in the classification of the
larvae of the genus QOdantacarus (family
Trombiculidae), in agreement with my earlier study of
Trombella where another group of correlated variates
was detined. This contrasts with similar analyses of
hard structures (such as bone) of vertebrates.
Additional morphological details are given for the
Australian species of Odontocarus, O. cookl (South-
cott, 1957), O. langani (Southcott, 1957), O, mathewi
(Southcott, 1957), and O southcotti (Womersley, 1944)
to help construct the key of the larvae and for O,
barrinensis (Womersley, 1945), Amended collection
data are provided for OC. seutheorii.
A key is given for the adults of Odontecaruy (all
known adults are Australian) and the adult QO swan
is described,
Acomatacarus reienius (Banks, 1916), known fren
only the original material from Victoria, is made the
type of Scopitrombium n. gzen., in the Lecuwenhoe-
kiinae. A lectotype is designated, and the species
redescribed.
Revised keys are given for the subfamilies of
Trombiculidae, and for the genera of the subfamily
Leeuwenhoekiinae for both adults and nymphs.
INTRODUCTION
The genus Odontacarus was founded by Ewing
(1929b, pp. 22, 188) with the following definition:
“Bach chelicera with a row of backwardly directed
teeth on the upper margin of chela, and some upturned
teeth on the lower margin (Fig. 16). Palpal claw
bifurcate, the two divisions unequal. Dorsal plate
without median anterior process and with only five
setae in addition to the pseudostigmalic organs, which
are flagelliform and. pectinate.
Type: Thombicula dentata Ewing”,
Ewing’s species was based upon two larvae:
“Described from two specimens; one, the holotype,
fram Sonora, Texas (Bish. No. 10681), taken by O. G-
Babcock, December 17, 1922 on a white-tailed deer and
the other from Balboa, Panama on a cotton rat by
L. H, Dunn”. He figured (1925a, p. 258) only the right
cheli¢era of the holotype (Cat. No. 888, U.S.N.M_.).
In fo'inding Odontacarus Ewing (1929b, p. 183)
compared it with the larvae then attributed to
Trombicula Berlese 1905 (described originally from the
adult) and to the larvae of Schdngastia Qudemans
1910, and saying “only the type species included”, Le.
T. dentata Ewing, 1925. He made nv reference to the
larval genus Leeuwenhoekia Qudemans, 1911.
In 1942 Ewing revised the taxonomy of some of the
American mites (‘chiggers’) then placed in the
subfamily Trombiculinae Ewing, 1929 (1929b, p. 22),
He (1942, p. 489) decided to divide the genus Leex wert
hoekia, and erected two new genera, Comatacarus and
Acomatacarus. The latter he founded for the larval 4.
arizonensis Ewing, 1942, defining it as:
“Most nearly related to Comatacarus |Ewing, /oc.
cil, p, 489] but differing from it as follows: Chelicers
with a row of teeth on upper margin instead of a single
170
jooth and both the first |palpal femorata] and second
[palpal genuala] palpal setae simple instead of one or
both being plumose or barbed".
The principal distinguishing characters of Conara-
carus (from Leeuwenhoekia) were (Ewing, 1942, p. 489)
‘in having the posterolateral setae of the dorsal plate
of the usual form and similar to the anterolateral,
instead of heing clavate, and in having a large number
of sessile setae on the dorsum of the abdomen, inslead
of a small number situated on rhe tubercles”,
Ewing (1946, p. 436), in a further revision of the
trombiculid mites (raised to the family Trombiculidae
by Ewing in 1944), redefined Odontacarus and stated:
“Qdontacarus is telated to Acomatacerus Ewing,
1942, from which it differs in having a single median
seta on the dorsal plate instead of two submedian setac.
lL is also related to Endotrombicula Ewing, 1931 [See
Wharton and Fuller (1952, p. 72) and Audy (L954, p.
160) for opinions on the present status of this
genus.],..%; he included in Odentacarus also O,
australis (Ewing, 1929), which was described (1929a,
p. 10) trorm several specimens collected on the lizard
Tropiduras peruvianus at Verrugas Canon, Lima, Peru,
15,iv.1928, R. C. Shannon, Cat. No, 987, U.S.NM.
Brennan (1959) stated that Trombicula dentata
Ewing, 1925 was based on a single teratological
specimen, possessing a single anteromedian scutal seta
instead of The mormal two, commenting that this
holotype specimen is in a distorted and damaged
condition, with most diagnostic features hard to
determine. Nevertheless he stated that “The AM [seta]
is slightly to the right of center”, and declared that
“here is no doubt whatsoever as Lu its generic status”,
even though its “specific identity ... is in question”.
With regard to the second larval specimen assigned
by Ewing originally to 7) dentefa, Brennan (1959, p. 2)
has commented that it is “damaged beyond recognition
and reclaim . .. It seems incomprehensible, hawever,
that this specimen could possibly have been regarded
as conspecific with the holotype”.
As far as O. australis (Ewing) is concerned, Brennan
(1959) redescribed the holotype briefly, without figures,
stating that this specimen also had a teratological
dorsal scutum—"In each [of the two holotypes}, the
left AM is lost; thus the specimens are freaks”. Two
other specimens, each labelled by Ewing as # “catype”,
were identified by Brennan (/ec, cin as Eutrombicula
alfreddugesi (Qudemans, 1910),
However, eventually O. denfatus holotype was found
not to be teratological alter all; Goff er al. (1972) were
able to see twa AM seta bases, nsing phase microscopy,
although the setae were not altached (Goff and
Loomis, 1977),
Wharton and Puller had earlier (1952, p. 103)
realized the probable synonymy of Gdentecerus wud
Aconmracarus, but had commented, with reference to
Odontacarus’ “Specimens in existence art loa badly
RECS, AUST. MUS. 19(12): 169-200
June, 1986
damaged to study satisfactorily’, These remarks
applied lo both O. denfaras and OQ, australis.
it appears strange that Ewing had not at this stage
seen the characteristic feature of the leeuwenhoekiinae
shield, ari anteromedian tongue-like projection to the
dorsal sculum, in fact stating specifically that it was
absent (see his definition of Odonlacarus above),
Although Oudemans had described this feature in
Leeuwenhoekia verduni (OQudemans, 1910) as early as
1910 and 1911, and again in his 19]2 monograph,
apparently it was not observed by Ewing and published
until his 1942 paper for Acomatacerus and
Comatacarus. Hawever, in his key to the larval
Trombiculinae, Bwing (1938, p, 291) showed that he was
aware of the anteromedian process in Leewwenhoekia
Oudemans, bul again failed to record it for Odon-
tacarus.
Comalacarus Ewing was reduced fa subgeneric
status (in Leeywenhoekia) by Wharton ef al. (1951),
followed by Wharton and Fuller (1952, p. 96), Gould
(1956), Loomis (1956), and Finley (1958), but restored
to full generic status by Reed (1973) and Goff and
Loomis (1974),
Acomatucarus Ewing, 1942, was resurrected by
Vercammen-Grandjean (1968, p, 122), and in its new
veneric status was divided into subgenera by the sare
author—Acomatacarus subgen,, and Oroehlorus
subgen. (Vercammen-Grandjean, /oc. cit).
Odontacarus has been divided inlo the subgenera
Tarsalucarus and Leagenius by Vercammmen-Grandjcan
(1968, pp. 120-121), Acomatacarus Ewing, 1942 includes
several subgenera: Yenodontdcarus Loomis and Gott
(1973) (a new name tor Xenacarus Greenberg, 1951, on
grounds of pre-occupation hy Yenavarus Kisbida,
1925), Meteearus Vereammen-Grandjean, 1956 and
Orochlorus Vercamimen-Grandjean (1968, (9. 122, The
last has been synonymized with 7Zarsalacarus by Gott
and Loomis (1977), since the type species,
Acoimatacarus micgheneri Greenberg, 1952, actually
belonged ta Qdontacarus, with the characters of
Tarsalacorus, which had page priority.
The genera Austracarus Lawrence, 1949 arid
Ayrecarus Lawrence, 1949, which were placed as
subpeners of Acomuraearys by Vercammen-CGrandican
(1957), are now considered by the sanve author (1¥73a},
without giving any explanation, ta be full genera in the
Leewenhoekiinae,
With the restriction of the genus OQdonfucarus and
its separation from Acomaflacerus, the only subgenus
of immediate concern here is Leozonius, in which the
Australia-New Guinea species considered here and O.
audyt (Radford, 1946) are placed (see Vercarnmen-
Grandjean, 1968; Goff, 1979a, b).
The genus Odontacarus as at present accepted is a
widespread one, with species in North and Sauth
America, Asia, New Guinea and Australia. In ihe
present paper the term will be Used in the sense of Gott
(1979a, b).
ODONTACARUS SWANS N.SP. "1
Most studies of Odontacarus species have been based
on larvae, as in that instar they are readily collectable
as parasites upon vertebrates, including, in some
instances, man, At times they are captured from a free-
living stale, e.g. upon cards, boots, fence posts, etc. In
a few Australasian species rearings have been achieved
from larvac to nymphs. These were Q. ausiraliensis
(Hirst), from Hollandia, former Dutch New Guinea
(now Dyayapura, West Irian), reared by C. B. Philip,
and O. longipes (Womersley, 1945) (synonymized with
O, novaguenea by Goff (1979a, p. 149)) from the
Dobadura area of New Guinea, and O. nova-guinea
(Wom,, 1944), from “New Guinea”, these Jas( two
having been reared by G. M, Kohls. These rearings were
recorded by Womersley (1945), and allowed him to
deline the post-larval characters of the genus (then
Acomatacarus), Later larva to nymph rearings have
been achieved by Domrow (1956) for O. australiensis
in Queensland, and by Nadchatram (1963) for O. auayi
(Radford, 1946) in Malaysia. The experimental rearings
reported in Womersley (!945) allowed him to assign
four species of adult trombidioid mites to the genus,
these being O. atfolas (Banks, 1916) from Sydney, New
South Wales, O. dromas {s. str.) (Womersley, 1939),
from Long Gully, South Australia, O. patrius (Wom.,
1945), from Murray Bridge and Burra, South Australia,
and QO, retentus (Banks, 1916), from Lal Lal, Sea Lake
and Ocean Grove in Victoria. Despite the widespread
geographical distribution of the genus, and the fact thal
the larvae may be at times locally numerous, so far all
adults that have been recognized as belonging to the
genus are of Australian origin. Hitherto there has been
no record of a correlation by rearing between an adult
at this genus and a larva.
{In 19441] was able to make records of oviposition
by adults of O. swani n. sp., collected in north
Queensland, and to rear larvae from these eggs, The
present paper describes these experiments and also the
adult, larva and ovum.
Since the great majority of taxa among the Leeu-
wenhoekiinae and other chigger mites of the family
Trombiculidae have been based on the larvae, one of
the reared larvae is selected as the type of the species.
J have taken the opportunity to examine the
taxonomic status of some species of the genus from
Queensland and New Guinea, and to give a key to the
larvae of Odentacarus known from New Guinea and
South-Bast Asia, However it should be pointed out that
this paper does not undertake a general taxonomic
review of the larvae of Odonlacearus, subgenus
Leogonius, bul considers only what is required to
establish the new taxon proposed, and for the majority
of the species considered in the key it relies on the data
of previous guthass.
One species of adult allotted to this genus by
Worersley (1945) is markedly distince from the others,
this being @ retentus (Banks, 1916), and for ic a new
genus Scopitrombium is proposed,
Revised keys are given for the classification, for
adults and nymphs, of the subfamilies of the Trom-
biculidae, and the genera of the Leeuwenkoekiinae,
All measurements are given in micrometres (um)
unless otherwise stated,
SYSTEMATICS
Odontacarus Ewing
Synonyery
(Note: Only a partial synonymy is given here. For
earlier and more complete lists of synonymies, see
Wharton and Fuller (1952), Audy (1954), Goff and
colleagues (1974-1979)),
Odontacarus Ewing, 1929b, p. 1B8; 1931, p. 6; 1946,
p. 436. Fuller, 1952, p. 228. Wharton and Puller, 1952,
p. 103 (ad part.). Vercammen-Grandjean, 1968, p. 120,
Loomis and Crossley, 1963, p. 38L. Nadchatram, 1963,
p. 535, Southeott, 1973, p. 46; 1976, p, 139; 1978, p. 16.
Reed and Brennan, 1975, p. 6. Goff and Loomis, 1977,
p. 370. Goff and Brennan, 1978, p, $0, Goff, 1979a,
p. 143; 1979b, p. 140.
Ceeuwenhoekia (Qudemans, [911) Hirst, [925,
p. 150. Womersley, 1934, p. 217; 1944, p, 103.
Womersley and Heaslip, 1943, p. [4i. Taylor, 1946, p.
227 PF.
Acomatacarus Ewing, 1942, p, 490. Womersley, 1945,
p. 98. Taylor, 1946, p. 225. Greenberg 1951, p. 525; 1952,
p. 473. Fuller, 1952. p. 229. Gunther, 1952, p, 39.
Wharton and Fuller, 1952, p, 94. Audy, 1954, p. 164.
Baker ef a/, 1956, p. 104. Domrow, 1956, p. 150.
Southcott, 1957, p, 146; 1973, p. 103; non Vercammen-
Grandjean, 1968.
Odontacarus swani no. sp.
Description of adult female (principally fram
specimen ACB240A, stide-mounted, somewhat com-
pressed; but supplemented by other specimens)
(Figs, LA-E; 2).
Colour in life red. Idiosoma (Fig, 1A) af normal
ovoid shape for a trombidioid mite, without waist, and
without division between prosoma and opisthosomia;
length on slide about 1400 ym {all measnrements in
micrometres), width about 900.
Prosoma carries dorsally a well-chitinized crista, with
an enlarged posterior end, carrying two sensillary setae
(Fig. 1B), The erista tapers anteriorly, to a blunt-
pointed rod, embedded in an arrow-shaped area of
chitinization (here named the “sagicla” since the term
“tectum” used by Crossley (L960) for this part of the
sculum is considered less appropriate.). The crista is
surrounded by the normal dorsal setation, sparser
around anterior end. The sagitta carries two long, thin,
ciliate setae (Figs. 1B, IE). Crista 343 long from its
blunted anterior point to rear edge of the posterior
sensillary boss; 423 long to tip of sagitta. Behind
posterior sensillary boss the crista continues for a
further 70, to end in a blunted point. Sensillary boss
94 across, distance between centres of sensillary setae
bases (ie, “SB") S57. Sensillary setae slender, long,
172 REC. 8. AUST. MUS, 19(12); 169-200
June, 1986
FIG. 1. Odontacarus swani sp. nay, Adult female, specimen ACB240, parent of the holotype. A Entire specimen, slide-mounted, seen
in transparency, B Crista and adjacent area. C Dorsal setae at posterior pole of idiosoma. D Left palp, medial aspect. E Chelicerae,
(All to nearest scale.)
filiform, 290 long. The chitin of crista, including that
of the area between the sensillary sockets, punctate.
The whole of dorsum of idiosoma is densely covered
with setae mounted on short papillae. The setae tend
to be separable into two distinct types—a shorter burr-
like, spindle-shaped, pointed, heavily ciliate group,
mostly 27-31 long, and interspersed among these are
longer, swordlike setae, in which the proximal part of
the scobillum is somewhat expanded and ciliate; but
the distal half is curved, swordlike, curved and pointed,
with adnate cilia along the edges: a batch of these setae
at the posterior pole of the idiosoma is illustrated in
ODONTACARUS SIFANI NSP. 173
Fig. JC. There is, however, some intergrading between
these two main types of dorsal idiosomalae; at the
posterior pole the longer swordlike setae are 64-73 pm
long, the shorter spindle-shaped setae 25-42 long. At
the centre of the dorsum the longer setae are 56-95
long, the shorter setae 27-44 long.
Eyes are not visible in any of the specimens of adults
available, and it must be concluded that they are absent
(despite being present and normal in the larvae),
Ventral surface of idiosoma is covered with a dense
selation Of spindle-shaped, tapering, heavily ciliate
setae, These tend to be uniform, similar to the shorter
dorsal idjosomalae, except (owards the posterior pole,
where they are interspersed with swordlike setae similar
to those of (he adjacent dorsum,
Genital aperture in usual position, about 390 long
by 2455 across, with three oval suckers along eaeh side,
Anus normal, 9! long by 46 across.
Legs long, but robust for a trombidioid nite,
moderately chitinized; trochanters generally rounded,
basifemoral, {clofemoral and genual segments
presenting as truncate cones, tibiae and tarsi approxi-
mately cylindrical. Leg lengths (including coxae and
claws): | 2660, 11 1760, [11 1770, 1V 2640. Each tarsus
bears two claws, Legs well covered with numerous fine,
pointed, lightly ciliate setae (scobalac), these being
inlerspersed in the distal segments with pointed,
specialized sensory setae (spinalae, or eupathidalae).
Tarsus | an elongate oval, 634 long by 214 across,
wider than tibia 1, which is 534 long by 178 across;
tarsus IT 397 long by 93 high; tibia TI 313 long by 121
high; tarsus 111 399 lang by 85 figh; Ubia 11 352~128
similarly; tarsus 1V 523114 similarly; dbia 1Y 627 «157
similarly, For other leg morphometri¢ data, se¢ Table I.
[ABLE L MEASUREMENTS (am) OF SOME CHITINIZED
PARTS OF EIGHT ADULTS OF GDONTACA RUS SHANI N. SP.
Charueter Range r Mean SLD.
Crista jength! 277-343 8 321,75 57,42
SH 44-63 & 52.87 23.89
Tal’ 585-634 7 603.14 4h. 36
Til 499-545 7 $17.29 46.17
Gel 332-368 7 344,14 42.45
Talv* 435-523 7 4838.71 72,9]
THY 506-627 7 574,00 44,95
GelV 370-427 7 400), 29 53,27
' Measured tram anterior end of chitinous rod to pasterlor pole of
sensillary bopy,
Qmiming claws and pedicle
Palp! comparatively slender, well covered with ciliate
setae (see Fig, 1D), Many of these scobalae along the
dorsal (jc, extensor) margins of the palpal genu and
trbia are cihate along their distal and outer aspects, the
cilia broad and flattened, curved, uniform and almost
apposed, as though capable of being touch receptors
utilizing a mechanism akin to stridulation; for these
setae the term peetinala is proposed (plural; pectinalae).
Palpal tibial claw stout, blunted, with four accessory
claws (thickened scobalae) on the medial to dersal
asnects,
Palpal tarsus an elongate ovoid, with numerous
ciliate setae (scobalae); (hese setae tend to resemble the
palpal bial and genual pectinalae, but are shorter.
Among them are also nude setae (spinalac) and pec-
tinalae. A cluster of shart sensory setae is present at
the tip of the palpal tarsus,
Chelicerae robust; fangs curved, pointed, aboul 135
long, with a row of about 1 fine retrorse teeth along
distal flexor edge; extensor edge smooth, Chelicera
basis about twice as long as fang. Rostrum (cone) of
mouthparts with mrumerous ciliated, tapering setae.
Taxonomic placing: In the key of Womerstey (1945,
p. LO) this adule keys to the caption for O, drarrus
(Wom., 1939), From that species O, swaei may readily
be separated as follows:
Colour of adult mite red. Longer dorsal idiosomal
setue with long cilia in the basal half, and with adnate
cilia, scarcely detectable, in the distal half of the seta.
Shorter dorsal idiosomal setae strongly ciliate, spindle:
shaped, tapering ....... QO swani n. sp.
Colour of adult mite white in life. Longer dorsal idis-
somal setae ciliate throughout length. Shorter dorsal
idiasomal setae slender, umlormly cibate —...__ -
. O a@romus (Wora,)
oF urther comments on the taxoniomy of adult leeu-
werihoekiine mites are made later in this article,
Description vf Ovum
(Fig, 2, 3A4-D) (from mounted material)
Colour in life not observed. Ovum nearly spherical
fo somewhat spheroidal, about 250-350 long in longest
diameter. Qvum has the usual lightly pigmented outer
layer (chorion), which is smooth, lacking any striate
markings in the available material, Minute tuber-
culations occur on the parts of the mounted material,
but these could be, at least to some extent, results oF
compression in eclosion, or rupture by other means.
The sole embryo available (Fig, 31D) is about 280 long
in its greatest diameter, by about 220 wide in its shortest
diameter. It contains a large number of rounded
granules, presumably of yolk, mainly concentrated in
the region opposite to that in Which the limbs are
developing, Two annull (seta bases) are present in (he
specimen, and two tooth-like structures, possibly shell-
piercers in eclosion (“4” Fig. 3D). From one annulus
there is a Vague indication of a developing seta shaft.
On the developing limb segments are smooth, rounded,
low tubercles, similar to those recorded for Trombelfiar
aipha developing nymph (Southcot! 1986a, Fig. SA)
One chorion (Fig, 3C) is unusual in being covered
randomly with small circular or oval lacunae, In
rupluring, presumably from eclosion, some of these
lacunae have been cut across transversely; it is clear that
they were present before rupture, and presumably in
life. No evidence of mould filaments or other possible
chulinolytic agents can be seen near them, and their
origin is unknown, Linder polarized light a few of them
174 REC. S. AUST. MUS. 19(12): 169-200 June, 1986
FIG. 2. Odontacarus swani sp. nov. Adult female, specimen ACB245, seen in transparency. The idiosoma contains nine eggs, of varying
sizes, some having been disrupted by the mounting process.
ODONTACARUS SWANT N.SP.
175
FIG, 3. Odontacarus swani sp. nov. Ova, slide-mounted specimens. A Ruptured ovum, showing chorion. B Chorion of another ovum
split into three pieces, and possibly some of the developing embryo. C A ruptured chorion with multiple lacunae (see text), D Developing
embryo, shed from its chorion, with early development of limbs and other structures: a annulus or seta base, t tooth-like projection
(see text), All specimens shown in transparency. (All to scale shown.)
show a typical Maltese-cross figure, indicating that
there is possibly a layer of optically active material at
the base of the lacuna, perhaps an artefact of the
mounting. The chorion shows only weak evidence of
optical activity under crossed polarizers.
Description of Larva (from reared specimen
ACB240B, holotype, and supplemented by other spe-
cimens) (Figs. 5A-D, 6A-C),
Colour in life red. Length of idiosoma (mounted on
slide) 207, width 197; total length of animal from tip
of cheliceral fangs to posterior pole of idiosoma 293.
Dorsal scutum slightly wider than long (including
nasus). Although most authors measure the length of
the dorsal scutum (“S.D.” or shield depth) of
Odontacarus larvae (and other genera of the Leeuwen-
hoekiinae in which there is a distinct beak-like “nasus”
to the scutum) excluding this “nasus”, a comparative
study of allied forms, e.g. Neotrombidium larvae, as
well as larval Trombe//a (see Southcott, 1986a), shows
that the “nasus” may range from a broad continuation
of the scutum, without clear point of differentiation,
to a distinct and sharply demarcated beak. The length
176 REC. §. AUST. MUS. 19(12); 169-214)
FIG, 4, Conventions of measurements of dorsal scutiim of a leeu-
wenhoekiinae larva, as used in the present article (see (exe).
(L) is taken here as including the “nasus”. However,
in order to make the measurermenls comparable
between different workers, measurements are offered
here of the shield length including “nasus” (L.) and
without “nasus” (LB) (see Fig, 4 for explanation),
Nasus well-developed, running into more or less
straight (slightly sinuous) anterior margin; lateral
borders slightly convex; posterolateral convex, of
varying curvature; AM setae tapering, pointed, lightly
ciliate; AL and PL similar. Sensillary setae normal,
ciliate in distal half or third. Sensillary sockets about
level with PL bases, sloping obliquely anterolaterally,
and with well-marked margins (“lids”).
The standard (and other) data for this specimen, as
well as other reared material, and the Type series and
other identified material in the South Australian
Museum Collection are given in Table 2.
Eyes 2+2, sessile, near PL angles of dorsal scutum,
oval. Anterior eye with maximum diameter of cornea
16, posterior |4,
Dorsal idiosomal setae long, moderately ciliate with
lightly projecting barbs; setae arranged 6, 12, ..., in
vague transverse rows, total in ACB240B, 57.
Venter with a pair of pointed setae between coxae
IT}, ciliate, 33 long. On opisthosoma ventrally about
37 further setae, pointed, well ciliate, 27-42 long, the
posterior selae longer and tending to be blunted at the
tip, resembling the posterior dorsal idiosomalae.
Coxalae 2, 1, 1. All coxalae long, pointed, ciliate;
lateral coxala | 51 long, medial 46, coxala 11 44, IIL —
(missing in holotype; 58 long in ACB240C),
Legs normal, 1379 long, Ll 322, LIL 380 (all lengths
include coxae and claws), Chaetotaxy of legs as figured,
Leg scobalae normal, pointed, ciliate. Trochanteral
formula L, t, 1; femoral 6, 5, 4. Leg specialized setae
as tallows; VsGel.70pd (“microseta”), YVsTil.87pd
(microseta"), SoGel 36nd, SoTIL.90d (i.e. slightly distal
to Vs), VsGell,70d, SoGell.30d, SoPilL.45d, SaTiL9td,
SoGellLL29pd. Tarsus | atid 1] have each a large dorsal
June, 1986
TABLE 2. MEASUREMENTS (um) OF REARED
ODONTACARUS SWANI N. SP. LARVAE
Character Holorype Range n Men S.D.
AW 65 68-73 8 69.13 3.271
PW 78 77-83 8 80,88 3.27)
SB 27 25.30 & 26.75 1.669
ASE 4y 49-S7 g S313 2.949
PSB 33 22-33 8 3025 3.536
L $2 77-87 8 83.38 3.462
LA 18 17-22 8 19.13 1,727
La 64 55-49 BR 664.25 4.432
LN 25 27-35 8 W285 = 3.495
Ww 6s R597 BR O62 3,739
A- 25 24.29 8 26.13 1.582
AM 41 35-43 8 38.88 2.900
AL 47 46-55 B 4863 3,159
PL 55 55-62 8 59.50 2,507
AMB 1 I1-13 8 12.00 (1.926
Sens 5\ 48-55 _ — —
PW/LB! 1.22 120-154 = = -
DS 46-47 (36-44)4(47-51) — -
(ids DS 36-44 (3644)-(44-51) 8 46257 2.1887
PDS 34-47 (38-464(47-51) § 49.13" 1.7277
Gel 46 41-51 R 47.25 3,012
Til 55 54.57 f 55,25 0,886
Gell a8 38-45 8 40.63 2,264
nu 46 44-54 8 47,50 2.264
Crelll 43 43-48 BR 46.15 1,38
Till 57 57-63 & 59.63 2.066
' This is the PW/SD of Womersley (1944, 1945) and the PW/L of
, Greenberg (1951) (see Fig. 4, and text explanation).
* For the maximum yalues of each variate,
solenoidala, FaTall present (see figure). Normal Sofalll
present, Tibia 11] with two mastisetae (=mastalac):
MaTilll.73d, MaTilll.76pd. Tarsus Hl with one
mastiseta: MaTilll.33pd. (In these estimates tarsal
length excludes claws and pedicle) Tarsus J 86 long by
32 across, Tarsus II] 85x24. Other specialized Ieg setae
as figured. Other leg measurements as in Table 2,
Tarsal claws normal, falciform, slender, with strong
cilia (onychotrichs) on anterior and posterior; middle
claw longer and more slender than the neolaterals, with
weak cilia.
Gnathosoma normal. Combined chelicerae bases 84
across, chelicerae 89 long from tips of fangs to
posterior pole of chelicerae bases, Cheliceral fang stout,
curved, pointed, with three of four retrorse teeth along
(voncave) flexor (dorsal) edge, and 4-5 tuberculalions
or blunted teeth on (convex) extensor (veritral) edge.
Galeala nude, 22 long. Gnathobagal setae (palpal
coxalae) curved, pointed, well ciliate, about 24 Jong.
Palpi compact. Palpal formula 1, 1, 3, 8, with palpal
formula B(b), N(7b), BNN, So+7 (Bor N), as figured.
Palpal tibial claw trifurcate. No supracoxala to legs or
palpi.
Origin af Adult Material Studied
[ collected ten adults in damp soil in rainforest at
Mt Hypipamee, Atherton Tableland, Queensland on
26,%, 1944,
Each was placed in a small tube with some damp
soil from the capture area, and observed periodically
(albeit somewhat irregularly, as | was engaged in
military duties). Laving larvae were observed in two of
the tubes from 20-24.911.1944 using a 28% Zeiss
microscope ocular reversed as a hand lens; in a third
ODONTACARUS SWANI N.SP. 177
FIG. 5, Odoniacarus swani sp. nov., larva, holotype, ACB240B. A Dorsal aspect, legs on right hand side omitted beyond trochanters.
B Cheliceral fangs and galeala from above (distal half of cheliceral fangs shown in transparency). C Posterior dorsal seta (x in A).
D Posterior ventral seta (y in Fig. 6A). (All to nearest scale.) In this and in subsequent figures the A sign indicates the seta is shown
in both dorsal and ventral figures.
178
REC. S. AUST. MUS. 19(12): 169-200 June, 1986
|
A |
| PaScTi3
|
d
OO
“PaScTi2 :
ie)
20
oS
100 ~
FIG. 6. Odontacarus swani sp. nov., larva, holotype, ACB240B. A Ventral view, legs on right hand side omitted beyond trochanters. B
Right palp, dorsal aspect. C Right palp, ventral aspect (B, C to scale on right). D Claws of leg LU, further enlarged (not to scale).
ODONTACARUS SWANI N.SP.
9
TABLE 3, DETAILS GF REARING EXPERIMENTS COMMENCING WITH A BATCH OF ADULTS OF ODONTACARUS SWANI
N..SP. FROM MT HYPIPAMEE, QUEENSLAND, CAPTURED 26 OCTOBER, 1944
Serial No: Sex Eggs Larvae Larvae Adult dead Remarks
ACR observed observed dead
238 — — — 22.x1.44-20_ sii .44
239 F (S.vii,45)* 20-23.xii.44 24-30.x1i,44 22.51.44-26.xi1.44
240 F (3.vii.45)* 20-23.xit 44 24-30.511-44 22_xi 44-20.) 44
241 — — — 2-6.xi44
242 a oa _ 22.xi.44=20.xii,44
243 F a _— — 17 2.44-22.xi.44 Gravid, laid no eggs, buy
body contains abour 50 egps
244 F (21.v.48)* (21-v.48)* — 22.61,44-20. 211.44
245 b - — _ da 6.41.44 Gravid (9 eggs), laid noe
246. — — — 22.81,.44-20, xii, 44
247 _ a — 22x. 44-20, xii 44
* These eggs oF larvae were found i) (he dried tubes, months or years after (he attempted rearings had been concluded
tube examined some years later (long alter all the tubes
had been allowed to dry out), larvae were also found,
dead and dry. Although living eggs were not observed
in these tubes, egg remnants were eventually found
when the opportunity to examine the tubes in detail
with a microscope occurred. Two female adults, on
being slide-mounted, were also found to contain eggs,
although no eggs or larvae were recovered from their
tubes, The details of these experimental animals are
summarized in Table 3,
Taxonomic Placing of Odontacarus swani nm. sp,
The larva of this species ts distinguished from others
of the genus by the following combination of
characters: (1) palpal tibial claw three-pronged, (2)
galeala nude, and (3) about 57 dorsal body setae
present, (4) two mastitibialae I] present, (5) one
mastitarsala LEE present, palpal genuala (PaScGe) and
the two palpal tibialae (PaScTil and PaScTi3) almost
nude. with only faint indications of barbs, cheliceral
fangs with a row of 3-4 dorsal and 4-5 ventral denticles.
Characters (1), (2) and (3) alone are sufficient to
distinguish this species from all others of the Australa-
sian to South-East Asian region (subgenus Leogonius),
The diagnostic characters of the adult have been
Biven above,
Nomenclature
This species 1s dedicated to Duncan Campbell Swan
(28.x1.1907-26,x11.1960), entomologist, who made field
observations on the ¢listribution of chigger mites in
South Australia (see Southcott, 1982).
Classification of the Larvae of the Australian, New
Guinean and South-East Asian Species of Odontacarus
About 20 species of Odontacarus have been
described from Australia and New Guinea (including
West Irian and Papua New Guinea), A single further
species, O, dadyi (Radford, 1946) has been described
fram South-East Asia, one, OL gyennodactyli (Ewing,
1925) (1925b) from India, and one, QO /ygasamae
(Dumbleton, 1947), From New Zealand. Since only
limited keys to the classification al these species have
been offered since Womersley’s (1944, 1945) papers, an
attempt to key the species of this region is offered here,
The most detailed studies of recent years for this
region have been done by Goff and Loomis (see refer
ences), who unfortunately deal only with [he species
from New Guinea, and incidentally with a few north
Queensland species, Where these occur also in New
Guinea,
In altempting to classify the larvae here allotted to
O. swanin. sp., as well as various other species from
northern Queensland and New Guinea, J have experi-
enced difficulty in using one of the main characters
used by Goff (1979a, b) in classification, this being
deciding on the number of setae on the dorsum of the
abdomen in the “first post-humeral row”, i.e, the row
of setae immediately behind the dorsal scutum, but
excluding the two (or four) long “humeral” or
“scapular” setae, These remarks apply not only to the
specimens of OQ. swani described here, but also
members of the type series of O. athertonensis
(Womersley, 1945), and of O. niceullochi (Worn., 1944),
which were used by Goff (1979a) in drawing up his key
to the species which occur in New Guinea.
For this reason, it has been decided not to use the
number of the setae in this row as a key character. It
should also be noted that students of the larvae of the
Trombidioidea and Erythraeoidea usually experience
difficulty in deciding which setae to allot to different
rows, as the arrangement is commonly irregular, and
the allotment of the setae is largely a mutter of
guesswork, On the other hand, the total number of
dorsal or ventral idiosomal setae appears (o be free
from these defects, even if the counting of them is ar
times tedious.
The following key to the larvae of Odentacarus for
this region is therefore proposed, based to a large exter!
upon already published descriptions, but supplemented
with some new data.
KEY TO THE LARVAE OF ODONTACARUS
(SUBGENUS LEOGON/US) OF AUSTRALIA,
NEW GUINEA AND SOUTH-EAST ASIA
L. Palpal tibial claw with morc than three prongs -
Palpal tibial claws with three prongs .-.--.--.
a) fs
180
10.
il.
L3.
14.
16,
17.
_ AL setue longer than PL
REC. 8. AUST. MUS. 19(12) 169-200
Palpal tibial daw with seven prongs
- , O. Hannae Goll, 1979
Palpal tibial claw with four prongs dus wo et
a os) peered eo eee ime ‘
QO. nadchamiont Gol, 1979
AL ‘setae “shorter ‘than PL. 4
Masuitibialae II] absent, Two mastitarsalae 111 present.
Dorsal idiosomal setae about 110 ,
eee, eee FP Oe Ste ee 6 QO, mitchelli Goll,
One or more : mastitibialae [Ul present .
One mastilibiala [1] present, Dorsal idiosomal =n
about 230 O. unisetosus Gott, 1979
Two mastitibialae [1] present to tibial IL... -, 6
Galeala nude. Palpal seta] formula B/B/BNN/So + 7B-
About 120 dorsal idiosomal setae present ,.....,.,
ne oC aieqnale te O. friselasus Goll, 1974
Galeala barbed . _ ae 7
Dorsal idiosomal ‘setae number about 54, “Ventral
idjosomal setae abou! 78. Palpa) setal formula B/B/
BBB/So + 7B. Cheliveral se 4-5d/d4-7¥ . 2.
O, zegleri Gott, 1979
Dorsal idiosomal setae number about 52-90, Veniral
idjosomal setae number about 46-54. Palpal setal
formula B/B/Bbb/So4 7B. Cheliceral denticles 7d/
G-By. cesses anne . O. aud? (Radford, 1946)
(Occasional apecimeris ‘of ©. wudyi have only three
palpal uibial prongs (see ncaa 1963)
Mastitibialae IIE lacking - V4e
Mastitibialae IIT present .,- ,~,-..--.--,----- il
Mastharsalae I]] lacking, About 42 dorsal idiosomal
seiae present, Cheliceral denticles 3-4d/3-4v. AM seta
stort and thickened, reaching about 2/3 across to the
AL seta base (Fig. 7A, B) -...-..-.---.---.2----
O. southcon’ (Womersley, 1944)
Mastitarsalae III present , 10
Cheliceral denticles 4-5d/ iv (Fig, 71C), DS to 22, yen fang
, . OY. langani (Southcott, 1957)
Cheliceral dentic les 4-4d/4-5d (Fig. 7D), DS 29-32 um
long OJ mathewi (Southeotl, 1957)
Tibia Uf with three mastitibialae. Palpal setal formula
B/B/BBB/So + 7B. Chehceral denticles 34/3v. One
mastitarsala [1] present . :
gt 2 GF miccutiacn! (Wamersley 1944)
Tibia Ww with two mastitibialae .-..--........, 2
About 30 opisthosomal ventral setae present . .
, O, adelaideae (Womersley, 1944)
More than 30 opisthosamal ventral setae present 13
PL/AL less than 1.20
tye ey ae, QO. HOve-guinea Womersley, 1944) and
O. lengipes LOR ET EY: 1945)
PL/AL greater than L.20 . ett erprec non dg
Over 75 dorsal idiosamal selae ‘present - fea
O. australiensis (Hirst, 1925)
and ©, hirsfi (Womersley, 1944)
Less than 70 dorsal idiosomal setae preseml .,.., 15
More than 58 ventral opisthosomal setae present,
Cheliceral denticles 3d/4v (Fig. 7B) ....-..-..--..
. O. cooki (Souchcou, 1957)
Less than 58 ventral opisthosomal setae present . bh
PSB/SB <1,00'. PW 87 pas] jam (Womersley, 1945)
Coe ee eee eee es OF, barrinensis (Womersley, 1945)
PSB/SB =1.00'. PW mostly less than 85 pm .... 17
PLYAL> 1.3, Ventral opisihosomal setae 48-54 in
number, Palpal setal formula B/B/BBB/So + 7B.
O. athertonensis (Womersley, 1945)
PL/AL<l, 3, Ventral Opisthosomal setae in range of
35-45 in number :
1981
piteaiag
bree bewte
srlisactpeetesger-
aS ims ti sli let Aire re wri ewer as
' For O. barrinensis and O. echidnus this stanstic is
based on Womersley's (1945) Figs. SF and 5D respec-
tively.
June, 1986
18. AW 34-61. PW/(A-P)<2,75-
O. echidnus (Womersley, 1945)
AW 65-73. PW/(A-P)=2.75. Palpal setal formula
B(b)/N{2b)/BNN/So » 7(B or Nj. Galeala nude —.,
dehe Meheafelerms Cee taieramr ccreitretues OF SWAN TR, Sp.
LLLE Lips im stew tmwenl
The following two species have been omitted from
the above key:
(1) Odontacarus gymnoduactyli (Ewing, 1925)
(1925b), described from India. This species was
doubtfully referred to 4comatacarus (Acomatacaris)
by Wharton and Fuller (1952, p. 98), who list the
previous synonymy. The descriptive details are
insufficient to place this species in the above key.
(2) Odontacarus lygosomae (Dumbleton, 1947),
from New Zealand, The descriptive details are
insufficient for the placing of this species in the above
key. Nevertheless it is quite a distinct species, with a
wide dorsal scutum, and flattened, broad, lanceolate
scutalae and dorsal idiosomal setae, the latrer
numbering about LOO,
Analysis af Marphometric Data of O. swani Larvae
In previous papers | have attempted to determine
correlations and other relationships between variates
that are used by taxonomists for classificatory
purposes, to see if they may be a useful basis for more
general studies such as whether the mechanisms
controlling size are interrelated in some way, eg.
genetic. Southeott (1966, pp. 736-738) reported on the
correlation of the standard data for the erythraeoid
larval mite, Charletonta swaztana (Lawrence, 1940), In
4 later paper (Sourheout, 1986a) a correlation study by
principal component analysis (Hotelling, 1933) was
presented for both seutal and leg variates for a
trombidioid larval mite, Trombella alpha Southcott. In
the present paper a Series of larval O, swani is similarly
analysed. Some of the same remarks made aboul
utilizing the data af Southcort (1986a) apply also to the
present paper. In attempting to elucidate relationships
we have here a group of larvae reared from conspecific
adults, collected over a few square metres of soil, near
the surface, on a single day. Additionally, the batches
of larvae were reared with damp soil in sealed tubes
in my baggaee, so that all the larvae of each batch may
be said to have experienced at least similar conditions
in their life-spans. Such conditions could therefore
possibly eliminate sources of variation from dissimilar
conditions of rearing.
In making statistical analyses, as was done with the
study of the larvae of Trombella alpha, we may restrict
the data for the dorsal setae to a single variate, in view
of the partial redundancy in using the variates DS, mid-
DS and PDS. We may also omit the data for Sens, since
these sensillary setae are slender, Mexible and difficult
to measure accurately; also these setae are commonly
absent In microscopi¢ mounts, whether from the
“Prom Wamerstey (L945),
181
ODONTACARUS SWANI N.SP.
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12 REC. S. AUST, MUS. 19(12): 169-200
capture, mounting or preservation procedures
themselves or possibly earlier traumas of life. Omitting
also PW/LB, which is a statistic derived from two of
the listed variates, we may therefore construct a & «23
table of variates from these data.
A correlation matrix of 23 = 23 values Gncluding L,
which is the sum of LA+LB, or ASB+ PSB) was then
calculated. There are only 2) independent measures in
the table of r, on 6 degrees of freedom (Table 4).
The table of the correlation matrix shows a total of
25 significant correlations (at P=.05 or at greater
significance),
From this we may calculate a principal component
analysis (Table 5), As there Were only eight specimens
in the series, it was considered the principal component
analysis should be restricted to only two (first and
second) components.
‘Table 5 for the principal components for the variates
selected for study in QO. swant shows that the first
principal components includes all 23 variates with the
exception of SB, PSB, LA, LB, A-P, AM, PDS, Til,
Gell and Till, indicatmg that this component may
represent a size variable affecting the majority of the
variates; and the second principal component includes
PSB, L, LB, and AM (also LA and Till, of opposite
sign of the direction cosines), thus indicating that these
measures are correlated, and that after allowing for size,
where one group is relatively large, the other is relatively
small.
Further Siudies on the Relationships of Variables
Used in Classification af Trambieulid Mite Larvae
Statistical analyses of chitinized parts of various
trombidioid, erythraeotd and other mites have been
increasingly used as an aid to classification, My use
of correlation analyses to determine the interrelation-
ships of these variates has been outlined above. In my
1986 (1986a) study | showed Lhat there is a correlation
belween a variate dependent upon a body character
(scutum) and of a leg character, in some cases, Above
it was found that in QO. swani larvae both principal
components (first and second) include both scutal and
leg dimension variates, these being positively correlated
in the case of the first, and both positively and
negatively (for the different variates) in the case of the
second.
In order to decide whether such interrelationships
are an expression Of some more general phenomenon
or phenomena, il was decided to do similar studies on
some other species of related trombiculid mites,
According to keys in existence before the present study
was commenced, it was thought that the larvae
described above as Odontacarus swani were classified
close to O. athertonensis and O. mecullaché, and
initially were classified as O. athertonensis, Both of
these species are included in ihe same couplet in the
key of Goff (1979a, p. 154). These two species of larvae
were therefore selected as being suitable for similar
dune, 1986
correlation studies, with the additional hope that if
difficulties in classification were to appear, the
correlation study might provide some guidance as to
the relationships between these specics. Accordingly
careful measurements were made on the available series
of these two species also. Ultimately, with further study,
it was found that the larvae described here as O, swuni
differed in several characters fram bath O.
athertonensis atid QO. mecullachi, soa that only @
athertonensis and O. swani appear close together in the
key above.
In the case of O. athertonensis a jong series of larvac
was available (28 specimens), and in the case of O
mecullochi, 10 specimens. These were measured for Lhe
same structures as in O swan. Nevertheless, it is clearly
as valuable to test for species widely separated from
each other as for closely related species, in a search for
such interrelationships.
Odontacsrus athertunensis (Womersley)
Synonyniy
Acomatacarus athertonensis Womersley, 1945, p. 104;
Taylor, 1946, p. 226; Fuller, 1952, p. 230; Gunther,
1952, p. 40; Wharton and Fuller, 1952, p. 97; Audy,
L954, p. 164,
Odontacarus aihertonensis Goff, 1979a, p. 144,
Source of the Larvae of Odontacarus athertonensis
Used in the Present Study
The larvae of O. atherfonensis studied for (he present
paper came from the following sources:
(1) The type series of 19 larvae, identification
numbers ACB732A-S, Wongabel, Q., October, 1944,
R. N. McC[ulloch], also two further larvae, idequ-
fication numbers ACB732T, U, with same locality, date
and collector information; South Australian Museum
collection, Specimen ACB732A slide is ooiated
“Syntype”; specimens ACB732B-S are notated as
TABLE 5. PRINCIPAL COMPONENT ANALYSIS OF THE
VARIATES OF ODONTACA RUS SWANS LARVAE
i 2
AW — 0.2846 0.1073
Pw ~ 0.2688 0.0357
SB —0,0810 01797
ASE ~O.3111 1144
PSE 0.0774 0.3769
1, ~ 0, 1860 0.2875
LA ~ 0.0884 — 0.4182
LG ~ 0.1108 0.3875
LN —0,3032 0, 1458
WwW ~ 01,2984 0.0569
A-P 0), 1496 ~ 0.1097
AM 0.1187 0.3843
Al, ~ 0, 2384 0.0396
PL ~ 0.2891 — 0.1461
AMB - 0.1852 0.0307
MDS —O.1783 - 0.0782
PDS 0.1625 0,0058
Gl — 0.2475 0.0756
Tl 0.0745 0.2512
G2 U.1476 0.0949
12 0.0850 — 0.2692
G3 — 0.2199 0, 1564
Tr —0,3011 0.548
ODONTACARUS SWANTE NASP. 183
Paratypes; specimens ACB732T, U are without type
notation.
Womiersley stated (1944, p. 105) that the species was
“described from 16 syntypes”. Specimen ACB732A,
which bears also a number N1981384, and was
presumably the one exanmined by Goff (1979a), is
hereby formally designated the lectatype,
(2) Six larvae, mounted individually on slides,
identification numbers ACB733A-F, Mt. Jukes,
Q[ucensland] [21°00'S, 148°57'E), 6.ix.1951, E. A.
Derrick (South Australian Museum collection), These
six slides were also examined by Goff (1979a), and bear
the notations G192, GI93, G197, G196, G206, G197
respectively, also numbers G1981385-G1981390,
respectively, Each slide also bears the comment “Was
marked paratype” (this marking has been removed).
These slides were clearly never pact of the type series.
Specimen ACB733C is a poor mount, unsuitable for
detailed measurement.
Omitting specimen ACB733C, three of the five
larvae kev directly to O. athertonensis by the key given
above, while for two others (ACB733 E & F) 0.
athertonensis is the nearest fit, there being a mismatch
iv each of PL/AL=1.225, Further studies of these
larvae have been presented elsewhere (Veitch and
Sourheott, L984; Sautheort, |Y8ob).
(3) ACB734, one larva, Lae Test Area, Papua New
Guinea, 24.iv.1944, R. N. McCulloch (S.A, Mus.). The
slide has the notation; Leenwenhoekia (Acomatacarus)
nr. athertanensis Wom. Lae Test Area. 24.4.44. R. N.
Mc, Det, H, Womersley. AW and PW Standard Data
longer. AW=75.6, PW=84.0, (This is a poor mount,
the dorsal scutum being damaged.)
By the key given above this specimen comes Lo
caption 18, but then does not fit the criteria of either
O, echidnus or O. swani. A further analysis of the
statisuical data of this specimen has been made
elsewhere (Veitch and Southcotl, 1984; Southcertt,
1986b),
I response to my request for locality data, Dr. R. N,
McCulloch has replied (pers. comm., 1983): “Lae test
arca was jungle just off the main road and where | did
stopping-time tests [see McCulloch 1944, 1946, 1947]
probably a mile towards the big air strip from 2/7
A, G, H,” (2nd/7th Australian General Hospital).
(4) ACB735, one larva, Lae Scrub, 24,ii1,.44, A, S. DL
(South Aust, Mus.), The slide is additionally notated:
“Teeuwenhoekia (Acomatacarus) nr. athertonensis
Wom. AW and PW Standard Data longer. AW=70.0,
PW=84,0, Det. H. Womersley.” This is another poor
mount.
By the key above this specimen is not identifiable,
owing to both legs III being damaged,
As no mite student or collector with the initials
A. 8. D. was known to me, | have written to Dr, R. N-.
McCulloch asking for further locality and other
information, He has replied (pers. comm., 1983) “ASD
means, I guess, Advanced Stores Depot and was on the
old disused civilian air strip which was between the new
[i.e. 1944] active air strip and 2/7 A. G. H.”.
TABLE 6. MORPHOMETRIC DATA (um) OF ODONTACARUS ATHERTONENSIS (WOMERSLEY) LARVAE
Character Lectotype Canspecificity
ACBI32A Queried
ACB732 ACB735
AW va 19 a5
PW aU 91 90
SB 25 uM 27
ASE 47 55 54152)
PSB MW 34(31)" 28(36)!
L 82 R9(B6)* §2(88)°
LA 16 20(25)' 18
La 46 69(61)" 6a(70)"
\ 25 30 29
Ww o4 96 97
A-P 25 32 27
AM 37 39 v
AL 41 46 46
PL 55 58 66
AMI 13 15 I
Sens 50 ‘= 4§
PW/LB {21 1,32 t.41
ps 29.42 33-64 36-67
mid-DS 29-40 33-38 34-44
PDS 36-40 42-46 33-42(-51)"
Gel 45 45 56
Til 60 55 67
Gell 40 37 il
TH 47 42 56
Gell a9 42(44)" 56(58)"
Tint 65 57 69(75)
Range for n Mean! §D.'
Type series
61-7 21 65.90 2.700
75-87 21 T7782 2.655
22-30 21 25.10 1413
47-56 21 53,00 2.490
25-35 21 28.90 2.719
76-87 21 81.90 3,113
16-23 7 19.81 2.040
§5-67 2) 62,10 2,897
25-32 21 29.05 2.202
81-96 21 87.14 4,246
25-32 2! 28.24 1.814
35-42 21 38.38 2.459
37-46 21 40.71 2.101
50-58 ra | 55.24 1.700
10-13 21 11.43 1,028
45-55 ? = -
1,17-1,45 al _ —
(29-35)442-56) 2 — —
129-35)-(35-42) 2 37.74" 2.385!
(35-38)-(38 46) 2 41.57 | .964'
42-49 Zt 45.76 1.670
53-60 20 §5.19 L662
36-42 at 33,19 1.662
41-49 2) 45,43 \ 886
44-50 21 4571 L.875
57-65 21 59.86 2.197
' The mean and the s.d. are calculated for the type series Only With nm» 21 (see text).
> In subsequent calculations of correlation the three missing values (dicated by (a), (b) and (c) were replaced «as follows: (a), AL for
ACBT732B: 4], (b) AM for ACB7325: 35, (c) Til for ACB732B; 53, after a study of the data. This refers ta the table of variates for
the type series held on file in (he South Australian Museum,
! Figures calculated from the maximal yalues only of thal variate.
+ Figures in brackets are a set Of Variant estimates made subsequent
ly and which differed by 2 wm or more from the earlier ones, and
used in calculations presented in Veitch and Southcott (1984) as a check on the reliability of analytical procedures. The specimens were the
most separated from the others using the variables selected. Specimens ACB734 and ACB735 were poor mounts, presenting difficulties in
Microscopic measurement,
REC. S. AUST. MUS. 19(12): 169-200 June, 1986
184
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OPONTACARUS SWAN] NSP, 185
A further statistical study of the data of this
specimen has been made elsewhere (Veitch and
Southcortt, 1984; Southeett, 1986b),
(5) ACB736, one larva, collection data as for
ACB734, The slide bears the notation; “Leewwenhoekia
(Acematacarus) nr, athertonensis Wom, Lae Test Area.
24.4.44. R. N, McC. AW and PW Standard Data
longer. AW=68.6, PW=84.0. Det. H. Womersley,”
Remeasuring gives AW about 65, PW about 75.
From obscurity in the mount the specimen is not
identifiable by the foregoing key, various key characters
not being visible, (The data of this specimen have been
omitted from the table given below (Table 4), as too
many values are unavailable for it to be capable of
being included in a correlation analysis.)
These three are the only specimens in the South
Australian Museum collection placed as or near @
athertonensis, trom Papua New Guinea. Although
Goff (1979a, p, 146) states that the three specimens
examined were from “Morobe Distr[.], Lae, 24.iv.1944,
collected in scrub”, only two slides bear the date
24.iv.1944, and the term “scrub” occurs on only one
slide; none of these slides carry any loan nuniber
designations, apart from that implied by my
identification oumbers.
Data
The measurements of the standard data and other
variates selected for study are shown in Table 6,
Results of the Correlation Analysis of the Variates
of Odontacarus athertonensis Larvae
in Table 6 the means and standard deviations of the
variates have been calculated from the type series only
(ACB732A4-U), The reason for this is that.a preliminary
discriminant analysis of the data given indicate that
the specimens previously considered as conspecific, as
well as those under the heading “Conspecificity
queried” differ significantly in various statistical
measures from the type series. This subject will be
analysed further in another paper* Accordingly,
correlation analyses here will be confined to the data
of the type series.
Proceeding as before, a correlation matrix of 23 x23
values is calculated; this is shown in Table 7,
There are 32 correlations between variates showfi in
Table 7 ai ithe 5% (or more significant) levels of
probability, These are marked in the table by asterisks,
as shown,
Examining the principal components as before, we
may select the first and second principal components
(in view of the sample size of 21 specimens). These are
shown In Table 8.
From Table 8 we see that there are two significant
groups of correlations. Component !, of AW, PSB, LB,
Til, Gell, Till, GellI, Till], together with (of oppasite
* Note added in proof: This has now been done; see Veitch and
Southcotr (1984).
TABLE 8, PRINCIPAL COMPONENT ANALYSIS OF THE
VARIATES OF ODONTACARUS ATHERTONENSIS LARVAE
/ 2
AW ~ 0.2232 0.1410
PW —0,0750 — 0,0299
SB — 0.0399 0.0587
ASB 0.2704 ~ 0.9856
PSB ~ 0.3569 ~ 0.2042
L 0.0954 0.4868
LA 0.2266 0.3494
LB 0.2621 0.2770
LN 0.2258 0.3471
WwW — 0.1864 0.1381
AP 0,2743 0.2177
AM 0.1562 0.0371
AL 0.0486 00,1865
PI 0.0007 0.1760
AMB ~ 0.1648 0.0630
MDS — 0.0869 0, L064
POS —(.1153 —0.2161
Gl 0.0827 0.0341
TI 0.2558 0.0222
G2 — 0.2074 0.0255
T2 ~O0.2151 ~ 0.1362
63 — 0.3325 0,0961
73 — 0.3248 D.0585
sign) ASB, LA, LN and A-P. This component thus
represents a mixture of size and shape variables,
Component 2, of ASB, PSB, L, LA, LB, LN and PDS,
and (of opposite sign) A-P, which is thus seen to include
mainly variables based upon the dorsal scutum. This
component again is a mixture of size and shape
variables,
Odontacarus mecullochi (Womersley)
Synonymy
Leeuwenhoekia mccullochi Womersley, 1944, p. 108.
Acomatacarus mecullochi Womersiley, 1945, p. 111.
Taylor, 1946, p. 226. Gunther, 1952, p. 40.
Acomatacarus (Acomatacarus) mecullocki Wharton
and Fuller, 1952, p. 99. Audy, 1954, p. 164
Odontacarus (Leogonius) mecullocht Goff, 1979a,
pp. 147, 154,
Material Examined
(All specimens in South Australian Museum
collection)
QUEENSLAND
Trinity Beach Area [Trinity Bay] 10.vii.1943, R. N-
McC[ulloch], on boots, edge of scrub, four specimens,
identification numbers ACB752A-D, A labelled Type,
B-D Paratypes,
Trinity Beach, Cairns, Q[ld.J, edge of scrub,
8.vi1.1943, name of collector not stated, identification
numbers ACH7S3A-D. (These slides are labelled
Paratypes, although according to Womersley's account
(1944, p. 109) they were not mentioned in the original
description.)
Emerald, Cfentral] Queensland], Nov. 1948, ex
Pomatostomus temporalis, H, Wlomersley|, two
specimens, identification numbers ACB754A, B,
Data
The measurements of the standard data and other
variates selected for study are shown in Table 9,
186
TABLE 9, MORPHOMETRIC DATE (um) FOR QDONTACARUS
MCCULLOCH] LARVAL
Type
Characier (A CB752A )
Runge n Mean &0D,
AW 62 56-féy 10 «$662.50 3.206
PW 77 74-83 1) 78,20 3,084
SB 24 23-27 10 24.20 1,229
ASB 50 46-59 ip 51.00 3.916
PSB 24 24-28 1D WA 1,270
iL. 74 73-87 10 FLL 4,358
LA 18 15-21 1 17.700 2.183
LB 56 56-66 1 59.20 3,360
LN 27 22-1 10 26.70 «4.098
Ww 86 78-93 tO R680) 4, 4902
A-P 32 24-35 1 =©30,90 3,315
AM 33 33-40 9 35.50 2,369
AL al 34-4) 10 37.80 2.394
PL 53 47-58 10 S270 3.621
AMB it O14 10 10.90 1 287
‘Sens 56 46-56 = ~ =
PW/LB 1.38 1.20-1,38 — = —
bs 41-56 (25-41}-(47-62) — — —
mid-DS 31-36 (25-41)-(31-36) 10 34.70! 1567!
PDS 36-40) (31-36)-(33-40) 10 «38.70! 2,406!
Gel 34 46-54 Y 49.70 2,584
Til 56 §5-62 9 57,70 2751
Gell 3% 38-46 10 42.20 7.741
Till 46 A3-§2 ww 47.20 D774
Gelll 49 45.54 1 48,00 2.467
Ti 2 57-63 10 =6b00 221)
1. Figures for the maximum values of rhe variate,
Analysis of Morphometric Data for Odontacar\s
mecullochi Larvae
The data in Table 9 were submitted fo the previously
described analysis, the variate Sens being omitted for
the reasons as before. Likewise the variates involving
the dorsal idiosomal setae were restricted to the
maximum values of the mid-DS and PDS variates,
Proceeding as before we thus have a 23 x 23 table on
8 degrees of freedom (Table 10).
Table 10 shows 28 significant correlations between
the variables, at the 5% or greater level of significance,
From these data we may perform a prineipal compo-
nent analysis, as before, This is shown in Table II.
From Table 1] i! can be seen that there are two
significant groups of correlations. Component 1, of
AW, PW, SB, ASB, L, LA, BL, LN, W, A-P, AMB,
Gel, Til and Till, ic. indicating largely that this is a
correlation induced by size factors, Cofmponent 2, of
AW, Til, Till, Gelll and Tilll, and (of oppasite sign}
LA, LN, A-P, AL, AMB, MDS and PDS; it is thus a
mixture of correlations of opposite signs, involving
variables derived from scucum, body setae and leg
dimensions.
Conclusions on Correlations Beiween Measurements
of Chitinized Idiosomal and Leg Parts in Three Species
of Larval Trombidiotd Mites
My earlier study on Trombella alpha (Southcott,
1986a) and the present study of three species of
trombiculids have defined in each species the lirst twa
(or three) principal components in a comparable series
of variates from specified chitinized parts of (he bady
and limbs of these mites. The findings are set out in
Table 12.
REC. S. AUST. MUS. 19(12): 169-200
June, 1986
It is apparent from the preceding analysis and Table
12 that the correlation patterns are different in each
species of mile stiidied, for the variates that have been
selected for study. For the first principal component,
there appears to be a greater similarity between the
patterns between O. mecullochi and O, swani, than
between either of these species with O, atherronensis.
The patterns of the first principal component in these
three species of Odentacarus resemble cach other more
than they do that of Trembella alpha,
This result suggests that all of these measurements
may be necessary to separate species of Odonfdacarus
and, by an extensian of this concept, species in other
genera of mites.
There is thus nol a compact set of correlations mani-
fested in these data. This is due in part to the small
sizes of the samples studied, although other causes of
these variations and comparative paucity of correla-
tions could be suggested, particularly nutritional or
other environmental factors,
Odontacarus southcotti (Womersley}
(Fig. 7A, B)
Synonymy
Leeuwenhoekia southcotti Womersley, 1944, p. 109,
Taylor, 1946, p. 232.
Acomatacarus sourhcont Womersley, 1945, p. LL.
Taylor, 1946, p. 226, Gunther, 1952, p. 40,
Acomatacarus (Acamatacarus) southecotti Wharton
and Fuller, 1982, p. 100; Audy, 1954, p. 164.
Material Examined
NORTHERN TERRITORY: Type series
ACBI69A1-7, South Australian Museum collection,
Adelaide River, 15,vi,1943, on skink. (These are the only
specimens if the S,A.M. collection; originally 1 had
submitted 10 specimens, on two slides, labelled
ACBI69A (8 specimens) and ACBI169B (2 spetimelis).
Remarks oan Collection Data
On 15.vi.1943 | collected 10 or more red larval mites
from the external auditory mieati of a small skink,
specimen R44 (my number), in the Adelaide River area,
about 12 km (8 miles) east of the 57 mile point (ie.
57 miles (91 km) souchwards along the main road from
Darwin; (his being the current military nomenclature),
This is about lat, 13°06'S, long. 131°14’E, There were
5 or more larval mites in each ear of the skink; but
none in the axillae or elsewhere on the external surface
of the skink, The skink was later identified as
Leiolepisma ?pectoralis by Mr F, J, Mitchell, South
Australlan Museum, but not kept, Another specimen
caught at the same lime and place (my R46) was
identified by Mr Mitchell as L. pectoralis, and is
preserved in the South Australian Museum collection
as R2703.
These notes supersede the collection data given by
Womersley (1944, p, 110), whith are partly wrong,
187
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188 REC. 8. AUST, MUS. 19(12): 169-200
TABLE il. PRINCIPAL COMPONENT ANALYSIS OF THE
VARIATES OF OQDONTACARUS MCCULLOCH! LARVAE
I 2
AW — 0.2326 0.1860
Pw ~ 0.2784 0,0693
SB 0.2094 0.1310
ASB ~ 0.3164 ~ 0.1233
PSB 0,0359 0.0428
L ~(),2730 ~ 0.0974
LA — 0.2598 — 0.2159
LB — 0.2155 ~ 0.0462
LN ~ 0.2920 0.1979
wW ~ 0.2796 — 0.0299
AP ~ 0.2459 0.2958
AM O.1781 0.0525
AL 0.0158 — U,3045
PL 0.0307 0.0012
AMB 0.1930 0.2467
MDS ~ (0.1259 - 0.2859
PDS 0.0017 0.4122
Gl ~ 1), 1934 0.1564
Ti ~ 0.2514 0,2677
G2 — 0). L009 0.1977
T2 0.2511 0,2536
i3 ~ 0.1443 0.2099
T3 0.1735 0.2212
Additional Notes on Morphology
This species has a broad, shallow dorsal scutum, with
short, rather thickened, AM setae, which reach about
two-thirds of the way across to the AL seta bases. The
cheliceral fang has 3-4 dorsal, and 3-4 ventral, denticles,
These features are illustrated in Figs. 7A, B.
Odontacarus barrinensis (Womersley)
Synonymy
Acomatacarus barrinensis Womersley, 1945, p. 106.
Taylor, 1946, p, 226. Gunther, 1952, pp. 41, 48.
Acomatacarus (Acomatacarus) barrinensis Wharton
and Fuller, 1952, p. 98. Audy, 1954, p. 164 (with
query).
Remarks on Collection Data
Womersley recorded (1945, p. 106) “Described from
five syn-types collected free, from Lake Barrine,
Queensland, 16 Nov. 1943 (RV.S.), and a single
specimen from man, Atherton Tableland, Queensland,
8 March 1944 (RV.S.)".
Reference to my field and other notes shows that the
first group (“five syn-types”) were my serial ACB188,
recorded originally by me as “ACBI88,..5
larval,,.Running over a log, Lake Barrine shores. Qld.
16.11.43.”
dune, 1986
With respect to the sixth specimen, stated to have
been collected “from man, Atherton Tableland,
8 March 1944”, this is my specimen ACB210A, This
was the sole specimen of Odontacarus among a batch
of mites which included many larval trombiculids
(ACB210B, C, D, ete.) and two small (larval) ticks
(ACC358, ACC359), collected parasitic on three
soldiers at Gilbey Creek, near Wondecla, Atherton
Tableland, Queensland (Map Ref. Herberton (1:63360)
392023), on 8 March 1944. The actual collector of the
specimens was Cpl. E. Grinham, A Company, 2nd/7th
Australian Field Ambulance, as | was away in another
area at the time. The other larval trombiculid mites
collected were later identified as Eutrombicula hirsti
(Sambon, 1927).
Remarks on the Subgeneric Classification of the
Larvae of Odontacarus Ewing
The genus Odontacarus (as well as a number of
others of the family Trombiculidae) was revised and
divided by Vercammen-Grandjean (1968), Definitions
were submitted not in the usual terminology of
acarological or even trombiculid description, but in
Vercammen-Grandjean’s own code, Such extensive use
of codes in the descriptions and definitions of
trombiculid mites has in fact been subject to crilicism
(see Brennan and Gol’, 1977).
The following is an account of the revision of Odon-
facarus made by Vercammen-Grandjean (1968, pp.
(20-121).
Definitions
Odontacarus Ewing, 1929
GT/ Trombicula dentata
Ewing, 1925
Leeuwenhoekiini of
medium to large size-
Remarks and interpretation
‘Type species, “genotype™
A new tribe introduced by
Vercammen-Grandjean on the
same page (p. 120); on p. L1Y
he retained the family Leeu-
wenhoekiidae Womersley,
1945, and subfamily Leeuwen-
hoekiinae Womersley, 1944,
Ip. Index pedibus, the sum of
the lengths of legs 1, [Land IH.
Despite its use by Vercammen-
Grandjean, it must be con-
sidered of little diagnostic
Value; @.u. Acomatacarus
(p. 122) is stated (o have an Ip
of 800-1020 (yum).
Ip = 700-1400
TABLE 12, ELEMENTS OF THE FIRST TWO (OR THREE) PRINCIPAL, COMPONENTS OF A PRINCIPAL COMPONENT
ANALYSIS OF COMPARABLE SERIES OF VARIABLES OF CHITINIZED IDIOSOMAL AND LEG STRUCTURES FOR THREE
SPECIES OF LARVAL TROMBIDIOILD MITES
Species Camponeni |
ASB, L., AeP, TUL
AW, PSB, LB, Til,
Gell, Till, Gelll, Ti,
-ASB, -LA, -LN, -+A-P)
AW, PW, ASB, L, LA,
Trombeltla alpha
Odontacarus
athertanensis
Odontacarus
meceullochi LB, LN, W, AP,
AMAR, Ciel, Til, Till
Odontacarus AW, PW, ASB, L, LN,
swani WwW, AL, PL, AMB
MDS, Gel, Gelll, TIL
Component 3
DS, lil
Component 2
AW, PW, PSB, AM, Gel
ASB, PSB, L, LA, LB,
PDS, -(A-P)
AW, Til, Till, Gell, —
Tilll, -LA, -LN, 4A-P),
-AL, -AMB, -MBS, -PDS
PSB, L; LH. “AM, -LA, =
Tin
Note: ASB=ASBa in Southeatt (9kbu); PSB ASB likewise,
ODONTACARUS SWANI N.SP. 189
FIG, 7. A, B Odontacarus southcotti (Womersley), holotype larva; A Dorsal scutum, B Tip of mouthparts, dorsal view. C O. langani
(Southcott), holotype larva, chelicerae, dislocated as on slide. D O. mathewi (Southcott), holotype larva, tip of mouthparts, dorsal
view. E O. cooki (Southcott), holotype larva, tip of mouthparts, dorsal view. All to scale shown. (Note: in each case the distal half
of the cheliceral fang is shown in transparency, so that the denticles on both the lateral and medial aspects are shown.)
190 REC, 5. AUST. MUS. 19(12): 169.200
This means there are two
anteromedian setae, two antero-
laterals, and (wo posterolaterals
to the scutum,
The term “2p” is not defined,
but presumably means that
two eyes are présent On each
side, each with a lens.
This is the code for the “palpo-
tarsal pilous formula”,
meaning here that there are
seven branched setae (plus a
sdlenoidala, stated to be in-
variably present) on the palpal
tarsus,
This means there are 2-4
prongs to the palpal tiblal claw
These are specialized setae
of the pedal tarsi. As accord-
ing to Vercammen-Grandjean's
slatement (hey may be absent
or present, and if present, then
branched or nude, they can
clearly be allotted no diag-
nostic significance.
Vercammen-Grandjean, in the
same Work, also uses § and T
With other meanings,
Scutum roughly penta-
gonal with a nasus, 2 AM,
2 AL and 2 PL.
fiye lenses, 2p.
fT=7B
and palpotibial claw,
Gr 2-4
Subterminala or parasub-
terminala more often
branched or absent than
nude,
Always al. least one mast-
tarsala or a solenidion (s)
on leg 3, Stigmatae (sic)
and tracheae present (5+T).
Ch - ¢,° or 2, mever 2, The code Ch refers to the cheli-
cerae, The notation ete., is
not explained, but presumably
refers to the presence or ab-
sence of dorsal (d) or ventral
(vy) teeth on the cheliceral fang,
The second is presumably
an error for >, as both dorsal
and ventral denticles are often
present to the lang in Qdanta-
cerus.
Vercammen-Grandjean (1968, p, 120) then proceeded
to divide Odontacarus into Odontacurus s. str., and two
new subgenera, Jaralacarus and Leogonius. He offered
no definition for Odontacerus §, str., commenting “The
subgenus Odontacarys s. str. is of no concern in this
study”, doubtless on the (implied) grounds that it was
not represented in the “Far East” (a term which was
used to include Australia, Malaysia, and eastern Asia
more generally), Odonjtacarus s, str. was also omitted
from the diagnostic key given on p. 53 of the same
work, Reference to that key shows that 7Jarsalacarus
is separated by being a section of Odontacarus sens.
lat. in which leg tarsus [Il has a dorsal solenidion
(solenoidala), while in Leogonius tarsus [II lacks such
a seta “but there is 1 maatitarsala and 2 mastitibialae”
to leg 111 of the larva. Additional characters are listed
June, 1986
for Tirsalacarus and Leogonius by Vercammen-Grand-
jean in that work (pp. 53, 121). Perusal of the characters
listed shows that many of (hese are shared in common,
or are Overlapping, ég. “Odontacurus of median (sic)
size” as against “Odontacarus of medium to large size",
or “Gr 2-4” (i.e, 2-4 claws to palpal tibia). The only
characters listed by Vercammen-CGrandjean as being
distinct between the two subgenera, apart from that
listed above, is Ch=O/v for Tursalacarus, and Ch=d/O0
for Leogonius, i. Tarsalacarus has ventral cheliceral
denticles but lacks dorsal ones, while the reverse
situiation is stated to apply in Leogonius, Onychotrichs
(cilia lo the pedotarsal claws) are not mentioned.
The subgenerie classification has been clarified and
corrected by Goff and Loomis (1977), who defined
Odontacarus s, str, as “Odontacarus laryae lacking
tarsala 111 fi.e. a solenoidalaj; mastitarsala [11 present
or absent”, and Jursalacarus as “Odontacarus larvae
with tarsala 111; mastitarsala I] absent; onychotriches
absent’, Leogonius was redefined by Goff (1979a), the
diagnosis including: onychotriches on both claws and
empodia; cheliceral blade with dorsal and ventral teeth;
2-3 mastitibialae II] present, with basal barbs; masti-
tarsala 11] present, with basal barbs.
By the above definitions therefore, the species (as
larvae) considered in the present paper come within
feogonius, as in fact all Australian, New Guinea and
South-East Asian species have been placed by previous
authors, since the subgeneric division of Vercammen-
Grandjean.
‘The characters of the subgenera af Odon/acarus are
shown in the following Table 13,
Classification of Adults and Nymphs of Odontacarus
As this is the first description of an adult
Odontacarus, confirmed by experimental rearing, it is
appropriate here to examine the status of other adults
that have been placed in this genus on the basis of
morphological similarity to nymphs reared from larvae,
So far seven nymphs have been allotted to Odontacarus
by such experimental rearing, these being:
(1) O a@rizonensis (Ewing), from North America,
reared by Crossley (1960, p. 196). This species has since
been replaced in Acomatacarus s. str. by Vercammen-
Grandjean (1968).
(2) O. plumosus (Greenberg, 1951). This binomen
was used by Brennan and Jones (1959), Loomis and
Crossley (1963) and Nadchatram (1963).
A separate genus name, Nenacarus Greenberg, 1951
(since replaced by Nenodontacarus by Loomis and
TABLE 13. DIFFERENTIATING CHARACTERS GF THE SUBGENERA OF GDONTACARUS (+PRESENT, -ABSENT)
(SEE TEXT FOR FURTHER EXPLANATION)
Maviitarsalae fit
(MaQTalti)
Mastitibialae if
Cade (MaTifit)
Subgenus
Odontacarus
Tarsalacarus
Leogonius
‘
‘
| |
Salenararsala ttt Onychatrichs Cheliceral denticles
fSoTalltl)
a 2 a/v
_— D/y
— ' ib’y
ODONTACARUS SHANI NS SP, EL
Golt, 1973, on grounds of pre-occupation) has been
allotted to this species, which was reared to a nymph
by Crossley (196t), p. 197). Since Loomis and Goff
(1973), as well as Vercammetn-Grandjean (1973a) confer
full generic status on this and related species, we may
exclude il from immediate consideration.
There are thus five species of trombiculid mites
whuvh have been reared experimentally from the larval
to the nymphal stage, which may be placed in the gequs
Oden/acarus in the sense of Goff (1979a, b). These are:
(3) O, (Leagonius) australiensis (Hirst, 1925) from
Australia and New Guinea, reared by C. B. Philip (in
Womersley, 1945) from New Guinea material, and by
Domrow (1956) from Australian material.
(4) O. (L.) adelaideae (Womersley, 1944), from
southeastern Australia, reared by the present author
(unpublished),
(5) O (L.) audyi (Radford, 1946), from India,
Thailand and Malaya, reared from Malayan material
by Nadchatram (1963),
(6) O. (L.) novaguinea (Womersley, 1944) which was
reared from larvae by G, M, Kohls in New Guinea in
April, 1944 Gin Womersley, 1945).
(7) © (LY) longipes (Womersley, 1945), which was
reared from larvae taken from Podargus sp, by G. M-
Kohls, Daobadura area of Papua New Guihea, 25 July
“1940” (probably 1944); recorded in Womersley (1945),
(This species was synonymized with ©. (L.) novaguinea
by Goff (1979a, p. 149). Nevertheless the nymphs were
stated by Womersley Lo differ significantly from those
of the latler species, aad this was not resolved by Golf
(low. cif), 1148 therefore praposed to retain these two
species 46 separate at present.)
Womersiey thus in 1945 had sound grounds to define
the post-larval characters of Odontacerus (as
Acumatacarus), and to allot to it several species of
adult mites whose previous taxonomic placings had
been largely conjectural, these being, as Womersley
stated (1945, p. 98) “RAyvrecholophus attolus Banks
1916, R. retentus Banks 1916, both considered
[Womersley] (1934) as Microtrombidium, and
Dromeathrambiurnt drorius Wor, 1939, the last now
being shown to be really two species” (Womersley, L945,
p. 109, erected Acomatacarus palrius to accommodate
this second species).
It is very curious that, despite the widespread
distribution of Odonracarus (s/., to include also
Acomatacarus) in North America, Asia and Australia,
adults of this genus have been recorded only from the
Australian continent. This would suggest, vifer alia
(when combined with the small amount of information
on their ecology), that the adults live obscure and
presiimably subterranean lives, Thus the adults of 0,
swan recorded here were found in sail in rain-forest,
and the original specimen of QO patrins (Wom., 1945)
was collected under a stong, in mallee country, about
7 wiles West of Murray Bridge, South Australia, on
254.1938 by the present author, and was. recorded as
“white in life” which would suggest that that imsrar al
least is spent in darkness, Four species, O. attolus
(Banks, 1916), O, dromus (Womersley, 1939), O
retentus (Banks, 1916) and QO. partrias have been
recorded as being associated with ants, and in fact these
are the only records of capture of these species apart
from the above-mentioned record for O. putrits.
All known species of larvae in the genus have 2+2
eyes, which is consistent with their mode of life as
ectoparasites upon terrestrial vertebrates. Goff (1979a)
considers that the New Guinea larvae are principally
ectoparasites of birds, although elsewhere there ate
records of these larval mites from mammals and
lizards. In the case of the adults, eyes are stated to be
absent in ©, swant {see earlier), QQ dromus (sec
Womersiey, 1945, p. 110), and O patrius (see
Womersley, fac, cit),
In the case of © altalus, Banks (1916, p. 225) staled
That this species had 14+1 eyes, hut Womerstey (L945,
p. IO8) stated that they “are not now visible"; he nade
a similar statement in 1934 after examining freshly
mounted specimens. Careful reexamination of the type
material using phase microscopy techniques (not
available to Womersley) also fails to reveal evidence of
eyes, nor was the writer able to see them in October,
1946, when the mite bad been remounted by Womersley
al same indeterminate time beforehand, Nt may there-
fore be concluded that this species lacks eyes.
The only adult species for which previous authors
have agreed on the presence of eyes is OL retentys
(Banks, 1916), for which Banks (1916, p. 116) recorded
“one eye each side”, whereas Womtersley (1945, p. 106)
recorded “Eyes present, 2+ 2, small, on distinct ocular
shields...”. Re-examination of the type material
confirms Womersley’s description,
This species differs also in having idiosomal setae
which are terminally forked or branched. Its taxonamic
placing has been even more uncertain than that of the
ather species menUioned by Womersley (see above). In
fact Womerstey placed it in Microtrombidium
{subgenus E&nemothrombium) in 1934, and in
Calothrombium in 1937,
For this species, | therefore erect a genus
Scopitrombium nov, (see further below) within the
Trombiculidac, subfamily Leeuwenhoekitnae,
Genus Scopitrombiom nov.
‘Type species (original designation) Rhyncholaphus
relenius Banks, 1916.
Definition: Adult of normal trambidioid shape, the
idjiosoma not constricted, Eyes 2+2, placed behind level
of middle of crista. Crista broad posteriorly, narrowing
anteriorly to end in a broad arrow-shaped expansion,
the sagiita, provided with (wo normal setae. Crista with
a pair of sensillary setae, filiform, at its posterior end,
Palp slender. Palpal tibia with strong spine with several
adjacent thickened setae, Dorsal body setac ciliate,
192 REC. S. AUST. MUS. 19(12): 169-200 June, 1986
terminally forked or branched. Legs rather long, longer an allusion to the branched and ciliate ends of the
than idiosoma. dorsal idiosomal setae, and trombi-, the root name of
Remarks: This species is known only from the type the family Trombidiidae.
species. It is a typical leeuwenhoekiine post-larval mite,
with a well-defined sagitta to the crista. As indicated
in the key below, it comes nearest to Hannemania, but Scopitr ombium retentum (Banks, 1916)
is easily distinguished by the character of the dorsal (Figs BA-C, 9A-F, 10)
setae. Its larval stage is unknown. Synonymy
Derivation of name: The name is derived from Rhyncholophus retentus Banks, 1916, p. 225.
scopi-, the root of scopae (f. pl.), a broom or besom, Microtrombidium (Enemthrombium—lapsus for
pet a St A
Imm
o-
H
re
‘
¥
2
ad .
‘ f
? ‘
)
_s,
‘a a
'
‘
,
external
! -} genitalia.
n
ney
'
} .
’ mn
‘ ete Oe
\ .
ia > 1 .
‘ "\e be- Toe --
Vp. afta of Seboe :
| f@ SN Grown f
FIG. &. Scopitrombium retentum (Banks), adult, lectotype. A Slide-mounted specimen seen in transparency. B Batch of posterior dorsal
idiosomal setae, C Anus. (All to nearest scale.)
ODONTACA RUS SWANI N.SP. 193
FIG. 9, Scopitrombium retentum (Banks). Adult. A Crista, eyes and adjacent part of dorsiim. B Crista and sagitta in lateral view, C
External genitalia, D Left palp, medial aspect. E Same, lateral aspect. F Chelicerae. (A, ‘C-F from lectotype; B from ACB748), (All
to scale shown.)
Enemothrombium) retentus Womersley, 1934, p. 193. | Rernarks: This species was described by Banks (1916)
Calothrombium retentus Womersley, 1937, p. 85. | and Womersley (1934, 1945). It was recorded originally
Acomatacarus retentus Womersley, 1945, p. 106. from three localities in Victoria, in each case with a
Gunther, 1952, p. 41. species of ant.
Microtrombidium retentum Thor and Willmann, Hirst (1928b) was the first to point out that this was
1947, p. 370. a trombidiid (sens. Jat.) mite, not an erythraaeid one
194
as originally described by Banks, and he assigned it
to Microtrombidium without formally giving if a
binomen. He commented (p, 1027) “This apecies is
closely related to M.. barringunense [Hirsl, 1928a], bul
the tarsus of the first leg is considerably shorier . ... It
is closely allied to M. affine [Hirst, 1928b] bul has a
short bul distinct nasal process",
Of the three specimens, each labelled “Cotype",
specimen ACB750 is the most suitable for further
description, and is therefore designated the lectotype
(see further below),
Redescriplion of Lectotype, Specimen ACB7SO
(Supplemented by the Two Paratypes)
The specimen in its present state is a squashed mass
mounted on a 75%25 mm slide in a transparent,
yellowish water-miscible medium, based an either gum
chloral or polyviny! alcohol, with picric acid used as
a stain (Womersley fayoured the use of picric-acid-
containing media in the later 1940-19505 and possibly
later). The squashed mass is the remains of the
idiosoma, and is about 2.7 mm long by about 2.25 mm
wide, indicating a considerably smaller original
specimen (Barks recorded the specimen(s) he described
as 1.3-1.5 mm long and “about one and a half times
as long as broad", i.e, ca 0.9-1.0 mm broad.
Crista is a rod set in a narrow tapering scutum,
widest posteriorly, and ending anteriorly in a broad
sapittas crista and sagitta with combined length of 306,
Sagitta bell- or probability-curve-shaped anteriorly,
borders smooth, posterior margin reflex-angled; sagitta
82 wm across by 52 deep, The corners of the sagitta
are sharp (i.e. Isteral angles). Sagitta bears two scabalae
which are both broken off, at 21, 25 length; they are
not broadened, but appear normal and almost smooth,
From specimen ACB748 these can be seen to be tapered
and pointed, 68 long. Sensilla bases normal, at
posterior end of erista, set in a sensillary boss, centres
37 apart; sensillary setae filiform, about 180 long,
The crista and sensillary boss are invested by a
narrow scutum, which tapers anteriorly, leaving the
anterior third of the crista without this. The scutum
is free of setae, but is set in the normal plicare idiosomal
skin with its characteristic setae. Behind the sensiflary
REC, S. AUST. MUS, 1912); 169. 20K)
dri LON
boss is a conical continuation of the erista, probably
subcuticular, about 50 long.
Eyes 2+2, each lateral pair set in a small oval plate
about 38 long by 25 across. Anterior eye considerably
the larger, Cornea 18 «20, posterior eye more or less
transverse, with cornea 164 Eyeshields set at the level
of the junction of the middle and posterior thirds of
the crista; centre of each eye shield about 171 distant
from the midline of Ihe crista,
Various dimensions of the type and paratypes are
shown in the following Table 14.
Dorsal idiosomal setae arise [rom a small papilla set
in a broad plate. They are heavily ciliate, more so
distally, and vary considerably in length, A clear
division between two distinct length-groups cannot be
made. The setae tend to splic or branch in about their
distal third, and these branches may branch again. As
all parts of the scobillum are heavily ciliate with coarse
cilia, the end of the more branched setae appears
commonly besom-like, these remarks applying parti-
cularly to the longer setae. In the anterior part of the
dorsum there are comparatively few of the longer setae,
except immediately around the sensillary area of the
crista, but they are more mumerous towards the
posterior part of the dorsum of the idiosoma.
Ventral surface (partly from ACB749) provided with
numerous but shorter ciliate setae, mare uniform, and
with only a slight tendency to branch terminally,
Genitalia oval, 235 lony by 165 wide, with three oval
suckers alongside the iitroitus on each side, anterior
sucker 57 long by 29 wide, middle 4834, posterior
4936. Each valve of the external genitalia with a single
row of tapering setae 32-45 long. No internal
chitinization seen to genitalia, so the specimen ACB749
is possibly 4 female (damage to the specimen making
this & little uncertain),
Anus oval, about 62 long by 51 wide, with (wo valves,
each valve with 4-6 ciliate setae 31-35 long.
Legs moderately long and thin, appearing normal
for a trombidioicd mite, | 1670 long, TH 1270, If
incomiplete, [V ca 1820 (lengths Including coxae and
claws). Legs with a normal degree of setation, Many
of the ordinary setae (scobalae) are unilaterally and
regularly ciliate, thus making them classifiable as
pectinalae (see above). Dimensions of leg segments of
TABLE 14. DIMENSIONS (um) OF SOME CHITINIZED STRUCTURES OF ADULTS OF SCGP/TROMBIUM RETENTUM
TYPE SERIES
Identification Number HWamersley
Srructure ACB750 ACB748 AOBT49 f1934) (1¥455
Crista (including 313 ca 285 cu Jos - —
sagilla)
SB 37 - Bb - 43
Tal 362 & 128 347 » LOS — Wi «up JHO ~ Lid
Til 339 334 — 314 430
Gel 245 248 — — =
TalV ca 300» 97" 278 @ 95 — -
TiV 402* 388 ~ . _
GelV a7a* MD — = =
DS 16.55 20-57 iRod6] 1050 ZU, 34
* This leg has been dislocated and damaged, buy | preswme that i has been correctly ident {ied.
Y Damaged specimen, and only a few of the lonwer setae are available
QDONTACARUS SWAN MSP, 198
lectotype: tarsus | 362 long by 128 high, tibia | 339,
yenus | 245, tarsus [1 252% 70, tibia HT 245, genu I 168,
tarsus [IL 265% 86, Uibia LLL 286, gen 111 195, tarsus
1V 300%97, tibia TV 402, genu TV 274 (all tarsi
measured without claws). (The preceding assumes |
have identified legs 111 atid [V correctly.)
Tarsal claws sickle-shaped, unciliate, slighUy blunted,
appearing of normal trombidioid facies,
Palpi rather slender, normally setose, Palpal tibial
claw slout, somewhat blunted, A blunted large dorso-
medial accessory claw (a thickened seta) alongside claw,
and several smaller thickened peg-like setae behind it.
One long whip-like seta present on lateral aspect of
tibia terminally, over-reaching claw. Palpal tarsus an
elongate oval, with usual setation of ciliate and sensory
scetac,
Cheliceral fangs slender with about 12 indistiner
teeth along the flexor (upper) border.
Material Examtined
The only material available is the three individually
mounted specimens of the original type series.
Womersley stated (1934, p. 193) “As the specimens have
been remounted for further examination. ..”, but
only tWo of the specimens have been remounted fram
the original balsam to water-miscible medium or
media. | suspect that the two slides remounted in Lhe
period 1933-1934 were remounted again in about
1944-1945, for reasons given below, Details of the three
specimens are as follows:
(1) Specimen ACB750. Lectotype, Specimen
mounted on a slide 75 mms*25 mm, squashed, in
yellow-stained medium, The only trace of what was
possibly the original labelling is a small pink label with
“Cotype” in red (LHS). On the RAS of the slide is a
label in Womersley’s writing which reads Calothrom-
bium/retentus (Banks)/Co-t [in red]/with Iridomyrmex
nitidus/OQcean Grove, Vic./No, 17 Lea Coll,
(2) Specimen ACB749. A slide similar lo the
preceding, with the mite squashed and disrupted into
three main parts and various lesser parts, in the same
medium, It is labelled with a small pink label “Cotype”
as in the preceding (LHS); on the RHS is a label in
Womersley's writing Calothrombium/retentus (Bks)/
Co-1 fin red} Qcean Grove, Vie./No. 17, Lea Coll,
This specimen is clearly a female, as abouf 20
spherical or spheroidal eggs can be counted in the
fragments. These have a normally pigmented chorion,
and measure abour 140-195 by 120-160 in the least
squashed specimens. There 1s no evidence of an embryo
in any of the eggs. These eggs appear as normal
Prostigmata-type eggs.
All legs are disrupted, and (the lower (separated) part
of the idiosoma retains only the trochanter 1 and
trochanter Ll on one side, and trochanter 1V on the
other,
This slide contains fine acicular crystals which would
confirm that it is mounted in a chloral hydrate-
containing medium, at least in part, AU the time When
Womersley remounted it in 1933-1934, he was using
exclusively gum-chloral media, However, a later
possible use of a polyviny! alcohol medium cannot be
excluded; there is clear evidence of two zones in the
mauntant in the slide, The slide contains also some
instar of an insect, possibly a prinitive one, and also
a rather stout and short trombidioid leg, different {rt
character trom the legs of Scapi/rambium refentune.
How this extra biological material got into the
Mountant is purely a matter for speculation.
In my opinion the specimen is in no condition to be
designated a lectotype.
(3) AC'B748, A slide of an almost undamaged adull
mite, Mounted on its side in balsam (thus indicating
that the other two specimens have been remounted
trom balsam to water-miscible media), with a cover-
glass of 19 mm diameter, It bears in the writing belicved
to be that of Nathan Banks (Dr E. G. Matthews, Senior
Curator of Insects, South Australian Museum, who is
familiar with the writing and labelling of A, M, Lea,
advises. that this writing is not Lea’s, so it may therefore
confidently be attributed to Banks.): (R.HL. label) Rhyn-
cholophus/retentus/Bks/Microthrombium [this word
in Womersley’s writing] /Ocean Grove/Victoria/ with
Iridomyrmex nitidus/Corype/No. (1 interpret this
figure as 14) i4/Lea coll; (L.H, label) (in Womersley's
writing): Calothrombiurm retentus (Bks)/Co-t {in red]
/Qcean Grove/ Vic. No, 17/Lea Coll,
As it is mounted on its side with its legs partly flexed
(see Fig. 10), it is clearly not (he specimen mainly used
by Banks for his original description. Despite the fact
that it is the only specimen in which the origirtal data
remain on the slide, it is not the most suitable to be
seleeted as the lecratyne,
Further Remarks on the Type Series and
Lecloivpe Designation
Banks (1916, p, 226) recorded three localities for thos
species: Lal Lal, Ocean Grove and Sea Lake in Victoria.
Despite this, all three slides are labelled “Ocean Grove,
Vic.” and “No. 17 Lea coll" in Womersley's Writing (see
above), There were clearly at least two original slides
(and probably three). The remaining balsam mount
(ACB748) has clearly never been interfered with. The
balsam mountant has yellowed in its more peripheral
parts.
According to Banks, the specimen Irom Lal Lal was
captured with the ants of Polyrachis hexacantha, while
the specimens from Sea Lake and Ocean Grove were
captured with J/ridormprntex rutidus. Accepting the
accuracy of the data on slide ACB748 in Banks's
writing, and if we aceepe that the notation of
Womersley on slide ACB750 about the species beine
“with Zridomurmex nilidus' is correct, it may be
deduced that this specimen, the lectotype, in taul came
from Sea Lake, and that specimen ACB749 in fact came
trom Lal Lal. Whether we are entitled ta place any
196 REC. S. AUST. MUS. 19(12): 169-200 June, 1986
RCL
0)
100
RTe Fel
La
LT. x\,
500
ym erie? ,
Rta fF ge
RT.
GY LT
p \
FIG. 10. Scopitrombium retentum (Banks). Adult, specimen ACB748, mounted on its side.
ODONTACARUS SWANT NSP 197
reliance Upon such a deduction is conjectural, in view
of Womersley’s other obvious inaccuracies with data
on the slide labels.
All three specimens have the distinetive type of
dorsal setation recorded, and the tabular data of the
chitinized parts also confirm that these three specimens
are conspecific, in addition to the opinions af Banks
and Wamersley, For re-description, a lectotype needs
to be selected_ For this purpose, all three specimens in
their present state have some defects. Specimen
ACB748 is the only one retaining a presumably original
label, but is unfortunately mounted on its side.
Specimens ACB749 and 750 have been badly damaged
by being remounted at some stage, undoubtedly by
Womiersley, and possibly at the time of his 1945 stucly,
Despite the serious damage these latter specimens are
more suitable for the usual standard descriptions used
in prostigmatic and other mites, as the crista, eyes,
dorsal setae and other parts remain deseribable.
Unfortunately Womersley was afflicted by poor
vision, and he was often extremely destructive to
specimens in-an effort to make them easier to see. He
likewise erred frequently in transcribing locality and
host information and often destroyed earlier labels,
One may therefore conclude that Womersley mis-
transcribed the lovality and other data tar the two
specimens ACB749 and 750, and presumably one of
these came fram Lal Lal, and the other from Sea Lake,
but it appears there is now na way of determming
which came Tram which.
In his writings upon this species, Womersley even-
tually (L945, p. 107) was aware of the three different
localities in Victoria from which this species was
collected, but this is unfortunately not reflected in his
slide notations, which were presumably made first for
his 1933-1934 study, but then re-transcribed for his 1937
study, in which he used the generic name Culothrom-
bium Berlese, 1918,
There is nothing in the description and figures of
Hanks which allows the selection of one specimen as
against the other for designation as a lectotype, Several
important diagnostic characters were overlooked or
misinterpreted by Banks. Thus he placed the species
in the Erythraeidae, instead of the Trombidiidae (as
then understood), He recorded only one eye on each
side instead of two, and tailed lo see the sagitta to the
crista. He gaye almost no morphometric data, and his
description of the species is such that, as far as present
evidence allows, it could equally well be applied to
either of these two specimens (or even three), as was
probably intended by Banks in any case.
IL is regrettable that Womersley re-smounted these
specimens so crudely, and mistranscribed at least some
of the data on the labels,
OF the choices available, | select ACB7S50 as the most
suitable for redescription, and hereby designate it che
lectotype.
Taxonomic Placing of Scapitrambium retentun
This species has had a rather chequered history of
taxonomic placement, Originally it was, placed by
Banks in the family Erythraeidae, Correctly transferred
to the Trombidiidae (5./.) by Womersley in 1934, it was
initially placed in Microtrombidium Haller, 1882, in
the subfamily Microtrombidiinae, and later (1937)
transferred to the genus Ca@lofhrombium in the
Tanaupodinae, In 1945 Womersley recognized its
alfinities with other Leeuwenhoekiinae, and transferred
it to this subfamily, which he elevated to a family, as
a member of Acomatacarus Ewing, 1942,
The suite of characters listed above, however, set this
species apart [rom the other members of the subfamily
Leeuwenhoekiinae, and distinguish it from the Neo-
trombidiinae Feider, 1955, whose systematic position
was clarified by Lindquist and Vercammen-Grandjean
(1971).
The following is a key to the adults and nymphs of
the subfamilies of the Trombiculidae (after Crossley,
1960; Lindquist and Vercammen-Grandjean, 1971) for
which adults or nymphs are known,
1. Sagitta of the crista with one seta or without setaey if
such a sagittal seta is present it is usually expanded
somewhat or well ciliate. Palpal femur with few setae,
equally numerous as, Or less numerous than, in palpal
gen, Sensillary setae of crista ciliate (except Blankaartia
Oudemans, 1911) ..-.... 0.00... ... Trombiculinae
Sagitta of the crista with two setae, Palpal femur with
numerous setae, more numerous than in genu,
Sensillary selae of crista filiform ......... rakes thmad?
2(1) Eyes 2+ 2. Body setae conspicuously triramous, Coxa
of af least the posterior leg with mosaic ornamentation.
Two pairs of elongate elliptical discs or suckers to
external genitalia In both nymph and adult ,,.,..
occ ete beets beep ae ey seas. Neotrombidiinae
Eyes 0+ 0, 1 +1, 0r2 +2. Body setae ciliate, and may
be branched or of other form, but nor conspicuously
triramous, Leg coxae withoul mosaic ornamentation.
Adult with three pairs of oval discs or suckers along
each side of external genitalia Leeuwenhoekiinae
Generic Classification of Adults and Nymphs
in the Subjamily Leeuwenhoekiinge
Recent classifications of the status of the various
genera and subgenera of a number of the Trombiculi-
dae, particularly the subfamily Leeuwenhoekiinae,
togeiher with the present study of the adult of Odonta-
carus, and the separation of Scopitrombium, allow
some revision of the generic classification.
The following is an attempt al a key to separate the
genera, for the adults and nymphs, of the Leeuwen-
hockiinae.
$c Byes; wbsertanaiisaisteree ele tent erties ite<teese 2
Eyes present ...,.,.-)..- ols’ 437 Pe ee
2(t) Posterior dorsal body setae in two distinct forms, rhe
shorter spindle-shaped, pointed, ciliate, the longer
sword-like, with ils proximal part ciliate ..-,..,..,
Ae NM ty hl wie gr agsta n{eped Odontacarus Ewing 1929
Posterior and other body setac tend to be locally uniferm
eee Perea ery ol poe ee ee ae |
198 RECS. AUST. MENS. 1902); 1A9-200)
3/2) Posterior dorsal setae leallike. Legs not longer than
body- Tarsus t pyciform. Cheliceral blades narrowed,
nearly shearlike .,.,,.,.., Mhartania Ewing, 1944
Posterior dorsal setae somewhat expanded, but not
leaflike, Cheliceral blades shearhke, nor fiarrowed .
~ 22 eee... Atvematacervs Ewing, 1942
a(loRves Pe) oe eee ee
Po aes eo Pe ee oe Pel ake Cee be Poe
5(4) Saviltal setae expanded. Cheliceral blades long and
dapgver-like Xenodonigcverus Loumis and Goll, 1973
Sagittal setae not expanded. Cheliveral blades nat long
and dagger-like ..... -.- pee ee ke,
4(5) Postenor dorsal setae not expanded, ciliate, and ending
in long attenualed tips, Crisla prominently punctate.
Lees longer ihan body. Tarsus | cylindrical, about 2.5
tinves as long as High .....-. Chaetia Brennan, 1946
Posterior dorsal setae Somewhat expanded, not ending
in long allenuated tips. Crista a harrow rod, not
prominenily punctate, Legs of normal length, Tarsus
| cylindrical, about 6.2 times ws long as high .
rete peters ce by Comatacarus Ewing, P42
7(4) Body constricted, to 4 figure-ol-eight shape. Posterior
dorsal selwe not expanded, rather elongate and ending
in a hook-like process. Tarsus | cylindrical -..-. -
Sey eee =, Hannemania Oudemans, LY.
Body not constricted, Posterior dorsal setae terminally
turked or branched. Tarsus 1 an elongate oval ,,.
Ob CUR erred ee Metee ten Scopitrombium gen, ney.
In preparing the above tabular key | have accepted
the separation of the Neotrombidiinae (or Neotrom-
bidiidae) from the Leeuwenhockiinae, proposed by
Lindquist and Vercammen-Grandjean (1971),
[ have also omitted the genus Parvithramisriumn
André, 1962 (whose sole species is RP crassifersale
André, 1962, from Angola), which its author placed
in the Leeuwenhoektidae, In the somewhat speculative
revisions of the classification of the Trombidioidea
which have been proposed in recent years ( Vercamimen-
Grandjean 1973a, ty; Feider 1979) the genus Purvi-
thrombium has been placed in the Anomalothrom-
biinae Feider, 1955 (later elevated lo Anomalothrom-
bidiidae and Asomalathrombidioidea by Feider, 1979)
along with Anormalothronibiugt André, 1936,
Anomalothromtiym adults, the only instar known,
possess a small “naso” with two setae somewhat
resembling the sagitta of the Leewenhoekiidae and the
Trombiculidag, This genus appears to reseynble the
genus. Spelueothrambiun Willmann, 1940, and as far
as descriptions allow (Andre, 1936, 1935, 1939, 1945,
1958; Meyer and Ryke, 1960 (for A. curiosetosum
(Meyer and Ryke})) Robaux, 1965) could be placed in
the subfamily Spelaeothrombiinae as redefined by
Robaux (1968, 1972).
It appears probable that the interrelationships
between these mites, at present in a state of taxonomic
instability, will be clarified only When the larva-adult
correlations are known, at least in the first Instance
ACKNOWLEDGMENTS
{f am greatly indebted to the South Australian
Museum for access to type and other material in the
acarological collection, | also thank Mr L. G. Veitch,
June, 1986
Principal Research Scientist, CSIRO Division of
Mathematical Statistics, for advice and assistance in
stalistival matters,
The National Health and Medical Research Council,
Commonwealth of Australia, provided financial
support for these studies.
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HISTORY OF THE DISCOVERY OF SPELEOGNATHUS AUSTRALIS
WOMERSLEY (ACARINA: TROMBIDIFORMES), WITH NOTES ON ITS
NATURAL HISTORY AND BEHAVIOUR
BYR. V. SOUTHCOTT
Summary
The early history of the discovery of Speleognathus australis Womersley, 1936, is recorded. In
Australia over 100 specimens of the mite were observed personally on the water surface of three
cattle- (and horse-) troughs between 1934 and 1939 at Glen Osmond, South Australia. Despite
hundreds of observations in the subsequent years on the fauna of cattle-troughs and other water
surfaces, including the original sites at Glen Osmond, and many other sites in Australia and Papua
New Guinea, no further specimens of this species have been discovered in Australasia.
HISTORY OF THE DISCOVERY OF SPELEOGNATHUS AUSTRALIS WOMERSLEY
(ACARINA: TROMBIDIFORMES), WITH NOTES ON ITS NATURAL HISTORY AND BEHAVIOUR
by
R, ¥. SOUTHCOTT
Honorary Research Associate, South Australian Museum, North Terrace, Adelaide, South Australia 5000
(Manuscript accepted 23 January 1985)
ABSTRACT
SQOUTHCOTT, R, V. 1986, History of the discovery of Speleognathus
australis Womersley (Acarinat Trombidiformes), with notes on
its natural history and behaviour. Rec. 8. Aust. Mus, 19(13):
201-212.
The early history of the discovery of Spe/eognathus
australis Womersley, 1936, is recorded, In Australia over
100 specimens of the mite were observed personally on
the water surface of three cattle- (and horse-) troughs
between 1934 and 1939 at Glen Osmond, South
Australia. Despite hundreds of observations in the
subsequent years on the fauna of cattle-troughs and
other water surfaces, including the original sites at Glen
Osmond, and many other sites in Australia and Papua
New Guinea, no further specimens of this species have
been discovered in Australasia.
This species has been recorded from two other
locations as nasal endoparasites of bovids, once being
found in the nasal fossae of Zebu cattle, Bos indicus
L., at Astrida, former Belgian Congo, Africa, and on
another occasion in the nasal fossae of the North
American bison, Bison bison athabascae (Rhodes),
near Cache, Oklahoma, United States.
Observations of mile behaviour on the water surface
are also described.
INTRODUCTION
Speleognathus australis was described in 1936 by
Herbert Womersley (1889-1962), Entomologist, South
Australian Museum. [n his original account Womersley
(1936) gave the following field data;
“Locality—Type, one of two specimens taken in moss
al Glen Osmond, Adelaide, South Australia, in July
1934 (R. V. Southcot); four other specimens from
same locality in January 1935 (RV.S.).”
All of the specimens collected had been caught
running over the surface of the water in three (or less)
cattle- (and horse-) troughs, at Glen Osmond, South
Australia, and this information had been passed on to
Womersley. At the time | had been supplying him with
large amounts of moss, leaf litter and soil from various
localities in South Australia and Victoria, for his
studies on the Collembola and Acarina. In view of the
volume of this material that Womersley was handling,
his initial mistake is understandable. As | was only 16
years old at the time, perhaps understandably he did
not check the accuracy of the locality record before
publishing,
However, this species of mite was unusual, not only
in its morphological similarity to the ereynetid mites
(Riccardoella) that are common upon slugs in the
Adelaide area, but also in its behaviour, Its manner of
running on the surface of water showed that it was
quite at home there, As far as | was concerned, there
was never the slightest question that every specimen
found in Australia up to that time (and subsequently)
had been collected only upon the water surface of these
catule-troughs. [ had found that in collecting mites,
Collembola and other anthropods, the surface of each
of these cattle-troughs (about 45 cm across by about
150-180 cm long) provided an area for sampling in
which all specimens were clearly visible, and at the
collector's mercy apart from the occasional water-
strider, water beetles, or other truly aquatic insect, and
could be captured and bottled in alcohol without
difficulty.
On the surface of the water, these light brown
Acarina stood out by appearing to be either fully
aquatic, or at least adapted to a life at the surface of
the water, Specimens were collected from 1934 onwards
and taken to Womersley, and repeated observations
were made upon their behaviour by myself, both in the
field and in the laboratory. These repeated observations
were described to Womersley. In fact, on one occasion,
I took Womersley, at his own request, to the troughs
in question, some time before the publication of his
1936 paper, and possibly as late as April-May 1936, the
exact date not being at present available. (Womersley
and | lived in adjoining suburbs at the time, and the
troughs were within easy walking distance},
It was recognized that the mites’ unusual mode of
life at the water surface indicated an adaptation to a
moist surface, and the possibility of their being
nasicolous parasites of cattle and horses was
considered. Following the joint inspection, Womersley
arranged for the noses of the caltle to be swabbed by
a veterinarian on 20 May 1936 (see further later), but
no mites were revealed.
When Womersley’s paper appeared in 1936, | again
advised him that all specimens had been collected at
the cattle-troughs and asked him then and on several
2
later occasions to correct the mistake al some time itt
the future. This Womersley declined to do.
In 1948 Elizabeth Boyd deseribed a secund species
of mite in the family Speleognathidae, which
Womersley had erected in 1936, and nanied it
Speleognathus sturn, since it had been obtained trom
the nasal passages of starlings (Sturnus vulgaris Lin
North America, (A specimen of the mite had also been
captured from the hoat-tailed grackle, Cassidly
mexicans).
In 1952 Lawrence deseribed an intranasal mire from
a South African toad, Bufo regularis, which he named
Riccardoella ewer) Lawrence, 1952, thus recognizing,
as Boyd (1948) had done, che ereynetid affinities of
these miles. (This species was later removed to
Lowrencarus by Fain in 1957, and as now knowl as
Lawrencarus ewert (Lawrence, 1952).)
In 1952 also, Crossley deseribed a species of
intranasal mite from a small number of specimens
Obtained from the domestic pigeon, Colambu flea
domestica, in Texas, L5.A., as Speleognathus stars.
Thal species is at present known as Opfthalmognathus
striams (Crossley, 1952) (see Fain 1963, Domrow
1969). Cunously enough, (he discussion in Crossley’s
paper (1952, p. 386) contains the following:
“The type species of the Family Spelcognathidae,
Speleagnarhus australis, was found in moss and has
never been reported tram a bird, In 1948, Boyd placed
in this gevus a nasal mite (S. s/urni) trom the starling,
The many similarities of the two mites justifled this
action. Dr Womiersley (sic) has suggested (private
correspondence) that the type species, &. australis, may
be a nasal mite and that drinking water may be the
vehicle of transmission, The author is in complete
agreement, Speleognathus striatus has a hydrophobic
culicle, enabling the mite to float on the surface of the
water, and also is able to run quite rapidly.”
The second Australian species of speleagnathid (or
speleognathing—the Speleognathidae are now
xenierally considered as only of subfamily status, as
Speleognathinae, by recent authors) mite to be
discovered was described by Womersley (1953), who
erected the genus Boydaia lor Spefeegnarius siuent
Bayd, 1948, and included in this new genus & ange/ae
Wormersley 1959, found in “mucus under the tongue
of a frog Liemaodynastes fasmaniensis Gunther var.”
in Adelaide, by Miss Laura Madeline Angel, M.Se., of
the Zoology Department, University of Adelaide, while
she was searching for internal parasites. This speeies
is now placed as Lawrencerus angelae (Womersley,
1953) (see Domrow (1961, p, 379)),
Miss Angel has informed me (pers. comumn., 1976)
that che mite was found stuck in the mucus under the
tongue of a frog, in whose oropharynx she was
searching for trematode parasites, Na mire behaviuur
was observed, This male frog had come front Meadows,
Mr Lofty Ranges, South Australia, in October 1952
(coll, ELE. M. Anyel). The frog was recorded as
REC, S. ALIST MUS, Paid}: 2ob-212
dune, 1986
Lo tasmaniensis vat, platveephalus, Miss Angel has
commented that she has examined many hundreds of
(rogs frum the Adelaide to lower Murray River regions
without finding any other specimens of speleoguathine
mites. Mr Michael Tyler, Department of Zoology,
University of Adelaide, advises (pers. comm, 1976) that
the variety plarycephalus is no longer recognized, and
the name of this species of frog remains al
Limnodynastes tasmaniensis Gunther, 1858.
Womersley also included in his genus Beavdaia the
North American species Speleagnuihus striatus
Crossley, 1942,
In his 1953 paper Womersley declared that 8.
qustralts hact been collecred by myself “in moss and alsa
on the surface of water in horse troughs at Glen
Osmond in 1934 and 1935...
“As all the specimens were females and tram the
habitat on horse troughs it was thought that in the carly
stages [hey may have been parasites in the nasal cavities
olf birds or catile drinking at the troughs.
"The swabbing of cattle and the examination «al
birds, however, failed to show any evidence of this.”
A further mite from the Anstralian speleognathine
fauna was described by Womersley in 1954, as Bovdaia
derricki: Womertsley, 1954, This was collected upon
Rattus assimilis in Queensland, Australia, and was the
first recorded speledgnathine mite from a rodent. At
the time Womersley believed thal the association was
“probably accidental’. Subsequent work by Fain
(1955b), Dotnrow (1961) and others has shown (hat
several species of speleagnathiiies are parasites of
rodents. This Species is now known as
Paraspeleognathopsis derricki (Womersley, 1954) (see
Fain 1963), Romrow (1961) has recorded this species
in Queeustand from Ratius conalius and Rathaus ratrus,
as well as Rattus assimilis, No further referenee to
Speleernanius australis was made by Womersley it that
paper (1954), which was his last contribution on the
speleognathines.
In 1954 Cooreman recorded a luriher species of
speleognathine mite, Speleognathopsis gall, as a new
genus and species, from the fasal cavities ol the
domestic fowl, Gallus gallus b., al Astrida, i the then
Belvian Congo. In the most recent revisions of the
Nomenclature (Faig 1963, Domrow 1969), this species
remains as Svefeognerhapsis galli Cooreman, 1954,
Fain (198Sa) described briefly, in an addendum, a
speleagnathine mite from Astrida, as Speleognathus
bovis Fain, obtained trom the nasal cavities of
“havid és” (species af vatlle were nol named). Fain later
{1956b) deseribed this species in more detail, Fain
(1956a, b) stared thar che maxillary and frontal sinuses
were the actizal sites for the mites, The cane were still
referred ta as “hovides”; recently (1983), in response
to the presene author's tequest, Dr Fain has advised
rhat the mites were found in the maxillary and frontal
sinuses of Zehu cattle, ie. of Kos indicus L.
DISCOVERY AND NATURAL HISTORY OF SPELEOGNATHUS AUSTRALIS
Later in 1956 Fain (1956e) was able to state that S,
bovis was a synonym of S. australis, from an
examination of authentic material which | collected
from Glen Osmond. Moreover, the previous separation
was based on inaccuracies in Womersley’s descriptions
and figures (Womersley's own admission, quoted by
Fain (1956b, p, 662)), Fain was also able to include a
statement from myself correcting the attribution of the
Australian material to moss, and il is also corrected
in Southcott’s (1957) and Domrow’s (1961) papers
referring to speleognathine mites.
Drummond and Medley (1964) recorded thal on one
occasion, in November 1961, near Cache, Oklahoma,
two out of three North American bison, Bison bison
athabaseae (Rhodes), examined had an infestation of
Speleognathus australis in the nasal cavilies, with
evidence of an abnormal sinus condition, in the form
of blackened areas of epithelium, and the sinuses. filled
with brownish fluid,
Thus the early suggestion that the original
speleognathine discovered, Speleognaihus australis,
Was an endoparasite upon domestic cattle was
substalitiated by the observations in central Africa, and
in North America.
In subsequent years mites of this subfamily have
been recorded from a wide variety of birds and
mammals as well as other vertebrates, in several
continents. In his review of the Speleognathinae Fain
(1963) listed over 40 species, divided among 8 genera
and further subgenera (omitting Lewrencarus and
Batracarus, placed by him in the subfamily
Lawrencarinae of the Ereynetidae), Since that time
further nasicolous ereynetid mites have been described
from Australia—see Donirow (1965, 1969, 1975)—and
more general reviews bave been published by Fain (1969,
1970a, b, 1971a, b), Fain and Aitken (1969), Fain and
Hyland (1970, 1975), O'Connor (1978) and Hyland
(1979), The nomenclature of these mites at present
appears to have reached a degree of stability.
IL is an interesting point that had jt not been for the
fortuitous circumstance that | used cattle-troughs in
m1
my area as a means of sampling and collecting small
arthropods, Speleognathus australis night never have
been known to occur in Australia at all. Moreover, had
there not been, in Womersley’s estimation, an affinity
or resemblance to the mesostigmatic Mite genus
Spelaeorhynchus Neumann, 1902, it is sale to say that
the history of the nomenclature of this subfamily of
mites would have been vastly different.
The literature of these mites is now extensive, but
it is Not proposed lo atlempt any general survey here.
Instead, the purpose of this paper is (6 place on record
a number of early observations upon the times of
occurrence, and certain details of the behaviour of the
mite Speleognathus uustralis Womersley, which have
either not been recorded at all, or else recorded scantily,
and in a somewhat corrupted form in the literature.
It is considered that this is justified, since all the
Australian observations upon this species have been
either in the notebooks Gr memory of one person, the
present author, for 40 years or more. The miles have
not been seen in Australia since 1939, and the
opportunity of making observations on the species in
Africa may well be limited for some time to core.
THE OCCURRENCE OF SPELEOGNATHUS
AUSTRALIS IN AUSTRALIA
(a) Locality
Over the years 1934 to 1950 (as well as somewhat
earlier and later), the area studied at Glen Osmond for
Speleognathus australis consisted of some open
paddocks upon the lower and north-west slopes of
Mount Osmond (Fig. 1) at MR656807 to 655806, at
a height of 183-19] m (600-625 [l) above mean sea level,
on map Adelaide, 1939, No, 810, Zone 6, Sheet
IS4M/LV SE & SW (Military Survey of Australia,
1:63,360). This area is shown in Fig. 2. The two main
paddocks were separated by a stranded wire fence and
a roadway (Fig. 3). Both paddocks were used by a Mr
Goldsack for agistment of small numbers of domestic
cattle (Bos faurus L,) (invariably cows, as no bulls or
FIG. |. General view of Mount Osmond, South Australia, from the horth-west, from a photograph taken on 19 August 1938, All of
the Matter land around (he base of (he mountain has been commonty called Glen Osmond, The three cattle troughs upon which
all specimens of Speleognathus australis have been collected in Australia are al the middle level of (he photograph. Trough A 1s
situated somewhat to the right of the centre, in the dark clump of trees. Troughs B and © are further to the right, behind the line of trees,
i/ OO
REC. 8. AUST. MUS. 19(13): 201-212
‘ [ \Golf Hause N
June, 1986
Zz
FIG, 2. Map of the sites of the three cattle troughs at Glen Osmond, South Australia, from which all Australian Speleognathus australis
Womersley, 1936 have been collected, A, B, C are the sites of troughs A, B, C respectively. The grid squares are 1000 yards square,
or 974 m Square.
steers were included), perhaps 10 to 20 at the most, and
usually many fewer, Horses were also occasionally kept
there. Mr Goldsack had no objection to his paddocks
and cattle-troughs being used for purposes of
biological surveys, as long as there was no interference
with or disturbance of stock, and would occasionally
ask what was being observed upon the cattle-troughs.
The troughs were of the usual galvanized-iron type
common in Australia; half-cylindrical, about 45 em
across, and with a float-operated cistern to replenish
the water from the suburban reticulated water supply.
Of the three troughs studied, one (trough A) (Figs 4
and 5) was actually on the ground surface, being
supported against damage from the cattle by being
banked up with the clayey soil along the sides. Some
grass and weeds at times grew alongside the troughs,
but the soil around each of them was largely bare from
the continual tread of the cattle and horses. The other
two (troughs B and C) (Figs 6 and 7) were close
together, to the south-west of trough A, and were
placed with the bottom of the trough perhaps 15-30 cm
above the soil. Troughs B and C were supported by
wooden posts at each end, and trough B also had
additional supporting posts at about the middle (see
Figs 6 and 7). Although the soil around troughs B and
C was as well trodden as that of trough A, some weeds
managed to grow underneath them and at the ends
where access to the animals was denied. Each of the
three tanks had some protective boarding over the
cistern, to prevent it being damaged by the cattle. This,
fo some extent, limited observation of the water
surfaces.
FIG. 3, The roadway, fences, and lines of Eucalyptus cladocalyx separating the paddock of trough A from that of troughs B and C,
Looking southwards from near trough A. Photograph taken in about 1938.
DISCOVERY AND NATURAL HISTORY OF SPELEOGNATHUS AUSTRALIS
205
FIG. 4. Trough A and its immediate surroundings, looking up the valley of the centre of the previous photograph (Fig. 3). One cow
is present, The trough is on the ground, and is banked up by earth. Photograph taken on 28 January 1938, at about 6 p.m.
FIG. 5, Trough A, looking to the west; with the same cow at the trough as in Fig. 4. The large trees are Eucalyptus cladocalyx. Photograph
taken 28 January 1938 at about 6 p.m.
FIG. 6. Trougn B, looking to the north-east. The trough is in an open paddock. Note the straw on the ground, and many patches of
dung. The trees are Eucalyptus cladocalyx, Photograph taken 28 January 1938,
Trough A was overhung by a row of sugar gum trees, Eucalyptus cladocalyx bordering the adjacent paddock.
Eucalyptus cladocalyx, which even in the 1930s must All of these trees appeared to be part of one planting,
have been about 15 m high, and presumably anumber and according to Gill (1905, p. 5), had been planted
of the psyllids and other insects found on the surface in 1895,
came from the foliage of these trees. The other two
At times birds were seen to drink at the troughs, and
troughs were also placed about 10 m from a row of
feathers would be found floating in the tanks, There
206 REC. 8. AUST. MUS, 19(13): 201-212
* - 7
June, 1986
a... #
fo aE ht Wr, Mt
> ey * es = 4 2
a ly es | tg aa ‘
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}
ee, f
iy
{i
al
FIG 7. Trough C, near trough B. Photograph taken 12 June 1938.
TABLE 1. NUMBERS OF SPECIMENS OF SPELEOGNATHUS OBSERVED* ON THE CATTLETROUGHS OVER 1934-1940
1 2 3 4 5 6 7 8 9 10 11 12 Total
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Year
1934 + of 2 0 2 0 0 0 4
1935 — 2 0 0 22 3 2 1 0 0 0 0 30
1936 0 2 6 14 0 0 0 (0) 0 0 0 l 23
1937 5 3 18 1 1 0 1 1 1 0 0 8 39
1938 1 0 0 0 0 0 0 0 0 0 2 0 3
1939 4 0 1 25 1 0 0 0 0 0 0 3 34
1940 —_ — 0 0 0 0 0 0
Totals 10 7 25 40 24 3 5 2 3 0 2 12 133
— No observation
.. Mites not recorded, presumably not observed
* Not all of these mites were taken. When they were numerous I was careful to take only a representative number, to preserve a presumed
breeding population.
appears to be little doubt that these troughs were used
freely by the usual range of birds found in suburban
Adelaide at the time. The tanks were observed
occasionally to have large insects such as small
grasshoppers, floating in the water. It was observed also
that bees, wasps, and other insects used these troughs
for drinking, either directly from the surfaces, or from
seepages.
No mammals other than cattle and horses were
observed to drink at the troughs, but dogs and cats
probably used them occasionally.
Among the local wild mammal fauna were possums
(Trichosurus vulpecula), bats, rats, and doubtless
others; there were no local kangaroos or wallabies.
Although a few snakes occurred in the vicinity (the
brown snake, Pseudonaja textilis (Dumeéril and Bibron)
being occasionally observed), there was no evidence
over the years that snakes actually came near the
troughs. However frogs undoubtedly used these waters
at times.
(b) Times of Occurrence
It was my practice for a number of years, to walk
each week from my home at Glenunga Avenue,
Glenunga, to the Glen Osmond sites for routine
collecting. These sites are mentioned in my work (1946)
on the Erythraeidae, and the location of the three
cattle-troughs is close to the “second situation”
mentioned in that paper (p. 7). I recorded my findings
systematically until 1940, the only exemptions being
when I was away on holiday. The numbers of mites
observed over the years 1934 to 1940 are shown in
Table 1.
Inspection of Table 1 shows that the numbers of
mites on the water surfaces of the troughs were
maximal in autumn to early winter (March to May),
but there was only one month (October) in which no
mites were found during the seven-year study.
Over the years I gained the impression that the most
likely time to find mites on the surfaces of the troughs
was during a spell of fine weather, after rain. It should
be remembered that all the observations were made at
a time when there was no knowledge of what the host
animal’s identity might be, or in fact if there were a
host species of animal.
BEHAVIOUR OF SPELEOGNATHUS AUSTRALIS
I propose to quote in this section a number of the
observations recorded in my notebooks, since similar
studies have apparently not been made upon this
species of mite in its only other known localities and
sites of occurrence, in the nasal cavities of cattle in the
former Belgian Congo, or of bison on Oklahoma,
North America.
28.11.1936, There were two animals [Speleognathus australis|
found, one on the surface of water on horsetrough A,
DISCOVERY AND NATURAL HISTORY Lit SPELEOCNAIAUS AUSTRALIS
and another on the surface of horsetrough C. [Through-
Gul my noles the term horsetrough is oflen used,
wthouwgh cattle were by far the major users of these
troughs.| The mites seemed to be quite at home on the
surface of the water; both were running quickly over it,
sopping al limes gear small objects as though in search
for (mod, The one on horsetrough C mounted the side
of the iron trough about 4 crm above the level of the waler
and then descended. Then ir cane upon solid matetial
(floatiig) it ran over jt, but only for a few seconds at
most,
14, iv. 1936, L revisited the horselroughs, The miles wert presen.
in wbundance on horsetrough By none was present on
either of horsetroughs A and ©. On horsetrough B13
mites were counted; these were all running quickly over
the walter surface, apparently either ii search of food
and/or males, or suitable places for egg-laying, They
were all tuning fairly quiekly, with to noticeable
difference in their average speed, All, (6 the naked eye,
were of the same size. One mite was observed to leave
the surface of the water, then (ov mount the sides of the
trough to 4 height of about 3 em above the water, then
to run dong the trough for about 20 em keeping nearly
it the same level, then to descend and (o run over the
Water surface once more, | again noticed the preference
ofthe animals for the water surface: most of them did
noc leave the surface we all except fo tun over sonic
floating prass ete. This was about 3 o'cluck im the
afternoon; there were about 10 cattle near the trough,
There were no horses, although last Saturday there were
about 4 borses there as well as cattle.
lhe last wo or three days have been fine and warm without
exceptions last Saturday (LLiv,36) was rather cool, and
tnly one mile was found (horsetrough B), This seemed
to be in a state of torpor or dead: it was. lying on its
back on the water
Jtappears that a warn spell is necessary to bring the nites
Qul; | have noticed this several limes,
Possthle conclusions:
Mites were all of the same sex
. Mites were seeking tu Lay eggs.
. Front the fact that it has only been found on the water
surface of these three troughs it may be that it 1s
farasific Tn some stage in callle, eg, bronchial
passages.
The continued search for clues as to the life history
ol Speleagnatius ausiralis led (to a number of
observations on the milutiae of behaviour. As none
of these has been previously recorded, even thaugh the
mite has now been described for over 40 years, and as
they fave not been duplicated in the scanty
observations On this species in Africa or North
America, it seems worthwhile to publish them.
61,1937, One of the three mites taken on 61.1937 was kepl
inv glass jac on [the] surface of waver for several days
Various inscers were thrown on to the surface but the
mite was not seen to feed on any, Picce[s] of grass ete
were placed on che water, and although the mite ran
readily over them it always came [6 rest On the water
It can move its legs about 10-15 tites per sec, lt would
run near (he side of the jar and vibrate iis legs rapidly
wilhoul changing its position, The purpese ts to me,
ubscure, for the animal could easily move up che curved
surfave of the water to the edge.
One other al the rice was drowned by immersion lor
duu 1 heur.
13.1,1937, Two were found GG trough A, These were quite
active, Neat day they were pur in a tube (without water)
oe oe
2?
{and taken on a journey). The day was hor At night chey
were found to be dead.
27,41,1937. The speed of the mites on the surface of (he water
in the troughs was estimated wl 1-3 cm/sec,
2.1,.1937. Three were seen on trough AL T'hese attacked a fly
(Musea) in the water of the trough simultaneously [for]
Pood? [To lay) Eggs? (It now seems more likely that the
My might have had contact with oro-nasal mucus fron
the calile, and thus provided a suitable aitraclani for
the mites.
26,11), 1937, Two specimens [Were] captured running over the
surface ef trough A, These were placed in a tube, ori
the surface of some water trom [Ihe] trough, together
with some of the inseets [that had] fallen into the trough,
and 4 specimen of Cypris (Crustacea: Ostracoda) from
the water. After two days the mites were recorded as
dormant, Jandy) exhibited no signs of jife on touching
witha needle. [They] remained like this for some time,
The mités remained (hus and were almost conpletely
decomposed by 20.1937, On 14vili.1997 the residual
scraps Of mile skins were removed and slide-mounted,
16,iv,1937, One decomposed mite was found on the trough.
10.Vii.1937. Another dead mite was founds first mite found
since 16.iv.1937,
16,xi,1937, One mite found on trough A, tipped over or its
back. It moved ils legs actively when firs! found, bul
was feeble a lirtle later. Mite taken.
17.20.1937. Another mile found and taken, active. [L died by
19.xi1.1937.
27.x1,1937. Trough A was examined al 4-5 pam, Three
specimens were present, One was dead and decomposed;
the otber two were active, plump, running actively aver
the surface of the water, The two came together, circled
rapidly, and met (anterior end to anterior end). They
were like (his for a second or wwo, moving, their legs. all
the lime: then they separated by 2 or 3 mm, aud were
blown apart by the wind . ~~
The trough Was re-exainined at 8 pun,, when the
decomposed mile was as before, One was moving its legs
slowly; one was dormant with its legs drawn in under it
On trough C at 4-5 p.m. there was one very plump
and acuive mite at the surface of the water. At & pan.
the (presumably same) mite was floating, tipped over
on its back.
On 11.1938 | again cxamined the three (roughs al
8p.m., looking for evidence of possible nocturval
characteristics of the mite. | made the following nate:
The evidence of the past few Gays suggests stronuly
that the Spe/. does not leave the water by climbing oul
Jof] rhe trough—it is possible that i leaves it by means
of the cattle drinking there.
On 2.1.1938 the troughs were examined at about 7 pom,
(sunset), On trough A there was one Speleognathus, very
plump, and running quickly over the water. This | took.
Troughs Band C had none. The animal was observed
on water ina dish, [It moves by moving only the last 4
joints [i.e. segrnents] of its legs. The tarsus is practically
al aright angle to the water surface when at rest, or when
moving, It does not move by moving his trochanter an
the coxa, bul by soving its farsus and. metatarsus [1.e.
obial backwards and forwards ina more or less vertical
lane. The anunal was drawn in oulline, and in its mam
characteristics, and then killed with Camoy’s uid, Na
trace of eggs could be seen in the bady, Shape ts as
figured, widest anteriorly, narrower posteciorly, with
sides becooning approximately parallel, (Figure is
shown jn Fig. 8)-
Speleognathus is delicate and cannot withstand much
handling ur shaking about—either on water or dry.
208
lit
huhdrnfl itm fIPe:
[. athe #gnimg of
On 28.i.1938 I photographed troughs A and B (see
Figs 4-7) and recorded in my notebooks on that date:
The cattle are shifted about constantly in the paddocks
containing troughs A, B and C, but rarely are any of
the paddocks containing the troughs without cattle for
more than a few days.
A few further observations were made during 1939
and 1940, but I find from my notebooks that all records
for 1940 were negative, and the last living mites seen
by me were collected in December 1939. (Previous
statements implying that the mites were seen as late as
1941 were based on faulty recollection.)
REC. S. AUST. MUS, 19(13): 201-212
“1a3S Tes ah A, :
ai }
“— ‘ Ls |
ie ke S40 % an & ate tan }
FIG. 8. Speleognathus australis Womersley. Living specimen from Glen Osmond, South Australia, 2 January 1938. Drawn freehand
through the microscope, on the surface of water. The main lines have been darkened with pencil for reproduction purposes, otherwise
the drawing is untouched. The reticular patterning of the legs is omitted.
June, 1986
On 29.1.1939 I observed four mites on trough A, but
none on B or C. One of the four was alive; three were
dead (one apparently not long dead, one with legs
flexed under it, and one decomposed). I made the
following comments in my notebooks:
Conclusions: The hot spell has brought out
Speleognathus, following as it does fairly wet
weather .. . Note: Dead Speleognathi are found on the
water,
1 observed the living Speleognathus for about half an
hour fon the trough]. When the... (just dead)
Speleognathus was put in its way it ran over it without
DISCOVERY AND NATURAL HISTORY OF SPELEUGNATHUS AUSTRALIS
stopping, The animal seems to give no preference to light
or shade. Ran for a length of a metre (with lrequent
stoppings, and many turns, and doublings back, and
assisted by the wind, and water currenis. Slops varied
from 0-10 seconds, rarely the latter, usually about 3
seconds.
li was put near the edge of the trough, but did not
go up acall. Also, it took no notice of algue just below
the surface of the water.
Wasps, bees [are] observed drinking here, Cows drink
here, and dogs sometimes; and birds (magpies) [This
ineans the white-backed magpie of south-eastern
Australia, Gymnorhina hypoleuca Gould, These
frequently nest in fall trees, Such as the rows of planted
Eucalyptus cladocalyx bordering the paddocks
containing the troughs.| live here), and some bird
feathers [were] seen in [the] trough, Many ants [fare]
about; some fall in the water. One butterfly (Danaus
archippus) [Now Danaus plexippus plexippus (1. 1758)
(see Common and Waterhouse, 1972, p, 221)] [was] seen
drinking.
Troughs B and C were swarrning with Cypris. Troughs
A, Band C were swarining with nematidiform (almost)
larvae. Small bugs run over the surface of the water:
Water dirty, much alga present.
A sketch was made of the path of the Speleognathus
on (he water, over a small area, fo indicate its many
twists and turns. This is reproduced in Fig. 9.
~
\ aa) A
ternal dyresre Mg h $0,
; of pas of Spbsnc tone
ste AP F329 4 Meine bel aie 4
f
A Gl Biend ees ,
Uk fhe oe Seen fae gf’ Trimgk
PIG. 8 Drawing made on 19 January 1939, of the path of a
specimen of Speleognathus australis om a limited area of the
water surface of trough A.at Glen Osmond, South Australia,
reproduced at original size The wind direction, and water
current directions are show?. Jt is apparent from this sketch
(hat the mile was gol under the control of either rhe wind or
water curren(s, and could move al will upon whe water.
On 9iV,1939 | observed 25 of these mites upon the
surface of trough B; there were none on troughs A and
c.
All of these were dormant with the legs serongly
flexed, Not one was seen to be in a state of
Uecoumposifion, Not one was seen to give any miavenient
al all, although Several were stimulated by touching and
208
several were taken (7 in all)—5 were taken as [hey were,
2 others put in aleohol—not seen to move at all), The
mites were in any position &g, tilted on their posterior
ends, their sides, or [i normal position, None was on
its back, The occurrence iy most pernarkable . . . the only
sunny day recently is today . . . no cattle were drinking
at the troughs, or present in the paddocks... on
9,\v.19399, The five unpreserved mites were examined f
hours later, but were unchanged, and were then preserved
for histological study,
Observations continued, but did not throw any
further light on the presumed host animals or ble-
history of these mites.
By 1940 I was systematically noting the various
species of birds seen in the vicinity of the troughs, but
without drawing any significant conclusions,
During 1941 1 was able to make only a few
observations, and thereafter, owing to military duties,
1 was not able to inspect the area again uutil 1946. By
then troughs A and B had been removed. Trough C
remained and was examined at irregular intervals from
1946 to 1949, and occasionally later, to 1953, bul na
further Speleognathus was found.
POSSIBLE HOSTS OF
SPELEOGNATHUS AUSTRALIS
The possibility that these mires. were endoparasites
of the nasal or other respiratory passages of mammals
and birds drinking at the troughs was entertained quite
early. Since the only residual local marsupials were
arboreal ones, such as Trichasurus vulpecula and
smaller species, it did not seem likely that marsupials
would prove (o be the hosts, The animals drinking mast
commonly ai the troughs were cattle, with a smaller
population of horses, but ii would seem likely, pritva
facie, that if such mites were parasites of the nasal
spaces of cattle and horses, they could hardly fail (o
have been detected in Europe, the obvious source of
the Australian domestic breeds of cattle and horses;
not to mention the other domestic animals in South
Australia, such as dogs, cats, and so on. Similar
considerations applied to introduced rodents. Bats
appeared to be another possibility, but rather a remote
one, although insectivorous bats were not uncommon
in the neighbourhood, even if mainly crepuseulac or
nocturnal.
Other possibilities that were considered, either then
or later, were that the mites could be tracheal parasites
of water insects occasionally seen upon the troughs.
Conceivably also small gastropods might serve as hosts,
in the same way as Riccardoella utilizes the common
introduced slugs in South Australla, because of the
abviaus resemblance between Speleagnathus and
Riccardoella. However, no solid evidence was found to
support seriously any suggestions of a non-vertebrate
host.
Since the mites were reasonably common at times,
and since the cattle were the commonest large mammal
ulilizing the troughs, it seemed worthwhile Lo
investigate further the possible role of the cattle
210
My notebooks record;
20..1934, Cattle drinking at troughs A, Band € ar Cilen
Osmond, South Australia, had cheir nostrils swabbed-.
No Speleog. were obtained.
This is not surprising, and probably the resull does
not mean yery much, since more than a month had
passed since any ol these mites had been seen on the
troughs. (Tests performed by C.S.0.R, officers, at
suggestion of H, Womersley (to them, that is).)
L was not a witness to the event, the information
having come fram Womersley (1953, p. 82) who referred
lo it briefly.
After this negative result, no further attempt was
made ta locate the mites in the nostrils of the cattle.
Subsequently the possible relationships remained
matters of speculation, as the various new species’ of
speleognathines were discovered in North America,
Africa, and other continents, The next species to be
discovered was by Elizabeth Boyd (1948), in North
American birds.
At the end of 1938, under the existing means of
study, I had summarized the possibililies of host-
relationships in my notebooks as follows:
(a) Nature of the appearances.of the animal en the water
surface
1. This cannot be fortuitous—lrom ease of jis
progression;
2, There are no streams left in this locality—oven iw
winter they do not run (although they do contain
water for a while)—and only remain with water
in them for a short (ime. The troughs are tram
1-25 m trom the creek bed,
(b) Fauna which might be related
(1) Non-aquatic
Birds (the word especially added in }
pencil at some later date, and the
word birds underlined) )
Cattle )
Insects including wasps, bees, bugs, etc
(2) Aquatic )
Trough contains a rich fauna of Cypris, In Dec, 1938
many nematodes (?) were present. (There is a good
collection Of fllamentous algae in each trough). ,
(c) Flora which might be related
(1) Non-aquatic—eucalypts, grass, weeds
(2) Aquatic—green algae mainly ) Spirogyra
Addendum 1i.1939
Only 3 Speleog. [were] Seen in L938 ¢and caught). (ep,
1935-with 26, 1936 with 23, 1937 with 34 (approx. figs.).
Tt may be that the continual taking of the Spel. has
reduced their number considerably.
That surmise was contradicted by the finding of a
large number in 1939; in fact the count of 25 for March
1939 was the highest for any individual month over the
whole period of the observation,
OTHER ATTEMPTS TO FIND SPELEOGNATHUS
IN AUSTRALIA AND PAPUA-NEW GUINEA
Over the last 40 years I have [ooked systematically
for Speleognathus australis (and related mites) when
travelling in Australia and Papua-New Gujnea, Na
cattle-trough encountered which contained water was
ever left uninspected, Despite searches in Papua-New
Guinea (two sojourns) aid in every Australian state
drink
here
REC, 8, AUST. MUS, 19(14): 201-212
Junie, 1986
except Western Australia, none has been found.
Additionally, a search has been made for intranasal
mites in birds and other vertebrates from £942 onwards,
as Opportunity permitted, but again all examinations
failed to reveal any speleogriathine mites,
After the distovery of S. sturni by Elizabeth Boyd
(1948) in North America, 1 decided to make more
systematic and [requent efforts to find these mites at
bird drinking sites. [ erected two troughs at my then
home at Unley Park, a southern suburb of Adelaide,
by cutting a 44-gallon drum in halves longitudinally,
One (D) was erected about a metre above ground level,
on the stump of a cypress tree, and the other (E) was
erected on a wooden frame about two-ihirds of a metre
above ground level amidst trees, bushes and other
vegetation. Birds were encouraged by suitable feeding
to drink at the troughs. Trough D was observed
regularly for nearly 4 years, and trough E for over 3
years. Observations were made daily, at times oftener,
ever much of this period. Despite these efforts,
however, no speleognathines were found,
During 1952 and 1953 1] carried out systematic
observations of the horse-troughs still located around
the streets, marks and other open spaces around
Adelaide. Thus 40 observations were made in March-
June, 1953, and observations conlinued into 1954,
when ihey were abandoned. Again, no speleognathines
were found.
DISCUSSION
It may reasonably be concluded from these many
hundreds of observations that Speleognathus australis
must be an uncommon species in Australia, If it were
a common parasite of cattle, it is reasonable to believe
that it would occasionally come to the notice af
veterinarians.. There is, of course, no evidence at present
that this species of mite is concerned with disease
transmission in-any way, or in fact that it causes serious
harm to its hosts. Nevertheless, the lesions recorded
bricfly in one specimen af North American bison by
Drummond and Medley (1944) show that this species
of mite is capable of causing some tissue damage.
Presumably ihe mechanism is similar to that in, for
example, Riccardoella linucum (Schrank) (Ereynetidae)
which feeds on slugs and is capable of killing them
under laboratory conditions of heavy infestations
(Baker 1970a, b). Baker (1971) studied the ereynetid
mite Xenaparcarus africanus Fain, Baker and Tinsley,
1969, which lives in the nasal passages of the African
clawed toad Xenopus laevis Daudin, and demanstrated
biood in the gut by histochemical tests. Both of these
species of mites have mouthparts which appear capable
of penetrating epithelium.
Baker (1973) discussed this further and commented
(p. SI): “The feeding of ereynetid mites, involving the
production of a stylostome, appears to be basically
similar to that already described in such trombiculid
mites as Neolrombicula zachvatkini Schluger and
Trombicula autummnalis Shaw ,, .”.
DISCOVERY AND NATURAL HISTORY OF SPELEQGNATHUS AUSTRALIS Zu
Other groups of invertebrates have exploited the
same niches, such as various mesostigmatid mites, and
the family Trombiculidae (sensw lato) of the
Trombidiformes.
The second majar question which w studem of the
distribution of Speleoegnathus australis will ask, is how
car we explain the greatly separated distribution of the
species, known only from three recorded localities:
Glen Osmond in South Australia, Astrida in (he former
Belgian Congo, and Oklahoma in North America. In
the African and North American cases there was
evidence of nasal parasitization of a bovid, while in
the Australian case there was evidence pointing to an
association with cattle Only one species is now
classified in the genus Speleognathus; all other
speleognathines, from a wide variety of vertebrate
hasts, are now placed in other genera (Fain 1963),
We may reasonably accept that Speleognarius
australis is a bovid-adapted nasicolous species,
The problem remains, however, of its extremely
disjunct distribution as at present known. Although
T stated (1963, p. 328) that this was “easily explicable
since a number of the original cattle of Australia were
of African origin", it now seems that that explanation
was too facile, particularly in view of the finding af
Speleagnathus australis in North America, In any case
the proposal did not explain the lack of reports of the
mite in southern Africa, nor tron New South Wales
avd other parts of Australia.
At this point we have to leave the study of its malogy
and distribution with much of the mite’s natural history
in Australia at last accurately recorded in some detaul
but with some major ecological questions sull
unartiswered,
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BAKER, R. A. 1970, Srudies on the life history at Xiccardoella
limacum (Sehrink) (Aeari-Trombidiformes) J. met Plist. 4:
SII-5E9.
BAKER, R.A, 1970b. The food of Riccaridoella (itmacunr (Schrank)
-Acari-Trombidiformes and its relationship with pulmonate
malluses. J. nes. Aisr. de 821-530.
BAKER, RK, A, 197). Observations on aspects of muirilion in
Nenopacarus africanus (Ereynetidac: Trombiditormes), J ved,
Entomul, 8: 307313.
BAKER, R. A, 1973, Notes on the internal anatomy, (he food
requirements and development in the family Ereynetidae
(lrombidiformes), lcaralagia 15. 43-52.
BOYD, Elizabeth, M, 1948. 4 new iie from rhe respiarory tact
of the starling (Acarina, Speleognathidae), Prac. Eni, Soo
Washington Stk 914.
COMMON, I. T. B., and WATERHOUSE, DF 1972, “Butlerfhes
of Australia’, Angus and Robertson, Sydney.
COOKEMAN, |. 1954. Sur ui acarien nouveau, parasite des fosses
fasales des oiseaux ef remarques sur la famille des
Speleognallndae. 4nm FPerusitol fini, comp. 29; 426-432,
CROSSLEY, BA, Jr 1Y52, Two new nasal miles frorn cotumbi torn
birds. J Purasial. 38> 385-390,
POMROW, R. 1961, The family Speleognathidae in Australia
(Acarina), Proc Drrn, Yau, NAM 85: 374981,
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birds, Aearclagia 7- 43-48.
DOMROW, RB. 1969. The nasal mites of Queernshuid birds (Acar:
Dermranyssidad, Ereynetidae, and Epidermoptidae). Prac. Linn,
Sav NSM 93, 297-426.
DOMROW, R. 1975. A new spevies of Paraspeleograrhopsis Fain
(Acari: Ereynetidaey from aly Austtalian dasyurid marsupial.
Awa, Bn Sov. 14: 97-99.
DRUMMOND, R. ©. and MEDLEY, §..G. 1964. Occurrence of
Speleounathus vustralis Womersiey (Acarina: Speleagnathidae)
in the nasal passages of bison, J. Parasitol SO: 655.
FALUN, AL 19SSa. Sur Un nouvel acarien parasite des fasses nasales
de la perdis au Ruane-Urund?: Boydara preraisis n. sp.
(Speleoguartidae). Rev. Zool Bar afr $2) 144-149,
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mammiféres eb les oiseaux par Tes acariens cle la famille
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vhez les chauve-souris. ela Soe, Belg, Méad, Trap. 35; 689-700.
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VAIN, ov. LOSGH. Presence dacanens de la famille Speleognathidae
Woamerstey dans les (tosses nasales de mammiféres. Descriptian
de treis espéces nouvelles. Ann Parasirol. hum. comp Mb:
155-148,
FAIN, A. 1956c. Nouvelles observations sur les acariens de la fanville
Speleognathidae parasites des fosses nasales ehez les batracwns,
les oiseaux et les mammiferes. Ann, Poresitel, hurt comp, AV
643-662,
FAIN, A. 1957, Sur la pasition systématique de Riceerdvella ewert
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FAIN, A. 1963. Chaetotaxie et classitication des Spelcognathinae
(Acarina: Trombidiforines), Bull, Inst. Ray. Sei. Nat. Bele BY:
1-80,
FAIN, A, (969. Nouvelles Speleognalhinue parasiles nasicoles
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369-376.
PAIN, A, 19708, Momenclacure des poils idiosomaux et description
de tris especes fouvelles dans la famille Ereynetidac
(Treanbidifermes). Acarofogu 12: 313-325.
FAIN, A. 1970b Acariens nasicoles d'aiscaux et de mammiféres de
Brésil, IV) Nouveaux Ereynetidae (Trombidiformes) el
Turhinoptidse (Sarcoputoarmes) de la region de Relem (Nord
Brésil). Acwralogiu (2 326-538.
FAIN, A, 197Ia. Notes sur les speleognathines parasites nasivoles
des mamnuléres (Ereynetidae: Trambidiformes), Acarologia 12;
509-521,
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(Ereynetidae: Trombidifarmes), tearo/ogig thy 9¥1)2.
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et des mammifeéres clu Greésil. 1t. Ereynetidae de la region de
Belem (Nord Bresil), Bull Ana, Soe, Rov. Entom, Belg, WS:
43-4.
FAIN, A., and HYLAND, K, 6. 1970. Acariens nasicoles des oisetux
du Mexique. Il, Pamilles Ereynetidae et firbinoplidae, Bull.
Ann. Sac. Roy, Enfam-. Bele. 6: 37-46.
FAIN, A. and HYLAND, K. & 1975. Speleognathinaeé collected fram
birds in North America (Acarina: Ereyneridae). 4. New rark
Ent, Soc, 83; 203-208,
GILL, PF. 1905, "The History and Topography of Glen Osmond”,
Vardon and Pritchard, Adelaide.
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in the Turbinoptidac, Cytoditidae and Ereynetidae living inthe
respiratory passages ol birds, Jn “Recent Advances in
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Sov. London 121; 747-752.
O'CONNOR, B, M, 1978. A redescriprien of Speleoredens
michivensis (Foard). comb. (Acari) Ereynetidae), a nasal mire
212 REC. S. AUST. MUS. 19(13): 201-212 June, 1986
of microtine rodents, with comments on generic relationships _WOMERSLEY, H. 1936. On a new family of Acarina, with
in the Speleognathinae. J. New York Ent. Soc. 86: 123-129. description of a new genus and species. Ann. Mag. nat. Hist.
SOUTHCOTT, R. V. 1946. Studies on Australian Erythraeidae (10) 18: 312-315.
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(1889-1962). Acarologia 5: 323-334. Aust. 77: 65-66.
DESCRIPTION OF ODONTACARUS VEITCHI SP. NOV.
(ACARINA: TROMBICULIDAE)
BY R. V. SOUTHCOTT
Summary
Odontacarus veitchi sp. nov. (Acarina: Trombiculidae) larva from central Queensland is described.
The classificatory key to the larval Odontacarus of the Australian, New Guinean and south-east
Asian region is revised.
DESCRIPTION OF ODONTACARUS VEITCHI SF, NOY, (ACARINA; TROMBICULIDAE)
by
R. V. SOUTHCOTT
Honorary Research Associate, South Australian Museum, North Terrace, Adelaide, South Australia 5000
(Manuscript accepted 13 January 1986)
ABSTRACT
SOUTHCOTT, R. V. 1986. Description of Odentacarus veitchi sp.
nov. (Acatina: Trombiculidae), Rec, 5. Aust Mus. ¥9(14)-
213-217,
Odontacarus veitchi sp, mov. (Acarina:
Trombiculidae) larva from central Queensland is
described, The classificatory key to the larval
Odontacarus of the Australian, New Guinean and
south-east Asian region is revised,
INTRODUCTION
The author recently (1986) described various stadia
in the life history of Odontacarus swani Southcott,
1986, and revised the classification of the larvae of
Odontacarus Ewing, 1929, for Australia and adjacent
parts of the southwest Pacific area. Difficulties in the
placement of O, athertonénsis (Womersley, 1945) and
related species were referred to. In a cognate* study
detailed statistical analyses of morphological features
of this group of larvae were presented (Veitch and
Southcott, 1984), which showed significant metric
differences between the species presenting classification
difficulties: O, mecullachi (Womersley, 1944), O.
athertonensis (Wom.), O. “species S”{ (now O, swani
Southeott), as well as some further specimens.
This last-named group, from Queensland and
Papua-New Guinea, included a few larvae in poor
condition and hence unsuitable for detailed
description, but also a series of six specimens from
central Queensland (identification numbers ACB733A-
F) of which five specimens were in good condition,
sujtable for detailed measurements and description.
These had been considered as conspecific with O.
athertonensis larvae by Womersley (slide
identifications), and by Goff (1979), Re-examination
of this series shows that it represents a separate species,
which can be distinguished from other species of
Odontacarus (subgenus Leogonius Vercammen-
Grandjean, 1968), particularly by leg metric characters.
Al] measurements are in zm, unless stated otherwise,
Seta and other terminology is as given by Southcott
(1986) and Veitch and Southcott (1984).
* It was originally planned that that study would follow the paper
here shown as Southcott (1986). However, unforeseen publication
schedules reversed the order of appearances.
7 Unrelated to an undescribed species mentioned as “Ss” by
Vercammen-Grandjean (1968, p. 121),
Odontacarus veitchi sp_ nav.
(Figs LA-3, 2,A-B)
Description of Holotype larva, N1981390
(ACB733F):
Colour in life not recorded, presumably red, Length
of idiosoma (mounted on slide) 237, width 203; total
length of animal from tip of cheliceral fangs to
posterior pole of idiosoma 328,
Dorsal scutum slightly wider than long (rasus
included); nasus well-developed, narrow, with rounded
lip and slightly sinuous sides, meeting the body of
shield at right angles. Anterolateral angles of scutum
rounded, lateral borders slightly concave, running Lo
rounded posterolateral angles, posterolateral borders
concave, posterior pole evenly rounded, projecting, AM
scutalae tapering, slightly pointed, lightly setulose; AL
and PL scutalse similar. Sensillary setae normal, well
setulose in distal half. Sensillary sockets slightly
anterior to level of bases of PL setae, set slightly
obliquely.
Standard and other metric data for type series as in
Table |,
Eyes 2+2, sessile, oval, conjoined, near PL angles
of scutum, Maximum diameter of anterior eye 1%,
posterior 13,
Dorsal idiosomalae normal, slightly tapering, lightly
setulose, pointed at tip, arranged 2 (‘post-humerals’),
6, then in vague rows across dorsum, of ca 10, total 58,
Ventral surface of idiosoma with a pair of pointed,
setulose setae between coxae III, ca 36 long. Venter of
opisthosoma bears 48 further setae, 24-36 long, with
outstanding setules; posterior setae longer and slightly
blunted, similar to posterior dorsal idiosomalae., Anus
(uroporus) oval, 23 long by 15 wide; about 23 setae
anterior to mid-level of uroporus, and 25 setae
posterior to same level, Urstizma oval, in normal site
between the contiguous coxa [ and II of each side, I8
long by 14 wide.
Coxalae 2, I, 1. All coxalae long, well setulose,
tapering to a fine point, Lateral coxala 1 60 long, medial
coxala | 60, coxala II $1, coxala Ll ca $5.
Legs well-developed, normal, 1 416 long, [I 368,
111 430 (all lengths include coxse and claws), Leg
scobalae normal, pointed, well setulose. Scobalar
formulae (including mastalae): trochantets |, t, J,
femora 6, 5, 4, genua 4, 4, 5, tibiae 7, 6, 6, tarsi 20,
17, 13.
214 REC, S. AUST. MUS. 19(14): 213-217
FIG.
June, 1986
Odontacarus veitcht sp. nov. A-E Larva, holotype, NJ981390. A Dorsal view, legs on left omitted beyond trochanters. B Tips
of chelicerae, seen in transparency, C, D Dorsal idiosomal setae (c, d in A, respectively). E Ventral idiosomal seta (e in Fig. 2a). All
figures to nearest scale. F Diagram to show metric characters of dorsal shield. The symbol A indicates that this seta is shown in both
the dorsal and ventral figures for this mite.
Leg specialized setae as follows: SoGel.32d (24 long),
VsGel.72d (4), SoGel.72pd (27), Sofil.55d (18),
VsTiI.82d (4), SOTIL.85d (16) ie. slightly distal, also
anterior, to VsTil.
SoGell.25d (20), VsGell.64d (3), SoTill.36d (16),
SoTill.88d (15), SoGell1.27d (22), SoTill1,35d (20),
Both tarsus I and II bear a large central dorsal
solenoidala, I 16 long, IT 18 long. On tarsus I is famala
FaTal, 3 long, level with proximal Sola!. On tarsus II
is Falall, 4 long, proximal to Solall (see Fig. 1A). Tibia
IIL has two long mastalae (mastisetae); tarsus III has
one mastala. Pretarsal formula 1, 1, 0.
Tarsus I 91 long by 30 across, Il 74x27, II] 97x24
(tarsal lengths exclude claws and pedicle). Tarsal claws
normal, falciform, slender, with strong setules
(onychotrichs) on anterior and posterior; middle claw
longer and more slender than neolaterals, with weak
setules,
NEW ODONTACARUS (ACARINA: TROMBICULIDAE) 215
FIG, 2.
palp of paratype NI981385. (Both to nearest scale).
Gnathosoma normal, well developed. Combined
chelicerae bases ca 80 across; chelicerae 91 long from
tip of fangs to posterior pole of bases. Cheliceral fangs
stout, curved, pointed, with 3 or 4 strong retrorse teeth
along concave, flexor (=dorsal) edge, and 2-5 blunted
denticles along convex, extensor (=ventral) edge.
Galeala 30 long, with one faint setule. Gnathobasal
setae curved, pointed, 25 long, with several long setules.
Palpi normal, strong. Palpal setal formula 1, 1, 3,
8, with palpal setal formula B, B, BNN, So+7B. Palpal
PaScTit
Odontucarus veitchi sp. nov., larva. A Ventral aspect of holotype; legs on left omitted beyond trochanters, B Ventral view of
tibial claw trifurcate. No supracoxalae to legs or
gnathosoma.
Material examined
Queensland: Mt. Jukes, 6.ix.1951, 6 larvae, collected
on card, E. H. Derrick. Holotype with South
Australian Museum registration number N1981390; it
bears on the right hand label ‘“Acomatacarus
athertonensis Wom,” and the locality information given
above, also “G197”. The word “Paratype”, which was
216 REC. 8. AUST. MUS. 19(14): 213-217 June, 1986
TABLE 1. METRIC DATA FOR TYPE SERIES OF LARVAE OF ODONTACARUS VEITCHT SP. NOV. IN pm*
N1981390
Specimen ACB733F N1981385 N1891386 N1981388 N1981389
number (Holotype) ACB733A ACB733B ACB733D ACB733E Means
AW 67 77 77 71 73 73.0
PW 76 89 89 77 80 82.2
SB 25 29 27 26 29 27.2
ASB 53 58 56 S1 52 54.0
PSB 28 29 30 29 30 29.2
L 81 87 86 80 82 83.2
LA 21 22 20 19 20 20.4
LB 60 65 66 6l 62 62.8
LN 31 34 35 29 29 31.6
W 86 96 99 857 85 90.2
AP 29 30 32 29 32 30.4
AM 34 40 36 38 4] 37.8
AL 40 4) 42 38 40 40.2
PL 49 60 64 52 49 54.8
AMB 9 11 12 10 13 11.0
Sens 51 51 48 38 — 47.0
PW/LB 1,27 1.37 1.35 1.26 1.29 =
DS 36-58 36-62 35-66 33-62 35-64 35.0-62.4
MDS 33-35 31-38 33-38 33-35 35-38 33.2-36.8
PDS 36-38 38-42 40-43 36-40 36-42 37.2-41.0
Gel 54 59 59 53 58 56.6
Til 61 66 66 60 62 63.0
Gell 46 49 48 47 46 47.2
Till 49 54 54 51 51 51.8
Gelll 50 55 55 49 59 53.6
Tilll 67 68 71 65 69 68.0
AW/TiIIL 1,00 1.13 1.08 1.09 1.06 —
PW/TillL 1.13 1.31 1.25 1.18 1.16 —
* See the criteria given in footnoie to Table 6 in Southcott (1986).
+ Another estimate of this variate is 82 ym (see preceding footnote, and Veitch and Southcou (1984).
clearly in error, has been obliterated with white ink.
On the left hand label is written “Was marked
‘PARAT Y PE’/ACB733F/Odontacarus veitchi
Southcott HOLOTYPE/N1981390.”.
The five paratypes, from the same source and
collector, have similar labels, with serial numbers on
them as follows: N1981386, ACB733B, G193; N1981387,
ACB733C, G197; N1981388, ACB733D, G196;
N1981389, ACB733E, G206; N1981385, ACB733A,
G192. The unofficial (personal) registration numbers
with an ACB prefix have been used by me in previous
publications (Veitch and Southcott, 1984; Southcott,
1986).
Systematic position
To distinguish this new species, criterion number 17
in my key (Southcott, 1986, p. 180) should be altered
to the following:
17 Ventral opisthosomal setae 48-54 in number ..
pbb bite eeebeday elt lel pedis thease ted ptBaye oy atta ar tee LTA
Ventral opisthosomal setae in range of 35-45
TPL PUTA ole ala le ia 0 tle iat hel fi 18
17A Longer dorsal idiosomal setae 42-56 long.
Genu | 42-49 long. Tibia III 57-65 long.
Palpal setal formula B, B, BBB, So + 7B....
O. athertonensis (Womersley)
Longer dorsal idiosomal setae 58-66 long.
Genu I 54-59 long. Tibia III 65-71 long.
Palpal setal formula B, B, BNN So+7B......
her we be tee wae chews O. veitchi sp. nov.
Remarks
Further details of the analyses of the statistical data
for this and related species are given in Veitch and
Southcott (1984).
Nomenclature
The species is dedicated to Mr L. G. Veitch, CSIRO
Division of Mathematical Statistics, in recognition of
his assistance in the metric studies of Australian
trombidioid mites.
ACKNOWLEDGMENTS
I thank the South Australian Museum for access to
these specimens.
Thanks are due to the National Health and Medical
Research Council, Commonwealth of Australia, for
assistance.
NEW ODONTACARUS (ACARINA: TROMBICULIDAE) 217
REFERENCES VEITCH, L. G., and SOUTHCOTT, R. V. 1984. Discriminant
analysis using metric data from larval specimens of three species
of Odontacarus (Acarina: Trombiculidae). Aust. J. Zool. 32:
EWING, H. E. 1929. A manual of external parasites. (Charles C.
Thomas: Springfield, Illinois and Baltimore, Maryland). 519-526. am : ‘
GOFF, M. L. 1979. The genus Odontacarus (Acari: Trombiculidae) VERCAMMEN-GRANDIJEAN, P. H. 1968. The chigger mites of
in New Guinea, with descriptions of four new species. J. med. the Far East (Acarina: Trombiculidae & Leeuwenhoekiidae). An
Entomol. 15: 143-154. illustrated key and a synopsis; some new tribes, genera and
SOUTHCOTT, R. V. 1986. The genus Odontacarus (Acarina: subgenera. (U.S. Army Medical Research and Development
Trombiculidae). II. Observations on the life history and Command: Washing D.C. 20315). +
morphology of Odontacarus swani n. sp., and related forms. WOMERSLEY, H. 1944, Notes on and additions to the
Rec. S. Aust. Mus. 19: 169-200. Trombiculinae and Leeuwenhoekiinae (Acarina) of Australia and
New Guinea. Trans. R. Soc. S. Aust. 68: 82-112.
WOMERSLEY, H. 1945. Acarina of Australia and New Guinea. The
family Leeuwenhoekiidae. Trans. R. Soc, S. Aust. 69: 96-113.
RECORDS oF THE
SOUTH AUSTRALIAN
MUSEUM
VOLUME 19 NUMBER 15 AUGUST 1986
No. 15 A CHECKLIST OF HELMINTHS FROM AUSTRALIAN
BIRDS
by PATRICIA M. MAWSON, L. MADELINE ANGEL and
S. J. EDMONDS
SOUTH AUSTRALIAN MUSEUM
North Terrace, Adelaide
South Australia 5000
A CHECKLIST OF HELMINTHS FROM AUSTRALIAN BIRDS
BY PATRICIA M. MAWSON, L. MADELINE ANGEL AND S. J. EDMONDS
Summary
This checklist includes all original published records (to the end of 1983) of the helminths occurring
in Australian birds, as well as undescribed specimens known to the authors to be held in museums
or other institutions.
A CHECKLIST OF HELMINTHS FROM AUSTRALIAN BIRDS
by
PATRICIA M. MAWSON, L. MADELINE ANGEL and 5, |. EDMONDS
Honorary Research Associates, South Australian Museum, North Terrace, Adelaide, South Australia S000
(Manuscript accepted 7 November, 1985)
ABSTRACT
MAWSON, P.M., ANGEL, £.M,, and EDMONDS, S.J. 1986. A
cheeklist of the helminths from Australian birds. Ree. 8, Asi.
Mus, V(15):. 219-325.
This checklist includes all original published records
(ta the end of 1983) of the helminths aceurring in
Australian birds, as well as undescribed specimens
known to the authors to be held in museums or other
institutions,
Taxonomic lists given are (1) bird hosts, numbered
consecutively, with the helminths from each bird noted
below it, with the locality of collection and the
authority for each record; and (2) helminths, with the
bird hosts listed below each helminth species,
There are also three alphabetical lists, one of the bird
families, one of the bird species and one of the
helminth species, all referring by number to the host
in List 1.
The incidence of helminths in birds dissected by uS
and by the late T.H. Johnston is noted and brief
comments made on the birds and on the helminths,
A complete reference list is also given,
CONTENTS
INTRODUCTION ........00...0-.-2---0520 05 219
ACKNOWLEDGMENTS ......-..-. wed othe 219
EXPLANATION OF FORMAT AND
CONTENTS ...0..0ecee ee: 2g. ae 6 220
COMMENTS ON THE BIRD HOSTS .-...-. 22]
COMMENTS ON THE HELMINTHS..,.... 22)
LIST 1. Birds in Taxonomic Order, and
their Parasites. ...,.-.-.--.----.-4.. 223
LIST 2, Helminths in Taxonomic Order,
and their Hosts ...6 2. sey eeu ee eeee 284
LIST 3. Index to Families of Birds......,,..- 308
LIST 4. index to Species of Birds.-.....-...- 306
LIST 5. index 10 Helminths......-....--:.:- 310
REFERENCES. ......-; 000 e cece ese tee eee 320
INTRODUCTION
No complete list of helminths from Australian
birds—not even a complete list of any major group of
helminths—has been published for over forty years.
During. this time a great deal of research has been done
on the taxonomy of helminths from Australian birds,
much of it by the present authors, and it is considered
timely to draw all these records together-
Comprehensive lists published earlier than [923 were
made by T.H. Johnston (1910, 1912b, 1918) and J.B.
Cleland (1922). Partial lists dealing with the parasites
Aueysl, 1986
in restricted geographical areas or with one group of
parasites were published by Bancroft (1889), Cleland
and T.H. Johnston (1910, 1912), TJH, Johnston (1916),
Breinl (1913), Nicoll (1914a,b), and T-H. Johnston and
Deland (1929). In 1939 M_R, Young pablished a list of
helminth parasites of Australia. Since then, restricted
lists have been issued by Mackerras (1962) on filarial
nematodes of Australian vertebrates and by Munday
and Green (1972) on all helminth parasites of
Tasmanian vertebrates.
Without these earlier works the compilation of the
present list would have been a formidable task, The
present authors are indebted to them and to the host
and parasite files started by T.H. Johnston and now
continued as part of the Australian Helminthological
Collection (at present housed in the South Australian
Museum),
Very little serious collecting of helminths from birds
has taken place in Australia. Most known helminths
have been found incidental to bird collection. Museum
ornithologists wha collect birds for taxonomic
purposes have usually no need to keep the carcases
when the skins are removed, so that unless a
helminthologist can arrange to obtain a carcase from
a bird collector, much valuable helminthological
material may be lost. We have been fortunate in having
such liaison with successive ornithologists (HT,
Condon and S.A, Parker) at the South Australian
Museum, and with officers of the (then) Animal
Industry and Agriculture Branch of the Northern
Territory Administration, who undertook extra work
in labelling and transmitting to us the frozen, skinned
torsos of birds which they had collected,
This list of helminths is concerned only with those
from native Australian birds; the parasites of
introduced birds, caged or feral, have not been
included. Nor have birds taken on Macquarie Island
or in the Australian Antarctic Territory been included.
We have found no case of interchange of helminth
parasites between introduced and native birds dissected
by ourselves except when the latter are in captivity.
Synganius trachea has been recorded by others from
Uncaged birds, as noted in the text.
ACKNOWLEDGMENTS
Inforrnation for this checklist was obtained from all
major Museums and collections in Australia and from
the British Museum (Natural History). We are grateful
for all the trouble taken by the staff of these institutions
in sending details of relevant material and, in some
cases, specimens.
220 REC. S. AUST. MLS. 19(15): 219-325
We wish to acknowledge special help from Mr Shane
Parker with ibe nomenclature of birds, Dr Gordon
Gross for advice on the arrangement of the mianuscript,
and to many helminthologists in Australian universities
for help in obtaining specimens and with references,
and identifications.
Our warm thanks go also to colleagues in the
Zoology Department of the University ef Adelaide and
the South Australian Museum, and to Friends in various
places, who have responded with patience and tolerance
to our frequent inquiries about Australian birds and
their helminth inhabitants.
EXPLANATION OF FORMAT AND CONTENTS
This checklist has been compiled from all published
records up to the end of 1983, which are known ta us,
and from unpublished material, lists of which have
been made avaijable to us, held in Museums avd other
institutions, Published work which is merely a
repetition of an earlier record is not included, Some
of the material listed is not fully identified. This applies
particularly to cestodes, as less work has been done in
Australia on these parasites than on other helminths;
and because none of the authors can claim expertise
in this field, All specimens, whether identified or not,
have been included, not only to show the range of
parasitism, but also in the hope that interested
taxonomists may wish to examine the specimens, We
have not ourselves seen all the material listed, and have
examined only that in the Australian Helminthological
Callection, that described in our own papers, and some
of that in the Veterinary School, University of
Melbourne and in the Commonwealth Institute of
Health, University of Sydney.
In this work the information is arranged in the
following, sections:
List 1. Bird species arranged systematically and
numbered consecutively, with the helminths
from each listed in taxonomic order under it;
the Jocality and authority are given for each
record. Synonymy is indicated where necessary.
List 2. Helminths arranged systematically, with bird
hosts under each, and with numerical
references to List 1.
List 3, Families of birds listed, in alphabetic order, with
page numbers (in this paper).
List 4. Bird species listed alphabetically, with nuimbers
referring to List |.
List 5. Helminth species arranged alphabetically in
each of the four helminth groups, with
numerical references to List 1. Synonyms
named in List | are included in alphabetic
sequence.
References.
J.B, Cleland (1922) gave a list of parasites from
Australian birds. He included, as well as a list of earlier
records, a section on birds examined by himself and
the parasites found in them, The helminths in this
section were seldom identified further than to one of
the four helminth groups. The present whereabouts of
many of these specimens is not known, but same have
August, 1986
been described since, and some are in the Australian
Helminthological Collection and are noted as such in
List 1.
Abbreviations used in the text:
Authors whose names appear frequently are referred
to, Where appropriate, by initials, as follows:
LMA L, Madeline Angel
STE Stanley J. Edmonds
SIJ Stephen J. Johnston
THJ TT. Harvey Johnston
PMM_ Patricia M, Mawson
The four major groups of helminths are referred ta
by their initials:
A. Acanthocephala
C Cestoda
N Nematoda
T Trematoda
Museums.and other Institutions which are referred
to as having bird helminths in their collections are
abbreviated as follows:
AHC Australian Helminthological Collection, at
present housed in the South Australian
Museum, Adelaide, South Australia
AM Australian Museum, Sydney, New South
Wales
BM(NH) British Museum (Natural History),
London, England
CiH Commonwealth Institute of
Helminthology, St Albans, Herts, England
CIHUS Commonwealth Institute of Health,
University of Sydney, New South Wales
DWRR_ Division of Wiidlife & Rangelands
Research (C.5.1,R.0.), Canberra, A\C.T,
NMV (Formerly National) Museum of Victoria,
Melbourne, Victoria
OM Queensland Museum, Brisbane,
Queensland
SAM South Australian Museum, Adelaide,
South Australia
UMYVS — University of Melbourne, Victoria, School
of Veterinary Science
UQDP University of Queensland, Brisbane,
Department of Parasitology
WAM ~ Western Australian Museum, Perth,
Western Australia
Localities tram which birds and parasiles are
recorded are given as the State In which they lie, though
more detail is usually available in the references cited-
State names are abbreviated as follows:
ACT Austrahan Capital Territory
EA Eastern Australia
NSW New South Wales, including Lord Howe Island
NT Narthern Territory
Qld Queensland, including tslands of the Great
Barrier Reef
SA South Australia, including Kangaroo J. and
Pearson 1,
Tas Tasmania, including the Bass Strait Islands
(King 1. & Flinders 1.)
Vie Victoria
WA Western Australia
If a host is known ta have been captive (cage, aviary,
etc.) the record is marked ‘cage’.
HEI MINTHS FROM ALISTRALIAN BIRDS au
COMMENTS ON THE BIRD HOSTS
Bird nomenclature, both English and setentific,
and the order in which the names are listed, follow that
of Sehodde et al, (1978), except for certain amendments
made since L978,
The birds in the host-parasite list have been
numbered 1-443, These numbers do not refer to any
previously published list of Australian birds, but are
given to lacihtate cross-reference from other lists in this
work to the first, which contains greater reference
detail,
In the first list, birds which migrate regularly to and
from other countries are noted as ‘migratory’, the term
Here not including those which migrate within
Australia, or those which occur outside Australia but
are not regular migrants. [f a bird occurs only in
Australia ir is noted as ‘endemic’. We have not included
helminths taken from nugratory birds, er from those
which oceur naturally in other countries, if they were
collected outside their Australian range, However, we
have included records from birds af distinctly
Australian origin in captivity in other countries, Any
record from a bird known to have been in captivity,
in Australia or elsewhere, is marked ‘(cage)’. As
mentioned earlier, helminths from birds introduced into
Australia since the advent of the white man have not
been included. It is hoped that this omission may be
rectified later,
The Cattle Egrei, Ardeola this, was not actually
introduced into Australia by man, It is thought to have
followed the introduced Water Buftala, alter this. had
become established in northern Australia. The Cattle
Egret was first noted by ornithologists to be present
in the late 1940s, and has now extended its range to
southern Australia.
The legends (numbers and letters) which appear
directly under the name of some birds im List | tefer
to autopsies undertaken by ourselyes or by Professor
T,H, Johnston, Careful reeards of these dissections
were kept in the Zoology Department, University of
Adelaide, by Johnston from 1922 to his death in 1951,
and since then by Angel and Mawson. [n these legends,
the figures indicate the number of birds
examined/number found with helminths, and the
letters refer to the group of helminth, the number after
cach letter being the number of birds infested wirh thal
group,
Seven hundred and six species of native Australian
birds, representing 82 bird families, are listed by
Schodde ef q/. (1978), Helminths are known to occur
in 384 (54%) of these birds; al least 55 (8%) other
species have been examined but have not yielded
helminths. This means thar about 40% of the
Australian birds are unexplored territory for internal
parasites. Bird families represented in Australia from
Which na helminths have been recorded are (with the
number of Australian species in brackets) Pandionidae
(1); Rostratulidae (1); Phalaropidae (3); Stercorartidae
(5); Opasittidae (1); Nectarinidae (1); Dicaeidae (1):
Atrichornithidae (2); and Sturnidae (1). In List 3 the
number of Australian bird species in each family is
noted afler the family name, tagether with the number
from which helminths have been recorded.
Before the studies of Rowley (1967, 1970) the
distribution and taxonomy of Australian corvids were
poorly Known or understood, Indeed it was not until
these studies were made that Cervus mellori (Liltle
Raven) and ©. /fasmaniensis (Forest Raven) were
recognised as species distinct from C ceronoides
(Australian Raven), Fora substantial number of early
records, the identity of the corvid hosts can be
established by recourse io the original (museum) skin,
or reasonably assumed from the locality involved. [or
other old records, however, where host specimens were
not retained and where the provenance falls within a
zone of overlap befween the now-recognised specics,
the identity af the corvid hast will probably always
remain uncertain. Recards from such uncerigain origins
are marked *? host sp;’.
COMMENTS ON THE HELMINTHS
In general, the helminths from Australian birds
resemble thase from orher parts of the world. There
are very few genera which are Jimited ta Australian
birds, allhough maty species and some sub-species are
so restricted, Some helminth species are found in
related birds in many parts of the world: such overlap
may become more evident with greater knowledge of
the various genera,
Helminths of the gut are rare in free-ranging
psittaciform birds in Australia Of about 160 such birds
dissected by us, cestodes were present in 5, nematodes
(minute larvae) in 2, and trematodes and
acanthocephalans in none (PMM, 1985, p, 199).
However, the tematode Platynosamum prexillicens
and the nematodes Ascaridia columbae aid A,
platyceri have been taken from native birds which have
been held in captivity in Australia or in other countries.
Two possible reasons for this may be that Ihe caged
bird is more vulnerable to infection (and many of the
infestations were very heavy and were said to have
caused the death of the host), or that it is exposed to
the egys or larvae of parasites through proximity to
other (7 introduced) caged birds or 16 domestic stock
such as. pigeons. Ascaridja platyceri has recently been
found in a feral colony of Agupernis roseicollis near
Cowell, SA (Coll,S, Parker, iderit,. PMM, in AHC}.
Birds dissected by ourselves had often been frazen
before we received them, a lreatment particularly
damaging to [at worms. However, rather than
disregard poor specimens we have listed them to record
their presence, and given whal classification js possible
[In this work we have not referred to sub-species,
elther of helminths ar hosts,
Trematodsa
The classification and nomenclature followed here
are for the most part those of Yamaguti (1971), The
main exception is in the Strigeidae, in which we follow
Dubois (1938 et seq.). Entries in the checklist are placed
according to the order im which families are placed by
Yamaguti (1971,p. 475), Families of Digenca from fish
also follow the order used by Yamaguti (1971, p. 19)
(Cysts of these trematodes are often recorded Irom Lhe
alimentary canals of birds)-
Since the number and arrangement of collar spines
is important in the identification of echinostomes, they
have been tecarded for incompletely indentified
specimens of this graup in one of the following ways,
as appropriate:
Unidentified echinostome: [spines (35(5))] indicates
that the total number of spines is 35, including 5
corner spines on each side.
Echinostomatidae: [spines missing].
Echmostomatinae: [spines? (35(5))]: total number 235,
including 5 corner spines on each side,
In Echinechasmus sp.: [spines 20); total number 20,
no camer spines.
lo Patagifer sp.: [spines 2 x (30(4))]: total number 60,
30 on each side including 4 corner spines.
In Patagifer sp.: [spines 2 x 7 (29(4))]: total number
758, including 4 corner spines on each side.
Cestoda
For the most part the names used for cestodes follow
the classification adopted by Yamaguti (1959), Other
more recent works consulted in special cases were Baer
and Bona (1960) and Bona (1975) for cestades of
Ciconiformes, Palmer (1981) for cestodes of the Black
Swan, Cyenus atratus, and Schmidt (1972) for some
cestodes from honeyeaters (Meliphagidae) and parrots.
None of the present authors is well versed in cestode
{axonomy} cestode lists here are the responsibility of
PMM,
li may be nated here that P.A, Maplestone deposited
four lots of cestodes in the Australian Institute of
Tropical Medicine (now the Commonwealth Institute
of Health, University of Sydney} under names which
apparently have never been published. All are labelled
as ‘“Tlew species, Maplestone”. The names are “Shipleva
lobivanellus” and "Monopylidium tobivanelli*) both
from “Venellus miles's “Hymenolepis variabilis". From
Cygnus atratus, and “H. eurysiomei from Eurystomus
orientalis, The explanation which occurs to us is that
Mapiestone examined this material while he was in
Queensland, decided that it belonged to these four new
species, and deposited paratypes in what is now the
CIHUS Collection before returning to England. After
his return, Maplestone published a number of papers,
same with Southwell, an Australian cestades, from the
same hosts and localities but with different scientific
names; ane such species is Monapylidiuni
macracanthum Maplestone and Southwell, 1923, which
may be the unpublished “M7, fabivanelli In one of the
papers. he identified Hymenolepis lanceolata (Bloch,
1782) from Cygnus airatus; the CLHUS material
labelled “A. variabilis Maplestone’ may belang to this
species,
In the Australian Helminthological Collection there
are shdes of the first two of these unpublished species,
presumably given to T.H.Johnstan by Maplestone when
they were both in Queensland, The AHC miaterial
labelled “Shipleya labivenellus Maplestone’ has been
examined by Schmidt (1972) who identified it as /nfule
burking Burt, 1939,
REC. 5. AUST. MUS, 19(15}; 219325
Aupusi, 1986
Nematoda
Nomenclature in this section follows on the whole
that adopted by Anderson, Chabaud and Willmott,
1974-1982, (CIH keys to the nematode parasites of
vertebrates) as far as these keys are completed. Some
points to be noted are:
Anisakis diomedeae (Linstow, 1888) is recorded from
a number of procellariform birds. The validity of this
species has recently beea upheld (Mawson 1983, p.
247), [tis the only species of the genus Anisakis known
from birds, but is commonly found in albatrasses and
petrels along the Australian coast. Linstow described
it, as Ascaris diomedeae, from material collected by
ihe Challenger Expedition in the narthern Pacific
Ocean, from Diomedea brachyura,
Contracaecum spicyligerum (Rudolphi, 1809),
commonly recorded from waterbirds was considered
by Hartwich (1964) ta be a junior synonym of C
microcephaluim (Rudolphi, 1809), and Hartwich
suggested that the specimens. hitherto allotted to
C spiculigerum belong cither lo C. riierocephalum or
ta C rudolph: Hartwich, 1964. However, in this
checklist the name C. spiculfeerumt is retained, pending
further review of the Australian species of
Contracaecum. Hartwich also suggested that
magnipapillatum Johnston and Mawson, 1941, is a
junior synonym of C variegatum (Rudolph, 1809} but
the lwo species appear to us to be quite distinct,
Three different species of Schiéstogendria have been
examined from various birds, but have not been
described. They are referred 10 by numbers; voucher
specimens are in the AHC.
Cyrrea spiralis Mawson, 1968d, is now considered
likely to belong to, or near, Microhadjelia Jogis, 1968,
and jis referred to in this work as ?Microhadjelia
spiralis.
Acanthocephala
The system of classification of acanthocephalans
used in this paper is that of Yamaguti (1963). However,
most of the changes in nomenclature riade since 1963,
especially those of Schmidt (1983) have been adopted,
No new genus of Acanthocephala from Australian
birds has so far been recorded.
The embryonated eggs of an acathocephalan are
shed with the faeces of the host. The first intermediate
hast in the life cycle of the parasite is ihought to be
always an arthropod, which is infected by eating the
eggs, Sometimes a second intermediate host (often
called a paratemic host or “Vhdéte d’attente”) is
necessary. A final or definitive host is infected when
it ingests an infected intermediate host. Consequently
adult acanthocephalans in birds accur mm those which
are partly or wholly carnivorous, This possibly explains.
why no acanthocephalans have so far been collected
from the Emu, Dromaius novaehollandiae, or from
psittaciform birds, which are principally seed eaters,
The life cyeles of Australian avian acanthocephalans
aré poorly known and none have been determined
experimentally. What information about life cycles is
available has been obtained by a comparison of the size
and armature of the introverts of larval and adult
HELMINTHS FROM AUSTRALIAN BIRDS 223
forms. Using such a method T.H. Johnston and
Edmonds (1949) considered that one of the larval stages
of Porrorchis (= Gordiorhynchus) hyvlae (Johnston,
1912), the adult stages of which occur in the birds
Podareus strigoides and Centropus phasianinus, is
found as a cyst in the mesenteries and muscles of a
number of frogs. The frog is infected, it is assumed,
by eating an infected crustacean or insect, Some light
has been thrown on the life cycles of two other ayian
Acanthocephala, The adult form of Polymorphus
biziurae T.H. Johnston and Edmonds (1948) occurs in
the small intestine of the Musk Duck, Biziura lobata.
T.H. Johnston and Edmonds (1948) considered that the
freshwater yabbie, Cherax destructor, is an intermediate
host of the parasite One of the commonest
acanthocephalans known to dissectors of birds in
Australia is a small cyst, 2-3 mm long and shaped like
a “grain of rice”, which occurs in the subcutaneous
tissues of the neck and thorax of a large number of
passerine birds, It was called Eehinorhynchus
pomatosiomi by T.H,Johnston and Cleland (1912).
Schmidt (1983) has found that the introvert of E.
pomatostomi matches that of an acanthocephalan
found in the intestine of dingos and feral cats in
Australia. From this information he has concluded that
the cyst known as £. pomatostomi is an early stage of
the acanthocephalan found in dingos and feral cats,
Since the adult form is an Oncicola and not an
Echinorhynchus, the parasite (including the cyst) is now
Oncicola pomatostomi (T.H. Johnston and Cleland,
1912). The carnivores are infected by ingesting the cysts
in their bird prey. The first host is probably a
scavenging insect, though at present there is no evidence
for this,
LIST 1, BIRDS IN TAXONOMIC ORDER, AND
THEIR. PARASITES
Explanation of the symbols used in this section are
given in “Comments on bird hosts” on p. 221,
Family DROMATIDAE
1, Dromaius noyachollandiae (Latham)
Emu (endemic)
1 3c
T. Philophthalmus sp,
NT: AHC
C. Cotugnia collini Fubrmann, 1909
EA: Fuhrmann, 1909, p.116
Colugnia sp.
NSW: AHC
Raillietina australis (Krabbe, 1869)
NSW: AHC
syn. Davainea australis (Krabbe, 1869)
NSW: THJ, 1909c, p.xxix
NSW, WA: THS, 1910, p.86
Qld; THS, 1916, p.45
syn. Taenia australis Krabbe, 1869
Copenhagen (cage): Krabbe, 1869, p.343
Raillietina sp.
NSW, SA: AHC
Vic: AHC, UMVS
Unidentified
Qld, NSW, Vic, SA, WA; AHC
N. Dromaeostrongylus bicuspis Lubimoy, 1933
Moscow (cage): Lubimov, 1933, p.173
Vic: Durette-Desset, 1979, p.1015
NSW, Vie: AHC
Trichostrongvius tenuis (Mehlis, 1846, in Creplin,
1846)
Aust: Arundel, 1982, pers, comm.
Family CASUARIIDAE
2, Casuarius casuarins (Linneé)
Southern Cassowary
C, Unidentified
Qld: Macgillivray, 1917, p.80
Family PODICIPEDIDAE
3. Podiceps cristatus (Linné)
Great Crested Grebe
5/5: Ti C4 N4
T. Schistosomatidae, unidentified
SA: AHC
Pelasiger australis TH, Johnston & Angel, 1941b
SA: AHC
Diplostomum — podicipinum
Niewiadomska, 1960
SA: Dubois & LMA, 1972, p,208
Schwarizitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972, p.205 (cysts)
C, Unidentified
SA: AHC
WA: WAM
Krefft (1873, pp.216,217) recorded Taenia paradoxa and
T. novaehollandiae from the “Little Grebe (Podiceps
australis)”. P. australis is wow placed as a synonym of
the Crested Grebe, (P. cristafus), and it is considered
that Krefft’s records refer to Tachybaptus
novaehollandiae, which was then called the Little Grebe
(q.v.).
N. Capillaria sp.
SA: THJ & PMM, 1949, p.64
Contracaecum podicipitis T.H, Johnston &
Mawson, 1949
SA: THJ & PMM, 1949, p.67
NSW, Vie: AHC
Tetrameres gubunovi Shigin, 1957
SA: PMM, 1979, p.180
Syneuaria sp.
ACT: PMM, 1982, p.21
Kozicka &
4. Poliocephalus poliocephalus
(Jardine & Selby)
Hoary-headed Grebe (endemic)
5/4; T3 C3 N3
T. Paramonostamum caeci Smith & Hickman, 1983b
Tas: Smith & Hickman, 1983b, p.86
Cyclocoelum jaenschi T,H, Johnston & Simpson,
1940b
C;
REC. 5. AUST, MUS. 19(15): 219-325
SA: THJ & Simpson, 1940b, p.273
Echinostoma sp. [spines (37(4))]
NSW: AHC
Patagifer sp. [spines 2x (26-27(3-4))]
Qld: AHC
Petasiger australis TH. Johnston & Angel, 1941b
SA: THJ & LMA, 1941b, p.285
Echinostomatidae, unidentified
NSW: Bradley, 1926, p.575; 1927, p.675
Psilochasmus oxyurus (Creplin, 1825)
Tas: S.J. Smith, 1981, p.181
Psilostomum sp, A, S.J. Smith, 1981
Tas: S.J. Smith, 1981, p.201
Psilostomum sp. B, S.J, Smith, 1981
Tas: S.J. Smith, 1981, p.214
Atriophallophorus coxiellae $.J. Smith, 1974
Tas: S.J. Smith, 1981, p.151
Levinseniella tasmaniue (S.J. Smith, 1974)
Tas: S.J. Smith, 1981, p.107
Maritrema calvertense S.J. Smith, 1974
Tas: S.J. Smith, 1981, p.64
Pachytrema sp.
Tas: S.J. Smith, pers. comm., 1979
Schwartzitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972, p.205 (cysts)
? Cephalogonimus sp.
SA: AHC
Dioecocestus sp.
Qld: BM(NH)
Gyrocoelia sp.
Aust: BM(NH)
Unidentified
NSW: Bradley, 1926, p.572, 1927, p.675
NSW: Cleland, 1922, p,105
Qld, NSW, SA: AHC
. Capillaria sp.
SA: AHC
Contracaecum praestriatum Moénnig, 1923
Qld: CIHUS, BM(NH)
Streptocara recta (Linstow, 1879)
SA: TH] & PMM, 194le, p.16
Unidentified
? loc: Bradley, 1927, p.675
5. Tachybaptus novaehollandiae
(Stephens)
Australian Grebe
13/10 : T8 CIO N8
Cyclocoelum jaenschi T.H. Johnston & Simpson,
1940b
SA: THJ & Simpson, 1940b, p.273
Petasiger australis T.H. Johnston & Angel, 1941b
SA: TH] & LMA, 1941b, p.285
SA: AHC
Echinostomatidae, very immature, [spines ? (37)]
SA: discarded
Maritrema odcystum (Lebour, 1907)
Qld: Deblock & Pearson, 1968b, p.459
Microphallidae, unidentified
SA: (believed lost)
Schwartzitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972, p.205 (cysts)
CG
Augusil, 1986
Dilepididae, unidentified
NSW: DWRR
Dioecocestus novaehollandiae (Krefft, 1873)
syn. Jaenia novaehollandiae Krefft, 1873
NSW: Krefft, 1873, p.216
syn. Taenia paradoxa Krefft, 1873
NSW: Krefft, 1873, p.217
Unidentified
NSW: AM
Qld, NSW, SA: AHC
. Contracaecum podicipitis T.H. Johnston &
Mawson, 1949
SA: TH] & PMM, 1949, p.67
Tetrameres gubanoyvi Shigin, 1957
NSW: PMM, 1979, p.180
Skrjabinoclava horrida (Rudolphi, 1809)
Qld: AHC
Streptocara recta (Linstow, 1879)
SA: THJ & PMM, 194le, p.260; PMM, 1955,
p.6
SA: AHC
Streptocara sp.
NSW: DWRR
Syncuaria sp.
NSW: DWRR
6. Podicipedidae
Unidentified grebe
. Unidentified
Qld: AHC
. Capillaria sp.
Qld: AHC
Family SPHENISCIDAE
7. Aptenodytes patagonicus Miller
King Penguin
. Tetrabothriidae, unidentified
Tas: AHC
8. Eudypies chrysocome (Forster)
Rockhopper Penguin
. Anisakidae
Tas: AHC (larva)
9. Eudyptes pachyrhynchus Gray
Fiordland Penguin
. Contracaecum sp.
Tas: AHC
10. Eudyptes schlegeli Finsch
Royal Penguin
. Unidentified
Tas: AHC
11. Eudyptula minor (Forster)
Little Penguin
4/4: T2 C4 N4 A2
HELMINTHS FROM AUSTRALIAN BIRDS
. Mawsonoirema eudyptulae Angel, 1973
SA: LMA, 1973, p.857
Vic: UMVS
Tas: AHC
Renicola sp.
Tas: AHC
Galactosomum angelae Pearson, 1973
SA: Pearson, 1973, p.361
. Tetrabothrius lutzi Parona, 1901
Tas: Prudhoe, 1969, p.185
Tetrabothrius sp.
NSW, Vic, SA: AHC
Tetrabothriidae, unidentified
Vic: AHC
Unidentified
SA: Cleland, 1922, p.105
NSW, Vic, Tas, SA: AHC
. Anisakis sp.
NSW, SA: TH] & PMM, 1[942a, p.94
Contracaecum spiculigerum (Rudolphi, 1809)
Vic, Tas, SA: AHC
Contracaecum eudyptulae T.H.
Mawson, 1942a
NSW, Vic, Tas, SA, WA: THJ & PMM, 1942a,
p.93
syn. unidentified nematode
SA: Cleland, 1922, p.107
Filarioidea, unidentified
SA: AHC (from heart)
Unidentified
Vic: UMVS
. Corynosoma sp.
SA: AHC
Johnston &
Family DIOMEDEIDAE
12, Diomedea exulans Linné
Wandering Albatross (migratory)
1/1:CN
’, Tetrabothrius diomedeae (Fuhrmann in Shipley,
1900)
syn. Prosthecocotyle diomedeae Fuhrmann in
Shipley, 1900
‘Pacific Ocean, NE of
Fuhrmann, in Shipley, 1900, p.557
Tetrabothrius sp.
NSW; THJ, 1912b, p.106
Unidentified
NSW, SA: AHC
. Anisakis diomedeae (Linstow, 1888)
NSW, Tas: TH) & PMM, 1942d, p.6
syn. Contracaecum diomedeae (Linstow, 1888)
Tas: THJ, 1938, p.15
Contracaecum sp.
SA: AHC (immature)
Seuratia shipleyi (Stossich, 1900)
NSW, SA: AHC
Stegophorus diomedeae (T.H. Johnston & Mawson,
1942d)
syn. Paryseria diomedeae T.H. Johnston &
Mawson, 1942d
NSW: THI & PMM,, 1942d, p.69
Austr.’:
N
to
i)
a
Stegophorus sp.
SA: AHC
13. Diomedea melanophrys Temminck
Black-browed Albatross
1/1:CN
. Tetrabothrius sp.
NSW: THJ, 1912b, p.106
Tas: AHC
Cyclophyllidea, unidentified
Tas: Munday & Green, 1972, p.4
Unidentified
NSW, SA: AHC
Capillaria convolutor Fourment, 1885
SA: THJ & PMM, 1945a, p.15]
Anisakis diomedeae (Linstow, 1888)
NSW, SA: THJ & PMM, 1942d, p.67
syn. Stomachus sp.
Tas: Munday & Green, 1972, p.9
Contracaecum pelagicum T.H. Johnston &
Mawson, 1942d
NSW, SA: THJ & PMM, 1942d, p.67
Anisakidae, unidentified
Tas: Munday & Green, 1972, p.9
Seuratia shipleyi (Stossich, 1900)
SA: THJ & PMM, 1942d, p.69
Stegophorus diomedeae (T.H. Johnston & Mawson,
1942d)
syn. Paryseria diomedeae T.H. Johnston &
Mawson, 1942d
SA: THI & PMM, 1942d, p.67
14. Diomedea bulleri Rothschild
Buller’s Albatross
. Seuratia shipleyi (Stossich, 1900)
NSW: AHC
15. Diomedea chrysostoma Forster
Grey-headed Albatross
4/4: Cl N4
. Unidentified
SA; AHC
. Anisakis diomedeae (Linstow, 1888)
SA: THJ & PMM, 1942d, p.67
Tas: AHC
syn, Sfomachus sp.
Tas: Munday & Green, 1972, p.9
Seuratia shipleyi (Stossich, 1900)
SA: THJ & PMM, 1952, p.33
Stegophorus diomedeae (T.H. Johnston & Mawson,
1942d)
syn. Paryseria diomedeae T.H. Johnston &
Mawson, 1942d
SA: THJ & PMM, 1942d, p.69; 1952, p.33
Stegophorus macronectes (T.H. Johnston &
Mawson, 1942d)
syn. Paryseria macronectes T.H. Johnston &
Mawson, 1942d
SA: TH] & PMM, 1942d, p.70
226 REC. S, AUST. MUS. 19(15): 219-325 August, 1986
Diomedenema diomedeae T.H. Johnston & C. Tetrabothrius sp.
Mawson, 1952 SA: AHC
SA: THJ & PMM, 1952, p.32 Unidentified
Qld, SA: AHC
16. Diomedea chlororhynchos Gmelin N. Capillaria conyolutor (Fourment, 1885)
Yellow-nosed Albatross SA: AHC
3/3: C3.N3 Anisakis diomedeae (Linstow, 1888)
SA: THJ & PMM, 1942d, p.67
C. Tetrabothrius sp. Phocascaris sp.
SA: AHC SA: THJ & PMM, 1942d, p.68
Unidentified Seuratia shipleyi (Stossich, 1900)
SA: AHC SA: TH] & PMM, 1942d, p.69
N. Anisakis diomedeae (Linstow, 1888) SA: AHC
SA: TH] & PMM, 1942d, p.67 Stegophorus macronectes (T.H. Johnston &
Contracaecum pelagicum T.H. Johnston & Mawson, 1942d)
Mawson, 1942d syn. Paryseria macronectes T.H. Johnston &
NSW: THJ & PMM, 1942d, p.67 Mawson, 1942d
Tetrameres certa (Leidy 1886) SA: THI & PMM, 1942d, p.70
syn. Tetrameres diomedeae T.H. Johnston & A. Unidentified
Mawson, 1942d SA: not kept
SA: THJ & PMM, 1942d, p.67
Seuratia shipleyi (Stossich, 1900) 20. Fulmarus glacialoides (Smith)
SA: THJ & PMM, 1942d, p.67 Southern Fulmar —
17. Diomedea cauta Gould C. Unidentified
Shy Albatross SA: AHC
1/1:CN N. Seuratia shipleyi (Stossich, 1900)
SA: AHC
C. Unidentified
SA: not kept
Tas: AHC 21. Daption capense (Linné)
N. Anisakis diomedeae (Linstow, 1888) ee ieee
SA: THJ & PMM, 1942d, p.67 :
ae oNS C. Unidentified
Tas: AHC SA: AHC
Contracaecum magnicollare T.H. Johnston & yt eer TY '
Mawson, 1942d N. Anisakis diomedeae (Linstow, 1888)
SA: THJ & PMM, 1942d, p.67 SA: THJ & PMM, 1942d, p.67
Tetrameres certa (Leidy, 1886) Seuratia shipleyi (Stossich, 1900)
syn. Tetrameres diomedeae T.H. Johnston & SA: THJ & PMM, 1942d, p.69
Mawson, 1942d Stegophorus pachyptilae (T.H. Johnston &
Tas: AHC Mawson, 1942d)
Stegophorus diomedeae T.H. Johnston & Mawson, . SA: AHC .
1942d Spirurida, unidentified
Tas: AHC Tas: Munday & Green, 1972, p.10
Diomedenema diomedeae T.H. Johnson &
Mawson, 1952 22. Pterodroma macroptera (Smith)
Tas: AHC Great-winged Petrel
18. Phoebetria palpebrata (Forster) N. Anisakis sp.
Light-mantled Sooty Albatross Tas: AHC
syn. Stomachus sp.
C. Tetrabothrius sp. Tas: Munday & Green, 1972, p.9
SA: AHC Seuratia shipleyi (Stossich, 1900)
N. Contracaecum sp. Tas: Munday & Green, 1972, p.l1
Vic: AHC Tas: AHC
Seuratia shipleyi (Stossich, 1900) Spirurida, unidentified
SA: AHC Tas: Munday & Green, 1972, p.9
19. Macronectes giganteus (Gmelin) 23. Pterodroma lessonii (Garnot)
Southern Giant Petrel White-headed Petrel
10/9: C2 N9 Al 3/3: Cl N3
HELMINTHS FROM AUSTRALIAN BIRDS 227
C. Unidentified 28. Pachyptila belcheri (Mathews)
SA: AHC Slender-billed Prion
N. Anisakis sp. 2/2:N
SA: TH] & PMM, 1942d, p.67 A
Seuratia shipleyi (Stossich, 1900) N, Anisakis sp.
SA: AHC SA: AHC (immature)
Stegophorus macronectes (T.H. Johnston & Seuratia shipleyi (Stossich, 1900)
Mawson, 1942d) SA: AHC
SA: AHC
29. Pachyptila turtur (Kuhl)
24, Pterodroma brevirostris (Lesson) Fairy Prion
Kerguelen Petrel 4/4: C3 N4
1/1: N
C. Unidentified
C. Tetrabothrius sp. SA: AHC
BA: AHC ; N. Skrjabinoclava sp.
N. Seuratia shipleyi (Stossich, 1900) Tas: AHC
SA; AHC Seuratia shipleyi (Stossich, 1900)
Stegophorus macronectes (T.H. Johnston & SA: AHC
Mawson, 1942d) Stegophorus pachyptilae (T.H. Johnston &
SA: AHC Mawson, 1942d)
25, Pachyplila vittata (Forster) ae
5. Pachyptila vittata (Forster f
Broad-billed Prion ms yer 4 fais SP.
6/5 ; C2 NS :
C. Tetrabothrius sp. 30. Procellaria cinerea Gmelin
SA: AHC Grey Petrel
Unidentified ;
SA: AHC C. Unidentified
N. Skrjabinoclava sp. NSW: AHC
Tas: AHC
Seuratia shipleyi (Stossich, 1900) 31. Puffinus pacificus (Gmelin)
SA: THJ & PMM, 1942d, p.69 Wedge-tailed Shearwater
Stegophorus pachyptilae (T.H. Johnston & 2/1:.N
Mawson, 1942d)
syn. Paryseria pachyptilae T.H. Johnston & T, Galactosomum renincolum Pearson, 1973
Mawson, 1942d Qld: Pearson, 1973, p.409
SA: THJ & PMM, 1942d, p.70 C. Unidentified
SA: AHC NSW: Cleland, 1922, p.105
26. Pachyptila salvini (Mathews) N. wa A (Stossich, 1900)
Lesser Broad-billed Prion 1
5/1 :N
32. Puffinus griseus (Gmelin)
N. Seuratia shipleyi (Stossich, 1900) Sooty Shearwater
SA: AHC
Stegophorus pachyptilae (TH. Johnston & C. Tetrabothrius sp.
Mawson, 1942d) NSW, Vic: AHC
SA: AHC Unidentified
NSW: AHC
27. Pachyptila desolata (Gmelin) N. Contracaecum magnicollare T.H. Johnston &
Antarctic Prion Mawson, 1941c
11/4:N Vic: AHC
Stegophorus pachyptilae (T.H. Johnston &
N. Anisakis sp., ? diomedeae (Linstow, 1888) Mawson, 1942d)
SA: THJ & PMM, 1942c, p.184 syn. Paryseria pachyptilae T.H. Johnston &
Stegophorus pachyptilae (T.H. Johnston & Mawson, 1942d
Mawson, 1942d) SA: TH] & PMM, 1942d, p.70
syn. Paryseria pachyptilae T.H. Johnston &
Mawson, 1942d 33. Puffinus tenuirostris (Temminck)
SA: THJ & PMM, 1942c, p.184 Short-tailed Shearwater (migratory)
SA: AHC 12/8 : C3 N7
T. Renicola sp.
NSW: AHC
Cryptocotyle sp.
Tas: AHC
C. Tetrabothriidae, unidentified
Vic: AHC
Unidentified
Tas: Cleland, 1922, p.l05 (AHC)
Tas: Munday & Green, 1972, p.4
EA, NSW, SA: AHC
N. Contracaecum magnicollare T.H. Johnston &
Mawson, 1941c
Vic: AHC
Seuratia shipleyi (Stossich, 1900)
Vic, Tas, SA: AHC
Bass Str: BM(NH)
Stegophorus pachyptilae (T.H.
Mawson, 1942d)
Tas, SA: AHC
Stegophorus stellaepolaris (Parona, 1901)
NSW, Bass Str: BM(NH)
Tas: AHC
Johnston &
34. Puftinus gavia (Forster)
Fluttering Shearwater
1/1:CN
T. Renicola sp.
Tas: AHC
C. Unidentified
SA: AHC
N. Unidentified
SA: not kept (pieces only)
Family OCEANITIDAE
35, Pelagodroma marina (Latham)
White-faced Storm Petrel
6/3: N
N. Seuratia shipleyi (Stossich, 1900)
Tas: AHC
syn. Seuratia marina T.H. Johnston & Mawson,
194le
Tas: THJ & PMM, 194le, p.259; TH) & PMM,
1944, p.62
SA: THJ & PMM, 1942d, p.69
syn. “nematode”
SA: Cleland, 1922, p.107 (AHC)
Family PELECANOIDIDAE
36. Pelecanoides urinatrix (Gmelin)
Common Diving Petrel
2/2:N
N. Spirurida, unidentified
Tas: AHC (larva)
REC. 8. AUST. MUS. 19(15): 219-325
August, 1986
Family PELECANIDAE
37. Pelecanus conspicillatus Temminck
Australian Pelican
12/12 : T12 C12 N12 Al
T. Ornithobilharzia sp.
SA: AHC
Dendritobilharzia sp.
SA: AHC
Schistosomatinae, unidentified
Qld, SA: AHC
Echinochasmus pelecani T.H. Johnston & Simpson,
1944
SA: THJ & Simpson, 1944, p.113
Qld: AHC
Echinostomatidae [spines missing]
SA: AHC
Renicola sp.
Qld: Pearson, 1979 (? 2 spp.)
Qid: AHC
Ascocotyle sp.
Qld: AHC
Haplorchis sprenti Pearson, 1964
Qld: Pearson, 1964, p. 639
syn. Haplorchis sp.
Qld: Pearson, 1960, p.93
Haplorchis paravanissimus Pearson & Ow-Yang,
1982
SA: Pearson & Ow-Yang, 1982, p.50
Haplorchis vanissimus Africa, 1938
SA: Pearson & Ow-Yang, 1982, p.50
Procerovum varium Onji & Nishio, 1916
Qld: Pearson, 1964, p.653
syn. Procerovum sp.
Qld: Pearson, 1960, p.93
Stictodora caballeroi Martin, 1955
Qld: Pearson, 1960, p.93
Clinostomum sp.
Qld: AHC
Mesostephanus haliasturis Tabangui & Masilungan,
194]
SA: Dubois & LMA, 1972, p.214
syn. M. minor Dubois & Pearson, 1965
Qld: Dubois & Pearson, 1967, p.202
Bolbophorus confusus (Krause, 1914)
Qld: Dubois & Pearson, 1965, p.95
SA: Dubois & LMA, 1972, p.205
Bolbophorus sp.
SA: THJ & LMA, 1942a, p.59
Diplostomum paryvulum Dubois & Angel, 1972
SA: Dubois & LMA, 1972, p.206
Diplostomum sp.
Qld: AHC
Posthodiplostomum australe Dubois, 1937a
SA: Dubois & LMA, 1972, p.212
Schwartzitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972, p.205 (cysts)
C, Hymenolepis murrayensis T.H. Johnston & Clark,
1948a*
SA: THJ & Clark, 1948a, p.77
“omitted by Yamaguti, 1959
HELMINTHS FROM AUSTRALIAN BIRDS
Hymenolepis jaenschi T.H. Johnston & Clark,
1948a*
SA: THJ & Clark, 1948a, p.79
Qld: AHC
Hymenolepis ellisi TH. Johnston & Clark, 1948af
SA: THJ & Clark, 1948a, p.81
Hymenolepis sp.
SA: AHC
Unidentified
Qld, NSW, Tas, SA, WA: AHC
N. Eustrongylides sp.
Qld: CIHUS (posterior end o, in gular pouch)
Capillaria jaenschi T.H. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p. 245
Contracaecum clelandi T.H. Johnston & Mawson,
1941c
WA: THJ & PMM, 194lc, p.113
Tas: AHC
Contracaecum micropapillatum (Stossich, 1890)
Qld: CIHUS
syn. Contracaecum bancrofti T.H. Johnston &
Mawson, 1941c
Qld, NSW, Vic, SA: THJ & PMM, 1941c, p,113
Contracaecum spiculigerum (Rudolphi, 1809)
Vic: AHC
syn. Ascaris spiculigerum Rudolphi, 1809
NSW: THJ, 1912a, p.74
Tetrameres greeni Mawson, 1979
Qld: PMM, 1979, p.180
syn. Tetrameres pelecani T.H. Johnston &
Mawson, 1942e, sensu T.H. Johnston &
Mawson, 1942¢
SA: THJ & PMM, 1942¢, p.185
Microtetrameres pelecani (T.H. Johnston &
Mawson, 1942e)
Qld: PMM, 1979, p.182
syn. Tetrameres pelecani T.H. Johnston &
Mawson, 1942e nec 1942c
SA: TH] & PMM, 1942e, p.72
Cosmocephalus jaenschi T.H. Johnston & Mawson,
1941le
SA: THJ & PMM, 1942e, p.185
Synhimantus sirry Khalil, 1931
Qld, Vic: PMM, 1982, p.23
syn. Dispharynx pelecani T.H. Johnston &
Mawson, 1942c
SA: THJ & PMM, 1942c, p.185
Filarioidea, unidentified
Qld: AHC (from eye)
A. Polymorphus biziurae T.H. Johnston & Edmonds,
1948
SA: AHC
Family SULIDAE
38. Sula serrater (Gray)
Australasian Gannet
6/4: T1 Cl N4
* listed by Yamaguti, 1959 to
Hymenolepididae
in appendix
ian
i?)
Z
Z
Z
. Galactosomum angelae Pearson, 1973
SA: Pearson, 1973, p.361
. Tetrabothriidae, unidentified
SA: AHC
. Contracaecum magnicollare TH. Johnston &
Mawson, 194lc
Tas: AHC
Contracaecum sp.
SA: AHC
39. Sula dactylatra Lesson
Masked Booby
. Unidentified
NSW: AHC
40, Sula sula (Linneé)
Red-footed Booby
1/1 iN
. Unidentified
Qld: AHC
41, Sula leucogaster (Boddaert)
Brown Booby
2/1: N
. Seuratia shipleyi (Stossich, 1900)
Qld, SA: AHC
Family ANHINGIDAE
42. Anhinga melanogaster Pennant
Darter
Maritrema oocystum (Lebour, 1907)
Qld: Deblock & Pearson, 1968b, p.459
Clinostomum australiense S.J. Johnston, 1917
Qld: SJJ, 1917, p.230
Qld: THJ, 1942b, p187
Mesostephanus haliasturis Tubangui & Masilungan,
194]
syn, M. minor Dubois & Pearson, 1965
Qld: Dubois & Pearson, 1967, p.202
Diplostomum auriculosum Dubois & Pearson, 1967
Qld: Dubois & Pearson, 1967, p.193
Schwarizitrema novaehollandiae Dubois &
Pearson, 1967
Qld: Dubois & Pearson, 1967, p.187
. Eustrongylides plotinus T.H. Johnston & Mawson,
194le
Qld: THJ & PMM, 1!94le, p.256
Contracaecum rodhaini (Gedoelst, 1916)
Qld: CIHUS, BM(NH)
Contracaecum sinulabiatum T.H. Johnston &
Mawson, 1941c
Qld: THJ & PMM, 194lc, p.114
syn. Ascaris spiculigerum Rudolphi, 1809
Qld: THJ, 1912a, p.74
Contracaecum tricuspe (Gedoels{, 1916)
Qld: TH] & PMM, 194Ic, p14
Qld: CIHUS, BM(NH)
SA: AHC
REC. S. AUST, MUS, 19(15): 219-325
syn. Ascaris sp. Krefft, 1873, p.213
Qld: Krefft, 1873, p.213
Synhimantus sp.
syn, Dispharynx sp.
Qld: THJ & PMM, 194le, p.257
Microfilaria sp.
Qld: THJ, 1912a, p.78
Family PHALACROCORACIDAE
43. Leucocarbo fuscescens (Vieillot)
Black-faced Shag (endemic)
8/8 : T4 C4 NB A3
. Echinoparyphium phalacrocoracis Yamaguti,
1939a
SA: AHC
Paryphostomum radiatum (Dujardin, 1845)
syn. PB tenuicolle (S.J. Johnston, 1917)
SA: THJ & LMA, 1942b, p.120
Petasiger exaeretus Dietz, 1909
SA: discarded
Renicola sp.
SA: AHC
Galactosomum sinuilacte Pearson, 1973
SA: Pearson, 1973, p,415
Galactosomum sp.
SA: AHC (immature)
Haplorchis vanissimus Africa, 1938
SA: AHC
Stictodora diplacantha T.H. Johnston, 1942a
SA: AHC
Hysteromorpha triloba (Rudolphi, 1819)
SA: THJ, 1942a, p.238
. Paradilepis scolecina (Rudolphi, 1819)
Tas: Prudhoe, 1969, p.190
Paradilepis sp.
Qld: CIHUS
SA: AHC
Unidentified
SA: AHC
. Eustrongylides phalacrocoracis T.H. Johnston &
Mawson, 194le
SA: THJ & PMM, 1942c, p.186
Tas: THJ & PMM, 1945a, p.149
Capillaria jaenschi T.H. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945a, p.151
Capillaria sp.
Tas: THJ & PMM, 1945a, p.151
Contracaecum spiculigerum (Rudolphi, 1809)
SA: THJS & PMM, 1941c, p.lI1
Tas, SA: AHC
Qld: CIHUS
Cosmocephalus jaenschi T.H. Johnston & Mawson,
1942c
Tas: THJ & PMM, 194Sa, p.143
A. Corynosoma clavatum Gass, 1940
SA: AHC
44, Phalacrocorax carbo Linné
Great Cormorant
7/7: T5 C5 N7
August, 1986
T. &chinoparyphium phalacrocoracis Yamaguti,
1939a
SA: THJ, 1942a, p.238
Paryphostomum radiutum (Dujardin, 1845)
syn. Paryphostomum tenuicolle (S.J. Johnston,
1917)
SA; THS & LMA, 1942b, p.120
Petasiger exaeretus Dietz, 1909
NSW, SA: THJ, 1942a, p.236
Qid: AHC
Haplorchis vanissimus Africa, 1938
SA: AHC (cysts)
Hysteromorpha triloba (Rudolphi, 1819)
NSW, SA: THJ, 1942a, p,238
Schwartzitrema pandubj (Pande, 1939)
SA: Dubois & LMA, 1972, p.205
. Paradilepis scolecina (Rudolphi, 1819)
SA: Clark, 1957, p.124
Unidentified
NSW, SA: AHC
. Eustrongylides phalacrocoracis T.H. Johnston &
Mawson, 194le
SA; TJH & PMM, 194le, p.255
Capillaria jaenschi TH. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p.245
Contracaecum spiculigerum (Rudolphi, 1809)
Qld, NSW, SA, WA: THJ & PMM, 194lc, p.111
Tas: Munday & Green, 1972, p.9
Tas: AHC
syn. Ascaris sp,
NSW; THJ, 1912b, p.108
Contracaecum sinulabiatum T.H. Johnston &
Mawson, 1941c
Qld: THJ & PMM, 194 lc, p.113
Procamallanus murrayensis TV.H. Johnston &
Mawson, 1940b
SA: TH] & PMM, 1944, p.64
Cosmocephalus jaenschi TH. Johnston & Mawson,
194 le
SA: THJ & PMM, 194le, p.259
Echinuria squamata (Linstow, 1883)
SA: THJ & PMM, 194te, p.257
SA: AHC
Synhimantus sp.
NSW: AHC
Microfilaria sp.
Qld: Mackerras, 1962, p.436
Unidentified
NSW: AM
45, Phalacrocorax yarius (Gmelin)
Pied Cormorant
$/5: TS Cl N4 A3
. Renicola sp.
SA: AHC
Galactosomum sinuilacite Pearson, 1973
Qld, SA: Pearson, 1973, p.415
Stictodora diplacantha T.H, Johnston, 1942a
SA: THJ, 1942a, p.239
SA: AHC
HELMINTHS FROM AUSTRALIAN BIRDS al
Cardiocephaloides ovicorpus Dubois & Angel, 1972
SA: Dubois & LMA, 1972, p.204
Strigeidae, unidentified
SA: AHC (larval)
Fellodistomidae, unidentified
syn. Steringophoridae, unidentified
WA: Goss, 1940, p.7
. Dilepis maxima Goss, 1940
WA; Goss, 1940, p.8
Trypanorhyncha, unidentified
SA: AHC (eysts)
Unidentified
SA: AHC
. Capillaria jaenschi T.H. Johnston & Mawson,
19456
SA: THJ & PMM, 1945b, p.245
Contracaecum spiculigerum (Rudolphi, 1809)
WA: THJ & PMM, 1941c, p11
SA: AHC
. Corynosoma clayatum Goss, 1940
WA: Goss, 1940, p,12
SA: AHC
46. Phalacrocorax sulcirostris (Brandt)
Little Black Cormorant
8/8 : T8 C6 N6 AS
. Echinochasmius sp. [spines (20)]
Qld: AHC
Echinoparyphium phalacrocoracis Yamaguti, 1939a
SA: THJ, 1942a, p.238
Paryphostomum radiatum (Dujardin, 1845)
Qld, Vic, SA, WA: THJ, 1942a, p.236
Qld: AHC
syn. P. tenuicolle (S.J. Johnston, 1917)
SA: THJ & LMA, 1942b, p.120
syn, P. phalacracoracis Goss, 1940
WA: Goss, 1940, p.2
Petasiger exaeretus Dietz, 1909
SA: THJ, 1942a, p.236
Qld: AHC
Echinostomatinae [spines missing]
SA, NT: AHC
Haplorchis sprenti Pearson, 1964
Qld: Pearson, 1964, p.639
Haplorchis paravanissimus Pearson & Ow-Yang,
1982
Qld: Pearson & Ow-Yang, 1982, p.50
syn. H. vanissimus of Pearson, 1964 (in part) not
Africa, 1938
Qld: Pearson, 1964, p.639
Haplorchis vanissimus Africa, 1938
Qld: Pearson, 1964, p.633; Pearson & Ow-Yang,
1982, p49
Stictodora diplacantha T,H, Johnston, 1942a
SA: AJC
Hysteromorpha triloba (Rudolphi, 1819)
NSW, SA: THJ, 1942a, p.238
SA: AHC
syn. Diplostamum granulosum Goss, 1940
WA: Goss, 1940, p.6
Posthodiplostomum australe Dubois, 1937a
SA: Dubois & LMA, 1972, p.212
QD: AHC
Diplostomidae, unidentified
Qld; AHC (immature)
Schwartzitrema pandubi (Pande, 1939)
Qld: Dubois & Pearson, 1965, p.89
SA: Dubois & LMA, 1972, p.205
Unidentified
SA: AHC
. Paradilepis minima (Goss, 1940)
SA: Clark, 1957, p.126
syn. Dilepis minima Goss, 1940
WA: Gass, 1940, p.10
Unidentified
Qld, Vic, SA: AHC
. Eustrongylides sp.
NT; AHC
Capillaria jaenschi T.H. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p.245
WA: BM(NH)
Contracaecum spiculigerum (Rudolphi, 1809)
Qld, SA: THJ & PMM, 194lc, p11}
SA: AHC
syn. Ascaris spiculigerum Rudolphi, 1809
Qld: THJ, 1912a, p.64
Streptocara recta (Linstow, 1879)
SA: AHC
Desmidocercella sp.
NT: AHC
Microfilaria sp.
Qld: THJ, 1912a, p.78
A. Corynosoma clavatum Goss, 1940
SA: AHC
WA: Goss, 1940, p.12
? Centrorhynchus sp.
NT; AHC
47. Phalacrocorax melanoleucos
(Vieillot)
Little Pied Cormorant
25/25 : T23 C24 N21 A9
. Echinoparyphium phalacrocoracis Yamaguti,
1939a
SA: THJ, 1942a, p.238
Paryphostomum radiatum (Dujardin, 1845)
Qld, NSW, Vic, SA; THJ, 1942a, p.233
SA: AHC
syn. Echinochasmus tenuicollis 5.3, Johnston,
1917
NSW: SJJ, 1917, p.206
Qld: THJ, 1918, p.212
syn. Paryphostomum tenuicolle (SJJ, 1917)
SA: THI & LMA, 1942b, p.120
syn. Paryphostomum phalacrocoracis Goss, 1940
WA: Goss, 1940, p.2
Petasiger exaeretus Dietz, 1909
NSW, SA: THJ, 1942a, p.236
SA: AHC
Echinostomalidae [spines (27(4))]
NSW: AHC
REC, S, AUST.
Prosthogonimus vitellatus Nicoll, i914a
SA: LMA, 1973, p.859
Haplorchis pumilia (Looss, 1896)
Qld: Pearson, 1960, p.93; 1964, p.618
Haplorchis sprenti Pearson 1964
Qld: Pearson, 1964, p.639
syn. Haplorchis sp.
Qld: Pearson, 1960, p.93
Procerovum sp.
Qld: Pearson, 1960, p.93
Stellantchasmus aspinosus Pearson, 1964
Qld: Pearson, 1964, p.668
syn. Stellantchasmus falcatus of Pearson, 1960,
not Onji & Nishio, 1916
Qld: Pearson, 1960, p.93
Hysteromorpha triloba (Rudolphi, 1819)
Vic, SA: THJ, 1942a, p.238
Posthodiplostomum australe Dubois, 1937a
SA: Dubois & LMA, 1972, p.212
Schwartzitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972, p.205
Dolichosacculus solecarius (S.J. Johnston, 1917)
syn. Dolichosaccus solecarius $.J, Johnston, 1917
NSW: SJJ, 1917, p.218
. Paradilepis minima (Goss, 1940)
SA: Clark, 1957, p.126
syn. Dilepis minima Goss, 1940
WA: Goss, 1940, p.10
Paradilepis sp.
SA: Clark, 1957, p.129
Microsomacanthus cormoranti (Ortlepp, 1938)
syn. Hymenolepis cormoranti Ortlepp, 1938
SA: Clark, 1957, p.131
Woodlandia phalacrocoracis (Woodland, 1929)
syn. Hymenolepis phalacrocoracis Woodland,
1929
SA: Clark, 1957, p.132
Unidentified
Qld, NSW, Vic, SA: AHC
. Eustrongylides phalacrocoracis T.H. Johnston &
Mawson, 194le
SA: THJ & PMM, 194le, p.255
Capillaria jaenschi T.H. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p.245
Contracaecum spiculigerum (Rudolphi, 1909)
Qld, NSW, SA: THJ & PMM, 194lc, p.111
Contracaecum sinulabiatum T.H. Johnston &
Mawson, 1941c
SA: THJ & PMM, 1941c, p.113
Tetrameres sp.
SA: AHC
Synhimantus sp.
SA: THJ & PMM, 1942e, p.71
SA: AHC
Chevreuxia sp.
SA: AHC
Streptocara recta (Linstow, 1879)
SA: THJ & PMM, 1942e, p.71
Desmidocercella sp.
SA: AHC
Microfilaria sp.
NSW: Cleland & THJ, 1912, p.430
MUS. 19(15): 219-325
August, 1986
A. Corynosoma clavatum Goss, 1940
Cc,
N.
WA: Goss, 1940. p.12
SA: AHC
Unidentifiable
Qld: AHC
48. Phalacrocorax sp.
. Renicola sp.
Vic: UMVS
Stictodora diplacantha T.H. Johnston, 1942a
SA: AHC
Posthodiplostamum australe Dubois, 1937a
NT: AHC
Family FREGATIDAE
49, Fregata minor (Gmelin)
Great Frigatebird
1/1: N
Tetrabothrius sp.
Qld: THJ, 1912b, p.108
Unidentified
Qld: AHC
. Contracaecum sp.
Qld, Vic: AHC
Seuratia shipleyi (Stossich, 1900)
Qld: AHC
50, Fregata ariel (Gray)
Least Frigatebird
1/1:CN
. Tetrabothrius polyorchis Nybelin, 1917
WA: Nybelin, 1917, p.15
. Seuratia shipleyi (Stossich, 1900)
Qld: not kept
Family PHAETHONTIDAE
51. Phaethon rubricauda Boddaert
Red-tailed Tropiebird
l/l: N
Contracaecum sp.
SA: AHC (immature)
Family ARDEIDAE
52. Ardea pacifica Latham
Pacific Heron
Distoma sp.
NSW: Krefft, 1873, p.213
. No record. Dendrouterina australiensis Baer &
Bona, 1960, p.13, was attributed to A. pacifica in
mistaken identification of ‘Grey Heron’
. Contracaecum spiculigerum (Rudolphi, 1809)
Qld: THJ & PMM, 1941c, p.111
Porrocaecum reticulatum (Linstow, 1899)
Qld: THJ & PMM, 194lc, p.115
Desmidocercella sp.
HELMINTHS FROM AUSTRALIAN BIRDS 253
Qld: AHC
‘Filarial worms’
Qld: Mackerras, 1962, p.435
53, Ardea novaehollandiae Latham
White-faced Heron
14/14 : T12 C12 N12
. Bucotvle sp,
Tas: AHC
Proacetabulorchis dogieli Belopolskaya &
Bychovskaya-Pavlovskaya, 1954
Qld: LMA & Pearson, 1977, p.130
Maritrema oocystum (Labour, 1907)
Tas: S.J. Smith, 1981, p.393
Haplorchis sprenti Pearson, 1964
Qld: Pearson, 1964, p.639
Procerovum varium Onji & Nishio, 1916
Qld: Pearson, 1964, p.654
Diplostomum galaxiae Smith & Hickman, 1983a
Tas: Smith & Hickman, 1983a, p.29
Posthodiplostomum australe Dubois, 1937a
Qld: Dubois & Pearson, 1967, p.201
SA: Dubois & LMA, 1972, p.212
Apharyngostrigea simplex (S.J. Johnston, 1904)
SA: Dubois & LMA, 1972, p.197
Vie, SA: AHC
syn. Holostontum simplex §.J. Johnston, 1904
NSW: SJJ, 1904, p.112
Parastrigea repens (Chase, 1921)
syn. Holostomum repens Chase, 1921
NSW; Chase, 1921, p.500
Schwartzitrema pandubi (Pande, 1939)
SA: AHC (cysts)
Unidentified
SA: AHC (cysts, metacercariac)
. Dendrouterina australiensis Baer & Bona, 1960
NSW, SA: Baer & Bona, 1960, p.13; Bona,
1975, p.91
Paradilepis urceina Bona, 1975
SA: Bona, 1975, p.503. ’Grey Heron’, quoted
by Baer & Bona, is Ardea novaehollandiae
(colloquial use)
Parvitaenia ardeae (TH. Johnston, 1913)
syn. Bancroftiella ardeae T.H. Johnston, 1913
Qld: THJ, 1913, p.85; 1912b, p.107
Parvitaenia clavipera Bacr & Bona, 1960
Qld: Baer & Bona, 1960, p.13; Bona, 1975,
p.244
syn. Bancroftiella glandularis of THJ, 1912b, (in
part) not (Fuhrmann, 1905)
Qld, NSW; THJ, 1912b, p.107
Parvitaenia paracyclorchida Baer & Bona, 1960
NSW: Bacr & Bona, 1960, p.3; Bona, 1975,
p.247
syn. Bancroftiella glandularis of THJ, 1912b, (in
part), not (Fuhrmann, 1905)
Qld, NSW: THJ, 1912b, p.107
Unidentified
NSW, Tas, SA: AHC
. Contracaecum spiculigerum (Rudolphi, 1809)
NSW: THJ & PMM, 1941c, p.111
Contracaecum sp.
SA: THJ & PMM, 194lIec, p15
Vic: AHC
Porrocaecum sp.
Qld: AHC
Tetrameres sp.
Qld, SA: AHC
54. Ardeola ibis (Linné)
Cattle Egret
. Microtetrameres egretes Rasheed, 1960
SA: AHC
55, Egretta alba (Linneé)
Great Egret
4/3: T2 C1 N3
. Apharyngostrigea sp.
Qld: CIHUS
Echinoparyphium oxyurum 8.J. Johnston, 1917
Qld: SJJ, 1917, p.201
Echinostoma sp.
Qld: THJ, 1912b, p.107
Qld; CIHUS
Patagifer fraternus 3.3. Johnston, 1917
Qld: SJJ, 1917, p.213
Renicola sp.
SA: AHC (cysts, metacercariae)
Haplorchis sprenii Pearson, 1964
Qld: Pearson, 1964, p.639
Proceroyum varium Onji & Nishio, 1916
Qld: Pearson, 1964, p.653
Heterophyidae, unidentified
SA: AHC
Posthodiplostomum australe Dubois, 1937a
SA: Dubois & LMA, 1972, p.212
Distoma sp.
NSW: Krefft, 1873, p.213
. Anomotaenia asymmetrica T.H. Johnston, 1913
Qld: THJ, 1913, p.81
Parvitaenia glandularis (Fuhrmann, 1905)
syn. Bancroftiella glandularis (Fuhrmann, 1905)
Qld: THJ, 1913, p.84
? Qld: CIHUS
, Contracaecum sp.
SA! TH] & PMM, 194Ic, p.114
Qld: CIHUS (immature)
Porrocaecum reticulatum (Linstow, 1899)
Qld: THJ & PMM, 194lc, p.115
Qld: CIHUS
Thelazia sp.
Qld: CIHUS, BM(NH)
. Unidentified
? loc: CIHUS
56, Egretia garzetta (Linné)
Little Egret
. Nephrostomum sp.
Tas: AHC
Echinostomatidae [spines (49(4))]
Tas: AHC
234
a
2
REC. S, AUST, MUS. 19(15); 219-325
Diplostomum amygdalum Dubois & Pearson, 1965
Qld: Dubois & Pearson, 1965, p.90
Apharyngostrigea simplex (S.J. Johnston, 1904)
Qld: Dubois & Pearson, 1965, p.79
57. Egretta intermedia (Wagler)
Intermediate Egret
Haplorchis pumilia (Looss, 1896)
Qld: Pearson, 1964, p.618
Haplorchis sprenti Pearson, 1964
Qld: Pearson, 1964, p.639
Procerovum varium Onji & Nishio, 1916
Qld: Pearson, 1964, p.655
Diplostomum amyzdalum Dubois & Pearson, 1965
Qld; Dubois & Pearson, 1965, p.90
Apharyngostrigea simplex (S.J. Johnston, 1904)
Qld; Dubois & Pearson, 1965, p.79
. None recorded. Baer & Bona (1960) and Bona
(1975) mistakenly quote this bird as host of
Anomotaenia asymmetrica THJ, 1913, and of
Bancroftiella glandularis (Fuhbrmann) of THJ, 1913,
but THJ gives host of both species as Herodias
timorensis Lesson, now Egretta alba.
58. Egretta sacra (Gmelin)
Eastern Reef Egret
. Austrobilharzia terrigalensis S.J. Johnston, 1917
Qld: Rohde, 1977, p.39
. Baerbonia parvitaeniunca (Baer & Bona, 1960)
Qld: Bona, 1975, p.166
syn. kalipora parvitaeniunca Baer & Bona, 1960
Qld: Baer & Bona, 1960, p.9
59. Butorides striatus (Linné)
Striated Heron
4/4.N
Contracaecum sp.
WA: WAM
60, Nycticorax caledonicus (Gmelin)
Rufous Night Heron
3/3; T2 C2 N3
. Haplorchis paravanissimus Pearson & Ow-Yang,
1982
Qld; Pearson & Ow-Yang, 1982, p.S0O
syn. A. vanissinius of Pearson, 1964 (in part) not
Africa, 1938
Qld: Pearson, 1964, p.633
Haplorchis vanissimus Africa, 1938
Qld: Pearson, 1964, p.633
Procerovum varium Onji & Nishio, 1916
Qld: Pearson, 1964, p,654
Stellantchasmus falcatus Onji & Nishio, 1916
Qld: Pearson, 1964, p.664
Clinostomum complanatum (Rudolphi, 1814)
syn. Clinostomum hornum Nicoll, 1914b
Qld: Nicoll, 1914b, p.123
Diplostomum amygdalum Dubois & Pearson, 1965
Qld: Dubois & Pearson, 1965, p.90
August, 1986
SA: Dubois & LMA, 1972, p,206
Posthodiplostomum australe Dubois, 1937a
NT: Dubois & LMA, 1972, p.212
. Parvitaenia ardeae (TH. Johnston, 1913)
SA: Bona, 1975, p.232
NSW: AHC
syn. Bancroftiella ardeae T.H. Jobnston, 1913
Qld: THJ, 1913, p.85
Hymenolepis sp.
NSW: T.H. Johnston, 1912b, p.107
Qld, NSW: AHC
Unidentified
Qld, NSW, SA, NT: AHC
N, Eustrongyvlides sp.
NT: AHC
Contracaecum microcephalum (Rudolphi, 1809)
Qld: CIHUS
Contracaecum nycticeracis T.H.
Mawson, 1941c
NSW: THJ & PMM, 1941c, p.111
Contracuecum spiculigerum (Rudolphi, 1809)
Qld, SA, NT; AHC
Desmidocercella sp,
NT: AHC
Johnston &
6]. Ixobrychus minutus (Linné)
Little Bittern
2/1: N
. Contracaecum sp,
SA: AHC (immature)
Desmidocercella sp.
SA: specimen lost
62, Botaurus poiciloptilus (Wagler)
Australian Bittern
Wvi;:TCN
. Clinostamum complanatum (Rudolphi, 1814)
syn, Clinostomum hornum Nicoll, 1914b
Qld: Nicoll, 1914b, p.123 (immature)
Posthodiplostomum australe Dubois, 1937a
Old: Dubois, 1937b, p.341
Posthodiplastomum oblongum Dubois, 1937a
Qld: Dubois, 1938, p.291
Schwartzitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972,
(metacercariae)
Hexangium sp.
Qld: CIHUS (? in food)
Gyliauchen sp.
Qld: CIHUS (? in food)
Unidentified
NSW: DWRR
p-2058
. Bancroftiella sp.
Qld: BM(NH)
Unidentified
Qld, NSW: AHC
SA: AHC (cysticercoids)
. Capillaria sp,
Qld: AHC
Contracaecum spiculigerum (Rudolphi, 1809)
SA; THJ & PMM, 194l]c, p11]
HELMINTHS FROM AUSTRALIAN BIRDS
Contracaecum sp., 2? microcephalum (Rudolphi,
1809)
Qld: CIHUS
. Arhythmorhynchus brevis Van Cleave, 1916
Qld: CIHUS
Family CICONIIDAE
63. Xenorhynchus asiaticus (Latham)
Black-necked Stork
11:TCN
. Chaunocephalus ferox (Rudolphi, 1795)
Qld: Nicoll, 1914b, p.117
’, Clelandia parva T.H. Johnston, 1909b
NSW: THJ, 1909b, p.146
. Contracaecum sp.
SA: THJ & PMM, 194l1c, p.114
Unidentified
SA (cage): AHC (larva, encysted)
Family PLATALEIDAE
64. Plegadis falcinellus (Linné)
Glossy Ibis
1/0
. Patagifer bilabus (Rudolphi, 1819)
Qld: Nicoll, 1914a, p.338
. Contracaecum spiculigerum (Rudolphi, 1809)
Qld: AHC
Physaloptera sp.
Qld: AHC
65. Threskiornis acthiopica (Latham)
Sacred Ibis
7/7: 75 C4 NS A4
. Echinostoma acuticauda Nicoll, 1914b
Qld: AHC
Patagifer acuminatus 8.J. Johnston, 1917
Qld: SJJ, 1917, p.210
syn. Echinostoma sp.
Qld: THJ, 1912b, p.107
syn. Patagifer bilobus of THY,
(Rudolphi, 1819)
Qld: THJ, 1916, p.47
Patagifer sp. [spines 2x (30(4))]
SA: AHC
Patagifer sp. [spines 2x (29(3))]
Vic: AHC
Platynotrema biliosum Nicoll, 1914b
Qld: Nicoll, 1914b, p.118
Hysteromorpha plataleae Dubinina & Dubinin,
1940
Qld: Dubois & LMA, 1972, p.208
Parastrigea sp.
Qld: Dubois & LMA, 1972, p.198
Strigea baylisi Dubois, 1937a
SA: Dubois & LMA, 1972, p.198
Unidentified
SA: AHC (cysts)
1916, not
Cc,
QO
A,
ie
oo
uw
Paradilepis urceina Bona, 1975
SA: Bona, 1975, p.503
Hymenolepididae, unidentified
SA: AHC
Unidentified
Qld, Vic, SA: AHC
. Capillaria sp.
SA: AHC
‘Nematoda (? hookworm)’
Qld: CLHUS
Skrjabinoclava sp. cf. S. alii Ali, 1968
SA: AHC
Syneuaria contorta (Molin, 1858)
Vic, SA: PMM, 1982, p.19
. Polymorphus biziurae T.H, Johnston & Edmonds,
1948
Tas, SA: AHC
66. Threskiornis spinicollis (Jameson)
Straw-necked Ibis
2/1: N
Echinostoma acuticauda Nicoll, 1914b
Qld: Nicoll, 1914b, p.110
Patagifer bilobus (Rudolphi, 1819)
Qld: Nicoll, 1914b, p.115
NSW: CIHUS
Strigea baylisi Dubois, 1937a
Qld: Dubois, 1937b, p.235
Opisthorchis obsequens Nicoll, 1914b
NSW: CIHUS
. Hymenolepis ibidis T.H. Johnston, 1913
Qld: CIHUS
Hymenolepis sp.
Aust: BM(NA)
Unidentified
Old: AHC
Vic: UMVS
Capillaria sp.
SA: AHC
Physaloptera sp.
Qld: THJ & PMM, 194le, p.257
67. Platalea regia Gould
Royal Spoonbill
Wl: A
Orchipedum sufflavum Nicoll, 1914b
Qld: Nicoll, 1914b, p.108
Patagifer bilobus (Rudoltphi, 1819)
Qld: SJJ, 1913, p.37]
? loc: CLHUS
. Cyclorchida omalancristrota (Wedl, 1856)
Qld: THJ, 1913, p.86; Bona, 1975, p.412
Unidentified
SA: specimens lost.
68. Platalea flavipes Gould
Yellow-billed Spoonbill
5/5: T5 C4 N2 A2
Patagifer sp. [spines 2x ? (29(4))]
SA: AHC
236
Patagifer sp. [spines 2x (27(? 3))]
Qld: AHC
Schwartzitrema pandubi (Pande, 1939)
SA: Dubois & LMA, 1972, p.205 (cysts)
Strigea baylisi Dubois, 1937a
SA: Dubois & LMA, 1972, p.198
. Paradilepis patriciae Baer & Bona, 1960
SA: Baer & Bona, 1960, p.17; Bona, 1975,
p.530
Hymenolepis ibidis T.H. Johnston, 1913
Qld: THJ, 1913, p.88
Hymenolepis sp.
SA: AHC
Unidentified
Qld, SA: AHC
. Contracaecum sp.
SA: AHC
. Polymorphus biziurae T.H. Johnston & Edmonds,
1948
SA: AHC
Family ANATIDAE
69. Anseranas semipalmata (Latham)
Magpie Goose
1/1:TCN
. Schistosomatidae (eggs, from nasal mucosa)
Qld: Blair & Otteson, 1979, p.984
Notocotylus attenuatus (Rudolphi, 1809)
Qld: Nicoll, 1914b, p.125
NT: AHC
Typhlocoelum reticulare S.J. Johnston, 1913
Qld: SJJ, 1913, p. 372
NT: AHC
Echinostoma revolutum (Froelich, 1802)
Qld: Nicoll, 1914b, p.115
Qld: AHC
. Angularella australis (Maplestone, 1921b)*
Aust: BM(NH)
Biuterina sp.
Aust: BM(NH)
Cloacotaenia megalops (Nitzsch in Creplin, 1829)
syn. Hymenolepis megalops (Nitzsch, 1829)
Qld: THJ, 1913, p.89
Passerilepis stylosa (Rudolphi, 1810)
Aust: BM(NH)
Sobolevicanthus terraereginae (T.H. Johnston,
1913)
syn. Hymenolepis terraereginae T.H. Johnston,
1913
Qld: THJ, 1913, p.89
Unidentified
Qld, Vic, NT: AHC
. Amidostomum anseris (Zeder, 1800)
Vic (cage): UMVS
Epomidiostomum sp.
NT: AHC
* Yamaguti, 1959, p.235, notes that this species cannot
be referred to Angularella (syn. Angularia) and
probably belongs to some other genus.
REC. S. AUST, MUS. 19(15): 219-325
August, 1986
Heterakis sp.
Vic (cage): UMVS
Tetrameres anseranas Mawson, 1979
NT: PMM, 1979, p.178
Vic (cage): UMVS
Tetrameres fissispina (Diesing, 1851)
Philadelphia Zoo: Canavan, 1931, p.221
Streptocara crassicauda (Creplin, 1829)
Vic (cage): UMVS
70. Dendrocygna arcuata (Horsfield)
Wandering Whistling-Duck
Schistosomatidae, unidentified
Qld: Blair & Otteson, 1979, p.984 (eggs from
nasal mucosa)
Notocotylidae, unidentified
Qld: AHC
Echinostomatinae [spines ? (35(5))]
Qld: AHC
. Ophiotaenia hylae T.H. Johnston, 1912e
Qld: Maplestone, 1921la, p.404 (? with food)
Cloacotaenia megalops (Nitzsch in Creplin, 1829)
Qld: CIHUS
Diorchis flavescens (Krefft, 1873)
Qld: Maplestone, 192la, p.403
Diplogynia oligorchis (Maplestone, 1922a)
syn. Coturnia oligorchis Maplestone, 1922a
Qld: Maplestone, 1922a, p.55
syn. Diploposthe laevis of THJ, 1913, not (Bloch,
1782)
Qld: THJ, 1913, p.91
Hymenolepis ibidis (T.H. Johnston, 1913)
Qld: CIHUS
? Passerilepis zosteropis (Fuhrmann, 1918)
Aust: BM(NH)
Unidentified
Qld: AHC
71. Dendrocygna eytoni (Eyton)
Plumed Whistling-Duck (endemic)
Schistosomatidae, unidentified
Qld: Blair & Otteson, 1979, p.984 (eggs from
nasal mucosa)
72. Cygnus atratus (Latham)
Black Swan (endemic)
19/19 : T14 C19 N13 A3
Trichobilharzia sp.
SA: THJ, 1941, p.276
Notocotylus attenuatus (Rudolphi, 1809)
Qld: Nicoll, 1914b, p.125
Notocotylus sp.
Tas, SA: AHC
Paramonostomum caeci Smith & Hickman, 1983b
Tas: Smith & Hickman, 1983b, p.86
Notocotylid, ? sp. B, S.J. Smith, 1981
Qld, Tas: S.J. Smith, 1981, p.256
Notocotylidae, unidentified
NSW, Tas, SA: AHC
Hyptiasmus magnus S.J. Johnston, 1917
C,
HELMINTHS FROM AUSTRALIAN BIRDS 237
Vie: SJJ, 1917, p.244
syn, Monostomuni sp.
Vie: THI, 1910, p.97
Cyclocoelidae, unidentified
NSW, Vic, SA; AHC
Echinoparyphiunt ellisi TA. Johnston & Simpson,
1944
SA; THI & LMA, 1949, p.25]
syn. Echinoparyphium sp.
? Old: Verma, 1936, p.1535; see THI & LMA,
1949, p.253
Echinoparyphium gizzardai Verma, 1936
Old: Verma, 1936, p.155; see TH] & LMA,
1949, p,253
” Echinostoma minimunt Verma, 1936
sce Verma, 1936, p,150. We consider the host
given by Verma as ‘Black Swan’ from Patna,
Behar, to be au error for Mute Swan, cited as the
host of Echinostoma longicirrum from. the same
locality (p.152)
Echinostomad revolutum (Froelich, 1802)
Qld: Nicoll, 1914b, p.1L5
SA: THJ & LMA, 194]e, p.321
Echinostoma sp. {spines (37(4-5))]
Tas; AHC
kermatrema longitestis (Verma, 1936)
Calcutta (? cage): Srivastava, 1972, (p.177
syn. Eupuryphium longitestis Verma, 1936
Calcutta (? Qld): Verma, 1936, p.156; see THI
& LMA, 1949, p,253
Echinostomatidac, unidentified
NSW: AHC
Echinostomatinae {spines 41(5))]
SA: AHC (immature)
Psilochasmus oxyurus (Creplin, 1825)
Tas: S.J. Smith, 1981, p.181
Psilostomum sp, A, S.J. Smith, 1981
Tas: SJ. Smith, 1981, p.201 (? this hast)
Psilostomum sp. B, S.J. Smith, 1981
Yas: SW Smith, 1981, p.214
Psilostonium sp.
SA: AHC
Cylindroirema cygni Angel, 1973
SA: LMA, 1973, p.854
Apatemon intermedius (S.J. Johnston, 1904)
SA: TH] & LMA, 1951, p.67
syn, Hemistomum intermedium 8.3, Johnston,
1904
NSW: SJJ, 1904, p.110
Carylurus magniacetabulus Dubois & Angel, 1972
SA; Dubois & LMA, 1972, p.204
Unidentilied
NSW: AHC
Armadoskrjabinia globosa (Szpotanska, 1931)
syn. Hymenolepis globosa Szpotanska, 1931
Aust: Szpotanska, 193], p.260
Qld: BM(UNH)
Austvraliolepis southwelli (Szpotanska, 1931)
syn, Aymenolepis sourhwelli Szpotanska, 1931
Aust: Szpotanska, 1931, p,260
syn. Echinorhyachotaenia nana Maplestone &
Southwell, 1922
Qld: Maplestone & Southwell, 1922b, p.193
Cloacotaenia megalops (Nivzsch in Creplin, (829)
Warsaw (cage): Kotechi, 1970, p.332
Qld: BM(NH), CIHUS
NSW: DWRR
Vic, SA; AHC
Diorchis spiralis Szpotanska, 1931
Aust: Szpotanska, 1931, p.258
Diorchis stefanski Czaplinski, 1955
Warsaw (cage): Kotechi, 1970, p.332
Drepanidotaenia lanceolata (Bloch, 1872)
Aust: Szpotanska, 1931, p.254
Qld: BM(NH)
NSW: WLRR
Vic, SA: AHC
syn. Aymenalepis lanceolata (Bloch, 1782)
Qld: Maplestone & Southwell, 1922b, p.197
Drepanidotuenia rapida (Szpotanska, 193))
syn. Hymenolepis rapida Szpotanska, 1931
Aust: Szpotanska, 1931, p.25]
Drepanidotaenia sp.
SA: AHC
Dicranotaenia coronula (Dujardin, 1845)
Warsaw (cage): Kotechi, 1970, p.332
Hymenolepis chenopis Palmer, 1981
Vic: Palmer, |981, p.126
Hymenolepis sp.
Bengal (cage); Southwell, 1916, p.11
Qld: BM(NH)
Hymenolepis (8.1) liophallos Krabbe, 1869
Europe (cage): Krabbe, 1869, p.291
“Hymenolepis variabilis n.sp. Maplestone”
Qld: CIHUS, We have found no record of
publication of this species by Maplestone,
Hymenolepis (sd,) micrancristrota (Wedl, 1855)
syn. Taenia micrancristrota Wedl, 1855
Hungary (cage): Wedl, 1855, p.6
Monosaccanthes curjiosa (Szpotanska, 1931)
syn. Mymenolepis curiosa Szpotanska, 1931
Aust: Szpotanska, 1931, p,252
Qld: BM(NH)
Monosaccanthes kazachsranica (Maksinova, 1963)
Vic: Palmer, 1981, p35
Monosaecanthes sp.
SA: AHC
Parabisaccanihes bisacculina (Szpotanska, 1931)
Vic: Palmer, 1981, p.130
Qld: BM(NH)
SA: AHC
syn, Drepanidotaenia bisacculina Szpotanska,
193]
Aust: Szpotanska, 1931, p.247
Tscherlkovilepis Krabbei (Kowolewski, 1895)
Warsaw (cage): Kotechi, 1970, p,332
Gastrotaenta sp,
SA: AHC
Nematoparataenia paradoxe Maplestone &
Southwell, 1922b
Qld: Maplestone & Southwell, 1922b, p.189
Vic, SA: AHC
Aust: BM(NH)
Cyclophyllidea, unidentified
SA: AHC
Unidentified
NSW: Cleland, 1922, p.105
Aust: AM
Qld, NSW, Tas, SA: AHC
Qld: CIHUS (larvae)
Vic: UMVS
N. Capillaria obsignata Madison, 1945
England (cage): Wakelin, 1963, p.38]
syn. Capillaria anatis (Schrank, 1790)
England (cage): Wakelin, 1963, p.38l.
Capillaria ellisi TH. Johnston & Mawson, 1945b
SA: THJ & PMM, 1945b, p.247
Amidostomum cygni Wehr, 1933
SA: THJ & PMM, 1947, p.550; PMM, 1980,
p.10
Tas: AHC
Amidostomum sp.
NSW: DWRR
Vic: AHC
Pseudamidostom sp.
NSW: DWRR
Trichostrongylus tenuis
England (cage): Wakelin, 1963, p.381.
Heterakis circunivallata (Linstow, 1906b)
Germany: Linstow, 1906b, p.251
Heterakis vesicularis (Froelich, 1791)
syn. Heterakis papillosa (Bloch, 1782)
Berlin Zoo: Schneider, 1866, p.69
Tetrameres australis TH. Johnston & Mawson,
194le
SA: THJ & PMM, 194le, p.262
SA: AHC
Echinuria uncinata (Rudolphi, 1819)
Vic: UMVS
Tas, SA: AHC
A. Polymorphus sp.
SA: AHC
Corynesoma sp.
SA: AHC
73. Stictonetta naevosa (Gould)
Freckled Duck (endemic)
19/11 T4 C7 N10
T. Notocotylidae, unidentified
SA: AHC
Echinoparyphium sp. [spines (41(4))]
SA: AHC
Psilochasmus sp.
SA: AHC
Psilostomum sp.
SA: AHC
C. Diorchis sp.
NSW; DWRR
Cloacotaenia sp.
NSW: DWRR
Unidentified
SA: AHC
N. Amidostomum acutum (Lundahl, 1848)
SA: PMM, 1980, p.9
Amidostomum ‘sp. @
NSW: DWRR
Tetrameres sp.
SA: AHC
REC. 8S. AUST. MUS. 19(15); 219-325
Z
August, 1986
Synhimantus. sp.
SA: AHC
74. Cereopsis novaehollandiae Latham
Cape Barren Goose (endemic)
. Psilochasmus sp.
Tas: AHC
. Fimbriarioides intermedia (Fuhrmann, 1913)
syn. Fimbriaria intermedia Fuhrmann, 1913
Calcutta (cage): Meggitt, 1933, p.153
Sobolevicanthus sp.
SA: AHC
. Amidostomum anseris (Zeder, 1800)
Tas: PMM, 1980, p.9
Heterakis vesicularis (Froelich, 1791)
SA: AHC
syn. Heterakis caudata (Linstow, 1906)
Calcutta Zoo: Maplestone, 1932, p.413
Heterakis chenonettae T.H. Johnston, 19]2a
Tas: AHC
Heterakis dispar (Schrank, 1790)
Bass Str: BM(NH)
75. Tadorna tadornoides (Jardine & Selby)
Australian Shelduck (endemic)
5/5: T2 C3 N3
. Echinoparyphium sp. [spines incomplete]
SA: AHC
Echinostoma revolutum (Froelich, 1802)
SA: AHC
syn. ‘echinostome larvae’) Bradley, 1927
NSW: Bradley, 1927, p.675
. Staphylepis lamellata (Woodland, 1930)
syn. Hymenolepis lamellata Woodland, 1930
London Zoo: Woodland, 1930, p.226
Hymenolepis sp.
Vic: AHC
Unidentified
NSW, SA, Vic: AHC
. Epomidiostomum sp.
NSW: AHC
Tetrameres sp.
Vic: AHC
Echinuria uncinata (Rudolphi, 1819)
Vic: AHC
Streptocara crassicauda (Creplin, 1829)
Vic: AHC
76. Tadorna radjah Garnot
Radjah Shelduck
1/1:N
N. Amidostomum acutum (Lundahl, 1848)
NT: PMM, 1980, p.9
77. Anas superciliosa Gmelin
Pacific Black Duck
19/19 ; TL] C13 N8
T. Trichobilharzia australis Blair & Islam, 1983
Qld: Blair & Islam, 1983, p.89
HELMINTHS FROM AUSTRALIAN BIRDS 2g
Trichobilharzia sp.
Qld: Blair & Otteson, 1979, p.983
Schistosomatidae (eggs, nasal mucosa)
Qld, NSW, Vic: Blair & Otteson, 1979, p.984
Notocotylus atienuatus (Rudolphi, 1809)
Qld: Nicoll, 1914b, p.127
Parumonostomum bursae Smith & Hickman,
1983b
Tas: Smith & Hickman, 1983b, p.95
Paramonostomum caeci Smith & Hickman, 1983b
Tas: Smith & Hickman, 1983b, p.86
Notocotylidae, unidentified
Qld, SA: AHC
Typhlocoelum sp.
Qld, Vic, SA: AHC
Echinostoma revolutum (Froelich, 1802)
Qld: SJJ, 1913, p.370
NSW, SA: THJ & LMA, 194le, p.321
Tas: AHC
syn, Echinostoma sp.
Qld, NSW: TH J, 1912b, p.108
Aypoderaeum sp.
Qld, SA: AHC
Patagifer sp. [spines 2x ? (26(3))]
NSW: AHC
Echinostomatinae [spines (37(5))}
Qld, SA: AHC
Echinostomatinae [spines (37(5))|
SA: AHC
Echinostomatinae [spines (45(5))]
SA: AHC
Psilochasmus oxyurus (Creplin, 1825)
Tas: S.J. Smith, 1981, p.181
Psilostamum sp.
Qld: AHC
? Prosthogonimus sp.
SA: AHC
Levinseniella tasmaniae (S.J. Smith, 1974)
syn. Microphallus tasmaniae S.J. Smith, 1974
Tas: S.J. Smith, 1981, p.107
Maritrema calvertense S.J, Smith, 1974
Tas: S.J. Smith, 198], p.64
Apalemon gracilis (Rudolphi, 1819)
Tas: Smith & Hickman, 1983a, p.23
Cotylurini, unidentified
Qld, SA: AHC
Diplostomidae, unidentified
SA: AHC
. Cloacotaenia megalops (Nitzsch in Creplin, 1829)
Qld; CIHUS
NSW: DWRR
WA: BM(NH)
syn. Hymenolepis megalops (Nitzsch in Creplin,
1829)
NSW: THI, 1912b, p.108
syn. Taenia cylindrica Krefft, 1873
NSW: Krefft, 1873, p.220
Diorchis flavescens (Krefft, 1873)
Qld: THJ, 1912a, p.66
Qld: CIHUS
NSW: AHC, DWRR
syn, Tuenia flavescens Kreflt, 1873
EA: Krefft, 1873, p.219
Drepanidotaenia lancealata (Bloch, 1782)
Qld: BM(NH)
Hymenolepis sp.
NSW: THJ, 1912b, p.108
Qld, SA: AHC
Microsomacanthus collaris (Batsch, 1786)
syn. Hymenolepis collaris (Batsch, 1786)
NSW: THJ, 1912b, p.108
syn. Tuenia bairdii Krelft, 1873
EA: Krefft, 1873, p.224
Fimbriaria fasciolaris (Pallas, 1871)
NSW: THJ, 1912b, p.108
syn. Tuenia pediformis Kretft, 1873
EA: Krefft, 1873, p.222
Fimbriaria sp.
SA: AHC
Diploposthe laevis (Bloch, 1782)
NSW: THJ, 1912d, p.12
Gastrotaenia sp.
NSW: DWRR
Unidennitied
NSW: Cleland, 1922, p.105
Tas: Munday & Green, 1972, p.4
Qld, NSW, Tas, SA, NT: AHC
Vic: MUVS
. Captillarta sp.
NSW: Cleland, 1922, p.105 (AHC)
Amidostonmum acutum (Lundahl, 1848)
Tas, SA: Mawson, 1980, p.9
Amidostomium sp.
NSW: DWRR
Epomidiostomum sp.
NSW: DWRR
Tas, SA, NT: AHC
Porrocuecum crassum (Deslongchamps, 1824)
Tas: AHC
Contracaecum microcephalum (Rudolphi, 1809)
NSW: THJ & PMM, 1941c, p.114
Tetrameres fissispina (Diesing, 1861)
SA: THJ & PMM, 1949, p.67
Tas, NT: AHC
Physaloptera sp.
NSW: THJ & PMM, 194]e, p.257
Echinuria sp.
NSW: DWRR
78, Anas gibberifrons S. Miller
Grey Teal
14/11 : TS C8 N6
Trichobilharzia sp.
NSW: Bearup, 1957, p.163 (eggs in nasal
mucosa)
Schistosomatidae
Qld, NSW, Vie; Blair & Otteson, 1979, p.984
(eggs in nasal mucosa)
Notocotylidae, unidentified
SA: AHC
Typhlocoelum sp.
SA: AHC
Cyclocoelidae, unidentified
NSW: AHC
Echinostoma sp. {spines (35(4 or 5))]
240
REC, 8, AUST. MUS, 19(15): 219-325
NT; AHC
Hypoderaeum sp.
SA: AHC
Echinostomatidae, unidentified
NSW: AHC
Echinostomatinae [spines (45(5))]
SA: AHC
Unidentified
NSW: AHC
. Cloacotaenia ? megalops
NSW: DWRR
Diorchis flavescens (Krefft, 1873)
Qld: Baylis, 1934b, p.129
NSW: DWRR
Hymenolepis robertsi Baylis, 1934b
Qld: Baylis, 1934b, p.129
Haploparaxis yeitchi Baylis, 1934b
Qld: Baylis, 1934b, p.129
Staphylepis lamellata (Woodland, 1930)
SA: AHC
Gastrotaenia sp.
NSW: DWRR
Unidentified
Vic, SA, NT: AHC, UMVS
. Strongyloides sp.
NSW: DWRR
Capillaria sp.
NSW: DWRR
NT: AHC
Amidostomum acutum (Lundahl, 1848)
SA: PMM, 1980, p.9
Epomidiostomum uncinatum (Lundahl, 1848)
SA: THJ & PMM, 1942e, p.73
Vic: UMVS
Epomidiostomum sp.
NSW: AHC
Tetrameres sp.
SA: THJ & PMM, 1942e, p.71
NSW: DWRR
Echinuria uncinata (Rudolphi, 1819)
NT: AHC
syn. Echinuria querquedulae T.H. Johnston &
Mawson, 1942e
SA: THJ & PMM, 1942e, p.71
Echinuria sp.
NSW: DWRR
Streptocara sp.
SA: THJ & PMM, 1942e, p.72
NSW: DWRR
79, Anas castanea (Eyton)
Chestnut Teal (endemic)
1/1:TCN
Hypoderaeum conoideum (Bloch, 1782)
Tas: AHC
Aypoderaeum sp.
NSW: AHC
Levinseniella tasmaniae (S.J. Smith, 1974)
syn. Microphallus tasmaniae $.J, Smith, 1974
Tas: S.J. Smith, 1974, p.202
Maritrema calvertense S.J. Smith, 1974
Tas: S.J. Smith, 1974, p.201
Cc.
N.
August, [986
Cloacotaenia mega/ops (Nitzsch in Creplin, 1829)
syn. Hymenolepis megalops (Nitzsch in Creplin,
1829)
NSW: THJ, 1912a, p.66
Diorchis flavescens (Krefft, 1873)
NSW: THJ, 1912d, p.15
Diorchis sp.
NSW: DWRR
Diploposthe laevis (Bloch, 1782)
NSW: THJ, 1912d, p.12
Microsomacanthus collaris (Batsch, 1786)
syn. Hymenolepis collaris (Batsch, 1786)
NSW: THJ, 1912b, p.108
Fimbriaria fasciolaris (Pallas, 1871)
syn. Taenia pediformis Kreffit, 1873
NSW: Krefft, 1873, p.222
NSW: THJ, 1912a, p.66
Unidentified
Vic: AHC
. Capillaria anatis (Schrank, 1790)
G. Brit. (cage): Wakelin, 1965, 0.293
Capillaria sp.
Tas: AHC
Epomidiostomum sp,
Tas: AHC
Tetrameres fissispina (Diesing, 1861)
Tas: AHC
Echinuria uncinata (Rudolphi, 1819)
Tas: AHC
80. Anas rhynchotis Latham
Australian Shoveler
5/5 :T2 C5 N5
. Schistosomatidae, unidentified
NSW: Blair & Otteson, 1979, p.984 (eggs from
nasal mucosa)
Echinostoma revolutum (Froelich, 1802)
SA: THJ & LMA, 194ic, p.321
Echinostomatidae, unidentified,
NSW: AHC
Echinostomatinae [spines (45(5))]
SA: AHC
. Cloacotaenia ? megalops (Nitzsch in Creplin,
1829)
NSW: DWRR
Diorchis flavescens (Krefft, 1873)
syn. Taenia flavescens Krefft, 1873
NSW: Krefft, 1873, p.219
Gastrolaenia sp.
NSW: DWRR
Unidentified
NSW: UMYS
SA: AHC
Capillaria sp.
SA: AHC
Amidostomum acutum (Lundahl, 1848)
NSW: UMVS
Tetrameres sp.
SA: AHC
Streptocara crassicauda (Creplin, 1829)
SA: AHC
Streptocara sp,
HELMINTHS FROM AUSTRALIAN BIRDS 241
NSW: UMVS
Acuariidae, unidentified
NSW: UMYVS
81. Malacorhynchus membranaceus
(Latham)
Pink-eared Duck (endemic)
3/2 : T2 C2 N2
. Paramonostomum sp.
SA: AHC
Notocotylidae, unidentified
Vic, SA: AHC
Cyclocoelum sp.
Vic: AHC
Echinostoma sp. [spines incomplete]
Vic: AHC (immature)
? Microphallidae, unidentifiable
SA: AHC
Unidentified
Vic, SA: AHC
. Unidentified
Vic, SA: AHC
. Capillaria sp.
SA: AHC
Acuariidae, unidentified
SA: AHC (larva)
82. Aythya australis (Eyton)
Hardhead
5/4: 73 C4 N2
. Schistosomatidae (eggs, nasal mucosa)
Qld, NSW: Blair & Otteson, 1979, p.984
Echinostomatidae, unidentified
NSW, SA: AHC
Cyclocoelidae
NSW: AHC
SA: AHC (eggs)
‘. Cloacotaenia, sp.
NSW: WLRR
Diorchis flavescens (Kretft, 1873)
NSW: THJ, 1912d, p.15
Diplogynia oligorchis (Maplestone, 1922)
Qld: CIHUS
Diplogynia, sp.
NSW: WLRR
? Sobolevicanthus sp.
NSW: WLRR
Diploposthe laevis (Bloch, 1782)
Qld: Maplestone, 1922a, p.60
NSW: TH4J, 1912d, p.4
Aust: BM(NH)
syn. Tuenia tuberculata Krefft, 1873
NSW: Krefft, 1873, p.215
Fimbriaria sp.
NSW: WLRR
SA: not kept
Gastrotaenia, sp.
NSW: WLRR
Unidentified
Qld, SA, NT: AHC
Vic: UMVS
N.
Capillaria sp.
SA: AHC
Epomidiostomum sp,
NSW: WLRR (‘sp. 3’)
Streptocara crassicauda (Creplin, 1829)
Tas: AHC
Streptocara sp.
Tas: Munday & Green, 1972, p.12
83. Chenonetta jubata (Latham)
Maned Duck (endemic)
Schistosomatidae, unidentified
Qld: Blair & Otteson, 1979, p.984 (eggs from
nasal mucosa)
Echinostomatidae, unidentified
NSW: AHC
. Heterakis chenonettae T.H. Johnston, 1912a
NSW: TH J, 1912a, p.71; THJ & PMM, 194]c,
p.l15
Amidostomum sp.
Qld, NSW: WLRR
Epomidiostomum sp.
NSW: WLRR
84, Nettapus coromandelianus (Gmelin)
Cotton Pygmy-Goose
Cyclocoelum sp.
Qld: AHC
85. Nettapus pulchellus Gould
Green Pygmy-Goose
2/1:CN
Notocotylus attenuatus (Rudolphi, 1809)
Qld: Nicoll, 1914b, p.125
Echinostoma revolutum (Froelich, 1802)
Qld: Nicoll, 1914b, p.115
. Unidentified
NSW: AHC
. Echinuria uncinata (Rudolphi, 1819)
NT: AHC
86. Oxyura australis Gould
Blue-billed Duck (endemic)
5/5: T4 C5 N2 Al
Notocotylidae, unidentified
Tas, SA: AHC
Echinostomatinae [spines ? (37(5))]
SA: AHC (immature)
Echinostomatinae [spines ? (40(5))]
SA: AHC
Apatemon intermedius (S.J. Johnston, 1904)
SA: Dubois & LMA, 1972, p.201
. Unidentified
Tas: Munday & Green, 1972, p.4
Tas, SA: AHC
. Capillaria sp.
SA: AHC
Epomidiostomum sp.
Tas: AHC
te
id
Streptocara crassicauda (Creplin, 1829)
Tas: AHC
. Unidentified
Tas: specimens lost
87. Biziura lobata (Shaw)
Musk Duck (endemic)
17/17 : Tl6 C13 N7 AQ
Notocotylidae, unidentified
NSW, SA: AHC
Echinoparyphium ellisi (T.H. Johnston & Simpson,
1944)
SA: THJ & LMA. 1949, p.248
Echinostoma revolutum (Froelich, 1802)
Qld, SA: THJ & LMA, 194lc, p.321
Echinostomatinae [spines (35(5))]
Qld, SA: AHC
Echinostomatidae [spines (37(5))]
NSW
Echinostomatidac, unidentified
NSW: AHC
Psilochasmus sp.
SA: AHC
Apatemon vitelliresiduus Dubois & Angel, 1972
Qid, SA: Dubois & LMA, 1972, p.199
Unidentified
NSW: AHC
. Taenia moschata Krefft, 1873
NSW: Krefft, 1873, p.223
(THJ, 1912d, p.3 noted that specimens lost)
Cyclophyllidea, unidentified
SA: AHC
Cloacotaenia ? megalops (Nitzsch in Creplin, 1829)
NSW: DWRR
Sobolevicanthus sp.
NSW: DWRR
Gastrotaenia sp.
NSW: DWRR
Unidentified
Qld, NSW, Tas, SA: AHC’
. Capillaria sp.
NSW: DWRR
Amidostomum biziurae T.H. Johnston & Mawson,
1947
SA: THJ & PMM, 1947, p.551; PMM, 1959,
p.152
NSW, SA: PMM, 1980, p.9
Amidostomum sp. (sp. 5)
NSW: DWRR
Tetrameres biziurae T.H. Johnston & Mawson,
1941e
SA: THJ & PMM, 194le, p.261
NSW: AHC
Streptocara recta (Linstow, 1879)
SA: AHC
Streptocara formosensis Sugimoto, 1930
NSW: AHC
. Polymorphus biziurae T.H. Johnston & Edmonds,
1948
SA: THJ & Edmonds, 1948, p,71
NSW, SA: AHC
REC. S, AUST. MUS.. 19(15): 219-325
August, 1986
Family ACCIPITRIDAE
88. Elanus notatus Gould
Black-shouldered Kite (endemic)
3/0
N. Porroccaeum circinum T.H, Johnston & Mawson,
1941b
Vic: AHC
89. Aviceda subcristata Gould
Pacific Baza
. Neodiplostomum spathula (Creplin, 1829)
syn. Neodiplostomum australiense Dubois, 1937a
Qld: Dubois, 1937b, p.337
. Centrorhynchus asturinus (T.H. Johnston, 1913)
NSW: THJ, 1918, p.215
syn. Echinorhynchus bazaé Southwell & Macfie,
1925
Qld: Southwell & Macfie, 1925, p.177
Unidentified
Qld: CIHUS
90. Milvus migrans (Boddaert)
Black Kite
3/1 CN
. Unidentified
SA: AHC
. Procyrnea mansioni (Seurat, 1914)
SA: AHC
91. Haliastur sphenurus (Vieillot)
Whistling Kite
9/5: T3 Cl NI
. Echinostomatinae [spines (45(5))]
SA: AHC (not mature)
Haplorchis paravanissimus Pearson & Ow-Yang,
1982
Qld: Pearson & Ow-Yang, 1982, p.50
syn. H. vanissimus of Pearson, 1964 (in part) not
Africa, 1938
Qld: Pearson, 1964, p.633; Pearson & Ow-Yang,
1982, p.50
Haplorchis vanissimus Africa, 1938
Qld: Pearson, 1964, p.633; Pearson & Ow-Yang,
1982, p.49
Haplorchis yokogawai (Katsuta, 1932)
Qid: Pearson, 1964, p.622
Procerovum yarium Onji & Nishio, 1916
Qld: Pearson, 1964, p.654
Stellantchasmus falcatus Onji & Nishio, 1916
Qld: Pearson, 1964, p.665
Mesostephanus haliasturis Tubangui & Masilungan,
1941
SA: AHC
syn. Mesosiephanus minor Dubois & Pearson,
1965
Qld: Dubois & Pearson, 1965, p.97
? Diplostomum spathaceum (Rudolphi, 1819)
SA: AHC
on
HELMINTHS FROM AUSTRALIAN BIRDS 243
Neodiplostomum spathula (Creplin, 1829)
Qld: Dubois & Pearson, 1967, p.196
NSW: Dubois, 1938, p.264
Neodiplostomum subaequipartiitnum Dubois &
Pearson, 1967
Qld: Dubois & Pearson, 1967, p.199
SA: Dubois & LMA, 1972, p.210
Strigea vlandulosa Dubois, 1937a
Qld: Dubois, 1937b, p.244
SA: Dubois & LMA, 1972, p.198
Allocreadiidae, unidentified
SA: AHC (metacercariae)
. Unidentified
SA, NT; AHC
. Capillaria sp.
SA: AHC
Contracaecum ceylanicum (Linstow, 1904)
Qld: CIHUS
Procyrnea mansioni (Seurat, 1914)
syn. Cyrnea mansioni (Seurat, 1914)
Qld: PMM, 1968d, p.749
Microfilaria sp.
Qld: Mackerras, 1962, p.436
92. Accipiter fasciatus (Vigors & Horsficld)
Brown Goshawk
10/8 : T2 N8& A3
Apatemon intermedius (S.J. Johnston, 1904)
SA: Dubois & LMA, 1972, p,.201
Unidentified
Qld, SA: AHC
. Microtetrameres paraccipiter Mawson, 1977
Qld, Tas, SA, NT: PMM, 1977, p.253
Microletrameres sp.
Vic: AHC
Spirurida, unidentified
NT: AHC (larva)
. Centrorhynchus asturinus (TH. Johnston, 1913)
Qld: Southwell & Macfie, 1925, p,l63
SA: THJ & Deland, 1929, p.148
NT: AHC
syn, Echinorhyachus sp.
NSW: THJ, 1910, p.100
93, Accipiler cirrhocephalus (Vieillot)
Collared Sparrowhawk
5/5 : T2 C1 NS
. Neodiplostamum spathula (Creplin, 1829)
Qld: Dubois & LMA, 1972, p.209
Tas: AHC
syn. Strigeidae, unidentified
Tas: Munday & Green, 1972, p.3
. Anomotaenia accipitris TH. Johnston, 1913
Qld: THJ, 1913, p.82
Unidentified
Tas: Munday & Green, 1972, p.4
. Physaloptera sp.
SA: AHC (immature Qs)
Microtetrameres sp.
Tas, SA: PMM, 1977, p.257
A.
Synhimantus sp.
Tas: AHC
Serratospiculum tendo (Nitzsch in Giebel, 1857)
Qld: Mackerras, 1962, p,432
? loc: CIHUS
Serratospiculum gutlatum (Schneider, 1866)
NT: AHC
Tas: Munday & Green, 1972, p.12
Hamatospiculum sp.
Qld: THJ & PMM, 1941b, p.35
Microfilaria sp.
Qld: Cleland & THJ, 1912, p.430
Qld: Mackerras, 1962, p.436
Spirurida, unidentified
Tas: AHC (encysted larvae)
. Centrorhynchus asturinus (T.H. Johnston, 1913)
Qld: Southwell & Macfie, 1925, p.163
Qld: CIHUS
NSW: THJ & Deland, 1929, p.148
94. Accipiter noyaehollandiae (Gmelin)
Grey Goshawk
. Porrocaecum circinum T.H. Johnston & Mawson,
194\b
Qld: THJ & PMM, 1941b, p.34
Thelazia aquilina Baylis, 1934a
Qld: Baylis, 1934a, p.150
Qld: THJ & PMM, 1941b, p.34
syn. ‘nematode’
Qld: Cleland, 1922, p.107
Pracyrnea mansioni (Seurat, 1914)
syn. Cyrnea mansioni (Seurat, 1914)
Qld: PMM, 1968d, p.749
Procyrnea ? leptoptera (Rudolphi, 1819)
? Qld: BM(NH)
Synhimantus fieldingi (Baylis, 1934a)
syn, Acuaria fieldingi Baylis, 1934a
Qld: Baylis, 1934a, p.144
Centrorhynchus asturinus (T.H. Johnston, 1913)
Qld: THJ, 1918, p.215
Qld: Southwell & Macfie, 1925, p.163
syn, Gigantorhynchus asturinus TH. Johnston,
1913
Qld: THJ, 1912b, p.108
95. Haliaeetus leucogaster (Gmelin)
White-bellied Sea-Eagle
1/1: T
. Scaphanocephalus australis 5.J. Johnston, 1917
NSW: SJJ, 1917, p.188
syn. Scaphanocephalus sp.
? NSW: SJJ, 1914, p.243
Neodiplostomum spathula (Creplin, 1829)
SA: Dubois & LMA, 1972, p.209
, Capillaria sp.
Tas: AHC
Thelazia aquilina Baylis, 1934a
Qld: Baylis, 1934a, p.150
Synhimantus sp.
Tas: AHC
REC. S. AUST. MUS. 19(15): 219-325
96. Aquila audax (Latham)
Wedge-tailed Eagle
4/1 :N
. Neodiplostomum spathula (Creplin, 1829)
SA: Dubois, 1982, p.109
Strigea glandulosa Dubois, 1937a
SA: Dubois, 1982, p.109
. Thelazia aquilina Baylis, 19341
Qld: Baylis, 1934a, p.150
Microtetrameres paraccipiter Mawson, 1977
Tas: AHC
‘Eye worms’
Qld: Nicoll, 1914c, p.243
97. Hieraaetus morphnoides (Gould)
Little Eagle
1/0
. Porrocaecum circinum TH. Johnston & Mawson,
1941b
SA: THJ & PMM, 1941b, p.30
98. Circus assimilis Jardine & Selby
Spotted Harrier
W1l:NA
. Cladotaenia feuta Meggitt, 1933
Calcutta Zoo: Meggitt, 1933, p.16l
Cladotaenia sp.
Vie: AHC
. Porrocaecum circinum T.H. Johnston & Mawson,
1941b
SA: THJ & PMM, 1941b, p.30
Procyrnea mansioni (Seurat, 1914)
NT: AHC
Microtetrameres circi PMM, 1977
NT: PMM, 1977, p.255
. Unidentified
NT; specimen not kept
99. Circus approximans Peales
Swamp Harrier
4/4: T2 C3 N2
. Opisthorchis sp.
SA: AHC
Neodiplostomum spathula (Creplin, 1829)
SA: Dubois & LMA, 1972, p.209
Parastrigea repens (Chase, 1921)
SA: Dubois & LMA, 1972, p.198
Strigea glandulosa Dubois, 1937a
SA: Dubois & LMA, 1972, p.198
*. Cladotaenia sp.
Vic: AHC
Unidentified
Tas, SA: AHC
. Capillaria sp.
SA: AHC
Porrocaecum sp.
Tas: AHC
. Centrorhynchus asturinus (1.H. Johnston, 1913)
August, 1986
Qld: THJ, 1918, p.216
NSW, SA: AHC
Family FALCONIDAE
100, Falco subniger Gray
Black Faleon (endemic)
1/0
. Neodiplostomum spathula (Creplin, 1829)
SA: Dubois & LMA, 1972, p.209
Strigea glandulosa Dubois, 1937a
SA: Dubois & LMA, 1972, p.199
>, Cladotaenia sp.
SA: AHC
. Serratospiculuim tendo (Nitzsch in Giebel, 1857)
SA: AHC
101. Falco peregrinus Tunstall
Peregrine Falcon
3/3: TI Cl N3
. Neodiplostomum spathula (Creplin, 1829)
SA: Dubois & LMA, 1972, p.209
Neodiplostomum sp.
NSW: CIHUS
Diplostomidae, unidentified
SA: AHC
Strigeidae, unidentified
SA: AHC
. Unidentified
SA: AHC
. Contracaecum sp.
NSW: AHC
Microtetrameres raptoris Mawson, 1977
SA: PMM, 1977, p.255
Physaloptera sp.
SA: AHC
Serratospiculum guttatum (Schneider, 1866)
SA: Bain & PMM, 1981, p.275
Serratospiculum tendo (Nitzsch in Giebel, 1857)
SA: Bain & PMM, 1981, p.275
NSW: AHC
Vie: UMVS
syn, S. guttatum (Schneider, 1866)
SA, NT: THJ & PMM, 1941b, p.34
Spirurida, unidentified
SA: AHC (encysted larva)
102. Falco longipennis Swainson
Australian Hobby
7/7: N7 Al
. Physaloptera alata Baylis, 1925
Qld: Baylis, 1925, p.115
Procyrnea falco (Mawson, 1968d)
Tas, SA, NT: AHC
syn. Cyrnea falco Mawson, 1968d
NT; PMM, 1968d, p.752
Tas: Munday & Green, 1972, p.10
Tétrameres sp.
NT: AHC
Microtetrameres raptoris Mawson, 1977
io
HELMINTHS FROM AUSTRALIAN BIRDS 245
NT: PMM, 1977, p.255
Geopetitia falco Mawson, 1966
NT: PMM, 1966, p.717
Synhumantus falco Mawson, 1982
WA: AHC
Serratospiculum guttatum (Schneider, 1866)
SA, WA: TH] & PMM, 1941b, p.34
Tas: Munday & Green, 1972, p.12
SA: Bain & PMM, 1981, p.273
Vic, Tas, NT: AHC
syn. S. attenuatum (Rudolphi, 1819)
Qld: Baylis, 1925, p.112
Serratospiculum tendo (Nitzsch in Giebel, 1857)
Qld: BM(NH)
SA, NT: AHC
Splendidofilariinae, unidentified
NT: AHC
‘Filarial worms’
Tas: Cleland, 1922, p.107
Spiriirida, unidentified
NT: AHC (encysted larvae)
. Unidentified
NT: specimen not kept
103. Faleo hypoleucos Gould
Grey Falcon (endemic)
. Unidentified
SA: AHC
. Hamatospiculum sp., near H.quadridens (Molin,
1858)
Qld: CIHUS, BM(NH)
Serratospiculum guttatum (Schneider, 1866)
NSW; BM(NH)
Microfilaria sp.
Qld: Breinl, 1913a, p.34 (CIHUS)
104. Falco berigora Vigors & Horsfield
Brown Falcon
13/11: CL NII A6
Episthmium prosthovitellatum (Nicoll, 1914b)
Qld; CIHUS
syn. Echinochasmus prosthovilellatus Nicoll,
1914b
Qld: Nicoll, 1914b, p.115
Psilochasmus sp.
Qld: CHHUS
Opisthorchis obsequens Nicoll, 1914b
Qid: Nicoll, 1914b, p.106
Strigea glandulosa Dubois, 1937a
syn. Strigea falconis Dubois, 1937b (not Szidat,
1928)
Aust. (? Qld): Dubois, 1937b, pp.244,247
Strigea sp., indeterminable
Qld: CIHUS
. Unidentified
Tas, SA, NT: AHC
Tas: Cleland, 1922, p.105 (AHC)
Tas: Munday & Green, 1972, p.4
. Capillaria sp.
Qld: CIHUS
Porrocaecum angusticolle (Molin, 1860)
Qld: CIHUS
Procyrnea falco (Mawson, 1968d)
Tas, NT: AHC
syn. Cyrnea falco Mawson, 1968d
SA; PMM, 1968d, p.752
Procyrnea mansioni (Seurat, 1914)
syn. Cyrnea mansioni (Seurat, 1914)
NSW: PMM, 1968d, p.749
Procyrnea ? leptoptera (Rudolphi, 1819)
Qld: BM(NH)
Tetrameres sp,
NT: AHC
Microtetrameres raptoris Mawson, 1977
SA, NT: PMM, 1977, p.255
Synhimantus flindersi (T.H, Johnston & Mawson,
1941b)
syn. Acuaria flindersi T.H. Johnston & Mawson,
194 1b
Tas: THJ & PMM, 1941b, p.3l
syn. ‘nematode’
Tas: Cleland, 1922, p.105 (AHC)
Synhimantus falco Mawson, 1982
SA: PMM, 1982, p.24
Tas; AHC
Physaloptera hieracidiae T.H. Johnston & Mawson,
1941b
Tas: THJ & PMM, 1941b, p.31
Bancroftinema dentatum T.H.
Mawson, 1941b
Qld: THJ & PMM, 1941b, p.33
Thelazia aquilina Baylis, 1934a
Qld: Baylis, 1934a, p.150
Qid: CLHUS
Serratospiculum guttatum (Schneider, 1866)
syn. Filaria gutiata Schneider, 1866
SA: Schneider, 1866, p.80
Diplotriaena falconis (Connal, 1912)
SA: Bain & Mawson, 1981, p.283
‘Eye worms’
Qld: Nicoll, 1914c¢, p.244
Johnston &
. Centrorhynchus falconis (TH. Johnston & Best,
1943)
syn. Gordiorhynchus jalconis T.H. Johnston &
Best, 1943
NT: THJ & Best, 1943, p.229
Centrorhynchus asturinus (T.H. Johnston, 1913)
Qld: Southwell & Macfie, 1925, p.164
NSW: THJ & Deland, 1929, p.148
SA, NT: AHC
Qld: CIHUS
105, Falco cenchroides Vigors & Horsfield
Australian Kestrel
7/5: Cl Ad
C. Unidentified
SA: AHC
N. Procyrnea falco (Mawson, 1968d)
SA, NT; AHC
Procyrnea_ paraleptoptera (T.H.
Mawson, 1941b)
syn. Cyrnea paraleptaptera T.H. Johnston &
Mawson, 194]1b
Johnston &
246
io)
. Mediorhynchus alecturae (T.H.
REC. S$. AUST, MUS. 19(15); 219-325
Qld: THJ & PMM, 1941b, p.32
Microtetrameres raptoris Mawson, 1977
SA: PMM, 1977, p.255
Synhimantus falco Mawson, 1982
SA: PMM, 1982, p.24
. Centrorhynchus asturinus (TH. Johnston, 1913)
Qld, SA: AHC
Family MEGAPODITDAE
106. Leipoa ocellata Gould
Mallee Fow! (endemic)
2/2: C2 N2
. Raillietina leipoae T.H. Johnston & Clark, 1948b
SA: THJ & Clark, 1948b, p.88
Raillietina sp.
SA: THJ & Clark, 1948b, p.90
. Leipoanema ellisi THJ & PMM, 1942e
SA: THJ & PMM, 1942e, p.73
107. Alectura lathami Gray
Australian Brush Turkey (endemic)
. Davainea sp.
Qld: AHC
Raillietina sp.
SA (cage): AHC
Unidentified
Qld: AHC
. Capillaria sp.
Qld: AHC
Syngamus sp.
Qld: AHC
Odontoterakis bancrofti (T.H. Johnston, 1912a)
Qld: AHC
syn. Heterakis bancrofti T.H. Johnston, 1912a
Qld: THJ, 1912a, p.72
Ascaridia catheturina (T.H. Johnston, 1912a)
Qld: THJ & PMM, 1942a, p.93
syn. Heterakis catheturinus T.H. Johnston, 1912a
Qld: THJ, 1912a, p.73
Gongylonema alecturae T.H. Johnston & Mawson,
1942a
Qld: THJ & PMM, 1942a, p.92
Lissonema sp.
Qld: AHC
Johnston &
Edmonds, 1947)
syn. Empodius alecturae T.H. Johnston &
Edmonds, 1947
Qld: THJ & Edmonds, 1947, p.557
syn. Echinorhyachus sp.
Qld: THJ, 1912b, p.106
. Oncicola pomatostomi (T.H.
August, 1986
Family PHASIANIDAE
108. Coturnix novaezelandiae Quoy &
Gaimard
Stubble Quail (endemic)
SALE
. Unidentified
NSW: AM
SA: AHC
109. Coturnix ypsilophora Bosc
Swamp Quail
10/9 : TI N8&
Echinostomatidae [spines missing]
Tas: discarded
. Metroliasthes sp.
Vic: AHC
. Teltrameres sp.
Tas: AHC
Cyrnea colini Cram, 1927
Tas: AHC
Cheilospirura gruveli (Gendre, 1913)
Tas: PMM, 1982, p.23
. Unidentified
SA (cage): AHC (cyst)
Family TURNICIDAE
110. Turnix castanota (Gould)
Chestnut-backed Button-quail (endemic)
2/2: Tl A2
. Skrjabinosomum mawsoni Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.124
Johnston &
Cleland, 1912)
SA: AHC (cyst)
lll. Turnix velox (Gould)
Little Button-quail (endemic)
4/1: N
. Acuariidae, unidentified
NT: AHC
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Oligacanthorhynchus pomatostomi
(T.H. Johnston & Cleland, 1912)
Qld: Hall Exped. Report, 1974, p.35 (cyst)
syn.
112. Turnix pyrrhothorax (Gould)
Red-chested Button-quail (endemic)
. Capillaria sp.
SA (cage): AHC
HELMINTHS FROM AUSTRALIAN BIRDS 247
Family PEDIONOMIDAE
113. Pedionomus torquatus Gould
Plains Wanderer (endemic)
VWL:A
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: THJ & Deland, 1929, p.148 (cyst)
Family RALLIDAE
114. Rallus philippensis Linné
Buff-banded Rail
4/1: T
T. Leucochloridium sp.
SA: AHC
A. Unidentified
Qld: AHC (part of worm only)
115. Rallus pectoralis Temminck
Lewin’s Rail
3/3: Tl N2
T. Unidentified
Tas: specimen not kept
N. Capillaria sp.
Tas: Munday & Green, 1972, p.6
Tas: AHC
116. Por :ana pusilla (Pallas)
*aillon’s Crake
3/3: Tl N2
T. ? Catatropis sp.
SA; AHC
Echinostoma sp. [spines missing]
SA: AHC
Echinostomatinae [spines (45(4-5))]
SA: AHC (immature)
N. Capillaria sp.
SA: AHC
Tetrameres globosa (Linstow, 1879)
SA: PMM, 1979, p.180
117. Porzana fluminea Gould
Australian Crake (endemic)
14/13 : T3 N13
T. ? Catatropis sp.
SA: AHC
Cyclocoelidae, unidentified
SA: AHC (larvae)
Echinostomatinae [spines missing]
SA: discarded (immature)
N. Capillaria sp.
SA: AHC
Porrocaecum ensicaudatum (Zeder, 1800)
SA: AHC
Tetrameres globosa (Linstow, 1879)
SA, NT: PMM, 1979, p.180
Acuariidae, unidentified
SA: not kept (larva)
118. Porzana tabuensis (Gmelin)
Spotless Crake
2/2: N
. Tetrameres globosa (Linstow, 1879)
SA: PMM, 1979, p.180
119. Gallinula mortierii (Du Bus)
Tasmanian Native-hen (endemic)
5/3: N
. Capillaria sp.
Tas: AHC
Tetrameres globosa (Linstow, 1879)
Tas: AHC
syn. 7eframeres sp.
Tas; Munday & Green, 1972, p.1l
120. Gallinula ventralis Gould
Black-tailed Native-hen (endemic)
7/4: T2 N2
. ? Catatropis sp.
SA: AHC
? Echinostomam hilliferum Nicoll, 1914b
Qld: AHC
Echinostomatidae, unidentified
SA: believed lost (immature)
. Unidentified
Qld: AHC
. Amidostomum tribonyx Mawson, 1980
SA: PMM, 1980, p.10
121. Gallinula tenebrosa Gould
Dusky Moorhen
12/7 : T7 C2 N2
. Catatropis gallinulae T.H. Johnston, 1928
SA: THJ, 1928, p.135
? Paramonostomum sp.
NSW: AHC
Echinostoma australe TH. Johnston, 1928
SA: THJ, 1928, p.138
Echinostoma bancrofti T.H. Johnston, 1928
Qld: THJ, 1928, p.140
? Echinostoma revolutum (Froelich, 1802)
SA: discarded
Echinostoma sp. [spines (37(5))]
Qld, SA: AHC
Unidentitied
SA: AHC (immature)
. Unidentified
Qld, SA: AHC
. Capillaria sp,
SA: AHC
Amidostomum sp.
NSW: WLRR
248
REC. 8, AUST. MUS. 19(15): 219-325
122. Porphyrio porphyrio (Linné)
Purple Swamphen
18/5: T4 NI
Echinostoma hilliferum Nicoll, 1914b
Qld: Nicoll, 1914b, p.112
Qld, SA: AHC
Cyclocoelum sp.
NSW: CIHUS
Echinostoma sp. [spines (35(5))]
SA: AHC
Echinostomatinae [spines 35(5))|
SA: AHC
Echinostomatinae [spines ? (27(4))]
SA: AHC
Distoma sp.
NSW: Krefft, 1873, p.213
. Unidentified
SA: AHC
. Capillaria sp.
? loc: AHC
Tetrameres globosa (Linstow, 1879)
SA: AHC
123. Fulica atra Linné
Eurasian Coot
3/3: Tl N2
Echinostoma hilliferum Nicoll, 1914b
Qld: Nicoll, 1914b, p.112
‘Echinostome larvae’
NSW: Bradley, 1926, p.575; 1927, p.675
Psilochasmus oxyurus (Creplin, 1825)
Tas: S.J. Smith, 1981, p.181
Atriophallophorus coxiellae S.J. Smith, 1974
Tas: S.J. Smith, 1981, p.181
. Hymenolepis sp.
Qld: AHC
Unidentified
NSW: Bradley, 1927, p.675
Qld, SA: AHC
. Amidostomum fulica (Rudolphi, 1819)
Vic: UMVS
SA: AHC
Pelecitus fulicaeatrae (Diesing, 1861)
Vic: UMVS
Unidentified
NSW: Bradley, 1927, p.675 (non-bursate)
Family GRUIDAE
124. Grus rubicundus (Perry)
Brolga
Allopyge antigones S.J. Johnston, 1913
Qld: SJJ, 1913, p.375
syn. ‘monostome’
Qld: Nicoll, 1914c, p.244
Echinostoma australasianum Nicoll, 1914a
Qld: Nicoll, 1914a, p.337
Prosthogonimus sp.
Qld: AHC
August, 1986
N. Porrocaecum serpentulum (Rudolphi, 1809)
syn. Ascaris serpentula Rudolphi, 1809
Germany (Mus.): Linstow, 1899, p.7
Ascaridia stroma (Linstow, 1899)
Qld: UQDP
125. Grus antigone (Linné)
Sarus Crane
. Ascaridia stroma (Linstow, 1899)
NSW (cage): AHC
Family OTIDIDAE
126. Ardeotis australis (Gray)
Australian Bustard (endemic)
1W1:c
. Philophthalmus sp.
NT: AHC
. Idiogenes sp.
Vic (cage): AHC
Ascometra sp.
Vic (cage): AHC
Unidentified
Qld, Vic (cage), SA (cage), NT: AHC
Vic (cage): UMVS
. Hartertia sp.
WA: AHC
Excisa dentifera (T.H. Johnston & Mawson, 1941d)
syn. Cyrnea dentifera T.H. Johnston & Mawson,
1941d
NT: THJ & PMM, 1941d, p.255
Family JACANIDAE
127. Irediparra gallinacea (Temminck)
Comb-crested Jacana
1/1:T
. ? Echinostomatidae, unidentifiable
NT: AHC (fragment)
Family BURHINIDAE
128. Burhinus magnirostris (Latham)
Bush Thick-knee
. Notocotylus atrenuatus (Rudolphi, 1809)
Qld: Nicoll, 1914b, p.125
Echinostoma ignavum Nicoll, 1914b
Qld: CIHUS
Platynotrema biliosum Nicoll, 1914b
Qld: Nicoll, 1914b, p.118
Platynotrema jecoris Nicoll, 1914b
Qld: Nicoll, 1914b, p.120
Strigea nicolli Dubois, 1937a
Qld: Dubois, 1937b, p.241
C, Angularella australis (Maplestone, 1921b)*
*Yamaguti, 1959, p.235, notes that this species cannot
be referred to Angularella (syn. Angularia) and
probably belongs to some other genus.
HELMINTHS FROM AUSTRALIAN BIRDS
syn. Angularia australis Maplestone, 1921b
Qld: Maplestone, 1921b, p,407
Choanotaenia sp.
Aust: BM(NH)
Paricterotaenia sp.
Aust: BM(NH)
Unidentified
Qld: AHC
Family HAEMATOPODIDAE
129. Haematopus longirostris Vieillot
Pied Oystercatcher
. Unidentified
Qld, SA: AHC
. Prosthorhynchus sp.
Tas: AHC
130. Haematopus fuliginosus Gould
Sooty Oystercatcher (endemic)
l/l:CNA
. Unidentified
SA: AHC
. Sciadocara sp.
SA: AHC
. Filicollis sphaerocephalus (Bremser, 1819)
Tas, SA: AHC
Family CHARADRIIDAE
131. Vanellus miles (Boddaert)
Masked Lapwing
11/11 : C7 N10 A2
. Notocotylus attenuatus (Rudolphi, 1809)
Qld: Nicoll, 1914b, p.125
Cyclocoelum obscurum (Leidy, 1887)
Qld: Hall Exped. Report, 1974, p.354
Haematrotrephus consimilis Nicoll, 1914b
Qld: Nicoll, 1914b, p.124
Plagiorchiidae, unidentified
syn. Lepodermatidae, unidentified
Qld: Nicoll, 1914b, p.106
Echinostoma ignavum Nicoll, 1914b
Qld: Nicoll, 1914b, p.114
Echinostoma sp. [spines ? (35(5))]
Tas: AHC
~. Angularella sp.
SA: AHC
Chounotaenia southwelli Fuhrmann, 1932
syn. Monopylidium macracanthum of
Maplestone & Southwell, 1923, not
Fubrmann, 1907
Qld: Maplestone & Southwell, 1923, p.322
Qld: CIHUS
Lapwingia adelaidae Schmidt, 1972
Qld: Schmidt, 1972, p.1085
Gyrocoelia australiensis Y.H. Johnston, 1910
Qld: Maplestone & Southwell, 1922a, p.61
Qld, SA: Schmidt, 1972, p.1089
syn. Gyrocoelia sp.
249
Qld: THJ, 1914a, p.108
Gyrocoelia sp.
NSW: CIHUS
Infula burhini Burt, 1939
Qld, SA: Schmidt, 1972, p.1038
syn. ‘Shipleva lobivanellus Maplestone‘
Qld: CIHUS, AHC
Unidentified
? NSW: Bradley, 1927, p.675
Tas: Munday & Green, 1972, p.6
Qld, Tas, SA: AHC
. Capillaria triloba Linstow, 1875
SA: PMM, 1968b, p.279
Tas: Munday & Green, 1972, p.6
Porrocaecum lobibicis Mawson, 1968b
SA: PMM, 1968b, p.284
Tas: Munday & Green, 1972, p.9
Tas, Vic: AHC
Tetrameres lobibicis Mawson, 1968b
SA: PMM, 1968b, p.300
Tas: AHC
Tetrameres sp.
Tas: Munday & Green, 1972, p.ll
Streptocara crassicauda (Creplin, 1829)
Tas: Munday & Green, 1972, p.12
Streptocara sp.
Tas: AHC
Pseudaproctella ? sp.
Qld: PMM, 1968b, p.302
Microfilaria sp.
Qld: Mackerras, 1962, p.436
. Prosthorhynchus sp.
Tas: AHC
132. Vanellus tricolor (Vieillot)
Banded Lapwing (endemic)
6/4: C3 NI Al
. Paricterotaenia zoniferae (T.H. Johnston, 1912c)
syn. Choanotaenia zoniferae T.H. Johnston,
1912c
NSW: THJ, 1912c, p.212
Unidentified
SA: AHC
. Unidentified
NT: specimen lost
. Unidentified
SA: specimen lost
133. Pluvialis dominica (P.L.S. Miller)
Lesser Golden Plover
Acanthoparyphium spinulosum S.J. Johnston,
1917
NSW: SJJ, 1917, p.198
Tas: AHC
syn, Echinostonia sp.
NSW: THI, 1910, p.93
Echinostomatidae, unidentified
Tas: Munday & Green, 1972, p.3
Levinseniella howensis S.J. Johnston, 1917
NSW: SJJ, 1917, p.221
NSW: Pearson & Deblock, 1979, p.32
250
a
REC. S. AUST. MUS, 19(15): 219-325
Levinseniella microovata Belopolskaya, 1958
Qld; Deblock & Pearson. 1970. bp, 777
Microphallus papillornatus Deblock & Pearson,
1969
Qld: Deblock & Pearson, 1969, 7.396
Microphallus vaginosus Deblock & Pearson, 1969
Qld: Deblock & Pearson, 1969, p.403
. Unidentified
NSW: AHC
134, Erythrogonys cinctus Gould
Red-kneed Dotterel
10/10 : T6 C10 Ni
Notocotylidae, unidentified
SA: AHC (immature)
Echinostomatidae [spines ? (35-37(5))]
SA: AHC (immature)
Echinostomatidae [spines ? (33(4))]
SA: AHC (immature)
Unidentified
SA: AHC (immature)
. Unidentified
SA: AHC
. Acuaridae, unidentified
SA: AHC (larva)
135. Charadrius rubricollis Gmelin
Hooded Plover (endemic)
2/2 : N2 A2
- Atriophallophorus coxiellae S.J. Smith, 1974
Tas: S.J. Smith, 1974, p.200 (immature)
Levinseniella tasmaniae (S.J. Smith, 1974)
syn. Microphallus tasmaniaé S.J. Smith, 1974
Tas: S.J. Smith, 1974, p.203
Maritrema calvertense §.J. Smith, 1974
Tas: 3.J. Smith, 1974, p,201
. Stellocaronema charadrii Mawson, 1968b
SA: PMM, 1968b, p.294
Skrjabinoclava horrida (Rudolphi, 1809)
SA: PMM, 1968b, p,292
. Plagiorhynchus charadrii (Yamaguti, 1939b)
syn, Prosthorhynchus charadrti Yamaguli, 1939b
SA: THJ & SJE, 1947, p.561
136. Charadrius mongolus Pallas
Mongolian Plover (migratory)
. Endocotyle incana Belopolskaya, 1952
Qld: Deblock & Pearson, 1968b, p.463
Gynaecolyla_ brisbanensis Deblock & Pearson,
1968a
Qld: Deblock & Pearson, 1968a, p.141
Levinseniella microovata Belopolskaya, 1958
Qld: Deblock & Pearson, 1970, p.777
Levinseniella monodactyla Deblock & Pearson,
1970
Qld: Deblock & Pearson, 1970, p.774
Maritrema eroliae Yamaguti, 1939a
Qld: Deblock & Pearson, 1968b, p.458
Microphallus papillornatus Deblock & Pearson,
1969
Cc,
August, 1986
Qld; Deblock & Pearson, 1969, p.396
Microphallus sp.
Qld: Deblock & Pearson, 1969, p.400
. Unidentified
Tas, SA; AHC
137. Charadrius bicinctus Jardine & Selby
Double-banded Plover (migratory)
2/2: Cl N2
. Unidentified
SA: AHC
. Capillaria sp.
SA; AHC
Streptovara crassicauda (Creplin, 1829)
SA: AHC
138, Charadrius leschenaultii Lesson
Large Sand Plover (migratory)
Acanthoparyphium marilae Yamaguti, 1934
Qld; CIHUS
Acanthoparyphium spinulosum §,J, Johnston, 1917
Qld: Bearup, 1960, p.219
Maritrema sp.
Qld: CIHUS
. Unidentified
Qld: CIHUS
139. Charadrius ruficapillus Temminck
Red-capped Plover (endemic)
17/14 : T2 C10 N10 AG
. Acanthoparyphium spinulosum §.J, Johnston,
1917
SA: AHC
Atriophallophorus coxiellae §.J. Smith, 1974
Tas: S.J. Smith, 1974, p.200 (immature)
Levinseniella tasmaniae (S.J. Smith, 1974)
syn, Microphallus tasmaniae S.J. Smith, 1974
Tas: S.J. Smith, 1974, p,203
Microphallidae, unidentified
Tas: Munday & Green, 1972, p.3
SA: AHC
Unidentified
Tas: Munday & Green, 1972, p.4
Tas, SA: AHC
. Capillaria sp.
SA: AHC
Tetrameres nouveli (Seurat, 1914)
SA: PMM, 1968b, p.297
Tetrameres sp.
Tas: Munday & Green, 1972, p.11
Echinuria heterobrachiata Wehr, 1937
SA: PMM, 1968b, p.289
Skrjabinoclava horrida (Rudolphi, 1809)
SA: PMM, 1968b, p,292
Tas: AHC
syn. Skrjabinoclava sp.
Tas: Munday & Green, 1972, p.12
Streptocata crassicauda (Creplin, 1829)
SA: PMM, 1968b, p.293
Tas: AHC
HELMINTHS FROM AUSTRALIAN BIRDS
syn. Streptocara sp.
Tas: Munday & Green, 1972, p.12
Spirurida, unidentified
SA: AHC
. Plagiorhynchus charadrii (Yamaguti, 1939b)
Tas, SA: AHC
140. Charadrius melanops Vieillot
Black-fronted Plover (endemic)
12/7 : T3 C3 NI
. Echinostomatidae [spines (37(5))]
SA: AHC (immature)
Echinostomatidae [spines ? (41(5))]
SA: AHC (immature)
Levinseniella tasmaniae (S.J. Smith, 1974)
syn. Microphallus tasmaniae $.J. Smith, 1974
Tas: S.J. Smith, 1974, p.203
Maritrema calvertense S.J. Smith, 1974
Tas: S.J. Smith, 1974, p.202
. Unidentified
SA: AHC
. Chevreuxia australis T.H. Johnston & Mawson,
194]e
SA: PMM, 1968b, p.292
141. Peltohyas australis Gould
Inland Dotterel (endemic)
. Unidentified
SA: AHC
. Unidentified
Aust: AHC (fragments)
Family RECURVIROSTRIDAE
142. Himantopus leucocephalus (Gould)
White-headed Stilt
1/1:CN
Trichobilharzia sp.
NT: CIHUS
Aust: Bearup & Langsford, 1966, p.523
Bilharziellinae, unidentified
NT: AHC
Haematotrephus adelphus $.J. Johnston, 1917
SA: SJJ, 1917, p.241
SA, NT: AHC
syn. Monostomum sp.
SA: THJ, 1910, p.94
Unidentified
Qld: CIHUS
. Davainea himantopodis T.H. Johnston, 1911
syn. Davainea sp.
SA: THJ, 1910, p.95
SA: AHC
Diorchis flavescens (Krefft, 1873)
NSW: AHC
Hymenolepis sp.
SA: THJ, 1910, p.95
Acoleus hedleyi T.H. Johnston, 1910
SA: THJ, 1912d
nw
Ww
syn. Tuenia rugosa Krefft, 1873
NSW: Krefft, 1873, p.223
Gyrocoelia australiensis (T.H. Johnston, 1910)
NSW: THJ, 1912d, p.28
syn. Dilepis australiensis T.H. Johnston, 1910
NSW: TH], 1910, p.95
syn, Tuenia coronata Krefft, 1873
NSW: Krefft, 1873, p.220
Infula burhini Burt, 1939
‘Aust’; Voge & Read, 1954, p.483
Unidentified
SA, NT: AHC
Qld: CIHUS
. Capillaria triloba Linstow, 1875
SA: PMM, 1968b, p.279
Amidostomum acutum (Lundahl, 1848)
NT: PMM, 1980, p.9
syn. A. chevreuxi Seurat, 1918
SA: PMM, 1968b, p.283
Contracaecum sp.
SA: PMM, 1968b, p.287
Tetrameres nouveli (Seurat, 1914)
SA: PMM, 1968b, p.297
NT: AHC
? Chevreuxia revoluta (Rudolphi, 1819)
NT: AHC
143. Cladorhynchus leucocephalus
(Vieillot)
Banded Stilt (endemic)
7/7: TS C4 N6
. Bilharziellinae, unidentified
SA: AHC
Notocotylidae, unidentified
SA: AHC
Haematotrephus adelphus S.J. Johnston, 1917
SA: AHC
?Omphalometridae, unidentified
SA: AHC
Acanthoparyphium sp. [spines (23(0))]
SA: AHC
’. Davainea sp,
SA (cage): AHC
Diorchis flavescens (Krefft, 1873)
NSW: AHC
Gyrocoelia sp.
SA: BM(NH)
Unidentified
SA: AHC (immature)
. Tetrameres cladorhynchi Mawson, 1968b
SA: PMM, 1968b, p.299
Echinuria heterobrachiata Wehr, 1937
SA: PMM, 1968b, p.289
144, Recurvirostra novaehollandiae
Vieillot
Red-necked Avocet (endemic)
4/4:T1 C4 N3
T. Notocotylidae, unidentified
SA: AHC
252 REC. 5, AUST, MUS, 19(15): 219-325 August, 1986
Cyclocoelidae, unidentified
WA: AHC
C. Hymenolepis sp.
SA: AHC
Davainia sp.
SA: AHC
Himantocestus sp.
SA: AHC
Unidentified
SA, WA: AHC
N. Capillaria triloba Linstow, 1875
SA: PMM, 1968b, p.279
Capillaria recurvirostrae Mawson, 1968b
SA: PMM, 1968b, p.280
Tetrameres nouveli (Seurat, 1914)
SA: PMM, 1968b, p.297
Chevreuxia revoluta (Rudolphi, 1819)
SA: AHC
Echinuria heterobrachiata Wehr, 1937
SA: AHC
Family SCOLOPACIDAE
145. Numenius madagascariensis (Linné)
(Linne)
Eastern Curlew (migratory)
T. AHimasthla harrisoni 8.J. Johnston, 1917
Qld: SJJ, 1917, p.195
syn. Echinostoma sp.
Qld: THJ, 1912b, p.107
syn. Echinostoma (Acanthochasmus) sp.
Qld: THJ, 1916, p.46; THJ, 1918, p.211
C. Unidentified
Qld, NSW, SA: AHC
A, Arhythmorhynchus johnstoni Golvan, 1960
syn. Arhythmorhynchus frassoni of THJ & SJE,
1951, not Molin, 1858
Qld: THJ & SJE, 1951, p.3
syn. Echinorhynchus sp.
Qld: THJ, 1912b, p.107; 1914a, p.110
Arhythmorhynchus sp.
Qld: CIHUS
146. Tringa glareola (Linné)
Wood Sandpiper (migratory)
3/3: Cl N3
T. Cyclocoelidae, unidentified
NSW: AHC
C. Unidentified
SA, NT; AHC
N. Tetrameres scolopacidis Mawson, 1968b
SA: PMM, 1968b, p.381
NT: AHC
147. Tringa brevipes (Vieillot)
Grey-tailed Tattler (migratory)
1/1, N
N. Skrjabinoclava sp,
WA: WAM
148. Tringa hypoleucos Linné
Common Sandpiper (migratory)
. Echinoparyphium sp.
? loc: CIHUS
Echinostoma sp.
? loc: CIHUS
Echinostomatidae [spines missing]
SA: discarded (very young)
Platynotrema sp.
Qld: CIHUS
Maritrema sp.
Qld: CIHUS
. Haploparaxis australis (T.H. Johnston, 1913)
Old: CIHUS
Unidentified
Qld: CIHUS
149. Tringa nebularia (Gunnerus)
Greenshank (migratory)
5/4: T1 C3 NI
. Haematotrephus sp.
SA: AHC
. Unidentified
SA: AHC
. Unidentified
SA: believed lost
150. Tringa stagnatilis (Bechstein)
Marsh Sandpiper (migratory)
2/2: C2 NI
. Paruterina rauschi Freeman, 1957
SA: AHC
N. Unidentified
SA: AHC (immature)
151. Tringa terek (Latham)
Terek Sandpiper (migratory)
W1:iNA
. Skrjabinoclava decorata (Solonitzin, 1928)
SA: AHC
Schistorophus cornutus Sobolev, 1943
SA: AHC
Viktorocara limosae Mawson, 1968b
SA: AHC
Viktorocara schejkini Gushanskaya, 1950
SA: AHC
. Unidentified
SA: specimen not kept
152. Gallinago hardwickii (Gray)
Latham’s Snipe (migratory)
2/2: C
. Haploparaxis australis (T.H. Johnston, 1913)
syn. Aploparaksis australis TH. Johnston, 1913
Qld: THJ, 1913, p.90
Unidentified
SA: Cleland, 1922, p.105
Qld: AHC
HELMINTHS FROM AUSTRALIAN BIRDS 253
153. Gallinago megala Swinhoe
Swinhoe’s Snipe (migratory)
C. Haploparaxis australis (T.H. Johnston, 1913)
Qld: AHC, CIHUS
Unidentified
Qld: AHC
154. Limosa lapponica (Linné)
Bar-tailed Godwit (migratory)
1/1:CN
T. Cycloceelum taxorchis S.J, Johnston, 1917
NSW: SJJ, 1917, p.239
Cyclocoelum sp.
Qld: CIHUS
Acanthoparyphium marilae Yamaguti, 1934
Qld: Bearup, 1960, p.219
Acanthoparyphium squatarolae Yamaguti, 1934
Qld: Bearup, 1960, p.219
Himasthla ? megacotyla Yamaguti, 1939a
Qld: CIHUS
Platynotrema sp.
Qld: CIHUS
Maritrema sp.
Qld: CIHUS
Heterophyidae, unidentified
Qld: CIHUS
C. Unidentified
NSW, SA: AHC
Qld: CIHUS
N. Streptocara sp.
SA: AHC
Schistorophus limosae Mawson, 1968b
Qld; Mawson, 1968b, p.287
Kiktorocara limosae Mawson, 196&b
Qld: PMM, 1968b, p.287
A. Arhythmorhynchus limosae Edmonds, 1971
Qld: Edmonds, 1971, p.58
Unidentified
Qld: CIHUS
155. Calidris canutus (Linné)
Red Knot (migratory)
8/7: T7 N7
T. Cyclocoelidae, unidentified
NT: AHC
Acanthoparyphium marilae Yamaguti, ‘1934
Qld: Bearup, 1960, p.219
Cloeophora sp. [spines ? (21)]
NT: AHC
Himasthlinae [spines ? (31(4))]
NT: AHC
N. Tetrameres calidris Mawson, 1968b
NT: PMM, 1968b, p.300
Sciadiocara umbellifera (Molin, 1860)
NT: PMM, 1968b, p.285
Schistorophus longicornis (Hemprich & Ehrenberg,
in Schneider, 1866)
NT: PMM, 1968b, p.285
Viktorocara sp.
NT: PMM, 1968b, p.289
156. Calidris tenuirostris (Horsfield)
Great Knot (migratory)
T. Acanthoparyphium marilae Yamaguti, 1934
Qld: Bearup, 1960, p.219
Acantheparyphium squatarolae Yamaguti, 1934
Qld: Bearup, 1960, p.219
Himasthla kusasigi Yamaguti, 1939a
Qld: CIHUS
Unidentified
Qld: CIHUS (? from pancreas)
157. Calidris acuminata (Horsfield)
Sharp-tailed Sandpiper (migratory)
32/15 ; T4 C12 N7
T. ? Notocotylidae, unidentified
SA: AHC (immature)
Echinostomatinae [spines (45(5))]
SA: AHC (immature)
Echinostomatidae [spines ? (37)]
SA: AHC (immature)
Endocotyle incana Belopolskaya, 1952
Qid: Deblock & Pearson, 1968b, p.463
? Microphallinae, unidentified
SA: believed lost
Unidentified
SA: AHC (immature)
Unidentified
SA: AHC (cysts)
C. Trichocephaloides sp.
SA: AHC
Unidentified
NSW: Cleland, 1922, p.10S (AHC)
NSW, SA: AHC
N. Tetrameres scolopacidis Mawson, 1968b
SA: PMM, 1968b, p.301
188. Calidris ruficollis (Pallas)
Red-necked Stint (migratory)
29/28 : C23 NIS Al
T. Microphallidae, unidentified
Tas: AHC
? Heterophyidae, unidentified
Tas: AHC
Trematoda, unidentified
Tas: AHC (cyst)
C. Unidentified
Tas, SA: AHC
N. Stellocaronema glarealae Mawson, 1968b
SA: AHC
Tetrameres scolopacidis Mawson, 1968b
SA: PMM, 1968b, p.301
Tas: AHC
Tetrameres sp.
Tas: AHC
Echinuria heterobrachiata Wehr, 1937
SA: PMM, 1968b, p.289
Skrjabinoclava sp.
SA: AHC
A. Unidentified
Tas: AHC
REC, &. AUST. MUS, 19(15): 219-325
159. Calidris ferruginea (Pontoppidan)
Curlew Sandpiper (migratory)
3/1 :C
C. Unidentified
SA: AHC
160. Calidris alba (Pallas)
Sanderling (migratory)
1/1: C
C. Unidentified
SA: AHC
161. Limicola falcinellus (Pontoppidan)
Broad-billed Sandpiper (migratory)
1/0
No helminths recorded
Family GLAREOLIDAE
162. Stiltia isabella Vieillot
Australian Pratincole (migratory)
2/2: C2 NI
. Unidentified
SA, NT: AHC
. Stellocaronema glareolae Mawson, 1968b
SA: PMM, 1968b, p.296
Family LARIDAE
163. Larus novaehollandiae Stephens
Silver Guli
36/36 : T22 C27 N& AY
T. Austrobilharzia terrigalensis S.J. Johnston, 1917
Qld: Rohde, 1977, p.39
NSW: SJJ, 1917, p.235
NSW: Bearup, 1956, p.471
SA: THJ, 1941, p.282
WA: Appleton, 1983, p.249
Vic, SA: AHC
Gigantobilharzia sp.
Qld: Rohde, 1978, p.40
Schistosomatidae, unidentified
NSW: CIHUS
Cloacitrema narrabeenense Howell & Bearup, 1967
NSW: Howell & Bearup, 1967, p.182
Parorchis acanthus (Nicoll, 1906)
SA: LMA, 1954, p,173
NSW: CIHUS
Philophthalmus burrili Howell & Bearup, 1967
NSW; Howell & Bearup, 1967, p.182
Acanthoparyphium spinulosum S.J. Johnston, 1917
NSW: Bearup, 1960, p.219 (exptl)
? Echinoparyphium sp. [spines (35(5))]
SA: AHC
Echinostoma sp. [spines missing]
SA: AHC
Stephanoprora sp.
Tas: AHC
Aueust, 1986
Echinostomatinae [spines (33(5))]
SA; AHC
Echinostomatinae [spines (37(4-5))]
SA: AHC
Prosthogonimus sp.
NSW: CIHUS
Gynoecotyla brisbanensis Deblock & Pearson,
1968a
Qld: Deblock & Pearson, 1968a, p.141
Levinseniella sp.
NSW: CIHUS
Microphallus papillornatus Deblock & Pearson,
1969
Qld: Deblock & Pearson, 1969, p.396
Pachytrema calculus Looss, 1907
NSW, SA: LMA, 1971, p.105
Vic: AHC
Galactosomum angelae Pearson, 1973
SA: Pearson, 1973, p.364
Galactosomum bearupi Pearson, 1973
Qld: Pearson, 1973, p.370
Galactosomum ussuriense Oshmarin, 1963
Qld: Pearson, 1973, p.429
Heterotestophyes sp.
Qld: CIHUS
Stictodora diplacantha T.H. Johnston, 1942a
SA: AHC
Stictodora lari Yamaguti, 1939a
NSW: Bearup, 1961, p.251 (exptl & natural)
syn. Sfictodora sp.
NSW: Bearup, 1958, p.219 (exptl)
Diplostomum spathaceum (Rudolphi, 1819)
syn. Diplostomum murrayense (T.H. Johnston &
Cleland, 1938)
SA: THJ & LMA, 194la, p.140
SA: Dubois & LMA, 1972, p.208
Diplostomum sp.
Qld: CIHUS
Neodiplostomum pricei Krull, 1934
Washington DC, USA (cage): Krull, 1934,
p.353
Cardiocephaloides hillit (S.J. Johnston, 1904)
SA: Dubois & LMA, 1972, p.201
Tas: AHC
syn. Holostomum hillii S.J. Johnston, 1904
NSW: SJJ, 1904, p.11]
‘. Tetrabothrius sp.
NSW: CIHUS
Anomotaenia hydrochelidonis Dubinina, 1954
WA: BM(NH)
Unidentified
NSW, ‘Tas, SA: AHC
Vic: UMVS
. Capillaria jaenschi TH. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p.245
Capillaria laricola Wassilikova & Gushanskaya,
1930
SA: THJ & PMM, 1947, p.549
Capillaria thomascameroni Mawson, 1969
SA: PMM, 1969, p.1104
Tetrameres sp.
Tas, SA: AHC
HELMINTHS FROM AUSTRALIAN BIRDS 255
Pectinospirura argentata Wehr, 1933
Tas: AHC
Eufilaria sp.
SA: PMM, 1969, p.1110
Filarial worm, unidentifiable
Tas: AHC
Microfilaria sp.
London Zoo: Scott, 1926, p.237
. Filicollis sphaerocephalus (Bremser, 1819)
SA: THJ & SJE, 1947, p.255
164. Chlidonias hybrida (Pallas)
Whiskered Tern
20/18 : T13 C7 N8&
. 2 Ornithobilharzia sp.
SA: AHC
Echinostomatidae, [spines (41-43(5))]
SA: AHC (metacercariae)
Prosthogonimus vitellatus Nicoll, 1914a
SA: LMA, 1973, p.859
Pachytrema calculus Looss, 1907
SA: LMA, 1971, p.105
Diplostomum spathaceum (Rudolphi, 1819)
syn. Diplostomum murrayense (T.H. Johnston &
Cleland, 1938)
SA: THJ & LMA, 194la, p.140
Cardiocephaloides musculosus (S.J. Johnston,
1904)
SA: Dubois & LMA, 1972, p.202
. Unidentified
SA: AHC
. Capillaria jaenschi TJH. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p.245
Contracaecum sp.
SA: THJ & PMM, 1947, p.551
Tetrameres sp.
SA: AHC
Chevreuxia australis T.H. Johnston & Mawson,
194le
SA: THJ & PMM, 194le, p.259
Cosmocephalus jaenschi T.H. Johnston & Mawson,
194le
SA: AHC
Acuaria (s.1.) sp.
SA: THJ & PMM, 194le, p.260
Streptocara recta (Linstow, 1879)
SA: THJ & PMM, 1942e, p.71
Spirurida, unidentified
SA: AHC (larva)
165, Gelochelidon nilotica (Gmelin)
Gull-billed Tern
1/1: N
. Habronematinae, unidentified
SA: PMM, 1969, p.1110
Tetrameres sp.
SA: PMM, 1969, p.1109
166. Hydroprogne caspia (Pallas)
Caspian Tern (migratory)
6/6: TI] C3 N5 Al
. Echinostoma sp.
Qld: CIHUS
Echinostomatinae [spines (45(S))]
SA: AHC (immature)
Galactosomum angelae Pearson, 1973
SA: Pearson, 1973, p.364
Galactosomum bearupi Pearson, 1973
Qld: Pearson, 1973, p.370
Galactosomum ussuriense Oshmarin, 1963
Qld: Pearson, 1973, p.429
Galactosomum sp.
Qld: CIHUS
SA: AHC (2 spp.)
Heterophyidae, unidentified
Qld: CIHUS
Diplostomum paryvulum Dubois & Angel, 1972
SA: Dubois & LMA, 1972, p.206
Posthodiplostomum australe Dubois, 1937a
SA: Dubois & LMA, 1972, p.212
Cardiocephaloides musculosus (S.J, Johnston,
1904)
Qld: Dubois & LMA, 1972, p.202
. Unidentified
Qld: CIHUS
SA: AHC
. Trichostrongylus incertus T.H. Johnston &
Mawson, 194le
SA: THJ & PMM, 194le, p.254
Streptocara pectinifera Neumann, 1900
SA: AHC
Acuariidae, unidentified
SA: TH] & PMM, 194le, p.260 (larva)
. Unidentified
SA: AHC (poor specimen)
167. Sterna sumatrana Raffles
Black-naped Tern
1/0
No helminths recorded
168. Sterna fuscata Linné
Sooty Tern
2/1:N
. Galactosomum bearupi Pearson, 1973
Qld: Pearson, 1973, p.370
Galactosomum renincolum Pearson, 1973
Qld: Pearson, 1973, p.412
Galactosomum ussuriense Oshmarin 1963
Qld: Pearson, 1973, p.429
. Seuratia shipleyi (Stossich, 1900)
Qld, NSW; AHC
te
REC. 5. AUST. MUS, 19(15); 219-325
169. Sterna albifrons Pallas
Little Tern
. Stictodora sp. cf. S. manilensis Africa & Garcia,
1935
Qld: Pearson, 1960, p.93
170. Sterna nereis (Gould)
Fairy Tern
2/1:A
. Unidentified
SA: AHC
171. Sterna bergii Lichtenstein
Crested Tern
5/4: T3 C3 N3
. Austrobilharzia terrigalensis (S.J. Johnston, 1917)
NSW: Bearup, 1955, p.955
Brachylecithum megastomum (S.J. Johnston, 1917)
syn. Lyperosomum megastomum S.J. Johnston,
1917
NSW: SJJ, 1917, p.225
Galactosomum angelae Pearson, 1973
SA: Pearson, 1973, p.36
Heterophyinae, unidentified
Qld: CIHUS
Cardiocephaloides musculosus (S.J. Johnston,
1904)
syn. Holostomum musculosum S.J. Johnston,
1904
NSW; SJJ, 1904, p.112
Maritrema sp. cf. eroliae Yamaguti, 1939a
Qld: CIHUS
. Tetrabothrius sp.
Qld, NSW (cage): AHC
Unidentified
SA: Cleland, 1922, p.105 (AHC)
NSW, SA: AHC
. Capillaria sp.
SA: AHC
Anisakis sp.
SA: AHC (immature)
Cosmocephalus jaenschi T.H. Johnston & Mawson,
1941d
Tas: AHC
Skrjabinocerca sp.
SA: AHC
Stegophorus sp.
SA: AHC
172. Sterna bengalensis Lesson
Lesser Crested Tern
. Galactosomum bearupi Pearson, 1973
Qld: Pearson, 1973, p.370
August, 1986
173. Anous stolidus (Linné)
Common Noddy
2/1, N
N. Contracaecum magnicollare T.H. Johnston &
Cc.
Mawson, 194l1c
Qld: THJ & PMM, 194lc, p.114
Coral Sea: PMM, 1969, p.1104
Seuratia shipleyi (Stossich, 1900)
Coral Sea: PMM, 1969, p.1108
174. Anous minutus Boie
Black Noddy
2/2 :N
. Galactosomum bearupi Pearson, 1973
Qld: Pearson, 1973, p.371
Galactosomum renincolum Pearson, 1973
Qld: Pearson, 1973, p.412
. Anisakis sp.
syn. Stomachus sp. (larvae)
Qld: THJ & PMM, 1951, p.294
Contracaecum magnicollare T.H.
Mawson, 1941c
Qld: AHC
Acuariidae, unidentified
Qld: AHC (larva)
Johnston &
Family COLUMBIDAE
175. Macropygia phasianella (Temminck)
Brown Cuckoo-Dove
Unidentified
Qld: AHC
176. Geopelia placida Gould
Peaceful Dove
§/t ire
. Unidentified
NT: not kept
177. Geopelia cuneata (Latham)
Diamond Dove (endemic)
5/0
No helminths recorded
178. Geopelia humeralis (Temminck)
Bar-shouldered Dove
4/1: N
. Ascaridia sp.
ACT (cage): AHC (poor specimen)
179. Chalcophaps indica (Linné)
Emerald Dove
. Unidentified
Qld: Cleland, 1922, p.104
ia)
ie)
HELMINTHS FROM AUSTRALIAN BIRDS 257
180, Phaps chalcoptera (Latham)
Common Bronzewing (endemic)
7/0
. Unidentified
Vic: AHC
. Ascaridia columbae (Gmelin, 1790)
Vic (cage): AHC
181. Phaps elegans (Temminck)
Brush Bronzewing (endemic)
2/1:C
. Unidentified
SA: Cleland, 1922, p.104
SA: AHC
182. Ocyphaps lophotes (Temminck & Laugier)
Crested Pigeon (endemic)
20/2: C
. Unidentified
SA: AHC
183. Petrophassa albipennis Gould
White-quilled Rock-pigeon (endemic)
1/0
No helminths recorded
184. Geophaps plumifera Gould
Spinifex Pigeon (endemic)
3/0
. Microfilaria sp.
London (cage): Plimmer, 1912, p.408
185. Geophaps scripta (Temminck)
Squatter Pigeon (endemic)
1/0
. Unidentified
Qid: AHC
186, Geophaps smithii (Jardine & Selby)
Partridge Pigeon (endemic)
. Unidentified
NT: AHC
187. Leucosarcia melanoleuca (Latham)
Wonga Pigeon (endemic)
Brachylaima pulchellum (S.J. Johnston, 1917)
Qld: AHC
syn. Harmostomum pulchellum S.J. Johnston,
1917
NSW: SJJ, 1917, p.227
Brachylaima sp.
? loc: AHC
. Davainea sp.
NSW: THJ, 1912b, p.106
Qld: AHC
Unidentified
Qld: AHC
N. Odontoterakis bancrofti (T.H. Johnston, 1912a)
syn. Heferakis bancrofti T.H. Johnston, 1912a
Qld: TH] & PMM, 1941d, p.251
Family CACATUIDAE
188. Calyptorhynchus magnificus (Shaw)
Red-tailed Black-Cockatoo (endemic)
1W1:cC
C. Unidentified
NT: specimens not kept
189, Calyptorhynchus funereus (Shaw)
Yellow-tailed Black-Cockatoo (endemic)
1/0
N. Syngamus trachea (Montagu, 1811)
Vic (cage): Harrigan & Arundel, 1978, p.360
190. Callocephalon fimbriatum (Grant)
Gang-gang Cockatoo (endemic)
N. Ascaridia platyceri Hartwich & Tscherner, 1979
SA (cage): AHC
191. Cacatua roseicapilla Vieillot
Galah (endemic)
13/0
C. Hemiparonia sp.
SA (? cage): AHC
Raillietina leptosoma (Diesing, 1850)
syn. Davainea leptosoma Diesing, 1850
Aust.: Fuhrmann, 1908, p.160
N. Ascaridia sp.
New Zealand: Weekes, 1982, p.457
192. Cacatua tenuirostris (Kuhl)
Long-billed Corella (endemic)
1/0
No helminths recorded
193. Cacatua sp.
A corella
N. Aprocta sp.
Qld: AHC
194, Cacatua sanguinea Gould
Little Corella
3/0
N. Microfilaria sp.
Qld: Mackerras, 1962, p.436
258 REC. S, AUST. MUS. 19(15): 219-325
195. Cacatua leadbeateri (Vigors)
Pink Cockatoo (endemic)
1/0
C. Hemiparonia cacatuae (Maplestone, 1922b)
syn. Hemiparonia merotomochaeta Woodland,
1930
Aust: Woodland, 1930, p.221
N. Heterakis gallinarum (Schrank, 1788)
syn. Heterakis gallinae (Gmelin, 1790)
NSW (cage): THJ & PMM, 1941d, p.251
196. Cacatua galerita (Latham)
Sulphur-crested Cockatoo
1/0
T. Platynosomum proxillicens (Canavan, 1937)
Fla, Wis, USA (cage): Kazakos et al, 1980,
p.788
syn. Dicrocoelium proxillicens Canavan, 1937
Penn, USA (cage): Canavan, 1937, p.478
syn. “Trematodes‘
Penn, USA (cage): Ratcliffe, 1933, p.21
syn. Platynosomum ventroplicatum Heidegger &
Mendheim, 1938b
syn. P. fallax Heidegger & Mendheim, 1938a
Germany (cage): Heidegger & Mendheim,
1938a, p.94; 1938b, p.674
Dicrocoeliinae, unidentified
Cal. USA (cage): AHC
syn. Dicrocoeliidae, unidentified
Cal. USA (cage): Koch & Duhamel, 1982,
p.1388
C. Hemiparonia cacatuae (Maplestone, 1922b)
Qld: Schmidt, 1972, p.1092
SA (cage): AHC
syn. Schizotaenia cacatuae Maplestone, 1922b
Qld: Maplestone, 1922b, p.305
Raillietina cacatuina (T.H. Johnston, 1913)
Qld: Schmidt, 1972, p.1088
Indonesia (cage): Sukarish, 1981, p.53
syn. Davainea cacatuina T.H. Johnson, 1913
Qld: THJ 1913, p.79
Raillietina paucitesticulata (Fuhrmann, 1908)
Aust: BM(NH)
Unidentified
NSW (cage): Cleland, 1922, p.105 (AHC)
Qld: AHC
N. Microfilaria sp.
Qld: Mackerras, 1962, p.436
Family PSITTACIDAE
197. Eclectus roratus (P.L.S. Miller)
Eclectus Parrot
C. Unidentified
Qld: AHC
N. Ascaridia columbae (Gmelin, 1790)
SA (cage): PMM, 1985, p.191
August, 1986
198. Geoffroyus geoffroyi (Bechstein)
Red-cheeked Parrot
N. Cardiofilaria dubia (T.H. Johnston & Mawson,
1940a)
syn. Carinema dubia T.H. Johnston & Mawson,
1940a
Qld: THJ & PMM, 1940a, p.357
syn. Filarial worm’
Qld: Cleland, 1922, p.107 (AHC)
Family LORIDAE
199. Trichoglossus haematodus (Linné)
Rainbow Lorikeet
4/0
C. Paronia trichoglossi (Linstow, 1888)
Qld: Schmidt, 1972, p.1092
Qld: CIHUS
Tas: Fuhrmann, 1908, p.159; 1932, p.283 (no
detail given, may be repetition of Linstow’s
record, with mistaken locality for ‘Cape
York’)
syn. Moniezia trichoglossi (Linstow, 1888)
Qld: THJ, 1913, p.78
syn. Tuenia trichoglossi Linstow, 1888
Qld: Linstow, 1888, p.14
Unidentified
Qld: Cleland, 1922, p.105
N. Microfilaria sp.
Qld: Bancroft, 1889, p.61
200. Glossopsitta concinna (Shaw)
Musk Lorikeet (endemic)
6/0
N. Microfilaria sp.
NSW: Mackerras, 1962, p.436
201. Glossopsitta porphyrocephala (Dietrichsen)
Purple-crowned Lorikeet (endemic)
4/0
No helminths recorded
202. Glossopsitta pusilla (White)
Little Lorikeet (endemic)
N. Microfilaria sp.
Qld: Cleland & THJ, 1912, p.430
Family POLYTELIDAE
203. Alisterus scapularis (Lichtenstein)
Australian King Parrot (endemic)
N. Ascaridia columbae (Gmelin, 1790)
Qld (cage): Mines & Green, 1983, p.279
Ascaridia platyceri Hartwich & Tscherner, 1979
Qld (cage), Vic (cage), SA (cage): PMM, 1985,
p.191
Cc,
T.
HELMINTHS FROM AUSTRALIAN BIRDS
204. Aprosmictus erythropterus Gmelin
Red-winged Parrot
. Ascaridia platyceri Hartwich & Tscherner, 1979
syn. Ascaridia sprenti Mines, 1979
Vie (cage); Mines, 1979, p.374
205, Polytelis swainsonii (Desmarest)
Superb Parrot (endemic)
1/0
. Ascaridia platyceri Hartwich & Tscherner, 1979
Tas (cage): PMM, 1985, p.191
206. Palytelis anthopeplus (Lear)
Regent Parrot (endemic)
Cotugnia polytelidis Burt, 1940
Ceylon (cage); Burt, 1940, p.65
. Ascaridia platyeeri Hartwich & Tscherner, 1979
Berlin (cage): Hartwich & Tscherner, 1979, p.64
syn. Ascuridia sprenti Mines, 1979
Vic (cage); Mines, 1979, p.374
Ascaridia sp.
New Zealand (cage): Weekes, 1983, p.457
207. Polytelis alexandrae Gould
Princess Parrot (endemic)
. Ascaridia columbae (Gmelin, 1790)
Qld (cage); Mines & Green, 1983, p.279
NSW (cage): THJ & PMM, 1941d, p.253
Ascaridia platyceri Hartwich & Tscherner, 1979
Vic (cage), SA (cage): PMM, 1985, p.191
syn, Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
208. Nymphicus hollandicus (Kerr)
Cockatiel (endemic)
3/0
. Filaria (8.1) sp.
NSW; THI & PMM, 1940a, p,36l
Baylisascaris sp.
lowa, USA (cage): Myers, Monroe & Greeve,
1983, p.1089.
Ascaridia platyceri Hartwich & Tscherner, 1979
Berlin (cage): Hartwich & Tscherner, 1979, p.64
SA (cage): AHC
Asearidia sp.
New Zealand (cage): Weekes, 1983, p.457.
Family PLATYCERCIDAE
209. Melopsittacus undulatus (Shaw)
Budgerigah (endemic)
2/0
Austrobilharzia terrigalensis 8.J. Johnston, 1917
NSW: Bearup, 1956, p.471 (exptl.)
Trichobilharzia sp.
NSW: CIHUS (exptl)
N,
a
Cupillaria sp.
Qld (? cage); AHC
Ascaridia columbae (Gmelin, 1790)
Qld (cage); Mines & Green, 1983, p.279
Ascaridia platyceri Hartwich & Tscherner, 1979
Vic (cage), SA (cage): PMM, 1985, p.191
Ascaridia sp.
New Zealand (cage); Weekes, 1983, p.457
Procyrnea incerta (A.J. Smith, 1908)
syn. Spiroptera incerta A.J, Smith, 1908
USA (cage): A.J. Smith, 1908, p.269
210, Lathamus discolor (White)
Swilt Parrot (endemic)
. Ascaridia columbae (Gmelin, 1790)
SA (cage): PMM, 1985, p.191
Ascuridia platvceri Hartwich & Tscherner, 1974
SA (cage): PMM, 1985, p.191
211. Platycercus caledonicus (Gmelin)
Green Rosella (endemic)
3/l1:N
. Unidentified
Tas: AHC (larva)
212. Platycercus elegans (Gmelin)
Crimson Rosella (endemic)
29/4: C3 NI
. Hemiparonia bancrofti (TH. Johnston, 1912¢)
SA: AHC
Unidentified
Vic: AHC
. Ascaridia platyceri Hartwich & Tscherner, 1979
Berlin (cage): Hartwich & Tscherner, 1979, p.64
Qld (cage): VQDP
Vic (cage), SA (cage): PMM, 1985, p.191
Unidentified
SA: AHC (larva in mesentery)
213, Platyeercus eximius (Shaw)
Eastern Rosella (endemic)
W/2:cC
Flemiparonia bancrofti (TH. Johnston, 1912¢)
NSW: Schmidt, 1972, p.1092
syn, Dilepis bancrofti T.H. Johnston, 1912c
Qld, NSW: THJ, 1912c, p.21)
Cotugnia seni Meggitt, 1926
Burma (cage): Meggitt, 1926, p.231
Cotugnia brotogerys Meggitt, 1915
? loc: cited by Fuhrmann, 1932, p.282, without
reference to his authority
Raillietina leptosoma (Diesing, 1850)
? loc: cited by Fuhrmann, 1932, p.282, without
reference to his authority
. Ascaridia platyceri Hartwich & Tscherner, 1979
Berlin (cage): Hartwich & Tscherner, 1979, p.64
Vic (cage); PMM, 1985, p.191
Procyrnea incerta (A.J, Smith, 1908)
26 REC. 8. AUST. MUS, 19(15): 219-325
syn. Spiroptera incerta A.J. Smith, 1908
Philadelphia Zoo: A.J, Smith, 1908, p.269
214. Platyeercus adscitus (Latham)
Pale-headed Rosella (endemic)
N. Microfilaria sp.
Qld: Mackerras, 1962, p.436
215. Platycercus venustus (Kuhl)
Northern Rosella (endemic)
3/1 :N
N. Splendidofilariinae, unidentified
NT: AHC
216. Platycercus icterotis (Kuhl)
Western Rosella (endemic)
C. Cotugnia platycerci Weerekoon, 1944
Ceylon Zoo: Weerekoon, 1944, p.155
N. Microfilaria sp.
London Zoo: Plimmer, 1912, p.408
217. Platycercus sp.
A rosella parrot
C. Raillietina polychalix (Kotlan, 1921)
NSW: Schmidt, 1972, p.1088
Hemiparonia bancrofti (T.H. Johnston, 1912c)
NSW: Schmidt, 1972, p.1092
N. Ascaridia sp.
New Zealand (cage): Weekes, 1983, p.457
Ascaridia platyceri Hartwich & Tscherner, 1979
SA (cage): PMM, 1985, p.191
218, Barnardius barnardi (Vigors & Horsfield)
Mallee Ringneck (endemic)
12/16
OQ
. Hemiparonia bancrofti (T.H. Johnson, 1912c)
NSW: Schmidt, 1972, p.1092
syn, ‘unidentified’
NSW: Cleland, 1922, p.105 (AHC)
Unidentified
SA (cage): AHC
N. Ascaridia columbae (Gmelin, 1790)
SA (cage): PMM, 1985, p.191
Ascaridia platyceri Hartwich & Tscherner, 1979
Qld (cage), Vic (cage), SA (cage): PMM, 1985,
p.191
syn. Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
Procyrnea incerta (A.J. Smith, 1908)
syn, Spiroptera incerta A.J. Smith, 1908
Philadelphia Zoo: A.J. Smith, 1908, p.269
219, Barnardius zonarius (Shaw)
Port Lincoln Ringneck (endemic)
TA: C
C. Unidentified
NT: specimen not kept
August, 1986
N. Ascaridia columbae (Gmelin, 1790)
SA (cage): PMM, 1985, p.191
Ascaridia platyceri Hartwich & Tscherner, 1979
Qld (cage), Tas (cage), SA (cage): PMM, 1985,
p.19L
220. Psephotus haematonotus (Gould)
Red-rumped Parrot (endemic)
13/0*
N, Ascaridia platvceri Hartwich & Tscherner, 1979
Berlin (cage): Hartwich & Tscherner, 1979, p.64
221, Psephotus varius Clark
Mulga Parrot (endemic)
9/0
N. Ascaridia platyvceri Hartwich & Tscherner, 1979
Vic (cage): PMM, 1985, p.191
222. Psephotus chrysopterygius Gould
Golden-shouldered Parrot (endemic)
N. Ascaridia platyceri Hartwich & Tscherner, 1979
SA (cage): PMM, 1985, p.191
223. Psephotus dissimilis Collett
Hooded Parrot (endemic)
1/0
N. Ascaridia platycert Hartwich & Tscherner, 1979
SA (cage): PMM, 1985, p,191
224. Northiella hacmatogaster (Gould)
Blue Bonnet (endemic)
6/0
N. Aeterakis sp.
SA (cage): AHC
Ascaridia platyvceri Hartwich & Tseherner,
SA (cage): PMM, 1985, p.191
syn. Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
1979
225. Northiella narethae (H.L. White)
Naretha Blue Bonnet (endemic)
3/0
No helminths recorded
226, Neophema bourkii (Gould)
Bourke's Parrot
(endemic) 5/0
N. Ascuridia coluwmbae (Gmelin, 1790)
Qld (cage); Mines & Green, 1983, p.279
*QOne body, without skin, head or legs, given to the
authors as that of Psephotus haematonotius, has been
omitted from consideration here, as the identification
is very doubtful. The record of Cyrnea sp. (PMM,
1968d, p. 755, Vic) for this host is now considered
incorrect.
HELMINTHS FROM AUSTRALIAN BIRDS
Ascaridia platyceri Hartwich & Tscherner, 1979
Berlin (cage): Hartwich & Tscherner, 1979, p.84
Vic (cage), SA (cage): PMM, 1985, p.191
syn. Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
227. Neophema chrysostoma (Kuhl)
Blue-winged Parrot (endemic)
2/0
No helminths recorded
228. Neophema elegans (Gould)
Elegant Parrot (endemic)
10/1: A
Z
. Ascaridia platyceri Hartwich & Tscherner, 1979
syn. Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
. Unidentified
SA: AHC (cysts)
>
229. Neophema petrophila (Gould)
Rock Parrot (endemic)
io)
. Raillietina polychalix (Kotlan, 1921)
SA: Schmidt, 1972, p.1088
230. Neophema pulchella Shaw
Turquoise Parrot (endemic)
Zz
. Ascaridia platycert Hartwich & Tscherner, 1979
syn. Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
231. Neophema splendida (Gould)
Scarlet-chested Parrot (endemic)
Oo
. Raillietina sp.
Qld (cage): AHC
. Capillaria sp.
SA: AHC
Ascaridia platyceri Hartwich & Tscherner, 1979
Qld (cage): PMM, 1985, p,191
syn. Ascaridia sprenti Mines, 1979
Vic (cage): Mines, 1979, p.371
Zz
Family CUCULIDAE
232. Cuculus pallidus (Latham)
Pallid Cuckoo
S/l:N
N. Microtetrameres coracinae Mawson, 1977
NT; PMM, 1977, p.253
233. Cuculus variolosus Vigors & Horsfield
Brush Cuckoo
1/1 :N
N. Microtetrameres cacomantis Mawson, 1977
NT: PMM, 1977, p.253
N.
26]
234. Cuculus flabelliformis (Latham)
Fan-tailed Cuckoo
11/4: T1 C2 N2
Pleuropsolus sp.
SA: AHC
. Unidentified
SA: AHC
. Capillaria sp.
Tas: AHC
Microtetrameres cacomantis Mawson, 1977
SA: AHC
Microtetrameres sp.
Tas, SA: PMM, 1977, p.258
235. Chrysococcyx basalis (Horsfield)
Horsfield’s Bronze Cuckoo
6/1 :N
. ? Microhadjelia spiralis (Mawson, 1968d)
NT: AHC
236. Chrysococeyx lucidus (Gmelin)
Shining Bronze Cuckoo
4/1 :N
. Ascaridia galli (Schrank, 1788)
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
Microtetrameres sp.
SA: AHC
237. Eudynamys scolopacea (Linné)
Common Koel (migratory)
1/0
. Spirurida, unidentified
Qld: CIHUS (larva)
238. Seythrops noyaehollandiae Latham
Channel-billed Cuckoo (migratory)
Allodapa suctoria (Molin, 1860)
syn. Subulura clelandi T.H. Johnston & Mawson,
1941d
Qld: THJ & PMM, 1941d, p.251L
239. Centropus phasianinus (Latham)
Pheasant Coucal
2/2: N2 A2
. Echinostoma emollitum Nicoll, 1914b
Qld: Nicoll, 1914b, p.111
. Ascaridia galli (Schrank, 1788)
Qld: BM(NH)
Ascaridia lineata (Schneider, 1866)
Qld: CIHUS
Ascaridia sp.
NT; AHC
Allodapa differens (Sonsino, 1890)
syn. Subulura differens (Sonsino, 1890)
Qld: THJ & PMM, 1942a, p.93
Torquatoides sp. (? T; torquata (Gendre, 1922))
262 REC. S. AUST. MUS, 19(15): 219-325
Qld: AHC
syn. Torquatella sp.
NT: PMM, 1968d, p.765
Aprocta australis (T.H. Johnston & Mawson,
1942a)
bagrifilaria australis T.H. Johnston &
Mawson, 1942a
Qld: THJ & PMM, 1942a, p.93
Lissonema rotundata (Linstow, 1903)
Qld; CIHUS, BM(NH), QM, AHC
syn. Aprocta rotundata (Linstow, 1903)
Qld: Mackerras, 1962, p.433
A. Porrorchis hylae (T.H. Johnston, 1914b)
Qld, NT: AHC
Qld: CIHUS
syn. Pseudoporrorchis hylae (T.H. Johnston,
1914b)
Qld: Edmonds, 1957, p.76
syn. Echinorhynchus bulbocaudatus Southwell &
Macfie, 1925
Qld: Southwell & Macfie, 1925, p.178
syn.
Family STRIGIDAE
240. Ninox rufa (Gould)
Rufous Owl
N. Hamatospiculum meneilli TJA. Johnston &
Mawson, 1941b
Qld: THJ & PMM, 1941b, p.35
syn. ‘nematodes’
Qld: Cleland, 1922, p.107 (AHC)
241. Ninex strenua (Gould)
Powerful Owl (endemic)
T. Strigea promiscua Nicoll, 1914a
Qld: Dubois & LMA, 1972, p.199
N. Allodapa sp.
syn. Subulura sp.
Qld: THJ & PMM, 1941b, p.34
Synhimantus affinis (Seurat, 1916)
Vic: UMVS
A. Centrorhynchus bancrojti (T.H. Johnston & Best,
1943)
syn. Gordiorhynchus bancrofti T.H. Johnston &
Best, 1943
Qld: THJ & Best, 1943, p.226
242. Ninox novaeseelandiae (Gmelin)
Southern Boobook
22/18 : Tl N15 A7
T. &chinoparyphium sp. [spines (41(4))]
SA: AHC
Echinostoma sp. [spines (45(4))]
Vie: AHC
Brachylecithum harrisoni (S.J. Johnston, 1917)
syn. Lyperosomum harrisoni S.J. Johnston, 1917
NSW: SJJ, 1917, p.226
Neodiplostomum brachyurum (Nicoll, 1914a)
SA: Dubois & LMA, 1972, p.209
syn. Hemistomum brachyurum Nicoll, 1914a
August, 1986
Qld: Nicoll, 1914a, p.346
Neodiplostomum lanceolatum Dubois & Angel,
1972
SA: Dubois & LMA, 1972, p.209
SA: AHC
Strigea promiscua Nicoll, 1914a
Qld: Nicoll, 1914a, p.347
SA: Dubois & LMA, 1972, p.199
C. Unidentified
Qld, Tas, SA: Cleland, 1922, p.105
N. Capillaria sp.
SA: AHC
Heterakis sp.
SA: AHC (? from prey)
Allodapa suctoria (Molin, 1860)
SA: AHC
Oxyuridae, unidentified
Tas, SA: AHC (? from prey)
Oxyspirura sp.
SA: AHC
Hartertia sp.
SA: AHC
Physaloptera sp.
NT: AHC
Excisa biloba Mawson, 1968d
Qld, SA, NT; PMM, 1968d, p.755
Procyrnea dollfusi (Mawson, 1968d)
syn. Cyrnea dollfusi Mawson, 1968d
NT: PMM, 1968d, p.750
Tetrameres sp.
SA: AHC
Microtetrameres raptoris Mawson, 1977
NT: PMM, 1977, p.255
Microtetrameres ninoctis Mawson, 1977
SA, NT: PMM, 1977, p.256
Synhimantus laticeps (Rudolphi, 1819)
SA: PMM, 1982, p.23
Synhimantus sp.
SA: AHC
Hamatospiculum meneilli
Mawson, 194la
Qld: THJ & PMM, 194]a, p.12
Qld: Hall Exped. Report, 1974, p.354
Qld: Ogden, 1967, p.506
Lissonema brevicaudata (Chow, 1939)
syn. Aprocta brevicaudata, Chow, 1939
Qld: Ogden, 1967, p.507
Qld: Hall Exped. Report, 1974, p.354
Lissonema sp.
NT: Bain & Mawson, 1981, p.271
Filaria (s.\.) sp.
Vic, WA: THJ, 1912b, p.109
A. Centrorhynchus bancrofti (T.H. Johnston & Best,
1943)
NSW, SA: AHC
syn. Echinorhynchus sp. THJ, 1912b
Qld: THJ, 1912b, p.109
syn. Centrorhynchus sp.
Qld: THJ, 1918, p.216
T.H. Johnston &
HELMINTHS FROM AUSTRALIAN BIRDS 263
243. Ninox connivens (Latham)
Barking Owl
N. Skrjabinura brevicaudatum (T.H. Johnston &
Mawson, 1941b)
syn. Seuratinema brevicaudatum T.H, Johnston
& Mawson, 1941b
Qld: THJ & PMM, 194ib, p.33
Filarial worm, unidentified
NT: NMV
Family TYTONIDAE
244, Tyto alba (Scopoli)
Barn Owll
1/§ : T2 C5 N3
T. Echinostomatinae [spines (41(5))]
SA: AHC
Petasiger sp. [spines (27(4))]
SA: AHC
WA: WAM
Neodiplostomum brachyurum (Nicoll, 1914a)
SA: Dubois & LMA, 1972, p.209
WA: SAM
Neodiplostomum spathula (Creplin, 1829)
Qld: Dubois & Pearson, 1967, p.197
C. Paruterina rauschi Freeman, 1957
SA: AHC
Unidentified
SA: AHC
N. Oxyuridae, unidentified
SA: AHC (? in food)
Microtetrameres tytonis Mawson, 1977
NT: PMM, 1977, p.257
Spirurida, unidentified
SA: AHC (encysted larva)
245. Tyto novaehollandiae (Stephens)
Masked Owl
2/1: N
T. Unidentified
Tas: AHC
N. Capilluria sp.
Tas: AHC
Spirurida, unidentified,
Tas: AHC (larva)
Family PODARGIDAE
246. Podargus strigoides (Latham)
Tawny Frogmouth (endemic)
14/13: TI) Cl NII A3
YT. Echinoparvphium sp. [spines ? (41(4))]
SA: AHC
Echinostoma elongatum Nicoll, 19l4a
Qld: Nicoll, 1914a, p.336
Himasthlinae [spines (35(5))]
Qld: AHC
Brachylecithum podargi Angel & Pearson, 1977
Qld: LMA & Pearson, 1977, p.122
Strigea flosculus Nicoll, 1914a
Qld: Nicoll, 1914a, p.348
Qld: AHC
C. Unidentified
Qld, NSW, Tas, SA: AHC
N. Capillaria sp.
SA: AHC
Ornithostrongylinae, unidentified
Qld: AHC
Allodapa suctoria (Molin, 1860)
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
Qld, SA, NT: AHC
syn. Subulura clelandi T.H. Johnston & Mawson,
1941d
Qld, WA: TH] & PMM, 194ld, p.25]1
Qld, SA, NT: AHC
Allodapa sp.
Qld: CIHUS (immature)
Skrjabinura sp.
syn. Seuratinema sp.
Qld: Ogden, 1967, p.S05
Excisa biloba Mawson, 1968d
Qid, SA, NT: PMM, 1968d, p.755
SA: AHC
Excisa excisiformis (Yamaguti, 1935)
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
Hadjelia acuariana (Gushanskaya, 1937)
Qld, SA, NT; PMM, 1968d, p.759
Qld: AHC
Viguiera chabaudi Mawson, 1968d
SA: PMM, 1968d, p.76l
Synhimantus laticeps (Rudolphi, 1819)
SA: AHC
Synhimantus podargi Mawson, 1982
SA: PMM, 1982, p.24
Microfilaria sp.
Qld: Bancroft, 1889, p.6l
Qld: Cleland & THJ, 1912, p.431
Qld: Cleland, 1915, p.35
A. Porrorchis hylae (T.H. Johnston, 1914b)
Qld, SA: AHC
syn. Pseudoporrorchis hylae (T.H. Johnston,
1914b)
Qld: Edmonds, 1957, p.76
syn. Gordiorhynchus hylae (T.H. Johnston,
1914b)
Qld: THJ & Edmonds, 1948, p.74
247. Podgarus ocellatus Quoy & Gaimard
Marbled Frogmouth
C. Unidentified
Qld: AHC
N. Excisa biloba Mawson 1968d
syn. ‘nematode’
Qld: Cleland, 1922, p.1J05 (AHC)
REC, 8. AUST. MUS, 19(15): 219-325 August, 1986
Family AEGOTHELIDAE
248. Aegotheles cristatus (J. White)
Australian Owlet-Nightjar
6/3 : N3 Cl
. Unidentified
SA: AHC
. Alainchabaudia aegotheles (T.H. Johnston &
Mawson, 194le)
SA: PMM, 1968c, p.743
syn. Habronema aegotheles T.H. Johnston &
Mawson, 194le
SA: TH] & PMM, 194le, p.256
Microtetrameres aegotheles Mawson, 1977
SA: PMM, 1977, p.253
Acuaria sp.
WA: WAM
Microfilaria sp.
Qld: THJ, 1916, p.51
Family CAPRIMULGIDAE
249, Eurostopodus mystacalis (Temminck)
White-throated Nightjar (migratory)
11:TCN
. Eumegacetes sp.
Qld: AHC
. Unidentified
Qld: AHC
. Allodapa suctoria (Molin, 1860)
WA, Qid: AHC
250. Caprimulgus argus (Hartert)
Spotted Nightjar (endemic)
6/1: N
. Allodapa suctoria (Molin, 1860)
WA: AHC
Family APODIDAE
251. Collocalia spodiopygia (Peale)
White-rumped Swiftlet (endemic)
. Unidentified
Qld: AHC
Family ALCEDINIDAE
252. Ceyx azurea (Latham)
Azure Kingfisher
3/1: N
. Spirurida, unidentified
NT; specimen not kept (broken)
253, Dacelo novaeguineae (Hermann)
Laughing Kookaburra (endemic)
16/6 : T3 N4
. Echinoparyphium sp. [spines (45(4))]
Tas: AHC
Brachylecithum dacelonis Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.122
Basantisia queenslandensis Deblock & Pearson,
1968a
Qld: Deblock & Pearson, 1968a, p.136
Diplostomum triangulare (S.J. Johnston, 1904)
Qld: Dubois & Pearson, 1967, p.190
syn. Hemistomum triangulare S.J. Johnston,
1904
NSW: SJJ, 1904, p.108
. Similuncinus dacelonis T.H. Johnston, 1909a
NSW: THJ, 1909a, p.246
Pseudophyllidea, plerocercoids
NSW (cage): AHC
Unidentified
Qld, NSW, Tas: AHC
. Protospirura sp.
SA: AHC (? from prey)
Skrjabinura magnum (T.H. Johnston & Mawson,
1941d)
syn. Seuratinema magnum T.H. Johnston &
Mawson, 1941d
NSW: THJ & PMM, 1941d, p.256
syn. ‘nematode’
NSW: Cleland, 1922, p.107
Alainchabaudia alcedinis Mawson, 1968c
NSW: PMM, 1968c, p.741
Tetrameres dacelonis Mawson, 1979
Qld, ACT: PMM, 1980, p.108
Spirurida, unidentified
Tas: AHC (larva)
. Centrorhynchus horridus (Linstow, 1898a)
Qld, SA: AHC
254. Dacelo leachii Vigors & Horsfield
Blue-winged Kookaburra
1/71: N
. Thelazia dacelonis (Breinl, 1913b)
syn. Filaria dacelonis Breinl, 1913b
Qld: Breinl, 1913b, p.42
? Schistorophus sp.
NT; AHC
255, Halycon macleayii Jardine & Selby
Forest Kingfisher
1/0
. Unidentified
Qld: Cleland, 1922, p.105 (AHC)
256. Halycon pyrrhopygia Gould
Red-backed Kingfisher (endemic)
1/0
. Hamatospiculum haleyonis TH. Johnston &
Mawson, 194la
HELMINTHS FROM AUSTRALIAN BIRDS 265
SA: THJ & PMM, 194la, p.14 Family CORACIIDAE
? Hamatospiculum sp.
NT: AHC 261. Eurystomus orientalis (Linné)
Dollarbird
257. Halycon sancta Vigors & Horstield bY N
her se aia C. ‘Hymenolepis eurystomi Maplestone’
cae at Qld: CIHUS. This appears to be another
. ope blish “ s
C. Unidentified ; ‘a denlified ished! SPECIES
N. Al aes sid lcedinis M 1968 Gi NS as
, ainchabaudld alceauius awson, Cc + ' ;
Qld, SA: PMM, 1968¢, p.741 N: mad RHC
Cheilonematodum halcyonis T.H. Johnston & Ha djelia truncatus (Creplin, 1825)
Mawson, 1941d ae apres
, , : PMM , p.758
Qld, NSW: TH] & PMM, 1941d, p.253 Ace 19680, B
Ancyracanthopsis sp. Ql d: Bancroft 1889, p.6l
a HS Qld: Cleland & THJ, 1912, p.431
Schistogendria sp. (2) Old: Cleland, 1915, p.33
SA: AHC . : sk
L c : Scott, 1927, p.189
Schistogendria sp. (3) ondon {cage}; Sco P
Qld: AHC
Hamatospiculum howense T.H. Johnston & Family PITTIDAE
Mawson, 1940a “
ayn, Pularia-sp. 262. Pitta erythrog: inck
maths . J vaster Temminck
‘ NSW: TH, 1912b, 9.109 Red-bellied Pitta (migratory)
Hamatospiculum halcyonis T.H. Johnston &
Mawson, 194la 34
. ; N. Thelazia pittae TH. Johnston & Mawson, 194le
SAD THY BEM bby Tals, pyl4 Qld: TH] & PMM, 194le, p.256
A. Centrorhynchus horridus (Linstow, 1898a) syn. Nematode’
Qld: THJ & Edmonds 1948, p.69 Qld: Cleland, 1922, p.107 (AHC)
syn, Echinorhynchus sp.
NSW: THJ, 1910, p.105 263. Pitta versicolor Swainson
Noisy Pitta
258. Halycon chloris (Boddaert)
Collared Kingfisher C. Unidentified
1/1/:N Qld: Cleland, 1922, p.105 (AHC)
N. Porrocaecum sp.
N. Contracaecum sp. Qld: AHC
WA: WAM Inglisonema typos Mawson, 1968a
Spirurida, unidentified Qld: PMM, 1968a, p.71
NT: AHC (larva) Microfilaria sp.
Qld: Breinl, 1913a, p.43
259. Syma torotora Lesson
Yellow-billed Kingfisher Family MENURIDAE
C, Unidentified 264. Menura novaehollandiae Latham
Qld: Cleland, 1922, p.105 (AHC) Superb Lyrebird (endemic)
N. Porrocaecum menurae T.H. Johnston & Mawson,
Family MEROPIDAE 1942b
NSW: THJ & PMM, 1942b, p.113
260. Merops ornatus Latham Vic: AHC
Rainbow Bee-eater A. Plagiorhyvnchus menurae (T.H. Johnston,
7/4:.N 1912b)
Vic: AHC
N. Porrocaecum sp. syn, Prosthorhynchus menurae (1H. Johnston,
NT. AHC 1912b)
Torquatoides balanocephala (Gendre, 1922) Qld: THJ & Best, 1943, p.226
Vic, NT; AHC syn. Echinorhynchus menurae T.H. Johnston,
syn, Torquatella balanocephala (Gendre, 1922) 19]2b
SA, NT: PMM, 1968d, p.764 Qld: THJ, 1912a, p.&3
Cc.
N.
T.
Cc,
REC. S. AUST. MUS, 19(15): 219-325 Aust, 1986
Family ALAUDIDAE N. Porrocaecum clelandi TH. Johnston & Mawson,
1941d
265. Miratra javanica Horstield SA: THJ & PMM, 1941d, p.252
Singing Bushlark Microtetrameres sp.
2/1 :N SA: AHC
Unidentified
Unidentified Qld: Hall Exped. Report, 1974, p.354 (from eve
SA: Cleland, 1922, p.106 sockel)
Microtetrameres mirafrae Mawson, 1977 A. Oncicola pomatostomi (TH. Johnston & Cleland,
NT: PMM, 1977, p.249 1912)
Acuaria mirafrae Mawson, 1972 syn. Oligacanthorhynchus pomatostomi (T.H.
NT: PMM, 1972, p.147 Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.354 (cyst)
Family HIRUNDINIDAE
266. Hirundo neoxena Gould
Welcome Swallow Family CAMPEPHAGIDAE
22/10 : T1O Cl
. 271. Coracina novaehollandiae (Gmelin)
Plagiorchis maculosus (Rudolphi, 1802) Black-faced Cuckoo-shrike
SA: LMA, 1959, p.265 5/3: NB
SA: AHC
Unidentified C. Unidentified
Qld, SA: AHC NSW: AHC
N. Capillaria graucalinum T.H. Johnston & Mawson,
267. Cheramoeca leucosternum (Gould) 1941d
Black and White Swallow (endemic) Qld: THJ & PMM, 1941d, p.250
: fo ? Microhadjelia spiralis (Mawson, 1968d)
. Unidentified NT: AHC
NSW: Cleland, 1922, p.105 (AHC) Microtetrameres coracinae Mawson, 1977
, sh _ SA: PMM, 1977, p.253
268. Cecropis nigricans (Vieillot) Paraprocta graucalina (T.H. Johnston & Mawson,
Tree Martin 1940a)
: syn. Carinema graucalinum T.H. Johnston &
. Brachylecithum parvum (S.J. Johnston, 1917) Mawson, 1940a*
SA: LMA & Pearson, 1977, p.118 Qld: THJ & PMM, 1940a, p.358
is Unidentified - ‘ Microfilaria sp.
Qld: Cleland, 1922, p.105 (AHC) Qld: Mackerras, 1962, p.437
269. Ceeropis ariel (Gould) 272. Coracina papuensis (Gmelin)
Fairy Martin (endemic) White-bellied Cuckoo-shrike
S/L:T W1:NT
. Plagiorchis maculosus (Rudolphi, 1802) T. Dicrocoeliidae, unidentified
SA: AHC ; ; NT: discarded
syn. Plagiorchis clelandi S.J. Johnston, 1917 N. Microtetrameres coracinae Mawson, 1977
NSW? S15; 3987 p26 NT: PMM, 1977, p.253
. Unidentified ‘ . A pat:
3 Diplotriaena tricuspis (Fedtschenko, 1874)
NSW: Cleland, 1922, p.105 Old: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
Family MOTACILLIDAE eee
. . Filaria sp.
270. Anthus novaeseelandiae (Gmelin) ;
Richard’s Pipit Qld: THJ, 1912b, p.109
12/3 : T1 C2 N3
Plagiorchis maculosus (Rudolphi, 1802)
syn. Plagiorchis spatulatus S.J. Johnston, 1917 * Carinema graucalinum was transferred to Paraprocta
Qld: SJJ, 1917, p.214 by Anderson (1957). The change was misrepresented
_ Unidentified in Zoological Record (Vol.94(6), p.108) as being to
Tas: Cleland, 1922, p.106 (AHC) Pseudaproctella. \t was later correctly quoted by
Qld, NSW, SA: AHC Mackerras (1962).
HELMINTHS FROM AUSTRALIAN BIRDS
273. Pteropodocys maxima (Riuppell)
Ground Cuckoo-shrike (endemic)
1/0
No helminths recorded
274. Lalage sueurii (Vieillot)
White-winged Triller
3/2: Cl N2
. Unidentified
SA: AHC
. ? Microhadjelia spiralis (Mawson, 1968d)
syn. Cyrnea spiralis Mawson, 1968d
SA: PMM, 1968d, p.753
Microtetrameres sp.
SA: PMM, 1977, p.258
275. Lalage leucomela
(Vigors & Horsfield)
Varied Triller
7/5: N5 Al
. Acuaria petterae Mawson, 1972
NT: PMM, 1972, p.144
. Oncicola pomatostomi (TH. Johnston & Cleland,
1912)
NT: not kept (cyst)
Family MUSCICAPIDAE
276. Zoothera dauma (Latham)
White’s Thrush
3/1: N
>, Unidentified
Qld, NSW, SA: AHC
. Porrocaecum clelandi T.H. Johnston & Mawson,
1941d
Qld: THJ & PMM, 1941d, p.252
SA: AHC
syn. ‘nematode’
Tas: Cleland, 1922, p.108 (AHC)
Microfilaria sp.
NSW: THJ, 1910, p.108
. Echinorhynchus sp.
NSW: THJ, 1910, p.108
Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi
Johnston & Cleland, 1912
Qld, SA: Cleland, 1922, p.108 (cyst)
syn, T.H.
277. Drymodes superciliaris Gould
Northern Serub-robin
’. Unidentified
Qld: AHC
Z.
267
278. Drymodes brunneopygia Gould
Southern Scrub-robin (endemic)
2/1: N
. Acuaria petterae Mawson, 1972
SA: PMM, 1972, p.144
279. Petroica phoenicea Gould
Flame Robin (endemic)
2/0
. Synhimantus sp.
SA: AHC
Filarial worms’
Tas: Cleland, 1922, p.107
280. Petroica multicolor (Gmelin)
Scarlet Robin
10/1: N
. Pseudaprocta copemani Bain & Mawson, 1981]
Tas: Bain & Mawson, 1981, p.266
syn. Aproctidae, unidentified
Tas: Munday & Green, 1972, p.12
281. Petroica goodenovii (Vigors & Horsfield)
Red-capped Robin (endemic)
14/2: N
. Hymenolepis sp.
SA: THJ, 1910, p.106
N. Spirurida, unidentified
SA, NT: AHC (larva)
Aproctidae, unidentified
Tas: Munday & Green, 1972, p.12
282. Melanodryas cucullata (Latham)
Hooded Robin (endemic)
9/1: A
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
SA: specimen not kept (cyst)
Unidentified
NT: AHC (cyst)
283. Eopsaltria pulverulenta Salvadori
Mangrove Robin
. Skrjabinura sp.
syn. Seuratinema sp.
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
284. Eopsaltria australis (Shaw)
Eastern Yellow Robin (endemic)
3/3: N
>. Unidentified
Qld: Cleland, 1922, p.105
. Microtetrameres eopsaltriae Mawson, 1977
SA: PMM 1977, p.253
Unidentified
SA: not kept (larva)
REC. 8, AUST. MUS. 19(15): 219-325
285. Eopsaltria griseogularis Gould
Western Yellow Robin (endemic)
1/0
No helminths recorded
286. Microeca flavigaster Gould
Lemon-bellied Flycatcher
2/0
No helminths recorded
287, Microeca leucophaea (Latham)
Jacky Winter
12/6: T1 N5
Echinoparyphium harveyanum S.J. Johnston, 1917
Qld: SJJ, 1917, p.204
Skrjabinosomum sp.
SA: LMA & Pearson, 1977, p.127
’. Unidentified
Qld: AHC
. Viguiera longicollis Mawson, 1968d
NT? PMM, 1968d, p.759
Microtetrameres mirafrae Mawson, 1977
SA, NT: PMM, 1977, p.249
Acuaria microeca Mawson, 1972
SA: PMM, 1972, p.145
Diplotriaena sp.
SA: AHC
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
Qld: Hall Exped. Report, 1974, p.355 (cyst)
288. Falcunculus frontatus (Latham)
Crested Shrike-tit (endemic)
2/1 :N
. Schistogendria sp. (1)
SA: AHC
289. Pachycephala olivacea
Vigors & Horsfield
Olive Whistler (endemic)
1/0
. Unidentified
Tas: Cleland, 1922, p.105 (AHC)
SA: AHC
290. Pachycephala inornata Gould
Gilbert’s Whistler (endemic)
T/1:N
. Microtetrameres sp.
SA: AHC
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: THJ & Cleland, 1912, p.112 (cyst)
syn,
N.
A,
August, 1986
291. Pachycephala pectoralis (Latham)
Golden Whistler
10/7: N
? Microhadjelia spiralis (Mawson 1968d)
syn. Cyrnea spiralis Mawson, 1968d
SA: AHC
Pseudaprocta copemani Bain & Mawson, 198]
SA: Bain & Mawson, 1981, p.266
Spirurida, unidentified
SA: AHC
292. Pachycephala melanura Gould
Mangrove Golden Whistler
. Unidentified
Qld: Cleland, 1922, p.105 (AHC)
293. Pachycephala rufiventris (Latham)
Rufous Whistler
7/3: Cl N2 A2
. Sphaeruterina punctata T.H. Johnston, 1914b
Qld: THJ, 1914b, p.76
Unidentified
Qld, NSW: Cleland, 1922, p.105 (AHC)
. Acuaria colluricincli Mawson, 1972
SA: AHC
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
NT: AHC (cyst)
294. Colluricincla harmonica (Latham)
Grey Shrike-thrush
14/10 : Tl N8& A4
. Unidentified
SA: AHC
. Unidentified
Tas, SA: Cleland, 1922, p.106 (AHC)
NSW: AHC
. Capillaria sp.
SA: AHC
Viguiera longicollis Mawson, 1968d
SA: PMM, 1968d, p.759
NT: AHC
Acuaria colluricinclae Mawson, 1972
SA: PMM, 1972, p.145
Schistogendria sp.
NT: AHC
Diplotriaena golvani Anderson, 1959
WA: Ogden, 1967, p.505
WA: Hall Exped. Report, 1974, p.354
Microfilaria sp.
NSW; Cleland, 1915, p.33
Mediorhynchus sp.
SA: AHC
Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Echinorhynchus pomatostomi T.H.
HELMINTHS FROM
Johnston & Cleland, 1912
SA: THJ & Deland, 1929, p,149 (cyst)*
295. Colluricincla megarhyncha
(Quoy & Gaimard)
Little Shrike-thrush
1/1: N
N. Diplotriaena sp.
NT: AHC
296. Oreoica gutiuralis (Vigors & Horsfield)
Crested-Bellbird (endemic)
6/6 : N6 Al
N. Microtetrameres sp.
SA, NT: PMM, 1977, p.258
Acuaria petterae Mawson, 1972
NT: PMM, 1972, p.144
Diplotriaena spratti Bain & Mawson, 1981
NT; Bain & Mawson, 1981, p.277
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
NT: specimen not kept (cyst)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
SA: Hall Exped. Report, 1974, p.355 (cyst)
297. Monarcha trivirgatus (Temminck)
Spectacled Monarch
N. Diplotriaena halli Ogden, 1967
Qld: Odgen, 1967, p.5S15
Qld: Hall Exped. Report, 1974, p.354
298. Arses kaupi Gould
Pied Monarch (endemic)
N. Diplotriaena sp.
Qld: Ogden, 1967, p.SO05
Qld: Hall Exped. Report, 1974, p.354
299. Myiagra rubecula (Latham)
Leaden Flycatcher
lili N
N. Rictularina spinosa T.H. Johnston & Mawson,
1941d
Qld: THJ & PMM, 1941d, p.254
Filarioidea, unidentified
NT: AHC
Microfilaria sp.
Qld: Bancroft, 1889, p.6]
Unidentified
Qld: Cleland, 1922, p.!07
* THJ & Deland referred to the host in this record as
‘Cinclosoma’ rufiventris presumably an error in
transcription for Colluricincla rufiventris, a synonym
of C harmonica, THS’s dissection record notes this
parasite from Colluricincla rufiventris trom the same
locality, Pt Lincoln, SA.
AUSTRALIAN BIRDS 269
300. Myiagra alecto (Temminck)
Shining Flycatcher
2/1:N
. Unidentified
Qld: Cleland, 1922, p.105 (larva, subcut.)
. Spirurida, unidentified
NT: not kept (larva)
301. Myiagra inquieta (Latham)
Restless Flycatcher
3/1 :N
. Acuaria petterae Mawson, 1972
SA; PMM, 1972, p.144
Filarioidea, unidentified
NT: :AHC
. ‘echinorhynch’
NSW: Cleland, 1922, p.108
302. Rhipidura fuliginosa (Sparrman)
Grey Fantail
8/4: T3 Cl NI
. Plagiorchis maculosus (Rudolphi, 1802)
SA: AHC
. Unidentified
SA: AHC
. Unidentified
SA; not kept (larva)
303. Rhipidura rufiventris Quoy & Gaimard
Northern Flycatcher
2/0
No helminths recorded
304. Rhipidura leucophrys (Latham)
Willie Wagtail
25/15 : T13 C5 NI Al
. Plagiorchis maculosus (Rudolphi, 1802)
SA: LMA, 1959, p.265
Prosthogonimus sp.
SA: AHC
Eumegacetes sp.
SA: AHC
Lecithodendriidae, unidentified
SA: AHC
. Unidentified
SA: AHC
. Mawsonofilaria rhipidurae (T.H. Johnston &
Mawson, 1952)
syn. Austrofilaria rhipidurae T.H. Johnston &
Mawson, 1952
SA: THJ & PMM, 1952, p.31
. Mediorhynchus sp.
SA: AHC
270 REC. 8. AUST, MUS, 19(15); 219-325 August, 1986
Family ORTHONYCHIDAE
305. Psophodes olivaceus (Latham)
Eastern Whipbird (endemic)
C. Unidentified
NSW: AHC
N. Schistogendria sp. (1)
Qld: AHC
Microfilaria sp.
Qld: Cleland & THJ, 1912, p.431!
A. Echinorhynchus sp.
NSW: THY, 1910, p.107
Qld: Cleland, 1922, p.108
306. Psephodes cristatus (Gould)
Chirruping Wedgebill (endemic)
2/0
No helminths recorded
307. Cinclosoma punctatum (Shaw)
Spotted Quail-thrush (endemic)
1/1 N
N. Porrocaecum clelandi T.H. Johnston & Mawson,
1941d
Tas: THJ & PMM, 194ld, p.252
308. Cinclosoma castanotum Gould
Chestnut Quail-thrush (endemic)
3/0
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn, Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: THJ & Deland, 1929, p.149 (cyst)
309. Cinelosoma cinnamomeum Gould
Cinnamon Quail-thrush (endemic)
‘8/4: N3 Al
N. Capillaria sp.
NT: AHC
Viguiera sp.
SA: AHC
? Microhadjelia spiralis (Mawson, 1968d)
NT: AHC
Acuaria petterae Mawson, 1972
NT: AHC
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
NT: AHC (cyst)
syn. Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: Cleland, 1922, p.108
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.354 (cyst)
Family TIMALIIDAE
310. Pomatostomus temporalis
(Vigors & Horsfield)
Grey-crowned Babbler
7/4: Cl A3
©. Paruterinae, unidentified
Qld: BM(NH)
Unidentified
Qld, NT: AHC
|. Skrjabinura brevicaudatum (T.H. Johnston &
Mawson, 1941b)
syn. Seuratinema brevicaudatum T.H. Johnston
& Mawson, 1941b
Qld: Inglis, 1967, p.130
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
Microfilaria sp.
Qld: Bancroft, 1889, p.61
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
NTE AHC
syn. Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
NSW, WA: THJ & Cleland, 1912, p.112 (cyst)
NSW: Cleland, 1922, p.108 (cyst)
syn. Echinorhynchus sp.
WA: THI, 1910, p.108
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
Qld: Hall Exped. Report, 1974, p.354 (cyst)
311. Pomatostomus superciliosus
(Vigors & Horsfield)
White-browed Babbler (endemic)
37/33 : T16 C2 N16 A22
Leucochloridium qaustraliense T.H. Johnston &
Cleland, 1938
SA: THJ & Simpson, 1940a, p.119
Plagiorchis maculosus (Rudolphi, 1802)
SA: LMA, 1959, p.265
Skrjabinosomum pomatostomi Angel & Pearson,
1977
SA: LMA & Pearson, 1977, p.126
Laterotrema sp.
SA: AHC
-, Paruterinae, unidentified
WA: BM(NH)
Unidentified
SA: AHC
. Capillaria pomatostomi TH. Johnston & Mawson
1945b
SA: THJ & PMM, 1945b, p.247
SA: AHC
Skrjabinura pomatostomi (T.H. Johnston &
Mawson, 1941d)
syn. Seuratinema pomatostomi T.H. Johnston &
Mawson, 1941d
NSW: THJ & PMM, 1941d, p.255
Spirurida, unidentified
SA: THJ & PMM, 194le, p.261 (larva)
HELMINTHS FROM AUSTRALIAN BIRDS 271
SA: AHC (larva)
syn. ‘nematode’
NSW: Cleland, 1922, p.l07
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
SA, WA, NT: AHC (cyst)
syn. Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
Qld, WA: Cleland, 1922, pp.107,108 (cyst)
syn. Echinorhynchus sp.
SA: THJ, 1910, p.107 (cyst)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.354 (cyst)
312. Pomatostomus ruficeps (Hartlaub)
Chestnut-crowned Babbler (endemic)
2/1: NA
N. Capillaria sp,
SA: AHC
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: THJ & Deland, 1929, p.150 (cyst)
SA: AHC (cyst)
Family SYLVIIDAE
313. Megalurus gramineus (Gould)
Little Grassbird
2/0
C. Unidentified
SA: AHC
314. Eremiornis carteri North
Spinifexbird (endemic)
2/0
No helminths recorded
315. Cisticola exilis (Vigors & Horsfield)
Golden-headed Cisticola
N. Diplotriaena tricuspis (Fedchenko, 1874)
Qld: THJ & PMM, 194la, p.14
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
316. Cinclorhamphus mathewsi Iredale
Rufous Songlark (endemic)
2/0
No helminths recorded
317. Cinclorhamphus cruralis
(Vigors & Horsfield)
Brown Songlark (endemic)
5/0
No helminths recorded
Family MALURIDAE
318. Malurus cyaneus (Latham)
Superb Fairy-wren (endemic)
14/3: N
. Choanotaenia taylor] TH. Johnston, 1912¢
SA: THJ, 1912¢, p.213
Unidentified
NSW: Cleland, 1922, p.106
Tas: Munday & Green, 1972, p.4
. Spnhimantus sp.
SA: AHC
Willmotiia australis Mawson, 1982
Tas: PMM, 1982, p.28
Diplotriaena sp.
SA: AHC
Filaria (s.1.) sp.
Tas: Cleland, 1922, p.108
319. Malurus splendens (Quoy & Gaimard)
Splendid Fairy-wren (endemic)
Unidentified (incomplete)
WA: WAM
. Unidentified
SA: AHC
320. Malurus lamberti Vigors & Horsfield
Variegated Fairy-wren (endemic)
8/1: N
. Unidentified
Tas: AHC
. Tetrameres sp.
NT: AHC
Diplotriaena delta T.H. Johnston & Mawson, 1940a
SA: THJ & PMM, 1940a, p.360
Qld, SA, WA: Ogden, 1967, p.511
Qld, SA, WA: Hall Exped. Report, 1974, p.354
Diplotriaena halli Ogden, 1967
WA: Ogden, 1947, p.SI5
WA: Hall Exped. Report, 1974, p.354
Diplotriaena sp.
Qld: AHC
Unidentified
NT: AHC (larva)
321. Malurus pulcherrimus Gould
Blue-breasted Fairy-wren (endemic)
2/0
No helminths recorded
322. Malurus leucopterus Dumont
White-winged Fairy-wren (endemic)
18/1: N
. Capillaria sp.
SA: AHC
Porrocaecum sp.
SA: AHC
Viguiera sp.
272 REC. 8. AUST. MUS. 19015): 219-325
SA: AHC
Acuaria microeca Mawson, 1972
SA: AHC
Filaria (s.L) sp.
SA: THS & PMM, 194la, p.I5 (cysts - ?
identification)
323. Stipiturus ruficeps Campbell
Rufous-crowned Emu-wren (endemic)
1/0
No helminths recorded
324, Amytornis goyderi (Gould)
Eyrean Grasswren (endemic)
6/0
N. Diplotriaena delta T.H. Johnston & Mawson,
1940a
SA: Bain & Mawson, 1981, p.281
325. Amytornis textilis (Dumont)
Thick-billed Grasswren
3/0
No helminths recorded*
326. Amytornis purnelli Mathews
Dusky Grasswren
4/0
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)*
Family ACANTHIZIDAE
327. Dasyornis broadbenti (McCoy)
Rufous Bristlebird (endemic)
1/0
No helminths recorded
328. Sericornis citreogularis Gould
Yellow-throated Scrubwren (endemic)
N. Larval nematode
Qld: AHC
329. Sericornis frontalis (Vigors & Horsfield)
White-browed Scrubwren (endemic)
2/0
A. Echinorhynchus sp.
SA: THJ & Deland, 1929, p.150 (cyst)
* The A. fextilis’ of the Hall Report was in fact A.
purnelli, (vide Parker, Emu 72, pp.157-166)
August, 1986
330. Sericornis pyrrhopygius
(Vigors & Horsfield)
Chestnut-rumped Hylacola (endemic)
C. Unidentified
NSW, SA: Cleland, 1922, p.105
SA: AHC
A. Oncicola pomatostomi (T,H. Johnston & Cleland,
1912)
SA: AHC
syn. Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: THJ & Cleland, 1912, p.112 (cyst)
331. Sericornis cautus (Gould)
Shy Hylacola (endemic)
S/1:A
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
SA: AHC (cyst)
332. Sericornis brunneus (Gould)
Redthroat (endemic)
2/1 :A
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi T.H,
Johnston & Cleland, 1912
SA: THJ & Deland, 1929, p.150 (cyst)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
syn.
333. Sericornis fuliginosus
(Vigors & Horsfield)
Calamanthus (endemic)
1/0
N. Diplotriaena alpha T.H. Johnston & Mawson,
1940a
SA: PMM, 1955, p.6
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
334. Smicrornis brevirostris (Gould)
Weebill (endemic)
S/1:C
C. Unidentified
SA: not kept (cysts)
335. Gerygone fusca (Gould)
Western Gerygone
1/0
No helminths recorded
HELMINTHS FROM AUSTRALIAN BIRDS 273
336. Acanthiza pusilla (Shaw)
Brown Thornbill (endemic)
16/2: N
T. Prosthogonimus sp.
SA: AHC
N. Spirurida, unidentified
SA: AHC (larva)
337. Acanthiza apicalis Gould
Inland Thornbill (endemic)
3/0
No helminths recorded
338. Acanthiza uropygialis Gould
Chestnut-rumped Thornbill (endemic)
1/0
No helminths recorded
339. Acanthiza reguloides
Vigors & Horsfield
Buff-rumped Thornbill (endemic)
5/0
No helminths recorded
340. Acanthiza iredalei Mathews
Slender-billed Thornbill (endemic)
4/0
No helminths recorded
341. Acanthiza chrysorrhoa
(Quoy & Gaimard)
Yellow-rumped Thornbill (endemic)
18/3: T1 NI Al
T. Unidentified
SA: AHC
N. Unidentified
SA: not kept (larva)
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
SA: specimen lost
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
342. Acanthiza lineata Gould
Striated Thornbill (endemic)
5/0
No helminths recorded
343. Aphelocephala leucopsis (Gould)
Southern Whiteface (endemic)
9/3: A
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Echinorhynchus sp.
SA: THJ, 1910, p.109 (cyst)
syn. Echinorhynchus pomatostomi
Johnston & Cleland, 1912
SA: Cleland, 1922, p.108 (cyst)
T.H.
344. Aphelocephala nigricincta (North)
Banded Whiteface (endemic)
N. Aprocta vestibulata (T.H. Johnston & Mawson,
1952)
syn. Austrofilaria vestibulata T.H. Johnston &
Mawson, 1952
SA: THJ & PMM, 1952, p.31
Family NEOSITTIDAE
345. Daphoenositta chrysoptera (Latham)
Varied Sitella (endemic)
5/0
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
WA: Hall Exped. Report, 1974, p.355 (cyst)
Family CLIMACTERIDAE
346. Climacteris leucophaea (Latham)
White-throated Treecreeper (endemic)
5/2: N
C. Unidentified
EA: AHC
N. Spirurida, unidentified
SA: AHC (larva)
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi
Johnston & Cleland, 1912
SA: THJ & Deland, 1929, p.151 (cyst)
syn. T:H.
347. Climacteris picumnus Temminck
Brown Treecreeper (endemic)
9/3: A
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
SA: Cleland, 1922, p.108; THJ & Deland, 1929,
p.150 (cyst)
syn.
348. Climacteris rufa Gould
Rufous Treecreeper (endemic)
3/0
No helminths recorded
349. Climacteris melanura Gould
Black-tailed Treecreeper (endemic)
N. Microfilaria sp.
NT: Mackerras, 1962, p.438
274
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
WA: THI, 1910, p.109 (cyst)
syn.
Family MELIPHAGIDAE
350. Anthochaera carunculata (White)
Red Wattlebird (endemic)
16/7 : C4 N7 Al
T. Lecithodendriidae, unidentified
SA: AHC
C, Dilepis sp.
NSW: AHC
Capiuterilepis australiensis Schmidt, 1972
SA: Schmidt, 1972, p.1089
Unidentified
SA: AHC
N. Oxyspirura anthochaerae (V.H. Johnston, 1912a)
syn. Ceratospira anthochaerae T.H. Johnston,
1912a
NSW: THJ, 1912a, p.80
syn. Ascaris sp.
NSW: Krefft, 1873, p.213
Viguiera longicollis Mawson, 1968d
SA: PMM, 1968d, p.759
Microtetrameres meliphagidae Mawson, 1977
SA: PMM, 1977, p.31
Geopetitia sp.
SA: AHC
A. Oncicola pomatostomi (T.H, Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
Unidentified
SA: AHC (fragments)
351. Anthochaera paradoxa (Daudin)
Yellow Wattlebird (endemic)
N. Diplotriaena sp.
Tas: Munday & Green, 1972, p.Il
352. Anthochoera chrysoptera (Latham)
Little Wattlebird (endemic)
10/3: Cl N2
C. Choanotaenia sp.
Qld: AHC
Unidentified
SA: AHC
N. Microtetrameres meliphagidae Mawson, 1977
SA: PMM, 1977, p.248
Filaria (s.L.) sp.
Qld: Bancroft, 1889, pp.60,61
Mtcrofilaria sp.
Qld: Bancroft, 1889, p.61
Qld: THJ, 1910, p.lll
Qld: Cleland & TH, 1910, p.108 (two species
noted)
REC. S. AUST. MUS. 19(15): 219-325
August, 1986
NSW: Mackerras, 1962, p.439
A. Mediorhynchus sp.
SA: AHC
353. Anthochaera sp.
a wattlebird
2/2: TI CI NI
T. Unidentified
SA: AHC
C. Capiuterilepis australiensis Schmidt, 1972
SA: Schmidt, 1972, p.1089
Hymenolepsis sp.
SA: AHC
N. Microtetrameres meliphagidis Mawson, 1977
SA: AHC
Acuaria petterae Mawson, 1972
SA: AHC
354. Acanthagenys rufogularis Gould
Spiny-cheeked Honeyeater (endemic)
26/12 : Cl N12
C. Unidentified
NSW, SA: Cleland, 1922, p.106
NT: AHC
N. Capillaria sp.
SA: AHC
Microtetrameres meliphagidae Mawson, 1977
SA: PMM, 1977, p.248
Acuaria petterae Mawson, 1972
SA: PMM, 1972, p.144
Diplotriaena zeta T.H. Johnston & Mawson, 1940a
SA: THJ & PMM, 1940a, p. 359
Diplotriaena smithi Bain & Mawson, 1981
SA, WA: Bain & Mawson, 1981], p.280
Filaria (s.1.) sp.
NSW: TH J, 1912b, p.111
Qld: THJ & PMM, 1940a, p.361
Unidentified
NSW: Cleland, 1922, p,108
A. Mediorhynchus sp.
SA: AHC
355. Plectorhyncha lanceolata Gould
Striped Honeyeater (endemic)
N. Microfilaria sp.
Qld: Cleland & THJ, 1912, p.433
356. Philemon argenticeps (Gould)
Silver-crowned Friarbird (endemic)
3/3 :N
N. ? Microhadjelia spiralis (Mawson 1968d)
syn. Crrnea spiralis Mawson, 1968d
NT: PMM, 1968d, p.753
Microtetrameres philemon Mawson, 1977
NT: PMM, 1977, p.249
Spirurida, unidentified
NT: AHC (larva)
HELMINTHS FROM AUSTRALIAN BIRDS 275
357. Philemon corniculatus (Latham)
Noisy Friarbird
C, Capiurerilepis meliphagicola Schmidt, 1972
Qld; Schmidt, 1972, p.L091
N. Diplotriaena smithi Bain & Mawson, 1981
Qld: Bain & Mawson, 1981, p.280
syn. Diplotriaena sp.
Qld: Mackerras, 1962, p.432
358. Philemon citreogularis (Gould)
Litthe Friarbird
3/3: N
C. Raillietina conopophilae (1H, Johnston, 1913)
syn. Davainea conopophilae T.H. Johnston, 1913
Qld: THJ, 1913, p.80
Hymenolepididae, unidentified
Qld: CIHUS
N. Oxvspirura bancrofti TH. Johnston & Mawson,
1941d
Qld: THJ & PMM, 1941d, p.255
Microtetrameres philemon Mawson, 1977
NT: PMM, 1977, p.249
Filaria sp.
Qld: THJ, 1912a, p.78
Microfilaria sp.
? loc: Cleland, 1922, p.9l
359. Entomyzon cyanotis (Latharn)
Blue-faced Honeyeater
5/2: N2 Al
C. Ratllietina conopophilae (TH. Johnston, 1913)
syn. Davained conopophilae T.H. Johnston, 1913
Qld: THJ, 1913, p.80
Anoncotaenta globata (Linstow, 1879)
NSW: Schmidt, 1972, p. L087 (S. mistakenly
gives locality as Qld)
syn. ‘Unidentified’
NSW: Cleland, 1922, p.106 (AHC)
Hymenolepididae, unidentified
? Qld : CIHUS
Unidentified
Qld: AHC
Qld: CIHUS (spargana)
N, ? Microhadjelia spiralis (Mawson, 1968d)
syn. Cyrnea spiralis Mawson, 1968d
NT: PMM, 1968d, p,763
Microtetrameres philemon Mawson, 1977
ACT, SA: PMM, 1977, p.248
Microfilaria sp.
Qld: Bancroft, 1889, p.6l
Qld, NSW: Cleland & THJ, 1912, p.434
Qld, NSW: Mackerras, 1962, p.439
London (cage): Plimmer, 1912, p.408
Spirurida, unidentified
NT: AHC (larva)
A. Unidentified
NT: specimen not kept (adult)
360. Manorina melanophrys (Latham)
Bell Miner (endemic)
T. Urotocus sp.
NSW (cage): AHC
361. Manorina melanocephala (Latham)
Noisy Miner (endemic)
10/3: N
N. Capillaria sp.
SA: AHC
Microtetrameres meliphagidae Mawson, 1977
ACT, SA: PMM, 1977, p.248
Filaria (8.1) sp. (2 Filarioidea)
Qld: Bancroft, 1889, pp.60, 62
Microfilarta sp.
Qld; Bancroft, 1889, p.61
Qld, NSW: Cleland & THJ, 1912, p.434
London (cage): Plimmer, 1912, p.408
362. Manorina flavigula (Gould)
Yellow-throated Miner (endemic)
10/4: T1 Cl N2
T. Skrjabinosomum inawsoni Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.l24
C. Unidentified
NSW: Cleland, 1922, p,106
NT: specimens not kept
N. Capillaria sp.
SA: AHC
Porrocaecuni sp.
SA: THJ & PMM, 1941d, p.253 (larva)
Microtetrameres meliphagidae Mawson, 1977
SA, WA: PMM, 1977, p.248
Pseudaprocta myzanthae T.H.
Mawson, 1940a
SA: THJ & PMM, 1940a, p.358
syn. ‘nematode’
SA: Cleland, 1922, p.108 (AHC)
A. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
Echinorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: AHC (cyst)
syn. Oligacanthorhynchus pomatostonu (TH,
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
Johnston &
syn.
363. Meliphaga lewini (Swainson)
Lewin’s Honeyeater (endemic)
3/0
C. Pseudochoanotaenia meliphagzidarum (T.H.
Johnston, 1911)
syn. Choanotaenia meliphagidarum T.H.
Johnston, 1911
NSW: THJ, 1911, p.5&
syn. Choandtuenia sp.
NSW: THJ, 1910, p.110
N. Svnahimantus lichenostomi Mawson, 1982
Qld: PMM, 1982, p.26
276 REC, $8. AUST.
. Pseudachoanotaenia
364. Stomiopera unicolor (Gould)
White-gaped Honeyeater (endemic)
2/0
No helminths recorded
365. Lichenostomus chrysops (Latham)
Yellow-faced Honeyeater (endemic)
3/0
No helminths recorded
366. Lichenostomus virescens (Vieillot)
Singing Honeyeater
26/6: Cl N6
. Hymenolepis sp.
SA: discarded
. Capillaria sp.
SA: AHC
Microtetrameres meliphagidae Mawson, 1977
SA: PMM, 1977, p.248
Acuaria petterae Mawson, 1972
NT: PMM, 1972, p.l44
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
367. Lichenostomus leucotis (Latham)
White-eared Honeyeater (endemic)
10/2: N
meliphagidarum (T.H.
Johnston, 1911)
syn. Choanotaenia meliphagidarum
Johnston, 1911
NSW: THJ, 1911, p.58
syn. Choanotaenia sp.
NSW; TH, 1910, p.110
kA.
. Microtetrameres meliphagidae Mawson, 1977
SA: PMM, 1977, p.248
368. Lichenostomus melanops (Latham)
Yellow-tufted Honeyeater (endemic)
. Unidentified
NSW: Cleland, 1922, p.106 (AHC)
369. Lichenostomus cratitius (Gould)
Purple-gaped Honeyeater
1/0
No helminths recorded
370. Lichenostomus keartlandi (North)
Grey-headed Honeyeater (endemic)
/O
No helminths recorded
19(15): 219-325 August, 1986
371. Lichenostomus ornatus (Gould)
Yellow-plumed Honeyeater (endemic)
8/3: T2 N1
. Pancreatrema meliphagae Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.129
. Unidentified
SA: Cleland, 1922, p.106
. Microtetrameres sp.
SA: AHC
372. Lichenostomus plumulus (Gould)
Grey-fronted Honeyeater (endemic)
4/1: N
. Pseudochoanotaenia meliphagidarum (T.H.
Johnston, 1911)
syn. Choanotaenia meéliphagidarum T.H.
Johnston, 1911
NSW: THJ, 1911, p.58
syn. Choanolaenia sp.
NSW: THJ, 1910, p.110
. ? Microhadjelia spiralis (Mawson, 1968d)
NT: AHC
Acuaria petierae Mawson, 1972
NT; PMM, 1972, p.144
. Oncicola pomatostami (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
WA: Hall Exped. Report, 1974, p.355 (cyst)
373. Lichenostomus fuscus (Gould)
Fuscous Honeyeater (endemic)
1/0
>, Unidentified
NSW: Cleland, 1922, p.l06 (AHC)
Qld: AHC
. Microfilaria sp.
Qld: Cleland & THJ, 1912, p.434
374. Lichenostomus penicillatus (Gould)
White-plumed Honeyeater (endemic)
21/2: C1 Ni
*, Unidentified
SA: AHC
. Synhimantus lichenostomi Mawson, 1982
SA: PMM, 1982, p.26
Schistogendria sp. |
SA: AHC
Unidentified
SA: Cleland, 1922, p.108
375. Melithreptus brevirostris
(Vigors & Horsfield)
Brown-headed Honeyeater (endemic)
4/0
No helminths recorded
HELMINTHS FROM AUSTRALIAN BIRDS
376. Melithreptus albogularis Gould
White-throated Honeyeater
N. Microfilaria sp.
Qld: Mackerras, 1962, p.438
377. Melithreptus lunatus (Vieillot)
White-naped Honeyeater (endemic)
4/0
No helminths recorded
378. Lichmera indistincta
(Vigors & Horsfield)
Brown Honeyeater
SIINT
T. Cyclocoelidae, unidentified
NT; AHC
Acanthoparyphium sp. [spines (23(0))]
NT: AHC
Himasthlinae [spines (31(4))]
NT: AHC
C. Unidentified
Qld: Cleland, 1922, p.106
NSW: AHC
N. Capillaria sp.
NT: AHC
Microfilaria sp.
NSW: Cleland & THJ, 1912, p.434
379. Phylidonyris pyrrhoptera (Latham)
Crescent Honeyeater (endemic)
9/0
No helminths recorded
380. Phylidonyris novaehollandiae
(Latham)
New Holland Honeyeater (endemic)
30/5 : C3 N2
C, Pseudochoanotaenia meliphagidarum (1.H.
Johnston, 1911)
SA: Schmidt, 1972, p.1038
syn. Choanotaenia meliphagidarum T.H.
Johnston, 1911
NSW: TH, 1911, p.58
syn. Choanotaenia sp.
NSW: THJ, 1910, p.111
Unidentitied
NSW, SA: AHC
N. Svahimantus falco Mawson, 1982
SA; PMM, 1982, p.24
Xenocordon patonae Mawson, 1982
SA: PMM, 1982, p.27
A. Echinorhynchus sp.
NSW: THJ, 1910, p.111
381, Phylidonyris nigra (Bechstein)
White-cheeked Honeyeater (endemic)
1/0
. Pseudochoanotaenia meliphagidarum (T.H.
Johnston, 1911)
syn. Choanotaenia meliphagidarum T.H.
Johnston, 1911
NSW: THJ, 1911, p.58
syn. Choanotaenia sp.
NSW: THJ, 1910, p.110
Unidentified
Qld: Cleland, 1922, p.106 (AHC)
NSW: AHC
382. Phylidonyris albifrons (Gould)
White-fronted Honeyeater (endemic)
T/LiN
>. Unidentified
SA: Cleland, 1922, p.106
. Microtetrameres meliphagidae Mawson, 1977
SA: AHC
383. Phylidonyris melanops (Latham)
Tawny-crowned Honeyeater (endemic)
T/12N
. Unidentified
SA: AHC (cysts)
384. Conopophila albogularis (Gould)
Rufous-banded Honeyeater
>. Raillietina conopophilae (T.H. Johnston, 1913)
syn. Davainea conopophilae T.H. Johnston,
1913*
Qld: THJ, 1913, p.80
385. Conopophila rufogularis (Gould)
Rufous-throated Honeyeater (endemic)
1/0
No helminths recorded
386. Acanthorhynchus tenutrostris
(Latham)
Eastern Spinebill (endemic)
3/0
No helminths recorded
* T.H. Johnston recorded the host of this cestode as
‘White-throated Honeyeater, Conopophila
albogularis’. It is possible that he was dealing with
what is now referred to as the White-throated
Honeyeater, Melithreptus albogularis (as listed in the
CIHUS), but, in view of the specific name given to
the cestode, unlikely. Entomophila’ the host genus
on the type slide, is a synonym of Conopophila.
te
REC. $. AUST. MUS. 19(15); 219-325
387, Certhionyx pectoralis (Gould)
Banded Honeyeater (endemic)
3/0
No helminths recorded
388. Certhionyx yariegatus Lesson
Pied Honeyeater (endemic)
3/0
No helminths recorded
389. Myzomela obscura Gould
Dusky Honeyeater
2/0
No helminths recorded
390, Myzomela sanguinolenta (Latham)
Scarlet Honeyeater (endemic)
. Pseudochoanotaenia meliphagidarum
Johnston, 1911)
SA: AHC
. Microfilaria sp.
Qld: Cleland & THJ, 1912, p.433
Qld: Cleland, 1915, p.33
Family EPHTHIANURIDAE
39]. Ephthianura tricolor Gould
Crimson Chat (endemic)
5/0
No helminths recorded
392. Ephthianura aurifrons Gould
Orange Chat (endemic)
4/0
No helminths recorded
393. Ephthianura albifrons
(Jardine & Selby)
White-fronted Chat (endemic)
5/1 :N
. Acuariidae, unidentified
SA: AHC (larva)
394. Ashbyia lovensis (Ashby)
Gibberbird (endemic)
1/0
No helminths recorded
Family DICAEIDAE
395. Dicaeum hirundinaceum (Shaw)
Mistletoe bird
2/0
No helminths recorded
(T.H.
Family PARDALOTIDAE
396. Pardalotus punctatus
(Shaw & Nodder)
Spotted Pardalote (endemic)
3/0
No helminths recorded
397. Pardalotus xanthopygus McCoy
Yellow-rumped Pardalote (endemic)
2/0
No helminths recorded
398. Pardalotus rubricatus Gould
Red-browed Pardalote (endemic)
W1:C
C. Unidentified
Qld: not kept
399, Pardalotus striatus (Gmelin)
Striated Pardalote (endemic)
23/7 : T2 C6 NI
T. Plagiorchis maculosus (Rudolphi, 1802)
SA: AHC
Unidentified
SA: AHC
C. Anonchotaenia arhyncha Fuhrmann, 1918
SA: Schmidt, 1972, p.1087
Hymenolepis sp.
SA: AHC
Unidentified
Tas, SA: Cleland, 1922, p.106
SA: AHC
N. Diplotriaena sp.
NT; AHC
Microfilaria sp.
Qld: Cleland & THJ, 1912, p.433
Unidentified
SA: Cleland, 1922, p.108
Family ZOSTEROPIDAE
400. Zosterops lateralis (Latham)
Silvereye
19/2: C
C. Zosteropicola clelandi T.H. Johnston, 1912c
NSW: TH J, 1912c, p.214
Unidentified
NSW, Tas: Cleland, 1922, p.106 (AHC)
Tas: Munday & Green, 1972, p.4
NSW, SA: AHC
N. Diplotriaena sp.
Qld: Ogden, 1967, p.SO5
Qld: Hall Exped. Report, 1974, p.354
Tas: AHC
Microfilaria sp.
Qld: Mackerras, 1962, p.438
August, 1986
qq
HELMINTHS FROM AUSTRALIAN BIRDS 279
Family PLOCEIDAE
401. Emblema temporale (Latham)
Red-browed Firetail (endemic)
8/0
No helminths recorded
402. Emblema bellum (Latham)
Beautiful Firetail (endemic)
3/0
. Acuariidae, unidentified
Tas (cage): AHC
403. Emblema pictum Gould
Painted Firetail (endemic)
. Dilepididae, unidentified
NSW (cage): AHC
404, Emblema guttatum (Shaw)
Diamond Firetail (endemic)
2/0
. Choanotaenia sp.
Vie: AHC
405. Neochmia phacton
(Hombron & Jacquinot)
Crimson Finch
7/0
No helminths recorded
406. Poephila guttata (Vieillot)
Zebra Finch
25/3: C
. Unidentified
SA: AHC
407. Poephila bichenovii
(Vigors & Horsfield)
Double-banded Finch (endemic)
1/0
. Choanotaenia sp.
Vic: AHC
Dilepididae, unidentified
NSW (cage): AHC
. Microfilaria sp.
London (cage): Plimmer, 1915, p.129
408. Poephila personata Gould
Masked Finch (endemic)
. Microfilaria sp.
London (cage): Scott, 1926, p.237
409. Poephila acuticauda (Gould)
Long-tailed Finch (endemic)
1/0
No helminths recorded
410. Poephila cincta (Gould)
Black-throated Finch (endemic)
. Dilepididae, unidentified
NSW (cage): AHC
Unidentified
Qld: AHC
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.H.
Johnston & Cleland, 1912)
Qld: Hall Exped. Report, 1974, p.355 (cyst)
411. Lonchura castaneothorax (Gould)
Chestnut-breasted Mannikin
3/0
No helminths recorded
412. Erythrura trichroa (Kittlitz)
Blue-faced Finch
. Unidentified
NSW (cage): AHC
Vic (cage): UMVS
. Acuaria skrjabini Ozerskaya, 1926
Vic (cage): AHC
413. Erythrura gouldiae (Gould)
Gouldian Finch (endemic)
9/0
No helminths recorded
Family ORIOLIDAE
414. Oriolus sagittatus (Latham)
Olive-backed Oriole
3/1 :N
. ? Microhadjelia spiralis (Mawson, 1968d)
syn. Cyrnea spiralis Mawson, 1968d
NT: PMM, 1968d, p.753
Microtetrameres oriolus Petrov & Tschertkova, 1950
NT: PMM, 1977, p.246
Microfilaria sp.
Qld: Bancroft, 1889, p.61
Qld: Cleland & THJ, 1912, p.435
415. Sphecotheres viridis Vieillot
Figbird
2/12 N
. Raillietina sphecotheridis (T:H. Johnston, 1914a)
Old: CIHUS
syn. Davainea sphecotheridis T.H. Johnston,
19l4a, p. 354
280)
REC. S, AUST. MUS. 19(15): 219-325
Qld: THS, 1914a, p.106
Railhetina sp.
Qld: Hall Exped. Rep. 1974, p.354
Dendrouterina sp.
Aust: BM(NH)
Biuterina sp,
Qld: BM(NH), CIHUS
. Microtetrameres sphecotheres Mawson, 1977
NT: PMM, 1977, p.253
Diplotriaena ? pungens (Schneider, 1866)
Qld: BM(NH)
Diplotriaena tridens (Molin, 1858)
? loc: Anderson, 1959, p.248
Onchocercidae, unidentified
Qld: CIHUS
Microfilaria sp.
Qld: Mackerras, 1962, p.439
Family DICRURIDAE
416. Dicrurus hottentottus (Linné)
Spangled Drongo
1/1: N
. Plagiorchis nisbetii (Nicoll, 1914a)
syn, Lepoderma nisbetii Nicoll, 1914a
Qld: Nicoll, 1914a, p.341
Prosthogonimus vitellatus Nicoll, 1914a
Qld: Nicoll, 1914a, p.345
. Viguiera chibiae Mawson, 1968d
NT: PMM, 1968d, p.763
Geopetitia chibiae Mawson, 1966
NT: PMM, 1966, p.716
Diplotriaena sp.
Qld: Mackerras, 1962, p.432
NT: AHC
Hamatospiculum chibiae
Mawson, 194la
Qld: THJ & PMM, 194la, p.l4
Microfilaria sp.
Qld: Bancroft, 1889, p.61
Qld: Mackerras, 1962, p.439
T.H. Johnston &
Family PARADISAEIDAE
417. Ptilonorhynchus violaceus (Vieillot)
Satin Bowerbird (endemic)
. Unidentified
Qld: Cleland, 1922, p.l07 (AHC)
. Microfilaria sp.
Qld: Mackerras, 1962, p.439
NSW: CIHUS
418. Sericulus chrysocephalus (Lewin)
Regent Bowerbird (endemic)
1/0
. Unidentified
Qld: Cleland, 1922, p.107 (AHC)
Qld: AHC
. Microfilaria sp.
Qld: Bancroft, 1889, p.6l
. Mediorhynchus corcoracis T.H.
August, 1986
419. Chlamydera maculata (Gould)
Spotted Bowerbird (endemic)
1/0
. Paricterotaenia chlamyderae (Krefft, 1873)
syn. Taenia chlamyderae Krefft, 1873
NSW: Krefft, 1873, p.224
Hymenolepis sp.
Qld: AHC
420. Ailuroedus melanotis (Gray)
Spotted Catbird
No helminths recorded
421. Ailuroedus crassirostris (Paykull)
Green Catbird
Lutztrema ailuroedi Angel & Pearson, 1977
Qld: LMA & Pearson, 1977, p.127
Qid: AHC
. Unidentified
Qld: Cleland, 1922, p.l07 (AHC)
. Filarioidea, unidentified
Qld: AHC (broken specimens)
422. Ptiloris paradiseus Swainson
Paradise Riflebird (endemic)
1/1:TCN
. Unidentified
SA (cage): not kept (eggs only, in pancreas)
. Unidentified
SA (cage): AHC
. Microtetrameres sp,
SA (cage): PMM, 1977, p.258
Family CORCORACIDAE
423. Corcorax melanorhamphos (Vieillot)
White-winged Chough (endemic)
10/8 : Tl C6 N2 A6
Leucochloridium australiense T.H. Johnston &
Cleland, 1938
SA: THJ & Simpson, 1940a, p.119
. Dayainea sp.
SA: AHC
Hymenolepididae, unidentified
NSW: CIHUS
Unidentified
Qld, NSW, SA: Cleland, 1922, p.106 (AHC)
Qld, Vic, SA: AHC
. Viguiera sp.
SA: AHC
Microtetrameres helix Cram, 1927
Vic, SA: PMM, 1977, p.245
Microfilaria sp.
Qld: Cleland & THJ, 1912, p.436
? NSW: Mackerras, 1962, p.439
Johnston &
Edmonds, 1951
HELMINTHS FROM AUSTRALIAN BIRDS 281
Qld, NSW, SA: THJ & Edmonds, 1951, p.1
Vic: AHC
syn. Echinorhynchus sp.
NSW, SA: Cleland, 1922, p.108
Unidentified
NSw: CIHUS
424. Struthidea cinerea Gould
Apostle bird (endemic)
1/0
. Microtetrameres sp.
SA: AHC
Splendidofilaria sp.
NSW: AHC
Microfilaria sp.
Qld: Cleland & THJ, 1912, p.435 (two species
noted)
Family GRALLINIDAE
425. Grallina cyanoleuca (Latham)
Australian Magpie-lark
27/19 : T10 C12 NIL A4
. ? Plagiorchis maculosus (Rudolphi, 1802)
SA: LMA, 1959, p.265
Echinostomatinae [spines?]
SA: AHC
Prosthogonimus sp.
Qld: Nicoll, 1914b, p.106
Lecithodendriidae, unidentified
SA: AHC
Neodiplostomulum sp.
SA: Dubois & LMA, 1972, p.212
(metacercaria)
Strigea nicolli Dubois, 1937a
syn. Strigea suttoni Dubois, 1937a
Qld; Dubois, 1937b, p.237
, Hymenolepis sp.
Qld: BM(NH)
Unidentified
SA: AHC
. Capillaria grallinae TH. Johnston & Mawson,
1945b
SA: THJ & PMM, 1945b, p.247
Microfilaria sp. Mackerras, 1962
Qld: Mackerras, 1962, p.437
. Echinorhynchus sp.
Qld: THJ, 1912b, p.110; 1914a, p.110
Mediorhynchus garruli (Yamaguti, 1939b)
NSW: CIHUS
Mediorhynchus sp.
SA: AHC
Oncicola pomatostomi (7.H. Johnston & Cleland,
1912)
NT: AHC (cyst)
Family ARTAMIDAE
426, Artamus leucorhynchus (Linné)
White-breasted Woodswallow
. Unidentified
Qld: Cleland, 1922, p.106 (AHC)
. Microfilaria sp.
Qld: Cleland & THJ, 1912, p.432
427. Artamus personatus (Gould)
Masked Woodswallow (endemic)
. Unidentified
SA: Cleland, 1922, p.106 (AHC)
. Microfilaria sp.
Qld: Mackerras, 1962, p.437
428. Artamus superciliosus (Gould)
White-browed Woodswallow (endemic)
. Unidentified
NSW: AHC
. Microfilaria sp.
London, UK. (cage): Plimmer, 1916, p.85
429, Artamus cinereus Viecillot
Black-faced Woodswallow
§/2:N
’. Unidentified
NSW: Cleland, 1922, p.106 (AHC)
. Acuaria petterae Mawson, 1972
SA, NT: PMM, 1972, p.144
Microfilaria sp.
Qld: Cleland & THJ, 1912, p.432
. Oncicola pomatostomi (T.H. Johnston & Cleland,
1912)
syn. Oligacanthorhynchus pomatostomi (T.A.
Johnston & Cleland, 1912)
Qld: Hall Exped. Report, 1974, p.355 (cyst)
430. Artamus cyanopterus (Latham)
Dusky Woodswallow (endemic)
4/1:N
. Unidentified
NSW: Cleland, 1922, p.106 (AHC)
. Microfilaria sp.
NSW: Mackerras, 1962, p.437
Qld: Cleland & THJ, 1912, p.432
Spirurida, unidentified
SA: AHC (larva)
431. Artamus minor Vieillot
Little Woodswallow (endemic)
2/0
No helminths recorded
282 REC, 8, AUST. MUS, 19(15); 219-325
Family CRACTICIDAE
432. Cracticus torquatus (Latham)
Grey Butcherbird (endemic)
5/3: T1 N2
T. Brachylecithum Jatius Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.123
C. Choanolaenia fieldingi (Maplestone & Southwell,
1923)
syn. Monopylidium fieldingi Maplestone &
Southwell, 1923
Qld: Maplestone & Southwell, 1923, p.318
N, Capillaria sp.
SA: AHC
Microtetrameres cractici Mawson, 1977
SA: PMM, 1977, p.251
Diplotriaena epsilon T.H. Johnston & Mawson,
1940a
Qld: TH] & PMM, 1940a, p.360
Filaria (s..) sp, (? Filarioidea)
Qld: Bancroft, 1889, pp.59,62
Microfilarta sp.
Qld: Bancroft, 1889, p.61
Qld; Cleland & THJ, 1912, p.432
Qld: Mackerras, 1962, p.438
433. Cracticus nigrogularis (Gould)
Pied Butcherbird (endemic)
7/3: ™N
N. PAysaloptera sp.
NT: AHC
Acuaria petterae Mawson, 1972
NT: PMM, 1972, p.144
Microfilaria sp.
Qld: Cleland & TH4J, 1912, p.432
Spirurida, unidentified
NT: AHC (larva)
434, Gymnorhina tibicen (Latham)
Australian Magpie
77/56 + T9 ClO N45 A4
T.. Plagiorchis maculosus (Rudolphi, 1802)
SA: LMA, 1959, p.265
Prosthogoninius vitellatus Nicoll, \914a
SA; LMA, 1973, p.859
Brachylecithum latius Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.123
Brachylecithum sp.
SA: LMA & Pearson, 1977, p.124
Strigea nicolli (Dubois, 1937a)
NSW: Dubois & LMA, 1972, p.199
Unidentified
NSW: lost
C. Dilepididae, unidentified,
SA: AHC
Unidentified
Qld; ACT, Vic, Tas, SA, WA: AHC
N. Capillaria gymnorhinae T.H. Johnston &
Mawson, 1947
SA: THJ & PMM, 1947, p.548
August, 1986
ACT, SA: AHC
Syngamus trachea (Montagu, 1811)
Vic: Harrigan & Arundel, 1978, p.360
Tas: Obendorf, 1984, p.7
Porrocaecum streperde T.H, Johnston & Mawsan,
1941d
ACT, SA: AHC
Porrocaecum wui Hs-f9-, 1933
ACT: BM(NH)
Oxyspirura streperae T.H. Johnston & Mawson,
1941a
ACT: BM(NH)
Microtetrameres gymnorhinae Mawson, 1977
ACT, SA, NT; PMM, 1977, p.251
Acuaria petterae Mawson, 1972
ACT, SA, NT: PMM, 1977, p.251
Qld: AHC
Xenocordon gymnorhinis
Robertson, 1983)
Qld, NSW, Vic, SA: AHC
syn. Cheilospirura gvninorhinis de Chaneet &
Robertson, 1983
SA, WA: de Chaneet & Robertson, 1983, p.143
Diplotriaena clelandi (TH. Johnston, 1912b)
syn. Filaria clelandi T.H. Johnston, 1912b
Qld: THJ, 1912b, p.110
Splendidofilariinae, unidentified
SA: AHC
Microfilaria gymnorhinae Gilruth, Sweet & Dodd,
1910
SA: Gilruth, Sweet & Dodd, 1910, p.236,
Mackerras (1962) notes that G, S. & D.
describe 3 species under this heading.
Microfilaria sp.
Qld; Bancroft, 1889, p.61
NSW: Cleland & THJ, 1910, p.107
NSW; Mackerras, 1962, p.438
London (cage): Plimmer, 1912, p.408
Spirurida, unidentified
SA: AHC (larva)
A. Prosthorhynchus cyvlindraceus (Goeze, 1782)
Vic, Tas: AHC
Oncicola pomatostomi (T.H, Johnston & Cleland,
1912)
SA, NT: AHC
syn. Oligacanthorhynchus pomatostomi T.H.
Johnston & Cleland, 1912
SA: Hall Exped. Report, 1974, p.355 (cyst)
(de Chaneet &
435. Strepera graculina (White)
Pied Currawong (endemic)
T. Dicrocoeliinae, unidentified
Qld: AHC
C. Hymenolepididae, unidentified
NSW: CIHUS
Unidentified
Qld: Cleland, 1922, p.107 (AHC)
Qld, NSW: AHC
N, Porrocaecum streperae T.H. Johnston & Mawson,
194]d
SA: AHC
Oxyspirura streperae TH. Johnston & Mawson,
194]a
HELMINTHS FROM AUSTRALIAN BIRDS
Qld: AHC
Acuaria streperina T.H. Johnston & Mawson, 1941d
Qld: AHC
Diplotriaena alpha T.H. Johnston & Mawson,
194la
SA: THJ & PMM, 1940a, p.359
syn. Filaria’
SA: Cleland, 1922, p.108
Paralemdana clelandi T.H. Johnston & Mawson,
1940a (sp. ing. in CIH keys)
NSW: THI & PMM, 1940a, p.356
syn. Filuria’
NSW: Cleland, 1922, p.108
Aprocta boulengeri Bain & Mawson, 1981
Qld: Bain & Mawson, 1981, p.269
Microfilaria sp.
Qld: Bancroft, 1889, p.6l
436, Strepera fuliginosa (Gould)
Black Currawong (endemic)
N. Oxyspirura streperae T.H. Johnston & Mawson,
1941a
SA: THJ & PMM, 194la, p.1l
Acuaria streperina T.H. Johnston & Mawson, 194]d
SA: THJ & PMM, 194Id, p.254
Diplotriaena sp.
Vic: AHC
Microfilaria sp.
London, U.K. (cage); Plimmer, 1912, p.408
437, Strepera versicolor (Latham)
Grey Currawong (endemic)
8/5: TL C2 N4
T. Echinostomatinae [spines ? (35(5))]
SA: AHC
Brachylecithum parvum (S.J. Johnston, 1917)
syn. Lyperosomum parvum §.J. Johnston, 1917
NSW: SJJ, 1917, p.223
C, Unidentified
NSW, SA: AHC
N. Porrocaecum streperae T.H. Johnston & Mawson,
1941d
NSW: THJ & PMM, 1941d, p.253
Geopetitia streperae Mawson, 1966
SA: PMM, 1966, p.715
Microtetrameres streperae Mawson, 1977
SA; PMM, 1977, p.251
Acuaria streperina T.H. Johnston & Mawson, 1941d
SA: TH] & PMM, 1941d, p.254
Family CORVIDAE
For note on host spp. in this family, see p. 221.
438. Corvus coronoides Vigors & Horsfield
Australian Raven (endemic)
2/2: Tl C2 N2
T. Prosthogonimidae, unidentified
Qld: AHC
Brachylecithum latius Angel & Pearson, 1977
SA: LMA & Pearson, 1977, p.123
C. Davainea sp.
NSW: THJ, 1912b, p.112
Unidentified
NSW: Cleland, 1922, p.107 (AHC)
EA: AHC (? host sp.)
SA: AHC
. Microtetrameres helix Cram, 1927
SA: PMM, 1977, p.245
Acuaria anthuris (Rudolphi, 1819)
SA: PMM, 1972, p.141
Aprocta bakeri Bain & Mawson, 1981
SA; AHC
439. Corvus tasmanicus Mathews
Forest Raven (endemic)
48/48 : T2 Cl4 N48 Al
. Echinostomatinae [spines missing]
Tas: AHC
Brachylecithum parvum (S.J, Johnston, 1917)
Tas: LMA & Pearson, 1977, p.118
Dicrocoeliidae, unidentified
Tas: Munday & Green, 1972, p.2
*, Unidentified
Tas: Cleland, 1922, p.107 (AHC)
Tas: Munday & Green, 1972, p.4
Tas: AHC
. Capillaria sp.
Tas: AHC
Syngamus trachea (Montagu, 1811)
Tas: Obendorf, 1982, p.7
Microtetrameres helix Cram, 1927
Tas: PMM, 1977, p.245
syn. Microtetrameres sp.
Tas: Munday & Green, 1972, p.1l
Acuaria anthuris (Rudolphi, 1819)
Tas: PMM, 1972, p.141
Tas: Munday & Green, 1972, p.1]
Spirurida, unidentified
Tas: AHC (larva)
. Mediorhynchus corcoracis T.H. Johnston
Edmonds, 1951
Tas: AHC
440. Corvus mellori Mathews
Little Raven
16/13 : T1 C7 Nil A6
. ? Tanaisiinae, unidentified
SA: AHC
Echinostomatinae [spines (37(5))]
Vic: AHC
’, Hymenolepis sp.
Vic: AHC
Unidentified
Vic, SA: AHC
. Capillaria sp.
Vic, SA: AHC
Porrocaecum sp.
Vic: AHC
Microtetrameres helix Cram, 1927
284
SA: PMM, 1977, p.245
Vic: AHC
Acuaria anthuris (Rudolphi, 1819)
SA: PMM, 1972, p.141
Vie: AHC
Aprocta bakeri Bain & Mawson, 1981]
Vie: AHC
Spirurida, unidentified
SA: AHC (larva)
A. Mediorhynchus corcoracis T.H. Johnston &
Edmonds, 195]
SA: AHC
44). Corvus bennetti North
Little Crow (endemic)
2/2 : N2
C. Unidentified
SA; AHC
N. Microletrameres helix Cram, 1927
NT: PMM, 1977, p.245
Acuaria anthuris (Rudolphi, 1819)
NT: PMM, i972, p.141
‘Filarial worms’
Qld: Mackerras, 1962, p.435
A. Mediorhynchus corcoracis T.H. Johnston &
Edmonds, 1951
SA: THJ & Edmonds, 1951, p.]
442. Corvus orru Bonaparte
Torresian Crow
8/6; Tl Cl N6
T. Dicrocoeliinae, unidentified
Qld: AHC
C. Ruillfetina corvina (Fuhrmann, 1905)
Qld: BM(NH)
Unidentified
Qld: Cleland, 1922, p.107 (AHC)
NT; AHC
N. Microtetrameres helix Cram, 1927
NT; PMM, 1977, p.245
Acuaria anthuris (Rudolphi, 1819)
NT; PMM, 1972, p.141
Qld: AHC
syn. A. corvicola T.H. Johnston & Mawson,
194la
SA: THJ & PMM, 1941a, p.10
Diplotriaena beveridgei Bain & Mawson, 1981
Qld: Bain & Mawson, 1981, p.278
Diplotriaena flabellata (Linstow, 1888)
Qld: Ogden, 1967, p.505
Qld: Hall Exped. Report, 1974, p.354
syn. D. beta T.H. Johnston & Mawson, 1940a
Qld: THJ & PMM, 1940a, p.360
Diplotriaena sp.
Qld: AHC
Aprocta bakeri Bain & Mawson, 1981
Qld: Bain & Mawson, 1981, p.269
Aproeta corvicola T.H, Johnston & Mawson, 1940a
SA: THJ & PMM, 1940a, p,358 (? host sp.)
Filaria (s.L.) sp. (? Filarioidea)
Qld: Bancroft, 1889, p.62
REC. $8. AUST. MUS, 19(15): 219-325
August, 1986
Microfilaria sp.
Qld: Bancroft, 1889, p.6l
Qld: Cleland & THJ, 1912, p.435
NSW; Cleland & THJ, 1910, p,108
443, Corvus sp.
C, Hymenolepis sp,
SA: AHC
N. Synhimantus sp.
SA: AHC
A. Mediorhynchus corcaracis T.H.
Edmonds, 1951
SA: AHC
Mediorhynchus sp.
NT: AHC
Johnston &
List 2. HELMINTHS IN TAXONOMIC ORDER,
AND THEIR HOSTS
In this section the host names are listed after the
name of each helminth. Synonyms of the helminths
are given if they have been cited elsewhere in the work,
but the list is in no way intended as a complete
synonymy of any species. Species are listed
alphabetically in their families or subfamilies, which
are in taxonomic order, or within each superfamily
within each order (Nematoda). Specimens not
identified further than to Trematoda, Cestoda,
Nematoda, or Acanthocephala have been omitted from
this list.
PHYLUM PLATYHELMINTHES
Class Trematoda
Order Digenea
Family SCHISTOSOMATIDAE Stiles & Hassall, 1898
Schistosomatidae, unidentified, Blair & Otteson,
1979,
Anseranas semipalmata, 69; Dendrocygna
arcuata, 70; D. eytont, 71; Anas superciliosa, 77;
A, gibberifrons, 78; A. rhynchotis, 80; Aythya
australis, 82; Chenonetta jubata, 83.
Schistosomatidae, unidentified.
Podiceps cristatus, 3; Larus novaehollandiae, 163.
Austrobilharzia terrigalensis S.J. Johnston, 1917,
Egretia sacra, 58; Larus novaehollandiae, 163;
Sterna bergii, 171; Melopsittacus undulatus, 209,
Bilharziellinae, unidentified.
Himantopus leucacephalus, 142; Cladorhynchus
leucocephalus, \43.
Dendritobilharzia sp.
Pelecanus conspicillatus, 37,
Gigantobilharzia sp.
Larus novaehollandiae, 163.
Ornithobilharzia sp.
Pelecanus conspicillanes, 37; Chiidonias hybrida,
164,
Schistosomatinae, unidentified.
Pelecanus conspicillatus, 37,
HELMINTHS FROM AUSTRALIAN BIRDS TRS
Trichobilharzia australis Blair & \slam, 1983.
Anas superciliosa, 77.
Thichobilharzia sp., Blair & Otteson, 1979.
Anas superciliosa, 77.
Trichobilharzia sp., Bearup, 1957.
Anas gibberifrons, 78.
Trichobilharzia sp., Bearup & Langsford, 1966.
Himantopus leucocephalus, 142.
Trichobitharzia sp., TH. Johnston, 1941,
Cygnus atratus, 72.
Trichobilharzia sp.
Himantopus leucocephalus, 142; Melopsitiacus
undulatus, 209.
Family NOTOCOTYLIDAE Liihe, 1909
Notocotylidae, unidentified.
Dendrocygna arcuata, 70; Cygnus atratus, 72:
Stictonetta naevosa, 73, Anas superciliosa, 77; A.
gibberifrons, 78; Malacorhynchus membranaceus,
81; Oxyura australis, 86; Biziura lobata, 87;
Erythrogonys cinctus, 134; Cladorhynechus
leucocephalus, 143; Reeurvirostra novae-
hollandiae, 144; Calidris acuminata, 157,
Notocotylid, ? sp. B of $.J, Smith, 1981.
Cygnus atratus, 72.
Catatropis gallinulae T.H. Johnston, 1928.
Gallinula tenebrosa, 121,
Catatrapis sp.
Porzana pusilla, 116; P. fluminea, 117; Gallinula
tenebrosa, 120.
Notocotylus attenuatus (Rudolphi, 1809).
Anseranas semipalmata, 69; Cygnus atratus, 72;
Anas superciliosa, 77; Nettapus pulchellus, 85;
Burhinus magnirostris, 128; Vunellus miles, 131;
Calidris acuminata, 157,
Paramonostomum bursae Smith & Hickman, 1983b.
Anas supercitiosa, 77.
Paramanostomum caeci Smith & Hickman, 1983b,
Poliocephalus poliocephalus, 4, Cygnus atratus,
72; Anas superciliosa, 77.
Paramonostomum sp.
Malacorhynchus membranaceus, 81; Gallinula
(enebrosa, 12).
Family CYCLOCOELIDAE Stossich, 1902
Cyclocoelidae, unidentified,
Cyenus atratus, 72; Anas gibberifrons, 78, Aythya
australis, 82; Leucosarcia inelanoleuca, 117;
Recurvirostra novaehollundiae, \44; Tringa
elareola, 146, Calidris canutus, 155; Lichmera
indistineta, 378,
Allopyge antigones $,.), Johnston, 1913, syn.
‘Monostome' of Nicoll, 1914c.
Grus rubicundus, 124.
Cyelocoelum jaenschi TH. Johnston & Simpson,
1940b,
Poliocephalus poliocephalus, 4; Tachybaptus
novaehollandiae, 5.
Cyelocoelum obscurum (Leidy, 1887).
Vanelluls miles, \31.
Cyclocoelum taxorchis §.J. Johnston, 1917.
Limosa lapponica, 154.
Cyclocoelum sp.
Malacorhynchus membranaceus, 81; Nettapus
coromandelianus, 84; Porphyrio porphyria, 122;
Limosa lapponica, 154.
Haematotrephus adelphus 8,5. Johnston, 1917, syn.
Monostomun sp., T.H, Johnston, 1910.
Himantopus leucocephalus, 142; Cladorhynchus
leucocephalus 143,
Haematotrephus consimilis Nicoll, 1914b,
vanellus miles, 131.
Haematotrephus sp.
Tringa nebularia, 149,
Hyptiasmus magnus §.J, Johnston, 1917, syi,
Monostomum sp., T.H. Johnston, 1910.
Cygnus atratus, 72.
Tvphlocoelum reticulare $.J. Johnston, 1913.
Anseranas semipalmata, 69,
Typhlocoelum sp.
Anas Superciliosa, 77; A. gibberifrons, 78.
Family EUCOTYLIDAE Skrjabin, 1924
Eucotyvie sp.
Ardea novaehollandiae, 53,
Tanaisiinae, unidentified,
Corvus mellori, 440,
Family PHILOPHTHALMIDAE Looss, 1899
Cloacitrema narrabeenense Howell & Bearup, 1967.
Larus noyaehollandiae, 163.
Parorchis acanthus (Nicoll, 1906),
Larus novaehollandiae, 163.
Philophihalmus burrili Howell & Bearup, 1967,
Larus noyaehollandiae, 163.
Philophthalmus sp,
Dramaius novaehollandiae, |; Ardeotis australis,
126.
Family BRACHYLAEMIDAE Joyeux & Poley, 1930
Brachylaima pulchellum (S.J, Johnston, 1917),
syn. Harmostomum pulehellum 8.J. Johnston,
1917,
Leucosarcia melanoleuca, 187.
Brachylaima sp.
Leucosarcia melanoleuca, \87.
kamily LEUCOCHLORIDIIDAE Poche, 1907
Leucochloridium australiense TH. Johnston &
Cleland, 1938.
Pamatostomus superciliasus, 311; Corcorax
melanorhamphos, 423,
Leucechloridiunt sp.
Rallus philippensis, W4,
Urotocus sp.
Manorina melanocephala, 360.
286 REC, S, AUST, MUS. 1913); 219-325
Family OMPHALOMETRIDAE Looss, 1899
Omphalometridae, unidentified.
Cladorhynchus leucocephalus, 143,
Dolichosacculus solecarius (S.J, Johnston, 1917),
syn. Delichesaccus solecarius 5,J. Johnston,
1917,
Phalacrocorax melanoleucas, 47.
Family PLAGIORCHIIDAE Liihe, 1901
Plagiorchis maculesus (Rudalphi, 1802), syn,
clelandt S.J, Johnston, 1917, PR spatulatus S.J.
Johnston, 1917.
Hirundo neoxena, 266, Cecropis ariel, 269;
Anthus novaeseelandiae, 270; Rhipidura
Sulizinosa, 302; R. leucophrys, 304, Pomato-
stomus superciliasus, 3)\; Pardalotus striatus, 399;
Grallina vvanoleuca, 425; Gymnorhina trhicen,
434.
Plagiorchis nisbetii (Nicoll, 1914a), syn, Lepodermu
nisbetii Nicoll, 1914a,
Dicrurus hottentottus, 416.
Plagiorchiidae, unidentified, syn. Lepodermatidae,
unidentified, Nicoll, 1914b,
Vanellus miles, 131.
Family ECHINOSTOMATIDAE Looss, 1899
Echinostomatidae, unidentified, Bradley 1926, 1927.
Poliocephalus poliocephalus, 4,
Echinostomatidac, unidentified, Munday & Green,
1972,
Pluyialis dominica, 133.
Echinostomatidae, unidentified-
Tachybaplus novaehallundiae, 5; Pelecanus
conspicillatus, 37; Phalacrocorax melanoleucos,
47; Egretta garzetta, 56, Cygnus atraius, 72; Anas
gibberifrons, 78; A. rhynchotis; 80; Avrhya
ausiralis, 82; Chenonetta jubata, 83, Biziura
lobata, 87: Coturnix ypsilophora, 109; Gallinula
ventralis, 120; frediparru gallinacea, 127;
Eryihrogonys vinetus, 134; Charadrius melanops,
140; Tringa hypoleucos, 148; Calidris acuminata,
(57; Chlidenias hybrida, 164.
Acanthoparyphium marilae Yamaguti, 1934,
Charadrius leschenaultii, 138; Limosa lapponica,
154; Calidris canutus, 185; Calidris tentuirasiris,
156,
Acanthoparyphium spinulosum 3.5, Johnston, 1917,
Pluvialis dominica, 133; Charadrius leschenaultii,
138; ©. euficapillus, 139; Larus novaehollandiae,
163.
Acanthoparyphium squatarolae Yamaguti, 1934.
Limeasa lapponica, 154; Calidris tenuirostris, 156.
Acanthoparyphium sp.
Cladorhynchus feucocephalus, 143; Lichmera
indistincta, 378.
Chaunocephalus férax (Rudalphi, 1795),
Xenorhynchus asiaticus, 63.
Cloeophora sp.
Calidris eanutus, 185.
Echinachasmus pelecani TH, Johuston & Simpson,
1944.
Auelesy, J986
Pelecanus conspicillatus, 37,
Echinechasnius sp.
Phalacrocorax sulcirostris, 46,
Evhinoparyphium ellist (TH. Johnston & Simpsan,
1944), syn. & sp., Verma, 1936.
Cyenus atratus, 72, Biziura lobata, 87,
Echinoparyphium gizzardai Verma, 1936,
Cvenus atratus, 72.
Echinoparyphium harvevanum S.J. Johnston, 1917.
Microece leucaphaea, 287,
Evhinoparyphium oxyurum §.).. fohnston, 1917.
Eeretta alba, 55.
Echinoparyphium phalacrecoracis Yamaguti, 193¥a.
Leucocarbo fuscescens, 43; Phalacracorax carbo,
44; P. sulcirostris, 46; RP melanoleucas, 47.
Echinonaryphium sp,
Stictonelta ngevosa, 73; Tadorna tadornoides, 75;
Tringa hypoleucos, 148, Larus novaehollandiae,
163; Ninox nevaeseelandiae, 242; Podaryus
strigoides, 246; Dacelo novaeguineae, 253.
Echinestoma acuticauda Nicoll, 1914b.
Threskiornis aethiopica, 65, T. spinicollis, 66.
Echinostoma australasianum Nicoll, 1914a,
Grus rubicundus, \24.
Echinostoma australe TH. Johnston, 1928.
Gallinula tenebrosa, 121.
Evhinostoma bancraftt T.A. Johnston, 1928.
Gallinula tenehrosa, 121.
Echinostoma elongaiunr Nicoll, 1914a,
Podareus strigoides, 246.
Echinostoma emollitum Nicoll, 1914b.
Centropus phasianus, 239.
Echinostoma hilliferum Nicoll, 1914b,
Gallinula ventralis, 120; Porphyrio porphyrio, \22;
Fulica atra, 123.
Echinostoma ignayvum Nicoll, 1914b.
Burhinus magnirostris, (28; Vanellus miles, 131.
Echinostoma minimum Verma, 1936,
Cygnus atratus, 72.
Echinostoma revolutum (Froelich, 1802), syn.
‘echinostome larva’, Bradley, 1927, Echinostoma
sp,, TH. Johnston 1912b,
Anseranas semipalmata, 69; Cygnus atratus, 72;
Tadorna tadornoides, 75: Anas superciliosa, 77;
A. rhynchotis, 80; Nettapus pulchellus, 85; Biziura
lohata, 87; Gallinula tenebrosa, \21.
Echinostoma sp., TA. fohnston, 1910.
Piuvialis daminica, 133.
Echinostoma sp., TH. Johnston, 1912b.
Egretta alba, 55,
Lchinostoma sp.
Pohocephalus poliacephalus, 4; Egretta alba, 55;
Cygnus atratus, 72; Anas gibberifrons, 78;
Malacorhynachus membranaceus, 81; Porzana
pusilla, 116; CGallinula tenebrosa, 12\; Porphyria
porphyrie, 122; Vanellus miles, 131; Tringa
hypolencos, 148; Larus novaehoallandiae, 163,
Hydraprogne caspia, 166; Ninox novaeseelandiae,
242.
‘Echinostome larva’ of Bradley, 1926, 1927.
Fulica wira, 123,
Echinostomalinae,. iinidentified.
Phalacrocorax suleirostris, 46; Dendrocyena
HELMINTHS PROM AUSTRALIAN BIRDS a7
arcuata, 70; Cygnus atratus, 72; Anas superciliosa,
77; A, gibherifrons, 78; A. rhynchotis, 80; Oxvura
australis, 86; Biziura lobata, 87; Haliastur
sphenurus, 91; Porzana pusilla, 116; P fluminea,
7; Porphyrio porphyrio, 122, Calidris
acuminata, 137; Larus novaehollandiae, 163;
Hydroaprogne caspia, \66; Tyto alba, 244, Grallina
cvanoleuca, 425; Strepera versicolor, 437, Corvus
fasmanicus, 439; ©, mellori, 440.
Episthmium prosthovitellatum (Nicoll, 1914b), syn.
Echinochasmus prosthovitellatum Nicoll, 1914b.
Falco berigora, 104.
Euparyphium longitestis Verma, 1936.
Cygnus atratus, 72.
Himasthla harrisoni S.1, Johnston, 1917, syn.
Echinostoma sp., T.H. Johnston, 191)2b,
Echinostoma (Acanrhochasmus) sp., T.W.
Johnston, 1916, 1918.
Numenius madagascariensis, \45.
Himasthla kusasigi Yamaguti, 1939a.
Calidris tenuirostris, \56,
Himasthla megacotvla Yamaguti, 1939a.
Limose lupponica, 154.
Himasthlinac, unidentitied.
Cualidris canutus, 155; Podargus strigoides, 246;
Lichmera indistincta, 378.
Hypoderaeum conoideum (Bloch, 1782),
Anas castanea, 79.
Hypoderaeum sp.
Anas superciliosu, 77; A, gibberifrons, 78; A,
castaneda, 79.
Nephrostomutm sp.
Fereia garzetta, 56.
Parvphostomun radiatum (Dujardin, 1845), syn.
Echinochasnius tenuicollis 5.3, Johnston, 1917;
Paryphostamumn tennicolle (S.J. Johnston,
1917); Paryphostomum phalacrocoracis Goss,
1940,
Leucocarbo fuscescens, 43; Phalacrocorax carbo,
44; P suleirostris, 46; PB melanoleucos, 47,
Patagifer acuminatus $.J, Johnston, 1917, syn.
Echinostoma sp., TH. Johnston, 19126,
Paiazifer bilabus of TH, Johnston, 1916, not
(Rudolphi, 1819),
Threskiornis aethiopica, 65,
Pautugifer bilobus (Rudolphi, 1819).
Plegadis faleinellus, 64; Threskiornis spinicollis,
66; Platalea regia, 67.
Patagifer Jraternus 8.J. Johnston, 1917.
Fvreita alba, 55.
Patagifer sp.
Poliocephulus poliocephalus, 4, Threskternis
aethiopica, 65, Platalea flavipes, 68; Anas
superciliosa, 77.
Petasiger australis T.A. Johnston & Angel, [941b,
Podiceps cristatus, 3, Poliocephatus
poliocephalus, 4; Tachybaptus novaehollandiae, 5.
Petusiger exaeretus Dietz, 1909,
Leucocarho fuscescens, 43, Phalacrocorax carbo,
44. P. sulvirosiris, 46; RP melanoleucas, 47.
Petasiger sp.
Tivo alba, 244.
Stephanoprora sp.
Larus novuehollandiae, \63.
Fermalrema loneiestis (Verma, 1936), Syn.
Euparyphium longitestis Verma, 1936.
Cyenus atratus, 72.
Family PSILOSTOMIDAE Looss, 1900
Psilochasmus oxyurus (Creplin, 1825).
Poliocephalus poliocephalus, 4 Cvgnus atratus,
72; Anas superciliosa, 77, Fulica atra, 123.
Psilochasmus sp.
Stictanetta naevosa, 73; Cereapsis
novaehollaundiae, 74, Biziura lobata, 87, Falco
herigora, \Od,
Psilostomum sp. A, of Sl. Smith, 198].
Poliocephalus poliovephalus, 4, Cygnus utralus,
72s
Psilostomum sp. B, Of S.J. Smith, 1981,
Poliocephalus poliocvephalus, 4; Cygnus atratus,
72.
Psilostomum sp.
Crenus atratus, 72; Sticfoneita naevosa, 73; Anas
superciliosa, 77.
Family PROSTHOGONIMIDAK, Lihe, 1909
Prosthogonimidac, unidentified.
Corvus coronaides, 438.
Prosthovonimus sp., Nicoll, 1914b,
Grullina cyanaleuca, 425.
Prosthogonimus sp., unidentitied.
Anas superciliosa, 77, Grus rubicundus, 124;
Larus novaehollandiue, 163; Rhinpidura
leucophrys, 304, Acanthiza pusilla, 336.
Cylindrotrema cveni Angel, 1973,
Cygnus arratus, 72.
Mawsonotrema eudvptulae Angel, 1973.
Fudyptula minor, 11.
Prosthegenimus vitellants Nicoll, 8ida.
Phalacrocorax melanoleucos, 47; Chlidanias
hybrida, 164; Dicrura horrentotrus, 416;
Gyinnorhina tibicen, 434.
Family DICROCOELHDAE Looss, 1899
Dicrocoeliidae, unidentified, Koch & Duhamel, 1982.
Caculua galerita, 196.
Dicrocoeliidae, unidentified, Munday & Green, 1972-
Corvus fasmanicus, 439,
Dicrocoeliidae, unidentified.
Caracina panuensis, 272.
Brachylecithum dacélonis Angel & Pearson, 1977.
Dacelo novaeguineae, 253.
Bruchylecithum harrisoni (S.1. Johnston, 1917), syn.
Lyperosomum harrisoni $A. Johnston, 1907,
Ninox novaeseelandiue, 242.
Rrachylecithum latius Angel & Pearson, 1977,
Cracticus forquatus, 432; Gymnerhina tibicer,
434, Corvus coronoides, 438.
Brachylecithum mezastomum (S.J. Johnston, }917),
syn. Lyperosomum megastomum S.J. Johnston,
1917.
288 REC. S. AUST. MUS, 19(15); 219.325
Sterna bergii, 171,
Brachylecithum parvum (S.J, Johnston, 1917), syn.
Lyperosamum parvum §.J. Johnstan, 1917.
Cecropis nigricans, 268; Strepera versicoler, 437;
Corvus tasmanicus, 439,
Brachylecithum podargi Angel & Pearson, 1977,
Podurgus strigoides, 46.
Brachylecithum sp.
Gymnorhina tibicen, 434,
Dicrocoeliinac, unidentified.
Cacatua galerita, 196; Strepera graculina, 435;
Corvus orru, 442,
Lutztrema ailuroedi Angel & Pearson, 1977.
Strepera versicolor, 421.
Pancreatrema meliphagae Angel & Pearsan, 1977.
Lichenostomus ornatus, 37\,
Platynosomum proxillicens (Canavan, 1937), syn,
Dicroceelium proxillicens Canavan, 1937;
Platvnosomum fallax Heidegger & Mendhein,
1938a; P ventraplicatum Heidegger &
Mendheim, 1938b.
Cacutua galerita, 196.
Platynotrema biliosum Nicoll, 19L4b,
Threskiornis aethiopica, 65;
magnirostris, 128.
Platynotrema jecoris Nicoll, 1914b.
Burhinus magnirostris, 128.
Plaivnotrema sp.
Tringa hypoleucos, 148; Limosa lapponica, 154.
Proacetabulorchis dogieli Belopolskaya &
Bychovskaya-Pavlovskaya, 1954.
Ardea novaehollandiae, 53.
Skrjabinosamum mawsoni Angel & Pearson, 1977.
Turnix castanota, 110; Manorina flavigula, 362.
Skrjabinesamum pomatostomi Angel & Pearson,
1977.
Pomutostamus superciliasus, 311.
Skrjabinosomum sp., Angel & Pearson, 1977.
Microeca leucophaea, 287.
Burhimus
Family LATEROTREMATIDAE Yamaguti, 1958
Laterotrema sp.
Pomatostomus superciliosus, 311.
Family EUMEGACETIDAE Travassos, 1922
Eumezacetes sp.
Eurostopodus mystacalis,
leucaphrys, 304.
249: Rhipidura
Family LECITHODENDRIIDAE Liihe, 1901
Lecithodendriidae, unidentified.
Rhipidura leucophrys, 304; Anthochaera
carunculata, 350; Grallina cyanoleuca, 425.
Pleuropsolus sp,
Cuculus flabellifermis, 234.
Family ORCHIPEDIDALE Skrjabin, 1913
Orchipedum sufflavum Nicholl, 1914b.
Platalea rexia, 67,
Aus! 1986
Family MICROPHALLIDAE Ward, 1901
Microphallidae, unidentified, Munday & Green,
1972,
Charadrius ruficapillus, 139; Calidris ruficollis,
158,
Microphallidae, unidentified.
Tachybaptus novaehallandiae, 5; Malacorhynchus
membranaceus, 81, Charadrius rufieapillus, 139,
Atriophallophorus coxiellae $.J. Smith, 1974.
Poliocephalus poliocephalus, 4; Fulica atra, 123;
Charadrius ritbricollis, 135; C. rufieapillus, 139.
Basantisia queenslandensis Deblock & Pearson,
1968a.
Dacelo novaeguinede, 253,
Endocotyle incana Belopolskaya, 1952.
Charadrius mongolus, 136, Calidris acuminata,
157,
Gynaecotyla brisbanensis Deblock & Pearson, 1968a.
Charadrius mangalus, 136, Larus novae-
hallandiae, 163.
Levinseniella hawensis 8.J. Johnston, 1917.
Pluyialis deminica, 133.
Levinseniella microovaia Belopolskaya, 1958.
Pluvialis dominica, 133; Charadrius inongolus,
136,
Levinseniella monodactyla Deblock & Pearson, 1970.
Charadrius mongolus, 136.
Levinseniella tasimaniae (3.1, Smith, 1974), syn.
Microphallus tasmaniae 3.J, Smith, 1974.
Poliocephalus paliacephalus, 4, Anas superciliosa,
77; Anas castanea, 79; Charadrius rubricollis, 135;
C ruficapillus, 139, C. melanops, 140,
Levinseniella sp,
Larus novaehollandtae, 163.
Maritrema calvertense 8.J, Smith, 1974.
Polivcephalus poliocephalus, 4, Anas supercitiosa,
77; A. castanea, 79; Charadrius rubricollis, 135;
C. melanops, 140.
Maritrema eroliae Yamaguti, 1939a.
Charadrius mongolus. 136; Sterna bergii, \71-.
Maritrema oocystum (Lebour, 1907).
Tachybaptus novuehollandiae, 3; Anhinga
melanogaster, 42; Ardea novaehollandiae, 53.
Maritremu sp.
Charadrius leschenaultii, 138; Tringa hypoleucos,
148; Limosa lapponica, 154; Sterna bergii, 171.
Microphallinae, unidentified.
Calidris acuminata, 157.
Micrephallus papillornatus Deblock & Pearson,
1969.
Pluvialis daminica, 133; Chanidrius mongolus,
136; Larus novaehollandiae, \63.
Microphallus vaginosus Deblock & Pearson, 1969.
Pluvialis dominica, 133.
Microphallus sp, Deblack & Pearson, 1969.
Charadrius mongolus, \36.
Family RENICOLIDAE Dollfus, 1929
Renicola sp,
Eudyptula mntinor, Wy Puffinus tenutrostris, 33; 8
gavid, 34; Pelecunus conspicillutus, 37;
HELMINTHS FROM AUSTRALIAN BIRDS 289
Leucecarho fuscescens, 43; Phalacracarax varius,
45; P. sp., 48) Eeretta alba, 55.
Family OPISTHORCHIIDAE Looss, 1899
Opisthorchis obsequens Nicoll, 1914b,
Threskiornis spinicollis, 66; Falco herigora, 104.
Opistharchis sp.
Circus approxrmans, 99,
Pachytrema caleulus Looss, 1907.
Larus novaehollandiae, 163, Chlidonias hybrida,
164.
Pachytrema sp. Sol. Smith, pers, comm,
Poliocephalus poliocephalus, 4,
Family HETEROPHYLDAE Ciurea, 1924
Heterophyidag, unidentified.
Eereita alba, 55; Limosa lapponica, 154, Calidris
ruficollis, 158, Hydroprogne caspia, 166.
Ascacoivle sp,
Pelecanus eonspicillatus, 37.
Crvplacatvle sp.
Puffinus lenutrostris, 33.
Galactosamum ungelae Pearson, 1973,
Eudyptula minor, \y Sula serrater, 38; Larus
novaehollandiae, 163, Hydroprogne cuspia, 166;
Sterna bergil, \7\.
Calactosamum bearupi Pearson, 1973,
Larus novaehollandiae, \63, Hydropragne caspia,
166; Sterna fuscata, 168; Sterna bengalensis, (72;
Anous minulus, 74,
Galactosomun) renincalian Pearson, 1973,
Pu/finus pacificus, 31; Sterna fuseata, 168; Anous
minutus, 174.
Cralactosonnin simuilacte Pearson, 1973,
Leucocarbo fuscescens, 43; Phalacrocorax yarius,
45.
Galactosomum ussuriense Oshmarin, 1963,
Larus novaehollandiae, 163; [ydroprogne caspia,
166; Srerna fuscuta, 168.
Galuctosomum sp.
Leucocurha Juscescens, 43; Hydroprogne caspia,
166,
flaplorchis puravanissimus Pearson & Ow-Yang,
1982, syn. 7/1. venisstinus of Pearson, 1964, in
parl, not Africa, 1938,
Pelecunus conspicillatus, 37; Phulacrocorax
sulcirosiris, 46; Nyericorax caledonicus, 60;
Haliastur sphenurus, 91.
Haplorchis pumilio (Looss, 1896).
Phalacrocorax melanoleucos, 47;
intermedia, 37,
Haplorchis sprenti Pearson, 1964, svn. Haplorchis
sp., Pearson, 1960.
Pelecanus conspicillatus, 37; Phalacrocorax
sulcirostris, 46; Po melanoleucos, 47; Ardeu
novaehollandiae, 33, Egretta alba, 55; E.
intermedia, 37.
Haplorchis vanissimus Africa, 1938.
Pelecanus conspicillatus, 37; Leucocarbo
Juseescens, 43, Phalacracorax carbo, 44; P
suleirostris, 46, Nyectivcerax caledonicus, 60;
Haliasitur sphenurus, 91.
Egretta
Haplorchis vokogawai (Katsuka, 1932),
Haliasiur sphenurus, 91,
Heterophyinae, unidentified,
Srerna hereit, \71.
Hererotestophyes sp.
Larus novaehollundiae, 163.
Proverovun yarium Onji & Nishio, L916.
Pelecanus conspicillatus, 37; Ardea
novuehollundiae, 53; Egretta alba, 35; E.
intermedia, 57; Nvericarax caledonicus, 60,
Haliustitr sphenurus, 9),
Procerovym sp., Pearson, 1960.
Pelecanus conspivillatus, 37. Phalucracorux
melanoleucos, 47.
Scaphanocephalus australis §.J. Johnston, 117, syn.
S. sp, Si. Johnston, 1914.
Haliueetus leucogaster, 95,
Stellanichasmus aspinosus Pearson, 1964, syn. S.
falcutus of Pearson, 1960, not Onji & Nishio,
1916.
Phalucrocorax inelanolencos, 47.
Srellanichasinus fulcatus Onqi & Nishio, 1916.
Nyetivorax calendonicus, 60; Maliastur sphenurus,
Ol.
Srictodora cuballeroi Martin, 1955.
Pelecanus canspicillutus, 37.
Stictodora diplacantha TH. Johnsten, 1942a.
Leucacarbo fuscescens, 43, Phalacrocorax varius,
45; P. sulctrostris, 46; P. sp, 46; Larus
novaehollandiae, 163.
Stictodora lari Yamaguli, 1939.
Larus novaehollandiae, \63.
Stictodora manilensis Alrica & Gareia, 1935,
Srerna albifrons, 169,
Srictodora sp., Bearuip, 1958.
Larus novaehollandiae, 163,
Suctodara sp., Pearson, 1960.
Sterna albifrons, 169,
Family CLINOSTOMIDAE Lihe, [90]
Clinostomum australiense §.J. Johnston, 1917,
Anhined inelanogaster, 42.
Clinostomum contplanatum (Rudolphi, (814), syn.
©. hornum, Nicoll, 1914b.
Nyeticorax caledontcus, 60;
poiciloptilus, 62,
Clinosramunty sp.
Pelecanus conspicillatus, 37,
Bolaurus
Family DIPLOSTOMIDAE Poirier, 1866
Diplostomidae, unidentified.
Phalacrocorax suleirostris, 46; nas superciliosa,
77; Falco peregrinus, 10\,
Bolbophorus confusus (Krause, 1914).
Pelecanus conspicillaius, 37,
Bolbophorus sp., TAH. Johnston & Angel, 1942a.
Pelecanus conspicillatus, 37,
Diplosramum amygdalum Dubois & Pearson, 1965.
Egretta garzetta, 56; E. intermedia, 57; Nvcticorax
culedonicus, 60,
Diplostomum auriculosum Dubois & Pearson, (947.
Anhinga melanogaster, 42,
290
REC, 5, AUST, MUS,
Diplostoniim galaxiae Smith & Hiekmuan, 1983a.
Ardea novaehollandiae, 53.
Diplostomum parvulum Dabois & Angel, 1972.
Pelecanus conspicillatus, 37; Hydroprogne caspia,
164.
Diplostomum poedicipinum
Niewiadomska, 1960,
Podiceps cristatus, 3.
Diplostomum spathaceum (Rudolph, 1819), syn.
Diplosiomum murrayense TAL. Joboston &
Cleland, 1938.
Haliastur sphenurus, 91; Larus nevaehollandiae,
163; Chlidonias hvbrida, 164.
Diplostonnon triangulare (S.1, Johnston, 1904), syn.
Hemistomum triangulare §.J. Johnston, 1904.
Dacela novaeguineae, 253,
Diplostomum sp.
Pelecanus — conspicillarus,
novaehullandiae, 163,
Aysteramorpha plataleae Dubinina & Dubinin,
1940,
Threskiornis dethiopica, 65.
Hysteromorpha triloba (Rudolphi, 1819),
Diplosiomun granulosum Goss, 1940.
Leycocarhe fuscescens, 43; Phalacracorax carbo,
44; P. sulcirasiris, 46; B melanoleucos, 47.
Neadiplostomum sp. Dubois & Angel, 1972,
Grullina eyunoleucu, 428,
Neodiplostonnim brachyurunl (Nicoll, 19144), syn.
Hemistomum brachvurum Nicoll, 19l4a.
Ninox novaeseelandiae, 242; Tvie alha, 244.
Neodiplostomum lanceolatum Dubois & Angel,
1972.
Ninas noveaeseelandiae, 242.
Neodiplostomum pricet Krull, 1934.
Larus novaehollandiae, 163,
Neodiplostamunt spathula (Creplin, 1829), syn,
Neodiplastomum australiense Dubois, 1937a,
Strigeidae, unidentified, Munday & Green,
1972.
Aviceda subcristata, 89; Haliastur sphenurus, 91;
Aveipirer eirrhacephalus, 93; Haliueetus
leucogaster, 95; Aquila audax, 96; Circus
approximans, 99) Faleo subniger, V0; Fi
peregrinus, \Ol, Tyte alba, 244,
Neodiplostomum subacquipartitunr
Pearson, 1967,
HMaliastur sphenurus, 91.
Neodiplasiomun. sp.
Falea peresrinus, 101,
Posthodiplostamum australe Dubois, 1937a.
Pelecanus conspicillatus, 37; Phalacrovorux
sulcirostris, 46; P. melanoleucas, 47; P. sp., 48;
Ardea novaehollandiae, 53; Egreita alha, 55;
Nyeticarax valedonicus, 60; Botaurys
poicilaptilus, 62; Hydroprogne caspia, 166.
Posthodiplostomum oblongunmi Dubois, 1937a.
Boltaurus poiciloptilus, 62,
Kavicka &
37; Larus
syn.
Dubois &
Family STRIGEIDAE Raillict, 1919
Strigeidae, unidentified.
Phalacrocerax varius, 45; Palco peregrinus, 101,
19(15); 219-325 AXES, LY86
Apatemon gracilis (Rudolphi, 1819).
Anas superciliosa, 77,
Apatemon intermedius (S.J. Johnston, 1904), svn.
Hemistomum intermedium S.J, Jolinston, 1904,
Cvenus airatus, 72; Oxyura australis, 86; Accipiier
fasciatus, 92.
Apatemon vitelliresiduus Dubois & Angel, 1972.
Bizivra lobara, 87,
Apharvagostrigea sintplex (S.J. Johnston, 1904), syn.
Holostamum simplex S.J. Johnston, 1904.
Ardeu novuehollundiue, 33; Egretia varsetia, 3G:
E. intermedia, 37-
Apharyngosiriged sp,
feretra alba, 55.
Cardiocephalaides hillii (S.J. Johnston, 1904), syn:
Holostomum hillii S.J. Johnston, 1904.
Larus novaechollundiue, 163,
Cardiocephuloides musculosus (S.J. Johnston, 1904),
syn. Holostomum musculosum 8.1. Johnston,
1904,
Chlidonius hybrida, 164; Hydroprogne caspia,
166, Sterna bergti, 171.
Cardiocephafoides ovicorpus Dubois & Angel, 1972,
Phalacrocerax varius, 45.
Corylurini, unidentified.
Anas superciliosa, 77.
Comlurus magniucetabulus Dubois & Angel, 1972.
Cygnus atralus, 72.
Parastrigea repens (Chase, 1921), syn. Holsrenuin
repens Chase, 1921,
Ardea novaehollandiae, 53, Circus appreximans,
99,
Parastrigea sp., Dubois & Angel, 1972.
Threskiornis aethiopica, 65.
Schwargitrema novuehollandiue Dubais & Pearson,
1967.
Anhinga inelanogasier, 42,
Schwarsirema pandubi (Pande, 1939).
Podiceps cristains, 3; Paliocephulus prolio-
cephalus, 4; Tachvhapius novaehollandiae, 5;
Pelecanus conspicillatus, 37, Phalacrocorax carbo,
44. P sulcirasiris, 46; PB melanoleucos, 47; Ardea
novuehollandiae, 53; Bataurus poicilaptilus, 62:
Platalea flavipes, 68,
Sirigea bavlist Dubois, 19397a.
Threskiornis aethiopica, 635; T. spinicollis, 66%
Platalea flavipes, 68,
Strigea flosculus Nicoll, 1914a,
Podargus strigoides, 246.
Strigea glundulosa Dubois, 1937a, syn, Srrigea
Jalconis, Dubois, 1937b,
Haliastur sphenurus, 91; Aquila audax, 96; Circus
aupproximans, 99; Falco subniger, 100; F) berigara,
104,
Strigew nicolli Dubois, 1937a, syn, Strigea suttoni
Dubois, 1937a.
Burhinus magnirostris, 128; Grallina cvanoleuca,
425; Gymnorhina tibicen, 434.
Strigea promiscua Nicoll, 1914a.
Ninox strenua, 241; N, nevaehollandive, 242.
Sirigead sp,
Falco berigora, 104.
HELMINTHS FROM AUSTRALIAN BIRDS 29
Family PROHEMISTOMIDAE Lutz, 1935
Mesostephanus haliasturis Tabangui & Masilungan,
1941, syn. Mesos(ephanus minor Dubois &
Pearson, 1965.
Pelecanus conspicillatus, 37; Anhinga
inelanogaster, 42; Haliastur sphenurus, 91.
Family FELLODISTOMIDAE Nicoll, 1909
Fellodistomidae, unidentified, syn. Steringophoridae,
unidentified, Goss, 1940.
Phalacrocorax varius, 45,
Family ALLOCREADIIDAE Looss, 1902
Allocreadiidae, unidentified.
Haliastur sphenurus, 91.
Family CEPHALOGONIMIDAE Looss, 1899
Cephalogonimus sp.
Poliocephalus poliocephalus, 4.
Family GYLIAUCHENIDAE Fukui, 1929
Gyliauchen sp.
Botaurus poiciloptilus, 62.
Family ANGIODICTYIDAE Looss, 1902
Hexangium sp.
Botaurus poiciloptilus, 62,
‘Trematoda’, including ‘Distoma’ sp. not further
identified have not been included in this list.
PHLYUM PLATYHELMINTHES
Class Cestoda
Order Trypanorhyncha
Trypanorhyncha, unidentified.
Phalacrocerax varius, 45,
Order Proteocephalidea
Ophiotaenia hylae T.H. Johnston, 1912e,
Dendrocygna arcuata, 70.
Order Pseudophyllidea
Plerocercoids, unidentified.
Dacelo novaeguineade, 253.
Order Cyclophyllidae
Cyclophyllidea, unidentified, Munday & Green,
1972.
Diomedea melanophrys, 13.
Cyclophyllidae, unidentified.
Cvenus atratus, 72; Biziura lobata, 87.
Family TETRABOTHRIIDAE Linton, 1891
Tetrabothriidae, Unidentified.
Aptenodytes patagonicus, 7; Eudyptula minor, \\;
Puffinus tenuirostris, 33; Sula serrator, 38.
Tetrabothrius diomedeae (Fuhrmann, in Shipley,
1900), syn. Prosthecocoryle diamediue
Fuhrmann, 1900.
Diomedea exulans, 12.
Tetrabothrius lutzi (Parona, 1901).
Eudyptula minor, 11.
Tetrabothrius polvorchis Nybelin, 1917,
Fregata ariel, 50,
Tetrabothrius sp., TH, Johnston, 1912b,
Diamedea exulans, 12; D. melanophrys, \3;
Fregata minor, 49.
Teirabothrius sp.
Eudyptula minor, \\; Diomedea exulans, 12; D.
melanophrys, 13; D. chlorarhyvachus, 16;
Phoebetria palpebrata, 18; Macronecies giganteus,
19; Prerodrama hrevirosiris, 24; Pachyptila vittata,
25; Puffinus griseus, 32; Larus novaehallandiae,
163; Sterna berevii, 171.
Family ANOPLOCEPHALIDAE Cholodkovsky, 1902
Subfamily Anoplocephatinae Blanchard, 1891
Paronia trichoglossi (Linstow, 1888), syn. Monieziu
trichoglossae (Linstow, 1888), Taenia tricho-
glossae Linstow, 1888.
Trichoglossus haematodus, 199.
Hemiparonia bancrofti (T.H. Johnston, 1912c), syn.
Dilepis bancrofti T.H. Johnston, 1912c.
Plaiycercus elegans, 212; P. eximius, 213; P. sp.,
217; Barnardius barnardi, 218.
Hemiparonia cacatuae (Maplestone, 1922b), syn.
Schizotuenia cacaiuae Maplestone, 1922b,
Hemiparonia merovitomochaeta Woodland,
1930,
Cacatua leadbeateri, 195; C. valerita, 196.
Hemiparonia sp.
Cacatua roseicapilla, 191.
Family DAVAINEIDAE Fuhrmann, 1907
Subfamily Davaineinae Braun, 1900
Coltugnia collini Fahrmann, 1909.
Dromaeus novaehollandiae, |.
Cotugnia brotogerys Meggitt, 1915.
Platycercus eximius, 213.
Cotugnia platycerci Weerekoon, 1944,
Platveercus icteratis, 216.
Colugnia polytelidis Burt, 1940,
Polytelis anthopeplus, 206.
Cotugnia seni Mezgitt, 1926.
Platycercus eximius, 213.
Colugnia sp.
Dromaius novaehollandiae, |.
Davainea himantopodis TH, Johnston, 1911.
Himantopus leucocephalus, \42.
292
Davainea sp., T.H. Johnston, 1910,
Himantopus leucocephalus, 142.
Davainea sp., TH. Johnston, 1912b,
Leucosarcia melanoleucus, 187;
coronoides, 43%.
Davainea sp-
Alectura lathaini, 107; Himentopus leuco-
cephalus, 142; Cladorhynchus leucocephalus, 143;
Recurvirostra novaehollandae, 144; Leucosarcia
melanoleucus, (87; Corcarax melanorhamphus,
423,
Raillietina australis (Krabbe, 1869), syn. Davainea
australis (Krabbe, 1869), Teenia australis
Krabbe, 1869.
Dromaius noyaehollandiae, 1.
Raillietina cucatuina (TH. Johnston, 1913), syn.
Davainea cacatuina T.H. Johnston, 1913.
Cacalua galerita, \96.
Raillietina conopophilae (T.H. Johnston, 1913), syn.
Davainea conopophilae T.H, Johnston, 1913.
Philemon citreogularis, 358; Entomyzon cyanotis,
359: Conopophila albogularis, 384.
Raillietina corvina (Fuhrmann, 1905),
Corvus orru, 442,
Raillietina leipoae V.H. Johnston & Clark, 1948,
Leipoa ovellata, 106,
Raillietinad leptosoma (Diesing, 1850), syn.
Davainea lepiasoma Diesing, 1850.
Cucatua roseicapilla, 191; Platycercus eximius,
213.
Raillietina paucitesticulaia (Vubrmann, 1908).
Cacatua galerita, \96.
Raillietina polvchalix (Kotlan, 1921).
Platvcercus sp, 217; Neophema petrophila, 229.
Raillietina sphecotheridis (VIA. Johnston, 1914a),
syn. Davainea sphecotheridis T.H.
Johnston, !9l4a.
Sphecotheres viridis, 415.
Raillietina sp., Hall Exped, Report, 1974.
Sphecotheres viridis, 415.
Raillietina sp., TH, Johnston & Clark, 1948b,
Leipoa ovellata, 106.
Raillietina sp,
Dromaius novaehollandiae, 1, Alectura lathumi,
107; Neophema splendida, 231.
Corvus
Subfamily Idiogeninae Fuhrmann, 1932
Idiogenes sp.
Ardeoius australis, 126.
Family DILEPIDIDAE Fuhrmann, 1907
Dilepididae, unidentified.
Tachybaptus novaehollandiae, $8; Emblema picta,
403; Poephila bichenovii, 407; PR cincta, 410;
Gymnorhina tibicen, 434.
Subfamily Dilepidinae Fuhrmann, 1907
Dilepis maxima Goss, 1940,
Phalacrocerax varius, 45.
REC. S. AUST. MUS. 19(15): 219-325
Auzusi, 1986
Dilepis sp.
Phalacrocorax melanoleucos, 47; Anthochuera
carunculata, 350.
Angularella australis (Maplestone, 19216), syn,
Angularia australis Maplestone, 1921b.
Anseranas semipalmata, 69; Burhinus
muenirostris, 128.
Angularella sp.
Vanellus miles, 13).
Anornotaenia accipitris T.H. Johnston, 1913.
Acceipiter cirrhocephalus, 93.
Anomotaenia asymimetrica TH. Johnston, 1913,
Egretta alba, 55,
Anomotaenia hydrochelidonis Dubinina, 1954,
Larus novaehallandiae, \63,
Buerbonia parvitaeniunca (Baer & Bona, 1960), syn,
Kalipora parvitaeniunca Baer & Bona, 1960.
Eereita sacra, 58.
Banceroftiella sp.
Botaurus poiciloptilus, 62.
Clelandia parva TH, Johnston, 1909b.
Nenorhvachus asiaticus, 63,
Cyelorchida omalancristrota (Wedl, 1856),
Platulea regia, 67.
Dendrouterina australiensis Baer & Bona, 1960.
Ardea novuehallandiue, 53,
Dendrouterina sp.
Sphecotheres viridis, 415,
lapwingia adelaidae Schmidt, 1972.
vanellus. miles, 13)
Paradilepis minima (Goss, 1940), syn. Dilepis
minima Gass, 1940,
Phalacrocorax sulcirostris, 46; Po melanoleucos,
47
Paradilepis patriciave Baer & Bona, 1960,
Platalea flavipes, 68.
Paradilepis scolecima (Rudolphi, 1819).
Leucocarbo fuscescens, 43; Phalacrocerax carbo,
44,
Paradilepis urceina Bona, 1975,
Ardea novaehallandiae, 53; Threskiornis
aethiapica, 65,
Puradilepis sp., Clark, 1957.
Phalacrocerax melanoleucos, 47.
Paradilepis sp.
Leucocarbo fuscescens, 43,
Paricterotaenia chlamyderae (Krefft, 1873), syn.
Tuenia chlamyderae Krefft, 1873.
Chiamydera rraculata, 419.
Puricterotdenia zoniferae (T.H. Johnston, 1912¢),
syn, Choanotaenia zoniferae T.H. Johnston,
191 2¢.
Fanellus tricolor, 132.
Puricterotaenia sp.
Burhinus magnirostris, 128.
Paryvitaenia ardeae (T,H. Johnston, 1913), syn.
Bancroftiella ardeae T.H. Johnston, 1913,
Ardea novaehollandiae, 53, Nycticorax
valedonicus, 60.
Parvitaenia clavipera Baer & Bona, 1960, syr.
Bancroftiella glandularis of T.H, Johnston
19)2b, in part, not (Fuhrmann, 1905),
Ardea novaehollandtae, 53.
HELMINTHS FROM AUSTRALIAN BIRDS 243
Parvituenia gldndularis (Fuhrmann, 1905), syn.
Bancroftiella glandularis (Fuhrmann, 1905).
eretta ulba, 35.
Parvitaenia paracyclorchida Baer & Bona, 1960, syn.
Bancroftiella glandularis (Puhrmann, 1903).
“Ardea novaehallandiae, 53.
Trichocephatloides sp.
Calidris acuminata, \57.
Subfamily Dipylidiidae Sules, 1896
Chounotaenia fieldingi (Maplestone & Southwell,
1923), syn, Monopylidium fieldingi Maplestone
& Southwell, 1923.
Cractious forquatus, 432.
Choanvtaenia southwelli Fubrmann, 1932, syn.
Monopvlidium macracanthus of Maplestone
and Southwell, 1932, not Puhrmann, 1907.
Fanellus miles, 131.
Choanotaenia taylori TA. Johnston, 1912c.
Malurus cyaneus, 318-
Choanotaenia sp.
Burhinus magnirostris, 128; Anthochoera
chrysoptera, 352; Emblema guttata, 404; Poephila
bichenavii, 407,
Pseudochoanotaenia meliphagidarum (V.H,
Johnston, 1911), syn. Choanotaenia
meliphagidarum TH, Johnston, 1911; C. sp.,
THJ, 1910).
Meliphaga lewint, 363; Lichenostome leucatis,
367; LL. plumula, 372; Phylidonyris noyuae-
hollundiae, 380; Po nigra, 381; Myzamela
sanguinolenta, 390.
Srniluncinus dacelonis TA, Johnston, 1909a.
Dacelo novaeguineae, 253,
Subfamily Parulerinae Pulirtnann, 1907
Paruterinae, unidentified.
Pomutostomus temporalis, 310; P. superciliosus,
31).
Anonchotuenia arhvacha Fuhrmann, 1918.
Pardalotus striatus, 399.
Anonchoalaenia glabata (Linstow, 1879), syn.
‘unidentified cestode’ Cleland, 1922.
Entamyzon cvanotis, 359.
Biuterina sp.
Anseranas semipalmata, 69; Sphecotheres viridis,
415,
Parulerina rauscht Freeman, 1957,
Tringa siagnatilis, 180; Tyto alba, 244,
Sphaeruterina punctata TH, Johnston, 1914b.
Pachycephalus rufiventris, 293.
Zosteropicola clelandi T.H, Johnston, 1912c.
Zosterops lateralis, 400.
Family HYMENOLEPIDIDAE Raillict & Henry, 1909
Hymenolepididae, unidentified,
Threskiornis aethiopica, 65; Philemon
citreogularis, 358: Entomyzon eyanotis, 359;
Corcorax melanorhamphus, 423, Strepera
graculina, 435.
Subfamily Hymenolepidinae Ransom, 1909
Armadoskrjahinia glohosa (Szpotanska, 1931), syn.
Hymenolepis globosa Szpatanska 1931,
Cyenus atratus, 72.
Australiolepis southwelli (Szpotanska, 1931), syn.
Hymenolepis southwelli Szpotanska, 193];
Echinorhynchotuenia nana Mapleston &
Southwell, 1922b.
Cyenus atratus, 72.
Cloacetaenia megalops (Nilzsch in Creplin, 1829),
syn. Hpmenolepis megalops Nitzsch in Creplin,
1829,
Anseranas semipalmala, 69; Dendracygna
arcuala, 70; Cygnus atratus, 72; -lnas superciliosa,
77; A. castanea, 79.
Cloacotdenia sp.
Cygnus atratus, 72; Sictonetia naevosa, 73; Anas
rhynchotis, 80; Avthya australis, 82.
Diorchis flavescens (Krefft, 1873), sya. Taenia
flavescens Krefft, 1873.
Dendrocy'gna arcuata, 10; Anas superciliosa, 77;
A. sibberifrons, 78; A. castanea, 79; A,
rhynachatis, 80; Aythye australis, 82; Himantopus
leucocephalus, 142; Cladorhynchus leucacephalus,
143,
Diorchis spiralis Szpotanska, 1931,
Cygnus atratus, 72.
Diorchis srefanski Czaplinski, 1955,
Cygnus atratus, 72.
Diorchis sp,
Stictonetta naevosa, 73; Anas castanea, 79,
Dicranotaenia coronula Dujardin, 1845.
Cyenus atratus, 72,
Diplogynea olizorchis (Maplestone, 1922a), syn,
Cotugnia oligerchis Maplestane, 1922a,
Diploposthe laevis of T.H. Johnston, 1913, not
(Bloch, 1782).
Dendrocyvgna arcuata, 70; Avthya australis, 82.
Diplopesthe laevis (Bloch, 1782), syn, Taenia
thberculata Kreftt, 1873.
Anas superciliosa, 77; A, castanea, 79; Aviva
australis, 82.
Cupiuterilepis australiensis Schmidt, 1972.
Anthochaera carunculara, 350; Anthochaera 3p.
353.
Capiuterilepis meliphagicola Schmidt, 1972.
Philemon corniculatus, 357.
Drepanidetaenia bisacculina Szporanska, 1931,
Cyenus atratus, 72.
Drepanidotaenia lanceolata (Bloch, 1782).
Cyenus atratus, 72; Anas superciliosu, 77.
Drepanidolaenia rapida (Szpotanska, 1931), syn-
Hymenalepis rapida Szpotanska, 1931.
Cygnus atratus, 72.
Drepanidotaenia sp.
Cygnus atratus, 72,
Haploparaxis australis (TH. Johnston, 1913), syn.
Aploparaksis australis TA. Johnston, 1913.
Tringa hypoleucos, 148; Gallinage hardwicki, 152;
G, megala, \53.
Haploparaxis veitchi Baylis, 19346,
Anas gibberifrons, 78.
204 REC S, AUST. MUS, [9(15): 219-325
Microsomacanthus collaris (Batsch, 1786), syn.
Hymenolepis collaris (Batsch, 1786), Tuenia
bairdti Krettt, 1873,
Anus superciliosa, 77; A. castanea, 79.
Microsomacantius cormoranti (Ortlepp, 1938), syn.
Hymenolepis cormorant’ Ortlepp, 1938.
Phalucrocorax melanoleucos, 47.
Monosaceanthes curiosa (Szpotanska, 1931), syn.
Hymenolepis curiosa Szepotanska, 1931,
Cvenus arrats, 72.
Monosaccanthes kdzachstanica (Maksinoya, 1963),
Cyenus arrarus, 72.
Monosaccunthes sp.
Cyenus atratus, 72
Parabisaccanthes bisacculina (Szpotanska, 1931),
syn. Drepunidolaenia bisacculina Szpotanska,
1931.
Crenus arrutus, 72.
? Passerilepis sivlosa (Rudolphi, 1810),
Anseranas semipalmata, 69.
” Passerilepis sosteropis (Fuhrmann, 1918).
Dendrocvenad arcuata, 70.
Sobolevicanthus terraereginae (TH. Johnston, 1913),
syn. (Lvmenolepis rerraereginae T.H. Johnston,
1913.
Anseranas semipalinata, 69.
Sobolevicanthus sp.
Cereopsis novaehollundiae, 74, Ayvihya australis,
82; Biziura lobaia, 47.
Stuphylepis lamella (Woodland, 1930), syn.
Hymenolepis lamellata Woodland, 1930.
Tudarna tadornoides, 75; Anas gibherifrons, 78.
Tschertkovilepis krabbei (Kowalewski, 1895).
Cyvents atratus, 72.
Moodlundia phalucrocoracis (Woodland, 1929), syn.
Hvymenolepis phalacrocaracis Woodland, 1929,
Phalacrocorax melanoleucos, 47.
Hymenolepis Chenypis Paliner, 1981,
Cygnus arrarys, 72.
Hymenolepis ellisi TAA, Johnston & Clark, 1948a,
Pelecunus conspicillatus, 37.
‘Hymenolepis curystonm Maplestone’.
Eurvstamus orientalis, 261.
Hymenolepis ibidis TH. Johnston, 1913.
Threskiornis spinicollis, 66; Platalea flavipes, 68;
Dendrocyzna arcuata, 70,
Hymenolepis jaenschi TH. Johnston & Clark,
1948a.
Pelecanus conspicillams, 37.
Hymenolepis liophallus (Krabbe, 1869).
Cyenus. atratus, 72.
Hymenolepis micrancrisirota (Wedl, 1855), Taenia
mucrancristrota Wedl, 1855,
Cvenus atratus, 72,
Hymenolepis murrayensis 1H. Johnston & Clark,
1948a.
Pelecanus conspicillatus, 37.
Hymenolepis roberts] Baylis, 1934b.
Anas gibberijrons, 78.
‘Hymenolepis variabilis Maplestone’,
Cyenus utrarus, 72.
Hymenalepis sp., TH. Johnston, 1910,
Aimuntopus leucdcephalus, 142;
goodenovii, 281.
Pefroieu
August, 1986
Hymenolepis sp., T.H. Johnston, 1912b.
Nyeticorax caledonicus, 60; Anas superciliosa, 77.
Hymenolepis sp,, Southwell, 1916.
Cygnus atratus, 72.
Tlymenoalepis sp.
Pelecanus conspicillatus, 37, Nyericorax
caledonicus, 60; Threskiornis spinicollis, 66,
Platalea flavipes, 68: Cvgnus atratus, 72:, Tadorna
tudarnoides, 75, Anas superciliosa, 77, Fulica atra,
123; Reeurvirostra novaehollandiae, 144;
Anthochaera sp., 353; Lichenostoma virescens,
366; Pardalotus striatus, 399; Chlampdera
maculata, 419; Grallina cyanoleuca, 425; Corvus
mellori, 440; C. sp., 443,
Subfamily Fimbriariinae Wolffhiigel, 1899
Fimbriaria fasciolaris (Pallas, \871), syn. Taenia
pediformis Krefft, 1873.
Anas superciliosa, 77; A. castaunea, 79,
Fimbriaria sp.
Anas superciliosa, 77; Aythya australis, 82.
Fimbriarioides intermedia (Fuhrmann, 1913), syii,
Fimbriaria intermedia Fulbemann, 1913.
Cereopsis noyaehollandiae, 74.
Family ACOLEIDAE Ransom, 1909
Acoleus hedleyi TH. Johnston, 1910, sya. Taenia
rugosa Keetlt, 1873.
Himantapus leucocephalus, 142,
Family DIPLOPOSTHIDAE Poche, 1926
Diplopasthe laevis (Bloch, Tuenia
tuberculata Krell, 1873.
Anas superciliosa, 77; A. castanea, 79; Avihva
gustralis, 82.
1782), syn
Family DIOECOCESTIDAE Southwell, 1930
Subfamily Dioecocestinae Southwell, 1930
Dioecocestus novaehollandiae (Kref{t, 1873), syn.
Juenia novuehollandiae Krellt, 1873, 7.
paradoxa WKrefll, 1873,
Tachybaptus noyaehollandiae, 5.
Dioecacestus sp.
Poliocephulus poliocephalus, 4.
Subfamily Gyrococlinae Yamaguli, 1959
Gyrocevelia australiensis (TH. Johnston, 1910), syn,
Gyrocoelia sp., TH. Johnston, 19l4a, Dilepis
australiensis TJ. Johnston, 1910, Jaenta
coronata Krefit, 1873.
banellus miles, 131: Himantopus leucacephalus,
142.
Gyreceelia sp.
Poliocephalus poliacephalus, 4; vanellus miles,
131; Cladorhynchus levicocephalys, 143.
tnfula burhini Burt 1939, syn. 'Shipleva lobivanellus
Maplestone’.
bunel/us niles, 131; Mimantopus leucocephulus,
142,
HLELMINTHS FROM AUSTRALIAN BIRDS 295
Family TAENIIDAE Ludwig, 1886
Cladotaenia feuta Meggitt, 1933,
Circus assimilis, 98,
Cladotaenia sp.
Circus assimilis, 98; C. approximans, 99; Falco
subniger, 100.
Family NEMATOPARATAENILLDAE Poche, 1926
Nematoparataenia paradoxa Maplestone &
Southwell, 1922b,
Cygnus atratus, 72
Gastrotaenia sp.
Cyenus atratus, 72; Anas superciliosa, 17; A.
vibberifrons, 78; A. rhynchotis, 80, Aythya
australis, 82; Bizinra lobata, 87.
PHYLUM NEMATODA
Order Enoplida
Superfamily DIOCTOPHYMATOIDEA
Eustrongylides phalacrocoracis TA, Johnston &
Mawson, 194 le.
Leucocarbo fuscescens, 43; Phalacrocorax carbo,
44; P. melanoleucos, 47.
Eustrongylides plotinus T.H. Johnston & Mawson,
194le.
Anhinga inelanogaster, 42.
Eustrongylides sp.
Pelecunus conspicillatus, 37; Phalacrocorax carbo,
44: P. sulcirosiris, 46; Nycticorax culedonicus, 60.
Superfamily STRONGYLOIDIDAE
Strongyloides sp.
Anas gibberifrons, 78.
Superfamily TRICHUROIDEA
Capilluria anatis (Schrank, 1790).
Cygnus atratus, 72; Anas castanca, 79.
Capillaria convolutor Fourment, 1885,
Diomedea pielanaphrys, 13; Macronectes
gigunteus, 19.
Capillaria ellisi TH, Johnston & Mawson, 1945b.
Cygnus arratus, 72.
Capillaria grallinae TH. Johnston & Mawson,
1945a,
Grallina evanoleuca, 425,
Capillaria graucaliny VW. Johnston & Mawson,
1941d.
Corueina novuaehollundiae, 27).
Capillaria eyninorhinde T.H. Johnston & Mawson,
1947,
Gyinnorhina tibicen, 434.
Cupillaria jaenschi T.H. Johnston & Mawson, 1945b.
Pelecanus conspticillatus, 37, Leucocarbo
fusvescens, 43; Phalacrocoray carbo, 44: PB varius,
45; P suleirostris, 46; P. melanoleucos, 47; Larus
novaehollandiae, 163, Chlidanius hybrida, 164.
Capilluria laricola Wassilikova & Gushanskaya, 1930.
Larus nevaehollandiae, 163.
Capillaria obsignata Madsen, 1945.
Cygnus atratus, 72.
Capillaria pomatostomi T.H, Johnston & Mawson,
1945b.
Pomatostomus superciliosus, 311.
Capillaria recurvirostrae Mawson, 1968b.
Recurvirostra novaehollandiae, \44.
Capillaria thomascameroni Mawson, 1969.
Larus novaehollandiue, 163.
Capillaria triloba Linstow, 1875.
Vanellus miles, 131; Himanropus leucocephulus,
142; Reeurvirostra noyaehollandiue, 144.
Capillaria sp., Munday & Green, 1972,
Rullus pectoralis, W5.
Capillaria sp., T.-H, Johnston & Mawson, 1945a,
Leucacarbo fuscescens, 43,
Capillaria sp, TH. Johnston & Mawson, 1949,
Podiceps cristatus, 3.
Capilluria sp.
Poliocephalus poliocephalus, 4, Tuchyvbaprus
novaehollundiue, 5; Podicepididae, unidentified,
6; Botuurus poicilaptilus, 62; Threskiornis
aethiopica, 6S; Threskiornis spinnicallis, 66, Anas
superciliasa, 77, A. gibbertifrons, 18, A. castaned,
79. 4. rhynchotis, 80; Malacarhynchus
membranaceus, 81; Aviva australis, 82; Oxvura
australis, 86; Haliastur sphenurus, 8, Haliauetus
leucogaster, 95; Cireus approximans, 99; Falco
berigora, \O4; Aleerura lathami 107, Turnix
pyrrhothorax, 112; Rallus pectoralis, 115; Porzana
pusilla, 116; PB fluminea, 7, Gallinula moruerii,
119: G. tenebrosa, 121; Porphyrio porphyria, 122;
Charadrius bicinera, \37, CL ruficupilius, 139;
Sterna bergil, 171; Melopsitiucus undulatus, 209;
Neophema splendida, 231, Cuculus /labelliformus,
234; Ninox novaeseelundiue, 242; Tylo
novachollandiae, 245; Podareus strigvides, 246,
Eurvsiomus orientalis, 261; Colluricinela
harmonica, 294; Cinclosoma cinnamonieutt, 309,
Pomatostomus ruficeps, 312; Malurus
leucopterus, 322; Alcanthagen\s clifogularis, 334;
Munorina nelanocephala, 361; MI. flavisila, 362;
Lichenostontus virescens, 366; Lichnrera
indistineta, 378, Cracticus torquatus, 432; Corviis
tusmranicus, 439; ©. mellori, 440,
Order Strongylida
superfamily STRONGYLOIDEA
Family SYNGAMIDAE Leiper, 1912
Syngamus (rachea (Montagu, I8t1).
Calyptorhynchus funereus, 188, Gyninorhina
tibicen, 434,
SVAeainus sp.
Alecrura lathanii, 107.
“? Hookworm
Threskiornis uethiopica, 65,
Supertamily TRICHOSTRONGY LOIDEA
Amidostramum acucun (Lindahl, i848), syn A
chevreuxi Seurat, 918.
Suctonetta nuevosa, 73; Tadorna radjah, 76; Anus
superciliosa, 77; A. gibberifrons, 78: A.
rhynchots, 80; Mimanteopus leucocephalus, 142.
Amidostomum unseris (Zeder, 1800).
Anseranus seinipalmata, 69;
novaehollandiae, 74.
Amidostomum biziurae TH. Jonnston & Mawson,
1947
Bisiura lobata, 87.
Anidostamum event Wehr, 1933,
Cygnus atraius, 72.
Amidastomunr fulicue (Rudalphi, 1819).
Fuliva atra, 123.
Alnidostomum tribonyx Mawson, 1980,
Gallinula ventralis, \20.
Amidostamum sp,
Cyenus atratus, 72; Stictonetta naevosa, 73, Anas
superciliosa, 77; Chenanetta jubata, 83; Bizinura
lobaia, 87; Gallinula tenebrosa, 121.
Dromaeostrongylus bicuspis Lubimov, 1933,
Dromaius novaehollandiae, \.
Epomidiostomum uncinatus (Lundahl, 1848),
Anas gibberifjrons, 78.
Epomidiastomum sp,
Anseranas semipalmata, 69, Cyenus atratys, 72;
Jadorna tadornoides, 75, Anas superciliosa, 17;
A. gibberifrons, 78; A. castanea, 79; Chenonetta
Jubata, 83; Oxvura dusiralis, 86,
Pseudamidostomumn sp.
Cygnus utratus, 72; Chenaonetla jubaia, 83,
Trichostrongylus incertus T.H. Johnston & Mawson,
194]e,
Hydroprogne caspia, |66.
Trichostrongylus tenuis (Mchlis, 1864),
Dromaius novaehollandiae, 1.
Ornithostrongylinae, unidentified.
Podargus strigoides, 246,
Cereopsis
Order Ascaridida
Superfamily ASCARIDOIDEA
Family ANISAKIDAE Railliet & Henry, 1912
Anisakidae, unidentified, Munday & Green, 1972
Diomedea melanophrys, 13.
Anisakidac, unidentified.
Eudyptes chrysocome, 8.
Anisakis diomedeae (Linstow, 1888), svn. Sromachus
sp.. Munday & Green, 1972, Contracaecum
diomedeae (Linstow, 1888).
Diomedea exulans, \2; 2. melanophrys, 13; DO.
chrvsostoma, 15; D. chlororhynchas, 16; D. cauta,
17; Macronectes giganteus, 19, Daption capense,
21; Pachyptila desolata, 27.
Anisakis sp., T.-H. Johnston & Mawson, 19424.
Budyplula minor, 11.
Anisakis sp., TwH. Johnston & Mawson, 1942c,
Pachyptila desolata, 27,
Anisakis sp. T.H. Johnston & Mawson, 1942d.
Pierodroma lessoni, 23.
Anisakis sp., syn. Stomachus sp. aucte,
Prerodroma macroptera, 22; Pachyptila belcheri,
28; Sterna bereii, 17; Anous minuttts, 174,
REC. S, ALIST, MUS. 1915): 219-325
ANS! LO86
Conitracaecum cevlanicum (Linstaw, 1904),
Haliastur sphenurus, 1.
Contracaecuim clelandi TH. Johnston & Mawsan,
194 1c,
Pelecanus conspicillatus, 37,
Contracaecum eudypiulae T.H. Johnston &
Mawson, 1942a,
Eudyvptula minor, 11,
Contracaecum maenicollare TH, Johnston &
Mawson, 194]c,
Diomedea eauta, \7; Puffinus wriseus, 32; P
tenuirostris, 33; Sula serrator, 38; Anous stolidus,
173; A. muinuius, 174-
Contracaecum tmicrocephalun (Rudolphi, 1809).
Nycricorax caledonicus, 60; Botaiirus
poicilopiilus, 62; Anas superciliosa, 77,
Coniracaecum micropapillatunr (Stossich, 1890),
syn. €. bancraft) T.H. Johnston & Mawson,
194 1c,
Pelecanus conspicillatys, 37,
Contracaecum nycticoracis TH. Johnston &
Mawson, 194 le,
Nyeticorax caledonicus, 60.
Contracuecum pelagicum TH. Johnston & Mawson,
1942d.
Diomedea melanophrys, 13; D. chlororhyncha, V6.
Contracaecum podicipitis T.H. Johnston & Mawson.
1949.
Podiceps cristaitus, 3; Tachybapius novae-
hollandiae, 5.
Contracaecum praestriatum Moénnig, 1923,
Poliacephalus paliocephalus, 4.
Contracaecum rodhaini (Gedoelst, 1916).
Anhinza melanogaster, 42,
Contracaecum sinulabiatum TM. Johnsion &
Mawson, 194le, syn, Ascaris spiculizerum
(Rudalphi, 1809) of TLH. Johnston, 1914a,
Anhinga melanoguster, 42; Phalacrocerax carha,
44; P, melanoleucos, 47.
Contracaecum spiculigeruim (Rudolphi, 1809), syn.
Ascaris spiculigerym Rudolphi, 1809; Ascaris
sp. T.H. Johnston, 1912b.
Eudyplula nunor, 11; Pelecanus conspicillatus, 37:
Leucoearbo Juscescens, 43, Phalicrocorax carbo,
44; PB varius, 45; P. sulcirostris, 46; 2
melanoleucos, 47; Ardea pacifica, 32; A-
novaehollandiae, 53; Nycticarax caledonicus, 60:
Botaurus pojeiloptilus, 62; Plegadis falcinellus, 64.
Contracaecum tricuspe (Gedoelst, 1916), syn Ascaris
spiculigerum (Rud.) of TH. Johnston 1912b, 4.
sp.. Krefft, 1873.
Anhinga melanogaster, 42,
Contracaecum sp., T.H. Johnstan & Mawson, 194 1c.
Ardea novaehollandiae, 53, Esrerta alba, 53;
Nenorhynchus asiaticus, 63.
Contracuecuin sp., T.H. Johnston & Mawson, 1947,
Chiidonias hybrida, \64,
Contracaecum sp., Mawson, 19G8b.
Himantopus leucacephalus, 142.
Contracdecum sp.
Eudyptes pachyrhynchus, 9; Diomedea exulans,
2: Phoeberria palpebrata, 18; Sula serrator, 38;
Fregata minor, 49; Phuethon rubricauda, 51;
HELMINTHS FROM AUSTRALIAN BIRDS 297
Ardea navaehollandiae, 53; Egretta alba, 35,
Butorides striatus, 59, Ixobrychus minutus, 61,
Botauriis poiciloptilus, 62; Platalea flavipes, 68;
Fulco peregrinus, 101; Halcyon chloris, 258.
Phocascaris sp. T.H, Johnston & Mawson, 1942d.
Macronectes viganieus, 19.
Family ASCARIDIDAER Baird, 1853
Baylisuscaris sp.
Nymphicus hollandicus, 208,
Porrocaecum angusticolle (Molin, 1860).
Falco berigora, 104,
Porrocaecum circinum T.H. Johnston & Mawson,
1941b.
Elanus notatus, 88; Accipiter novaehollandiae, 94:
Hieraaetus morphnoides, 97; Circus assimilis, 98.
Porrocaecum clelandi TH. Johnston & Mawson,
1941d, syn, ‘nematode; Cleland, 1922,
Anthus novaeseelandiae, 270; Zoothera dauma,
276; Cinclosoma punctaium, 307.
Porracaecum crassunt (Deslongchamps, 1824).
Anas superciliosd, 77,
Poarrocaecum ensicaudatum (Zeder, 1800).
Porzanu fluminea, \17.
Porrocaecuin lobibicis Mawson, 1968b.
Vanellus miles, 13).
Porrocuecum menurae T.H. Johnston & Mawson,
1942b.
Menura novachollandiae, 264.
Parrocaecum reticulatum (Linstow, 1889).
Ardea pacifica, 52, Egreita alba, 55:
Porrocaecum serpentulum (Rudolphi, 1809), syn.
Ascuris serpentula Rudolphi, 1809.
Grus rubicundus, 124.
Porrocaecum streperae TH, Johnston & Mawson,
194!d.
Gymunorhina tibicen, 434; Strepera graculina, 435,
S. versicolor, 437.
Porrocuecum wui Hsti, 1933.
Gymunorhina tibicen, 434.
Porrocaecum sp., TH, Johnston & Mawson, 1941d,
Manorinu Jlavieula, 362.
Porrocaecum sp.
Ardea novuehollandiae, 53; Circus approximans,
99 Merops ornatui, 260; Pilta versicolor, 263;
Malurus leucopterus, 322, Corvus mellori, 440,
Superfamily SEURATOIDEA
Family SEURATIDAE Hall, 1916
Riciularina spinosa T.H. Johnston & Mawson,
1941d.
Muyiagra rubecula, 299,
Skrjabinura magna (TH. Johnston & Mawson,
4d), syn. Sewratinema magnum T.A.-
Johnston & Mawson 1941d.,
Dacelo novaeguineae, 253,
Skrjabinura pomatosiomi (TH. Johnston &
Mawson, 941d), syn. Seurafinema
pomatostomi T.H, Johnston & Mawson, 1941d.
Pomatostomus superciliosus, 311.
Skrjabinura brevicaudatum (VA. Johnston &
Mawson, 1941b), syn, Senratinema
brevicaudutum (TH. Johnston & Mawson,
194 1b).
Ninox cannivens, 243; Pomalostomus lemporalis,
310.
Skrjabinura sp., syn. Seuratinema sp,, Ogden, 1967-
Podargus strizoides, 246; Eopsaltria pulverulenia,
283,
Family SCHNEIDERNEMATIDAE Freitas, 1936
Inglisonemad typos Mawson, 1968a.
Pitta versicolor, 263.
Superfamily HETERAKOIDEA
Family HETERAKIDAE Railliet & Henry, 1912
Heierakis ¢henonetiae 1,H, Johnston, 19)2a.
Cereopsis novaehollandiae, 74; Chenonetta
jubata, 83.
Heterakis cireumvallata (Linstow, 1906b).
Cyenus atratus, 72.
Heterakis dispar (Schrank, 1790),
Cereopsis novaehallandiae, 74,
Heterakis gallinarum (Schrank, 1788), syn, Hererakis
gallinae (Gmelin, 1790),
Cacatua /eadheateri, 195.
Heterakis vesicularis (Froelich, 1791), syn. ff,
papillosa (Bloch, 1782), 17, cuudara (Linstow,
1906).
Cygnus atratus, 72; Cereopsis novoehollandiue,
74.
Heterakis sp.
Anseranas semipalmaia, 69; Northiella
huematogaster, 224; Ninox novaeseelandiae, 242.
Odontolerakis bancrofti(T.H. Johnston, 1912a), syn.
Heterakis bancrofii; YH. Johnston, 1912a.
Alecrura luthami, 107, Leucosarcia meldnoleuca,
187.
Family ASCARIDITDAE Travassos, 1919
Ascuridia catheturina (T.H. Johnston, 1912a), syn.
Heterakis catheturina TH. Johnston, 1912a.
Alectura lathamti, 107.
Asearidia columbae (Gmelin, 1790),
Phaps chaleoptera, \80; Evlectus rorarus, 197,
Alisterus scapularis, 203; Polytelis alexandriae,
207; Melopsittacus undulatus, 209; Lathamus
discolar, 210; Burnardius barnardi, 218, 8.
zonarius, 219; Neophema bourkei, 226.
Ascaridia galli (Schrank, 1788).
Chrysococeyx lucidus, 236; Cenrropus phasianus,
239.
Ascaridia lineatu (Schneider, 1866),
Centropus phasianus, 239.
Ascuridia plarveeri Hartwich & Tscherner, 1979, syn.
Ascuridia sprenti Mines, 1979,
Calocephalon fimbriatum, 190; Alisrerus
scapularis, 203; Aprosmicrus erythropterus, 204;
Polyrelis swainsoni, 205, P. anthopeplus, 206; B
alexundrae, 207; Nvimphicus hollandicus, 208;
298
Melopsitiacus undulatus, 209: Lathamus discolor,
210; Platycercus elegans, 212; P eximius, 213; P.
sp., 217; Barnardius burnardius, 218; B. zonarits,
219; Psephorus haematonotus, 220; P. varius, 221;
P. chrysopterygius, 222: P. dissiinilis, 223;
Northiella haematogaster, 224; Neophema
bourkii, 226; N. elegans, 228; N. pulchella, 230;
N. splendida, 231,
Ascaridia stroma (Linstow, 1899).
Grus rubieundus, 124, G, antizene, 125.
Alscaridia sp,
Geopelia humeralis, \78; Cacatua roseicapilla, 191;
Polyteles anthapeplus, 206; Nymphicus
hollandicus, 208; Melopsittacus undulatus, 209:
Platycercus sp, 217, Centrapus phasianus, 239.
Superfamily SUBULUROIDEA
Family SUBULURIDAE Travassos, 1914
Allodapea differens (Sonsino, 1890), syn, Subulura
differens (Sonsino, 1890).
Cernlropus phasianus, 239.
Allodapa suctoria (Malin, 1860), syn. Subulure
clelandi T.H. Johnston & Mawson, 1941d,
Scrvthraps novaehollandiae, 238: Ninox
novaeseelandiae, 242; Podargus strigoides, 246;
Eurostopodus inystacalis, 249; Caprimulaus argus,
250.
Allodapa sp., syn, Subulura sp., TH. Johnston &
Mawson, 1941b.
Ninox strenua, 241.
Allodapa sp.
Podargus strigoides, 246.
Leipoanema ellisi TH, Johnston & Mawson, 1942a.
Leipoa océlluta, 106.
Superfamily OXYUROIDEA
Oxyuridae, unidentified.
Ninox novdeseelandiae, 242; Tyro alba, 244,
Order Spirurida
Spirurida, unidentified, TJH, Johnston & Mawson,
1941e.
Pomutostomus superciliosus 311,
Spirurida, unidentified, Munday & Green, 1972.
Daption capense, 2); Pterodroma inacraptera, 22.
Spirurida, unidentified,
Pelecanoides urinatrix, 36; Accipiter fasciatus, 92;
A, cirrhocephalus, 93; Falco peregrirus, 101; F
longipennis, 102; Ervthrogenys cinerus, 134:
Charadrius ruficapillus, 139 Chlidonias hybrida,
164; Eudynamys scolopacea, 237; Tylo alba, 244;
T, novaehollandiae, 245; Cevx azurea, 252; Dacela
novaeguinege, 253; Halcyon echloris, 258; Petroica
voedenovii, 281; Pachvcephala pectoralis, 291;
Miyiagra alecto, 300; Pomatastonius superciliosus,
311; Acanthiza pusilla, 336; Climacteris
leucophaea, 346; Philemon argenticeps, 356;
Entomyzon cyanotis, 359; Artamus cyanaplerus,
430, Cracticus nigragularis, 433; Gynmerhina
tibicen, 434; Corvus tasmanicus, 439: C mellori,
440.
REC. 5. AUST, MUS, 19(15); 219-325
Ajugust, 1986
Superfamily ?, Family ?
Bancroftinema dentatum T.H, Johnston & Mawson,
1941b,
Falco berigera, O04.
Superfamily CAMALLANOIDEA
Family CAMALLANIDAE Railliet & Henry, 1915
Procamallanus murrayvensis TH. Johnston &
Mawson, 1940b.
Phalacrocorax carbo, 44.
Superfamily PHYSALOPTEROIDEA
Family PHYSALOPTERIDAE (Railliet, 1893)
Physaloptera alata Baylis, 1925,
Falco longipennis, 102.
Physaloptera hieracidiae T.H. Johnston & Mawson,
194 1b.
Falco berigora, \04.
Physaloptera sp., T.H. Johnston & Mawson, 194 Ie.
Threskiornis spinicollis, 66; Anas superciliosa, 77.
Physaloptera sp.
Plegadis fulcinellus, 64; Accipiter cirrhocephalus,
93; Faleo peregrinus, 101; Ninex novaeseelandiae,
242, Cracticus nigroguluris, 433.
Superfamily THELAZIOIDEA
Family THELAZIIDAE Skrjabin, 1915
Thelazia aquilina Baylis, 1934a.
Accipiier novaehallandiae, 94: Hualiaeetus
leucogaster, 95; Aquila audax, 96; Falco berigora,
104.
Thelazia dacelonis (Breinl, 19136), syn.
dacelonis Breinl, 1913b.
Dacelo leachii, 254,
Thelazia pittae T,H. Johnston & Mawson, 1941¢, syn
‘nematode’, Cleland, 1922,
Pitta ervthrogaster, 262,
Thelazia sp.
Evretta alba, 55.
Oxyspirura anthochaerae (TH. Johnston, 1912a),
syn. Ceralospira anthochaerae T.H. Johnston,
19124; Ascaris sp., Krefft, 1873,
Anthochaeru carunculata, 350.
Oxyspirura banerofti TAH. Johnston & Mawson,
194d.
Philemon citreogularis, 358.
Oxyspirura streperae ‘V.H. Johnston & Mawson,
194ia,
Crvninorhina tibicen, 434; Strepera graculina, 435;
S. fuliginosa, 436.
Oxyspirutd sp.
Ninox novaeseelundiae, 242.
Filaria
Superfamily SPLRUROIDEA
Family GONGYLONEMATIDAE Hall, 1916
Gongvlonemed aleeturae TH. Johnston & Mawson,
19424,
Alectura lathami, \O7.
HELMINTHS (ROM AUSTRALIAN BIRDS 299
Protospirura spi.
Dacelo novacguineae, 253,
Family HARTERTHDAE Quentin, 1970
Alainchabaudia uegetheles (TH. Johnston &
Mawson, l94le), syn. Alabronema aegotheles
T.H. Johnston & Mawson, 194le,
Aegotheles cristatus, 248,
Alainchabaudiu alcedinis Mawson, 1968c.
Dacelo novaeeuineae, 253; Halcyon saneta, 257.
Harlertia sp.
Ardeotis australis, 126; Ninox novaeseetandiae,
242.
Superfamily HABRONEMATOIDEA
Family HABRONEMATIDAE Chitwood & Wetir,
1932
Habronematinae, unidentified, Mawson, 1969.
Geochelidon nilotica, 165,
Cyrnea colini Cram, 1927.
Coturnix ypsilophora, 109.
Excisa biloba Mawson, 1968d,
Ninox rovaeseelandiae, 242; Podareus strigoides,
246; P. ocellaius, 247,
Exeisu dentifera (T.H. Johnston & Mawson, 1941d),
syn. Cyrnea dentifera T.H. Johnston &
Mawson, 1941d,
Ardeotis australis, 126,
Excisa excisiformus (Yamaguli, 1935),
Podareus strigvides, 246,
Procyrnea dollfusi (Mawson, 1968d), syn, Crrnea
dollfusi Mawson, 1968d.
Ninox novaeseelandiae, 242.
Procyrnea fJalea (Mawson, 968d), syn. Cyrnea faleo
Mawson, 1968d,
Faleo longipennis, 102; Ff) berigera, 4; #
cenchroides, 105,
Procyrnea incerta (A.J. Smith, 1908), syn. Sprroptera
incerta A.J. Smith, 1908,
Melopsitiacus undulatus, 209; Platycercus
eximivs, 213; Barnardius barnardi, 21%.
Procyrnea mansion’ (Seurat, 1914), syn, Cyrnea
mansion’ (Seurat, 1914).
Milvus migrans, 90; Haliastur sphenurus, I;
Accipiter novaehollandiae, 94, Circus assimilis, 98;
Falco berigora, \04,
Procvrnea paraleptoptera (TH. Johnston &
Mawson, 1941b), syn, Cyrnea paruleploptera
T.H. Johnston & Mawson, 1941b.
baleo cenchroides, 105,
Procyrned sp., ? leptoptera (Rudolphi, 1819).
Aceipiter novaehollandiae, 94; halco berigora, 104,
Hudyelia acuariana (Gushanskaya, 1937).
Podareus strigvides, 246.
Hadjelia truncata (Creplin, 1825).
Eurystomus orientalis, 261.
Miguierd chabaudi Mawson, 1968d.
Podaraus sirigvides, 246,
Viewlera chibiae Mawson, 968d,
Dierurus hottentottus, 416.
Viguiera longicallis Mawson, 1968d,
Microeca leucophaea, 287; Colluricinela
harmonica, 294; Anthechaera carunculara, 350,
Viguiera sp.
Cincloseina cinnumomeum, 309; Malurus
leucopterus, 322; Corcorax melanorhamphus, 423.
Srellocarenema glareolue Mawson, 1968b.
Calidris ruficollis, 158, Stiltia isabella, 162.
Stellacaronema charadrii Mawson, 19686,
Charadrius rubricollis, \35.
Torquatvoides balanocephala (Gendre, 1922), syu.
Yorquatella balanocephala (Gendre, 1922).
Merops ornatus, 260,
Torquatoides sp., ? 7. torquata (Gendre, 1922), syn.
Torquatella sp,, Mawson, 1968d,
Centropus phasianus, 239.
Family TETRAMERIDAE Travassos, 1914
Geopeltitia chibiue Mawson, 1966,
Dicrura hoitentoitus, 416.
Geopetitia falco Mawson, 1966.
Faleo longipennis, 102.
Geopetitia streperue Mawson, 1966,
Strepera versicolor, 437.
Geopetitia sp.
Anthochaera carunetlata, 350.
? Mierohadjelia spiralis (Mawson, 1968d), syn,
Cyrnea spiralis Mawson, 1968d.
Chrysococeyx basalis, 235; Coracind novae-
hollandiae, 271; Lalage sueurii, 274; Pachycephala
pectoralis, 291; Cinclosma cinnamomeum, 309;
Philenton argenticeps, 356, Entomyzon evanotis,
359; Lichenvstomus plumulus, 372; Oriolus
sagittatus, 414,
Microtetrameres aevotheles Mawson, 1977.
Aegotheles cristatus, 248.
Microtetrameres cacomantis Mawson, 1977,
Cuculus variolosus, 233, C. flabelliformis, 234.
Microtetrameres circi Mawson, 1977.
Circus assimiilis, 98,
Microtetrameres caracinae Mawson, 1977.
Cuculus pallidus, 232; Coracina navaehollandiae,
271; C, papuensis, 272.
Microtetrameres cractici Mawson, 1977.
Cracticus (orqualus, 432.
Microtetrameres egretes Rastieed, 1960,
Ardeola ibis, 54.
Microtetrameres eopsaltriae Mawson, 1977.
Eopsaltria ausiralis, 284.
Microtetrameres gyrinorhinae Mawson, 1977.
Gymnorhina tibicen, 434.
Microtetrameres helix, Cram, 1927, syn.
Microtetrameres sp., Munday & Green, 1972.
Coercerax melunorhamphos, 423; Cervus coro-
noides, 438; C, rasmanicus, 439; C mellori, 440;
C. bennerti, 441; © orrti, 442.
Micretetrameres meliphagidaue Mawson, 1977
Anthochaera caruneulata, 350; A. chrysoprera,
352; Ll. sp., 353; Acunthagenvs rufogularts, 354;
Manorina melanacephala, 361; M, flavigula, 362,
Lichenastomius virescens, 366; L. lencotis, 367;
Philitonyris albifrans, 382.
300 REC, 5. AUST. MUS.
Microtetrameres mirafrae Mawson, |977.
Mirafra javanica, 265; Microeca leucophaea, 287,
Microtetrameres ninoctis Mawsan, 1977,
Ninox novaeseelandiae, 242.
Microtetrameres orialus Petroy & Tschertkova, 1950,
Oriolus sagi(tutus, 414.
Micratetrameres paraccipiter Mawson, 1977.
Accipiter fasciatus, 92; Aquila audax, 96.
Microtetrumeres pelecani (TH. Johnston & Mawson,
1942¢), syn. Tetrameres pelecani TH. Jatnstan
& Mawson, 1942¢.
Pelecanus conspicillatus, 37,
Micratetrameres philemon Mawson, 1977,
Philemon arventiceps, 356; P citreogularis, 358:
Entomvzon cyanotis, 35%.
Microtetrameres raptoris Mawson, 1977,
Faleo peregrinus, 101; Fo longipennis, \02; F;
berigara, 104; FL cenchroides, (05; Ninox
novaeseelandiae, 242,
Microtetrameres sphecotheres Mawson, 1977.
Sphecatheres viridis, 415,
Microtelrameres sireperae Mawson, 1977-
Streperd versicolor, 437,
Microtetrameres. tvionis Mawson, 1977.
Tyra alha, 244,
Microtelrameres sp., Mawsan, 1977.
Accipiter cirrhocephalus, 93, Cuculis
flabelliformis, 234; Lalage sueurit, 274; Grevica
gutluralis, 296, Pliloris paradiseus, 422.
Micretetrameres sp,
Accipiter fasciatus, 92; Chrysococeyx lucidus, 236;
Anthus novaeseclandiae, 270; Pachycephala
inornata, 290; Lichenostomus ornatus, 37\;
Struthidea cinerea, 424,
Tetrameres anseranas Mawson, 1979.
Anseranas semipalmata, 69.
Teirameres australis T.H. Johnsion & Mawson,
194le,
Crenus atrarus, 72.
Tetrameres biziurae TH. Johnston & Mawson, |94le.
Biziura lobata, 87,
Tetrameres calideis Mawsan, 1968b.
Calidris canutus, 155,
Tetrameres certa (Leidy 1886), syn. 2. diomedeae
T.H, Johnston & Mawson, 1942d,
Diomedea chlororhynchus, 16; DB. cauta, 17.
Tetrameres cladarhynchi Mawson, 1968b,
Cladorhynchus leueocephalus,, 143.
Telrameres dacelonis Mawson, 1979,
Dacelo nevaeguinede, 253.
Tetrameres fissispina (Diesing, 184l).
Anseranas semipalmata, 69, Anas superciliosa, 77,
A. castanea, 79.
Tetrameres globosa (Linstow, (879), syn, Fefraimeres
sp, Munday & Green, 1972,
Porzana pusilla, 116; P. fluminea, 117; P. tabuensis,.
118; Gallinula mortierii, 9; Perphyrio perphyrio,
122,
Teirameres #reent Mawson, 1980, syn. TD pelecanr
T.A. Johnston & Mawson, 1942e, in part.
Pelecanus conspicillarus, 37.
19(75): 219-325 August, 1986
Tetrameres eubanovi Shigin, 1957.
Podiceps — crixfarns, 3;
novachollandiae, 5.
fetrameres lobibicis Mawson, 1968b.
vanellus miles, 131.
Terrameres nouvelt (Seural, 1914),
Charadrius ruficapillus, 139;
leucocephalus, 142;
novaehollancliae, |44.
Terrameres scolaopacidis Mawson, \96%b.
Tringa glareola, 146; Calidris acuminata \57, ©
ruficollis, \58,
Tetrameres sp., Munday & Green, 1972,
Gallinula mortierii, 119; Vanellus miles,
Charadrius rificapillus, 139,
Tetrameres sp., Mawson, 1969,
Geochelidon nilotica, 165
Tetrameres sp., TH. Johnstan & Mawson, 1942e,
Anas gibberifrons, 78,
Tetrameres sp.
Phalucrocorax melanolencas, 47; Ardea
novaehollandiae, 53; Stlictonetta nuevosa, 73;
Tidorna tadarnoides, 75, Cygnus atratus, 72;
Anas gibberifrons, 78; A. rhynchotis, 80; Falco
langipennis, 102; #& berigaru, 104; Caturnix
ypsilophera, 109; Calidris ruficollis, \58; Larus
novuehollandiae, 163; Chlidonias hybrida, 164;
Ninox novaeseelandiae, 242; Malurus lambert,
320.
Tachybaplus
Himantopus
Reclirvirostra
131;
Superfamily ACUARTOLDEA Sobolev, 1949
Family ACUARIIDAE Railhliet, Flenry & Sisoif, 1912
Acuariidae, unidentified, T.H. Johnston & Mawson,
194 le.
Hydroprogne caspia, 166.
Acuariidae, unidentified.
Anas rhynchetis, 80; Malacorhynchus
membranaceus, 81; Turnix velox, 111; Porzana
Jluminea, WT, Ervthrogonys cinetus, 134, Anous
minutus, 174; Ephthianura albifrons, 393;
Emblema pieta, 402.
Subfamily Acuariinae Railliet, Henry & Sisoff, 1912
Acuaria aunthuris (Rudolphi, 1819), syn. Acuaria
corvicola T.H. Johnston & Mawson, [941a,
Corvus coronoides, 438; C. tasmanicus, 439; C.
mellori, 440; C, bennerti, 441, C. orru, 442.
Acuaria calluricinclae Mawson, 1972.
Pachvcephala rufiventris, 293; Colluricinela
harmonica, 294.
eleuaria microeca Mawson, 1972.
Microeca leucophaea, 287; Maluris leucopterus,
322.
Acuaria mirafrae Mawson, 1972.
Mirafra javanica, 265,
Aecuaria petterae Mawson, 1972.
Lalage leucomela, 275; Drymodes brunneapy gia,
278; Oreoica gutturalis, 296, Mviagra inquieta,
301; Cinclosoma cinnamomenm, 309;
Anihochaera sp., 353; Acanthagenys rufogularis,
354; Lichenostomus virescens, 366, L. plumulus,
HELMINTHS FROM AUSTRALIAN BIRDS Mil
372; Artamus cinereus, 429, Cracticus
nigrogularis, 433; Gymunorhina tibicen, 434.
Acuarta skrjabint Ozerskaya, 1926,
Erythrura trichroa, 412.
Alcvaria streperina T.H. Johnston & Mawson, 1941d.
Strepera graculina, 435; S. fuliginosa, 436; S.
versicolor, 437,
Aevaria sp., TH. Johnston & Mawson, 194le.
Chlidonias hybrida, 164,
Cheilospirura gruvelt (Gendre, 1913).
Corurnix yepstlophora, \O9,
Chevreuxia australis TH. Johnston & Mawson,
19d 1b,
Charadrius melanops, \40; Chlidonias hybrida,
lad,
Chevreuxia revoluta (Rudolphi, 1819),
Himantopus leucocephalus, 142; Reeurvirostra
novaehollandiae, 144,
Chevreiuxia sp.
Phalacracorax melanoleucos, 47.
Cosmocephalus juenschi TH. Johnston & Mawson,
194]e,
Pelecanus conspicillatus, 37; Leucocarba
Juscescens, 43; Phalacrocorax carbo, 44;
Chlidonias hybrida, 164; Sterna bereli, 171.
Echinuria heterobrachiata Wehr, 1937,
Charadrius rujficapillus, 139; Cladorhyachus
leucocephalus, 143; Recurvirastra
novaehollandiae, 144, Calidris ruficollis, 158.
Echinuria squamaia (Linstow, 1883),
Phalacrovarax carho, 44,
Echinuria uncinata (Rudophi, 1819), syn. Echinuria
querquedulae TH. Johnston & Mawson, 1942.
Cygnus arrurus, 72; Tadorna tadernoides, 75;
Anas gibberifrons, 78; A. castanea, 79%, Nettapus
pulchellus, 85,
Kehinuria sp.
Anas superciliosa, 77.
Pectinespirura arzentata Wehr, 1933.
Larus novaehollandiae, 163.
Skrjabinocerca sp.
Sterna beret, 171.
Skrjabinoclava decorala (Solonitzin, 1928).
Tringa terek, 15).
Skrjabinoclava horrida (Rudalphi, 1809), syn.
Skrjabinoctava sp., Munday & Green, 1972,
Tachvbapius novaehollandae, 5; Charadrius
rubricollis, 135; C. ruficapillus, 139.
Skrjabinoclava sp., ef.-alit Ali, 1968.
Threskiornis aethiapica, 65.
Skrjabinocla\va sp,, Munday & Green, 1972.
Charadrius rufieapillus, 139.
Skrjabinacliva sp,
Pachyptila vittata, 25; B turtur, 29, Threskiernis
uethiopica, 65; Tringa brevipes, 147; Culidris
ruficollis, 158,
Syneuaria contorta (Molin, 1858).
Threskiornis dethtopica, 65,
Syneuaria sp.
Padiceps erisfatus, 3;
novaehollandiae, 5.
Svnhimuntus affinis (Seurat, 1916),
Ninox strenua, 24),
Tachybaptus
Synhimantus falea Mawson, 1982.
falco berizora, 104; F. cenehrotdes, 105;
Philidonyris novaehollandiae, 380.
Synhimanius flindersi (TH. Johnston & Mawson,
1941b), syn. Acuaria /lindersi T.H. Johnston &
Mawson, 1941b; ‘nematode’, Cleland, 1922,
Falco herigora, 104.
Synhimantus fielding! (Baylis, 1934a), syn, Acueria
fielding Baylis, 1934a.
Accipiter novaehkollandiae, 94,
Synhimantus luticeps (Rudolphi, 1819).
Ninex novaeseelandiue, 242; Podurgus strigoides,
246.
Svnhimantus lichenastomi Mawson, 1982,
Meliphaga lewint, 363; Lichenostomus
penicillatus, 374.
Svnhimantus podargt Mawson, 1982,
Podargeus strivoides, 246.
Svnhimantus sirry Khalil, 1931, syn, Dispharvnan
pelecani T.H. Johnston & Mawson, 1942c.
Pelecanus conspicillatus, 37.
Svnhimantus sp., T.H, Johnston & Mawson, 194]e.
Anhinga melanogaster, 42,
Svahimantus sp., TH. Johnston & Mawson 1942¢.
Phalacrocorax melanoleucos, 47.
Svnahimantus sp,, syn, Dispharvax sp,
Phalacrocorax carbo, 44; P. melanoleucos, 47;
Stictonelta naevosa, 73; Accipiler cirrhocephalus,
93; Haliaeetus leucoguster, 98; Ninos
novaeseelandiue, 242; Petroica phoenica, 279;
Malurus cyaneus, 318, Corvus sp., 443.
Willmottia australis Mawson, 1982,
Malurus cyvaneus, 318.
Xenacordon patonae Mawson, 1982.
Phylidonyris novuehollandiae, 380,
Xenocordon gymnorhinis (de Chaneet & Robertson,
1983), syn, Cheilospirura gyninorhinis de
Chaneet & Robertson, 1983.
Gymnorhina tibicen, 434,
Subtamily Seuratiinae Chitwood & Wehr, 1932
Cheilonematodum falevonis TWH. Johnston &
Mawson, 1941d.
Halcyon sancta, 257.
Seuratia shipleyi (Stossich, 1900), syn. Seuraria
marina T.H. Johnston & Mawson, 194le.
Diomedea exulans, 12; D. melanaphrys, 13; D.
bulleri, 14, D. chrvsostonta, 15; BD chlororhyncha,
16; Phoebetria palpebrata, 18; Macronectes
bivanteus, 19; Fulmarus glacialoides, 20, Daptior
capense, 2\, Prerodroma macroptera, 22; P
lessoni, 23; P brevirastra, 24, Pachyptila vitrata,
25; P salvini, 26, P belcheri, 28; Po turiur, 29;
Puffinus pavificus, 3); & tenuirostris, 33;
Pelagodroma tnarina, 35: Sula leucogvasrer, Al;
Frevata minor, 49; F ariel, 50; Sterna fuscata, 168;
Anous stolidus, \73.
Stregophorus diomedeae (T.H. Johnsion & Mawson,
1942d), syn, Purvseria diomediae TH. Johnston
& Mawson, 1942d.
Diomedea exulans, 12; D. melamophrvs, 13. 2
chrysostoma, iS; D. canta, 17-
302
Stegophorus muacronectes (TH. Johuston &
Mawson, 1942d), syn. Parvseria mucronectes
TH. Johnston & Mawson, 1942d.
Diomedea chrysostoma, 15; Macronectes
viganieus, 19; Plerodroma lessoni, 23, RP.
brevirostris, 24.
Stezophorus pachyptilue (TH, Johnston & Mawson,
1942d), syn. Paryserta pachyptilae V.H
Johnston & Mawson 1942d.,
Duprion capense, 21; Pachyptila vittata, 25; P
salvini, 26; P. desolata, 27; PB turtur, 29; Puffinus
griseus, 32; P. tenuirosiris, 33.
Stezophorus stellaepaluris (Parona, 1901).
Puffinus tenuirostris, 33,
Stegophorus sp.
Diomedea exuluns, 12; Sterna bergii, 17),
Streptocara crassicauda (Creplin, 1829).
Alnseranas semipalmata, 69; Tadorna ladornoides,
75; Anas rhynchotis, 80; Aythya australis, 82;
Oxyura australis, 86; Vanellus niles, 131;
Charadrius bicinctus, 137; C. ruficapillus, 139-
Streplocara formosensis Sugimato, 1930.
Biziura lobata, 87,
Streptocura pectinifera Neumann, 1900.
Hydroprogne caspia, 166,
Streptocara recta (Linstow, 1879).
Poliocephalus poliocephalus, 4; Ptachybaptus
novdehollandiae, 5; Phalacrocerax sulcirastris, 46,
P melanoleucos, 47; Bizivra lobata, 87;
Chiidonias hybrida, 164.
Streptocara sp., TH. Johnston & Mawson, 1942e.
Anas gzibberifrons, 78,
Streptocara sp.. Munday & Green, 1972,
Avthya australis, 82; Charadrius rufieapillus, 139,
Streplocarad sp.
Tachybaptus novaehollandiae, 5; Anas gibheri-
rans, 18; A. rhyachotis, 80; Vanellus niles, 131;
Limosa lapponica, 54.
Subfamily Schistorophinae Travassas, 1918
Aneyracanthopsts sp.
Haleyon sancta, 257.
Schistogendria sp. 1.
Fuleunculus Jrontatus, 288; Psephodes olivaceus,
305; Lichenostomus penicillatus, 374.
Schistogendria sp, 2,
Halevon sanela, 257.
Schistogendria sp. 3,
Halcyon saneta, 257.
Schistogendria sp.
Colluricincla harmonicu, 294,
Schistorophus cornutus Sobolev, 1943
Trinwa terek, (54.
Schisioraphus limosde Mawson, 1968b.
Limosa lappanica, 154-
Schistorophus longicernis (Aemprich & Ehrenberg,
1866).
Calidris canutus, 155,
Schistorophus sp.
Dacelo leachii, 254,
Sciadocara umbellifera (Molin, 1860).
Calidris cunulus, 158.
REC. S. AUST, MUS. 1915): 219-325
Akeust, 1986
Sciadocara sp.
Haematopus fulizinosus, 130.
Viktorocara limosue Mawson, 1968a,
Tringa lerek, 151; Limosa lapponica, \54.
Vikrorocara schejkini Gushanskaya, 1950,
Tringa terek, 151.
Viktorocara sp., Mawson, 1968b.
Culidris canutus, 1558.
Superfamily DIPLOTRIAENOIDEA
Family DIPLOTRIAENIDAE Skrjabin, 1916
Diplotriaena alpha V1. Johnston & Mawson, 1940a,
syn, ‘Filaria sp.’, Cleland, 1922.
Sericornis Juliginosus, 333; Strepera graculina,
435,
Diplotriaena heveridge’ Bain & Mawson, 1981.
Corvus arru, 442.
Diplotriaena clelandi (TJ, Johnston, 1912b), syn.
Filaria clelandi TH. Johnston, 1912b.
Gymnorhina tibicen, 434,
Diplotriaena delia TH. Johnston & Mawson, 1940a.
Malurus lainberti, 320; Amytornis goyderi, 324.
Diplotriaena epsilon TH, Johnston & Mawson,
1940a.
Cracticus lorquatus, 432.
Diplorriaena falconis (Connal, 1912).
Falvo berigora, 104,
Diplotriaena flabellata (Linstow, 1888), syn,
Diplotriuend beta T.A. Johnston & Mawson,
1940a,
Corvus orru, 442.
Diplotriaena golvani Anderson, 1959.
Colluricinela harmonica, 294,
Diplotriaena halli Qgden, 1967.
Monarcha trivirgata, 297; Malurus lambert, 320,
Diplotriaena pungens (Schneider, 1866),
Sphecotheres viridis, 415.
Diplotrigaena smithi Bain & Mawson, J981, syn.
Diplotriaenu sp. Mackerras, 1962,
Acunthagenys rufogularis, 354; Phileman
corniculatus, 357.
Diplotrivena spratti Bain & Mawson, L98I.
Oreoica gutturalis, 296,
Diplotriaena tricuspis (Fedschenko, 1874),
Coracinu papuensis, 272; Cisticola exilis, 315.
Diplotriaena tridens (Molin, 1858).
Sphecotheres viridis, 415.
Diplotriaena zeta T.H. Johnston & Mawson, 1940a.
Acanthagenys rufagularis, 354,
Diplotriaena sp., Mackerras, 1962,
Dierurus hottentotius, 416.
Diplotriaena sp., Munday & Green, 1972.
Anthochaera paradoxa, 35),
Diplotriaena sp., Ogden, 1967.
Arses kuupi, 298; Zosterops lateralis, 400.
Diplotrigena sp.
Coracina papuensis, 272; Micraéca leucophaea,
287; Colluricinela megarhynchus, 295; Malurus
cyaneus, 318; M. lamberti, 320; Pardaloius
sirratus, 399; Zosterops lateralis, 400; Dicrurus
hottentottus, 416; Streperu filiginasus, 436;
Corvus orru, 442.
HELMINTHS FROM AUSTRALIAN BIRDS 403
Hamatospiculum chibiae TH. Johnston & Mawson,
1941a.
Dicrurus hottentottus, 416,
Hamatospiculum halevonis TAH. Johnston &
Mawson, 194la.
Halevon pyrrhopygius, 256; HH. sancta, 257.
Hamatospiculum howense TH. Johnston &
Mawson, 1940a, syn. Filariasp., TH. Johnston,
19]2b.
Halcyon sancta, 257,
Hamatospiculum meneilli TH. Johnston & Mawson,
1941b, syn, ‘nematode’ Cleland, 1922, Ninox
rufa, 240, N. novaeseelandiae, 242.
Hamatospiculun sp,, ef, quddridens (Molin, 1858).
Faleo hypoleucos, 103.
Hamatospiculunmt sp., TAH. Johnston & Mawson,
1941b,
Avcipiter cirrhocephalus, 93.
Hamatospiculum sp,
Haleyon pyrrhopyetus, 256.
Serratospiculum eguttatim (Schneider, 1866), syn.
Serratospiculum atienuatum (Rudolph, 1803);
Filaria gutiata Sctineider, 1866.
Accipiter cirrhucephalus, 93; Falco peregrinus,
101; A longipennis, 102; E Avpoleucos, 103; F:
herigora, \04,
Serratospiculum tendo (Nitzsch in Giebel, 1857), syn.
S. gudtalum (Schneider, 1966) in part,
Accipiter cirrhocephalus, 93; Falco subniger, 100;
I. peregrinus, \O1; F longipennis, 102.
Superfamily APROCTOIDEA
Vamily APROCTIDAE Yorke & Maplestone, 1926
Aproctidae, unidentified, Munday & Green, 1972.
Pefroica multicolor, 280; P xoadenavii, 281,
Aprocta australis (TH. Johnston & Mawson, 1942a),
syn. Maerifilaria australis TH, Johnston &
Mawson, 1942a,
Centropus phusianus, 239.
Aprocta hakeri Bain & Mawson, (981.
Corvus coronatdes, 448; © mellori, 440; C. orru,
442.
Aprocta boulengeri Bain & Mawson, 1981.
Srrepera graculina, 435.
Aprocta corvicala TAL, Johnston & Mawson, 1940a.
Cervus orru, 442.
Aprocta vestibulaia (TH. Johnston & Mawson,
1940a), syn. Austrofilaria vestibulata T.H.
Johnston & Mawson, 1940a.
Aphelocephala nigricincia, 344.
Aprocta sp.
Cucatya sq., 193.
Lissonema brevicaudaia (Chaw, 1939), syn. Aprocta
brevieaudata Chow, 1939.
Ninox novueseelundiae, 242.
Lissonema rotundalum (Linstow, 1903), svi.
Aprocta rorundara Linstow, 1903,
Centropus phasianus, 239.
Lissonema sp., Bain & Mawson, 1981.
Ninex novaeseelandiae, 242,
Lissonema sp.
Alectura lathajri, \O7.
Mawsonfilaria rhipidurae (T.H. Johnston &
Mawson, 1952), syn, Austrofilaria rhipidurae,
T.H. Johnston & Mawson, 1952.
Rhinidura leucophrys, 304.
Pseudaprocta copemani Bain & Mawson, 1981, syn,
Aproctidae, unidentified, Munday & Green,
1972.
Petroica multicolor, 280; Pachycephala pectoralts,
291,
Pseudaprocta myzanthae T.H. Johnston & Mawsun,
1940a, syn. ‘nematode’, Cleland, 1922.
Manorina flavigula, 362.
Desmidocercella sy.
Phalacrocorax sulcirostris, 46; P. melanvleucos,
47; Ardea pacifica, 52; Nycticorax caledonicus, 60;
Ixobrychus ininutus, 6),
Diomedenema diomediae T.H. Johnston & Mawson,
1952,
Diomedea chrysestamu, 15; D. cuuta, 17.
Superfamily FILARIOIDEA
Unidentified filarial worms cited by:
Bancroft, 1889,
Anthochaera chrysoptera, 352; Matorinu
melanocephala, 361; Cracticus torquatus, 432;
Corvus orru, 442.
Cleland, 1922.
Falco longipennis, 102; Petroicu phoenica, 279,
Malurus evyaneus, 318; Geoffravus geoffroyi, 1981.
T.H. Johnston, 19) 2a.
Philemon citreoguluris, 358.
T.H. Johnston, 1912b.
Ninox novaeseelandiue, 242; Coracina papuensis,
272; Acanthagenys rufogularis, 354.
T.H. Johnston & Mawson, 1940a,
Nymphieus hollandieus, 208; Acan(thagernys
rufogularis, 354.
T.H. Johnston & Mawson, 1941la.
Mulurus leucopterus, 322.
Mackerras, 1962.
Ardeu pacifica, 52; Corvus bennetti, 442.
Plimmer, 1912.
Ailuroedus melanotis, 420.
Unidentified filarial worms:
Budyptula minor, \\; Pelecanus conspicillatus, 37;
Larus novaehollandiae, 163; Ninox connivens,
243; Myiagra rubeculus, 299; M. inguteta, 301;
Alluroedus crassirostris, 421.
Family ONCHOCERCIDAE Leiper, 1911
Onchocercidae, unidentified.
Sphecotheres \iridis, 415.
Cardiefilaria dubia (TH, Johnston & Mawson,
1940a), syn. Carinema dubia T.M. Johnston &
Mawson, 1940a, ‘Filarial’ worms, Cleland, 1922.
Gea/froyus geoffrovi, \98.
Eufilaria sp,, Mawson, 1969.
Larus novaehollandiae, 163.
Paralemdana clelandi TH. Johnston & Mawson,
1940a,
Strepera graculina, 435,
304
REC, 5, AUST. MUS, 19(15): 219-325 Augusl, 1986
Pelecitus fulicaeatrae (Diesing, 1861).
Fulica atra, 123.
Pseudaproctella sp., Mawson, 1968b.
Vanellus miles, 131,
Splendidofilaria sp,
Struthidia cinerea, 424.
Splendidofilariinag, unidentified,
Falco longipennis, 102; Platyeercus venustus, 215;
Gymnorhina tibicen, 434.
Microfilaria gymnorhinae Gilruth, Sweet & Dodd,
1910,
Gyninorhina tibicen, 434.
Microfilaria sp.
Prilonorhynchus vieluceus, 417,
Microfilaria sp. have been recorded by the following:
Bancroft, 1889,
Trichoglossus haemalodus, 199; Podargus
strigoides, 246, Eurystomus orientalis, 261;
Myiagra rubeculus, 299; Pomatostomus
temporalis, 310; Anthochaera chrysoptera, 352;
Entamyzon cyanotis, 359; Manorina
melanocephala, 361; Orielus sugittatus, 414;
Dicrurus heattentorttus, 416; Sericul/us
chrysocephalus, 418; Cracticus torquatus, 432;
Gymnorhina tibicen, 434; Strepera graculina, 435;
Corvus orru, 442.
Breinl, 1913a.
Falca hypoleucas, 103; Pitta versicolar, 263,
Cleland, 1915.
Podargus strivoides, 246, Eurvstomus orientalis,
261; Colluricinela harmonica, 294, Myzomela
sanguinolenta, 390),
Cleland, 1922.
Philemon citreogilaris, 358,
Cleland & T.H. Johnston, 1910.
Anthochaera chrysoptera, 352; Gymnorhina
tibicen, 434; Corvus arru, 442,
Cleland & T.H. Johnston, 1912.
Phalacrocorax melanoleucas, 47; Accipiter
cirrhocephalus, 93; Glossupsitia pusilla, 202;
Podareus strigoides, 246; Eurystomus orientalis,
261; Psophedes olivaceus, 305, Plectorhyacha
lanceolata, 355; Enlomyzon cvanotis, 359;
Manorina melanocephala, 361; Lichenostoiius
Juscus, 373; Lichmera indistincta, 378; Myzamelu
sanguinolenta, 390; Pardaloius striatus, 399;
Oriolus Sagitlalus, 414; Corcorax
melanorhamphos, 423; Struthidea cinerea, 424;
Arramus leucorhynchus, 426; A. cinerea, 429; A.
cyanopterus, 430, Cracticus torquatus, 432; C.
nigrogularis, 433, ©. orru, 442.
T.H, Johnston, 1910,
Zoothera dauma, 276; Anthochaera chrvsoplera,
352.
T.H,. Johnston, 1912a,
Anhinga melanogaster, 42; Phalacrocorax
sulctrasiris,
46,
T.H. Johnston, 1916.
Aegotheles eristatus, 248.
Mackerras, 1962,
Phalacracorax carbo, 44, Haliastur sphenurus, 91;
Accipiter cirrhocephalus, 93, Vanellus miles, 131,
Cacatua sanguinea, 194, C. walerita, 196;
Glossepsitta concinna, 200; Platycercus adsictus,
214; Coracina novaehollandiae, 271; Climacteris
melanura, 349; Anihachuaera chrysoptera, 352;
Entomyson cyanotis, 359; Melithreptus
alboxularis, 376, Zosterops lateralis, 400;
Sphecotheres viridis, 415, Dicrurus hottenrortus,
416; Ptilonorhynchus violaceus, 417; Corcorux
melanorhumphus, 423; Grallina cvanoleuca, 425;
Artamus personatus, 427; A, cvanopterys, 430;
Cracticus torqualus, 432; Gymneorhina tibicen,
434,
Plimmer, 1912.
Geophaps plumifera, 184; Platvcercus icleratis,
216; Entamyzon cvanetis, 359; Manorina
melanocephala, 361; Gymnorhina tibiven, 434,
Strepera fulizinosa, 436,
Plimmer, 1914.
Geophaps plumifera, V4; Platycercus icterolis,
216.
Plimmer, 1915,
Poephila bichenovii, 407.
Plimmer, 1916.
Artamus superciliosa, 428.
Scott, 1926.
Larus novaehollandiae, 163; Poephila personata,
408.
Seort, 1927.
Lurystomus orientalis, 261.
Nematoda not further identified are not inchided in
this list.
PHYLUM ACANTHOCEPHALA
Order Echinorhynchidea
Family FILICOLLIDAE Petroschenko, 1956
Filicollis sphaerocephalus (Bremser, 1819).
Haematopus fuliginosus, 130; Larus
novaehollandiae, 163.
Family PLAGIORHYNCHIDAE Golvan, 1960
Plagiorhvachus charadrii (Yamaguli, 1939b), syn,
Prosthorhynchus charadrii Yamaguti, 1939b.
Charadrius rubricollis, 135; C. ruficapillus, 139.
Plagiorhynachus menurue (TH. Johnston, 1912b),
syn, Prosthorhynchus menurae (TH: Johnston,
1912b), Echinorhyvnchus menurae T.H,.
Johnston, 1912b.
Menurae novaehollandiue, 264,
Plagiorhynchus sp,
Pachyptila turrur, 29,
Family POLYMORPHIDAE Meyer, 193]
Polymorphus biziurae V.H. Johoston & Edmonds,
1948,
Pelecanus conspicillatus, 37; Threskiornis
aethiopieus, 65; Platalea flavipes, 68; Biziura
lobuta, 87,
HELMINTHS FROM AUSTRALIAN BIRDS
Polymorphus sp.
Cyenus atratus, 72.
Corynasoima clovatum Goss, 1940.
Leucocarbo fiscescens, 43; Phalacrocorax varius,
45: P. sulcirosiris, 465 P. melanoleucos, 47.
Corvnosoma sp.
Budyptyla minor, Uy Cygnus atraius, 72.
Arhyvthmorhynchus brevis yan Cleave, 1916,
Botuurus poiciloptilus, 62.
Arhythmorhynchus johnstoni Golvan, 1960, syn.
Arivitmorhynachus frassoni of T.H. Johnston
& Edmonds, 1950 not (Molin, 1858),
Numenius madagascariensis, 145,
Arhythmorhynchus limosae Edmonds, 1971,
Limosa lapponica, 154.
“Arhythmorhynechus sp.
Numenius madagascariensis, \4§,
Order Gigantorhynchidea
Family CENTRORHY NCHIDAE van Cleave, 1916
Centraryhnchus asturinus (TH. Johnston, 1913),
syn. Echinorhynchus bazae Southwell & Macfie,
1925; syn. Giventorhpachus asturinus TA.
Johaston, 1912.
Aviceda subvristuta, 89; Accipiter fasciatus, 92; A,
cirrocephalus, 93; A. novaehollandiae, 94; Circus
approximins, 99, Faleo berigora, \04; FF
cenchroides, 105,
Centrorhynchus bancrofti (TH. Johuston & Best,
1943), syn. Gordiorhynachus bancrofti T.H.
Johnston & Best, 1943,
Ninox strenua, 241; N, noyueseelandiae, 242.
Centrorhynchus falconis (TH. Johnston & Best,
1943), syn. Gordiorhynchus Jfalconis T.H.
Johnston & Best, 1943.
Fulco berigora, \O4,
Centrorhynchus horridus (Linstow, 1898a).
Dacelo novaeguineae, 253; Halcyon sanctus, 257.
Centrarhvachus ap. T.A. Johnston, 1918b.
Ninox novaeseelandiue, 242
Centrarhvachus sp.
Phalacracorax sulcirostris, 46,
Family GIGANTORHYNCHIDAE Hamman, (892
Mediorhynchus alecturae (TH. Johnston &
Edmonds, 1947), syn. Bpodius aleciurae T.A.
Johnston & Edmonds 1947,
vlectura lathami, 107,
Mediorhyachus coreoerucis TH. Johnston &
bdmonds, 1951, syn. Echinarhynchus 4$p.
Cleland 1922.
Corcorax melanorhamphus, 423; Corvus
fasmunicus, 439, ©, mellori, 440; C. bennetri, 44);
Corvus sp,, 443,
Mediorhynchus garruli (YVamaguti, 1939b),
Grallina cyanoleuca, 425.
Mediorhynchus sp.
Colluricincla harmontcu, 294; Rhipidura
leucophrys, 304; Anthochoera chrysaplera, 352;
Acunthagenys rufogularis, 354; Grallina
evanoleuca, 425; Corvus sp., 443.
3s
Family OLIGACANTHORHYNCHIDAE Southwell
& Macfie, 1925
Oncicola pomatostom) (TH. Johnston & Cleland,
1912) (eneysted stage), syn. Echinorhyachus
pomatostomi T,H. Johnston & Cleland, 1912;
syn. Oligacanthorhynchus pamatostomi (T,F,
Johnston & Cleland, 1912).
Tirnix castanola, 110; T. velox, Vb, Pedionamus
terquatus, 13; Anthus novaeseelandiae, 270;
Lalage leucomela, 275; Zoothera dauma, 276;
Melanodryas cucullara, 282; Microeca
leucophaea, 287; Pachycephala inornata, 290, P
rufiventris, 293; Colluricincla harmonica, 294;
Oreoica gutluralis, 296; Cinclosoma castanotum,
308; C cinnaimameum, 309, Pomatostomus
temporalis, 310; FP. superciliosus, 311; PB ryficeps,
312; Amptornis purnelli, 326; Serivornis
prrrhopyeius, 330; S. cautus, 331, 8. brunneus,
332; S. fuliginosus, 333; Acanthiza chrysorrhoa,
341; Aphelocephala leucopsis, 343; Duphoenositta
chrvsoptera, 345; Climacteris leucophaea, 346; C.
picumnus, 347; C. melanura, 349; Anthochoera
carunculata, 350; Manorina flavigula, 362;
Lichenosromus virescens, 366; L. plumulus, 372;
Poephila cincta, 410; Gralline eyanoleuca, 423,
Artamus cinereus, 429; Gymnorhina libicen, 434.
Family PROSTHORHYNCHIDAE Petroschenko,
1956
Porrorchis hylae (T.H. Johnston, 1914b), syn.
Eehinorhynchus bulbocaudatus Southwell &
Mactlie, 1925, syn. Gordiorhynehus hylae (T.H.
Johnston, 1914b), syn. Pseudoporrorchis hylae
(TH, Johnston, 1914b).
Centropus phasianinus, 239, Podargus strigoides,
246. 131,
Prosthorhynchus cylindraceus (Goeze, 1782).
Gymnorhina tibicen, 434,
Prosthorhynchus sp.
Haematopus longirostris, \29; Varnellis miles, (31.
LIST 3. INDEX TO BIRD FAMILIES REFERRED
TO IN THIS CHECKLIST
In this list the number of bird species listed by
Schodde e a/. (1978) for each family is given after the
family name, followed by the number of birds from
which helminths have been recorded, Introduced
species are omitted.
Acanthizidae, 41/12 p.272
Accipitridae, 12/17 242
Aegothelidae, 1/1 264
Alaudidae, 1/1 266
Aleedinidae, 10/8 264
Anatidae, 21/1] 236
Anhingidae, 1/1 22
Apodidae, 5/1 264
Ardeidae, 16/11 232
Artamidae, 6/5 281
Burhinidae, 2/1 248
306
Cacatuidae, 11/6
Campephagidae, 7/4
Caprimulgidae, 3/2
Casuariidae, 1/1
Charadriidae, 16/11
Ciconiidae, 1/1
Climacteridae, 7/3
Columbidae, 22/1]
Coraciidae, 1/1
Corcoracidae, 2/2
Corvidae, 5/5
Cractividae, 8/5
Cuculidae, 13/8
Dicaeidae, 1/0
Dicruridae, 1/1
Diomedeidae, 48/23
Dromaiidae, 2/1
Ephthianuridae, 5/1
Falconidae, 6/6
Fregatidae, 3/2
Glareolidae, 2/1
Grallinidae, 1/1
Gruidae, 2/2
Haematopodidae, 2/2
Hirundinidae, 6/4
Jacanidae, 1/1
Laridae, 26/12
Loridae, 7/3
Maluridae, 18/6
Megapodiidae, 3/2
Meliphagidae, 66/28
Menuridae, 2/1
Meropidae, 1/1
Motacillidae, 5/1
Muscicapidae, 52/65
Neosittidae, 1/1
Oceanitidae, 6/1
Oriolidae, 3/2
Orthonychidae, 10/5
Otididae, 1/1
Paradisaeidae, 13/5
Pardalotidae, 3/2
Pedionomidae, 1/1
Pelecanidae, 1/1
Pelecanoididae, 2/1
Phaethontidae, 2/1
Phalacrocoracidae, 5/5
Phasianidae, 4/2
Pittidae, 4/2
Plataleidae, 5/5
Platycercidae, 26/18
Ploceidae, 22/7
Podargidae, 3/2
Podicipedidae, 3/3
Polytelidae, 6/6
Psittacidae, 2/2
Rallidae, 17/10
Recurvirostridae, 3/3
Scolopacidae, 33/17
Spheniscidae, 10/5
Strigidae, 4/4
Sulidae, 4/4
Sylviidae, 9/2
REC. 5, AUST. MUS, [9(15): 219-325
257
Timaliidae, 4/3
Turnicidae, 7/3
Tytonidae, 4/2
Zosteropidae, 3/1
August, 1986
270
246
263
278
LIST 4. INDEX TO BIRD SPECIES REFERRED
TO IN THIS CHECKLIST
Bird name Bird number
Acanthagenys rufogularis Gould 354
Acanthiza apicalis Gould 337
chrysorrhoa (Quoy & Gaimard) 341
iredalei Mathews 340
lineata Gould 342
pusilla (Shaw) 336
reguloides Vigors & Horsfield 339
uropygialis Gould 338
Acanthorhynchus tenuirostris (Latham) 386
Accipiter cirrhacephalus (Vieillot) 93
Sasciatus (Vigors & Horsfield) 92
novaehollandiae (Gmelin) 94
Aegotheles cristatus (J. White) 248
Ailuroedus crassirostris (Paykull) 421
melanotis (Gray) 420
Alectura lathami Gray 107
Alisterus scapularis (Lichtenstein) 203
Amytornis govderi (Gould) 324
purnelli Mathews 326
textilis (Dumont) 325
Anas castanea (Eyton) 79
gibberifrons S. Muller 78
rhynchotis Latham 80
superciliosa Gmelin 77
Anhinga melanogaster Pennant 42
Anous minutus Boie 174
stolidus Linneé 173
Anseranas semipalmata (Latham) 69
Anthochaera carunculata (White) 350
chrysoptera (Latham) 352
paradoxa (Daudin) 351
sp. 353
Anthus novaeseelandiae (Gmelin) 270
Aphelocephala leucapsis (Gould) 343
nigricincta (North) 344
Aprosmictus erythropterus Gmelin 204
Aptenodytes patagonicus Miller 7
Aquila audax (Latham) 96
Ardea novaehollandiae Latham 53
pacifica Latham 52
Ardeola ibis (Linné) 54
Ardeotis australis (Gray) 126
Arses kaupi Gould 298
Artamus cinereus Vicillot 429
cyanopterus (Latham) 430
leucorhynchus (Linne) 426
minor Vicillot 43]
personatus (Gould) 427
superciliosus (Gould) 428
Ashbyia lovensis (Ashby) 394
Aviceda subcristata Gould 89
Aythva australis (Eyton) 82
HELMINTHS FROM AUSTRALIAN BIRDS
Barnurdius barnardi (Vigors & Horsfield)
zonarius (Shaw)
Biztura lobata (Shaw)
Botaurus poiciloptilus (Wagler)
Burhinus magnirostris (Latham)
Butorides striatus (Linne)
Cacatua galerita Latham
leadheateri (Vigors)
roseicapilla Vieillot
sanedinea Gould
fenuirostris (Kuhl)
sp.
Calidris acuminata (Haorsfield)
alba (Pallas)
canutus (Linneé)
Jerruginea (Pontopiddan)
ruficollis (Pallas)
tenuirostris (Horstield)
Callocephalon fimbriatum (Grant)
Calyptorhynchus Junereus (Shaw)
magnifieus (Shaw)
Cuaprimulgus areus (Hartert)
Casuarius casuarius (Linne)
Cecropsis ariel (Giould)
nigricans (Vieillot)
Centropus phasianinus (Latham)
Cereopsis novaehollandiae Latham
Certhionyx pectoralis (Gould)
Certhionyx variegatus Lesson
Ceyx azurea (Latham)
Chalcophaps indica (Linne)
Charadrius bicinetus Jardine & Selby
leschenaultil Lesson
melanops Vieillot
mongolus Pallas
rubricollis Gmelin
ruficapillus Termmink
Chenonetta jubata (Latham)
Cheramoeca leucosternum (Gould)
Chlamydera maculata (Gould)
Chlidonias hybrida (Pallas)
Chrysocoecyx basalis (Horsticld)
lucidus (Grnelin)
218
219
87
62
128
59
196
195
19]
194
192
193
157
160
155
159
158
156
190
189
188
250
>
269
268
239
74
387
388
252
179
137
138
140
136
135
139
83
267
419
164
235
236
Cinclorhamphus cruralis (Vigors & Horsfield) 317
mathewsi Iredale
Cinclosoma castanoitum Gould
cinnamomeuin Gould
punctatuin (Shaw)
Circus approximans Peales
assimilis Jardine & Selby
Cisticola exilis (Vigors & Horsfield)
Cladorhynchus leucocephalus (Vieillov)
Climacteris leacophaea (Latham)
melanura Gould
picumaus Temminck
rufu Gould
Collurivincla hurmonica (Latham)
megarhyncha (Qnoy & Gaimard)
Collocalia spodiopvgia (Peale)
Conopophila albogularis (Gould)
rufogularis (Gould)
Coracina novaehollundiae (Gmelin)
papuensis (Gmelin)
316
308
309
307
99
OR
315
143
346
349
347
348
294
295
251
384
385
27\
272
Corcorax melanorhamphus (Vieillot)
Corvus bennetti North
voronoides Vigors & Horstield
mellori Mathews
orru Bonaparte
fasmanicus Mathews
sp.
Coturnix novaezelandiae Quoy & Gaimard
ypsilophora Bose
Cracticus nigrogularis (Gould)
forquatus (Latham)
Cuculus pallidus (Latham)
Hlabelliformis (Latham)
variolosus Vigors & Horsfield
Cygnus atratus (Latham)
Dacelo leachii Vigors & Horsficld
novaeguineae (Hermann)
Daphoenositla chrysoplera (Latham)
Daplion capense (Linneé)
Dasvornis broadbenti (McCoy)
Dendrocyena arcuata (Horstield)
eytoni (Eyton)
Dicuewm hirundinaceum (Shaw)
Dierurus hottentottus (inne)
Diomedea bullert Rothschild
cauta Gould
chlororhyachos Gmelin
chrysostoma Forster
exuluns Linne
melanophrys Temminck
Dromaius novaehollandiae (Latham)
Drymodes brunneopygia Gould
superciliaris Gould
Eclectus roratus (P.L.S. Mueller)
Eeretta alba (Linné)
garzetta (Linné)
intermedia (Wagler)
sacra (Gmelin)
Elanus notatus Gould
Emblema bellum (Latham)
guttatum (Shaw)
pietum Gould
temporale (Latham)
Entomyzon cyanotis (Latham)
Eepsaltria australis (Shaw)
griseogularis Gould
pulverulenta Salvadori
Ephthianura albifrons (Jardine & Selby)
aurifrons Gould
tricolor Gould
Eremiornis carteri North
Erythrura gouldiae (Gould)
trichroa (Kittlitz)
Erythrogonys cincius Gould
Eudynamys scolopacea (Linne)
Eudyptes chrysacome (Forster)
pachyrhynechus Gray
schlegeli Finsch
Eudyptula minor (Forster)
Eurostopodus mystacalis (Temminck)
Eurystomus orientalis (inne)
Falco berigora Visors & Horstield
cenchroides Vigors & Horsfield
308
hypoleucos Gould
longipennis Swainson
peregrinus Tunstall
subniger Gray
Falcunculus frontatis (Latham)
Fregaia ariel (Gray)
minor (Gmelin)
Fulica atra Linné
Fulmarus glacialoides (Smith)
Gallinago hardwicki (Gray)
megala Swinhoe
Gallinula mortierii (Du Bus)
tenebrosa Gould
ventralis Gould
Gelochelidon nilotica (Gmelin)
Geoffroyus geoffroyi (Bechstein)
Geopelia cuneata (Latham)
Aumeralis (Temminck)
placida (Gould)
Geuphaps plumifera Gould
seripta (Temininck)
sinithi (Jardine & Selby)
Gerveone fusca (Gould)
Glossepsitta concinna (Shaw)
porphyrocephala (Dietrichsen)
pusilla (White)
Grallina cvyanoleuca (Latham)
Grus antigone (Linne)
rubicundus (Perry)
Gymnorhina tibicen (Latham)
Haematopus fuliginosus Gould
longirostris Vieillot
Haleyon chloris (Boddaerl)
inacleavii Jardine & Selby
pyrrhopygia Gould
sancta Vigors & Horsfield
Haliaeeius leucogaster (Gmelin)
Haltustur sphenurus (Vieillot)
Hierawetus morphnoides (Gould)
Himantopus leucocephalus Gould
Hirundo neoxena Gould
Hydroprogne caspia (Pallas)
Trediparra gallinaceu (Temminck)
Ixobrychus minutus (Linne)
Lalage leycomela (Vigots & Horsfield)
sueurii (Vieillot)
Larus novaehollandiae Stephens
Lathamus discolor (While)
Leipoa ocellata Gould
Leucocarbo fuscescens (Vieillot)
Leucosarcia nelanoleuca (Latham)
Lichenostomus chrysops (Latham)
cratitius (Gould)
Juscus (Gould)
keartlandi (North)
feuecotis (Latham)
melanops (Latham)
ornatus (Gould)
penicillatus (Could)
plumulus (Gould)
virescens (Vieillot)
Lichmera indistincta (Vigors & Horsfield)
Limicola falcinellus (Pontopiddan)
REC. S. AUST. MUS, 19(15): 219-325
103
102
101
100
288
50
49
123
Limosa lapponica (Linné)
Lonechura castaneotharax (Gould)
Macrenectes giganteus (Gmelin)
Macropyeia phasianella (Temminck)
Malacorhynchus membranaceus (Latham)
Matlurus cyaneus (Latham)
lamberti Vigors & Horsfield
leucopterus Dumont
pulcherrimus Gould
splendens (Quoy & Gaimard)
Manorina flavigula (Gould)
melanocephala (Latham)
melanophrys (Latham)
Megalurus gramineus (Gould)
Melanodryas cucullata (Latham)
Meliphaga lewint (Swainson)
Melithrepius albogularis Gould
brevirastris (Vigors & Horsfield)
lunatus (Vieillot)
Melopsittacus undulatus (Shaw)
Menura novaehollandiae Latham
Merops ornatus Latham
Microeca flavigaster Gould
leucophaea (Latham)
Milvus migrans (Boddaert)
Mirafra javanica Horsfield
Monarcha trivirgatus (Temminck)
Myiagra alecto (Temminck)
inquicta (Latham)
rubecula (Latham)
Myzomela obseura Gould
sanguinolenta (Latham)
Neochmia phaeton (Hombron & Jaequinot)
Neophema bourkir (Gould)
chrysostomu (Kuhl)
elegans (Gould)
petrophila (Gould)
pulchella Shaw
splendida (Gould)
Nettapus coromundelianus (Gmelin)
pulchellus Gould
Ninox connivens (Latham)
novaeseelandiae (Gmelin)
rufa (Gould)
strenua (Gould)
Northiella haematogaster (Gould)
narethae (H.L. White)
Numenius madagascariensis (Linne)
Nyvelicorax caledonicus (Gmelin)
Nymphicus hollandicus (Kerr)
Ocyphaps lophates (Yemminck & Langier)
Oreoica guituralis (Vigors & Horsfield)
Oriolus sagittatus (Latham)
Oxyura australis Gould
Pachycephala inornata Gould
melanura Gould
olivacea Vigors & Horsfield
pectoralis (Latham)
rufiveniris (Latham)
Pachyptila beleheri (Mathews)
desolata (Gmelin)
salvini (Mathews)
turtur (Kubl)
Auuusl, 1986
390
405
22f
227
228
229
230
231
84
85
243
242
240
24]
224
225
145
40
208
182
296
414
86
290
292
289
291
293
28
27
26
29
HELMINTHS PROM AUSTRALIAN BIRDS
vittaia (Forster) 25
Pardalotus punctalus (Shaw & Nodder) 396
rubricatus Gould 398
striatus (Gmelin) 399
xanthopygus McCoy 397
Pedionomus tarquatus Gould 3
Pelagodroma marina (Latham) 38
Pelecanoides urinatrix (Gmelin) 36
Pelecanus conspicillatus (Temminck) 37
Pelitohyas australis Gould 141
Petroica goodenovii (Vigors & Horstield) 281
multicolor (Gmelin) 280
Phoenicea Gould 279
Petrophassa albipennis Gould 183
Phaethon rubricauda Boddaert 51
Phalacrocorax carbo Linn’ 44
melanoleucos (Vieillot) 47
sulcirostris (Brandt) 46
varius (Gmelin) 45
sp, 48
Phaps chalcoptera (Latham) 180
elegans (Temminck) 181
Philemon argenticeps (Gould) 356
citreogularis (Gould) 358
corniculaius (Latham) 357
Phoebetria palpebrata (Forster) 18
Phylidonyris albifrons (Gould) 382
melanops (Latham) 383
nigra (Bechstein) 38]
novaehollandiae (Latham) 380
pyrrhoptera (Latham) 379
Pitta erythrogaster Temminck 262
versicolor Swainson 263
Platalea flavipes Gould 68
regia Gould 67
Platycercus adscitus (Latham) 214
caledonicus (Gmelin) 2
elegans (Gmelin) 212
eximius (Shaw) 213
icterotis (Kuhl) 216
venustus (Kuhl) 15
sp. 217
Plectorhyncha lanceolata Gould 355
Plegadis Jalcinellus (Linne) 64
Pluvialis dominica (P.,8, Miller) 133
Podargus ocellatus (Quoy & Gaimard) 247
strigoides (Latham) 246
Podicepididae, wnidentitied 6
Podiveps cristatus (Linne) 3
Poephila ucuticauda (Gould) 409
bichenoyil (Vigors & Horsfield) 407
cincla (Gould) 410
guttata (Vieillot) 406
personata Gould 408
Poliocephalus poliocephalus (Jardine & Selby) 4
Polvielis alexandrae Gould 207
anthopeplus (Lear) 206
swainsoni’ (Desmarest) 205
Pomatostomus rificeps (Hartlaub) 312
superciliosus (Vigors & Horsfield) 311
temporalis (Vigors & Horsfield) 310
Porphyrio porphyrio (Linne) 122
Porzana fluminea (Gould) 17
pusilla (Pallas)
tubuensis (Gmelin)
Procellaria cinerea Gmelin
Psephatus chryseplervetus Gould
dissimilis Collett
haematonotus (Gould)
varius Clark
Psophodes cristatus (Gould)
olivaceus (Latham)
Pterodroma brevirostris (Lesson)
lessani (Garnot)
macroptera (Smith)
Pteropodocys maxima (Ruppell)
Ptilonarhynchus violaceus (Vieillot)
Priloris paradiseus Swainson
Puffinus gavia (Forster)
griseus (Gmelin)
pacificus (Gmelin)
tenuirostris (Temminck)
Rallus pectoralis Temminek
Philippensis Linné
Recurvirostra noyaehollandiae Vieillot
Rhipidura fuliginosa (Spartman)
leucophrys (Latham)
rufiventris Quoy & Gaimard
Scythrops novuehollandiae Latham
Sericornis brunneus (Gould)
cau(us (Gould)
citreogularis Gould
Jrontalis (Vigors & Horstield)
fulizinosus Vigors & Horsfield
pyrrhopygius (Vigors & Horsfield)
Sericulus chrysocephalus (Lewin)
Smicrornis. brevirostris (Gould)
Sphecotheres viridis Vieillot
Sterna albifrons Pallas
bengalensis Lesson
bergii Lichtenstein
Juscata Linné
nereis (Gould)
sumatrana Rattles
Stictonetta naevosd (Gould)
Suiltia isabella Vieillot
Stipiturus ruficeps Campbell
Stomiopera unicelor (Gould)
Strepera fuliginosa (Gould)
graculina (White)
versicolor (Latham)
Struthidea cinerea Gould
Sula dactvlatra Lesson
leucoguster (Boddaert)
serrator (Gray)
sulu (Linneé)
Syma toratoro Lesson
Tachybaptus noyaehollandiae (Stephens)
Tadorna radjah Garnot
ladornoides (Jardine & Selby)
Threskiornis aethiopica (Latham)
spinicollis (Jameson)
Trichoglossus haematodus (Linne)
Tringa brevipes (Vieillot)
glareola (Linné)
Aypoleucos Linnie
y0 REC, S, AUST, MUS, 1915); 219.328
nebuluria (Gunnerus) 149
stagnalilis (Bechstein) 150
terek (Latham) 151
Turniix castanota (Gould) 10
pyrrhothorax (Gould) 112
velox (Gould) 1]]
Tyto alba (Scopoli) 244
novaehollandiae (Stephens) 245
banellus miles (Boddaert) 131
tricolor (Vieillot) 132
Nenorhynchus asiaticus (Latham) 63
Zoothera dayma (Latham) 276
Zosterops lateralis (Latham) 400
LIST $5. INDEX TO HELMINTHS
Jn this section the helminths are arranged in
alphabetical order within each of the taxa Trematoda,
Cestoda, Nematoda, and Acanthocephala.
Species names cited in List | as junior synonyms of
other species are included in this index in their
alphabetical sequence. The number(s) following each
helminth name is that of the host(s) in which it has
been found. As these are consecutive in the hosl-
parasite list (Lest 1) the bird name may easily be found.
Class Trematoda
Acanthoparyphium marilae Yamaguti, 1934: 138, 154,
155, 156
Acunthoparyphium spindlosum §.J. Johnston, 1917;
133, 138, 139, 163
Aeanthoparyphinm squatarolae Yamaguli, 1934: 154,
156
Acanthoparvphiumn sp. 143, 378
Allocreadiidae, unidentified: 91
ANopyge antigones S.J. Johnston, 1913: 124
Apatemon gracilis (Radolphi, 1819); 77
Apatemon intermedius (5.3. Johnston, 1904); 72, 86, 92
Apatemon vitelliresiduus Dubois & Angel, 1972: 87
4Aphuaryagostrigea simplex (S.J. Johnston, 1904): 53,
56, 57
Apharyngostrigea sp. 55
Ascocoiyle sp. 47
Atriophallophorus coxiellae §.J. Smith, L974: 4, 123,
135, 139
Austrobilharzia terrigalensis §.J. Johnston, 1917: 58,
163, 171, 209
Basantisia queenslandensis Deblock & Pearson, 1968a;
253
Bilharziellinae, unidentified: 142, 143
Bolbophorus confusus (Krause, 1914); 37
Bolbophorus sp., TAH, Johnston & Angel, 1942a; 37
Brachylaima pulchellum (S.J. Johnston, 1917): 187
Brachylaima sp. V87
Brachylecithum dacelonis Angel & Pearson, 1977: 253
Brachylecithum harrisoni (S.J. Johnston, 1917): 242
Wuust, 1986
Brachvlecithum latius Angel & Pearson, 1977: 432, 434,
438
Brachylecithum mesastomuim (3.4, Jolnston, 1917); 171
Brachylecithum parvum (S.J. Johnston, 1917): 268, 437,
439
Brachylecithum podaret Angel & Pearson, 1977; 246
Brachylecithun sp., Angel & Pearson, 1977; 434
Cardiocephaloides hillii (S.J. Johnston, 1904); 163
Cardiocephaloides museulosus (S.J. Johnston, 1904):
164, 166, 171
Cardiocephaloides ovicorpus Dubois & Angel, 1972; 45
Catatropis gallinulae TH, Johnston, 1928: 121
Catarropis sp. 116, WW7, 120
Cephalogonimus sp.: 4
Chaunocephalus ferox (Rudolphi, 1795): 63
Clinostomum australiense 8.3. Johnston, 1917: 42
Clinostamum complanatum (Rudolphi, 1814): 60, 62
OClinestamum hornum Nicoll, 1914b: 60, 62
Clinostomun sp. 37
Cloacitrema narrabeenense Howell & Bearup, 1967: 163
Cloeophora sp. 155
Cotylurini, unidentified: 77
Corylurus magniacetabulus Dubois & Angel, 1972: 72
Cryptocotyle sp. 33
Cyclocoelidae, unidentified: 72, 78, 82, 117, 144, 146,
(55, 378
Cyelocoeluin jaenschi TAH. Johnston & Simpson,
1940b: 4, 5
Cyclocoelum obscurum (Leidy, 1887): 131
Cyclocoelum taxarchis 5.J, Johnston, 1917: 154
Cyclocoelum sp. 81, 84, 122, 154
Cylindrotrema cygni Angel, 1973; 72
Dendritobitharzia sp. 37
Dicrocoeliidae, unidentilied, Koch & Duhamel, 1982;
196
Dicrocoeliidae, unidentified, Munday & Green, 1972:
439
Dicrocochidae, unidentified: 272
Dicrocoghinae, unidentified: 196, 435, 442
Dicrocoelium proxillicens Canavan, 1937; 196
Diplostomidae, unidentified; 46, 77, LOI
Diplostomum amygdalum Dubois & Pearson, 1965: 56,
57, 60
Diplostomum auriculosum Dubois & Pearson, 1967; 42
Diplostomum galaxiae Smith & Hickman, 1983a: 53
Diplostomum granulosum Goss, 1940: 46
Diplostomum murrayense (TA, Johnston & Cleland,
1938): 163, 164
Diplostomum parvulum Dubois & Angel, 1972: 37, 166
Diplostomum podicipinum Rozicka & Niewiadomska,
1960: 3
Diplostomum spathaceum (Rudolphi, 1819): 8, 163,
164
Diplostomumr triangulare (S.J. Johnston, 1904): 253
Diplostomuni sp. 37, 163
Distoma sp., Krefft, 1873: 52, $5, 122
Dolichosacculus solecarius (S.1. Johnston, 117): 47
Dolichosaccus solecarius S.\, Johnston, 1917: 47
Echinochasmus pelecani T.H. Johnston & Simpsou,
1944; 37
Echinochasmus prosthovitellatus Nicoll, 1914b: 104
Echinochasmus tenutcollis S.\, Johnston, 1917: 47
Evhinechasmus sp,; 46
TELMINTHS FROM AUSTRALIAN BIRDS 1T
Echineparyphium ellist (TAH. Johnston & Simpson,
1944) 72, &7
Echinoparyphium gizzardal Verma, 1936: 72
Echinoparyphium harveyanum 3.5. Johnston, 1917:
287
Echinoparyphium oxyurum S.J. Johnston, 1917: 55
Echinoparyphiuim phalacrocoracis Yamaguti, 1939a;
43, 44, 46, 47
Echinoparyphium sp., Verma, 1936: 72
Evhinoparyphium sp; 73, 75, 148, 163, 242, 246, 253
Echinostoma (Acanthochasmus) sp., T,H. Johnston,
1916, 1918: 145
Echinostoma aculicauda Nicoll, 1914b: 65, 66
Echinostomu australasianum Nicoll, 1914a; 124
Echinostoma australe T.H. Johnston, 1928: 121
Echimostoma bancrofii TH. Johnston, 1928: 121
Echinosioma elongalum Nicoll, 1914a: 246
Eehinostama emollitum Nicoll, 1914b: 239
Fchinostoma hilliferum Nicoll, Wl4b: 120, 122, 123
Fehinostoma ignavum Nicoll, 1914b: 128, 131
Echinosioma minimum Verma, 1936: 72
Echinostoma revolutum (Froelich, 1802); 69, 72, 75,
77, 80, 85, 87, 12)
Eehinostoma sp., TH. Johnston, t910: 133
Echinosiama sp., TH, Johnston, 19126; $5, 65, 77, 145
Echinestama spe 4, 55, 72, 78, 81, 116, 121, 122, 131,
}d8, 163, 166, 242
Echinostomatidae, unidentified, Bradley, 1926, 1927: 4
Echinostomatidae, unidentified, Munday & Green,
}972; 133
Echinostomatidac, unidentified: 4, 5, 37, 47, 56, 72,
78, 80, 82, 83, 87, 109, 120, 127, 134, 140, 148, 157,
164
Echinostematinae, unidentified: 46, 70, 72, 77, 78, 80,
86, 87, 92, 116, 117, 122, 157, 163, 166, 244, 425, 437,
439, 440
‘Echinostome larvae’, Bradley, 1926, 1927: 75, 123
Endocotvle incana Belopolskaya, 1952: 136, 157
Episthinium prosthovitellaria (Nicoll, 19146): 104
Eucotvle sp. 53
Eumegacetes sp.: 249, 304
Euparyphium longitestis Verma, 1936: 72
Fellodistomidae, unidentified: 45
Galaciosamum anvelae Pearson, 1973; 11, 38, 163, 166,
17]
Galactosoniunt bearupi Pearson, 1973: 163, 166, 168,
172, 174
Galactosomum renincolum Pearson, 1973; 31, 168, 174
Galactosontum sinuilacie Pearson, 1973: 43, 45
Galaciosomum ussuriense Oshmarin, 1963: 163, 166,
16s
Galactosamum sp.2 43, 166
Giguntobilharzia sp., Rohde, 1978: 163
Gyliauchen sp.r 62
Gynoecotvla brishanensis Deblock & Pearson, 1968a:
136, 163
Huemarotrephus adelphus §.3, Johnston, 1917: 142, 143
/luematatrephus consimilis Nicoll, 1914b: 131
Haemaioirephus spa 149
Haplorchis paravanissimus Pearson & Ow-Yany, 1982:
37, 46, 60, 91
Haplorchis pumilia (Looss, (896): 47, 37
Haplorchis sprenti Pearson, 1964: 37, 46, 47, 53, 55, 57
Haplorchis vanisstmus Alrica, 1938: 37, 43, 44, 46, 60,
9]
Haplorchis vanissimus of Pearson, 1964, not Alrica,
1938: 46, 60, 91
Hapltorehis vokogawai (Katsula, 1932): 91
FHuplorchis sp., Pearson, 1960; 37, 47
Harmostomum pulehellum $.J, Johnston, 1917: 187
Hemistomum hrachyurum Nicoll, 14a; 242
Hemistomum intermedium S.J. Solnston, 1904: 72
Hemistomuni rrianguldre 8.3. Johnston, 1904: 253
Heterophyidae, unidentified: 55, 154, 158, 166
Heterophyinae, unidentified: 171]
Heterotestophves sp: 163
Hexangiuin sp. 62
Himasthla harrisoni S.J. Johnston, 1917: 145
Himasthla kusasizi Yamaguti,, 1939a; 156
Himusthla megacotvla Yamaguti, 1939a; [54
Himasthlinae, unidentified: 155, 246, 378
Holostomum Aili) $1, Johnston, 1904: 163
Hotostomum musculosum St. Johnston, 1904 171
Holostomumn repens Chase, 1921: 53
Holostomun: simplex S.J. Johnston, 1904: 53
Hypoderaeum conoideum (Bloch, 1782): 79
Mlypoderaeum sp. 77, 78, 79
Hyptiasmus magnus §.). Johnston, 1917: 72
/ysteromorpha plataleae Dubinina & Dubinin, 1940;
65
Hysteromorphd triloba (Rudolphi, 1819); 43, 44, 46, 47
Lateretrema sp. 311
Lecithodendriidac, unidentified: 304, 350, 425
Lepoderma nisbelii Nicoll, |914a: 416
Lepodermatidae, unidentified: Nicoll, 1914b; 13]
Leucochloridium australiense TJA- Johnston &
Cleland, 1938: 311, 423
Leucachloridium spa i4
Levinseniella howensis S.J, Johnston, 1917: 133
Levinseniella. microovata Belopolskaya,, 1958: 133, 136
Levinseniella monodactyla Deblock & Pearson, 1970:
136
Levinseniella tusmaniae (S.J. Smith, 1974): 4, 77, 79,
135, 139, 140
Leyinsentella sp. 163
Lutztrema ailuroedi Angel & Pearson, 1977: 421
Lyperosomum harrisoni S.J, Johnston, (917; 242
Lyperosomum megastomum §.J. Jotinston, 1917 171
Lyperosomum parvum SA. Johnston, 19172 437
Maritremu calvertense S.J. Smith, 1974: 4, 77, 79, 135,
149
Maritrema eroliae Yamaguti, 1939a; 136, 171
Maritrema ocystum (Lebour, 1907): 5, 42, 53
Mariirema sp.; 138, 148, 154, 171
Mawsonatrema eudyptulae Angel, 1973: U
Mesostephanus haliasturis Tubangui & Masilurigan,
194]: 37, 42, 91
Mesostephanus minar Dubois & Pearson, 1965: 37, 42,
1
Micraphalhidae, unidentified, Munday & Green, J972:
139
Microphallidae, unidentified: 5, 81
Microphallidae, unidentified: 157
Microphallus pupillornatus Deblock & Pearson, 1969:
133, 136, 163
312 REC. S. AUST. MUS. 19(15): 219-325
Microphallus tasmaniae 5.3, Smith, 1974: 4, 77, 79, 135,
139, 140
Microphallus vaginosus Deblock & Pearson, 1969: 133
Miecrophallus sp., Deblock & Pearsan, 1969: 136
‘Monostome’, Nicoll, 1914c: 124
Monostomum sp,, T.-H, Johnston, 1910; 72,142
Neodiplostomulum sp., Dubois & Angel, 1972; 425
Neodiplostomum australiense Dubois, 1937a: 8&9
Neodiplestomum brachyurum (Nicoll, 1914a): 242, 244
Neodiplostomum lanceolatum Dubois & Angel, 1972:
242
Neodiplastomum pricei Krull, 1934: 163
Neodiplostramum spathula (Creplin, 1829): 89, 91, 93,
95, 96, 99, 100, 101, 244
Neodiplostomum subaequipartitum Dubois & Pearson,
1967; 91
Neodiplostomum sp, 101
Nephrostomum sp; 56
Notocotylid, sp. B, S.J. Smith, 1981: 72
Noatocotylidae, unidentified: 70, 72, 73, 77, 78. 81, 86,
87, 134, 143, 144, 157
Natocotylus attenuaius (Rudolphi, 1809): 69, 72, 77,
85, 128, 131, 1S7
Notocorylus spa 72
Omphalometridae, unidentified: 143
Opisthorchis absequens Nicoll, 1914b: 66, 104
Opisthorchis sp. 99
Orchipedum sufflavuim Nicoll, 1914h: 67
Ornithobitharzia sp.: 37, 164
Pachytrema calculus Looss, 1907: 163, 164
Pachytrema sp., S.J. Smith (pers. comm.), 1979: 4
Pancreatrema meliphagae Angel & Pearson, 1977: 371
Paramonostamum bursae Smith & Hickman, 1983b; 77
Paramonostomum caeci Smith & Hickman, 1983hb: 4,
72, 77
Paramoanostamum spo Sl, 121
Parastrigea repens (Chase, 1921): 53, 99
Parastrigea sp, Dubois & Angel, 1972: 65
Parorchis acanthus (Nicoll, 1906): 163
Paryphostomum phalacrocoracis Goss, 1940; 46, 47
Paryphostamum radialum (Dujardin, 1845): 43, 44, 46,
47
Paryphostomum tenuicolle (S.J. Johnston, 1917): 43,
44, 46, 47
Patagifer acuminatus 3S, Johnston, 1917: 65
Patagifer bilobus (Rudolphi, 1819): 64, 65, 66, 67
Patagifer bilabus of T,H, Johnston, 1916, not
(Rudolphi, 1819); 65
Patagifer Jraternus S.J, Johnston, 1917; 55
Patagvfer sp.: 4, 65, 68, 77
Petusiger australis T.H. Johnston & Angel, 194th: 3,
4,5
Perasiger exaeretus Dietz, 1909; 43, 44, 44, 47
Petasiger sp. 244
Philophthalmus burrili Howell & Bearup, 1967: 143
Philophihalmus sp; 1, 126
Plagiorchiidae, unidentified: 131
Plagiorchis clelandi 8.3, Johnston, 1917; 269
Plagiorchis macu/osus (Rudolphi, 1802); 266, 269, 270,
302, 304, 311, 399, 425, 434
Plagiorchis nishetij (Nicoll, 1914a): 416
Plagiorchis spatulaius §.J. Johnston, 1917. 270
Platynosomum proxillicens (Canavan, 1937); 196
August, 1986
Platynosomum fallax Heidegger & Mendheim, 1938a:
196
Platynosomum ventroplicatum
Mendheim, 1938h; 196
Platynotrema biliosum Nicoll, '914b: 65, 128
Platynotrema jecoris Nicoll, 19146; 128
Platynotrema spi 148, 154
Pleuropsoalus sp.: 234
Posthodiplostomum australe Dubois, 1937a: 37, 46, 47,
48, 53, 55, 60, 62, 166
Posthodiplostamum eblongum Dubois, 1937a: 62
Proacetabulorchis dogieli Belopolskaya &
Bychovskaya-Paylovskaya, 1954; 53
Pracerovum veriuin Onji & Nishio, 1916; 37, 53, 55,
57, 60, 91
Pracerovien sp., Pearson, 1960: 37, 47
Prosthogonimidae, unidentified! 438
Prosthagonimus vitellatus Nicoll, 1914az 47, 164, 416,
434
Prosthogonimus sp., Nicoll, 1914b: 425
Prosthoganimus sp.: 77, 124, 163, 304, 336
Psilochasmus oxyurus (Creplin, 1825}; 4, 72, 77, 123
Psilachasmus sp.: 72, 73, 74, 87, 104
Psilastomuim sp. A, S.J, Smith, 1981: 4, 72
Psilostomum sp. B, S.J. Smith, 198i: 4, 72
Psilostomum sp.: 72, 73, 77
Renicola sp.: 11, 33, 34, 37, 43, 45, 48, 55
Scaphanocephalus australis SJ. Johnston, 1917: 95
Scaphanocephalus sp., $.J. Johnston, 1914: 95
Schistosomatidae, unidentified, Blair & Otteson, 1979:
69, 70, 71, 77, 78, 80, 82, 83
Schistosomatidae, unidentified: 3, 163
Schistosomatinae, unidentified: 37
Schwarizitrema novaehollandiae Dubois & Pearson,
1947: 42
Schwartzitrema pandubi (Pande, 1939); 3, 4, 5, 37, 44,
46, 47, 53. 62, 68
Skrjabinosomum mawsoni Angel & Pearson, 1977: 110,
362
Skrjabinosomun pomatostomi Angel & Pearson, 1977:
311
Skrjabinosamum sp, Angel & Pearson, 1977: 287
Stellanivhasinus aspinosus Pearson, 1964: 47
Stellantchasmus falcatus Onji & Nishio, 1916: 60, 91
Stellanichasmus falcatus of Pearson, (960, not Onji &
Nishio, 1916; 47
Slephanoprora sp. 63
Steringophoridae, unidentified, Goss, 1940: 45
Stictedora caballeroi Martin, 1955: 37
Sticlodora diplacaatha TH. Johnstan, 1942a: 43, 45,
46, 48, 163
Stictedora lari Yamaguti, 1939a: 163
Stictodora manilensis Africa & Garcia, 1935: 169
Stictodora sp,, Bearup, 1958: 163
Srictedora sp., Pearson, 1960: 169
Strigea baylisi Dubois, 1937a: 65, 66, 68
Strigea falconis Dubois, 1937b: 104
Sreigea floseulus Nicoll, 19)4a: 246
Strizgea glandulosa Dubois, 1937a: 91, 96, 99, 100, L04
Strigea nicalli (Dubois, 1937a): 128, 425, 434
Sirigea proniscua Nicoll, 1914a: 241, 242
Sirigea suttoni Dubois, 1937a; 425
Strigea sp.: 104
Heidegyer &
HELMINTHS FROM AUSTRALIAN BIRDS si
Strigeidae, unidentilied, Munday & Green, 1972: 93
Strigeidae, unidentified: 45, 101
Tanaisiinae, unidentified: 440
Trichobilharzia australis Blair & Islam, 1983: 77
Trichobilharsta sp., Bearup, 1957: 78
Trichobilharzia sp., Bearup & Langsford, 1966; 142
Trichobilharzia sp. Blair & Otteson, 1979: 77
Trichobilharzia sp., T.H. Johnston, 1941; 72
Trichobilharzia sp. 142, 209
Tiphlocoelum reticulare §.J. Johnston, 1913; 69
Tiphlocoelum sp 77, 78
Uretocus sp.: 360
Fermatrema lonsitestis (Verma, 1936): 72
Trematodes, unidentified, Ratclrffe, 1933; 196
Trematodes, unidentified; 46, 53, 62, 65, 72, 78, 81, 87,
115, 121, 134, 142, 156, 157, 158, 245, 294, 319, 341,
353, 399, 422, 434.
Class Cestoda
Acoleus hedlevi T.H. Johnston, 1910; 142
Angularella australis (Maplestone, 1921b): 69, 128
Anegularella sp.: 13)
Angularia australis Maplestone, 1921b: 128
Anomotaenia accipiiris TH. Johnston, 1913; 93
Anomotaenia asvnmetrica TAA. Johnston, 1913: 55
Anomotaenia hydrochelidonis Dubinina, 1954: 163
Anonchotaenia arhyncha Fuhrmannn, 1918: 399
Anonchotaenia globata (Litstow, 1879); 359
Aploparaksis australis TAH, Johnston, 1913: 152
Armadoskrjabinia globosa (Szpotanska, 1931): 72
Ascometra sp. 126
Australiolepis southwelll (Szpotanska, 1931): 72
Baerbonia parvitueniunca (Baer & Bona, 1960); 58
Baneroftiella ardeae TH, Johnston, 1913: 53, 60
Bancroftiella glandularis (Fuhrmann, 1905); 55
Bancrofriella glandularis of TH. Johnston, 1912b, not
(Fuhrmann, 1905); 53
Bancroftiella sp.: 62
Biuterinu sp.: 69, 415
Cupiuterilepis australivnsis Schmidt, 1972: 350, 353
Capiuterilepis melinhagieola Schmidt, 1972: 337
Choanaraenia fieldingi (Maplestone & Southwell,
1923); 432
Choanotaenia meliphagidarum T.H. Johnston, 1911:
363, 367, 372, 380, 381
Choanotuenia southwelly Pulyrmann, 1932) 131
Choanataenia favlori TH. Johnston, 1912¢: 318
Choanotaenia zoniferae VH. Johnston, 19120: 132
Chownotaenia sp. TH. Johnston, 1910: 363, 367, 372,
380, 381
Choanotaenia sp. 128, 352, 404, 407
Cladataenia feura Meggitt, 1933: 98
Cladotauenia sp.. 98, 99, LOO
Clelandia parva T.H. Johnston, 1909b; 63
Cloucotaenia negalops (Nitzsch in Creplin, 1829); 69,
70, 72, 77, 78, 79
Cloacolaeniu sp. 82
Cotuenia brotogerys. Meggitt, 1915; 213
Cotmenia collini Fuhrmann, 1909: L
Cotugnia vligerclhis Maplestone, 1922a: 70
Cotugnia platycerci Weerekoon, 1944: 216
Cetugnia polvielidis Burt, 1940; 206
Tolugnia Sent Meggitt, 1926: 213
Cotuentia sp.: |
Cyclophyllidea, unidentified, Munday & Green, 1972;
13
Cyclophyllidea, unidentified: 72, 87
Cyelorchida omalancrisirota (Wedl, 1856): 67
Duvainea australis (Krabbe, 1869): 1
Davainea cucatuina T.H, Jofnston, 1913: 196
Davainee conopophilae TH. Johnston, 1913: 358, 359,
384
Davainea himantopodis T.H. Johnston, 1911y 142
Davainea leptosoma Diesing, 1850: 191
Davainea sphecotheridis TH. Johnston, 19l4a; 415
Davainea sp., TAH. Johnston, 1910: 142
Davainea sp., T.H. Johnston, 1912b: 187, 438
Davainea spi 107, 142, 143) 187, 423
Dendrouterina austriliensis Baer & Bana, 1960: 33
Dendrauterina sp. 415
Dicranotaenia coronula (Dujardin, 1845); 72
Dilepididae, unidentified: 403, 407, 410, 434
Dilepis australiensis V.H, Johoston, 912d: 142
Dilepis hancrofti TH. Johnston, 1912c: 213
Dilepis maxima Goss, 1940; 45
Dilepis minima Goss, 1940: 46, 47
Dilepis sp.. 46, 350
Dinecocesins novaehollandiae (Krefft, 1873): 3
Dioecoceslus sp. 4
Diorchis flavescens (Kreilt, 1873): 70, 77, 78, 79, 80,
82, 142, 143
Diorchis spiralis Szpotanska, 1931; 72
Diorchis stefanski Czaplinski, 1955: 72
Diorchis sp, 73, 79
Diplogynea oligorchiy (Mapiestone, 1922a); 70
Diplozvnea sp. &2
Diploposthe laevis (Bloch, 1782); 77, 79, 82
Diploposthe laevis of TH. Johnston, 1913, not (Bloch,
1782): 70
Drepanidataenia bisacculina Szpotanska, 1931: 72
Drepanidotaenia lanceolata (Bloch, 1782): 72, 77
Drepanidotaenia rapida (Szpotanska, 1931); 72
Drepanidotuenia sp.c 72
Echinorhynchotaenia nana Mapleston & Southwell,
1922b: 72
Fimbriaria fasciolaris (Pallas, 1871): 77, 79
Fimbriaria intermedia Fuhrmann, 1913; 74
Fimbriaria sp; 17, 82
Fimbriarioides intermedia (Fuhrman, 1913): 74
Gustrotuehia sp. 72, 77, 78, 82, 87
Gyrocoelia australiensis (T.H, Johnston, 1910): 131, 142
Gyroceelia sp., TH, Johnston J914az 131
Gyrocoelia sp.: 4, 134, 143
Haploparaxis australis (TH. Johnston, 1913): 148, 152,
153
Hapleparaxis veitch{ Baylis, 1934b: 78
Hemiparonia hancrofii (TH. Johnston, 1912¢); 212,
213, 217, 218
Hemiparonia cacafuae (Maplestone, 1922b): 195, 198
Hemiparonia merotomochaeta Woodland, 1930: 195
Hemiparonia sp. 191
Himantocestus sp; 144
Hymenolepididae, unidentified: 65, 358, 359, 423, 445
Aymenolepis chenapis, Palmer, 1981: 72
Aymenolepis colluris (Batsch, 17&6): 77, 79
ald REC, 8, AUST MUS. J9U15); 2/9-325
Hymenolepis cormoranti Ortlepp, 1938: 47
Hyntenolepis curiosa Szpotanska, 1931; 72
Hymenalepis ellisi TH. Johnston & Clark, 194%a: 37
‘Aymenolepis eurysiomi Maplestane’: 261
Hymenolepis globosa Szpotanska, 193]: 72
Hymenolepis ibidis T.H, Johnston, 1913: 66, 68, 70
Hymenolepis jaensehi TH. Johnston & Clark, 1948a:
37
Hymenolepis lamellata Woodland, 1930; 75
Hymenalepis lanceolata (Bloch, 1782): 72, 77
Afymenolepis liophallas Krabbe, 1869: 72
Hymenolepis meealaps (Nitzsch in Creplin, 1829): 69,
77, 79
Ayvmenalepis micrancristrota (Wedl, 1855): 72
Aymenolepis murrayensis TH. Johnston & Clark,
1948a; 37
Flymenolenis phalacracoracis Woodland, 1929: 47
Aymenolepis rapida Szpotanska, 1931; 72
Fivmenolepis robertsi Baylis, 1934b; 78
Hymenolepis southwelli Szpotanska, 1931; 72
Hymenolepis terraereginae TH. Jotinston, 1913: 69
‘Hymenolepis variabilis n, sp. Maplestone’: 72
Hymenolepis sp., T.H, Johnston, 1910: 142, 281
Hymenolepis sp., T.H, Johnston, 1912b: 60, 77
Hymenolepis sp., Southwell, 1916: 72
Hymenolepis sp.: 37, 60, 66, 68, 72, 75, 77, 123, 144,
353, 366, 399, 419, 425, 440, 443
Idiogenes spo 126
Infula burhini Burt, 1939: 131, 142
Lapwingia adelaidae Schmidt, 1972: 131
Metroliasrhes sp. 109
Microsomacanthus collaris (Batsch, 1786); 77, 79
Microsamacanthus cormoranti (Ortlepp, 1938): 47
Moniezia trichoglossi (Linstow, 1888): 199
Monopylidium fieldingi Maplestone & Southwell, 1923:
432
Monopylidium macracanthum, ol Maplestone &
Southwell, 1923, not Fuhrmann, 1907; 131
Monosaccanthes curiosa (Szpotanska, 1931): 72
Monosaccanthes kazachstanica (Maksinava, 1963); 72
Monasaccanthes sp. 72
Nematoparataenia parudoxa Maplestone & Southwell,
1922b: 72
Ophiataenia hylae TH. Johnston, 1912¢: 70
Parabisaccanthes bisacculina (Szpotanska, 1931); 72
Paradilenis minima (Goss, 1940): 46, 47
Poaradilepis patriciae Baer & Bona, 1960: 68
Paradilepis scolecina (Rudolphi, 1819): 43, 44
Paradilepis urceina Bona, 1975, 53, 65
Paradilepis sp., Clark, 1957: 47
Paradilepis sp.. 43
Paricterotaenia chlamyderae (Krefft, 1873); 419
Paricterotaenia zoniferae (TH. Johnston, 1912c): 132
Paricterataenia sp... 128
Paronia trichoglossi (Linstow, 1888): 199
Peruterina rauschi Freeman, 1957: 150, 244
Paruterinae, unidentified: 310, 311
Parvitaenia ardeae (T.H. Johnston, 1913): 53, 60
Parvitaenia clavipera Baer & Bona, 1960: 53
Parvitaenia glandularis (Fuhrmann, 1905); 5§
Paryitgenta paracyclorchida Baer & Bona, 1960; 53
Passerilepis stvlosa (Rudolphi, 1810): 69
Passerilepis zostéropis (Fuhymann, 1918): 70
August, JY8A
Prosthecocotvle diomediae Fulrmanon, tn Shipley,
1900: 12
Pseudochoaneluenia meliphagidarum (T.H. Johnston,
1911}: 343, 367, 372, 380, 381, 390
Pseudophyllidea (plerocercoids): 253
Raillietina australis (Krabbe, 1869): L
Raillietina cacatuing (TH. Johnston, 1913): 196
Raillietina conopophilae (TH. Johnston, 1913): 358,
359, 384
Raillielina corvina (Fuhrmann, 1905); 442
Raillietina leipoae T.H, Johnston & Clark, 1948; 106
Raillietina leptesoma (Dicsing, 1850): 191, 213
Raillierina peuciiesticulata (Fuhrmann, 1908): 196
Raillielina pelychalix (Kotlan, 1921): 217, 229
Raillietina sphecotheridis TH, Johnston, Wl4a; 415
Raillietina sp., T.H. Johnston & Clark, 1948b; 106
Roillietina sp., Hall Exped. Report, 1974: 415
Raillietina sp.; 1, 107, 231
Schizotaenia cacatuae Maplestone, 1922b: 196
Similuncinus dacelonis VW. Johnston, 1909a: 253
‘Shipleva lobivanellus Maplestone’: 131
Sobolevicanthus terraereginae (TH. Johnston, 1913);
69
Sobolevicunthus sp.: 74, 82, 87
Sphaeruterina punctata TH, Johnston, 1914b: 243
Staphylepis lamellata (Woodland, 1930): 75, 78
Taenia australis Krabbe, 1869: |
Tuenia bairdti Krelli,, 1873; 77
Tuenia chlamyderae Kretft, 1873: 419
Tuenia coronata Kreffi, 1873; t42
Tuenia evlindrica Kretlt, 1873: 77
Taenia flavescens Krefft, 1873; 77, 80
Tuenia ricrancristrova Wedl, |8S5: 72
Tuenia moschata Kreffi, 1873; 87
Taenia novaehoNandiae Krettt, 1873; 5
Taenia puradoxa Krefft, (8732 5
Taénia pediformis Kreltt, 1873: 77, 79
Taenia rugosu Kretti, (873: 142
Taenia trichoglossi Linstow, 1858; 199
Taenta tuberculata Krettt, 1873; 82
Tetrabothriidae, utadentified: 7, Ul, 33, 38
Tetrabothrius diomediae (Fuhrmann, in Shipley, 1900);
12
Tetrabathrius lutzi Parona, 901s 1
Tetrabothrius. polyorchis Nybelin, 1917: 50
Yerrabothrius sp,, TH, Johnston, 1912b: 12, 14, 49
Tetrabothrivs sp.z \1, 13, 16, 18, 19, 24, 25, 32, 163, 171
Trichocephaloides sp. 157
Trypanorhyncha, unidentified: 45
Tschertkovilepis Krabbei (Kowalewski, 1895): 72
Falipore parvituerniunca Bacr & Bona, 1960: 58
HWoadlandia phalacracoracis (Woodland, 1929); 47
Zosrerapicola elelandi TH, Johnston, (912e: 400
Unidentified cestodes have been reported by:
Bradley, 1926: 4
Bradley, 1927: 4, 123, 131
Cleland, 1922; 4. 11, 31, 33, 72, 77, 104, 152, 157,
171, 179, 181, 196, 199, 218, 242, 255, 259, 263,
265, 267, 268, 269, 270, 284, 289, 292, 293,294,
300, 318, 330, 354, 359, 462, 348, 371, 373, 378,
381, 382, 399, 400, 417, 418, 421, 423, 426, 427,
429, 430, 435, 438, 439, 442
HWELMINTHS FROM AUSTRALIAN BIRDS WS
Macgillivray, 1917; |
Munday & Cireen, 1972: 33, 77, 86, 93, 104, 131, 139,
318, 400, 439
Unidentified cestodes are held in collections from: 1,
2, 3, 4, 5, 6, 10, 11, 12, 13, 15, 16, 17, 19, 20,
2], 23, 25, 29, 30, 32, 33, 34, 37, 43, 44, 45, 46,
47, 49, 53, 60, 62, 65, 66, 68, 69, 70, 72, 73,
75, 77, 78, 79, 80, 81, 82, 85, 86, 87, 90, 91, 92,
99, 101, 103, 104, 105, 107, 108, 120, 121, 122,
123, 126, 128, 129, 130, 131, 132, 133, 134, 136,
137, 138, 139, 140, 141, 142, 143, 144, 145, 146,
147, 148, 149, 152, 153, 154, 157, 158, 159, 160,
162, 163, 164, 166, 171, 175, 176, 180, 181, 182,
185, 186, I87_ 188, 196, 197, 212. 218, 219, 234,
244, 246, 247, 248, 249, 251, 253, 257, 261, 266,
270, 271, 274, 276, 277, 287, 289, 294, 302, 304,
305, 310, 311, 313, 319, 320, 330, 334, 346, 350,
352, 354, 359, 362, 373, 374, 378, 380, 381, 398,
399, 400, 406, 410, 412, 418, 422, 423, 425, 428,
434, 435, 437, 438, 439, 440, 441, 442.
Phylum Nematoda
Acvaria anthuris (Rudolphi, 1819): 438, 439, 440, 441,
442
Acuaria calluricinclae Mawson, 1972: 293, 294
Aevaria corvicola TH, Johnston & Mawson, [941a: 442
Acuaria fieldingi Baylis, 1934a; 94
Acuaria flindersi T.H. Johnston & Mawson, 1941b: 104
Aeuaria microeca Mawson, 1972; 287, 322
Acuaria wrafrae Mawson, 1972: 265
Acuaria petierae Mawson, 1972: 275, 278, 296, 301,
309, 353, 354, 366, 372, 429, 433, 434
Aeuaria skrfabini Ozerskaya, 1926: 412
Acuaria streperina T.H. Johnston & Mawson, 194]d:
435, 436, 437
Acuaria sp. T.H. Johnston & Mawson, 1941le: 164
Acuaria sp.: 248
Acuariidae, unidentified. T.H- Johnston & Mawsan,
194le: 166
Acuariidae, unidentified; 80, 81, [11, 117, 134, 174, 393,
402
Alainchahaudia aegotheles (TH. Johnston & Mawson,
194]e): 248
Alainchabaudia alcedinis Mawson, 1968: 253, 257
Allodapa differens (Sonsino, 1890); 239
Allodapa suctoria (Molin, 1860): 238, 242, 246, 249,
250
Allodapa sp., TH, Johnston & Mawson, 1941b: 241
Allodapa sp. 246
Amidostomum acutum (Lundahl, 1848): 73, 76, 77, 78,
RO, 142
Amidostomum anseris (Zeder, 1800); 69, 74
Apnidastomum biziurae T.H. Johnston & Mawson,
1947: 87
Amidostomum chevreuxt Seural, 1918: 142
Amidoastomum eygni Wehr, 1933: 72
“Amidostanum fulicae (Rudalphi, 1819): 123
Amidostamum tribonyy Mawson, 1980; 120
Amidostomum sp. 72, 77, 78, 83
Ancrracanthopsts sp.: 257
Anisakidae, Munday & Green, 1972: 13
Anisakidae, unidentified: 8, 13, 149, 174
Anisukis diomedeue (Linstow, (888). 12, 13, 18, 16, 17,
19, 2L, 27
Anisakis §p., T.H. Johnston & Mawson,
Anisakis sp., T.H. Johnston & Mawson, 1942¢: 27
Anisakis sp., T-H. Johnston & Mawson, 1942d: 23
Anisakis sp., TH. Johnston & Mawson, 1951; 174
Anisekis sp.) 22, 28, 171, 174
Aprocta australis (T.H. Johnston & Mawson, 1942a):
239
Aprocta bakeri Baiti & Mawson, 1981; 438, 440, 442
Aprocvta boulengeri Bain & Mawson, 1981; 435
Aprocta brevicaudata Chaw, 1939: 242
Aprocta carvicela T.A. Johnston & Mawson, 19404;
442
Aprocta retundata (Linstow, 1903): 239
Aprocta vestibulata (T.H. Johnston & Mawson, 1940a):
344
Aprocta sp.) (93
Aproctidae, Munday & Green, 1972; 280, 281
Ascaridia catheturina (T.H. Johnston, L912a): 107
Ascaridia columbae (Gmelin, 1790); 180, 197, 203, 207,
209, 210, 218, 219, 226
Aseartdia galli (Schrank, 1788); 236, 239
Ascaridia lineata (Schneider, 1866): 239
Ascaridia platycert Harlwich & Tscherner, 1979: 190,
203, 204, 205, 206, 207, 208, 209, 210, 212, 213, 217,
218, 219, 220, 221, 222, 223, 224, 226, 228, 230, 231
Ascaridia sprentt Mines, 1979: 204, 206, 207, 218, 224,
226, 228, 230, 23)
Ascaridia stroma (Linstaw, 1899): 124, 125
Ascaridia sp. 178, 191, 206, 208, 217, 239
Ascaris serpentula Rudolphi, 1809: 124
Ascaris §piculigera Rudolphi, 1809: 37, 46
Ascaris spiculigera of T.H. Johnston, 1912b, not
Rudolphi, 109; 42.
Ascaris sp., Krefft, 1873: 42, 350
Ascaris sp., T.H. Johnston, 1912b; 44
Austrofilaria rhipidurae T.H. Johnston & Mawson,
1952; 304
Austrofilaria vestibulata T.H. Johnston & Mawsen,
1940a: 344
Bancroftinerna dentaium T.H. Johnston & Mawson,
194ib: 104
Baylisascaris sp. 208
Capillaria anatis (Schrank, 1790); 72
Cupillaria convolujor Fourment, 1885: 13, 19
Capillaria ellisi TH. Johnston & Mawson. 1945b: 72
Capillaria grallina TA. Johnston & Mawson, 1945a:
425
Capillaria graucalina TA. Johnston & Mawson, 1941d;
271
Capillaria gymnorhinae T.A. Johnston & Mawson,
1947: 434
Capillaria jaénsehi T.H. Johnston & Mawson, 1945b:
37, 43, 44, 45, 46, 47, 163, 164
Capillaria lartcola Wassilikova, 1930; 163
Capillaria obsignata Madsen, 1945: 72
Cupillaria pomatostomi T.H. Johnston & Mawson,
1945b: 311
Capillaria recurvirostrae Mawson, 1968b: 144
Capillaria thomascameroni Mawson, 1969; 163
Cupillaria triloba Linstow, 1875: 131, 142, 144
1942a: 1]
316 REC. S, AUST. MUS. 19115): 219-425
Capillaria sp., TV.H. Johnston & Mawson, 1945a: 43
Capillaria sp,, TH, Johnston & Mawson, 1949; 3
Cupillaria sp., Munday & Green, 1972; 115
Capillaria sp.; 4, 6, 62, 65, 66, 77, 78, 79, BO, 81, 82,
86, 9L, 95, 99, 104, 107, 112, ILS, 116, 117, 119, 121,
122, 137, 139, 171, 209, 231, 234, 242, 245, 246, 261,
294, 309, 312, 322, 354, 361, 362, 366, 378, 432, 439,
440, 442,
Cardiofilaria dubia (TM. Jotinston & Mawson, 1940a):
198
Curinema dubia T.A, Johnston & Mawson, 19404; 198
Carinema gravcalinum TH. Johnston & Mawson,
1940a; 271
Cerafospira anthochaerae TH. Johnston, 1912a: 350
Cheilonematodum halevanis T.H. Johnstan &
Mawson, 194]d: 257
Cheilospirura gruveli (Gendre, 1913); TO9
Cheilospirura eymnorhinis de Chaneet & Robertson,
[9832 434
Chevreuxia australis T,H, Johnston & Mawson, (94) b:
140, 164
Chevreuxia revoluta (Rudolphi, 1819); 142, I44
Chevreuxia sp. 47
Contravaecum bancrofti TH. Johnston & Mawson,
1941e: 37
Contracaecum ceylunicum (Linstow, 1904): 91
Contracuecum clelandi T.H, Johnston & Mawson,
194le: 37
Contracaecum diomedeae (Linstaw, 1888): 12
Contracaecum eudyptulae V.A. Johnston & Mawson,
19424: Ll
Contracaecum magnicollare T.H. Johnston & Mawson,
194le: 17, 32, 33, 38, 173, 174
Contracaecum microcephalum (Rudolphi, 1809); 60,
62, 77
Contracaecum micropapillatum (Stossich, 1890); 37
Contracaecum nycticoracis T.H, Johnston & Mawson,
L94ie: 60
Contracaecum pelagicum TH. Johnston & Mawson,
1942d: 13, 16
Contracaecum podicipiris TA. Johnston & Mawson,
1949; 3, 5
Cantracaecum praestriatum Monnig, 1923: 4
Contracaecum rodhaini (Gedoelst, 1916); 42
Contracaecum sinulabjatym T.H. Johnston & Mawson,
194le: 42, 44, 47
Contracaecum spiculigerum (Rudolphi, 1809); 11, 37,
43, 44, 45, 46, 47, 52, 53, 60, 62, 64
Contracaecum tricuspe (Gedoelst, 1916): 42
Contracaecum sp., T.H. Johnston & Mawson, 1941c;
53, 55, 63
Contracaecum sp., T.H. Johnston & Mawson, 1947: 164
Contracaecum sp., Mawson, L968b: 142
Contracuecum sp. 9, 12, 18, 38, 49, 51, 53, 55, 59, 41,
62, 68, 101, 258
Cosmocephalus. jaenschi T,H, Johnston & Mawson,
194le: 37, 43, 44, 164, 171
Cyrnea calini Cram, 1927: 109
Cyrnea dentifera T,H. Johnston & Mawson, 194ld: 126
Cyrnea dallfusi Mawson, 1968d; 242
Cyrnea falea Mawson, 168d: 102, 104
Cyrnea mansion? (Seurat, 1914): 91, 94, 104
August, 1986
Cyrnea paraleptoplera TH. Johnston & Mawson,
1941b; 108
Cyrnea spiralis Mawson, 1968d; 274, 291, 356, 359, dl4
Desmidocercellu sp. 46, 47, 52, 60, 61
Diomedenemu diomedeae T.H, Johnston & Mawson,
1952; 1S, 17
Diplotriaena alpha T.H. Johnston & Mawson, 1940a:
433, 435
Diplotriaena beta TW. Johnston & Mawson, 940a:
442
Diplotriaena beveridgei Bain & Mawson, 1981: 442
Diplotriaena clelandi (1H. Jahnston, 1912b): 434
Diplotriaena delta TA. Johnston & Mawson, 1940a:
320, 324
Diplotriaena epsilon 1H. Johnston & Mawson, 1940a:
432
Diplotriaena faleonis (Connal, 1912): 104
Diplotriaena flabellata (Linstow, 1888): 442
Diplotriaena golvani Anderson, 1959: 294
Diplotriaena halli Ogden, 1967: 297, 320
Diplotriaena pungens (Schneider 1866): 415
Diplotriaena smitht Bain & Mawson, 1981: 354, 357
Diplotriaena spratti Bain & Mawson, 1981: 296
Diplotriaena tricuspis (Fedtschenko, 1874): 272, 315
Diplotriaena tridens (Molin, 1858): 415
Diplotrigena zeta T.H. Johnston & Mawson, 1940a; 354
Diplatriaena sp., Qgden, 1967; 298, 400
Diplotriaena sp., Mackerras, 1962: 357, 416
Diplotriaena sp., Munday & Green, 1972: 351
Diplotriaena sp.: 272, 287, 295, 318, 320, 399, 400, 416,
436, 442
Dispharynx pelecani T.H. Johnston & Mawson, 1942c;
37
Dispharynx sp, TH. Johnston & Mawson, 194fe: 42
Dromaeostrongylus bicuspis Lubimoy, 1933; 1
Echinuria heterabrachiata Wehr, 1937: 139, 143, 144,
158
Echinuria querquedulae TA. Johnston & Mawson,
1942¢: 78
Echinuria squamata (Linstow, 1883): 44
Echinuria uncinata (Rudolphi, 1819}: 72, 75, 78, 79, 85
Eehinuria sp. 77, 78
Epomidiostontm uncinatum (Landahl, 1848), 78
Epamidiostomum sp.: 69, 72, 75, 77, 78, 79, 83, 86
Eu/filaria sp., Mawson, 1969° 163
Eustrongyvlides phalacrecoracis V.A. Johnston &
Mawson, 194le: 43. 44, 47
Eustrangylides plotinus TH. Johnston & Mawson,
194le: 42
Eustrong ylides sp,: 37, 46, 60
Excvisa biloba Mawson, 1968d: 242, 246, 247
Excisa dentifera (TH, Johnston & Mawson, 1941d): 126
Exvisa excisiformis (Yamaguti, 1935): 246
Filaria clelandi TMH, Johnston, 1912b: 434
Filaria dacelonis Breinl, 1913: 254
Filaria guttata Schneider. 1866: 104
Vilarial worms, (‘Filaria' or ‘Filarioidea’, etc.) recorded
hy:
Bancroft, 1889: 352, 361, 432, 442
Cleland, 1922: 102, 198, 279, 318, 435
TH, Johnston, 1912a; 358
T.H. Johnston, 1912b; 242, 257, 272, 354
T.H. Johnston & Mawson, 1940a: 208, 134
HELMINTHS FROM AUSTRALIAN BIRDS AT
T.H. Johnston & Mawson, l94la: 322
Mackerras, (962: 52, 200, 271, 376, 415, 425, 427,
430, 432, 434, 441
Filarioidea, unidentified: 11, 37, 163, 243, 299, 301, 421
Geopetitia chibiae Mawson, 1966: 416
Geopetitia falco Mawson, 1966: 102
Geopelitia streperae Mawson, 1966: 437
Geopetitia sp. 350
Gongylonema alecturae TH. Johnston & Mawson,
1942a: 107
Habronernatinae, unidentified: Mawson, 1969: 165
Habronema qaegatheles T.H. Johnston & Mawsan,
194]e: 248
Hadjelia acuariana (Gushanskaya, 1937): 246
Hadjelia truneata (Creplin, 1825); 261
Hamatospiculum chibiae T.A. Johoston & Mawson,
194]a: 416
Haumatospicilum haleyouis TH. Johnston & Mawson,
1941a: 256, 257
Hamatospiculum howense T.H. Johostan & Mawson,
1940a; 257
Hamatospiculum meneilli TH. Johnston & Mawson,
194)b; 240, 242
Hamatospiculum sp. T.H. Johnston & Mawson, 1941b;
93
Hamatospiculum sp. cf. quadridens (Molin, 1858): 103
Hamatospiculum sp.: 256
Aartertia sp; 126, 242
Heterakis bancrofti T.A. Johnston, 1912a; 107, 187
Heterakis catheturina TH. Johnston, 1912a: 107
Heterakis caudata (Linstow, 1906): 74
Heterakis chenonettae TH. Johnston, 19)2a: 74, 83
Heterakis circumvallata (Linstow, 19066): 72
Heterakis dispar (Schrank, 1790): 74
Heterakis gallinae (Gmelin, 1790); 195
Heterakis gallinarum (Schrank, 1788): 195
Heterakts papillosa (Bloch, 1782); 72, 74
Heterakis vesicularis (Froelich, 1791): 72, 74
Heterakis sp.: 69, 224, 242
° Hookworm': 65
Inglisonema typos Mawson, 1968a; 263
Leipoanema ellist TA. Johnston & Mawson, 1942a; 106
Lissonema brevicaudata (Chow, 1939): 242
Lissonema rotundata (Linstaw, 1903); 239
Lisscnema sp. Bain & Mawson, 1981: 242
Lissonema sp.t 107
Mawsonofilaria rhipidurae (TA. Johnston & Mawson,
1952): 304
Microfilaria gymnorhinae Gilruth, Sweet & Dodd,
1910: 434
Microfilaria sp, recorded by:
Bancroft, 1889: 199, 246, 261, 299, 310, 352, 359, 361,
414, 416, 418, 432, 434, 435, 442
Breinl, 1913a: 103, 263
Cleland, 1915: 246, 261, 294, 390
Cleland, 1922: 358:
Cleland & TH. Johnston, 1910: 352, 434, 442
Cleland & T.H. Johnston, 1912; 47, 93, 202, 246, 261,
305, 355, 359, 361, 373, 378, 390, 399, 414, 423,
424, 426, 429, 430, 432, 433, 442
TH, Johnston, 1910; 276, 352
T.H. Johnston, 1912a; 42, 46
T.H. Johnston, 1916: 248
Mackerras, 1962: 44, 91, 93, 131, 194, 196, 200, 214,
271, 349, 352, 359, 376, 400, 415, 416, 417, 423,
425, 427, 430, 432, 434
Plimmer, 1912: 184, 216, 359, 361, 434, 436
Plimmier, 1955; 407
Plimmer, 1916; 428
Scott, 1926: 163, 408
Scotti, 1927: 261
Mierofilaria sp: 242
? Microhadjelia spiralis (Mawson, 968d): 235, 27L,
274, 291, 309, 356, 359, 372, 414
Microtetrameres aegotheles Mawson, 1977: 248
Microtetrameres cacomantis Mawson, 1977: 233, 234
Micratetrameres circi, Mawson, 1977: 98
Microtetrameres coracinae Mawsan, 1977: 232, 271, 272
Microtetrameres cracrici Mawson, 1977: 432
Microtetrameres egretes Rasheed, 1960: 54
Microtetrameres eopsaltriae Mawson, 1977: 284
Microtetrameres gymnorkinae Mawson, 1977; 434
Microtetrameres helix Cram, 1927: 423, 438, 439, 440,
441, 442
Microtetrameres mirafrae Mawson, 1977: 265, 287
Microtetrameres meéliphazidae Mawson, 1977: 350, 352,
353, 354, 36], 362, 346, 367, 382
Microletrameres ninactis Mawson, 1977: 242
Microtetrameres ariolus Petrov & Tschertkova, 1950):
414
Microtetrameres paraccipiter Mawson, 1977; 92, 96
Microtetrameres pelecant (T.H. Johnston & Mawson,
1942¢): 37
Microletrumeres philemon Mawson, 1977; 356, 358,
359
Microtetrameres raptoris Mawson, 1977: LOL, 102, 104,
105, 242
Microtetrameres sphecotheres Mawson, 1977: 415
Microtetrameres sireperae Mawson, 1977: 437
Microtetrameres tvtonis Mawson, 1977: 244
Microtetrameres sp., Mawson, 1977: 93, 234, 274, 296,
422
Microltetrameres sp., Munday & Green, 1972: 439
Microletrameres sp.: 92, 236, 270, 290, 371
Odontoierakis bancrofti (TH, Johnston, 1912a): 107,
187
Onchocetcidae, unidentified: 415
Ornithostrongylinae, unidentified; 246
Oxyspirura anthochaerae (T.H. Johnston, 1912a); 350
Oxyspirura bancrofti T,.H, Johnston & Mawson, L941d:
358
Oxyspirura streperae T.H. Johnston & Mawson, 194la:
434, 435, 436
Oxyspirura.sp.t 242
Oxyuridae, unidentified: 242, 244
Paralemdana clelendi 'T.H. Johnston & Mawsori,
1940a> 435
Puaraprocta graucalinum (T,H. Johnston & Mawson,
1940a): 271
Parvseria diomedeae T.H. Johnston & Mawson, 1942d:
12, 13, 15
Paryseria macronectes T.H, Johnston & Mawson,
1942d: 15, 19
Paryseria pachyptilae T.H. Johnston & Mawson,
1942d; 25, 27, 32
Pectinospirura argentatu Wehr, 1933: 163
AIK REC, §. AUST. MUS, 19(15): 219-325
Pelecitus fulicaeatrae (Diesing, 1861); 123
Phacascaris sp., V.H. Johnston & Mawson, 1942d: 19
Physaloptera alata Baylis, 1925: lO2
Physuloptera hieracidiae T.A. Johnston & Mawson,
194]}b: 104
Physaloptere sp-1,H. Johnston & Mawson, 194Le: 66,
77, 93
Physaloptera sp.: 64, 93, LO1, 242, 433
Porrecuecum angusticolle (Molin, (860): 104
Porrocaecum circinuin T.A, Johnston & Mawson,
1941b; 88, 94, 97, 98
Porrocaecum clelandt T.H. Johnston & Mawson,
194d; 270, 276, 307
Porracaecum crassum (Deslongchamps, 824): 77
Porrocaecum ensicaudarum (Zeder, 1800): 117
Porrocaecum lobibicis Mawson, !9868b; 131
Porrvcuecum menurae T.H. Jahnston & Mawson,
1942b: 264
Porrocaecum reticulatum (Linstow, 1889): 52, 55
Porrecaecum serpentulum (Rudolphi, 1809): 124
Porracaecum Streperge T.H. Johnston & Mawson,
1941d: 434, 435, 437
Porracaecum wui Hsu, 1933: 434
Porrocaecum sp., T.H, Johnston & Mawson, 1941d: 362
Porrocaecuin sp.: 53, 99, 260, 263, 322, 440
Procamallanus murrayensis TH, Johnston & Mawson,
1940b: 44
Procyrnea dallfust (Mawson, 1968d); 242
Procyrnea falco (Mawson, 1968d): 102, 104, 105
Procyrnea incerta (A.J, Smith, 1908); 209, 213, 218
Procyrnea ? leptoplera (Rudolphi, 1819); 94, 104
Procyrnea mansioni (Seurat, 1914); 90, 91, 94, 98, 104
Procyrnea paraleploplera (TH. Johnston & Mawson,
1941b), JOS
Protospirura sp.t 253
Pseudamidostomum sp.. 72, 83
Pseudaprocia copemant Bain & Mawson, 1981: 280,
291
Pseudupracta pivzanthae TH. Johnston & Mawson,
[940a: 362
Pseudaproctella sp,, Mawson 1968b: 131
Rictularina spinosa T.H. Johnston & Mawson, |941d;
299
Schistogendria sp. 1: 288, 305, 374
Schistogendria sp, 2: 257
Schistogendria sp, 3; 257
Schistogendria sp. 294
Schistorophus cornurls Sobolev, (943: 151
Schistoraphus limosae Mawson, 1968b: 154
Schistorophus longicorais (Hemprich & Ehrenberg,
1866): 155
Schisierophus sp.c 254
Seiadocara umbellifera (Malin, 1860): 155
Sciadocara sp. 130
Serratospiculum uilenuatum (Rudolphi, 1803); 102
Serratospicului guttarum (Schneider, 1866): 93, 101,
102, 103, 104
Serratospiculurm tendo (Nitesch, 1857): 93, 100, LOI, 102
Seuratia marina TH. lohnston & Mawson, 194de: 35
Seuratia shipleys (Stossich, 1900); 12, 13, 14, 15, 16, 18,
19, 20, 2), 22, 23, 24, 25, 26, 28, 29, 31, 33, 35, 41,
49, 50, 168, 173
August, 1986
Seuratinema brevicaudatum T.H, Johnston & Mawson,
1941b; 243, 310
Seuratinema magnum T.H. Johnston & Mawson,
194d; 253
Seuratinema pomarostami TH. fohnston & Mawson,
194)d; 311
Seuratineme sp., Ogden, 1967: 246, 283
Skrjabinacerca sp.: 171
Skrjabinoclava decorata (Solonitzin, 1928); (51
Skrjabinoclava horrida (Rudolphi, 1809): 5, 135, 139
Skrjabinoclava sp., cl. S. alii Ali, 1968; 65
Skrjubinoclaya sp., Munday & Green, 1972: 139
Skrjabinoclava sp. 25, 29, 635, 147, 158
Skrjabinura brevicaudata (TH. Jotnston & Mawson,
1941b): 243, 310
Skrjabinura magna (T.H. Johnston & Mawson, 1941d);
253
Skrjabinura pomatosionn (7H. Johnston & Mawson,
194]d): 311
Skrjabinura sp.. 246, 283
Snireptera attenuata Rudolphi, 1803: 102
Spiroptera incerta A.J. Smith, 1908; 209, 213, 218
Spirurida, unidentitied, TH. Johnston & Mawson,
194le: 31]
Spirurida, unidentified, Munday & Green, 1972: 21, 22
Spirurida, unidentified,: 36, 92, 93, 101, 102, 134, 139,
164, 237, 244, 245, 252, 253, 258, 281, 291, 300, 311,
336, 346, 356, 459, 430, 433, 434, 439, 440
Splendidofilaria sp.c 424
Splendidofilariiae; unidentified: 102, 215, 434
Stegophorus diomedeae (1H, Johnston & Mawson,
1942d); 12, 13, 15
Slegzophorus macronectes (T.H, Johnston & Mawson,
1942d): 15, 19, 23, 24
Stegophorus pachyptilue (TH. Johnston & Mawson,
1942d): 21, 25, 26, 27, 29, 32, 33
Stesopharus stellaepolaris (Parona, 190): 33
Stegophorus sp. 12, 17)
Stellocaronema vlareolae Mawson, 1968%b: 158, 142
Stellocaranema charadrii Mawson, 1968b; 135
Stomachus sp., Munday & Green, 1972; 13, 1S, 22
Stomachus sp., T.H. Johnston & Mawson, 1951: 174
Streptocara crassicauda (Creplin, 1829); 69, 75, 80, 82,
86, 131, 137, 139
Streplacara fermosensis Sugimoto, 1930: 87
Streptocara pectinifera Neumann, 1900: 166
Streptocara recta (Linstow, 1879): 4, §, 46, 47, 87, 164
Streptocara sp., T.-H, Jahnsten & Mawson, 1942e: 78
Streptocara sp., Munday & Green, 1972: §2, 139
Streplocera sp. 5, 78, 80, 131, 154
Strongyloides sp.z 78
Subulura clelandi TA. Johnston & Mawson, 1941d:
238, 246
Subulura differens (Sonsina, 1890); 239
Subulura sp., TuH. Johnston & Mawson, |941b: 241
Svncueria contorta (Molin, 1858): 65
Spneuuria spe 3, 5
Svngamus trachea (Montagu, 1811): 189, 434, 439
Syngamus sp.: 107
Synhiinantus affinis (Seurat, 19b6). 241
Svrahimantus falco Mawson, 1982: 102, 104, 105, 380
Svahimanius flindersi (TH Johnston & Mawson,
1941b): 104
HELMINTHS FROM AUSTRALIAN BIRDS 319
Synhimantus fielding (Baylis, 1934a): 94
Synhimantus laticeps (Rudolphi, 1819): 242, 246
Synhimantus lichenostomi Mawson, 1982: 363, 374
Synhimantus podargi Mawson, 1982: 246
Svnkimantus sirry Khalil, 1931: 37
Synhimantus sp., T.H. Johnston & Mawson, 1941e: 42
Synhimantus sp., TH. Johnston & Mawson, 1942e; 47
Synhimanius sp.c 44,47, 73, 93,95, 242, 279, 318, 443
Tetrameres anseranas Mawson, 1979: 69
Tetrameres australis TH, Johnston & Mawson, 1941e:
72
Tetranreres bizitirae TH, Johnston & Mawson, 194le:
87
Tetrumeres calidris Mawson, 1968b: 155
Tetranieres ceria (Leidy, 1886): 16, 17
Tetrameres cladorhynchi Mawson, }968b:; 143
Tetrameres dacelonis Mawson, 1979: 253
Jetrameres diomedeae TH. Johnston & Mawson,
1942d: 16
Tetrameres fissispina (Diesing, 1861): 69, 77, 79
Tetrameres globosa (Linstow, 1879): L16, 117, 118, 119,
122
Tetrameres greeni Mawson, 1980; 37
Tetrameres gubanovi Shigin, 1957: 3, 5
Teirameres lobibicis Mawson, 19648b; 131
Tetrameres nouveli (Seurat, 1914): 139, 142, 144
Tetrameres pelecani TH. Johnston & Mawson, 1942c:
37
Tetrameres scolopacidis Mawson, 19686; 146, 157, 158
Tetrameres sp., T.H. Johnston & Mawson, 1942e: 78
Tetrumeres sp., Munday & Green, 1972: 119, 131, 139
Teirameres sp., Mawson, 1969: 165
Teftrameres sp.; 47, 53, 72, 73, 75, 78, 80, 102, 104, 109,
158, 163, 164, 242, 320
Thelazia aquilina Baylis, 1934a: 94, 95, 96, LO4
Thelazia dacelonis (Brein], 19136): 254
Thelazia pittae TH. Johnston & Mawson, 194le: 262
Thelazia sp.; 55
Torguatella balanocephula (Gendre, 1922): 260
Torquatella sp., Mawson, 1968d: 239
Torguatoides balanocephala (Gendre, 1922): 260
Torquatoides torquata (Gendre, 1922): 239
Torquatoides sp.z 239
Trichostrongylus incertus T.H, Johnston & Mawson,
194le: 166
Trichostrongylus renuis (Mehlis, 1864): 1, 72
Vagrifilaria australis T.H.. Johnston & Mawson, 19424;
239
Viktorocara limesae Mawson, 1968a; 151, 154
Viktorocara schejkini Gushanskaya, 1950: 151
Viklorocara sp., Mawson, L¥68b: 155
Vigniera chabaudi Mawson, 1968d: 246
Viguieru chibiae Mawson, 1968d: 416
Viguiera longicollis Mawson, 1968d: 287, 294, 350
Misuiera sp.: 309, 322, 423
Willmoitia australis Mawsan, 1982: 318
Xenocordon patonae Mawson, 1982: 380
Xenocordon gvmnorhinis (de Chaneet & Robertson,
1983); 434
Unidentified nematodes in published records, not nated
above:
Bradley, 1927; 4, 123
Cleland, 1922: 11, 35, 77, 94, 104, 240, 247, 253, 262,
276, 299, 311, 354, 362. 374, 399
Hall Exped, Report, 1974: 270
Nicoll, 1914e: 96, 104
Unidentified nematodes not previously recorded: 11,
34, 39, 40, 44, 63, 65, 132, 149, 150, 211, 212,
284, 302, 320, 328, 341, 383
Phylum Acanthocephala
Arhythmorhynchus brevis van Cleave, (916: 62
Arhythmorhynchus frassoni of T.H, fohnston &
Edmonds, 1951, not (Molin, 1858): 145
Arhythmorhynchus johnstoni Golvan, 1960: 145
Arhythmorhyachus limosue Edmonds, 1871; 154
Arhvthinarhynachus sp. 145
Centrarhynchus asturinus (TH. Johnston, 1913): 89,
92, 93, 94, 99, 104, 105
Centrorhynchus bancrofii (T.H. Jahnston & Best,
1943); 241, 242
Centrorhynchus falconis (T.H, Johnston & Best, 1943):
104
Cenlrorhynchus horridus (Linstow, 1898a): 253, 257
Centrarhynchus sp., T.H. Johnston, 1918; 242
Centrorhynehus spo 46
Corynosoema clavutum Goss, 1940: 43, 45, 46, 47
Corynosoma sp.. 11, 72
Echinorhynchus bazae Southwell & Macfie, 1925; 89
Echinorhynchus bulboeaudatus Southwell & Macfie,
1925: 239
Echinorhynchus menurae TH. Johnston, 19)2b: 264
Echinorhynchus pomatestomi TH. Johnston &
Cleland, 1912: 113, 276, 290, 294, 308, 409, 310, 3Lt,
312, 350, 332, 343, 346, 347, 349, 362
Echinorhynchus sp. Cleland, 1922: 305, 423
Echinorhynchus sp., 'T.H. Johnstan, 1910; 92, 257, 276,
305, 310, 311, 343, 380
Echinorhynchus sp., TH. Johnston, 1912a; 107
Echinorhynchus sp., TH. Johnston, 1912b: 145, 242,
425
Echinorhynchus sp,, T.H, Johnston, 19l4a: 145, 425
Echinorhynchus sp., T.H. Johnston & Deland, 1929:
329
‘Echinorhynch’, Cleland, 1922; 301, 423
Empodlius alecturac T.H. Johnston & Edmonds, 1947>
107
Filieellis sphaerocephalus (Bremser, 1819): 130, 163
Givantorhvnchus asturinus TH. Johnston, 1913: 94
Gortiorhynchus buncrofti TH. Johnston & Best, 1943:
24\
Gordiorhynchus faleonis TH. Johnston & Best, 1943:
104
Gordiorhynchus hvlae (TH. Johnston, 1914b): 246
Medtorhyachus alecturae (TH. Johnston & Edinonds,
1947): 107
Mediorhynchus corcorucis T.H. Johnstan & Edmonds,
1951: 423, 439, 440, 441, 443
Mediorhynchus earruli (Yamaguti, 1939b): 425
Mediorhynehus sp.: 294, 304, 352, 454, 425, 443
Oligacanthorhynchus pomatestomi (TH, Johnston &
Cleland, 1912); 11), 270, 287, 296, 309, 310, 311, 326,
332, 333, 341, 345, 350, 362, 366, 372, 410,429, 434
320
Oncivala pomatostami (TH. Johnston & Cleland,
19]2): 110, 111, 113, 270, 275, 276, 282, 287, 250, 293,
294, 296, 308, 309, 310, 311, 312, 325, 330, 333, 332,
333, 341, 343, 345, 346, 347, 349, 350, 342, 366, 372,
410, 425, 429, 434
Plagiorhynchus charadrii (Yamagutt, 1939b): 135, 139
Plagiorhynchus menurae (T,H. Johnston, 1912b): 264
Plagiorhyachus sp,. 29
Polymorphus biziurae TH. Johnston & Edmonds,
1948; 37, 65, G8, 87
Polymorphus sp. 72
REC. S. AUST, MUS. 1915): 219-324
Andeust, 1986
Porrerchis hylae (7.H, Johnston, 1914b); 239, 246
Prosthorhynchus charadrii Yamaguri, 1939b; 135
Prosthorhynchus evlindraceus (Goeze, 1782): 434
Prosthorhvachus menurae (TH. Johnstan, 1912h): 264
Prosthorhynchus sp,: 129, 131
Pseudoporrarchis hylae (T.H. Johnston, 19146): 239,
246
Unidentified specimens; 19, 47, 55, 67, 86, 89, 98, 102,
109, 114, 132, 141, 151, 154, 158, 166, 170, 282, 350,
359, 423
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Acanthocephala.’ Interscience Publishers Inc., New York. 423 pp.
YAMAGUTI, S. 1971. Synopsis of digenetic trematodes of
vertebrates. Keigaku Publishing Co., Tokyo. Vol. 1, 1074 pp.;
Vol. II, 349 pp.
YOUNG, M.R. 1939. ‘Helminth parasites of Australia.’ Imp. Bureau
Agriculture, St Albans, England. 145 pp.
ZEDER, J.G.H. 1800. ‘Erster Nachtrag zur Naturgesichte der
Eingeweidewtirmer.’ Bamberg. 432 pp.
RECORDS oF THE 6
PA
WS
SOUTH AUSTRALIAN E—™® a
MUSEUM By
VOLUME 19 NUMBERS 16-19 NOVEMBER 1986
No. 16 ANATOMICAL NOTES ON THE LAND SNAIL BOTHRI-
EMBRYON (PULMONATA: BULIMULIDAE) FROM SOUTH
AUSTRALIA AND WESTERN AUSTRALIA
by RON KERSHAW
No. 17 THE EPHEMEROPTERA (MAYFLIES) OF SOUTH
AUSTRALIA
by P J. SUTER
No. 18 FIRST REPRESENTATIVE OF THE ORDER
MACROSTOMIDA IN AUSTRALIA (PLATYHELMINTHES,
MACROSTOMIDAE)
by RONALD SLUYS
No. 19 A REASSESSMENT OF THE PAPUAN SUBFAMILY
ASTEROPHRYINAE (ANURA: MICROHYLIDAE)
by THOMAS C. BURTON
SOUTH AUSTRALIAN MUSEUM
North Terrace, Adelaide
South Australia 5000
ANATOMICAL NOTES ON THE LAND SNAIL BOTHRIEMBRYON
(PULMONATA: BULIMULIDAE) FROM SOUTH AUSTRALIA AND
WESTERN AUSTRALIA
BY RON C. KERSHAW
Summary
Comparative notes on the anatomy of the South Australian land snail Bothriembryon mastersi (Cox,
1867) from different localities are presented. Comparisons are also made with Bothriembryon
barretti Iredale, 1930 (Nullarbor Plain), Bothreimbryon melo (Quoy & Gaimard, 1832) type of the
genus (Western Australia), Bothreimbryon tasmanicus (Pfeiffer, 1853) (Tasmania) and seven other
Western Australian species. Features of the anatomy together with the protoconch sculpture of B.
mastersi and B. tasmanicus suggest a clinal distribution pattern. Therefore the recognition of a
subgenus Tasmanembryon Iredale, 1933 (Breure, 1979) is here suggested to be unjustified.
ANATOMICAL NOTES ON THE LAND SNATL BOTHRIEMBRYON (PULMONATA: BULIMULIDALY
FROM SOUTH AUSTRALIA AND WESTERN AUSTRALIA
RON C. KERSHAW
Honorary Research Associate, Queen Victoria Museum and Art Gallery, Launceston, Tasmania 7250,
(Manuscript accepted 20 June 1985)
ABSTRACT
KERSHAW, KR. C, 1986. Anatomical notes at the kid snail
Bothrienihryon (Pulmonate. Bulimulidae) trom South Australia
aid Western Australia, Ree. S. Awsy. Mus. 9(16)) 327-437.
Comparative notes an the anatomy of the South
Australian land snail Bothriembryon masterst (Cox,
1867) from different localities are presented.
Comparisons are also made with Bothriembryon
barreiti \redale, 1930 (Nullarbor Plain), Bothriembrvan
melo (Quoy & Gaimard, 1832) type of the genus
(Western Australia), Bothriembrvon rasmanicus
(Pfeiffer, 1853) (Tasmania) and seven other Western
Australian species, Features of the anatomy together
with the protovonch sculpture of B wtastersi and B.
lasmanicus suyvest a clinal distribution pattern.
Therefore the recognition of a subgenus
Tasmanembryan \redale, 1933 (Breure, 1979) is here
Sugeested to be unjustified.
INTRODUCTION
Specimens of the snail Bothriembryon mastersi
(Cox, 1867) collected by members of the Malacological
Society of South Australia from St Francis Island
(Nuyts Archipelago: 32°31'S, 133°18’E) and Vents Bay
(33° 11'S, 134°40'E) were dissected and compared with
specimens of this species from Port Lincoln and
Plitders Island, South Australia.
Study of (he Nullarbor Plain species Bathiriembryon
barretti Iredale, 1930 from two localities together with
Bothriembryon melo (Quoy & Gaimard, 1832) (Albany,
Western Australia), seven other Western Australian
species and the Tasmanian species Bothriembryan
tasmanicus (Pleitter, 1853) hus enabled presentation of
camparative notes on the anatomy and relationships
within the genus.
SEM micrographs taken by Dr A. Solem of the lield
Museum of Natural History, Chicago of the shell
protoconch sculpture of & mastersiand &, fasmuiicus
provide new information on the nature of this feature,
The recent evaluation by Breure (1979, pp. 91-96) of
the stalus of veneric level taxa suggested recognition
of Borhriembryon Pilsbry, 1894 and Tiesmanendrvon
lredile, 1933 as subgenera on the basis of differences
in the spermatheval duct and the protoconch sculpture,
The anatomical and protoconch data presented in this
paper permit further comment on fhe status of
Tasmanembryon,
The holotype of Bathriembryon mastersi (Cox, 1867)
described from Part Lincoln, Eyre Peninsula, South
Australia, has not been located, but an apparent
syntype from Flinders Island, 170 km to the north west
within the Great Australian Bight, is preserved in the
South Australian Museum (1.11341). The status of
another Eyre Peninsula species Bothriembryon
angusianus (Pfeiffer, 1864) which predates B masters!
has yet to be determined; matter, however, beyond the
scope of this paper, Full study on the South Australian
species and their shells must depend on fycture research.
Very little has been published on the anatomy of
Bothriembryon. Early camments and illustrations of
ecnitalia and buccal organs were presented by Semper
(1870) and Hedley (1889). Pilsbry (1946) gave more
detail and discussed the affinities of Bothriembryan
with New World taxa. Breure (1978b, 1979) revising the
Bulimulinac, has provided some descriptive and ulus-
tralive information. Breure (1978a) provided illustra-
tions of part of the radula of both A me/y and B.
(UsimManicus.
The following abbreviations have been used in this
paper: SAM, South Australian Museum, Adclaide;
NMV, Museum of Victoria, Melbourne; TM,
Tasmanian Museum, Hobart; QVM, Queen Victoria
Museum, Launceston; FMNH, Field Museum of
Natural History, Chicago; Ak, anterior ocsophagus:
AG, albumen gland; BM, buccal mass; D1, digestive
wland; E, prostate; EP, epiphallus; F, flagellum; GA,
genital atrium; HD, hermaphrodite duct; |, intestine;
KD, kidney; O, Free oviduct; P, penis; PC, pericardium;
PM, penial muscle; PV, pulmonary vein; R, rectum,
RM, penial retractor; S, stomach; SD, bursa duet; SL,
salivary gland; SO, spermoviduct; SP, bursa copulatrix;
T, talon; U, uterus; V, vagina; VD, vas deferens; SB,
site of bursa copulatrin,
MATERIALS AND METHODS
This work is based on study of more than 200 South
Australian and 100 Western Australian specimens from
the collections of the Museum of Victoria, the
Tasmanian Museum and the Queen Victoria Museum,
One “fossil” specimen of Bathriembryan barretti
Iredale, 1930, collected by Miss Karen Gowlett ai Petrel
Cove, St Francis Island, 22 January 1982, trom a sand
328 ROC. 5. AUST. MUS. 19 (16): 327-337
layer in the South-west clifl, 1s in Miss Gowlett’s
collection.
Selected Tasmanian specimens are included for
comparative purposes from a study of Bothriembryon
tasmanicus (Pfeiffer, 1853) (Kershaw, unpublished),
The results from dissections presented are taken from
the following material:
Bathriembryan mastersi (Cox, 1867)
St Prancis Island, Nuyts Archipelago (32°31'S,
133°18'E); collected 24 January 1982 by K. Gowlett
and R. Brown, 2 live and 12 dead specimens;
deposited as follows: SAM D 17089, 1 spirit; D
17091, 6 dry specimens; QVM 1 spirit, 4 dry
specimens,
Venus Bay, West Coast, Eyre Peninsula (33°11'S,
134°40'E); collected January 1982 by R. Brown, 14
live specimens; deposited as follows: SAM D 18090,
8 spirit specimens; QVM 6 spirit specimens.
Flinders Island, west coast Eyre Peninsula (33°43'5,
134°30'B); collected T. Castle, 6 March 1968, 5 spirit
specimens; deposited: SAM D 17089,
Port Lincoln, Eyre Peninsula (34°44'S, 135°52’E)
west side Spalding Cove; collected by B, J. Smith,
5 November 1969; NMV series spirit specimens.
Bothriembryon barretti redale, 1930
Wilson Bluff, Nullarbor Plain (31°40'S, 129°06’E):
collected by T. A. Darragh, March 1969, deposited:
NMYV series spirit specimens.
Eyre Highway 44 miles east of SA-WA boarder
(30°35'S, 129°20'B); collected by T. A. Darragh, 9
November 1973; deposited: NMV series spirit
specimens.
Bothriembryon melo (Quoy & Gaimard, 1832)
Near Albany Western Australia (35°02'S, 117°43'B);
6 animals collected by G. W. Kendrick, 22 January
1972; author’s collection courtesy 8, Slack-Smith,
Bothriembryon bulla (Menke, 1843)
Clarence, Western Australia (and other localities); 8
specimens collected by G. W, Kendrick, 15 May 1971:
T. M. Dartoall collection,
Bathriembryon glauerti lredale, 1939
Bluff Knoll, Stitling Range north from Albany; 6
specimens collected by B. R. Wilson, 28 May 1971:
T. M. Dartnall collection.
Bothriembryon indurus (Menke, 1843)
Walyunga National Park, Darlinp Range; 4
Specimens: T. M, Dartnali collection.
8oathriembryon serpentinus Iredale, 1939
Bickley, Darling Range; 7 specimens vcallected by
G. W. Kendrick, 10 July 197); T. M. Dartnall
callection.
November 1986
Bothriembryon savi (Pleilter, 1847)
Kudadup 200 m south of Jewel and Moondyne
Caves, 8 km north of Augusta; collected by Anne
Paterson, 6 July 1971; 6 specimens: T. M. Dartnall
collection.
Bothriembrvan leeuwinensis (Smith, [894)
Kudadup near Jewel and Moondyne Caves, § km
north of Augusta; collected by Anne Paterson, 6 July
1971; 6 specimens: T. M. Dartnali collection.
Bothriembryon kendricki Hill, Johnson and Merri-
field, 1983
Booragoon west side of Blue Crum Swamp; collected
by G, W. Kendrick, 13 June 1971; several specifnens
im T. M. Dartnall collection.
Bothriembryon kingil (Gray, 1825)
East of Wilson Inlet (35°00'S, 117°28'E); collected
by R. C. Kershaw, 12 May 1978; 3 live and many dead
specimens: QVM,
Bothriembryon tasmanicus (Pfeilfer, 1853)
Maria Island (42°38'S, 148°05'B); collected by R. H.
Green, 1 May 1969: OVM.
Cooks Beach (42°13'S, 148°18'E); collected by B.
Moore, 27 May 1971: TM E 8914,
All dissections were done by the author using a Zeiss
dissecting microscope and a Wild dissecting microscope
with drawing arm. The drawings and photographs are
by the author. Scanning electron micrographs were
done by Dr Alan Solem (FMNH), The specimens
selected from the listed collections, were coated with
a 150 Angstrom units gold coating in a sputter coater
and then studied with a Cambridge SR-10 slereoscan
microscope. These specimens are stored as follows:
St Francis |. and Venus. Bay: QVM; Maria 1,: QVM;
Bicheno: TM & 13642; Near Eagle Hawk Neck: NMV,
The dissection data are presented on a species and
locality basis within South Australia by comparison
with B masters! rom St Franeis |, and between all
other species and the South Australian species using
the St Francis |. morph as the basie concept,
OBSERVATIONS
While the distribution of B mastersi (Cox) is
confined within South Australia that of B barretti
Iredale extends into Western Australia on the Nullarbor
Plain, Both South Australian species haye an elongate
pallial structure (Pigs 2, 6, UL, 17) which does not differ
significantly from the generic features as described by
Breure (1979), Each species has a pale orange jaw with
16 transverse ribs usually of darker orange colour (Figs
2, 17). Apart from gross size differences there are no
important external bady features requiring comment
in this paper.
ANATOMICAL NOTES ON BOTHRIEMBRYON 329
inn
LULL
ws
FIGS. 1-6. Bothriembryon mastersi (Cox, 1867); 1, shell: (a) St Francis Island (natural size 21 mm), (b) Venus Bay (natural sive 17 mm);
2. St Francis Island, (a) pallial anatomy, (b) jaw; 3, St Francis Island, (a) whole genitalia, (b) detail of talon; 4, St Francis Island,
(a) view of lower female tract, (b) taternal view of penis; 5, Venus Bay, animal; 6, Venus Bay, (a) pallial anatomy, (b) jaw.
Bothriembryon mastersi (Cox, 1867)
Shell (Figs 1, 9)
Measurements of many specimens failed to reveal
more than minor racial differences in the adult from
the several localities. The sculpture and superficial
appearance is as describd by Cox (1867) and Iredale
(1937). The dimensions of shells from the dissected
animals are presented in Table 1.
Protoconch (Figs 24a, b)
The sculpture is usually described as pitted or pit-
reticulate. The optical microscope suggests ridges
crossing. obliquely. Anastomosing results in some
irregularity of the pattern, Shells from Port Lincoln,
Elliston, Wallanippie, Fowlers Bay, Streaky Bay, Venus
Bay and St Francis I, provided minor differences. In
some cases the basic wrinkled pattern as seen tn B.
lasmanicus can be seen near sutures.
Genital Anatomy
St Francis Island Material (Figs 3, 4).
Hermaphroditice duct thin, coiled, cream in colour,
expanded slightly before entering the talon laterally.
aH
Talon (Fig. 3b) 0.5 mm long, 0.3 mm wide, resting on
albumen gland surface, then descending into albumen
anteriorly (o join with albumen gland duct and enter
uterine passage, Albutnen gland pale orange. ca 7 mm
jong. Spermoviduct with uterus translucent greyish
amber in colour, prostate cream coloured slightly
brownish hear free oviduct. Latter short, 1.5 mm long,
inflated, internally with wide fleshy ridges entering
uterus, Vagina (Fig. 4a), ca 6mm long, internally with
broad low ridges just above atrium, apically becoming
low rounded ridges crossed by transverse lines of
pustules which continue as thin raised ridges into bursa
duct. Free oviduct opening into bursa-vaginal channel
is small and laterally oriented. Bursa duct ca 22. mm
long, the bursa copulatrix 3x2 mm, somewhat
globular, elongated and embedded above pallial apex
next to albumen gland, kidney and digestive gland
lobes.
Prostate narrowing to form thin, white tube of vas
delerens (Fig, 3a) which emerges from [ree oviduct ta
vagina surface to atrial region then ascending partly
free of penis to epiphallus to enter at base of flagellum.
Latter short with a speckled grey surface, Chamber of
penis (Fig. 4b) with 5 high subrounded. folded longi-
tudinal ridges modified into epiphallus. Atrium narrow
with relatively very small lobes.
Dimensions and relative sizes presented in ‘fable 2.
Venus Bay Material (Figs 5, 7, &).
Apparently at a similar stage of maturity the
anatomy closely resembles St Francis |. animal. Talon
larger, 0,5 mim long, 0.5 mm wide, albumen gland less
developed, 6.5 mm Jong of deep cream colour, uterus
translucent pale yellowish cream, Free oviduct longer,
2.2 mm, Vagina shorter, 4.5 mm. Internally the latter
(Fig. 8a) has Fine Ughtly folded ridges passing into
bursa duct. Similar oblique lines of raised pustules are
present.
Elongate globular bursa copulatrix appressed ta
uterine lobes. (Pig. 6a) adjacent dorsal crop surlace, not
embedded as in the St Francis J. animal, This is the
only mstance observed of this nature in South or
Western Australian animals and the desree of incidence
or relationship to maturity is not known. The bursa
duct coils through the uterine lobes normally.
The penial chamber (Fig. 8a) has 7 broad tightly
folded longitudinal ridges abruptly very thin with
epiphallus, Vas deferens appressed to penis-epiphallus
throughout,
Dimensions and relative sizes presented in Table 2.
The complexly folded distinctly everted atrium has
two large White lobes visible (Fig. 8b).
Flinders Island Material (Figs 14, 15).
A more mature larger animal (Fig. 14) with some
differences {Table 2), Talon (Fig. !Sb) large, 0.9 mm
long, 0.6 mm wide, albumen gland ca # mm long
greyish in colour, ulerus (ranslucent greyish but prostate
a5 in St Francis f. animal. Free oviduct short, 1.5 mm
REC. 5. AUST. MUS, 19 (16): 327-337
Navember 1986
TABLE |. DIMENSIONS OF MALE AND FEMALE GLNITALIA
OF DISHECTED SPECIES (MEASUREMENTS OF MALE
ORGANS AND SPERMOVIDUCT IN MM, PROPORTIONS OF
FEMALE ORGANS EXPRESSED AS PERCENTAGE OF
SPERMOVIDUCT),
ae
Species Male genitalia Female genitalia
P EP FE sO SD QO Vv
length length = %y My ™
RB, mtastersi
St Francis {- 3.0 20,0 3.5 24.5 90 O« 24,5
Venus Bay 25 [5.0 3.0 265 79 #8 (7.0
Flinders |. 25 WS 45 3h WW oS 95
Port Lincoln 25 Wa 3 20 Rl 12 15.0
B. burretit
Wilson Blur 10.0 «14.0 98.0 328.0 128 4h 12.5
hyre Highway 8.0 13.0 74) 3bO 1223 124) ido
B. meta 30 2234) 35 2.0 ay IS 15.0
By bulla 0 26.8 32 22.0 WW) IA 90)
B. wlaverti 60 24.0 16.0 WO [RO 7 140
B. durns 35 18S 100 3s oth OY 5.0
BR. serpentinus 4.0 16.0 1.0 22.0 6 9 6.5
B, suyt 2.0 8.0 80D 22.0 k20 OF 7.0
B. leeuwinensis 6.0 27.0 190 42.0 WO & acm
B, king 1.0 17,4 450 GS ce | 13.0
B. tasmanicus
Maria |, 5.0 19.0 65 19.U 53° «8 13.0
Cooks Beach §.0 265 90 21.0 sl ol4 (OG
with internal fleshy ridges, vagina short, 3 mm, with
internal folds entering bursa duct as very fine close
pustulose ridges. Bursa duct and bursa copulacrix as
in St Francis |. animal.
Penis chamber ormamence of broad flatly convex
ridges variably folded similar to Venus Bay animal. The
somewhat longer flagellum is normally very tightly
coiled. Dimensions in Table 2.
Port Lineolh Material (Figs 12, 13).
Animal of similar size and maturity to St Francis
I. specimen. ‘Talon (Fig. 12b) 0.5 mm long, 0.4 nim
wide, coloured pale cream, albumen gland 6mm Jong.
uterus translucent off white, tree oviduct longer,
2.5 mm, than in the other animals but only 1.7 pum
Neshy inflated.
Vagina, 3,2 mm, short as in Flinders 1. animal, the
internal pustulose ornament (Fig. 13) passing into the
bursa duct which, at 17 mm length, is relatively longer
than in the other animals,
Penial chamber ornament (Fig. 13) high rounded
ughtly folded ridges resembling the St Francis 1.
specimen.
Dimensions and relative sizes presented in Table 2,
Bothriembryon barrett |recdale, 1930
Shell (Fig, 16)
This is a large species with a distribution on the
Nullarbor Plain just west of Fowlers Bay in South
Australia, possibly as far as Balladonia in Western
Australia, 360 km trom Eucla and Wilson Bluff at the
border, The young shell has a very thin very pale brown
epidermis usually Jost in the adult, Descriptions are
provided by lredale (1930, 1937, 1939).
The “lossd” specimen (Fig. 6b) collected by Miss
Gowleti on St Francis |. appears toa be very similar to
ANATOMICAL NOTES ON BOTHRIEMBRYON Aw
PIGS, 7-9. Borhriembrvon niastersi (Cox, 1867); 7, Venus Bay, Whole genitalia: 8, Venus Bay, (a) internal view of terminal genitalia,
2 . op ' . / a .
(b) atrial complex, detail of folding: 9, Port Lincoln, shell (natural size 16 mm).
PIG. 10. Bothriembryvon angasianus (Pleiffer, 1864), Port Lincoln; shell (natural size 21 mm).
FIGS. 1-13. Bothriembrvon mastersi (Cox, 1867), Port Lincoln; 11, pallial anatomy; 12, (a) terminal genitalia, (b) detail of talon: 13,
internal view of terminal genitalia.
this species, the aperture proportions being almost
identical with the holotype figure (Iredale 1930), The
age of the specimen is unknown. Its appearance
suggests early Recent (subfossil?) or possibly Late
Pleistocene age. If correctly identified the range of the
species formerly extended further east than is now
apparent, The dimensions of this specimen and the
shells of dissected animals are presented in Table 1.
Protoconch
The optical microscope reveals radial oblique ridges
crossed by vertical or oblique ridges resulting in
somewhat squared pits. Descriptions (Iredale 1930)
refer to pits not distinguished notably from B. mastersi
(Cox).
Genital Anatomy
Wilson Bluff Material (Figs 18, 19).
Talon (Pig. 18b) somewhat elongate, 0.7 mm long,
0.4 mm wide, the elongate cream-coloured albumen
gland ca 12 mm long, uterus translucent off-white,
prostate cream. Free oviduct short, 3 mm, with internal
swollen fleshy ridges through half the length, vagina
November, 1986
REC. S. AUST. MUS. 19 (16): 327-337
1Omm
Wwuwpt
17b
TIGS. 14-15. Bothriembryon mastersi (Cox, 1867), Flinders Island; 14, animal showing genitalia in siru; 15, (a) whole genitalia, (b) detail
of talon.
FIG, 16, Bothriembryon barrett; Iredale, 1930; (a) Wilson Bluff near Eucla (natural size 30 mm), (b) St Francis Island “fossil” (natural
size 29.6 mm),
FIGS. 17-19, Bothriembryan barrett Iredale, 1930 Wilson Bluff; 17, (a) pallial anatomy, (b) jaw; 18, (a) whole genitalia, (b) detail of
talon; 19, internal view of terminal genitalia.
ANATOMICAL NOTES ON BOTHRIEMBRYON 433
(Fig, 19) slightly longer, 3.5 mm, with usual raised
pustular ornament together with fine ridges entering
bursa duct,
The bursa copulatrix is embedded above the pallial
apex asin B mnastersi {rom St Francis |. (Pig, 2a) and
Port Lincoln (Fig. 11) but the bursa duct is longer, 36
mim, not shorter than the spermaviduct,
Penis (Fig. 18a) very long, almost as. long as
epiphallus without a clear junction. The clearly
constricted region (Fig, 19) between the atrium and
penis, much more evident than in B mastersi, 1s
internally lined with thin folds, Penial chamber
(Fig. 19) ornament bold rounded raised weakly folded
ridges become thinner within epiphallus. The vas
deferens emerges [ram free oviduet wall tather Uran at
vawina junction as in B eastersi, and is then appressed
through to flagellum base. Flagellum, 9 mm, propor-
tionately much longer than in B, ¢mastersi (Table 2).
Eyre Highway Material (Figs 20, 21).
Talon (Fig. 20b) rounded. 0.6 mm long, 0,6 mim wide,
with relatively short hermaphrodite duct entering 3 mm
From apex, the narrowed exit duct passing deeply into
pale cream-coloured 7.5 mm long albumen gland.
Spermoviduet closely resembling Wilson Bluff animal
but {ree oviduct shorter, 3 mm, vagina longer. Vas
deferens (Fig. 21) within free oviduct narrows from
prostate to form a ridge continuous with one of Ileshy
free oviduct ridges. Vagina (Fig. 21) internal pustular
arnament is very close packed and becomes elongate
enrering the bursa duct, The latter, 34 mm long, is not
voiled within the uterine lobes mm the usual manner.
The penis-epiphallus dimensions are shorter than the
Wilson Bluff animal but the proportions are similar,
Dimensions and relative sizes in both these animals
are presented in Table 2.
Bathriembryon melo (Quay & Gaimard, 1832)
Shell
This shell is clearly described by Kendrick & Wilson
(1975) and no additional comment is needed, The
genital anatomy is described on a comparative basis
with dimensional data of this and other Wesiern
Australian species tabulated (Table 2).
Genital Anatomy (Figs. 22, 23).
This specimen is considered tepreseniative of the
several dissections made, Talon, 0.6 mm long, 0.5 mim
wide, differs in appearance from & masrersi with
hermaphrodite duct entering relatively basally.
Albumen gland short, ca 5 mm, cream-coloured, uterus
very pale translucent greyish white, prostate very pale
cream, free oviduct (Fig, 23), at 3.5 mm to4.5 mm in
length, relatively long with small fleshy ridges lining
most of the chamber internally. Vagina (Mg 23),
3.6 mm, shorl compared to St Francis. 1. B. masters?
but longer than some Western Australian species, The
internal ornament of oblique irregular low clangate
pustular liries becoming fine ridges in bursa duct,
resembles &. masiersi.
Bursa duet (Fig. 22) much longer than spermoviduct
bul globular bursa copulatrix embedded exactly as in
B. mastersi ftom St Francis |. (Fig. 2a) and Port
Lincoln (Pig. 11).
Penis (Fig. 22) short, its chamber (Fig. 23) lined with
about 6 close rounded folded ridges reducing in
number and size within relatively long epiphallus. Vas
deferens (Fig, 22) free from vagina surface in part and
penis, becomes appressed at epiphallus. Penial complex
proportions resemble those of B. mastersi.but the
oviduct and bursa duct are similar to B barrett
(Table 2).
The most significant feature noted in the other
Western Australian species dissected is [hat the bursa
duct length varies from 61% (B, serpentinus) to 191%
(B& bulla) of the spermoviduct (Table 2).
Breure has figured the genitalia of BR indurus
(Menke) (Darling Range) (1978a, Fig, 5; 1978b, Fig.
346), B gluuerti lredale (Darling Range) (1978b, Fig.
345) and B. onslowi (Cox) (Shark Bay) (1978b, Fig.
350),
Bothriembryon tasmanicus (Pfeiffer, 1853)
Shell
Complete animal and shell descriptions will be
presented elsewhere (Kershaw, unpublished). Selected
dimensions are presented in Table 1,
Protocanch (Figs 24¢-!),
The optical microscope reveals oblique wrinkles as
noted by Breure (1979). Spiral lines were reported by
Breure (1979, p, 95) but not by Pilsbry (1900, pl, 4),
These spiral lines have not been detected in very many
examples studied and their absence is supported by the
micrographs figured, It is suggested that the spiral
increment observed is due Lo the spirals observed in the
adult sometimes having a visible origin usually low on
the last protoconch wharl.
Genital Anatomy
The spermoviduct (Breure 1978a, Fig. 6; 1978b, Fig.
351) is broadly similar to the South Australian and
Western Australian species. The essential differences
are seen in the internal free oviduct and vagina
ornament with fleshy glandular development greatly
reduced in B& fasiunicus, The vagina has no pustular
ornament and that of the free oviduct is distinct, The
bursa duct is shorter than various Western Australiao
species and & Aarre?fi, but it is not shorter Lhan that
of B mastersi nor that of B. serpentinys (lable 2).
In Tasmania the range im length expressed as a
proportion of spermoviduer length af 40% to 100%,
can also imelude & suvi and .B, kingil, These data were
unknown lo Breure (1979) whose Tasmanian material
4344 KLC. S. AUST. MUS. 19 (16): 327-337
TABLE J SHELL MEASUREMENTS (MM) OF SOUTH
AUSTRALIAN AND TASMANIAN SPECIES
Spevies Whor) Heizht = Diainetei Aperture
Number Height
A. mastersi (Cox)
Si raticis. t-
dissected 4p. 4.75 18.2 11.3 1.1
largest seen 3.375 T1.0 13.4 Vs
Venus Bay 4,75 16,8 12.) 1.4
Vhinders t. 3.0 20.0 {2.5 1.0
Port Lincoln $875 16.0 10.0 9.2
R. barreiti Iredale
Wilson Bluth
disseered 4p. S.R75 29.1) 15,0) 14.3
larwest secu 6.00 31.2 16.8 (6.0
Lyre Highway S75 24.1 14.4 14.2
St Frans [.
fossil sp, 5,4 19 16.49 18,9
RB. fasinaniens (Preitfer)
Matia f. 4.75 22.4 13.7 14.1
Cooks Beach 4.75 21.0 12.5 13.0
came tram Coles Bay. Six specimens from this locality
huve a mean of 53% of spermoviduct length.
The significant distinguishing feature of the female
genitalia and spermoviduet is that the bursa copulatrix
is always appressed Lo (he uterus adjacent the base of
the albumen. gland. Ir is never embedded above the
pallial apex as in the Southern and Western Australian
species. Only the one specimen from Venus Bay has
any resemblance.
Compared with & melo and B mastersi the penis
is longer and (he internal penial ornament is not folded.
The flagellum is longer than in those species but is
comparable with & barrett? and other species (Table 2).
B& tasmanicus has a rounded pallial region compared
with the elongate western forms. The shallow atrium
has no constriction preceding the penial lomen. The
species does form a very distinct entity but the features
suggest along period of isolation rather than consistent
subgerieric features, OF the features used by Breure to
define Tasntanembryon only the cadula may he
consistent.
DISCUSSION
The basic data on variation in the material studied
are summarized in Table 1, Shell dimensions. of the
South Australian species. aré give ii Table 2. The
dimension of the largest specimen seen in each species
is alsa given for comparison, From these data it can
be Seon that the shell of B. wiuszersi compare in size
to 8 tasmunicus but the aperture of the latter is nearer
lo B barreiti. Despite differences in shell heighe,
hawever, {here is surprising similarity in spermoviduct
length, The most obvious differences are in the propor-
Tionate lengths aF the bursa duct and secondly in the
penial complex,
Genital Anatomy
B. barretti and &. kingii both have a much enlarged
penis bul the flagellum differs. B wasters/, B, melo,
B. indutus, 8. serpentinus and B. savi all have a small
penis bur the Iagellum in the first two is rather short
Navewrber 19a8e
such as that found in & bulla. & tastanicus and others
tend to be intermediate, The bursa duet is signifieanrly
longer than other species in & barren’, 8 ntelo: B. bulla
and & glauerti.
The South Australian & masters? from the localities
studied, hasa duct length (mean of 80%) that is inter-
mediate between the long ducts (mean 11870) of several
Western Australian species (see also Breure 1978h, pp.
203-209; 1979, pp, 92-96) and the duct of B rasnrnicus
(mean 63%). The length of the duct in the latter species
varies considerably (Kershaw, unpublished) the range
in different morphs as already noled being inclusive
of that noted for B&B mustersi and other species,
The South Australian B masters/ is thus not strongly
differentiated [rom the Western Australiari species with
which it shows clear affinity but could be part of a
clinal gradient across southern Australia, More species
need study to allow further evaluation of this theme;
but the similarities observed in & fasinanicus together
with the relative dimensions given appear Lo support
the concept,
Protoconch
Few scanning electron micrographs have been
published of bulimulid species. Breure (1978a, 1979,
Pls |-3) has provided several including B. tasmmanicus
and &. onslowi (Cox), He does not include micrographs
of other genera he considers related to Boshriembryon
nor does he expand on his data other than to Separate
Tasmanembryan ‘trom Bothriembryon, Since the
structural details are cormplex, subject to change
because of surface wear, and difficult to illustrate
oplically, more detailed study is desirable,
The photographs presented here of Borhriembryon
masiersi and B. tasmanicus ave the first ta enable
comparison both on a species and locality basis (Figs
24a-f) logether with some different paris of the
protoconch. Breure’s (1979, PI, 2) very useful work
illustrated only part af one specinien of each af B
fasmanicus and B onslowi.
The specimen of 8 mastersi trom Venus Bay is
unworn, with both apical and post apical sculpture well
defined (Fig. 24a). Note that the radial ribs on the apex
are mostly continuous, with only a few anastomosing,
or ate interrupted along their length. In the worn
specimen from St Francis Island (Fig, 24b) note that
the apical sculpture is absent just abave the periphery
and the lower part of the first whorl has the sculpture
worn down into minute “pits”, although the typical
sculpture can be seen on the right margin of the
photograph. The post-apical sculpture is sinvilarly
degraded in comparison with that of Venus Bay
Specimen (Fig. 24a).
The specimen from Maria Island (Figs 24e, d), the
probable type locality of & sasmanicns, is unworn, The
apical sculpture consists ol generally continuous radial
ribs With some anastomosing (Pig. 24d) at the sutures,
While the post-apical sculpture is relatively inconspi-
cuous on the upper spire: Another unworn specimen
ANATOMICAL
NOLES ON BOTHRIEMBRYON 335
FIGS, 20-21. Bathriembrvoan barrett Ieedale, 1930 Eyre Highway; 20, (a) whole genitalia, (b) detail of talon; 21, internal view of terminal
penitahia.
PIGS. 22-23. Borhriembrven melo (Quoy & Gaimard, 1432) Albany; 22, Whole genitalia; 23, internal view of terminal genitalia.
from Bicheno (Fig, 24e) shows interruptions in the
apical radials, more frequent anastomosing, and much
more prominent early spire sculpture, A worn example
{rom just north of Eagle Hawk Neck, south-eastern
Tasmania, shows increased irregularity of the apical
radials and more interruptions (Fig. 24f),
None of the Tasmanian specimens examined with the
SEM show the spiral sculpture as claimed by Breure
(1979, p, 92). Lt is possible that the points of irregularity
visible (Figs 24e, f) in these specimens could be
misinterpreted as spiral lines during optical viewing.
The effect is noticeable above the suture in the Bicheno
specimen (Fig, 24e), But Breure’s figure of B.
rasmanicus, presumably from Coles Bay, does show
someé trace of spiral joining of irregularity points on
the lower part of the whorl (1979, pl. 2, Pig. 4). The
possibility (hat adult spirals sometimes develop in the
lawer protoconch and sometimes do not has been
suggested in comment on the species above,
In addition the illustration of B onslowl (Breure
1979, Pl, 2, Fig, 3) shows traces of the irregular radials
near the surure seen in some South Australian
specimens. The apical sculpture is otherwise as seen
in the western species.
Since (he dissections reported on above show an
apparent clinal variation in proportionate bursa duet
length, and the SEM photographs of South Australian
and Tasmanian species of Bofhriembryon illustrate
apical shell sculpture composed of the same basic
elements rather than distinctive elements, the evidence
does nol support subgenerie distinction, Thus Breure’s
(1979) suggestion of subgeneric status lor Tasyan-
embryon Iredale, 1933 requires further evidence i it
is lo be supported, The present evidence suggests that
only one generic unit, Bofhriembrvon Pilsbry, 1894,
is recognizable for the Australian Bulimulidae.
ACKNOWLEDGMENTS
This report was made possible by the provision of
specimens from St Francis Island and Venus Bay for
which gratitude is due to Miss K. Gowlett and Messrs
N. J. C, Holmes and R. Brown, Gratitude is also due
for specimens and loan of collections which
comtributed to the full study. Mrs Shirley Slack-Smith
provided specimens of B, me/o collected by Mr G.
Kendrick and Mr Wolfgang Zeidler (South Australian
Museum) loaned specimens from Flinders [sland.
Callections studied were provided by Dr B. J. Smith
(Museum of Victoria), Miss Alison Green and Mrs B,
Turner (Tasmanian Museum), Messrs C. B, Tassell and
R, H, Green (Queen Vietoria Museum). Dr A. J. and
336 REC. S. AUST. MUS. 19 (16): 327-337 November, 1986
FIG. 24. Scanning electron micrographs of Bothriembryon apical sculpture. (a) B. mastersi, Venus Bay, whole apex = 15.7; (b) B. mastersi,
St Francis Island, worn sculpture x 48.8; (c) B. tasmanicus, Maria Island, whole apex x 17.4; (d) B. tasmanicus, Maria Island, whole
first stage x 88.5; (e) B. tasmanicus, Bicheno, whole apex x 18.1; (f) B. fasmanicus, near Eagle Hawk Neck, whole apex x 20.1.
ANATOMICAL NOTES ON BOTHRIEMBRYON 337
Mrs J. A. Dartnall kindly made available material,
drawings and manuscripts through the courtesy of the
Tasmanian Museum.
Dr Alan Solem (Field Museum of Natural History,
Chicago) made the SEM photographs used and
provided very valuable discussion on the results. He
also made valuable comments on earlier drafts of this
manuscript and suggested an outline for the present
approach. Dr Brian Smith provided many suggestions
and discussion during the study and commented on
the manuscript. Gratitude is due to reviewers for
encouragement and some useful comment on presen-
tation, The Science and Industry Endowment Fund has
provided equipment and support critical to the success
of my research for which gratitude is expressed to the
Trustees.
REFERENCES
BREURE, A. S. H. 1978a. Taxonomical, ecological and zoogeo-
graphical research on Bulimulidae (Gastropoda, Pulmonata).
Malacologia 18: 107-114.
BREURE, A.S. H, 1978b, Notes on and descriptions of Bulimulidae
(Mollusca: Gastropoda). Zool. verh, Leiden No. 164: 1-255.
BREURE, A, S. H, 1979. Systematics, phylogeny and zoogeography
of Bulimulinae (Mollusca), Zool. verh. Leiden No, 168: 1-215.
COX, J. C. 1867. Characters of five new species of Australian land-
shells. Proc. Zool. Soc. Lond. 1867: 39-40,
HEDLEY, C. 1889. Anatomical notes on the Helicidae. Part ILL. Prac,
R. Soe. Qiland. 6: 249-251.
IREDALE, T. 1930. Notes on some desert snails. Viet, Nat. 47:
118-20.
IREDALE, T. 1933. Systematic notes on Australian land shells. Rec.
Aust. Mus. 19: 37-59.
IREDALE, T, 1937, An annotated checklist of the land Mollusca
of South and Central Australia. S. Austr. Nat. 18: 6-59.
IREDALE, T. 1939. A review of the land Mollusca of Western
Australia. J. R. Soc. Western Australia 25: 1-88.
KENDRICK, G. W. and WILSON, B. R. 1975, Nomenclatural notes
on the land snail genus Bothriembrvon Pilsbry, 1894
(Pulmonata, Bulimulidae), with descriptions of the type and
two other species. Rec. West. Aust. Mus, 3; 295-325.
PFEIFFER, L. 1853. Description of fifty-four new species of Helices,
from the collection of Hugh Cuming, Esq. Proc. Zool. Soc.
Lond,, 1851 Part 19: 252-263.
PFEIFFER, L. 1864, Description of ten new species of land-shells
from the collection of George French Angas, Esq. Proc. Zool.
Soc. Lond., 1863: 526-529.
PILSBRY, H. A. 1894. Note on Liparus. The Nautilus, 8: 35-36.
PILSBRY, H. A. 1900. Manual of Conchology, (2). 13. Philadelphia,
Academy of Natural Sciences of Philadelphia, pp. 253.
PILSBRY, H, A. 1946. Notes on the anatomy of Australian and
Galapagos Bulimulidae (Mollusca, Pulmonata), Notol. Nat, No.
168.
QUOY, J. R. C. and GAIMARD, J. P. 1832. “Voyage de decouvertes
de l’Astrolabe Zoologie” 2. Paris Tastu.
SEMPER, C. 1870. Reisen im Archipel der Philippinen. Wiesbaden:
Kreidel.
ADDENDUM
Since the above text was written Ludbrook (1984: pp.
301-302, Fig. 82) (“Quaternary Molluscs of South
Australia”. Department of Mines and Energy, South
Australia, Handbook No. 9) has described and figured
fossil Bothriembryon barretti barretti \redale from the
Semaphore Sand Formation and the Upper Member
Bridgewater Formation. The St Francis Island specimen
resembles her Figure 82a and b in form and boldness
of sculpture, but appears to be an older specimen.
THE EPHEMEROPTERA (MAYFLIES) OF SOUTH AUSTRALIA
BY P. J. SUTER
Summary
In a study of the Ephemeroptera of South Australia, thirteen species have been recognised. All
species have been described from both adult and nymphal material and keys enabling their
identification are included. Five new species have been recognised, Atalophlebia auratus sp. nov.,
Nousia pilosa sp. nov., Ulmerophlebia pipinna sp. nov., Cloeon paradieniensis sp. nov., and
Centroptilum elongatum sp. nov.; and one species transferred from each genus Atalophlebia and
Atalonella to Nousia. (Nousia inconspicua (Eaton) comb. Nov. and Nousia fuscula Tinnyard comb.
Novy.). The first associated nymphal descriptions of five previously described species Nousia
inconspicua (Eaton), N. fuscula (Tillyard), Baetis soror Ulmer, Cloeon fluviatile Ulmer, and
Tasmanocoenis tillyardi (Lestage) are also made. Two species have been redescribed from South
Australian material; Atalophlebia australis (Walker), and A. australasica (Pictet) and one species of
Tasmanophlebia is described but not formally named.
CHE EPHEMEROPTERA (MAYEFLIES) OF SOUTH AUSTRALIA
by
P. J, SUTER!
Department of Zoology, University of Adelaide, Adelaide, South Australia S000
(Manuscript accepted 2 July 1985)
ABSTRACT
SUTTER Bo. 1986. The Ephemeroptera (Maytlies) of South
Australia. Ree, So lad Mus, 19(07): 339-397,
In astudy of the Ephemeroptera of South Australia,
thirteen species have been recognised, All species have
been described from both adult and nymphal material
and keys enabling (hetr identification are included, Five
new species have been recognised, Afalophlebia auratus
sp. nov., Nousia pilosa sp, noy,, Ulmeraphlebia pipinna
sp, noy., Cloeon paradieniensis sp. nov., anc
Centroplilum elongatum sp. nov; and ane species
transferred from each genus Afvalophlebia and
Atalonella to Nousia, (Nousia inconspicua (Eaton)
comb, nov, and Nousia fuscula Tillyard comb, novy.),
The first associated nymphal deseriptions of five
previously deseribed species Nousia inconspicua
(Baton), N Juscula (Tillyard), Raetis soror Ulmer,
Cloeon /luyiatile Ulmer, and Tusmanacvenis tillvardi
(Lestage) aré also made. Two species have been
redescribed from South Australian material;
Atalophlebia australis (Walker), and A. australasica
(Pictet) and one species of Tasmanophilebia is deseribed
but not formally named.
INTRODUCTION
The first species of mayfly in South Australia was
recorded by Eaton in 1871 when Leplophlebja incon-
spicua was described from Adelaide. In subsequent
papers 1883-1888 Eaton placed this species into the new
genus Allglophlebiu, Since that date only two other
records of the Ephemeroptera in South Australia have
been made, Harker (1954) recorded A/alophlebia
australasica from Tillyard’s 1934 collection near Mount
Gambier; and Timms (1974) recorded a “Caenis sp.”
in Valley Lake, Mount Gambier, and L. Edward near
Millicent, Both Williams (1968) and Rick (1970)
acknowledge the presence of mayflics in South
Australia, but neither mention any specific families or
genera.
With the exception of the species described by Eaton
(1871) the other records of Ephemeroptera are from the
wet South East of South Australia. It appears that
because South Australia is the driest State in Australia,
the existence of freshwater in regions other than the
South East had becn ignored, leaving a large gap in
' Present address: State Water Laboratory, Engineering and Water
Supply Departement, Private Mail Bag, Salisbury Post Office South
Australia 5108.
our knowledge of Ausiralian freshwater invertebrales,
and zoogeographical relationships of these animals.
Preliminary collections from the Fleurieu Peninsula,
the Mount Lofty Ranges and the Flinders Ranges
showed that mayflies were abundant in all permanent
freshwater streams, waterholes and in many dams and
reservoirs. Further collections from the freshwater
habitats in South Australia have led lo the recognition
af 13 species of Ephemeraoptera, all but one of which
have nymph and adult associations confirmed in the
laboratory or in the field,
AIL 13 species have been described from both adult
and nymphal material and keys to their identification
ate included. Five new species have been recognised,
Atalophlebia auratts, Nousia pilosa, Ulmerophlebia
pipinna, Cloean paradieniensis and Ceniroplilurm
elongatum; one species transferred from each genus
Atalophlebia and Atalonella to Nousia (Nousia
inconspicua (Eaton) comb. nov. and-Nousia fuscula
(Tillyard) comb. nov.) and the first associated nymphal
descriptions of five previously described species;,
Nousia inconspicua (Eaton), N, fuseula (Tillyard),
Baetis soror Ulmer, Cloeon fluviatile Ulmer and
Tasmanocoenis tillyardi (Lestage) made. Enlarged
descriptions of Afalophlebia australis (Walker), and A,
australasica (Pictet) are also given, A single species of
Tusmanophlebia was also recorded and is described,
but since a revision of the Siphlonuridae is being
prepared by Dr I, C, Campbell material has been sent
to him to include in his more detailed studies af this
group.
MATERIALS AND METHODS
Collection of nymphs was by dip-net with mesh-pore
size of 500 um, or by hand of nymphs clinging to the
under-surface of rocks or bark in streams. Adults were
collected by beating the vegetation along river banks
with an insect net, or by sweeping the net through a
swarm. Specimens were preserved in 70% ethyl alcohol
with 5% glycerol.
Specimens for study were dissected under a Wild M5
stereoscopic microscope, and (he appendages (legs and
wings of adulls; legs, mouth-parts and gills in nymphs)
were mounted on glass shdes using “Buparal” or
Polyvinyl lacto-phenol mounting media. The sterna,
and nymphal abdominal terga, were prepared using the
techniques of Tsui and Peters (1972, 1975), and the
340 RECS. AUST. MUS. 19 (17): 339-397
nymphal tentoria were studied by the
by Hudson (195)),
Genitalia and eggs were cither mounted on slides or
prepared for the Scanning Electron Microscope
(S.E.M.) using eritical-point drying which eliminated
both shrinkage and distortion. They were then mounted
and coated with Au/Pd and examined using an E‘T.B,C,
Autoscan with un operating volilave of 5, 10, or 20 KY,
Hlustrations of wings and body colour-patterns were
made using a Wild M3 or M5 stereoscopic microscope
with an attached drawing head. Legs, mouthparts, gills
and high magnification (>100») illustrations were
drawn using a Wild compound microscope and camera
lucida.
Wing venalion terminology is based on Tillyard's
(1932) scheme, and as illustrated in Figures la and Ib
used by Peters & Edmunds (1964, 1970, 1972) and
methods given
Edmunds, Jensen & Berner (1976), Each segment of
the fore, middle and hind legs of the nymph and male
imago is compared to the length of the fernur, and is
expressed as a ratio, The absolute length of the fernur
is given last in parenthesis,
In figures of the labium the method used by Peters
& Edmunds (1964, 1970, 1972) 1s follawed, with the
ventral surface shown on the right hand side of the
illustration, and the dorsal surface on the left.
Comparative measurements of the segments af the
labial palpi and maxillary palpi are expressed as ratios,
compared with the proximal segment length, which is
given im parenthesis. All measurements are given in
mullimetces.
KEYS TO THE SOUTH AUSTRALIAN
EPHEMEROPTERA
The following keys will serve to distinguish the male
imago and mature nymphs of the species of Epheme-
roplera recorded in South Australia. Figures are
included with eavh couplet to give illustrated examples
of the key characters, although oceasionally these are
not required (eg. “terminal filament present” as
compared with “terminal filament absent”).
The key to the imagos is primarily for male
specimens because species identification of the female
imagos ts often very difficult as external morphological
characteristics of the female show generic, rather than
specific affinities, Only direct association with nymphs
can allow accurate identification of females, using
external characters. However, with the limited South
Australian mayfly fauna, the females have also been
included in the key and van be distinguished by external
morphological characters in all genera with the
exception of Naysia. The external characteristics are
useful to distinguish this genus, bul species Separation
is difficult without direct nymiphal association, or a
study of the morphalogy of fertilized eggs. Identili-
cation of subimaygos is not as successful, Generic
Segregaling characters ale present in subimagas, and
generic identification can be made using the imago key,
November 1Y8e
For specific identification it is necessary to refer to (he
subimago characterislics listed in the description of
cach species.
KEY TO SOUTH AUSTRALIAN ADULT
EPHEMEROPTERA
Ja Hind wings comparatively large, about half as lone
as the fare wing (Figs 28a, b). Male penes long ane
tubular (Pigs 30a, b) , Siphlonuridae
Only one represen(alive in South Australia: Tesmana-
Phlebia sp.
Ih Hind wings small ar entirely absent ..2..2,.., 2
2a(1) Fore wings with many cross-veins: hind wings small,
also with nvany cross-veins (Figs la, by 3a, b; Sa, b;
Ra, b; on b; i2a, b, ¢; 17a, by... ALY Big tot
tree Leprophlebiidac 3
2b) «Fare wings with few eross-veins, clear open
appearance, hind wings present or absent, if present
reduced, with cross-veins few or absent (Figs 2la, b;
23a, b; 25a; Sia, 2)... Soprigrta:d Fees 2c 9
Yarsal claws of each pair similar, elongate, slender,
hooked distally (Figs 1g, 3u; Sh; Be; 10g; 12%); fore
wings <3.9 » longer [han wide (Figs la; 3a; Sa; 8a;
Ws “Feak tt ce ay th ee De! ee
3b Tarsal claws of each pair dissimilar, one clongate,
slender, hooked distally, one blunt with a distal tooth,
club shaped (Fig, 17h); fore Wiles >3.5 » longer
than wide. Male gertilalia (Figs 17¢: 19a, b) - :
Ulmernphiebia
Only one species in South Australia: Of pipinna sp.
noy.
Large species, body length > 8 mm; fore wing 7-13
mm; Se of hind wing reaches wing margin at 9/10
wing length (Figs lb; 3b; 5b)... Afolophlehia 5
db Small species, body length < 8mm; fore wing 5-8.5
mm, 5¢ ol hind wing reaches wing marin at 3/4 of
wing length (Pigs 8h; lb; 12by...-.. Nausia 7
Males and females with terminal filament; body
colour yellow-brown (orange): female with ninth
abdominal sternite deeply ingised (Figs les Si) , 6
Sh Males without terminal filament (present in females):
body colour black. Male genitalia (Fis 3c, dj 7e, d);
female With ninth abdominal srernite with shallow
incision (Fiz. Je) _.._, Atalophiebia wustralasica
Fore Wings with darker pterostigmatic region, cross-
veing Of costal and subcostal spaces shaded with
black, hind wings and rest of tore wings hyaline (Fig.
la, b); bady yellow with black thorax. Apices of penes
widely separated and divergen( (Pies le, dy; Ta, b)
AL. australix
Petnatese
3a(2)
OUR eh eee eee eee pee ee te tee
4a(3)
Sa(4)
6a(5)
6b Pore wings linged entirely with yellow, pterostigmati¢
region darker than rest of wing; hind wing erey-
brown, darker than fore wing (Fig. 5a, b). Apices of
penes convergent (Figs 56, di Fe, 1),
ween tens 8 Ege tereeteet-tte” A. nateiitils sp. NOV:
Two halves of penes distinctly separated forming a
U-shape (Figs le, dy 15e, d); with a small vriangular
spine on inner margin (visible using transmitied light,
Tal4)
Hig. 10c}e..- 45% oiasc thames Nousia fuscula
Tb ‘Two Halves of penes held! close Lovether, without spine
f-Heuaiat crepe Searieo ek stard: fh doe
Ba(7) Penes with wo ventral lobes near apex (Fins 8a; 15a,
Mag: pe Megas i sgrie N. incaunspieud comb, TOY
&b Penes withour ventral lobes (Pius 12d. 0: (Se, Fy),
N. pilosa sp. We,
i ce ae oar a ae er rer
EPHEMEROPTERA OF SOUTH AUSTRALIA
9a(2) Males with turbinate eyes (females without dorsal
compound eves); marginal intercalaries short,
developed between the ends of the main veins (Figs
Da; 23a; 25a, 2); hind wings present or absent
Seti Baeridae 10
Males and ‘females with small lateral eyes, dorsal
compound. eyes absent; na marginal inrercalaries
present (Pig. 31a); hind wings absent. Penes fused
will apical indentation, forceps sirongly bowed (Figs
Silo; SHAY = oe cat tit Caenidae
Only one species in South Australia: Tusmanocdenis
tillvareli
10a(9) Hind wings present... ...... seer oe !
Wb = Hind wings whsenr —..- - he cnc: Cloeon 12
1 1u(10) Marvinal intercalanies of tore wing paired (Fig. 21a).
Male genitalia (Pig. (9c; 2ld)... 2.0.4) --- Bactis
Only one species in South Australia: B serar
Marvinal intercalaries of lore wing single (Fig. 23a),
Male venilalia (Figs 19d; 23d). ...- Centroptifiumn
Only one species in South Australia: C. elongatum
8p). OW.
12a(10) Males will jurbinale eves, yellows terminal segment
of foreeps icangular (Figs 19% 2S5i), females with
costal atid subvostal spaces al wings shaded
red/ brown; hody length > 7 mm
Clacon purucieniensis sp, now,
hn
oertrt ey enn
(hb
13b Males with lurbinale eves, sepia: lerminal segment
of loreeps small and globular (Figs 9c; 25d); females
with castal and subcostal spaves of wings apace,
milky; body length < S mm ....,-- ©. fluviatile
KEY TO THE NYMPHS OF SOUTH AUSTRALIAN
EPHEMEROPTERA
la Head prognathous; tail filaments with whorls of setac
al apex ol each segment, body dorsa-ventrally
fattened. Families Leprophlebiidac; Caenidae., 2
Ib Head hypognathous; tail filaments fringed laterally
with long line setae, body usually cylindrical, Families
Baetidae; Siphlomutidae,,....0. 0.2.0 eee 9
2a(!) Seven pairs of gills inserted laterally on abdomen,
each gill consisting of a pair of lamellae ,..,., 3
2b Six pairs oF gills, Ist very small mono lilament, 2nd
enlarged to form an elytriform gill cover, remaining.
pairs bearing long tracheal filaments (Fizs 32b, c,d)
Caenidae: Yasinanovoenis
Only one species in South Australia: To ¢i//varedi,
Gill base broad with one apieal Hlament, lined with
fine setae (Fig, Ie), legs and body covered with long
fine setae (Fig. 20) Ulmerophlebia
Only one species in South Australia: ( pipintia sp.
nov.
Gills lanceolate or linear, with one apical filament
(Figs 9: He; 13e), apex with muluple tracheal
filaments (Pigs 2c; dc), body not covered with lang
line setae
3a(2)
ee
Gills lanceolate or linear, nat branched at apex into
tracheal filaments (Pigs Sey Ley 13c).. Nousia 5
Gills broad al apex, with each lamella subdivided into
tracheal filaments (Figs 2c; 4c) - Atlophlehia 7
Gills lanceolate (Figs Ye; 13c); proximal segment of
labial palpi narrow, 2 » longer than wide (Figs 92;
TOR) . ts zs adored Pea rere) eB os 6
Gills linear (Fig. Me)s proximal sexment of labial
palpi broad, 1,67 longer (han wide (Pig. Weds
prostheca of teh mandible robusl with serrated apex
(Fig. li); dorsal abdominal markings irregular black
and white (Fig. Ide). N. fuseula
5a(4)
Sh
341
6a(5) Dorsal abdaminal marking with a broad median (ight
stripe (Fig, Idd); labrum rectangular, 2 wider than
long (Fig. 9d); femora with lew fine setae (Fig, 9a)
N. [rcenspicua comb. fy.
Dorsal abdominal marking wilh narrow light regions
on sezments 4-10 (Fig. 14; labrum almost square
(5 ~ wider than long (Mig. 13d); femora lined with
numerous Jong fine setae (Fig. 13a)
N_ pilosa sp. Way.
Apex of gills with each lamella trifurcate (Fig. 2c)
. Atalophlebia australis
Apex of gills multifurcale with more than three
tracheal filamenrs (Fig. 4e) 22220 22 ee ee 8
Ninth abdominal segment with only one backward
pointing, spine (Fig. 14c); no dark markings an legs
(FR BB) 6 oa Be ence waco apace bencerph Al. auratus sp, mov.
Ninth abdoniinal segntent with two backward
pointing spines (Vig. 14b); femur and (ibja with broad
bands of black (Fig. 4a) A. australasica
Four pairs of gills present, first pair elytriform,
covering last three pairs. Postero-lateral margins of
abdominal segments produced into backward
pointiig spies (Fig. 28g)
Jeers att Siphlonuridae: Tuspranophlebia
Only one species in South Australia: Jasimanophilebia
Sp).
Seven pairs of gills present, postero-lateral margins
of abdominal sevinerits ror produced into backward
pointing spines Baetidae 1)
Gill lamellae double of abdominal segments 1-6,
single on 7th (Fies 26e; 27e); hind wing sheaths absent
Cloean SI
Gill lamellae single on abdominal segments 1-7 (Pigs
Qi; 24d); hind wing sheaths present
H1a(10) Paraprocis with 27-30 spines (Fig. 27d); maxillary
palpi with terminal spines on distal segment (Fig.
28f); without definite abdominal colour patteri. .
Cloeon parddieniensis sp. nov,
Paraproets with 17-22 spines (Fig. 26d); manillary
palpi without terminal spines on distal segment (Fig.
261); abdominal. pattern as in Pig. 260
6b
7Ta(4)
7b
pew penne
9a(1)
Yb
pine teet cw tao tan de
10a(9)
10b
Ib
CC. Sluviariile
(2a(10) Labrum with deep median V-shaped coneavity with
a tooth on each side of the lateral margins of the
coneavity, near the apex (Pig. 24¢); tarsal claws very
long and slender (Fiz. 24a),..,. es al ee
Ceniroplilim elongaltuni sp. Hoy.
Labrum rounded, with shallow U-shaped concavily
(Fig. 21)); tarsal claw short (Pig. 211) Beetis sarar
FAMILY LEPTOPHLEBITDAE
Peters and Edmunds (1964, 1970) recorded the
systematic history of the family and. listed
characterizations based on Ethiopian and Eastern
Hemisphere material,
GENUS ATALOPHLEBIA Eaton 1881
Burmeister, 1839; 800 (In Buetis); Pictet, 1843; 189-19)
(in Baetis) B ausiralasicu; Walker, 1853: 538 (In
Ephemera) EF. australis; 1853: 559-561 (In Baetis) B.
australasica, 8 costalis; Eaton, 1871: 78-81 (In
Leptophlebia), \88l: 193-194 (Type species Al, australis),
1884; 83-91; Ulmer, 1908: 40-46; 1916; 2-17; 19L9: 16-23;
Needham & Murphy, 1924; 34-36; Tillyard, 1926: 63-64;
(934: 1-1; 1936; 30-49) Harker, 1950: 8-17; 1954:
et eee ee eee
342
243-252; 1957: 63-68; Kimmins, 1960; 294; Riek, 1970:
239; Tsui & Peters, 1975; 542-544,
ivpe Species: Atalophlebia australis (Walker).
Detailed characteristics of the genus are given in
Suter (1980),
Alalophilebia australis (Walker) 1853
Ephemera australis Walker, 1853: 538; Lentophlebia
australis Eaton, 1871: 78; Atalophlebia australis Eaton,
1881; 193-194; 1884; 86; Tillyard, 1934: 1-16.
This species was fully described by Tillyard (1934)
in a study of the type species of Asa/ophlebia.
Therefore much of the detailed description is
unnecessary, bul measurements and ratios have been
included. A transparency of the genitalia of the
lectotype designated by Tillyard (held in the British
Museum of Natural History) has been examined, but
the actual specimen has not been seen, The following
description is based on South Australian representa-
tives, and includes measurements mot recorded by
Tillyard.
Male Imago
x SD oA Range
Body Length 10.69 1.38 [4 &.80-12,50
Notal Jength 3.65 0.39 18 3,20- 4,20
Mesonotal Width 1.93 0,27 15 1.52- 2.28
Pronotal Widih 1.50 0.46 5 1.24 1.64
Fore Wing Length 11.84 0.87 9 9,93-12.48
Hind Wing Length 3.45 0.37 10 2,52- 3,72
Cerci Length 28.00 2.86 § 25,00-32,.60
Terminal Filament Lengthy 23.35 1.40 4 22.00-25.00
Thorax: Fore leg femur length 1.18 » middle leg
fernur length, and 1,07 « hind leg femur length. Ratios
of leg segments (Note; in middle and hind legs, second
ratio is tibia + Ty length: femur length): fore leg
1.00 2 1.34: O11: 0.51 20.44: 0.34: 0,21 (2.34 mm);
middle leg J.00:1,00;-+: 0.11: 0.10 30,09 : 0.19
(1.99 mm); hind leg 1.00 : 1.04 1 - + 0,10 10,09 > 0.08 ;
O17 (2.18 mm), Mesuste: num: basislernum length 1.13
» width, 0.62 * furcasternum Jength, fureasternum
length 0,77 x width, posterior margin with a short
Iriangular indentation, lateral marvins of fureasternum
strangly sclerotized and dark brow,
Wings: Fore wing 2.75 « longer than wide. Hind
Wing (Fig, 1b) 1.62 » longer than wide.
Abdomen: yellow with dark brown markings
dorsally (Pig, |h). Segments 8, 9, LO yellow-brown,
ventrally yellow with light brown markings (Fig. li).
Genitalia: (Figs Ie; d) forceps yellow-brown. Penes
broad, with a V-shaped median indentation, ventral
sperm duct openings obvious (Figs 7a, b),
Mature Male Nymph (Fig. ta)
¥ SDP on Range
Head Width 2.22. 0.06 7% Z.42- 2.28
Notal Length 3.2) 0.07 7 3,12- 3.34
Pronotal Width 3.14 (LiR 7 1.80. 2.30
REC, 5, AUST. MLS. 19 (17): 339-397
Noverther, 1986
Mesonotal Width 2.42 0.09 7 224 2.48
Cerci Length 16.03 0.86 4 15,05-17_10
Terminal Filament Length 13.54 0.96 45 12.31-14.54
Thorax: Pronotum width 0.96 » head width.
Mesonotum 109 « wider than head, Legs (Fig, 2a)
yellow-brown with brown bands on cach segment.
Tarsal claws short and curved with 20-25 peg-like
ventral denticles (Fig. 2b), Remora of lore.and middle
leg equal in length, hind leg longest, 1,12 * fore femur
length, Ratios of leg segments: fore leg 1.00 : 1.05 + 0.53
(1.88 mm); middle leg. 1,00 : 0.97 : 0.43 (1.90 mm); hind
leg 1.00 ; 1,02 + 0.43 (2.10 mm). Femur length to width
ratios: tore leg 3.16, middle leg 3.26, hind leg 3.54,
Sternum: prosternum width equal to anterior width of
mesobasisternum. Mesosternum: basisternum length
equal to or slightly longer than furcasternum, sterna-
costal suture present. Metasternum) basisternum 4-5
« wider than long, widih slightly less than
mesofurcasternum.
Abdomen: (Fig. 14a) posterior margins of tergites
with large singular spines, with smaller spines basally.
Cerei well developed, longer than terminal filament.
Gills, seven pairs, each consisting of a pair of lamellae
with three tracheal filaments per lamella (Fig, 2¢),
Mouthparts: labrum (Fig. 2d) 2.78 x wider than
long, mid-anterior indented, rugose, with 4-5 rounded
tubercles (Pig, 2e). Left mandible (Fig. 2h); outer
incisors with four teeth, inner with three, prostheca
robust (Fig, 21), Right mandible (Fig. 2)); outer incisors
With three apical teeth, with five serrations on inner
margin ol third tooth, inner incisors with two teeth,
inner lateral margin with up to five spines, prostheea
Jong and slender with four spines along length (Fig.
2k). Hypopharyny (Fig, 2f). Maxillae (Fig. 21):
proximal segment of palp 2.24 » longer than wide,
segment ratios; 1.00: 1.22; 0,66 (0.32 mm). Labium
(Fig. 22); proximal segment of papl 1.81 + longer than
wide, segment ratios; 1,00: 0.80 : 0.60 (0.37 mm).
Female lniage
Colour similar to male, ninth abdominal sternite
with deeply incised posterior margin (Fig, le), seventh
sternite produced posteriorly, slightly hooked when
viewed laterally (Fig. if), Fore legs shorter (han in male.
Sternum broader than male, mesobasistcroum length
O.85 « width, mesofureasternum 0.65 » width.
Subimagzo
Black to dark brown, wings shaded grey with hyaline
lambda (A) marking complete (see Tillyard, 1934).
Female Nvwinph
Similac to but larger than male, lackine dorsal
compound eyes.
EPHEMEROPTERA OF SOUTH AUSTRALEA yay
Diagnostic Characteristics
1. Genitalia of male imago (Figs ie, d and Figs 7a, b).
2. Distinctive yellow-brown (orange) colour described
by Walker (1853) and Villyard (1934) as “ved”, and
“rufopiceous above; venter dull light burnt-umber,
approaching, rusty brown” by Eaton (1884),
Distinctive patiern of subimayo,
_ Nymphial gills with uifureate filaments (Mig. 2c).
_ Form and shape of mandibles, incisors and
prosiheeae (Figs 2h, 1, j, kK).
to el
History and Discussion
In 1853 Walker briefly described Liphemera australis
{rom Dr Hooker's 1842 collection trom Tasmania, This
species was later placed in the genus Lepraphlebia
(Eaton I871) and subsequently (Earon 1881) was
designated the genolype for the new genus
Atulophiebia. Walker referred to the “red” abdomen,
and Eaton referred to rufo-piceous coloration, “an
unusual character for this wenus” (Tillyard 1934),
Tillyard's extensive descriptions ol the male imago,
subimago and nymph of A. ausrra/is enable relatively
easy recognition of all stages of this species, but the
coloration in the living material differs from that of
dry pinned specimens. Newly caught and alcohol-
preserved adults have a distinctive Orage colour, not
red, as is {he case in dry specimens.
Material Bxanined
SOUTH AUSTRALIA. South East: Brown Lake,
Bakers Range Drain, Eastern Division Diversion Drain,
Mi. Hope Drain, Reedy Creek Drain, Sutherlands
Drain and Valley Lake. Mt, Lofty Ranges; Marne R.,
Para R., Torrens R. Southern llinders Ranges: Back
Creek, Julia Creek, Rogky River.
VICTORIA, Clunes, larango Dam, Neerin, Lake west
of Harrow, L, Hatlah, L. Wendouree, Ballarat,
Konongwootong Reservoir, Rocklands Reservoir,
Surrey R,
TASMANTA. Break O'Day R,, Lagoon of Islands, L.
Leake.
Atalophiebia australasica (Pictet) 1843
Bueris australasicu Picter, 1843: 89-190; Walker, 1853:
559: Leplaphlebia australasica Gaton, W871 78-79;
Alalophiebia ausiralasicu Baron, 1884; 86-87; Ulmer,
1916: 2-3: Harken 1YSQ0r 28; [954: 248-249.
Male linase
at SD on Runge
Body Length) 10,10 0.86 26 8 40-11.46
Notal Length 3.13 O<:.24 8 2.84 3.60
Fronotal Width 109 O10 8 2.84- 3.60
Mesonotal Width V4 0.13 8H Lidde 1,80
Tore Wing Length 10,33 1.10 Tb 9.23-12.83
Hind Wing Length 3.2 O41 1 2.40- 3,96
Corel Lenurh 25.20) 2.72 #R 24.00-31.00
Head: dark brown to black. Dorsal compound eyes
dark red-grey, lateral eyes dark grey.
Thorax: shiny black, Pronotum narrower than head.
Mesonotum without markings. Legs: fore legs black
with two darker bands on femur, one mid-, ane distally,
T, partially fused to tibia. Middle and hind legs
shorter than fore leg, light yellow-brown with two bhick
bands on femara, one mid, one distal, [ibiae with one
proximal band, T, fused to tibia with joint apparent,
Fore leg femur length 1.20 » middle leg femur lenguh,
and 1.06 hind leg femur length, Ratios of leg
sepments: fore leg 1.00 + 1.39: 0.13 : 0.53 = 048-0 39 |
0.20 (2.26 mm); middle leg 1.00: 0,93 ; 0,08 + O11 :
0,09 : 0.08 2 0.16 (88 mom); hind lee 1.00; 1.01: 0.08 :
0.09: 0.08 :0.07 015 (2.13 mm). Mesostermim:
basisternum length 1.12 » width, 0.66 » fureasternum
length, lateral margins aileriorly expanded, posterior
margin truncated, furcasternum length 0.71» width,
posterior margin with a triangular indentation.
Wingss fore Wing (Fig. 3a) 2,77 » Jonger than wide.
pterostigmal region brown, costal and subcostal cross-
veins broadly shaded with black, bulla in Se
surrounded by black marking. Hind wing (Fig. 3b) 1.60
« longer than wide, costal space with 1-3 proximal and
6-10) distal cross-veins,
Abdomen: light brown with darker markings
dorsally (Fig. 3h) ventrally grey with light patches on
sexments 3-9 (Fig. 3i). Cerei stout, black-brown with
the last 3-5 mm bulf, lerminal filament absent,
Genitulia? (Figs 3c, dj. penes broad at base, with
concave lateral margins and bulbous distal region,
fused, centrally, giving a triangular apes (Pigs Te. cl).
Mature Male Nvinph (Fig, 4b)
x SD on Range
Head Width 2.38 Git 9 2,2k- 2.60
Notal Lenuth 341 O41 9 3.H0- 4.372
Pronotal Width 744 015 9 2.28 2.76
Mesonotal Width 2.70 0.22 9 Q,dk- 3.12
Cerci Length 12.74 — 2 12.)4-13.34
Terminal Filament Length (2.82 0.35 3 12.42-13.40
Head: dark brown-black, Dorsal compound. eyes
red-black, lateral eyes black, Antennae yellow-brown,
longer than head ts wide.
Thorux: dark browo with pale mid-longitudinal line.
Pronotum wider or narrower than head (geographi-
cally variable). Mesonotum L.1t = wider than head.
Legs yellow brown with black bands on each seemenl,
femora with two bands (one mid-, one distal), tibiae
with fwo bands (one proximal, one distal), farsi with
a broad band covering proximal half (Pig. 4a), Pore
femur 1.05 » longer than middle femur, hind femur
1.15 ~ longer than fore femur. Tarsal claws short,
eurved, with 15-20 peg like denticles (Fig, 4b), Ratios
of leg seements! fore leg 1.00 : 1.02 2 0.50 (2.24 mm);
middle lez 1.00 : O98 + 0,37 (2.21 mm); hind leg 1.00 =
1.06 : 0.36 (2.42 mm). Pemur length to width ratios:
S44}
fore leg 3.18, middle leg 3.42, hind leg 3,41,
Mesosiernum: basisternum length equal to
furcasternum Jength, sternacostal suture present,
Metasternum> basisternum 3.7-5 » wider than long,
width equal to width of mesoturcasternum.
Abdamen: brown, patterned (Fig. 14b), lateral
Mange of segment 9 with two spines, outer largest,
inner rounded; posterior margin of tergum with spines
arranged singularly. Cerei well developed, terminal
filament longer than lateral filaments. Gills: multi-
furcate (Fig. 4c), each gill consistitig of a pair of
lamellae with 8-20 tracheal filaments per lamella.
Mouthparts; labrum (Fig, 4d) 3-41 » wider fhan
long, mid-anterior margin indented, rugose (Fig. 4e).
Left mandible (Fig. 4h): outer incisors with three large
(veth, inner with three large teeth, prastheca robust,
crenulated with 6-8 blunt teeth (Fig. 4i). Right
mandible (Fig. 4j): outer incisors with Five feeth, inner
incisors with two teeth and four small spines on inner
lateral macein, prostheca, slender, with crenulations
on Outer margin of 2-4 tubercles, distally with a long
spine (Fig. 4k), Hypopharynx (Fig, 4f), Maxillae (Pig.
41): proximal segment of palp 2.80 « longer than wide,
segment ratios, 1,00; 1.19 + 0,69 (0.41 mm), Labtum
(Fig. 4g): proximal segment af palp long and broad,
1.97 « longer than wide, segment rating 1.00 + 0.67 :
0.68 (0,42 mm).
Female Imago
Larger than male, similar colour and dorsal markings,
9th abdominal sternite with shallow incision on
posterior margin, 7th sternite produced posteriorly
forming a bulbous projection (Figs 3e, f). Fore legs
shorter than in male, sternum and thorax broader than
male, mesobasisternum length of O81 » width, 0.63
« fureasternum length, furcasternum length 0.6]
width. Terminal filament present.
Sudprage
Dull, black-brown, wings uniformly shaded arey, no
“lambda” pattern, terminal filament present,
Diagnostic Churacteristics
1. Genitaha of male imago (Figs 3c, d; 7c, d).
. Lack of terminal filament in adult male,
. Subimaga with uniform grey wings.
. Nymphs with multifureate tracheal gill filaments
(Fig. 4c).
. Oth abdominal segment with two spines on the
posterolateral margin.
6. Form and shape of mandibles, incisors and
prosthecae (Figs 4h-k),
Bet
History und Discussion
Atalophlebia australasica, originally described by
Picter (1843), has been redescribed by Eaton (1871,
REC. &. AUST. MUS. 19 (17): 339.397
Navermber 1986
1884), by Ulmer (1916) and Harker (1954). The
description given by Piclet was not extensive enough
fo enable recognition of this species as. more Australian
Species were described. Eaton (1871, 1884) redeseribed
this species and illustrated the wings and geniralin.
Ulmer (1916) designated material from northern
Queensland as A. australasica, and described (he male
and female imagos, and the subimago. Ulmer noted
differences in the venation of the hind wing of these
specimens when compared with Eaton's (884)
iustration, but did not comment on the apparent
differences in the genitalia as illustrated by Eaton,
Harker (1950) recorded that Ulmer (1919) had placed
A. austtalusicad as a synonym of A. vastalis
(Burmeister). In 1954, Harker noted, after examining
material in the British Museum, that A, australasica
as designated by Eaton was distinctly different from
AL. castalis as determined by Tillyard, and redeseribed
all stages of the species,
Although the male imago description is of the
holotype (as inferred by the statement “fore legs are
missing in holotype and paratypes”), Harker included
a record of a well-developed terminal filament. Pictet’s
(1843) description included only one caudal Silament
measurement, consistent with the accompanying illus-
tration, and of descriptions of species lacking the
terminal filament, Eaton (1871, 1884) also included
only one measurement. Ulmer (1916) recorded only the
lateral cerci “Die Schwanzborsten (2 beimc) sind
schwarzbraun ...” bur recorded the presence of the
terminal filament in the male subimago. Harker (1954)
also noted that material examined from Mt Gambier
(from Tillyard's 1934 collection) was. consistent with
her recognition of this species. Adult males of ¢1.
austrdlasica from south-east Sonth Australia and the
Fleuricu Peninsula lack the terminal filament, consis-
tent with the type description. Harker's record
mentions no variability of expression of the terminal
filament (as described for A. wustrolis by Tillyard 1934),
Therefore, Since subsequent collectians, as well as the
fype material all lack this filament, its presence ay
indicated by Harker must be suspect,
A comparison of kigs 3¢, and 7e, with the illustration
al the A, australasica genitalia by Harker (1954) shows
hittle resemblance. The South Australian. material of
iis species, examined in this study, was initially
considered different from previously described species.
However, a comparison of an air-dried specimen and
a critical-point dried specimen revealed (wo different
penes characteristics. The air-dried specimen closely
resembles the illustration presenied by Harker, from
a dry, pmned preparation. The lateral lobes of air-dried
specimens curl in towards the mid-line, producing a
long narrow structure, The critical-point dried
specimens represent more closely the living, or alcohol-
preserved, characteristics of the genitalia, with the
lateral lobes of the penes maintaining their lateral
pasitian,
EPHEMEROPTERA OF SOUTH AUSTRALIA
Material Examined
SOUTH AUSTRALIA. South East: Cress Ck, Deep
Ck, Bight Mile Ck, Hitehcock Drain, Jerusalem Ck,
Mt. Lofty Ranges: Black fellow Ck, Brownhil Ck, Bull
Ck, Cudlee Ck, Currency Ck, Dam at Carey's Gully,
Dam at Ashton, Deep Ck, First Ck, Pourth Ck, Little
Para R., Morialta Ck, Onkaparinga R., Sturt R.,
Torrens R., Wakefield R. Fleurieu Peninsula:
Carcickalinga Ck, Coolawang Ck, The Deep Ck,
(Delamere), Finnis R., Gold Digging Swamp,
Hindmarsh R,, Inman R., Kangarilla Ck, Myponga
Ck, No Where Else Ck, Tookayerta Ck, Yankalilla R.
Kangaroo Island; Breakneck R,, De Mole R,, Middle
R., Rocky R., South West R.
VICTOR/A, Crawlord R,, Eumarella R,, Fitzroy R,,
Glenelg R., Shaw R.
Atalophlebia auratus sp, nov.
Holotype Male naga
Body Length 7.95 mm
Notal Lengitli 2,60 mim.
Pronotal Width 1,04 mm
Mesonotal Width 1.27 mm
Fore Wing Length 7.44 mm
Hind Wing Length 2.23 mm
Cerci Length (9.67 mm
‘Terminal Filament Length 15.57 mm
General colour yellow-brown, fore wings yellow, hind
wings grey.
flead; dark brown. Antennae short, basal segment
0.12 mm long, 1.4 » longer than wide, 2nd segment
0.1 mm long, 2.5 * longer than wide, flagellum
0.24 mm long. Dorsal compound eyes pink-brown,
lateral eyes dark grey, Ocelli dark brown laterally, white
anteriorly.
Thorax: shiny black. Pronolum narrower (han head,
Legs: fore legs long, femur and tibia dark brown, tarsal
segments dark brown, no banding on any segment, Ty
partially fused to tibia, Middle and hind legs shorter,
light brown without banding, larsal segrnents 1-4 with
distal spine, T; fused to tibia, join visible. Fore leg
femur length 1,03 < middle leg femur length, and 0,95
» hind leg femur length. Ratios of leg segrnents: fore
leg 1.00) 1.145 0,09 : 0.40 20.402 0.28 20.19
(1.84 min), middle leg 100 | 0.77; - 2 0,10: 0.08 : 0.16
(1.78 mm); hind ley 1.00 + 0,98 + © 0.08 « O10: 0.07 :
0.17 (1.94 mim). Sternum dark black-brown, Prasternum
iriangular, Tonger thon wide. Mesosternum;
basisternum length 117 « width, 0.78 « fureasternum
length 0.89 « width, lateral margins of median
longitudinal invagination divergent, posterior margin
concave (Fig. Se),
Wings: fore wing (Vig, 5a) tinged throughout with
yellow-brown, veins yellow-brown, pterostigma darker
than rest of wing, Cubital region shaded grey, length
345
3,35 =» Width, pterostigmatic crass-veins forked in lett
wing, simple, not forked in right wing, cross-veins
present in proximal half of costal region, Hind wing
(Fig. 5b) grey-brown, darker than fore wing, shaded
completely, length 2.06 « width, costal hump not large,
costal space with nine cross-veins.
Abdomen: yellow-brown dorsally, lighter yellow
ventrally, dorsal tergites with dark brown-black
markings laterally, and light mid-longitudinal stripe
along, all segments (Fig, 51), Sternites yellow-grey, with
light red-brown circular markings on mid-line (Fig. 5g).
Cerci long, terminal filaments well developed but not
as long or robust as cervi.
Geniialia (Figs 5c, d; Te, 0: foreeps with long
proximal segment, broadest proximally, narrows half
way along length, 2nd segment short, globular, distal
segment longer, ovoid, rounded apically, Penes broad
at base, tapering, towards apex with apices held vlose
together, curving upwards in profile.
Mature Female Nymph (Pix, l4e)
Body Length 9,20 mm
Head Width 2.03 joan
Notal Length 2,22 min
Pronotal Width 1.90 min
Mesonotal Width 2,19 mm
Cerci Length 15.20 min
ferminal Filantent Length 14,66 mm
General colour brown.
Head: brown. Lateral eyes black, ovelli black,
Tentorial bady almost square, length 0.95 « width.
Antennae 4.8 mm long.
Thorax: pronotum brown without markings, width
0.95 «x head width, 2 stout spine setae on anterior
margin, no setae on lateral margin. Mesonotum brown,
broad, width 1.08 » head. Legs light brown without
banding (Fig, 6a). Tarsal claws with 31-35 peg-like
denticles (Fig. 6b), segment ratios: fore leg 1,00 ; 0,98
/049 (1.8L mm); middle leg 1,00: 0.96: 0.48
(1.81 mm); hind jeg 1.00 : 0.96: 0.47 1.97 mm), Fenvur
length to width ratios; lore Jeg 3.77, middle leg 4.11,
hind lee 4.48. Sternum: prosternum, length 0.73
width, Wider than anterior margin Of mesosternurm,
Mesosternum: basisternum rectangular, width 0.68 <
length. Metasternum: basisternum short, width 5.7L «
length.
Abdomen: brown dorsally with black patches on
lateral flanges. of segments 1-7, segments. LS otherwise
brown without markings, segments 6, 7, 8, 9 with
central hight stripe, segment 10 light brown (Fig, 14e),
Segment 9 with a single postero-lateral spine Cerei atid
terminal filament well developed, cerci longer. Gills:
multifureate, with numerous fine tracheal lilaments of
cach lamella.
Mouthparts, \abrum (Fig, 6c) length 0.40 « width,
anterior margin with median concavity lined with five
rounded tubercles (Fig. 6d). Left mandible (Fig, 6g):
346 REC. 8S. AUST. MUS, 19 (17): 339-397
outer incisors with three apparent (four actual) teeth;
inner incisors with three teeth, prostheca robust with
8-10 pointed teeth on external margin (Fig. 11h). Right
mandible (Fig. 67): outer incisors with three teeth and
a fourth shoulder-like ridge on third tooth, two small
tubercles on mesal margin, inner incisors with three
teeth, inner most with two small tubercles laterally,
prostheca long, narrow with one small external spine
and two terminal spines (Fig. 6j). Hypopharynx (Fig.
6e). Maxillae (Fig. 6k). Segment ratios of palp
1.00 : 1.10 : 0.67 (0.30 mm). Labium (Fig. 6f): proximal
segment of palp broad, length 1.61 » width, distal
segment triangular, segment ratios 1.00: 0.80 : 0.62
(0.38 mm).
Female Imago
Reared in laboratory. Wing and body coloration
resemble male, body robust, filled with eggs. Ninth
abdominal segment with a deep ventral cleft (Fig. 51);
sternite of seventh abdominal segment slightly
produced posteriorly (Fig. 5j).
Subimago
Similar to male imago, wing colour dull yellow-grey,
without marking.
November, 1986
Male Nymph
Smaller than female, head with red-brown compound
eyes.
Diagnostic Characteristics
1. Genitalia of male imago; shape of forceps segments
two and three and shape of penes (Figs Sc, f; 7e, f).
. Wing coloration in both fore and hind wings.
. Lack of banding on legs of adults.
. Multifurcate gills of nymphs.
. Only one postero-lateral spine on abdominal
segment 9.
6, Shape of mandibles, incisors and prosthecae (Figs
6g-j).
7. Lack of banding on legs of nymphs (Fig. 6a).
8. Dorsal colour pattern of nymph and adults (Fig.
14c; and Fig. 5f).
Type Locality
ty Bh Wo to
Bakers Range Drain, west of Penola, South
Australia. Grid Reference on | : 250 000 map series,
Penola Sheet: 357393. Collected 22 November 1977 by
D. N. Suter and P. J. Suter.
TABLE 1, COMPARISON OF QUALITATIVE CHARACTERISTICS OF ATALOPHLEBIA AUSTRALIS,
A, AUSTRALASICA AND A. AURATUS FROM SOUTH AUSTRALIA
Character A. australis
Male Imago
Body colour
Fore wing colouration
Yellow-brown
Pterostigma only
(yellow)
Clear
Separate apically
Hind wing colouration
Penes
Legs Femur banded
Fusion of tarsal
Segment 1 of fore leg Partial
Fusion of tarsal
Segment | of middle Fused
and hind legs
Terminal filament
Nymph
Present-absent
Dorsal eye Colour oo Sepia
Lateral eye colour Black
Legs: banding Femora 2 bands
Tibiae 2 bands
Tarsi 1 band
Postero-lateral spines on Single
abdominal segment 9
Abdominal tergite spines Large singular spines with
smaller basal spines
Gills Trifurcate
Left mandible
Incisors Outer 4
Incisors Inner 3
Prostheca Robust-serrated
Right mandible
Inctsors Outer
Incisors Inner 2
Prostheca Long, slender with
4 spines
A. australasica A, quraius
Yellow-brown
Totally yellow
Black-brown
Pterostigma only
(brown)
Clear Grey
Fused apically with lateral Fused apically without
lobes lateral projections
Femur banded No banding
Partial Partial
Fused Fused
Absent Present
Red-black Pink-brown
Black Black
2 bands Absent
2 bands Absent
1 band Absent
2 spines Single
Single spines with smaller
basal spines
Singular spines
Multifurcate Multifurcate
3 4
3 3
Robust-serrated Robust-serrated
3
3
Long, narrow
mw
Slender-serrated
FPHEMEROPTERA OF SOUTH AUSTRALIA
Type Specimens
A auratus is only known from the type locality,
Holotype male and paratypes are placed in the
Museum of Victoria. The wings and legs of the
holotype male are mounted on slides, and the
mouthparts, legs and gills of the female nymph are alsa
mounted on slides, The genitalia and body of the
holotype are maintained in ethanol.
Tipe Habitat
Near (he source of Bakers Range Main Drain, which
33 a man-made drain, draining the swamps of Bakers
Range in the South East of South Australia. The water
at the type locality was. evaporating rapidly during
Noveniber 1977, when the type collection was made.
A. ausiralis was also present at this locality,
Ewimology of Specific Epithet
The spevifie epithet auratis (L), ornamented with
gold, refers to the golden coloration of the forewings
which make this species distinct from all other
described Atulophlebia species.
Affinities
In adult characters Lhe wing colarahoen, genitalia and
lack of banding of the legs distinguish this species front
other described species in the genus Alglophlebia, The
nymph, however, resembles the nymph of 4.
ausiralasica in possessing multifurcate gills, Although
smaller than A, austra/asica (a character to be used
with great care) the lack of banding of the legs, the
presence of only one postero-lateral spine on the 9th
abdominal segment, the mandible incisors aad
prosthecae all distinguish A, aurarus from A.
uustralasica. A comparison of qualitative
characteristics which distinguish the three 4ral/ophlebia
species found in South Australia is presented in Table: 1,
GENUS NOUSIA Nayas, 1918
Navds, 1918: 213: 1925; 308; Ulmer, 1919; 20 (In
Atalophlebia), Needham & Murphy, t924: 35-37;
Lesiage, 1931; 52: Traver, 1946: 420; Harker, 1950:
30-32; 1954: 242-243; 1957; 69-71) Riek, 1970: 239;
Peters & Edmunds, 1972: 1411; Tsui & Peters, 1975:
540-542; Pescador & Peters, L985: 91-123.
Tipe Species: Nousia delicuta Navas by original
desiznation, Detailed characterisation of this genus is
piven by Pescador & Peters (1985) and Suter (1980).
There has heen considerable discussion of the validity
of the genus /lra/onella in Australia (sui arid Peters
1972; Suter 1980) and following the work of Pescador
& Petors (1985) the South Australian species are placed
in the genus Nousia, Although differences between the
South American and Australian species exist (ag. eae
structure) until a full revision is performed all
Atalonella species should be considered as belonging
(o the genus Nousia.
347
Nousia inconspicua (Eaton) 1871 comb, nov.
Leptophlebia inconspicua Eatan, 1871: 79-80;
Atalophlebia incanspicua Eaton, 1884 8&7; Ulmer,
1908: 43-44; Tillyard, 1936: 31; Harker, 1950: 28; 1954;
265.
Male Imago
*¥ SD pn Range
Body Length 7.27 0.50 26 6,07- 7.95
Notal Length 2.26 (15 26 2.02- 2.62
Pronoral Width 096 0.09 16 0.84 1.08
Mesonoltal Width 1.19 0.08 26 1.04 1,32
Fore Wing Length 7.25 0.52 26 5.74 8.36
Hind Wing Length 1.43 0.14 26 1.20- 180
Cerci Length 10.60 O.80 11 9.411214
Terminal Filament Length 12.67 1.0L 10) 11.11-14.02.
General colour black, with light transparent regions
between adbomijnal segments, giving a banded
appearance,
Head: black. Dorsal compound eyes light brown,
lateral eyes grey
Thorax: dark brown-black. Legs: lore legs dark
brown, without banding, Middle and hind legs shorter,
light brown, without bands. T, fused to tibia, suture
apparent. Ratios of leg segments: fore leg 1.00: 1.46 :
0.14: 0.48 : 0.452033: 017 (2.13 mm) middle leg
1.00 :1.07 ; 0.06 = 0,08 } 0.08 0,072 0,16 (1.44 mm);
bind leg 1.00: 1.13 : 0.06: 0.08 2 0.08 : 0.07 : 0.16
(1.49 mm),
Wings: fore wings (Fig. 8a) 3.29 » longer than wide,
plerostigmal region slightly opaque, With simple crass-
veins in distal 1/3 only, proximal region of subcostal
space without cross-veins, distal region with very fmint
cross-veing, cubital and anal regions with few cross-
veins. Hind wing (Fig, 8b) 1.55 » longer than wide,
4-5 cross-veins in distal region of costal space, absent
in proximal hall, subeostal space wiih 3-5 cross-veins.
Abdomen: black with light central marking dorsally
(Fig. 8h), all segments light brown vevtrally (Fig. 81).
Genitalia (Migs 8c, d): distal segment of loreeps
globular. Penes broad, fused along entire length,
extending beyond constriction of proximal segment of
forceps, ventral lobes triangular with base separate
anteriorly, sperm ducts open on mid line (Figs 15a, b).
Mature Male Nemph (Vis. td)
*¥ SD on Range
Head Width 1.59 0.04 13 L.S4-1.70
Notal Length 1.95 0.08 13 1.80-2.12
Pronatal Width 1.50 (0.08 13 1.38-1.56
Mesonoral Widih 1.58 0.07 13 1.50-1.72
Cerci Length 9.92 — 1 =
Terminal Vilameor Length W777 — 1 _—
Flead; dark brown, Dorsal compound eyes dark
reddish brown,
Thorax: legs brown, no banding (Fig. 9a), Ratios of
leg segments; fore leg 1,00: 0,92: 0.52 (1.39 mm);
middle leg 1.00 : 0.92: 0.44 (1,39 mm); hind leg 1.00;
348 REC. S, AUST. MUS. 19 (17): 339-397
0.98 : 0,36 (1.49 mm), Femur length Lo width ratio: fore
leg 3.17, middle leg 3.20, hind leg 3.35,
Abdomen; brawn dorsally with a light stripe down
mid-line (Fig. 14d), posterior margins of terga with
large singular spines, with smaller spines between them,
Gills lanceolate (Fig. 9c), lamellae with numerous
tracheal branches,
Mouthparts: labrum (Fig, 9d) 2. « wider than long,
median cavity without obvious rounded denticles
(Fig. 9e). Left mandible (Fig, 9h); outer incisors with
three teeth, inner with three teeth, prostheca long and
slender, apically with lwo rounded teeth (Fig. 9i). Right
mandible (Fig. 9j): outer incisors with three teeth, inner
with 2-4 teeth, prostheca simple, narrow with one apical
spine (Fig. 9k), Hypopharynx (Fig. 9f), Maxillae (Fig,
9): proximal segment of palp 2.06 » longer than wide,
segment ratios 1.00: 0.92: 0.80 (0.22 mm), Labium
(Fig. 9g); proximal segment of palp 1.98 » longer than
broad, distal segment with 3-4 short apical spines,
segment ratios) 1,00 : 0,79 + 0.64 (0.35 mm).
Female Imago
More robust than male, uniform brown. Fore wigs
with cross-Veins along entire costal and subcostal
spaces. Hind wings with cross-veins in radial, median
and cubital sectors.. Anal plate with a deep V shaped
incision (Fig, 8e), na ovipositor (Fig. Sf), Ege, oval,
0.12 mm long, 0.09 mm wide with a polar cap of twa
tings of tubular projections on each apex (Fig. |6a).
Subimagvo
Black with uniformly grey wings,
Diagnostic Characteristics
|. Genitalia: penes shape, broad, fused, with triangular
ventral lobes (Figs 8c, d and Figs. 15a, b).
2. Lack of cross-veins in proximal regions of C and
Sc spaces of fore wings (Fig. 8a).
34. Egg morphology, polar caps with 2 tows of tubular
processes (liz, 16a).
4. Nymphal gills lanceolate (Fig, 9¢),
5. Mandibles, incisors and prosthecae shape (Figs
9h-k),
6, Proximal segment of labial palp long and narraw
(Fig. 9g).
7. Dorsal white stripe on abdomen (Fig, 14d),
History and Discussion
Untd this study NM. inconspicua ad the distinction
of being the only species described from South
Australia, Eaton (1871) described and placed it in
Leprophlebia, noting the bluck and White patrerning
of the abdomen. His ilhistration of the genitalia differs
from (he scanning eleciromicrograph (Figs 1a, b) as
he showed separate penes lobes. This separation of the
Navember, 1986
two lobes of the pénes is observed with air-dried
specimens, and may occur with slide-mounted material.
Living, critical-point dried, and alcohal-preserved
Specimens possess genitalia with the two lobes held
close together as illustrated in the micrograph. The type
locality of this species is Adelaide, but the river or
stream fram which the holotype was collected is
unknown. In 1884 Eaton placed Nousia inconspicua
in the new genus Afalophlebia where it has remained
until the preserit stady, Ulmer (1908) added to Eaton’s
description from material from the South-West of
Western Australia. He included illustrations of male
genitalia, both wet-preserved im alcohol and dried,
showing the separation of the paired penes lobes in the
dried preparation. From the illustrations of the wings
(the presence of cross-veins in the proximal half of the
C and Sc regions of the fore wing) and genitaha it is
difficult to know if (he species described by Ulmer is
N, incenspicua or a different species.
Tillyard (1936) recognised two groups within the
genus Alalophlebia, and he included N. incanspleua
in the group with the smaller species ie, “smaller
species, expanding from one-half to three-quarters of
an inch (fore wing from 5 to 8 mm long)". Since this
species is not recorded in Tasmania, no description was
given, and in fact it is only referred to in the adult key.
All the other species from the smaller sized group were
later placed in the genus 4/g/onella by Harker (1954),
but \, Jiconspicua was left in Atalophilebia.
The nymphs and adults of this species, as described
above, have characteristics which distinguish them from
the genus Aralophlebig, but are consistent with the
generic characteristics of Nousiv, Consequently N.
inconspicua is now formally placed in the genus
Nolisid,
Material Examined
SOUTH AUSTRALIA, Mi. Lofty Ranges: Aldgate Ck,
Black fellow Ck, Brownhill Ck, Bull Ck, Currency Ck,
Deep Ck, Fifth Ck, Finnis R., Fourth Ck, Little Para
R,, Marne R., Marialta Ck, Onkaparinga R., Scoit Ck,
Sturi R., Torrens R. Southern Flinders Ranges: Back
Ck, Neetar Brook Dam, Rocky R,, Schumacher Ck,
Skillogalee Ck, Spring Ck, Wakefield R, Fleuricu
Peninsula; Anacotilla Ck, Carrickalinga Ck,
Coolawang Ck, The Deep Creek (Delamere), Gold
Digging Swamp, Hindmarsh R.. Kangarifla Ck,
Myponga Ck, Yankalilla R. Kangaroo (sland:
Breakneck R., Cygnet R., De Mole R., Grassy/Sheep
Ck, Middle R., North East R., Rocky R., South West
Bay R., South West R., Stunsail Boom R,
Nousia fuscuta (Tillyard, 1936)
Alalophlebia fuscula Tillyard, 1936: 44-47;
Atalophlebia fuscula Harker, 1950; 28; Atalonella
Juseula Harker, 1954: 242-243, 264; Scholes, 1961:
31-33.
EPHEMEROPTERA OF SOUTH AUSTRALIA
Male Imago
y¥ SD on Range
Body Length 6.58 0.32 23 $§,82-7.01
Notal Length 2.02 O41 2b 1.B4- 2.20
Pronotal Width O87 0.06 LO 0.74- 0.92
Mesanotal Width 1.02 0,09 2 Q),82- 1.12
Fore Wing Length 6.41 0,28 22 5.90- 6.97
Hind Wing Length 1.29 0.09 23 1,12- 1.40
Cerei Length 10.32 0.62 12 9,23-11.11
Terminal Filament Length 11,67 0.76 8 10.77-13.00
Head: black, Dorsal compound eyes, brown-grey,
Thorax: black. Fore legs black without banding.
Middle and hind legs brown without banding. Ratias
of leg segments: lore leg 1,00 ; 1.29 ; 0.14: 0.49 + 0.44:
0.28 : 0.16 (1.64 mm); middle leg 1.00: 1.15: -: O11:
0.09: 0,09 : 0.15 (1.38 mm); hind leg 1.00: 1.20: -:
0,09 ; 0,08 : 0,09} 0.15 (1.58 mm). Fore leg femur
length 1.19 « middle leg femur length, and 1.04 » hind
leg femur length,
Wirigs: fore wing (Fig. 10a) 3,13 « longer than wide,
plerostigmal region slightly tinged with brown, cross-
veins slanted, simple, costal space proximal (o bulla
with 3-7 very faint cross-veins, subcostal space with 2-4
crass-veins in proximal half, 7-10 in distal half, Hind
wing (Fig. 10b), 1.52 ~ longer than wide, 3-4 cross-
veins in distal region of costal space, absent in proximial
half, subcostal space with 4-6 cross-veins,
Abdomen; black, With brown and light brown
markings (Fig. 10h); ventral pattern (Fig. 101).
Genitalia (Figs 10c, d): distal segment of forceps
globular. Penes lobes widely separated, cylindrical,
constricted near apex, apex rounded, lobes. apparently
sheathed, inner margin wilh a small spine hidden
within sheath, visible in mounted preparations
examined using transmitted light (Fig. 10j), but not in
the Scanning Electron Micrographs (Figs 15c, d).
Mature Male Nvinph (big, 14e)
x SD «# Range
Head Width 1.44 0,06 6 — 1.36-1.50
Notal Length 1.71 0.07 4 1 64-1.80
Pronotal Width 1.28 0.04 4 1,22-1.32
Mesonotal Width 1.33 0.06 4 — 1,26-1.40
Cerci and terminal filamen| danyaged in available specimens.
Head: dark brown, Dorsal compound eyes reddish
brown.
Thorax: pronotum with spine setae on antero-lateral
margins. Lees brown, not banded (Fig. Ma), Ratios of
lez segments: fore leg J.002 0.87 20.55 (1.18 mm),
tniddle Jez 1,00; 0.84:0,39 (119 mm); hind leg
1,00 : 0,91: 0.38 (1.40 mm), Femur length to width
ralios, fore leg 2.74, middle leg 2.82, hind leg 3.11.
Abdomen: colour paliern irregular (Fig. I4e), Spines
on posterior margins ol terga occur either separately
or in pairs. Gills (Fig. Lic); linear, lamellae lacking
tracheal branches, or, if present, very short
a4y
Mouthparts: labrum (Fig. Jld) 2 » wider than long;
median cavily with lour to six rounded denticles (Fig.
Le), Left mandible (Fig. 1h), outer incisors with three
apical (eeth, inner incisors with three teeth, prostheca
broad, robust, outer and apical margin serrated (Fig.
11i)- Right mandible (Fig. 11j): outer incisors with three
apical teeth, inner incisors with two teeth, prostheca
simple, long and narrow with one apical spine (Fig
11k). Hypopharynx (Fig, If), Mavxillae (Pig. 111),
proximal segment of palp TIL « longer than wide.
Segment ratios: 1.00 : 0.71: 0.78 (0.21 mm), Labium
(Fig. 11g): proximal segment of pal 1.57 » longer than
broad, segment ratios; 1.00 2 0.71 : 0.69 (0.29 mm),
Female Imaze
Mare robust than male, wings similar, hind wings. with
more cross-veins in radial, median and cubital sectors.
Anal plaje with a deep V shaped incision (Fig, 10¢),
no ovipositor (Fig, 10f), Exes oval 0142 mmx
(0.09 mm, polar caps with three rings of tubular
processes (Fig. 16b).
Subimazo
Dull black, wings opaque, dark grey,
Diutgnosiic Charaeteristics
1. Male penitalia, laleral lobes of penes widely
separate, small spine on mesal margin of lobes (Figs
10c, d and Fius 15c, d).
2, Cross-veins in proximal region of costal and
subcastal spaces (Fig. 10a).
3. Egg polar cap with 3 coronae of tubular processes
(Fig, 16b),
4, Nymphs have linear gills with few tracheal branches
on lamellae (Fig, le),
5. Mandibles, shape of incisars and prosthecae (Figs
Wh-k),
6. Proximal segment of labial palpi broad (Fig. 112),
7. Abdominal colour pattern irregular (Fig. le).
History and Discussion
Tillyard (1936) deseribed the adults (male and
female), subimago, and nymph of N, fuseuda placing
it in his distinet group of small species of the genus
Aldlophlebia. Warker (1984) recognisd that all the
species Tillyard placed in this second graup ol smali-
sized Species belonged in the genus Afalonella. These
species now belong to rhe genus Nausia, and No fuscula
is farmally included in this. genus,
The male genitalia are distinctive in N, fiyseule, with
a small spine on the mesal margins of the penes lobes,
but the associated nymph in South Australia differs
from the description and illistration given by Tillyard
(1936). The gill illustrated by Tillyard shows a more
lanceolate lamella with numerous tracheal branches,
both shape and tracheation are inconsisient with the
associated nymphs of South Australia. Nymphs of
Tillyard’s description were not found at Tookayerta
450
Creek (the only locality where N, fxscula is present on
the Fleurieu Peninsula), but one of a similar description
to his was recarded from Deep Creek, east of Port
Macdonnell in the South East of South Australia.
Similar nymphs from the Grampians, Victoria, were
bred through and N. pi/osa sp. nov. was the assaciated
adult. Field observations suggest that N, fuscula
emerges before N, pilosa, therefore leaving N. pilosa
nymphs in the stream, This observation was repeated
by the author in the South East of South Australia,
the Grampians, Victoria, and in Tasmaria, The nymphs
of N. fuseula from Tasmania (collected in February
1978 and associaled with adults) were similar to those
in South Australia, From (hese observations it appears
that Tillyard indirectly associated the nymph in the
stream with the flying adults and emerged subimayos
rather than by breeding through the nymphs. and
directly associating these with the resulting adults.
Material Examined
SOUTH AUSTRALSA, South East: Cress Ck, Deep
Ck, Eight Mile Ck. Fleurieu Peninsula: Tookayerta Ck,
VICTORIA. Aire R., Albert R., Beehive Ck, Crawford
R,, Cumberland Falls (Marysville), Darlots Ck, Genoa
Ck, Howqua R., Jimmy’s Ck, Little R,, McKenzie R.,
Met. Zero Channel, Stony Ck (Halls Gap), Tanjil R,,
Tarwin R,, Toorongo Falls (Noojee).
NEW SOUTH WALES. Leatherbarrel Ck, Styx R.,
Wallagaraugh R.
TASMANIA, Break O'Day R., Dee R., George R.,
Great Forester R., Isis R., Macquarie R., Rostrevar Ck.
Nousia pilosa sp. nov.
Holotype Mule
Body. Length 7.50 mm
Notal Length 1.90 mm
Pronotal Width 0.93 nem
Mesonotal Width 1.17 mm
Fore Wing Length 7.67 mm
Hind Wing Length 1,23 mm
Cerci Length 9.05 mm
Terminal Filament Lengrh 12.00 nim
Head: black-brown. Dorsal cyes pink/brown,
Therax: pronotum black, narrower than head, Fore
leg without banding, femur dark brawn, tibia and tarsi
light brawn, Ty) partially fused to tibia; middle and
hind legs with dark brown femora each with one black
band at 3/4 of length, tibiae and tarsi light brown, T,
fused to tibia. Ratios of leg segments: fore leg 1.00 :
1.52: 0.08 : 0.48 2 0.5L: 0.38 : 0.15 (1.92 mm); middle
leg LOO: 1.22:- : 0.07 : 0.07 : 0,07 + 0.11 (1.62 mim);
hind leg 1.00: 1.35 :-:0.09: 0.08 : 0,07: 0,13
(1.67 mini).
Wings: fore wing (Fig, 12a) 3.3 » longer than wide,
plerostigmal region with simple, slanted eross-veins,
proximal 2/3 of costal space without ¢ross-veins,
subcostal space without proximal crass-yeins, distal
REC. AUST, MUS. 19 (17): 339.397
November, 1986
region with seven. Hind wing (Fig. 12b, ¢) 1.52 «
longer than wide; 2 cross-veins in distal region of costal
Space, absent in proximal hall, sub costal space with
3 cross-veins,
Abdomen: dark brown with mid-dorsal light brown
region on segments 2-7, segments 8-10 dark brown (Fig.
12g); ventrally light brown (Fig. 12h). Base of each
segment of caudal filaments tinged with brown giving
a banded appearance,
Genitalia (Figs. 12d, ¢): second segment of forceps
short, ovaid, distal segment globular. Penes fused, apex
with 2 lobes, ventral surface lacking lobes (Figs 12d,
ISe, f),
Mature Male Nvinph (Pig. 14)
Head Width {.20 mm
Notal Length 2,30 mm
Pronotal Width 1.09 mm
Mesonoial Width Lal mm
Caudal Filaments Damayed in type
Head; light brown, light marking in centre af frans.
Dorsal eyes red-brown,
Thorux: pronotum width 0,91 x head width, few
spine setae present on antero-ateral margin. Legs light
brown, without banding, margins of segments lined
with long fine setae (Fig. 13a). Tarsal claws with 12 pee-
like teeth (Fig- 13b), Ratios of leg segments: fore leg
1,00 ; 0.96 : 0.40 (1.70 mm); middle leg 1.00 : 0.95 =
0.3L .77 mm); hind leg 1.00: 1.01 0.29 (1,93 mim).
Femur length to width ratios: fore leg 3.13, middle leg
3.39, hind leg 3,70,
Abdomen; dark brown with light central marking
(Fig, 14f), Gills (Fig. 13c) lanceolate with single
terminal tracheal filament, lamellae with tracheal
branches obvious.
Mouthparts: labrum (Fig. 13d) 1,5 «wider than long;
anterior margins with a median cavity with six rounded
denticles (Fig. 13¢), one row of setae behind median
cavity. Left mandibles (Fig. 13h); outer incisors with
ihree apical teeth, inner incisors with three apical teeth
and two ridges on inner margin, prostheca broad
basally, long and slender distally with three terminal
teeth (Fig. 13i). Right mandible (Fig. 13)); outer incisors
with three apical teeth, inner incisors with two Leeth;
prostheca slender, elongate with a long terminal spine
(Fig, 13k), Hypopharyns (Fig, 131), Mavillae (Fig. 131);
palpi, proximal segment 1.79 sx longer than wide,
Segment ratio 1,00 ; 0.68 : 0.68 (0.27 mm), Labium
(Fig. 13g); palpi, proximal segment 2.26 » longer than
broad, distal segment; apex with four shor( teeth,
segment ratios; 1.00 : 0.77 : 0.49 (0.44 mm),
Female Inagzo
Unknown,
Diuenostic Characteristics
1. Genitalia of male imago (Figs 12d, e and Figs |5e, [),
2. Gills of nymphs lanceolate, similar to N,
inconspicua (Fig. 13¢),
LPHEMEROPTERA OF SOUTH AUSTRALIA 351
3. Lubrum long (width 1,5 » greater than length) (Pig.
13d).
4, Prostheca shape of left and right mandibles (Figs
13i, k).
5, Legs lined with numerous long fine setae (Fig. 13a).
Type Locality
‘Wpe material was collected from Second Wannon
River on the road from Halls Gap to Dunkeld,
Grampian Mountains, Victoria. Grid Reference on
1:250 000 map series Ballarat Sheet; 547396,
Collected 25 November 1977 by D. N. and P. J. Suter.
Type Specimens
Holotype male, and nymphal exuvium are placed in
the Museum of Victoria. The wings and legs of the
holotype are mounted on slides, and the body and
genitalia are maintained in ethanol, The nymphal
exuyiuim is mounted on slides. Three paratype males,
two mature males and three nature female nymphs are
included in the type series. Slides of a nymph, and adult
male jmago [rom Hitchcock Drain, South East South
Australia, are also included.
Type Habitat
The Second Wannon River in the Cirampians
Mountains, is a moderately fast-flowing stream over
cobble-sized rocks,
Liymology of Specific Epithet
The specific epithet pilosa (L) for hairy refers to the
long, fine setae which line the margins of the legs
distinguishing this species from N. inconspicua and N.,
Juseula.
Affinities
In the adult characters N. pilosa resembles closely
N. inconspicua, with the penes lused, and lacking
spination, N. fuscula is distinctly different with
V-shaped penes, and internal spines. Although super-
ficially similar, the penes of N. ineonspicud and N.
pilosa are distinct, with N. inconspicua with obvious
ventral lobes, and separated basal halves of the penes
(Figs 15a, b). N, pilosa lacks the ventral lobes, and the
penes are fused along their entire length.
In nymphal characters N. pilosa resembles N,
inconspicud, possessing lanceolate gills, but lacks the
obvious dorsal white stripe on the abdomen, character-
istic of N, inconspicua. The mouthpart structure,
especially the labrum (Fig. 13d) and prosthecal
structure (Figs 13i, k), and the tine setae on the legs,
clearly distinguish WN. pilosa trom both N, inconspicua
and N. frsculer,
A comparison of qualitative characteristics of (he
three species of Nousia from South Australia is given
in Table 2,
History and Discussion
As mentioned previously the deseription of N,
Juscula nymphs given by Tillyard (1936) is similar to
that of N. pilosa and the associated nymph of N,
JSuscula differs (rom the deseription given by Tillyard.
N. fuscula and N, pilosa appear ecologically separated
by the timing of maturation and the imago mating
flights. Therefore, it is possible that Tillyard indirectly
associated the nymphs and adults collected from the
River Shannon, Tasmania. Consequently it appears
that (he nymph of N, pilosa was indirectly ascribed to
N. fuseula. The present descriptions are based on
associated material, and distinguishes the nymphs. ol
these two co-occurring species.
TABLE 2, COMPARISON OF QUALITATIVE CHARACTERISTICS OF NOUS14 JINCONSPICUA, N. FUSCULA AND
N, PILOSA FROM SOUTH AUSTRALIA
Character
N. ineonspicua
Male Imago
Lees: banding
Penes
Nymph
Legs: banding
Gills
Lamellae trachea
Lel| mandible
Incisors Quier
Incisors Inner
Right mandible
Incisors Outer
Incisors Inner
Prostheca Left
Prostheca Right
Labial palpi
Labrum
Hypopharyns
Absent
Fused apically
Ventral lobes present
No internal spines
Uniform brown
Broad luneveolate
Plentiful, branched
3
3
row
Slender, (wo terminal spines
Slender, two terminal spines
Slender
Width + 2 « length
Nol divided
N. fuscula
N. pilosa
Absent
Separate apically
Ventral lobes absent
Internal spines present
Absent
Linear
Few, simple
Robust, apex serrated
Slender, two terminal spines
Broad
Width = 2 » length
Deeply divided
Present
Fused apically
Ventral lobes absent
No internal spines
Present
Broad lanceolate
Plentiful, branched
3
2
Slender with lateral comb
Slender apex with three spines
Slender
Width <2
Not divided
length
GENUS ULMEROPHLEBIA Demoaulin [955
Demoulin 1955: 228-229) Tsui and Peters 1975; 538.
Male Imago
Fore wings 3-3.5 » longer than Wide, with numerous
crass-veins, those in the subcostal space are upright and
parallel. Hind wing with narrow costal region with
numeraus cross-veins apically, S¢ joins wing margin al
3/4 of wing length, MA forked, single intercalary
present, Tarsal claws dissimilar, one blunt club-shaped,
ane long, slendet, hooked distally (Fig. 17h). Forceps
three-segmented, basal segment very long, Penes much
shorter than basal segment of forceps, not reaching
narrowing of this segment. Cerci longer tan terminal
filament.
Type Species: Ulmerophlebia mjabersi,
Majure Nvinph
Head without tusk-like projections, Jabrum with a
small canvex projection on mid-anteérior margin, Labial
palpi three-segmented. Maxillary palpi three-
segmented, distal sezment very small. Gills on
abdominal segments 1-7 paired, each consisting al a
pair of broad, avate lamellae, with apical tracheal
filaments fringed with long fine hairs. Body and legs
fringed with long fine setae. Abdominal segments 6-9
with postero-lateral margins produced into backward-
pointing projections.
FRistory and Discussion
Ulmerophlebju was deseribed by Demoulin (1955) to
include a species described by Ulmer (1916) in the genus
Ewphyrus as &. mjobergi. Subsequently Ulmer (1920)
plaved this species in Deleatidinm and, Harker (1953)
accepted this determination. In 1955, Ulmer noted that
the 2/CUA were parallel, the hind wings had an Sc
which joined the wing margin at 3/4 of wing length
und resembled 4ta/one/la, However, the short pointed
penes of D. mijoberzi separated this species from
Deleatidiuin and Alalonella, and therefore a new
generic designation was made. The only generic
description is made by Ulmer (1916) in his species
deseription and, therefore the characteristics of the
imago are included here,
The nymph of W/merophlebia has never been
formally described and, therefore the generic character-
istics are also given here.
In the nymphal charaeteristics Ulmerophlebiq
resembles very closely nymphs of Jappa Harker, The
only major distinguishing feature is the lack of frontal
harns characteristic of this latter genus. Riek (1970)
also made note of this similarity although indirectly,
by stating that the eastern States’ nymphs of the genus
Jappa lack frontal horns. Tsui & Peters (1975) examined
the thoracic morphology of nymphs of Uhneraphlebia
and, although the nymphs had not been described or
associated with adults they found only 4 of the 16
REC. S. AUST. MUS. 19 (17): 339-197
Nayember 1986
character states that they examined differed from
nymphs of Juppa, These, plus the abseice of frontal
horns are the only dillerences in the nymph.
In the adults the major differences are the length of
the penes, which in Jeppa are almost equal to the
length of the basal segment of the forceps and, the
shorter Se vein in the hind wing which extends almost
to the apex of (he wing in Jappa. Tsui and Peters (1975)
reeorded | of 6 character states that differed.
Clearly Jappu and Ulmerophlebia are closely related
and future research may demonstrate that the two are
congeneric. However, on the basis of the limited South
Australian material, this is not possible-as part of this
work.
Ulmerophiebia pipinna sp. nov,
Holotype Mule lmao
Body Length 9.44 mim
Notal Length 2.54 nin
Pronotal Width 0.92 mm
Mesonotal Width 1.33 mm
Fore Wing Length 9.3) mm
Hind Wing Length 2.03 mm
Cerei Length 14.10 mm
Terminal Filamen( Length 11.97 mm
Body colour reddish brawn, abdomen darker dorsally,
Head: dark brown, with a white patch between ocelli,
Antennac shart, 1 mm long, basal segment twive as
long as wide, flagella 0,87 mm long.
Thorax: light brawn dorsally with white patches on
Scutosculellum, laterally with patches of white and
pink. Legs light brown, femora with two dark bands,
one distally, and one at 2/3 of length, T, partially
fused to tibia, join apparent. Ratios of lez segments:
fore leg 1.00: 1.74: 0.07 ; 0.65 : 0,60 : 0.47 ; 0.19
(1.84 mim); middle leg 1,00 ; 1.28 > 0.04 - 0.09: 0.09
0.06 7 0.20 (1.49 mm); hind lez 1.00 2 1.09 : 0.04 + 0.08 :
0.06: 0.06} 0.15 (1.80 mm). Sternum dark brown
(Fig. 17e). Prosterniim with heavily scleratized base.
Mesasternum: basisternurn length 1.46 » width, 0.79
~ furcasternum length, posterior margin rounded;
furcasternum length 0.87 » width, lateral matgins of
median longitudinal invagination divergent posteriorly,
posterior margin slightly concave.
Wings, hyaline. Fore wing (Fig. 17a), length 3.5»
width, cross-veins in plerostigmal region simple, costal
cross-veing extending along entire length, slightly
shaded with grey, cross-veins af subcostal space alsa
shaded with grey. Hind wing (Fig. 17b), length 2 =
width, costal hump not exaggerated, costal space with
5 cross-veins distally. Ry straight, Rs joins MA in
centre of wing, MA straight, MP branched in proximal
half of wing, anal tevion without cross-veins,
Genitalia: forceps light brown, proximal segment
long, 0.74 mm, broad at base, narrows approximately
hall way along length, second segment almost square,
distal segment longer, narrow proximally, Penes paired,
very short, extending half way to harrowing of proximal
segment of forceps, separate, mesal margins divergent,
lobes simple (Figs 17e, d; 19a, b).
EPHEMEROPTERA OF SOUTH AUSTRALIA
Mature Female Nymph (Fig, 20)
Head Width 1,96 mm
Notal Length 3.12 mm
Pronoral Width 2.32 mm
Mesanotal Width 2.36 inm
Cerei Length 8.25 mm
Terminal Filament Length 9.71 mm
Heud: brown. Lateral eyes. black, Antennae 3.08 mm
long wilh whorls of setae at apex of cach segment,
Tentorial body; width 2.33 »« length,
Thorax: brown. Pronotum, brown with black-brown
median marking, lateral margins lined with long fine
setae, Legs brown, with ove mid and one distal black
band on femora; tibiae and tarsi not banded (Fig, 18a),
Tarsal claws short and curved, with 12-16 small rounded
ventral denticles (Fig. 8b), Femora of fore and middle
legs almost equal in length, hind leg longest, 1.34 *
fore fermur length. Ratios of leg segments: fore leg
1,00 : 1.04: 0.37 (1,66 mm); middle leg 1.00: 0.94:
0.33 (1,60 tm); hind leg 1.00: 0,87 : 0.26 (2,14 mm).
Femur length to width ratios: fore ley 2.18, middle leg
2.22, hind leg 2.68, Sternum: prosternum triangular
wilh apex truncated, sternacostal suture absent,
Mesosternum: basislernum, almost square, length 0.92
= width, and equal to furcasternum length,
sternacostal suture present. Metasternum basisteraum
narrower than mesofurcasternum, width 4.75 « length.
Abdomen: dark brown wilh light central stripe on
segments 4-7, segments 8, 9, and 10 dark brown
(Pig. 20). All segments with long fine setae dorsally.
Caudal filaments well developed, terminal filament
longer (han cerei. Gills (Fig. 18e).
Moutlparts: labrum (Vig, 18d) 2.12 » wider than
long, lateral margins angular lined with long fine setae,
mid anterior margin with seven tubercles, three
tubercles on each side of a large sharp central
projection (Fig. 18e), dorsal surface covered with long
setae, Lelt mandible (Pig. 18h); lateral margin lined
with long setac, invisors displaced mesally, outer
incisors with two large teeth and four smaller teeth on
mesal margin, inner incisors with three apical teeth and
one small lateral tubercle, prostheca narrow with six
spines (Fig. 18i). Right mandible (Pig. 18j); outer
margin lined with long setae, incisors displaced mesally,
outer incisors rugose, with three apical teeth, with four
lateral (ubercles, inner incisors with two apical teeth
and one lateral tubercle, prostheca long, spinous (Vig.
I8k), Hfypopharyns (Pig 18f). Maxillae (Fig. 181) galeo-
Javinia rectangular, row of sixteen rake setae on ventral
surface, rake setae also interspersed within apical brush;
mesal corner with one large rake spine, inner margin
lined with lone fine pinnate sctae; palpi longer than
galeo-lacinia, proximal segment 2.33 x longer than
wide, second sezment long and broad, 1.88 « longer
than wide, distal segment very short, triangular, covered
with long setae, apex with two short spines, segment
ratios 1.00: 0.94 + 0.29 (0.36 mm). Labium (Pig. 182);
proximal segment of palpi 1.71 longer than wide,
segment ratios 1.00 ; 0.65 » 0.58 (0.41 mm).
Female linagoe
Unknown.
Subimago
Unknown,
Male Nymph
Unknown,
Diagnostic Characteristics
1, Genitalia of male imago, penes very short and
simple (Figs 17e, d; 19a, b),
2. Wing length 3.5 x width (Fig. 17a),
3, Femora of imago with two black bands.
4, Nymph without lrontal lobes (Mig. 20).
5. Labrum with one mid anterjor projection and three
denticles on cach side (Figs 18d, ¢).
. Maxillary palpi three segmented (Fig, 181),
. Lateral projection of glossae of labium (Fig. 18g).
. Mandibles, shape and form ol incisors and
prosthecae (Figs [8h-k).
mu Ind
Tipe Locality
Second Wannon River, Grampian Mouutains,
Victoria, Grid Reference on 1} 250 000 map series
Ballarat Sheet: 547396. Collected 25 November 1977
by P. J. and D. N. Suter.
Tvpe Specimens
The holotype male is deposited in the Museum of
Victoria. Two paratype imagos are also placed in the
Museum of Victoria. The wings and legs of the
holotype male are mounted on slides, and the genitulia
and body are in ethanol. The female nymph is mounted
on slides,
Type Habitat
The Second Watnon River in the Grampian
Mountains, is 4 moderately fast-flowing stream over
large cobble-size racks. The nymphs were collected
using a kick-sample technique, and therefore there is
no certainty of the habitat being occupied by this
species.
Etymology of Specific Epithet
The penes of L. pipinna are very short and simple
hence the specific epithet pipinna (L) for small penes.
Affinities
Uhe male imago of LU! pipinau can be distinguished
from U, mjoberei by the short simple penes. In U.
mioberai the penes are short and boot shaped, having
a lateral angular projection, absent in U. pipinna.
454
Material Examined
SOUTH AUSTRALIA. South East: Cress Ck, Bight
Mile Ck, Hitcheocks Drain.
VICTORIA, Crawford R,, Fitzroy R,, Fyans Ck, Shaw
R,,, Wannon R., Second Wannon R.
FAMILY BAETIDAE
This family occurs on every continent, and is
represented in Australia by the genera Baetis Leach,
Bungona Harker, Centroptilum Eaton, Cloeon Leach
and Pseudocloeon Kiapalek.
Characterization of the Baetidae can be found in
Edmunds, Jensen and Berner (1976),
GENUS BAETIS Leach [815
Leach, 1815: 137; Burmeister, 1839: 800 (&. cosialis later
placed in Atalophlebia); Pictet, 1843; 189-191 (B,
australasica later placed in Alalophlebia), Walker, 1853:
559-561 (B. australasiva, B. castalis); Eaton, 187i: 110;
1881; 196; 1885: 156-158; Ulmer, 1908: 44-45; Tillyard,
1926; 64; 1936; 50-53; Harker, 1980: 21-24, 29: 1954:
263-264, 266; Scholes, [96]: 36-38; Rick, [970: 235.
Type Species: Baetis bieculatus (Linn, Fabr.),
The first record of Baetis trom Australia was made
by Ulmer (1908) When & seror was described from
Western Australia, Tillyard (1936) described A. frater
from Tasmamia, and Harker (1950) added two further
species B, baddamsae and 8. canfluens trom New
South Wales. In 1954, Harker described B sogerensis
from Port Moresby, New Guinea, but this species has
not been recorded on the Australian mainland
Miiller-Liebenau (1969, 1973) recorded and discussed
the characteristics ised in the revision of the European
species of Bueris; these characteristics have been found
fo be useful in the present study. The following
redeseription of the male adult, and the original
description of the nymph of & sorer Ulmer, mclude
the characteristics which Miiller-Liebenau (1969, 1973)
found to be species specific.
Baetis soror Ulmer 1908
Baelis sorar Ulmer, 1908: 44-45; Tillyard, (926. 64:
Harker, 1950: 29; 1954: 266,
Male Imago
¥ SD wn Range
Body Length 5.45 0.45 18 4.40- 6.00
Notal Length 1.59 0,21 |4 1,36- 2.00
Pronotal Width 0.62 0.08 12 U,54- 0.76
Mesonotal Width O.88 0.08 12 0.74- L.00
Fore Wing Length 4.93 USI 30 4,20- 4.82
Hind Wing Length 1.0] 0.16 30 O.80- 1.26
Cerci Length 11.57 O92 5 40.25-12.50
General colour brown.
Head: dark brown, Dorsal turbinate eyes yellow,
oval, lateral eyes black,
REC. 8, AUST. MUS. 19 (17); 339-397
Noyentber, 1986
Thorax: light brown, Pronotum narrower than head.
Legs; buff, slender. Fore legs longer than middle and
hind Jegs, fore leg femur length 1.23 « middle leg
femur Jength, and 1.25 » hind leg femur length, middle
and hind legs with four tarsal segments. Ratios af leg
segments: fore leg 1.00: 1.80 : 0.08 ; 0,78 ; 0.62 : 0,37:
0.18 (0.93 mm); middle lez 1,00 : 0.98: 0,16 = 0.13 ;
0,07 | 0.20: - (0.75 mm); hind leg 1.00: 0.97 = 0.16:
0.13 5 0.07 ; 0,20 > - (0.76 mm). Tarsal claws dissimilar,
one blunt, club-shaped, one slender with a terminal
hook.
Wings: fore and hind wings hyaline with light brown
venation, Fore wing (Fig. 21a) 2.67 ~ longer than wide,
pterosligmal region slightly opaque (milky) with 7-10
cross-veins, anastomosed, branched or incamplete,
proximal region of costal space without cross-veins.
Hind wing With triangular costal projection, three
longitudinal veins, second forked with one intercalary,
one proximally located cross-vein in costal space (Fig.
2b).
Abdomen: brown, with black marking (Fig. 2c).
Cerci long, terminal filament reduced to a basal stump,
Genitalia: proximal segment of forceps cylindrical,
second segment bulbous, fused Lo third segment, third
segment long and slender, distal segment short but 3 x
longer than side, rounded, bulbous distally. Cavers of
penes broad, extending beyond bulbous second
segment ol forceps, bluntly pointed with divergent
apices (Figs 2ld; 19c).
Mature Male Nymph (Fig. te)
¥ SD on Runge
Head Width 0.97 ON7 22 0,86-1.06
Notal Length 1.50 (14 17 1.30-1-74
Pronotal Width 0.87 0.09 17 0.76-L.00
Mesonotal Width 1.27 0.13 17 — 1.06-1.46
Cerci Length 3.43, 0.50 7 2.60-4,16
Terminal Filament Length 2.29 0.24 7 2.00-2,460
Body cylindrical, hight brown.
Head: brown, dorsal compound eyes sepia, lateral
eyes black. Antennae long withou! apical projection
on basal segment.
Thorax; mesonotum brown with an oxbow shaped
white marking beside mesonotal suture, Legs bul grey
with darker femurtibia joint (Fig. 211). Tarsal claws
short curved with ventral peg-like denticles (Fig. 21g).
Ratios of leg segments: fore leg 1,00: 0.72: 0.64
(0.77 mm); middle leg 1.00 : 0.71 : 0.88 (0.79 mm): hind
leg 1,00 : 0.68 : 0.54 (0.82 mm), Femur length to width
rarios: fore leg 3.15, middle lee 3.45, hind leg 3.70.
Abdomen: brown, without definite markings,
Posteriar margins of terga with short sharp spines.
Paraprocts curved, lined with 17-24 spines (Figs 21h;
22a, b). Gills (Figs, 21i), margins serrated With one fine
bristle alternating with each serration (Fig. 211), first
gill small without clear trachea, gills 2-7 with black
branched trachea, 3,4,5, largest, ovate, 1,7 narrower,
EPHEMEROPTERA OF SOUTH AUSTRALIA 4545
Mouthparis: labrum (Vig. 21) ovoid, length 0.58 =
width, with deep coneavily in centre of anteriov margin
(Fig. 21k}, hyo small median denticles within concavity.
Left mandible (fig. 21m) outer incisors with three teeth,
outer tooth broad and robust, inner incisors with one
long central tooth and three shorter lateral teeth,
prostheca robust with one large curved apical tooth
with 3-4 spines in coneavity of tooth (Fig, 2lo). Right
mandible (Fig. 2p); outer incisors with first tooth
robust with two small inner teeth, inner incisors with
1-2 long central teeth with two (one each side) lateral
teeth, prostheea long and narrow with outer margin
lined with 6-7 setae (Fig. 21q). Hypopharynx (Fig. 211),
Maxillae (Pig. 2/1); galeo-lacinia long and narrow with
pointed apex, lined with four stout teeth, ventrally with
a line of pinnate setae anda line of curved setae below
leeth, palpi longer than galeo-lacinia, (hree-segmented,
segment ratios 1.00: 1.45 :0.36 (0.10 mm), distal
segmenl with a short tooth, all segments with short fine
setae, Labium (Fig. 21m); palpi, length of proximal
segment 2,06 ¥ width, inner margin af second segment
produced forming a lobe, segment ratios 1.00: 0.71 :
0,33 (0.19 min),
Female Imago
Wings similar, lacking dorsal compound eyes, lore legs
shorter than male, body colour cream, abdomen broad.
female Nyniph
Similar to male, lacking dorsal compound eyes, lateral
eyes black,
Diagnostic Characteristics
1, Hind Wings With three longitudinal veins, second
forked (Fiz, 21b).
. Third and fourth segments of forceps elonvated
(Figs 2ld; and Pig. 19¢),
3. Dorsal compound eye yellow in imago,
4. [ncisors and prosthecae of Jeft and right mandibles
(Figs 21n-q).
. Number of spines on paraprocts (Figs 21h; 22a, b).
tw
Ww
History and Diseussion
Ulmer (1908) described Baetis soror from south-west
Western Australia. This species resembles the South
Australian species in hind wing venation and forceps
structure. Ulmer recorded that "the costal and subcostal
region of the fore wings are weakly tanned (browned)”
sic, translation fram Ulmer, 1908, but the South
Australian specimens possess a milky costal and
subcostal region. Miuiller-Liebenau (1973) stated “the
colour of the plerostigma can be useful, especially
when one separates two related species in the same
material”. This colour difference is usually associated
with other character differences (genivalia and hind
wing venation) and is not used as a primary specific
character, Until further material from Western
Australia ig made available, it is. considered that this
single feature which differs from the type description
is not enough fo validate erection of a new species for
the South Ausfralian material. This material is there-
fore designated as Baeris soror Ulmer,
Malerial Examined
SOUTH AUSTRALIA, South East: Eight Mile Ck:
Deep Ck, Jerusalem Ck, Mosquito Ck, Hitchcock
Drain. Mt. Lofty Ranges: Deep Ck, Little Para R-
Fleurieu. Peninsula: The Deep Ck (Delamere), Finnts
R., Hindmarsh R., Inman R., Tookayerta Ck,
Yankalilla R, Southern Flinders Ranges: Nectar Brook
Ck, Spring Ck. Northern Flinders Ranges: Baleanoona
Ck, Brachina Ck, Bunyeroo Ck, Elatina Ck, Emu Ck,
Wirrealpa Ck,
GENUS CENTROPTILUM Eaton 1869
Eaton, 1869; 131-132; 1871: 107-108; 1885: 174-175;
Harker, 1957: 75-76; Rick, }970; 235,
Type Species: Centroptilum liiteoliint Eaton,
Cenfroptilum js also.a cosmopolitan genus, but it
was not until Harker (1957) deseribed C. callendum
from Kuringae Chase, New South Wales, that an
Australian species was recognised. A new species C
elongatum sp. nov, which is recorded in South
Australia, is described from associated material {rom
the Wannon River, Victoria. © e/ongaruim has a limited
distribution in South Australia and although many
nymphs are present in the collections there are relatively
few adult specimens. Consequently the type series was
taken from a much larger collection made in Victoria,
from which some. assessment of variation of character
expression could be made,
Centroptilum clongatum sp. pay,
Flalotype Male
Body Length 8.20 mm
Nota! Length 2.01 mm
Pronotal Width 0.95 mm
Mesonotal Widih 1.33 mm
Fore Wing Length 7.05 mm
Wind Wing Length 1.72 mm
Cerci Length 14.02 mm
Terminal Filament absent,
Head: light brown. Dorsal eyes turbinate with upper
portion brown/orange, oval, lateral eves grey,
Thorax: pronotum brawn. Metanolum dark brown
with median backward-produced projection (Fig. 23¢).
Lees; fore Ieg femur light brown, Gbia and tarsal
scxments darker brown, middie and hind legs light
brown, tarsal segments darker brown, T, fused to tibia
in all legs, Fore leg longer than middle and hind legs,
fore leg femur length 119 » middle leg femur length,
and 1.16 » hind leg femur length. Ratios of leg
segments (second value is the combined tibia + Ty
356 REC, $ AUST. MUS. 19 (17): 439-397
length to femur length): fore leg 1.00; 1.18 :-: 0,50:
0.44 + 0.26 | 0.16 (1.60 mm); middle leg 1,00: 0,75 ; -:
0.19 10,10: 0.08 | 0.16 (1,34 mim); bind Jeg 1.00 : 0.74 -
~7 O19; 0.10 20.07 ° 0.16 0.38 mm). Sternum hight
brown (Fig. 23e).
Wings: hyaline, Fore wing (Pig. 23a), 2.78 » longer
than wide, costal and subcostal spaces shaded with
yellow, pierostigma with 10-14 cross-veins, some forked,
one faint cross-vein in costal space present or absent,
subcostal space with six cross-veins, Hind wing (Fig.
23b); 2.15 % longer than wide, with a curved costal
projection, three longitudinal veins, secand forked with
one intercalary, two cross-veing between first and
second longitudinal veins.
Abdomen: brown dorsally with red tinges in patches
on segments 2-4 and 6-9, segment | dark brown
(Fig, 23c), Light brown ventrally with paired brown
inarkings on each side of median line.
Genitalia (Pig. 23d; Fig. 19d): forceps lour-
segmented, proximal segment rectarigular 1.6 » longer
than wide, second segment fused to third segment, third
segment long and slender, bowed slightly, distal
segment shorter, rounded apically. Pene covers paired,
rectangular, rounded apically, divergent, extending
beyond apex of proximal segment of forceps.
Mature Male Nymph (Fig. 231)
Body Length 7.68 mm
Head Width 1.32 mm
Notal Length 2.12 mm
Pronotal Width 1.16 mm
Mesonotal Width 1.6% mir
Cera Length 4,20 mm
Terminal Filament Lengeh 3.44 mm
Body colour brawn.
Heid: brown, Darsal compound eyes red-brown.
Thorax: brown. Legs yellow-brown with black
patches distally on femur about 2/3 of length (Fig.
24a); tibia and tarsi yellow-brown, nol marked. Tarsal
claws very long and slender, half tarsal length, lined
proximally with 13-20 fine deuticles, distal half smooth
and tapering, Ratios of leg segments: fore lee 1,00 :
0.63 : 0.66 (1.28 mm); middle lez 1.00: 0.625 0.54
(1.30 mm); hind leg 1.00 : 0.62 : 0.52 (1.30 nm), Femur
length width ratios: fore leg 4.92, middle and hind ley
5.42,
Abdomen brown, with tinges of red-brown,
patrerned a4 in Fig, 23f, Posterior margin of tervites
with long spines and smaller minute spines between
them, Hind margin of sternites similarly lined with long
spines and only minute inner basal spines. Paraprocts
rounded, with 25 large spines on internal and apical
margins (Fig. 24c; Fig. 22d); hind margins of ninth
Sternite with developing foreeps, separated by a concave
depression with 24 spines (Fig. 24b; Fig. 22c¢). Gills with
black, branched trachea (Fig, 24d), first gill sniallest,
hall-moon-shaped, lilth and sixth largest, margins
Naveinber 1986
serrated with one short fine bristle in each depression.
Mouthparts: labrim rectangular (Pig. 242), length
0.77 ». width, with a broad, deep, V-shaped concavity,
with truncated apex, and lateral denticles near apex
(Fig. 24f). Left mandible (Fig. 241), ourer incisors with
four large teelh apically and four smaller teeth along
inner margin, outer margin with one long spine, inner
incisors with three large apical teeth and 3-4 small
tubercles on ifmer basal margin, prostheca lany,
broadest proximally with paired apical projections, one
long and slender with one apical spine seta, the other
shorter, curved and blunt, with:a sharp opposing tooth
(Big. 24). Right mandible (Fig, 24k): outer melsors
with three large teeth, outer margin with one long
Lapering spine, inner incisors with two contiguous teeth,
prostheca strap-like, long and slender with 2-3 minute
spines on apex, anid two long setae near mid region
(Fig. 241). Hypopharynx simple, median lobe with a
large, bulbous, apical tubercle (Fig, 24h). Mawillae (Fig.
24m): palpi three-seemented, longer than galeo-lacinia,
basal segment long and slender, 3.67 » longer than
wide, lined with short fine sctac, segment ratios 1.00 :
0.55 + 1.09 (0.22 mim), Labrum (Mig. 24e¢): palpi three-
sepmented proximal segment 2.43 « longer than wide,
second segment narrow proximally, broad distally,
distal segment short, broad with concave apical margin,
segment ratios 1.005 0.71 20.35 (0.34 mm),
Feniale Imago
Resembles male, but lacks dorsal turbinate eves, Body
length 7.95 mm, fare wing length 7.45 mm, hind wing
length 2.87 mm. Thorax grey, abdomen red-brown.
Coastal margin ol tare wing browa. Fore leg shorter than
in male. Sub-atral plate with paraproets lacking spies.
Femule Nvmph
Similar lo male, lacking dorsal compound eyes, lateral
eyes black. Ninth abdominal sternite hind margin
square and lined with spines.
Diugnostic Charecteristies
}. Genitalia, shape of last seement of forceps (bik.
23d; Fig. 19¢).
Hind wing lacks acute vostal projection (Fig, 23h).
Number of spines on paraprocts of nymphs (Figs
24b, cy Figs 22e, d).
Incisors and prosiheea of mandibles (Figs 247-1).
Labrum shape (Fiz. 24e).
Saddle-like colour pattern on abdomen (Fig. 231).
Ww bs
DAS
Type Locality
Wannon River just above Wannon lalls, near
Wannon, Western Vicloria, Grid reference | : 250 (00
map senes Flamilton Sheet: 353482. Collected 30
October, 1977 by P. and A. Suter and A. Wells.
EPHEMEROPTERA OF SOUTH AUSTRAL DA
Type Specimens
The holotype male and nymphal type are located in
the Museum of Victoria, Five paratype male imagos
and paratype nymphs are also placed in the Museum
of Victoria. and five paratype male imagos and nymphs
are placed in the South Australian Museum,
Type Habitat
The nymphs were found in fast-flowing water about
500 m above the Wannon Falls. Adult males and
females were swarming in the afternoon sun above a
grass covered bank about 10 m above the water's edge,
The habitat records of this species in South Australia
are quite variable, but may reflect only the season and
(low conditions of the streams [rom which the collec-
tions were made, Collections from Kangaroo Island,
and Mosquito Creek in South Bast South Australia
during spring, indicated thal C. e/ongatuin nymphs
occupied a similar habitat to that observed in Victoria.
However, in Carrickalinga Creek on the Fleurieu
Peninsula, a collection was made in November 1977
when the creek was in the process of drying, and pools
were being formed, ti was from one of these stationary
Juncus lined pools, that the only record af
elongaliunr on the Fleurieu Peninsula, was made, At
times of higher water levels and discharge, C
élongatum may be found occupying the faster-flowing
waters, rather than the stationary pool habitat recorded
in November 1977.
Evvmoloey af Specific Epithet
The specific epithet elongatun refers to the
elongated distal segment of the forceps of the male
imago. This long segment distinguishes C, e/ongatumn
from C. collendum Harker, the only other species in
this genus deseribed from Ausiralia.
Affinities
OF the diagnostic features, the genitalia and hind
wing characteristics clearly distinguish C. elongaruim
from C. collendum Harker. Harker's (1957) description
records that the nymphs of C. collendum have long
tarsal claws: “about equal in lengch to the tarsus itself”.
C efongatum has much shorter tarsal claws, being
about half the tarsal length, Other differences between
the nymphs are not obvious from Harker's description,
and the nymphal morphotype of C, collendum has not
been cxamined.
Material Examined
SOUTH AUSTRALIA. South East: Mosquita Ck.
Fleurieu Peninsula: Carrickalinga Ck. Kangaroo
Islind: Breakneck R., DeMole R,, Rocky R., South
West R., Stunsail Boom R.,. Western R,
MICTORTA, Jimmy's Ck, Mt. Emu Ck, Wannon R,
357
GENUS CLOEON Leach 1815
Leach, 1815: 137; Baton, 1868: 87-88; 1871: 102; 1885:
179-181]; Klapadlek, 1905; 106-107; Ulmer, 1916: 17; 1919:
54: Tillyard, 1926: 64; L93@: 53-55; Harker, 1950: 24,
29; 1954; 266; 1957: 72-73; Scholes, 1961: 38-39; Rick,
1970: 236,
Type Species: Cloeon dipterum (Lian. Fabr.).
Alistory and Discussion
The genus Cloeon was erected in 1815 by Leach, to
include C, dipterum (inn, Fabr,), Eaton (1868) noted
“A species (1 specimen in British Museum) is reputed
to be from 5S. Australia’., and in 1885 he included
Australia in the distribution of this cosmopolitan
genus,
The first confirmed record of Cloeon in Australia
was made by Ulmer (1916) when he recorded C. virens
Klapdalek (incorrectly spelt as C. viridis Klap. by Ulmer
and Jater by Tillyard, 1926) [rom the Kimberley district,
NM. Australia. This species was originally described
from Java by Klapdlek (1905). A further species C.
/luviatile Ulmer was described by Ulmer (1919) fram
New Guinea, and was later recorded at Armidale
(N.S.W.) by Harker (1950), wha added the description
of the egg.
Tillyard (1936) recorded C. tasynanive from the
Macquarie R,. Tasmania, and described the male and
female imagos, and the subimago., Harker (1957)
described the first Cloeon nymph in her deseription
of ©. nandirum from Townsville, Queensland.
The present study recognises one new species, and
C. fluviatile Ulmer from South Australia, and the
descriptions melude the characteristics used by Miiller-
Liebenau (1969, 1973) in her revision of Beetts, The
nymph of C fluviatile is described from South
Australian material.
Cloeon Fluviatile Ulmer 1919
Cloeon fluviatile Ulmer, 1919; 54-57: Harker, 1950: 24,
29; 1954; 266,
The following description is based on one male
imago from rhe Third Spring on the Oratunga Loop,
Flinders Ranges, South Australia, collected 9 April
1977 by PL J. Suter.
Body Length 4.12 mm
Notal Length 1.40 mm
Pronotal Width 0.56 mm
Mesonotal Width 0.80 mm
Pore Wing Length 4,32 mm
Cerci Length &.53 mm
‘Terminal Filament absent.
Heud: dark brown, Antennae light brown 0,72 mm
long. Dorsal eyes turbinate, upper portion yellow-
brown, lateral portion lighter brown.
388 REC. §, AUST. MUS. 19 (17): 339-997
Thorax: brown, notal sutures black, Legs; buff. Ty
fused in middle and hind tibia, only partially fused in
fore tibia. Ratios of leg segments: fore lex 1.00: 1.76:
0.07 : 0.68 ; 0.49: 0,27 ; 0.20 (0,82 mm). middle lex
1.00: 0.82: - | 0.28 50,15 = 0.08 ; 0.18 (0.78 mm); bind
leg 1,00 : 0.86 > - : 0.262 0.12 : 0.07 : 0.19 (0.86 mm),
Sternum brown (Pig, 25c),
Wings: hyaline. Fore wing (Pig. 25a) length 2.70 »
Width, costal and subcostal region opaque, cream-
coloured, milky in pterostizmal region, prerostizma
with 2-4 cross-veins, no cross-veins in proximal region
of costal space, subcostal space wilhoul cross-veins,
Abdomen; red-brown with light brown median
markings on segments 1-7, segments 8, 9, 10 red-brown
(Fig, 25b), Light red-grey ventrally, Cerei long, white
with red-brown joints every fourth segnrent, lerminal
filament absent,
Genilalia (Figs 25d-f, 19e): Forceps proximal segment
rectangular, length 0.69 xwidth, second segment
slender, partially fused with third segment which is long
and slender, distal segment short, with a basal stalk
and globular apex. Penes covers large and obvious,
extending to middle of second segment of forceps,
separate, rectangular, apices divergent. Subgenital plate
with a posterior brown projection which separates
proximal segments ol forceps.
Mature Male Nymph
Body Length 508-560
Head Width 1.02-1.08
Notal Length 1.60-1. 74
Pronotal Width ().92-0.98
Mesonotral Width 1.38-1.46
Cerci Length 3. 40-5 40
Terminal Filament Leneth 1,62-3.80
Body cylindrical, general colour light brown.
Head: brown. Dorsal compound eyes red-brown.
Antennae long, 3 xlonger thai head width,
Thorax: pronotum width 0.91 « head width, brown
with light markings. Mesonorum width 1.35 » head
width, uniformly light brown. Legs cream with brown
banding, one band on distal 1/3 of femur, one
proximally on tibia, tarsus wilh one proximal and one
distal band (Pig. 26a). Tarsal claws long and slender
with two ventral rows of peg like denticles (Fig. 26b).
Ratios of leg segments: fore lez 1.00: 0.73 : 0.63
(1.0L min); middle leg, 1,00 : 0.74 ; 0.54 (1.03 mm); hind
lex 1,00; 0.68 : 0.53 (1.16 mm), Femur length-widih
ratios: fore leg 4.87, middle lee 5.01, and hind leg, 5.27.
Abdomen: brown, with dark brown rectangular
patches on cach segment, edged with light brown, 4
central light stripe on each segment (Fig. 26c), lateral
flanges af segments 7-10 lined with spines, J-2 spines
on postera-lateral margins. Posterior margins of tergites.
with large single spines, Paraproeis with 16-22 spines
Gn inner and apical margins, larges! spine at apex,
smaller mesally (Fig. 26cl). Gills (Fig. 26e) with paired
lamellae on segments I-6, single on segment 7, black
Novenilier, 1986
branched trachea, margins serrated with short fine
bristle in each depression,
Mouthparts: labrum (Fig. 26f) rectangular, 1.94 =
broader than long with a U-shaped concavity in centre
of anterior margin (Fig. 26g). Left mandible (Fig, 26));
robust, ouler imeisors with 3-4 large teeth, inner incisors
with 4-5 teeth, prostheca with three apical denricles atid
two sharp spines (Fig. 26k), Right mandible (Pig. 261)
robust, outer incisors with three apical teeth, inner
incisors. with three large and one small teeth, prostheca
long and slender, mesal margin near apes with two
short spines and one small denticle (Mig. 26m).
Hypopharynx (Fig, 26n), Maxillae (Mig, 261) galeo-
lacinia slender, with three well developed apical teeth,
lined mesally with 12-15 large spine setae, palpi longer
than galvo-lacinia, proximal segment 5.83. « longer
than wide, segment ratios; 1,00: 0.80 : 0.82 (0.15 num),
distal segment lacks apical spines. Labium (Fig. 26h);
palpi: length af proximal segment 2,20 ~ width, apical
margin of distal segment concave lined with short spine
selue, segment ratios; 1,00; 0,61 ; 0.80 (0.22 mm);
glossae shorter than paraglossae.
Female Imago
Lacks dorsal compound eyes, lateral eyes black, fore
leg shorter than male, otherwise similar to male imago,
Female Nymph
Similar to male nymph, lacks dorsal compound eyes,
lateral eyes black, thorax broad, wider than head,
Diagnostic Characteristics
1. Distal segment of forceps minute, globular (Figs
25d-t; !9e),
2. Turhinate eyes. yellow/brown,
3. Incisors and prostheca al lett and right mandibles
(Figs 26j-m).
4. Distal segment af maxillary palpi without apical
spines (Fig, 261),
3, Paraproct spination (Fig. 26d)-
History and Discussion
C. fluviatile was deseribed in 1919 by Lilmer from
Specimens from New Guinea. Harker (1950) recorded
this species from Armidale, New South Wales, and
deseriked the egg. Since the type material is held in the
Berlin Museum (Ulmer 1919) or the Stockholm
Museum (Harker 1950, 1954) it was not available for
his study, The South Australian specimens have po
features which distinguish them Irom the species.
described by Ulmer, and until the type material is
examined they. are placed as ©. fhviitile Ulmer.
Material Examined
SOUTH AUSTRALIA, Mi, Lolry Ranges: Torrens R,
Souther Hhnders Ranges: Ohlenmeyer Reservoir,
EPHLEMEROPTERA Gh SOUTH AUSTRALIA
Rocky R., Wild Dog Ck. Northern Flinders Ranges:
Baleanoona Ck, Bendieuta Ck, Brachina Ck, Elatina
Ck, Emu Ck, Erevunda Ck, Kanyaka Ck, Marolana
Ck, Mt. Chambers Ck, Parachilna Ck, Stubbs
Waterhole, Willigan Ck, Creek in Warren Gorge,
Cloeon paradieniensis sp. noy,
Holotype Male
Body Length 7.80 mm
Notal Length 2,96 nim
Pronotal Width 0.92 mm
Mesonotal Width 1.20 mm
Fore Wing Length 7.05 mm
Cere) Length 15.73 mm
Heal: brawn, Antennae short, 1,02 mm, buff.
Darsal eyes turbinate, yellow dorsally, brown laterally.
Thera brown, pronotum narrower than head. Legs;
white-cream, fore leg longer than middle and hind legs,
fore leg femur length 1.06 « middle leg femur length,
and equal to hind leg femur length. T; of middle and
hind legs fused to tibia, Ratios of leg seyments; fore
leg 1.00 : 1.68 : 0.05 : 0.70: 0.50 : 0.28 : 0.18
(1.48 mm); middle leg 1.00: 1.0L; - + 0,29: 013 : 0,06;
0.16 (1.40 mm); hind leg 1,00: 0,99; -:;0,26:012:
0.06; 0.15 (1.46 mm),
Wings: hyaline (Fig. 25g); 2.89 » longer than wide,
veins buff, transparent, pterostigmal region with 3-4
cross-veins, well separated, proximal region of costal
space with two cross-veins, sub-costal space with two
cross-veins in distal half,
Abdomen: red-brown with a light brown dorsal
stripe, segments 8 and 9 dark red-brown, 10 lighter (Fig.
25h), Cerci long, terminal filament reduced ta a
vestigial stump,
Genilalia (Figs 25i-k; 19f): forceps proximal segment
short and broad; second segment shart, narrower than
proximal segment, almost fused with third segment;
third segment long and narrow, bulbous apically; distal
segment short and angular, conical. Penes covers broad,
extending beyond fusion of second and third segments
of forceps, bluntly pointed, apices divergent,
Mature Male Nyrnph
Head Width 1,31 mm
Notal Length 1.96 mm
FPronotal Width 1.15 oun
Mesonotal Width 1.64 mim
Cera’ Length 6.23 mm
ferminal Filament Length 4.59 mm
Body cylindrical, red-brown dorsally, light brown
ventrally.
Head: dorsal compound eyes red-brown (sepia).
Antennae long, 4.92 mm, proximal and second segment
brown, llagellae buff,
Thorax: pronotum width 0.88 « head width, brown
with median longitudinal white stripe, Mesouotum
width 1.25 head width, brown with light median
359
longitudinal stripe. Sternum light brown, with little
sclerotization, Legs buff without markings (Fig, 27a).
Tarsal claws long and slender, with two ventral rows
of peg-lke denticles, Ratios of leg segments: fore leg
1.00: 0.67 : 0.58 (1.44 mm); middle leg 1.00 : 0.67 :
0.51 (1,56 mm); hind leg 1.00 : 0.79 : 0,60 (1.68 mm).
Femur length to width ratios: fore leg 5,18, middle leg
6.05, hind leg 6.51.
Abdomen: red-brown dorsally, yellow-brown
ventrally, without definite colour pattern, overlap of
segments darker brown, Lateral margins o! segments
7-10 lined with spines, postero-lateral margins with 2-3
spines, posterior margins ol tergites with large and
small irregularly placed spines, Paraprocts broadly
triangular, lined on mesal margin with 27-30 large
spines (Fig. 27b). Cerci long, terminal filament shorter,
every fourth segment red-brown, giving banded
appearance, Gills; lamellae paired on segments 1-6 (Fig.
27e), seventh single, margins of gills serrated with a
single fine bristle.
Mouthparts: labrum (Fig, 27c) rectangular, 1.50 ©
broader than long with a deep concavity in centre of
anterior margin (Fig. 27d). Left mandible robust (Fig.
271), outer incisors with four teeth, inner incisors with
3-4 teeth, prostheca robust with a corrugated apex of
5-6 rounded teeth and lwo long spines mesally (Fig.
27)). Right mandible (Fig, 27k) robust, outer incisors
with four teeth, innerincisors with two large teeth and
two smaller teeth, prostheca robust with apex af 6-8
tooth-like ridges (Fig. 271). Hypopharynx (Fig. 272)
simple, median lobe rounded with a small median
bulbous projection. Maxillae (Pig. 27f) galeo-lacinia
long and narrow, with three well developed sharp teeth
apically, palpi longer than galeo-lacinia, proximal
segment 5.33 ~ longer than wide; segment ratios
1.00 : 0.69 = 0.88 (0.24 mim), distal segment fringed
with fine setae, and two small terminal teeth. Labium
(Fig, 27h) palpi three segmented, lenvth of proximal
segment of palpi 2.74 » width, apical margin of distal
segment slightly concave, segment ratios
1.00 50.56: 0.48 (0.33 mm); glossae shorter than
paraglossae.
Feinale Imago
Walhout dorsal compound eyes, fore lees shorter than
male. Fore wing: costal and subcostal spaces shaded
with red-brown, twelve crass-veins in costal space, Body
calour red-brown.
Female Nymph
Similar to mate, Jacks dorsal compound eyes, lateral
eyes black, thorax broader than male, pronotum wider
than head.
Diagnostic Characteristics
1. Distal seement of forceps conical shape (Figs 281-
ky 19f),
2. Turbinate eyes yellow.
360 REC. S. AUST. MUS. 19 (17): 339-397
3. Female with costal and subcostal spaces shaded
red-brown.
4, Incisors and prostheca of left and right mandibles
(Figs 27i-l).
. Paraproct spination (Fig. 27b).
6. Maxillary palpi with terminal spines on distal
segment (Fig. 27f).
an
Type Locality
Little Para River at Paracombe, Mt, Lofty Ranges,
South Australia, Grid Reference | : 250 000 map series,
Adelaide Sheet: 179695. Collected 20 October, 1976 by
J. H. Diener and P. J. Suter.
Type Specimens
Holotype male and nymphal type and allotype
female are located in the Museum of Victoria. This
short type series is because this species is only known
from two collections from the type locality, and one
is the drain system of the South East. Further material
has not been collected.
November, 1986
Tvpe Habitat
The nymphs were found in a non-flowing pool
system in a culvert beside the Paracombe road. The
pool was overgrown with Nasturtium sp. and Lemna
sp. Adults were raised in the laboratory from mature
nymphs collected from the type locality.
Etymology of the Specific Epithet
The specific epithet paradieniensis refers to the river
(Little Para River), and is in recognition of J. H. Diener
whose collection from the Little Para River was the first
of this species.
Affinities
Cloeon paradieniensis resembles all the described
Australian species, but it can be readily distinguished
by the conical shaped distal segment of the forceps of
the male imago. The nymph can only be compared with
C. nandirum Harker and C. fluviatile Ulmer, the only
Australian species associated with their nymphs. The
shape of the labrum clearly distinguish the nymphs of
TABLE 3. TABULATED COMPARISON OF ALL DESCRIBED SPECIES OF CLOEON IN AUSTRALIA — DATA
COLLECTED FROM THE PRESENT STUDY AND FROM PUBLISHED DESCRIPTIONS BY KLAPALEK (1905),
ULMER (1919), TILLYARD (1936) AND HARKER (1957)
Character Cloeon jluviatile C. paradieniensis C. nandirum C. virens C. tasmaniae
Male Imago
Body length (mm) 4.12 7.80 7 6 7
Fore Wing length (mm) 4.32 7.05 4 6 6
Fore Wing width (mm) 1.60 2.44 1.6 2.27 2.4
Cerci length (mm) 8.53 15.73 = 11-12 12
Eye colour Sepia Yellow Orange — Bufl-pink
Pterostigmal cross-veins 2-4 3-4 5 4-5 5
C/Sc basal cross-veins 0 2-5 0 6 —
Sc/R cross-veins 0 2-6 0 7 —
Costal colouration Milky Milky Milky Emerald Green Cream
in females.
Genitalia
Forceps, terminal Short and narrow Triangular Long and narrow Short and narrow Globular
segment
Penes covers Rectangular, Pointed apically ? ? ?
flat apically
Nymph
Body length (mm) 5.08-5.60 — 6 Unknown Unknown
Head width (mm) 1.02-1.08 1.31 a
Cerei length (mm) 3.50-5.40 6.23 —
Terminal filament length 1.62-3.80 4.59 oe
Legs Banded Not banded Not banded
Left Mandibles
Outer incisors 3-4 teeth 3-4 teeth _
Inner incisors 4-5 teeth 5-6 teeth _
Prostheca 3 denticles + 5-6 teeth + —
Right Mandibles
Outer incisors
Inner incisors
Prostheca
Maxillary palpi
ae eer ee ee Oe
2 sharp spines
3 teeth
3 large, ] small
Long and slender,
2 short spines +
1] denticle apically
No terminal teeth
2 long spines
4 teeth
2 large, 2 small
Robust, 6-8
tooth-like ridges
2 terminal teeth
-EPHEMEROPTERA OF SOUTH AUSTRALIA
C. paradieniensiy and CG nandirum, but as the type
material is in the British Museum, and the description
given by Harker (1957) was not comprehensive enough,
no further character comparisons are possible.
The nymphs of C. puradieniensis can be
distinguished trom C. fluviatile Ulmer initially by size,
the litter species being less than 6 mm, the former
greater than 7.5 mimi, The number of spines on the
paraprocts, the lack of bands on the femora, spines on
the distal segment of the maxillary palpi, and ihe shape
of the prostheca of the right mandible also distinguish
the twa species found in South Australia, Tabulated
comparisons of all Australian Cloeon species 15 given
in Table 3, with data taken from the present study, and
frow published descriptions by Klapalek (1905), Ulmer
(1919), Tillyard (1936) and Harker (1957),
FAMILY SIPHLONURIDAE
A full revision of this Family is at present being
prepared by Dr I, Campbell, Chisholm Institute of
Technology, Victoria.
GENUS TASMANOPHLEBIA Tillyard 1921
Tillyard, 1921: 409-412; 1926: 62; 1933; 12-13; 1936; 27;
Lestage, 193Sa: 132; 1935b: 350-353 (in part as
Tasmanophlebioides), Harker, 1950; 29; 1954; 267;
Riek, 1955; 268-269; 1976; 235; Scholes, 1961: 21-23.
Type Species: Tusmanophlebia lacustrus.
As mentioned above, Dr 1, Campbell is revising the
Siphlonuridace and therefore material of the single
species of Tusmanophlebia recorded in South Australia
has been forwarded to him for comparison with other
Australian species. No specific designation has been
made for the South Australian species,
Tasmanophiebia sp.
The following description is of one male imago from
Tookayerta Creek, Mleurieu Peninsula, South Australia..
Adult specimens are rare in collections from South
Australia, and for this reason the mean, ranges and
standard deviations are based on only three animals,
Male Imago
¥ SD n Range
Body Length 10.40 1.03 3 9.40-11.46
Fore Wing Length 9.54 0.90 3 §&.53-10.26
Fore Wing Width 2:90 0.43 3 2.42- 3,25
Hind Wing Length 4.53 0.6) 3 3,86- 5,06
Hind Wing Widih 251 0.26 3 2.23- 2.74
Cerei Length 4.71 (other cerci damayed)
Terminal Filament Length 0.24 0.14 3 0.08 0.32
Head: light brown. Dorsal region of compound eyes
burgundy:
Thorax: brown, Fore leg dark brawn, longer than
middle and hind legs, fore lee femur length 211»
middle leg femur length and 1.87 « hind femur length,
Middle and hind legs light brawn, lirst tarsal segment
301
fused to tibia. Ratios of lee segments: fore leg 1.00:
0,67 + 0.63 | 0.47 2 0.52 2 0.47 = 0.28 (2.24 mm); middle
leg 1.00 : 0.84 2-2 0.17 > O17; O14: 0.29 (06 mm);
hind leg 1,00; 0.803 -1 01712017: 012.028
(1.20 mim).
Wines; fare and hind wings hyaline, tinged with
yellow, veins brown. Fore wing (Fig. 28a) 3.31 x longer
than wide, costal and subcostal region shaded with
brown, radial and proximal regions of median, cubital
and anal veins tinged with yellow. Hind wing (Fig. 28b);
1.79 « longer than wide, half as long as. forewing,
subcostal space shaded with brown, costal, subcostal
and proximal regions of the radial and anal veins linged
with yellow.
Abdomen; brown, speckled with black, segments 8
and 9 with median black stripe and two convex lateral
Stripes, segment 10 black (Fig, 28c), Cerci long, dark
brown, terminal filament vestigial, of 1-4 segments,
Genilalia (Figs 28d, @; 30a, b): forceps three-
segmented, subgenital plate broad forming forceps
base, proximal segment very lang and narrow, middie
segment shorter, but elongated, distal segment just
shorter than middie, rounded apically. Penes long,
arrow, extending to mid proximal segment of larceps,
lobes tubular, almost fused, rounded at apices.
Mature Male Nviriph (Fig. 282)
y¥ SD on Range
Head Width 17! O12 & 1.53- 1,84
Body Length 12,64 1.00 4 11.18-13,40
Notal Length 2,91 0.21 6 2.58- 3.20
Pronotal Width 177 O12 6 J.58- 1.88
Mesonotal Width 2.26 0.16 6 2.00- 224d
Cerei Length 5.09 0.29 3° 4.71. 5,41
Terminal Pilament Lengih 4,68 0,13 3 4.52- 4.83
Colour mottled sandy brown.
Head: small, light brown, Compound eyes large, red-
brown dorsally, black laterally. Antennae 1,30 mim long,
Thorax: pronotim as Wide as head, miortled grey-
hrown. Mesonotum width 1.32 « head width, mottled
brown. Legs yellow-brown, femora with a brown patch
on posterior margin, joints of tibia and tarsi dark
brown, tarsi with brown bands glory length, equivalent
to Larsal segments of adult (Fig. 29a). Tarsal claws long
and slender 0.50-0.71 « length of tarsus, smooth
without denticles. Ratios of leg segments: fore leg
1.00 >.0.46:.0.85 (1.25 mm), middle leg 1.00: 0.41.
0.67 (1.30 mm), hind Jeg 1.00: 0.44 ; 0,74 (1,32 mm),
Femur length ta width ratios similar; fore lee 3.44,
middle leg 3.25, hind leg 3.56.
Abdoinen: dorso-verrally flattened, with medial
dorsal crest of curved pasteriarly directed projections
on segments 1-7; process on segment | small, 2, 3,
largest, becoming less prominent from segment 4-7
(Fig. 28g), Lateral flanges of each abdominal segment
semi-transparent, postero-lateral margin sharply
produced. Paraprocts separate, smooth, developing
362 REC. 5. AUST. MUS, 19 (17): 339-397
forceps large (Figs 22c, f), Gills; on segments 1-4, first
pair broadly ovoid, opereulate, other three pairs. with
paired lamellae, transparent, with well-developed gill
lamellae lined with fine setae (Figs 29b-e).
Mouthparis: jabrum (Fig, 29f) rectangular, length
0.44 x width, anterior margin smooth (Fig. 29g), Lefi
mandible (Fig. 29k) rabust. incisors widely separate,
outer group with three apical teeth, anda ventral row
af short setae, inner incisors with three apical teeth and
arow of short setae, prostheca broad at base, anlerior
margin tapers to torm a long narrow projection with
3-4 short spines (Fig, 291). Right mandible (Fig. 29m)
rabust, incisors widely separate, ouler group with two
apical teeth, and a ventral row af short selae, inner
incisors with three apical teeth and a ventral row of
short setae, prosthcea broad at base, curved, apex
divided into (wo separate lobes, posterior lobe largest
(Fig. 29n). Hypopharynx (Fig. 291): median lobe deeply
bifid. Manillae (Fig. 29}); apical angle of galeo-lacinia
with three or four spine setae, palpi three seemented,
longer than galeo-lacinia, segment ratios; 1.00 : 0.86 :
0.77 (0.35 mm). Labium (Fig. 29h): palpi three-
segmented, length of proximal segment 1.3 » width,
segment ratios; 1.00 ; 0.86; 0.51 (0.37 mm), elossae
with one small pointed tubercle distally.
Diagnostic Characteristics
1. Genitalia; shape of forceps and penes (Figs 28d, e;
30a, b)..
. Dorsal crest of nymph with curved posteriorly
directed projections on abdominal segrnents }-7
(Fig. 28g).
3. First abdominal gill ovoid, rounded posteriorly (Fig,
29b),
4, Lateral flanges of abdominal segments narrow.
5. Shape of incisors and prosthecae at mandibles (Figs
29k-m).
tw
Material Examined
SOUTH AUSTRALIA. Fleurieu Peninsula: Tookayerta
Ck, Yankalilla R.
VICTORIA, Gawkers Ck, Stakes Ck,
FAMILY CAENIDAE
The Caenidae was recognised as a distinct group of
mayflies by Eaton (1883) when he included the genera
Tricorythus, Leptohyphes and Caenis in Section 7 of
his Reyisional Monograph, Banks (1900) erected the
tribe Cacnini and Thew (1960) states that “according
io the Copenhagen decision of the International
Commission on Zoological Nomenclature, Banks
should be credited with the authorship of the family.”
Lestage (1930, 1938) refers lo the Caenidae as. the
Brachycercidae, but with the exception of Demoulin
(19556), the Brachyceridae has not been recognised by
authors working on the Australian Ephemeroptera,
Tillyard (1936), Harker (1950, 1954, 1957), Thew (1960),
November, 1986
Riek (1970), Soldan (1978) and Suter (1979, 1984) have
all recognised the family Caenidae,
In 1978 Soldan described a new genus of caenid from
Australia (Psewdocaenis) trom nymphal material only,
but Suter (1984) demonstrated that this genus was a
synonyin of Tasmanocoenis,
GENUS TASMANOCOENIS Lesiage 1930
Puthz, 1975; 412; Saldan, 1978: 128; Suter, 1979: 82:
Suter, 1984; 105.
Type Species; Tasmnanacoenis tonnoiri.
The genus Jasmanacoenis has been reviewed by
Suter (1984) and further discussion is not included here,
In the present study all caenid material from South
Australia was Tasmanocoenis lillvardi (Lestage).
Tasmanocoenis tillyardi (Lestage) 1938
Caenis scott Villyard, 1936: 56-58; Coens tillvarei
Lestage, 1938: 320; Caenis scot(i Harker, 1950: 24-26,
29; Caenis tillvardi Harker, 1954: 266; Tus manocoenis
Hillyard’ Demoulin, 1955b: 4; Harker, 1957: 77: van
Bruggen, 1957: 33; Thew, 1960: 202; Scholes, 1961:
39-41; “Caenis” scott Williams, 1968: 169.
Male Imago
¥ SD A Range
Body Length 3.25 O14 7 3,12- 3.52
Nolal Length 1.45 G07 7 1.32- 1.52
Pronotal Width U.68 0.06 7 O.55- 0.72
Mesonotal Width G84 0.08 7 0,67- 0,90
Fore Wing Length 3.15 0,14 7 2,96- 3.36
Cerei Length 130 — = —
Terminal Filament Length 12.00 — — —
Colour dark black-brown.
Head; brown, With light brown epieranial sutures,
Compound eyes lateral, black. Antennal base brawn,
one-segmented, Magellae long, 0.54 mm.
Thorax: robust, dark black-brawn (Fig. Jb).
Pronotum narrower than head. Mesonotum Wider than
head. Legs slender, pale brown-grey; fore legs longer
than middle and hind legs, fore leg femur LIS
middle leg femur length and 1.28 x hind femurlength.
Ratios of lez segments: fore leg 1.00: 1.93: 0.09 :
0.57; 0.26 7 0.25 20.16 (0.69 mm); middle leg 1.00 :
0.55 : 0.08 : 0.05 : 0.05 : 0.07 : 0,08 (0.60 mm); hind
leg 1,00 ; 0,61 : 0.08 ; 0.05 : 0.05 : 0.07 : 0.08
(0.54 mm). ‘Targal claws similar in fore leg, both blunt,
club-shaped, dissimilar in middle and hind legs, one
blunt club-shaped, one slender and sharp. Sternum
(Fig. 3lc); prosternum triangular, apex truncated,
lateral margins separated anteriorly, slightly longer than
broad. Mesosternum dark black-brown, basisternum
lengih 1.19 « maximum width, sterna-costal suture
well deVeloped, furcasternum length 0.65 « width, and
0,65 » basisteruum length, posterior margin straighe.
Wings (Fig. 31a): short and broad, length 1.72. «
EPHEMEROPTERA OF SOUTH AUSTRALIA 463
width, hyaline with milky-opaque plerosigma,
venation reduced, simple, almost lacking ¢ross-veins,
posterior margms may be lined with fine setae.
Abdomen: short, cylindrical, segmenrs 1-5 very
short, light brown, and speckled with black, segments
8 and 9 lighter. Cerci long, transparent, terminal
filament longer, both tipped with long fine setae,
Genitalia (Figs 3)d; 30c): forceps one-segmented,
bowed, sharply pointed with ventral mesal groove,
penes lobed, fused wilh a small apical indentation,
sclerotized basally,
Mature Male Nymph (Fig, 3le)
y¥ SD a Rarige
Head Width 0.96 0.04 21 0.90-1.04
Nolal Length 1.62 O10 21 1,.40-1.76
Pronotal Width 1.00 0.06 21 O.78-1.04
Mesonotal Width 1.14 0.05 21 1.00-1.20
Cerci Length 3.03 0.16 3 2.92-3.22
Terminal Filament Length 3,39 (18 3 3,28-3,60
Body calour brown.
Head: dark brown, Antennae light brown, basal
sezment 0.16 mm long, flagellum 1.46 mm (Fig. 32e).
Tentorial body rectangular, length 0.79 « width,
Thorax: pronotum brown, lateral flanges lighter
semi-transparent, anterior margins with spine setae,
0.96 « wider than the head, Mesonotum dark brown.
Lees light brown, margins lined with spine Setae
(Fig, 32a), ‘farsal claws short, curved with 4-6 small
ventral denticles, otherwise smooth. Ratios of leg
segments: fare leg 1.00 ; 0.73 ; 0.65 (0.68 mm); middle
leg 1,00 : 0.70 30.59 (0.68 mm); hind leg 1.00 : 0.76:
0,60 (0,74 mm), Femur length to width ratios: fore leg
2.81, middle leg 2.86, and hind leg 2.95.
Abdomen; brown, with square patierns of brown on
each side of median line. Operculate gill of Secand
segment covers segments 3-S, segment two with median
backward-projecting spine, poslero-lateral margins
produced forming backward pointing projections.
Cerci and terminal filament dark brown, well
developed, Gills, on segments 1-6, first pair single,
filarnentous with indistincl segments, lined with Line
selae (Pig, 32b), second pair onerculate with raised
triangular region dorsally, mesal ridge with few setae,
outer ridge not reaching posterior margin of gill covers,
margins lined with long setae (Figs 32¢; 30d); third-
sixth pairs laminate with 40-50 tracheal fringes, single
or bifid, few trifid, third gill largest (Pig, 32d),
Mouthparts; labrum (Fig. 32f) rectangular, width
2.34 ~ length, anterior margin with slight median
concavity with 2-3 small denticles (Mig. 32g), Left
mandible (Fig. 32j): outer incisors with three apical
teeth and one shorter mesal tooth, inner incisors with
three apical teerh, prostheea robust with apical brush
of setae (Fig. 32k). Right mandible (Fig. 321); outer
incisors with 2-3 apical teeth, inner with two, prostheca
robust wilh apical brush of setae (Fig, 32m). Hypo-
pharynx (Fig. 32i) with square median labe, slightly
concave anteriorly, Maxillae (Fig. 32n); galeo-lacinia
short and narrow, with 3-4 robust apical spines, mesal
margin lined with stout spine setae, palpi three-
segmented, longer than galeo-lacinia, segment ratios
1.00 0.70 : 1.07 (0.12.mm), Labium (Fig. 32h): palpi
three-segmented, proximal segment length 1.47. »
width, segment ratios 1.00; 0.78; 0.53 (0.13 mm);
glossae rectangular.
Female Imaga
Similar to male, sternum and notum broader, tarsi four
segmented, tarsal claws, each pair dissinular, one blunt,
club-shaped, one curved and sharp.
Female Nymph
Body shape similar to male, more robust than male,
ic. head width of last instar greater than male, wing
sheaths longer, second abdominal operculaie gill
longer, covering segments 3-6, fore, middle and hind
femora langer.
Diugnastic Characteristics
1. Genitalia of male with curved sharp foreeps and
fused penes with a small apical indentation, nor
extending beyond apices of forceps (Figs 31d; 30c).
2. Labrum rectangular (Fig, 321),
3. Structure of mandibles, incisors and prosthecac
(Figs 32j-m).
History and Discussion
Tillyard (1936) described Caenis scotti from the
South Esk River at Clarendon, Tasmania. Lestage
(1938) noted that C. scotti was preoccupied by a species
described by Ulmer in 1924 (referred to by Thew, 1960)
and renamed the Tasmanian species Coenis tllardi.,
Harker (1950) apparently was unaware af this name
alteration and described a nymph which she assigned
to Cuenis scorti, Subsequently in 1954 she recognised
the mame change, and maintained the generic
recognition as Caenis, Demoulin J955b) reviewed the
genus Jasmanoccenis and recognised that Cgenis
vilvardi belonged in the genus Tasmanocoenis, an
observalion validated by the review of the Caenidae by
Thew (1960),
Williams (1968) noled that Ceenis and
Tasmanocoenis in Australia were probably
synonymous, and illustrated gills of “Caenis” scvtti
after Harker (1950). Rick (1970) noted, as had
Demouli (1955b) and Thew (1960), that 7asmiuno-
coenis was the only Australian genus in the Caenidae
but records of nymphs ol Caenis sp, were made by
Timms (1974) in a benthic study of three South
Australian voleanic lakes. This record and all olhers
from South Australia belong to the one species, T
tillvardi.
364 REC,
Material Examined
SOUTH AUSTRALIA, South East: Drain L, Drain k,
Eastern Division Diversion Drain, Eight Mile Ck,
Hitchcock Drain, Mosquito Ck, Mt. Hape Drain,
Sutherland's Drain, Mt. Lofty Ranges: Deep Ck,
Eleanor R., Sturt R., Torrens R., Waite Institute Pond,
Fleurieu Peninsula; Anacotilla Ck, Carvakalinga Ck,
Deep Ck, Hindmarsh R., Inman R., Kangarilla Ck.
Lake Alexandrina: Tookayerla Ck, Yankalilla R,
Kangaroo Island: Breakneck R., Cygnet R,,
Grassy/Sheep Ck, North-East R,, Tin Hut/Bullock Ck,
South West R., South West Bay R. Southern Firoders
Ranges: Broughton R., Nectar Brook Ck, Ohlenmeyer
Reservoir, Rocky R., Schumacher Ck, Spring Ck.
Northern Flinders Ranges} Arkaba Ck, Arkaroola Ck,
Balcanoana Ck, Bendicuta Ck, Brachina Ck, Bunyerao
Ck, Elatina Ck, Bmmu Ck, Enorama Ck, Eregunda Ck,
Hot Springs (Paralanaj, Kanyaka Ck, Marolana Ck,
Maunt Chambers Ck, Nepouie Ck, Old Wirrealpa
Springs, Oraparina Ck, Oratunga Ck, Parachilna Ck,
Stubbs Waterhole, Willigan Ck, Teatree Ck, Warren
Gorge €C, Wilpena Ck, Wockerawirra Ck, Wooden-
dimna Ck, Eyré Peninsula: Old Woolshed Dam.
VICTORIA, Crawtord R,,, Darlors Ck, Gawkers Ck,
Glenelg R., Lake Wendouree (Ballarat), Mougl Emu
Ck, Pigeon Hole Ck, Rocklands Reservair, Stoke Ck,
Surrey R., Wando R., Wannon R., Wennicott R.
TASMANIA. Elizabeth R., Lagoon of Islands,
Macquarie R., Rileys Creek Reservoir, Geeveston.
ACKNOWLEDGMENTS
This work was supported by a Commonwealth of
Australia Post Graduate Research Award and by
research funds of the Zoology Department, University
of Adelaide Special thanks must go to my supervisor,
Dr J, E, Bishop for his valuable advice, encouragement,
and constructive criticism throughout this study.
Thanks also to Prof, W, D, Williams for his encourage-
ment and guidance, and to all people who assisted in
collecting material from throughout South Australia.
] would also like to thank my wife, for her support
during the study.
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TILLYARD, R. J. 1921. A new genus and species of May-Fly (Order
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TILLYARD, R. J. 1932. Kansas Permian insects. Pt. 15. The Order
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365
TILLYARD, R. J. 1936, The trout-food insects of Tasmania. Part
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366 REC. 8, AUST, MUS, 19 (17); 339-397 November, 1986
pa ao ore
SE ae ae ¢
FIG. 1. Atalophlebia australis. a-d, male imago: a, fore wing; b, hind wing; c. genitalia, ventral view; d, genitalia, lateral view, e-f, female
imago: e, abdominal segments 7-10, ventral view; f, abdominal segments 7-10, lateral view. g-i, male imago: g, fore claws; h, dorsal
abdominal colour pattern; i, ventral abdominal colour pattern. Scale line: a-f, h-i, | mm; g, 0.1 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 367
PIG. 2. Atulophlebia australis, mature nymph: a, fore legs b, fore claw; ¢, third abdominal gill; d, labrum, dorsal view; ¢, antero-median
emargination of labrum, enlarged; f, hypopharynx; w, labium, dorsal (left) and ventral views; h, left mandible, ventral view, i, left
incisors and prostheca, enlarged; j, right mandible, ventral view; k, right incisors and prostheca, enlarged; |, right maailla, ventral
view. Scale lines: a, c, d. f, g, hy j, 1, 0S mm; b, e, i, Kk, 0.1 mm,
368 REC. S. AUST, MUS. 19 (17); 339-397 November, 1986
YH
FIG. 3. Atalophlebia australasica, a-d, male imago: a, fore wing; b, hind wing; c, genitalia, ventral view; d, genitalia, lateral view. e-f,
female imago: e, abdominal segments 7-10, ventral view; f, abdominal segments 7-10, lateral view. g-i, male imago: g, fore claw; h,
dorsal abdominal colour pattern; i, ventral abdominal colour pattern. Scale line: a-f, h-i, | mm; g, 0.1 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 369
ee
FIG. 4. Atalophlebia australasica, mature nymph: a, fore leg; b, fore claw; ¢, third abdominal gill; d, labrum, dorsal view; ¢, antero-median
emaryination of labrum, enlarged; f, hypopharyny; g, labium, dorsal (left) and ventral views; h, lett mandible, ventral view; 1, lett
incisors and prostheca, enlarged: j, right mandible, ventral view; k, right incisors and prastheca enlarged; |, right maxilla, ventral
view, Scale line: a, ¢, d, f, 2, hy j. 1, 0.5 mm: b, eg i, ky O.L mm.
370 REC. S. AUST, MUS. 19 (17): 339-397 November, 1986
——_5 4 } —__
FIG. $. Atalophlebia auratus. a-h, male imago: a, fore wing; b, hind wing; c, genitalia, ventral view; d, genitalia, lateral view; ¢, thoracic
sterna; f, dorsal colour pattern; g, ventral abdominal colour pattern; h, fore claw. i-j, female imago: i, abdominal segments 7-10,
ventral view; j, abdominal segments 7-10, lateral view, Scale lines: a-g, i, j, | mm; h, 0.1 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 371
VIG, 6, Atalophlebia auratus, mature nymph: a, fore leg; b, fore claw; c, labrum, dorsal view; d, antero-median emargination of labrum,
enlarged; e, hypopharnyx; f, labium, dorsal (left) and ventral views; g, left mandible, ventral view; h, left incisors and prostheca,
enlarged; i, right mandible, ventral view; j, right incisors and prostheca, enlarged; k, left maxilla, ventral view. Scale lines: a, c, e,
f, 2, i, k, 0.5 mm; b, d, h, j, 0.1 mm.
372 REC. S. AUST. MUS. 19 (17): 339-397 November, 1986
e _ |
FIG. 7. SEM micrographs of genitalia of Ata/lophlebia male imagos. a-b, Atalophlebia australis: a, genitalia, ventral view; b, penes, ventral
view, enlarged. c-d, Ataloniilebia australasica: c, genitalia, ventral view; d, penes, ventral view, enlarged. e-f, Atalophlebia auratus:
e, genitalia, ventral view; 1, penes, ventral view, enlarged. Scale lines: 100 um.
EPHEMEROPTERA OF SOUTH AUSTRALIA 373
FIG. 8. Nousia inconspicua. a-d, male imago: a, fore wing; b, hind wing; c, genitalia, ventral view; d, genitalia, lateral view. e-f, female
imago: e, abdominal segment 10, ventral view; f, abdominal segments 7-10, lateral view. g-i, male imago: g, fore claws; h, dorsal abdominal
colour pattern; i, ventral abdominal colour pattern. Scale lines: a, h, i, | mm; b-f, 0.5 mm; g, 0.05 mm.
374 REC. 8. AUST. MUS. 19 (17); 339-397 November, 1986
P1G, 9. Nousia inconspicuva, mature nymph: a, fore leg; b, fore claw; ¢, third abdominal gill; d, labrum, dorsal view; e, antero-median
emargination of labrum, enlarged; f, hypopharynx; g, labium, dorsal (left) and ventral views: h, left mandible, ventral view; i, left
incisors and prostheea, enlarged; i right mandible, ventral view; k, right incisors and prostheca, enlarged; I, left maxilla, ventral view.
Seale lines: a, c, O.S mm; b, d, f, g, hy, j, |, OL mm; e, i, k, 0,0S mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 375
SSS SS
iG. 10, Nousia fuscula. a-d, male imago: a, fore wing; b, hind wing; c, genitalia, ventral view; d, genitalia, lateral view. e-f, female imago:
e, ventral view of abdominal segment 10; f, abdominal segments 6-10, lateral view. g-j, male imago: g, fore claws; h, dorsal abdominal
colour pattern; i, ventral abdominal colour pattern; j, penes enlarged, ventral view. Scale lines: a, h, i, 1 mm; b, ¢, d, e, f, 0.5 mm;
J, 0.1 mm; g, 0.05 mm.
376 REC. S. AUST, MUS, 19 (17); 339-397 November, 1986
FIG. 11. Nousia fuseula, mature nymph: a, fore leg; b, fore claw; ¢, third abdominal gill; d, labrum, dorsal view; e, antero-median emargination
of labrum, enlarged; f, hypopharynx; v, labium, dorsal (left) and ventral views; h, left mandible, ventral view; i, left incisors and
prostheca, enlarged; j, right mandible, ventral view; k, right incisors and prostheca, enlarged; 1, left maxilla, ventral view. Scale lines:
a, ¢, 0.S mm; b, d, f, g, hy j, 1, O.L mm; e, i, k, 0.05 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 377
FIG. 12. Nousia pilosa. a-h, male imago: a, fore wing; b, hind wing; c, left hind wing, enlarged; d, genitalia, ventral view; e, genitalia,
lateral view; f, fore claws; g, dorsal abdominal colour pattern; h, ventral abdominal colour pattern. Scale lines: a, b, g, h, | mm;
c-e, 0.5 mm; f, 0.1 mm.
378 REC, S. AUST. MUS. 19 (17): 339-397 November, 1986
FIG, 13, Nousia pilosa, mature nymph: a, fore leg; b, fore claw; c, third abdominal gill; d, labrum, dorsal view; e, antero-median emargination
of labrum, enlarged; f, hypopharynx; g, labium, dorsal (left) and ventral views; h, left mandible, ventral view; i, left incisors and
prostheca, enlarged; i left prostheca, enlarged; j, right mandible, ventral view; k, right incisors and prostheca, enlarged; k', right
prostheca, enlarged; |, left maxilla, ventral view. Scale lines: a, c, d, f, g, h, j, 1, 0.S mm; b, e, i, k, 0.1 mm; i, k, 0.05 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 379
FIG. 14. Dorsal colour patterns of Aralophlebia and Nousia mature nymphs. a, Atalophiebia australis; b, Atalophlebia australasica; ¢,
Ajalophlebia auratus; d, Nousia inconspicua; e, Nousia fuscula; {, Nousia pilosa. Scale lines: | mm.
380 REC. S. AUST. MUS. 19 (17): 339-397 November, 1986
FIG. 15. SEM micrographs of genitalia of Nousia male imagos. a-b, Nousia inconspicua: a, genitalia, ventral view; b, penes, ventral view,
enlarged. c-d, Nousia fuscula: c, genitalia, ventral view; d, penes, ventral view, enlarged. e-f, Nousia pilosa: e, penes, ventral view,
enlarged; f, genitalia, ventral view. Scale lines: a, c, f, 100 wm; b, d, e, 10 wm.
EPHEMEROPTERA OF SOUTH AUSTRALIA
b ne
FIG. 16. SEM micrographs of eggs of a, Nousia inconspicua; and
b, Nousia fuscula, illustrating the general similarity of
morphology, but distinct polar caps of each species. Scale line:
10 um.
REC, S, AUST. MUS. 19 (17): 339-397
tiG, 17, Ulmerophlebia pipinna, male imago
sterna
go: a, fore wing; b, hind wing; c, genitalia, ventral view; d, geni atera +B
na; f, dorsal colour pattern; g, ventral abdominal colour pattern; h, fore claws. Scale lines: a-g, 1 mm; h, 0.5 mm
EPHEMEROPTERA OF SOUTH AUSTRALIA 383
PIG. 18. Ulnerophlebia pipinna, mature nymph: a, fore legs b, fore claw; ¢, third abdominal gills d, labrum, dorsal view; e, anlero-median
emarginaltion of labrum, enlarged; f, hypopharynx; g, labium, dorsal (left) and ventral views; h, left mandible, ventral view; 1, left
incisors and prostheea, enlarged and prostheca, enlarged; j, right mandible, ventral view; k, right incisors and prostheca, enlarged
and prostheca, enlarged; 1, left maxilla, ventral view. Scale lines: a, ¢, dy f, 2, hy J, 1, 0.5 mm; b, ei, k, 0.1 mm; prostheca, 0.05 mim.
384 REC. S. AUST. MUS. 19 (17): 339-397 November, 1986
FIG. 19. SEM micrographs of some Australian mayflies. a-b, U/merophlebia pipinna: a, genitalia, ventral view; b, enlarged ventral view
of penes. c-e, ventral view of genitalia of c, Baetis soror; d, Centroptilum elongatum, e, Cloeon fluviatile; f, Cloeon paradieniensis.
Scale lines: 100 pm.
FIG. 20.
EPHEMEROPTERA OF SOUTH AUSTRALIA
Dorsal colour pattern of mature female nymph of U/merophlebia pipinna. Scale line: | mm.
385
386 REC, S, AUST. MUS, 19 (17): 339-397 November, 1986
FIG, 21, Baeris soror. a-d, male imago: a, fore wing; b, hind wing; c, dorsal colour pattern; d, genitalia, ventral view. e-1, mature nymph:
¢, dorsal colour pattern; f, fore leg; g, fore claw; h, paraprocts, ventral view; i, third abdominal gill; i, margin of gill, enlarged; j,
labrum, dorsal view; k, antero-median emargination of labrum, enlarged; |, left maxilla, ventral view; m, labium, dorsal (left) and
ventral views; n, left mandible, ventral view; 0, left incisors and prostheca, enlarged; p, right mandible, ventral view; q, right incisors
and prostheca, enlarged; r, hypopharynx. Scale lines: a, b,c, e, 1 mm; d, f, i, 0.S mm; g, h, i, j-r, 0. mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 387
PIG. 22. SEM micrographs of the paraprocts of aymplis of Bueris soror (a, b); Centropiiium elongutum (e, d) and Tasmanophiebig sp.
(c.f), Seale lines: 100 ym.
388 REC. S. AUST. MUS. 19 (17): 339-397 November, 1986
SS ~
FIG. 23. Centroptilum elongatum. a-e, male imago: a, fore wing; b, hind wing; c, dorsal colour pattern; d, genitalia, ventral view; e, thoracic
sterna. f, mature nymph: dorsal colour pattern. Scale lines: a, b, c, e, f, 1 mm; d, 0,5 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 389
FIG. 24. Centroptilum elongatum, mature nymph: a, fore leg; b, paraprocts, ventral view; c, paraprocts, enlarged; d, third abdominal
vill; ¢, labrum, dorsal view; f, antero-median emargination of labrum, enlarged; g, labium, dorsal (left) and ventral views; h, hypopharynx;
i, left mandible, ventral view; j, left incisors and prostheca, enlarged; k, right mandible, ventral view; |, right incisors and prostheca,
enlarged; m, right maxilla, ventral view. Scale lines: a, b, d, 0.5 mm; ¢, d, e, f, 2g, h, i, k, m, 0.1 mm; j, 1, 0.05 mm.
390 REC, 8S. AUST. MUS. 19 (17): 339-397 November, 1986
VIG. 25. Cloeon fluviatile. a-f, male imago: a, fore wing; b, dorsal colour pattern; c, thoracic sterna; d, genitalia, ventral view; e, genitalia,
dorsal view; f, genitalia, lateral view. Cloeon paradieniensis. g-k, male imago: g, fore wing; h, dorsal colour pattern; i, genitalia,
ventral view; j, genitalia, lateral view; k, genitalia, dorsal view. Scale lines: a, b, c, g-k, | mm; d, e, f, 0.5 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 39]
FIG, 26. Cloeon fluviatile, mature nymph: a, fore leg; b, fore claw; c, dorsal abdominal colour pattern; d, paraprocts; e, third abdominal
gill; f, labrum, dorsal view; g, antero-median emargination of labrum; h, labium, dorsal (left) and ventral views; i, left maxilla, ventral
view; j, left mandible, ventral view; k, left prostheca, enlarged; 1, right mandible, ventral view; m, left prostheca, enlarged; n, hypopharynx.
Scale lines: ¢, 1 mm; a, e, 0.5 mm; b, d, f-j, 1, n, 0.L mm; k, m, 0.05 mm.
392 REC. S. AUST. MUS. 19 (17); 339-397 November, 1986
FIG, 27. Cloeon paradieniensis, mature nymph: a, fore leg; b, paraprocts; c, labrum, dorsal view; d, antero-median emargination of labrum;
e, third abdominal gill; f, right maxilla, ventral view, with enlarged apex of terminal segment of the palp; g, hypopharynx; h, labium,
dorsal (left) and ventral views; i, left mandible, ventral view; j, left incisors and prostheca, enlarged; k, right mandible, ventral view:
|, right incisors and prostheca, enlarged. Scale lines: a, e, 0.5 mm; b-d, f-i, k, 0.1 mm; j, 1, 0.05 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA
pac eae PN
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wa es 8 oes ae
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Pi
FIG. 28, Tasmanophlebia sp. a-f, ma
le imago: a, fore wing; b, hind wing; c, dorsal abdominal colour pattern; d, genitalia, ventral view;
e, genitalia, lateral view; f, fore claws. g, mature nymph: dorsal colour pattern. Scale lines: a-e, g, 1 mm; f, 0.1 mm.
394 REC. S. AUST. MUS, 19 (17); 339-397 November, 1986
FIG. 29, Tusmanophlehia sp. mature nymph: a, fore leg; b, first abdominal gill; c, second abdominal gill; d, third abdominal gill} e, fourth
abdominal gill; f, labrum, dorsal view; g, antero-median emargination, enlarged; h, labium, dorsal (left) and ventral views; i, hypopharynx;
j, left maxilla, ventral view; k, left mandible, ventral view; I, left incisors and prostheca, enlarged; m, right mandible, ventral view;
n, right incisors and prostheca, enlarged. Scale lines: b-e, | mm; a, f, h-k, m, 0.5 mm; g, |, n, 0.1 mm,
ft)
EPHEMEROPTERA OF SOUTH AUSTRALIA 395
FIG. 30. SEM micrographs of Tasmanophlebia sp. and Tasmanocoenis tillyardi. Tasmanophlebia sp. a, b, genitalia of male imago, ventral
views. Tasmanocoenis tillyardi. c, genitalia of male imago, ventral view; d, second gill of mature nymph. Scale lines: 100 um.
396 REC. 8. AUST. MUS. 19 (17): 339-397 November, 1986
FIG. 31. Tusmnanocoenis tillyardi, a-d, male imago: a, fore wing; b, dorsal colour pattern; c, thoracic sterna; d, genitalia, ventral view.
e, mature nymph: dorsal colour pattern. Scale lines: a-c, e, | mm; d, 0.1 mm.
EPHEMEROPTERA OF SOUTH AUSTRALIA 397
FIG. 32. Jasmanocoenis tillyardi, mature nymph: a, fore leg; b, first abdominal gill; c, second abdominal gill; d, third abdominal gill;
e, basal antennal segments; f, labrum, dorsal view; g, antero-median emargination of labrum, enlarged; h, labium, dorsal (left) and
ventral views; i, hypopharynx; j, left mandible, ventral view; k, left incisors and prostheca, enlarged; |, right mandible, ventral view;
m, right incisors and prostheca, enlarged; n, left maxilla, ventral view. Scale lines: a-e, 0.5 mm; f-n, 0.1 mm.
FIRST REPRESENTATIVE OF THE ORDER MACROSTOMIDA IN
AUSTRALIA (PLATYHELMINTHES, MACROSTOMIDAE)
BY RONALD SLUYS
Summary
A new species of macrostomid flatworm is described, Promacrostomum palum sp. nov., forming
the third member of its genus and being the first representative of the order Macrostomida to be
reported for Australia.
FIRST REPRESENTATIVE OF THE ORDER MACROSTOMIDA IN AUSTRALIA
(PLATYHELMINTHES, MACROSTOMIDAE)
by
RONALD SLUYS
Instituie of Taxonomic Zoology, Universily of Amsterdam, P.O, Box 20125, 1000 HC Amsterdam,
The Netherlands.
(Manuscript accepted 6 January 1986)
ABSTRACT
SLUYS, R, 1986. First representutive of the order Macrostomida tn
Australia (Platyhelminthes, Macrostomidae). Ree. 8. dust, Atus.
1918); 399-404,
A new species of macrostomid flatworm is described,
Promacrostomum paluim sp, nov,, forming the third
member of its genus and being the first representative
of the order Macrostomida to be reported for Australia.
INTRODUCTION
Macrostomid flatworms have been reported from all
major parts of the world, except for Australia and New
Zealand (cf, Ferguson 1939, Map 2; Ferguson 1954,
Table 1; Williams 1980, p. 52). The majority of the
species within the family Macrostomidae belong to the
large genus Macrosiomum ©. Schmidt, 1848. The
present paper desvribes a new macrostomid species,
which was found in Australia. The species belongs ta
a venus which has a different and more complex female
copulatory apparatus than is the case in Macros/omum.
Recent literature on the Macrostomidae is rather
scattered but the papers of Ferguson (1939-40, 1954)
still represent a useful introduction, whereas those of
Papi (1953), Luther (1960) and Young (1976) are some
of the larger papers among more recent publications,
SYSTEMATIC SECTION
Family MACROSTOMIDAEL Van Beneden, 1870
Genus Promacrostomuim An-der-Lan, 1939
Promacrostomum palum sp. noy.
Material Exumined
Holotype: South Australian Museum, Adelaide,
V3973, Elizabeth Springs, South Australia (29°21.36
S, 136''46.30’E), 27.11.1983, coll, W. Zeidler, W. Ponder,
sagittal sections an two slides,
Paratypes: SAM, V3974, ibid., horizontal sections
on one slide; SAM, V3975, ibid., transverse sections
on one slide; Australian Museum, Sydney, W197775-1,
Elizabeth Springs, South Australia, 5,09.1983, coll. W.
Ponder, E. Hershler, D. Winn, sagittal sections on one
slide; AM, W197775-2, ibid., horizontal sections on one
slide,
The holotype was sectioned at intervals of 5 ym; the
paratypes at 8 pm. All sections Were stained in Mallory-
Heidenhain.
Etymology
The specifie epithet is from the Latin pala (= spade)
and refers to the shape of the hind end of the body,
Description
External Features
The preserved specimens measured 2,38-3,5 mm itt
length and 0.75-1 mm in diameter. In some specimens
of sample AM W197775 the front end of the body was
pointed, but in others and in specimens from SAM
V3973-75 it was broadly rounded (Figs 1, 2), The hind
end of the body is of a peculiar shape. [In preserved
Specimens the posterior lateral margins give rise lo a
dorsally directed ridge at cither side of the bady; the
posterior margin of the body shows a convex middle
section (Figs 1, 2, 3). The preserved animals-are devoid
of pigment, Eyes were not visible in preserved
specimens but only in animals cleared in clove oil.
Epidermis and Subepidermal Musculature
The height of the epidermal cells is about 11.5 pm;
the cells are provided with numerous cilia which have
a length of about 9 pm, Numerous and well-developed
packages of rhabdites pierce the body wall, whereas
rhabdite-tracks (“Stabchenstrassen”) are present at the
front end. “Haftpapillen” are absent, A cyanophilous,
granular secretion is discharged through the ventral
epidermis at the posterior tip of the body. The gland
cells are situated in the parenchyma.
The subepithelial musculature consists of outer
circular muscle fibres and inner longitudinal fibres.
Nervous System and Eyes
The brain lies just in front of the pharynx and is
closer to the ventral than to the dorsal body surface.
The two eyes are situated just on top of the brain.
Alimentary System
The thin epithelium lining the simple pharynx bears
numerous well-developed cilia, Erythrophilous gland
cells surround the pharynx where it communicates with
the intestine, The intestinal cells are provided with long,
but not very conspicuous, cilia, The inmestine ts
underlain with a well-developed row of circular muscle
400 REC. S. AUST. MUS, 19 (18): 399-404 November, 1986
FIGS 1-6. Promacrostomum palum sp. nov. 1, Dorsal view of preserved specimen from sample W197775. 2. Dorsal-lateral view of preserved
specimen from sample V3973-75. 3. Lateral view of the hind end of the body. 4. Sagittal reconstruction of holotype to show the position
of the male and female copulatory organs and the intestine. 5. Horizontal section of V3974, 6, Horizontal reconstruction of the male
copulatory organs of V3974.
ORDER MACROSTOMIDA IN CAUSTRALIA 401
fibres which is bounded by an outer row of longitudinal
libres. The sac-shaped ftestine extends backwards to
the male copulatory apparatus, and (hus runs over the
female copularory system (Pig. 4).
Male Reproductive System
The two elongated testes are situated ventrally and
shortly behind the pharynx, ane follicle af either side
of the body (Fig. 5). The vasa delerentia open behind
the posterior tip of the intestine, into a large false
seminal vesicle (Pig, 6). The false seminal vesicle
communicales wilh the (rue seminal vesicle by means
of aconsiderable, muscularized constriction. This true
seminal vesicle lies partly underneath the false seminal
vesicle; the former is ati clongated sae which is provided
with a very thick muscular wall. A short aid narrow
intervesicular duct connects the irue seminal vesicle
with the rounded granular vesicle which receives the
vranular secretion of glands that lie distributed in the
parenchyma. The wall of the yvramilar vesicle js
provided with a thin layer of muscles.
The proximal portion of the “vhitinized” penis stvlet
14 allached to the granular vesicle and consists of a
broad cone that runs parallel to the body surlace and
Which tapers into the much narrower distal section of
the stylet. The distal section shows-a pronounced bend
towards (he ventral body surface (Fig. 7). Sagicral
sevlions already sugested the presence of a lateral
flexure in the very distal portion of the styler. This
Hlesure did indeed show up in the transverse sections
(Fig, 8). No unequivocal information could be oblamed
on the apening of the stylet. The sagittal sections
suggested that the tip was closed such that there was
a subrerminal opening. On the other hand, the
horizantal sevlions suggested the presence of a lerminial
opening, whereas in the transverse sections the tip of
the stylet could not be discerned.
Female Reproductive Systen
The ovaries lie direcUy behind the testes and are
situated ventrally; they are rather small (ig. 5). The
oviduets do not extend backwards but run directly
towards the mid-line of (he veurral body region, where
they unite info a short common oviduct, Che latter
opens into the female anirum. There is a [ree
connection berween common oviduct and femule
antrum; a so-valled “Verschlussapparat” is absent
(Migs 9, 10).
The female antrum is lined with more or less
cuboidal cells which bear long cilia. In specimen V3975
a Jatge and thick mass of cells was allached to the
dorsal surface of the antrum leaving at one place a
small opening lor (he common oviduer. Sperm were
allached to the clump of cells (Fig, 9). The antrum
opens via a ciliated stalk into (he ventral exterior. The
lining epithelium of the stalk is penetrated by humerous
openings of erythrophilous cerment glands which are
distributed in the parenchyma around the lemale
antrum. Che posrerior wall of the female antrum meets
a 40-called senimal bursa. Communication. between
anirum and bursa may iake place either via a
considerable narrowing, 48 is the case, for example, in
specimen W197775-1 (Fig. 11) and V3975, of via a
niuch wider opening (Fig. 10). The seminal bursa 1s
lined with tall, vacuolated, cells, whereas its lumen may
contain sperm, The bursa communicates via a narrow
veniral pore with the exterior, Female antrum ane
seminal bursa dre surrounded by a well-developed layer
of muscles.
COMPARATIVE DISCUSSION
The senus Promacrostomuim at present contains
only three species, inchiding the one deseribed in this
paper. The type species BP puradoxnin was described
from Lake Ohrid, Yugoslavia (An-der-Lan 1939).
On the basis of a description by Gieysztor (1931),
Ferguson (1939-40) deseribed the species Mucrosromurh
gievsziori which he showed to be different trom Ad
eracile (Pereyslawzewa, 1892; Grall, 1905) to which the
first-mentioned author had assigned (he specimens
collected. Papi (1951a) transferred the species to the
genus Pramacrostamiun, but Ferguson (1954) erected
the genus Axia for the species gievssrart. The new
generic name “Lara, however, has not been accepted
since iis not used io recent literature. ? gievssrarl has
been found in Spain and Italy; it is a thermophilic
freshwater species (Ferguson 1939-40, 954; Papi
195th),
P vievsziori has a distinct common oviduct Which
empnes. into a structure (hat possesses two openings
to the exterior (ef. Papi 1951b, Fig, 54; Ferguson 1954,
Vig, 23), The aiterior portion of this body is rather
spacious and seerelion of cement vlands is discharped
into its vertral opening, The major part of this anterior
structure is lined with cells bearme long cilia. Because
of its shape, distinet cilidtion, and the presence of
cement glands around its pore, | consider this anterior
section of the female copulalory apparatus of P
gievszrarito be homologous on the one hand with the
fermale antrum of Maerostamiuin species, and on the
other hand with the antrum of PB palin, As a
consequence, (he most anterior pore in A gievssreri
corresponds with the female gonopore.
The second opening of the female reproductive
sysiem in P vieysctori lies posteriorly (o the female
sonopore and leads into the narrower posterior portion
of the female apparatus, of which (he ventral section
is lined with cells having long cilia. The dorsal portion
of this posterior part of the female apparatus is
conneeted ta the dorsal portion of the female antrum
by means of a curved duct. The non-ehated lining of
this wide duct varies in height and vonsists af a
syneviial cell mass; the duct is surrounded by a rather
iick layer of circular muscles (cl. Papi lY51b, Viv, 34),
Iris because of structural and positional similarities
that | cansider this. duet jo be homologous with the
seminal bursa of 2 palin. For the same reasons |
consider the posterior female pore of B gievsstary to
be hamologous with the bursal pore of Ro peli,
402 REC. S. AUST, MUS. 19 (18): 399-404 November 1986
in
| inna $ fi]
Pretty Lh |
Jhigt
bpo sp sb fgo
FIGS 7-1. Promucrostomum palum sp. nov. 7. Sagittal reconstruction of the male copulatory system of W197775-1. 8. Transversal
reconstruction of the penis stylet of V3975. 9. Transversal reconstruction of the female copulatory apparatus of ¥3975 at the level of
the female antrum, 10, Sagittal reconstruction of the female copulatory apparatus of the holotype. I]. Sagittal reconstruction of the
female copulatory organs of W197775-1.
ORDER MACKOSTOMIDA IN AUSTRALIA
Urifortunately, less detailed histological information
is available on the female copulatory apparatus of P
paradoxunt, In this species too, the female apparatus
has two ventral Openings to the exterior. The posterior
pore leads into a well-developed lemale antrum. The
anierior pore leads into a section of the female
copulatory apparatus that communicates anteriorly
with the intestine and posteriorly with the female
antrum. The connection wilh the lemale antrum goes
via 4 rather spacious “middle section” of the female
apparatus. This “middle section” is surrounded by well-
developed circular muscles, The female antrum and
“middle section” are lined with a relatively tall, almost
synevtial, epithelium (An-derLan 1939; Ferguson
1954),
It is evident from An-der-Lan's account that he
cansidered the female antrum of PB paradoxum to be
homologous with the atrium af Macrostemum and
considered the posteriorly located pore of the former
to be homologous with the single female genital pore
in Afacrosfamunt species. I agree with An-der-Lan's
conclusion, although mo data are available on the
presence or absence al cement glands around the
posterior female genital pore of PR patadoxuni or on
the cilianhon of the female antrum, and oviducts were
absent in rhe specimens examined by An-der-Lan, My
view on this subject is based only on the position and
the overall shape of the female antrum (cf, An-der-Lan
1939, Fig. 3). Nevertheless, | postulate homologous
relationships berween the posteriorly located female
gonopore in & parudoxum, the anterior female genital
pore in P gievssrori and P palin, and the single fernale
gonapore in Macrostomum. Such relationships are, of
course, also hypothesized for the female antra into
whieh these pores lead,
It is even more difficult to evaluate the possible
relationships of the anterior pore and the “middle
section” of the female copulatory apparatus in P
purudoxunmt with structures in other macrostomids.
Because of the poor slate of thal particular section of
the preparations, An-derLan (1939) was unable to
provide informarion on the histology of rhat portion
of the female copulatory apparatus which communi-
cates with the anterior pare, Therefore, it is only
because of ils pasition that J consider this anterior pore
in P. partdoxum to be dilferent (ram the posterior
female pore in 2 vievszrarcon (he one hand, and from
the bursal pore in R pa/um an the other hand.
IF the above interpretation is correct, then bath &
paluinvand P eieysztari differ fram PB paradaxum in
thal in both the first-mentioned species the “secandary”
pore ts sttualed behind the lemale ponopore, whereas
in the latter it lies in. front of the “primary” female pore
(“primary” pore refers to that opering into which
cement vlands discharge — as in Mucrosroniunt — no
phylogenetic meaning js. implied),
P palum differs trom the two other members of the
genus Promacrosiemun ia the shape of its tail, stylet
Ws
and that of the ferriale copulatory apparatus (ef. Papi
195]b, Pigs 52-54; Ferguson 1954, Figs 20-24).
Apart from similarities in the female copularory
apparatus, 2 palumand FP giepsstori also agrce in the
absence of “Haltpapillen” and in the lact that the
Jntestine runs over the female copularory apparatus (cf.
Ferguson 1939-40, 1954; Papi 195|/b). In B paradoxum
the intestine terminates in front of the female
copulatory apparatus and the species possesses well-
developed “Haftpapillen” (An-der-Lan 1939).
From the account given above, one could conclude
that PB gievstlor? and P palum baye many more
morphological similarities in common than either of
them shares with P paradoxun). In that respect the
genus name “xia, as proposed by Ferguson (1954), may
form indeed a plausible allernalive lor expressing the
differences between P gleyszrari and P palum on the
one hand, and 2 paradoxum on the other hand.
However, absence of detailed histological information
on the lemale system of A paradoxum prevents a
proper assessment of homologies. Pending such
histological information it seenis best, for rhe moment,
to assign P gievsztori and P palin to the same genus
as P paradoxurn.
‘The structure of the female copulatory apparatus in
Promacrostomunt inyates a comparison with quite
another member of the Macrostomida, via. a represen
lative of the family Micrastamidae, In contrast to all
other Microstoniam species, the female apparatus of
M. spiriferuny Westblad, 1953 opens ta the exterior hy
means Of two ventral pores, These openings lead into
two separate ducts, both connected with the single
ovary. The most anterior duct ts ciliated and opens inta
a small, ciliated chamber which is separated From the
ovary by a sphincter. The posterior duet is non-ciliated
and leads into a small seminal bursa which opens inte
the ovary (cf Westblad 1953, Fig. 40). Aceording to
Westhlad (/.¢,) the anterior duct carresponds with the
usual vagina but functions in AZ. spiriferuin only as
a discharging duct for the fertilized eygs. The posterior
female duct would function as copulatory duet. Ie may
be that the same functional distinction between both
female pores holds truc at least for P pola and
P gievsztori, If such ts the case, then it is clear that
the same end result has been reached through parallel
evolution.
It is evident from Papi’s (1951b) account thar he
assumed such a functional distinction 10 be presene in
P. vievsztori, Papi described the female antrum of 1s
species under the name of “antrum ovipositorum"™,
Whereas he designated the female gonopore as “poris
ovipositorius”, The posterior female pore in
gievsstor/ leads, according to Papi, into the “vagina”.
Although [ do not question the plausibiliry of such a
funclienal distinenon and even think if to be very likely,
| have avoided terms such as vagina and antrum
ovipositorum in the deseription of either B palin ot
P paradexyum and P vievsslerr. In the nomenclature
404 REC. S. AUST. MUS. 19 (18); 399-404
of the various morphological features of the female
apparatus I tried to use the same name for structures
considered to be homologous in Promacrostomum and
Macrostomum.
Ball (private communication) during his extensive
collecting of freshwater planarians in Australia
(1982-1984) has recorded specimens of macrostomids
from caves in Yanchep National Park, Western
Australia but not in surface water. Therefore, it is
Noveniber, 1986
interesting that the first unequivocal record of a
macrostomid turbellarian in Australia, Promacro-
stomum palum sp. nov., is also from a specialized
habitat, viz. freshwater springs. Whether this situation
represents an artifact or a real phenomenon remains
an open question since in Australia scarcity of serious
records for both freshwater and marine macrostomids
also results from a lack of specialized interest in this -
group of animals.
REFERENCES
AN-DER-LAN, H. 1939. Zur rhabdocoelen Turbellarienfauna des
Ochridasees (Balkan). Sitz. Ber. Akad. Wiss. Wien, Mat.
-nalurw, Kl. Abt. 1, 148: 195-254,
FERGUSON, . PF. 1939, A monograph of the genus Macrosfomum
O. Schmidt, 1848, Parts LV. Zool Anz. 126; 7-20, 127: 131-144,
128: 49-68, 128: 180-205, 128: 274-291,
FERGUSON, F. lk. 1940, A monograph of the genus Macrostomum
O. Schmidt, 1848. Parts VI-VITL, Zool, Anz, 129: 21-48, 129:
120-146, 129: 244-268.
FERGUSON, F. F. 1954. Monograph of macrostomine worms of
Turbellaria, Trans. Amer. Microsc. Soc. 73: 137-164.
GIEYSZTOR, M. 1931. Contribution 4 la connaissance des
Turbellaries Rhabdoceles (Turbellaria Rhabdocoela) d’ Espagne.
Bull. Acad. Polon. Sc. Lettr. Cl. Sc. Mathem, Nat., Ser. B: Se.
Nat., (LI): 125-153 (not seen in original, cited in Ferguson
1939-40, 1954).
ABBREVIATIONS IN THE FIGURES
bpo bursal pore
cel cement gland
cod common oviduct
fa female antrum
fgo female genital pore
fsy false seminal vesicle
gl gland
gv granular vesicle
in intestine
ivd intervesicular duct
If lateral flexure
mgo male gonopore
od = oviduet
ov ovary
ph pharynx
ps penis stylet
sb seminal bursa
se secretion
sp sperm
sv serninal vesicle
te testes
vd vas deferens
LUTHER, A, 1960. Die Turbellarien Ostfennoskandiens |. Fauna
Fennica 7: 1-55.
PAPI, F, 195la, Sulle aflinita morfologiche nella fam. Macrostomidae
(Turbellaria). Boll, Zoologia 17 (Suppl.): 461-468.
PAPI, F. 1951b. Richerche sui Turbellari Macrostomidae. Arch. Zool.
Tral. 36: 289-340 + 1 pl.
PAPI, PF. 1953. Beitrage zur Kenntnis der Macrostomiden
(Turbellarien). Acta. Zool. Fenn, 78: 1-32,
WESTBLAD, E. 1953, Marine Macrostomida (Turbellaria) from
Scandinavia and England. Arkiv for Zoologi Bd. 4, nr, 23:
391-408,
WILLIAMS, W. D. 1980. Australian Freshwater Life. MacMillan
Comp., Melbourne. 321 pp:
YOUNG, J. O. 1976. Systematic studies on limnic Macrostomuni
species (Turbellaria, Macrostomida) from East Africa, Zool, Ser,
5; 49-60.
A REASSESSMENT OF THE PAPUAN SUBFAMILY ASTEROPHRYINAE
(ANURA: MICROHYLIDAE)
BY THOMAS C. BURTON
Summary
Information obtained from examination of the osteology, myology and external morphology is used
in a phylogenetic analysis to assess the relationships in the Asterophryinae, a subfamily of
terrestrial and fossorial microhylid frogs restricted to the Papuan Subregion, and to assess the
relationships between the Asterophryinae and the other Papuan subfamily, Genyophyrninae. While
the Asterophyrinae is monophyletic, no evidence of monophyly of the Genyophryninnnae is found.
Taxonomic changes are made and all genera redefined in light of the phylogenetic analysis: four
new tribes are erected to accommodate monophyletic groups of asterophryine genera; the genus
Mantophyne Boulenger is resurrected to accommodate three species currently included in
Phrynomantis; and Xenorhina doriae is transferred to Phrynomantis.
A REASSESSMENT OF THE PAPUAN SUBFAMILY ASTEROPHRYINAE (ANURA: MICROHYLIDAE)
THOMAS C. BURTON
Department of Zoology, University of Adclaide, Adelaide, South Australia 5000
(Manuscript accepted 1 April 1986)
ABSTRACT
BURTON, 7. C. A peupssessment of the Papinor sublamily Astero-
phrvinne (Anira: Microhylidae). Ree. S, Aust Mus. L919):
405-450.
Information obtained from cxamination of the
osteology, myology and external morphology is used
in a phylogenetic analysis to assess the relationships
in the Asterophryinae, a subfamily of terrestrial and
fossorial microhylid frogs restricted to the Papuan
Subregion, and to assess the relationships between the
Asterophryinae and the other Papuan subfamily,
Genyophryninac. While the Asterophryinae is mono-
phyletic, no. evidence of monophyly of the
Genyophryninae is. found. Taxonomic changes are
made and all genera redefined in light of the
phylogenetic analysis: four new tribes are erected to
accommodate monophyletic groups of asterophryine
genera; the genus Mantophryne Boulenger is
resurrected to accommodate three species currently
included in Phrynemantis; and Xenorhina doriae is
transferred to Phryvnumantis,
FIG, 1. Distribution of the Microhylidae (adapted front Savage. 1973).
INTRODUCTION
The Microhylids of the Papuan Zoegeogruphic
Subregion
The frog family Microhylidae is distributed
predominantly in the tropical areas of Asia, Africa,
South America and the Australian Region, but with
representatives also in (he adjacent temperate areas of
Asia, Alrica and the Americas (Fig. 1). Frost (1985)
lists 281 mierohylid species, 61 genera and nine
subfamilies, and the mean numbers of species per
genus and of genera per subfamily are lower than in
any other large family ol lrogs, The morphological
diversity suggested by these figures reflects in part the
ecological diversity found within the Microhylidae.
There are fossorial, terrestrial, arboreal and aquatie
species (Tyler 1976a), sharing the microhylid features
of a firmisternal pectoral girdle, expanded sacral
diapophyses, palatal folds, posteriorly directed process
of the lower jaw (Roux 1944), .M. rectus abdominis pars
anteroflecta (Burton 1980) and a characteristic jawless
tadpole (Parker 1934).
The subfamilial classification of the Papuan
microhylids has long been contentious, In his
monograph of the microhylids, in which he established
the basis of the modern classification of the family,
Parker (1934) revognized two Papuan subfamilies,
Aslerophryinae and Sphenophryninae, He recognized
that these subfamilies were closely related and the only
microhylids sharing the characteristic of direc!
development; as the distinctions between them were
blurred by exceptions he expressed misgivings about
separating them taxonomically: (a) while all spheno-
phrynines possessed a procoelous backbone, and nearly
all of the asterophryines possessed a diplasiocoelous
backbone, one aslterophrying, Genvophryne thomsoni
exhibited the sphenophrynine condition; (b) while the
maxillae of all sphenophrynines were separated
anteriorly by the premaxillae (the “eleutherognathine”
condition), and nearly all of the asterophryines were
symphygnathine, i.e, the two maxillae were fused on
the midline anteriorly to the premaxillae, the three
members of the genus Mefopostira, Matra
(=Barveenvs atra), M. kopsteint ( +» Phrynomantis
kopsteini) and M. ocellata (= Nvlaphorhus rufescens))
and G. thomsoni exhibited the sphenophrynine
condition, Parker considered, but rejected the
advisability of classifying G. thomsoni as a
sphenophrynine. G. thamseni is a heavily built [rog
(as most asterophryines but few sphenophrynines are)
and Parker considered the tongue of G. thomsoni 10
resemble the pasteriarly adherent tongue of the astero-
phryines more than the spbhenophrynine tongue, which
is [ree posteriorly,
Zweifel (1971) and Savage (1973) independently
tackled the problem of the unsatisfactory distinction
between Asterophryinae and Sphenophryninae.
Savage’s solution was to declare the taxonomic
distinction between these two subfamilies of direct-
developing lrogs “invalid", At the same time, echoing
Noble (1931), he claimed that the Asian genus Callvella
406 REC. 8. AUST. MUS. [9 (19); 405-450
(which is not direct-developing) “appears to be a
primitive genus from which the more highly evolved
asterophryines [=Asterophryinae + Sphenophryninae]
may have developed", and so he included the
Sphenophryninae and Cul/vella (sensu Inger, 1967) in
an enlarged Asterophryinae,
Zweifel’s solution was to examine the morphology
of the animals, especially G, (Aurisoni, the species
which had provided much of the heterogeneity in
Parker’s Asterophryinac. He found thar in tongue
morphology as in other features, Genyophryne
resembled (the sphenophrynines more than the
asterophryines, and he transferred G. fhomsont into
Sphenophryninae. Recognition thal G, (Aomsoni was
more closely related to the sphenophrynines allowed
new diagnoses of two subfamilies which were
consistently distinct for two characters: (1)
Asterophryinae diplasiovoglous with adherent tongues,
(2) Genyophryninae (Sphenophryninae plus Genyo-
phryne) procoelous with tongues at least one-quarter
free posteriorly; and mostly consistent for two others
(almost all asterophryines with symphygnathine
maxillae and dentaries; all genyophrynines with
eleutherognathine maxillae and dentaries). (Zweilel
1981) and subsequent authors continued to refer to the
augmented Sphenophryninac as Sphenophryninac, but
Dubois (1985) indicates that Genophryninae Boulenger
1890 has priority over Sphenophryninae Noble 1931.)
Zweifel (1972) considered Genyophryninae the parent
group of the Asterophryinae. This implies that
Genyophryninae is nol a monophyletic sister-group but
a paraphyletic parent-group. Subsequently, Tyler (1979)
added to Zweilel’s list of characters separating the
subfamilies a character associated with the superficial
submandibular musculature, viz., overlap of the Mi.
interhvaideus and intermandihularis which he found
present in all but one of the asterophryine genera and
in no genyophrynines (seasu Zweifel, 1971) (Table 1),
Tyler (1979) concurred. with Zweitel’s view that the
Asteropliryinae arose oul of the Genyophryninae, and
cited his study of the submandibular muscles (Tyler
1974) in support of this position.
TABLE |, CHARACTERS TO DISTINGUISH THLE ASTERO-
PHRYINAF PROM THE CGCENYOPHRYNINAE (ALTER
TYL PR 1979)
Charueter Asterophryinae Genyophryninae
Olten overlapping
premaxillae, maxiliae,
and usually in
contact
Maxillac Not overlapping pre-
manillae. Never ity
contact medially
Denhanes In contitel anteriorly Not in comlitel
fexcept in /Tyla-
phorhus)
Di plisiocaelors Prowovlous
Oval, hallstree
behind, lacking
edie PUPOwW Avie
posterion pouch
Vertebral cola
Tongue Subeisvular, critirely
adherent), olen with
a median tirrow and
posterior poe
Does not underlie
intermeanelibularis
Mnlerfvaideus
Mmiscle
Anenoarky underlies
mrerniauadibularis
texeent in Ahi
PHOPAUS)
November, 1986
in this study [ follow Zweifel’s classification and the
terms “Asterophryinac” and “asterophryine” are used
in Zweitel’s restricted sense, The species and genera
which constitute the Asterophryinae are listed in
Table 2,
The Asterophryinae currently comprises 43 named
taxa in seven) genera. These species occur exclusively
within the Papuan Subregion, majnly on the island of
New Guinea, but some occur on islands from Seram,.
Amboijna and Halmahera in the west lo the Louisiade
Archipelago in the east. The Genyophryninae is also
centred upon the New Guinea mainland but ranges far
more widely, from the Philippines in the north to
tropical Australia in the south, and trom Sulawesi and
the Lesser Sunda Islands in the west to New Britain
and the Louisiade Archipelago in the east, All of the
six genyophryniné genera (Table 3) occur on the island
ol’ New Guinea, and only three of these (Cophixalus,
Oreophryne and Sphenophryne) occur elsewhere. OF
approximately 70 species, 50 have been recorded from
New Guinea (Zweifel and Tyler 1982).
TABLE 2. THE ASTEROPHRYINAE
Asteraphrys Gunther 1858
turpicula (Schlegel) 1837
Baryeenvs Parker 1936
arra (Oiinther) 896
cheesmanve Parker 1936
exsul Zweifel 1963
Slavivalaris Aweifel 972
maculata Mendies and Tyler 1977
nana Zweilel 1972
parvula Zweilel Hsu
[ylapharhus Macleay 1878
rufescens rufescens Miucleuy i878
hr eNtNUN AWwellel 1972
ro mvapicus Aweifel 1972
Pherohapses Aweilel 872
mrensiest Zweifel 1972
Phrynomantis Peters 1867
hoetigeri (Mehely) 190t
duhia (Bociigery 1895
en bdacivla Zweite) Y72
fusca Peters 1867
alandulosa Zweitel YT
lranveola humicola Zweilel 1972
hh. compte Aweilel 1972
infulata Aweitel 1972
kopstei (Mertens) 1930
leivratis (Boulenger) 18Y7
fowisiadensis (Parker) 1984
persondia “weilel 1972
rabiisia (Boulenger) [ROR
stutery (Loveridue) 1955
sHetnuaster éweilel 172
withelmana (Loveridge) 148
VYenobarrachus Peters and Doria (R78
Hidens (van Kampeny 1909
figunteus (van Kampen) 1915
meeropy (var Rarpen) TY0y
MeAelvs (Boulenger) [ROS
ohesus. Zweifel [972
ucedlatus (van Kampen) 13
ophicodon Peters aid Doria 1878
rasirars Mebely [S98
suberoceus Mengies and Tyler 1977
Nenorhing Peters 1863
houwerst (de Witte) 1934)
doriae (Boulener) ISs¥
mrinrog (Parker) 1934
oxveephula (Schlewel) 1858
parkerorunt Zweitel 1972
sony (Aweilel) 1s6
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 407
TABLE 3, GENERA OF THE GENYOPHRYNINAE Zweifel (1972) undertook a major revision of the
Chosrophaye van, Kampen 1915 asterophryines (sensu Zweifel, 1971). In that paper,
Cophixalus Boettger 1892 Zweifel used characters of cranial osteology and
Copiula Mehely 1901 t | hol d ted t It ti
Genyophryne Boulenger 1890 external morphology, and erecte wo alternative
Oreophryne Boettger 1895 _ phylogenetic trees to indicate intergeneric relationships
Sphenophryne Peters and Doria 1878
(Fig. 2). These proposed phylogenies agree in
Hylophorbus Barygenys Phrynomantis Pherohapsis Asterophrys Xenorhina Xenobatrachus
prootic arcade vomerine spikes
frontoparietal
crest
nasals fused additional vomerine growth
support
squamosal more developed
broadened parasphenoid broadened parasphenoid
skull sculptured
eyes smaller eyes smaller
vomers broadened vomers broadened
maxillae
symphygnathine
dentaries
symphygnathine
Hylophorbus Phrynomantis Pherohapsis Asterophrys Barygenys xXenorhina Xenobatrachus
erie frontoparietal worigrine, Spikes
crest
maxillae secondarily
eleutherognathine
nasals fused additional vomerine
growth, support
squamosals more
developed
eye smaller
skull sculptured
| broadened parasphenoid
vomers broadened
maxillae and dentaries
symphygnathine
FIG. 2. Alternative proposed phylogenies of the Asterophryinae. Redrawn from Zweifel (1972, pp. 430, 431).
48
suggesting that Hylophorbus is the most primitive
aslerophryine genus, an hypothesis supported on the
basis of features of the submandibular musculature by
Tyler (1979), The proposed phylogemes differ in the
placement of Bur\genys, which is regarded as either
a primitive genus which has undergane much parallel
evolution with Yerorhina and Xenehatruchus, or a
genus closely related phylogenetically to the other two,
bat which has undergane a reversal lo partial eleu-
therognathy.
The Mvalogy and Ostealogy of the Pepuan
Microhvlidsy
The only published study of the entire musculature
of any Papuan microhylid is that of Burton (983a).
lylec (1974, 1979) and Emerson (19764) examined the
superficial throat niusculature of some Papuan
microhylids; Horton (1982) included in her review of
longue musculature one genyoaphrynine (Sphenaphryne
robusta); Trewavas (1933) examined the hyoid
musculature of one genyophrynine (Greopliryvne
célebensisy, Jones (1933) examined the pectoral
inusculature al fowr genyophrynines (Cophixalus
verrucosus, Genvophryne thamsoni, Oreophryvne
variahilis. and Sphenophryae carautay, and Pry (1917)
and Zweifel and Allison (1982) examined aspects of the
musculature of Cophixd/us pansus,
While asteological features were used by Bouleuger
(1882) and van Kampen (1923), che main osteological
studies of Papitan microhylids are thase of Mehely
(1901), Wandollek (1910) and Zweifel (1971, 1972),
Important contributions have also been made by Noble
(1931), Parker (1934, 1936), Brongersma (1933) and
Menzies and Tyler (1977),
Mehely (1901) deseribed in detail the osteology and
in particular cranial ostealogy of the following Papuan
microhylids. (Mchely’s names in parentheses):
Phrynomantis (Mantophryne) lateralis, P.
(Gnathophrvne) boetiger, P (G.) dubia, P (G.)
robusta, Hvlaphorbus rufescens (Metapostira océllata),
Nenobatrachus rostrdtus (Nenorhind rostrata),
Nenorhing oxvcephala, Cophixalus (Phryxinalus) biroi,
C (2) montanus, Copiula fistulans (Co oxyrhina),
Oreaphryne (Sphenophryne) biroi, O. velebensis, O.
moluecens[s (QO. senckenbersiana), Sphenophryne
mehelyt (Chaperina fusea) and 8S. (C) pelysticra.
Wandollek (J910) deseribed the osteology and
iNustrated the cranial bones and hyatds of
Choerophryae (Copiula) rastellifer, Oreophryne biroi
(Mehelvia affinis and M. lineata), Sphenaphryne
cormutd, S. mucrorhynche (Chaperina quatuerlabata)
and S. sellaginhaufeni).
Noble (1931) and Parker (1934, 1936) included a small
Number of Skeletal characters in their generic and
subfamilial diagnoses; Brangersma (1953) described the
skeleton of Asterophrys turpicula; Zweitel (1971) used
some skeletal characrers in his analysis of the
relaponships of Genvoptirvere and Menzies and Tyler
(1977) deseribed osteological features of a number af
RECS AUST. MLS. 19 (19): 405-450)
Naverrher, 1986
burrowing Papuan microhyhds of the genera
Barygenvs, Choerophryne, Capiula and Xenobatre-
chus,
in his revision ol the Asterophryinae, Zweilel (1972)
relied to a large extent on cranial osteology, He
described general features of the skeletons af astera-
phryines, the diagnastic features of each genus in
#eneri¢ accounts, and particular features of same
Species in species acvounis, Zweilel’s account of the
cranial skeletons of asterophryines is comprehensive
but scattered, and the contributions of the presene
study 16 the knowledge of the cranial skeleton are the
addition of a few previously unrecorded features, and
also information oo a number of species not examined
by #weifel, This permits reinterpretation of some
cranial characters.
There are problems in the interpretation of the
cranial skeletons of asterophryines and in the selection
of characters for phylogenetic analysis, The problems
of interpretation arise from a number of factors: in all
asterophryines (here are some bones that are fused; in
some asterophryines many bones are fused) and in
some asierophyrines the task of discriminating bone
margins is further complicated by exastosis,
The most problematic bone of the microhylid skull
is the product of fusion of the yomer and the palatine.
Mchely (1901) interpreted a narrow anterad projection
of this bone skirting the medial margin of the choana
as a vomer, and the remainder as a palatine, Noble
(1931) and Parker (1934) referred to the entire bone as
a prevanmier and Zwelfel (1972) asa vomer. The presence
of odontoids.on this. bone conyinced Noble and. Parker
(1926) thar the bone was not a palatine, and they
mterpreted cases of separation of the anterad process
from the rest al the bone as division of the prevamer,
father than separation of the prevomer from the
palatine. | preter to fallow Trueb (1973) in regarding
the identity of the bone as insoluble uptil ontogenetic
data become available, and follow her tn referring to
this bone aga “vornero-palatine”,
The most problematic species is Pherahapsiy
mernsiesi, the skull of which is. heavily fused and
exostased, and particularly difficult to interpret in the
absence of ontogenetic dala. As myolopical and
external similarines of Pherohapsis to Hvlaphorhus,
three species. of Phrynomantis and to a lesser extent
Asterophrys indicate a close relationship with thase
(taxa, J interpret (he cranial structures of Pherohupsis
as homologous with structures in those frogs. For
example, | interpret the prootic arcade of Pherohapsis,
a ribbon of dermal bone which forms an arch between
the lrontoparietal and [he squamosal (Fig. (7), as a
mediad extension of the posrerad projection of the
7ygzomatie ramus of the squamosal exhibited anly by
Asteroplirvs, Hvlophorbys and the three PArynomantis
species, Similarly, although the dorsal surface of the
ote capsule is so fused and featureless thar ir is
imtpassible to identify rmdividual bones, the
THE PAPUAN SUBFAMILY ASTEROPHRYINAF Bila
conformation of that surface and the adjacent broad
medial {lange of the squamosal shalt is consistent with
the interpreranion that at least part af the bone covering
the otic capsule isan otic ramus of the squamiosal. The
dorsal surface of this atic ramus is continuous with the
anterior surface of the medial flange of the squamosal
shatt, This interpretation is comparable to the
condition in Asterophrys, Aylophorhus and the three
Phrynomanrlis species,
The post-cranial skeleton of asterophryines has been
considered too uniform to be informative of relation-
ships within the sublamily (Zweile] 1972, p. 428). In
contrast, the post-crunial skeleton provides characters
fundamental La the separation af the Asteroptiyrinae
und Genyophryninae (nature of the vertebral colurnn),
and to the diagnosis of genyophrynine penera (nature
of the peetaral girdle),
Following Zweifel’s revision af the Asterophryinae,
a number of papers have appeared focussing on
yanation in the post-cranial skeleton of frogs. Trueb
(1973) provided a useful survey of skeletal characters
and subsequently demonstrated the use af measure-
ments of skeletal features (Trueb 1977). Andersen
(1978) surveyed variation of the manus.and pes, and
Tyler (19766), Emerson (1979), Emerson and DeJongh
(1980) and Emerson (1982) demonstrated and evaluated
characters of the pelvic girdle. As a result of (hese
studies more information and hypotheses relating to
post-cranial skeletons are available to. systematists.
Ecology of the Papuan Microhylidsy
The Papuan microhylids are diverse ceologically,
Menzies (1975) uses four categories to accommodate
Ihe comimon Species; to these Zweilel and Tyler (1982)
add a Tilth lo accommodate some less eammon species,
These categories are (a) fossorial, the frogs are
normally found below ground from whence they call,
rarely or Hever encountered on the surface: Barveenys,
Xenobairachus aud probably Xenerhina (Menzies’
placement of Xenorhine among the terrestrial genera
is based on the only common Xenorhina species,
X, doriae, which is shown ut this study to be more
appropriately assigned jlo PArynomantis),
Choerophrvne and Copiula (Menzies and Tyler 1977),
and probably Geavephryne (Zweilel (971); (b)
terresinial, hiding under the gronod by day, but moving
abou! of the ground at nighl: Asleroplrys,
Hvlophorbus, Pherohupsis, Phrynomantis, some
species olf Cophixalus, cg. Co pansus (Zweilel and
Allison 1982), some species of Sphenophrvne: (c)
scansorial, climbing low vegetation to two or three
metres; some species of Cophixalus and Sphenophrvne,
(d) arboreal, climbing high into the trees; some species
of Cophixalus, Oreophrvne; (ve) aquatic: Sphenophryne
palinipes (Zweitel 1956).
Menzies stresses that there is overlap between rhese
categories, and that i} 1s arbitrary ta.a degree. Marcover,
little is Known of the ecology of the majority of species}
but it is clear that the asterophryines are far less diverse
ecologically than the genyaphrynines.
The terrestrial and fossorial frogs of both subfamilies
burrow head first (Menzies and Tyler 1977), The frogs
] observed in captivity (Barveenys flavizularts,
Phrynomantis. lateralis, PB wilhelinana, Copiula
Jistulans, Cophixalus kaindiensis, Sphenophryne Jryi,
S. schlaginhaufent) conform with this mode of
burrowing, the driving power coming mainly from the
hind legs, the arms being used mainly to part leaf litter
and moss. This mode of burrawing contrasts with that
of ather frogs which are described as burrowing head
first, Hemisus marmoratus (Emerson 1976b) and
Arenophryne rotinda (Tyler etal, 1984), both of which
propel themselves largely by arm movements (Emersan
1976b, pers, obs.),
Aims of this Study
This study arose out of the observation by Tyler
(1974) of diversity in the superficial throat musculature
of the asterophryines. li was believed Uhat a detailed
comparative myological study might not only aid in
{he evaluation of Zwielel’s (1972) phylogenetic hypo-
theses but also provide characters with which to
evaluate the hypotheses of relationships between the
two New Guinea subfamilies. Moreover, as differences
were observed between Zweilel's. (1972) drawing af the
lium of Phrynonrantis louisiadénsis and the ilia of
several genyophrynines slored as alizarin preparations
in the Department af Zoology, University of Adelaide,
the potential usefulness of examination of the post-
cranial skeleton was realized, Finally, unremarked
heterogeneity in #weifel’s diagrams of skull
morphology suggested that reassessment of skull
characters. might also yield useful characters.
The aims of this Study have been (o determine the
relationships af the asterophryine genera, using
characters of myology and osteology, and to contribute
tO the assessment of the merits of uniting the
asterophiryines and genyophrynmes mto a sinule
sublamily.
MATERIALS AND METHODS
Muterial Available
The course of this study was dictated in part by the
availabilily of material, Specimens ol adult asiero-
phryines available for dissection ate often difficult or
impossible to obtain. GF the 40 taxa examined by
Zweitel (1972), 18 were represented in collections by
fewer than JO specimens cach, and the three species
named subsequently are known only by their lype
series. Although additional specimens of some of the
very rare species have been collected since 1972, none
has become commonly represented in collections. As
a result there are a number of species Unavailable for
dissection and skeletal preparation.
As aslerophrymes undergo direct development ana
eggs at most species are hidden in leal litter, under
4
moss or under logs, the discovery of eggs or larvae is
rare and accidental, As asterophryines are yery difficult
to rear from the eggs ta adulthoad, identificahon of
epys penerally is impossible unless an attendant adull
is found; even then the evidence of identity is
circumstantial. There are vio published accounts. of
complete life histories of asterophryine larvae (Tyler
1976a), and larval material is so limited thal a
comparative study is impossible.
Techniques such as gel clectrophoresis, karyology
und microcomplement fixation are possible only when
Freshly killed specimens are available, It was impossible
to obtain adequate samples from New Guinea to use
any of these techniques.
Material Examined
A total of 268 specimens of preserved adult
microhylids were examined, including 196 specimens
of 33 species or subspecies of asterophryines,
representing all seven genera, OF the 268 specimens,
256 were examined externally, 159 specimens (14
aslerophryines) were dissected, at least partially, for
myological examination, 5 specimens (82 astero-
phryines) were cleared and stained for osteological
examination, 16 dried skeletons (10 asterophryines)
were prepared, and x-rays were obtained of 37
specimens (all asiterophryines). Three specimens (AUZ
A206, 207, 208) had previously been cleared and
stained,
A total of 23 specimens of preserved adult ranaids
were examined externally, and partially dissected for
myological examination, These specimens. represent two
families, ]O genera and 14 species. All are uncatalogued
specimens housed in the Department of Zoology,
University of Adelaide; Ranidae — Cacasrernum sp.
(L specimen), Hemisns mmarmoratus (1), Natalo-
batrachus sp. (1), Platymantis papuensis (1),
Pyxicephalus sp. (2), Rana fuscigula (1), R. gravi (1),
R. grisea (5), R. papue (1); Hyperooliidae — Afrixalys
sp. (2), Flyperolins marmoratus (2), A. (uberilinguis
(I), Aussina sp. (2) and Lepropelis sp. (2).
The specimens were preserved in 65% or 70%
ethanol, Most were lent by the institutions listed.
Others collected on a field trip are lodyed in the
collection of the Department of Zoology, University
of Adelaide; These specimens. were killed in a 3%
chloral hydrate solution, fixed in 3% formalin, and
preserved in @5%) ethanol,
Specimens Examined
Abbreviations: AA: Collection of Dr A. Allison,
Wau Evology Institute, PNG; AM: Australian
Museum, Sydney; AMNH: American Museum of
Natutal History, New York; AUZ! Department of
Zoology, University of Adelaide (iinregistered apart
from skeletal preparations); BPBM: Bishop Museum,
Honolulu; FMNH: Field Museum of Natural History,
Chicavo, MCZ. Museum Comparative Zoology,
Harvard; RMNH: Rijksmuseum yan Natuurlijke
REC. S. AUST. MUS, 14 (19): 405-450)
Noventber, 1986
Historie, Leiden; SAMA: South Australian Museum,
Adelaide! UPNG: Biology Museum, University of
Papua New Guinea, Port Moresby.
ASTEROPHRYINAE
Asterophiys turpicula (9) Vogelkop: RMNIL 16655;
Olsobip; UPNG 1548; Megalsimbip, Ok Menga:
UPNG 6739-45.
Burygenys uira (8) Lejo via Popondetia: UPNG 383),
3832, 3836, 3837, 3957, 3958, 5475, 5476,
Barveenps exsul (1) Alotau; UPNG 5201,
Barvuenys flaviguluris (7) Mi Kaindi: AUZ A729;
B726a, B726b, B728, D741, SAMA R23851, UPNG
5134,
Rarygenys maculata (V1) Agaun; UPNG 509|-99,
S101, 5102.
Barveenvs nae (4) Purigoi, Kaironk Valley, Schrader
Mis: AM 22802; Kaironk Valley, Schrader Mts: UPNG
3245, 3247: Elimbari: LIPNG 3249,
Baryeenys sp, noy. (6) Mt, Missim; AA 11171, 11172,
BPBM 9366-69,
Aviopharhus r. rufescens (1) Baiyer River: AUZ
A722, D738; SAMA R23844; Madang: UPNG 2285,
2286, 2288; Agaun,; UPNG S041, 5042, 5044; Manga,
Huon Pen; UPNG 5714; Go River, Huon Pen: UPNG
5732.
Hylophorbus r. extinus (1) Mt Riu, Sudest [,:
AMNH 60092,
Hylophorbus r. myopicus (1) Kulumadau, Woodlark
L: AMNH 59988,
Pherohapsis menziesi (6) Sogeri: UPNG 1865, 1970,
2093, 2578, 2579, 5196.
Phrynamantis eurvdacivla (2) Kahilfon, Bultem:
UPNG 5306-07,
Phrynomartis fusca (1) Rohua, $8. Seram: UPNG
5257.
Phrynomantis ho Aumicola (8) Kotuni, Mt Otto:
AMNHE 66266-70 (2 specimens); Daulo Pass: MCZ
52970-81, 52983-86.
Phrynamantis fh. compta (3) Kaironk Valley,
Schrader Mts: SAMA R9387 (3 specimens).
Phrynomantis infulata (3) Arau, Kratke Mts:
AMNH 66685, 66699, 66670.
Phrynomantis: lateralis (13) Taraka vig Lav: AUZ
A730, B724a, B724b, 9737, SAMA R23838; Lae; MCZ
59000; 16 km $ af Popondettas MC'Z 87535; McDowell
t,, Purari R,; UPNG 2499, 2500; Alotau: UPNG 2619,
2621, 2622, 5202.
Phrynamantls louisiadensis (5) Mt Rassel, Rossel L.:
AMNEL 60135-43 (2 specimens); Rossel lL: AMNH
$9117; Abaleti, Rossel lL: AMNH 69347, UPNG 5689,
Phrvnoniantis personate (2) Lumi: AMNEL 78092;
Rauit: UPNG 4087.
Phrynomantis rahusta (4) Derongo: MCZ. 1688:
SAMA RIOS80; Siagara, Misima [.. UPNG 4295;
Bwagaoia, Misima |; UPNG 4303.
Phrynamantis stietagester (17) Lita Patrol Post:
MCZ 59908-16; Okapa: SAMA R20886-93,
Phrynomantis wilhelmuna 16) Wahei-Sepik Divide:
THE PAPUAN SUBFAMILY ASTEROPHRYINAE a
AM R16825: Daulo Pass: AM R66747; Eastern slopes
of Mt Wilhelm: AMNH 65868-86 (2 specimens);
Tomba; AUZ A727, B721 (a), B721(b), B723, B731; MI
Giluwe: SAMA R23849; Kogi, Suai Ra: MCZ 5989)-96,
Nenobatrachus giganteus (4) Eipomek Valley: UPNG
5346, 5347, SATY, SGKO.
Xenobatrachus mehelyi (6) Derongo: MCZ 81673,
81674; Imigabip: MCZ 81675, 81676; Tabubil; UPNG
4790; no data: AUZ,
Xenobatrachus obesus (3) 20 kan NE of Lumi:
AMNH 78187-207 (2 specimens); Amanab: LIPNG
2822.
Nenobatrachus rosiratus (1) Kaironk Valley, Schrader
Mts: SAMA R9386 (4 specimens), UPNG 3240, 3244,
3342, S014; Exqpomek Valley; UPNG 5681, 5682.
Xenobatrachus subcroceus (5) Lae: UPNG 4390-93,
4143,
Xenorhina bauwensi (10) Rigonmendip, Ok Sibil
Valley, Star Mis: RMN 16657 (5 specimens), 16658
(5 specimens).
Xenorhina doriae (4) Mt Lamington: AM R9604;
Camp [ff, Nimi Rx MCZ 64405; Bomai: SAMA
R6284; Alotaus UPNG 2608.
Nenorhina minima (2) Fipomek Valley! UPNG 5677,
5737.
Nenorhina oxveephala (3) 8. coast-of Flurnbold! Bay:
RMWNH 5032; Fak-Fak: RMNH 17017; Eipomek Valley:
UPNG 5678.
Xenorhina parkerarum (1) Halalinja, Nipar UPNG
5827,
Xenorhina similis (1) Lake Habbema, Bele R,, 18 km
N: AMNH 43726.
GENYOPHRYNINAB
Choerophryne rostellifer (i): Moiyokabip Village,
Bultem: UPNG 4410,
Cophixalus darlingtoni(2) Tomba, Mi Hagen; AUZ
B735, SAMA R23844,
Cophixalus kaindiensis (1) Mt Kaindi; AUZ.,
Cophixalus neglectus (3) My Bellenden Ker, Qld:
AUZ A744, A747, 8749.
Cophixalus ornatus (3) S$, Bell Peak, Malbon
Thompson Ra, Qld; AUZ A720, B725(a), B725(b),
D740, SAMA R23845,
Cophixalus pansus (3) Bulldog Rd, Waur AUZ (2
specimens), AUZ A209.
Caphixalus parker (2) Okapa: SAMA R5604 (2
specimens),
Cophixalus ripurius (4) Okapa: SAMA RS216 (4
specimens).
Cophixalus shellvi (2) wo data: AUZ. (2 specimens),
Cophixalus variegatus (8) Mt Kaindi: AUZ B719,
B743, D739, SAMA R23834-43.
Cophixalus
specimesis),
Copinha fistulans (8) Lae: AUZ A723, b742, SAMA
R23836, R23837; Avenchambo via Popondetia: SAMA
R14241.
verrucosus (3) Sogeris AUF (3
Genvophrynae thomsoni (4) Agaun: AUZ (|
specimen), UPNG 5118, 5120, 5130.
Oreophryne biroi (l) Karimui: SAMA R10899,
Oreophryne insulana (2) Ma-u R., Camp lt: AUZ
B745, UPNG 3556,
Sphenophryne cornuta (1) Kigonmendip, Sibil
Valley; SAMA RIL599,
Sphenophryne Jrvi (8) S. Bell) Peak, Malbon
Thompson Ra, Qld: AUZ A746, 8734, D736, SAMA
R23654-55,
Sphenophrvne robusta (4) Boonjie, 16 km SE of
Malanda, Qld; AUZ (1 specimen), AUZ A208; Millaa
Millaa, Qld: AUZ. (1 specimen), AUZ A207.
Sphenophryne sp, (2) Wau: BPBM 9879, 9882,
Sphenophryne sehlaginhau/seri (2) Trauna Ridge, 13
km NE of Baiyer River, AUZ B733, SAMA R23852.
BREVICIPITINAE
Breviceps mossambicus (2) Bronkhorstbrust,
S. Afriear AUZ,
Breviceps sp. (1) Durban N., S. Africa: AUZ.
DYSCOPHINAE
Calluella guitulata (1) Kuala Tahan, Pahang, Malaya:
FMNH 143960,
MICROHYLINAE
Chaperina fusca (1) Deramakot, Kinabatangan Dist,
N. Borneo; FMNH 77253.
Elachistocleis sp. (4) lunapuna, Trinidad: AUZ {3
specimens), ALLZ B748.
Glyphoglossus mmolossus (1) Sakaerat, Ampho Pak
Thong Chai, Nakhon Ratchasima Prov., Thailand)
FMNH 182650,
Kalophrynus pleurostigma (1) Nanga Tekalit Canip,
Mengiong R., Kapit Dist, Sarawak: FMNH 138052.
Kaloula pulchra (\) Siracha, Chalemlarb, Chon Buri,
Thailand: FMNH 175952.
Microhyla hevmonsi (1) Bukit Lanjan, Selangor,
Malaya: FMNE 186029,
Microhyla pulehra (1) Sakacrat, Amphoe Pak Thong
Chai, Nakhon Ratchasima Prov,, Thailand: FMNH
183064.
Methods: Morphologs
External examination included the taking of
standard measurements of snour-veut length (S-V),
head width (HW), eye diameter (EF), eye to naris
distance (BN), internarial span (UN), tympanum
diameter (T), and tibiofibula length (TL). The
mvasurements were taken with Mitutoyo dial calipers
according to the methods deseribed by Zweifel (1972),
In 188 specimens the following additional measure-
ments were taken: (a) head length (ML), the distance
from the tip of the snout to the angle of the jaw (Fig.
3A); (b) mouth width (MW), the distance between the
corners of the mouth (Fig. 3B); (¢) mouth length (ML),
the distance between the most anterior point of the
mouth and the posterior corner (Fig. 3C),
4)2 REC. 5. AUST. MUS. 19 (19): 405-450
‘dies
FIG. 3. Methods of measurement: A, head length; B, mouth width;
C, mouth length
Myological dissections were carried out with the
assistance of topical applications of the iodine-
potassium iodide solution of Bock and Shear (1972).
In eases where muscles were obscured by connective
tissue, the tissue was bathed in 30% nitric acid until
the connective tissue was removed, and the nitric acid
was then aspirated, In 134 specimens all of the skeletal
musculature was examined. In 32 specimens of rare
species, dissection was prohibited, but partial
examination of the muscles was carried out through
pre-existing incisions made by previous workers, One
rare specimen, PArynomantis fusca UPNG 5257, was
partly dissected through straight incisions in the skin,
which was then folded back, and muscle groups
November, 1986
previously known to be taxonomically significant were
examined,
The tongues and associated musculature of six
specimens were examined histologically. Transverse
sections were stained with Mayer’s haecmatoxylon and
eosin.
Muscle descriptions follow the terminology of Ecker
(1889) as modified for the throat by Tyler (1971), the
jaw by Starrett (1968), the hyolaryngeal apparatus by
Trewavas (1933), the pectoral girdle by Jones (1933), the
forearm and manus by Gaupp (1896), Lhe pelvic virdle
by Dunlap (1960) and cutaneous muscles by Burton
(1980).
One hundred and one specimens were cleared and
double-stained for skeletal examination by the Alcian
blue-Alizarin red technique of Dingerkus and Uhler
(1977). Seventeen specimens were cleared and stained
with Alizarin red by the technique of Davis and Gore
(1947) to reveal bones. Sixteen specimens were flensed,
cleared of adherent soft tissue by application of sodium
hypochlorite solution, and allowed to dry slowly.
Thirty-seven specimens, most of (hem too rare to
dissect, were radiographed from several aspects and
prints obtained on a Rank Xerograph in positive mode,
As the drying process led to some collapse of skulls
and caused some bones which are not articulated in
cleared and stained specimens to come into contact,
the osteological descriptions are based primarily on
cleared and stained specimens. Osteological descrip-
tions follow the nomenclature of Trueb (1973).
The skeletal preparations were surveyed in order to
evaluate the potential for taking measurements in the
manner of Tueb (1977), This survey indicated that the
features with the best potential for yielding information
regarding relationships. were the angle between the
vertebral column and the leading edge of the sacral
diapophysis, the sacral expansion and the angle
between the ilial shafts. These were measured using a
goniometer attachment on a Wild M5 stereomicro-
scope, care having been taken that the features being
measured were lying in a horizontal plane.
All myological and osteological drawings were
prepared with the use of a Wild M5 stereomicroscope
with an attached camera licida.
Methods: Phylogeny
The approach adopted in this study is that of Hennig
(1966) as delined by Wiley (1976), This involves the
attempt to falsify competing hypotheses of recency of
common ancestry of groups of taxa, using as evidence
the distribution among the taxa of synapomorphies,
that is, shared uniquely derived character states, The
myology, osteology and external morphology af the
asterophryines, genyophrynines and the other micro-
hylids examined provided the characters that were the
basis of the phylogenetic analysis.
There are a number of problems associated with the
recognition of synapomorphy; first, the determination
of primitive and derived character states, second,
THE PAPLAN SUBFAMILY ASTEROPHRYINAL
distinction of cases where a shared derived character
Stale has arisen once in a common ancestor (i.e. are
“homologous” sensu Bock, 1963) from cases where
sitnilar character states have arisen independently in
different lineages (i.e, are "homoplasious” sensu Bock,
1963), and third, how to handle cases of conflicting
evidence of synapomorphy.
The problem of character slate polarity has been
addressed frequently in recent papers, e.g., Crisci and
Stuessy (1980), De Jongh (1980), Stevens (1980, 1981),
Amold (1981), Bock (1981), Watrous and Wheeler
(1981) and Wheeler (1981). In these papers many criteria
have been assessed and, while a consensus has not
emerged, the case favouring the use of outgroup
analysis alone is compelling, That is, useful
information regarding the direction of change can only
be derived from analysis of the distribution of those
States between the group under study and an outgroup.
A character state shared by members of the graup
under study and the outgroup is considered primitive
relative 10 a character state found only in some
members of the group under study, The character state
shared by members of the sludy group and the
outgroup is assumed to have arisen in a cammon
ancestor of (he two groups, while the restricted
character state 1s assumed to have arisen more recently
in an ancestor common only to those species which
share the state.
The outgroup method is by far the most widely used
method of assessing character state polarities. Studies,
including those of Marx and Rabb (1970), Lundberg
(1972), Lynch (1973, 1975a, b, 1978), Moffat (1973),
Heyer (1975), Heyer and Liem (1976), Wiley (1976),
Hecht and Edwards (1976), Michener (1977), ard
Enghoff (1981) have relied primarily on outgroup
analysis.
In consideration of the asterophryime genera, the
most appropriate outgroup is the Genyophryninac,
whose status as a group closely related to the Astero-
phryinae has been established by a synapomorphy,
direct development of larvae (Parker 1934),
lt is hot necessary for the Genyophryninae ta be
recognized as a distinct laxan in order for it to bean
outgroup. Watrous and Wheeler (1981) defined the
concept of a functional outgroup and demonstrated
that an out-group need not be taxoriomically distinct.
This concept is similar in application to, though
presented more rigorously than, Kluge’s (1976) use of
the mgroup, wea, that a character state widespread
among taxa related at the next higher taxonomic level
“that ornerwise have little in common” js primitive, a
position supported also by Arnold (1981). This is not
the same as the “commonality principle” based ingroup
analysis, Where the more cofnmon character staie
within the group is coded as primitive, a position
criticized by a number al workers, including Moffat
(1974), Stevens (1980), and Watrous and Wheelcr (1981),
who demonstrated the logical shortcomings af the
ANT
commonality principle, All cladagrams based on the
commonality principle nvust root near the middle, and
no three-taxan hypothesis can be solved by the
commonality principle, since any shared character siate
must be the more common, hence primitive, amd so
there can be no shared, derived character states, While
it is necessary to demonstrate chat the Asterophryinae
is a natural group whose monophyly (sensu Hennig,
1966) is attested by autapomorphy and that it is
therefore possible to postulate its evolution, it is rat
necessary ta the study of the asterophryine genera fo
establish the naturalness.and taxonomic viability of the
Genyophryninae.
In the search for evidence of synapomorphy to
suppart the hypotheses af the monophyly of the
Asterophryinae and of the Genyophryninae, the
outgroup consists of ranoid frogs and microhylids of
other subfamilies, listed previously. Myological, skeletal
and external morphological data obtained fram
specimens representing 10 species in the subfamilies
Brevicipilinae, Dyscophinae and Mierobylinae
(Onental and Neotrapical) and 14 ranoid species are
used to assess whether either subfamily possesses
aulapomorphies,
It is considered impossible by many authors fo make
a priori the judgement that the sharing of a given
derived character state results from homoplasy: such
a judgement can only be made when evidence [ram
different characters is showa to conflict (le Quesne
1969; Cracraft 1981; Whecler 1981; but Boek, 198],
presents 4 contrary view). When two characters supperl
incompatible hypotheses of common ancestry, one of
therm is homoplasious, i, the character has been
subject to convergence (sensy /a/o) or reversal, Which
of the two characters is the homoplasious one remains
to be determined. ;
The approach almost universally advocated, eg. by
Camin and Sokal (1965), Kluge and Farris (1969),
Lundberg (1972), Eldredge and Cracraft (1980), Nelson
and Platnick (1981), for resolving such conflicts is to
favour the cladogram supported by the greatest number
of characters, or “the hypothesis that has been rejected
the least number of times” (Wiley 1976), and then 10
reject those characters which do not contribute to the
cladogram, on the grounds that they are homoplasics.
That is, same form of numerical analysis is suggested
in order that the hypothesis of relationships depicted
in a cladogram be the most parsimonious, on the
grounds that while evolution may not be parsimonious.
scientific hypotheses of evolution, as of anything else,
miist be (Kluge and Farris 1969; Wiley 1975; Cracraft
1979; contra Inger. 1967).
However, numerical approaches must be applied ta
cladistic analyses with caulion, as they are based on
a number of questionable assumptions (Panchen 1982).
First, when ane hypothesis of relationships is preferred
to another because it is rejected by fewer characters,
the notion ol what constitutes a character is crucial,
A character appears in practice to be any feature of
44 REC, S AUST. MUS, 19 (19); 405.450
a group of taxa which is perceived to be relatively
constant within taxa but variable between them. It is
assumed in using most numerical techniques that all
characters used in.a cladistic analysis are of cqual value
to the assessment of relationships, But this need not
be the case, For example, characters obtained from
different parts of the phenotype may be correlated.
Such correlation may be obviaus: characters relating
to the length of a bone ard the site of origin of a
Muscle on that bone may well be correlated and be
treated better as a single character. But correlation may
be less obvious, For example, leg length and jongue
morphalogy may well be correlated in a particular case
because both may be adaptations to capture of the
same kind of food. Hecht and Edwards (1976) showed
that radically different interpretations of frog
phylogeny resulted from different perceptions af the
associations of characters of tadpole morphology. An
assessment of character correlation appears to be one
of the legitimate roles of functional analysis, which
could not be carried out for this study, beyand
observations of living specimens during the field trip.
Whether a suite of correlated characters be weighted
highly, as suggested by Heeht and Edwards (1976),
Hecht (1977), or be given equal weight with other
characters is problematical, While the Hecht and
Edwards (1976) weighing scheme seems arbitrary, and
Hecht’s numerical value scheme mare so, the principle
that some form of assessment of characters to
determine their relative “credibility” in the case of
conflict (Schlee 1975) seems valid, Moller-Anderson's
(1978) caveat regarding the difficulty of applying
Schlee’s scheme notwithstanding,
A second problem with the use of numerical analyses
is the likelihood of high levels of homoplasy in
morphologically uniform animals like birds or frogs
(Bock 1963; Hecht and Edwards 1976). Such animals
are constrained in their evolution by similar develop-
mental possibilities, for example, the physical require-
ments far Flight severely limit the possibilities of
evaluuion of the avian body, form, This is particularly
so al low taxanomic levels when the animals are
genetically similar, and might therefore be expected ta
evolve independently similar character states under
similar environmental constraints. But the history of
frog taxonomy gives testimony to a high rate ol
homoplasy even at family level. Classifications based
on features of, for example, the feeth (Gtinther 18585
Boulenger (882) and nature of the pectoral girdle
(Noble (931), both characters now believed to have
evolved convergently, and incompatible phylogenies of
frog Families erected by, for example, Hecht (1963) and
Inger (1967) on one side and Kluge and Parris (1969)
and Lynch (1973) on the other, indicate the high
frequency of homoplasy and the difficulty of
discerning it even at the family level.
The use of numerical techniques to resolve conflicts
between characters presupposes that homologies
outnumber homoplasies, or that “evolution is narmally
Navernher, 1986
divergent” (Panchen 1982; Friday 1982). This
proposition is dubjous, for the reasons indicated inthe
previous paragraph.
One means of resolving these conflicts is to relate
observed structures to their functions. Gans (1966)
points out that it Is mot possible to deduce function
simply from morphology, and few direct studies of the
relation of form and function in living trogs have been
caccied out, mainly studies of the muscles involved in
tangue action, ag., Gans (1962), Gans and Gorniak
(1982) and of the museles involved in locomotion
(Emersan and De Jongh 1980). However, a second
approach based on the correlation al morphologies
with particular modes of burrowing (Emerson 1976b,
Sanders and Davies 1984) has provided dramatic
examples of convergence in muscle complexes, as
groups of muscles of (dentical form adapted for similar
burrowing techniques are found in frogs from different
families.
Other morphological patterns related with particular
functions have been identified but not treated. Liem
(1970) and Anderson (1978) identity as an adaptation
to climbing the division of the M. palmaris longus into
several slips with separate tendons of insertion, which
their comparative studies have shown to be
characteristic of treetrogs of several families. Liem
based his division of the firmisternal treefrogs (the
Rhacophoridae and the Hyperoliidae) into separate
families partly on the grounds that the different
patterns of division of this muscle indicated that the
division had been acquired by different evolutionary
steps. That is, it is likely that tree climbing had evolved
independently in the two groups. This if tur implies
that other derived characters shared by these groups
that are recognisable adaptations to tree climbing, e.g,,
the possession of finger discs and of intercalary
cartilages, must be seen ag likely convergences,
This is selevant ta the asterophryine frogs, all of
which spend part of cach day underground, and some
of which seldom or never emerge above ground, the
genera listed in the Introduction as fossorial. Evidence
in this group of the adaption of different evoluvonary
strategies in the adaptation to the fossorial mode may.
be indireet evidence of convergence in other shared
adaptations to the same mode.
Convergence may also be suspected as the source of
conflict between apparent synapomorphies in cases
Where a particular apparently derived morphology is
observed to recur in distantly related groups, even if
the function is unknown. For example, the reduction
of the pectoral girdle in some genera of microhylids
and ranids is certainly due to convergence, Possession
of a reduced pectoral girdle by ditlerent microhylid
venera does not seem to be a reliable character in
phylogenctic analysis as it has been shown to be liable
ta convergence, aud in cases where this. character
conflicted with other apparent synapomorphies it
would be considered of low value, Panchen (1979, 1982)
THE PAPUAN SUBPFAMILY ASTEROPHRYINAE 415
indicated a third problem: that the number of
synapomorphies detected in a systematic analysis is
unlikely to be the complete set of synapomorphies. The
assumption must therefore be made in numerical
analyses that the ratio of “true” to “false”
synapomorphies revealed in the study of a limited
number of characters cquals the ratio of “true” to
“false” synapomorphies in the whole set,
In this study, characters are deemed to be of equal
weight unless there is reason to believe that characters
are correlated, and then the suite of correlated
characters is regarded as equal to one character, on the
grounds that such a suite may be deemed fo have
resulled from but one evolutionary event, In the case
of contlicts between characters, judgements as to the
relajive likelihoods of the relevant character state
transformations are made whenever possible. The use
of numerical procedures is restricted in this study to
the illustration rather than the evaluation of the
conflicting hypatheses of relationships in a case where
the data confliet intractably.
RESULTS
laxenonuc Recommendations and Nomenclature
In order to avoid confusion and tedious repetition
in the following character state analysis, | foreshadow
faxoriomic recommendations which derive from the
analysis, and which | make formally in the systematies
section of this paper. Two of the asterophryine genera,
Phrynomantis and Nenorhina (Table 2) are
heterogeneous assemblages, and the following changes
are necessary. (a) Removal of Phrvnomuntis lateralis,
FP infulata and P louisiadensis from Phrynamantis.
These species share a number of character stales with
Asterophrys, Hylopherbus and Pherehapsis which
indicate thar it is with those genera that their affinities
lic. Their removal trom Phrynomantis makes this genus
amuch more natural assemblage supported by auta-
pomorphies. The three species removed [rom
Phrynomantis do not fit neatly into any of the other
existing genera. | recommend resurrection of Manto-
phryne Boulenger 1897 (type species, MZ /areralis) to
accommodate them, and subsequently refer lo these
species as Mantophryne luteralis, M, infulata and M.
louistadensis. (b) Xenorhina cdoriae does not conform
morphologically with other members of its genus,
showing affinities rather to Phryromantis (sensu
stricto), and sharing, apoinorphies with the other
members of thal genus. Xevorhing becomes a much
more uniform group without AL deride, and the
remaining members of the reduced genus share many
derived characters not shared by .¥. doriae. |
recommend that AL derfge be transferred to
Phrynomantis and subsequently refer to if as
Phrynomantis doriae,
As well as these asterophryine genera, (he senyo-
phrynine genus Cophixalus is heterogencous to the
extent that it is impossible to make general statements
species comprising C
abou! it in the following discussions. Cophixalus
darlinetoni and C. variegatus are considered as an
entity distinct from Cophixalus. As the term
"yariega/us-group” was used by Menzies (1975) ta
denote a group of four or more small (S-V length
approximately 12 mm) cryptic species, | employ the
term “darlingioni-group” to refer lo the group al
darlingioni and the ©
variegatus-group. Myologically and osteologically, the
darlingtoni-zroup is uniform and is clearly more clasely
related to Choerophryne than (o other Cophixalus, as
it shares many unusual characters with Choerophrvne
alone. Whether the darlingtoni-group should form a
new genus as a sister-group to Choerophryne as
suggested by Zweitel (/a /ite, 19 May, 1982) or the
definition ol Choerophryne be broadened slightly to
accommodate the derlingtoni-group (which may ‘be
paraphyletic) is beyond the scope of this study, The
removal of the durlingtoni-graup reduces the
heterogeneity of Cophixalus. Un this study, (he term
“Cophixalus” refers to the genus Cophixalus, but
excluding the darlingtoni-group. As no Choerophryne
specimens were available for compleie dissection, only
ihe superficial musculature of the venter, pectoral girdle
and throat and the jaw musculature were examined in
this genus and details of the osteology were derived
from Menzies and Tyler (1977),
FIG, 4. Supplementary slips to the WW. ortermandibularts in
asterophryines. Abbrewerians 0 Stare (0) single origin Via a
tendon; 1, State ()) oneins via a lendon ane direet from the
dentarys Py Ste (UY origins from the ventral surface of phe
angulosplenial; 2, State (2) origins from adjacent parts of the
ventral surtice al the angulosplenial.
46
REC. 8. AUST. MUS. 19 (19): 405-450
November, 1986
FIG, 5 Superticial mandibular musculature of (A) Cophixulus kaindiensis AUZ; (B) Copiuila fisuilanys AUZ D742; (C) Genyophryne
thomsoni UPNG 5130, Abbreviations: D, dorsal slip-of the M4. inflerhyoideus; WH, WZ inferhyoideus; IM, M. intermandibularis; (Ms,
supplementary slip of M. intermandibularis, S, M. submentalis,
Character State Analysis
The characters discussed are ones which vary in such
a way as to be of potential use in a phylogenetic study
of the asterophryines or to shed light on relationships
between the Asterophyrinae and Genyophryninae.
Characters which are numbered are those whose States
are distributed in such a way as to warrant subsequent
discussion, The distribution of the states of these
characters are listed in Table 4. States designated 0 are
primitive states. States designated 1, 1', Ll" etc. are
derived, but the relationship between these derived
states is unknown, States designated 1, 2, 3, etc, are
derived with the polarity | —- 2 — 3 ete, States
designated by letters are states of unknown polarity,
ie, the primitive state cannot be identified,
The M. submentalis; Variation in the attachments
of this muscle is in part related to the shape of the
mentomeckelians. In the Asterophryinae alone the
mentomeckelians are posterior to the dentaries and
deflected posteroventrally. Consequently,
mentomeckelians are more prominent sites of
attachment of the M. submentalis im asterophryines
the
than in other microhylids, and in at least some species
of Xenorhina the mentomeckelians are the only site of
attachmenL. I treat this involvement of the mento-
meckelians in asterophryines as a character of the
mentomeckelians, below,
Among the asterophryines there is wide variation in
the posterior extent of the M. submentalis reaching an
extreme expansion in Barygenys utra (Fig. 6), but this
variation is partly independent of generic classifi-
cations and so cannot be employed in a phylogenetic
analysis at the generic level.
Character I. Nature of the supplementary slips of
the M, intermandibularis. Four states occur among the
asterophyrines (Fig. 4): (0) a single supplementary slip
arising via a tendon (4/ylophorbus, Pherohapsis and
Mantophryne),; (1) two supplementary slips, the
anterior arising via a tendon, the posterior direct from
the dentary (PArynomantis) (Burton, 1983): (1') two
supplementary strap-like slips from the ventral surface
of the angulosplenial, the posterior slip inserting on
the median aponeurosis of the M. intermandibularis
(Asterophrys, Xenobatrachus and Xenorhina) (Fig. 6);
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 4)
FIG. 6. Superficial mandibular musculature of (A) Barygenys flavigularis AUZ D741, (B) Barygenys utra UPNG 3836, left posterior
supplementary slip removed; (C) Nenobatrachus rostratus SAMA R9386a.
and (2') two supplementary slips from the
angulosplenial, the posterior slip inserting via a narrow
tendon on the AZ. genioglossus basalis (Barygenys)
(Fig. 6).
State (0) occurs also in the Genyophryninae (Burton,
1984), and is primitive among the Asterophryinae. State
(1) is likely to have evolved directly from State (0), and
State (2') may have evolved from State (1'), but it is not
clear whether State (1') evolved [rom State (1) or directly
from State (0). The polarities are either 0 ~ 1 —
I — 2) or2)?— 1 ~0— 1.
Character 2, Occurrence of a dorsal sheet of
posteromedially directed fibres on the M,
intermandibularis. Two states occur among the Papuan
microhylids: (0) dorsal fibres present (Genyophryninae)
(Burton 1984, Tig. SA), These fibres are not always
visible from the ventral surface (Figs 5B, SC); and (1)
dorsal fibres absent (Asterophryinae),
State (0) is shared by the other microhylids examined,
and is primitive among the Papuan microhylids.
Character 3. Overlap of the Mdm. intermandibularis
and interhyoideus. Two states occur among the Papuan
microhylids: (0) no overlap (Genyophryninae (Burton
1984, Fig. 5), except Cophixalus pansus and ©, riparius;
Xenorhina bouwensi); and (1) overlap of the muscles,
the anterior fibre of the M. inferhyoideus lying on the
ventral surface of the posterior fibres of the M,
intermandibularis (Asterophryinae (Burton 1983,
Fig. 6) except Xenorhina hbouwensi; also Cophixulus
pansus and C. riparius).
State (0) is shared by most of the other microhylids
examined, and is primitive. Overlap occurs also in
Calluelila and Kalould, but it is different in form from
State (1) as it is the M. infermandibularis which
overlaps on the ventral surface of the M. inferhyoideus
in these two genera, The possession of State (0) by
Papuan microhylids appears to be related to small size,
Xenorhina bouwensi (17,9, 21.3, 20.7 mm S-V) is the
smallest asterophryine examined, while Cophixulus
riparius is the largest genyophrynine examined, and C.
pansus among the most densely muscled. Similarly,
4th REC. 8, AUST, MUS. 19 (19); 405-450
Novernber, 1986
FIG. 7. (A) Deeper musculature of the throat of /tvlopherbus nr rufescens AUZ D738, left M. geniohvoldeus lateralis externus severed
{o reveal the M.g.l. infernus origin from the hyale; (B) Tongue muscularire of Nenobairachus eigunteus UPNG 5680; (C) Deeper
throat musculature of Spherophryne schluginhaufeni AUZ R733; (D) Baryvenys flavigularis AUZ D741, the left posterior supplementary
Slip to Whe AL. iermandibuluris not removed. Abbreviations: GB, Ad. geniogloysus basalisy Gl, ML geninhyoldeus lateralis interniis;
GM, ML. genvohvoideus medialis; HG, M. hvoglassus.
Calluella and Kaloula are the largest of the other
microhylids examined.
Character 4, Occurrence of the M. geniohyoideus
medialis. Two states occur in the asterophryines; (0)
M.g, medialis present (Barygenys) (Fig. 7C), (1) Mg.
medialis absent (other asterophryines (Fig. 7A, B)),
State (0) is shared by all genyophrynines, and is the
primitive state for the asterophryines.
Character 5, Origin of the M, geniohyoideus lateralis
internus. Three states occur in the asterophryines: (A)
origins from the dentary and the hyale (PArynomaniis)
(Burton 1983); (B) origin from the dentary only
(Barygenys (Fig, 7C), Xenobatrachus and Xenorhina);
and (C) origin from the hyale only (Asferophrys,
Alvlophorbus (Fig. 7A), Mantophryne aud
Pherohapsis).
AILof these states occur among the genyophrynines
and so polarities cannot be assigned by out-group
analysis.
Character 6. The M. genioglossus, Three states occur
among the Asterophryinae; (0) the Mie. basalis
labiform, and the M7.g. dorsalis comprising two strap-
like muscles (Aslerophrys, Hylophorbus,
Mantophryne, Pherohapsis and Phrynomantis) (Fig.
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 419
FIG. &. (A) Ventral view of the M1. genioglossus of Barygenys atra UPNG 3836, M.g. basalis reflected to the left to reveal M.g. dorsalis;
(2) Ventral view of the deepest muscles of the tongue of Barygenys flavigularis AUZ D741; (C) Ventral view of the M, genioglossus
of Yenobatrachus viganteus UPNG 5680, Vg. basalis reflected to the right to reveal M4.g. dorsalis; (D) Dorsal view of the Miz, basalis
of Phrynomantis stictogaster SAMA R20886; (E) Dorsal view of the Mg. basalis of Genvophryne thomsoni UPNG $130, Abbreviations:
GB, M. genioglossus basalis; GD, M.g. dorsalis; HG, M. hvovlossits.
TA); (1) the Mg. basalis bearing a posterad cultriform
process, and the M.g. dorsalis comprising two strap-
like muscles (Xenobatrachus and Xenorhina) (Figs 7B,
8C); and (1') the Ag. hasalis lamellate and folded, and
the M.z. dorsalis fused and ensheathed in connective
tissue (Barygenys) (Figs 8A, 8B).
State (0) is the usual state among microhylids
including all genyophrynines except the unique
Genyophryne, and is the primitive state. In
Genyophryne (Fig. 8D) the M. genioglossus is short
and bilobular and unlike that of any other microhylid
examined. There is no evidence to indicate the
relationship between states | and 1.
Character 7, Occurrence of a deep slip of the M.
hyoglossus. Two states occur in the Papuan
microhylids: (0) slip present (Genyophryninae, except
Genyophryne), and (1) slip absent (Asterophryinae and
Genyophryne).
State (0) appears to be the usual microhylid
condition (Trewavas 1933), and is probably the
primitive state of the Papuan microhylids. Its absence
in Kaloula is interpreted as a case of parallel evolution,
Barygenys possesses a unique state of the AZ.
hyoglossus (Fig. 8A), but the reduction and division
of this muscle into three narrow, discrete sections
appears to have evolved in association with the
lamellate M. genioglossus basalis, which excludes the
M. hyoglossus from access to the tongue except at three
sites: the two lateral folds of the M.g. basalis and the
posterior notch in the M.g. basalis. As the forms of
these two muscles are so intimately related, they cannot
be considered separately.
420 REC. 8. AUST. MUS. 19 (19): 405-450
Noveniber, 1986
FIG. ¥, Insertions of the Atm. petrahyeidel posteriores in Papuan inicrohylids, (A) Stare (0) iwa Mim. p. posteriores, We pastenor ot
which inserts an the tip of the postero-medial process of the hyoid and the hyocricojd ligament; (B) State (1) two Mim. p, pusteriores,
the posterior of which inserts only on che tip of the posteromedial process of the hyoid; (C) State (1') three Min. p. posteriores.
Abbreviations: I, Wf. petruhvoideus posterior \; Vl, M.p. posteriar Uy U1, Mijn postertor UL.
Character 8. Number and insertions of the Mm.
petrohyoidei poslerivres, Three states occur among the
Papuan microhylids (Fig. 9): (0) two Mfm, p.
posteriores, the posterior of which inserts on the
epicondyle of the posteromedial process of the hyoid
and the adjacent hyocricoid ligament (Genyophryninae,
except Genyvophryne) (Fig. JOA); (1) two Mn. p.
posteriores, the posterior of which inserts on the
epicondyle alone (Barygenys) (Pig. 10B); and (1') three
Mim. p. posteriores, of which the Mp, posterior \\
inserts On the hyocricoid ligament, and the M.p,
PIG, 10, Dorsal view of the larynx of (A) Cophinalus ripurius SAMA
R5216a: (B) Barygenvy afra UPNG 3836, Abbreviations: 1, MW.
petrohyoldeus posterior \; Wl, Mp. posterfor M1,
posterior 1)) on the epicondyle (Asterophryinae except
Barygenys; Genvophryne) (Burton 1983),
State (0) is shared by Ca/luella, the Oniental
microhylines and Gastrophryne (Trewavas 1933) and
is primitive among the Papuan microhylids, The
occurrence of State (1') in Elachistocleis is interpreted
as a case of convergence. While State (1') appears to
have derived directly from State (0) by division of the
muscle, it 1s unclear whether State (1) derives from State
(0) or from State (1'),
Character 9, Two states of the origin of the M.p.
posterior [If occur in the Asterophryinae: (0) origin
from the otic ramus of the squamosal and the adjacent
exoccipital (Barygenyvs, Xenobatrachus and Xenorhina)
(Fig. 11C), and (1) origin from the zygomatic ramus
of the squamosal (Hylophorbus, Phrynomantis,
Mantophryne infulata and M. louisiadensis) or its
posterad projection (Asterophrys, Pherohapsis and
Mantophryne lateralis) (Figs (WA, B).
State (0) is shared by the genyophrynines (Fig, 11D)
and is primitive among the asterophryines,
Character /0. Extent of the anterior origin of the
M, depressor mandibulae. Two states occur among the
asterophryines: (0) origin from the entire ventral margin
of the tympanic ring, and in some small specimens also
from the adjacent epimysium of the M. udductor
mandibulae externus superficialis (Phrynomantis,
except RB dorive) (Fig. 11A); and (1) from the posterior
1/2 only of the ventral margin (Asterophrys,
Baryeenys, Hylophorbus, Pherohipsis, Xenobatra-
chus, Xenorhina, Phrynomantis doriae and Manto-
Dhryne (Pig, IC),
State (0) is shared by the genyophrynines (Fig. 11D),
and is primitive.
Character 1], Extent of the slip of the WW. depressor
mandibulae from the otic ramus, Two states occur
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 42)
FIG. 1. A. Right lateral view of the jaw musculature of Phrvnomantis h. humicala MCZ 52970, M. adductor miandibulae externus
supersicialis removed to reveal the position of the mandibular branch of the trigeminal nerve; B. Dorsal yiew of (he jaw musculature
of Phrynomuntis wilhelmana AM R66747; C. Left lateral view of the jaw musculature of Xenobarrachus giganieus UPNG 5680;
D. Right lateral view of jaw muscles of Sphenophryne schlaginhaufeni AUZ B733. Scale bar — 5 mm- Abbreviations: N, trigeminal
nerves IIL, origin ol M. petrohyoideus posterior.
among the asterophryines: (0) origin relatively small,
significantly less bulky than the slip from the dorsal
fascia, or absent (Asferephrys, Barygenys, Hylo-
phorbus, Mantophryne, Pherohapsis and Phryno-
mantis); and (1) origin relatively extensive,
approximately equal in bulk to the slip from the dorsal
fascia (Xenobatrachus, Xenorhina).
State (0) is shared by all genyophrynines, except
Choerophryne and the darlingtoni-group, which share
an unusual conformation of the depressor musculature,
which does not occur in asterophryines. State (0) is
considered primitive among the asterophryines.
Character 12. Development of the M. adductor
mandibulae posterior longus, Three states occur among
the asterophryines: (0) the fibres passing directly from
their origins on the surfaces of the frontoparietal,
prootic and exoccipital to a tendon placed antero-
laterally in the orbit, the fibres not organized into
discrete segments (//ylophorbus, Mantophryne, Phero-
hapsis and Phrynomantis) (Figs 11B, 12A); (1) the fibres
from the frontoparietal passing laterally and the fibres
from the prootic and exoccipital passing anteriorly, sa
that the two sets of fibres form segments (Asterophrys);
and (2) the fibres organized into segments as in State
(1), some fibres from the more posterior origin inserting
on a superficial tendon which unites distally with the
usual deep tendon of insertion (Barygenys, Xeno-
batrachus and Xenorhina) (Fig. SiC).
State (0) is shared by the genyophrynines (Fig, 11D)
and is primitive. States (1) and (2) appear to be by-
products of the massiveness of the adductor muscles
of Asterophrys, Barygenys, Xenobatrachus and
Xenorhina. The direction of evolution 1 — 2 is asso-
Ciated in part with a diminution of the relative size of
the skull, and a consequent tendency of the jaw muscles
to bulge in Barygenys, Xenobatrachus and Xenorhina.
The exclusion of the M.a.m. posterior longus from
an exostosed or otherwise adorned skull in
Aslerophrys, Hylophorbus, Mantophryne and Phero-
hapsis is considered below as a character state of the
skull.
Characier 13. Grigin of the M. adductor mandibulae
externus superficialis. Two states occur among the
asterophryines: (0) origin from the zygomatic ramus
of the squamosal, with linle or no origin trom the
fascia between the anterior tip of the zygomatic ramus
422 REC. S. AUST. MUb, 19 (19): 405-450.
FIG, 12. A. Dorsal sjew of (he jaw musentature of Martuphryie
lateralis AUZ DAT; B. Anterior muscles of the shunk of
Mantophryne fatralis AU. 19737. Seale bar = 5 mm.
and the eye (.4sferophrvs, Hyvlophorbus, Pherahapsts,
Mantophryne and Phrynomuntis) (Figs 1A, B, 12A);
and (1) an extensive origin from the fascia anterior to
the anterior tip of the zygomatic ramus (Barygenys,
Nenobatrachus and Nenorhina) (Vig. UC).
State (0) is shared by the genyophrynines (Fig, 11D)
and is primitive. State (1) is related to the long exparise
between the tip of the zygomatic ramus and the eye
in Barveenys, Nenobatrachus and Xenorhina, which
in turn is related to the smallness of the eye in these
two genera, and consequently this character cannot be
considered independently of Character 47: eye-size.
The origin of this muscle in Pherohapsis is modified
as a result of the connexion between the zygomatic
ramus and the maxilla, but although the May,
exrernus superficialis is entirely deep to this sheet of
bane, its origin is from the zygomatic ramusas in other
asteropliryines,
The Trigeminal nerve: The position of the mandi-
bular branch of the trigeminal nerve is too inconsistent
to be useful as a character in phylogenetic analysis.
Character 1/4. Insertion of the M, longissimus dorsi:
Two states occur among the asterophryines: (()
insertion on the exoccipital (Asterophryinae except
Barvgenys) (Figs UB, 12A); and (1) insertion partly on
the dorsal fascia (Barvsenys),
Stare (0) is shared by eenyophrynines, and is
primitive among the asterophryines.
The ML ileolumbaris: Two states of the At,
dedlumbaris occur among the asterophryines! origin
lram the tip of the ilial shall (4A steroplirys, Barygenvys,
Phrynomantis, Nenobatrachys, Xenorhina and Muanto-
pliryne louisiedensis), and origin trom a site well
posterior to the tip (/i/ephorbus, Pherehapsis,
Mantophryne infilata and MM. lateralis). Both states
occur inthe genyophrynines, The former state occurs
November, 1986
in species with the ilio-sacral articulation type IA of
Emerson (1979), and the lavier in species with a type
| articulation. This character cannot be considered
independently of Character 46: “iha-sacral articulation
type”.
Character 15. Tendinous inscriptions in the M. recruis
abdeminis. Two states occur among the microhylids;
(A) one abdominal tendinous inscription (Astero-
phryinae, Genyophryiinae and Breviceps); and (B)
three abdominal tendinous inscriptions (Microhylinae
and Ca//ueillu).
Neither state occurs among the ranoids in which Lwo
Stales occur: four abdominal tendinous inscriptions
(Most genera); and twa abdominal tendinous
inscriptions (Hemisus), It is impossible from such data
to determine the primitive state in the microhylids,
Character 16. Extent of fibres. of the Adm. obliqui
abdominis externus and transversus dbdominis, Two
states occur among the asterophryines: (0) ventral
Insertions On a broad tendon which covers the ventral
abdomen (Asierophrvinae, except Barygenys; and (1)
libres from the two sides meeting on the mid-ventral
Surface of the abdomen, al least in part (Burvgenvs)).
State (0) is shared by the genyopbrynines and is
primilive among the asterophryines.
The origin of the WW rhemboideus anierior. Two
states Occur among the asterophryines: orivin entirely
from the exoceipital (sferopArys, Pherohapsisy, and
origin partly from the dorsal fascia (Barveenrs,
Hylaphorbus, Mantophryne, Phrvnomantis, Xeno-
batrachus and Xenorhina) (Figs 1B, C, 12A).
Both states occur among the genyophrynines, The
condition of this muscle in .dsferophrys and
Pherohapsis could be related (o modifications of the
skulls of these genera. The genyophrynines which
possess a partial orgin fram (he dorsal fascia are the
more heavily muscled terrestrial or fossorial species
(Cophixalus neglectus, ©. pansus, Copinla,
Genvophrvne and Sphenophryne), and this condition
may be related to the utilization of an additional site
of origin in the face of crowding by muscles of the
limited skull surface, This character will not be
considered further.
Character 17, Insertion of the M. serratus medins.
‘Awo stales occur among the asterophryines: (0) two
insertions On the suprascapula: one dorsal avd one
ventral ta the site of insertion of the AZ levator
scapulde superior (Asterophryinae, except Barvgerys);
afid (2) one insertion posterior to tie insertians af’ the
Afi. levatores (Barvgenys).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines,
Character 18, Origins of the M, levalor scapulae
inferior, VWwo states oecur among the asterophryines:
(0) origin partly from the ventral surtaces of the two
anterior yeriebrae (Asterophrvs, Hylophorbus,
Mantophryne, Pherohapsis and Phrynomuntis)
(Burton 1983); and (1) origin partly from the ventral
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 424
FIG, 13, Ventral superficial museles of right magus of Barveertys
alra UPNG 3836, Abbreviation: L, M. luntbricalis brevis diviti
IV, Seale bur - J mm.
Surfaces af ihe three anterior vertebrae (Barveeny's,
Xenobatrachus and Xenorhina).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines.
Character 19. Development of a deep slip of the M,
pecioralis sternalis, Wo states occur among the
asterophryimes; (0) deep slip poorly developed and
uniting distally With the superficial musele
(Asterophrys, Hylopharbus, Mantophryne,
Pherohapsis and Phrynomantis); and (1) deep slip well
developed, oblique to the superficial muscle and
inserting separately (Baryeenys, Xenobatrachus and
Nenorhina),
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines,
Character 20. Origin of the Mt. honbricalis brevis
digiti \V af the manus. Two states of the origin of the
fedial slip occur among the asterophryines: (0) origin
[rom a superficial tendon, musele slender (Barvgenys)
(Fig. 13); and (1) origin via a short, stout tendon from
the centrale postaxiale; muscle large and fusiform
(4Aslerophryinae, except Baryzenys) (Burton 1983).
State (0) oceurs in Cophixalus riparius and
Genvaphryne thomsani among the genyaphrynines. In
all genyophrynines this js a slender muscle arising from
a long narrow jendon, usually from the palmar apo-
neurosis, State (1) does not occur among the genyo-
pbrynines, State (0) is considered primitive among the
asterophryines.
Character 21. Position of division of the M. rbialis
anticus longus inta bellies. Two states occur among the
asterophryines: (A) within the proximal 2/3 of the
muscle (Barygenys, Pherohapsis, Phrynomantis,
Xenobatruchus and Xenorhina) (Burton 1983); and (B)
division within the distal 1/4 of the muscle (Asteno-
phrys, Avlophorbus and Mantophryne) (Fig. 128).
As both states occur among the genyophrynines, no
polarities can be ascribed to the states of this character,
which may well be correlated with Character 22.
Character 22, Origin of the M. tibialis antious brevis,
Two slates occur among the asterophryines: (0) origin
entirely or partly within the proximal 2/3 of the
libiofibula (Barygenys, Phrvnomantis, Xenobatrachus
and Xenorhina), and (1) origin entirely within the distal
1/4 of the tibiofibula (Asterophrys, Aylophorbus,
Mantophryne and Pherohapsis).
State (0) is shared by the genyophrynines, aud is
primitive among the aslerophryines.
Character 23. Origin of the M. opponens halluets.
‘Two states occur among the asterophryines: (0) origin
from the tarsalia (Asteraphryinae, except Barygenys)
(Burton 1983); and (1) origin trom the dorsal surface
of the plantar aponeurosis (Burygenys),
State (0) is shared by the genyophrynines, and is
primitive.
Character 24, Union of the Mm. /umbricales breves
digiterum IV and V. Two states relating lo the degree
of fusion of the lateral slip of the A@.Lb. digiti 1V and
the medial slip of the Mb, digit? VY occur in the
aslerophryines: (0) separation proximal, much less than
1/2 the medial slip of the M./>. digiti V involved in
fusion (Barvgenys); and (1) separation distal, the MLA,
digili V almost entirely fused to the MLLB. digiti TV
(Asterophrvinae, except Baryeenys),
State (0) is shared by the Genyaphryninae, and is
primitive among the asterophryines,
Character 25, Relative breadth of the frontoparietals,
Two stales occur among the asterophryines; (0) lronta-
parictals broad, length approximately 2« breadth of
the combined frontoparietals (Bar\eenvs) (Fig. 14A);
and (1) frontoparietals relatively narrow, length
approximately 3» the breadth (Asterophryinae, except
Barygenys) (Figs |14C, )SA, C, E, 16A, C),
State (0) is shared by the genyopbrynines (Figs 17A,
I8A) and is primitive among the asterophryines,
Character 26. Oceurrence of parasagittal ridges on
the frontoparietals, Twe states ovcur among the
Papuan micrehylids: (0) parasagittal ridges lacking
(Barycenys, Phrynomantis, Xenobalrachus, Xenorhina
und Genyophryninae except Genvophryne) (Figs |4A,
©, ISA, 18A); and (1) parasagittal ndges present
(Hvlophorbus, Mantophryne, Pherohapsis arid
Genyaphryne) (Fiz. 16C).
State (0) oceurs in the other microhvlids and is
primitive in the Pupuan microhylids. The frontoparie-
tals differ in form among (hose microhvlids exhibiting
State (1), In Hylophorbus and Mantaphryne infylata
the ridges are small, and the area between the ridges
is unadorned. In the other taxa the area between the
ridges is rugose, and the effect is that the mid-dorsal
424
REC. 8. AUST. MUS. 19 (19): 405-450
November, 1986
FIG. 14. A, Dorsal and B. ventral view of the skull of Baryvgenys atra UPNG 3836; C. Dorsal and D. ventral views of the skull of Yenubarrachus
glizanteus UPNG 5680. Scale bar = 5 mm.
surface of the cranium appears as a rugose plateau.
It is assumed that the two conditions described are
alternative forms of the one state, The state of
Asterophrys (Fig. 16A) is not clear (see discussion of
Character 27, following).
Character 27. Occurrence of a sagittal crest on the
cranium. Two states occur among the asterophryines;
(O) sagittal crest Jacking (Asterophryinae except
Asterophrvs and some specimens of Xenobatruchus
obesus and Phrynemantis doriae); and (1) sagittal crest
present (Asferophrys and some specimens of
NXenobatrachus obesus and Phrynomantis doriae) (Figs
I5SC, 16A).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines. The development
of a strong sagittal crest in large specimens of
Phrynomantis doriae and Xenobatrachus abesus
clearly results from an ontogenetic effect — progressive
lateral compression of the frontoparietals and raising
of a sagittal crest as the frontoparietals are forced
against each other. This may result from the dispropor-
tionate development of adductor muscles during
ontogeny. The crest of the small specimen of
Asterophrys that | examined was very similar to that
of the largest specimen of P. doriae, but the specimen
figured by Zweitel (1972) possesses an exostosed plateau
on the mid-dorsal cranium similar to that of
Pherohapsis, Mantophryne lateralis, M, louisiadensis
and Genyophryne, but narrower. Whether the evolution
of the sagittal crest of Asferophrys followed a pattern
similar to the ontogeny of the crest in PR doriae, or
occurred by lateral compression of an exostosed plateau
is unknown.
Character 28, Fusion and expansion of {he vomero-
palatine. Though many states occur in the Microhylidac
(Parker 1934; Carvalho 1954) (Fig. 19), two appear
relevant to this study: (0) vomero-palatine not forming
a large plate extending from a median suture to the
maxillae (Microhylidae, except Asterophryinae and
Genyophryninae); and (1) vomero-palatine a large plate
extending from a median suture to the maxillue
(Asterophryinae and Genyophryninae) (Figs 14B, D,
15B, D, F, 16B, D, 17B, D, 18B),
Though the primitive state of the microhylids is
THE PAPUAN SUBFAMILY ASTEROPHRYINAEL 425
FIG. 15. A, Dorsal and B. ventral views of the skull of Phryvomuntis sticlogaster SAMA R 20886; C. Dorsal and D. ventral views of
the skull of Phrynamantis doriae SAMA R6284, E. Dorsal and F. ventral views of the skull of Mantaphryne lateralis AUZ 173).
Seale bar « 5 mm.
unclear, it is unlikely that il is the unique state exhibited
by the New Guinea subfamilies, and State (1) is
considered derived,
Character 29. Lateral expansion of the vormero-
palatine. Two States occur in the Papuan microhylids:
(0) vomero-palatine not expanded close to its Jateral
articulations (Genyophryninae, except Cophixalus
pansus and Genyophryne) (Fig. 18B); and (1) yomero-
palatine expanded laterally (Asrerophryinae, C pansus
and Genyophryne) (Figs 14B, D, 15B, D, F, 14B, D,
17B, D).
As the vomero-palatine of other microhylids are
reduced relative to the Papuan microhylids, the
unexpanded condition, State (0) is likely (o be primitive,
Character 30. Extent of median expansion of the
vomero-palatine. Two states occur among the astero-
phryines: (0) expansion moderate (Asterophrys,
Hylophorbus, Mantophryne, Pherohapsis and
Phrynomantis) (Figs 15B, D, F, 16B, D, 17D); and (1)
expansion large (Barygenys, Xenobutrachus and
Xenorhina) (Figs 14B, D).
State (0) is shared by the genyophrynines (Figs 17B,
18B), and is primitive among the asterophryjnes.
Character 31. Occurrence of spike-like odontoids on
the vomero-palatine. Two states occur among the
asterophryines; (0) spikes absent (As/erophrvinae,
except Xenohatrachus); and (1) spikes present
(Xenabatrachus) (Fig. 14D),
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines.
Character 32. Width of the cultriform process of the
parasphenoid, Two states occur among the Papuan
microhylids: (A) cultriform process relatively narrow,
less than 1/2 of the width of the frontoparietals
(Asterophryinae, except Barygenys, Xenobatrachus and
Xenorhina, Genyophryninae, except Genyophryne)
(Figs 15B, D, F, 16B, D, 17D, 18B); and (B) cultriform
process broad, more than 2/3 of the width of the
frontoparietals (Barygenys, Xenobatrachus, Xenorhina
and Genyophryne) (Figs 14B, D, 7B).
Both states are shared by the other microhylids
examined, and it is not possible to assign polarities to
this character.
Character 33, Extent of the articulation of the
pterygoid with the prootic, Two states occur in the
Papuan microhylids; (0) articulation short (Genyo-
phryninae, except Genyophryne) (Fig, 18B); and (1)
articulation long (Asterophryinae and Genyophryne)
(Figs 14B, D, 15B, D, F, 16B, D, 17B, D).
State (0) is shared by the other microhylids examined,
and is primitive among the Papuan microhylids,
Character 34, Development of the quadratojugal.
Two states occur in the Papuan microhylids; (0)
quadratojugal poorly developed, articulation with the
maxilla brief or lacking (Genyophryninae, except
Genyophryne) (Fig. 18B); and (1) quadratojugal well
developed, articulation with the maxilla long (Astero-
phryinae and Genyophryne) (Figs 14B, D, 15B, D, F,
16B, D, 17B, D).
State (0) is Shared by the other microhylids examined,
and is primitive among the Papuan microhylids.
Characler 35. Relationship between the squamosal
and the maxilla. Two states occur among the Papuan
microhylids; (0) no contact between the zygomatic
426 REC, S$. AUST, MUS, 19 (19): 405-450
November, 1986
FIG. 16. A. Dorsal and B. ventral views of the skull of Asterophrvs rurpicula RMNH 16655; C. Dorsal and D. ventral views of the skull
of Hylophorbus r rufescens AUZ D738, Seale bar ~
tamus and the maxilla (Genyophryninae, except Genyo-
Phryne; Asterophryinae, except Asterophrys and
Pherohapsis); and (1) ventral margins of the zygomatic
ramus and the squamosal shaft in contact with the
dorsal margins of the quadratojugal and the maxilla,
and the lateral surface of the resultant sheet of bone
exostosed (Asterophrys, Pherohapsis and Genyo-
phryne) (Figs ISA, 17A).
State (0) is shared by the other microhylids examined,
and is primitive among the Papuan microhylids. The
small specimen of Asterophrys that I examined lacks
this connexion (Figs 16A, B) but Zweifel (1972) reports
and figures it in other specimens.
Character 36, Occurrence of a posterad extension of
the zygomatic ramus of the squamosal. Two states
occur among the asterophryines: (0) no posterad
extension, or else a slight flange on the posterior
surface of the base of the otic ramus (Barygenys,
Hylophorbus, Phrynomantis, Xenobatrachus,
Xenorhina, Mantophryne infulata, M. louisiadensis);
3mm,
and (1) posterad expansion of the zygomatic ramus
(Asterophrys, Pherohapsis and Mantophryne lateralis)
(Figs 1SE, 16A, 17C),
State (0) is Shared by the genyophrynines, and is
primitive among the asterophryines.
Character 37. Nature of the otic ramus of the
squamosal, Four states occur among the Papuan micro-
hylids: (0) dorsal surface of the otic ramus a flat plate
continuous with the anterior surface of the medial
flange of the squamosal shaft; ramus short, not
extending to articulate with the crista parotica (Genyo-
phryninae, except Choerophryne, the darlingtoni-group
and Genyophryne) (Fig. 18A); (1) as in State (0), but
ramus longer, overlying the crista parotica (A stero-
Phrys, Hylophorbus, Mantophryne, Pherohapsis,
Xenohatrachus, Xenorhina, Choerophryne, the
darlingtoni-group and Genyophryne) (Figs 14C, 15E,
16A, C, 17C); (1') dorsal surface of the otic ramus
continuous with the lateral shaft; ramus extending
medially; medial flange of shaft reduced
THE PAPUAN SUBFAMILY ASTEROPHRYINAL W27
PiG. 17. A. Dorsal and B. ventral views of the skull of Genvoplryne thomsont UPNG 5130;-C. Dorsal and D. venttal Views of the skull
iW Pherohupsis menzies’ LINC) 2579, Seale bur Simm,
(Phrynomantis) (Figs 15A, C); and 1") dorsal surface
of the otic ramus vontinuous with the anterior surface
of the medial flange; ramus not a flat plate, bur folded
60 (hat it forms a sheet covering the anterior surfaces
of the olic capsule; just extending to articulate on the
anierodorsal margin of the crista parotica (Burygenys)
(Fig. 14A),
State (0) occurs in the other microbylids and appears
to be primitive among the Papuan microbylids. State
(1) is likely to have been derived direetly from State (0),
bul is unlikely (o have formed an intermediate stage
in the evolution of State (1') as State (1) involves an even
greater expansion of the medial lange of the
squamosal shalt than does State (0), and State (1°)
involves a reduction of the medial flange to a ridge on
the shaft. In State (") the medial flange is expanded
and this stale may have been derived fram cither State
(0) or State (1). The polarity of this character is either
t-OLor Ph -~O-—1—1"
Character 38. Occurrence o! an anterior connexion
between the partes faciales of the maxillae. Four states
occur among the Papuan microhylids (Pig, 20): (0)
partes faciales produced antero-medially to overlap the
premaxillae slightly; direct ligamentous connexion
between the tips of the partes laciales Jacking
(Genyophryninac, except Genyophryne), (1) partes
faciales produced anteromedially to overlap the
premaxillae slightly, bul more thati in State (0); dense
ligamentous connexion between the tips of the partes
faciales (Genyophryne and #yvlophorhuys; (2) partes
faciales broadly overlapping the premaxillae; connexion
by dense ligaments or by a median suture (Astere-
phryinae, except Baryeenys and Hylopherbus; and (1)
no anleronmiediad projection of the partes faciales, no
overlap ol the premanillae, no ligamentous connesxtan
(pace Zweifel, 1971); premaxillae narrow, compressed
between the maxillae (Barvgernys).
State (0) occurs in other microhylids, and ts primitive
among the Papuan micrahylids. State (1’) is more likely
to have been derived from State (0) than from State
(1) or (2), as State (1, which is associated with three
thickened ridges of skin on the snout corresponding
428 REC. $8, AUST, MUS, 19 (19): 405-450
FIG. 18. A. Dorsal and B. ventral views of the skull of Cophixulus
riparius SAMA RS2l6a. Scale bar = 5 mm.
with sites of articulation of the mazxillae and
premaxillae, represents a radically different mode of
snout reinforcement from that represented by States (1)
and (2). State (1) is likely to have been derived from
State (0) by extension of the anteromediad processes
of the partes faciales and the establishment of a
ligamentous connexion between them, The polarity of
this character is thus I' — 0 — 1 — 2,
| regard States (1) (possessed by GenyophAryne and
Hylophorbus) and (2) (symphygnathy) as differing only
in degree, and consider the distinction between them
somewhat arbitrary, Mehely (1901), Parker (1934) and
Zweifel (1971, 1972), on the other hand, regarded the
distinction as, to varying degrees, crucial. The
difference between my appraisal and that of Mehely
November, 1986
FIG, 19. Ventral views of anterior bones of the skull of various
microhylids, with the vornero-palatine and palatine shelf of the
maxillary shaded, (A) Calluella guttalata; (B) Chaperina fusca:
(C) Elachistocleis sp; (D) Glyphaglossus molossus; (E)
Kalophrynus pleurostigma; (F) Kaloula pulchra; (G) Microhyla
pulchra (partly after Parker, 1934).
and Parker may arise in part from my having access
to a large number of cleared and double-stained
specimens, in which the presence or absence of
ligamentous connexions is more obvious than in dried
skeletons. Zweifel (1971) noted ligamentous connexions
in Genyophryne (and implied their presence in
Barygenys and Hylephorbus), but decided that the
degree of closeness of contact of the maxillae is a more
important indicator of relationships. I contend that
closeness of contact has been overemphasised, as in
some “symphygnathine” species, eg., Mantophryne
infulata, the distance between the partes faciales (up
to 0.3 mm) approaches that of Genyophryne and
Hylophorbus, and in Genyvophryne and Hylophorbus
the partes faciales are closer than in those genyophry-
nines exhibiting the State (0) condition, Whether or not
my contention that the condition of Genyophryne and
Hylophorbus is close to that exhibited by those
asterophryines of State (2) condition is accepted, the
possession by Genyophryne and Hylophorbus of a
state different from State (0) and intermediate between
States (0) and (2) is indisputable.
Character 39. Occurrence of a posteromediad
process of the anterior margin of the palatal shelf of
the maxilla, Two states occur among the astero-
phryines: (0) process lacking (Asterophrys, Barygenys,
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 429
VIG, 20. Anterior view of the bones of the snouts of (A)
Phrynomantis humicola compta SAMA. R938Ta; (B)
Hylophorbus r. rufescens AUZ. D738; (C) Barygenys atra UPNG
3836; (D) Xenobatrachus giganteus UPNG 5680; (EB)
Mantophryne lateralis N\UZ D737; (FP) P infulata AMNH 66685.
Seale bar = 5 mm.
Hylophorbus, Pherohapsis, Mantophryne and
Phrynomantis), and (1) process present (Nenobatrachus
and Xenorhina) (Fig. 14D).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines.
Character 40. Calcification of the nasal capsule. Two
states which occur among the asterophryines are
FIG, 21, Anterior views of the mandibles of (A) Ranu enisen Aus;
(B) Genvophryne thomsoni UPNG 5130; (C) Sphenophryne
schaginhaufeni AUZ. B733; (D) Kaloula puichra FMNH 175952:
(E) Barygenys sp. nov. AA LILTL. Scale bar 5 mm,
considered: (0) anterior portion of the nasal capsule
cartilaginous (Asterophryinae, except Barygenys); and
(1) anterior portion of the nasal capsule calcified to
form a calcified are between the septomanxillaries
(Barygenys) (Fig. 20C). State (0) is shared by the
genyophrynines, and is primitive among (he
asterophryines.
Character 4]. Relations of the dentaries and
mentomeckelians. Two states occur in the Papuan
microhylids: (0) dentary and mentomeckelian not
fused; mentomeckelians lying on the rim of the
mandible between the dentaries; angle between the long
axes of the mentomeckelians obtuse (Genyophryninae)
(Fig. 21B, C); and (1) dentary and mentomeckelian
fused; mentomeckelians largely or entirely excluded
from the rim of the mandible by the dentaries; angles
between the long axes of the mentomeckelians acute
(Asterophryinae) (Fig. 21E).
State (0) is shared by ranoids and other microhylids
(Figs 21A, D), and is primitive among the Papuan
microhylids. State (1) embraces the condition in
Hylopharbus and Mantophryne infulata, in which the
dentaries do not meet, and that of other asterophryines
in which they do (the symphygnathine condition), as
I consider this difference a matter of degree only, and
not as indicative of relationships as the similarities of
430
(IG. 22. Right humerus of (A) Phrvnemantis humicola comple
SAMA R9387a; (B) XYenobutrachus eizanteus UPNG S680; (C)
Barvgenvs atra UPNG 3836; (D) Asferophrys lurpicula RMNH
16655; (LE) Genvephryne thamsani UPNG 5130; (F) Cophixatus
ripdrius SAMA RS52l6a. Scale bar = 5 mm.
the conformation of the mentomeckelians and the
relationship between the dentary and the mentomecke-
lian, Which are unique to the Asterophryinae.
Character 42. Degree of development of the
humerus. Two states occur among the Papuan micro-
hylids: (A) humerus relatively straight; crista ventralis
moderately developed (Asterophryinae, except
Baryeenys, Xenobatrachus and Xenorhina:
Genyophryninae, except Genvophryne) (Figs 22A, D,
F) (B) humerus curved; crista ventralis well developed
(Barygenys, Xenobatrachus, Xenorhina and Genvo-
phryne) (Figs 22B, C, E).
Both states occur among the other microhylids, and
so polarities cannot be assigned, No noticeable sexual
difference oecurs in this character,
Character 43, Reduction of the pectoral girdle. Four
states Occur among the Papuan microhylids (Fig. 23):
(0) procoracoids present; clavicle extending Jaterally to
articulate with scapula (SphenopAryvae), (1)
Procoracoids present; clavicles not extending as far
REC. 5. AUST, MUS, 19 (19): 405-450
November, 1986
laterally as the scapula (Oreophryne); (2) procoracoids
present; clavicles absent (Genvephrine); and (3)
procoracoids absent; clavicles absent (Astcrophryinae,
Choerophryne, Cophixalus, Copiufa and the
darlirgtoni-group).
Reduction of the pectoral girdle occurs in other
microhylid subfamilies (Fig. 23) and in the Ranidae,
and is believed to have occurred independently several
times. A complete pectoral girdle is generally regarded
as primitive, eg., Trucb (1973), Laurent (1979).
Certainly the reacquisition of elements of the pectoral
girdle lost in the evolutionary history of the
genyophrynines is less likely than the alternative, a
progressive loss of elements of the pectoral girdle.
Character 44, Nature of the vertebral column. Two
states Occur among the Papuan microhylids; (A) all
presacral vertebrae procoelous (vertebral colunin
procoelous} (Genyophryninae); and (B) all presacral
vertebrae procoelous except the eighth, which is
opisthecoelous (vertebral column diplasiocoelous)
{Asterophryinae). Both states occur in ranoids
(Duellman 1975) and other microhylids, and although
State (B) is the more common (Parker 1934) it is not
possible to ascribe polarities to this character on the
basis of outgroup analysis. Parker (1934) and Carvalho
(1954) demonstrated variation in this character within
the Microhylinae. Whether this indicates that this
character is of little significance in the Microhylidae
as a whole (Savage 1973) is unclear.
Character 45, Occurrence of a dorsal crest on the
ilium. Two states occur among the aslerophryines
(Fig. 24): (0) ilial shaft Jacking a dorsal crest (Barvgenvs
and Phrynomantis); and (1) ilial shaft bearing a dorsal
crest (Asterophrvs, Hylophorbus, Mantoaphryne,
Pherohapsis, Xenobatrachus and Xenorhina),
State (0) is shared by the genyophrynines, and is
primitive among the asterophryinae,
Character 46. Nature of the tlo-sacral articulation.
Two states occur in Papuan microhylids: (A) articula-
tion indirect, a dorsal ligament connecting the ilial
shafts; this is the type I articulation of Emerson (1979)
(Hvlophorbus, Pherohupsis, Manlophryne infulata, M.
lateralis, Choeraphryne, Cophixalus, Copiula, Qreo-
Phryne and the darlingtoni-group); and (B) articulation
by a ligament attaching to the dorsal surface of the
sacral diapophysis close to its base; this is the type ITA
articulation of Emerson (1979) (Asrerophrys,
Barygenys, Phrynomantis,; Mantophrvne louisiadensis,
Xenobatrachus, Xenorhina, Genyophryne and
Sphenophryne).
Both states occur among the other microhylids, and
it is not possible to ascribe polarities to this character.
Character 47, Eye-size. Two states occur among
the Papuan microhylids: (0) eye relatively large;
E:S-V > 0.090 (Asterophryinae, except Baryeenys,
NXenobatrachus, Xenorhina and Phrynemantis doriae
— see below: Genyophryninae, except Copiula and
Genyophryne); and (1) eye relatively small; E: SV <
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 43)
FIG, 23, Ventral views of the pectoral girdles of (A) Cophixalus ornatus AUZ D740; (B) Genyophryne thomsoni UPNG 5130; (C) Oreophrvne
birot SAMA R10899; (D) Sphenaphryne cornuta; (E) Calluella guttulata; (F) Chaperina fusca; (G) Elachistocleis sp.. (H) Kaloula
pulchra, () Kalophrynus pleurostigma (D-1 modified after Parker, 1934).
0.090 (Barygenys, Xenobatrachus except X. ocellatus,
Xenorhina, Copiula and Genyophryne).
The eyes of the genera exhibiting State (1) are
generally smaller than those of the other microhylids
examined and State (0) is regarded as primitive among
the Papuan microhylids.
Zweifel (1972) characterizes Phrynomantis doriae as
a small-eyed frog, with a relative eye size conforming
to the ratios seen in Xenorhina. The sample sizes for
populations | examined are too small to document
conclusively an ontogenetic trend to smaller relative eye
sizes within asterophryine species, but a trend to
smaller relative eye size in larger species within genera
is apparent. Figures 25 and 26 show the logs of eye
diameter to snout-vent length plotted against snout-
vent length in Phrynomantis and Xenorhina and
Xenobatrachus combined, with the ratios of specimens
of P. doriae plotted on both graphs. Clearly, although
the eye of P doriae is smaller that that of other
Phrynomantis, it conforms better to the trend to
smaller relative eye size in larger PArynomantis better
than to the similar trend in Xenorhina and
Xenobatrachus.
Character 48. Development of subarticular tubercles.
Two states occur among the asterophyrines: (0)
subarucular tubercles poorly developed or absent
(Barygenys, Phrynomantis, Xenobatrachus and
Xenorhina); and (1) subarticular tubercles well
developed (Asterophrys, Hylophorbus, Mantophryne
and Pherohapsis).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines.
Character 49. Adherence of the tongue. Two states
occur among the Papuan microhylids: (0) tongue free
posteriorly, for at least 1/4 of its length (Genyophry-
ninae); and (1) tongue adherent posteriorly (Astero
phryinae).
Although both states occur among the other micro-
hylids, only State (0) appears to be found among the
ranoids. For this reason I regard State (0) as primitive
among the Papuan microhylids.
Character 50. Uniformity of the tongue surface. Two
432
FIG. 24. A. Right lateral and B. dorsal views of the pelvic girdle
of Phrynomantis stictogaster SAMA R20886; lateral view of
right ilium of (C) Mantophryne fateralis AUZ D737; (D)
Xenobatrachus giganteus UPNG 5680; (E) Hylophorbus: r.
rufescens AUZ D738; (F) Barygenys atra UPNG 3836. Scale
bar = 5 mm,
states occur among the Papuan microhylids (Fig. 27):
(0) tongue uniformly pitted and glandular (Genyophry-
ninae, except Cophixalus riparius); and (1) tongue
divided into an anterior smooth, non glandular section
and a posterior glandular section (Asterophryinae and
Cophixalus riparius).
State (0) occurs in other microhylids, and is primitive
among the Papuan microhylids. C. riparius is the
largest of the genyophrynines examined, and the occur-
rence of this character may be related to ontogeny, as
it is lacking in juvenile specimens of PArynomantis h.
humicola.
Character 5], Nature of the glandular surface of the
tongue. Two states occur among the asterophryines: (0)
REC, 5, AUST, MUS. 19 (19): 405-450
November, 1986
~ nas!
ro] e
ls
i
i
~
'
a
z
2
(inaa—inoNs © 44a) 9m
'
3
B
Vt in Whe 17 fait as
LOG SNOUl = VENT LENGTH
FIG, 25, Graph of log of eye to snout-vent length ratio against log
of snout-vent length of PArynomantis species. Means and
standard deviations are shown for each species. Legend: a,
Phrynomantis doriae, ¢, P humicola compta; e, R eurydactyla;
f, PB fusca; h, RA. humicola; rt, RP robusta; s, P. stictogaster sl,
P. slateri; w, PR. wifhelmana.
7.00: |
- 1.05) | |
anons + 343) 907
~ 1,40 |
—1As
(ana
~ 1,20 Ss
— 1.25
LOG SNOUT— VENT LENGTH
FIG, 26. Graph of the log of eye to snout-vent ratio against the log
of snout-vent length in species of Xenobatrachus and Xenorhina,
including PhArynomantis doriae for comparison. Means and
standard deviations are shown for each species. Legend: g,
Xenobatrachus giganteus; m, X, mehelyi; 0, X. obesus; 1, X.
rostratus; 8, X. suberoceus; b, X. bouwensi; i, X. similis; n, X.
minima; p, X. parkerorum; x, X. oxycephala; a, Phrynomiantis
doriae.
glandular surface more or less uniformly pitted
(Asterophrys, Barygenys, Hylophorbus, Mantophryne,
Pherohapsis and Phrynomantis) and (1) glandular
surface bearing deep longitudinal striae
(Xenobatrachus and Xenorhina) (Fig. 27).
State (0) is shared by the genyophrynines, and is
primitive.
Character 52. Possession of a pair of warts on the
chin. Two states occur among the asterophryines: (0)
warts absent (Barygenys, Hylophorbus, Phrynomaniis,
Xenobatrachus and Mantophryne infulata); and (1) two
warts or wart-like protrusions on the chin (Asterophrys,
Pherohapsis, Mantophryne lateralis, M. louisiadensis).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines.
Character 53. Number of denticles on the posterior
prepharyngeal fold. Two states occur among the
asterophryines: (0) large number of denticles, usually
far more than 10 except in a few specimens
(Asterophrys, Hylophorbus, Mantophryne,
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 33
\
FIG, 27. Borsal surfaces of phe tongues of (A) Barygenvs atrra UPNG 3836; (B) Xenobatrachus viguntens UPNG S680.
Pherohapsis and Phrynomantis); and (1) small number
of denticles, less than 10 (Berygenys, Xenobartrachus
and Xenorhina).
State (0) is shared by the genyophrynines, and is
primitive among the asterophryines.
Character 54. Life history. There are many states of
life history among the Microhylidae (Parker 1934), but
two are considered here: (0) larva possessing an
operculum; metamorphosis usually completed outside
the egg capsule (Microhylidae, except Asterophryinae
and Genyophryninae), and (1) larva lacking an
operculum; metamorphosis completed within the egg
capsule (Asterophryinae and Genyophryninae).
State (0) is the usual condition of [rogs, and is
primitive,
Occurrence of digital grooves and discs: Zweifel
(1972) noted that the occurrence of prooves and dises
varies within genera, particularly PAryvnomantis,
Xenabatrachus and Xenarhina, The changes | propose
in the composition of the genera do not affect this
variability. Zweilel (P. 429) chose to use the
development of discs as a phylogenetic character
despite identifying “separate parallel trends towards
reduction of digital dises" within these genera, He
coded Phryvnomantis and Xenobatrachus and
Xenorhina differently on the grounds that some
Phrynomantis species which possess discs possess
broader discs than Xenobairachus and Xenorhina
species. | regard this character as too variable for use
in a phylogenetic analysis and will not consider it
further,
Cladistic Analysis
Here hypotheses are presented relating to the
monophyly of groups of microhylids, and the evidence
for and against these hypotheses, leading to the
establishment of a cladogram, Throughout this section
reference is made to the polarities of characters
discussed in the previous section summarised in
Table 4, and numbers in parentheses are references to
the number ascribed fo a particular character in the
previous section,
Hypothesis 1, Monophyly of the Papuan microhylids
The monophyly of the Papuan microhylids has not
been questioned sinve Parker’s (1934) synthesis, Though
I attempted to identify autapomorphies of the Papuan
microhylids, | was able to add only one character of
dubious polarity (15, number of tendinous inscriptions
of the M. rectus abdominis) to the two characters (28,
development of the vomero-palatine; and 54, life
history and larval form) presented by Parker (1934).
On the other hand there is no evidence of conflicting
synapomorphies suggesting that either Papuan
subfamily forms a natural group with any other
subfamily examined,
Hypothesis 2(a). Monophyly of the Asterophryinae
and of the Genyophryninae
Zweifel (1972) and Tyler (1979) present the opinion
that the Asterophryinae descended from a genyophry-
nine ancestor, Implicit in this belief is the conclusion
that the Genyophryninae form a paraphyletic group,
and that the Asterophryinae are monophyletic.
This study yielded no unequivocal evidence of the
monophyly of the Genyophryninae, O! the four
characters in which the two subfamilies differ
consistently, three present evidence for the monophyly
of the Asterophryinae, and one is of doubttul polarity,
The hypothesis of asterophryine monophyly is
supported by; (a) lack of a set of posteromedially
434
REC. S. AUST. MUS. 19 (19): 405-450
Navember, 1986
TABLE 4, SUMMARY OF CHARACTER STATES
Character a 2
number % b, & i s
x < 2 = s a & a 2 iS Y é =
Ss » SF Fg g§ & § < fF ¢¥ s *£ =
& § a iS) ~ < ~~ = iS) v 4
Fess &§ F —§§ F§ FER FF FEB
< ¢ § ££ £ SF F € FF KF OF SY & 4 s
1 V 2! 0 0 1 I lV’ 0 0 0 0 0 + 0 4 0
2 1 1 I 1 1 1 1 1 0 0 tt) 0 0 0 0 0
3 ] 1 | 1 1 1 O/1 I 0 0 0/1 0 0 0 0 0, +
4 1 0 ] ] ! 1 ! 1 0 0 0 0 0 0 0 0
5 Cc B Cc Cc A B B tS Cc Cc A B B A B A,B
6 0 I! 0 0 0 1 1 0 — 0 0 0 0 0 0 0
7 1 1 1 1 1 ! | | — 0 0 0 l 0 Q 0,1
8 I I I' I 1’ 1’ 1 1’ — 0 0 0 1’ 0 0. 0,1
9 I 0 1 1 1 0 0 1 _ 0 0 0 0 0 0 0,1
LO 1 1 ] ! O/1 ! 1 ] 0 0 0 0 0 0 0 0
a 0 0 0 0 0 ! 1 0 + + 0 0 0 0 0 0
12 ! 2 0 0 0 2 2 0 0 0 0 0 0 0 0 0
13 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0
4 0 | 0 0 0 0 0 0 — 0 0 0 0 0 0 0
15 A A A A A A A A = A A A A A A A,B
16 0 l 0 0 0 0 0 0 = 0 0 0 0 0 0 0
17 0 1 0 0 0 0 0 0 — 0 0 0 0 0 0 0
18 0 1 i) 0 0 I | 0 — 0 0 0 0 0 0 0,1, +
19 0 1 0 0 0 ] i 0 — 0 i) 0 0 0) 0 0
20 ] 0 l I 1 ! 1 1 = 0 0 0 0 0 0 0
21 B A B A A A A B — A A B A A B A,B
22 1 0 i i} 0 0 0 | _ 0 0 0 0 0 0 0
23 0 1 0 0 0 0 0 0 — 0 0 0 0 0 0 0
24 ] 0 1 1 1 1 1 ] _ ) 0 0 0 0 0 0
25 l 0 1 1 1 1 1 | — 0 0 0 0 0 0 0
26 0 0 1 | 0 0 0 1 _ 0 0 0 1 0 0 ()
27 1 0 0 0 0/1 0/1 0 0 _— 0 0) 0 0 0 0 0
28 1 1 1 | 1 1 1 ] — 1 1 1 l 1 1 0
2 ! 1 i} 1 1 | 1 1 _ 0 0/1 0 ] 0 0 0
30 0 1 0 0 0 1 I 0 + + 0 0 0 0 0 0
31 i) 0) 0 0 0 1 0 0 _ 0 0 0 0 0 0 0
32 A B A A A B B A — A A A B A A A,B
33 1 1 1 1 I 1 1 1 = 0 0) 0 1 0 0 0
34 1 1 1 1 1 l 1 1 —_ 0 0 0 ] 0 0 0
35 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0
36 1 0 0 1 0 0 0 0/1 — 0 0 0 0. 0 0 0
37 1 sf I 1 l 1 ] 1 1 1 0 0 i] 0 0 0
38 2 }! | 2 2 2 2 2 — 0 0 0 1 0 0 0
39 0 0 0 0 0 | 1 0 — 0 0 0 0 0 0 0
40 0 i 0 0 0 0 0 0 — 0 0 0 0 0 0 0
41 1 l 1 1 1 1 ] | — 0 0 0 0 0 0 0
42 A B A A A B B A A A A B A A A,B
43 3 3 3 3 3 3 3 3 3 3 3 3 2 | 0 0-3
44 B B B B B B B B A A A A A A A A,B
45 1 0 ! 1 0 1 ] 1 — 0 0 0 0 0 0 0
46 B B A A B B B A/B oS A A A B A B A,B
47 0 1 0 0 0 l 1 0 0 0 0 1 I 0 0 0
48 I 0 ! 1 0 0 0 l 0 0 0 0 0 0 0 0
49 1 1 1 1 1 1 l ! 0 0 0 0 4) 0 0 0.1
50 1 1 1 1 1 1 1 l 0 0 0/1 0 0 0 0 0
51 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0
52 1 0 0 1 0 0 0 O/1 0 0 0 0 0 0 0 0
53 0 | 0 0 0 I | 0 — 0 0 0 0 0) 0 0
54 ] ] 1 1 1 1 1 1 1 | 1 | I 1 1 0
See text for discussion of State numbers. (—) indicates information is not available; (+) indicates a derived state not discussed in
the foregoing character analysis; only states shared with the Asterophryinae and the Sphenophryninae recorded in the “Other Microhylidae”
column,
oriented fibres on the dorsal surface of the M.
intermandibularis (2); (b) a unique structure of the
mandible (41); and (c) a tongue that is adherent and
superficially complex (49 and 50).
The procoelous vertebral column (44) of genyo-
phrynines may or may not represent an autapomorphy.
However, while the hypothesis of paraphyly of a group
may be rejected on the evidence of a single
autapomorphy, the absence of an autapomorphy does
not disprove monophyly: it may reflected the inability
of the worker or of the techniques employed to detect
autapomorphy. The monophyly of the Genyophryninae
can only be disproven by autapomorphies which
conflict with the hypothesis of monophyly. Such would
be an autapomorphy supporting the monophyly of one
or more of the genyophrynine genera with the Astero-
phryinae.
If, in Figure 28, a genyophrynine genus (G) and the
Asterophryinae form a monophyletic group, then the
Genyophryninae (S+G) would consist of descendants
of a common ancestor (C), but excluding one
descendant of (C), the Asterophryinae. That is, the
THE PAPUAN SUBFAMILY ASTEROPHRYINAE
Genyophryninae Asterophr yinae
———
S G
FIG. 28 Phylogeny of the New Guinea microhylids postulated in
hypothesis of monophyly 2(b), Abbreviauons: C, common
ancestor of the Asterophryinae and Genyophryninae; G,
Genyophrytine genus sharing synapomorphies with the
Asterophryinae; §, other genyophrynine genera. Horizontal
strokes through lines indicate acquisition of derived character
slates.
Genyophryninae would be united only by plesio-
morphies and be, by Hennig’s (1966) definition,
paraphyletic.
Hypothesis 2(b), Monophyly of the group formed by
Genyophryne and the Asterophryinae
The monotypic genus Genyophryne has had a
complex history (Introduction), as Genyophryne
(homsoni possesses a mixture of genyophrynine and
asterophryine character states, plus a number of states
which do not occur elsewhere in the Microhylidae (e.g.
odontoids on the dentary). Consequently, since Parker's
(1934) monograph it has been allied with the astero-
phryines (Parker 1934), the genyophrynines (Zweifel
1971), and has been the cause of amalgamation of the
subfamilies (Savage 1973).
There are three characters of which Genyophryne
shares with the whole of the Asterophryinae the derived
State: lack of a deep slip to the M. Ayoglossus (7); an
extensive articulation of the pterygoid with the prootic
(33); and a well developed quadratojugal with a long
articulation with the maxilla (34),
Though these characters appear to provide strong
evidence of the monophyly of Genvophryne plus the
Asterophryinae, there are three lines of argument that
may be marshalled against this hypothesis.
(i) Conflict with relationships within
Asterophryinae.
There are two characters of which Genyophryne
shares the derived condition with all of the
asterophryines except Barygenys: possession of three
Mm, petrohyvoidei posteriores (8); and extension and
the
FIG, 29. Postulated relationships of Genpophryne, Barygenys atl
the other Asterophryinae, A. Common ancestor (C) of
Genvophryne (G), Barvgenys (B) and the other Asterophryinae
(A-B) possesses the derived states of characters 8 and 38, and
Burygenys undergoes reversals, indicated by crosses, in these
characters; B. Common ancestor possesses the primitive states
of characters 8 and 38, and the derived states are acquired
independently in Genyophryne and the Asterophryinae except
Barygenys, which possesses the primitive states.
ligamentous connexion of the maxillae (38). If
Genyophryne and the Asterophryinae form a
monophyletic group, either their common ancestor had
evolved three Mm. petrohyoidei posteriores, and
extension of and connexion of the partes faciales which
subsequently underwent reversal in Barygenps
(Fig. 29A), or else the common ancestor maintained
the genyophrynine condition, and parallel evolution
occurred, Genyophryne and Asterophryinae except
Barygenys acquiring the apomorphous states
independently (Fig. 29B).
It is necessary to assess the relative likelihood of
these two models. The Mm. petrohyoidei posteriores
of Barygenys differ from both the asterophryine and
genyophrynine conditions, and haye evolved either by
loss of the entire M.p. posterior 11 of an asterophryine-
like ancestor, or by reduction of the posterior slip of
a genyophrynine-like ancestor. As indicated earlier in
43g
the discussion of this character, | can see to reason to
favauc either hypothesis.
On the other hand, the conformation of the snout
of Barygeny's seems. unlikely to have evolved from the
conditian shared by the asterophryines and Genyo-
Phryne, as such as evoluvionary step would involve the
loss by a burrowing animal of one mode of
reinforcement of the snout (by extension of the
maxillae and their connexion anteriorly to the
premaxillae) and acquisition of a radically different
mode of reinforcement, involving reduction of the
premaxillae, and the buffering of the points of
weakness in the snout (the gaps between the bones) by
ridges of thickened skin (Fig. 20). Such a course of
evolution, from one specialised burrowing confor-
mation to another radically different, seems far less
likely than the de nevo acquisilion of the different
burrowing adaptations in the different lineages from
a peneralised genyophrynine- or microhyline-tike
ancestor,
Thus, while a common ancestor of Genyophrvne,
Barygenys aud the ather asterophyrines possessing the
asierophryine condition of three Min, petrohpoidei
posteriores may be envisaged, | cannot envisage an
ancestor of Sarygenvs possessing {he snout
conformation shared by Genyophryne and the
asterophryines, and | favour the hypothesis that this
state evalved convergently (sensu lato) in Genvophryne
and the Asterophryinae, which raises the possibility
that the other similarities of Genyophrvne and the
Asterophryinac are hkewise convergences.
(ij) Conflict with apparent synapomorphies between
other genyophrynines and the asterophryines,
Some characters indicate that if the Asterophryinae
indeed evolved [rom a genyophrynine aficestor,
Cophixalus rather than Genophryne forms the sister
froup to the Asterophryinae, the position implied by
Zweifel (1972) and Tyler (1979), presumably on the
grounds. that Cophixalys and the Asterophryinae share
the derived state of a reduced pectoral girdle (43). If
a case can be made to establish Caphivalys as a
possible sistegroup of the Asterophryinag i would
decrease the plausibility of the apparent autapa-
morphies of the group Genvophryne-Asterophryinae,
and, provided the case for a natural group Cophixulus-
Asterophryinae were nol toa compelling, weaken the
credibility of the hypothesis that the Asterophryinae
evalved from any genyophrynine ancestor,
Two additional characters support (he monophyly
of Cophixalus and the Asterophryinae: overlap of the
Min. inlerhyoideus and inlermandibularis (2); and
complex conguc-surface (50),
The characters supporting Cophixalus-Astero-
phryinae monophyly are not compelling, The pectoral
girdle has undetgoue reduction many times in the
Microhylidae and Ranidae, and there is a reasonable
likelihood that reduction took place independently in
Cophinxalus and the Asterophryinae from an ancestor
REC 8. AUST. MUS, 19 (19): 405-450
November, 1986
TABLE 5. GROUP SHARING DERIVED CHARACTERS WITH
GENYOPHRYNE
Group Character
Barygedys, Nenurhina 32 Broad culinform process of
the parasphenaid (probably
derived),
42 Well-developed crisia
ventalis of the humerus
(probably derived).
Barveenys, Xenorhina, 47 Small eye,
Cupinta
Algrerapheys, Aylophoartys, 47 Olie ramus of the
Pherotiapsis, Xenorhing, squamosal overlying the enista
Mantophryue paratien.
Asterapheyinge, Cophixatus: 29 Laterally broudened
pansus nalaline-prevomer
Asleraplirys; PRherahapals 44 Connection of the zygzo-
matic ramus ef the
squamosal and the
maxillary.
26 Parasagittal ridge
deluniting the exient ol
adductor museles on the
(rontoparictals.
Hiloptiorbus, Pherohapsis,
Monflephryae
with the condition exhibited by Genvyophryne.
Moreover, | suggest that as small and juvenile
specimens of asterophryines exhibit the genyophrynine
states of characters (2) and (50), and only large
specimens of Cophixalus exhibit the asterophryine
states of these characters, (hese may be states related
to degrees of devclopment, rather (han states which
evolved in a common ancestor shared only by Cophix-
dlls and the Asteraphryinac.
I conclude that the evidence for Cophixalus-Astero-
phryinae monophyly is too weak to threaten the case
tor Genvophryne-Asterophryinae monophyly:
(iii) Cases of canvergence involving Gernyvephrvne
and members of the Asterophryinae.
There are Seven derived states shared by Genvo-
phryne and a small number of asterophryines, and
sometimes some genyophrynines (Table 5), I many
cases the distribulions of these states support
incompatible natural groups, and the derived staies of
characters 29 and 47, which would support very odd
natural groups, ace highly unlikely to have evolved only
once,
Importantly, of the 12 characters shared by Genyo-
pPhrvne and some or all of the Asterophryinae, cight
(characters 26, 29, 32, 33, 34, 35, 37 and 38) relate to
the enlarging and remforcement of bones of the skull,
and with two others (42, development af the humerus:
and 47, reduced eve size) are likely burrowing
adaptations. Lf these ten characters were reduced to
one: “adaptation to a head-first burrowing mode of
life’, this may. better express the relationstip between
Genyophryne atid the asterophryines. The two shared
derived states (7, loss of a deep slip to the M,
hyeglossus; and &, possessian of three Adm.
petrohyoidei pasteriares) which are not at least
notionally related ta the burrowing way of life are states
which have evolved independently in the Microhylinae
(in Aaloula (7) and Elachistocleis (8)).
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 437
I therefore regard the hypothesis of monophyly of
Genyophryne and the Asterophryinae, although
apparently supported by many characters, only weakly
supported when the characters are examined. If the
monophyly of (his group is only weakly supported,
then the hypothesis of the monophyly of the Genyo-
phryninae is not falsified but sul an open question.
Hypothesis 3, Monophyly of the genera and
suprageneric groups within the Asterophryinae
Having established the monophyly of the Astera-
phryinae and having discussed iis relationship with the
Genyophryninae, | now establish the monophyly of
groups within the Asterophryinae,
(a) Monophyly of Barveenps.
The monophyly of Burpgenys is supported by the
following autapomorphies: posteriar supplementary
slip of the M, iatermandibularis from the ventral
surface of the angulosplenial lo the ventral surface of
the M. genioglossus basalis (i); lamellate M.
venioglossus basalis (6); two Adm. petrohyoidei
posteriares, the pasterior of which inserts on the
epicondyle of the posteromedial process of the hyoid
only (8); insertion of the M. /ongissimus dorsi on the
dorsal fascia (14); the Mm obliqui.and transversi of
each side riceting on the ventral surface of the
abdamen (16); the M. serratus medius with only one
insertion (17); the Af. opponens hallucis from the dorsal
surface of the plantar apancurosis (23); the olie ramus
of the squamosal averlying the anterior and dorsal
surfaces of the otic capsule (37) no overlap of the
maxillac anterior to the premaxillae; compression of
the premaxillae between the maxillae; possession of
tliree cutaneous ridges of the snout (38).
Baryeenys is a very uniform genus and of the above
nine character stales, none is shared by any other
microhylid examined.
(b) Monophyly of Phrvnorantis.
The monophyly of PArvnomantis (including
Phrynomartis doride) is supported by two
aulapomorphies: two supplementary slips to the M,
intermandibularis, the anterior ftom a deep tendon,
the posterior clirect from the dentary (1); and the dorsal
surface of the otic ramus of the squamosal continuous
with the laleral surface of the squamosal! shaft; the
medial [ange of the shaft reduced (37).
Of these states the first (1) is unique, and the jatter
is shared only by microhylines with very small otic rami
(eg. Microhyla), far smaller than those of
Phryvnament/s,
(c) Monophyly of XNerobatrachus and Xenorhina.,
The monophyly of the group comprising Xenobarra-
chus and Aenorhina is supported by the following
aulapomorphies: the MW, gentog/ossus basalis
possessing a posterad cultriform projection (6); a major
component of the Af. depressor mandibulae arising on
the otic ramus of the squamosal (11); posteromedial
projection of (he anterior margin of the palatine
process of rhe maxillary (39); posteriar section of Lhe
longue bearing deep longitudinal striae (51).
OF these four autapomorphies, three (6, 39, $1) are
states which occur only in these two genera, and Lhough
the M, depressor mandibulae arises mostly from the
oti¢ ramus in Choerophryne and the darlingtoni-group,
the form of the M, depressor mandibulae in those taxa
is very different, as there is no slip fram the dorsal
fascia overlying the otic ramus, In subsequent
discussion | refer to this pair of genera collectively as
the Xenarhina-group.
(d) Monophyly of Asferaphrys,
Mantophryne and Pherohapsis.
The monophyly ol this group is supported by two
autapomorphies: distal organ of the M. (7bialis andieus
brevis (22); and large subarticular tubercles (48). A
third shared state which is unique among the Astero-
phryinae and is probably derived is the origin of the
M. geniohyoideus lateralis internus entirely from the
hyale (5),
Several other characters which are derived or
probably derived are shared by members of this group:
distal separation of the M, (/bialis anticus longus into
bellies (4sterophrys, Hylophorbus and Mantophryre}
(21); parasagittal ridges on the frontoparietals
(Hylophorbus, Mantophryne and Pherohapsis) (27),
exostosed sheet of bone between the squamasal and
the maxillary (Asterophrys, Pherohapsis) (35); posterad
projection of ihe zygomatic ramus of the Squamosal
(Asteraphrys, Pherohapsis, Mantophryvne lateralis)
(36); type | ilio-sacral articulation (Alplaphortus,
Pherohapsis, Mantophryne infulata and M. lateralis)
(46); and wart-like protrusions of the skin of the clin
(Asierophrys, Pherohapsis, Muntophryne lateralis and
M, lauisiadensis) (82).
Neither of the autapomarphies of rhis graup (22, 48)
occurs in the other microhylids examined. This group’s
state of the M. geniohyoideus lateralis internus is
shared by the aberrant darlingroni-group and
Elachistocleis.
The hypothesis of manophyly of this group is in
conflict with the hypothesis of monaphyly of
Asterophrys, Xenobatrachus and Xenorhina, The latter
hypothesis is supported by lwo apparent aulapomor-
phies: (1) supplementary slips of the M. intermandi-
buluris arising [rom posteriorly on the ventral surface
of the angulosplenial: and (12) segmentation of the
M. adductor mandibulae posterior longus. The ventral
origin of the supplementary slips has occurred probably
independently in three other Papaan microhylid genera:
Barvaenys, Genyaphryne and Sphenophryne.
Segmentation of the M. adductor mandibulae posteriar
longus has probably arisen independently in Barygenys,
[ regard neither af these characters as reliable an
indicator of relationships as characters (5), (22) and
(48).
I regard AsteropaArys, Hylopherbus, Mantaphryne
and Pherohaysis as forming a monophyletic group
which has undergone extensive radiation, reflected tn
Hylaphorhus,
4aH REC. 5. AUST. MUS. [9 (19): 405450
its disposition by Zweifel (1972) into four wenera, three
of them monotypic. In subsequent discussion in this
section this group ts referred to as the Asterophrys-
group, and the taxonomic implications of us
manophyly are deferred,
Hypothesis 4. Monophyly of PArvynomantis, the
Xenorhinu-graup and the Asierophrys-group,
In four characters,
asterophryings exhibits the plesiomorphic condition,
and Phrynomantis, the Xenorhina-group and the
Asterophrys-zroup share the apomorphic condition:
lack of the M. geniohyaideus medialis (4); the M.
lubricalis brevis diziti 1V medial slip (of (he manus)
a stout muscle arising via a short tendon trom the
centrale postaxiale (20); distal separation of the Mn.
lumbricales breves digiteruin VW and V of the pes (24);
and the lateral margins of the frontoparietals strongly
down curved, so that the braincase is relatively narrow
(25).
This apparently well-supported hypothesis conflicts
with another apparently well-supported hypothesis: Lhe
monophyly of a group formed by Barvgenys and the
Xenorhina-group. The derived character states shared
by these genera are the M. adductor inandibulae
posterior longus segmented with a superficial tendon
of insertion (12); the Maen. exiernus superficialis
arising parily from the fascia (13); origin of the M.
levaror Scapulee infertor in part from the ventral
surfaces of the first three vertebrae (18); a large deep
slip of the M, pecroralis sternalis (19); mesial expansion
of the vomero-palatine well developed (30); and
reduction in the number of denticles on the posterior
palatal fold (53).
Inaddition to the shared and derived states exclusive
to this group, a4 number of other states which .are
probably derived are shared also by other non-
asterophryine taxa: cultrilorm process of the
parasphenoid broad (Genvophrvnae) (32); crista
ventralis of the humerus well developed (Genpophrvne,
Eluchistacleis, Glyphoglossus and Kaloula) (42); and
eyes small (Copiula and Genyophryne) (47).
Despite the preponderance of apparent autapomor-
phies supporting the concept of Barveenys-Nenarhina-
group monophyly, and falsifying the hypothesis of
Phrynoniantis- Xenorhina-graup Asterophrys-group
monophyly, the latter hypothesis is preferable for the
following reasons.
(a) The monophyly of Barygenps and the Xerorhina-
group implies that the sister-group of Burveenys and
the Nenorhina-group is cither Phrynemarntis, the
Alyieraphrys-group, or a hypothetical common ancestor
otf those two groups (Fig. 30). Therefore the
hypothetical ancestor of Barvgenvs and the Xenorhina-
group would be postulated to possess the states of
characters4, 20, 24 and 25 possessed by the Xenarhina-
group aud the other asterophryines, and the states
possessed by Barygenys (M. geniohyeideus mediulis
present; genyophrynine-like conditions of the Mv
Barygenys alone of the
Noavernber (986
Renarhine Borygenys —Xenarhind
Barygenys
Phrynamans Asteniphrys~ group
Asrropnrys-
qroup Pn yromantis
Xenorhina
Asteruphrys-
group
Phrynaments PBorygenys
FIG, 30. Possible cladogramis of the Asterophryinag, UssumMing
Barvaenvs-Nenarhing group monyphyty
lumbricalis brevis VV of the manus and Mim. lubricates
hreves 1V and V of the pes; broad, relafively [lat
frontoparictals) to be reversals. ta the genyophrynine
condition, The implications of this are;
(i) Barygenys must have reacquired a muscle, the
M. geniohyoideus medialis (4) not present in its
ancestral species. While this is possible, it is less likely
than the situation in which a common ancestor of
Bar\veenys and the other asterophryines possessed the
muscle, and that it subsequently was lost in the ancestor
of the other asterophryines but retained in Barygerys.
(ii) The Asterophryines are all either terrestrial or
fossorial, whereas the genyophryines occupy a variety
of niches: scansorial, terrestrial, fossorial and aquatic.
The reversion of the specialised fossorial Barveenvs to
the generalised condition of hand and foot musculature
of the diverse genyophrynines (20, 24) appears unlikely.
(iii) As has been discussed above, the loss of
maxillary extension and connexion and acquisition of
a radically different mode of adaptation of the snout
to burrowing, as are implied by Lhe postulated common
ancestry of Barveenvs and the Xenorhina-yroup,
appear unlikely evolutionary events.
(b) Of the nine shared and derived characters
supporting the hypothesis of monophyly of Barveervs
and the Nenorhind-sroup, three relate to the
strengthening of the arm action (18, 19, 42), lwo to the
reinforcement ol the skull (30, 32), and two to the
reduction of size of the eye (13, 47), These all relate.
at least notionally, to the burrowing habit. The arm
action of Barvgenyvs when walking is extreme (as the
forearm is pulled through about 180°, from the vicinity
of the eye to the flank — personal observation), The
arms are involved in pullmg Barygenss through the
moss and leaf litter, and Characters 18, 19 and 42
appear to relate to this action. Acquisition of a similar
mode of burrowing in the Nenorhina-aroup, al whieh
Udid not observe living specimens, would be expected
1o lead to similar adaptations. That thvee of these
shared churacter slates (32, 42, 47) occurin unrelated
THE PAPUAN SUBFAMILY ASTEROPHRYINAL 439
fossorial species indicates. the possibility of convergent
acquisition of these states in the two fossorial genera
of asterophryines also.
(c) While differences in characters do not indicate
distance of relationship, | accept Arnold’s (1981)
argument that evolution in two taxa of different
solutions to common problems is indirect evidence that
the taxa may be distant. As well as differences in the
mode of reinforcement of the premaxillary region,
Barygenys and the Xenorhina-group differ m same
other forms of skull reinforcement: in Baryezenys alone,
there js a calcified are between the seprornaxillaries (4),
in the Nenerhina-group alone the anterior margins of
the palatine shelves of the maxillae project postero-
mesially towards the anterad process of the vomer-
palatine (39),
(d) The Asterophrys-group and the Nenarhina-group
share a derived state, possession of a dorsal crest of
the ilium (45). This state, which does not occur in other
microhylids is an apparent autapomorphy supporting
ithe hypothesis of monophyly of the Asrerophrvs-xroup
and the Nenorhina-group, which is incompatible with
Buryeenys-enorhina-group monophyly.
The evidence, therefore, favours the hypothesis that
Phrynamantis, the Xenorhina-group and the Astero-
phrys-group form a natural group, and that Barygenys
and the Xenarhina-graup do not.
Hypothesis 3. Monophyly of the Xenorhina-graup and
the Asteraphrys-proup,
The salution of the three-taxon problem of relation-
ships between As/erophrys-group, Phrynontantis and
the Xenorhina-group is modified by the establishment
of Barygenyvs as the sister-graup of the three taxa.
However, the estimation of polarities of character states
is not made easier. As the states of character (10), (12),
(13), (18), G9), (30), (32), (42), (47) and (53) exhibited
by Barvgenys conflict with those exhibited by the
genyophrynines, the fact that Barygenys joins the
Genyophryninae in the role of out-group to the three
taxa simply obscures the polarities of these characters,
The only unequivocal autapomorphy supporting the
monophyly of the Yenerhina-group and the Astera-
Phrys-group is the possession of the ilial crest (45),
Another shared and probably derived character is the
Asteraphr ys-
group
Xenariina = Parynamoantis
Barygenys
PIG. 31. Cladopram of the groups of mareropheyines.
extension ol the otic ramus medially to overlic the crista
parotica (37). This state is shared by Genvophrvne, and
a similar but not identical extensian is seen also in
Choerophryne and the darlingtoni-group. This state
has been subject to convergence and is not a reliatle
indicator of relationships,
This hypothesis of monophyly is in cantlict with (he
hypotheses that Phrynomantis and the Nenorhina-
group form a natural group, for which there is no
evidence, and that the Asferophrys-group and
Phrynomantis farm a natural group, which [ now
consider.
Hypothesis 6 Manophyly of Paryaomuaris and ihe
Asteraphrys-group,
This hypothesis is supported by one apparent
autapomorphy: origin of the At petrohyoideus
posterior 111 from the zygomatic ramus or its posterad
projection (9), This state occurs alsa in Elachistacleis,
There appears Lo be no way of deciding be{ween the
hypotheses 5 and 6 of monophyly presented here; Bath
characters (45) supporting monophyly of ihe
Xenorhina-group atid the Asterophrys-group and (9)
supporting monophyly of the Xenorhina-group and
Phrynomantis appeat equally valid. Given the present
state of knowledge, the best coufse appears to be to
admit that the relationship between the three groups
is unresolved, and best expressed on a cladogram as
a Irichotamy (Fig, 31),
Phylogeny of the Asterophrys-group: Here | attempt
to establish a phylogeny of the Asferophrys-group, This
exercise is undertaken an the assumption that the
Asterophrys-group is monophyletic (sensu Hefitiig,
1966), ie, that this group comprises all of the
descendants of a common ancestor. Phryvnamuntis
kopsteini was unavailable for dissection (only three
specimens are known), and as this may well be a
member of the Asterophrys-gtaup, conclusions
concerning the phylogeny of this group can only be
conditional. Evidence that P. kopsteini is a member of
the Asterophrys-group is that (1) the “subarticular
tubercles are strong” (Zieifel 1972, p. 479: contra
Mertens, 1930), and (b) “the anterior process of the
maxillae are only narrowly separated at the midline,
$0 the condition is virtually symphygnathine" (Zweitel
1972, p. 479). This condition of the maxillae appears
similar to that of Mantophryne infulata, in which ihe
maxillae are separated by between 0.2. and 0.3 mmm in
the three specimens available.
As has been pointed oul above, the Asreroplirvs-
group is very diverse, and within this group relation-
ships are obscure. Conflicting trends occur, eg.,
towards exostosis of the skull — none in Hv/opharinus
and Mantophryne infulata, slight in M. louisiactensis,
moderate in M. lateralis, heavy in Asteraphrys and
Pherohapsis — and towards lightening of the skull,
expressed in the lack of symphygnathy in Hyv/apharbes
and the marginal symphygnathy of Md. infilare,
Moreover, in a phylogenetic analysis, “trends” are
440 REC. 5S, AUST. MUS. 19 (19); 405-450
: Mantaghryne
Asterophrys Pherchopss feferalis M lowsjadensis
Minho
Aylaphertus
Montonaryne
Asrerophrys Pherahapsin lateralis Miouisiadesis Munfulote Mplapkorbus
27
$6
Mantaphryne
Hylophorbus infiuiati
Miloterais Asferaphrys Pherehapasis Mloulsjadensis.
FIG, 32. Three of the most parsimonious cladograms of the Astero-
phrys-eroup produced by the WISS methad. Horizontal strokes
through lines indicate acquisition of a derived character state;
crosses indicate reversals. Several cladoprains as parsimonious
as (B) can be produced with different treatments of character 46,
uninformative unless shared, derived character states
can be identified.
The establishment of character state polarities is also
complicated by uncertainty as to the appropriate sister-
group of the As/erophrys-group (Pig. 30), resulting
(rom any inability to resolve the Phrynomantis-Xenor-
hina-group-A sterophrys-group trichotomy. The sister-
group, the most appropriate functional outgroup for
this group of species, could be Barygenys or the
Xenorhina-group or Phrynomantis, depending on how
the trichotomy were resolved. Consequently, only six
characters potentially informative of relationships in
this group can be assigned polarities, and the polarity
of Character 38, symphygnathy of the upper jaw, is
November, 1986
TABLE 6. CHARACTER STATES OF THE
ASTEROPHRYS-GROUP
Character
Species 21 ey oT «6
Asterophrys turpicula | Ald ! et ]
Iivlophorbus rufescens | B 0 0 B | 0
Pherohapsis mensiesi 0 B 1 | A | |
Muantephryne infulata | B ) 0 A 1 ()
M. laieralts | RB 0 | A Ll ]
M. luuisivdensis l B 0) 0 A 0 1
' The relationship between the sagittal crest of Asreraphrys and the
_ Parasayittal ridgey of other species is unknown,
~As there is conflict in the information fram the outgroup,
polarities cannot be assigned,
unknown. On the other hand, the polarity of Character
46 is resolved, type I ilio-sacral articulation being the
derived state. The polarities are listed in Table 6,
The other characters given conflicting evidence of
relationship: Character 21 (division of the M, tibialis
anticus /ongus) supports the monophyly of
Asterophrys, Hyvlophorbus and Mantophryne;
Character 35 (connexion between the squamosal and
maxilla) supports the monophyly of Asterophrys and
Pherohapsis, Character 36 (posterad projection of the
zygomatic tamus) supports the monophyly of
Asterophrys, Pherohapsis and Muntophryne lateralis,
Character 46 (nature of the iliosacral articulation)
supports the monophyly of Aylophorbus, Pherohapsis,
Mantophryne infulata and M. lateralis; and
Character 52 (chin warts) supports the monophyly of
Asterophrys, Pherohapsis, Mantophrynre lateralis and
M. louisiadensis. There is no reason to believe that any
of these characters is more reliable than the others: the
derived states of Characters 35 and 46 have evolved
convergently within the Genyophryninae; the
homology of the prootic arcade of Pherohapsis with
the posterad process of As/erophrvs and Mantophryne
lateralis (36) is unproven (Introduction); and Character
21 may well be related in this group to the relative
length of the tibiofibula, which is relatively less in
Pherohapsis (han in other members of the Asterophrys-
group, and is variable within genera (Zweifel 1972), [In
order to illustrate graphically the possible relationships
Figs 32-34 show the most parsimonious cladograms
constructed by the Weighted Invariant Step Strategy
of Farms, Kluge and Eekhardt (1970) (Fig, 32), the
Waener Tree method of Lundbery (1972) (Fig, 33) and
a phylogenetic tree constructed by the Character
Compatibility Analysis method of Meacham (1981)
(Fig, 34), These demonstrate the ambiguities arising
from the uncertain polarity of Character 38 (cf. Figs
A and B of each Figure) and from the paucity of
characters. However, | conchide trom (hese figures the
following:
(1) Monophyly of the groups Asferophrys-Phero-
hapsis and Asterophrys-Pherohapsis-Mantophryne
luteralis appear to be best supported by the data,
(ii) No support is given to the hypothesis of the
monophyly of Manrophryne, M. lateralis appears to
be related lo Pherohapsis and Asterophrys, the
THE PAPUAN SUBFAMILY ASTEROPHRYINAE 44|
Monfapnrynre
intuloee
Miowsindensis Wiarerahs Pregrahapsis peleronneré
a
2) =
7 6
Ayvlapharbus
Manrognryne '
Ha AM nwsindensis Miuteraliz Phefanapas — ASneropnr vs
Aplnpnorbus
Man rophryne-
Jowsraden sts Mt jefulote M laterals Pherchaows Asteraphrys
I
38 45
at
Mylapnarbus
FIG. 33. Three of the most parsimonious cladograms of [he
Asterophrys-group produced by Lundberg's (1972) method.
relationships of M. /ouisiadensis are obscure, and the
close phenetic relationship A, infwlata bears to
Hylophorbus may or may not be the result of
symplesiomorphy.
(iii) Parallel acquisition of the type I ilio-sacral
articulation by Aylopherbus, Pherohapsis, M. infulata
and M. /aferalis and parallel reversal to the type ITA
articulation by Asterophrys and M. louisiadensis are
equally parsimonious hypotheses.
lt is clear that the data are insufficient for many
confident statements to be made about relationships
within this group.
Hypothesis 7. Monophyly of Xenorhina and
Xenobatrachus
The one feature distinguishing these genera is the
occurrence in Xenobairachus of the derived state of
Character (31), presence of one or more spike-like
odontoids on the vomero-palatine. I conclude that
Xenorhina is paraphyletic and Xenobatrachus arose
from a Xenorhina-like ancestor.
DISCUSSION
Taxonomic Implications of the Phylogenetic Analysis
Some of the data presented in this study point
inescapably to the necessity for taxonomic changes
which involve redefinition of the asterophryine genera,
The changes to be made will involve decisions related
to the philosophy of classification,
Few would disagree that a classification should be
useful. Yapp (1981, p. 245) encapsulated the functions
Prerahapsis Asteraphrys Pherohapsis Aterophrys
er 27
46 el AG
Mar tophryne
faferayis
Mapraparyne
laterals
Hy/opherbus
38
M intulara
infulota 4g
“v/opherbus
Ancestor Ancestor
A. B.
FIG. 34. ‘Two of the most parsimonious cladograms of the
Asterophrys-group produced by the Character Compatibility
Analysis method,
and properties of a useful classification: “The funda-
mental object of classification is to facilitate
economical statements. If we can agree that bats, cats
and rats are mammals, we say, ‘Mammals have hair,’
instead of ‘Bats... cats ... rats have hair,’ and if a
newly discovered kangaroo is a mammal, it too has
hair. A secondary value is to enable a museum curator
to put his specimens in appropriate cupboards.” Yapp
implies that whatever its basis, a classification must
establish “groupings about which generalizations can
be made” (Mayr 1981, p, 511), and so have maximal
predictive value, and, because its information content
is high, be stable in the face of new information.
Each of three schools of classification, the phenetic
(eg., Sneath and Sokal 1973) the evolutionary (e¢g.,
Mayr 1969; Michener 1978; Mayr 1981) and the
cladistic (¢.g., Farris 1977; Eldredge and Cracraft 1980)
claims to incorporate maximum information content
and predictive value into its classifications. Each claims
to seek out “natural groups”. However, the meaning
of “natural group” differs between the schools.
Pheneticists define natural groups as groups based on
overall similarity (Sneath and Sokal 1973, p. 24):
evolutionists define natural groups as groups sharing
greatest genetic similarity (Mayr 1969); and cladists
define natural groups as monophyletic (sensu Hennig
1966) or as groups sharing particular defining
characters (Nelson and Platnick 1981, p. 328). In the
recognition of natural groups, cladistic relationship is
either irrelevant,
“\.. this measure of similarity does not carry with it any
necessary implication as to the relationship by. ancesiry, but
does imply exhaustive estimates of similarity of jhe
phenotypes.” (Sneath and Sokal 1973, p. 28)
relevant, but not the whole story,
“a relationship, in the evolutionary sense is determined by
442 REC_S, AUST, MUS, 19 [19}: 405-4511
both processes of phylogeny, namely, branching tind subsequent
divergence.” (Mayr 1969, p. 70)
or all,
The prime goal of syaematics, according wo clidisis, iy (he
deliniiion and recognition of monophylene groups." (Eldredge
und Cracrafi 1980, p. LO)
The schools are not manolithic. For example, Holmes
(L980) reviewed the diversity of the evolutionary school,
and well publicised differences of opinion occur
between practitioners of transformed cladistics (1.c.,
cladistics divorced from evolutionary theory, eg.,
Platnick 1980; Patterson 1980; Nelson and Plathick
1981) and Hennmigian phylogeneticisls, who relate
phylogenctic patterns 10 evolution (e.g., Eldredge and
Cracralt 1980; Ball 1981; Wiley 1981; Beatty 1982),
However, though fot monolithic, the schools are well
delineated, and a systematist revising and redelining
Taxonomic groups must either knowingly or
unknowingly follaw the tenets of one of the schools.
The phenetic schoal claims that the most useful,
objective and stable classification arises out of an
analysis of as large a selection of characters as possible,
in order to gain a measure of overall similarity, without
reference to the evolution of the taxa. The stability of
phenetic classifications has been thrown inio daub by
the studies of Mickevich and Johnson (1976) Mickevich
(1978) and Schuh and Polhemus (1980),
Tt has long been known that different clustering
methods to discover groups of most similar taxa
produce different results, and this has been
demonstrated conclusively by Presch (1979). Though
Sneath and Sokal (1973, p. 31) saw a virtue in this
divergence:
"Some types of dillerences in resulls may Uiemselves be
of great (nteresr leading to new insights into the nature of the
organisms Gr GO! Lhe (axa bein seadied”,
in practice different results from different cluster
techniques appear rather to cause contusion. For
example, Blake (1973, p, 123) in his revision of the
myobatrachine frogs simply rejected the results of One
of his two clustering methods on the grounds that it
yielded results “inconsistent with other [unspecified]
indicators Of relationship’S Key (1976, p. 28), lacking
“any canvinceing ground of principle” for choosing
between eight different classifications derived fram the
same data by eight clustering techniques, adopted the
previously Tecognised species-groups as ihe basis for
his classification of the morabine grasshoppers. When
such actions are taken it is difficull to accept the
efficacy of phenetic classifications,
Some studies, notably that of Blackith and Blackith
(1967) on ortlopteroids, demonstrated convordance in
classifications based on phylovenetic and pheneric
methods. However, though they attribute this
concordance to their use of a large number of
characters, it appears more likely to be the result of
the high taxonomic level at which this study was carried
out. The study of Schuh and Polhemus (1980) on the
Leptopadomorpha indicate that al lower taxonomic
November, 1986
levels agreement between the results of phylogenctic
and phenelic methods is low.
Pheneticists sacrifice the information content of
phylogeny for objectiviry and stability. On both counts
phenetic techniques appear to fail, as groups claimed
lo be natural because they are based on overall
similarity must be chosen by a subjectively chosen
technique, and appear ra be susceptible to change if
new information is.added. The sense in which such a
vroup can be said fo be natural is obscure,
The other rwo svhools utilize phylogenetic data, but
differ in that the classifications of evelulionists
incorporate anagenetic information, as they claim that
this action increases the information content and
predictivity of their classilications, a claim disputed by
Farris (1977), Al the heart of the dispute between these
schools is the treatmen| of paraphyletic groups,
Evalutionists, e2,, Ashlack (1971, 1974), Mayr (1974),
Hrothers (1975), Bock ([977) and Michener (1977,
1978), consider paraphyleric groups natural and worthy
of recognition; phylogeneticists and “transformed”
cladists, g2., Hennig (1975), Nelson (1978), Eldredge
and Cracraft (1980), Nelson and Plaimick (1981) and
Wilcy (1979, 1981) consider only monophyletic (sensu
Hennig) groups natural.
The arguments.ot both sides ta this question appear
to have meril, [1 is argued by evolutionists that if it is
assumed thal, for example, fleas evolved fram a fly
species and therefore, flies are paraphyletic (Michener
1978), aname is needed for fleas and Mies to distinguish
them, as useful geteralizations can be made about each
group. Mayr (1974) argues also that paraphyletic groups
may be considered natural if they have retained
morphological similarities owing to their fidelity to an
adaptive zone (Ge. the reptiles), because this
morphological similarity may be assumed to be the
result of genetic similarity, and it is che relative genetic
uniformity of a group that makes it natural.
Against this view if is argued that paraphyletic
froups such as the Flies do nor necessarily represent
a group of most closely related species: amoug the flies
there is a Sisler-group to the fleas. That group of flies
is more closely related to the Meas than to the other
flics, and theretore paraphyletic groups should not be
considered natural: they are defined simply by lack of
the autapomorphies of the Meas. Indeed the assump-
tion that retained morphological similarity is
necessarily related to a retained genetic similarity is
questionable. Patterson (198ia) cites the study of Bruce
and Ayala (1979) lo show that, despite their morpho-
logical similarity to Pongo, the African apes. a
paraphyletic group with the taxonomic removal of man,
are verictically far closer to mau than to Pongo: and
Patterson shows that carp, a member of the para-
phyletic Osteichthyes, are much closer penetically ta
man and ta chickens, to which they are phylogenetically
closer, than to sharks whieh they resemble morpho-
logically. As assumed genetic affinity is the basis lor
THE PAPUAN SUBFAMILY ASTEROPHRYINAT 443
belie! in the naturalness of paraphyletic groups,
evolutionary classifications are al dubious value.
The demonstrated stability of cladistic classifications
(Mickevich 1978; Schuh and Polhemus 1980), che
necessity Gf cladistic classifications in some methods
of historical biogeovraphy (Brundin 1966, 1968; Rosen
1978; Patterson 198Ib), their high information content
(Farris 1977), and the unaninjous belief of evolutionists
and phylogeneticists (hat monophyletic groups are
nalural, suggest the desirability of adopting a
phylogenetic classification. However, as | pointed out
above, paraphyleic zroups, to the extent that they ure
useful, Le, ta the extent that userul generalizations can
be made about them, need to be recognized.
The Annotated Linnea Hicrarehy of Wiley (1979,
1981) is based on only monophyletic groups, bul can
accomnrodate paraphyletic and polyphletic groups, and
alsa groups of unknown affinities, Jt appears to offer
the kind of compromise that is needed. In Wiley's
scheme, laxa forming an asyminetrical part ata phylo-
generic tree may be placed al the same categorical rank
and sequenced in their phylogenetic order of origin,
thereby relaxing the Hennigian requirement of a
separate (axonomic rank for every fork of sucha tree.
Trichotomies are recognized by the term “seeds
multabilis” (o indicate the sequence of the (hree taxa
in the hierarchy is interchangeable. Groups that ure
paraphyletic, polyphyletic or of unknown status are
denied formal rank, and are labelled by inverred
commas and the term ‘“incerige sedis", As the
Annotated Linnean Hierarchy appears to offer a
phylogenetic classilication (hat has ihe capacity to
invorporate addijional useful anageneric information
without loss ol rigour, | use it in this revision of
asteraphryine taxonomy, wilh the modificanoen that as
the genus is a mandalory calegory, Lhe paraphyletic
venera, though labelled as such in the hierarchy, have
formal tank,
Jaxononic Reconmendatians
A. Genyopliryninae
The monaphyly of the Cenyophryninae has not been
dentonstrated, Nor has its paraphyly, The status of this
subfamily must await.a phylogenetic analysis of the
subfamilies of the Microhiylidac, and the action of
amalgamating the Genyophryninae and the Astero-
phryinae would be premature.
RB Phrveamantsa dariae
Phrynomantis doriae lacks the autapomorphies of
Ihe Xenorhing-groupy the supplementary slips of the
M, uvermandibularis arising from the angulosplenial;
the cultriform process of the AL. genioglossus busalls:
the striated tongue; and the posteromedial expansion
of the anterior corner of the palatine process of the
maxillary: If also lacks many other derived states shared
by Xenorhina and other genera (Characters 1, 13, 18,
19, 30, 47, 53).
P. doriae possesses ihe autapomorphies of
Phrvnomarttis: the two supplementary slips to the M7.
interinandibularis, the anterior ftom a deep tendon,
the posterior direct from the dentary; and the medial
Nange of the squamosal shaft reduced, the dorsal
surface of the otic ramus instead continuous with the
lateral surface of the shaft.
Therefore, | recommend that Xenorhina doriae be
transferred to tbe genus PArynromantis, as P dotiae
(Boulenger).
C. Nenorhina and Xenobatrachus
Nenorhina ws paraphyletic, differing from
NXenabatrachus only in Jacking one or more large
odontoids on rhe vomero-palating, There appear to be
{wo taxonomic options! Nenohatrachus could be
referred to the synonomy of Xenorhing Peters (which
has priority); or the genera retain their idevtities with
Nenorhina annotated to indicate its paraphyly.
The advantage of the first course is the elimination
of the Xevarhinu-Xenobalrachus dichotomy with its
unlortunate concomitant, the obliteration of the
relationships of the throat muscles when the diagnostic
odontoids are sought, The advantages of the secand
course are thal current nomenclature is conserved; aud
the identities of two monophyletic graups, Venobartra-
chus and the Aevtorhina-group are expressed, As the
second course appears to prodice a more informative
classification, | adopt il.
D. Mduntophryvne
The three species Mantophryne infulaia, M. lateralis
and MM. /avisiadensis show alTinities to the Aslerophrvs-
group rather than to PAryrnomanris. When these species
are removed, PArvnomantis beeomes a much more
homogeneous genus Supported by the autopomorphies
listed in the precediig discussion of P doriage. The
derived characters shared by these three species with
other members of the -4terophrys-group are: distal
origin of the MZ. tibialis anticus brevis (22), crested dium
(45); large subarticular tubercles (48); and the possible
apomorphy: orgin of the Ad. geniohvoeideus lateralis
internus entirely from {he hyale (5), It js therefore
appropriate ta remove these species [ram Parynd-
mantis. | recommend resurrection ef Mantophrvne
Boulenger 1897, type species At lateralis, 10
accammodate the three species, as there are no
compelling data to associate any of the species with
any of the existing Asieraphrvne-group genera. No
autopomorphy of Manfophrvve was found, so its
monophyly is unvertain. The status of this venus is
discussed Further in the next section.
E. The 4sierophevs-group
The Asferophrvs-group comprises three divergent
monolypic genera (As/eropArys, Mvlopherbus and
Pherohapsis) and Munrophryne, which is a group of
uncertain affinities and uneertain monaphyly,
444 REC, 8. AUST, MUS, 19 (19); 405-450
Although Avlopherhus ts phenotypically distinctive,
the polarity of its one defining generic character (38:
separated maxillae) is dubious, and thus its status as
a gerius is dubious, The taxonomic choice lies between
(a) inclusion of the six species into a smegle genus
(Asterophrys) and (b) retention of As/erophrys and
Pherohapsis, resurrecting Mantophryne Boulenger
1897 toa accommodate three of the species, and
regarding Aylophorbus. and Mantophryne as “incertae
sedis”, i.e, paraphyletic or of uncertain monophyly, and
“sedis mutabilis’, i.e., of doubtful order of arising.
] adapt the latter course of action because (a) it
conserves much of the current classilication; (b) as the
species of this group are very divergent, a single generic
name would be af little heuristic value; and (¢) the
monophyly of the entire group and the uncertainty of
the status of A/ophorbus and Manrophrvne are
expressed.
The convention of placing sistet-groups on the same
rank necessitates the insertion of a rank intermediate
between the subfamily and the genus, which have
traditionally been the only ranks between the family
and the species in the Microhylidae, J follow Parker
(1940) and Lynch (1971) who use the tribe as an
intermediate rank tn the Leptodactylidae,
The classificanon | favour is:
FAMILY: MICROHYLIDAE
SUBFAMILY: Asterophryinae
TRIBE: Barygenyini
GENUS: Barygenys
TRIBE: Asteraphryini sedis siutabilis
Asterophryinit incertue sedis “Hylo
phorbus" sedis mutabilis
Asterophryim incertae sedis “Manto
phryne” sedis mutahilis
GENUS: Asterophrys
GENUS: Pherohapsis
TRIBE: Phrynomantini sédi murebilis
GENUS: Parynananiis
TRIBE; Xenorhinini sedis mrutabilis
Xenorhininoi incertue sedis
“"Nenorhina”
GENUS: Xenobatrachus
F. Phrynomantis &. humicola and PA. compta
In distinguishing these subspecies, Zweifel (1972)
Stressed fCwo features: occurrence of a pale orange
postocular stripe in RA. compla (lacking in PA.
humicola), and possession by RA. Aumicoela of
relatively longer legs than those of Rf. comput.
According io Zweifel’s data the ranges of these taxa
are essentially allopatric: PA. conip/a is a western form
with jis range centring on the Madang and Western
Highlands Provinee, and extending eastwards into
Chimbu Pravince; PA. Aumicota is an eastern form
with its range centring On the Eastern Highlands.
Province and extending westwards into Chimbu
Province. Zweifel (1972) reported one case of sympalry
at lgindi, Chimbu Province, but was unwilling to name
November, 1946
Ph, humicofa and BA, compra as separate species until
evidence of reproductive isolation was available, as the
two (axa are very Similar morphologically,
These two taxa differ anatomically in two respects:
(a) Ph. camipra possesses a slip of the MW. depressar
mandibulae arising from the posterior margin af the
tympanic fing, Ph, Aumicala and same Barygen\s
Species are the only asterophryines to lack this slip. (b)
in PA, campra the mandibular branch of the trigeminal
nerve passes anicralaterally between the Mor.
adductores mandibulae posterior longus and anterior
longus. It passes laterally around the antenor margin
of the M.a.m. posteriar longus and posteroventrally
across the Jateral surface of that muscle until the nerve
reaches the mandible. RA, fumicola is the only
asterophryine which does hot conform to this pattern,
tmstead the nerve passes directly lateroventrally from
the braincase, penetrating the Mam. posterior langus.
It emerges on the lateral surface of the Ada. posterior
longus close to that muscle’s insertion on the mandible,
and passes ventrally on the lateral surface of the muscle
until it reaches the mandible.
The specimens examined are from allopatric sites.
Examination of sympatric specimens could provide
evidence of introgression or genetic isolation,
Systematic Accounts
ASTEROPHRYINAEB (part.) Parker, 1934
Asterophryinae Zweifel, 1971
Asterophryidae Giinther (1858) p. 51
Xenorhinidae Mivart (1869) p, 289
Symphygnathinae and Eleutherognathinae (part.)
Mehely (1901) pp. 172-189
Cacopinae (part.) Noble (193!) p. 531
Diagnostic definition: (2) posteromedially directed
dorsal sheet of fibres an M. in/ermandibularis lacking;
(3) overlap of Mm. interhyoideus and
intermandibularis (except Xenorhina bouwenst), (7)
deep slip of M, hvoglassus to hyoid lacking; (15) three
tendinous inscriptions of M_ rectus abdominis; (28),
(29) palatine and vamer fused and expanded both
laterally and mesially where each combined bane meets
its. fellow; (33) broad contact of medial ramus of
pterygoid with prootie; (34) broad contact of
quadratojugal with axilla; (41) dentary fused ta
anterior surface of mentomeckelian arid usually in
contact anteriorly with its fellaw (except MM vlopherbus),
mentomeckelians in contact at an acule angle; (43)
pectoral girdle lacking clavicles, procoracoids and
omosternum; (44) vertebral column diplasiocactous,
(49), (50) tongue adherent and divided superficially ito
anterior and posterior sections; (54) larva undergoing
metamorphosis in egg capsule, and lacking open gill
clefts, operculum and spiracle, respiration performed
through non-muscular, vascular taj] in known species.
Additional states not referred to in previaus secrions
also oceur; maxillary and vomerine teeth absent; ear
fully developed.
‘The states of characrers (2), (41) and the combination
THE PAPUAN SUBLUAMILY ASTFROPHRYINAE a4
of the states Of (49) and (50) are diagnastic of the
Asterophryinac. The Asterophryinae comprises four
tribes.
ASTEROPHRYINI New Tribe
This tribe accommodates the genera Asleroplirys,
Hylopherbus, Mantophryne and Pherohapsis.
Diagnostic definitian (4) M. gemohvoideus medialis
absent; (5) Ad. geniohyoideus lateralis internus arising
from hyale only: (6) M. geniog/lossus basalis labiform;
(8) three Mii. petrohyoidei posterioress (9) M,
petrohiyoideus posterior WL arising: (rom zygomatic
ramus of squamosal; (10) .M. depressor mandibulae not
arising [rom anterior 1/2. of ventral margin of tympanic
ring; (11) origin of Mf. depressor inanhdibulae from atic
ramus slender: (12) superficial tendon of M. adductor
mandibulae posterior longus lacking; (13) M, adductor
mundibulae externus superficialis arising entirely from
zygomatic ramus; (14) Mt. lonwissinins dorsi noi
inserting on dorsal fascia; (16) Me. lransversi
abdominis and Mm. obliqui abdominis externi
terminating on broad ventral aponeurases; (17) two
sites oF insertion of M, serra(us medius; (V8) origin of
M. levator scapulae inferior partly from first (wo
vertebrae: (19) deep slip to M. pectoralis siernalis
slender, (20) medial slip of ML dusibricalis brevis digiti
IV of manus cylindrical, arising from centrale
postaxiale; (22).M. (hialis enticus. brevis arisifig from
distally on tibiofibula; (23) AL opponens hallucis
arising from a distal tarsale; (24) Mim, lambricales
breves digitorum \V and V of pes fused along most
of their length; (25) frontoparietals. 3~ as long as
broad; (30) median expansion of yomtero-palatine
moderate; (32) culttiferin process of parasphenoid
narrows (37) dorsal surface al otic ramus of squamosal
continuous with anterior surface of medial flange al’
squamosal shalt; (38) maxillae overlapping premanillae
and connected by ligament or suture, no ridges on
snout; (39) mesial expansion of anterior margin af
palatine shelf of maxilla lacking; (40) anterior margin
Of nasal capsule pot calcified: (42) humerus relatively
Straight, crest moderately developed; (45) ilium bearing
a moderate dorsal crest; (47) eye large (Er S-Y > 0.090);
(48) subarficular tubercles large and prominent; (51)
surface Of posterior section of tongue pilted uniformly;
(53) posterior pre-pharangeal fold) bearing Jarge
number oF denticles (rarely fewer than ten).
OF these states, (5), (22) and (48) are diagnostic of
the Asterophryini
GENUS Asterophrys TSCEIUDI, 1838
Asteraphrys Tschudi (1838) p. 82
(type species Cerarophrys turpicofe Schlegel
[1837 (1838-1844)|
“lsierophirys (part), Parker (1934) p. 66
Asterophrys Zweite] (1972) p, 432
Diagnostic definition: (1) Wwo stipplementary slips 10
M. intermondibularis, cach avising from ventral margin
of aungulosplenial; (12) AL adductor mandible
ran
posterior longus divided into segments; (21) Md. Ubialis
anticus longus divided distally; (26) parasagittal ridges
on frontoparietals lacking) (27) sagittal crest on
cranium well developed; (35) squamosal and maxilla
meeting in large specimens; (36) posterad extension of
zygomatic ramus well developed; (38) maxillae meeting
anteriorly; (46) direct ligamentous connection between
ilium and sacrurn; (52) warls on chin well develaped.
The states of (1), (£2), (27) and (52) are peculiar to
Asteraphrys among the Asterophryini.
Content: A. turpicula (Schlegel).
GENUS Hylophorhus MACLEAY 1878
Avlophorbus Macleay (1878) p. 136
(type species H. rufescens Macleay)
Mantophryne (part,) Boulenger (1897) p. 12
Metopostira Mehely (901) p. 239
Hylopharhus Zweilel, 1972
Diagnostic definition: (1) one supplerneniary Slip Lo
M. intermandibularis via tendon trom deep on
angiilosplenials (12) ML adductor mandibulae posterior
longus unsegmented; (21) M4 fibiulis anticus lonzus
divided distally; (26) parasagittal ridges present; (27)
sagittal crest lacking; (35) squaniosal and maxilla nol
meeting; (36) posterad extension of zygamatic ramus
a slight lange only; (38) maxillag well separated (about
lL mm) anteriorly; (46) ligamentous connexion between
ilium and sacrurh lackinws (52) chin warts lacking.
The State of characler (38) is peculiar fo
Hylophorbus among the Asterophrytni.
Content: There are (hree subspecies of Hy/ophorias
rufescens Macleay: Hr rufescens: Hur, exiimus Zweilel
1972; Hin, myopicus Aweilel 1972.
Remarks: This genus may be paraphyletic as the
polarity of character (38) is dubious.
GENUS Mantophryne BOULENGER 1897
Mantophryne Boulenger (1897) p. 12
(type species M. /ateralis Boulenger)
Mantophryne Vogt (1911) p, 427
Hylophorbus Fry (1913) p, 48
Asteroplirys (part,) Parker (1934) pp. 62, 63
Phrynamantis (part.) Zweitel (1972) pp, 476-474,
480-489
Diagnostic definition: CG) one slip to MM,
intermendibularts via a tendon from deep on
angulosplenial; (J2).M. adductor mandibulae posteriar
fongus unsepmenied; (21) ML ribialis anticus longus
divided distally; (26) parasagittal ridges present or
absent; (27) sagittal erest absent; (35) squamosal and
maxilla not mecling; (36) posterad projection of
zygomatic ramus well or poorly developed; (38)
maxillae meeting of only narrowly separated
(maximum 0.3 mm) unteriorly; (46) ligamentous
connexion between ihium and savrum present or absent;
(52) chin warts present or absent-
Content: M. infulata (Zweilel) 1972; MM laterults
Boulenger; WW. dowisiudensis (Parker) 1934,
Remarks: Vhis is probably a paraphyletic genus, and
is relaiionships with the other genera of the
AAG
Asterophryini are uncertain. te differs from
Hylophorbus in Character 38, from Asterophrys in
Characters |, 12, 27 and $2, and from Pherolidpsis in
Characters 21 and 36, J was not able to examine
Phrynomarts kopsteini Mertens, which may be closely
related to the species Which comprise Mantephryne,
GENUS Pherohapsis ZWEIPEL 1972
Pherohapsis Zweifel (1972) p. 456
(\ype species BR mensziesi Zweifel)
Diaghostic déefinitiens (1) one slip to M
iniermandibularis, via tendon from deep on
angulosplenial; (21) A, rihidlis antieus. forges divided
relatively proximally: (26) parasagittal ridges present,
(27) sagittal crest lacking; (35) squamosal and maxilla
meeling; (34) posterad extension of zygomatic ramus
extended medially to meet frontoparictal to which
fused (o form proatic arch; (38) maxillae meeting
anteriorly; (46) ligamentous cannexion between ilium
and sacrum lacking; (52) chin warts poorly developed.
The slates of characters (21) and (36) are peculiar
to Pherohupsis among the Asterophryini.
Content: P. menziesi Zweitel.
BARYGENYINI New Tribe
This tribe accommodates the zenus Bar\genys,
Diagnostic definition: (L) wo supplementary slips to
M. intermandibularis ‘rom the ventral surface of
angulosplenial, posterior inserting on M. venlog/ossus
hasalis by narrow tendon; (4) M. geniohvatdeus
medialis present or absent; (5) M, geniahvoideus
lateralis internus arising from mandible only; (&) M4,
venioglassus basalis lamellate; (8) two Mim,
petrohvoidei posteriares; (9) M. petrohyoideus
posterior 1 arising from exoccipital and oli¢ ranius;
(10) M4. depressor niandibulue not arising from anterior
1/2 of ventral margin of tympanic nog; (11) origin of
M. depressor mandibulae (rom the otic ramus absent
or slender: (12) AZ, adductor mandibylae posterior
longus: superficial tendon present; segmented in larger
species; (13) anterior origin of M4. adducior mandibulue
exrernus superficialix (ram fascias (4) M. longissinins
dorsi inserting in part on dorsal fascia; (16) Adm,
transverst abdvininis meeting on ventral abdamen,
Min, obliqui abdominis exterai meocting on ventral
abdamen; (17) only one site of insertion of M. serratus
medius; (18) ongin of AW levator scapulue inferior
partly trom first three vertebrae; (19) deep slip of M,
pectoralis sternalis well develaped; (20) medial slip of
M. lumbricalis brevis digit’ VV of manus a narrow sheet
from superficial tendon; (22) M, tblalis anticus brevis
arising relatively proximally; (23).M, opponens hallucis
ansing from palmar aponeurasis; (24). Mm. /unbricales
brevey digitortin VW atid V separated proximally; (25)
fronloparielals twice as long a& broad) (30) median
expansion of yomero-palatine well developed) (32)
cultriform process of parasphenoid broad; (37) dorsal
surface of otic ramus of squamosal continuous with
anterior surface al medial Mange of squamosal shaft,
REC. S$. AUST. MUS. 19 (19): 405-450
Noveniher 1986
and overlying anieriar surface of proatic as well as
dorsal surface; (38) premaxillac compressed between
maxillae: maxillae not meeting; three vertical, dermal
tidges on snout; (39) mesial expansion of anterior
margin at palatine Shelf of maxilla lacking; (40)
anterior margin of nasal capsule calcified; (42)
humerus curved, crest well developed; (45) ilial crest
lacking; (46) direct ligamentous attachment between
ilituro and sacrum; (47) eye small (E: S-V < 0,090); (48)
subarticular (ubercles absent or poorly develaped; (51)
surface of posterior part of tongue pitted unvoruily;
(53) posterior pre-pharyngeal told with < 10 denticles,
OF these states, (1), (4), (6), (8), (4), (16), (17), (29),
(23), (24), (233, (37), (38) and (40) are peculiar to
Baryeenyini among the Asteraphryinag,
GENUS Barveenvs PARKER 1936
Baryveenvs Parker (1936) p. 73
(type species & cheesmanae Parker)
Baragenvys |sic] Zweitel (1956) p, 9
Didsnostic definition; As above for the tribe
Barygenyini
Content: & atira (Gunther), 1896; B cheesmanae
Parker, 1936; & exsu/ Zweifel, 1963; 8 /levigularis
Zweilel, 1972; B maculata Menzies and Tyler, 1977; B.
nana Zweilel, 1972,
Remarks: A new species from Mt Missum 1s bemg
deseribed (Allison and Burton, i prep,).
PHRYNOMANTINI New Tribe
This tribe accommodates the genus Phryromantis,
Diagnastic definition: (1) two slips to M.
infermundibuluris, anterior via narrow tendon,
posterior direct from dentary; (4) Mi. geniohyoideus
thediulis absent, (S) M. wenlohyoideus lateralis internus
arising from both mandible and hyale; (6) M.
venioglassyus basalis labiform (8) three Adar.
petrohyoidei posteriores; (9) M. pétrohyoideus
posterior UW from zyvomatic ramus; (10) Mf. depressor
mandibulae with origin usually from entire ventral
margin ol tympanic ring, rarely pasteriar t/2; UL)
origin of Mt. depressor mandibulue (rom otic ramus
slender or moderate; (12) M. vdduetor manditulae
posterior longus unsegmented: (13) anterior origin of
M adductor mandibulae externus superficialis entirely
from zygomatic ramus; (14) M. longissimus dorsi not
inserting on dorsal fascia; (16) Marr transverst
abdominis and abliqui ahdominis externi terminating
on ventral abdominal aponeuroses; (17) two sites of
insertion Of M. serratus medius; (8) origin ol) AV,
levator seapylae inferior partly from first (we vertebrae;
(19) deep slip of M. peeloralis sternalis poorly
developed; (20) medial slip of AX /umbricalis brevis
diviti (V of manus cylindrical, arisine from centrale
postaxiale, (22) M. ribialts anticus brevis arising
relajively proximally; (23). M. opponens hallucis ansing
from a distal tarsale; (24) Men, lumoricales breves
divitorum LV and V fused along much of length; (25)
frontoparietals 3» ds long as broad; (30) median
expansion of yomero-palaline moderately developed;
THE PAPUAN SUBFAMILY ASTEROPHRYINAF 442
(32) culliform process al parasphenoid narrow; (37)
dorsal surface of otic ramus continuous with lateral
surface af squamosal shaft, medial flange poorly
developed; (38) maxillae meeting anteriorly, no ridges
On soout; (39) mesial expansion of anterior margin of
palatine shelf of maxilla lacking; (40) anterior margin
of nasal capsule not calcified; (42) humerus relatively
straight, chest moderately developed; (45) ilial cresl
lackinw; (46) direct hgamentous attachment between
ilium and sacrum; (47) eye large (FE; SY > 0,090
usually); (48) subarticular tubercles poorly developed;
(51) surface of posterior part af tongue pitted
unitormly; (53) posterior pre-pharyngeal fold hearing
rarely fewer than ten denticles,
OF these states, (1), (10).and (37) are peculiar to the
Phrynomantini among the Asteropliryinae,
GENUS Phrvnamantis PETERS 1867
Phrynomanris Peters (1867) p, 35
(lype species by subsequent designation [Noble
(1926) p. 20) P fusea Peters)
Callulops Boulenger (1898) p. 345
Gnathophryvne Mehely (1901) p, 177
Pomataops Barbour (1910) p. 89
Phrynomantis Zweilel (1972) p, 460
Diagnostic definition: As above for the tribe
Phrynomaatiqn.
Conrent: PB boettger; (Mehcly) 1901; PB doriae
(Boulenger) 1888; 8 dubia (Boetiger) 1895; P
eurvdacila Zwelel, 1972; P fusca Peters, 1867; P
slandilosu Zweilel, 1972; 2 Aumicola Zweitel, 1972:
P kopsieini (Mertens) 1989; P sfateri (Laveridge) 1955;
P stictogaster Zweifel, 1972; PB wilhelmana (Loveridge)
1948.
Remarks: | have not seen & boettgeri, PB dubia, P
glundulosa or BP kopsreini. While it is likely that the
first three Species are correctly assigned to PhAryao-
mons, the large subarlicular tubercles and narrawly
separated maxillac of 2 Avpsteint (Zweifel, 1972)
indicate (hat it may be more appropriately assigned to
Mantophryne. Two subspecies ol PB humicola (Ph.
hunicola and Ph. compta) have been described
(Zweilel, 1972), 1 have added (above) characters
emphasizing their phenotypic distirictness,
XENORHININI New Tribe
This tribe accommodates the penera Nenohatrachus
and Xenorlina.
Diagnesue definition: (1) bwo supplementary slips ta
M. intermandibularis from the ventral margin of
aiguloasplenial, pasterior inserting on median
aponeurosis; (4)... gentahyardeus niedialis absent: (5)
M~ geniohvoideus lateralis arising from mandible only;
(6) M. ventoglossus basalis with cultritorm posterior
projection; (8) three Vine. petrolvoidei posteriares; (9)
Mf. petrohvaideus posterior UU arising from exoccipital
and otic ramus; (LO) M, depressor mandibulae lacking
origin fram anterior 1/2 of ventral margin of trypanic
ring; (11) origin of ML depressor mandibulae trom atic
ramus broad, well developed; (12) At adductor
mandibulae posterio’ longus seamented, bearing
superficial tendon ol insertion; (13) anterior origin of
M. adductor mandibulue externus superfictalis {rom
fascia; (14) M. /ongissimus dorsi not inserting on dorsal
fascia; (16) Mim. rrensverst abdominis and obliqui
abdominis exierni terminating an ventral abdorninal
aponeuroses, (17) two sites of insertion of M. serraries
medius; (18) origin of At. levator scapulie inferior
partly from first three vertebrae; (19) deep slip of M.
pectoralis slernalis well developed; (20 medial slip of
M. lumbricalis brevis digit? WV of manus cylindrical,
arising from centrale postaxiale; (22) M, tibialis anticus
brevis arising relatively proximally; (23) 4. opponens
hallucis arising from a distal tarsale; (24) Mori,
lumbricales: breves digitorum \WV and V of pes fused
along much of length; (25) frontoparietals 3« as long
as broad; (30) median expansion of yomero-palatine
Well developed; (32) cultriform process of parasphenoid
broad; (37) dorsal surface of otic ramus continuaus
with antecior surface of medial flange of squamosal
shall; (38) maxillae meeting anteriorly, no ridge on
snout, (39) mesial expansion of anterior margin of
palatine shelf of maxilla well developed; (4) anterior
margin of nasal capsule nol calcified; (42) humerus
curved, crest well developed; (45) ilium bearing
moderately developed dorsal crest; (46) direct
ligamentous altachment between ilium and sacrum;
(47) eye small (E; SV < 0.090 usually); (48)
subarticular tubercles absent or poorly developed; (51)
surface of posterior part of tongue bearing deep
longitudinal striae (53) posterior pre-pharynyeal tald
bearitre few denticles (usually < 10).
Of these states, (6), (11), (39) and (51) are peculiar
to the Xenorhinini among the Asterophryinac.
GENUS Neénobatrachus PETERS aud
DORIA 1878
Xenobatrachus Peters and Doria (1878) p, 432
(type species X. ophivdan Peters and Doria)
Choanacantha Mehely (1898) p. 175
Diagnostic definition: (1) one or more spike-like
odontoids on the vomero-palatine.
Canrent: X, bidens (van Kampen) (909: 4. giganiens
(van Karnpen) 3915; 3. macrops (van Kampen) 1909;
NX. mehelyi (Boulenger) 1898; X. obesus Zweifel 1960;
AX ocellatus (van Kampen) 1913; ¥, aphiodon Peters
and Daria, 1878; X. rosrratus (Mehely) i898; x,
subcroceus Menzies and Tyler, 1977.
GENUS NXenorhina PETERS 1863
Nenorhina Peters (1863) p, 82
Ufype species, Bontbinaror oxycephaius
Schlegel)
Pseudengystoma de Witte (1930) p, 132
Asterophrys (part.) Parker (1934) p. 58
Xenorhina Zweitel (L972) pp. 529
Diagnostic definitions AS aboye for the isthe
Xenorhinini,
Content: X. bouwensi (de Witte) 1930) X~ riininea
(Parker) (934, AL oxveephals (Schlegel) 1858; XN,
44% REC. S. AUST. MUS. 19 (19); 405-450
parkerorum éweifel, 1972; X. similis Zweifel, 1956.
Remarks: This is a paraphyletic genus closely related
to Xenobutrachus.
ACKNOWLEDGMENTS
This project was supported in part by grants ram
the Australian Museum and the University of Adelaide.
The Department of Science and Industry supparted
field work In Papua New Guinea and the Department
of Biology, UPNG, supported my visa application, In
PNG | was assisted in many ways by Dennis and Krista
Black, Roy and Margaret Mackay, Rupert Stocks and
Allen Atlison..
Specimens were provided by the following institu-
tions: American Museum of Natural History (R. G.
“weilel); Australian Museum (H. G. Cogeer); Museum
of Comparative Zoology (J. Rosado), Rijksmuseum
van Natuurlijke Historie, Leiden (M. Hoogmoed);
South Australian Museum (M. J. Tyler); University of
Papua New Guinea (L. Hill, D. Black). Keith
MeDonald provided live specimens from Queensland,
Ross McDonald took radiographs of specimens
deemed too precious lo dissecl, Mike Tyler suggested
and supervised this project and provided access to
specimens so that | could get it under way, Mike was
an unfailing sauree of support and encouragement
throughout the project. He read and criticised dratts
of the manuscript. Margaret Davies aided me by
instruction and encouragement with artwork and in the
divers arts of osteology, and critically read drafts of
the manuscript, Philippa Hortan led me back to the
long-forgotten realms of histology, and helped me with
proof-reading. | awe much to the generosity and
forbearance of these three. Terry Schwaner, Jenny
Gardner, Graeme Brawning, Alice Wells, Cherie Burton
and Mary Meredith lent their ears and gave useful
advice and ideas when | needed them, Linda Trueb gave
meé invaluable advice On andtomical drawings, and
made useful suggestians for this paper:
Rev. Br. John Marks, my former employer, gave me
time off te complete this project with generous
vonditions, and some al iny calleazues, particularly
Mary Meredith aud Peter Taylor, bore added burdens
as a result ol my absence.
This project could not have bepun withour the
pioneering work of Dr R, G. Zweifel, who placed the
taxonomy of the Asterophryinae ona firm basis. He
was pronipe and generous in the provision of
Specimens, transparencies, advice and encouragement.
It was my wife, Cherie, who Suggested that I take
lime off work to finish this projeet, | owe much to her
tolerance, constant support and encouragement,
Heather Kimber and Sundrmi Lawson typed the
manuseript and Ruth Evans drew Fizs 28, 32-34.
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