RECORDS
OF
THE
SOUTH
AUSTRALIAN
MUSEUM
VOLUME 21 PART1.
MAY 1987
CONTENTS:
EDMONDS, §. J.
Obituary of I. M. Thomas
EDMONDS, 8. J.
Echiurans from Australia (Echiura)
HERCUS, L. A.
Looking for Ditji-mingka
JAGO, J. B. & PLEDGE, N. S.
Obituary of B. Daily
KERZHNER, I. M.
Nabidae (Heteroptera) of Vanuatu
LEE, D. C.
Introductory study of advanced oribate mites
(Acarida: Cryptostigmata: Planofissurae) and a
redescription of the only valid species of
Constrictobates (Oripodoidea)
PATTERSON, C. & RICH, P. V.
The fossil history of the emus, Dromaius (Aves: Dromaiinae)
RILEY, J. & SPRATT, D. M.
Further observations on pentastomids (Arthropoda)
parasitic in Australian reptiles and mammals
SUTTON, P.
From horizontal to perpendicular: two recent books on
central Australian Aboriginal painting
TINDALE, N. B.
Kariara views on some rock engravings at Port Hedland,
Western Australia
TRIGGER, D. S.
Inland, coast and islands; traditional Aboriginal
society and material culture in a region of the
southern Gulf of Carpentaria
WATTS, C. H. S.
Revision of Australian Berosus Leach
(Coleoptera: Hydrophilidae)
ZEIDLER, W.
The scaled-squid, Lepidoteuthis grimaldii Joubin, from
southern Australian waters
Volume 21(1) was published on 22 July 1987.
Volume 21(2) was published on 24 December 1987.
ISSN 0081-2676
PAGES
61-63
119-138
149-156
65-68
29-33
35-42
85-117
139-147
161-165
43-59
69-84
1-28
157-159
REVISION OF AUSTRALIAN BEROSUS LEACH (COLEOPTERA :
HYDROPHILIDAE)
BY H. S. WATTS
Summary
The Australian members of the hydrophilid genus Berosus are revised and redescribed. A key to
species is give. Thirty-two species are recognised of which 18 are described as new: B. amoenus, B.
arcus, B. aquilo, B. dallasae, B. gibbae, B. josephenae, B. juxta discolor, B. reardoni, B.
macropunctatus, B. sadieae, B. nicholasi, B. niger, B. quadrapunctatus, B. trishae, B. timmsi, B.
veronicae, B. ralphi and B. vijae.
REVISION OF AUSTRALIAN BEROSUS LEACH (COLEOPTERA: HYDROPHILIDAE)
©. 1.5, WATTS
WATTS, ©. HS. 1987, Revision of Australian Berosus Leach (Coledptera: Hydrophilidae). Ree:
Se lua Mis. 211) 1-28.
The Australian members of (he hydrophilid genus Berosyvs are revised and redeseribed. A
key [Oo splecies is 2iven. Thirly-fWe species are recognised of which 18 are descrihed as new: 2
umoonus, Burens, B aquily, B dallasae, B. gibhae, B. josephenae, Bo justadiscolar, B. reardoni,
B wnecropuncratus, B sacieae B. nicholas) B niger, B. quadrupuncralns, B trishae, B tiniest,
B, vervnicae, B, ralphi and B. yijee,
The following synonymies are proposed: B /lindersi Blackburn
B. sinnlans Blackburn — B&. stigitatico/lis Fatrmaine: = B. auriveps
B. approximans Pairmaire, Bo quartinus d'Orchmenr -
B. queenslandicus Blackbur; B wrayis Blackburn
B ovipentis kainmaire
Wlackburn = & blackhurnt Zain
B. discolor Blackburn:
B wustratige Mutsarit.
Co bLOS. Waus, Sourh Australian Museum, North Terrace, Adelaide. Soul Australia S000
Manuseripr received 24 July J98s.
The hydrophilid fauna of Ausiralia js relatively
rich in species and numbers of individuals. They
form uw conspicuous part of tle aquatic fauna. In
general they are well-colleeted and well-represented
in vollections, Despite this they have received litle
recent taxonomic attention.
A major difficully in dealing with Berasus Leach,
IS? in Australia was the inacequacies af existing
keys which reflected a weak taxonomic base exens-
plified by the facet that some [8 of the 32 known
species were undesenbed, including the commonest
southern species. At the same time some widespread
species were deseribed under as many as five
different names.
The last biajor atlemipt jo revise Australian
Berosus was by Blackburn in 1898,
Some characters used in the key, such as leg
colour and details of Lhe punctation, are nat totally
reliable, but the male genitalia have provided good
characters i all cases.
The genus Berasus is a member of |he subfamily
Berosinae which is characterized by a markedly
deflexed head, seven- or cighteseemented antennae
and meso- and metasternae without a continuous
common keel,
Berosus can be separated trom the other
Australian genera in the subfamily (Regéinbartia,
Gloharia and Amphiopsy by possessian af
protruding eyes, seven-segmented antennae, and
striae on elytra, although in some species these are
partially masked by strong elytral punctations.
Adults of all bur a few al the smaller species are
yellow-brown in colour They are found mainly
among vegetaiion at the boron and sides of ponds
and pools. Adulbs are often taken at light sometimes
some distance from water. The larvae are distinetive
witha series of lateral filamentous appendages ard
live in pools in similar situations to the adules. |
know of no detailed description of the larvae of any
Australian species,
Berosus is a large cosmopolitan genus and many
attempts have been made to subdivide it, none of
which appear particularly satisfactory to me. In
particular, the subgenus /¥ygrorrophus W. Macleay
which includes only the species & (4.) nutans and
B (H.) devisi, both Australian species, seems
unwarranted and to be based on an undue emphiasts
on the punetation of the upper surface.
Within Australian species there are five more or
less distinctive groupins.
Group | species are small, have the midline of
the mesosternum produced downwards in a distinet
ridge, head black, metacoxal process weakly to
moderately lobed laterally, last abdominal seement
notched and the second elytra! stria 1/4-the length
of the elytron (& discolor, B. approximans,
B, reardoni, B. juxtadiscolor, B. limmsi).
Group 2 species are small, with the midline of
Ihe mesosternum produced downwards in a distinct
pillac, head black, metacoxal process strongly lobed
laterally, the last abdominal segment notched, the
second elytral stria about 1/2 the length of elytron,
and with strongly developed medial ridge on first
abdominal segment (& macropunctatus,
8 quadrapunctatus, 8. trishae),
Group 3 species are similar to Group 2 specics
except for a less developed medial ridge on first
abdominal! segment, strongly humpbacked which
is associated with the presence of a space between
elytral striae 7 and 8 and/or 8 and 9 in the middle
of the elytron (A involutus. AL niger, & areus).
Group 4 species are similar to Ciroup 2 but are
te
larger and have the second clytral sta reaching
about two-thirds the lenpih of the eélytran
(B duplopunctatus and B. queenslandicus).
Group 5 species are generally larger with brown
heads, midline of mesosiernum with a weak central
keel, und metacoxal process withaul lateral lobes
(BO nutans, B, pulchellus, B dallasi,
8B. macumbensis, B. imwittipennis, B. anoenus,
8, josephenae, & gibbae, B. majusculis,
B. veronicae, B. aquila, B. australiae, B, decipiens,
B sadieae, B nicholasi, B. vijae, B. subovatus,
B. ralphi and B. debilipennis),
Groups 1-4 have similarities with cach other and
roughly correspond to the subgenus Berosus Hope.
Group 5 species correspond to the subgenera
Lnoplurus Hope and AHyzrotrophus W. MacLeay.
Since Berosus is a world-wide genus, Surther
consideration of subgeneric groupings should in-
clude extra-Australian species and is beyond the
scope ol this revision.
The collections from whieh specimens were
examined are listed under the following
abbreviations:
AM Australian Museum, Sydney,
ANIC Australian National Inseet Collection,
Canberra,
BM(NH) &ritish Museuni (Natural History).
London,
OW Private collection of author.
NMV National Museutn of Vivtoria,
Melbourne,
NSW DA New South Wales Department of
Agriculture, Sydney.
NTM Northern Territory Museum, Darwin,
MNHN Museuni National d'Histoire
Nalurelle, Paris.
SAM South Australian Museum, Adelaide,
WAM Wesiern Australian Museum, Perth.
OP! Queensland Department of Primary
Industry, Mareeba.
KEY TO AUSTRALIAN SPECIES OF BEROSUS
1, Midhine of mesosternum produced downwards
in a distinct pillar (Fig. 10), Aor projecting
backwards heryeen metavoxue. Mead black
imetallic), Metacoxal process lobed laterally
(rudimentary in &. discolor). Last abdominal
segment notched, Small (< 7mm). --.. ..2
| Midhine of mesosternum with weak central keel,
projecting weakly forward between mesoconac
Head completely or partially brown except
B pulchellus whieh lacks elytal stfiae and
B amoenus whieh has a predominantly black
Pronolum and elytra also. Melacaxal process
irvangular (Pig. 1) 22... + + ay,
2,(1) Second elytral soja about 4 feneth ‘of elvinontd
2, Second elylral stra 1/2 length of ebyiron
©. WS. WATTS
3, 42)
3.
4. (3)
4,
. (4)
§,
b. (2)
6,
7, (5)
7.
B. 16)
R.
9. (4)
9.
10. (9)
10,
1) (Ry
i.
Rugose portion at metalentur <1/2 teneth of
fomutr (Fig. 2), Mesosternul pillar flat.on ventral
COG evens | tppceee & discolor Blackburn
Rugose portion ol metalenmuy S12 lengil of
fomtur. Mesosiernal pillar weakly concave of
veritral edge rete:: |
Riuizose portion of inesofernuir 12 2-2/3 length of
fa) Ue ee er, awe |
Rugose portion of mesofemur <1 /2 length ot
femur . ot B approximans bairmane
Rugose portion of (feta: and meso-lemora pale,
same colour as rest of leg. B reardoni spnov.
Rugose portions of meta- and meso-femora
much darker |Man vest of leg 2.0, 2.2... 7
Second elyiral sija about 1/2 length of vlytiron,
fugose portion of metafemur S1/2 length ot
Femina ge sire ft iigaees. 9 cake, a)
Second elytral seria aul least 2/3 lenerh af elytron,
rigose poruion of metafemur <1 /2 lengih of
PRAMS s SoS ae Oo =~ -e TF
Punclures on sides af pronorum never in size,
Interstvial pupclures small, subobsolete, hind
lemurof male with triungilar bulge an hind edge
we Banta’ sp.nev
Panatiiver on 1 sittes of prenewum evenly sized with
ouly a few smaller anes, inferstrial punctures
moderately marked, hind femur of male normal
ctr teers e ees B ytvtadiseoler sp.nov
Space be} ween ely(ral striae 7 and 8 and/or 8 and
9 prealer in middle than at ends of elytron (Fig.
8), Medial ridge on firsr abdominal seyment
1/4 as deep as long, wilhoul backward projection
on ventral edge
Elytral striae 7, 8 and ¥ subparallel with each
other, Medial ridge on fest abclontinal segment
deep | 3-1/2 as deep-as long, wilh a backward
projection on ventral edge -—,-.220,2--,,, 0
Elyrral siria 8 raised and cacinatein middle and
towards apex. Distance between elyiral striae 8
and Y greatly enlarged i iniddle (Fig, 8),
Interstrial punctures weak, subobsolete. Northern
SEPRCIES Scare oicjciem O07 tan ae Dever aoe tll
Elvtral stria 8 ot raised, ‘Interstria \ae 7 Ras Wide
or wider than 8-9 (Fig, 9). Laterstrial punctures
moderately strong, Southern species -- 46
B. invalutus Macleay
Rowose por tion of metafemur approximately 1/2
length of femur. Panetures on dise of pranotum
ol Ovo sizes, larger predominating. Posrerior
partion of carinate eight) imtersta at right
angles co edge of elytron. Black, except for
interstria in apical 1/4 of elytron...........
litt ty eeeet corer e B Riger spTDy,
Ruwrense purhon of metafentur ‘tenehes 2/3 length
of femur on rear edge, Punetures on diye ol!
pronolum of abe size, posterior portion of
catinate eighth imtersiria of approximately 45°
to edge of elyiron (Fig. 8). Barcus sp.nov.
Pronotum smooth, shiny, moderately covered
With large puncitires often with spaces preater
than 1/2 diameter of puncture or erealer between
punctures. Pronoltum vellow with three
longilidinal black Stripes... peace AD
Head and proratiim rugase, densely. covered with
large punctures, those an provotuny separated
15, (14)
18,
16, (15)
16,
Vy, (14)
7,
8. (17)
REVISION OF BEROSLIS
by narrow nilges, Promotumt Black except far
extreme lateral margins
BR, IMACFOPUNCLALUS sp.Nov.
} Rugose portions of mesofemur a little under 1/2
length of femur. . 8, quadropunctalus sp.m0y,
Ruwose portions af mesofemur Ly 2-2/3 lengthy
CU MANOR ese pe oe B. trishae sp.nov.
Ponctures On prononiny af Iwo distinct sizes.
Areas between elytral striae with shallow
punctures... .8 daplopunctatus Blackburn
Punctures on pronotum vary little in size. Areas
belween elytral striae virtually impunctale ..,
B. queensiandieus Blackburn
Punctures on elytron densely und evenly
distributed, each with a short seta, Punctures in
clyERAl striae ditfieull to distinguish froin (hose
WM WMICPRIe wee Rete ne 15
Punctures on elytra mostly without setae.
Phnetures if clytral ilersttiac seatrered, distines
from those in striae
Punetures on pronatuint expanded faterally
forming a-dense network of closely spaced short
striae orientated across the pronotum (Fig, 5)
; BL niteans W, Macleay
Punciures ¢ on n prongtum MOTMA eee eee 16
Small (< 3.0 nod), elytvon without striae Head
metallic black... . B. pulehellus W. Macleay
Larger | >3.6%m), elytra wilh deeply impressed
striae, Rear of head with dark markings ———.
; B. dallasi sp.nov.
Interstrial purnctu ures towards sides of elyira not
as mimcrous as as those on disc, olfen arranged
TB SIMRO TOW ie ee ee ee eee 18
lorerstrial purtetures towards sides-al elytra mare
numerous and larger (han (hose on disc, never
arranged in single lines... .
~B macumbensis Blackburn
Outer anical spine on elytron tong and (hin
(Fig. 3) Interstrial punetures laterally
subobsolete, arranged in one row aver most of
elyiron. Rugese portion of meiafemur approni-
mately 1/3-2/3 length of femur Pale -----
a Re bean Bo munitipennis Blackburn
Nor with above combination of characters
(Xi8; SV veep aecorresurat etogeo rere teres LS
19. (18)
19,
of pronotum of one size (eg. Fig. 6)...
20, (19) Head black.
20, Head pale brown,
21, (20)
2),
22. (19) Rugose portions of mesc
22.
23. (22)
stiria >22. Rupose portion of
abdominal segment notched .,.
23
VEFONIEE vor yyy ss t fast
24,
”
Punotures in jntersiria 3 of two sizes, with small
size predominwing (Fig. 11), Punetures on dise
of pronotum of two sizes mostly large with some
smaller ones (ug. Fig. 7) 2.) 0-22.) 02... , 22
Punctures ininterstria 3 of one size, orwith only
an occasionally smaller one. Punctures on dise
20
Interstrial punctures on elytron
approximately the same size as those in striae
Pi eb eee clavate mcm B. amoenus sp.nov.
Inierstrial punetures on
elytron smaller than (hose in striae... ,, 2)
Rugose portions of femore dark-brown —....
5 -B. Josephenue sp wov.
Rupose Portion of femare pale-brown...,...
. B. gibhde sp.nov
Rugose portions of meso- and meta-lemore
noticeably darker than rest of femursee Table |.
Rugase portions of meso- and metafemore of
same colour as rest of femur... oc... 23
larger (>3.9 mm). Number of punctures iw
second ely(ral interstria lo level of end of second
mesotemur
approx. 1/3 length of lenitir. Male with apical
iB. nemageulds Blackburn
Sonaller (<6.5 mm). Number of punetures in
second elyiral iniersiria back to level of end of
sccond sifia usually <26. Rugase portion of
inesofemur >1/2 length of femur except in B,
NOC 24
Rugase portian of nievolerwur 2 1/2. Apical
spines on elytra moderately developed (hig. 4)
B veronicae sp.nov.
Rugose purticrn of mesofemur >1/2 lengiti of
femur ate Table 2.
SYSTEMATICS
Berosus discolor Blackburn
(Figs 2, 13, 18)
Berosus discolor Blackburn, !888(1889), p. 829
Berosus flindersi Blackburn, 1888(1889), p. 831, syn.
Noy,
TABLE L. Distinguishing characters for four species of Berasus,
B. aquila sp. nov.
B. australiae Mulsant
B sadiae sp. nov.
; ilo sp. . I s & decipiens Blackburn j . 5
3.3-5,0 mm long
Tips of elytron usually
without spines.
Third segment of male
prolars) approximately
gare length a5 segment,
Elytral punctures weak,
Strial punctures arranged
in one line, over most of
elviron.
Coastal Norihern
Territory,
6.5-9.0 mm long
Tips of elytron with well
developed spines.
Third segment of male
protarsi shorter than
second segment.
Elytral punctures usually
strong, sirial punctures
not artanged in one line
inwards from. siria 7.
Australia-wide.
6,1-8.0 mm long
Tips of elytron with well-
developed spines.
Third segment of male
length of second
segment.
Elytral pufierures usually
weak, Strial punerures
usually arranged in one
line over most of elytron.
Wyndham to Cairns.
§-5.0 mm long
Tips of elytron usually
with well-developed
spines.
Third sewment of male
length of second
segment.
Elytral punetures sually
weak, strial punctiires
usualy arranged th one
line aver most of elytran.
Coastal Northern
Territory.
©. HS, WATTS
TABLE 2. Distinguishing characters for five species oF Heresies,
B, debilipennis
Blaekburn
4.0-6.0 mm long
Apex of elytron
rounded or with
weak spines.
Punctures in Ist
elytral interstria,
scatiered, not
arranged in a row.
Lateral punctation
on elytran
moderale to
STON.
Tip of posiconul
process Ii same
plane as rest of
underside,
10-18 punctures in
second clyrral
Strie.
First segment of
male protarsi much
B. subovatus
Knisch
3.0-3.0 iM lone
Apex of elytron
rounded of weakly
truneated,
Punctures in 1st
elytral interstria,
arranged ina
Single row,
Lateral punctation
on elytron weak.
Tip of posicoxal
process in same
pling as rest of
underside.
6-7 punciures in
second elyiral
suria,
First segment of
male protarsi
B. yiyae sp. nov.
3.0-4.5 mm long
Apex of elytron
with weak spines,
Pufietures in Ist
elytral interstria,
arranged ina
Single row,
Lateral punctation
on elytron
moderate to
strong.
Tip of postvoxul
process in same
Plane as rest of
underside.
8-13 punetures in
second elytral
stra,
First segment of
male prolarsi equal
B. nicholasi
sp. nov.
5.0-6.5 mm long
Apex of élytron
with weak to
moddenale spines.
Punetures in tor
elyinal interstria,
sealtered, not
arranged in a row.
Lateral puneciarion
on elytron small
and Weak.
Tip of postcasal
provess i Same
plunge as rest of
woiderside,
LO<1R punchires in
second elyiral
stria.
First sepment of
male protarsi equal
B, ralphi sp. noy.
3.5-4,.5 mm long
Apex of elyiron
truncated or with
weak spines,
Punetures in Ist
elytral interseria,
seattered, fol
arranged ina
single row,
Lateral punclation
on elytron small
and weak,
Tip of posicoxal
process bent
downwaruds av 45)
to rest of
Underside.
Tel3 punctures in
second ¢lytral
Atria.
First segment of
male provars)
longer than
second.
longer than
second. second,
in lenpth to
longer than
second.
in length to
second.
55
Thpes
Berosus discolor: Holotype, male, Pr Lincoln, §.A.,
in BM(NH), seen.
Berosus_/lindersi; Holotype, male, Pt Lincoln, S.A.,
in BM(NH), seen. Paralype, female, in SAM, seen.
Descriplion (number examined 131)
Length 2.8.8 mm. Oval, elytra moderately
humped, highest behind middle where it is
approximately 1/3 » height alshoulder: Apex of”
ely(ron rounded, in some subavure and suggestive
of a thick broad spine, Head relatively narrow,
black, shiny, metallic, Pronotum black with sides
broadly brown, often with small central brown spot
or band behind front edge, shiny, metallic, Elytron
shiny brown, with striae, punctures and a few spots
black, Ventral surface black, appendages brown,
except for extreme lips Of lapial palps and rugose
porlion of meso- and metafemur which ean range
from light-brown to black. Punctures small, dense.
Punctures on head strong, regular, well-impressed,
most less than 1/2 puncture width apart. Punctures
on pronotum similar but slightly less dense. Elytral
striae well-impressed. Second stria on elytron 1/4
length of elytron, Punctures in and between striae
similar, about same size as on pronotum, strongest
laterally, those on extreme humeral angles and
belween first stria and suture smaller. Mesosternal
keel weakly to moderately well-developed, pro-
jecting backwards. Metacoxal process sharply
produced backwards in niidline, lateral lobes raised,
sides diverging towards lront, small narrowly oval
shiny depression in midline. Keelin midline of first
abdominal segment weak, reaching only |/4 to 1/3
of width. Rugose portion of metafemur approxr
mately 1/3 length of feriiur. Meso- and meta-libia
strongly spined. Apical abdominal seament with
small weak noteh.
Male: Protarsi foursegmented, first seament
moderately expanded,
Remarks
Less. common in collections than B approximans.
Readily separated from those other Berosus with
a well-developed mesosternal keel and short second
elyiral stria by the small area ol rugosity on the
femora.
Distribution (Fig. 79)
Coastal southern Australia from Perth, W.A. ta
Sydney, N.SW., and Tas,
Berosus approximans Fairmaire
(Pigs 10, 12, 17)
Berosus approximans Fairmaire, 1879, p. $2.
Berosus ovipennis Fairmaire, 1879, p82, syn. nev,
wn
REVISION OF BEROSUS
She.
Testa
> BG) ost at %.5
$e lo 8 Gre Fe ess oy,
FIGURES I-11. 1, Midventral region of B. nicholasi; 2, midventral region of B. discolor, 3, apex of elytra of B.
munitipennis; 4, ditto, B. munitipennis; 5, pronotum of B. nutans; 6, pronotum of B gibbae showing punctation
on disc; 7, pronotum of B arcus showing punctation on disc; 8, lateral yiew of elytron of B. arcus; 9, lateral view
of elytron of A involutus; 10, lateral view of B. upproximans showing pronotal pillar (shaded); 11, elytron ot
B. majusculus showing punctation of disc,
Cc. H. S. WATTS
|
KY
18
16
ART
See
al
—
ie)
17
ie
22 3
:
SHE? Ee
27 28 29 30 31
FIGURES 12-31. 12, dorsal view of male genitalia of B. approximans; 13, ditto, B. discolor; 14, ditto, B. juxtadiscolor;
15, ditto, B. reardoni; 16, ditto, B. timmsi; 17, lateral view of male genitalia of B. approximans; 18, ditto,B. discolor:
19, ditto, B. juxtadiscolor; 20, ditto, B. reardoni; 21, ditto, B. timmsi; 22, dorsal view of aedeagus and right paramere
of B. duplopunctatus;, 23, ditto, B. queenslandicus; 24, dorsal view of aedeagus of B. arcus; 25, ditto, B. niger; 26,
ditto, B. involutans;, 27, lateral view of aedeagus of B. duplopunctatus; 28, ditto, B. queenslandicus; 29, ditto B. arcus;
30, ditto, B. niger; 31, ditto, B involutans.
25 26
REVISION OF BEROSUS 7
yea a
38 39 40
=
sega
0
ede
-
ot
46
=
52
Gao
\S
sae
Ca
49 5 51 53
FIGURES 32-53. 32, dorsal view of mate genitaha of B dallasi; 33, ditto, B. gibbae; 34, ditto, B. aquilo; 35, ditto,
B. vijue; 36, ditto, B. subovatus; 37, B. trishae; 38, lateral view of male genitalia of B. dallasae; 39, ditto, B. gibbae;
40, ditto, B. aquilo; 41, ditto, B. vijae; 42, ditto, B. subovatus; 43, ditto, B trishae; 44, dorsal view of male genitalia
of B. debilipennis (thin form); 45, ditto, B. debilipennis (thick form); 46, ditto, B sadieae; 47, ditto, B. decipiens;
48, ditto, B. pulchellus; 49, ventral view of male genitalia of B. debilipennis (thin form); 50, ditto, B. sadieae; 51,
B. debilipennis (thick form); 52, ditto, B. decipiens; 53, ditto, B, pulchellus,
Q
0
C,H. S.
55 \ |
56
54
hy
| | )
60 61 Re
68 69
WATTS
58
57
59
ity
70
FIGURES 54-70. 54, dorsal view of male genitalia of B, majusculus; 55, ditto, B. josephenae; 56, ditto, B. amoenus:
57, ditto, B. nutans; 58, ditto, B. munitipennis; 59, ditto, B. macumbensis; 60, lateral view of male genitalia of B
majusculus; 61, ditto, B. josephenae; 62, ditto, B. amoenus; 63, ditto, B. nutans; 64, ditto, B. munitipennis; 65, ditto,
B, macumbensis; 66, dorsal view of male genitalia of B. nicholasi; 67, lateral view of male genitalia of B. veronicae;
68, dorsal, B. veronicae; 69, lateral, B. nicholasi; 70, lateral, B. macropunctatus.
72
FIGURES 71-74. 71, lateral view of male genitalia of B. australiae; 72, ditto, B. ralphi; 73, dorsal view of male genitalia
71
of B. ralphi; 74, ditto, B, australiae.
74
REVISION OF BEROSUS 9
Berosus stigmaticollis Fairmaire, 1879, p, 82, syn.
nov.
Berosus simulans Blackburn, 1888(1889), p. 832,
syn. nov,
Berosus auriceps Blackburn, 1889(1890), p. 447,
syn. nov.
Berosus blackburni Zaitz, 1908 p. 358, nom. nov.
for B. auriceps.
Types
B. approximans. Holotype, male, Peak Downs, N-T.
in MNHN, seen.
B. stigmaticollis, Holotype, female, Peak Downs,
N-T. in MNHN, seen.
B. ovipennis, Holotype, female, Pt Mackay, in
MNHN, seen.
B. auriceps. Holotype, male, Northern Territory, in
BM(NH), seen.
B. simulans. Holotype, male, Rivoli Bay, S.A, in
SAM, seen,
Description (number examined 134)
Length 3,0-5.2 mm. Narrowly oval, elytra
moderately humped, higher behind middle where
it is 1/3 higher than at humeral angle. Apex of
elytron rounded or produced into a small, widely
triangular spine. Head black, shiny, metallic.
Pronotum brown with broad central panel darker,
dark portion usually not reaching front or rear
edges and often with lighter area in midline, shiny.
Elytron brown, with striae, punctures and a few
spots, black, shiny. Ventral surface black, append-
ages lighter. Rugose portions of femora variable
from light to dark-brown. Punctures on head
strong, regular, well-impressed, most less than 1/2
puncture width apart. Punctures on pronotum simi-
larly sized but less dense. Weaker on disc. Elytral
striae well-impressed. Second stria on elytron 1/4
length of elytron. Punctures in striae and between
them approximately the same size, as strong laterally
as on disc. Reticulation of elytron variable, virtually
absent in some, in others very strong and almost
masking punctures, especially laterally. Ventral
surface densely rugose punctate. Mesosternal keel
strongly produced downwards forming a pillar.
Front edge almost perpendicular to body, ventral
edge straight, or concave in some specimens, with
the front portion projecting downwards as much
or further than rear portion. Lateral lobes of
metacoxa raised, bluntly pointed, midline produced
backwards in a point, small oval depression in
midline devoid of sculpture. Keel in midline of first
abdominal segment well-marked, reaching 1/4-3/4
segment. Rugose portion of metafemur 1/2-2/3
length of femur. Meso- and meta-tibiae strongly
spined. Apical abdominal segment weakly and
widely notched.
Male: Protarsi four-segmented, first segment
moderately expanded. Head narrower than in
female,
Remarks
A southern species slightly larger than B. /lindersi.
Readily separated from that species by the larger
area of rugose sculpture on the femora. The notch
on the last abdominal segment is also broader and
weaker in B&B. approximans. Differs from
B. juxtadiscolor and B. reardoni from northern
Australia by having a smaller amount of rugose
sculpture on the femora than in these species, in
the generally strongly elytral punctures and in
characters of the aedeagus. Its wide distribution,
variability in size and in the strength of the dorsal
punctation has led to it being described a number
of times by earlier authors.
Distribution (Fig. 76)
Southern coastal Australia from Kimberley, W.A.,
to Rockhampton, Qld.
Berosus reardoni sp. nov.
(Figs 15, 20)
Description (number examined 7)
Length 3,2-4.5 mm. Narrowly oval, elytra moder-
ately humpbacked, highest behind middle where it
is 1.4 x height at shoulders. Apex of elytron bluntly
pointed. Head shiny, black. Pronotum yellow-brown
with two longitudinal black strips adjacent to
midline, dark portions not reaching front or hind
margin. Elytron yellow-brown with striae and many
punctures on head strong, regular, well-impressed,
most less than 1/2 a puncture-width apart. Punc-
tures on pronotum moderately dense and strong
laterally, weaker and sparser on disc. Dorsal surface
weakly to moderately reticulate. Elytral striae well-
impressed, less so on disc, second stria on elytron
a little less than 1/4 length of elytron. Punctures
in striae somewhat larger than those in interstriae,
shallower and slightly larger laterally. Mesosternal
pillar long, well-developed, ventral edge concave.
Ventral surface densely rugose-punctate. Metacoxal
process broad, lateral lobes lowered slightly,
diverging towards rear, deeply and broadly pitted
in centre, weakly triangularly produced backward
in midline. Apical abdominal segment notched.
Rugose portion of metafemur 2/3 length of femur,
that of mesofemur between 1/2-2/3, that of
profemur about 1/3 length of femur on hind edge.
Male: Protarsi four segmented. First segment
moderately expanded.
Remarks
This northern member of the approximans species
group closely resembles B, juxtadiscolor, but differs
in being lighter in colour, particularly the rugose
portions of the femora, and in the shape of the
aedeagus and parameres.
Distribution (Fig, 75)
Northern N.T.
My tC HOS WATTS
pes
Halatype male ‘Katherine, NT, at light 9.11.68
-L oA. L. Warson’, in ANIC.
Berosus jovtrdiscolar sp. noy.
(Pigs I4, 19)
Description (umber exited 18)
Length 3,5-5,0 mm. Narrowly oval. Elytra weakly
humpbacked, bighesr behind middle where itis 1.2
» fleizht at shoulder, Apex of elytron rounded
Head shiny black. Pronotuin dark yellow-brown
will two central longitudinal stripes darker. Elyiron
darkish yellow-brown with punctures and strine
darker, in thany individuals elytron almost com-
pletely dark. Ventral Surface dark-brown, append
ages other than pugose portion of femurs somewhat
lighter. Punctures on head strong, revue and well-
impressed, most less than 142 au puneture-width
apatt. Punecures on pronoeruot moderately dense
and strony laterally, weaker and sparser on disc,
Dorsal surtace Weakly to moderately reticulare,
Elytral strive well-impressed, less so on dise. Second
siria on elyfron a little less (han a 1/4 length of
clyiron. Punerures in strive sliglitly larger thar chose
in interstriaé Punetures stronger towards sides.
Interstriae not arranged ina single row, Mesosternal
pillar long, well-developed, veritral cdee slopes
downwards towards rear, sharply pointed on ventral
rear ungle. ventral surface densely but shallowly
rugose-punctate. Metacoxal process broad, square
ish), lateral lobes weakly lowered, deeply and broadly
pitted jn cenrre, triangularly produced backwards
in midline. Midtine keel in first abdominal seument
niodenuely developed in frant hall. Apical
abdominal segment ootched. Rugose partion of
metafemtur 2/3 length of fenur, that of mesofermur
a little over |) 2, thar of profemur about 1/3 on hind
ede,
Male: Protarst foar-seemented, First sexnmient
noderacely expanded. second seamient small chine
segment long aid thin abour lengrh of firse and
second semen,
Remurks
Chis species closely resembles B reardoni but has
a more vomiplexly shaped paramere and dillerently
Shaped aedeagus. It seems to be a darker species
than B reardant which has the rugase portions of
the femora the same colour as rest of leg, Only a
lew speeiniens of & reardoriare known, A clearer
idea of the relationships between these two species
must awail (he colleciion of further dyalecial. The
preater extent of rugosily an the legs separates i
from the otherwise quite similar 2 disealon
Types
Holotype, mile, “LL 09'S 132-00, Black Point,
Cobourg, N.T., 15-23 beh, 1977. 4. A. Weir’, in
ANIC, Paratynes; six, same data as holotype, in
ANIC: ong, same data as trolorype, in CW
two.'Howard Sprites, WT, light 27.68, JO8. 1,
Watson’, in ANIC,
Distribution (Fig, 77), At present Known only from
the coastal areas of the NVD,
Berasns timunsi sp. nov.
(Figs 16, 21)
Description (dumber exanined seven)
Length 3.4-4.0 nim. Oval, elytron weakly humped,
highest behind middle where it is approximately 1.2
» heht of shoulder, Apex af elytvon founded er
weakly trunedred. Head black, shiny, metallic
Pronotun yellowish with central panel dark-brown,
shiny, metllic. Elytron shiny yellow-brown, with
sina, punetures and a few spots black, Ventral
surface dark-brown to blak, appendages brown
except for rugose portions of meso and metafemurs
which are darker, Purciures on ventral surface mod
vrately large, dense. Punctures an head strong.
regular, well-iinpressed, mos at rear af head less
than 1/2 puncturewidth apart, Punetures on pre-
noruni strong, variable in size particularly lateralls
less dense (han on head, well-impressed. Elytral
striae well-impressed, with strong tendeney tor the
inner edve of striae to he sharper than outer, Second
strig an elytron 1/4 lenath of elytron, Punciuresin
étriae strong and little larger (han largest on pro-
nouuim, nlerstrial punctures small, subobsolete over
most of elytron, Mesosternal keel moderacely
developed, pillar well-developed, concave and dent:
volate ou veutral cdee front and rear projections
largest. Metacoxal process projecting backwards in
midline, lateral lobes weakly raised, pointed behinu,
weukly converging Lowards front, a wide shiny dig-
mond shaped depression in midline, Keel in nividline
of first abdominal segment moderately strone
reaching weross mos! of seament. Rugose portian
of metalemut approximately 2/3 length of femur,
that an mesofemur approximately 1/2, that on pro-
femur 1/2-173 length of femur Meso- and
metatibiae strongly spined. Apical abdominal see-
nent With song deep wide noteh lo about 1/2
distance of tiormally visible partion of segment.
Mule; Protwarsi foursegmenred, first seunent
stromgly expanded. Merafemur with distinet trie
angular bulge in lower Wind margin midway alone
rugoase poriian.
female. Metafemur of sme speciniens show
slight thickening in same arca as ip nyale.
Remarks
This species resembles the nrere widespread
B Juntadisculor with whieh i t& sympatric, but
differs from it (and all other Berasus) by the shape
REVISION OF BEROSUS MN
of the mule metafemut which is reminiseent of the
femora of some male Dyriscids of the genus
Antfiporus. The variably-sized pronotal punctures
and very small interstrial punctures also separate
it from B fuxtadiscolor and B. reardani,
Disrribution (Pig. 82)
Known only from the wpe locality at the tip of
Cape York, Old,
Tepes
Holotype, male, 'Bamayza, Old, 29/6/83, Timms’,
i SAM. Seven pararypes, same data as holotype;
one male. one female, in ANIC; one male, three
females in CW.
Berusus niger sp. nov.
(Figs 25, 30)
Description (number examined seven)
Length 3.3-4.0 mm. Oval. Elyira parallel-sided,
humpbacked, hivhest just behind tbe middle where
iris 18 ~ the height at che shoulders. Apex of
elytron nol spined. Black, portions of elytral inter-
siviae in apical hall and appendages of head, dark
yellow-brown. Head and pronotum with large,
dense, moderately-impressed punctures. Promoturm
with some scattered much smaller punctures in
spaces between larger punctures. Strial punctures
large, shallow. luerstilal punetures weak, ill
defined, lavking lalerally and apically, Second striae
approxinuuely 1/2 length of clyiron. Spaces be-
tween elytral striae 8 and Y, 9 and 10, and 10 and
11 greater in the middle of elytron than elsewhere.
Elytral striae 1, 3, 4, 5, 6, 7, 8 and Il ridged,
pronouncedly so an stra &. The stronely-ridged
stria 8 bends sharply towards apex where if is abOUL
90° to the edve of elyrron when viewed lalerally,
Ventral surface rugose-punctate. Mesosteroal pillar
long, ventral edge weakly concave, anrerior ventral
wogle wilh very small spine, Melacoxal provess
broad, squarish, with large pit in nidling; projecting
baekwatds sharply in midline, continued torwards
in midline by well-marked ridge, Midline keel on
first abdominal sezment sirongly raised for whole
length of seyment. Apical abdominal segment
moderately natvhed, lateral Manges moderately
developed. Rugose portion of metafemur reaching
1/2 way on posterior edge and a little less on
anterior edge, rugose portion of mesofemur about
13 length of lemut, that on profernur very small.
Male: Protarsi four-seemented. Basal segments
not expanded, first seement somewhat longer than
seeoud and third which are subequal in size.
Reatarks
A north-eastern highlands species. Readily sep-
araled [rom similar species by the almost completely
black colour and the fori of the lateral striae on
the elytron.
Distribution (Fig. 7S)
Nountalnous arcas on east coast north of Towns-
ville, Old,
Types
Halotype, male, ‘Star Valley loaokoul & 3 km W of
Paluma Qld, 3.ii.67, at light, LG. Brooks’, in
ANIC. Paralypes: one, ‘13.6M up Whitfield Ra, Rd,
Cairns, 3/11/70, J. G, Brooks, in ANIC, one,
‘Tinaroo Uk., NO. 1/73 AW.H.' in ANIC; one ‘MI,
Spey, N.Q, 11/73 G.B.. in ANIC, one, “7.8M. NW
Paluma, NO 5.11.73 J, G, Brooks, in ANLC; two,
‘Ausiralia N.Qid. Windsor Tableland via Mt,
Carbine 11-12 Jon 1980 R. |. Store’, in QOPI.
Berosus favolatus (W. Macleay)
(Figs 9, 26, 31)
Hyzrorrophus involutus W. MacLeay, 1873 p. 132.
Types
There are four specimens [rom Gayndah, Old in
ANIC (on loan from the Macleay Museum), label-
led ‘Syntypes’, Two of these are B palchellus and
(wo B approximans. The original description does
not indicate how many specimens MacLeay had be-
fore him. Apart Irom the loealily there is little to
connect these insects with a type series of
RB favo/itus. Two specimens labelled *Hygrotrophus
involutus M, L. W, Gayndah’, mounted on the same
card, are in Lhe AM labelled ‘Holotype’ and num-
bered K1953). They agree well with the bret des~
cription and belong to the species that subsequent
workers such as Blackburn and Lea have identified
as B involutus | therefore herein designate the lett-
hand specimen as the leclotype of Hygratrophus
invalulus W. Macleay, 1873.
Description (number examined 475)
Lengih 3.5-4.5 mim. Oval, elytra humpbacked,
highes! just behind middle where about 1.5 +
height! al shoulder. Apes of elytron pounced. Head
and most of pronotum black, with metallic sheen.
Front and side margins of pronotum yellow-brown,
ovcasionally lacking, elytron brown, punctures and
striae and variable number of markings black.
Ventral surface black. Appendages light-brown, tip
of labial palpi and rugose portions of meso- and
metafemora black, Head and pronorum with derise
lane evenly-sized nearly confluent pilpetures,
Interstrial punctures on elytton Jaree and well-
marked, almost touching, those lowards sides and
humeral angles larzer and squarer. Interstrial
puncture sinall, shallow, strongest on disc, obsolete
laterally. Second sina approximately half length of
I2 CPL S. WATTS
elyiron. Distance between striae seven and eight and
eight and nine often slightly ereater in middle, other
singe approximately equidistant! fram each other.
Ventral surface rugose-punctate. Mesosternal keel
in form Of a long pillar, with a small downward
point at tront edge, Metacoxal process broad, sides
raised, Subparallel, deeply and broadly pilted in
middle, pit devoid af sculpture, projecting backward
sharply for short distance in midline, lobed. Midline
keel of first. abdominal segment stronyly raised for
whole width of segment, Apical abdontirial sexment
broadly notched with well-developed lateral Nanges,
a pair of very Small tubercules on edge of middle
of notch. Rugose portion of metafemur dark,
reaching 1/2-2/3 along femur on anterior edge, not
as far on posterjor edge
Male; Prolarsi four-se¢emented. Seemenis nol
expanded, basal ttirge subequal in size.
Remarks
A common species in clear sireams and pools in
the Great Dividing Range from Vie, to north Qld.
Many specimens from the higher-areas of Vic, and
N.S.W. are darker in colour and lack the distinctive
yellow margios on the pronotum. Separated from
the two related Species, B niger and Berens, by
(he lack of strongly raised and bowed seventh elytral
strid, tis also noticeably larger than B ares and
is different in colour trom & niger. [tis only in the
northern partion of ils range that B invelufus is
sympatric with these two similar species.
Distribution (Fiz, 76)
Coastal eastern Australia (fom southern Gane York
to south-eastern Vie.
Rervsus areus sp, nov,
(Figs 8, 24, 29)
Descriptian (number examined 149)
Length L-§-3,4 mm. Oval, elyira humpbacked,
highest just behind middle where about 1.2 »
heighr at shoulder. Apex of elytron without spines,
Black, elytral interstriae, edges af pronotum apart
from central portion of rear margin, apperidages
apart from tip of labial palpi ajd rugose portions
of meso and metalemora yellow-brown. hlead and
pronotum with dense strong evenly-sized und spaeed
punctures. Strial punetures on elyiron large, alnrose
touching. Literstrial punetures large bur very
shallow, stronger on dise than laterally. Second stria
on elytron approximately 1/2 length of elytron.
Distance between striae eight and nine greatly
enlareed in middle to about twice distance between
slmae nine and ten, which is also slightly enlarged
in middle. Stria eight moderately carinate in middle
section where distance from stria nine greatest.
Ventral surface rugose-punclata Mesosternal keel
in wide pillar, anterior ventral angle of pillar sharply
produced downward inte small sharp point. Mera-
coxal process broad, squarish, sides raised, sub-
parallel, deeply and widely pitted in midline,
projecling backwards sharply in midline, Midline
keel on first abdominal s¢ement strongly raised for
whole length ol segment, Apical abdominal seg-
ment broadly notched wirh well-developed lareral
flanges. Rugeose portion of metalemur reaching
nearly 2/3 along posterior edge of femur and 1/3
On anterior edge, thal on mesofemur reaching a
little over 1/2 length of femur an pasterior edge,
1/3 on anterior edge, that ou profemur 1/3 on
posterior edge and less than. 1/4 on anterior edge,
Mole: Protars| fourseamented, basal Uiree
segments subequal afi size.
Reniarks
A northern species confined to the sireanis and
niverside pools of the ranges around wand to the
north of the Atherton Tableland, Smaller than
B involutus and & niger and readily separated trom
them by colour arid the form oF the eigtith elytral
stra.
Distribution (Fig, 79)
Ranges of soucher Cape York and Atherton Table-
land, Qld,
Tipes
Holotype, male, ‘S mi. N. Bloamfield Rn., N,O, 7-9.
May 1970S, K, Curtis’, in ANIC. Paratypes, 11
same data as Holotype, in ANIC; one same data.
ii CW.
Berosus trishae sp. nov,
(Figs 37, 43)
Hescription (umber examined 39)
Length 2.4-4.5 mm, Oval, Elytra parallel-sided,
humpbacked, highest just behind middle where it
is 1.20.2 higher than at shoulders. Apex of elytron
rounded. Yellow-brown, head, three pough broad
longitudinal patches on pronotum, about a dozen
small patehes on elytron and vague areas of ventral
surface darker. Head and pronotim moderately
covered with large strong punctures, A single even
row of slightly smaller punccures along lront and
rear edges of pronotum. Strial punctures on elytron
very large, almost confluent, stranpest laterally
where they are ulmost square, Interstrial areas shiny
lacking punctures except apically in interstria 3.
Second elytral stria a litthe more than 1/2 length
of clytron, Interstrial areas raised somewhat lowards
apex of elV¥tron. Ventral surface moderately
punctate, Mesosternal keel produced in robust pillar,
ventral edge of pillar convex. Metacoxal process very
broad, parallel sided, central portion with wide pit,
lateral lobes project downwards at about 35°,
narrowly and triangulacly produced backwards in
REVISION OF BEROSUS i
inidliie. Midline keel on lirst abdominal segorernt
strongly raised for whole length of segment. Apical
abdominal segment widely and deeply notched with
well-developed lateral flanges, Rugase porion of
metafemur abou! 2/3 length of femur thal on
mesofemur |/2-2/3, and that on profemur aboul
1/2 length of femur,
Male: Protarsi lour-segmented, basa] segments
Hot expanded
Remarks
A northern Species, Separated (rom the quite simular
& quadrapuncratys by ie ereater extent of rugose
sculpture on the temura and characters of the
aedeagus. The sculpture on the head is a Tit(le
rougher than in & quadrupunciatus and the
punctures on pronotum and élytra are smaller. Al-
thouwh this size difference is clearly evident in direct
comparisons of most specimens, | have been Unable
16 quantil’y it to enable the character to be used to
separate the species. Most specimens are < 3,3 mm
long bul a female from Charters Towers, Qld, in
NSWD, is considerably larger at 4.5 mm, f ean
derect no other difference from N-T. specimens.
Distribution (Fig, 80)
Darwin area, ST, Atherton
Charlers Towers Qld.
Tableland and,
Tppes
Holotype, fenvile, ‘NT. Lake Bennett area c. 25 km
SE. of Manton Dam. 29-30 Dee, 1979, M, B.
Malipatil’, in NTM, Paratypes; 20, ‘N.T., Ck. in
Flolmes Jungle 131.1980 M, Malapatil’, in NTM,
iwo same data in CW
Berosus macropunctaius sp. Wav.
(Fig. 70)
Description (wmumber examined twa)
Length 3.0-3.6 mm. Oval. Sides of elytra
subparallel, humpbacked, highest about 1/3 dis-
lance front apex of elytra where it is abaut 1.2
height at shoulders. Apex of clytron squarish.
Yellow-brown, head and propotum black except for
narrow yellowish band om lateral and hind edges,
elytron wilh scattered darker patches, Head and
pronotum strongly rugose-punckale, Stelal pune-
(iires on clytron large, larger and squartsh towards
sides, Interstrial areas shiny, impunetate exeept for
a row of very small setae-beariny, punctures towards
apex of third interstria. Lateral three interstrial areas
raised into weak ridges, fnterstrial areas at extreme
apex weakly raised, Ventral surface stromaly rugose-
punctace. Mesosternal keel produced into a narrow
pillir, Ventral edge of pillar slopes downwards
rowards rear and has a small downward pointing
spine an ventral anterior angle, Metadoxal pravess
very broad, lateral lobes parallel sided, rather
narrow, separated by wide pit in centre ol process,
open towards rear, lateral lobes slope downwards
at about 20%, broadly triangularly produced
backwards in midline. Midline keel of first abdomi-
val segmeni strongly developed over Whole length
of segment, veotal edge serrale, apical abdominal
segment deeply and widely norehed with well:
developed lateral flanges. Rugose portion of
metufemur 3/4 length of femur, (hat of mesolemut
a little less than 3/4, thal on profemur 1/2 length
of femur.
Male: Protarsi four-segmented, basal segments
not expanded,
Remarks
A distinetive species readily separated from &
rishae and B quadrapuncrarus by the strongly
rugose-punctate black head and pranolum.
Distribution (Fig. 76) (N-T. (ype localities only,)
Types
Holotype, inale, ‘Burrell’s Ck Stuart H’way NT, 24
Nov, 1972 D.H. Colless'!, in AIDC. Pararype, one,
13°LS'S, 131°06'E, Adelaide River, NT, b6,x-72,
M. 5. Upton’, in ANIC.
Berosus quadrapunctatus sp. nov.
Deseriplion (number examined 4)
Lenerh 3.2-3.9 mm. Elytra humpbacked, highest
just behind niddle where it is 1.3 height at
shoulders, Apex of elytron rounded, weakly
flanged. Yellow-brown With head black and three
broad vague longitudinal patches on pronetum,
scailered patches on ¢elytron and patches in the
midline ventrally, darker, shiny. Head and pronotum
moderately covered with large well-impressed
punctures. Elytron with strial punctures very strong,
particularly laterally Where they are square
Punerures in adjacent laleral striae alnyost (Ouch.
Interstrial areas shiny, lacking punctures excepl for
some miuute setae-bearing ones Lowards apex if
intersiria three, Second clytral stria reaches a little
over 1/2 length of elytron, luterstrial areas towards
extreme apex become strongly raised, Ventral
surtace covered With numerous large bul separate
rugose punctures. Mesosternal keel produced inti
a narrow’ pillar, concave on ventral edge: Metacoxal
process very broad, parallel-sided with large pit in
midline, narrowly triangular, projecting backwards
in midline, lateral lobes slope downwards at about
45" Keel on first abdominal segment strongly raised
for Whole leneth of segment, serrate on ventral edpe
Apical abdominal segment broadly notched, with
14 Cry Ss;
well-developed lateral flanges. Rugose portion of
metafemur a little over 1/2 length of femur, that
on mesofemur a little under 1/2, that on profemur
1/3 length of femur.
Male: Protarsi four-segmented, basal segments
not expanded.
Remarks
A little known species differing from B. trishae by
the less well-developed rugose sculpture on the
femora and in the more uniform sculpture on head.
A female specimen from Charters Towers, Qld.,(in
NSWDA) may belong to this species. It is longer
(4.5 mm), the rugose portion on the femora is a little
greater, the sculpture on the head is rougher with
some scattered very small punctures as in B&, frishae.
In these characters it is intermediate between B
trishae and B. quadrapunctatus.
Distribution (Fig. 80)
Known only from the type series from McArthur
River, N.T,
Types
Holotype, male, ‘McArthur River, 16°47'S
135°45'E, 14 km S by W of Cape Crawford, NT.,
25 Oct. 1975 M. S. Upton, in ANIC. Paratypes,
three, same data as holotype, in ANIC.
Berosus queenslandicus Blackburn
(Figs 23, 28)
Berosus queenslandicus Blackburn, 1898, p. 223,
Berosus quartinus d’Orchymont, 1943, p. 6, syn,
nov.
Types
B. queenslandicus, holotype female, Qld., in
BM(NH), seen. Paratype, female damaged, in SAM,
seen. B. quartinus, holotype, male, and paratype,
in MNHN.
Description (number examined 42)
Length 4.2-6.3 mm. Oval, elytra humpbacked,
approximately twice as high just behind middle as
at shoulder. Apex of elytron rounded. Head black
with metallic sheen. Pronotum shiny, widely black
or dark-brown in midline, brown at sides. Elytron
shiny, brown with dark-brown to black mottlings.
Ventral surface mottled dark-brown and black,
appendages brown, apical segment of labial palpi
darker, rugose portions of meso- and meta femora
dark-brown to black. Punctures on head large,
subequal except for clypeal margin where they are
smaller, those on disc separated by about 1/2 their
width, a few scattered very small punctures in some
specimens. Head with small midline ridge on back,
sometime lacking. Punctures on pronotum as on
head, closer together laterally. Punctures in elytral
WATTS
striae well-marked almost confluent in any one
stria, stronger towards front. Interstrial areas vir-
tually impunctate except for small area of large
punctures on humeral angles. In some specimens
some very shallow small punctures can be seen in
some lights in interstriae on disc. Striae weakly
impressed on disc, stronger laterally and apically.
Second stria on elytron reaching 2/3 length of
elytron. Striae approximately equidistant from each
other. Ventral surface rugose-punctate. Mesosternal
keel in form of high narrow pillar, ventral edge con-
cave, keel on midline of mesosternum forward of
pillar weak. Midline keel on first abdominal seg-
ment well-marked, reaching almost completely
across segment but weakening rapidly towards rear.
Metacoxal process broad with well-marked lateral
lobes, sides subparallel, with depressed shiny im-
punctate area in midline. Rugose portion of meta-
femur about 1/3 length of femur, ending in
relatively straight edge across most of femur, that
on metafemur 1/3, that on profemur 1/3 length of
femur on hind edge. Apical abdominal segment
deeply notched in midline with lateral flanges to
notch.
Male: Protarsi four-segmented, basal segments
subequal not expanded.
Remarks
Separated from the superficially similar B.
duplopunctatus by the slightly larger and differently
shaped rugose portions on femora, the virtual lack
of smaller punctures on head and pronotum and
the virtual lack of interstrial punctures. Some
specimens have some minute punctures in between
the large ones on the head and to a lesser extent
on the pronotum, but these are much smaller and
fewer than those in B. duplopunctatus. Synonomy
based on d’Orchmont’s description and illustration
of male genitalia.
Distribution (Fig. 78)
Coastal south-eastern Australia from Caloundra,
Qld., to Adelaide, S.A.
Berosus duplopunctatus Blackburn
(Figs 22, 27)
Berosus duplopunctatus Blackburn, 1888 (1889), p.
828.
Types
Holotype, female, Pt Lincoln, S.A. in BM(NH),
seen, Paratype, female, Pt Lincoln, S.A. in SAM,
seen,
Description (number examined 109)
Length 4.5-6,4 mm. Oval, elytra humpbacked, at
highest point just behind middle about 1.5 x height
REVISION OF BEROSUS 1s
at shoulder, Apex of elyiron rounded. bead
relatively wide, black, shiny with metalhe sheen.
Midline of pronotim black, sides brown. Elytron
predominantly brown with punctures outlined in
black plus some black spots, extent of black varies
between specimens and is the daminani colour in
many. Ventral surface black, appendages light-
brown, lip of apical segment of labial palpi black,
pupose portions of meso- and metafemora black,
Head densely punetare with both large and small
purictures. Pronotum with large and small
punctures, less dense than on head. Punetures in
elytral striae well-impressed, almost confluent,
stronger towards front, about as sirong on cise as
laterally, Interstrial punctures shallow, small,
numerous, scattered, very weak taterally. Stnae
moderately impressed, slightly weaker on dise
Striac approximately equidistant apart. Second stria
onelyvon reaches to 3/4 length of elytron. Hurneral
angle of elytron serrate due to large punctures close
to cdee. Ventral surfaee strongly ruose-punetate,
Mesosternal keel in form of high narrow pillar,
weakly concave on ventral cdae, keel in front of
pillar weak. Midline keel on first) abdominal
segment well-marked in front half, weaker im hind
half of segment. Apical abdominal segment deeply
notched, notch with lateral Tlanges. Metacoxal
process with well marked Jateral lobes, converging
towards rear and wilh small shiny inipurectate
depressed urea in midline, Rugose portion of
mesolemur about 1/3 length of femur on hind edge,
ubout L/4 Jenerh on front edge of lemur, that on
hind edge of metalemur 1/4, thaton profenmtue b/4
lenerh af femur.
Male: Protarsi foursepemented, basal three
subequal not expanded.
Remarks
Reudily separated from the superticially similar B
queeusiaidious by the puncrace elytral intersinae
ald the presence of Small punerures on the
pronoun,
Distribucian (Fix, BO)
Coustal easrern Australis from Sydney, N.S.W. (0
Kangaroo Island, SA Tas.
Berosus nutans (W. Macleay)
(bigs 5, 37, 63)
Hyerotrophus mitans W. Macl vay, 1873, p. 132.
Berosus pallidulus Faatoaire, (379, pos, syn, aller
Blackburn, 1898, p, 222,
Types
Two synivpes fabelled “Gayadah’ and mounted on
the sane card are in the ANIC, The leti-hand side
specimen, a female, is here designated as leerory pe.
Descriplion (qumber examined 440)
Length 4.2-8.8 mm, Elongate oval, hot
humpbacked, Apex of elytron rounded or obliquely
truncated, Dorsal surface brown with vague darker
markings particularly on head aud pronotum.
Elytron with steiae darker, Ventral surface dark
brown bo black, appendages brown to dark-brown
except for darker tip of Tabial palpi. Apex of
abdomen occasionally brown, Punelures on head
well-marked, weaker tawards front, transversely
elongate. Sculpture on pronotum consists of a close
nelwork of Vine lransyerse wrooves which cover most
of the surtave, all but obscuring, punctures which
themselves tend to be transversely clongate as an
head, A fine reticulation ts also present on some
specimens. Scutellum seulptyred as pronotum,
Elytron with siriae lacking or only Weakly im-
pressed, rheir position allen oullined by rows of
small crapsversely expanded. darkly pigmented
spors. Elytrot densely and evenly covered with short
rather stout setac, each seta arising from a small
shallow transverse groove. Ventral surface shallowly
rugose-punclale with small setae arising trom each
puncture. Mesosternum with weak keel in midline,
extended quite strongly buckwards between imeso-
voxae. Midline of first abdominal segment with
moderale keel in front 1/4. Metadcoxal process
raised, sharply triangular with elongate oval area
in midline devoid of sculpture, Rugese portion of
metafemur 3/74 length of femur, that of mesofemur
2/3, that of profemur 1/2 length of femur,
Male: Provarsi toursegmented, basal segment
expanded, aboul as long as second and third
segment combined, scvond segment expanded, same
length as simple third segment.
Reniurks
A common and widespread inland species, readily
recognised by the unique sculpture of the pronotum.
Distrihutian (Fig. 78)
Coastal and southern inland Australia.
Berosus pulehellis WS. Nacleay
(Figs 48, 33)
Beresus pulchellus W. 5. Macleay, 1525, p. 35.
Fiyeratraphus devisi Blackburn, L898, p, 225, svi
alter d'Orehymont 1943, p. 421.
Types
B. pulchellus, Java, lovation unknown, Bo deviat
fiolotype, ? female, Old., in BM(NH), seen,
Description (number examined 233)
Length 2,8-4.8 mii. Elongate oval, elytra tot
humpbacked. Apex of elyiron founded, Head
16 C. H.S.
black, shiny. Pronotum light-brown with front edge
narrowly black and usually with wide central panel
black, black area not reaching rear border and often
with portion of midline narrowly light-brown.
Elytral striae often darker. Ventral surface black,
appendages light-brown except extreme apex of
labial palpi which is darker. Head evenly punctate
with moderately dense small punctures, punctures
somewhat weaker towards front. Pronotum moder-
ately and evenly covered with small punctures.
Elytron strongly and evenly covered with similarly
sized punctures, each puncture with a well-marked
seta. Elytral striae lacking or very weakly impressed.
Punctures in elytral striae same size as interstrial
punctures and difficult to distinguish from them,
Ventral surface evenly, shallowly rugose-punctate.
Midline of prosternum with weak but well-marked
keel, weakly projecting backwards between meso-
coxae. Midline of first abdominal segment weakly
raised in front half, Metacoxal process bluntly
triangular, raised, midline with narrow oval area
devoid of punctures. Apical abdominal segment
deeply notched. Rugose portions of meta and meso-
femora 3/4 length of femura, that on profemur
2/3-3/4 length of femur.
Male: Protarsi four-segmented, basal two
segments moderately expanded, basal segment
about twice length of second which is about same
length as third.
Remarks
This widespread and common species is readily
separated from all other Australian Berosus by the
small size, black head and virtual lack of elytral
striae. It is most commonly collected from the sandy
pools formed during the dry season in the beds of
the major northern and eastern river systems. It has
recently been collected in numbers in sewerage
settlement pits at Whyalla, S,A., an unusual habitat
and a long way from its normal range.
Distribution (Fig, 82)
Tropical Australia; Whyalla, S.A.; Asia.
Berosus dallasae sp. nov.
(Figs 32, 38, 70)
Description (number examined 80)
Length 3.8-6.8 mm. Elongate oval. Elytra not
humpbacked. Head and pronotum shiny, elytron
rugosely reticulate. Apex of elytron produced into
two spines, outer one usually strongly and widely
developed, inner often small or even vestigal. Dorsal
surface light yellow-brown, rough patches on head,
pronotum and elytron darker, elytron stria darker.
Ventral surface dark-brown, appendages lighter, tip
of lapial palpi dark, Head evenly and strongly
punctate, punctures much larger than eye facet,
much weaker towards front, Pronotum evenly
WATTS
punctate with large strong regular punctures except
for some smaller areas along front and rear margin.
Punctures on disc have a tendency to become longi-
tudinally elongate. Elytral striae strongly impressed.
Stria 2 with punctures a little smaller than those
on pronotum and about twice size of those in
adjacent interstriae. Interstrial punctures evenly,
densely but shallowly impressed over whole elytron,
each puncture bearing a small seta. Ventral surface
rugose-punctate. Midline of mesosternum weakly
and broadly carinate, moderately projected back-
wards. Midline of first abdominal segment weakly
carinate in front 1/4. Metacoxal process broadly
triangular, midline weakly carinate in front of a
small diamond-shaped area in centre, devoid of
sculpture and bounded on front sides by weak
edges. Rugose portion of metafemur 3/4 length of
femur, that on mesofemur 2/3-3/4 that on
profemur 1/2-2/3 length of femur.
Male: Protarsi four-segmented. Basal two seg-
ments moderately expanded, first segment about
1.5 x length of second which is a little larger than
the narrow third segment,
Remarks
This widespread species of northern Australia is
readily separated from the two other, equally wide-
spread, species with setose elytral punctures, by the
lack of strongly transversely elongate pronotal
punctures (from & nutans), and lack of a black
head (from B. pulchellus).
Distribution (Fig. 75)
Tropical Australia.
Types
Holotype, male, ‘| km N. of Millstream, W.A,.
(21°35'S 177°04'E), 9-10.iv.1975, M. S. Upton’, in
ANIC. Paratypes, nine same data as holotype, in
ANIC; one same data as holotype, in CW.
Berosus macumbensis Blackburn
(Figs 59, 65)
Berosus macumbensis Blackburn, 1896, p. 259.
Types
Holotype, male, Macumba Creek, S.A. in BM(NH),
seen.
Description (number examined 150)
Length 5.6-7.8 mm. Elongate oval, not
humpbacked. Apex of elytron with two spines, the
inner usually small and the outer as long as the
distance between the two spines. Elytra shiny.
Dorsal surface light brown to brown with dark mot-
lings. Elytral striae and punctures on disc darker.
Ventral surface dark-brown to black. Appendages
brown except for extreme tip of labial palpi which
REVISION OF BEROSUS 17
7 7
@ B. majusculus e B. involutus
a B. vijae © B. macropunctatus
A B. dallasae a B. josephenae
oO B. niger Oo B. approximans
4 B. reardoni
2 2
77 W 78 W
e 8B. australiae e B. nutans
a B. juxtadiscolor a B. gibbae
gs B. ralphi A B. decipiens
B. queenslandicus
FIGURES 75-78. Distribution maps for Australian Berosus spp.
18 C. H. 8. WATTS
3
80 W
7
@ B. arcus e B. debilipennis
m B. discolor a B. duplopunctatus
A B. sadiae mw 6B. quadropunctatus
0 B. aquilo © B. trishae
a B. nicholasi
y
82 Vv)
8
a 8B. munitipennis @ B. pulchellus
@ B. macumbensis O B. aemoenus
A B. veronicae A B. timmsi
4 B. subovatus
FIGURES 79-82. Distribution maps for Australian Berosus spp. (cont.).
REVISION OF BEROSUS )¥
is durker. Head strongly and evenly punctate except
towards front where punctures are considerably
smaller and weaker, Punctures on pronolulh uneven
in-size and often with a rather patchy distribution,
A rew of very Small punctures along frontand rear
margins. Elytral striae weakly impressed, lor the
mosi part reduced to a row of Well-impressed pune-
tures about the size of the larger punetures on
pronatum. Inerstral punctures on disc markedly
symaller than (hose on sides of elytra; in no case is
there a tendeney for interstrial punctures to farm
a single row, Interstrial punctures on dise tend to
be uneven in size. Underside densely rugose-
punctate, Midline of mesosrernum with weak rather
rounded keel, projecting alittle distance backwards.
Midline of first abdominal seament with litle or
no keel, Metacoxal process raised, triangular, a
narrow diarnond or oval-shaped area devoid of
sculpture io the midline. Rugase portion of meta-
femur t/2 to 2/3 Jength of femun that on
mesofemur.a little over 1/2.and on profermura 1/3
toa 2 length of femur.
Male: Pratarsi four-segmented, basal two
sepmenrs expanded, basal segment larger than
second, sevond and third segment subequal in
length,
Remarks
A widespread inland species readily recognized Irom
the other large Berosus by the pattern of punctation
on the elytra, Where the outer punctures are much
stronger than the inner punetures,
Distribution (Fig, 81)
Inland oAustraha.
Berostis minitipennis Blackburn
(Figs 3, 58, 64)
Berosus munitipennis Blackburn, 1895, p. 30.
Livper
Holotype, ?mate,
RM(NH), seen,
Luke Callabonna, S.A., in
Deseripnon (umber examined 48)
Length 4,8-4.6 nim. Elongate oval, not
humpbacked. Apex of elytron with yery long
narrow ouler spine aud shor joner spine, outer
usually Jonger than distanee between spine bases..
Light-brown, pranotum and head slightly darker
than elytra. Suture fines and some punctures usually
black on elyrron disc, Ventral surface dark-brown
to black, appendages brown, ip of labial palpi
darker. Head moderately fo quite densely covered
with punctures, large and well-marked in basal halt,
smaller towards froor. Elytron shiny or with weak
to moderate reliculation, Pronotum sparsely or
moderately and father unevenly covered with
variably sized, well-marked punctures, front and
fear margins with anly a few shallow small. punc-
tures. Elytral striae weakly to moderately impressed,
otien reduced oyer much of disc toa row of punc-
tures. Serial punctures about same size as larger
ones on pronotum. Interstrial punctures sparse,
shallow, lacking completely over atuch of elytra in
some specimens, where present tend to be arranged
in single row, Second elyirae stm less than 1/4
length of elytron. Ventral surface shallowly rugase
punctate. Midline of mesosternum weakly carinate.
weakly extended backward, Midline of first
abdominal seement weakly carinate in basal 1/2 oF
1/4. Mctacoxal process [riangularly-raised, with
patch in midline withour sculpture, pateh bounded
at rear by a slight V-shaped ridge extended back-
wards in midline to tip af process. Rugose portion
of metalemur |/3-2/3 leneth of femur, that of
mesoferur I/4-1/2, thal on profemur 1/3 lengtt
of fernur along ventral edge.
Mule: Basal two seanmients of protarsi moderately
expanded, First segment about 2 « lengih of
second, which is.about the same length as the third,
Remarks
An inland species readily reeognized by the long,
thin, outer elytral spines. and virtual lack of an inner
ong and the weak development of sculpture on the
elytra, This latter charaeter tends to be more marked
in central and noerthert specimens compared with
those from Vie. The eontrast of the pale upper side
and black ventral surface in most specimens is quite
striking. The extent of ruvosity an the femora ts
particularly yariable in this species.
Distriburian (Fig. 81)
Inland Australia south of Barrow Creek, NLT.
Berosus amoenns sp. mov.
(Figs 56, 62)
Distribution (number examined nine)
Length 3.3-4.5 mm, Elongate oval. Weakly
reticulate in some specimens. Elytra nol hump-
backed. Apex of elytron truncated, spines vestivial,
Dorsal surface dark-brown to black, edges of pro-
notum arid elytron yellow-brown, sides of elyiren
with three pale patches extending inwards from
sides, (he apical ones more distinct, Dise of elytron
variegated yellow and brown-black.. Ventral surface
dark-brown, appendages a little lighter, apex of
labial palpi dark-brown. Head sparsely covered wilh
strong punctures, about. 1.5 x size of eye facet, those
towards frant much smaller. Pronotum moderately
but unevenly covered with strang punctures, a little
larger than those on head, punctures tending to be
20 ©. 1 5. WATTS
elongate longitudinally, a few isolated smaller
punctures along front and rear margins, elytral
striae moderately lo slrongly impressed. Second
sina on elytron with 11 to 16 punctures. Punetures
in striae, between siriae and on pronotum subequal
in sizvé, Interstrial punctures lateral ro siria 9
apyoximately arranged in one line. Veriral surtace
rugose-punctate, Midline of mesosternum weakly
raised, moderately projecting backwards. Midline
of first abdominal seement weakly carinate in front
1/4, Metacoxal process raised, triangularly pro-
duced backwards with a shor! narrow keel in
midline at extreme rear apd small narrowly oval area
in widdle devoid af sculpture. Rugose portion of
metalemur 2/3 « length of femur, that of
mesoferur 1/2-2/3 that of profemur 1/4-1/2
length of femur,
Male; Protarsi four-segmented, basal two
segments broadly expanded, basal longer than
second which is a litle longer that the small third
seginent, Apical abdominal segment entire.
Remarks
A tare species from coastal NCT., readily recognised
by the black head and predominantly black
pronotum. The size and black head resemble the
& approximans aroup of species bit the tinlobed
mietacoxal process and lack of mesosternal pillar
separate il from that group.
Distribujian (Fig, 82)
Coastal NT.
Tipes
Holotype, male. 12°52'S, 132°50'E Koonearra,
\Skm E of Mt, Cahill, NT, 15.41.1972, M, §.
Upton’, in ANIC, Paratypes; two, ‘16°28'S,
136°09'E 46 km SSW of Borroloola, NV. 28 Ot,
1975. M.S. Upton’, in ANIC; six, ‘King River NUT,
W. McLennan Pres. H. L, White 17,1016", in NMV,
Berosus josephenae sp, nav,
(Figs 55, 41)
Descripron (nuriber exaniined 19)
Length §.5-6.0 mm. Elongate oval. Elyrra wor
humpbacked. Apex of elytrou weakly to quite
strongly produced backwards, ending in rwo spines,
the outer well-develaped, the inner much less so,
Dorsal surface light-brown, with patches on head,
centre of pronotum, elywon, inner striae and pune-
tures an elyiron darker. Ventral surface dark brown-
black, appendages lighler, rugose portions of meso
and metafemora and tip of lapial palpi dark brown-
black. Head strongly punctate, punctures larger
that eye facets, punctures weakening towards lront.
Pronotum strongly densely and cvenly punctate,
punctures ol one size execpt fora few stnaller ones
on extreme front and rear edges; Elyival sitlae
strongly impressed. Siria 2 with 15 to 23 punctures
about size of thase an pronotum and 1.5 to 3 +
size of {hose in adjacent interstrial area. Inferstrial
punctures relatively large, uniform in size, well im-
pressed, arranged in more than one row over most
of elytron, larger but shallower towards sides. Ventral
Surface strongly rugose-punciate, Midline of meso-
sternum weakly to moderately carinate, moderately
projecled backwards, Micline of first abdominal
segment raised in front third, Metacoxal process
raised, tritumgularly produved backwards, weakly
ridyed in midline except for « broadly diamond-
shaped area in centre which is devoid of sculpture.
Rugose portion of mejafemur 2/3-3/4 length of
femur that ol mesofemur 1/2-2/3, thal of
profemmur about 1/2 length of femur.
Male: Protarsi four-segmented, basal seaments
moderately expanded, basal segment niuch longer
than second seginent which is aboul same length
as much narrower third segment, Apical abdominal
segment entire,
Remarks
As yet known only by 4 few speciinens from coastal
NT, this rather clongale species is distinguished by
ils Strong even punctration on the pronatum ane cle
black rugase portions of the femora.
Distribution (Fig. 76)
Coastal N.T. and Cape York, Qld.
Tepes
Holotype, male, 12°23'S 132°57'E § km NNW of
Cahills Crossing, NT. East Alligator River, vi. 73,
Upton & Foehan’, in ANIC, Pararypes; six, two
same data as holotype, in ANIC: one, ‘Mareeba
22.1976 Ko & EF. Carnaby’ in CW. one, NUT,
Junction of Arnhem Hwy & Genpelli Road MV.
light 26-27 Sune 1980 M, B. Malipahi’, in NTM;
one, NT, Lake Bennelt area © 25 km SE of
Manton Dam 29-30 Dec, 1979. M. B. Malipatil’,
in NT,
Berosus gibhae sp. nov.
(Figs 33, 39)
Description (gumber examined '7)
Length 4,0-5,0 mm, Elongate oval, Elytta jot
humpbacked. Shiny but with elytvon of many
specimens with a line-meshed reticulation. Apex af
elytron wilh a stout moderately-sized outer spine
and a virlually absemt inner one, Dorsal surface
hzht-brown, elytral punctures and many striae and
vague patches on head, pronotum and elytron
darker. Ventral surface dark-brown, appendages
lighter, except tip Of labial palpi which is dark-
brown. Head moderately densely covered with
REVISION OF BEROSUS 21
strong punctures, strong, 2 x size of eye facets
except towards front where they become much
smaller, Pronotum evenly and quite densely covered
with strong punctures, the same size as those at rear
of head, a few smaller ones along front and rear
margins. Elytral striae moderately impressed, with
punctures the same size or slightly smaller than
those on pronotum. Second elytron stria with 12
to 18 punctures. Punctures in interstriae smaller
than those in striae, larger but shallowly impressed
towards side, over most of elytron arranged in one
or two irregular lines, often bearing a single seta
particularly those towards sides. Punctures in
interstria 3 of uniform size, Ventral surface densely
but finely rugose-punctate. Midline of mesosternum
strongly raised, moderately projecting backwards.
Midline of first abdominal segment weakly carinate
in front 1/2-1/4. Metacoxal process raised, tri-
angularly produced backwards, with a broad
diamond-shaped area in middle devoid of sculpture.
Rugose portion of metafemur 2/3-3/4 length of
femur, that of mesofemur 2/3, that of profemur
about 1/2 length of femur on rear face.
Male: Protarsi four-segmented, basal segments
weakly expanded. Basal segment longer than
second, second and third segments subequal in all
dimensions. Apical abdominal segment broadly but
shallowly notched.
Remarks
This widespread northern species is rare in
collections. The peculiar fine sculpture of the elytra,
which is either irregular or with a fine grained
reticulation, and the presence of a large number of
setae- bearing punctures, are reminiscent of B,
nutans and B. dallasae which show these characters
to a greater extent. The more normal pattern of
elytral punctation readily separates it from these
species, in which the elytra are densely and evenly
covered with punctures. The round rather than
elongate pronotal punctures also separate it from
these otherwise quite similar species. The light-
coloured rugose portions of the femora separate it
from B, josephenae.
Distribution (Fig. 78)
Coastal northern Australia.
Types
Holotype, male, ‘N.T. Katherine low level Native
Park, 25 Ap. 1980 M. B. Malipatil’, in NTM.
Paratypes; six, two same data as holotype in NTM;
four ‘NT. N. P. Korlonjotlok Stream 18 Nov. 1979
M. B. Malipatil’, in NTM,
Berosus majusculus Blackburn
(Figs 54, 60)
Berosus majusculus Blackburn, 1888 (1889), p. 824.
Types
Holotype and paratype in BM(NH), seen. Paratype,
male, Adelaide, S.A. in SAM, seen.
Description (number examined 336)
Length 6.0-8.7 mm. Elongate oval, elytra not
humpbacked. Apex of elytron with two well-
developed spines, the outer longer than the inner.
Dorsal surface brown with darker mottlings. Elytral
punctures and striae black, strongly so on disc,
weaker or absent on sides. Ventral surface black.
Appendages brown, tip of labial palpi darker. Head
relatively narrow, evenly strongly and quite closely
punctate, punctures much weaker towards front.
Pronotum unevenly covered with moderately dense,
well-marked punctures, variable in size, a little
stronger laterally with a row of small punctures
along front and rear margins. Elytral striae well-
impressed, second stria 1/4 ‘length of elytra.
Punctures in striae well-marked, same size or larger
than those on pronotum, interstrial punctures not
in one line except in some lateral interstriae, well-
marked, a little smaller than those in striae,
particularly on humeral angles. Reticulation of
elytron absent to moderately strong. Ventral surface
densely and evenly rugose-punctate, Midline of
mesosternum tending to form weak keel, weakly
projecting backwards. Midline of first abdominal
segment with weak keel in anterior 1/4. Metacoxal
process raised, narrowly triangular. Midline with
long narrow area devoid of sculpture. Rugose
portion of metafemur 1/2 to 2/3 length of femur,
that on mesofemur a little over 1/3 along ventral
face, that on profemur about 1/3 length of femur
along ventral face.
Male: Protarsi four-segmented, basal two
segments dilated. Basal segment about twice length
of subequal second and third segments. Apical
abdominal segment notched.
Remarks
This common southern species is readily separated
from the rather similar B australiae by lack of black
rugose, femore. It is more difficult to separate from
B. veronicae, with which it is generally sympatric,
but it is larger and has a greater number of
punctures on stria 2. The notch on the last
abdominal segment of the male also separates this
species from others of its size.
Distribution (Fig. 75)
Southern coastal Australia from Perth to Sydney
areas.
Berosus veronicae sp. nov.
(Figs 67, 68)
Description (number examined 209)
22 CORLS.
Lengitt 35-64 mm. Elongate oval, elytra not
humpbacked. Apex of elytron with two Weak blunt
spines. Dorsal surface light-brown with darker
mottlings. Elytral punctures and striae black,
strongly sa on dise in many specimens, Weakly or
not ut all lacerally, Ventral suclace black, append-
ages light-brown, tip of labial palpi a little darker.
Head evenly, strongly and quite closely punctale
Punctures toward front weaker. Pronolum evenly.
strongly and quite closely punctate, svatered
smaller punctures particularly on disc and front and
rear margins, weakly reticulate in some. Elytron
variably reticulate fram none to moderate, Elytral
striae well impressed. Second stra about |/4 length
of elytra. Strjal punctures about size of targer
punctures on proootum. Interstriae | to S scattered,
those between more lureral striue tending to be in
one row except for extreme lateral edge. Ventral
surface densely pugose-puretate, Midline ol’ pro-
‘sterniudy Weakly and evenly keeled, projecting
backwards in weak prosrernal process. Midline of
first abdominal segment weakly keeled in anterior
1/4. Metacoxal process raised, triangular, midline
with diamond-shaped area devwid of sculpture in
middle and shght keel to the rear of this. area.
Rugose portion of merafemur berween 1/3 and 1/2
length of femur, that on mesofemur 1/3, that on
profermur 1/4 length of femur.
Male: Protarsi four-segmented, basal two
segments moderately expanded. First sexment about
twice length (and size) of second, which isa little
longer than the narrow third segment, Apical
abdominal segment entire,
Remarks
A cammon species in southern Australia, this
species has frequently been mis-ideritfied in
collections, usually as & majusculus, The un-
notched apical seyment in the male separates it from
this species. The smaller size and fewer punctures
in the area between the lirst two elytral striae will
separate all but the occasional specimens. Balbipes
Fauvel is another name rhat has been altached to
specimens of this species but this isa species from
New Caledonia with differen aedeagus and other
characters,
Distribution (Fig, 8)
South-eastern Australia, ‘Tas.
Tipes
Holotype, male, ‘Goulburn’, in SAM. Paratypes,
same data as the holowpe, in SAM; two, ‘Albury
NUSW. 1/61 CW", in CW.
Berosus austratiae Mulsant
{Figs 71, 74)
Berosus ausrratiae Mulsant, 1859, 1p. 38,
WATTS
Berosus exlernespiaosus Pairmaire, i879, p. Rl sven.
aller D’Orehymont, 1943_
Berosus gravis Blackburn, 1888 (89), p 826, syn,
nov,
Tipes
Berosus australiae, not loeated (Hape, Dept of Ent.
Oxford, revords i as having been sent ta BM(NH)
in 1925). Type locality ‘Australia’,
Berosus zravis; Holotype, male, S.A. in RM(NH),
seen. Paratype, malein SAM, seen. Paratype, male,
in SAM, seen. Type locality (Paratype), Murray
Bridge, S.A.
Berosus externespinosus, nol lovaled (?7MHNHI.
lype locality, Rockhampton, Old.
Description (number examined 455)
Length 6.5-9.0 tam. Elongate oval, not tiump-
backed, Apes of elyLron with two spines, the outer
strong and usually abour two times length of inner,
Dorsal surface brown to dark-brown with darker
motlings. Elytral striae and punctures outlined in
black except for punctures laterally, Ventral sur
face black, Appendages brown, rugose portion of
femora black or dark- brown. Punctures ov head
nioderately strong and dense, much Weaker towards
front, Pronotun) moderately covered wiih well-
marked variable sized punctures, front and reat
edges with rows of elasely spaccd small punctures.
Elytron with sinae moderately impressed, strial
punetures relatively small, aboul same size as those
on pronotum. Interstrial punctures seatreredt,
smaller, those in more lateral incerstriae tending to
form a single line to varying degrees, Blyton surface
shiny or Weakly to moderately reticulate with a fine
reticulation. Ventral surface densely rugose-
punctate, Midline of mesosternuin raised into weak
keel, weakly projecting backwards. Midlinpe of first
abdominal segment weakly keeled in front 1/4,
Metacoxal process yaised, iiangular, with diamond
Shaped area devoid of sculpture in midline. Rugose
portion of metalémur 2/3-3/4 leneth of femur that
on mesofemur a little more (han 1/2. that on pio-
femur about 1/3 length of femur.
Mule: Protarsi four-segmented, basal two scx
ments strongly expanded, basal segment about 1.5
* length of second, whicl is tach longer than the
small third segment, Apical abdominal segment
entire.
Remarks
In southero and eastern Australia this large
common spevies 18 readily recognized by the black
rugose portions of the femora, In northern
Australia some specimens have the nigose portions
dark-brown and not greatly darker (han the rest of
the les, and closely resemble B sadiewe and &
decipiens, Separation of these three species is hest
REVISION OF BEROSLS 44
done on aedeagi as. although & wivstraliae is
nenerally larger, aud generally has stronger elytral
purictupes, this is nor always so,
Distrifuition (Fig, 77)
Virtually Australia-wide, more common ii sauth-
cast than elsewhere, rare in W.A., apparently absent
from centre and north of thar state.
Berosuy decipiens Blackburty
(Fips 47, 52)
Berasus decipiens Blackburn, |S88, p. 827.
Types
Holotype and pararype in BM(NH), seen, Paratype
in SAM, seen.
Descriplion (number examined (8)
Length 6.1-8.0 mm, Elongale oval, nor
hinmpbacked. Apex of elytrow with two weul spines
placed close together. Reddish-brown with elytral
siriae and punetures on elytron and pronotum
black, particularly on dises, black areas on elywron
coalesced in. some spevimens lo give extensive black
areas on disc, Ventral surfaee black, appendages
reddish-brown, rugosé portions of lemora darker
Head moderately punctate with rather shallow
punctures, those in front half smaller. Propotuin
sparsely and rarher unevenly punclale wilh srvall
ro Mmoderately-sized rather shallow punctures. Front
and rear boarders with row of shallow small! punc-
sures, Elytral striae weakly impressed especially on
dise where for most part reduced toa line of punc-
lures. Sina! punctures about same size as the large
punctures on pronotum, Interstrial punctures sparse
and shallow, those on disc smaller (usually much
smaller) than those ii adjacent striae, weaker
laterally, strong tendency for interstrial punetures
to be arranged in one line over most of elytron, im
intersiriae 3 and 5 there are some seatiered larger
punctures. Venrral surface rugose-punctate. Midline
of mesoslernum quite strongly and sharply carinute,
weakly projecting backwards. Midline of first
abdominul segment without keel. Metacoxal process
raised, triangular wilh rather bowed sides, Midline
weakly carinate hehind, with narrow area in middle
unsculpcured. Rugose portion of metafemur
1/2-2/3 lenyth of femur, thal of mesofemur about
1/2, that of profemur a liltle over 1/2 length af
femur
Male: Protarsi four-segmented, basal twa
segments moderarely expanded. First segment about
1.5 « length of second segment which is a little
shorter than the narraw third segment,
Remturks
&. decipiens tsa northerm coasu species resembling
B, australiae, \t differs (ram that spevies by generally
being a hittle smaller and more clongate, and by the
weaker and less numerous punctures on elytra and
pronotum., The aedeagus and the much larger third
segment of che male prorarsi separate it fram B
ansiraliee. \ have fourid no characters that separate
this species from B, sadieve other than the aedeagus.
Distritnition (Fig, 78)
Coastal northern Australia from Wyndham, W,A,,
to Kuranda, Qld.
Berosus sadleae sp, nov,
(Figs 46, 50)
Description (umber examined 2)
Length 4.5-5.0mm Narrowly oval. Shiny. Elytrou
nol humpbacked. Apex of elytron with two weak
spines) the inner one smaller (han the outer, Dorsal
surface light-brown, punctures on elylron and
pronotym, elytral striae and patches on pronotum
and ¢lytrou darker. Rear of head dark, centre and
front lighter. Ventral surface dark-brown, append-
ages lighter, rugose porlions of meso and meta-
femora somewhat darker than rest of legs.. Tip of
Jabial palpi dark, Head relatively narrow, relatively
sparsely covered With well impressed punctures,
about size of eye facet, smaller towards front.
Pronotum sparsely covered with punetures, most
about 1.4% puncture width apart, towards midline
and in front punctures vary in size, Elytral strive
weakly lo moderalely impressed, strial punctures
about size of the larger ones on pronotuim,
Intersirial punctures smaller. In interstriae 3 and 5
there are some scattered larger punctures. Punctures
lateral to about stria 7 weak and arranged in one
row, those inward of about stra 7 stronger and tend
to be seartered. Stria 2 with I to 1S punctures which
are about /wice as Jarge a5 these in adjacent inter
strial areas, Ventral surface densely punctate.
Midline of mesosternum raised, moderately pro
jecied backwards, Midline of trst abdominal seg-
ment weakly raised in front 1/3, Melacoxal process
raised, broadly triangularly produced backwards,
with srnall wide diamond shaped portion in middle
devoid of sculpture. Rugose portion of metalemur
2/3 length of femur, that of mesofemur 1/2-2/3,
that of profeniur 1/2 length of femur
Male: Protarsi four-segmented, basal segment
weakly expanded, first segment larger chan second
which is sume leneth but-a little wider than third.
Las| abdominal segment entire.
Reimiurks
A northern species Known for certain only from the
iwo inale types. These are indistinguishable [rom
B. deviniens excep for the distinctive aedeagus,
Both species have been taken together at Howard
Springs, N-T., and Lake Bennet, NC Further work
a4 C1. S. WATTS
is heeded to clarily the relationship of this northern
group which also includes & wusfralive and B.
aquilo, A series af specimens from Lake Bennet,
NT. Dec. 1979, in NTM, includes both &..swcdieae
and & decipiens but most are teneral which
precludes. extraction of an identifiable aedeagus,
hence J carol be sure to which species various
individuals belong.
Disiriburian (Big, 79)
Known only from the wpe localiries,
Types
Holoiype, wale, “Howard Springs, NIL ar light,
27.1.68, E. Matthews’, in ANIC. Paratype, male
‘NT, Lake Bennett area & 25 km SE of Manton
Dam, 29-30 Dec, 1979, M. B. Maliparil al MW,
light’, in NTM
Berosus aguila sp. nov,
(Figs 34, 40)
Description (number examined 11)
Length 3.3-5.0 mm- Elongate oval. Elytra not
humpbacked. Shiny. Apex of elytron with two
spines, outer weakly lo moderately developed, inner
one often lacking, Dorsal surface light-brown,
punctures and striae on elytron darker, elycran with
one or twa darker patches, rear of head darker
Ventral surface dark-brown and black. Appendages
lighler, rugase portions of femora dark-brown lo
black. Head moderately covered with strong pune-
jures about the saine size us facets of the eye or
slightly larger, much smaller towards front. Pronotal
puncttifes same size a8 those al rear OF head, rather
unevenly distribuled, all approximately the same
size except for a few smaller ones along extreme
front inargin aid on disc. Elytral striae moderately
impressed, stronger towards sides anc apex, Inter
strial punctures smaller than those in striae,
arranged in one line aver most of elylron, except
between striae 1 to Zand 4to 5. Area bel ween striae
2 and 3 with a few larger punctures. Second elytral
stria with 10 10 16 punctures which are about 3
size of those in adjacent interstrial areas, Ventral
surface densely rugose-punctate. Midline of meso-
sternum raised, moderately produced backwards,
Midline of first abdominal segment carinate in {ront
third, Metaconal process.raised, broadly triangularly
produced backwards, midline weakly caripate, a
sal] diamond-shaped area in centre devoid of
sculpture, Rugose portion of metafermur 3/4 length
of femur, that af mesolemur 2/3, (hat of prefemur
1/2-2/3 length of femur.
Male: Protarsi four-seymenied, basal two
segments strongly expanded, basal seament larger
than second which is about same length as the
unexpanded thin.
Remarks
Acsmall dark species from voastal NV The only
smiall species in this grouping with the rupese
portions of the femora darker (han the pest of the
femur, and usally almost black.
Distribution (Fig, 79)
Known only from rhe type locality. and fram Daly
River Missian, NE (ANIC).
Types
Holotype, orale, ‘Coastal Plains Rescarely Station,
CS.LBR.O. Darwin, NT. at light, 30.66 E.C.B.
Langfield’, in ANIC, Pararypes, ain saric data as
holotype, seven in ANIC, two in OW,
Berosus vijae sp. nov.
(Figs 35, 41)
Description (number examined 31)
Length 3.0-4.5 mm. Narrowly oval, nat
humpbaecked, apex of elytran with two weak broad
spines, Yellow-brown, rear of head, areas adjacent
to eyes, clytral striae, 2-3 vague patches behind
imiddle of elytron, ventral surface except for
appendages darker. Shiny. Head spursely punctured
with variably sized punctures, sparser and weaker
towards front and centre, Pronotum sparsely punc-
tured with yatiably sized but quite well impressed
punctures, sironger and denser laterally, front and
rear margins with a single row of small punetures
irregularly spaced, Elytral striae moderately to quite
strongly impressed, particularly towards apex of
elyiron, Punctures in striae well marked, about same
size as the larger ones on pronotum. Intersrrial
punctures a liule to much smaller than those in
stride, arranged in single row oxcepr in interstrial
2to 3 Which has scattered large and small punctures.
Second stria aboul 1/4 length of elytran with 9 to
14 punctures, Mesosternal keel weakly raised. Meta-
coxal provess Iriangular with apex towards rear, a
small approximately oval area in midline devoid of
sculpture. Ventral surface, rugose-punctate with
smallish punctures. First abdominal segment with
weak midline keel in front 1/4, Rueose portion of
metalemur 3/4 length of femur, that of mesofemur
2/3-3/4 that of profemur about 1/2 on frant edge
and a little greater on hind edge,
Male: Protarsi four-segnented. Mirst three segarents
subequal in length, lirst moderately expanded,
second rather less 50 and third net at all. Apical
abdominal segment very weakly norehed with weak
protruberence at each side of woteh.
Remarks
A small widespread porthern species closely
resembling & ralph| and & veronicue, separated by
range of Characters given in Table 2, Generally with
REVISION OF BEROSUS
larger elyival pluretures and more strongly intpressed
striae allhough (hese are weakly impressed on (he
dise in some specimens.
Distribution (Fig, 75)
Northern Australia, predoyninantly che western half,
Types
Holorype, male, “Tindal, NT 14=3PS, $32°22'R.
1-20 Dec, 1967, light trap. W. J. M- Vesijens*, in
ANIC. Paratypes, four same data as holotype, in
ANIC.
Berosus ralphi sp. nav,
(Figs 72, 73)
Deseviptian (number examined 8)
Length 3.54.35 mm, Oval, Not humpbacked, Apex
of elytron iruncat or very weakly produced into
Iwo broad short spines. Yellow-brown, purictures on
elytron and a few small spats in apical 1/2 of elytron
darker, Ventral abdominal segments orher (han
upical are blotched darker. Shiny, Eltron alten
moderately reticulate, Punetures of head moderately
impressed, sparse, smaller towards [root and venter.
Punctures of pronotum moderately impressed,
sparse, variable ii size, With Yow of small punctures
along extreme lront and rear margins, Elytron striae
moderately to weakly impressed, Punctures on
striae close, well fmopresscd, ubout size ol larger
pronotal punctures, Unterstrial pupetures much
Smalley than those in strlacand yeattered (in iler-
sirtae One lo five, strong tendency to be arranged
in a single row in interstriae lateral to striu 6.
Interstria 3 with punctures of Uwe sizes. Stria 2
short, often poorly marked with 7 to 13 punctures.
Mesosternal keel inoderately raised. Metavoxal
provess, sharply trangularly produced backwards
and downwards, a small slightly raised area iu
midline in centre. Ventral surface rugose-punctate.
First abdominal sezment with raised midline keel
in tront L’4, Rugese portion of metafemur 3/4
length of femur, thal On mesalemur 1/2-2/3, thal
on prolemmur t/4-1/2 length of femur.
Male; Apical abdominal segment sinuate on hmd
margin. First (wo segments of protarsi expanded,
third segment not expanded, Firsi three seements
progressively smaller in length,
Remurks
Similar in general facies to & vijae and B veronicae
bur differing fron then) in the wrearer number and
smaller size of the punctures in the first elytral
juterstriae, Where they are arranged in more ihan
one line, and in chargeters of the aedeagus. In all
specimens so far examined the tip of rhe nietacoxal
process is sharply deflexed downwards, 4 character
not seen in other Australian Berosus.
fh
w
Distribution (Fiz, 77)
Coustal porth-westen) Australia and Queensland.
Types
Halotype, male, ‘Wyndham W.A, S. Stephens
20,2.01', in SAM. Paratyes 17; seven same data as
hololype, i S.A.M, one ‘Derby NWA, W. DL
Dodd’, in SAM; seven ‘Derby WIW.A,’, in SAM? two
‘Queenstand J.8261', in SAM.
Berosus subavatus Kaisch
(Pigs 36, 42)
Berosus stictieus Fairmaire, 1879, yp. 82.
Berosus sibovaTey Kuisch, 1924 (not Boheman,
1859)
Type
Holotype, female, Peak Downs NT, in MNHN,
seen,
Descriplian (number examined four)
Length 4.6-3.4) mm. Elongate oval. Elytra not
humpbucked. Shiny. Apex of clyiron rounded o1
weakly truncated, Dorsal surface light-brawn,
punctures aud striae on dise of elyrron and vague
patches on head, pronotum and elytron darker.
Punetures in lateral striae on elytron with large
squarish darkish subsurface markings. Ventral sur
face dark-brown to black, appendages lighter,
rugose porhions Of lemiura wilh a slightly darker hue
thar rest of leg, Head with scaltered relatively small
punctures, (hose at back of head a little larger than
size of eye facet, those towards ventre and front of
head smaller, Pronotum éparsely and unevenly
covered wilh punelires, of twa sizes, larger aboul
same size aS those ai rear of head, Elytral striac
weakly impressed, over most of elytran little more
than series of punetures. Second elytral stria with
6 to 7 punctures, about same size ora little smaller
than the larger ones of pronoium 2 © size of those
in adjacent interstrial areas, Punctures in interstriae
3 and 4 of two sizes. Inorerstrial puuctures lareral
ro stria 6 arranged iu one line, Thase towants sides
and apex small and weakly impressed. Ventral sure
face finely rugose punctate. Midline of mesoster-
num moderately and evenly raised, moderarely
projected backwards. Midline of first abdominal
segment weakly carinate in front 1/4, Metacoxal
process raised, trlangularly produced backwards,
midline carinate in rear 1/4 @ Targe roughly
diamond-shaped area in middle, devoid of seulp-
ture. Ruwose portion of metafemur 2/3 length of
femur, that of metafemur 2/3, that of profemur
about 1/2 length of femur
Male; Protarsi four-segmented, basal two
segments moderately expanded Basal segment
larger than second which is larger than third.
26 tH & WANS
Renwrks
The relatyely Small pumiber of punctures in the
sevond elyiral stria. strong lendency for punctures
in literstriae to be arranged in one ling, and sparse
and variably-sived pronotal and head punctures
separate this species from related species, B vijae
and B, raiphi are dit ficult to definitely separate trom
this species apart from the form af the aedeagus
which is vousiderably shorter than the parameres
in B subovetus,
Distribution (Fig, 814
Known from ouly two localities: Station Creek in
north Old. (in ANIC), and Katherine, ST. fin
ANIC). (The type is labelled s\ustral’)
Berosus nicholasi sp. nav,
(Figs 66, 69)
Description (number examined $1)
Lengih 5,0-6.5 mm. Glongate oval, bly(ra not
humpbacked. Apex of elytron weakly extended wilh
two small spines, inser one often small or lacking.
Shiny, often moderately reticulate with a very fine
even reticulation. Dorsal surface light-brown with
punctures, elylral stride and patches on head, pro-
notum and elyiron dark-brown, Ventral surface
dark-brown to black with lighter movtlings.
Appendages brown, tip of labial palpi darker. Houd
relalively narrow, evenly and quite densely covered
with slrang punctures, punctures weaker towards
front. Pronotim moderately bul unevenly covered
with strongly impressed punctures of 1wo general
sizes with the larger ones predominating, rows of
small punctures along front and rear margins.
Elytral striae quite strongly impressed except
towards sides and back. Srrial punctures relatively
small, a little smaller than the larger ones on the
pronotum. Interstrial punctures small, relatively
sparse, those laleral (o stria 8 arranged in-one line,
amaller than strial punctites aver most of elytron,
Some of the more central interstriae with a few large
punctures with shorl setae. Ventral surface densely
rugose-punctare, Midline of mesosternum raised,
moderately projecting backwards, Midline of first
abdominal segment carinate only in extreme front,
Metacoxal process raised, triangularly produced
backwards, small elongate Oval area in centre devoid
of sculpture. Rugose porrion of metafemur 2/3-1/4
length of metalemur, that of mesofemur 1/2-2/3,
that of profemur 1/3-1/2 length of femur.
Male: Protarsai four-segmented. Basal segments
subequal, quite strongly dilated. Apical abdominal
segment not notched.
Remarks :
One of a group of very similar northern specie
which can only be confidently separated by
characters of the acdeagys, Ip all but a few cases,
jarger than other specles in the group ocher than
B, dehilipennis, which usually has much sironger
puncrures On the elytron, See Table 2,
Distribution (Fie, 79)
Coastal northern Australia from Derby, W.A. To
Tawnsvifle, Qld,
Types
Holotype, male 12° 25'S 131'03'R Howard Springs.
NC 24 ki S of Darwin 10.41.72. rainfovest, at light,
E. Britton’, in ANIC. Paratypes, 12 same data ax
holotype; L) in ANIC, one in CW.
Berasies dehilipennis Blackburn
(Figs 44, 45, 49, 51)
Berosus dehilipernis Blackburn, 1898, pr. 223.
Tivpe
Holotype, temale, Cape York, Qld, in BM(NEL),
seen.
Description (number examined 74)
Length 4.0-6.0mm. Klongate oval. Elyira nor
humpbacked, Apex of elytron rounded or with (wo
weak spines. Dorsal surface light-brown with pune-
tures on pronotum and elytron, elytral striae and
patches on pronotum and elytron darker, Rear of
head dark-brown, centre and front lighter. Ventral
surface dark-brown, appendages hghter, lip of labial
palpi dark-brown, Mead wiilt sparse to moderately
denise, strong purictures, a little larger than eye
facets, grading 10 obsolete towards front. Pronotiun
rather unevenly bur moderately covered with strong
punctures the size of hose at rear of head, a few
smaller punctures towards midline and front, Blytral
striae quite strongly impressed parucularly laterally
and towards rear. Second elytral striae with 10 to
18 punctures, 2x the size of those in adjacent inter-
strial areas, Interstrial punctiires scattered ner
arranged in a single row except in same places
towards sides and rear, A few large punctures in
interstfiae 3 and 5, Veritral surface densely rugose-
punctale. Midline of mesasternum weakly raised,
moderately projected backwards, Midline of first
abdominal segment weakly carivate at extreme
front. Metaucoxal process raised, narrowly triangue
larly produced backwards, with a narrow diamonel.
shaped area in. middle unseulptured. Rugose partion
of imetafemur 2/3-3/4 length of femur, that of
mesofenyur 1/2-2/3, thal of profeniur [/3-1/2
lenath of femur,
Male; Protarsi four-segmented, First and second
seymenrs moderately expanded, lirst much longer
than sceand, which is about the same length as
third, Apical abdominal segment broadly bur yeey
weakly notched,
REVISION OF BEROSUS 27
Remarks
A widespread northern species which I initially con-
sidered to be composed of at least two species.
Further study may show that it inchides several
closely related species. In particular, individuals can
be placed in one of two groups according to the
shape of the aedeagus. One group has the parameres
thin and the aedeagus roundly hooded at the tip,
the other has thick parameres and has a transverse
ridge on the upper surface behind the roughened
end portion of the aedeagus, These characters are
to some extent variable and intermediates exist, eg.
notably a specimen from Cunnamulla, Qld., in
SAM. This species can be separated from the rather
similar B micholasi by the usually much stronger
elytral punctures and the relatively smaller second
segment of the male protarsi (Table 2).
Distribution (Fig. 80)
Coastal northern Australia from Derby to Caims
region.
CHULCKLIST OF AUSTRALIAN BEROSUN
(in alphabetical order)
8. australiaeg Mulsant
= externespinosus Fairmaire
= gravis Blackburn
B. amoenus sp. nov.
B. arcus sp. nov.
B, approximuans Fairmaire
= B. auriceps Blackburn
B. blackburni Zatz
= B. avipennis Fairmaire
- B. simulans Blackburn
~ B. stigmaticollis Fairmaire
aquilo sp. nov.
dallasae sp. nov.
debilipennis Blackburn
decipiens Blackburn
discolor Blackburn
~ B, flindersi Blackburn
duplopunctatus Blackburn
egibbae sp. nov.
invalutus (W. MacLeay)
Josephenae sp. nov.
B. juxtadiscolor sp, nov,
l
Reah boB&aS
ACKNOWLEDGMENTS
The Curators of the collections listed earlier are thanked
for the free and rapid access to their collections afforded
to me. Dr E. Matthews kindly read portions of the
manuseript and greatly improved it. Mrs P. Kidd’s expert
typing of the various drafts is greatly appreciated as is
Miss J, Thurmer’s drawings of elytrae and other structures,
Mrs. Marianne Anthony, Librarian of the S.A. Museum,
ferreted out obscure references without which progress
would hitve been much slower
. mlacropunciatus Sp, NOV,
} macumbensis Blackburn
. majusculus Blackburn
munitipennis Blackburn
nicholasi sp. nov.
niger 8p. nov.
nutans (W. MacLeay)
= B. pallidulus Fairmaire
B. pulchellus W. MacLeay
— B, devisi Blackburn
8. queenslandicus Blackburn
= B, quartinus d’Orchymont
quadrapunclatus sp. nov.
ralphi sp, nov.
reardoni sp. nov.
Sadieae sp. nov.
suboyatus Knisch
= 8B. sticticus Fairmaire
(rishae sp. nov.
timmsi Sp. Noy.
} veronicae sp. nov.
. vijde sp, nov,
Dehra nan
PaREtBe PRRDHA
REFERENCES
BLACKBURN, T, [888 (1889), Notes on Australian
Coleoptera with descriptions of new species. Proc. Linn.
Soc. N.S.MO (2/1011, 1S88(1889); 805-875.
BLACKBURN, T. 1889 (1890). Notes on Australian
Coleoptera, with descriptions of new species. Prove.
Linn, Soc. N.S.W. (2)1V: 445-482.
BLACKBURN, T. 1898. Further notes on Australian
Coleoptera, with descriptions of new genera and species.
Trans. R. Soc. 8. Aust, XXII, 1898: 221-223.
BLACKBURN, T. 1896. Coleoptera (exclusive of the
Carabidae) in Report of the Horn Scientific Expedition
28 in ALES:
to Central Australia, Part Il — Zoology: 254-263.
Melville, Mullen & Slade, Melbourne.
BLACKBURN, T. 1895. Further notes on Australian
Coleoptera, with descriptions of new genera and species.
Trans. R. Soc. S. Aust, X1X: 27-60.
WORCHYMONT, A. 1925. Contribution a l'étude des
hydrophilides II. Annals Soc. Ent. Belg. LXV: 139-169.
d’?ORCHYMONT, A. 1943, Neue oder interessante
Sphaenidiien und Hydrophilinen der Malayischen
region. Treubia Ill: 416-421.
d@ORCHYMONT, A. 1943. Notes sur la tribus Berosini
Bedel (Coleoptera Palpicornia Hydrophilidae). Bull.
Mus. Hist. Nat. Belg. 19(42): 1-12.
FAIRMAIRE, L. 1879. Déscriptions de Coléopteres
WATTS
nouveaux ou peu connus du Musée Godeffroy
(Hydrophilidae). J. Mus. Godeffroy XIV: 80-83.
KNISCH, A. 1924. ‘Col. Cat. Part 79: Hydrophilidae. 304
pp. Junk, Berlin.
MAcLEAY, W. 1825. New Coleoptera from Java. Annul.
Jay, 1825, 35.
MacLEAY, W. 1873. Notes on a collection of insects
from Gayndah. Trans. Ent. Soc. N.S.W. 11: 79-205.
MULSANT, 1859. Australian Hydrophilidae. Opuse. Ent.
IX: 58.
ZAITZ, 1908. Catalogue des Coléoptéres aquatiques des
familles des Dryopidae, Georyssidae, Cyathoceridae,
Heteroceridae et Hydrophilidae. Horae Soc. Ent. Ross.
XXXVIII, 1908; 282-359.
NABIDAE (HETEROPTERA) OF VANUATU
BY I. M. KERZHNER
Summary
Six species are listed, one of them (Stenonbis nitidicollis Kerzh.) is new from the Vanuatu fauna.
Keys to adult Nabidae and to fifth instar larvae of Arbela occurring in Vanuatu are given.
NABIDAE (HETEROPTERA) OF VANUATU
1, M, KERZHNER
KER/ZHNER, |, M. 1987, Nabidue (Heteroptera) of Vanuutu, Rec, S. Ausr, Mus. 211) 29633,
Six species are listed, one of them (Srenonabis nilidicollis Keren.) is new from the Vanuatu
fauna. Keys to adult Nabidae and {o fifth instar larvae of Arbela occurring in Vanuatu are given.
1. M. Kerzhner, Zoolowical Institute, Academy of Sciences of the U.S.5.R., Leningrad 199034,
U.S.S.R. Manuscript received (7 Pebruary 1986,
Previous records of Nabidae from Vanuatu
(formerly the New Hebrides) were based on the
material in the Museum National d'Histoire
Naturelle, Paris (Reuter 1908); the British Museum
(Natural History), London (Harris 1938, 1939;
Kerzhner 1970a); and the Universitetets Zoologiske
Museum, Copenhagen (Kerzhner 1970b), In all, five
species have been recorded.
The material received from the South Australian
Museum, the B. P. Bishop Museum and the U.S.
National Museum of Natural History (thanks to
the kindness of Dr G. F, Gross, (he late Dr J. L.
Gressitt and Dr Th. J. Henry) contains all of the
species already known from Vanuatu and one
species new for the fauna.
In the distribution lists below, only localities and
collecting dates are mentioned, The data on
expeditions, collectors and deposition of material
for corresponding years are as follows:
1943 — collector Knight} deposited in the U.S.
National Museum of Natural History, Smithsonian
Institution, Washington, D.C,, USA;
1950 and 1970 — colleetor N. L. H. Krauss;
deposited in the B. P. Bishop Museum, Honolulu,
Hawaii, USA;
1957 — collector J, L. Gressitts the same
museum,
1958 — collector Borys Malkin; the same
museum;
1960 — collector W. W. Brandt; the same
museum
1964 — collector R, Straatman; the same
mMuUscUmm;
1965 — Biospeleological Expedition, collector
G, P. Gross; deposited in the South Australian
Museum, Adelaide, South Australia;
1967 — collectors J. & M. Sedlacek; deposited
in the B, P, Bishop Museum;
1971 — Royal ‘Society and Perey Sladen
Expedition; collectors — P. Cochereau (on
Malekula J.), G. F. Gross (on all other islands),
Masing Andrew (who assisted G, F, Gross on
Tanna Jj); deposited in the South Australian
Museum, but duplicates from large series will be
sent 10 British Museum (Natural History), Museum
National d'Histoire Naturelle, Paris, or will be
retained in Zoological Institute, Leningrad;
1973 and 1976 — collector N, L, H. Krauss;
deposited in the U.S, National Museum of Natural
History.
SYSTEMATICS
Gorpis simillimus Harris
(Figs 1-3)
Previous records
MALEKULA, Harris 1939: 150-I51.
Material examined
ESPIRITU SANTO: Apouna R. Camp 2, 146
m, 30.VIU,1971, at light, 1G) above Namatasopa,
400 m, 30. VILL.1957, 1 9; Hill EB, of Luganville,
100 m, Macaranga, 10.1X.1957, 1 o.
General distribution
Vanuatu (type locality) and Solomon Islands.
Remarks
Previously (Kerzhner |970a) | mentioned that
there were some colour differences between
specimens from Vanuatu.and the Solomon Islands.
These differences are in the main confirmed by the
new mutterial, In addition, shght differences in male
and female genitalia (Figs 1-3) have been found.
However the material at hand is insufficient for a
study of individual variability and therefore |
refrain from description of a new subspecies for
the Solomon Islands population.
Arbela immista Harris
Previous records
MALERULA. Harris 1938; 579; Kerzhner 1970a:
298; 1970b; 192,
0 L M. KERZHNER
FIGURES 1-3, Gorpis simillimus Harris, 1, male from
Vanuatu (Espiritu Santo), aedeagus; 2, female from
Vanuatu (Espiritu Santo), vagina in ventral aspect; 3,
female from Solomon Jslands (Guadalcanal), the same.
Material examined
ESPIRITU SANTO: Baldwin Bay, 17,VILL, 1958,
} 2; 25 km NE Luganville, 12.1V.1964, 1 of, 1 9.
PENTECOST (NE): 200-500 m, 27, 111.1964, 1 o.
MALEKULA: Amok, 17.1X.1958, 1 0% South West
Bay, 11 and 13.X,1971, by beating trees and sweeping
grasses in the forest and along the river of the forest,
9 of, 10 9, 4 larvae,
General distribution
Vanuatu (type locality) and the Bistmarck
Archipelago,
Arbela costalis Stal
Previous records
BANKS, MALEKULA, ERROMANGA. Harris
1938; 581; Kerzhner 1970a: 298.
Material examined
API (‘Epi’): Vaemali, 10,VIII.1967, 1 oc.
ERROMANGA: Ipota, 0-100 mi, March 1970, 2
@; Ipota, vicinity of Ipota, and River Camp
between Ipota and Nuankao, 4, 5, 7, 12. VII1.1971,
4 o, 3 9. TANNA: Lenakel, 0-150 m, March
1970, 1 9; between Lenakel and Bethel,
28.VIL1971, 1 larva), ANEITYUM: vicinity of
Analgahaut, 19.V11.1971, 1 o, 1 larva.
General distribution
Fiji (type locality), Samoa, Vanuatu and
Solomon Islands,
Arbela nitidula (Stal)
Previous recards
MALEKULA., Reuter 1908: 127; Harris (938:
568,
Material examined
ESPIRITU SANTO: Segond Channel, Santo,
Aug. 1950, 2 9; Namatasopa, 300 m,
28.VII1.1957, 1 9; below Namatasopa, 250 m, 2
and 3.1N.1957, 1 o, 1 9; Tasmalum, 3 m, bush,
4.1X.1957, | or; Luganville, 23-28, V1I.1958, 1 or;
Baldwin Bay, 28-30.1X.1958, 1 9; Narango, 90
m, May-June 1960, 2 9; Apouna R. Camp 2,
146.3 m, 2, 3.[X.1971, 10%, 7 9, 3 larvae; Malao
Village, Big Bay Area, 23.VIII.1971, | larva.
MALEKULA: Amok, 17,1X.1958, 3 o, 4 9,1
larva; South West Bay, 13-14.X.1971, beating
small trees and sweeping grasses along the river to
the forest and on swamp, 4 oc, 2 9, EFATE: Maai
(Mat, Ambryn Vill.), 3 m, 1S.VIIL.1957, 1 9;
Limestone, Plateau N of Maat, 100 m, 18,
20. VIIL.1957, 2 0; Vila, 0-100 m, Jan. 1976, 1 @,
ERROMANGA: Ipota, 5.VIII.1971, 3 9,
TANNA: between Lenakel and Balhel,
28.VIL1971, 1 o, 2 9. ANEITYUM: vicinity of
Analgahaut, 18-20.VI1,1971, 3 ©,
General distribution
From India and south China to Vanuatu,
Stenonabis nitidicollis Kerzhner
(Figs 4-6)
Material examined
ESPIRITU SANTO: Nokowoula, 1100 m,
14.1X.1971, by sweeping, 1 brachypterous female,
General distribution
Australia, Vanuatu (new record).
Remarks
S. nitidicollis was described (Kerzhner 1970a)
from a macropterous female, collected in New
South Wales, Mrs N. Strommer (formerly N,
Winkler), now in Heathmount, Melbourne, kindly
informed me that she has examined macropterous
and brachypierous males and females of S.
nitidicollis trom Queensland. | compared the
NABIDAE OF VANUATU 3
female (Fig, 4) from Vanuata with the holotype of
S. nitidicollis.
The yagina in the Vanuatu female (Figs 5, 6) is
slightly larger (width 1.00 mm, in the holotype 0.93
mm), the dark coloration of the body is more
pronounced and the hind lobe of pronotum has an
intermediate longitudinal brownish stripe on each
side berween the medial and lateral stripes, the
antennae are longer (length of the first segment 0,93
mm, of the second 1.33 mm), and also the legs and
rostrum are slightly longer. All these differences
do not surpass the level of individual and
geographic variability in related species.
In the Vanuatu specimen, head width 0.76 mm,
yertex width 0.34 mm, hind lobe of pronotum
length 0.43 mm, width 1.29.mm, scutellum of equal
length and width, body length 6.7 mm, width of
abdomen 2.0 mm.
Nabis (Tropiconabis) kinbergii Reuter, 1872
Nabis nigrolineatus (Distant, 1920). Nabis
tasmanicus Remane, 1964. Nabis capsiformis auctt.
(non Germar, 1838), part,
Previous records
‘NEW HEBRIDES’ (as Reduviolus capsiformis).
Reuter 1908; 114, TANNA (as Nabis nigrolineatus).
Kerzhner 1970a; 355.
FIGURES 4-6. Stenonabis nitidicollis Kerzhner, brachyplerous female from Vanuatu. 4, dorsal aspect; 5, vagina in
dorsal aspect; 6, vagina in ventral aspect.
32 1M, KERZHNER
Material examined
ESPIRITU SANTO: tho exact locality, Oct,
1943, 1 @; Segond Channel, Santo, Aug. 1950, 1
©; Luganville, 20, 23-28.VIL1958, 17 o,2 9, 4
larva; 10 km W. Luganville, JOJ1, 1964, 1 9)
Santo, 13, 14.X 11.1965, 1 o, 19; Malao Village in
Big Bay Area, 28VIILI97I, 1 9, MALEKULA:
Amok, 17.1X.1958, 1 9, | larva; Lakatoro,
16-17,%, 1971, 1 o, EPATE: Vila, Aug. 1950, 1
©; Vila, 0-100 m, Feb. and March 1970, | o, |
©; Vila, 0-200 m, Feb, 1973, 1 ©; Vila, 0-100 m,
Jan. 1976, 2c; SE corner, JONVILA971, 1 oy 1
3, Plantation Gaillarde nr. Tagobe, 11. VIL.1971,
1 9, ERROMANGA: IL km W. of lpota, 100-200
m, Feb. 1970, L ony tpota, 0-100 m, Mareh 1970,
1 o; Ipota and vicinity of Ipota, 5, 6, 10,
12. VII.1971, 5 o&, 9G, TANNA: East Coast, 450
m, 8.011.1964, 1 o, | ©, Some specimens are taken
at light.
General distribution
Australia and Pacific islands. The most remote
records are Ryukyu and Bonin Islands, New
Guinea, Samoa, Society Islands and New Zealand.
Remarks
The species was known formerly as N.
tasmanicus and then as N. nigrdlinealus. However
some doubts existed on {he last name because
W, L, Distant described the species in Reduviidae
(genus Sestrapuda) and mentioned a number of
characters not appropriate to Nabidae. | examined
the type series of N. kinbervii (Naturhistoriska
Riksmuseet, Stockholm), which included | female
from Sydney, belonging to N. nigrolineatus, and
2 females from Buenos-Aires, belonging to N,
cupsiformis. | designated the specimen from
Sydney as the lectotype of N, Ainbergii and this
name should be used for the species occurring in
Australia and surrounding islands (Kerzhier 1981;
Woodward & Strammer 1982).
KEY TO ADULT NABIDAE OCCURRING IN VANUATL
Note, The key is intended only for determinaiun of
Vanuatu! material, therefore same ol the characters
mentioned are not applicable to all species of included
genera nor (o all populations of the included species,
1, Fore coxal cavities closed behind. Fore coxae slender,
greatly clongated, Fore tibiae curved, shorter than fore
femora... 2... -. 0. Grorpis Stal
One. species _ GC. simiflimus Harris
— Fore conal cavities open behind, Fore caxae nearly
canical, less than twice as Jong as (hick, Fore libiae
straight, subequal in lengtl) to the fore femora...
it 2
2. Nore and middle femora and tibiae with long slender
spines, Ocelli vonliguous lrhela Sifl (see 3)
—~ Femora and tibiae withouw ae spines. Ocelli well
separated . . ee a oe a, Oe Ae,
3. Pronotum dirty yellow wilh (wo longitudinal brown ar
black stripes, seldom completely light coloured. Hind
lobe of pronotum not dull, bur Horse strongly shining
as the fore lobe, Hind tibiae of lhe male wilh a densely
pilose sub-basal thickening «0... 4. mit/dula (Stal)
— Promotum (except in distinctly toneral specimens)
complercly black or wilh only corners or sides of the
hind lobe yellow. Hind lobe of pronotum either velvety
dull, or as strongly shining as the fore lobe. Hind tibiae
of the male without thickening, ...... 4... 000.04
4. Hind lobe of pronetum cull.
vipepo ced. Immista y Harris
- Hind lobe of peonotum strongly sev afl Os
nfotefeten..ecfry-ommatet-tetam " jas LAL castalis sia!
5. Hind lobe of pranotum punctured. Connerxivum below
not separated from the venter by a suture. Short- or
long-winded. -----..,))......Slenonabis Reuter
One species — S$. nitidienllis Kershner
- Hind tobe of pronotum without punetures,
Connexivun) below separated trom ihe yenter by a
suture lying in a deep impression. Long- winged .
One: species ~ N. Kinbergii Peder
Kiy TO FLETH INSTAR LARVAE OF ARRBEI 4
OCCURRING IN VANL ATL
|. Wing pads black wuh the cai third white -
esas t0 por. ' --A immista Harris
— Wing pads wnfealorous , REREAD SEER RIOCE Per-
2. Wing pads black . 4 hires 4 Costalis Stal
— Wing pads light votoured. - A. nitiduta (Sia
ACKNOWLEDGMENTS
Iam very thankful to Dr G, PF, Gross, to the late
Dr J. Le. Gressitt and to De Th. J. Henry for the interesting
material and to Dr N. Strommer for the information on
Stenonahis nitidicollis..
REFERENCES
HARRIS, H. M, 1938.. The genus_4rhe/a Stal (Hemiptera,
Nabidae). Man, Mag. rat, Hist, (ser, 11), te 561-584.
HARRIS, H. M. 1939, 4 contribution ta our knowledge
of Gorpis Stal (Hemiptera: Nabidae). Philipp. J. Sev.
69(2): 147-155.
KERZHNER, |. M. 1970a, Neve und wenig bekannte
Nubidae (Heteroptera) aus den tropisehen Gebieten der
Alten Well. Acta ent. Mus. nain. Pragae 38 (1969);
279-359.
KERZHNER, I. M. 1970b. Some Heieroptera Nabidae
NABIDAE OF VANUATU 33
(Hemiptera) from the Southern Philippines and the © WOODWARD, T. E. & STROMMER, N. 1982. Nabis
Bismarck Islands. Ent. Medd. 38: 117-194. kinbergii Reuter, the current name for Tropiconabis
KERZHNER, I. M. 1981. Fauna SSSR. Nasekomye nigrolineatus (Distant), and its Australian distribution
khobotnye, t. 13, nr. 2. Poluzhestkokrylye semeystva (Hemiptera: Nabidae). J. Aust. ent. Soc. 21: 306.
Nabidae. Leningrad. 327 pp. (In Russian.)
REUTER, O. M. 1908. Bemerkungen iiber Nabiden nebst
Beschreibung neuer Arten. Mém. Soc. ent. Belg. 15:
87-130.
INTRODUCTORY STUDY OF ADVANCED ORIBATE MITES (ACARIDA) :
CRYPTOSTIGMATA : PLANOFISSURAE) AND A REDECSCRIPTION OF
THE ONLY VALID SPECIES OF CONSTRICTOBATES (ORIPODOIDEA)
BY D. C. LEE
Summary
The study of advanced oribate mites (Planofissurae, new name) is introduced as a further part of an
ongoing study of sarcoptiform mites from South Australian surface soils. Morphology is considered
with reference to a unified notation for hysteronotal chaetotaxy, notal pores, the form of leg
trochanters, acetabula and apodemes. Constrictobates lineolatus Balogh and Mahunka is redescribed
from South Australian material, the generic diagnosis 1s modified, Constrictobatinae (Fenicheliidae)
is newly synonymised with Pseudoppiinae (Oribatulidae), and the superfamily Oripodoidea is
considered.
INTRODUCTORY STUDY OF ADVANCED ORIBATE MITES (ACARIDA; CRYPTOSTIGMATA:
PLANOFISSURAK) AND A KEDESCRIPTION OF THE ONLY YALID SPECIES OF
CONSTRICTOBATES (ORIPODOIDEA)
D, C. LEE
LEE, D. ©, 1987, Introdictory study of advanged oribate mites (Acarida> Cryptostigmata:
Planotissurde) and a redeseription of the only valid species of Cunsirictabates (Oripodoidea).
Ree, & Aust Mus. 211): 35-42,
The study of advanced oribate miles (Planotissurae, new name) is intmodticed asa further part
of ant ongoing study of sareoptiform mites from South Australian surface sails, Morphology
is considered with reference to a unified notalion for hysteranotal chaetotaxy, notal pores, the
form OM ee trochanter’, adetabula and apodemes, Constricrabates lineolarus Balogh and Mahunka
isvedesenbed from South Australian material, the generic divgnosis is modified, Constrictobatinae
(Penicheliidaed is newly synonymised with Pseudoppiinae (Oribalulidae), and (he superfamily
Onpodajded is considered.
Bb, C. Lee, South Australian Museum, North Terrace, Adelaide, South Australie 5000, Manuseript
received 2] October (986,
This is a further part of an ongoing study of
sarcoptifarm miles from surface soil sampled from
nine florally diverse South Australian sites. The
primitive oribate mites have been considered
elsewhere (Lee 1981, 1982, 1985), and here the study
of advanced oribate mites {is introduced, The
majority of the oribate mites sampled belong to this
group which, because of morphological changes,
requires a consideration of homology and notavon.
Furthermore, because a new diagnostic character
stale for these miles is recognised, and (he opinion
that they should be unnamed (Lee 1985) is revoked,
they are rediagnosed and dealt with under a new
name (Planotissurae),
The description of the primitive oribate mites
in this sludy has been thorough but time-
consuming. Balogh & Mahunka (1983) sugges!
that ‘painstaking scrutiny, using some recently
discovered features’ is not worth doing for only
some members of a genus, Whilst appreciating this
point, the paucity of a cammon denominator
description is so limiting for many oribate mite
groups thal a more subsranual level had to be
undertaken, but gol to such an extent as in my
previous work. The dorsal and ventral aspects af
the sonta and the shape of the leg segments have
been desvribed, but not the gnathosternum or the
chaelotaxy and lorm of the hairs on the legs.
The first superfamily to be considered is the
Oripodoidea, partly because \t is a diverse and
dominant group within the well-established Poro-
hotae, atid partly because i is not only important
in the study of soil zoology, but some of its
members are intermediate hosts of anoplocephalid
tapeworms, being infected by the cystercerooid
(bladder worm) stage, The most recent work on the
Oripodoidea is by Balogh & Balogh (1984), referring
to it.as the “Oribatuloidea’ as well as excluding the
Mochlozeidae and Parakalummidae, It includes 20
families in the superfamily, of which hall are listed
ag new, The work gives great importance to whether
or nor the hysteronoral foramina are nulliporose,
sacculate or a mixture of both those character states.
The keys and diagnoses use few character’ states,
which for Constrietobatinae (the only family group
so far considered) are in part inaccurate. Despite
this, the work is valuable on the basis for studying
oripodoids,
Constrictobates was selected as an example,
because it is unique within the Oribatuloidea in
having |5 pairs of hysteronotal setae on the adult,
This is only one pair less than in the holotrichous
stale amongst ‘primitive oribate mites and is
therefore valuable in homologizing the hystcronotal
chactotaxy of the two groups.
The South Australian mites examined are
deposited in the South Australian Museum; the
types have been returned to the Hungarian National
Museum.
MORPHOLOGY
flystéronotal chaetotaxy
There are three regularly used notational systems
for ihe hysteronotal chaetotaxy, depending on
whether they have a full complement of either 16,
1§ or 10 pairs of setae, The multiplicity of systems
is based on uncertainties of homology, | introduced
another system (Lee 1981) for the primitive oribate
mites (16-pairs chaetotaxy) with the intention of
applying it 1o all sarcoptiform mites, The chaetotaxy
Ab DC. LEL
of oripodoid mites, for which both the [5-pairs and
10-pairs chaetotaxies huve been used, is
homologized here with the l6-pairs system. as
illustrated (Figs |, 2). Because most advanced
oribate mites have 15 pairs of hysteronotal setae on
the trtanymph, both systems have sometimes been
used for one species, Certainly, with the substantial
vhange in form between. the nymphal and adalr
stages, Any proposals of homology are uncertain,
On the other hand, | consider that the heuristl
advantage of a uniform notation oulweighs the
disadvantage of using an uncertain homology,
Previously (Grandjean 1954, Lee 1984) ir has heen
stated (hat all Planoligsurae (= Circumdehiscentiae)
lack seta J4 (</'1). The loss of J4 (atid often #2 and
J/3) occurs m the most primitive subsection of the
Planofissurae, the Pherenotac, with the exceprion
of the Hermaniellidae where such setae are very
small under the hysteronotal scalp, but there js a
derived reversal of this loss and (he seta (hat is
absent from the 15-pair system of the oripadoid
mites 15 $2,
The possible Joss of setae in handling specimens,
when only a few are available, can make jt uncertain
as to whether a selal base locates a vestigial or a
broken off seta. In either case, it will be regarded
as present in the chactotaxy.
Notal pores
The slit-like pores (A/1-A/6) and the pore leading
to the hysteranotal gland duct (#G/) are treated as
before, but the notation avd signatures for the ureac
porosae, sacculi and reduced sacculi are changed,
There 35 little doubt thal these three slructures
s Z J $ Zz J
oa c2 co te
1 s es s x r x 1
la da te
4, “ie 2 x s xX 2
Im ant aT)
3 s e x 3
\p op ms
4 s . x 4
ha na nt ra i2 "
5 «® a s 4
ps3 ps2 ps! ng 2 my
ve J s 6
FIGURES | AND 2. Eqpivalent norations lor hysteremetal
chaetomaxy fsed in this study (signdtires a) periphery with
capital Tellers lor files and dumibers for evks}, 1, 13-pairs
avstem (e2 ete); 2, 1D-parrs system (fw ele). % = scta absent,
would be present in Tull complemenl of 16 setal pairs.
(samelimes referrec! to as Octotasic organs because
four pairs are often conspicuous on the hystero-
notum) are homologous, but their funerion is
uncertain, They dre olten regarded as respiratory
(Wallwork 1969), but they may be elther respiratory
or glandular (Hammen 1980), Here they are referred
to as Joramina (singular: foramen), and as either
muliporose, sacculate, or uniporose. The term
‘foramen’ has been used for a thin patch bearing
the Infracapitular gland orifice on the anatho-
sternum (Hammen 1983), but this is considered as
the unneeessary commitment of a valuable teenr to
a uivial structure. The comnionest state of ihe
foramina is multiporose, which is regarded as
primitive. The hysteronotal foramina may be
sacculale or uniporose, which are regarded as
derived. Their signature is ‘f° and the foramina are
numbered depending on positinn: lateral
proteronotal (/1), dorsosejugal (F2¢/), laterosejueal
(P24), the four hysteronolal pairs (3, Fa, FS, PG),
a postanal strip (f7), A particular hysteronatal
foramen may be divided into two paris. which are
then giveo the signature suffix of ether ‘a’ or “bt
Form ef leg-segments (particularly the trochanter)
and the acetabular cavity
Mite leg segments are primitively subevlindrical,
with some tapering, and with only moderate
variation in segment size, This is the ease jn
primitive oribate mite groups except that the coxa
is merged inte (he podosaria. Diversification in
shape and sive has been derived in al lease iwo
suborders (Protissurida, Comatida). It is diffieuh
to rigidly categorize the various shapes, but the
derived segment shape is termed pedunculule, The
pedunculale segment has a caput, stalk and
pedestal. The proximal stalk varies in relative size,
as to Whether i! is gradually or abruptly delineated
from the caput and as to whether its axis is
continuous With or at an angle fo that of the caput,
The pedestal may be an inconspicuous lared out
base to the stalk or il may be larger than (he slalk
and capul combined (Fig. 6 — nole (rochantera |
and IJ, state found in all Planelissurae),
The acelabulum is a socket in the coxite (merged
coxa with podosoma). Io most primitive oribate
mites the socket edge is level with the rest of the
prosomati¢ exoskeleton, but there may be a derived
state where (he raised edge partly eficloses {he
trochanter (eg, PAyl/hermunnia euselose ee, 1985:
Fig, 5), In contrast, the tbrochanteral pedestal ol (he
Planofissurae is totally encompassed within an
acetabular cavity (not illustrated since 11s Internal),
which has a proximal ace/ebuluen and an external
aperture, termed the acetabular access. In the case
of legs) and [1, the trochanter is represented mainly
by its pedestal, and is nearly absent externally.
Apodemes extend from the proximal walls of
PLANOFISSURAE MITES, CONSTRICTOBATES
ed
“1
100m
3
FIGURES 3 AND 4. Constrictobates lineolatus, female soma. 3, notum; 4, idiosternum.
acetabular cavities to bases on the prosternal
integument. The full complement of prosternal
apodeme bases is five pairs (I, IL, sejugal, III, 1V).
There is also a pleural apodeme associated with the
sejugal furrow, which merges dorsally with the
bothridium to seta zl, termed the bethridial
apodeme.
The trochanter is illustrated in this paper (Fig. 6)
in order to demonstrate a synapomorphy af the
Planofissurae, but when it is similar to this in later
works it may not be figured, Although hairs of the
legs are not normally considered, they may be
illustrated in some cases in relation ta segment
shape as with the long, flagelliform solenidia on
tubercles or a ventral setae near a flange a femur //,
Measurements
Measurements are in microns (um) and applied
as before (Lee 1981: 201), except that gnathosternal
appendages are not considered. Also, because the
trochantera are either abscured in the soma or have
an angled stalk, they are excluded from the leg
measurements. The length for a leg 1s the total of
the lengths of each of the distal four segments
(femur, genu, tibia, tarsus) disregarding the
pretarsus, and the breadths are the greatest width
(usually the height) of the tibia.
SYSTEMATICS
Section PLANOFISSURAE new name
Diagnosis (Adults)
Comalida. Holosomatina. Gnathosternum with
dicoxal fissure usually present. Leg trochantera
pedunculate with pedestal (on legs I and II
comprising most of segment) encompassed within
acetabular cavity. Tracheae usually opening into
38 DG) BE
a b c
7
FIGURES 5-8. Constrictobates lineolatus, female. 5, integumental striations and structures around setae /2, J3/2Z4,
Sal; 6, right legs I-IV, femora/tarsi — posterior aspect, trochantera — dorsal aspect; 7, various positions of pore and
seta beside left anal shield; 8, proteronotal sensory seta (z2) with capitate (a) and clavate (b) forms.
some acetabular cavities. Leg genu usually less than
quarter volume of tibia. Hysterosomal dehiscence
line circumnotal.
Remarks
The Planofissurae is a well established taxon also
known as the Circumdehiscentiae (Grandjean, 1954)
or the Euoribatida (Balogh & Mahunka, 1979). I
have previously considered it as an unnamed taxon
(Lee 1985: 50), preferring to use the slightly more
extensive Holosomatina (= Brachypylina: Balogh
1972), regarding the fusion together of the coxites
into a single shield in the latter taxon as a more
important evolutionary event. Also, the Plano-
PLANGFPISSURAB MITES. CONSTRICTOBATES "
fissurae has been dilficull ta diagnase because the
form of the hysteronoral dehiscence line is obscure
(unless an exuvial ‘scalp’ is present), the genu is
sometimes large and the dicoxal fissure is sometimes
secondarily lost or masked by a gnathosternal
teetum. The recognition of the errcompassing of the
trochanteral pedestal as a diagnostic charaeter state
makes easy the identificatian of members of the
Planofissurae, and even if the somal integument is
onaque, the external absence of any substantial part
of troehanters | or UW is asceriainable, Therefore,
] now prefer to have this monophyletic taxon as a
named group within the classification, There is a
considerable disparity between its large size (over
100 families) and the size of the Clinofissurae (Lee
1985: 40 — 6 families), the only other section within
the Holosonvarina. Jt has been wiven a new name
to match that of its Sister group and to emphasize
irs new sower rank. The Cireumdehiscentiae is
currently regarded elsewhere as one of seven cohorts
within the Cryptostigmata (Johnston 1982), ar as
the Buoribalida, being one of twa suborders of
Cryplostizmata (Balogh & Mahunka 1979).
The Planofissurae generally includes two major
groups; che Gymnonota (Pycnonaolicae or Aptero-
vastrina) and Poronota (Poronoticae or Piero-
fastrina), Previously | (Lee J985: 49) reintroduced
the use of Pherenotae for same members of the
Gymnonolae, The Gymnonotae |s further divided
here so that the Planolissurae includes the following
five subsections with their characteristic saper-
families: Pherenotae (Gymnoudamaeoidea); Cari-
notae (Liacargidea); Gymnonotae (Oppioidea);
Pliconotae (Tectocepheaidea); Poronotae (Oripo-
doidea). This is intended to undermine a tendency
in past Classifications to tepresenr the evolution of
the Planofissurae as.a linear progressian fram the
primitive Pherenolae to the advanced Poronotae.
it is possible that three distinct lineages or
subsections were derived from the Pherenotae, two
of them monophyletic (Carinotae and Gymno-
nolae), whilst the Pliconotae may have given rise
to the Poronolae, Such a model may prove inade-
quate, but it currently aids the search for alternatives
lo using the tivminonotae as a large, probably
polyphyletic taxon.
Superfamily ORIPODOIDEA Jacor
Onbatuloidea Thor Woolley, 1958; 870 (part).
Oributuloidea Thor: Balogh & Balogh, 1984: 257.
Excentroselcrosae Grandjean: Trave, 1970: 209.
Nominotypical family-group; Oripodinac Jacor,
1925; 277,
Remarks
Dr R.A. Norton (State University af New York,
Syracuse) has pointed out (pers. comm.) that Oripo-
didae is the senior valid name for any family
grouped in the Oribatuloidea: Balogh & Balogh,
1984, The principal of co-ordinatian (Article 36,
current Code of Zoological Nomenclature) requires
that Oripodoidea is the senior available supertamily
name and is valid for the taxon,
The Oripadoidea is ane olf the biggest super-
families in the Cryptostigmata. No clear delineating
diagnosis has been made for the adults. They
usually have small fixed pteromorphs, but these
structures may be either large and hinged or absent,
and even the foramina (diagnostic of the Poranutae)
may be absent, On the other land, immatures may
be diagnosed by being ‘nymphes & microselérites'
(Grandjean 1954) later referred to the ‘Excentro-
sclerosac” (Grandjean 1959), The replacing of
‘Excentrosclerosae’ by a superfamily name was
considered by Grandjean (1959; 473), but he was
cancerned about the similaricy of the included
Mochlezetes ta members of the Ceratozeraidea.
Later, However, he considered (Grandjean 1960) that
Mochlozetidae Grandjean 1960 were similar to
Sellnichitdae Grandjean, 1940 and alsa, Trave (1970)
added Parakalummidae Grandjean, 1934 to the
‘Excentrosclenosac’.
Sublamily PSEUDOPPUNAE Mahunka
Pseudoppitnae Mahunka, 1975: 293,
Constrictobatinae Balogh & Balogh, 1984: 280,
Type-venus! Psevdoppia Mahunka, 975,
Diagnosis (Adults)
Planofissurae. Poronotae, Oripodaidea, Oribaluli-
dae, Proteronotal seta zl shorter than). Lamellae
usually absent (exteption: Cansrric/obates),
Hysteronolum pear-shaped, sejugal furrow
extending anterior to bothridium (around seta 22),
may he evanescent or absent across mid-line.
Hysieronoral seiae in 11-15 pairs (always 67), setose
(may have small cilia), not as Jong as distance
between bases. Two pairs of hypertrophied slittike
pores (4/3, fd) present, Either two, three or four
pairs of stnall multiporose foramina present. Ptero.
morphs, if present, inconspicuous. Intercoxite
apademes transverse, short, not crossing midsternal
line or reaching margin of genital aperture, Genital
shield with two or three pairs of setae (exception;
Phauloppiella with 47Zg).
Remarks
Balogh & Balogh (1984) ignored the part of the
onginal diagnosis stating ‘One pair of very small
4)
irca porosae’ were present and regarded
Constrictobates as Sacewlonotic’, whilst this paper
records three pairs of multiporose foramina (=
arcae porosae), Therefore, Consirictobates should
be srouped in the ‘Poronorie’ Gribarulidae rather
ihan the Penivheltidde. The options were to cither
naintain the Cansfrictohates im the monotypical
Constrictobatiniae, 10 group il in the Pseudoppiinae
or to disregard these subfamilies and place it in the
lacge, diverse Oribaiulinae, Relationships in the
Oribatulidae are poorly understood and the matn-
tenance of many srrall subtamilies is nol advanta-
geous, yet there 18.4 group of venera including small
species with few genial setae, an anterior extension
af the hysteronowwm into the proénanotal region
and many hysteronotal setae In order to cecognise
this group, Pseudopplinae is considered valid and
the Consirictobatinae fts junior synonym. The
delineation of this more extensive concept of
Pseudoppiinae is open to argument, To follow the
weighting by Balogh & Balogh (1984), the number
of genital selae would be regarded as the most
imporiant diagnostic characteristic of the subfamily,
but genera such as Diphanloppia Balogh & Balogh,
1984.and Paraphauloppia Harmer, 1967 with only
two or three pairs of genital setae are excluded,
Whilst Phauloppiella with tour pairs al such setae
is included. This results from a weighting in
preference of the number of hysteronatal setae
because of differences in chaetotaxy (for example
compare Paraphauleppia-3J, 5Z, 28 with Senari-
bule-2/, 6Z, 3S) as. well as total number The
following five valid genera are included in the
Pseudoppiinae: Constrictabutes (sce following);
Phowlonpiella Subias, (977 (Spain-Pm);
Pseudopoia Pérez-Inigo, 1966 (central Spain, east
Pyrenees~Pm); Svaiphaulappia Balogh, 1972
(Bulgaria-Pm), Sevoeribulu Mahunka, 1975
(Senegal-EBe near Pmj. The three genepa
(Pseudoppia, Senorihula, Symphauloppia) origi-
nally making up the Pseudoppilnae are more similar
to each other han tothe distinctive Consirictobates
and Phaulappiella.
Genus Constrictobates Balogh & Mahunke
Consifictobates Balogh & Mahunka, 1966; 559,
Type-species; Consirictobates lineolaivs Balogh &
Mahunka.
Diagnosis (Adults)
Psxeudoppiinae. Hysteronotum with (4 pairs of
setae (S2 absent), 3 pairs of multiporose foranyna
U9, #4, F5), Sejugal furrow continuous ackoss rid-
dorsal line. Lariellae present but Mlattened tseto
zl-72). Mmute, honaantal pleromorph bearing setae
7) and Sl. Three prosiernal apodenre bases (f W,
we,
LEE
sejugal) present, Anterior inargin of genital aperture
and acetabulum IV transversely level. Two pairs al
setae on bath genital (2/2 e) anc anal (2/242) shields,
Legs short (order of decreasing leneth: 1, Vv, 0, 1b
tarsi only about twice gen length, three pretarsal
claws,
Remarks
Constriciabales includes one species previously
recorded only from Western Australia, I] was-estab-
hshed in the Oribatulidae and, after being grouped
in the Feneheliicae by Balogh & Bulogh (1984), ix
wow returned to the Oribatulidae (see Remarks on
Pseudoppiinac), The material from South Australia
is described here and cormpared with type material
lo correct original description inaccuracies and
indicate intraspecific variations. The South
Australian specimens are smaller and usually have
a few character states differing fron) the type series,
but hot it all cases.
Coustricfobates Hineolutus Balogh & Mahunka
{Figs 3-8)
Constriciobates linedlatus Balogh & Matuoka,
1964; S56),
Female
General appearance and measurements: Minute
to small, dull yellow brown, cerotegument incon-
spicuous (sometimes evident as a granular, truns-
lucent layer partly detached around leg bases or
posterior somal inargin), Soimal setae, other than
proteronotal file / and 2, medium length bul fine
and inconspicuous, Somal integument covered in
fine furrows forming 4 reticulate pattern (Fig, S$),
but superticially appearing as longitvdinal striae,
with only limited smooth patches (rostrum, lamella
region anda pletiral strip On ventral shreld Iaterul
(o s¢ia Sal), Idiosomal length 207 (25 ex Tamhoore,
183-217); 213 (1 ex Fernics-McDonald Reserve); 228
(holotype and three paratypes). Appendage lengths
(for 215, ex Tamboore)-i 88, If 72, 111 68, 1V 79;
tibial heights-I 15, U9, WI 10, IV ta,
Prosrerruen: Mentotectum width variable, harrow
(Fig. 4) to broad (—setal base distance 1-1), Firsi
apodeme base conspicuous, beside apodeme bases
some faint external ridges between coxite zones.
Coxisternal setae with seven puirs on flat ntid-venter
(/2 large) and two pairs on lateral tecta (4
pedotectum 1, /#3-discidium),
Proteronotum; Rostral margin tripartite. Lamella
flattened (regardable as cosiula) extends anterior
lo sela z1 and posteriorly notched beiween it anc
circumbothridial ridge, whieh has hyaline flap
ventral 10 seta 22, Three stout setae (/l,/2, al) all
ciliate, conspicuous proximally on abaxial surface
of /], inconspicuous on /2.and 21. Seta /2 finer than
PLANOFISSURAE MITES, CONSTRICTOBATES 4
al, but somerimes similar tn lenge (not as Fig. 3)
Fine furrows restrieted to near seta /2 and shallow
groove bebween setae /2-/2 (Fig, 5), Seta fl atl
antwrion end of shor ridge, Seta s2 inconspicuous,
fing about @ third length of /2, ventral to 22, The
Sensory seta 22 Varies from clavate to capitate (Fig.
8), Wilh cilia im longattidinal files, and often fewer
on globose head of capitate stale, also transverse
profile may be either circular or oval.
Opisthosternum: Genital shield only slightly
anteriorly transposed into region between leg
acetabula, Substantially unpatterned lateral strip on
ventral shield (Fig. 5) abaxial (o sets Sul, Consider-
able yariation in relative positions of seta Sa2 and
pure Sa/ (Wigs 4, 7), with commonest state on South
Australian material being Saf parallel to and hardly
extending anterior to anal aperture, whilst base of
sia Sa2 about level with posterior end of pore (Fig,
Aj. Posterior setal pair (Sa3) 00 protrusion of ventral
whield.
[Tysteranuruenr: Reticuilate pattern of fine furrows
(Fig. 5) divided inte inid-dorsal and two lateral
aones by & more substantial furrow (Pig. 3) which
merges anteriorly with the sejugal furrow, Some
vaniution uy length ef LS pairs of hysteroworal setae,
seta Sl longest and stoutest, sometimes more than
(wice length of 21, Pteromorph minute, sometimes
blunt (Fig. 3}, sometimes pointed, base of seta $1
central or biased. Three pairs of multiporose
foramina small (4 largest), raised rim, ventral
‘blister’ with 7-13 pores clustered in middle (Fig, 5),
foramina JS unusual amongst oribatuloids in
posilioning anteriar not posterior lo seta 25,
Auitcnioe hypertrophied sit-like pore {4/3}
conspictious, posterior pore (476) as long, only
small pare visible from above (Fig. 3).
Leys; Lees short, tongest (1, femur-larsus) 41%
jdiosomal tenth Tarsi particularly shore (all less
than tibia, Fig. 6), thick (breadth more than $0%
length), Tarsus U) with distinctive bulbous base All
femora antecaposterlorly flattened with shallow
ventral flange. no. bearing ventral setae few femur
HW, seta vy, Fig. &). Long flagellium solenidium on
tibia Pand 1. Pretarsal central claw longer and more
robust than lateral claws.
Semel inclusions: Oviposior involuted tube
length, 65 (some 215), three lobes length 20. Bears
iS setuc, subequal in size tO each Other, longer and
Tore robust than proteronotal seta /2, proximal
selue (2) unusually distal im position with lips
reaching bases of distal setae {dg), No eges
observed, One to three boli in cach female, some-
lumes trauslucent and granular (? bacteria), ocea-
sionally unrecognisable tragments of spherical
spores, rarely niulticellular fragments.
Mule
Meusuremiertts atid spertiaposiar (atherwiste as
female): Idiosomal leayrh 193 (25 ex Tamboore.
177-204), Zih (Aelotpe), Spermapositor breadth
santerior-posterior, length shorter) -16, sclerorized
bilobed shield length -13, bearing 12 very small
(shorter than proreronotal seta #2) subequal serac.
Material examined
Twenty-five feroales (N1986193-N1986217) and
twenty-five males (NI986218-N1986242), leaf litter
under banksia shrubs (Bunksia ornate), Tantoore
(35°57'S, 140°29'E), 4.71974, D.C. Lee, Holotype
male dnd three paratype females (Hunparian
Natural Aystory Museum), teal litter under dry
sclerophyll woodland (Eucalypius marginata and
E_ calaphyila), Mi Toolbrunup (34723'S, 118°03'E),
Stirling Ranges, 7.6.1953, G. FE Bornemissza,
Oistriburion
Australia fAa). Western Australia; Stirling
Ranges, dry seleraphyll open-woodland, 13 adults
(types). South Australia; Ferries-McDonald
Reserve, mallee-heath tall opef-shrubland, 1
female/} of 8 » 25 em?; Tamboore, mallee-heath
(all open-serubland, 39 females, 17 males /4 of 8
« 25 em? 469 adults in one 25 cm? sample).
Remarks
Initially, the South Ausiralian maternal was
though to be a sew species. It tras been established
(hat this was because of inaccuraces in the orivinul
description of the only valid species and because
of morphological intraspecitic variation. The South
Austrahan specimens are smaller than the type
gvaterial bur are similar cnough ro be grouped in
& lineolatus,
inportant differences between the specimens and
the original description are as follows:
Proteronotum with shart rostrum (seta /} close to,
rather than well separated from zl); sejugal furrow
completes 15 pairs of hysteronotal setac (/5 and 25
overlooked); three palrs of mulliperose foramina
{Iwo pairs overlooked); two pairs of hypertrophied
stlu-hke pores (4/6 overlooked); hysteronotal seta SI
twice as long and stout as Zl (described as
subequal),
Intraspecific variations anyongst South Augtialian
speclinens (sometimes berween fight and left sides)
ay follows: pore (Saf) and seta (S@2) beside anal
shield moytly as first \lusbraled (Fig. 4), varies lo
three olher positions (Fig, 7) of which one is as
rypes (Fig. 7e); sensory proreronotal seta 32 usually
capitate (Fig, 8a), sometimes clavate as on types
(Fig. 8h); size of @ number of setae varies (72, 52,
Sl), usually stnaller than on types,
ACKNGOW| EDOMENTS
(ami fdebiwd to Mr George Pajak for the drnaenigs (Pies
4-8) andi the Australian Biological Resources Suaely lee
funding his salary. Thanks are also due to Ms Jenni
Thurmer for the diagrams (Figs | and 2) and for setting
up the notation on the other figures, as well as to Dr 8.
Mahunka, Hungary, for arranging the loan of type
material.
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LEE
LEE, D, C, 1981. Sarcoptiformes (Acari) of South
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S. Aust. Mus, 18; 327-359.
LEE, D. C. 1984, 5.16. A preliminary revised classification
for oritbate mites (Acari: Cryptostigmata), Pp. 241-248.
In D. A, Griffiths & C. E. Bowman (Eds), ‘Proc. 6th
Int. Congress of Acarology, Vol. I’. Ellis Horwood,
Chichester: 645 pp,
LEE, D. C. 1985. Sarcoptiformes (Acari) of South
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(Cryptostigmata) with an extensive, unfissured
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MAHUNKA, S. 1975. Neue and interessante Milben aus
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principal Families of the Oribatei (Sarcoptiformes:
Acarina). Ent. News 69: 85~92.
KARIARA VIEWS ON SOME ROCK ENGRAVINGS AT PORT HEDLAND,
WESTERN AUSTRALIA
BY NORMAN B. TINDALE
Summary
This paper describes, through an analysis of rock engravings at Port Hedland in Western Australia,
aspects of Aboriginal economic life and material culture in the area. The carvings were recorded on
a 1953 expedition and interpretations are given based on work done then with Kariara and other
Aboriginal informants in the region.
KARIARA VIEWS ON SOME ROCK ENGRAVINGS AT PORT HEDLAND,
WESTERN AUSTRALIA
NORMAN B. TINDALE
TINDALL FR, NORMA B. 1987. Kuriara views on some rock engravings at Port Hedland, Westerti
Australia, Rec. 8. Aust Adus. 211): 43-39
This paper describes, through an analysis of rock engravings at Port Hedland in Western
Australia, aspects of Aboriginal economic lile and material culture in the area. The carvings
were recorded on a 1953 expedition and interpretations are given based on work done then with
Kariara and arher Aboriginal informants in the region.
Norman & Tindale, Nonorary Researeh Associare, South Australian Museum, Sorllt Terrace,
Adeluide, SA. S000. Manuseripr receivecl 22 Sepleniber 19R6.
‘This paper is an example of ethno-archaeological
salvage rescarch in that it attempts to preserve the
opimans of some Aboriginal people of the Kariara
(Kariera), Ngarla, Kurama, Pandjima and Njanga-
marda wribes of north-western Australia abour
some of the many rock carvings or engravings
present at Port Hedland. These were cut into the
rock by earlier generations of Aborigines whose
ideas appear to have been.sulficiently close to those
still held today that Kariara people of the present
generation are able to make Interpretations of the
intentions behind the ancestral work,
Simple images were eut into the flat bard Lime
rock surtaces which had been eroded during former
higher seas, The subjects cover varjous aspects of
Pre-Kariara people's cultural life, their economy,
some of the animals upon winch they depended far
food, and also some dangerous ones faced during
their endeavours to win a livelihood from. the sea
which confronted them with its relatively great daily
tides. In the time available for study (recreation
intervals over two months), a limmfted number of the
hammered-in outline engravings were copied and
there were same opportunities for discussions about
then with che Kariara and other Aboriginal people
of the wider Hedland area,
To them the old-time engravers were the Minju-
bururu or Axe-people, This we can link with the
Njangamarda term mtinjurure applied to the black
volganic rock used for their own axes,
Two types of hafted stone tools appear among
the envravings, both of which were recdenised by
the Kariara as still in use i 1953, Both are discussed
not only for their modes af use bur also for their
special interest (0 archacolowises, The stone axes are
discussed at some length beth as presumptively
Pleistocene tools and as ones used today. Resin-
hafted discotdal flake stone knives, also still in use
and having a history extending from over 30 000
yeas§ ago until rhe present time, are given particular
attention.
Evidence points to the particular engravings ay
Port Hedland as being post-Mid-Recent in origin,
so that the Pre-Kariara artists might have begun
their work less than 3 500 years ago, It is reasonable
lo view the makers af these engravings as much like
the Kariara of today. Rapid growth of Port Hedland
as a shipping port since 1953 is leading to the loss
of some engravirig sites, and the survivinig Kariara
folk are losing many of their links with the past,
Thus this paper seems warranted.
On 2 May 1953 members of the University of
California at Los Angeles and Adelaide University
Anthrapological Expedition with Protessor J, B.
Birdsell and this wriler, haying concluded anthro-
pomelric and associated anthropological work at
Marble Bar in Western Australia, journeyed by car
to Port Hedland on the west coast to set up further
field stations among the Kariara and other loca!
tIribespeople, At a point near Strelley we searched
a granite knoll tor Aboriginal campsites and imple-
ments but found nothing, save the remains of a
baler stiell near a yery temporary rainwater pool,
already dry. Continuing towards Hedland the
spinifex 7riodia pungens plain gradually yielded
pride of place to beath-like types of shrub. Port
Hedland was reached across a landscape of old
estuaries and salt marsh lakes, many filled with
mangroves, There are also older dune range shore
lines with banks of lime-cemented sand,
The party was met by Mr Harvey Tilbrook,
Native Welfare Officer for the Hedland area who,
after We had settled In, took us to see some
Aboriginal rock carvings on a site about 1.5 km
beyond the then little town and situated on some
of the smoothly-eroded hardened fossil dune banks
10 be seen opposite the sniall local Native Hospital,
as it Was then known,
The consolidated dune beds there are approxi-
mately three metres above the present-day normal
highest tide mark and appear to have been swept
over |n the past by seas from both sides so that they
fa % FE TINDALE
now form flat, smooth-contoured lime-hardened old
coral or dune banks. Usually there are two parallel
lines of them close together, with a depression
between often filled with brown earth. In other
places they appear conjoined asa single line of old
dune, There are Aboriginal deposits in the brown
earth, chiefly of pelecypod shells, identified by the
late Bernard C, Cotton as Tegillarca besalis (Ire-
dale), These are present usually directly on the hard
surface of tbe dune. Where this brown layer has
been eroded away or is absent, the smooth rock
surfaces, extending for nearly 3 km along the line
if these dunes, are covered with engravings made
by Aborigines. Usually they are shown as fines
hammered in with a partly grooved pecking
technique as rhough first marked with a line of
punctures and then connected to make continuous
grooves, These grooves usually are from about 1.3
to 2,0 em in diameter and in general depict the
outlines of the subject, sometimes with additional
lines to indicate decoration. The grooves are not
very deeply cut and often have well-worn edges.
Only in one small area were there any deeply cut,
very [resh-looking ones. which might have been Jong
protected by an earchy covering, Figure 39 could
be an engraving of the kind of tool used by the ald
engravers.
The carvings historically may not be very old and
the efigravings conform to the present contolirs of
the surface, well exemplified in the case of one
particular turtle figure where a slight erosion gurer
has heen incorporated into and affects the outline
of its head. The subjects appear to belong toa pre-
European period since there are neither figures of
ships, nor other designs which could be assessed
as foreign, with one possible exception, a female
figure, described later in this paper,
The existence of the rock engravings at Port
Hedland has been lang known. Herbert Basedow
(1925) showed some human figure carvings, and
Frederick C, G. Rose (1950) depicted others. A
detailed account by Frederick D, McCarthy (1962)
reported on a great frany of these engravings,
working from an archaeological point of ew, and
hus account should be ¢onsuilted for detail, Similar
rock engravings are described in Hester Australian
Museum Special Publication No. 2 (Ride &
Neuman 1964), It is devoted in great part to the
story of Depuch [sland off the coast near Port
Hedland, I, M. Crawford therein illustrates rock
engravings from that island, some of which are
similar to the Hedland ones, but others are styled
as stick figures. Some of the Depuch engravings
represent humans in action, often carrying spears.
Such designs were not generally evident in the area
of the present study, which tend to he of static
figures, I is unfortunate that there were no
Aborigines surviving on Depuch to help Crawford
in his understanding (he engravings, They may not
be contemporaneous with the Port Hedland ones.
A drawing shown by Crawford (1964: Fig. 7)
described as an ‘elaborate opening scene, Hunter’s
Pool’ seems to me to match closely the description
of a dugong-spearing as given independently by my
informants, li shows the tong spears, some barbed,
and the ring of armed men about an already speared
animal. The marine hunting way of life thus appears
to have been miuch the same as today.
THE Port HEDLAND ENGRAVINGS
At first, work on copying the rock markings was
confined to the area near the Native Hospital where
we began anthropometric work and a rough (abula-
tion was made of one area, but circumstances led
(o some random copying of interesting designs
elsewhere, and of ones to which our attention was
directed by Aborigines, The following list (hus is
based only on the majority of the recognisable
designs present in an area roughly estimated as some
50500 m, made irregular by the presence of several
lemporary Aboriginal camps for people attending
the Nalive Hospital, In the list relattve numbers of
carvings could be of some interest as showing Lhe
general range of the various artists’ thoughts:
Boomerang (plain silhouette) (is Fig. 19) 18
Boomerang (decorated) as Fig. 28 ete, 15
Spearrhrower (decorated) (av Fig. 14)
Spearthrower (plain)
Shield (deeorared) (as Fig. 1)
Shield (simple meander figure am it)
Spear tas Fig. 7 etc.)
Snake-like desian (as Fig. 80)
Turtle (as Fig. 73)
Shark (as Wig, 84)
Ray (as Pig. 75)
Hird (as Fiz, 59)
Bird tracks Jas Fig. 66)
Kangaroo or euro track (as Fig. ¥9)
Wallaty track (as Pig. 94)
Man (as Fig. 48)
Bar and lives (as Fig 99
Designs, complex and indeterminshe
das Figs 102, 103, JIL etc)
Axe-sharpening marks
Meander of 2 tines
Meanders of 3 lines
Meanders of 4 lines
Concentric circles (as Fig. 103)
Parallel lines (as Pig- 12)
‘Scraped sticks’ (as Fig, 107)
Circle (crossed) (a4 Fig. 113)
Fat bodies ot fish (as Fig. 76)
Human foot tracks 4-toed
Human foot tracks 3-toed
Human foot tracks 6-rocd
(Fig, 31)
Human foot tracks (indeterminalel
——wuUuvte—Uupe-nsese Seow eos we w—o—te
PORT HEDLAND BOCK ENGRAVINGS 48
As nored. (ne eneraved desiens which ar illus-
tated in this paper were nul all obtained from the
one area near che Native Hospital. Some were
selected from several wider areas within the
township liniits, There was One strange engraving,
that ola female figure (Fig, 132) which appeared
io be ‘foreign’, It is unlike any others | haye seen
from anywhere in Australia, Strange elements in it
include emphasized female breasts with the nipples
duplicated, perhaps implying views from two
directions, It could bea late accession, Publicarian
ol the figure could be considered of dubious value,
but perhaps If shown might lead ta discovery of
(urther data on its origin, The design was clear cut
and the edges of the cits were sharper than in the
pencrality of the other rock carvings, hence there
vould be room for debale as to [ts Uime of making,
Sou anwhiropotogists perhaps would wish to ignore
such w foreigner’ — but it did exist in March 1953
and should be noted, Local opinion asgribes nat
fo Aborigiries, bul la sonicone ota small aroup of
American soldiers belonging toa Navajo specil
unit that was camiped close by during World War
1, [tis the only eneraving considered possibly to
he of nea-Aboriginal design,
KARIAIA COUNTEHY AN OUTLINE OF THEIR Lire
Kariara country extends along the coast from
north of Port Hedland west and south fo beyond
the Yule River, Intand it goes south for at mast 150
km lo Where, in the Yule headwaters, the peaple
meel te Njanial tlbesfolk. For further details of
Uiwir boundaries. see Tindale (1974: 244}
As far hack as Kariara tradilion goes they haye
lived at Port Hedland, and very old men talk about
eatly events Which had come down to them, of a
time before white men had first appeared from the
northeast. Malayan fishermen had been working
along the coast, seeking trepang even hetore the First
white men had arrived. Accarding to local informa-
tion, {he nearest Malayan anchorages had been in
the vicinity of La Grange, particularly at False Cape
Bossut in Karadjan country, This was over 400 km
away to the norih-east. Same Malayan vessels had
ovade visits to fhe Por! Hediand area but there had
been little Kariara contact with them.
In terins of the far earlier postulated arrivals of
men in Australia, down the ancient corridor across
the lowland Sahul Shelf from New Guinea during
nenods of lowered sea level in the Late Pleistocene,
the land of the Kanara was about as remote from
the ancient Asian doorway [o Australia as could be
envisaged. However, according to their own stories
(hey had beer subject to pressure from the eastern
Desert interior. People now kaown to them as the
Njamal had forced their way to rhe sea near Strelley,
thereby cutting the Kariars off from their kindred
folk, the Ngarla, who live along the De Grey River,
Although wow separated they still speak kindred
languages and are alike in not having succumbed
to the taltiatory rites of circuntelsion and sub-
Incision brought 1A by the Njamal and urged on
them with great force by the newcomers. In spite
of jlis pressure both Nyarla and Karlara had
resisted adoption of any of the secret rites common
fo these peWweoomers out Of the Desert to the east.
From a quick reading of the evidence in the present-
day Naciara way of life, [heir social organization
and even thei Janguage reflect their isolation, bur
jt is clear they still have had to defend themselves
apainst invading customs.
Some of these |deas were already old if looked
at in terms of social organization to be seen in some
eastern parts of Australia, The Kariara had adopted,
for example, along with many of the other people
ilving on the western coast, the four-seetion system
of social organization, and had abandoned some
ol phe ideas which continued in the extreme south-
west, s\ich as the still litth-understood kinship
systein that prevailed near Perth. This switch to the
four-section system had been present long enough
for iL ta become locally adapted in many tribes, with
some well-rooted changes, so much so that, even
within the northern and southern hordes of the
Karlara, there were special adaptations, details
unnoticed by Radcliffe-Brown (1930: 208) who had
come to the view that it was indeed the model for
the Four-class system. Actually the model he get up
is really the form of the system practised today by
the adjoining Ngarla tribespeaple. Re that as it may,
the Kariara in the days When white people first
appeared were defending (heir mode of initiating
their youths into adult life without the secrecy of
the rites of (he men of the Njamal and other inland
desert tribes. These desert tribes had adopted the
rite of circumcision as a first mte performed in
secrecy, away from their women, followed by a
further, ultra-secret, second rite of subinecision.
These two initiation practices seem to have spread
\ndependently frorn the east as they found their way
across Ue desert Interior from the north-east The
second rite of sub-Incision has usually caupht up
with the lirst, a6 among the Njanval.
The Kariara were faced with resisting the full
impact of the Njamal tribe ways of initialion,
However, despite their despised position in terms
of their Njamal neighbours, they had been able to
resist (he new ways for some generations. Their
southern kindred, the Indjibandi, had suecumbest
Lo the first or circumcision rite under pressure from
people like the Bailgu who also, coming from the
eas}, had attempted to exert pressure on the coastal
people, The full significance of these fundamental
life differences is well shown by the lines on uhe 1974
iribal map published by this writer
46 N. B. TINDALE
As weapons of defence, the Kariara have only
barbed spears with barbs affixed with resin and/or
with sinew lashings. These are also their principal
hunting weapons. The desert Njamal folk more
often use simple, often slightly swollen-headed
throwing clubs for most of their hunting needs,
spinning them as they attack their prey, reserving
their spears, of which they possess many, for use
in contention and in acts of aggression. Often the
spears are armed with resin-affixed terminal stone-
heads instead of the merely affixed lateral barbs of
the Kariara spears. The Njamal also use a different
spearthrower. On the basis of the engravings on the
Hedland rocks it would seem that the Kariara
inherited inferior types of spears from their Pre-
Kariara ancestors.
Although thus disadvantaged, both the Ngarla
and Kariara have been able to resist adoption of
the dreaded secret rites, retaining the simpler, openly
displayed ['tjilimindi]', or arm-binding, and hair-
plucking ceremonies of their past to mark the
changeover of their youths from boyhood to
manhood. However, their southern kin, the light-
skinned and blond-haired Indjibandi of the
Fortescue River, had been forced, or enticed, in
recent years to accept the rite of circumcision. One
old Kariara man complained bitterly to us that
lately, by making offerings of brides, Indjibandi
men had enticed one or two Kariara youths to
accept the first secret rite, thus depriving the Kariara
of their own men.
Openly displayed as rock engravings at Hedland
are some of the decorative symbols, concentric
circles and spiral designs, held as secret by inland
tribes along with such tools as ['tjimari] or resin-
hafted, discoidal knives as used in their initiatory
operations. The last named, the ¢t/imari, is the
instrument for their circumcision rite and is con-
sidered to be a particularly important secret object
to be produced only at men’s ceremonies. Openly
displayed as the symbols are at Port Hedland, they
cause worry among Njamal, Njangamards, and
Bailgu tribes visitors who had discovered them.
They might be seen by their women. To the Kariara,
however, such engravings remind them merely of
openly used paraphernalia of their own initiation
days. In discussion they suggest that the initiation
rites of the Pre-Kariara artists must also have been
open to viewing by both sexes.
Until the changeover to employment as cattle
station hands, and more recently as gatherers of rare
earth minerals, the Kariara of Port Hedland area
were exploiters of the life they found in the sea. They
had no watercraft, and walking in water up to their
armpits was a limiting factor in their efforts. Fish-
spears, held javelin-like, were used, providing one
of the principal ways of obtaining their fish foods.
Sometimes a man’s javelin-like weapon was linked
to his body by rope made from human hair, one
end tied around his waist and loops caught under
it, ready to be played out as the fish, or other marine
creature, was lanced and, trying to escape, dragged
the fisherman about until it succumbed. Daily tidal
changes enabled the Aborigines to use several other
special ways of fishing. One method involved the
use of special fish-killing weapons of heavy wood,
of boomerang form but biconvex in section. These
could be thrown at their prey, usually one of the
larger fish, even one as large as a shark, or at a
marine mammal, such as porpoise or dugong,
especially at times of high tide. If a cast was
unsuccessful, the weapon could be retrieved readily
at low tide. Nets for catching fish were hard to come
by and in the Kariara area came usually in trade
from the northeast. According to my informants
this was only for lack of local fibres suitable for
string-making. Women’s hair was the only useful
and ready source of fibre for string. An important
way of trapping fish, as substitute for netting them,
was by using teams of men and women to push
walls of beach vegetation through the water sur-
rounding their prey, generally at particular places
in estuaries and channels where there were suitable
coves or banks against which to trap the animals,
THE KARIARA PEOPLE AND THE ROCK ENGRAVINGS
Kariara interest in the rock markings was
apparent when we met our first Aboriginal
informant, an old man named ['Kundjin]. He
immediately recognised an engraving I had copied
(Fig. 92), calling it ['kadarabaga], a whale. Then
he discussed two boomerangs (Fig. 18 and 19),
saying they were ones used as fish-killers, and that
the human figures depicted (Fig. 45 and 48) were
of men; others were of women. He recognised the
saw-shark as [‘irawari], noting that this big fish
came into marine lagoons and water channels at
high tide and could be speared or killed with the
aid of boomerangs. Twelve circles clustered together
(actually 13) were a nest of emu eggs (Fig. 69).
Figure 14 was the engraving of a decorated spear-
thrower ['walbara], of the kind he called ['bilbinj].
With the help of this man and other Kariara it
became possible to interpret and give names to
many of the rock engravings.
Kundjing ascribed the engravings at Port
Hedland to ['Minjubururu], the stone axe people,
who were ‘people like ourselves’. Traditionally the
Minjubururu came from the south, long ago, and
looked for water at Port Hedland. Many died there.
Some Minjubururu went to Depuch Island. He said
that ‘Kariara Aborigines cannot get there now, but
in the afternoon one can see them on the island.
We can’t get to them.’ The last statement was
obviously in his imagination, an allusion to the
PORT HEDLAND ROCK ENGRAVINGS 47
former existence of Aborigines on that island as
evidenced by smoke and other signs of living.
Like the Kariara, the Pre-Kariara artist-folk can
be inferred to have been a shore-dwelling people
who obtained much of their sustenance from the
sea. They were relatively sedentary since the demand
of the rise and fall of the tides, here relatively great,
must have determined their daily round of taking
fish and other foods from the sea and from the tidal
mangrove swamps that surrounded them. They tell
the story of these activities in the engravings they
placed on the surfaces of their seaside dwelling
places. Their artistic attention was focused to a
relatively great extent on their weapons for defence
and on the ones for the gathering of food. This
interest also extended to the most spectacular of the
marine creatures, ranging from whales, kadarabaga,
the dugong [‘njamina], sharks ['ira’dananga], and
even to the fresh-water perch [‘jurda], which they
could take in the Yule River. Other than marine
products, emus and their nests of eggs, and the birds
which they struck down by launching boomerangs
at passing flights, provided ready subjects for their
artistry.
There are interesting touches from which we can
learn, such as the frequent depiction of the fat-
bodies on ray-like fish reflecting a special desire for
fat, similar to the cravings of Central Desert men,
like the Pitjandjara and Ngadajara, for fat bodies
of kangaroos and the fatty meat of dingo pups.
Undoubted similarities are to be seen between
some of the present-day drawings (Figs 115-131),
made for us by Kariara and by kindred tribesmen,
and the rock art figures. Indeed it seems safe to
assert that there has been a continuity of tradition
of some duration, perhaps even for more than a
millenium of past residence there by Kariara-like
people,
COMMENTS ON THE ENGRAVINGS
Figures 1-6 were identified by Kariara informants
as being of shields, or ['jata]. Present-day ones are
often carved from the relatively soft wood of
Brachychiton and may also be from Adansonia
trees. The wavy patterns carved on the engraved
shields are still displayed on both Kariara and
Pandjima jata (see Figs 123 and 128). Woods
suitable for weapons are everywhere hard to get in
Kariara country and one of the first tasks of a newly
initiated young man was to begin a search for
suitable trees from which to take his weapons.
Readily accessible trees in clan territory were all
claimed as potential sources by others, hence he had
to search in remoter parts of the tribal territory for
suitable hard woods for clubs, spears and boome-
rangs, and for lighter woods to make shields.
Disputes were readily aroused when an older man
claimed that this or that tree or shrub had already
been touched, and thus claimed as his own.
Kundjing, our Kariara informant, made a drawing
of a Kariara shield as Fig. 123 and Ngoera, a
Pandjima man, made one of his own people’s
[‘jandijiri] shields from the Hamersley Ranges
(Fig. 128).
Spears of the Kariara that were used in serious
fighting could be armed singly or serially with
stingray barbs and spines of other fish set in resin.
Wounds inflicted with them were considered to be
poisoned, and in their traditions, all things from the
sea were considered dangerous. They have a saying
about this: ["Kutunuru 'kapa 'meida wiriwani] (sea/
from/do not/play with it).
Several kinds of spears were used in fishing, both
in the Yule River and in the tidal estuaries. These
constantly active tidal streams carried deep water
inland because of the high tidal range, and the
spearing of dugong or njamina gave them their
most favoured source of meat. Taking advantage
of these tides, men armed with extra long pole-like
spears watched the water as it flowed up an inlet.
They tested the water flow by throwing leaves on
to it. When the inflow ceased at highest tide they
set nets, called ['parubaru], bracing them with the
poles while men armed with barbed spears awaited
the chance of spearing an entrapped animal. At low
tide they could readily recover and repair their nets.
This description of dugong hunting, developed
from Aboriginal descriptions, was later noticed to
closely fit a rock engraving which was recorded by
Crawford (1964; Fig. 7, p. 59) on Depuch Island off
the Kariara coastline.
Kundjing could see little difference between the
engraved spears shown as Figs 7-13 and the familiar
ones of his own people, for which he could
remember no fewer than seven different names,
chiefly depending on the types of armature. Figures
115-122 are of pencil drawings he made and brought
to me to illustrate his own recollections of and
names of the several Kariara types. Although they
may seem different in our eyes, this may be due to
our interpretation of differences in what he saw as
resemblances.
According to other Kariara helpers the spears
shown in Figs 7-13 were similar to ones they threw,
using spearthrowers as aids in launching them.
Theirs were multibarbed and the barbs were
attachments. The latter could be of stone, shell,
wooden points, or stingray and other fish spines.
The resin ['waruba], derived from spinifex (Triodia),
held the barbs in place. Stingray tail spines were
used as tips for some spears.
Bifacially worked pressure-flaked points called
[‘tjimbila] came, on rare occasions, as traded objects
from the north-east. The informant did not know
where they came from, but they had for long been
AN N. B. TINDALR
=
= SS
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5 }
!
‘ 4}
FIGURES 1-19 Pott Hedland rock engravings. 1-6,
shields, yafay \ length 130 em) 2, not measured; 3, levgrl
81 cm; 4, lemetl Bf cm; 4, lenert 66 cm; 6. 33 em, 7-12,
spears, fd/amiaras 7, ener) 110 orm) 8, leng(h 150 ern; 9-]t,
Not measureds (2. length 9) om; 13, spearheads 14-17,
spearthrowers, weltaru also called Aithin; 14 lenerh
Stem; 15, nor measured, 16, length 7 em: tT, not
ficasured, 18-19, fish-killer wardward: length 53 emand
38 em,
ascribed to ['Waijuparil, a mythical man who also
came from the same easterly direction, A few
Kariara men had placed them on their spears, bul
‘only for show’, When white men first arrived,
appearing from the same direction, they fell heir
to che tame Waijungari, and Kundjing recalled the
beginning of a song abour the pressure-flaked
blades: [Tjimbila:na waijuyari) (repeat).
Linbila blades were not shown on any of the
Pre-Kariara engravings of spears, her¢e there is a
possibility they began to arrive only atter Pre-
Kariara days. A Njangamarda man said that in their
country, further to the northeast, (/mbila points
came regularly in trade. With them came a special
very long spearthrower known as a ['jabalinj], also
called ['nabalin), and illustrated in Figs 129-130,
Tjimbila-armed spears launched by Agabalins were
much dreaded — the reason: ‘One cut and you bled
to death’.
In the days When white men were fighting with
Aborigines, this informant said, the police ordered
all Yimbila, many of them made from inteoduced
bottle glass. to be destroyed, and they confiscated
all spears armed with these Kimberley blades.
li should be noted that in South Australia among
the Jaralde the name Waljungari 1s (hal of a skilled
ancestral spearsman, son of their preat being
Nepele, Waijungari, newly iuittated, escaped with
two of his father’s younger wives and fled to the
heavens, a5 told jo 4 detailed myth (Tindale 1935),
At the present time it is dilficult to imagine any
direct link bel ween these two far-separared uses of
the name Waijungari among people living over
2500 km apart. If proof of substantive
relationships were to emerge, I] would surely imply
that the earlier counterparts of Lhe well-known
Molongge, the Kurangara, and similar travelling
ceremonies, such as chose which accompanied baler
shell omaments for spearthrowers from Cape York
Peninsula to Winton in Queensland, and even as
tar as the Lower Diamantina area, had the power
to carry ideas and words with them. Pearl shells
fromthe vicinity of Broome, similarly, but perhaps
id less organized Fashion, are carried evenas far as
Ooldea (Tindale, in press). Such Inter-tribal
exchanges perhaps have had greater influence ou
cullural shifls and introduction of new ideas than
some have suspected.
Spearthrowers depicted by the Pre-Kariara
people, as in Figs 14-17 were recognised by Kariari
men as being similarly shaped, with a peg orien-
tation the same as in their awn spearthrowers or
welbura. It called for the use of a particular grip
in holding spear and spearthrawer in position for
launching the Weapon, For this the handle end of
the wa/bara is Held between the thumb and index
linger and firmly gripped agains the palni of the
hand by liogers 3, 4and 5. Tips-of the thumb and
index finger support the spearshaft so thar the pew
ol the spearthrower rests firmly in the hole in the
butt of the spear, This may be known as Method
A, th is widely used in southecn Australia, The
method they do not use, but which is widespread
in the northern parts of Australia, involves the
holding of the spearthrower belween the index
finger and the middle one. This requires not only
a different shaped beady for the spearthrower but
also a different orientation of Lhe peg, This second
atip | am calling Method B, The latter is linked in
complex tastion with the use of composite spears
with head, shalt and slightly angled butt piece, cach
juncuion lashed with kangaroo sinews and resin
voaritigs. [ts employment enables the apear user jo
keep his forefinger free to take part in) an action
assisting the development of spin in Lhe spear as
it travels towards its largel, Having had
opportunlties to study (he methods and distribution
ol these spear-projecting ways over much of
Australia, | have a study im preparation giving
support tor this preliminary statement about spear-
throwers and their use. As is well-shown in my
unpublished Jilm on Kimberley blade-makiig and
PORT HEDLAND BOCK ENGRAVINGS dy
se al Moulabulla and the parallel published
description, Tindale (1985), the Method B grip is
presentamoug the Djaru and Kilja vibespeaple and
widely used also in the northern interior of Western
Australia, Figure 124 drawn by Kundjing shows the
Kariara Spearthrower, and Pigs 129-130 by Mekata
of the Njangamarda tribe shows (he long-shafled
ngahalin) they receive by trade trom rhe Mangala,
Who in turn recetve then fron. the Walmadjari
people of the Christmas Creck area, whom they
know as the Tiiwaling folk, Lhe mgabalin) requires
the use of Grip B
Boomerangs as weapons were evidently impor-
tant to the early Pre-Kariara artists as also 10 our
Karivra informants who considered that among the
boomerang engravings they could recognise several
dilferent (ypes similar to ones they use, The main
one used for fighting was then ['wirba], .a
distinguishing feature of which seems to be its
having the end held while throwing 1, shahtly longer
afid less Wide than the forward end, Such a boorier-
ang would be idenlilied by marks the owner placed
PIGURES 20-35, Hedland boomerang engravings, W/rba,
20, leneth 4 cn 21, lengiy §3 chy; 22, length 41 am; 23,
lish-killor, wer ward, leneth 4] om; 24, leneth 48 em: 25,
nol measured; 26, lengli: 53 cn; 27, Cehekillen, waruwens,
length 46 em) 28, left-handed returner, wakundy, lenguh
46 cm 29, length 25 cm; 30, lengli 48 em circular holes
are subsequent; 31, fish-killer, wardéwark, length 3% om,
32, left-handed returner wukundi; 33, lish-killer.
waruwaru, lengitt 43 cm; 34, lett-handed returner,
wakunai, 35, nght-handed returner, length Si om.
on (t, My informant recognised among the Hedland
engravings right-handed wirhe, like Flys 33-and 35,
and left-handed ones, Figs 32 and 34. Men who
went secking a fight might carry an anmlul of werdee
to combat. Figure 20 was suggested as indicating
such a parcel of boomerangs, A comment about
this set was that the left-tanded owner had not
placed his identifying marks on the bundle.
Bird-hunting boormerangs |'wakundi) were made
so they would return to rhe thrower if they missed
their target when thrown into a flock of birds
circling to land, 1 allowed the quick retrieval of
such weapons, Opinion was |hat Figs 28 and 3) were
engravings respectively of left- and right-handed
wokundi,
A special boomerang called |'waruwaru] by the
Kariara was employed in (ishing and could also be
wsed #8 a weapon in close ‘arm to arm” (their
wording) quarrels, The waruwaru, also known as
|'waruku:ndi], was thrown at large fish, such as
sharks, and other animals when scen to the water.
[t had special characteristics and was important in
marine exploifalion, It was thrown fo enter the
water spinning vertically, and was bilaterally
symmetrical, ic. had a biconvex section. Such
boomerang-like weapons were made of the heaviest
of woods go that they would sink and not drill away.
They could be reclaimed when the massive tides fell,
leaving the area dry, Figures 23, 27 and 3] were ones
considered likely to have been intended as represen-
tations of warwwurnu, and Kundjing drew, as Fig.
126, his concept of their own Kariara one, Similar
heavy wooden boomerangs for fishing have been
used as far away as at Porl Lincoln in South
Australia, among the Pangkala tribespeople. In
post-contact time examples made of metal cut-outs
have been used,
Representations of hafted stone axes appear often
umonge the Port Hedland engravings. Perhaps they
were highly treasured objects, Kariara men con-
sidered them good depictions of their own ['bulbul
which they said (in English) were always made of
‘stee] stone’. The present-day Kariara hatchet has
a split withy wrap-around handle bound with
kangaroo leg sinews |'talbara], since dingo sinews
are too short to be Useful. The stone was fixed in
the loop of the haft and the lashings held with
waruba resin which was gathered from spinifex. Tir
obtain this resin the Kariara beat matured clumps
of the grass on a hard surface and sweep up the
resin fakes and winnow. By holding burning twigs
over the dish of flakes they carefully melt the resin
at the lowest possible temperature, A bluish smoke
indicates over-heating, rendering it brittle and
useless. The melted resin 1s gathered up ona stick
to form a ball ready for use,
Kanara axe stone came to them by trade from
whe northeast beyand Njamal country. Bach block
AO N. B. TINDALE
had to be edgeground to make a cuoing edge by
rubbing it on hard rock surface such as wranite, It
happened that Windaru, an aged Njangamarda
than from further forth, who was visiling the
Native Hospital, was with us on ene oceasion and
heard Joseph Birdsell remark on the seeming
shortness of the handle of an ake depicted in one
of the engravings (Fig. 37} that we had just noticed,
The old man’s response, Using his awn name (ur
the axe, was brief and to the point — ‘Heavy
|'mardi], short haridle’. The present-day stone
tomahawks tend to be smaller and have long
handles. Another Niangamarda man of the coastal
country further north said tbat their maedi axes
were made from |'minju'ru:ru], litcrally his ‘black
stane’, which came by way of trade fram the hills.
southeast of [\Jalajala] or Bil Fil Spring. Thus it
may have been traded to Port Hedland from more
than 300 km away, The engravings of axes unfortu-
nately do not fell us much about the work dowe on
the axe heads during their making, and whether or
not the hafting was held in.a groove. The presence
of heavy axes with short handles might suzgest that
more and heavier timbers than grow nearby today
were available locally in Pre-Kariara times.
Figures 40 and 4] were pointed aut to us as
representing resin-hafted diseojdal stone knives
called ['tjimari) or |'jiman). One was depicted as
larger than ones used by the Kariara people today.
Both sexes could use them Jor general purposes as
unifacially knapped knives with resin Handles, Since
Zood chert was not locally available, women often
used instead a girfilarly shaped Cyrena-like bivalve
shell for their food preparation work. The situation
was different arnong the inland tribes to the east
and south, particularly the Njamal, Njangamarda,
Niabali, Indjibandi, Kurama, Pandjima,.and Wan-
man who, using, the stone Nake knives in their secret
rite of male circumcision, have came to regard the
Vimarias one not to be seen by women. [n several
papers, particularly Tindale (1957: 15; [96S: 143;
and L985: 28) I have reported the discovery of the
various Surviving uses for this knife, which is of
Spevial interest because it is recognisable as one of
the principal tools found archaeologically in
horizons as far back in time as.the 30 000 BP era
in eastern Australia, as well as being one of the
dominant tools of the Newer Tasmanian culture, tr
was one of the cutting tools of the Tartangan culture
phase (of Hale & Tindale 1930), There is further
discussion of this (/imari tool in the concluding part
of this paper,
Ie will be noticed that while very many of the rock
engravings are of tems and subjecté of interest
particularly to men, a few relate to implements used
by women. Perhaps there was.someé bids since the
informants did not recognize all of the designs. Lt
happened that they all were males, One engraving
a
Ce hy
—— ee .
39 a0 “4
FIGURES 36-44. Hedland tool-engiwings. 36-39, halted
edge-eround axes. bulby; 40-41, resin-falted discoida!
Make knives, témari; 40, diamever 18 cm; 4t, diumeter 9,3,
of resin handle 10 em; 42, wooded dish for winnawing;
43, beater for opening termire moonds, length, 74.em; 44,
throwing clubs waribu (Nuarla tame), length @f em,
(Fig, 44) was. recognised by them as.a [‘lumba] or
beating-club such as used by women in breaking
into the numerous tall meridionally-oriented
termitaria present on the Yriodia grass-eovered
plains situated a little inland from the coast,
Termites, called [‘maladja], were one of the foods
Bathered by women, using wooden dishes, called
[barduj, to separate termites from the soil by
rocking and winnowing techniques. Figure 42 was
interpreted as an oval-shaped (/ardu with riffles cut
into the surface to assist in the effeets of the
tapping, recking and other manipulations which
help clean the producr and yield a whitish mass of
crushed termites used as food. Such dishes were
used also by men in winnowing resin for their
weapons.
Human figures were not common in |he carvings
of the area near the hospital and did noi seem to
display (he ame considered detail of some other
carvings. Figure 45 was thought by Kariara infor
mants probably to represent a youth. Taking note
of the differences in the arms of the boy the
informants believed tim to be in the stage of
initiation when he was undergoing an arm-binding
rite. His Jef arm was interpreted as being badly
swollen by too much pressitre while the right was
lean and firm-muscled, as was proper, Following
up this Jead we obtained details of the Kariara
inihation. When a youth isaboul 14 or 15 years of
age he is seized as a ['malulu] by a group of men
who became his ['mamia]. In their four-secrion
system of sacial organization, if the boy is a Paljeri
ora Purungu Wis maria are of the opposite moiety
(Karimara and Panaka), who call him [’piruway].
PORT HEDLAND ROCK ENGRAVINGS 51
The men of ihe boys own, the inactive morery,
heeome his ['kayu] and they call him [‘walara),
The mamia take (heir piruwany ona lour through
thet country during which he is told much about
the history and (he names of” places and he also
learns of che responsibilities he will be assuming
asaman, At this.stage the boy wears a small pearl
sfiell ornament ['pira'pira]l, suspended [rom his
neck, and anolher over his pubic area. Messengers
with him ask peuple of other hordes of the (ribe
(met during the maduiw journey) to assemble at an
arranged time and place for dancing displays.
Meanwhile the Kang men remain at home working,
particularly through their women, building up food
supplies against the day when the marmia party
returns and other invired people arrive. Then there
is a ['pundulul, ar feast, with singing. Men seat
themselves in the centre holding the youth (ar
youths if, as t§ usual, more than One faces initiation
at the one time). Women dance around the men on
the otter side of the circle. This is the climax of
their share in the proceedings, Then the sramia
continue ona second stage in thelr travels.
The youth during further travel has to underga
the beginning of the |'tjilimindi] or arm-binding
ordeal. During this ceremony the men who are his
maria sitip.
wawila ton? kulbarba tanai
winding blood swelling
(Tyilimindt minds
Armbands
Kulbarta
swelling
tana:
Suings made from the fur of the ['wadjiwara],
or kiipgaroo tat, are wound lightly around both of
(he boy’s upper arms and are left for ‘six days’ and
then taken aff. If the arnig sWell too much the
bindings are slackened a little, During this stage he
is a |'wamuli), His hands become numb but he is
compelled (6 keep travelling on’. He is not allowed
16 swim. When they come to water the youth is
carried across it, for the arm bands must not be
allowed to become wel. When these bands are
removed his muscles are supposed to be small and
solid and he is se strong than he can throw a
boomerang very tar, The youth is painted with red
ochre and is shown the secret of the bullroarer
|‘kulisdi]. He is now |'maijaga), a man, He makes
a triumphant return to his people.
In the main there was litle comment [rom ibe
Kariara about the other human figures. Figure 46
was considered to be that of a woman and 4 snake,
It was a visiting Njamal man who expressed the
view thal the artist had giver an action picture of
a ['kolibiri] or poisonous snake attacking a woman.
This chances to be one of the rare Pre-Kariara
carvings in which the engraving is not of a static
situation bur of 4 real life one. Aimong the other
FIGURES 45-57, Hedland engravings of humans, 48,
man, considered a youth, undergoing the t/ilimindi arm
hinding rite daring initiation; 46, Woman, mand and
kolihiru, @ paisonous snake: 47, woman; 48, man,
muliaeza; 49, girl ohild, 80, hand, mare, 51-57, foorprints,
t]tnet,
carvings Fin. 49 was believed to be that of a female
child, The genital area happened to be carved
around. a pre-existing hole in the rock,
(n the human figures and in the buman tracks
the depicted hands and feet have fingers und toes
shown with much Variation as to number, ranging
from four to six, The Kariara had no explanation
for these variations. In the case of footprints there
seemed to have been no preference in depicting
either right or left foot imprints. From the way the
Kariara discussed the depicted foot tracks | gained
the impression that they thought each of the
engraved footprints was registering something
special about some individual, recording peculiari-
lies of gait and toe position that were important
as aids to identification of individuals which could
affect their own daily lives. What mother-in-law
would willingly place ber own track in juxtaposition
to some registering movements of her daughter's
husband, with whom she could never openly meet?
One human and interesting point came out. Some-
times a distinctly shuffling gait, destroying the
legibility of footprints, was a precaution taken as
it woman neared a ‘dangerous place’, one where
such kin were likely o have left visible tracks,
The Pre-Kariara artists depicted birds, usually ul
the larger and more edible species, and their eps,
1} received some help in identification, Figure 58
showed a crane ["jaingaral, and its tracks were
considered to be as shown in Fig, 62, while similar
tracks, Figs 61, 64, 65, were thought to be those of
ibis, more than one species of which live in the
swamps, Owls, of which we found two engravings,
are in some fashion linked with present-day stories,
bul I failed to follow wp because of pressure of other
recording. Kundjing, our first Kartura informant,
revopnized the subjects of Fig, 66 a9 u set of the
eggs of the emu [‘tjanguna]. In egg-laying season
the emus were nol molested, but (he eges were
taken, At other times of the year emu and kangaroo
were hunted, using fire to drive the animals toward
strategically placed and concealed spearsmeén, The
fe Bh 71
FIGURES S872, Bird cnhgravines, 58, crane, jungure,
Tenth 6 em; 59. owl, height 58 em; 60, owl, length 28 em;
61, ibis track, lengif 10 cm; 62, crane, line of four tracks,
each about 20 cm; 63, 25 em lone, fol recognised, 64,
ibis tracks, eroup of four each, 7 em: 65, ibis, part of line
of (ey imprints! 66, emu tracks, /jangurc, set of four,
length 95 cm, showing pecked harnmerings; 67, duck eggs,
diameter of setting 28 em; 68, emu eggs, nes) ser of fifteen;
A9, emu eggs, thirteen, diameter of nest space about 50
tm; 70, emo track, length. 1K em; 71, emu tracks ser of
two, 25 em apart.
52 N. B. TINDALE
eges shown tn Pig, 67, occupying a space some 30
vem in diameter, were believed to be those of ducks,
but | did not follow the discussion as to which kind
they might be, Emu tracks are commonly depicted
(Pigs 66, 70 and 71) and one solitary track, Fig. 72,
was [hought to be that of the bustard |'ranguritjif.
The neighbouring Njamal people, who had access
to better fibres for string, trapped the birds, making
nets 16 enclose native plum trees, the fruit of whieh
Were very attractive to ihe birds. The Kariara were
never able to do (his netting for lavk of the
necessary string for making nets.
All food animals played important roles in the
thoughts and lives of the early artists of Port
Hedland, We chanced not to copy any engraving
of che large nionitor lizards which were commonly
depicted, OF |he smaller lizards there were seemingly
very few representations ainong the varvings. Sueh
vreatures ioday are chiefly gathered hy women and
children hence were perhaps not subjects for men’s
(thoughts,
AL all times these lesser animals provide a sub-
stantial part of the Kariara children’s diet. With
reference to the larger lizards, it should he
mentioned (hat there 1s 4 carved wooden club |i
the South Australian Museum collection, specimen
No, A 46268, received wilhoul data other than a
general indication of having come from Western
Australia. Ir came in 1964 from the estate ol Mr
H, Savage, Figures carved on its shaft include one
of a monitor lizard, two human figures, and also
one of an owl, one thal is virtually identical with
Pig. 59. These are so sinilar to the rock engravings
being described here that it is diffieult 10 believe
that the club could have been carved by other (han
a Kariara mun, one personally familiar wilh the very
engravinus present at Port Hedland itsell. The wark
was done on the hardwood surface of the club, (irst
by incision, and then finished by a burning tech-
nigue. Perhaps this club should be ragarded as
providing an indication (hal (he Hedland carved
designs include some thal are not very old, or
alternatively that the old artistic lradition continued
unlil @ time as recenl as a generation or two ago.
Snakes were lrequently depicted by the Pre-
Kariara artist, The supposed action picture shown
in Fig. 46 has already been discussed. Figure 79 was
interpreted as that of a python called |’ maniara] and
deseribed as possessing a striped pattern and a black
head, This type lives in swampy.areas, There was
another python known as ['palgumaral] living in
rocky places around permanent sources of fresh
water,
Turtle figures Were commonly engraved. Figure
73 shows one of normal size. [lL should be noted
that, jn general, designs adhere to expected sizes,
however, Fig, 74 does depict a very large turtle,
measuring lar larger than usual. Although executed
PORT HEDLAND ROCK ENGRAVINGS 33
—_—5
/
cate on
* r
jj Mee = |
ay we |
—./ ——e |
oo a 2 -—
#
82 ¢
—"
[\
|
lls
FIGURES 73683. Reptile and marine-animal engravings
al Port Hedland, 73, turtle, satjaroka, overall length
83 cm; 74, turtle, large, length 1-65 mz 75, stingray, bocly
length 58 em; 76, fal body of ray 13 crm; 77, stingray, body
length Gt em; 78, fish, 60 cm; 79, python, AlaAiara,
Idendlied by its banding and black on bead, length 3.35 m;
80, snake, nol identified, length 94 cn emerging from hole;
81, fish, not identified, length 80cm; 82, marine mammal,
length of body 61 em; 83, shark, maritjawarin of Nyarla
tribe, length 2.4 m, supposedly benign.
by artists not in any way linked to each other, these
two turtle figures are remarkably similar in execu-
tion. Both were considered to be |'tjatja'roka] by
Niibiri, a man of the adjoining Negarla tribe, He
said that his own people were skilled fishermen, able
10 sWim Up to a turtle, take hold of the shell at its
neck, overturn the animal, and then wrestle with
it in a swim back to shore.
Kariara identification of some marine creatures
Was fot always positive. Recent generations largely
have had to abandon the gathering of marine foods
in favour of work with Europeans. But some details
they remember seern important, They appreciate the
value slingrays had, both for the early artists and
for their own activities, These and other fish
provided them with spines for spear lip armatures,
the serrated tailspine is still used for various cutting
purposes, While their fat is much sought after as
food and for rubbing on their bodies.as protection
avainst the sun and for decorative purposes,
Kundjing drew particular attention Lo the picturing
of (hese internal fat bodies as if they were externally
visible, and to other examples engraved separately
on a large seale, as in Fig. 76. The Port Hedland
artists, when drawing fish and other marine
animals, seemed habitually to show both eyes. An
exceplion is in the unidentified fish seen in Fig. 81
where an unusual side view of the creature is given.
Figure 82 appears to represent a pinniped
mammal and it is possible that the engraving
records the existence of fur seals in the Hedland area
during the times of the Pre-Kariara people, Figure
83 was considered by a Ngarla nian (o be a small
and benign shark, good for eating, and he named
it as [‘Maritja'warin], The same edible quality was
ascribed to the saw-shark shown in Fig. 84. Inci-
dentally, in the Fitzroy River, further north, such
saw-sharks during flood seasons may go far inland
for nearly 200 km, and may become permanent
inhabitants of the large freshwater lagoons often
present there. One such saw-shark was shot by
Joseph Birdsell while on a hunting trip during our
stay at Liveringa Station. Figures 85 and &6 possibly
FIGURES 84-92. Fish and marine mammals. 84, saw-
shark, irawart, lengily 160 em; 85-86, fish, Australian
salmon-like, lengths of both 106 cm); 87, fish, gurnard-
like, 76 cm; 88, not identified, leneth 160 em; 89, unidenti-
fied, perhaps shark, lenge 100 em; 90, Dolphin-like,
length 335 em; 9), doubtful, ‘saltwater fish" of Jabili, body
length 86 cm; 92, whale, Auderabava of Kundjing, the
purvkulana of the Ngarla, lengih 2.8 m, kangaroo tracks
in place for eyes are a subsequenr addition.
Sq NOB TINDALE
represent a salrhonelike fish of large size, and Fig,
87, unidentilied, seems fo fave long pectoral fins
much like a gurnard. Descriptions ol dolphins, for
which they could not at the time recall the Kariara
name, applied to Fig. 90, while Fig, 91 was of a fish,
perhaps @ hexanthid shark. Figure 92 represents the
whale engraving which Kundjing spoke about when
we had our first conversation with him on the Port
Hedland carvings. He called i) Aadarahage in bis
southern Kariara dialect, and other men afterwards
used [‘kararabuka}, the Port Hedland version of the
same name, Ii will be noticed that in the whale
engraving a pair of kangaroo foot tracks were cut
over the area where eyes origittally might have been
present. Younger meér expressed regrets rhat feasts
of stranded whale had not taken place during their
time, Evidently older people had fond memories of
such feasts.
Kangaroo tracks frequently were the subjects of
the old artists, often, as in Fig. 93, emphasizing the
deep impressions of the hind legs made in places
favourable for tegistering such prints. Figure 94
depicts another set of prints in which the forepaws
suggest a moment of slow feeding (at such a time
the animal would bea better target). In the latter
example there was a human footprint carving and
also a long spear-like groove 2.8 m long. The juxta-
position of these three elements (may suggest that
they had been placed there to record a hunting
episode but, as emphasized elsewhere in this report,
such combinadoas implying action are rare as
compared with isolated static Images.
Entirely dilferen| are the impressians on the
ground representing human beings, as in Figs 95
and 96. They both register the Imprint made by a
squatting or seated person, ‘The first represents a
man, and the samewhal ore formal one happens
to indicate # woman, as registered by the digging
stick Which is shown in her lap, Figure 98 was
interpreted as a scated maw with bird down or
feather ornamentation fastened to his body with
blood, as might he the case when he was decorated
tor a dancing display. There was dispute as to
Fig. 97 which was carved much too large to be a
normal-sized inan’s sitting imprint,
For lack of opportunities some of the smaller
drawings were not identified, bul more than one
informarit sugested that Fig, 99 represented a fur
string genital cover, cither of the fur of the kangaroo
fat or of the opossum, ‘This type of pubic cover was
used by Kariara persons of both sexes, especially
when Mies were tiove irritating than usval,
The original of Pye. 100 was thought by Rundjing
ro represen! @ necklace made of jleces of shell stuck
amo a hair string base using resin. Later, when
working with Jammi, @ Kurami man from the
Hamersley Ranges, far inland, we learned (hal
during secret ceremonies initiated men there often
*-e _
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E \ ne fa!
» Mf ly
“Ls, Ww we 14 a
FIGURES 93-114. Lmpressions, decorative designs, anc
symbols, 93-94, Kangaroo, werineura, Tool (racks; 95,
buttock imprint of man (small carving, width only 23 em);
96, woman with digging stick (small carving, width only
13 cm);97, not recognised, 90 em Jong; 98, bullock pri
of feather-decorated man, width 20 em; 99, fur strinu
modesty shield, width 23 em; 100, teekler, lengrh 38 ems
11, much debated shield or skin design, length 84 cin,
102-103, body designs; 04105, apifal and cireular designs
representing totenvie homes or places, dlameter 25 enn, 1h,
notrecagniseds 107, whitewood ornament, length 23 cm;
108, body design, lengeh 30cm, 109, lizard, Jongth 25 cre;
1L0- 113, Hot reeoyniseds Id shield with mand murks,
length 9S erm and circular (otemic design myertry minnie ts
newly mcised Ky Jahili (white tame Dick) of Njangamarda
tribe, on (9 May 1953,
wore wecklets made of pieces of pearl shell fastened
to a fur string banc, or to a bair string choker, by
being affixed with porcupine grass resin. The
Kuramia tect for the necklace was |'tjitjarliri} and
the pearl shell came to them in trade from the
Ngaluma (ribespeople of the Roehourne area by
way of the Indjibandi, The Kurama, who in earhee
days had no knowledge of the sea, gave mulga wood
speats in exchange for the shell ornaments. Good
quality mules (Acacia uneura) country oceurred in
their tribal ateas and continued south for a grear
distance, Their mulga shrubs, called ['kuritjarda],
supplied spear shafts which were better than thase
avallable to the Indjibandi. Thus there always was
a demand lor Kurama spears.
MOC) HeDLAND ROCK ENGRAVINGS
Figure WTowas much disputed, Ta same it was
a shield. dts dimensions were much the same as
those of ther own shields. but oo one made then
with [inant] marks like the ones decorating this ane
Later, When we were in Halls Creek, an old man
named Neepal of the Whadjuk tribe Thom near
Perth, on being shown my drawing, had his own
Thoughts about i Me had been a lifelong resident
in the northwest of Australia, following extradition
for some long-forgetten youthful offence against
while men. He elumed that marks on the strange
shield-like carving were similar to the designs on
the inner side of deeorated skin cloaks such as those
worn by his people when he was a boy in his own
country. Virtually forgotten, Ngepal in 1953
probably was the last living person of Full descent
from (he Whadjuk tribe of Perth, He provided me
with a vovabulary and other details lor study
(Tindale 1974 260),
Inthe engravings ac Port Hedlanct che early artists
frequently used concentric ereles and spiral designs
similar (o those shown in Figs 14 and 105. To the
Kariara Chey were ceremonial pallens, known as
['kuri], painted on the ground and on objects tor
display iu various ways during dances in which both
sexes could take part, Visiting members of the
adjoining inland tribes used similar designs, which
were somelimes quite elaborate, bul only for secret
decorations displayed during initiation ceremonies
from which women were excluded, Such visiting
tribesmen did not appreciate the open display of
the markings on the rocks al Port Hedland, and
they feared that their women might see then
Figure 107 was thought to be an engraving of a
| Mayeur'mangar}, or scraped stick ornament, as
made and used by the Kariara when decorating thelr
hair or bodies to take part in displays of dancing
before their women and children, The Kariara had
no sectel ceremonies. These scraped white wood
sucks ‘looked like flowers’, They were mace by
scraping masses of shavings frony sticks of the
Alalave shrub which haya very white wood, Masses
of the soft curled shayines became part of wip-like
hair dressings Wery offen the shavings were left
attached (4 8 tapered base stick as they dried they
curled up, givitig the look of a devorated hairpin.
Ip the sawie manner the shavings still attached to
slicks could become other forms of decorated
appurtenances. A Ngarla man, who said his peaple
also used these manggar, supplied a yarety of
nancy lor them according 10 their particular uses.
A fully dressed Ngarladanger might wear wp to four
short decorated pins called ['tjural in his hair with
4 longer one |'katjiri] thrust laterally through a
rolled or gathered bundle of hair un the back of
his head, Stull other such ornaments, called
[ihitbiri], would be worn around his arms just abave
the elbow. with one or more, knows as ['wambi],
oar)
aT
thrnst undera bell around tis waist. In dancinu he
ought flourish several longer sdeks with gatherings
of shavings at intervals along thew length, These
were called |'piluj, Depending on the subject of his
perlomance, he might catry a boormerane or a
spear in one hand and a [‘karubinj], or shield, in
the orhen
Rarly in our stay t had been copying engravines
near the hospital wher | had a visivwr, an old man
named Jablli from Wallal, a Npaneamarda of the
group living near the sea, Re sal and watehed for
awhile and then volunteered sane ideud fications,
claiming that he vould ‘read them all’, He (hought
he was still in Ngarla country atid said that people
in his own country alsa made such marks, He
recognised both emu and ibis tracks. Nearby, as it
happened, there was a design (shown herein as
Fig, 108). This design and similar marks he claimed
were placed on the bodies of dancers, and lie began
to demunstrate, rubbing some limestone an the
rock, wetting it with saliva, and painting iron his
chest with his index finger. Pointing at a drawing
I had already on my card, le identified Pig. 103 as
a man's body wilh the same kind of design he had
placed on his own chest. Il seemed that these might
be considered us personal identification marks. We
talked of shiclds and he said that his people’s shields
always had ['maini or ['mani) marks on them, and
with a piece of stone he lightly cut parallel vivzag
lines on the rock, T made a sketch of this modern
addition lo the engravings, as Fig, 114, We had been
discussing orher carvings when | noticed that also
he had cut. circle within circle design, Perhaps ii
was an absent-minded cutting of a ceremonial
mark, He called it |'‘jorinj mimbura] for which 1
was Unable ta get a translation, He was returning
10 Wallal and unfortunately 1 Was nor able tu see
hiny again,
ILUISTRATIVE DRawives By My INFORMANTS
in my discussions with the Abongines who
helped me to undersiand their ideas about the Port
Hedlatid engravings we covered other aspects of
their life nol enlarged upon on in this brief study.
More than one of them who had to actual model
or ntifacts ro show, chose to use drawings to
illustrate their thoughts abour the subjects we had
heen discussing. A few of these drawings are
included jy this paper as Figs 115-131, Others have
been used as part of the backeround data for this
paper, They could warrant a lurther paper on
Various ways of lite along the coastal country of
Weslern Australia, togelherwith results of enquiries
into kinship, language, and story details gleaned as
avcessory parts of the fieldwork plan on our 195%
expedition.
56 N.. B, TINDALE
5 116 7
FIGURES 115-122. Drawings of Karian spears by
Kundjing in illustration of his discussion aboul Port
Hedland engravings. 115-16, called Aoruparta; 117, karta;
118, walekari; 119, (jaran; 120, winda, 121, wadekutare:
122, woeljin
Figures 115-122 illustrate the thoughts of
Kundjing in support of his remarks about the
similarities between the Pre-Kariata spears and
those of the Kariara tribe. They are named:
Pig. US ‘karuparta Fig. 119 'tjaran
Fig, 116 ‘karuparta Fig, 120 ‘winda
Pia. 117 ‘karte Fig. 121 ‘wadakutara
Fig. 118 'walakari Fig. 122 ‘wadjir
He pictures the harpoon-like spear in Fig. 122 as
having a wooden barb integral with the shaft, At
first he could not recall its name, wad/ir, It had a
knob at the butt end to help in securing a human
hair rope-line, the other end of which was tied about
the lisherman's waist. Ln his drawings the butts of
the other weapons were marked, somewhat out of
proportion, to indicate that the butts had holes into
which spearthrower pegs would fit.
Figures 123-127 were drawn also by Kundjing to
illustrate other artifacts of the Kariara, including
their jata shield, their wa/bara spearthrower, used
with the grip between thumb and forefinger
(Method A), their club ealled ['pilbin], used in close
fighting, and in Pig. 126, the waruwarn boourerang
for fish killing, which he stressed was cut from the
heaviest available hardwood. A group of three
boomerangs (Fig. 127) shows the several shapes
preferred for their returning ones, the wakundi.
Another Aboriginal illustrator, Nguno, an ini-
tiated man of the Pandjima tribe of the Hamersley.
Ranges, provided two drawings for (his paper. His
people were leaders in the spreading of the secret
rites of initiation for their young men, and
according to their traditions, they lived so lar to the
east in olden days that they had no knowledge of
the existence of the sea. Figure 131 gives his
impression of a newly initiated ['malbu] or
Pandjima man. Figure 128 pictures their shield,
called [‘jandijiri], which shows a very close
resemblance to the engraved ones on the rocks at
Port Hedland,
Mekata, a member of the Njangumarda Kundal
tribe, about 200 km east of the Kariara country,
took a leading part in some of our discussions and
provided two drawings illustrating long spear-
throwers, known as [jabalinj], shown here iu
Figs 129-130. These are used in the throwing ol
124
FIGURES 123-131, Further illustrations by Aborigines.
123-127, drawn by Kundjing, of Kariara objects; 123,
shield, jaca; 124, spearthrower, walbara; (25, club, pilbin
for close fighting; 126, boomerang for fishing, waruwaru,
also used while standing up ‘arm in arm’; 127, outlines
Of three wekundi returning boomerangs; 128, shield,
Jandijiri of Pandjima tribe of Hamersley Ranges by man
Ngoera; 129-30, long spearthrowers ngabalinj traded Lo
Njangamarda tribe through Mangala from Walmadjari
people whom they call ‘Tyiwaling; they are tised with the
index and middle finger grip; 131, concept by Ngoera of
a newly initiated Pandjima ma/bu or man.
PORT HEDLAND ROCK ENGRAVINGS
FIGURE 132. Srrange figure, nor recognised by the
present-day Kariara — possibly of foreign origin.
spears While employing (he grip (my Method B) in
which the spearthrower is held between the [ore-
and middle lingers. These are obtained by trade
from the Mangala tribespeople who in turn were
said to get them from the Tyiwaling people still
further east, Later al Chrisimas Creek Station we
learned that Tjiwaling was a western name for the
Walroadjari who were in contact with the Kitja and
the Djaru of the Hall Creek area. Thus there were
indications of a trade route for ngabalin] of about
900 kim, This seems to have provided a, route by
which cultural novelties came to the Karjara,
CONCLUSIONS
There are few direct clues as to the age of the Port
Hedland rock carvings. Present-day Kariara Abori-
vines acvep! them as being made by people similar
to themselves and consider that they understand
Their meanings and che motives intlueneing the
earlier aruists who made them. Kundjing, one of
the oldest living Kariara men, recalled a tradition
o! people like themselves coming from the south
und making some of the engravings.
Geologically (he situation of the freshly planed
ol? rock surfaces on which Lhe engravings are
concentrated, standing as they do only Lhree (o four
metres above the present highest tde marks of the
bay, yields a limiting date falling near the end of
the latest Peronian eustatic higher sew level at
nt
~
around 3 500 to 3 700 BP, The relative absence o!
signs of aerial erosion tends-alsa to place a further
limit on the age of the carvings. The Kariara
recounise the principal types of their own weapons
as being present among the engravings, and they
have ready explanations for the meanings of the
designs, The presence of engravings of hafted edge-
ground stone axes is important. Mulvaney (1975:
194) concluded that there may have been some
season for a gap between the older Pleistocene stone
axes of the tropical paris of Australia and the far
younger basaltic rock types which appear se
commonly in much of Australia front Mid-Recent
time on. In parts of Australia where stitaless honey
bees ovcur, the present-day axes are consianily being
employed in the chopping outof thew nests in the
hollows of trees. In the Kimberleys of Western
Australia both carlicr and later axe types have beer
found, Mulyaney (1975: 147) shows a photograph
ofa peeked, grooved, older type axe from Stonewall
Creek, and another well-worked one was found by
us (Tindale: 1981: 1772, Fig. 5, right) at Moolabulla,
near Halls Creck, with similar groove on a quartzite
core, Aller some use the Moolabulla one had been
adapted by knapping to form » chopping tool of
the type considered to be characterisne of Karran
limes in the earlier ball ol the Wisconsinan or Last
Ice Age,
In earlier paragraphs references were made to the
i/imari knife ag present among the engravings at
the Port, Figure 133 shows a living culture Njanga-
marda example obtained by J. B. Birdsell and P J
Epling during an curly stage of the lieldwark at
Marble Bar. lt anay be considered very characteristic
of ones widely known today from the western part
of Australia. In some (ribes (here (jimari are still
in general use and in others they are kept, m secret,
away from. wooten, and used only in. their initiadon
rites for young men. The discoidal form of such
flake Knives becomes well developed in use and
generally as the resull of one or more retrimmings:
The knapped paris exiend around aboul one-half
of ihe perimeter, After retrimming, some of them
are developed with @ partly coneave margin when
needed for trimming such objects as shafts of spears
and those of other weapons such as clubs. The back
of the knife, covered by the resin handle from the
very conception af the flake as a tool is offen
enurely udtrimmed and ioay be untidily iregular,
which may be useful in strengthening the bond
between handle and knife. Liarchaealogical speci-
tens this contrast of uirinimed and worked parts
leads rather readily to their identilicanon as yenaert,
as does also the (requent presence of rollet-pressure
fine-trimming preserved on the cutting edge af the
knife (Tindale 965)
In eastern Australia. particularly in the riverine
carridor lying west of the Great Dividing Range,
Sk NB TINDALE
FIGURE 193. Kesin-hafted unifacially worked flake knife.
tiimary, ot erypto-crystalline gray chert from Karduma
Hill, cast of Warrawagine, received from Mudjing of the
Njangamarde tribe, May 1953, diameter 77 em.
the use of “imuri-like tools was replaced un the Mid-
Recent by other tools, ones of the so-called Micro-
lithic Culture Phase Which appeared suddenly, full-
fledged, and spread widely just after the Mid-
Recent, rather quickly in the east and more slowly
across the desen interior to the west, Hale & Tindale
(1930) found the original evidence for the change-
Over int Miineralised beds at Tartanga, on che Lower
Murray River, an Aboriginally-named midstream
island bank, Hale & Tindale selected the name to
represent a cultural phase older and geologically
separate from a great depth of Aboriginal occupa-
tional debris, recording three named phases in an
adjoining rock shelter, them Known as Devort
Downs, but now correctly recognised by its Abari-
ginal ame, Neatngaut,
At Vartanga the change-over in stone tools is
ecologically recorded from older and larger flake
ones to the newer microliths, which appear only
above a waterlaid sterile blue-black clay horizon
(thet F) of relatively great thickness, now koown
to be the silt laver deposited during the many
centumes of the Peronian high sea levels between
6 000 and 3 500 BP, The few tools found in the
post-Peronian beds above it matched [hose found
it rhe three named phases found in the sx meue
fill of Oeeupational deposit in| Neautngaut rock-
shelter. Thus it happened to be that it was the latest
phase of the Tartangan culture phase that Was
pamied. Its tools were shown in detail (Hale and
Tindale 1930: 149, Fig. 4) and from the geological
section (their Fig, 4), it is clear that the separation
from the microlithic tools of the Upper Beds G (o | is
real and complete, From (he begining if was recog
oised that the Tartangan type sile tools were in dn
area Where stone for tools was not ecadily available
and other richer sites were soon of record in the
South East of South Australia and elsewhere, After
interest in Australian archaeology developed else
Where it unfortunately happened that misinterpreta-
von ol the Tartangan data as published, led to che
ignoring of the clear-cut geological evidence, This
seems in part 10 have been dhe to an older view that
Australian stone implements could not be typed and
that all kinds were present only by chanee, There
was little appreciation of the Jong time-span of the
occupation of Australia, Today it is clear that the
Tartangan stone tools can be recognised as domi-
nating the Australian cullure scene from as early
as the Pleistocene occupational horizons of the
30 000 BP era at Lake Mungo, in New Sourh Wales
down to around 6 000 BP in eastern Australia, with
survivals of some of its tools in a slowly progressing
transitional phase, spreading towards Lhe west and
leading. to the deminance of the Microliifr tool type
still in progress in the far west. Thus the presence
of the Vimar’ as the key too! of the long-lasting.
Tartangan phase in Australia and Tasmania aod
surviving, in part even today in the northwest, 1
important, Very isolated tribes, such as the Nakako,
who first care Into contact with Europeans only
in the 1960s, live more (han 900 km south-east of
the north-west coust, They alsa tad the same
Tartangan-type knife.
Mulvaney, unfortunately in disregard of the great
priority in our nomenvlature, has tended to avoid
recognising the reality of a Tartangan culture phase
and its range of key tools, tending to use instead
a vague term such as “Tasmanoid’, to deseribe his
own finds at Kenniff Cave and elsewhere. An
anthropologist with geological training should re
PORT HEDLAND ROCK ENGRAVINGS ag
examine the site on Tartanga Island, Having had
training in Pleistocene geology mysell, it is my
opinion that the geological realities of the original
Tartangan site should be studied again, when the
original work there will be cleared of the errors in
interpretation which have occurred. The priority
and status of the term Tartangan also will be
verified,
ACKNOWLEDGMENTS
This wuihor acknowledges Hie support in his researches
by the Members of the Board of the South Australian
Museum and its Directars over the more than sixty-five
years of his association with the orgam2aion,
Permission to do fieldwork was granted by the then
Department of Native Welfare of Western Australia Lo
members of the University of California at Los Angeles
and the University of Adelaide Anthropological
Expedition af 1952 to 1084. Professor Joseph B, Birdsell
and this writer (accompunied for several months by the
late PJ. Epling) made anthropometric records ane
vondocual anthropological enquiries among the
Aborigines of Western Austria,
Members of the Expedition had the full co-operation
ac all stages of (he Native Welfare officers, and for the
particuliir work at Port Hedland District, Officer Harvey
Tilbrook was mest helpful in stimulating the interest of
Reregh
BASEDOW, H. 1925. The Australian Aborginal’ Preece,
Adelaide.
CRAWFORD, LM, 1964, The cnaravings of Depuch
Island, HW. Aust, Mus. Special Publication No, 2.
W. DL. Ride & A, Neumann (Eds.). Pp, 23-63,
HALE, H. M, & TINDALE, N, B. 1930. Notes on some
human. remains in the Lower Murray Valley, South
Australia, Ree. S. Aust Mus, 4(2); 145-218,
McCARTHY, F. DB, 1962. The rock engravings of Port
Hedland, northwestern Australia. Kroeber Anthrapo-
logical Papers No, 26, Berkeley,
MULVANEY, D. J. 1975 (orig. 1969), ‘The Prehistory of
Australia”, Pelican Books, Melbourne.
RADCLIFFE-BROWN, A, R, 1930, Social organization
of Australian tribes. Oceania 1; 34-63, 206-246;
322-441; 426-456.
RIDE, W. D. L.& NEUMANN, A. (Eds.) 1964. Depuch
Island, M4 Aust. Mus, Special Publication No, 2. Perth,
ROSE, F.C. G. 1950. An interpretation of some Abori-
zinal rock carvings and paintings in north western
Australia, Man 50; 13,
TINDALE, M, B. 1935, The legend of Waijungari, Jaralde
tribe, Lake Alexandrina, South Australia and the
the Aborigines in the work of the Expedition. In the course
of the eleven indnths spent in Western Australia more than
a thousand Aborigines helped in the studios,
Joseph B, Birdsell and | have been associated in such
fieldwork since 1938 and | appreciate continuing periodic
discussions with him. My wife Muriel Nevin Tindale has
been helpful in the ordering of this paper, and Mrs Eloise
Hardman has had the onerous task of deviphernine my
almost impossible writing. |am indebted to Peter Sutton
and Christopher Anderson who saw This paper through
the press while | wats absent in California, There may be
some errors Of which Lam unaware, aod some omissions
of dati still in my extensive field journals; for these | must
be held responsible.
ENDNOTES
1 ln this paper when individual Aboriginal words and
names are mentioned for the first time they are transeribed
in International Phonetics as set. out in my book on
Australian tribes (Tindale 1974; 2) and placed in-square
brackets. When tsed again they are given in close
conventional form, with place names in particular
following, the mandates of the Geographic U system of
spelling,
2. [mii is their normal word for blood, por tarda,
NCRS
phonetic system employed in its transeription, Ree. 8,
Aust. Mus, 12(5): 261-274,
TINDALE, N. B. 1957, Culture succession in southeastert
Australia from Late Pleistocene to the present, Ree. 9.
Aust, Mus. T3(1): 1-49.
TINDALE, N. B. 1965, Stone implement making among
the Nakako, Ngadadjara and Pitjandjara of the Greal
Western Desert, Rec, S, Aust. Mus, W5() 131-164,
TINDALE, N. B. 1974. ‘Aboriginal Tribes of Australia’.
Univ, Calif, Press, Berkeley.
TINDALE, N, B, 1981, Prehistory of the Aborigines! some
interesting considerations. /n A. Keast. (Ed.), Ecological
Biogeography of Australia’, Junk, The Hague, Pp, 1761-
1797,
TINDALE, N, B. 1983. Woakwine ‘Terrace in the South
Hast of South Australia and indications of the very early
presence of man. 7n N.C. Fleming (Ed.). ‘Quaternary
Coastlines and Marine Archaeology’, Academic Press,
London. Pp, 543-600.
TINDALE, N. B, (In press), Celestial lore of Australian
Aborigines.
OBITUARY IFOR M. THOMAS 1 APRIL 1931 [I.E. 8 JULY 1909] - 14
AUGUST 1985
BY S.J. EDMONDS
Summary
Ifor Morris Thomas was born on 8 July 1909 at Aberdare, south Wales and died at Adelaide on 14
August 1985. He was the fourth and youngest son of Thomas and Jane Thomas (née Watkins). His
father was Headmaster at Aberdare Grammar School. Ifor received his primary and secondary
education locally and in 1928 went on to study science at the University of Wales at Cardiff. He
graduated B.Sc. in 1931 and B.Sc. (with Honours) in 1932. In 1933 he undertook postgraduate
research on crustaceans under Professor Tattersal and at the end of the year gained the degree of
M.Sc. After graduation he acted as Demonstrator in Zoology, gave a series of lectures on Biology to
the Workers’ Educational Association of Wales and, much to his liking, spent some time on board a
trawler in the North Atlantic as a collector of marine specimens. His Atlantic experience
undoubtedly influenced the future course of his life, serving to stimulate a keen and life-long
interest not only in marine biology but also in boats, ships and the lore associated with them.
OBITUARY
IFOR M,. THOMAS
| April 1931 - 14 August 1985
lfor Morris Thomas was born on 8 July 1909 at
Aberdare, south Wales and died at Adelaide on 14
August 1985, He was the fourth and youngest son
of Thomas and Jane Thomas (née Watkins). His
father was Headmaster at Aberdare Grammar
School, Ifor received his primary and secondary
education locally and in 1928 went on to study
science at the University of Wales ai Cardiff. He
graduated B.Sc. in 193] and B.Sc. (with Honours)
in 1932, In 1933 he undertook postgraduate research
on crustaceans under Professor Tattersal and at the
end of the year gained the degree of M,Se. After
graduation he acted as Demonstrator in Zoology,
gave a series of lectures on Biology to the Workers’
Educational Association of Wales and, much to his
liking, spent some time on board a trawler in the
North Atlantic as a collector of marine specimens.
His Adlantic experience undoubtedly influenced the
future course of his life, serving (o stimulate a keen
and life-long interest not only in marine biology but
also in boats, ships and the lore associated with
them,
He came to Australia in January 1939 to take up
a lecturing post in the Department of Zoology at
the University of Sydney, at that time under the
direction of Professor W. J. Dakin, His arrival
happened to coincide with a huge bushfire which
endangered many districts along the coast of New
South Wales and covered Sydney with a pall of
smoke and ashes. He used to relate how dismaying
were his first impressions of Australia, He soon
found himself, however, fully occupied in his new
post, where his duties included teaching and
organising classes in physiology, lecturing on
invertebrates and assisting in {he running of a small
collecting vessel called the ‘Thistle’. When World
War II broke out in 1939, Ifor and a colleague, Allan
Colefax, became involved in devising ways of
camouflaging objects at sea and on land. They
specialised on the construction and use of different
kinds of netting for use in concealing things on the
ground,
In 1947 Ifor joined the staff of the Department
of Zoology of the University of Adelaide as
Lecturer and became Senior Lecturer in 1950, He
remained in Adelaide for the rest of his life and
occupied a senior post in the Department until he
retired at the end of 1974, In 1973 and 1974 he acted
as Head of the Department. In 1947 the lecturing
staff of the Department consisted only of him and
Professor T, H. Johnston. During an association
of almost 30 years with the University of Adelaide
he lectured to students at all levels and on almost
all aspects of zoology. Most of his early teaching
was to second and third year classes but as the
number of post-graduate students increased and the
number of the staff grew, more of his attention was
given to those reading for Honours, Masters and
Ph.D. degrees. His chief interest was marine
zoology. His third year ‘unit’ course in marine
zoology was always a popular one; students found
the lectures and practical work interesting and
62 S. J. EDMONDS
stimulating. For many years he used to take a party
of his students for a couple of weeks each year to
the CSIRO Marine Laboratory at Cronulla, New
South Wales, where they made good use of facilities
that were not available in Adelaide. The trips were
a great success and much liked by the students.
Some of them, although now old and sere, still talk
about them when they meet. Most of his advanced
and post-graduate students took for their research
topic some aspect of the biology of a common,
South Australian marine or freshwater animal.
Some of the animals studied were the freshwater
mussel, rock lobster (crayfish), whiting, garfish,
‘Coorong’ mullet and black bream. Consequently
his influence in the fields of freshwater and marine
research in South Australia was very considerable.
Many of his former students today occupy posts,
some senior, in the State Fisheries Department, the
State Water Laboratory, the South Australian
Museum and Australian Universities. Without
doubt most of them were encouraged to pursue their
careers as marine zoologists or ichthyologists as a
result of attending his classes.
His own research was concerned with: (1) the
taxonomy and certain aspects of the physiology of
enteropneusts (Hemichordata), and (2) the zoology
of the intertidal zone. He published papers on both
topics. His best paper was probably that on the
accumulation of radioactive iodine in the endostylar
region of the cephalochordate Amphioxus. By using
autoradiographic techniques he showed that certain
regions of the endostyle of Amphioxus were able
to take up radioactive iodine to form a mucin-like
substance that contained iodine. His results lent
support to the idea that the endostyle of the lower
chordates was the forerunner or homologue of the
thyroid gland of cyclostomate fishes and so of early
vertebrates. He wrote two papers on intertidal shore
life. He enjoyed nothing more than working along
the coast, especially during the summer. On such
occasions his ‘outfit’ was special — a very large,
wide-brimmed straw hat (held in position by an
elastic chin-strap), slightly ‘oversized’ shorts, a khaki
shirt, long stockings (he sunburnt easily) and white
sand shoes. To a collecting bag, held in place by
a strap around his shoulder, were fastened by plastic’
tapes a pair of scissors, a pair of forceps, a hand
lens, a large pocket knife and a pencil. For good
measure his spectacles were sometimes attached to
the lapel of his shirt. So engrossed was he in his
work, however, that he was never aware of the
slightly puzzled looks that he sometimes drew from
quizzical bystanders.
After retiring from the University at the end of
1974, he continued an active, scientific life as an
Honorary Associate attached to the Marine
Invertebrates Section of the South Australian
Museum. His work was now directed mainly along
two lines. Firstly, he acted as advisor to the Electricity
Trust of South Australia on the thermal pollution
of marine environments caused by the building of
power stations in the State. Secondly, he acted as
co-editor of one of the recent publications of the
Handbooks of the Flora and Fauna of South
Australia, namely ‘Marine Invertebrates of
Southern Australia, Part I’. The book is a very
useful one because it helps to fill some of the gaps
in the knowledge of the invertebrates of the State.
Part II was almost completed when he died.
During his life in Adelaide he took an active
interest in a number of societies. He served as
Secretary, Librarian, Councillor and President of
the Royal Society of South Australia and acted as
Assistant Editor of its Transactions for a number
of years; in 1977 he was made an Honorary Fellow,
a rare honour. He was a member of the Royal
Zoological Society of South Australia, the
Australian Marine Sciences Association, the Nature
Conservation Society, and the National Trust of
South Australia, He was also a member of the
Institute of Biology.
To those who knew him well he was a good friend
and a valued colleague. He could be courageous,
too, Not many people know that about four years
ago, by challenging an apparently armed man, he
foiled the robbery of a suburban bank.
Ifor was well-liked by both his students and those
with whom he worked, and was always approach-
able, reasonable and helpful. His contribution to
marine zoology in South Australia was a most
valuable one.
He is survived by his wife (née Patricia Mawson)
and three sons.
BIBLIOGRAPHY
THOMAS, I. M. 1936, Diastase in rabbit saliva. Nature,
Lond. 138: 1015.
THOMAS, |. M. 1949. The adhesion of limpets. Aust.
J. Sci. WW: 28-29,
THOMAS, I. M. 1949. A sex-intergrade. Aust. vet. J. 25:
294-297.
THOMAS, I. M. 1950. Craspedacusta sowerbyi in South
Australia, with notes on its habits, Trans. R. Soc. S.
Aust. 74: 59-65,
THOMAS, I. M. 1950. The Medusa Craspedacusta in
Australia. Nature, Lond, 166: 312,
THOMAS, 1. M. 1950. The reaction of mosquito larvae
to regular repetitions of shadow stimuli. Aust, J. Sci.
B, 3: 113-123.
THOMAS, I. M. 1955. The Council for the promotion
of Field Studies. S. Aust. Nat. 29: 67-69.
THOMAS, I, M. & EDMONDS, S. J. 1956. Chlorinities
of coastal waters in South Australia, Trans, R. Soc. S.
Aust. 79: 152-166.
OBITUARY — IFOR M. THOMAS 63
THOMAS, 1, M. 1956. The accumulation of radioactive
iodine by Amphioxus. J, mar. biol. Assac. U.K. 35:
203-210.
THOMAS, I. M. 1958. Some notes on the fauna of South
Australia. Jn R. J. Best (Ed.). ‘Introducing South
Australia’, ANZAAS, Adelaide: 111-126.
THOMAS, I. M. 1962. Some aspects of the evolution of
thyroid structure and function. 7m ‘The Evolution of
Living Organisms’. R, Soc, Vict,, Melbourne: 166-172.
THOMAS, I. M. 1968. Two species of Saccoglossus
(Enteropneusta) from South Australia. Trans. R. Soc.
§. Aust, 92; 73-84.
BYE, J. A. T., TAMULY, A. & THOMAS, I. M. 1968,
Cruise no, H.L.C. 3. Horace Lamb Centre for
Oceanographic Research, Adelaide. 50 pp.
SHEPHERD, S. A. & THOMAS, I. M. 1969, Pearson
Island Expedition 1969. I. Narrative. Trans. R. Soc. S.
Aust. 95: 321-122,
THOMAS, I. M. & DELROY, L. B, 1969. Pearson Island
Expedition 1969. IV. The Pearson Island Wallaby, Trans.
R. Soc. S. Aust. 95: 143-145,
OTTAWAY, J. R. & THOMAS, I. M. 1971, Movement and
zonation of the intertidal anemone Actinia tenebrosa
Farg. (Cnidaria; Anthozoa) under experimental
conditions. Aust. J. mar. & freshw. Res, 22: 63-78.
THOMAS, I. M. 1972, Action of the gut in Saccoglossus
otagoensis (Hemichordata: Enteropneusta). N. 2, J.
mar. & freshw, Res. 6: 560-569.
WOMERSLEY, H. B. S. & THOMAS, |. M. 1976.
Intertidal ecology. Jn C. R. Twidale, M. J. Tyler, &
B. P. Webb (Eds.). ‘Natural History of the Adelaide
Region’. R, Soc. S. Aust, Adelaide: 175-186.
THOMAS, I. M. & EDMONDS, §. J. 1979. Intertidal
invertebrates. Jn M. J. Tyler, C. R. Twidale, & J. K. Ling
(Eds.). ‘Natural History of Kangaroo Island’. R. Soc.
S. Aust., Adelaide: 155-166,
THOMAS, I, M. & SHEPHERD, S. A. 1982. The marine
environment. Jn S. A. Shepherd & 1. M. Thomas (Eds.),
‘Marine Invertebrates of Southern Australia, Part 1’.
Govt Printer, Adelaide: 11-25.
THOMAS, I. M. & SHEPHERD, S. A. 1982. Sea
anemones (Order Actiniaria, Zoanthidea and
Corallimorpharia). Jn S. A. Shepherd & I. M. Thomas
(Eds.). ‘Marine Invertebrates of Southern Australia, Part
I’. Govt Printer, Adelaide: 160-169.
THOMAS, I. M., AINSLIE, R. C., JOHNSON, D. A.,
OFFLER, E. W. & RED, P. A. 1986, The effect of
cooling water discharge on intertidal fauna in the Port
River Estuary, South Australia, Trans. R. Soc. S. Aust.
110: 159-172.
S, J. EDMONDS, South Australian Museum, North Terrace, Adelaide, South Australia 5000,
OBITUARY BRIAN DAILY 1 APRIL 1931 - 6 MARCH 1986
BY S.J. EDMONDS
Summary
Dr Brian Daily died in his home in Adelaide on 6 March 1986, after a courageous battle with
cancer, during which he continued to work right up until his death. For five years in the 1950s he
had worked in the South Australian Museum.
OBITUARY
BRIAN DAILY
1 April 1931 - 6 March 1986
Dr Brian Daily died in his home in Adelaide on
6 March 1986, after a courageous battle with cancer,
during which he continued to work right up until
his death. For five years in the 1950s he had worked
in the South Australian Museum.
Brian was born in Adelaide on 1 April 1931.
Following schooling at the Marist Brothers School,
Norwood, and Sacred Heart College, Somerton
Park, Brian attended the University of Adelaide
where he completed his B.Sc.(Hons.) in 1952 with
a thesis on the Tertiary sequence of the Noarlunga
Basin. He continued on to his Ph.D, on the Cam-
brian stratigraphy and palaeontology of South
Australia. This work, for which he was awarded the
1956 Tate Medal of the University of Adelaide, has
been the basis of most subsequent studies on the
Cambrian of South Australia. The faunal assem-
blages that Brian established have been used as a
reference succession for Australian Lower Cambrian
sequences.
While still a student in 1954, he joined the late
C. P. Mountford’s National Geographic Society
expedition to Arnhem Land, and made the first
comprehensive collections of fossils from Bathurst
and Melville Islands. Mostly ammonites, these are
still being studied.
Brian completed his Ph.D. late in 1955, and
joined the South Australian Museum in January
1956, as Assistant Curator of Fossils. Subsequently
he became Curator of Fossils and Minerals, demon-
strating his versatility and gaining valuable
experience for his later career. Brian was indefati-
gable in the field, as many of his students can attest.
While at the Museum, he collected widely through-
out the State: Tertiary plant fossils on the Arcoona
plateau in 1956 and again in 1960; Pleistocene
mammals at Hookina Creek in 1956; Cambrian
trilobites from Kangaroo Island in 1957, and Pre-
cambrian fossils at Ediacara in 1957, 1958 and 1959,
He also collected from the Cambrian and Triassic
66
of the Flinders Ranges in 1959, and from the Cam-
brian and Ordoviglan of central Australia in 1959
and 1940,
Jn 1957, Brian joined Professor R.A. Stirton of
the University of California, Berkeley, for 4 three
inonth expedition into the Lake Eyre region in
search ot Tertiary marsupials al Lake Palankarinna
and adjoining areas, discovering the richly fossili-
ferous deposic of Pliocene Mampuwordu Sands in
the Lawsan-Datly quarry, and making an important
contribution to the understanding of the geology
Of (he area. His interest in the region continued
through the years, and he and his students used the
southern shores of Lake Byre North for fieldwork.
During these years, he began to take on a consul-
tancy role, examining bore material for the South
Australian Mines Department and petroleum
explorers, Thus he studied the fossils of the Minla-
ton and Stansbury Stratigraphic Bores ou Yorke
Peninsula, and later, the Gidgealpa No_ ! Well.
Wich several fellow students, he had been a
founding member of the Cave Exploration Group
of South Australia in 1955, and was influential iy
having that society closely affiliated with the South
Australian Museum so that all specimens encoun-
tered, both fossil and living, became part of che
State collections, Thus he was involved jn a pumber
of excavations in Naracoorte caves [a 1957 and 1959,
mainly for the (then) puzzling Thvlacales or marsu
pial ‘lion’, This interest in vertebrare fossils was
maintained despite his first Jove — the Cambrian
— and communicated itself subsequently to a
number of his students, including one of us (NSP).
Brian resigned trom the South Australian
Museum in January 196), and was subsequently
made an Honorary Associate of the Palacontology
Section, He was appointed a Lecturer in (he Depart-
ment of Geology and Mineralogy, University of
Adelaide, in 1961; was promoted lo Senior Lecturer
in 1964 and to Reader in (974, a pasitioti he held
until his death. Brian was an en|husiastic and
dedicated teacher, particularly of fleld work. He was
an excellent field observer With a wide general
geological knowledge which he passed on lo fis
students. During his time at Adelaide University,
Brian supervised more Honours Degree projects
than any other person in the department and also
supervised a number of suecessful M.Sc. and Ph,D.
projects, mainly on the Adelaidean and Cambrian
geology and biostratigraphy of the Flinders Ranges.
Brian’s main research activities were on the
Precambrian and Cambrian stratigraphy and sedi-
mentology of South Australia. He made substantial
coninbutions lo the study and understanding of \Ive
Precambrian/Cambrian boundary, both in Aus-
tralia and overseas. This work included visiting
important Precdmbrian-Cambrian boundary
sections in North America, Argentina, China,
Morgeco, Burope and Siberia, We was a valiable
member of 140,P. project 7, “South-West Pacific
Basement Correlations’, the results of which were
published in G.S.A, Speeial Publication No. 9 In
clucidating the sttareraphy and structure of the
Kanmanioo Group, Brian, in association with Dr
A. R. Milnes, made a major contribution to the
understanding of the gcolugy of Fleuriew Peninsulit
and Kangaroo Island. Unfortunately much of
Brian's detailed and pioneering work on Le
Adelaidean and Cambrian sequences of the Flinders
Ranges was unpublished at che time uf his death.
In addition to his professional cuties, Brian wave
much voluntary service both to geology and science
in general. In the period 1957-57 he was successively
programme secrelary, council mentber, vice-
preside and president of she Royal Society of
‘South Australia, He was chairman of the S.A.
Division of ANZAAS in 1981-1983 and was the
ANZAAS correspondent tor some years. His
interest in geological education is shown by his
ating as chairman of the S.A, Public Examination
Board Geology subject comimiitice and also a chiel
examiner ol matriculation geology in South
Australia over a number of years. Arian was a
member ol the S.A. Divisional Commitice of the
Geological Society of Australia in 1965-66, vice-
chairman in 1967-48 and chairman in 1968-69 and
was a proxy member of the federal execulive of the
G.S.A_in 1980-8) He served ou borlrilte S.A. Strati-
graphie Nomenclature and Geological Manuments
Subcommittees.
Both asa friend aiid a colleague Brian will be
sorely missed. Deepest sympathies are extenued ja
his family
BialioorAryy
DAILY, B. 1956. Phe Cambrian in South Australia. fn ‘tl
Sistema Cambrico, su Paleagcogsalia y el Problema de
sU base’. Rep Jrternat, Geal. Congr 20h Mexiee 2,
31-147,
DAILY, B. 1957, The Cumbrian in Soullt Ausiraliq. frp
‘The Canihnan Geology of Australia’. Aan Aton.
Resour, Geol, Geaphes, Aust. Bull, 4%; 91-147,
HORWITZ, R. LC, & DAILY, B ISSR. Yorke Peninsola
in MF Glaessner & L.W, Parkin (Eds.), The Geolopy
of South Ausiralia’ < Geol, Soc, lust. 5: 46-50.
GLAESSNER, M. F & DALLY, B. 1959. The geology and
Late Precambrian fauna of the Ediacara Fossil Reserve,
Reo S. Aust, Mus. 13: 369-401,
DAILY, B. 1960. TAylacoleo, the extinct marsiipial lion,
Aust. Mus. Mag, 13: 163-166.
DALLY, B. 1963. The fossififcrous Cambrian successing
on Fleurieu Peninsula, Sourl Atusicalia, Ree, 8. Aust
Mis. Id: 579-601.
BONYTHON, Cc. W..& DALLY, B. 1963, ‘Report of Sourh
Australian Subcomimines, Nacional Parks Committes’,
Alist. Acad. Scj., Canberra, 63 op., 6 maps,
BONY THON, ©, W, & DAILY, ik 1945, Nationa! Parks
and Reserves in South Austeslia, Brae. R. Geogr Sue.
Aunt, S Aust. Branch aS: 5 1S_
DAILY, B 1966, Delhi Santos Gidgealpa No. | Well, South
Australia. Appendix 3, Part 8, Palacontological report
on. cores nos, 12 to 32) 1, 23, 24, 27, 29 and JO. Bun
Adin, Res. tush Pein Search Subs, lets, Publ, 13:
95-111,
IWIDALE, C. R., DAILY, B. & FIRMAN, LB. 1967,
Bustaric and climatic history of the Adelaide area, South
Australia: a discussion, J Gea/, 7S: 247-242.
DALLY, B, PWIDALE, C. BR. & ALLEY, N. 1969
Oecurrence of Lower Cambrian sediments in Wilpena
Pound, central Plinders Ranges, South Australia. Awse
J. Str. AV A 307.
DAILY, B & FORBES, & & 1969. Notes on the Proero-
yoie and Cambnan. sourhern and central Minders
Kariwes, Soultt Austria fy B. Daily (Ed.). ‘Geological
Excursians Handbook” ANZAAS, Adelaide, Section
3, 23-30.
DALY, & 1969. Fossiliferous Cambrian sedinicnrs and
low-grade metamorphics, Fleurigu Peninsula, South
Australia, dn BL Daily (Ed) ‘Geological Excursions
Handbook’. ANZAAS, Adelaide, Seation 3, 49.354.
OLIVER, Ro L & DAILY, B. 1969. Proictozoic and
Permian glacigene Weposits near Adelaide, South
Australia, Ji W. Daily (Ed.).. Geological Excursions
Handbook: ANZAAS, Adelaide, Section 3, 63-66.
CRAWFORD. AR. & DAILY, B, 197], Probable non-
synchraneily of Late Precambrian glanalons. Nature,
Lund, 2b WT-112
DAILY. B. & MILNES, A. R, 197). Stranigraphic mores
on bLawer Cambrian fopsiliferous fetaeseditnents
hetween Campbell Creek and Tunkalilla Beach in the
Type section of the Kanmanioo Group, Fleurieu
Peninsula, South Australia. Trans, RK. Sac. §. Aust. 95:
[00-214,
DAILY, 8 & MILNES, A. R. IST). Diseovery af Late
Precambrian ulliles (Sturt “iroup) and younger
melasediments (Mafia Group) on Daidley Peninsula,
Kangaroo Island, South Australia. SeareA 2: 431-453,
DAILY, B. 1872. Aspects of carbonale sedimeniaton in
the Cambrian of South Austratia, Geol. Sac. Aust.,
Abstracts Joint Speviulists Groups Meetings, Canberra,
Feb,, C1001.
DAILY, 8. t972, The base of (he Cambriay| and the first
Cambrian faunas. Spee. Pub. No |. Univ Adelaide,
Centre for Precambrian Research: 14 41,
DAILY, K.& MILNES, A, R, 1972. Sigititicance of basal
Cambrian metasediments of andalusice grade, Dudley
Peninsula, Kangaroo Istand, Seuch Australia. Search
SB: 89-90).
BAILY, B.& MILNES, A. BR. 1972, Revision of [he
siralipraphic nomenclature of dye Cambrian Kanman-
too Group, South Australia. Geo’ Sac, Auge, 19
197-202,
PORBES, B. G,, COATS, R, BP. & DAILY, B. 1972, The
lrure Voleanius. Q Nuves S.A, Depi. Mines 44) 1-5.
IAGO, J.B, REID, K.O., QUILTY, PG. GREEN, G, R.
& DAILY, B, 1972. Mossiliferous Cambrian limestone
fiom Within te Mi, Read Voleanics, Mi. Lyell Mine
Arta, ‘Tasmania. 2 Geal Soc, Aust, 1%: 379382,
DAILY, B, GOSTIN, Vi & & NELSON, C. A. 1973.
Tectonic origin far an assumed (ave Proterozoic glacial
07
pavement in South Ausivalia vt Geel Soe, clus. 20:
75-78.
DAILY, B.& MILNES, A. R, 1973. Stratigraphy, structure
and metamorphism of the Kangaroo Group (Cambrian)
in ats type section east of Twnkalilla Beach, Sourh
Australia, Trans, R Soe. 8 Aust. 97: 213-242.
DAILY, B. 1973. Discovery and significance of basal
Cambrian Uratanna Formation, MI, Scott Range.
Flinders Range, South Australia, Search 4: 202-205.
DAILY, B., IAGO, J.B. & MILNES, vA. R. 1973, Large-
soale horizontal displacement within Australo-
Antarctica jn the Ordovician, Nature Physical Science
244: 41-f4.
DAILY, B., TWIDALE, ©. R. & MILNES, A. R. 1974,
The age of he lateritized Summit surface On Kangaroo
Island and adjacent areas of South Australia. &. Geol
Soe. Aust, 21. 387-392.
JAGO, |, B.& DAILY, B, 1974. The trilobile Clavaznustuy
Howell from the Cambrian of Tasmania. Palaeontology
17: 95-109.
DAILY, B. & JAGO, J. B. 1975. The (ribolile Lejopyge
Hawle and Gorda and (he middle-upper Cambrian
boundary. Palgeontolaxy 1B: 527-350.
DAILY, B 1976. Novye dannye ab osnovanii Kembriya
y Yulmoy Australi, (New dara on the base of the
Cambrian in South Australia). /2v. Akad. Nowk, SSSK.
Ser, Geol. 1976 (3); 45-52 din Russian).
DAILY, B. & COOPER, M. R. 1976. Clastic Wedges and
patlerned ground inthe Late Ordovieian-Early Silurian
Lillites of South Africa, Sedimenialagy 23: 271-283.
DAILY, B, FIRMAN, |. B, FORBES, BG. & LINDSAY,
J, M. 1976, Geology, In C. R. Twidale, M. J. Tyler &
BH. P. Webb (Pas.). ‘Natural History of the Adelaide
Region’. R. Soe. S, Aust,, Adelaide: 5-42.
DAILY, I, THOMSON, B, P., COATS, R, P. & FORBES,
Ik G, 1976. ‘Late Precambrian and Cambrian gcology
of the Adelaide “Geosyneline” and Stuart Shell, South
Australia’, Excursion Guide Na. 332A, 25th International
Geological Congress.
MILNES, A. R., COMPSTON, W, & DAILY, B 1977. Pre
to syn-tectonic emplacement of early Palacozoie granites
in south-eastern South Australia, J. Geol Sec. Ase
24: 87-106,
DAILY, U6, MILNES, A. R,.& TWIDALE, CR, 1979.
Geology aid Geomorphology. dr M. J. Tylor, C. Ry
Twidale & J. K. Ling (Eds. ‘Natural History of
Kangaroo Island’. R. Soc. 5, Ausi,, Adelaide: 1-38.
DAILY, B, MOORE, PS. & RUST, BL R. 1980. Terres-
triul-marine wansition im the Cambrian Rocks of
Kangaroo Island, South Atistralia.. Sedimentology 27:
379-399,
DAILY, G., JAGO, J. B, & JAMES, P. R, 1982, ‘Lower
Cinbran sediments, Precambrian-Cambrian boundary
and Delamerian tectonies of southern Fleurieu
Peninsula’. 1V In(, Symposium on Antartic Earth
Sciences, 1982. Liniv. Adelaide, Guide to Excursions
BI-Ba: 30-41,
JAGO, J, B. & DALLY, EK 1982. Rewional sequences, Sourh
Australia, Jn R. A. Comper, & G. W. Grindley (Eds).
‘Late Proterozoic to Devonian sequences of soulheastern
Australia, Antartica afd New Zealand and their
correlauion”. Geol. Soe. Aust Syrec. Publ, 9: 6-12,
GRINDLEY, G. W., COOPER, R.A, DAILY, 8 &
OORBETT, KD. 1982. Principal imer-regional varre-
68
lations and geotectonic events. Jn R. A. Cooper, & G. W.
Grindley (Eds.). ‘Late Proterozoic to Devonian
sequences of southeastern Australia, Antarctica and
New Zealand and their correlation’. Geol. Soc. Aust.
Spec. Publ. 9: 57-71.
JAGO, J. B., DAILY, B., VON DER BORCH, C. C.,
CERNOSKIS, A. & SAUNDERS, N. 1984. First
reported Trilobites from the Lower Cambrian
Normanville Group, Fleurieu Peninsula, South
Australia. Trans. R. Soc. S. Aust. 108: 207-211.
JAGO, J. B., GEHLING, J. G. & DAILY, B. 1986.
Cambrian sediments of the Sellick Hill-Carrickalinga
Head area, Fleurieu Peninsula, South Australia. Eighth
Aust. Geol. Convention, ‘One Day Geological Excur-
sions of the Adelaide Region’. Geol. Soc. Aust.: 67-81.
J. B. JAGO, School of Applied Geology, South Australian Institute of Technology, P.O. Box 1, Ingle Farm, South
Australia 5089, and N. S. PLEDGE, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Photo
courtesy A. R. Milnes.
RECORDS
I)
THE
SOUTH
AUSTRALIAN
MUSEUM
VOLUME 21
MAY 1987
ISSN 0081-2676
CONTENTS
I
29
35
43
61
CVHYS: WARES
Revision of Australian Berosus Leach (Coleoptera: Hydrophilidae) :
I. M. KERZHNER
INE Voy (oteVom Ge (akceejolcae-) elma cteretctanl
Di @aREE.
Introductory study of advanced oribate mites (Acarida: Cryptostigmata: Plano-
fissurae) and a redescription of the only valid species of Constrictobates
(COyat exexeCeyreler9)
N. B. TINDALE
Kariara views on some rock engravings at Port Hedland, Western Australia
NOTES
S. J. EDMONDS
Obituary of I. M. Thomas
J. B. JAGO & N. S. PLEDGE
Obituary of B. Daily
RECORDS
OF
THE
SOUTH
AUSTRALIAN
MUSEUM
VOLUME 21 PART 2
NOVEMBER 1987
INLAND, COAST AND ISLANDS : TRADITIONAL ABORIGINAL
SOCIETY AND MATERIAL CULTURE IN A REGION OF THE SOUTHERN
GULF OF CARPENTARIA
BY D. S. TRIGGER
Summary
This paper compares aspects of traditional Australian Aboriginal societies in inland, coastal and
island settings, in a region of the southern Gulf of Carpentaria. It focuses on similarities and
differences in material culture after summarising comparative data on language, emic
environmental classification, kinship and social organization, and genetics. As ‘saltwater people’
occupying what was regarded as ‘saltwater country’, coastal mainlanders were part of the island
cultural bloc in significant respects, while being in other respects incorporated within a large
cultural bloc which extended inland to the west and southwest. The paper proposes that the coastal
mainlanders’ relationship with the North Wellesley Islanders in particular, reinforced the
maintenance of cultural differences between coast and inland society on the mainland. Despite
arguments for the relative isolation of the South Wellesley Islanders, the paper describes a complex
situation of overlapping cultural forms throughout the region, with no area emerging as completely
distinctive.
INLAND, COAST AND ISLANDS: TRADITIONAL ABORIGINAL SOCIETY AND MATERIAL
CULTURE IN A REGION OF THE SOUTHERN GULF OF CARPENTARIA
D. 8S. TRIGGER
TRIGGER, D. S. 1987, Inland, coast and islands: traditional Aboriginal society and material
culture in a region of the southern Gulf of Carpentaria, Rec. S. Aust. Mus. 21(2): 69-84.
This paper compares aspects of traditional Australian Aboriginal societies in inland, coastal
and island settings, in a region of the southern Gulf of Carpentaria. It focuses on similarities
and differences in material culture, after summarising comparative data on language, emic
environmental classification, kinship and social organization, and genetics. As ‘saltwater people’
occupying what was regarded as ‘saltwater country’, coastal mainlanders were part of the island
cultural bloc in significant respects, while being in other respects incorporated within a large
cultural bloc which extended inland to the west and southwest. The paper proposes that the coastal
mainlanders’ relationship with the North Wellesley Islanders in particular, reinforced the
maintenance of cultural differences between coast and inland society on the mainland. Despite
arguments for the relative isolation of the South Wellesley Islanders, the paper describes a complex
situation of overlapping cultural forms throughout the region, with no area emerging as completely
distinctive.
D. S. Trigger, Department of Anthropology, University of Western Australia, Nedlands, Western
Australia 6009. Manuscript received 10 August 1986.
This paper presents comparative ethnographic
data in a reconstruction of aspects of cultural
diversity among Australian Aboriginal societies in
an area of the southern Gulf of Carpentaria (Fig. 1).
It is concerned particularly with the situation of
coastal mainland dwellers located adjacent to two
major off-shore island populations, and apparently
subject to influences from diverse cultural
traditions. These are traditions which will be glossed
as those of ‘inland society’ on the one hand and
islands society’ on the other hand, although both
labels themselves encompass a degree of cultural
diversity.
The paper investigates the ways in which a
mainland coastal group’s relations with off-shore
island societies may have influenced the former’s
relations with other mainland societies (particularly
inland ones). It will be argued that while the coastal
mainland society can be seen in some ways to be
included within a broad cultural bloc extending
inland to the west and south-west, in other respects
it is clearly on the periphery of that bloc. The paper
proposes that the influence of ongoing relations
with North Wellesley Islands societies can be seen
as having kept coastal mainland society on the
periphery of the cultural bloc extending inland to
the west and south-west. Hence, rather than
focusing directly on the degree of isolation of the
islands society, my approach is to ask whether
major off-shore islands society exercises important
influences over immediately adjacent mainlanders,
thereby reinforcing tendencies towards cultural
differences between the latter and those further
inland,
Cultural differences between coast and inland in
the Gulf of Carpentaria context have been noted
in previous literature. Spencer & Gillen (1904: 634)
mention the ‘very considerable uniformity’ in
material culture in inland areas on the southwest
side of the Gulf, ‘until we come to the true coastal
tribes, amongst whom we naturally meet with
certain objects not present amongst the inland
tribes’. For the south-eastern side of the Gulf,
Tindale (1974: 121-122) notes the ‘great contrasts
between the life and economies of the inhabitants
of the coastal mangrove and saline shore flats and
the ways of the scrub covered upland dwellers’.
Tindale also refers to the Aboriginal terms given
in Curr (1886: 303) for the Leichhardt River mouth
area, designating distinctive peoples of saltwater,
freshwater and scrubland domains. Detailed
material for the Cape Keerweer area on the east side
of the Gulf, has been given in Chase & Sutton
(1981). Those people living along the coastal strip
are described as not having interacted closely with
hinterlanders, Their territories are much smaller
than those of inlanders with correspondingly higher
population densities. Coastal people also possessed
higher social segmentation, a richer ritual tradition
and greater linguistic diversity (Chase & Sutton,
1981: 1835).
A considerable amount of literature is available
for the study region dealt with in this paper. This
is especially the case for work on the cultural
traditions of the Wellesley Islands, Together with
the results of my own research, this literature
provides a sound basis for a broad comparative
study of Aboriginal society in different environ-
70 D. S. TRIGGER
WELLESLEY ISLANDS
3 ISLANDS GROUP
wd OF
I Hore”
yor pot
1. Bayley Pt,
2. Bayley Is.
3, Pt. Parker
4. Forsyth Is.
FIGURE 1. Linguistic territories from Aboriginal perspective during the study period.
INLAND. COAST AND ISLANDS 7
menis, While athe: publicauions are planned, the
present paper focuses mainly on a comparison of
aspects of material culiure across inland, coastal
and island settings, Several museum collections of
material culture from the region have proved a
valuable resource in reconstructing aspects of
traditional life. However, to establish the hroader
context for the data on material culture, | will first
briefly discuss several more general isswes: (1) the
linguistle setting; (2) emic environmental classifi-
cations; (3) kinship and social organisation; and (4)
genetic similarities and differences through the
region.
THE LINGUISTIC SETTING
This paper primarity concerns four definable
areas Within the range of languave proups shown
in Fig, } Gatswa and Waanyi in the inland area
to the west and south-west of the Wellestey Islands:
Gangealida’ (and Gananggalinda) on the coast
adjacent to the islands; Layardilda* and Yanegzaal
on Mornington, Denham and Forsyth Islands
(constituting the rain islands of the North
Wellesley group); Gayardild on Bentinck and Sweers
Islands (constituting the main islands af the South
Wellesicy group). Several points conceming the
linguistic territories of the region warrant brief
comment.
Contemparary Aboriginal opinion makes it clear
that “Garawa people” were nat Saltwater people’,
Mapping of traditional Garawa estates which I have
completed indicates che northern mast of them to
be no closer than 2 km from the coast (ef.
Tindale's (1974; 228) figure oF 63 km). Across most
of the northern limit of Garawa teritory. the coastal
area is Yaniyula country. However, it remains unclear
which language group was predominant in the
coastal area eastwards fron the Calvert River to the
west side of Massacre Inlel (the weslern boundary
of coastal Ganggalida country). The language name
‘Nyangga' has been applied to this area in the
literature, and | Haye discussed elsewhere (Trigger
1985; 340-349) whether there Was ln fact such a
distinctive group in dialectal, territorial or
sociopolitical lerms, The igsue is raised here to note
that in comparing aspects of the inland Garawa
traditions with the coastal Ganggalida traditions,
1 am most likely pot comparing territorially
adjacent linguistle groups,
We should also gate the question of territorial
proximity between Waanyl and Ganggalida
traditions, s far ay U have been able 16 reconstruct
them (Trigger 1982, Map UU), iradibonal Waanyi
estates extended eastwards al least 30 km into
Queensland from the Northern Territory border.
The most precise data concern am eastern-most
Waanyi estate on the Nicholson River This does
not indicate close proximity betwoen Waaryl and
Ganggalida as it ig doubtful whether Ganggalida
speakers were predominant along the Nicholson
River in Queensland; the minimal but most rehable
information (a hand-written note on a word list by
Roth [/900]) suggests that Neuburinji speakers were
predominant in this area (cf. Keen 1983; 193).
Linguistic work indicates that Neguburinji was
mutually intelligible with Ganggalida, and in fact
that these two, together with Yanggaal (on Forsyth
tsland) and Gayardild (on Bentinck Island), were
dialects of one language: Layardilda (on
Mornington Island) was ‘ different bur closely
related language’ (Reeri 1983; 192; and see Evans
1985: 3), Pwo further points are thus significant.
Firstly, the nature of Unguistic relations demon-
strates the incorporation of two substantial linguis:
lic territories of the malnland (Ganggalida on the
coast and Nguburinji further Inland) within the
“Tangkic sub-group’ (Evans L985) extending
throughout the Wellesley Islands, ] use the nation
of ‘incorporation’ of the islands and a section of
the mainland deliberately to emphasise the lack of
linguistic isolation of the island societies; Tindale’s
discussion of ‘the relative wolation' (J9@2a7 278; and
see also 1977; 256-7) of the language of the
Bentinck Islanders is clearly inadequate (ef, Evans
§985: 9)" Secondly, Ganggalida speakers quite
likely extended inland from the coast for #
considerable distance (possibly up to 100 km)
however, in the comparative analysis presented in
this paper it is aspects of coastal Ganggalida society
which constitute the significant wnit-
The final point in discussing linguistic aspects of
the region concerns the area on Fig, t labelled
‘Gananggalinda’. This term was known during my
research by only a few senior Ganggalida people.
[t was applied to a group of people said to have
‘belonged beach side’ in the area between Bayley
Point and Point Parker (i.e, the coastal mainland
area immediately opposite bath the North and
South Wellesley Islands, and ineorporating a
distance along the coastline af approximately 20
kra). These are people said to have routinely visited
several small islands comparatively close to the
mainland (damely Bayley and Pains Islands
approximately 3 km and 4,5 km off Bayley Point
respectively, and Allen lsland approximately $5 km
off Point Parker). One senior knowledgeable
Ganggalida woman has defined Gananggalinds as
a form of speech very similar to Ganewalida, bur
with a different accent. However, it is probable that
the term was essentially applied to a group of people
rather (han to a dialect.4
tn establishing the setting for (the further
comparisons in this paper, it is approprale te
recognise these Ciananggalinda people as a
mainland group which was regarded by fellow
72 D. S, TRIGGER
coastal mainlanders (at least those to the west) as
particularly closely oriented towards the
immediately adjacent small islands to the north and
east. Indeed, it is probable that these people spent
substantial time on the close off-shore islands, in
regular contact with Yanggaal speakers based on
Forsyth Island. They also had occasional contact
with Layardilda people from Mornington Island,
and perhaps even sporadic contact with Bentinck
Islanders who may have travelled across to Allen
Island. The latter contact would have involved safely
traversing a distance of approximately 13 km by
watercraft, and Tindale (1962b: 298-301) gives
accounts of two attempts (in 1940 and late 1946 or
early 1947, prior to major intervention by
Europeans) where lives were lost. In the second
attempt, 14 out of 19 persons attempting the
crossing were drowned, and there is little doubt that
such trips would not have been undertaken without
knowledge of the great danger. Yet, it is likely that
the factors reportedly leading to the 1940s crossing
attempts (particularly quarrels and fights), among
others, would have, in earlier days, led to sporadic
successful crossings by small numbers of people
from Bentinck Island to Allen Island. Tindale
(1962a: 273, 290-291) found oral traditions among
the Gayardild which recounted occasional hostile
encounters with mainland Aborigines on Allen
Island. Evans (1985: 15-16) suggests that ‘whatever
contacts there were, that did not end in death or
exile, must have been separated by decades of
isolation’; he also mentions that nothing was
obtained by the Bentinck Islanders via trade.
Having mentioned this distinctive ‘beach side’
mainland group, I will nevertheless subsume them
within the category of coastal Ganggalida people
throughout the paper. In fact, there is little
remaining specific knowledge of these people, and
among the reasons for this is an apparent revenge
killing carried out on Bayley Island by Yanyula
people from the northwest. Historical records (The
Queenslander 10 April 1897: 774) indicate that this
occurred in 1897, and according to Aboriginal oral
tradition a large number of people were killed (see
Trigger 1985: 140-141).
THE DISTINCTIVENESS OF THE MAINLAND COASTAL
AREA: THE ABORIGINAL VIEW
Various kinds of distinctive environments
throughout the entire study region are distinguished
in Aboriginal classification (Trigger 1985: 63).
However, none are regarded as so significantly
different as ‘saltwater country’. Ganggalida
terminology for the coastal area is shown in Table 1,
As the English translations indicate, the environ-
ment consists of long sand ridges (supporting open
woodland) typically parallel to the beach and dune
areas. These raised ridges are separated by flat sandy
stretches of ground varying considerably in width.
In some areas, ‘saltpans’ or saline coastal flats
separate the sand ridges, but the widest area of
saltpan is usually between the most inland sand
ridge and the beginning of sharply defined open
sclerophyll woodland which extends inland to the
south (Fig. 2).
From the Ganggalida perspective the sand ridges
are termed ‘islands’ (murndamurra). The same term
is used for the small islands immediately off-shore.
In the Aboriginal view, the ‘land’ or ‘mainland’
(wambalda) thus begins at the inland limit of the
saltpan, which also marks the inland limit of what
is generally known as ‘saltwater country’. Figure 3
shows this limit approximately, designating a coastal
strip of ‘saltwater country’ extending southwards for
varying distances between 3 km and 10 km from
the beach. At times during the wet season, much
of the saltpan areas are said to be covered with water
(probably from tidal surges as well as from the flow
of fresh water from the inland), leaving the ‘sland’
sand ridges isolated with water on all sides.
In the Aboriginal view then, the coastal strip is
regarded as environmentally distinctive and
different from further inland, and in fact is
TABLE 1. Schematic representation of Ganggalida classification of environmental zones in the
mainland coastal area.
<a----- North
malara /mirlaja/mala/lilu ngarnda dumuwa gabara gin.gara
‘sea’ /go back/sea/north ‘beach’ . ‘sand ‘sand ... ‘Ssaltpan’ ‘flat country’
ridge’ ridge’
[inter-tidal zone] .. [multiple ridges] ... [on other side
of saltpan]
area containing
<---- murndamurra---->
‘islands’
<----=-------------- =~ == -------- ‘Saltwater country’ ------------------------------------ > ‘mainland’
‘land’
wambalda
INLAND, COAST AND ISLANDS 73
FIGURE 2. The physical environment in the coastal
mainland area (Ganggalida country). The long sand ridges
parallel to the coast-line are evident from the lines of
vegetation. The area of saltpan in the background
separates the coastal strip containing ‘islands’ (‘saltwater
country’) from the ‘mainland’, in the Aboriginal perspec-
tive. The saltpan and mangroves in the foreground occur
only occasionally along this stretch of coastline. (Courtesy
of Connah/Jones Aerial Archaeology, University of New
England,)
conceived (in general terms at least) as containing
areas equivalent to the close off-shore islands.®
However, this narrow coastal strip should not be
considered as socially isolated in any way. Figure 3
indicates quite precise boundaries for the coastal
estates, to the east and west; the boundaries are
hence the sea to the north, and creeks and salt-arms
to the east and west. But the boundedness to the
south (i.e. in an inland direction) is much less
precisely defined. All estates except one in the area
shown on Fig. 3 extend across the saltpan to the
‘mainland’ to include one or more fresh waterholes.
In the case of the exceptional estate (F), it includes
site Fl (Gunamula), a large fresh water lagoon near
the mouth of Cliffdale Creek, which is said to
contain surface water for a substantial part of the
dry season. Seasonal movements across the saltpan
to waterholes, occurred particularly during the
middle and late stages of the dry season, not so
much because of a lack of fresh water in ‘saltwater
country’ (for this is said to have been always
available by digging ‘soaks’), but in order to obtain
certain material resources apparently not at that
time readily available on the ‘islands’ (e.g. water
lilies, Nymphoea sp. and Nymphoides sp.). During
parts of the dry season people are also said to have
come to the coastal strip from a considerable
distance inland; however, they apparently rarely
remained there during the wet season because of
their lack of tolerance of the increased number of
mosquitoes then. The significant point here is that
apart from the recognition of the environmental
distinctiveness of ‘saltwater country’, people are said
Example of named island -—--—>|sland"
- vicinities
“Wambilbayi
Island”
FIGURE 3, Ganggalida and Gananggalinda coastal estates showing the Aboriginal designation of the inland limit
of the saltpan areas as the beginning of the ‘mainland’ (wambalda). Certain fresh water holes on the ‘mainland’ are
shown for each estate.
4 DS TRIGGER
10 have moved in and out of it constantly,
Nevertheless, it was viewed as the distinctive domain
appropriate to 3 Conceptually separable group of
‘sullwaler people’.
KaysHip AND SOCIAL ORGANISATION
An Aranda (ype kinship system appears lo fave
operated traditionally throughout the study regien.
See Trigger (J985} for some discussion of its natitre,
and also comments by Warner (1933; 68), Sharp
(1935: 160-161) and Hale (1982). Evans (1985: 17-18,
48)-492) elaborales the Gayardild system of
Bentinck Island and it is clearly of the same type
as both others throughout the Wellesley Islands and
also as Ganggalida, Garawa and Waanyi on the
mainland; Tindale’s failure to consider the \ssue of
this similarity (1977; 258-260) again renders his
general suggestions about the lengthy isolation of
the Bentinck Islanders jnadequate. Evans (L985:
21-22) in fact suggests “a relatively short period of
isolation’ of the Gayardild (500 to L000 years), based
partly on this matter of kinship system similarity,
However, Tindale (1977: 258-259) does refer
correctly to 1he marked lack of any system of tamed
sovial categories (or ‘class’ system as he puts it) on
Bentinck [sland in comparison with elsewhere
throughout the region, though his expression of this
fact Is at best imprecise in implying that the
surrounding groups had ‘sections’. Evans’ expression
of the comparison also lacks precision in suggesting
that the Mornington Islanders and those on {he
adjacent mainland had moieties and sections (Evans
1985: 17). What has operated throughout the region,
except on Bentinck Island, is a system of named
subsections (Trigger L985; 69-71, 350-340), though
this system can be said to be ‘organized as unnamed
patrilineal semi-moieties and moieties’ (Sharp 1935:
159; 1939; 455),
Putting aside the distinctiveness of the Bentinck
Islanders in this respect, and without discussing
social organisation in detail here, we may simply
note the situation of the coastal mainlanders
(Ganggalida), They were included within the
mainland blov stretebing to the Southwest in havine
both male and female subsection terms, and were
only partly similar to the inhabitants of the North
Wellesley Islands im thar the latter had male
Subsection terms only (Sharp 1935: 192, fn. 4). They
lacked the named senii-morety terms which
operated within at least che western parts of Garawa
country (Sharp £935, 1939; Reay (962; Trigger
1987a),
The coastal Gangyatida alse apparently lacked
the |nstitutionalisad role distinction of “ownership
and ‘managership’ in relation to land and ritual
property, which operated to the sourhwese (Thigeer
(985: Table 6). Although: it is difficult co reconstruct
thelr pre-contact ceremonial life because of more
recent influesices from the west, it is most likely that
the coastal Garpgalida also lacked the major cult
teremonies in whith the ownership/managership
distinction ts so important. In these respects, they
were identical to the inhabitants of all the Wellesley
Islands,
GENETIC EVIDENCE
Several publications have described similarities
and differences of a genetic nature between North
Wellesley Islanders, South Wellesley Istanders and
mainlanders, Among other findings, Simmons ef
a/, (1962) discuss the unusually high B blood
group gene frequencies among the Bentinek
Islanders, as compared with groups in the North
Wellesley Islands, However, in later work, Simmons
ef aj, (1964) did flirther comparisons with nearby
malnlanders and concluded thal the ‘Karawa’ also
had a high frequency of this gene, and in fact have
been ‘a main source in the spread of the B gene in
the mainland tribes of this remote area’ (p, GR).
This discussion about certain genetic features of
the Garawa people is nor without methodological
problems.’ However, the relevant point w be
considered here concerns claims based on this
genetic evidence by Tindale (1977: 254-255), He
asseris (hat the ‘Karawa’ constituted a ‘small
separate and {solated tribal envlave', and lived ir
a ‘perhaps refuge area’. These assertions fit with
Tindale’s suggestion that the ‘Karawa’, like the
Bentinck Islanders (and one other very distant north
Queensland group) witty whom they apparently
share certain blood type characteristics, are ‘relicts’
of 4 previously more widespread population with
this blood-type, Elsewhere, Tindale (1974; 122)
himself notes that his speculation that the name
‘Karawa' can be (ranslated as ‘uplanders” or ‘hills
people’, is ‘hazardous’, This is also the case tor his
other speculations about this group. Neither the
sociocultural evidence oor the mapped distribution
of Garawa estates im ny data, support the
Suggestion that Gatawa people have been in any way
isolated, although oral tradition must be reearded
as nal necessarily providing precise information for
historical periods greater than perhaps four
generations, Reay’s (1962: 95) comments about there
‘always’ having been ‘a great deal of contact,
invluding inter-marriage, between Garawa and
Anyula |[Vanyula]’, are similarly based on
ethnographic data. However, ipter-marriage
between these two language groups has also been
indicated jn genetic studles carrled oat by White
(1978: 42: 1979; 439, 445).
To summarise the discussion so far, the coastal
mainianders have been shown to be part of a
linguistic sub-group extending throughout the
Wellesley Islands, and imland for a substantial
distarice (at least 1D) ke southwards to the
INLAND, COAST AND ISLANDS 75
Nicholson River). To the west and south-west,
neither Garawa nor Waanyi (nor Yanyula on the
coast) are within this linguistic sub-group. A narrow
coastal strip adjacent to the Wellesley Islands is
classified in the Aboriginal view as containing areas
which are environmentally similar to close off-shore
islands, and is conceived as the separate domain
appropriate to ‘saltwater people’. Nevertheless,
substantial movement occurred between coast and
adjacent inland, as well as between the coast and
the North Wellesley Islands. The fact that the
Gayardild in the South Wellesley Islands shared
language and kinship system with others on the
mainland and in the North Wellesley Islands, means
that it is problematic just how isolated these
particular islands were. The same type of kinship
system extends throughout the study region,
however there are differences in the presence of
systems of named social categories. Finally,
published genetic data assert various differences and
similarities between populations in the study region.
While there has been a suggestion that the
Gayardild (Bentinck Islanders) and the Garawa may
be ‘elict’ groups of a previously more widespread
population, ethnographic and certain genetic data
indicate a lack of isolation of the Garawa.
Thus, my evidence certainly does not indicate any
simple pattern of relationships between the various
groups and cultural forms treated so far. We are
confronted with a complex pattern of overlapping
cultural forms extending throughout inland, coastal
and island societies. The paper now turns to a
detailed consideration of some aspects of material
culture. The evidence here indicates major
differences between coastal mainland and the inland
Garawa/Waanyi traditions. In a number of respects,
the coastal mainland traditions resemble those of
the off-shore islands (particularly the North
Wellesley Islands), in their differences from Garawa
and Waanyi traditions.
DIVERSITY IN INLAND, COASTAL AND ISLANDS
MATERIAL CULTURE
In this section I discuss a form of decorative
marking, the use of (and attribution of meaning
to) shell material, spearthrowers, spears, fishing
artefacts and technology, and watercraft.
Decorative marking
The marking known as jinanggliyari in Garawa/
Waanyi is said to be absent from items traditionally
produced in ‘saltwater country’, including both the
mainland coastal area and the off-shore Wellesley
Islands. The marking is a longitudinal fluted pattern
made by an end of the adze-type tool known as
biynmala. This tool consists of a gently curved stick
of varying length with stone blades attached at both
ends. One end is typically said to be a broad blade
(known as gubija), while the more pointed blade
is known as jinangyi, hence the name given to the
mark it makes.
Roth (1904: 20) refers to this tool as a ‘native
gouge’. He presents figures showing it and the
manner of its use (1904: Plate XIV, Figs 101, 105),
and notes the Lawn Hill [Station] name’ as ‘gobija’.
He does not, however, distinguish between the two
ends of the implement; although the figure he gives
elsewhere (1897, Plate XII, Fig. 235) for areas to
the south of the study region, possibly indicates the
blade at one end as broader than the other. In
dealing with this implement for areas to the west
of the study region, Spencer & Gillen (1904:
636-640) describe considerable variation in the size
and shape of the attached stone flakes, and discuss
the consequent different sizes of fluting made by
the implement. Contemporary Garawa and Waanyi
older people explain that the jinanggliyari mark was
commonly applied to the convex side of two kinds
of boomerangs: juguli, near-symmetrical shaped for
hunting (QM QE6218), and man.guburana, hook-
shaped for fighting (QM QE4274). The marking
was also applied to wooden coolamons, jugiva (QM
QES51), and sometimes to other items such as spears,
clubs and shields (Fig. 4). It is said to have helped
ensure that a weapon would ‘fly straight’, parti-
cularly when combined with the right song (cf.
Roth’s [1897: 146] comment concerning spears).
From the inland perspective, the mark described
above distinguished the item as belonging to the
network of language groups stretching to the west
from, and including, Garawa and Waanyi country;
although it is clear from Roth’s account, that it was
FIGURE 4. Three items showing the longitudinal fluting
mark known as jinanggliyari, which is absent from items
both on the coastal mainland and the Wellesley Islands,
(Courtesy Queensland Museum.)
16 DS. TRIGGER
also used in areas well to the south of the study
regioli (see also Mulvaney’s (1976: 81] summary map
showing ils distribution to the south and west of
the study region). [tems with this mark may have
been traded to the coastal ‘saltwater country’ (and
to the North Wellesley Islands) but the mark was
apparently not used by coastal artisans. Nor js iL
likely that coastal people commonly used the tool
which made the mark. Lt is absent From Meniniott’s
(19794) list of the Mornington [sland artefact
repertaire. Merimmott (1979a: 111) does state the
contemporary claim by some “Lardil’ people thal
the practice of imseribing fluting on hooked
boomeranes ‘was obtained from the mainland in
recent history’, and coastal mainlanders may also
have partially incorporated the marking during
post-contact times. Memmott (1979a; 113) alse
notes the wooden coolamon as of possible recent
mainland ongin, and like the ‘Lardil’, coastal
Ganggalida people appear to have only made
coolamons from Mefalewor bark (Roth’s ‘pleat-lype’
[1904: 30]), in pre-contact times.? The adze tool
which was absent from the islands and the coastal
mainland repertoires, was the implement typically
sed in inland areas fp gouge out wooden
covlamons, as well as to inseribe the fluted mark
on them,
Shell material
Shel! material is of course plentiful jn the coastal
environment, and the fourteen Ganggalida
classifications that | have recorded represent only
a portion of such knowledge. By contrast, the
intand perspective appears tc regard ‘saltwater shell”
as belonging to two broad size categories only:
raluinve — ‘big one’ and malduwa — small one’.
An example of the former is the large ‘bailer shell’
(Melo Amphora) used for carrying water leg QM
QE593, obtained from Allen Island), Roth (1904:
29) notes that on the Wellesley Islands ‘the ventral
curface of the last whorl of the. . ..shell is pierced
for insertion of the thumb during transport’.
Garawa and Waany! people are said to have not
customarily used this item. Inlanders know that
coastal people used both malduwa and rabunye
saltwater shells lo construct small and larger
(respectively) cutting and scraping implements, and
this is seen as having been necessary and
appropriate due to their lack of sujtable stone
material. Freshwater mussel shell wag apparently
used similarly, but only to a limited extent, in the
Inland areas. Shellfish also formed a more sub-
‘stantial part of the coastal diel, as compared with
the inland diet,
However, the difference in use of shell ntajenal
that inland people today remark upon, is the coastal
people's lack of shell pendants known by Inianders
as jaramara (OM QEI809, obtained from the Lawn
Hill-Nicholson River area in 1900), This item is
pearl-shell (Pinctada siaxitna), as are the others in
several museum collections identified by an inland
man as jaramara. These pearl-shell pendants
were received by Garawa and Weanyi people from
areas to the west and were considered items of great
significance. Thelr general Iniportance appears to
have been asa marker of certain formal social rales.
For example, two individuals ‘promised’ to each
other as marriage partners wore Jaraniara (for 9
short time at least) on the occasion of the promise
being formally and ritually recognised, A further
example isa woman wearing the item following her
child’s death, particularly in any assoctated rituml
settings. The /aramara thus marks its wearer as the
focus of ritual related to an |mportant social
situation, Occasionally, there may be a scratching,
or engraving on it which may indicate the tie to #
particular person, For example, such a scratching
on the QM specimen noted above was Interpreted
as representing the hand and extended fingers of
its custodian. These items are inherited from
parents, grandparents and other kin depending on
circumstances, While not always considered highly
secret, access to them Is restricted.
It is thus the exclusion of coastal people froms the
customary teceipt of these items from west of the
study region, Which distinguishes them from the
inlanders. Senior Garawa and Waanyi people now
say that Lhe items were not passed on to Ganggalida
people ‘because they didn't know’. In these respects,
the situation of the coastal mainianders was
identical with that of the Wellesley Islanders."
Spearthrowers
While inland artisans made three types of
spearthrower, the coastal people along with all the
occupants of the Wellesley Islands, made only one
of these. It is known in Garawa, Waanyi and
Ganggalida as miurnigu, and Menmott (19793; ILD)
reports the same lerm (‘nrurraku’) for “Lardil’, Wis
cut from one straight piece of cylindrical wood so
that the distal end has a raised notch which slips
into the hole typically at the back of spears (QM
QEL6/113, see Fig, 5), Roth (1909: 200) describes
it in some detail and refers to it as ‘a very primitive
form of implement met With in the Wellesley
Islands, and on the adjoining mainiand in the
neighbourhood of Burketown , , .’, and illustrates
itin his Plate LVI, Fig. 14 (and see also Roth 1902:
15). As Roth mentions, this is a very light
implement; contemporary people say it is
commonly made from the farrgwida tree (Thespesia
pozgminecides), or one of several other light woods.
Presumably Roth designates it as ‘primitive’ because
of the notion that its construction is simple relative
INLAND, COAST AND ISLANDS 77
QE Als}
mere
FIGURE 5. Three types of spearthrower made in the study
region. The bottom one (QE 16/1113) was made
throughout mainland coast, island and inland areas, while
the other two were made only in the inland areas.
(Courtesy Queensland Museum.)
to other types, or at least somehow less developed.
This is further made clear when he notes (1902: 15)
that it is identical (except in its size) with a ‘toy
wommera’ used commonly throughout north
Queensland by boys emulating the fighting
behaviour of men. However, both coastal and
inland people state its advantage over other types
of spearthrower to be that there is no problem with
a separate attached peg coming loose. It can not
be seen as under-developed in terms of functional
performance.
The two further types of spearthrowers made and
used by Garawa and Waanyi people, but not in the
coastal mainland area or on the islands, are termed
ngaliga and wujula. The former is again a
cylindrical straight piece of wood, but with a
separate wooden peg (known in Garawa/ Waanyi as
ngurru) attached to the distal end, usually with gum
(QM QE11/53, obtained from Turn Off Lagoon on
the Nicholson River). The latter type is flat and
linear with a large notch forming a spatulate
proximal end. The peg is attached to the distal end
at an acute angle to the face (Robins 1980: 58), (UQ
1813; see Fig, 5). Roth (1909: Plate LXI, Figs 1-4)
describes these two types and notes (pp. 200-201)
them as ‘occasionally to be found in the area around
Burketown, but certainly not of local
manufacture, being brought in from eastward’.
However, I am sure the editor’s footnote replacing
‘eastward’ with ‘westward and southward’ represents
what Roth intended; though it can be noted that
the editor’s correction was overlooked in McCarthy
(1939: 419). My information is certainly that these
two latter types were made and used in Garawa and
Waanyi territory to the west and southwest of
Burketown.
Spencer & Gillen (1904: 668-670) describe and
illustrate the ngaliga type as often found to the west
of the study region, and state that it is traded
‘eastwards towards the Gulf’. The one they illustrate
is in fact from the ‘Anula [Yanyula] tribe’ so the
type apparently reached the saltwater domain to the
west of the coastal Ganggalida area. It is shown
with a human hair-string tassel at the handle end,
and, although Roth (1909: 201 and Plate LXI) also
shows such a tassel, this is now said not to have
normally been added by Garawa and Waanyi
people. In another publication, Spencer & Gillen
(1969 [1899]: 578-582) give the name for this type
(said to be made by ‘the Waimbia [Wambaya] tribe’)
as ‘Nulliga’, clearly the same term as ngaliga given
above. In both publications, Spencer & Gillen also
show the wujula type in areas to the west of Garawa
and Waanyi territory.
While neither of these latter two types are now
said to have been customarily manufactured by
coastal people or Wellesley Islanders, some coastal
incorporation of these items and the skills to make
them did occur during post-contact times. One
museum specimen (UQ 1814) of the ngaliga type
obtained in the 1940s, is recorded as having been
made by a man at Mornington Island who learned
about the style from mainland sources. 2
Spears
Four types of spear can be distinguished. In
general, spear shafts are now commonly known as
mugura, but are distinguished according to the kind
of tree they are cut from. However, spear types are
distinguished mainly according to the nature of
their heads.
The first type was used throughout inland, coast
and island settings; it is known by the word for
prong in all languages, and usually two or three
prongs are attached to a shaft (QM QEII/53, see
Fig. 6). The manner of construction described and
illustrated by Roth (1909: 190-191) for the Wellesley
Islands, appears applicable to both the coastal
Ganggalida and the inland Garawa and Waanyi.
(However, with the incorporation of European wire,
the prongs have been commonly attached by
wedging them into a hole made in the end of the
shaft.) To summarise the earlier technique, the
wooden prongs are fitted neatly against grooves cut
along the shaft and then tied with string. These
spears were used mainly for fish, but also for
animals such as goanna in some circumstances, in
both coastal and inland contexts.
The second type of spear head (babagana
[Garawa] and jimindi [Waanyi], but not known by
a term in Ganggalida) is stone, and the spear was
used to hunt large animals like kangaroo and emu
(QM QE4278, obtained in the Burketown area), The
stone items are said not to have been manufactured
in the coastal area, but were apparently procured
by coastal people from various inland sources. They
were thus used by inland and coastal people both
as knives and as spear heads fitted to shafts (cf.
78 D. 8. TRIGGER
serrerses
FIGURE 6. Four types of spear made in the region. The
QE 4766/1 type was not used in inland areas, while the
stone flakes for the QE4687 type had to be imported into
the coastal area. (Courtesy Queensland Museum.)
Spencer & Gillen’s [1969 (1899): 592-593; 1904:
640-656] accounts of the manufacture of these
stone blades and their multiple uses, upon which
McCarthy [1976: 35] appears to rely). Roth (1904:
18, 22) notes that the stone blades were made ‘along
the ranges up the western border north from Lawn
Hill, etc.’, and that small bundles of them rolled
up in tea-tree bark were bartered. He specifically
mentions (1909; 190) ‘the stone-spears of Burke-
town, Point Parker [ie. coastal Ganggalida
country], and the ranges along the Queensland
Northern Territory Border [i.e. inland Garawa/
Waanyi country]’; and, it may be noted, he also
comments (1904: 22) on a large import’ of such
items occurring ‘from the Northern Territory, across
the border from Wollogorang [Station] southwards’
to his North-West-Central districts.
Tindale (1974: 122, 228) states that ‘Karawa’
people traded stone knife blades to the east
generally, and that Mornington Islanders sent young
marriageable girls in return for these items. He also
mentions McCourt’s claim (published soon after
[McCourt 1975: 80, 108]) to have discovered the
origin of the long stone spear points found by
Tindale ‘on Bentinck Island . . . and on the adjacent
[mainland?] coastline’; McCourt says these were
quarried and made in the Wollogorang-Calvert Hills
Stations area, on the Calvert River, near Redbank
Mine, etc, — that is, in Garawa territory.
Contemporary old Garawa people certainly state
that a number of places, including this area, were
sources of stone used for both long knives and spear
points. Indeed, they take pride in the Garawa
knowledge of the manufacture of various kinds of
stone tools.
However, how far the stone blades made in
Garawa territory reached, remains an open
question. Although a comment from Roth (190la:
4) states that strong shell tools take the place of
knives and scrapers on Bentinck Island, ‘as on the
mainland’, his statement (quoted above) about the
presence of stone spear-heads in coastal Ganggalida
country is quite unambiguous. Yet none of Roth’s
data describe these items for the Wellesley Islands,
nor does Memmott’s (1979a) recent work with
contemporary ‘Lardil’ people mention them. Thus,
Tindale’s assertion that the stone spear-blades
reached Mornington Island may be incorrect.
Tindale (1977: 267) himself notes that such stone
items were never used on Bentinck Island, though
he was shown one on Sweers Island which had been
brought by visiting mainland Aborigines accom-
panying European men,
The third type of spear is known in Garawa and
Waanyi as ngarrgidigidi (ngarrgadaba =to spear),
(eg QM QES56/1, obtained from Turn Off Lagoon).
The long wooden head is barbed on one or both
sides, making it difficult to remove. This is a
fighting spear, said to have been used in a variety
of conflict and dispute contexts. Coastal
Ganggalida people are said to have similarly used
this spear. Memmott (1979a: 110) also describes it
for the ‘Lardil’.
The final type!3 was apparently not made or
customarily used by the inland people. It consists
of a light wood shaft attached to a long smooth
hardwood point of approximately the same length
as the shaft (QM QE4766, Nos 1-7). The method
of attachment is apparently unusual (R. Robins,
Queensland Museum, pers. comm.), being the
hafting together of the two sticks whose ends have
been sliced at much the same angles. The join is
bound in a distinctive fashion and no gum is used.
This method of attachment is quite different from
that customarily used by inland people. The spear
was used for large marine animals such as dugong
and sea turtle and is known in Ganggalida as
miyalda. Memmott (1979a: 110) describes the same
spear for the ‘Lardil’ on Mornington, where it is
known as miya.
Fishing and other methods of obtaining aquatic
foods
Both inland and coastal people used pronged
spears and nets to obtain a wide range of aquatic
foods. A large amount of knowledge is particular
to certain environments, and this is especially so
in relation to ‘saltwater country’. Yet coastal people
clearly also used the same techniques as inlanders
when ranging into the vicinity of fresh waterholes,
for example, ‘poisoning’ techniques involving the
placing of particular plants in the water, which
cause fish to float to the surface where they can be
easily obtained. As well, not all practices are the
result of environmental determination. For example
the ‘squeezing’ of stingray flesh in the coastal area
or swordfish flesh in both saltwater and freshwater
INLAND, COAST AND ISLANDS 79
areas, appears to be done solely because the taste
of the meat is preferred without the ‘sticky stuff?
which comes out when it is squeezed.
Nets were used by both inlanders and coastal
people. Despite size variation they were all made
from string which was produced by rolling together
either strands of certain grasses (QM QE1912), or
strands of bark from certain trees — the bark of
Hibiscus tiliaceus is particularly useful for this
purpose in coastal areas, but other trees such as
kurrajong (Brachychiton sp.) or black wattle
(Acacia hemsleyi) are used throughout the study
region. There are two methods of net usage in both
coastal and inland areas. The first involves four
people, one at each corner of a big net, dragging
it so that two corners are held near the bottom and
two at the surface, and the two ends are moved so
that the net forms part of a circle, eventually
dragging its contents on to the shore. In the second
method, two sticks are attached to two sides of the
net which is moved vertically through the water
while other people frighten fish into it; when a
weight is felt against the net, it is taken to the
shore. !4
It remains uncertain whether line and hook
fishing was done in this region. Roth (1903: 5) states
that ‘before the advent of the Europeans, such
articles in any material (shell, bone, etc.) were
unknown’. Some Aboriginal people today agree
with this view. Some remain unsure, and others
claim that hooks and lines were in use prior to
colonisation. Indeed, a shell type of hook certainly
exists, obtained in 1882 from the Settlement Creek
area (QM QE5924, see Fig. 7): a grass string line is
attached to a groove cut around the top of the shank
of a shell hook, and what are designated in the
associated documentation as several shell sinkers,
are tied to the line. Roth’s (1901b: 20) general
comment for other areas where he found usage of
fish hooks, that weights or sinkers are never used,
is thus also problematic. It seems most likely that
the hook shown in Fig. 7 was an isolated recent
import into the region, possibly originating from
the Torres Strait Islands. !5
The term wardugu has been given by several
contemporary old people to refer to fishing lines
complete with hook (cf. Keen’s [1983: 294]
translation of the term as ‘fish, hook, wire, line’),
and while the term bala is given for the hook itself,
it is more generally applied to any kind of forked
shape, whether as part of an implement or simply
the fork formed by tree branches. Some people
today, both inlanders and coastal people among
them, state that hooks were also made from emu,
kangaroo and pelican bone. One specific account
from two older Garawa men is that ‘goanna collar
bone’ and ‘catfish jaw bone’ are suitably hook
shaped and were used as such by Garawa people.
‘molt 2 S & S06 Gee Boyt) fe es
FIGURE 7, Fishing line with shell hook and sinkers,
collected in 1882 in the Settlement Creek area. (Courtesy
Queensland Museum.)
Various kinds of grub, frog, insect and shellfish are
now used as fish bait, as well as the fruit from the
fig tree (Ficus racemosa). If line fishing has indeed
developed since the early phases of European
impact, it represents an innovation which has
occurred throughout inland, coastal and island
settings.
Of particular interest in this region is a large
complex of stone fish traps on the Wellesley Islands
and the adjacent mainland coast. Stone walls as
traps are said to have been constructed across inland
watercourses at narrow places, and these were
mainly used during ‘flood time’ (the wet season)
when the stream ran strongly due to the extensive
wet season rains. More temporary sapling fences
were also constructed across both inland and coastal
watercourses; in the coastal area stakes were stuck
in the bed at an angle so that the receding tide
would push deliberately placed bushes and other
debris against them. However, the inland traps
appear to have been much less important in the
food production process than their coastal
equivalents.
The extensiveness of the stone traps throughout
the islands and adjacent mainland coast, suggest
their importance in the food production process.
Over approximately 470 km of coastline, 334 traps
have been recorded, one trap per 1.4 km of coast-
line (although there is considerable variation
in the density of traps throughout the islands and
adjacent mainland); (see Robins et al. [n.d.] for
details of the physical features of the traps, their
distribution and relevant ethnographic data).
Figure 8 shows one of the mainland trap sites (at
Bayley Point), This series of traps extends along the
shore-line for approximately 400 m, and from the
shore-line up to approximately 150 m out on to
mudflats, to the edge of an extending rock platform.
The receding tide would leave contained (if not
RO DLS. TRIGGER
FIGURE &. Stone fish trap complex ar Bayley Point on
the mainland coast opposite the Wellesley Islands.
(Courtesy Connah/Jones Aerial Archaeology, University
of New England.)
exposed in some instances) fishes (including shark
and stingray), and at times sea turtle and dugong,
On a visit to these traps in 1983, Ganggalida people
also used a pronged spear to obtaln crabs from
Within crevices under the trap wall, The stone wall
traps represent a highly developed (and apparently
durable) form of food production technology,
which is distinctive of the coastal economy of the
mainland and the Wellesley Islands,
Watercraft
The distinctive triangular shaped raft of the
Wellesley Islands area (known in ‘Lardil’ as walpa)
has been described in various sources (eg.
Memmiott 19792; 111), [1 is clear from contemporary
accounts that these were also used by enastal
mainlanders opposue the islands who refer to it ax
walbywe. Roth (1908: 161) confirms this fact, and
describes the call as:
formed of atimerous logs of while Mangrove’ (ied
toxelher al the baits as well as ar the extremities, with
the result that it is much narrower foreward than al the
stern, On top is placed some sea-weed, a sort of cushion
tor (he voyager to sit upon. With such frail craft the
[Aborigines] will not only visit island and istand, but
even cross over (o the mainland, usually on the one
course, making fora spol somewhere in the vicinily of
Poine Parker.
Elsewhere, Roth (1903: 3) notes again that: ‘There
is no doubt whatever that Communication goes on
here [irom Forsyth Island] between the mainland
via Raines [later to be called Paines] and Bayley
Islands and Bayley Point’.
Davidgott (1935: 40), 10 his survey of Aboripinal
watercraft, states that apart from in this relatively
linnted area of the southern Gulf of Carpentaria,
this type of raft existed elsewhere only on a section
of the north-western Australian coast, and the
question of its advantages and disadvantages for
those who adopted it remains an interesting research
issue. Davidson (1935; 39-45) reconsteucts an
liistori¢al developmental sequence for watercraft
where the triangular raft was invented earliest, but
had then ‘given way’ along most of the northern
Australian coast to the supposedly more sophistl-
cated and preferred sewn bark and dug-out canoes,
He makes no direct comment on why the raft was
retained tm two regions: However, a cautionary
comment is surely required here against the notion
that the rafts would haye necessarily been inferior
to the canoes, The pointis similar to the one made
above concerning Roth’s attribution of primitive-
ness to the coastal (and islands) spearthrower. While
the raft may appear to involve less sophisticated
construction techniques than the canoes, an
assumption that the canoes were better suited than
rafis for travel in all circumstances may well
overlook certain advantages possessed by the latter;
For example, .a feature of the rafts, particularly the
larger ones, may have been greater stability which
may have enabled safer transportation avross
shallow sandbanks and channels, of larger numbers
of children and valued items (including burning
cinders to be used to re-kindle fires}.
Within the study region of this paper, Ganggalida
people on the coast just io the west of the Wellesley
[slands are said to have used the raft, but to a much
more limited extent than the Gananggalinda people
directly opposite the islands, [t was not maniufac-
tured or used by the inlanders. Garawa people are
said to have been familiar with the manutacture of
the sewn bark canoe (termed by them wulganyi)
from the bark of the messmate tree; however, they
only used it occasionally, most likely when moving
over wide sections of the lower reaches of such
watercourses as the Robinson and Calvert Rivers
and Settlement Creek, While Gangpalida has a
similar term for this type of canoe (wu/gunda), the
coastal people are said to have used it even less than
Garawa people. No suitable messmate bark |s
apparently available in the coastal country. The
sewn bark canoe was thug used only te a limited
extent east of Yanyula territory, and the above data
fill the gap mentioned by Davidson (1935: 35)
concerning |ts distribution west of the Wellesley
Islands,
It was the Yanyula people from the coastal areas
far to the northwest of the Wellesley Iskands, wha
are now regarded throughout the region as having
been the experts with bark canoes. It was they also
who came from the northwest in dug-out canoes,
which had apparently diffused as far eastwards in
the Gulf as the vicinity of the Sic Edward Pellew
Islands in Yanyula territory, having been initially
INLAND, COAST AND ISLANDS 81
introduced into northern Arnhem Land by
Macassan visitors well prior to the European
presence in the Gulf (Davidson 1935: 20-24). Dug-
outs are now known generally by the same term,
muwarda, as that given to all European-style boats
regardless of their size. Whether Keen (1983: 278)
is correct in asserting that this term (her ‘muwata’)
derives from the English word ‘motor’, it is most
likely that dug-outs were known in the study area
before motor-driven boats were brought in by
Europeans. Contemporary coastal Ganggalida old
people know how dug-outs were made from large-
girthed trees of certain kinds (for example,
Canarium australianum). However, while such trees
may be found in their country, only a small number
of dug-outs were apparently made here following
the introduction of steel axes and the increased
eastwards migration of Yanyula people with dug-
outs after colonisation.
Material culture — a Summary
In summary, a longitudinal fluting mark was
used in decoration by Garawa and Waanyi artisans,
but not in the Ganggalida mainland coastal area
nor in the Wellesley Islands. This mark is now
recognised as distinguishing artefacts as belonging
to the cultural bloc extending from Garawa and
Waanyi territories to the west and southwest. The
adze-type implement used to make the fluting was
not manufactured in the coastal or island areas, and
hence the wooden coolamon could also not be
produced, There was no established stone tool
working tradition on the coast or in the islands,
whereas Garawa and Waanyi people speak with
pride of the stone tool production techniques used
in earlier times. Shell cutting tools appear to have
taken the place of stone tools along the mainland
coastal strip and in the islands. Pearl shell pendants
were received by Garawa and Waanyi people
through trade from far to the west and southwest,
but these were not passed on to coastal Ganggalida
people or to Wellesley Islanders.
Coastal mainlanders and Wellesley Islanders
made only one type of spearthrower, whereas the
inlanders produced two further types as well. Of
the four spear types in the study region, two were
used across inland, mainland coast and island areas,
one was produced only on the coast and islands,
and one (with a stone blade as the head) was
produced only among inlanders, although mainland
coastal people (and possibly North Wellesley
Islanders) received the stone blades through trade
to a certain extent.
Aquatic resources were obtained by use of spears
and nets throughout all parts of the study region
(and line fishing appears to have been adopted
across the region, with the probable exception of
the South Wellesley Islands, from early phases of
European colonisation). Large stone wall fish traps
were used only in mainland coastal areas and
throughout the islands. Unlike both inlanders and
Yanyula coastal people to the northwest, coastal
Ganggalida people and Wellesley Islanders used
only rafts as watercraft.
CONCLUSION
This paper has compared aspects of inland,
coastal and island societies. The most detailed data
have been given for traditional material culture, and
this has indicated a substantial degree of similarity
between coastal mainland and islands societies,
compared with nearby inland society to the west
and southwest. As ‘saltwater people’ occupying
‘saltwater country’, the coastal mainlanders can be
regarded as part of the island cultural bloc in
significant respects. Yet the material culture of
coastal mainland society has simultaneously
emerged as similar to that of inland society in
certain respects, and if further aspects of material
culture were to be studied closely, a number of
differences between mainland coast and the islands
could well become evident.!© While the material
culture repertoires of the mainland coast and islands
are strikingly similar in significant ways, it would
be inaccurate to define them collectively as a
homogeneous material culture tradition completely
separate from that of the inland.
The same point was made when summarising the
comparative data for the region on language,
kinship and social organisation, and genetics. The
paper has described a complex pattern of over-
lapping cultural and other forms, with no area
emerging as completely distinctive. This general-
isation includes the situation of the Bentinck
Islanders, despite the literature in support of their
alleged lengthy isolation, At least wN% ffdemon-
strated otherwise, we must recognise that the differ-
ences between Gayardild society and that on the
North Wellesley Islands and the coastal mainland,
are apparently no more substantial than the
differences between the mainland coast and inland.
The main concern of the paper has been to
consider the situation of coastal mainland dwellers
adjacent to major off-shore islands society. The
generalisation of this paper is that the relationship
with the North Wellesley Islands societies in
particular, has reinforced the maintenance of
cultural differences between coast and inland
societies on the mainland. This is not to argue for
uni-directional influence from the islands to the
coastal mainland. Indeed, some of the data indicate
major areal influence from the mainland to both
the North and South Wellesley Islands (e.g. the
existence of an Aranda type kinship system
RZ B.S, TRIGGER
throughout the study region).. However, the North
Wellesley Island societies should pot be viewed as
without influence on mainland coastal socrety, In
this sense, 1 am arguing thal those mainfand-island
relations which entail regular contact should be
viewed as similar to relations among peaples on the
mainland, It may be that the envirorimental simi-
larities in this region between the mainland coast
and parts of the off-shore islands, particularly
engender the cultural similarities dealt with im this
paper (especially in the aspects of material culture
traditions Which have been considered), However,
the implication for studies in other regions is that
when comparing island and mainland societies in
Aboriginal Australia, particular attention should be
paid to the environment and society of the
immediately adjacent mainland coast, as well as to
societies located further inland..
ENDNOTES
|. The name of this language has been given as “Yukulta’
in the literature (Keen 1983), However, the term
‘Ganggalida’ was predominantly tised for the language and
a group of people at the lime of my research, The larter
term can be translated as meaning language’ or ‘talk’ (see
Trigger (L985: 340-49} for further discussion of ihe
relationship between these twa verms),
2. The language of the Mornington Islanders is otore
conventionally spelt ‘Lardil’ (see eg, Memmott 1979p).
However, except where quoling from works using this
spelling, | will use the mainland Aboriginal expression
‘Layardiida’
4. Elsewhere (Trigger 1987b), | have dealt if detail with
the problematic issues entailed in using Janpuage names.
in designate tr(bal” Units if this region, In this paper such
cxpressions represeat a predominant View among
Aboriginal residents which glosscs separable areas of land
and associated bhadiles of tradnion by the use of language
wares.
4 Indeed, Evans (1985; 7-8) describes the “basic
‘fsolation™” (\.e. distinctiveness) of the broader ‘Tangkie
subgroup' of languages, meluding those throughout the
islands and Ganggatida and Neuburinli oo (he maintand,
when vompared with other Aboriginal languages.
5. Evans (pers. comm.) has speculated that the clymolagy
of this term may be derived from kanatva — to buen’
and kafitva = Yo junp” (sce Keen 19R3: 272) fence, these
people may have been known by a term for ‘ares
jumping up’ or bushfire at times visible fram (he vantage
point of other coastal peaple,
6, This is not to imply thal coastal mainlanders vagarded
even the small close off-shore islands (Bayley and Pains)
as withoul environmental features differentiating them in
more specific respects frown (he ‘islands’ in maintand
Saliwater country’ Certainly, the larger offshore istinds
are much more diverse environments than both the smaller
off-shore islands and the sand ridge ‘island’ areas within
the mainland ‘saltwater country’; (see Memmott [1979b:
45-65) for a discussion of the land forms and ecology for
Mornington, and Tindale |)962a; 280-288] for relevant
information concerning Bentiack). [It would be interesting
fo investigate whether the larger off-shore islands are
designated as ‘slands’ (murndarmurra) in the coastal
mainland perspective. Minimal data indicate that they are
in fact regarded as wambalda (or and), hough coastal
mamlanders would presumably identify as siurndamurra,
areas within the larger off-shore islands similar to the sand
ridges In mainland ‘saltwarer country’,
7, Simmons e/ @/, (1964: 75) themselves point out for the
whole population tesied in this region, that ‘about one-
half... were of mixed tribes and, of course, (heir earlier
history ts unknown! Elsewhere (Trigger L987b), | have
discussed the necessity of examining the linguistic (or
tribal’) aljiations of individuals’ four grandparents if
this matter is to be adequately researched. The basis on
which individuals state primary affiliation with, say,
father’s language rather than mother’s, involves a variety
of social factors. Hence the possibility of defining clearly
a relatively enduring endogamous population in ‘Wibal’
terms, from the slalements of individuals, is fraught with
diffieulries.
8. Throughout the paper, items located in museum
collections are identified by placing their registiation
identifications iF parentheses, The letters ‘QM! indicate
that the item isin the Queensland Museum collection, and
UQ' indicates the University of Queensland Anthropology
Museum cotlection,
9 Roi obtained wooden coolamons at Burketown in 1901
aod Forsyth and Mornington Islands in (903 (items 13359,
13360, 1336) respectively, in the Australian Museum
register af the Roth Collection), however it is mast likely
thal (hese would have been manufactured in inland areas.
LO. Mr Tommy George, of Waanyi and Qarawa descent,
inspected museum collections in Brisbane and Canberra
9 1980-ayid [) Brisbane during March 1984; some of the
information he provided js confained in a transcription
of an audio tape made alter his 1984 work (George 1984),
11, Several authors (see Mutlvaney’s (1969: 96; 1976: 83)
summary maps) have documented peart-shell as an rem
traded over very long distances, and by (hese accounts iL
is possible that jt alsa came into the study region from
af caglerly direction via Cape York Peninsula, However,
itis rather the accounts of It coming originally (hur the
very distant Kimberley coast (eg, McCarchy 1939: 96-98)
which aré consistent with contemporary Aboriginal
opinion in the study region,
1, Menmott (1979b; Table 7) notes ‘wuijala'as the name
given by ‘Lardil' people to a spearthrower of the typical
wert Cane York kind with attached shells atthe proximal
end, and he says they reacted Morpington Island only in
recent times. While it appears the “Lardil’ have thus
borrowed his term. both mainland coastal and inland
groups remain unfamillar with rie Cape York type and
INLAND, COAST AND ISLANDS RS
would mast likely simply referto it as being fram ‘another
commry’,
13, Rowh (1902; 15) refers to a kind of toy spear used in
games fy young boys: ‘so far observed . , . only amang
the coastal blacks (o {he west of Burketown, and at
Wollogoning (Northern Territory border)’, However while
the few peaple of whom f have enquired stale thal it was
used, | have no dala to compare with Roth’s description
or ii,
14. The large ‘dugong net’ described for the “Lardil! by
Memmoitt (1979: 112, 118) does fot appear to be known
by contemporary Ganggalida people, though whether it
was Once used in the coastal mainland area remains
unvertain,
15, The hook collected from the region (Pig. 7) appears
to be of the ‘bent pin’ type which is discussed by Massola
(95: LL, 15) as having been possibly intraduced from the
Torres Stralt, Anell (/955: 14-115) says that all fish hooks
in Australla Were ‘undoubtedly’ intraduced from che Torres
Sinait, and suggests that this accurred ‘relatively dale’, that
16 during the recent past.
(6. Memmott's (1985) ongoing work comparing ‘Lardil’
and ‘Kaiadili' material culture repertoires indicates some
significant differences between aspects of the material
culture of the North and South Wellesley Islanders, the
most striking being (hat (he Gayardild had a much smaller
number of artefacts. Lt is most likely that further research
would indicate that the Gayardild also diftered tran the
mainland coastal Ganggalida in this respect,
ACKNOWLEDGMENTS
Research In the study region from early 1978 to 1983
was supported by the Department of Anthropology and
Sovuidlogy, University of Queensland, and the Australian
Institurce of Aborizinal Seudies, The staff of the
Queensland Museum, particularly Mr Richatd Robins and
Ms Julia Findlay, and of the University of Quoensland
Anthropolagy Museum, parneulacly Ms Linely Allen,
generously assisted in the location of items in the
calleclions. Mr Richard Robins, and Dr Paul Memmott
(Department of Architecture, University of Queensland)
have provided helpful comments. Mr Tommy George, of
Waanyi and Garawa descent, took particular interest in
this research, and provided much Information during visits
to museum collections in Canberra and Brisbane,
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SIMMONS, R‘T., GRAYDON, J.J. & TINDALE, N.B.
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Forsyth Islands and the mainland, Gulf of Carpentaria,
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SPENCER, B. & GILLEN, FG. 1969 [1899]. ‘The Native
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TINDALE, N.B. 1962b. Some population changes among
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people of Bentinck Island, Gulf of Carpentaria,
Queensland. /n J. Allen, J. Golson & R. Jones (Eds.).
‘Sunda and Sahul: Prehistoric Studies in Southeast Asia,
Melanesia and Australia’, Pp. 247-273. Academic Press,
London.
TRIGGER, D.S. 1982. ‘Nicholson River (Waanyi/Garawa)
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217-238.
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437-454,
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES :
DROMATINAE)
BY C. PATTERSON & P.V. RICH
Summary
The oldest known emu is Dromaius gidju n. sp. from the medial Miocene Kutjamarpu fauna at Lake
Ngapakaldi in northern South Australia. This form, based on a partial hind limb, is smaller and has
relatively shorter and less mediolaterally compressed hind limb bones, and less reduction of the
medial and lateral digits than in the living form. D. gidju thus appears to be less specialized for a
cursorial lifestyle, being somewhat intermediate between the forest dwelling cassowaries and the
highly cursorial living emu, D. novachollandiae. Fossils from the Late Miocene and Early Pliocene
may be allied to D. gidju, but more material is needed to allow confident assignment. D. ocypus
from the medial Pliocene Palankarinna fauna at Lake Palankarinna, northern South Australia, is
intermediate in size between D. gidju and D. novaehollandiae. In addition, its tarsometatarsus is
decidely shorter relative to width than that in D. novachollandiae, thus indicating it is not as highly
adapted for cursorial life as the living emu. Essentially all other emu fossils, Late Pliocene-Recent,
appear to belong in D. novaehollandiae including : D. ‘patricius’, D. “gracilipes’, and ‘Metapteryx
bifrons’, all defined originally by C.W. De Vis. The only exceptions are the King Island emu (D.
ater) and the Kangaroo Island emu (D. baudinianus). Whether there was greater size variability in
Pleistocene emu populations and whether a separate species of emu once inhabited Tasmania are
problems yet to be resolved once larger collections of both living and fossil emus can be measured
and analyzed.
‘THE FOSSIL HISTORY OF THE EMUS, DROMATUS (AVES: DROMAIINAE)
C. PATTERSON & P, V, RICH
PATTERSON, ©, & RICH, PV. 1987. The fossil history of the emus, Bronraius (Aves:
Dromaiinae). Rec. S. Aust. Mus. 21(2); 85-117.
The oldest known emu is Dramaius gidju n. sp. from the medial Miocene Kutjamarpu fauna
at Lake Ngapakaldi in northert: South Australia, This form, based on a partial hind limb, is
smaller and has relatively shorter and less mediolaterally compressed hind limb bones, and less
reduction of the medial and lateral digits than in the living form. D. gidju thus appears to be
less specialized for 4 cursorial lifestyle, being somewhat intermediate between the forest dwelling
cassowaries and the highly cursorial living emu, D. novaehollandiae, Fossils from the Late Miocene
and Early Plidcene may be allied to D. gidju, but more material is needed to allow confident
assignment. 2. ocypus from the medial Pliocene Palankarinna fauna at Lake Palankarinna,
northern South: Australia, is intermediate in size between D. gidju and D. nevachallandiae, In
addition, its tarsometatarsus is decidedly shorter relative to width than that in 2, nevaehollandiac,
thus indicating that it is not as highly adapted for a cursorial life as the living emu, Essentially
all other emu tossils, Late Pllocene-Recent, appear to belong in D. nevaehollandiae including:
D. putricius! D. ‘eracilipes; and Metapteryx bifrons; all defined originally by CW. De Vis. The
only exceptions are the King Island emu (D. afer) and the Kangaroo Island emu (.D. bavdiniunus),
Whether there was greater size variability in Pleistocene emu populations and whether a separate
species of emu once inhabited Tasmania are problems yet to be resolved once larger collections
of both living and fossil emus can be measured and analyzed.
C. Patterson, Department of Zoology and P. V. Rich, Departments of Zoology and Earth Sciences,
Monash University, Clayton, Victoria 3168. Manuscript received 19 August 1986.
The living emu (Dromaius novaehallandiae) is B.P, Years belore present
the second largest living ground bird, exceeded only c. Cranium, crania
by the ostrich in size. Today and in the past, emus Cor. Coracoid i).
have been restricted to Australia, and their origins CSIRO =~ Commonwealth Scientific and
Industrial Research Organization,
Division of Wildlife and. Rangelands
Research, Canberra
are nol understood.
The fossil record of emus begins in the Miocene,
with two now extinct species occurring one each in d Distal
the Pliocene and the Miocene of northern South diapop. Diapophbyses
Australia, The Quaternary King Island and est, Estimated
Kangaroo Island emus seem to belong in two EB Femur
separate species. All other fossil emus, mainly Fib, Fibula
Pleistocene, however, are very similar fo and most HM Hunterian Museum, Glasgow
probably conspecific with the living D. novae- Aurn, aon ;
hollandiae. It is very likely, however; that the history hd ~_ f
of emus on the Australian continent is much older M Mandible
than curtently understood because of the general MM Geological and Mining Museum,
lack of a pre-Miocene terrestrial record. Sydney
Although the Pleistocene emus are currently NMY Museum of Victoria, Melbourne
indistinguishable from the living emu, the Tertiary p Proximal
species are distinct, The hlnd limb of the single Ph. Phalanx, phalanges
Miocene form is not as cursofially adapted. This postzyg. Postzygapophyses 7
species hag a tarsometatarsus that is shorter and QM Queensland Museurn, ageing
more robust, avd the lateral and medial digits of QVM ace Victoria Museum and Art
: = ery, Launceston
the foot are not as reduced as in the living emu, Rir} Right
This paper reviews fossil emu material and SAM South Australian Museum, Adelaide
outlines the major evolutionary trends demon- SIAM — Smithsonian Institution - American
strated by the dromaiines during the last 20 million Museum of Natural History
years, Expedition Field Numbers,
The following abbreviations are used: Washington, D.C. and New York
AM Australian Muscum, Sydney Sk. Skeleton(s), many skeletal elements
AMNH = American’ Muscum of Natanil History, St. Sternum
New York Syn, Synsacrum
86 C, PATTERSON & PV. RICH
T2, 73, T4 Trochleae (1, [t, LV
Tih,
Tiblotarsus
Tr. Tarsometatarsus
Ku Vertebrate)
UCMP = University of California, Museum of
Paleontology, Berkeley
WAM Western Australian Muscumt, Peeth
PRewous Work
There is surprisingly little in the literature
concerning the fossil emus of Australia {see Table
1). The first reference to a specimen supposedly
related ta emus was, in fact, a moa, ‘Dinornis
queenslondiae’, described by De Vis (1884) from the
Darling Downs, Queensland, Some later workers
considered [his specimen (e.g, Hutton 1293, Miller
1963) to be related to the emus and cassowaries-
Scarlett (1969), however, Found the fossil could be
assigned ro Pachyornis elephantopus, probably
collected from a midden on South tsland, New
Zealand, and thus it is not a valid Australian record,
and certainly not an emu.
In 1888 De Vis described a new species af emu,
Dromeius petricius, from a proximal end of a right
tibiotarsus (QM 5547) and the distal end of
another tibiotarsus (QM F5548), In the same paper
he provisionally referred a left coracoid (QM FiL20)
to the same species, These three fossils were from
King Creek, Darling Downs, in south-eastern
Queensland, De Vis (1892) considered the whale of
the Darling Downs sediments to be much the same
age, but it is now known that these fossil-bearing
deposits represent a range of ages. The Chinchilla
fauna is likely to be of Late Pliocene age. On the
other hand, the Darling Downs fauna of the eastern
part of the Downs, including King Creek, is of Late
Pleistocene age (Woods 1960, Stirton ef af 1968,
Rich 1979), Later De Vis (1892, 1905) also referred
a femur fragment. three tarsometatarsi, and a
partial synsacrum (QM F5549) to D. potricins. The
referral of the synsacrum |s especially poteworthy,
De Vis considered that because of its size, the
fragment must have been from a cassowary er an
emu, but: ‘as no extinct cassowary Is known yet in
Australia, it seems almost necessary to attribute the
present fossilio the emu DB. parriciis’ {De Vis 1905:
5).
In. 1892 De Vis set up another species of emu,
Dromaius. gracilipes, based on a distal left
tarsometatarsus (QM F1142)- In the description De
Vis omitted to note the location from which the
Specimen was collected, but the museum Label
associated With the specirneri Indicates that [t was
from the Darling Downs, In this article De Vis
(1892) also described a supposed kiwi, ‘Metapferyx
bifrons', again without giving a location
Spencer (1906) described minor of King
Island, Bass Sirait, Tasmania. The previous exist-
ence OF a Separate species of emu on this island was
almost simullancously made by Legge (1907), but
he withdrew the name. D. minor was tedefined by
Spencer & Kershaw (1910) as more specimens
became available, and recently the taxonomic status
of this species has been discussed by Parker (1984),
The status of the extiner Tasmanian emu is an
as yet unresolved problem, Emus were introduced
from the mainland in the 1800s, and interbreeding
may have occurred (Howchin 1926). Le Souef (1903)
gave the Tasmanian emu the specific name 2
diemenensis. Ridpath & Moreau (1966) considered
it a subspecies of D newaehollandiae. The onty
fossils and recent specimens of D diemenensis
collected allve which are known to exist include 4
fernut, @ synsaccum, three tibintarsi, two tarso-
metatarsi, a ces vical vertebra, and a leg lacking the
femur and part of digit II (all at OVM) (Scott 1924,
1932), and three eggs (in private collections)
(Campbell (900, Le Souef 1903, Spencer & Kershaw
1910, Dave 1926).
Anderson (1997) deseribed an emu sternum,
which is much thickee than those of the living
DB. navachallandiae, from the Wellington Cayes,
New South Wales. He suggested it might possibly
belong 16 B patricius.
Miller (1962) restudied Caswarites lpdekkeri,
discussed earlier by Rothschild (1911). The type of
the species is a distal right tibiotarsus (AM
MEF!268). The type locality has been variously piven
as Queensland, Cooma und Wellington Caves
(Miller 1962), but its provenance is uncertain, lus
preservation, however, is very unlike that of fossils
from Wellington Caves, It ts clearly a-cassowary,
however, and not an emu,
Miller (1963) described a new species of emu
Dromivetus (=Dromaius) ocypus based on an
essentially complete tight tarsometatarsus (SAM
P13444) from the Pliocene Mampuwordu Sands,
Palankarinna fauna, Lake Palankarinna, South
Australia, It is smaller than D, neveehollandiae, The
tarsomietatarsug is eVidently the one referred to by
Miller In Stirton e¢ af, (1961) as a new species of
em with ‘proportions of the bone... intermediate
between those of the emu and the cassowary’.
Miller also assigned four phalanges (UCMP
56849, 60563, 94679, 94680) from Lake Kanunka
(UCMP V-5772, Katipiri Sands or possibly Tirarl
Formation) to the Dromornithidae, possibly
Genyornis newioni (Stirton et af. 1961). As noted
by Rich (1979), however, they actually belong in the
genus Drormalas; Thus, no dromornithids are
known from the Pliocene-Pleistocene Lake
Kanunka fauna, and this adds another record for
emus,
Rich (1979) refers a left femur (SAM PL7104),
from Brother's sland, South Australla to Genvornis
newiori, but it conformis in all respects to Dromoafus
and should be transferred to that taxon,
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE)
TABLE 1. Australian localities producing fossil emus (Dromaius).
Locality
Fossil Elements
Rock
Fauna
Age
87
References
Leaf locality, Lake
Ngapakaldi, South
Australia
Bullock Creek,
Northern Territory
Alcoota (including
Rochow locality),
Northern Territory
Lawson-Daily
Quarry, Lake Palan-
karinna, South
Australia
Lake Kanunka,
South Australia
Chinchilla,
Queensland
Darling Downs,
Queensland
King Creek,
Queensland
Thorlindah, Paroo
River, Queensland
Bingara, New South
Wales
Lake Menindee, New
South Wales
Lake Tandou, New
South Wales
Wellington Caves,
New South Wales
Wombeyan Quarry
Cave, New South
Wales
?Baldina Creek, near
Burra, South
Australia
Brothers Island,
South Australia
Tmt., d Tib., Pes
Dromaius gidju
Tmt., Tib. Dromaius
sp.
Tmt. frags., Phs.
Dromaius sp.
Tmt.
Dromaius ocypus;
Tib., F.
Dromaius cf. ocypus
F., Ph., R., Tib., V.
Dromaius novae-
hollandiae, Dromaius
sp.
Syn., F., Tmt.
Dromaius novae-
hollandiae
Cor., F., Tib., Tmt.
Dromaius novae-
hollandiae
Dromaius novae-
hollandiae
Tib.
Dromasius novae-
hollandiae
V., Syn., Tib.
Dromaius novae-
hollandiae, Dromaius
sp.
Ph., Tib.
Dromaius novae-
hollandiae
Dromaius sp.
Tib.
Casuarius lydekkeri;
St., Tib., Tmt.
Dromaius novae-
hollandiae, Dromaius
sp.
Tib., Tmt.
Dromaius novae-
hollandiae
F.
Dromaius sp.
F,
Dromaius novae-
hollandiae
Wipajiri Fm.
Kutjamarpu
Miocene
Camfield beds Bullock Creek Late Miocene
Waite Fm.
Mampuwordu
Sands
Katipiri Sands
Chinchilla
Sands
Unnamed
Unnamed
Unnamed
Unnamed
Unnamed
sand lunette
Unnamed
Unnamed
cave sediments
Unnamed
cave sediments
Unnamed
Unnamed
aeolianite
Alcoota
Palankarinna
Kanunka
Chinchilla
Darling
Downs
King Creek
Unnamed
Bingara
Unnamed
Unnamed
Unnamed
Unnamed
Unnamed
Unnamed
Late Miocene
Pliocene
Late Pliocene
or Early
Pleistocene?
Early to
Middle
Pliocene
Pleistocene
Late
Pleistocene
Pleistocene
Pleistocene
Late
Pleistocene
Pleistocene
Quaternary
Late
Pleistocene
Quaternary
Quaternary
Stirton et al.
1967, 1968; Rich
1979.
Rich 1979
Woodburne 1967,
Stirton et al.
1968, Rich 1979,
Rich et al. 1982
Miller 1963, Rich
1979
Rich 1975, 1979
Woods 1960,
Stirton et al.
1968, Rich et al.
1982
Woods 1960, Rich
1975, 1979
Baird 1986
Etheridge, 1889,
Rich 1975, 1979
Anderson 1889,
Rich 1975, Marcus
1976
Tedford 1967
Rich 1975
David 1950, Rich
1979, Dawson
pers. comm.
Hope 1971, Rich
1975
S.A.M. Museum
label
Rich 1975, 1979
88
C. PATTERSON & P. V. RICH
—_——————————————————————————————————————————————————
Locality
Cooper Creek,
(includes Katipiri
Waterhole and
Wurdulumankula),
South Australia
Kangaroo Island,
South Australia
(Several localities)
Lake Callabonna
(lower stratigraphic
level), South
Australia
Lake Kittakittaooloo,
South Australia
Naracoorte
(Henschkes Bone Dig
and Victoria Fossil
Cave) South
Australia
Salt Creek, South
Australia
Warburton River,
South Australia
(includes Green Bluff
locality and
Kalamurina).
Bone Cave, Western
Australia
A cave north of
Moore River,
Western Australia
Irishtown, Tasmania
King Island, Bass
Strait, Tasmania
Mole Creek,
Tasmania
Moybray Swamp,
Smithton, Tasmania
Scotchtown Cave,
Tasmania
Fossil Elements Fauna Age References
F,M Tmt., Syn., V. Katipiri Sands Malkuni Pliocene- Stirton et al.
Dromaius sp. Quaternary 1961, Rich 1975
Dromaiinae
Sk.. Unnamed Quaternary Morgan & Sutton
Dromaius baudinianus 1928, Rich 1975,
Parker 1984
Ci, Sym, Vi5F., Tib, Lake Pleistocene Stirling & Zeitz
Dromaius novae- Callabonna 1900, Rich 1975,
hollandiae 1979
Tmt, Katipiri Sands Malkuni Quaternary S.I.A.M. Museum
Dromaius novae- label
hollandiae
M., V., R., Hum., Unnamed Pleistocene van Tets & Smith
Syn., F., Tib., Tmt., cave sediments 1974
phs.
Dromaius novae-
hollandiae, Dromaius
sp.
F. frag. Unnamed Quaternary Rich 1975
Dromaius cf. novae-
hollandiae
Syn., Tib., Tmt. Katipiri Sands Malkuni Quaternary — Rich 1975
Dromaius sp.
Tib., Tmt. Unnamed Quaternary
Dromaius novae- cave sediments
hollandiae
Tmt. Unnamed Quaternary —_ Rich (unpublished,
Dromaius novae- 1971, field notes)
hollandiae
Tib. Unnamed Quaternary Scott 1924
Dromaius diemenen-
sis. Needs review
Sk. Unnamed Quaternary Spencer 1906,
Dromaius minor sand rock and Spencer &
Kershaw 1910,
Jennings 1959,
Parker 1984, Rich
1975
Tib. Unnamed Quaternary Scott 1932
Dromaius diemenensis.
Needs review
Syn., V., F., Tib., Unnamed Quaternary Scott 1932
tmts.
Dromaius diemenensis.
Needs review
Le Unnamed Quaternary Gill & Banks,
Dromaius diemenensis. 1956
Needs review
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE)
89
a
Locality Fossil Elements Rock
Lancefield, Victoria Tib., Tmt., Ph. Unnamed
Dromaius novae-
hollandiae
McEachern’s Cave, Many skeletal Unnamed
Victoria elements
Dromaius novae-
hollandiae
Buchan Caves, Tmt. Dromaius Unnamed
Trogdip Cave area, novae-hollandiae
Victoria
Fauna Age References
Unnamed Pleistocene Gillespie et al.
(26 000 B.P.) 1978
Unnamed Quaternary McNamara pers.
comm.
Unnamed Quaternary = Rich 1975.
The bird remains in the Riversleigh fauna (Carl
Creek Limestone) identified only as close to
‘Dromiceius’ in Tedford (1967) have been determined
by Rich (1979) to belong to a dromornithid,
Barawertornis tedfordi, and thus are not a record
of emu.
STRATIGRAPHY
(see Table 1)
Only a few fossil sites producing emus have been
found thus far, and most are of Pleistocene age.
Fossils of a new species, Dromaius gidju, proposed
in this paper, have been found at the Leaf Locality
(UCMP V-6313) on the eastern shore of Lake
Negapakaldi, eastern Lake Eyre sub-basin, South
Australia (Stirton et al. 1967). The sediments that
outcrop here, known as the Wipajiri Formation,
contain the Kutjamarpu fauna. Diprotodontid
marsupials in this fauna are considered more
primitive than those in the Beaumaris,
Palankarinna, and Alcoota faunas, and have closest
affinities with forms in the older Ngapakaldi fauna,
known from localities listed in Stirton et al. (1968),
in the Lake Eyre sub-basin and thought to be of
medial Miocene age (Rich et al. 1982).
The Camfield beds at Bullock Creek (Bullock
Creek fauna), Northern Territory, of probable Late
Miocene age, have produced Dromaius sp. currently
under study by P. V. Rich.
The Rochow locality (UCMP V-6349) at Alcoota,
Northern Territory, near Alice Springs, has
produced Dromaius remains that may be Miocene
in age. But, as discussed by Rich (1979), the Waite
Formation, which contains the Alcoota fauna, is
not well dated at present. The diprotodont
marsupials from this locale suggest a date younger
than that represented by the Kutjamarpu fauna but
older or contemporaneous with the Hamilton
fauna. An unnamed rock unit containing the
Hamilton fauna is capped by a basalt, which has
been dated as 4.35 + 0.01 my. B.P. (or Early
Pliocene) by Turnbull ef a/. (1965) and Turnbull &
Lundelius (1970). The sequence is underlain by
marine sediments of the Grange Burn Formation
assigned to the Kalimnan stage. At present Alcoota
is viewed as Late Miocene in age.
Dromaius ocypus was recovered from the
Lawson-Daily Quarry (or Lawson Quarry; UCMP
V-5769) at Lake Palankarinna, eastern Lake Eyre
Basin, South Australia (Miller 1963). The enclosing
rocks, the Mampuwordu Sands, contain the
Palankarinna fauna, and are overlain by the Tirari
Formation and the Late Pliocene or Early
Pleistocene Katipiri Sands containing the Malkuni
fauna at Lake Palankarinna. An age of Middle to
Late Pliocene is established by marsupial fossils, in
particular Zygomaturus, which are more advanced
than zygomaturines from Awe, Beaumaris, and
Alcoota and yet more primitive than Pleistocene
forms (Stirton ef al. 1968).
Lake Kanunka (UCMP V-5772) in the eastern
Lake Eyre sub-basin, South Australia, has also
yielded Dromaius fossils. The Katipiri Sands or
possibly Tirari Formation (see Rich 1979: 61)
contains the Kanunka fauna dated as Pliocene or
Early Pleistocene (Stirton et ai. 1961, Rich et al.
1982, Tedford pers. comm. 1985).
The Chinchilla locality, south-eastern
Queensland (Chinchilla Sands, Chinchilla fauna)
also contains Dromaius. Several elements of the
marsupial fauna appear more primitive than those
in the Pleistocene eastern Darling Downs, and
Woods (1960) assigned it a Pliocene age. Rich ef al.
(1982) consider Chinchilla to be Early to Middle
Pliocene in age.
The distal part of a tarsometatarsus (AM F
58087) of an emu was found in the Australian
Museum’s ‘old collection’ and labelled ‘mixed plus
some from Lord Howe Island’. The fossil is very
incomplete and appears to be from a juvenile
individual. There are no reliable stratigraphic or
locality data available for this form.
All of the other known fossiliferous sites
producing emus are Pleistocene in age. For these
o) © PATTERSON & Py, RICH
Pleistocene sites, as might be expected, some dates
are better established than others, Depasition, tor
instance, of the Darling Downs sedirnerts in
Queensland may have occurred al several diflerent
times (Rich 1979) during the Pleistocene, and
definite ages for specific sites are difficult to
determine,
Thorlindah, on the Paroo River, Queensland, is
thought by Rich (1979) to be Probably Pleistocene
.-. the bird remains were collected along with
fragments of ‘kangaroos’ and Diprofaden (Stirling
& Zeitz 1900; 44) ina well 20 feet deep’, Diprolodon
appears to be restricted ta the Pleistocene in all
precisely dated situations. Emu material indistin-
guishable from the living forms is known from
Thorlindah,
Vertebrate fossil-bearing localities at Lake
Menindee adjacent to the Darling River and its
major anabranch, western New South Wales, have
been radiocarbon dated at 26 300 + 1500 B.P, and
1§ 800 B.P, (Tedford 1967), Fossils of Diprotedan,
Thylaceieo, Phkaseolanus, Protemnedon and
mracropodids have been recovered. UCMP Jocalities
V-5371, Y-7185, V-67186 and Y-67187 have produced
Dromaius fossils, Hope (1978) discusses the
Slratigraphy of the Menindee area in some detail,
with reference to the problem of dating the
Pleistocene megafauna extinctions, At present the
emu fossils from Menindee appear to be Late
Pleistocene in age.
At Lake Tandou, New South Wales, several
Dromaius fossils were found in archaeological
excavalions. Hope (pers. comm.) states that: ‘there
is NOW 4 reasonable stratigraphy for the lumette [at
Lake Tandoul], and a lot more cates; the oldest are
in the order of 22 000-25 000) and lie ac the base
of the uppermost scratigraphie unit. The problem
..- i8in Working out where Harry's [Harry Allen,
who collected the specimens while doing research
toward a Ph.D. thesis) matefial came from’, Tenta-
Lively, a Pleistocene age seems appropriate for these
fossils,
Bingara in New South Wales has produced
verlebrae and a tibiotarsus of Dromaius. The bone
bed occurs in a fluviatile clay deposit about 39-N)
em thick on the western side of Myall Creek,
Remains of Diprotodon indicate a Pleistocene age
(Anderson (889),
Also in. New South Wales,. the Wombeyan Quarry
Cave has yielded Dromaius Fossils. This is not the
same cave as Broom Cave or Guineacor Cave, also
in the vicinity of Wombeyan. The Wombeyan
Quarry Cave has not been radiocarbon dated, but
Hope (1982) beheves that it is of Late Pleistacerie
age, It seems likely that the quarry deposit is older
than the ‘Broom breccia’, but bath appear te be of
Late. Pleistocene age. Other fossils recovered from
Wombeyan Quarry Cave include Protemnadon,
Sthenurus, Zygomaturus, Palorchestes, Thylacolea
carrifex, Sarcophitus faniarius, and Progura
gallinavea,
There are several bone producing caves in the
Wellington Valley area of New South Wales.
Different levels and different caves may have
trapped animals at various times in the Pleistocene
to Recent, perbiaps even prior to this (L. Dawson
pers, comm.) depending on when they were opened
and rescaled (David 1950, Tedford 1967), Emu
fossils have been recavered from caves in this area.
Rich (1979: 58) states that Dromiaius remains were
recovered from Cuddie Springs (Mara Creek, SSE
of Brewarrina, 16 km ESE of Gilgoin), New South
Wales, Anderson & Fletcher (£934) do not mention
Dramaius in their, admittedly incomplete, list of
fossils tecuyered from this, site. Wilkinson (1884)
stated that; ‘bones of Diprotodon, Stherurts,
Macrapus titan, latge wormbats, large birds probably
emus, cracodiles and a gigantic camivorous lizard,
Nofiosaurus .—. are found only within a few yards
of the centre of the spring* Unfortunately, he does
not describe or figure these bones, and the large
birds may be Genpornis, specimens af which were
later recovered by Anderson and Fletcher. MM F
19420, onlabelled when found in an old collection,
has ‘the style of preservation [suggesting] that it
comes from Cuddie Springs* {Pickett pers. comm.)
but is too large to be Brorraius. lt appears, instead,
to be the internal condyle of # titiotarsus of a
dromornithid, perhaps Geryarnis. We have been
unable to relocate (he specimens Rich (1979)
assigned to Drorpeivs.
Two Pleistocere cave deposits producing
Droinaius fosals are Known in Westeen Australia,
A cave norit of East Moore, Western Australia, has
produced a tarsometatarsus of a juvenile emu
(unregistered WAM). Bane Cave, near Jewel Cave,
has produced an emu tibiotarsus and tarsorneta-
tarsus.
Four Pleistocene Dromaius tocalities are known
from Tasmaria, Scott (1924, 1932) reported a
tibiotarsus from Irishtown, 2 tibiotarsus from Mole
Creek, and several elements (a synsacrum, Jemur,
tibiatarsus, two farsemetatarsi, and a cervical
vertebra) fram Mowbray Swaynp, near Smithton,
in western Tasmania. The Mowbray Swatnp fossil
site has been rachocarbon dated ait greater than
37 780 BLP. (Gall & Banks 1956), Another Mr Scott
found bones of the Tasmanian emu at Scotchtown
Cave in association with ‘Notorherium
tasmanicum', Thylacoleo carnifex, and Palorchestey
(Gill & Banks L9SA).
DPromaius minor is known from the Bass Strait
island, King Island, Tasmania. Anderson (1914) was
of the opinion that the original Fossil matrix was
a fairly hard, coarse, red-brown sand rock of
shallow manne origin. Jennings (1959) stated thar
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 91
the fossils occurred in windblown sand dunes of
Pleistocene to Recent age and that finds from
different geological horizons had likely been
brought together by winnowing. Separate from the
King Island form, the now extinct Dromaius
baudinianus, is known from Kangaroo Island
(Parker 1984).
Several localities along the south coast of
Kangaroo Island (Cape du Couedic, Kelly Hill,
Eleanor River, and The Brecknells) have produced
fossil material (Morgan & Sutton 1928). Rich (1975)
states that the age is Pleistocene.
Three Victorian sites, all Pleistocene in age, have
produced fossil emus. A partial tarsometatarsus is
known from Trogdip Cave, part of the Buchan
Caves said by Rich (1975) to be Pleistocene because
of the nature of the marsupial fauna also preserved
in it.
Many fossils of the Australian megafauna have
been recovered from a swamp near Lancefield,
Victoria. As well as emus (less than 1% of the
bones), Macropus giganteus, Protemnodon,
Sthenurus, Diprotodon, and a dromornithid,
probably Genyornis, were found. A sample of the
bones themselves was radiocarbon dated at 19 800
+ 450 B.P., while charcoal in the channel fill in and
upon which the fossil deposit rests provides a
maximum age for the bones of 26 000 + 500 B.P.
(Gillespie ef al. 1978).
A third Victorian site which has produced emu
fossils is McEachern’s Cave in western Victoria.
According to Wakefield (1967, 1969), due to the
funnel shape of the entrance the cave has acted as
a death trap for terrestrial animals. Gravitational
movement, movement of trapped animals and water
action were responsible for considerable mixing of
cave sediments. The fossils are Late Pleistocene to
Recent in age. A sample of mammal bones from
the top layer of the Pleistocene sediments gave a
radiocarbon date of 15 200 + 320 B.P. Extinct
Pleistocene species found in the cave include
Sarcophilus laniarius, Zygomaturus trilobus,
Thylacoleo carnifex, Sthenurus spp. and
Protemnodon cf. brehus.
The remaining sites from which fossil emus have
been recovered are all South Australian. From
Brothers Island, Coffin Bay, about 50 km WNW
of Port Lincoln, a femur fragment SAM P17104,
referred to Genyornis newtoni by Rich (1979) but
actually Dromaius, was found in an unnamed
aeolianite of sand and shells. As similar deposits
on the island have produced Sthenurus cf. brownei
(Tedford in Rich 1979), a Pleistocene age is
indicated.
A number of Pleistocene localities collected by
J. W. Gregory (1906) and later by joint expeditions
from the University of California and the South
Australian Museum, occur in the eastern Lake Eyre
basin. The fossils were found as ‘float’ or in place
in the Katipiri Sands, which contain the Malkuni
fauna. Also collected by Gregory and later
expeditions of the University of California and the
South Australian Museum, are several localities on
the Warburton River, including Green Bluff Locality
(UCMP V-5771), Lookout Locality (UCMP Y-5776)
and Kalamurina. The Warburton River is in the
eastern Lake Eyre sub-basin, and has produced
fossils from the Katipiri Sands. A Smithsonian
Institution-American Museum (SIAM) expedition
in 1970 recovered a tarsometatarsal fragment (SIAM
75) of an emu from the Katipiri Sands (Malkuni
fauna) at Lake Kittakittaooloo.
The Smithsonian Institution-American Museum
Expedition and later a Museum of Victoria-
Australian Army Expedition also recovered a
number of Dromaius fossils from the lower level
of Lake Callabonna in South Australia. This
stratigraphic unit producing the emus also
contained Genyornis newtoni, Diprotodon
optatum, Phascolonus gigas, Sthenurus, Protem-
nodon, and Macropus (Stirling & Zeitz 1900, Rich
1979) and has been dated at greater than 40 000 B.P.
(Tedford 1967), but sometime during the Pleisto-
cene.
An incomplete femur (SAM P17103) bearing the
museum label: ‘ ?Genyornis. Pleistocene locality
unknown, possibly Baldina Creek near Burra,
South Australia’ is actually Dromaius. If the
location is in fact Baldina Creek, a Pleistocene age
is suggested by the occurrence of known Genyornis
newtoni (Rich 1979) and Diprotodon at this site
(Stirling & Zeitz 1900).
Several fossils of Dromaius were collected near
Burra, South Australia by Mr R. E. Ireland and
forwarded by the police department on 12 March
1935 to the South Australian Museum, They were
found in a sandhill in association with Aboriginal
(Homo sapiens) bones, SAM A25805 (information
from museum label). The Aboriginal remains
suggest a Pleistocene to Recent age.
Two caves near Naracoorte (about 320 km SE of
Adelaide near the Victorian border), South
Australia, have produced Dromaius: Victoria Fossil
Cave (van Tets & Smith 1974) and Henschke’s Bone
Dig. Sediments producing the fossils in Henschke’s
Bone Dig have been radiocarbon dated at about
33 800 B.P. (van Tets 1974). Smith (1971) stated that
the bones in Victoria Fossil Cave are most abundant
in the top 15 cm of the damp, friable, light brown
earth forming the floor of the cave. She also states
that the abundance of sthenurines, diprotodontids,
and Thylacoleo suggests that the deposit was
formed sometime during the Pleistocene and sealed
before the Recent. Wells ef al. (1984) provide a
complete discussion of current dates from this site.
92 C. PATTERSON & P. V, RICH
SYSTEMATICS
Only those features exhibited by the fossil
specimens are discussed. For measurements see
Table 2.
Family CASUARIIDAE Brisson
Members of the Casuariidae have a pterygoid
that is inflated where it contacts the palatine; a
palatine with a short shaft and an expanded medial
plate; a long vomer; a palate that lies ventral to the
parasphenoid rostrum and makes contact with the
braincase only at the basipterygoid processes;
maxillopalatines that are cone-shaped and open
posteriorly; the cervical vertebrae are antero-
posteriorly compressed; the atlas possesses lateral
spines or occasionally complete vertebrarterial
canals; the sternum is longer than wide with lateral
margins concave laterally and has short, dorsally-
directed sternocoracoidal processes and no sternal
notches; the costal margin forms about 50% of the
lateral margin; the antitrochanter of the synsacrum
is located at the anteroposterior mid-point of the
synsacrum; the ilium, ischium and pubis are
subequal in posterior extent; neither the pubes nor
the ischia are fused posteriorly along the mid-line;
the ischium is deeper than the pubis; the ilium
dorsal to the acetabulum is deep; the trochanter and
head of the femur are subequal in proximal extent;
the external condyle extends only moderately distal
to the internal condyle; the popliteal fossa is
elliptical and of moderate width; the distal depth
and width of the femur are subequal; the posterior
margin of the proximal articular surface is highly
concave anteriorly; the external condyle and fibular
condyle are subequal in breadth or the fibular
condyle is broader; the cnemial crests of the
tibiotarsus are little compressed mediolaterally; the
inner cnemial crest extends far proximally to the
proximal articular surface; the external articular
surface extends far laterally; the margin of the
external condyle is semicircular in lateral view; the
tibiotarsus lacks a supratendinal bridge and also
lacks an intercondylar eminence; the hypotarsus of
the tarsometatarsus is narrow and centrally located;
the hypotarsus extends decidedly further proximally
than the intercotylar prominence; the internal cotyla
is deeper than the external; the posterior shaft
surface is deeply grooved; the anterior metatarsal
groove is deep and extends the length of the shaft;
trochlea IV extends distal to trochlea Il; trochlea
III extends distal to trochleae II and IV; the
phalangeal count for digits I, III, IV is 3-4-5; of
the proximal phalanges that of digit III is longest;
that of digit IV is shortest; the unguals are generally
claw-like, except for the elongated ungual of digit
II in Casuarius.
Subfamily DROMAIINAE Vieillot
Within the Casuariidae there are a number of
characters which reliably distinguish Dromaius from
Casuarius, the only other member of the family.
In Dromaius the mandible is broad and rounded
distally, not narrow and pointed distally; the
mandibular articulation of the quadrate is step-
shaped, with the external facet decidedly more
excavated (in Casuarius the facets of the mandibular
articulation of the quadrate are subequal); the
pterygoid is not excavated dorsally; and the palatine
and vomer are decidedly shorter than in Casuarius;
the semicircular notch in the prearticular surface
of the atlas is shallow and narrow, not deep and
broad; the axis is longer, and the hypapophysis not
as deep as in Casuarius; the cervical vertebrae
possess long, not short, styloid ribs, which come
to a point distally, and are not rounded; the neural
canals and vertebrarterial canals are small; the
excavation of the neural arch posterior to the
prezygapophysis is shallow (from the eighth cervical
posteriorly in Casuarius the excavation of the neural
arch is deep); the thoracic vertebrae are similar to
those of Casuarius, but the neural canals are
smaller; the neural canals of the caudal vertebrae
are small, with an elliptical cross-section, whereas
in Casuarius the neural canals are large and
triangular in cross-section; the sternum is only
slightly longer than wide, not much longer than
wide as in Casuarius; the costal processes lie in an
almost horizontal plane, whereas in Casuarius they
lie on a downward curve (antero-posteriorly); the
sterno-coracoidal processes are moderately long,
not very short as in Casuarius; the coracoidal sulci
are short and overlap medially, whereas they are
long and do not overlap in Casuarius; the body of
the sternum is weakly concave dorsally, but in
Casuarius it is strongly concave dorsally; the depth
of the sternum anteriorly is shallow, not deep; the
costal margin is long, whereas in Casuarius it is
shorter; the supratrochanteric ridge is broader; the
pre-acetabular synsacrum tends to be shorter than
the post-acetabular synsacrum in Dromaius while
the opposite condition exists in Casuarius;
proximally and posteriorly the femur bears a large
pneumatic foramen, lacking in Casuarius; in
anterior view, the external condyle extends decidedly
further proximally than the internal condyle, while
the two condyles are subequal in anterior proximal
extent in Casuarius; in medial view, the internal
condyle is semicircular in outline, while in Casuarius
it is triangular; the diameter of the head and the
minimum diameter of the shaft at its proximo-distal
mid-point are equal, whereas in Casuarius the head
diameter is less than the shaft diameter; the shaft
is almost straight, being more curved in Casuarius;
the proximal extent of the cnemial crest is not as
great as in Casuarius; anteriorly the external
VHE FOSSIL HISTORY OF THE EMUS, OROMAIUS (AVES! IROMAIINAR) "
condyle is rounded proximally, and it extends
further proximally and is more pointed in
Cusuarlus; aboye the anterior intercundylar fossa
is. a. small ridge trending dorsally and laterally from
the mid-line and ending in a small foramen, while
in Casuarius this cldge is absent, but the foramen
still exists; the tarsometatarsus and tibiotarsus are
subsqual in length, unlike in Casuarius in which the
tarsometatarsus is decidedly shorter; the secand
trocidea is much more reduced than in Cagiutrllss;
the intercotylar prominence is low and tends to be
flat, while in Casvarius the intercotylar prominence
is higher and convex dorsally; a distal foramen,
which completely pencirales (he tatsometatarsus
(antere-posteriorly), and a groove (occasionally a
completely roofed-over forainen) running proximo-
distally, are present, both absent in Cuswarius, the
condyles af the phalanges of the foot tend to be
greatly divergent plantarly; In Caswarius the
condyles tend to be only moderately divergent
plantarly; in distal view, the intercondylar fossa
tends to be only slightly notched in a step-shaped
fashion dorsally, while in Casvarius this notels rencds
10 be deeper and more V-shaped; the ungual of digit
I11 is longest, and that of digit TV shortest, while
in Casvarius the ungual of digit IL is longes!, and
that OF digit IV shortest,
Dromaiirs novuelollandise (Latham)
Type
Casuarius noveehdllandiae (Latham)
Type Locality
New Holland (Sydney, New South Wiles,
Australia) (Table 1),
Measurements
Tables 2-13.
Referred Fassil Marerial
Bingura, New South Wales — , MM F16786,
dorsal vertebra (¥.24-26°), neural spine, pre- and
postzygapophyses, diapophyses and prearticular
surface damaged; MM FI6797, dorsal vertebra
(H24-267), neural apine and diapophyses net
preserved, Tie, MM F16775, distal end and distal
half of shaft. Pletstocene.
Bone Cave (wear Jewel Cavel, Western Australia
— Tih, WAM6S,5,34 (in part), shaft only. Tint,
WAM 68,5.94 (in part), shaft only, Quaternary.
Brothers Islan, South Ausirelia — &, SAM
PI7I0d, proxiinal ead and proximal Ewo-lhinds of
shaft, head and trochanter damaged, Quaternary.
Chinchily, Queenstand — Yar. OM F143 din
part), third trochlea only.
Cooper Creck, South Australia — Spa, UCMP
56233, acetabular complex, (site 2, LICMP V5378).
Ff, HM B775/869, entire, (Lower Cooper,
locality 4). Tint,, UCMP 56313, distal end, fourth
trochlea not preserved, (site 3, UCMP V5379). Late
Pliocene or Early Pletstocene.
Darting Downs, Queensland — &, QM FI43 (in
part), distal, popliteal fossa region only (eastern
Darling Downs). Tif, AM A97I3, proximal, see
Figure 1; QM FSS47, proximal, figured (De Vis
1589); OM FSS48, distal, figured (De Vis L889), OM
F1652, proximal end, most of cnempal crests noc
preserved, (Condamine River, fear Dalby), Tors,
QM F112), proximal frag.; QM F135, figured (De
Vis, 1892), distal, juvenile; QM F142, distal Irae.
QM F1143 {in part), distal end (eastern Darling
Downs), Pleistocene,
Lake Callabonna, South Australia — AMNH
9478, Vi, second cervical, posterior left side; third
cervival, left side: Fourth cervical (articulates with
third cervical), neural spine and ribs not preserved;
sivth cervical’, ribs and right side postarticuliar
surface not preserved; seventh cervical’, (articulates
with sixth cervical), ribs and right stde
prezygapophysis not preserved; ninth cervigal’,
postarticular surface, ribs, right side
postzygapophysis not preserved. SIAM 61, Si,
fragments. AMNE 9677. F, distal end, internal
vondyle damaged, AMNH 9676. Tif, entire, see
Figure 2, Pleistocene,
Lake Kanunka, South Australia — K, UCMP
$6854, dorsal vertebra (#22 of 23), (UCMP 5772).
F, UCMP RHTLO64, trochanter, condyles, and
head partly eroded and crushed, (site 1, UCMI*
V5772). Tib, UCMP 56845, distal end, most of
internal condyle not preserved and remainder highly
eroded (UCMP V5772), Ph, UCMP 56849, first
phalanx, second digit, (UCMP V5772, UCMP
94679, first phalanx, third digit; UCMP 94680,
second phalanx, third digit (UCMP V5772). Late
Pliocene or Early Pleistocene
Lake Menindee, New South Wales — Eggshell,
UCMP 55948. Tih, UCMP 53825, two distal
tibiorarsi with the same number, Tin, UCMP
53835, distal end most of third trochlea and second
and fourth trochleaée not preserved. PR, UCMP
$3832, dark colour (presumably burne), first
phalanx, second digit, distal end; Ungual phalanx,
second or fourth digit} flest phalanx, second digit,
proximal end; second phalanx, Fourth digit (UCMP
V5371), LICMP 53833, first phalanx, fourth digit
and ungual (site I. VA7ISS UCMP 55983, first
and second phalanges, third digit (UCMP Y67186),
Late Pleistocene
Lancefield,. Vietorta — Tid, NMV P43037, distal
shaft; NMV P43041, distal shaft, juvenile: NMV
Pd40l1, entire but articular surface worm; NMV
PISO0(3, distal, Tw, NMV LS, distal; NMV
P44012, proximal artitular surface eroded; NMV
P44013, hypularsus eroded; NM¥ P4404; NMV
P44015, proximal articular surface worn, sevond
and fourth trochleae nol preserved, NMV P44016,
04 C. PATTERSON & PV. RICH
proxtnial articular surface wort; NMV P4sD17,
distal shaft; NMY P44018, distal; NMY P44019,
distal; NMY P48392, sécond and part of third
trochleace not preserved; NMV P150014, distal,
fourth trochlea not preserved. Ph: NMV P4319,
first phalanx, third digit, articular surfaces worn;
NMY P43200, second phalany, third digit, proximal
end. Late Pleistocene, 26 000 BP,
MecWachern'’s Cave, Vieloria — C, NMV
P157345, posterior fragment; NMY P157350, lower
jaw, distal; NMY 9157353, posterior tragment. h,
NMV PI57346, 2Jst or 22nd vertebra, neural spine,
diapopbysis, might side prezygapophysis, part of
centrun aod prearticular surface not preserved;
NMV P175349, 23rd or 24th vertebra, neural spine,
pre- and postzygapophyses not preserved,
postarlicular surface worn: NMY PI57351, seventh
cervical?, juvenile, ribs not ankylosed) NMY
P157352, lith, !2th or 13th cervical, juvenile; NMV
P15735%, third cervical, right side rib not preserved;
NMV P1IS7364, 25th or 26th vertebra, leit side of
centrum with prezygapophysis and diapophysis, but
postarticular surface not preserved; NMV P157367,
20th or 23st verjebra, diagpophysis, part of neural
arch, postzygzapophyses, and right side
prezyzapophysis only, juvenile; NMV PI57368,
about lth vervioal, prearticular surtace worn, left
side prezygapophysis and ribs (not ankylosed) nat
preserved, juvenile; NMV P157369, 22nd to 26th
vertebra, right side of centrum only, juvenile St,
NMV P157347, incomplete; NMV P15735§, entire.
Syn, NMV P157361, fragment. Tih, NMV P157356,
proximal; NMV P157357, distal; NMV P5736,
distal; NMV_ P157365, proximal, Fih, NMYV
P157363, proximal end. Tit, NMV P15S7344, distal,
trochleae nol preserved, Quaternary.
Cave sorth of Moore River, Western Australia
— Trt, WAM-190 unregistered, (not seen, data
from Pat Rich’s 197! field notes), Quaternary.
Naracoorte, (Henschke's Bone Dig and Victoria
Fossil Cave), South Australia — C, SAM PL7234,
lower jaw, distal end only, (Henschke’s A3, 40"),
KK, SAM PI7589, Léth cervical, ribs nol preserved;
SAM P18246, fourth cervical; SAME P1I8247, 22nd
to 24th vertebra, part of centrum and left side
diapoplrysis, (Henschke'sk SAM PIS673, 15th to
1th cervical? vertebra, prezygapophyses and
centrum damaged, (Hensehke's area X4, depth 17");
SAM Pi8830, 26th vertebra, prezygapophyscs,
prearticular surface and diapophyses damaged,
(Henschke's area X6, 15-30 em). Syn, unregistered
SAM; SAM PI6501, acetabular complex only,
(Victoria Fossil Cave, 0-10+RI0-0-12"), see Figure 3;
SAM P17747, parts of alium, ischia and pubes not
preserved, (Henschke’s A3, 39-42"); SAM PL&s00,
distal tight ischium only, (Henschke's Af, 32-36"),
J, SAM P2281? (in past), condyles badly erodod:
unregistered SAM, intemal condyle darmaged. Tih,
SAM PI7L49, distal end, internal condyle worn;
SAM PI8829, distal, part of condyles not preserved,
(Henschke's area X64, D-15 cm). Tint, SAM PL7B16,
distal, (Henschike’s Al, 30-33" SAM P18693, 2
pieces, proximal, with articular surface badly
eroded, and distal (Henschke's area A4, 150 om,
western wall), Ph, SAM Pfs059, first phalanx,
second digit (Henschke’s area A3, 33-36"); SAM
P18248, second phalanx, third digit, (Henschke’s);
SAM P18249, first phalanx, Fourth digit,
(Henschke’s}; SAM Pi8252, first phalanx, fourth
digit, (Hensclike's). Pleistocene,
Salt Creek, South Australia — &, SAM P1710),
proximal shaft-only, but head and part of trochanter
not preserved. Quaternary,
Thorlindah, (Paroo River), Queensland — 77b.,
MMF 12074, figured (Etheridge 1889), a cast (AM
£516) has been made, distal end, condyles worn,
Pleistocene.
Trogdip Cave, Buchan Caves, Victoria — Trt.,
NMY Pi57343, shaft only. Pleistocene.
Warturton River, South Australis — i, UCMP
56642, 2ist or 22nd vertebra, centrum only (Green
Blut lovality, UCMP V-5775). Syn, UCMP 5647,
fragment, fused sacral vertebrae only. F, HM
801/934, distal, moat of internal condyle nol
preserved, {Kalamurina), see Figure 4. Ter, SAM
P1318, distal end, second trochlea not preserved.
(Stony crossing of Warburton, Six road miles west
of new Kalamurina Station.) Quaternary.
Wellingion Caves, New South Wales — 0b, AM
‘Ri’, distal end, external condyle missing; AM
F10949, distal half, (J. Mahoney, in a note on the
back of the museum label, disputes this locality),
Tint, unregistered AM. proximal] articular surface
badly worn; AMC’, proximal; AM F18935, discal
end and part of shaft, second trochlea not
preserved, other trochlea pitted, no distal foramen,
juvenile, AM MBF7T7I, distal. Quaternary.
Wombeyan Quarry, New South Wales — Tih,
AM P58025, distal end and distal one-third of shalt.
Tint, AM PS8026, proximal, Late Pleistocene,
Comments and Descriplion
Dromeius nevaehollandiae is the only extart
species. A number of subspecies have been
suggested (Condon 3975), but little is known of
their ranges or morphological distinctness. The
osteological characteristics of the species have been
described above. There is a suggestion that a slightly
smaller, as well as a larger form of Drowiaius
novaehollandiag existed during the Pleistocene, The
flame gracilipes proposed by De Vis for the smaller
form was applied to a juvenile D. noveehollandiae
{see below). Hence the smaller form, if real, is yer
unnamed. As the evidence is limited, we have
chosen nol ( create @ separate specific or
subspecific names. De Vis alsa described a larger
THE BOSSIL HISTORY OF THE EMUS, BROALAIUS (AVES: DROMAIINAE) 95
species of emu, B patricius, which we have also
synionymized with B. navaehollandiae {see below},
Dromaius patricias (De Vis)
Lecrolype (here designated)
QM F847, proximal right tibiotarsus, King
Creek, Darling Downs, south-eastern Queensland,
Pleisiocene,
Measurerients
Tables 9 and WJ,
Referred Material
De Vis assigned a coracoid, and a proximal and
4 distal tibiotarsus to this species without naming
a type specimen. The left coracoid, OM FI120, was
only provisianally referred to parriciis (De Vis L888;
1291). In actual fact itis not even a bird bore. The
bone is not hollow, and it projects too far lateral
to the point taken by De Vis for the glenold facet
to conform with an emu coracoid, Additionally, it
lacks a pneumatic foramen, [t 15 too large, heavy,
and robust to match any bird, ft is probably part
of a mammalian pelvis.
The distal ead of the left (not right as De Vis
states) tibiolarsus, QM £5548, which De Vis
assigned to D. pa(ricius is not distinguishable from
D, novaehollandiae. De Vis also stated (p. 1290)
that: ‘the rotular surface is relatively longer fore and
all to a considerable extent and less concave
transversely’, but he admitted (p. 1291) that this: ‘is
perhaps in some measure due to abrasion’. The
difference in the ‘eminences and ridges for niuscle
ingertions’ anteriorly are also as De Vis states
{p. 1291): ‘scarcely of specific value’, This specimen
is within the size range of 2 noveehollandiae.
Hence we designate the proximal right tiblo-
tursus, QM F5547, as the lectotype of Dramans
patricius, It is in most respects trivially different,
if al all from BD novaehollandiae. The proximal
width (of the articular surface) is greater than any
0 sovaekollandiae in our sample (57.6 mrn vs a
maximum of 55.6. mm for xevaehollandiae, sample
size, 2 =9), The inner cnemial crest (> 90.2 mm) is
unfortunately not entirely preserved. lL may have
exceeded (he maximum of our sample a! D. novae-
hollandiae (103.1 mm), Concerning other points
raised by De Vis, the fibular crest does oot attach
more proximally, but the bone is thicker at the most
proximal point of this crest, the external enemtal
eresr does descend more distally and a groove
between the external and internal cnemial crests
does exist that is larger than is present in DB, novae-
Hollandiae, We feel that the variability exhibired by
D, patricius would not fall outside that of a large
samtple af che lying emu,
De Vis (1905) also referred a synsacral fragment
(consisting of (He neiral canal of several syasacral
vertebrae), OM FS549, to D, pareicins, though itis
so incomplete as lo render diagnosis difficult. It is
within the size range of D. nevaehollandiae and
probably could be referred to that species, We have,
however, chosen to assign it only to Casuariidae
indeterminate, [t was collected from Wurdulu-
mankula, a Cooper Creek tocality,
De Vis (1892) referred a part of a distal end of
a femur (likely to be QM F143, in part), the
proximal third of a tarsometatarsus (likely QM
F121), the ‘valcaneal region of another metatarse'
(apparently lost subsequently) and a distal
tarsometatarsus (likely QM FIl43 in pari) to
D. patricius. As De Vis did not figure or describe
these specimens in any detail, we are assuming chat
the QM specimens listed are those referred to In his
{892 paper, They agree with the (limited)
description, were collected in the Darling Downs
according to the museum labels accompanying the
specimens, and appear to bear (on the fossils
themselves) De Vis' handwriting. OF these, only the
distal tarsometatarsus is. described, t is stated to
be larver in almost all of its dimensions than the
living Emu. It is, indeed, wider than any in our
small sample both in the shaft and im ita lrochlear
eXpansion, and trochlea 3 is deeper than those in
our sample of B noyvaehollundiae. The proximal
tarsometatarsus is very worn; it lacks both the
internal and external cotyla, the intercotylar region,
and the hypotarsus. It is within the size range of
the living D, nevaehollandiae, The femur is. also
very frazmentary; only the popliteal region is
preserved, and it, too, lies Within the range of
D. novaechollandiae.
Etheridge (1889) referred a distal right tibiolarsus,
MM F 12074 (and cast AM L516) to D. parricins,
but contrary to his assertion, the specimen is net
lareer than nor of a different shape to that element
in the modern emu. I should be reterred to BD.
novaehollandiae.
Thus, some of the specimens that have been
considered to be D. patricius are indistinguishable
from DB. novaehallandiae. Others may he slightly
larger in some measurements, but we doubt that
D potricius deserves, on that account, specific
status, because our sample of the living emu is still
small. We, therefore, synonymize D, patricins will
D, navachallandiae.
Dromaius gracilipes (De Vis)
Holotype
Dnt. QM Fil4d2, Darling Downs, Queensland,
Pleistocene,
Measurements
Tables 10 and 11.
Deswvription
De Vis based this species on a distal left
tarsometatarsus, QM P1142, with the second and
6 © PATTERSON & PV RICH
fourth trochieae not preserved and che margins of
the Uvird rrochlea very eroded, The characteristics
which De Vis used to distinguish thas fram 2
novachollonidiee are the lack of w distal foramen
and associated miuscle canal, infecior size of the
distal end, antero-posterior compression of the
Shalt, and disproportionate size of the mesial
trachlea, These are all juvenile characteristics, De
Vis also states that the width of the third trochlea
taken from centre of the lateral depressions (i.e, the
ligamental pits) is greater in D, gracilipes than in
D, novachollandiae. This is not the case. We
therefore synonymize 2 gracilipes with D. novae-
Aullendize,
Metapterys bifrons (De Vis)
The
Tint. QM FII3S, locality not given, but
presumably Darhng Downs, Queenstand,
Pleistacene,
Measturemnents
Table 1.
Deseriprian
De Vis erected this genus and speries on the basis
of a lef distal tarsometatarsus, QM F1I35, and
allied it with the kiwis because: ‘the trochlea appear
10 be borne on the ends of moderately long stalks’
(De Vis 1892: 449), the lateral trochlear processes
(i.e. the second and fourth trochleae) are almost
equal in length, the medial trochlea extends beyond
the orhers, the posterior surface of the shaft shows
the lines of junction between the coalesced
segments, and it lacks a distal foramen which
perforales the shaft. De Vis considered the
possibility that these might be juvenile
characteristics, but unfortunately dismissed this
idea. De Vis was of the opiiiion that the fourth
trochlea Was shorter (ihal it was a righr
tarsomelatarsus), but actually the second trochlea
is shorter. The comparatively large intertrochlear
notches, the rough pitted vaps on the trochleae, and
the points raised by De Vis (presence of epiphyseal
lines, lack of distal foramen) are indications thal
the specimen is of a juvenile bird. As De Vis himself
noted, the fossil does not have any articulation for
the hallux possessed by kiwis, and the middle
trochlea is too large for that of a kiwi. Metapteryx:
bifrons in all respects conforms to a juvenile
individual of 2 navaehollandioe, and we
synonymize it with that species
Dromaius ocypus (Miller)
Holotype
Tint, SAM Pi3444, Lawson-Daily Quarry, Lake
Palankarinna, easfem Lake Eyre Basin, South
Australia, Mampuwordo Sands, Palankarinna
fauna, Phooene.
Measurements
Tables 8, 9, 10 and 11,
Referred Material
F UCMP RASS5176, condyles, trochanter, and
mast or head not preserved. Th, UCMP RASSI82,
distal, Same locality as type,
Description
Miller (1963) established this species from an
essentially complete, but somewhat distorted and
cracked right tarsometatarsus, SAM P13444, The
overall length, width across the distal end, depth
across the internal cotyla and proximal width are
all less than similar measurements in living and
fossil D, novaehollandiue. Additionally, the
curvature of the intercotylar region is. moré
pronounced (convex dorsally) than in D, novae-
hollandiae, As noted by Miller (1963), the trachleae
have already allained the size and proportions of
2 novaehollandiae. Vhe width across the distal end
is smaller, in part because the intertrochlear notches
are narrower than in D, noveehollundiae,
The femur and Ubiotarsus are provisionally
assigned (0 D, acypus, although they lie within the
range of D. novaehallandiae in those parts which
are preserved, because they were found |i the
locality of the type specimen. The tarsometatarsus
of D. acypus differs most noticeably from D. novae-
hollandiae in its shortness. Unfortunately, the
referred specimens are incomplete, and their length
cannot be ascertained,
Dramatus gidjur i. sp.
Holotype
SAM P26779. Associated Jett leg elements,
Type locality
Leaf locality (UCMP V6213), Lake Ngapakaldi,
eastern Lake Eyre Basin, South Australia, Wipajirl
Fm. Kuljamarpu fauna, Medial Miocene.
Descripiion
An incomplete left lez, consisting of the distal
fragment of the ubiotersus (originally UCMP
71397), the proximal part of the tarsometatarsus
(originally UCMP 71398), and a complete pes
(originally UCMP RASS234}, The tibiotarsal
fragment articulates with the tarsometatarsal
fragment, The pes is complete but does not arti-
culate with the tarsometatarsal fragment as the
trochleae of the tarsometatarsus are not preserved.
All fragments were found in close proximity and
the assumption is made here thal they are from one
individual, Dr R.A. Stirton in his field notes of 19
July 1962 assumed thal the leg and foot elements
were all from one individual and assigned them a
single field number RAS 5234.
THE FOSSIL HISTORY OF THE EMUS, DROMA/US (AVES: DROMAIINAB) 97
Etymalogy
From an Aboriginal word meaning ‘small’
(Anonymous 1965, language not specified).
Meashrements
‘Tables 9, (0 and 12.
Referred material
None,
Diagnosis
A small emu with a slender antero-posteriorly
compressed tibiotarsus and tarsometatarsus, The
anterior lip of the Intercotylar region is convex
dorsally in D. gidju (as in D. ocypus compared tu
nearly Hatin D. novaehollandiae). The intercotylar
region does not extend far proximal to the articular
surface as it does in Casuarius, however, The lateral
lip of the external cotyla is noticeably convex
laterally in D gidju (weakly so in D, novae-
hallandiae), The width and depth of the proximal
articular surface are much less than in D. oeypus
and D. novaehollandiae. The D, gidju tibiotarsus
is much smaller than those In our D. novee-
hollandiae sample, but is similar in general
appearance and proportions of the distal end,
Anteriorly the tarsometatarsus proximal to che
condyles is somewhat crushed, The external
ligzamental prominence above the external liga-
mental pit is not as well defined in D, gidju as it
is in D, eavoehollanciae. The anterior ligamental
fossa appears proportionally larger and deeper than
in D, novaehollandiae. The phalanges (except the
second phalanx of Lhe second digit) are smaller than
those ia & novaehallandiae, Digit 11 1s compara-
tively longer relative to the other digits than in
D, novaeliallandine (64% of the length of digit ILI
versus 57,5-460.5% for our sample of De nevege-
hollandiae (see Tables 2 and 12); the first phalanx
of digic It is 88% of the proximal depth of the first
phalanx of digit [11 versus 76.5-78% for our sample
of 2D novaehollandiae, and 66% of the proximal
width of digit LL yersus 52-53% for our sample
of D. novaehollandiae), Digit 1V is also Compara-
tively longer but less change has occurred (68% of
the length of digit IL versus.62-64% for our sample
of D, novaehollandiae, the first phalanx of digit 1V
is 78% of the depth of the first phalanx digit IIE
versus 73-75% in our sample of 2 novae-
hollandiae). Excepting the proximal phalanges, the
ratio of maximum proximal depth to width ts
greater in 2 gidju; thus, the phalanges of A novue-
hollondiae are more dorso-ventrally compressed
than in DO gid/u. The ungual phalanx of digit IT
in D gidjn is longer than the ungual of digit tH]
(which ts poorly preserved). This is not duc to an
clongation of the ungual of digit Hin gidje,as
in Casuarlus, but the weak development of the digit
JIL unpual.
Comment
From what 1s known of its hind limb structure,
i) would appear that D. gidju was less cursorially
adapted than D. novaehollandiae. This is based on
the greater length of digits Wand LY relative to dapat
Wiin D. gidjv as compared to D. novaehollandiae.
This. foot structure is presumably adapted for
Breater maneuverability in forested or less open
conditions and greater ability Lo qove over regions
of a somewhat unpredictable nature.
Dromaius sp. indet.
Several specimens because of their fragmentary
nature and/or Wnusual proportions could pot be
assigned (o species level, Other specimens were
referrable only to Casuarlidac indet. (sce following
section).
Referred Material
Tables 3-12
No Locality Data — Jini, AM F38087_ distal end,
trochleae not preserved,
Alcoota, Northern Territory — Tint, QM
QA205, third trochlea only, QM QASOS, distal,
fourth trochlea not preserved, UCMP RASS397 (in
part), a third trovhica only. PA, OM QASO4, first
phalanx, third digi; UCMP RAS $397 (in part),
second phalanx, second digit. The Afvoota
specimens are close fo & gidyu. Late Miocene.
Baldina Creek? (near Burra). South Australia —
Ff, SAM P17103, shalt only, partially reconstituted
in plaster, Quaternary,
Cooper Creek, South Australia — Tit, QM
FIN21, proximal, badly eroded articular surface:
Could be Pliocene-Recent in ave,
Lancetield, Victoria — Tht. NM'V P35578, distal
shaft, Pleistocene in age, dated at about 26 000 BP.
(Qillesple ez af, 1978).
Warburton River, Suuth Austratia — Db, OM
F6668,, distal, condyles not preserved, (‘Kalie-
murina?’ ts pencilled on the bane). Tit, QM F667),
proximal, incomplete fusion of metatarsals dorsally,
(atsal cajy not preserved, juvenile, Pleistoccne-
Recent in age
Wellington Caves, New South Wales — Si, AM
F25218, figured (Anderson, 1934), incomplete, Thur,
AM !A' distal, trachleae nor preserved; AM FLO8S0
(cf. DL nevaehollandiae), proximal end plus
proximal part of shaft, articular surface eroded.
Pletslocene.
Dramaiinac inset.
Referred Material
Tables 4-12.
No Location Data — Syn,, OM £6673, taecnent,
fused vertebrae only.
98 ©, PATTENSON & PV. RICEL
Bingara, New South Wales — Syn, MM FI6795,
fused vertebrae only, Pleistocene,
Couper Creek, South Australia — , HM BT76,
dorsal vertebra, 24th-26th?, centrum and left
postzygapophysis only, (Lower Cooper, locality 2).
Ff, HM 8777, part of a lelt internal condyle and
internal popliteal fossa region anlyy UCMP 60532,
popliteal fossa region only, (Karipiri Waterhaole,
UCMP site 9, V-586J), The locality suggests all
Speciniens are Oromiais sp. Pleistocene.
Lake Kanunka, South Ausiralia — r, UCMP
60560, third or fourth vertebrae?, proximal (dorsal)
only, (Site |, UCMP V-S772). Probably Dramatus:
other Dramaius dements known here Probably
Late Pliocene, possibly Pleistocene.
Lake Kittakittanolow, South Australia — Thr,
SLAM 75, fourth trochlea only, pitted, juvenile. Late
Pliocene, possibly Pleistocene,
Mebachern's Cave, Victoria — KF or NMYV
P157354, dorsal lrazmenls NMY PIS7358, dorsal
fragment; NMV P157362, dorsal fragment; NMV
P)57366, dorsal fragment, one facet not preserved,
Probably Dromains. Quaternary,
Naracoorte, South Australia — Kr, SAM P1807,
third vertebrae?, dorsal Fragment, (Heuschke's Bone
Dig); SAM PL8251, dorsal fragineut only but facers
not preserved, (Henschke’s Bone Dig); SAM
P8784, third vertebrae?, dorsal fragment
(Hensthke’s Bone Dig}i SAM P22812 {in pari), third
or fourth vertebrae?, dorsal fragment, The lovaliry
sugpests these speciniens are Dvevrtuins. Quaternary,
probably Late Pleistocene.
Warburton River, South Adstralia — Syn.,
UCMP 56647, acetabulum and pectineal process
only, Quaternary.
Wombeyan Quarry, New South Wales — Tih,
AM. P58027, ef, Bromaius, distal end. Late
Pleistocerie,
Aves indet,
(previously assigned to Dromraius)
Referred Material
Tables 10 and 11
Kalamurina, South Australia — Tier, SAM
P11552, distal end, second trachlea not preserved,
large. Perhaps Dromornithidae (Table Li),
Quaternary,
Nu Locality Data — 7/b,, SAM P1748, proximal
shall anly, articular surface and cnemial crests not
preserved. Possibly net avian — the bone js rather
dense.
Dromaius ater and D. bauddinianus are pot
reviewed here as Parker (1984) has. recently revised
their taxonomy,
Discussian
Al least One species of emu, 2B gidju, was presen
in central Australia in the mid-Miocene. Itis known
from the Lake Ngapakaldi jn northern Sout
Australia and associated with the Kutjamvarpu
fauna. It does not differ suffictently from other
emus to require erection of a new genus. While the
intercotylar region of the fossil tarsometatarsus
resembles the condition found in Casuariys in that
is if not markedly flattened as in the Ilving
Promaius, this character stale is also Inve of B
ocypus, an undoubted emu from the Pliovene. The
shape of the margin of the internal cotyla of the
fossil ps similar to that of Dromaius and dissimilar
to that of Casuarius, which is more excavated
posteriorly. The posterior surface of the
tarsometatarsus of D. gidju is unfortunately
chipped and cracked, but the shape of the remaining
fragment of the hypotarsus and Larsal cap appears
more emu-like Lhan cassowary-tike. Comparing the
pes of the Miocene fossil with recent emus it is
obvious that lhe foor structure has undergone
change through the last few million years, The
second.alid, (oa lesser extent, the fourth toes haye
undergone a reduction in size, This (rend is parallel
with several other ratites (the ostrich and some of
ihe Dromornithidae; Rich 1979), and would appear
to be a cursorial adaptation similar to the reduction
and eventual loss of Jateraland medial digits within
horses (Equidae), The pes of D giejy is more
cassowary-like than that of any known living or
other fossil emus, implying that in the Miocene,
emus were not as highly adapted to an open plains,
cursorial lifestyle as they are now. The pes does not,
however, contain the specialized ungual spike on
digit U, which so characterizes the cassowaries, D,
gigju appears to bea species that may be close to
forms which gave rise to both emus and cassowarles.
Based on material now ayailable, D gidju has a few
specialized characteristics that seem to ally it with
Dromaius, But, as our records of this form increase,
there may be sufficient reason Lo separate it from
both genera within this family as an early, quite
Unspecialized form.
A small emu of Miocene age, is also known from
Riversicigh, Queensland. This material is currently
under study by Waller Boles (Australian Museum),
By Late Miocene or Barly Pliocene times a species
near in size to D gid/u is kwown in the Alcoota
fauna. It is represented by tarsometatarsal and pedal
fragments. This form may be referable to DL gidju,
The two phatanges dilfér slightly from those of the
Lake Ngapakaldi form, which we have referred to
D) gidju, bin are within the range expecred for
intraspecific variation. Until more complete
miaienal is available from Alcaota, however,
Assignment (o Dremelus sp, indet, is preferable. It
TIE FOSSIL HISTORY OF THE BMUS, BROMATLS (AVES: DROMAIINAE) 99
is certainly not referrable to Cisuarits, as the
second trochlea ig much more reduced relative to
the other trochleae, similar to the condition in
Dramatis.
By mid-Plioceng, a second species of emt,
D. ecypus Miller, intermediate in size between
D, gidjue and D. novaehollandiae. existed. It 1s
known from a right larsometatarsus atid part of a
fenjur and Gbiotarsus. The femur and tibiotarsus
are, in those parts which are preserved, not unlike
PD. novaehallandiae, but if complete would probably
be shorter than the corresponding elements in
Db novyuehollandiae,
In any event, 2 novaehollandiae existed by the
Late Pliocene or Early Pleistocene. Since then only
D, novaehollandiae has been present on the
mainland, [thas probably fluctuated slightly in size,
presumably as a result of a host of selection
Pressures such as climate (both temperature and
rainfall), diet, predation pressure and competition.
The species restricted in Recent times to King
Island and now extinct may possibly extend Into the
Pleistocene. Localities on King Island are as yet nol
carefully dated, The emus of King Island (2 runor)
and Kangaroo Island (D bavdinianus) appear to
be separale species (Parker 1984), The populations
oo both of these are most likely relics isolated by
rising sea Jevels atthe end of the Jast glaciation of
&@ population that perhaps Wag once more
widespread, Other than their smaller size the King
and Kangaroo Island emus differ but little from the
mainland emu. The main osteolagical difference is
in the shape of the skull (Spencer & Kershaw 1910,
Morgan & Sutton 1928). Possibly also the distal
foramen of the tarsometatarsus differs (Shane
Parker pers, comm.), but there is considerable
variability in this character in the mainland emu,
The degree to which whe groove for che musculus
adductor digiti 1V is roofed over by bone would
appear to be age related. In juvenile crus the arch
is almosi completely lacking, whereas in some adult
specimens it is completely formed ln bone. Thus,
this character is aot taxonomically significant for
emus, Some Australian mainland fossil emus are
within or jusr larger thaw the size range for D, miner
tabulated in Rich (1979). At Lake Menindee, two
distal tibiotarsi (UCMP 53825, includes RHT6 and
RHT25) lic within the range of DB eninor for the
width across the condyles and depth of external
condyle but exceed DB minor in the depth of the
internal candyle — both specimens measure
37.0 mm in depth of the internal condyle; the range
of asample of 50 2. minor was 26.2-36,1 mm, the
mean was 30.4 (Rich 1979, Table 33). What the
relationship of these fossils is to 2 minor \s
unresolved and will remain so until a much larger
fossil sainple of mainland birds is at hand.
De Vis (892) did deserjbe a smaller mainland
Pleistocene species, D, gracilipes, but hls type
specimen is undoubtedly an immature PD, novae
hallandiae, Nevertheless, smaller emus did exist in
Australia In the Pleistocene, OF specimens listed in
the systematics section (above) the following lic
below the range for modern emus in one ar mote
measurements: AM — unregistered tarsomete-
tarsus, “A', Fi0949, MF773; HM B775/869; QM
Fit2l; SAM — P1318, PI8099,; UCMP — 53825,
53832, 35983, 60532, 79510, RHTIO64.
The presently known fossils of mainiand emits
smaller than the living 2 sovaehellandine are
unfortunately few. We do not believe that they are
represeniatives of D minor or D. haudinianus,
because of the age of the fossils on the islands ly
Late Quaternary. We favour the idea that speciation
on King and Kangaroo Island could have taken
place in very iittle (ime geologically speaking,
Strong selection for dwarfism quite likely occurred
alter chese emus became isolated al the beginning
of the last interglacial (Le. the Holocene).
The mainland emu, 2 nevarkallandiae, may
have been at any One time in the Pletstocene bouk
larger or smaller than at present. However, there
is a possibility that the differences seen in fossil
samples are more apparent than real, since the
sample size of modern emus is still fairly small and
some of the emus in osteological collections were
recovered from zoological gardens, Whether or not
extant wild emu populations differed significantly
in size is largely unknown, As our sample did not
contain representatives from the Northern Terntory
or Western Australia, it is also unavoidably biased
geouraphically.
Periodic dwarfing of the mainland form may
have been caused by the same selective agents which
produced dwarfing in the island forms, We were
unable to test che hypothesis that size changes were
related 10 palaeoclimate or other environmental
variables because there are coo few reliably dated
emu specimens in the Quaternary collections.
De Vis recognized 2 pairicius as a separate,
larger species of emu, but we can see no significant
size difference from B raveettollandiae, Perhaps
D. patricius differed in its proportions from
D. noveehollandiae, As so few complete bones are
known, however, this is difficult to assess, Por
example, a tibiatarsus SIAM 6) was found to have
a smaller length to distal width ratio than most
modern emus, bur this difference did not prove
statistically significant (9>O0.405, t-lest)-
There was a mass extinction of the Australian
megalauna fihe larger macropodids,
diprotodontids, dromornithids, ete just before the
Holocene (Tedford 1967, Gillespie er al 1978))
suggesting a widely acting selective agent against
large size,
There is a suggestion thar the Tasmanian emu,
UX) (~ PATTERSON & BP. RICH
D. diemenensis, averaged slightly smaller than the
mainland form, This idéa stenimed from the known
eggs of the Tasmanian emu measuring slightly
smoaller than those of the mainland emu (Dove
1926), and from the recollections of Legge (1907),
Who saw the Tasmanian emu as a boy. On the other
hand, Spencer & Kershaw (1910) report that the Rev,
Knopwood captured an ‘Emew 60 lbs. weight’ on
9 October 1804 in Tasmania, Scoll (1924) gives the
dimensions of a leg of the Lismanian emu collected
by Gunn in the 19th century; it ts as large as those
of the mainland, The fossils of the Tasmanian emu
are large (Scott, 1924, 1932) indicating thal the
larger Pleistocene form of the mainland also
reached Tasmania, presumably ata tinie when Bass
Strait did not exist — during a glacial period of
lowered sealevel, Concerning the extinct Recent
Tasmanian emu, the best evidence supports the view
thal it was about the same size as (he mainland emu,
Condon (1975), following Ridpath & Moreau
(1966), treated the Tasmanian emu as a subspecies
of D. novaehollandiae, Kathryn Medlock
(Tasmanian Museum) is currently reviewing the
status of this form,
ACKNOWLEDGMENTS
Our thanks go to many individuals and
Histitutions that provided specimens utilized in this
study ; W, Boles (Australian Museum), G, George
(Sir Colin McKenzie Wildlife Sancinary, Hewles-
ville), 1, Hope (formerly of the Australian National
University), A. MceBvey (Museum of Victoria), M-
McKenna (American Museum of Natural History),
K, McNamara (Western Australian) Museum), R.
Molnar (Queensland Museum), 8. Parker (South
Australian Museum),.J. Pickett (Mining Museum),
N_ Pledge (South Australian Museum), T. Rich
(Museuny of Victoria), W, D. 1, Rolfe (Aunterian
Museum), G, F van Tels (CSIRO, Division of
Wildlife and Rangclands Research, Canberra), and
D, Vernon (Queensland Museum), R. F. Baird, S,
Parker, T. Flannery, F. Szalay, N, S, Pledge and G.
F, van Tets critically read (he manuscript, and we
are graletul for their time,
We are sincerely grateful to all those field parties
Which collected the original material, in particular
those led by the fate R, A, Stirton (University of
California, Berkeley) and R_ H. Tedford (Americar
Museum of Natural History)
Photography of specimens was provided by R.
Smart, &, Coffa and E, M. Thompson (Museum
of Victoria) and &. Morton and T. Zylstra (Monash
University). Drafting was provided by D, Gelt, P,
Hermansen, J, Muir and B, Shea typed varlous
versions of the manuscript. Funding allowing the
complelion of this project was provided by the
National Geographic Sociery, the Australian
Researeh Grants Scheme, the Commonwealth
Education Department, and the Ingram Trust. We
owe them our deep gratitude, Both the Garth
Sciences and the Zoology Departments, Monash
University, provided the venue for our work.
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TABLE 2. Statistical summaries of the extant emu Dromaius novaehollandiae (X, mean; s, standard
deviation; n, sample size).
Range (mm) x s n
Skull
Length 140 -165 154 6.3 16
Width 58.7- 76.5 68.4 4.1 21
Depth 44.7- 50.7 48.0 1.6 19
Diameter of Foramen Magnum 9.3- 13.1 11.0 1.0 25
Length of Lower Jaw 131 -155 145 6.5 21
Symphysial Length of Lower Jaw 16.4- 23.0 21.0 2.4 23
Sternum
Maximum Length 114 -164 143 11.4 25
Maximum Width 104 -141 125 8.2 25
Number of Costal Processes a eS 4 0.49 28
Width of First Costal Process 98.7-134 116 8.2 24
Width of Last Costal Process 76,4-112 § 96.8 8.2 24
Length of Costal Margin 42.4- 66.9 56.0 7.2 24
Length of Sternocoracoidal Process 16.3- 44.7 33.3 8.2 24
Width of Coracoidal Sulci 40.2- 62.2 51.9 5.9 24
Anterior Depth 14.4- 20.4 16.7 2.0 23
Scapulacoracoid
Proximal Width 38.9- 55.6 45.9 4.9 20
Maximum Length 151 -187 168 11.3 18
Scapular Length 98.4-127 114 8.7 18
Minimum Width of Coracoid 13.1- 22.0 16.1 2,2 20
Minimum Width of Scapula 5.9- 8.9 73 1,0 19
Clavicles
Length 35.2- 53,3 44.0 3.1 12
Maximum Width 3.6- 8.1 5.5 1.2 12
Depth 2.0- 4.9 3.4 0.8 12
Humerus
Length 83.1- 98.7 90.3 4.2 20
Proximal Width 5.4—- 8.3 6.4 0.8 20
Proximal Depth 6.2- 7.9 6.9 0.5 20
Ulna
Length 57.5- 73.0 64.9 4.5 20
Proximal Width 3.6- 4.8 4.2 0.3 20
Proximal Depth 3.3- 4.8 4.0 0.4 20
Radius
Length 55.2- 68.9 63.1 4.1 20
Carpometacarpus
Length 36.6- 50.6 43.6 3.6 20
Proximal Width 7.7- 12.3 10,0 1.2 20
Proximal Depth 4.6- 6.5 5.5 0.5 20
Distal Depth 2.5- 4.4 3.4 0.5 19
Manus
Pl, Length 10.0- 26.6 13.6 4.1 13
Pl, Proximal Diameter 4.2- 6.9 5.0 0.8 Il
P2, Length 4.8- 8.0 6.3 1.4 5
P2, Proximal Diameter 1.9- 3.7 3.0 0.7 5
P3, Length 6.1- 14.9 10.4 2.7 6
P3, Proximal Diameter 14.0- §.2 2.8 1.3 6
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 103
Range (mm) x s n
Synsacrum
Length 337 -412 378 24.7 17
Diameter of Acetabular Foramen 12.3- 21.0 17.3 2.5 23
Width across Antitrochanter 90.9-109 101 5.3 22
Maximum Depth 88.7-109.2 99.3 4.2 23
Femur
Length 175 -218 203 10.1 22
Proximal Width 62.0- 68.0 64.9 2.0 10
Proximal Depth (Trochanter) 55.8- 63.4 59.7 Zid 10
Diameter of Head 23.5- 29.7 26.4 ss 26
Distal Width 65.4- 79.2 yf Wes 3:2 26
Depth of External Condyle 64.0- 75.4 69.3 2.9 26
Tibiotarsus
Length 340 -432 401 21.8 18
Diameter of Shaft, Minimum 19.5- 27.0 23.1 1.8 23
Diameter of Shaft, Maximum 24.6- 34.2 28.1 2.2 23
Proximal Depth 86.2-103 96.2 5.6 9
Proximal Width 47.4- 55.6 52.4 2.8 9
Length of Fibular Crest 74.6-110 90.4 120 9
Depth, Internal Condyle 38.7- 47.4 42.7 2.8 9
Depth, External Condyle 36.5- 45.9 41.8 2.1 26
Width, Distal End 38.9- 49.1 45.9 2;2 25
Fibula
Length 231 -305 272 20.9 12
Proximal Width 13.9- 19.0 17.1 1.5 22
Proximal Depth 35.2- 48.7 38.7 pt 22
Tarsometatarsus
Length 332 -422 383 18.0 22
Minimum Diameter of Shaft 11.6- 17.3 14.7 15 23
Maximum Diameter of Shaft 16.8- 23.1 19.9 1.5 23
Proximal Width 47.2- 54.0 50.0 2.1 25
Depth of Internal Cotyla 25.4- 27.6 26.6 0.8 9
Depth of External Cotyla 19.9- 23.7 22.0 12 9
Depth of Hypotarsus 36.0- 41.3 38.5 LSS) 8
Distal Width 47.4- 54.6 51.1 2.0 8
Tarsometatarsus, trochleae
Width T2 9.0- 11.1 10.0 0.8 9
Width T3 21.9- 28.9 24.9 2.0 24
Width T4 12.2- 14.9 13.6 1.0 9
Depth T2 13.0- 17.6 15.4 1.4 9
Depth T3 19.0- 24.3 22.2 1.8 9
Depth T4 14.3- 17.2 15.6 IQ 9
Pes
DII P1 Length 40.7- 52.8 47.4 2.8 14
Proximal Depth 16.3- 21.4 18.5 1,7 15
Proximal Width 13.2- 16.4 14.8 0.8 15
DII P2 Length 17.1- 22.4 19.5 2.1 13
Proximal Depth 11.6- 13.3 12.3 0.6 13
Proximal Width 12.4- 15.6 14.0 1.0 13
DII P3 Length 21.4- 28.2 25.9 22 9
Proximal Depth 9.4- 12.2 11.0 0.8 10
Proximal Width 10.4- 12.9 11.5 0.9 10
DIII P1 Length 58.1- 65.7 60.5 1.9 14
Proximal Depth 21.3- 27.3 23.6 155 15
Proximal Width 25.3- 31.1 275 2.0 15
DIII P2 Length 33.4- 42.8 38.8 2.3 13
Proximal Depth 15.0- 18.9 16.9 0.9 15
Proximal Width 20.3- 25.4 22.5 1.6 14
104
C. PATTERSON & P. V. RICH
Range (mm) x s n
DIT P3 Length 19.5 — 29.3 23.6 2.8 12
Proximal Depth 12,1- 14.1 13.1 0.7 13
Proximal Width 17.2- 20.8 19.0 0.9 13
DIII P4 Length 26.7- 34.4 30.4 2.1 9
Proximal Depth 11.7- 14.9 12.9 1,0 ll
Proximal Width 14.8- 17.7 15.6 0.9 11
DIV P1 Length 33.7- 41.2 38.5 2.0 14
Proximal Depth 16.0- 19.8 17.3 0.8 15
Proximal Width 16.0- 19.2 18.0 0.8 15
DIV P2 Length 14.9- 18.2 17.0 1.0 13
Proximal Depth 11.1- 13.2 12.5 0.6 13
Proximal Width 14.2- 16.0 15.1 0.5 13
DIV P3 Length 10.5- 14.4 12.0 1.1 11
Proximal Depth 9.9- 12.1 11.0 0.6 12
Proximal Width 12.5- 14.9 13.2 0.6 12
DIV P4 Length 6.4— 12,3 9.7 1.6 ll
Proximal Depth 8.7- 10.7 10.1 0.6 ll
Proximal Width 10.4- 13.4 12.0 0.8 ll
DIV P5 Length 19.2- 24,3 22.2 1.6 8
Proximal Depth 9.3- 11.8 10.5 0.7 10
Proximal Width 10.1- 11,5 10.9 0.5 10
Vertebrae
Cl Length, Ventral 5.0- 7.9 6.1 0.8 12
Depth of Hypopophysis $.3- 6.1 5.8 0.3 11
Maximum Width across Arch 12,.8- 16.5 14.5 1.4 9
Depth 14.9- 17.7 16,1 0.8 12
Prearticular Surface 6.2- 8.5 Vl 0.7 12
Postarticular Surface 9.0- 11.3 10.4 0.8 10
Dorsal Length §.8- 7,3 6.4 0.5 10
C2 Depth 25,7- 30.9 28.1 1.4 16
Width across Postzygapophyses 20.4- 24.9 22.3 1,3 16
Width across Diapophyses 12.2- 15.5 13.8 1.0 16
Width across Postarticular surface 6.3- 8.7 7.3 0.7 16
Width across Prearticular surface 9.7- 11.4 10.5 0.5 14
Centrum Length 15.9- 20.5 18.3 1.5 15
Length from Pre- to Post Zygapophyses 14.1- 17.9 16.6 Ll 12
C3 Depth 21.0- 26.1 23.7 1.3 15
Postzygapophyses 22.7- 28:2 25.4 1.4 16
Diapophyses 17.7- 23.5 21.0 1.4 15
Postarticular Surface 6.9- 9.6 8.4 0.7 16
Prearticular Surface 8.1- 10.9 9.1 0.8 16
Centrum length 16.5— 21.2 18.9 1.4 16
Pre-postzygapophyses 22.3- 27.8 29.8 1.7 16
C4 Depth 18.1- 22.6 20.2 1.3 16
Postzygapophyses 22.1- 26.4 24.5 1.3 16
Diapophyses 21.6- 26.8 23.6 1,3 16
Postarticular Surface 9.5-— 13.0 11.3 1.2 16
Prearticular Surface 7,9- 12.0 10.3 0.9 16
Centrum Length 20,.5- 25.3 22.7 1.6 16
Pre-postzygapophyses 26.4- 32.3 29.4 19 16
C5 Depth 14.9- 19,3 17,1 1.2 17
Postzygapophyses 12.8- 19.5 17.0 1.6 17
Diapophyses 25,0- 28.6 26.2 1.1 l7
Postarticular Surface 13.6- 19.7 16.0 1.6 17
Prearticular Surface 10.5- 15.7 13.3 1.6 17
Centrum Length 22.4- 26.7 25.2 13 17
Pre-postzygapophyses 30.8- 40.6 36.0 2.5 17
C6 Depth 13.9- 19.7 16.5 1.3 17
Postzygapophyses 11.6- 16.4 13.6 1.2 17
THE FOSSIL HISTORY OF THE EMUS, DROMAJUS (AVES: DROMAIINAE)
Range (mm) ¥ s n
Diapophyses 17.4- 31.1 27.8 3.1 17
Postarticular Surface 16.8- 20.6 18,7 1,3 17
Prearticular Surface 14.5- 19.9 17.4 1.7 17
Centrum Length 25.3- 29.4 27.8 1.3 17
Pre-posizygapophyses 35,.7- 42.2 38.8 1.8 17
C7 Depth 15.0- 19.0 16.9 1.2 17
Postzygapophyses 11.5- 17.9 14.0 1.8 17
Diapophyses 27.8- 32.9 30.4 1.7 17
Postarticular Surface 1§.2- 20.1 18.0 1.6 17
Prearticular Surface 18.9- 24.1 21.0 1.4 17
Centrum Length 28.5- 32.3 30.7 1,3 17
Pre-postzygzapophyses 35.7- 41.4 38.4 1.5 17
C8 Depth 16.2- 19.9 18.0 1.1 17
Postzygapophyses 12.1- 19.7 16.8 2.2 17
Diapophyses 18.4- 34.2 30.4 3.6 17
Postarticular Surface 14.4- 19,1 16.9 1,4 17
Prearticular Surface 16.8- 32,2 20.8 3.5 17
Centrum Length 23.6- 35.6 32.9 2.8 17
Pre-postzygapophyses 36.5- 40.5 38.6 1.3 17
C9 Depth 18,2- 22,7 20,2 1,2 17
Postzygapophyses 16.3- 25.0 20.8 2.0 17
Diapophyses 28.4- 34.5 31.2 1.8 17
Postarticular Surface 13.4- 17.9 15.4 1.3 17
Prearticular Surface 16,1- 21.7 19.0 1.4 17
Centrum Length 34.2- 38.7 36.2 1.4 17
Pre-postzygapophyses 38.2- 45.3 40.4 1.9 17
C10 Depth 19.7- 23.6 21.7 1.2 17
Postzygapophyses 19.8- 27.2 22.9 1.6 17
Diapophyses 28.8- 34.7 31.0 1.8 17
Postarticular Surface 12.6- 16,7 14.8 1.2 17
Prearticular Surface 15.2- 21.1 17.5 1.7 17
Centrum Length 36.0- 40.8 38.4 1.6 17
Pre-postzygapophyses 40.5- 50.6 44,2 2.3 17
Cll Depth 21.0- 24.9 23.2 1.2 17
Postzygapophyses 21.2- 27.5 23.4 1.5 17
Diapophyses 29.4- 34,7 31.5 1.6 17
Postarticular Surface 13.8- 18.4 15.4 1.4 17
Centrum Length 37.7- 43,2 40.7 1.7 17
Pre-postzygapophyses 43.6- 53,2 47.3 2.2 17
Prearticular Surface 15.1- 19.9 16.9 1.6 17
C12 Depth 23.0- 27.1 24.6 1.3 17
Postzygapophyses 20.2- 27.5 23.8 1.7 17
Diapophyses 30.1- 35.0 25.0 1.5 17
Postarticular Surface 14.6- 19.4 16.5 1.5 17
Prearticular Surface 15.4- 23,1 17.9 2.3 17
Centrum Length 39.5- 44.4 42.4 1.5 17
Pre-postzygapophyses 44.9- 53.7 49.7 2.3 17
C13 Depth 23,.3- 28.2 25.7 i 17
Postzygapophyses 20.3- 27.0 23.8 1.9 17
Diapophyses 30.7- 37.3 33.7 1.7 17
Postarticular Surface 16.2- 20.6 17.8 1.4 17
Prearticular Surface 15.5- 22.6 18.9 1.9 17
Centrum Length 40.4- 45.9 43.6 1.6 17
Pre-postzygapophyses 47.7- 54.4 51.9 1,9 17
C14 Depth 24.3- 29.7 26.9 1.5 17
Postzygapophyses 20.7- 26.4 23.9 1.7 17
Diapophyses 32.0- 38.5 35.2 1.6 17
Postarticular Surface 17,3- 22.2 19.2 1.4 17
Prearticular Surface 17.1- 23.4 20.4 1.9 17
105
106
C. PATTERSON & P. V. RICH
Range (mm) x s n
Centrum length 41.0- 46.5 44,3 1.7 17
Pre-postzygapophyses 48.6- 56.6 §2.8 2.3 17
C15 Depth 25,6- 32.2 28.6 2.0 17
Postzygapophyses 21.4- 27.4 24.1 1.6 17
Diapophyses 34.5- 40.9 37,1 2.0 17
Postarticular Surface 19,0- 23.5 21.2 1.4 17
Prearticular Surface 17.6- 25.9 22.0 2.1 17
Centrum Length 41.2- 47.4 44,9 1.8 17
Pre-postzygapophyses 49.8- 58.4 53.6 2.5 17
C16 Depth 27.5- 33.5 30.5 2.1 16
Postzygapophyses 22.4- 28.1 25.4 1.7 16
Diapophyses 36,9- 45.7 40.7 2.6 16
Postarticular Surface 20.5- 25.7 23.3 1.6 16
Prearticular Surface 20,4- 29.2 24,1 2,2 16
Centrum Length 41.9- 48.5 45.3 1.9 16
Pre-postzygapophyses SL.1- 60.8 55.0 2.8 16
C17 Depth 29,3- 41.1 33.8 3.2 17
Postzygapophyses 25.1- 30.4 27.2 1.9 17
Diapophyses 41.8- 51.4 45.7 3.2 17
Postarticular Surface 20.9- 27.6 24.5 1.9 \7
Prearticular Surface 23.5- 32.4 26.5 2.2 17
Centrum Length 42.5- 48.7 45.5 2.0 17
Pre-postzygapophyses 50.0- 61.2 55.6 2.8 v7
C or V18 Depth 34.0- 56.3 41.2 5.5 16
Postzygapophyses 25.3- 32.7 28.4 2.0 16
Diapophyses 46.5- 68.4 53.6 5.7 16
Postarticular Surface 21.9- 29.0 25.3 1.8 16
Prearticular Surface 22.4- 31.1 27.6 2.6 16
Centrum Length 42.3- 48.2 45.1 1.8 16
Pre-postzygapophyses 48.4- 60.6 55.4 3.0 16
V19 Depth 43.0- 66.4 50,7 6,3 17
Postzygapophyses 25.2- 31.3 28.3 1.7 17
Diapophyses 56.5- 70.6 61.7 4.0 17
Postarticular Surface 21.6- 27.8 24.9 1.8 17
Prearticular Surface 23.4- 33.1 27.4 2.5 17
Centrum Length 40.9- 47.5 44.4 1.9 17
Pre-postzygapophyses 51.4- 58.4 54.2 3.1 17
V20 Depth 50.7- 77.1 62.6 5.7 7
Postzygapophyses 23.7- 30.2 26.5 1.9 17
Diapophyses 62.1- 81.8 67.5 4.6 17
Postarticular Surface 20.5- 25.6 23.3 1.5 17
Prearticular Surface 23.7- 34.0 27.6 2.7 17
Centrum Length 40.4- 47.6 42.8 1.8 17
Pre-postzygapophyses 49.7- 58.3 54.2 2.4 17
V21 Depth 56.6- 78.1 68.0 5.5 17
Postzygapophyses 23.4- 29,1 26.2 1.6 17
Diapophyses 61.3— 82.0 68.2 4.8 17
Postarticular Surface 19.6- 26.5 22.7 1.5 17
Prearticular Surface 21.4- 29.4 25.3 2.0 17
Centrum Length 37.2- 46.3 42.1 2.4 17
Pre-postzygzapophyses 47.9- 57.0 51.9 2.7 17
V22 Depth 57.1- 74.9 64.7 6.1 17
Postzygapophyses 22.4- 30.5 26.7 2.0 17
Diapophyses 61.0- 76.8 67.0 3,7 17
Postarticular Surface 19,8- 29,1 23.3 2.3 17
Prearticular Surface 21.9- 29.1 24.5 1.9 17
Centrum Length 38.3- 45.5 41.5 2.2 17
Pre-postzygapophyses 47.2- 57.8 50.7 2.9 7
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE)
a
Range (mm)
V23 Depth 55.7- 72.9
Postzygapophyses 25.8- 33.1
Diapophyses 60.9- 74,4
Postarticular Surface 21.0- 26.7
Prearticular Surface 20.3- 29.0
Centrum Length 37.8- 43.7
Pre-postzygapophyses 43.8- 51.6
V24 Depth 60.2- 79.9
Postzygapophyses 26.1- 33.8
Diapophyses 62.1- 75.5
Postarticular Surface 19.8- 28.4
Prearticular Surface 21.3- 28.8
Centrum Length 37.5- 46.1
Pre-postzygapoplyses 43.2- 52.1
V25 Depth 63.7- 85.9
Postzygapophyses 27.9- 40.2
Diapophyses 43.4- 76.2
Postarticular Surface 20.3- 27.3
Prearticular Surface 20.4- 30.3
Centrum Length 36.8- 44.4
Pre-postzygapophyses 42.3- 52.9
V26 Depth 68.5- 87.8
Postzygapophyses 26.2- 40.9
Diapophyses 59.7- 75.6
Postarticular Surface 23.0- 30.4
Prearticular Surface 23.1- 38.7
Centrum Length 33.3- 49.4
Pre-postzygapophyses 41.2- 50.7
x
nw
n
VA NENOwNY
ND RFPANROR Ww
“
~
NN es
ee
*
a
New pw NNN ANY NN?
CAwWWORUONMN NOUCIBON $C
wi
~~
TABLE 3. Measurements (in mm) of skull material of fossil emus (Dromaiinae) from Australia.
SPECIMEN
NMV P157345
NMV P157350
NMV P157353
SAM P17834
Diameter,
Foramen
Width Depth Magnum
75.0 (est.) 47.4 10.8
>65.2 48.5 10.7
Lower Jaw,
Symphysial
Length
21.5
24.5
107
108
C. PATTERSON & P. V. RICH
TABLE 4. Measurements (in mm) of vertebrae of fossil emus (Dromaiinae) from Australia.
SPECIMEN
MM F16786
MM F16797
NMV
P157346
NMV
P157349
NMV
P15735]
NMYV
P157352
NMV
P157359
P157367
NMV
P157368
NMV
P157369
SAM P17589
SAM P18246
SAM P18673
SAM P18830
SAM P18247
AMNH 9678
AMNH 9678
AMNH 9678
AMNH 9678
AMNH 9678
AMNH 9678
UCMP 56642
UCMP 56855
Length
of
Centrum
37,7
38.8
39.3
36.0
24.5
31.2
Length
across
Zygopop.
46.3
42.1
46.3
Vertebral
Number
V24?
V26 or 25?
V21 or 22
V23 or 24
c7? (juv)
C11-13? Guy)
Measurements
See ene SS NN SS Oe
Width
Posterior Anterior
Depth Postzyg. Diapop. Articulation —_ Articulation
_ 42.2 (est.) = 25.5+ 30.0 (est.)
-- 31.8 — 27.0 27.2
_ 23.6 _ 20.2 23,2
— — 62.8 (est.) 21.4 22.2 (est.)
14.3 9.7 24.6 16.9 12.8
18.2 17,7 — 11.9 12.4 (est.)
22,5 26.1 21.7 8.7 9.2 C3
— = 68.0 (est.) — — V25 or 26
_ 19.8 46.8 (est.) _ _ V20-21 (juv)
16.6* 16.6* 11.0 16.2 (est.) ?Cl1_— (juv)
- _— _ 21.4 (est.) 26.0 (est.) V22-26 (juv)
29.7 23.1 39.7 21.8 23.1 C16 (15-17?)
19.9 23.9 22.5 10.7 9.8 C4
30.5. 27.5 37.9* 20.6 18.8 + C15-C17
_ 33.4 _ 27.4 > 25.6 V26?
— _ 68.0 (est.) 24.8 V22 or 23
_ 26.0 (est.) -- _ — C2
24.2* 30.0 (est.) 22.0 (est.) = 11.2 (est.) C3
>20.2 29,1 25.9 12.6 11.7 C4
18.0 16.8 32.9 22.8 19.5 C6?
19.1 16.3 34.9 23.1 23.3 C7?
21.7 oo 36.4 = 22.5 C9?
_ _ _— _ — V21 or 22
_ _ — 23.3 25.7 V22 or 23
—eeeee_
TABLE 5. Measurements (in mm) of vertebral ribs of fossil emus (Dromaiinae) from Australia.
Seen eee
Width of Facets
>36.8
~34.3
> 28.0
SPECIMEN
NMV P157354
NMV P157358
NMV P157362
SAM P18107
SAM P18784
SAM P22812 (in part)
UCMP 60560
SE
29.3
34.1
38.6
34.8
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 109
TABLE 6. Measurements (in mm) of sterna of fossil emus (Dromaiinae) from Australia.
Width Width Length Length Width
Number of of first of last of of Sterno- of Cora-
Maximum Maximum _ Costal Costal Costal Costal coracoidal coidal Anterior
SPECIMEN Length Width Processes Process Process Margin Process Sulci Depth
D. novaehollandiae
NMV P157347 _ 124 3 118 — — — ~52,6 14.6
NMV P157355 — 112 4 109 90.9 49,7 16.4 — —
D. sp.
AMP 25218 147 118 4 127 109 64.2 —_ 70.5 17.9
Ow
TABLE 7. Measurements (in mm) of synsacra of fossil emus (Dromaiinae) from Australia.
ee ee eee ee ee
Diameter of Length of Depth of
Acetabular Width across Acetabular Acetabular
SPECIMEN Foramen Antitrochanter Complex Complex
D. novaehollandiae
NMV P157361 — >83.6 56.3 _
SAM Unreg. 15.5 _ —
SAM P16501 18.0 108 — —_
SAM P17767 12.2 104 60.7 33.8
UCMP 56333 17.4 — 62.3 36.8
TABLE 8. Measurements (in mm) of femora of fossil emus (Dromaiinae) from Australia.
ee
Proximal Depth of
Proximal Depth Diameter Distal External
SPECIMEN Length Width (Trochanter) of Head Width Condyle
D. novaehollandiae
HM B775/869 190 oe — — 59.6 61.2 R
HM B801/934 — — — — — 76.2 d,R
RHT 1064 190 65.1 — 27.6 — — R
SAM Unreg. 7 _— _ 27.4 >76.3 76.7
SAM P17104 _ ~67.2 ~61.2 ~28.0 _ — pL
SAM P22812 204 67.4 61.4 25.5 68.8 — L
AMNH 9677 _— — — — 83.3 70.7 dR
D. cf. ocypus
UCMP RASS176 190 59.5 — — — — pb
oe
110 C. PATTERSON & P. V, RICH
TABLE 9. Measurements (in mm) of tibiotarsi of fossil emus (Dromaiinae) from Australia.
Length, Depth, Depth, Width,
__Diameter of Shaft. proximal Proximal Fibular Internal External Distal
SPECIMEN Length Minimum Maximum Depth Width Crest Condyle Condyle End
D. novaehollandiae
AM 49713 — _— —_— 113 57.8 — a a — p,R
AM ‘B’ — — _— — a —_— 45.1 — 46.6 d,L
AM F10949 — — _— — = — >35.0 31,2 38.1 d,L
AM P5802 — — _ _ = 41.0 42.1 47.3 d,R
MM F16775 —_ — — _— — _ 44.9 44,3 49.5 d,R
NMV P44011 = 22.1 26.6 >76.0 >44.1 93.1 _ — — R
NMV P150013 — 24.9 32.7 — a 95 44.7 40.2 50.4 d,L
NMV P157356 —_— _— _— >92.2 47.4 _ — — p,R
NMV P157357 — — os — — _ 39.1* 39.1 43.3 d,L
NMV P157360 — — = — ~ — >39.3 39.3 41.8 d,L
NMV P157365 — 21.1 29.2 >91.5 49.2 96.4 _ _ — »p,R
QM F1652 — — oa — 56.1 >80.5 — a — p,L
SAM P17149 _— — _ — — — 42.7 — 43.6 d,R
SAM P18829 _ _ — — — > 38.8 41.3* >37.5 d,R
SIAM 61 384 23.3 32.3 102 56.9 88.6 45.8 43.9 51.8 L
UCMP 53825
(RHT6) —_— —_ _ _— — os 37.0 35.1 37.4 d,L
UCMP 53825
(RHT25) _ a a = — _— 37.0 34.9 35.6 d,L
UCMP 56845 -- = = — = _ — 38.8 40.6+ d,L
WAM 68.5.34 ve 21.6 26.3 — = 107 — _ _— L
D. patricus’”
MM F12074 o — — —_— a —_— 35.5 34,3* 41.3 d,R
QM F5547 - — — — 57.6 >90.2 — a — p,R
QM F5548 — ~ _— a — — 46.2 45.7 48.6 d,L
D. cf. ocypus
UCMP RASS5182 _ _ — _ — _ 43.0 40.8 46.0 d,R
D. gidju
SAM P26779 = = — — — _ 31.8 31.3 32.0 d,L
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 11
TABLE 10. Measurements (in mm) of tarsometatarsi of fossil emus (Dromaiinae) from Australia.
Depth, Depth,
_ Diameter of Shaft = Proximal Internal External Depth, Distal
SPECIMEN Length Minimum Maximum Width Cotyla Cotyla Hypotarsus Width
D. novaehollandiae
AM Unreg. _— — a 56.7 25.9 24.5 42.5 — p,R
AM “C’ _— — — 53.9 25.2 21.3 40.5* — pL
AM F771 — 11.8* 20.5 — — — — 47.5 d,R
AM F18935 — 14.0 19.4 — — — — >46.2 dL
(juv)
AM P58026 — _ 53.0 23.2* 19,1* 39.5* — p,R
NMV L5 — — — —_— a = — 54,17 d,L
NMV P44012 — 14.1 21.9 >42.3 = _— >34.0 52.4 R
NMV P44013 — 16.1 22,2 48.3 26.3 22.5! >35.1 55.9 R
NMV P44014 — 14.9 19.8 49.8 24.2 21.4 37.5? §2.2 R
NMV P44015 _ 14.0 19.5 46.5 24.0' 19.8 >37.0 — R
NMV P44016 — 15.5 20.4 — 23.0 >21.8 > 38.2 51.2 L
NMV P44017 _ 13.3 21.7 — — - >50.5 dL
NMV P44018 — 13.8 23.1 — — = -- 54.0° d,L
NMV P48392 _— 12.2 21.0 — — — — — dR
NMV P150015 _ 16.7 23.6 —_— — — — — d,R
NMV P157343 oo 13.0 19.3 >39.0 = a >32,4 — L
NMV P157344 _ 14.2 19.6 = — — — - d
QM F1143 (in part) = — 15.4 25.7 _ oe = — 56.1 L
SAM P13118 a 12.6 -- _ = — — — 4d,R
» SAM P17816 — 16.0 23.4 — _ — — 53.5 d,L
SAM P18693 _ 16.7 23.9 49.0 (est.) = a — 56,3 L
UCMP 53835 13.9 18.7* — _ — — 39.0 d,R
UCMP 56313 — 13,57 os — — — — 4d,R
WAM Unreg. 190 — — a 33.8 — — — — (juv)
WAM 68.5.34 >330 16.6 18.2 —- —_— _ os _ L
D. gracilipes’
QM F1142 _ 10.9 21.5 = - — _ — 4d,L
(juv)
‘Metapteryx bifrons’
QM F1135 _ — os — = _— 34.1 d,L
(juy)
D, ocypus
SAM P13444 330 — — 47.1 21.2 20.9 35.5+ 53.0 R
D. gidju
SAM P26779 — — = 35.0 18.9 14,.8* 26.9+ — pL
Dy sp.
AM ‘A’ aa 10.3 17.1 os — _ == d
(juv)
AM F10850 — — _ 46.4 ~ — = — pL
NMYV P35578 — 16,2 28.0 — — —_ _ — 4d,R
Aves, indet.
SAM P11552 at 21.2 28.0 _ _ — — 63.7 dL
um
112
C. PATTERSON & P. V. RICH
TABLE 11, Measurements (in mm) of the distal ends of tarsometatarsi of fossil emus (Dromaiinae)
from Australia.
—_----—————e—”:,—O0W0—_?ee—— —
Width, Width, Width, Depth, Depth,
re T3 T4 T2 T3
D. novaehollandiae
AM F771 9.1 23.8 12.2 12.1 20.3
AM F18935 a 24.1 12.3 os 24.1
QM F1135 5.9 15.7 45. 8.8 13.8
QM F1143 (in part) 10.9 27.2 14.3 15.3 25.0
QM F1143 (in part) _— 23.0 a _ 22.6
NMV L5 > 10.5 24.8+ 13.2+ > 14.0 25.2+
NMV P44012 > 9.2 24.3 12.8+ 13.1+ 22.3*
NMV P44013 9,8+ 24.3 14.5+ 15.1 23.8
NMV P44014 10.9 26.4 13.2 14.8 22.5
NMV P44015 _ — = _— 21.47
NMV P44016 10.3 25.9 13.3 13.8 21.4
NMV P44017 > 8.8 >23.3 >13.1 14.8 23.3
NMV P44018 > 9.0 25.0+ 14.8 13.4+ 20.4+
NMV P48392 — —- >10.5 >21.5
NMV P150014 1133 28.3 — 15.0' 24.0
NMV P150015 14.9% 25.1 _— 11.6 29.8
SAM P13118 _ _— 11.9 — 20.2 +
SAM P17816 10.0 27.8 >13.3 15.0 23.1
SAM P18693 12.6 29.8 15.6 >15.3 25.6
UCMP 53835 — > 18.4 — _
UCMP 56313 10.7 27.9 — 15.3 23.1
D. §racilipes’
QM F1142 — 16.6 = — 16.5
D. ocypus
SAM P 13444 10.6 27.8 13.5 14.7 21.9
D. sp.
AM ‘A’ —_ 16.5 _ —_ -14.7
QM QA205 _ ~16.1 _— — >14.4
QM QA416 13.3* — _ 23.9* _
QM QAS505 8.5 17.5 _— 11.9 16.8
UCMP RAS5397
(in part) — 15.7+ — _— 14.0*
Aves indet.
SAM P11552 — >26.3 > 14.5 — >27.5
* perhaps T4
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 113
TABLE 12. Measurements (in mm) of phalanges of fossil emus (Dromiinae) from Australia.
Proximal Proximal
SPECIMEN Phalanx Length Depth Width Element
D. novaehollandiae
SAM P18059 P1,DII 45.0 20.0 14.8 L
SAM P18248 P2,DII 37.6 17.8 23.8 R
SAM P18249 P1,DIV 38.2 17.6 18.8 R
UCMP 53832 P4,DII or DIV 20.6 10.6 10,5
UCMP 53832 P1,DII a 16.7 13.4 p
UCMP 53832 P2,DIV 16.3 12.3 14.1 L
UCMP 53833 P1,DIV 34.2 16.1 17.8 L
UCMP 53833 P5,DIV? 20.0* 9.6 8.6+
UCMP 55983 P1,DIII 53.2 20.8 25.7 L
UCMP 55983 P2,DIII 36.2 13.9 21.3 L
UCMP 56849 P1,DIU 43.0 18.9 17.0 L
UCMP 94679 P1,DI1 48.4 18.9 24.0
UCMP 94680 P2,DIII 37.0 18.8 21.4 L
D. gidju
SAM P26779 P1,DII 33.1 14.2 12.9 L
SAM P26779 P2,DII 23.5 11.8 11.6 ie
SAM P26779 P3,DII 19.3 10.1 8.9 L
SAM P26779 P1,DII1 45.1 16.7 19.5 L
SAM P26779 P2,DIII 31.7 12.9 15.2 L
SAM P26779 P3,DII 22.5 10.5 13.0 L
SAM P26779 P4,DIII 18.4 9.8 10.5 L
SAM P26779 P1,DIV 29.2 13.0 14.0 L
SAM P26779 P2,DIV 16.1 10.6 10.8 L
SAM P26779 P3,DIV 10.8 9.2 9.4 L
SAM P26779 P4,DIV 8.8 7.9 8.2 L
SAM P26779 P5,DIV 15.6 8.8 7.7 L
D. sp.
QM QA504 P1,DIII 41.5 15.3 16.8 R
UCMP RAS 5397 P2,DII 19.8 11.9 10.9 L
TABLE 13. Measurements (in mm) of the fibula of fossil emus (Dromaiinae) from Australia.
SPECIMEN Proximal Width Proximal Depth Element
NMV P157363 17.0 36.4 p,L
114 C, PATTERSON & P. V. RICH
“a tem
FIGURE 1. Dromaius gidju, n. sp. Type from the Wipajiri Fm. Leaf Locality, Lake Ngapakaldi,
Kutjamarpu fauna, Miocene. A,B, stereo pair of pes, dorsal view. C,D,E,F, tarsometatarsi in
posterior (C), anterior (D) and proximal (E,F, stereo pair) views. G,H,I,J,K, distal left tibiotarsus
in anterior (G), posterior (H), distal (I), internal (J), and external (K) views. Scale indicates 1 cm.
THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMALINAE) 1s
TT
FIGURE 2, Mid-Cainozoic emu fossils from northern South Australia and living casuariids.
A-D, posterior views of tarsometatarsi of (A,B) the extant Dromaius novaehollandiae, (C)
D. ocypus (SAM P13444), and (D) the extant Caswarius unappendiculatus (from Miller, 1963).
E,F, stereo pair in anterior view, tarsometatarsus, the type specimen of Dromaius ocypus, SAM
P13444, Pliocene, Lawson-Daily Quarry, Mampuwordu Sands, Lake Palankarinna, Palankarinna
fauna, G, distal tibiotarsus in anterior view of D. cf. ocypus, RAS 5182, Pliocene. H,!, femur
of D. ef, ocypus, in posterior (H) and anterior (I) views, RAS 5176, Pliocene,
116 C. PATTERSON & P. V. RICH
Tem
FIGURE 3. A variety of late Cainozoic emu fossils. A, sternum of Dromaius sp. in dorsal view,
AM F25218. B, pelvic fragment of D. novaehollandiae in lateral view, SAM P16501. C, distal
left femur in posterior view of D. novaehollandiae, HM B801/B934. D,E, proximal right
tibiotarsus in lateral (D) and proximal (E) views. F, left tibiotarsus in anterior view, SIAM 61.
7
tem
tem
FIGURE 4. A variety of late Cainozoic emu fossils. A, distal right tibiotarsus in anterior view, ane of the original
type specimens of Dromuius putricus'De Vis, QM F5548. B, QM F120 considered by CW. De Vis to be a coracoid
af D. patricus, but non-avian. C, distal left tarsometatarsus of ‘Metapteryx bifrons' De Vis in anterior view, QM
F1135, originally designated a kiwi but actually a juvenile D. novaehollandiae. D, distal tarsometatarsus of D. gracilipes’
De Vis in anterior view, QM F1142, a juvenile D. novaehollandiae. E, distal left tarsometatarsus of Aves indet. ef.
dromornithid in-anterior view, SAM P1552, assigned previously as emu. F,H, tarsometatarsus fragment of a small
D. novaehollandiae in anterior (F) and posterior (H) views, AM ‘A’, no locality or age data available, G,I,J, second
and third trochleae of a small Dromuwius in posterior (G), distal (1), and posterior (J) views, QW QA505. K-N, phalanges
of D. novaehollandiae in dorsal views. (K) phalanx 1 digit Lf, UCMP 94679; (L) phalanx 2, digit Ll, UCMP 94680;
(M) phalanx 1, digit Il, UCMP 56849; and (N) ungual phalanx 4, digit 111, UCMP 60563.
ECHIURANS FROM AUSTRALIA (ECHIURA)
BY S. J. EDMONDS
Summary
Seventeen species of euchiurans [i.e. echiurans] are recorded from Australia of which nine are
redescribed, and none are new. One, Anelassorhychus porcellus adelaidensis, is given new status.
Listriolobus bulbocaudatus Edmonds, 1963 is now considered a junior synonym of L. brevirostris
Chen & Chen Chang, 1958, Ochetostoma myersae Edmonds, 1963 a junior synonym of O. baroni
Greeff, 1879 and Austrobonellia mjoebergi (Fischer, 1921) a junior synonym of Pseudobonellia
biuterina Johnston & Tiegs, 1919. A key to the Australian species is given.
ECHIURANS FROM AUSTRALIA (ECHIURA)
S, J. EDMONDS
EDMONDS, S. J. 1987. Echiurans from Australia (Eehinra). Reo. S. Aust. Afus, 52(2) 196138.
Seventeen species of euchiurans are recorded from Australia of which ning are redescribed, and
none are new. One, Anelassorhynehus porcellus adelaidensis, is given new status. Lisiriolohus
bulbocaudatus Edmonds, 1963 is now considered a junior synonym of L. brevirostris Chen &
Chen Chang, 1958, Qchelostorne myersue Edmonds, 1963 a junior synonym of O. baron Greeff,
1879 and Austrobonellia mjoebergi (Fischer, 1921) a junior synonym af Psexdoboneliia hiuterina
Johnston & Tiegs, 1919. A key to the Australian species is given.
5. J. Edmonds, Honorary Associate, South Australian Museum, North Terrace, Adelaide, South
Australia 5000, Manuseript reeetved 2) October 1986,
Echiurans are soft-bodied, protostomous,
coclomate, worm- to sausage-shaped, marine Inver-
lebrafes thal resenible annelids and sipunculans.
Because they are largely subtidal and occur in
burrows and protected places echiurans are not
always easy to find. They have, however, been
collected in tropical, temperate and polar seas and
are known from the littoral to the ultra-abyssal
resions of the oceans. More than 150 species have
heen described.
Records of Australian echiurans are few and
scattered (Haswell 1885, Whitelegge 1899, Augener
1903, Hedley 1906, Dakin 1916 and 1952, Fischer
1919 and 1921, Johnston & Tiegs 1919 and 1920,
Monro 1931, Edmonds 1960, 1963, 1966 and 1982,
Nielsen 1963, Dartnall 1970 and 1976) and only 16
species have so far been reported, of which three
are well known, In Australia echiurans have been
found in burrows in mud and sand, in limestone
rocks and in coral, in tangled roots of sea-grasses,
under stones and in cracks and fissures in rocks,
Some have been dredged. The best known
Australian echiurans (and ones that can be readily
collected) are Merabonellia haswelli, Anelassorhyn-
chus parcellus adelaidensis and Ochetostoma
australiense. Scuba divers report the presence of
Jarge numbers of the first in shallow water at
Encounter Bay, S,A,, between Wright J]. and the
Bluff and from the islands comprising the Banks
Group in Spencer Gulf, S.A, and of the second
species from Coobowie and Bdithburg in St Vincent
Gulf, S.A. Large numbers of O. austra/iense occur
at Caloundra and Dunwich, Qu., the species feeding
from the surface of intertidal mud flats at low water,
In other parts of the world they have alsa been
found in the empty shells of molluscs and in sand
dollars, Most of the specimens examined in the
present study were found intertidally by collectors
or subtidally by divers and are now in the collections
of Slave Museums,
The classification used in this paper ts that
outlined by Stephen & Edmonds (1972), which in
tum is based on that of Fisher (1946).
LIST OF AUSTRALIAN ECHIURANS
The species marked ‘*' have not been examined
by the author. Records of species marked “?" are
considered doubtful.
Family Bonelliidae
Metabonellia haswelli (Johnston & Tiegs)
Pseudobonellia biuterina Johnston & Tiegs
7* Archibonellia michaelseni Fischer
+ Zenkevitchiola brevirosiris Murina
* Sluilerina alba Murina
* Vitjazema ultraabyssalis Murina
* Protohonellia papillasa Murina
Family Echiuridae
Anelassorhynchus porcellus porcellus Fisher
Anelassorhynchus partellus adelaidensis
Fdmonds 7, statis.
”* Anelassorhynchus vegrandis (Lampert)
Arhynchite hiscocki Edinonds
Listrialobus hrevirosiris Chen & Yeh
Chen-Chang
7* Listriolabus sorbillans (Lampert)
Ochetostoma baroni (Greeff)
Ovheiostoma australiense Edmonds
Thalassema sydniense Edmonds
Family Ikedaidac
Ikeda sp.
The following abbreviations are used in tis
paper} AMS (Australian Museum, Sydney), MV
(Museum of Victoria, Melbourne), SAM (South
Australian Museum, Adelaide), WAM (Western
Australian Museum, Perth), TMH (Tasmanian
Museum, Hobart), N.SW, (New South Wales), Qu.
(Queensland), S.A. (South Australia), Tus,
(Tasmania), Vie. (Victoria), WA. (Western
Australia,), The anatomy of a ‘generalised’ echiuran
is shown in Fig. 1.
120 8. J. EDMONDS
Key To GENERA OF AUSTRALIAN ECHIURANS
The doubtful genus Archibonellia has not been included
in the key; it would key near Pseudobonellia,
I Erpbeyscis fifty seth o ei ic eee pes | 2
Proboscis not bifid. .-.--,..............02.. 3
2. One nephridium, with distally placed, stalked
nephrostome..,............4- Metabonellia
Two nephridia with a small sac carrying a male
between them, ,....,.....,. Pseudobonellia
bands ..... J ovur: Eile nga e 5.9 MONTE edbs-t 4
Longitudinal musculature of body wall not grouped
Hato Mandy): ppt ok. stay hte nate hh wens 6
4. Nephridia less than 10 and arranged in pairs; trunk
Jong pr SHOP 5 ws: ASC L9b Dee. 300k ee 5
Nephridia very numerous and not in pairs; trunk very
Jong, up to 400 mm... ..... 0.2. eee Ikeda
5. Intervals between longitudinal muscle bands traversed
by small bundles (fascicles) of oblique muscles
Bhd § Seay. 5 5 SRC hae. 38 Sb oo Ochetostoma
Intervals between longitudinal muscle bands not
traversed by small bundles of oblique muscles
+ Annet tiveeeerereesaeeess Listriolobug
6. Nephrostome distal ................. ‘Vitjazema
Nephrostome basal ........0....0000e00s cease 7
7, Ventral setae none........ 2... cece cee eee 8
Ventral setae tWo os eid ee eee eu eesewaene 9
8. Anal glands long and slender; posterior region of
proboscis surrounded by a collar Zenkevitchiola
Anal glands bushy or feathery; posterior region of
proboscis modified to form a cup... Sluiterina
9, Nephrostomal lips long and spirally coiled...,...
ann! $0-seaee eevee eees es Anelassorhynchus
Nephrostomal lips long or short but not spirally coiled
Le ee oon oe ring t ae nibiitdeteebictis tenet 10
10. Anal glands tubular or sac-like.,.....,...... il
Anal glands feathery or bushy .... Protobonellia
11. Proboscis with expanded or fan-like anterior extremity
Pticleite ati p tinge eters gpg bt we teen 8 Arhynchile
Proboscis without expanded or fan-like anterior
CREPES sey ate Mey celI ae om Thalassema
FIGURE 1. A generalized diagram to show some of the
internal anatomy of an echiuran; dorsal view. Most of
the much coiled intestine has been omitted. av, anal DESCRIPTION OF GENERA AND SPECIES
vesicle; c, caecum; cf, ciliated funnel; cg, ciliated groove:
cl, cloaca; dy, dorsal vessel; 2, gonad; i, intestine; in, Genus Metabonellia Stephen & Edmonds
interbasal muscle; Ib, lateral vessel; m, median vessel; n,
nephridia; ne, nerve cord; nl, nephrostomal lips; nv, _Afefahonellia Stephen & Edmonds, 1972, p. 394,
neurointestinal vessel; 0, oesophagus; p, pharynx; rv,
ring vessel; s, siphon; vy, ventral vessel. Type-species: Bonellia haswelli Johnston & Tiegs,
1920, (Bonellia gigas Nielsen, 1963, which was
named as type by Stephen & Edmonds (1972) is now
AUSTRALIAN ECHIURANS 11
considered to be 4 junior synonym of B. haswelli).
The genus contains only one known species.
Diagnasis
Female of medium to large size, with proboseis
bifid, grooved and ciliated on ventral surface. Pale
to dark green. Two vevitral setae. Single nephridium
with nephrostame on a short stalk placed about two
chirds of way along vephridium towarils its distal
extremity. Anal vesicles branching. Intestinal siphon
present, Male worm-like, as long as 20 mim and
without setae, Pound in nephridium of female,
Motaboneiia haswelli (lohnston & Tiegs)
(Figs 2-4, 18)
Bonelfig fleswelli Johnston & Tiegs, 1930, pp.
73-76; Ednionds, 1940, pp. 95-96.
Bonellia gigas Nielsen, 1963, pp. 41-67,
Bonellia rasmanicu Dartnall, 1970, pp. 69-74.
Metabonellia haswelli Stephen & Edmonds, L972,
pp. 394-395; Darthall 1976, pp. 1041-1043;
Edmonds 1982, pp, 314-316,
Halowpe: AMS Gll22 and paratype O)261. Type
locality, Sydney Harbour, NSW, ‘under stones just
above the limit of law water’.
Previous Australiun reeards. Jolnston & Tiegs
(1926), Edmonds (1960), Niclsen (1963), Durtnall
(1976).
Description of femule
trunk: Medium to liurge, shape vartable (sausage
{a sub-ovoidal), light to dark green, length
20-80mm (Nielsen 1963; length 80-120mm,
maaimum width 40mm), Skin smooth to rough,
usually covered anteriorly und posteriorly with near
circular rows of rather Flattened papillac. Thickness
of body wall variable, longitudinal jusculature
CoOmLMUOUS,
Proboseis: Firmly attached, long, bitid and
cupable of great extension, maximum length in fixed
specimens 260 oum- Anns shore and usually of
abour equal length, 20-30 mm. Lateral edges tend
to roll inwards, veritral surface ciliate, dorsal surface
smooth,
Setoe Two, together with reserve setae, ventrally
placed and posterior to mouth. Setal but no
interbasal muscle present
Alimentary Cunal: Mouth at base ef preboscis,
canal much coded and fastened to body wall by
numerous strands of muscle. Consists of (1) foregut
(divided into pharynx, oesophagus, crop and
gizzard), (2) midgut qwider and longer than
foregut), associated for most of its length with a
coll#leral litestine or siphon and (3) hindeut and
gloaca, Siphon jubular, of smaller diameter than
lutestine and arising near beginning of midgut. A
ciliated groove hes in wall of much of posterior
intestine. No precloacal caecum, Faecal matler
forms pellets. Anus at posterior extremity of trunk,
Nephridium: Single, attached to coelomic wall of
(rink just posterior to level of setae, Size variable,
depending on reproductive condition of animal.
Wall of anterior and posterior regions usually
thicker than thin, often transparent, middle region,
which seems capable of much extension and even
sacculation, Nephrostonie distal, situated about
three-quarters to two-thirds length of nephridium
away from sephridiopore. Nephrostomal lips frilled
or cremated, situated at end of short stalk-
Nephridium holds eggs and/or a male (two males
in one specimen). Eggs develop in coelom along a
mesentery assoviated with ventral blood vessel and
nerve cord. Diameter of largest eggs 0.50-0,55 mm,
Vascular System: Thin walled ventral blood
yessel, Single neurointestinal vessel, arising
posteriorly Tram two short arms on each side of
anteriormos! region of intestinal siphon, joins
ventral vessel in anterior half of trunk. Thin walled
dorsal vessel] fuses with intestine near posterior
exlremily of foregul. Anterior continuations of
ventral and dorsal vessels extend into probasels.
Neuroimestinal vessel often vesiculated or
superficially roughened and coloured dark yellow
to orange.
Anal Vesicles; Two, much branched, tufted and
each a complex of tubes and tubules, Johnston &
Tiegs (1920, p. 75) described them thus: ‘Into the
cloaca open fwo anal vesicles ... Into each open
about 18 tubules, some quite short, others much
longer. These tubes give aff smaller or larger
numbers, al times very large numbers, of secondary
nephridial [? excretory] rubes, cuch ending in a
narrow heck Which bears a circular disc with the
nephrostome opening in iis cenlre. The dise is
composed of a ring of compressed elongated cells,
with strongly staining nuclei and fringed with a ring
of cilia’, In larger specimens the vesicles are larger
and the branches more tufted.
Descriplion of male (based on four stalned and
mounted specimens)
Located In wephridia of females but not
permanently aitached like male of Pseudobanellia
diuierina, Long, thin or flat but swollen or rounded
anteriorly, lendifiy to taper posteriorly; largest about
20 mm long, maximum ‘width 1,2, lacking setae,
Body ‘wall very thin, Outline of whal might be a
rudimentary gut tuns through most of animal; body
cavity contains developing sperm morulae. Cilia on
some regions of body surface,
No males were found by Johnston & Tiegs (1920).
The male af Bonellia gigas Nielsen, 1963 is 19 mun
long and 0,5-(,0mm wide, lacking setae bur
122 5, J. EDMONDS
possessing a posterior sucker or clasper, The male
of Bonellia tasmanica Dartnall, 1970 is 7 nim long,
0.8 mm wide, tapering posteriorly and lacking both
setae and clasper.
Systematics
Bonellia gigas Nielsen, 1963, described from a
very large bonelliid collected at Western Port, Vie.,
was considered to be different from & fuswelli
chiefly because it lacked an intestinal siphon,
Although I have not been able to dissect the type
of B gigas, [ have dissected one specimen trom Port
Phillip Bay and another from Flinders. Both have
siphons. Further, Fig. 2 of Nielsen, 1963, clearly
shows a siphon attached to part of the pur,
especially the posterlor part. Nielsen apparently
mistook the siphon for a vontinuation of the
neurointestinal vessel and labelled it ‘intestinal
blood vessel’
Dartnall (1970) described B tasmanica trom
northern Tasmania, atguing that it differed from
& haswelli because it lacked a siphon and from beth
B& haswelliand & gigas because its nephridia were
sacculated. It is probable that what Dartnall
described as ‘an intestinal vessel, Which runs closely
along the wall of the intestine for about the
posterior two thirds of its length’ and then ‘leaves
the gut and joins the ventral vessel’ is in part a
siphon and in part a neurointestinal vessel. The
sacculated condition of the nephridium of &
tasmanica is'a doubtful character and was probably
caused by temporary nvuseular contractions of the
organ at the time of fixation, [n one specimen trom
S.A, the thin walled part of the nephridium is
constricted near its middle so-as to form twa sacs.
From a study of male and female specimens of
RB haswelli, B. gigas and & fasmanica it is concluded
that the three species are synonymous, the first
name having priority, Stephen & Edmonds (1972)
transferred the species to a new genus Metetionellia
on account of the distal position of its nephrostome.
Johnston & Tiegs’ specimens from N.SW, were
small but Nielsen’s from Victoria were very large.
Specimens examined and localities
New South Wales; Fairlight (near Manly) (1)
AMS W56i2 and (3) AMS W4702 (two of these
clearly show (the relationship between the
neurointestinal vessel and the siphon); Camp Cove,
Sydney Harbour (1) AMS W8703; locality unknown
(1) SAM E1404.
Victoria: Port Phillip Bay near Porr Arlington
(2) MV coll. and at Mornington (1) MV coll;
Flinders, Western Port Bay, SAM EL400 (1) and type
male and female of B& gigas MV G2696 and 2697;
Port Phillip Survey area 31 ‘inside buoy’ (1) SAM
E1407.
Tasmania; Jacobs Boat Harbour (north
Tasmania), paratype of B /asmanica, TM K226,
South Australia; Spencer Gulf — Boston f. (near
Port Lincoln) (2) SAM EI401; at following islands
in Banks. Group, Marum tl. (3) SAM E1475, (3)
SAM E1492, Lusby 1, 0) SAM E1488, Langton 1,
(1) SAM E1504, Hareby 1. (1) SAM E508; Se
Vincent Gulf — Edithburg jetty (near base af outer
piles) (2) SAM E1403, (1) SAM El4§7, (3) SAM
1458 (3) SAM Bi502; Marino Rocks (3) SAM
E1406; Aldinga Reef (i) SAM E1402; Victor
Harbour - near Rosetla Head (3) SAM B145], (3)
SAM E1483, (3) SAM E1466, (5) SAM Et45], (3)
SAM El4%) and near Whalers Wharf (3) SAM
E1474; St Francis t. (Nuyrs Archipelago) (4) SAM
1469,
Western Australia: Mistaken 1. (King George
Sound) (1) WAM 37-85; Garden I, (1) WAM 10-73;
off Carnac 1. (1) WAM 71-75; off Dansborough (LU
WAM %73; Houtman Abrothos (Ac N. end of
Morley |, Easter Group) (1) WAM 279-85,
Distrifurian
Known trem south-western, southern and south-
eastern Austratia, from the Abrolhos J. in Western
Australia to Sydney Harbour in New South Wales.
Usually collected subtidally, occasionally
intertidally. No records atfrer than from Australia.
Habitat
‘In South Australia individuals of this species live
in crevices between rocks and Under rocks in
Sheltered, calmer water where there isa deposition
of fine, muddy silrs, such as occurs on the lee side
of the Blu/f at Victor Harbour and at Edithburg
jetty on Yorke Peninsula. The gteatest concentration
appears to he at the perimeter of the rubble-reef
‘area, especially where the latter abuts open flat
where sea-prasses grow, The proboscis extends only
at night and then over the bottom adjacent to the
burrow for a radius of about one nietre' (NW. Holmes
pers. comm).
Genus Pseudobonellia Johnston & Tiege
Pseudehonellia Johnston & Tiegs, 1919, pp.
213-230; Stephen & Edmonds, 1972, p. 401; Datta
Gupta (976 p. 115,
Type-species: Psevdobonellia buiterina Jobnsion &
Tiegs
Diugnasis
Female with bifid proboscis Trunk with twa
ventral setae, Two nephridia (gonoducts) with
distally placed mephrostomes; nephrostomal lips
crcnated. Anal vesicles branching, Male carried in
small blind tube thal projects into coclam between
nephridiopores, Type species: Pseudobonellia
AUSTRALIAN ECHIURANS
FIGURES 2-4, Metahonellia haswelli, 2, anterior vegion dissected. 3, seta (scale line = 0.3 mm),
4, portion of anal vesiele,
hiuterina Johnston & Tiegs, 1919, The genus
contains only one species.
Pseudobonellia biuterina Johnston & Tiegs
(Figs 5-8, 19)
Pseudobonellia hiuterina Johnston & Tiegs, 1919,
pp, 213-230, pls 9-11; Monro 193], p, 33; Fisher
1948a, p. 856; Edmonds 1960, pp. 96-97, fig. 5;
Stephen & Edmonds 1972, p, 401.
Archibonellia mjoebergt Fischer, 1921, pp, 6-8;
Austrobonellia mjoebergi Fisher, 1948a, 856;
Edmonds 1960, p. 97.
Holotype: AMS G477; type locality, North West
[slet (Capricorn Group), Qu.
Description of female
Trunk: Small, maximum length 23 mm (mostly
6-14), maximum Width 3-6, pyriform, sub-ovoidal
to globular, pale io dark green. Body wall thin
(sometimes transparent) except in anterior and
posterior region, usually thinnest on dorsal side.
Surface usually wrinkled by large numbers of
closely set papillae. Nephridiopores and opening of
male tube on anterio-ventral surface often very
noticeable,
Proboscis: Long, bifid, adherent and capable of
much extension; in fixed specimens 1-10 times as
long as trunk. Arms shorter (3-10 mim), normally
about equal length. Ventral surface ciliated, lateral
edges tending to curve inwards. Mouth at base of
proboscis,
Setae: Two (in addition smaller reserve setae
usually present), golden brown, slightly iridescent,
with free end forming a weak hook. Johnston and
Tiegs state that larger hooks are 2-3 mm long and
smaller 0,7-0.8 mm and that a strong muscle pad
joins their internal ends ‘evidently serving to impart
fo them a lateral pincer-like movement’, The
description aptly fits the structure and function of
an interbasal muscle. Although there are well
developed setal muscles in the present specimens,
none clearly show the presence of an interbasal
muscle,
Nephridia (gonoducts or uteri): One pair, size
variable, usually prominent, tapering distally, each
with a slightly frilled, distally placed nephrostome
borne on a short stalk. Largest eggs in nephridium
0.25-0.35 mm (Johnston & Tiegs 1919 give 0.[1
mm). Nephridiopores on each side of nerve cord
in anterior trunk region,
Male tube (androecium): Small, 1,5-3.0 mm long,
opening externally near nerve cord at about level
of nephridiopores. Opening often very noticeable.
Tube encloses small, wormlike male attached to
distal end of tabe.
Alimentary Canal: Foregut short; midgut long,
thin walled and associated for much of its length
with an intestinal siphon; hindgut short. No
precloacal caecum. Faeces form sub-ellipsoidal
pellets.
124 5 1. EDMONDS
Vascular System; Thin walled ventral vessel runs
alongside of nerve cord, Thin walled dorsal bioad
vessel (fastened to foregut and body wall by
mesenteries) fuses with fut near junction of fore-
and mid-gut, Single nevrointestinal vessel, arising
from wall of midgut just posterior to anteriamast
extremity of siphon, joins ventral vessel in anterior
half of trunk. lotestinal extremity of neurointesunal
vessel usally arises from two short roots which lie
on each side of the siphon and in close contact with
it, Neurolntestinal yessel sometimes vesiculated,
Anal Fesicles; Fwo branching, tuft-like masses of
fine tubes, arising on each side of posieriormost
section of hindgut, In this respect they differ from
the type description ia which ‘each anal tree consists
af masses af very delicate, simple, cylindrical tubes
Opening separately into the rectum’. Edmonds
(1960, p, 96) remarked that ‘the anal vesicles do not
seem to communicate With the cloaca as simply as
described by Faohnston and Tiegs’and (hat the tubes
branch to some extent. The anal vesicles of dissected
specimens in the present collection fram
Queensland and Western Australia are branched.
In one of the paratypes (AMS 6477), the vesicles
arise fram about! 12 shar! tubes which branch and
sometimes rebranch into simple cylindrieal rubules
that open to the coelom through slightly dilated
funnels fringed with cilia,
Ovaries: Jotinston & Tiegs (1919) slate thal the
ovaries arise from the peritoneum lining the
muscular strands (hat hold the posterior portion
of the rectum in position and that they |le
transversally on the frenulae. In the present
specimens developing egys lie more longitudinally
in the posterior third or half of the body cavity in
close association with the ventral vessel,
Description af male
Accarding ta Johnston & Tiegs (1919) the male
lives permanently in the androecium of Ihe female,
the two being fused distally. Without definite mouth
or anus, although a rudimentary canal ls presen,
No setae. Two seminal vesicles present. How the
male performs its sexual function is not known.
Johnston & Ticgs (1919) suggest that ‘the sperms
may be liberated into the cavity of the androecium
whence they reach the exterior through its canal and
enter the adjacent openings. ft 18 possible, however,
that the male may be protruded through the canal
of the androecium and actually liberate eperms in
the female apercure’.
Systemulics
Although the specimens examined 17 the present
collection from North West |. and Heron 1, differ
in a few respects from the lype description, they are
considered to be ? biuierina.
No satisfactory character has been found tat
distinguishes any of the Western Australian
bonelhids from Aiaterine. Those collected from
the CSIRO laboratory a} Waterman Bay (from
under pots standing on sand in an indoor aquarkum
through which sea Water was conlmually passed)
Were pale green while Lhose collected at Heron 1
were dark green. How important colour di flerences
are is noc known, Agius & Jaccarini (1981) have
shown that the unpigmented trochophores of
Bonellia viridis when kept under constant
Wlumination develop into unpigmenied adults,
Whether the depth of colour of diverting depends
on the amount of light received is Unknown.
Twe other bonelllids closely related to P biuterina
haye been described from W.A., Archiborellia
michaelsen Fischer, 1919 and Austrehanellia
myoebergi (Fischer, 1921), the former from Rottnest
1, the latter fram Broome, both localities being
places where # bivéerina has been found. Bath
Fischer's species were described from single
speciinens.
Anstrobonellia mineberg) Fischer, (921, is now
being placed ip the synonymy of FB biuterina
Johnsten & Tiegs, (919 Fischer described his
specimen thus; trunk oval, 45 mm lung, light prey
and |ransparent; proboscis short, 18 mim long, with
two arms of unequal length, Setae two with
recurved tip, no interbasal muscle. Nephricia tava,
thin walled (containing eggs) with distally plaved
néphrostames and a median, unpsired, smailler,
thick-walled ‘Segmentalorgan’ or ‘Uterns’, with a
basully placed nephrostome, Intestine lone and
convoluted, Anal vesicle dise-like and expanded
With J2e15 deniritic main stems the branches af
which possess lateral funnels.
Lf Fischor's Segmentalorgan’ is a male tube and
if injury accounts for the inequality in the length
of the probosels arms, then the description of A_
thioebergi fits very well that of P divrerine.
Fischer's other specimen from W.A., Austrabonellia
michaelseni, differs iy other characters and is
Possibly a separate species,
Specimens exvarined and locglities (these are
additional to those recorded in Edmonds, 1960)
Queensland; Heron 1, (1) AMS W3719; North
West F (3) AMS WISO7, (l) AMS W691;
Whilsunday Group (2) AMS W3029: One Tree 1.
(1) AMS W9275
Westen Australia! Barrow f, (2) WAM 139-8);
Riddell Pt, Broome (3) WAM 50-85; Roebuck Bay,
Broome (2) WAM 47-835; Abrainos I, (1) WAM
102-79; Yanchep Reef (2) WAM 98-79; Waterman
Bay (CSIRO Manne Laboratory) 110) SAM
E1439-1441; Garden I, (1) AMS W3720; Albany (S.
side of Princess Royal Harbour) (2) WAM 147-81;
Lookour Pt, Cheyne Bay (1) SAM E1482.
Distribuwlion
Queensland: Phe Crear Rariice Reet, from
AUSTRALIAN ECHIURANS 125
Capricorn Group in the south to Low I. in the
north.
Western Australia: From Barrow LI. in the north
to Cheyne Bay in the south. Whether the species
extends from Queensland to Western Australia
through Torres St is not known. It is not known
from S.A., Vic., N.S.W. or Tas.
Other record: New Caledonia (Stephen 1976).
Habitat
Specimens have been collected intertidally in
coral and limestone reefs; also from under stones
and objects resting on sand.
Genus Archibonellia Fischer
Archibonellia Fischer, 1919, p. 283; Fisher 1984a,
p. 856.
Type-species: Archibonellia michaelseni Fischer,
1919.
Diagnosis
Female with bifid proboscis. Trunk with two
ventral setae. Three nephridia, two being paired,
lateral and very small and the third median, large
and unpaired. Position of nephrostome not known.
Intestine very short with globular caecum. Male not
described.
i
FIGURES 5-8, Pseudobonellia biuterina, 5, anterior region, digestive and vascular systems; 6,
nephridia (gonoducts) and male tube; 7, small portion of anal vesicle; 8, setae from different
specimens (scale line = 0.5 mm).
126 8. J EDMONDS
?* Archibonellia michaelseni Fischer
Archibonellia michaelseni Fischer, 1919, pp,
283-285; 1926, p, 207; Fisher 1948a, p, 856;
Edmonds L960, p. 97.
Type-specimen; Not Known, type locality: west coast
of Rottnest J,, near Fremantle, W,A,
Description of female (after Fischer 1919) (Male not
kriown)
Trunk: About 12 mm long, grey in life, Proboscis
anteriorly forms two lappets. Setae two, with reserve
pair and an interbasal muscle. Two small, paired
nephridia (‘Segmentalorgane’) placed below a large
unpaired ‘Uterus’, Intestine very short, consisting
of a single loop in anterior half of trunk and a bew-
shaped tubular part in posterior half. Globular
caecum present, Ovary lies along posterior region
of nerve cord, Anal vesicles arise as single stems
an either side of rectum and branch at tip. Position
of nephrostome not known.
Systematics
A, michaelseni resembles FP bititerina Johnston
& Tiegs in many respects, especially in the
possession of an unpaired, median ‘Uterus’ lying
between paired lateral nephridia, Moreover, Fischer
(1919) thought that he saw a miale in the median
structure, In PR bivterina, however, (1) the lateral
nephiridia are much larger than the medial tube; (2)
the alimentary canal is very long and convoluted
and not very short (incredibly short) as shown in
Fischer 1919, fig. 6; and (3) neither caecum nor
interbasal muscle is present, While the two species
may be synonymous the described differences
between them are considerable and until the type
of more specimens became available it ig probably
best to consider them as dilferent.
Genus Protobonellia [keda
Protobonellia Ikeda, 1908, p, 259; Fisher 1948a, p,
854; Datla Gupta 1976, p.1L5.
Type-species; Profohonellia mitsukurli Ikeda, 1908,
Didgnosis
Proboscis of female long, tubular, non-bifid,
Ventral setae two, One nephridium. Nephrostome
stalked, fimbriated, basal, Anal vesicles tong,
dentrine. Male not known.
* Protobonellia papillosa Murina
Protobonellia papillase Murina 1978, pp. 112-113,
fig, 4.
Description (atter Murina, 1978)
Trunk 28 mm long, 16 mm wide, bearing rounded
papillae 0,25-1,25 mm in diameter, densest
anteriorly and posteriorly, Proboscis light grey,
width 5.5 mm, distally blunted; basal part (near
mouth) has form of oval collar with thick, wavy,
pigmented borders and two long processes laterally.
Setae two, golden-yellow, bent or curved, 0.5 mm
long, 0,15 wide, Nephridium single, rounded, lying
oo right side of nerve cord; nephrostome short,
basal. Anal vesicles form dense bushes on each side
of rectum. No clearly visible anal rosette.
Specimen and locality
Described from one female specimen collected
during cruise of ‘Dmitrii Mendeleef', Stn 1245, 30°
24'S, 161° 57'E near Lord Howe 1, al 1200.m. No
other record,
Genus Sluiterina Monro
Sluiterina Monro, 1927, p. 618; Murina 1976, p. 840.
Type-species: Mamingia sibogae S\uiter, 1902,
Diagnosis
Proboscis of female non-bifid; lateral edges turin
inwards piving structure a tubular appearance; edges
fuse near mouth to form a cup, Nephridium single,
nephrostome basal. Anal vesicles bushy or brush
like. Male unknown,
* Sluitering alba Murina
Sluiterina alha Murina, 1978, p. 11, fig. 3.
Description (after Murina 1978)
Trunk 44 mm long, 8 mm wide, posterior region
damaged, Body wall white, thick and not
transparent. Proboscis 12 mm long, 6 mm. wide
(distal part damaged), with lateral margins folded
inwards making i tubular in form. Nephridium
single, suc-like, 7 mm long, 3 mm wide, with
centrally located nephrostome, About 50 white eggs
with diameter 0.12-0,13 mom in cavity of body.
Between posterlor coils of gut are bunches of bight
yellow material, probably remains of anal vesicles,
Specimen and lacality
Described from a single female collected during
eruise of ‘Dmitrit Mendeleef', Stn. 1373, Great
Australian Bight, 33° 48S, [27°07°E at 1080-
1100 m. No other record,
Genus Vitjazema Zenkevitch
Viljazema Zenkevitch, 1958, p, 193; Datta Gupta
1976, p. 115.
AUSTRALIAN ECHIURANS 12)
‘Type-species: Vitjuzema ul/raabyssalis Zenkevitch,
1958.
Diagnosis
Probosics of female non-bifid; anterior region,
however, expanded into a shghtly widened ‘head’
with thickened festoons along anterior border;
under festoons are triangular flaps directed inside
a ventral gutter. Setae two. Nephridia two,
nephrostome distal, Anal vesicles sac-like. No male
known,
* Vitjazema ultraabyssalis Zenkevitch
Vitjazema ultraabyssalis Zenkevitch, 1958, pp.
195-197, fig, 3; Murina 1978, p. 115.
Description
Trunk green, length I4-15 mm. Proboscis
9-27mm Jong with deep funnel on ventral side;
anterior region widened with festoon-like border
consisting of 5-6 triangular lappets, Setae 2, large
with bent blades and blunt tips. Nephridia one pair,
wilh distal nephrostomes at end of long tube. Anal
vesicles unbranched, covered with small funnels,
Specimens and localities
‘Two female specimens collected during cruise of
‘Dmitrii Mendeleef’, Stn 1365, Great Australian
Bight, 34° 25°S, 128° 12° 5B at 3880 m.
Distribution
Kurile - Kamchatka Trench (at 5560-9700 m);
Marianne Trench; Great Australian Bight (at 3880
m).
Genus Zenkevitchiola Murine
Zenkeviichiola Murina, 1978, p, 108,
‘Type-species: Zenkevitchiola brevirostris Murina,
1978,
Diagnosis
Proboscis long, non-bifid. Trunk without setae.
Single pephridium, nephrostome basal. Anal
vesicles, two, long, slender, filamentous. Male not
known,
* Zenkevilchiola brevirostris Murina
Zenkevitehiola hrevirostris Munna,
108-109, fig. 1.
1978, pp.
Description (alter Murina, 1978)
Trunk 68 mm long, 28 mm wide, anterior third
and posterior quarter covered with low, rounded
pupillae 1.5 x 0.8 mm. Coils of gut visible through
body wall, Proboscis white, transparent with lateral
margins turned up or folded, length 65 mm, width
5-7 mm, distal extremity curved and widened,
proximal extremity forms cup-with a slit on ventral
side. Nephridium single, }imm Jong, 5mm wide,
located on right side of nerve cord; anterior region
swollen and filled with eggs 0.25-0.3 mm in
diameter, posterior half with thicker walls and na
eggs. Nephrostome basal, stalked and with rosette
at distal end, Anal vesicles two, dark brown,
tapering distally. Gut coils 10-12. Anus forms weak
rosette, surrounded with small papillae.
Specimen and lacality
Described from one female collected during
voyage of ‘Dmitrii Mendeleef’, Stn 1345, near
southern Tasmania, 43° 47'5"S, 147° 51°E at 755
m. No other record.
Genus Anelassorhynehus Annandale
Anelassorhynchus Annandale, 1922, p. 148; Fisher
1946, pp. 221-22; 1949, pp. 480-481; Stephen &
Edmonds 1972, pp. 443-444,
Type-species; Thalassema branchiorhynchus
Annandale & Kemp, 1922.
Diagnosis
Proboscis well developed, usually long, never
bifid. One pair of ventral setae, Longitudinal,
circular and oblique musculature of body wall
continuous. Nephridia, |-3 pairs, Nephrostomal
lips long and spirally coiled (thus differing from
genus Thalassema},
KEY TO AUSTRALIAN SPECLES OF
ANELASSORH YNCHUS
|. Nephridia, (wo pairs and post-setal........... 2
Nephridia, three pairs and post-setal.., A, vegrandis
2. Trunk globular to ovoidal, sandy-grey to light
brown in colour ..... A. poreellus porcellus
Trunk sausage-shaped to clongate, green in
colour... 20. veeey Ay porcellus adelaidensis
Anelassorhynchas vegrandis (Lampert)
Thaslassema yegronde Lampert, 1883, p. 341;
Monro 1932, p. 33.
Anelassorhyachus vegrandis Visher, 1946, p, 222:
1949, p. 481.
Dipe-localin Philippines,
128 5. J, EDMONDS
Description (atler Lampert 1883)
Proboscis lacking. Nephridia three pairs and
post-selal. Nephrostomal lips spirally coiled, Anal
vesicles long and without ciliated funnels.
Remarks
Monro’s specimen Irom the Barrier Reef was in
poor condition and his identification wag made with
some reservation, The species is not well known
Anelassorhynchus porcellus porcellus Fishvei
(Fig, 20)
Anelassorhynchus porcellus Fisher, 1948b, pp.
274-277, figs la-d; Edmonds 1960, pp. 91-92, pl, le.
Type-specinren: US. Nat. Mus., Washington D.C..
Type-locality, Honolulu, on reef south of hartrour.
Description
Yrunk: Globular to ovoidal, colour sandy grey
to light brown, length 25-40 mm, maximum width
15-29 mim} skin rather thick but wrinkled with
tumerous flat papillae; musculature continuous,
Selae two, golden-brown, lying posterior to mouth;
no interbasal muscle present,
Proboscis; Fleshy, readily deciduate, usually
tapering antetiorly, 8-20 mm long.
Alimentary Canat: Very long and fragile, tilled
wilh fragments of coral, small shells and coral sand
(which usually rupture the thin gut wall as soon as
one tries to free the intestinal coils); presiphonal
secon of mid-gut very long.
Vascular System: Consists of dorsal vessel, ring
vessel, 1WO neuiro-intestinal vesgels and a ventral
vessel.
Serae: Two pairs, post-setal, with nephrostomal
lips long, slender and coiled.
Anal Vesicles: Two long, with small, utstalked
funnels, Intestinal siphon present bul no precloacal
caccum.
Sysfematics
The specimens from Heron I, correspond closely
with two of Fisher’s specimens of A. porcellus from
Kakaoha Reef, Hawaii (U.S. Nat, Mus, part 26423).
Fisher was unable to recognise any ciliate funnels
on the anal vesicles of his specimens. In the
Australian specimens the funnels, though small and
sparse, are definitely present. One of Fisher's
Speciinens when dissected was found ro possess
three pairs of nephridia,
Specimens Exarrined
Qu; Heron I, (Capricorn Group) (3) SAM E1425;
North-West 1, (Capricorn Group ) (1) AMS W214;
Ingram J, (1) SAM ElL43]; Brockhurst Reef off
Townsville (1) SAM ELdY4,
Distribution
Western Pacitie Ocean at Hawaii and Great
Barrier Reef, Qu,
Anelassorhynchus porcellus adéelaidensis Edmonds
fi. stat,
(Figs 9-U, 21)
Anelussarhynchys udelaidensis Edmonds, 1960, pp.
92-93, pl, 2a.
Anelassortynchus porcellus (in part) Edmonds,
1982, p. 316.
Type-specimen; AMS; type locality Aldinga Beach,
S.A,
Desvription
A number of specimens Which have previously
been called 4. adelaidensis and A, porcellus (in
part) are now being referred to as a new subspecies,
A. porcellus adelaidensis. The new subspecies
differs from the nominate form in size, colour and
distribution,
Thank: More elongate than nominate subspecies,
length 15-90 mm, maximum width 10-40 mm,
always lighi to dark green, surface wrinkled and
made verrucose by many, large, flat, white papillae
(most Numerous anteriorly and posteriorly); secretes
copious amounts of mucus making animal very
slippery to held,
Probescis, Fleshy, readily deciduate, up to 37 mm
long, lateral edges. may be wavy but never with
processes,
See: Two golden brown, 2.8-5 mm long, no
interbasal muscle (setae lost in some specimens).
Nephridia: Two post-setal pairs (occaslonally an
extra single nephridiufm or pair); nephrostomal lips
filamentous, weakly lo sirongly coiled,
Alimentary Canal; Very long and much
convoluted, with very long presiphonal section of
mid-gut; faecal matter not in form of pellets;
intestinal siphon present but no caecum.
Anal besicles: Two, long, thin walled, tubular bul
usually expanded basally, and attached (especially
basally) to body wall by numerous mesenteries or
muscles; coelomic surface bears sparsely distributed
ciliate funnels and numerous brown spots which
appear to be aggregates of very small pigmented
granules,
Vascular system: Dorsal, ring, two
weurointestinal- and ventral vessels, Hand cut
sections of proboseis show two lateral and one
median vessel, Well-developed ventral nerve cord
extending Into proboscis.
Specimens and lacalities
Vicioria; Port Phillip Bay, S4M E429, Brighton,
SAM B1432..
AUSTRALIAN ECHIURANS
FIGURES 9-11. Anelassorhynchus porcellus adelaidensis. 9, dissected specimen; 10, setae (scale
line = 1.0 mm); II, ciliated cup from anal vesicle.
130 §. |. EBMONDS
South Australia; St Vincent Guil-Porl Willunga,
SAM E1427 (1); Aldinga Reef, SAM E1428 (1); Cape
Jervis, SAM E1430 (1); Coobowie, SAM E1454 (1),
SAM E1455 (1), SAM E1456 (5), SAM El46l (3),
SAM E1462 (1); Brighton, SAM E1463 (5). Spencer
Gulf-at following islands al Banks Group, Winceby
1_, SAM E1476 (3), Marum £., SAM 61477 (2), SAM
E1486 (1), SAM E1493 (3), Lushy lL, SAM E1489
(3). Eyre Peninsula-Port Lincoln, SAM E1479 (2);
Port Turton Jetty, SAM B1S03 (1); Venus Bay (under
jetty), Blanche Pt (Streaky Bay), Smokey Bay, (coll,
K. Gowlett-Holmes)..
Systematics
The internal anatomy of the South Australian
specimens corresponds very closely with that of 4.
porcellus, Their shape, however, is more elongate
and their colour green. Fisher does not mention the
colour of his specimens bul specimens of A,
porcellus collected by the author at Heron I., Qu,,
were light brown to sandy grey, The green subspecies
is commonly collected by divers in S.A. but no
brown forms have yet been found in the State. The
distribution of the two subspecies consequently
seems different. A. gangae Bisewar, 1984, recently
described from Natal, South Africa and A,
porcellus adelaidensis are closely related species.
Distribution
Southern Australia from Port Phillip Bay ( Vie.)
to Streaky Bay (Eyre Peninsula) in S.A,
Genus Arhynchite Sato
Arhynchite Sato, 1937, pp. 142-143; Fisher 1946,
p. 485; Stepheri & Edmonds (972, p. 414,
Type-species: Vhalassema arhynchile \keda.
Diagnosis
Proboseis long slender, often ribbon-ike,
sometimes deciduate; anterior extremity expanded
into @ fan-like structure, Two ventral setae with
strong interbasal muscles. Nephridia two, with
nephrostomal lips expanded into a leaf-like
structure. Anal vesicles long, thin walled and
unbranching, Vascular system with or without ring
vessel
Arhynchite hiscocki Edmonds
Arhynchite hiscocki Edinonds, 1960, pp. 9U-91, Ng.
3, pl. 1, fig. 1b; 1966, p. 178; Stephen & Eximonds
1972, p. 417,
Holotype: AMS W3714; type locality: Dunwich,
Qu., ‘dug from sand pit, 18" below surface’,
Description
Trunk: Elongate, slender, pencil-like, length
100-120 mm, width 4-6 mm, fixed specimens
yellow-brown to zrey green. Surface made verrucose
by numerous near rows of elevated papillae, slightly
larger at anterjor and posterior extremities.
Masculature continuaus,
Proboscis: Delicate, slender, about 30 mm long,
{.5-2.5 mm wide, still attached to trunk in holotype;
anterior extremity flattened and fan-like. In one
specimen from Victoria proboscis is shorter,
deciduale, with antenor extremity more spoon-like,
Selae: Two, connected to bady wall internally by
strong radiating muscles and to each other by strong
interbasal muscle.
Nephridia: Two, sub-cylindrical, slender, length
variable; postsetal, Nephrostome basal was
expanded, frilled or leaf-like lip.
Alimentary Canal: Midgut with siphon; no
precloacal caecum.
kascular System: Dorsal blood vessel fuses with
foregul at posterior extremrty of latier;
neurointestinal vessel conneets with anterior section
of midgut near anterior extremity of siphon, No
Ting vessel observed.
Anal Vesicles: Two, very slender, brown, about
one third to a quarter as long as trunk, fastened
(hroughout their length to posterior region of
alimentary canal but to body wall over last quarter
of their length, Numerous ciliated furmels scattered
over their surface.
Systematics
Sato (1937) erecled the genus Arhynchile for a
group of echiurans Jacking a proboscis, Fisher
(1949), having found cwo species possessing a long
deciduate proboscis, redescribed the genus. The
proboscis of the helotype of 4. hiscocki is still
attached but that of one of the Victorian specimens
is detached.
The genus contains six species collected from
places bordering the Pacitic Ocean; A, californicus
-Monterey {U.S.A,), A. inamoenus - Monterey
(U.S.A), A, pugettensis Puget Sound (U.S.A.), -4-
rugosus -Shantung (China), 4, arhivachite —Japan.
Some of the spevies are closely related and difficult
to distinguish, 4, Aiscacki is not a well known
species und needs re-examination and revision when
more species are found.
Specimens and localities
Queensland: Stradbroke |, (Edmonds 1960),
AMS W714 (2)
Victoria; Port Phillip Bay (Editiends 1966) VM
coll, (2)
AUSTRALIAN ECHILIRANS it
South Australia; Spencer Gulf, north of Port
Lowly, SAM E1524 (1B).
Genus Listriolobus Spengel
Listriolobus Spengel, $912, p. 316; Fischer 1926,
p. L10; Fisher 1946, p, 233.
Type-species: Listriolobus bahamensis (Fischer),
(designated by Fisher 1944),
Diagnosis
Proboscis of variable length, truncate but never
bifid, Two setae with interbasal muscle,
Longitudinal musculature of trunk wall grouped
into bands (not always well developed in young
specimens), Oblique musculature not banded or
fasciculated as in Ochetosiama. Nephridia two to
three pairs, nephrostomal lips long and spirally
coiled. Anal vesicles sac-like to tubular and without
branches.
Kry 10 AUSTRALIAN SPECIES OF LISTRIOLORBUS
|. Longitudinal musculature in 7 bands L.. brevirostris
Longitudinal musculature in 13 bands 1. sorbrllans
?”* Listriolobus sorhillans (Lampert)
Thalassema serhillans Lampert, 1883, pp. 340-341;
Augener 1903, p. 349.
Listriolebus sorbillans Fisher, 1946, p. 234.
TWpe-locality: Philippines.
Australian record: Sydney (call. Dr Schutte, 1876)
in Augener 1903, p, 349.
Description (after Lampert 1883)
According to the type descriprion (based on a
single specimen) the trunk is 65 mm long and
proboscis 24. Longitudinal musculature in 13 bands,
Nephridia three pairs with spirally coiled
nephrostomal |ips, first pair presetal in position,
Setae small, Anal vesicles long, brown, bearing
tmicroscoupic ciate funnels, Small rectal caecum.
Augener’s description of his single specimen is
brief. Trunk about 42 mm long, proboscis 18 mm,
Whole body covered wirh papillae which are
smallest in mid-trunk and largest posteriorly, Anal
vesicles about two-thirds length of trunk,
Remarks
This Australian record needs conlirmation. If the
oblique musculature of Augener’s specimen was
fasciculated then it might have been Ocherostoma
australiense,
Listriolobus breviewstris Chen & Yeh Chen-Chang
(Figs 12-13, 22)
Listriolobus brevirostris Chen & Yeh Chen-Chang,
1958, pp. 273-278, fig. 7 A-D; Stephen and
Edmonds 1972, p, 424,
Listriolobus bulbocaudatus Edmonds, 1963, pp-
243-244, pl. 1, Fig. 1.
Type-locelion, Kiao-chow Bay, Shantung, China.
Description
Trunk: Sub-cylindrical to cigar shaped, length
2-85 mm, maximum width 10-20 mim, fixed
specimens light to dark pink, Surface cavered with
while papillae, lying almost in rows. Posterior
extremity way sometimes be modified and
expanded Into a fleshy, bulbous, conical structure:
bearing three or four rows of prominent, pointed
or mamillate, white or pink papillae. Longitudinal
rnusculature arranged in bundles, often difficult to
discern externally. Dissected specimens show seven
(eighi in one specimen) well spaced longitudinal
bands, occasionally weakly developed where the
body wall is thin. Oblique musculature between
bands of longitudinal musculature continuous and
not in fascicles.
Proboscis; Non-deciduate in all specimens, In
fixed condition short, stout, 10-16 mm long, 6-12
mm wide. Small papillae on dorsal surface. Lateral
margins wrinkled, folded, indented or crenated. Ne
lateral processes as in Anmelassorhynchus
branchiorhynchus Ammandale & Kemp.
Selae: Two (with smaller reserve setae), length
(measured in straight line from base co tip) up to
7.2mm, strongly hooked and sickle-shaped. Strong
interbasal muscle.
Nephridia: Two post-setal pairs, in one specimen
three nephridia on one side and two on other.
Length variable, some extending almost to posterior
extremity of trunk. Nephrostamal lips long and
much coiled,
Alimentary Canali Long, much coiled,
Presiphonal section of midgut long. Precloacal
caecum present.
vascular System: Dorsal vessel expands into a
thin walled saccular vessel or heart. Well-developed
ring vessel at Junction of fore- and midgut gives off
two neurointestinal vessels which join before they
reach the interbasal muscle and then bifurcate to
form a loop around the muscle Ventral vessel
pressed close to nerve cord and terminates in the
cloacal caecum.
Anal Vesicles; Long, thin walled, brawn, swollen
basally in most specimens, bearing numerous,
small, ciliated cups some on very short stalks.
Anteriorly placed cups more sparsely distributed.
132 5. J. FDMONDS
Svstematics
These Australian specimens closely resemble
Ochetostorna septemvotum Datta Gupta, Menon
& Johnson, 1963 from Quillon, India, In none of
the Australian specimens, however, has the oblique
musculature of the body wall been found to form
fascicles, like that shown for Ochetostoma
acionyotum by Fisher 1946, pl. 23, fig. 2, and pl.
24 (in the spaces between muscles labelled MYL,
ML and MDL) or like that shown in the transverse
sections of Ochelostoma bombayense by Mathew
1976, fig, 5,
Because fasciculation of the oblique musculature
is a character of Ochetostoma, the Australian
spetimens are considered different from O.
septemyvoruin,
Edmonds (1963) originally described the
specimens from Queensland as Listriolobus
bulbacaueatus. At the time he was unaware of 1,
brevirostris Chen & Yeh Chen-Chang (1958) from
Kiao-chow Bay, Shantung, China. At a laler date
Stephen and Edimorids (1972, p. 424) considered the
two species were distinguished by three or four rows
of prominent papillae and a bulbous structure both
present at the posterior region of L. bulbocaudatus.
More recently the author has examined three
specimens from Queensland, in which the rings of
prominent papillae and the bulbous structure are
much reduced, This information brings the
specimens. within the range of Z, brevirastris.
Consequently L. bulbocaudatus \s now considered
to be a junior synonym of LZ, breviresiris Chen &
Yeh Chen-Chang, (1958),
Speciniens examined and localities
Queensland; Yeppoon, SAM E434 (1); Mud t.,
Moreton Bay, SAM E1433 (1); Round Is, Hervey
Bay, SAM E1436 (1); Bramble Bay, SAM EL460 (1);
? Bramble Bay, SAM E1435 (4). Dredged from mud.
Distributian
China at Shantung, Australia: Queensland.
Genus Ochetostoma Leuckart & Rueppell
Ochetosioma Leuckart & Rueppell, 1828, pp. 7-8;
Fisher 1946, p. 240; Stephen & Edmonds 1972,
p, 426,
Type-species: Ochetostoma erythragrammon
Leuckart & Rueppell, 1828.
Diagnosis
Proboscis long, Capable of much extension, non-
bifid. Trunk medium to large with longitudinal
musculature lying in well-defined bands. Intervals
between bands crossed by numerous fascicles ar
small bundles of inner oblique musculature (Fisher
12 7 13
FIGURES 12-13, Lisirialobus brevirnstris, 12, anterior
region dissected; 13, seta
1946, pl, 23, fig, 2), Nephridia in one to seven pairs,
with long spirally coiled nephrostomal lips. Setae
two, with or without interbasal museles. Anal
vesicles long, more or less tubular, unbranched
Rectal caecum usually present.
KEY TO AUSTRALIAN SPECIES OF OCHETOSTOMA
1, Longitudinal musculature in /1-\3 (\1-14) bands; three
pairs of nephridia, One pair presetal, (wo pairs
postsetal. Fasciculation of oblique musculature
usually well developed. No intérbasal muscle
ii todelgh hers eege tevy tera Ch yustentivose
Longitudinal musculature in 17-19 (17-21) bands; two
pairs of postsetal nephridia. Fusciculation of
oblique muscualture well developed, Strone
interbasal muscle.) 1.00.60. cove OQ baroné
Ochetostoma australiense Edmonds
(Figs 14-15, 23)
Ochetostoma australiense Edmonds, 1960, pp.
93-94, tig. 4, pl. 2b; Datta Gupta & Menon 197],
pp. 177-178, figs. 2c, 2e.
Tipe-specimen: AMS; type loeality, mud flats at
Dunwich (Stradbroke [.), Qu.
Description
Trunk: Usually large, sausage-like, cigar shaped
or elongate, pale to dark red, length 40-130 mim,
maximum width 15-30. Thickness of body wall
variable, sometimes Very thin, Surface, especially
in anterior and posterior fegions, covered by
numerous small, fal, fleshy to wartlike papillae.
Usually 12-13 (11-14) well developed longitudinal
muscles best counted in dissected specimens. Jn 2¢)
dissected specimens the maximum number of bands
was 14 in 3, 13 in 10, 12 in 6 and J) in one, Two
AUSTRALIAN ECHIVRANS 35
bands, one on each side of the nerve cord, lie very
close together and may appear to be one. Oblique
musculature between longitudinal bands usually
grouped into numerous fascicles, which may be
weakly developed or even absent in parts of some
animals.
Proboseis: Jn living animals is highly extensible
(150-200 mm), pale and ribbon-like. Ln. fixed
specimens shorter, fleshy, up to 60 mm long, with
lateral margins rolled inwards on yeritral side.
Usually adherent or non-deciduate, Anterior
extremity may be flattened somewhat and lateral
Margins may be slightly wrinkled. No lateral
PIPOCeASES,
Serae: Two, 2.5-3,1 mm long (measured in a
Straight line from tip to midpoint of base), golden,
encased in sheath connected to body wall by a
nuniber of sera] muscles. No interbasal muscle.
Nephridia: Three pairs, the first presetal, others
postsetal, Length variable, sometimes over half
length of trunk. Nephrostomal lips elongate and
spirally coiled, although Sometimes only weakly, Jn
one specimen only five nephridia present.
Digestive System: Mouth at base of proboscis.
Foregut short, midgut very long and much eoiled,
Presiphonal section of midgut long and traversed
for part of its length by cilrale groave, Siphonal
section of gul also long, Well developed caecum
present. Gut contents largely mud and sand; oo
faecal pellets.
Vasenlar System: Consists of dorsal blood vessel
(sometimes well expanded), ring vessel or sinus (may
also be expanded), two long neurointestinal vessels
and a ventral blood vessel. Two neurountestinal
vessels fuse to form one short vessel which joins
the ventral vessel at about the level of the setae,
Posterlorly ventral vessel gives off a branch to
caecum,
Anal vesicles: Two, long, slender, thin walled,
light to dark brown, opening into cloacal region of
intestine. Posses small, unstalked ciliate funnels.
Systematics
Edmonds (1960) considered! these specimens with
11-14 muscle bands to be different from O.
ervrhrograminod Leuckart & Rueppell which
possess 14-18 muscles. @ erytArogrammon and O,
auslraliense. however, are closely related, The
neurointestinal vessel of all the Australian
specimens exaniined in the presen study is double
for most of its length, Datta Gupta & Menon (1971)
state that the corresponding vessel in their
specimens Of & ervthragrammon is single, [f this
difference always exists, ir further distinguishes the
lwo closely allied spevies, Sato (1999, fig. 9),
however, shows two ceuroiptestinal vessels for his
specimens af C2. ecvifrogriniunert.
Stephen & Edmonds (1972) list nine species of
Ocherostoriu that possess three pairs of nephridia
and in which the number of longitudinal muscles
varies from 12 to 22. Wesenberg-Lund (1939) and
Sato (1959) considered that they were conspecilic.
If they are correct then O. australiense would
become ©. ervthrogrammon.
Specimens examined and localities
Queensland: Dunwich (in mud flats-in front of
cemetary) SAM E1410 (4) and B14] (1); Caloundra
(dug at low tides from mud flat; opposite a small
mangrove island) SAM El415 (7); Myora (in mud
flats at law ude) SAM ES413 (20).
New South Wales; Goodwood [. (near mouth of
Clarence River) AMS W3186, W3375, W387;
Brunswick Heads (1) AMS coll,
Habitat
At Goodwood 1., the worms are found ‘between
high and low water marks in rather dark sand
Situated close to some small mangrove clumps. The
proboscis is white and fleshy and protrudes from
a hole in the sand. [t lies along the surface of the
sand and is about 6” long and 1/2” wide. In this
condition it appears to be quile flat (like a ribbon)
and does not appear to take ona tube-like shape
as in preserved specimens. The body of the worm
is Solt and bright red in colour. Twelve longitudinal
muscles show up clearly’ (P, Durie pers, comm.)
Distribution
Eastern Australia from Caloundra (Qu.) to
Goodwoad Et. (N,S.W.), Andaman 1. (Datta Gupta
& Menon, 1971). An inhabitant of intertidal mud
f}ats.
Ochelostoma barani (Greeff)
(Fig. 24)
Thalassema barenit Greeff, 1879, pp. (41-192, pl.
6, figs 62-67-
Ochetostoma baronij Mackie, 1951, p. 247; Stephen
& Edmonds 1972, p, 429; Amor (974, p, 123-124.
Ochetostoma nyersae Edmonds, 1963. pr. 245-246,
pl. 1, fig. 2.
Previous Ausiralian record: N.SANV, (Edmonds 1963),
Description
Trunk: Small to moderately large, sac~ sausage-
or cigar-shaped, length 21-70 mm, maximum width
9-25 mm; anterior region rounded, posterior
sometimes almost pointed. Green. Surface covered
with soft, almost white, shghtly clevated, wari-like
papillae, largest in posterior region of trunk.
Longitudinal musculacure in 18-19 (1 in one
specimen) bundles, Oblique musculature between
134 S. J. EDMONDS
longitudinal muscles forms fascicles which
sometimes are only weak.
Proboscis: In preserved specimens about half to
fifth length of trunk, either deciduate or non-
deciduate. Lateral margins tend to roll inwards so
as to form a tube. Plump and almost concial in
largest specimen.
Setae: Two, up to 3.1 mm long, golden, connected
by strong interbasal muscle.
Alimentary Canal: Mouth at base of proboscis.
Gut much coiled and filled with coral and shell
fragments; obviously animal is able to ingest larger
particles than Ochetostoma erythrogrammon and
Bonellia viridis (Chuang 1962, Jaccarini & Schembri
1977).
Nephridia: Two post-setal pairs, nephrostomal
lips long, weakly or strongle coiled. Largest ova
0.09-0.11 mm in diameter.
Anal Vesicles: Two, very large, slender, tapering
distally and bearing numerous, small, brown
unbranched ciliate cups or funnels.
Vascular System: Dorsal blood vessel, ring vessel,
two neuro-intestinal vessels and ventral vessel.
Neurointestinal vessels long but fusing to form one
short vessel which joins ventral vessel at about level
of setae.
Systematics
Ochetostoma myersae Edmonds, 1963, was
described from N.SW. as possessing 18-21
longitudinal muscles, two pairs of post-setal
nephridia and unbranching ciliate funnels, At the
time the species was considered to be different from
O. baronii (Greeff) in which the ciliate funnels were
described as being branched, a fact confirmed by
Fischer (1895: 19).
Amor (1976, p. 123), however, after studying
specimens collected at Canary Is (type locality),
Brazil and Galapagos Is. found that ‘amongst the
38 specimens examined there did not exist any
branched outgrowths in the anal vesicles’, The
examination of three specimens of O. baroni from
Arrecife, Canary Is, collected by A.K. Totten and
identified by A.C. Stephen (B.M. 11.7.7.37).
confirms Amor’s finding that the fuhnels are
unbranched,
In view of this evidence (especially as Amor
examined 38 specimens) the statements of Greeff
and Fischer about the branching of the ciliate
funnels are questionable. Consequently the chief
reason given by Edmonds (1963) for separating O.
myersae and O. baronii is now invalid and O.
myersae becomes a junior synonym of O. baronii.
Amor (1976: 123) also considers O. edax Fisher,
1946 and O. kefersteini (ten Broeke, 1925) as junior
synonyms of O. baronii.
O. punicea (Dartnall, 1976) is very closely related
to O, baronii if the ciliated funnels of the latter are
unbranched. O. punicea has 18-19 longitudinal
muscles, fasciculated oblique musculature, setae
about 2.25 mm long, two pairs of post-setal
nephridia with spirally coiled lips and anal vesicles
with unbranched ciliate cups. No interbasal muscle,
however, is present and the left anal vesicle is not
symmetrically placed in relation to the right.
Whether the last character is taxonomically
significant is doubtful. The species was described
from Great Tulear Reef, off south-west Madagascar.
Specimens examined and localities
New South Wales: Long Reef (near Sydney) (1)
AMS 3357; Collaroy, AMS W3368 (1); Minnie
Waters, ‘intertidal region of low tide’ AMS coll. (1).
Queensland: Bird I. (Moreton Bay) (1) SAM
E1417.
Distribution
Eastern Australian from Sydney (N.SW.) to
Moreton Bay (Qu.).
Extra Australian, wide: Atlantic Ocean (Canary
I., Bermuda, West Indies, Senegal, Florida, Brazil);
Indian Ocean (Zanzibar, Amboina); Pacific Ocean
(Papua, Loyalty I., Galapagos).
FIGURES 14-15. Ochetostoma australiense. 14, anterior
region dissected; 15, setae (scale line = 1.0 mm).
AUSTRALIAN ECHIURANS 135
Genus Thalassema Lamarck
Thalassema Lamarck, 1801, p. 28; Fisher 1946, p.
230; Stephen & Edmonds 1972, p. 452.
Type-species: Lumbricus thalassemus Pallas, 1776
= Thalassema thalassemum (Fisher, 1946).
Diagnosis
Echiuridae with well developed, long, non-bifid
but usually truncated proboscis Two ventral setae;
lacking anal setae. Longitudinal, circular and
oblique musculature continuous, Nephridia in one
or LWo pairs; nephrostome basal and nephrostomal
lips neither elongated nor spirally coiled.
Thalassema sydniense Edmonds
(Figs. 16-17)
Thalassema sydniense Edmonds, 1960, p. 89-90,
figs. 1-2, pl. la.
Holotype: AMS G11219; off Watson Bay, Sydney,
N.S.W.
Description (based on four specimens reported in
Edmonds, 1960 and two additional ones)
Trunk; Small, grey-brown, sausage to sub-
ovoidal; length 2.5-8 mm (most about 5), width
1-2.7, Surface covered with papillae, lying almost
in tows and largest posteriorly. Musculature
continuous.
Proboscis: Firmly attached, as long as trunk or
less, becoming narrower anteriorly.
Setae; Two, golden brown, 1.0-1.1 mm long, with
strongly recurved tip; strong interbasal muscle and
well developed system of setal muscles,
Nephridia: Two pairs, post-setal. Nephrostome
on short stalk near proximal extremity of
nephridium; lips expanded but not elongate or
spirally coiled. One specimen with only 3 nephridia.
Alimentary Canal: Very long, intestinal siphon
present but no caecum.
Anal Vesicles: Two, expanded towards base;
surface with ciliated cups.
Systematics
These specimens resemble Thalassema steinbecki
Fisher, 1946, which occurs along the Pacific coast
of North America from California to Ecuador and
which has also been reported from the Indian Ocean
(Datta Gupta 1975), T sydniense differs from T.
steinbecki because its nephrostomes are on short
stalks or peduncles. 7. sydniense is known only
from six specimens, four of which are very small.
it is not a well known species and needs
redescription when more specimens become
available. Whether the species is a small one or
whether the specimens so far collected are simply
small ones is not known.
Specimens examined and localities
New South Wales: Watson Bay, AMS G11219 (4).
Victoria: 40° 39°0"S, 144° 56’E (Bass St Survey)
MV G3386 (2).
No other records.
Q
\ 17
FIGURES 16-17. Thalassema sydniense, 16, entire animal
(scale line = 1,00 mm); 17, seta (scale line = 0.25 mm),
Genus Ikeda Wharton
Ikeda Wharton, 1913, pp. 260-261; Fisher 1946, pp.
220; Stephen & Edmonds 1972, pp. 471-472.
Type-speécies: Thalassema taenivides |keda, 1904,
Diagnosis
Trunk very long with longitudinal musculature
thickened to form bands, Proboscis very long, non
bifid. Nephridia very numerous and unpaired.
Ikeda sp,
The yery long proboscis of an echiuran has been
noticed and collected a number of times by divers
136 8. J, EDMONDS
in St Vincent Gulf, S.A. [Edmonds 1982, pl. 23 (4)].
They report that the organ Is able to extend for more
than 1.5 m, The echiuran itself, however, has proved
very difficult to collect on account of the depth of
iis burrow and the problem of digging in sand at
depths of 6-10 m. So far only one specimen has
been collected. Unfortunately, it was considerably
damaged so that only a limited amount of
information can be given about it, Several intact
probosces of other specimens have been collected,
The worm resembles in some respects /keda
taenioides (Ikeda, 1904), known from six Japanese
specimens. A specific identification of the specimen
from S.A_18 nat possible on accotint of the damage
to its nephridial and anal regions. 1 am, however,
tentatively assigning it to the genus Ikeda,
Description
Trunk: Long, slender, worm-like, rather flat jn
preserved condition; 290 mm long, 7-11 mm wide,
pinkish-brown when collected but dark brown when
fixed. Langitudinal musculature grouped in 5 bands
prominent externally; numerous. small sub-globular
papillae cover surface of much of trunk.
Proboscis; Flat, about. 400m long, 5-10 wide, with
margins in fixed specimens slightly frilled; one
surface cream-brown in colour marked with almost
transverse brown-black stripes; posterior region of
proboscis (near mouth) modified to form cup-like
Structure,
Seftae: Two, about 10 mm long, with well
developed setal muscles. Nephridial region much
damaged,
Anal Vesicles: Missing,
Alimentary Canal: Very long and convoluted,
Some eves with maximum diamrer 0.35-0,38 mm
entangled im gut.
Specimens examined end localities
‘Kemps Ground’, off Glenelg, St Vincent Gulf,
S.A., at 9-10 m, 13 March, 1986, ane spec. coll, N.
Holmes and S. Parker; SAM E1509; separate
probosces of other specimens SAM E1587, Coffin
Bay (near Black Spririgs), Eyre Peninsula, S.A-
(probosels only).
ACKNOWI EDGMENTS
Thanks are due to the following far help with specimens:
from W.A,, Mrs L. Marsh, Mra S$, Slack-Smith, Dr E.
Hodgkin and Dr B. Wilson: fram 8.A4., I.M. Thomas, Mrs
K, Gowletl-Holmes, N, Holmes, Miss H. Kald and W,
Zeidler; fran) Vic, Dr B, Smith and Mry H, Black; from
Tas., Dr A. Dartnell and Dr A. Green; from N.SW., Dr
P. Hutchings, Miss P, Wearne, Miss E. Pope and Miss E.
Bennett, trom Qu,, Prof W. Stephenson, W. Green, S.
Cook, ©. Kelly and Mrs M. Specht. Mr P. Kenipster
(University of Adelaide) took the photographs and Dr 1.
Beveridge (Institate of Medical and Veterinary Sciience,
Adelaide) made sections of the body wall of one apecies,
Mr R. Sims and Mr E, Easton (British Musuem of Natural
History, Londen) and Dr M. E. Rice (Smilhsonian
Institute, Washington D.C.) kindly sent specimens on loan,
Dr E, Matthews (SAM) helped with translations {rom
Russian.
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FIGURES 18-24. 18, Metabonellia haswelli; 19, Pseudobonellia biuterina; 20, Anelassorhynchus
porcellus porcellus; 21, Anelassorhynchus porcellus adelaidensis; 22, Listriolobus brevirostris;
23, Ochetostoma australiense; 24, Ochetostoma baroni,
FURTHER OBSERVATIONS ON PENTATOMIDS (ARTHROPODA)
PARASITIC IN AUSTRALIAN REPTILES AND MAMMALS
BY J. RILEY & D. M, SPRATT
Summary
A collection of adult and nymphal pentastomids representative of at least four genera is described.
Two species of Raillietiella and one species of Parasambonia from snakes (Pseudechis australis,
Pseudonaja textilis and Cryptophis nigrescens) are probably new, but more specimens are required
before their status can be confirmed. Mature and immature Waddycephalus longicauda, W.
superbus and W. punctulatus (all Riley & Self 1981) are identified from snake hosts but specific
determination of five lots of specimens was not possible. Evidence endorses an earlier suggestion
that there may be two species of Waddyccephalus in tiger snakes ; W. scutata from island
populations and an unnamed species from mainland populations of the Notechis scutatus/ater
complex. Nymphal specimens of Waddycephalus from marsupials (Parantechinus apicalis and
Dasykaluta rosamondae), a snake (Cryptophis nigrescens), a gecko (Heteronotia binoci), a skink
(Hemiergis decresiensis) and frogs (Ranidella remota and Palmatorappia solomonis) all bear
characteristic double hooks. The accessory spine above the hook arises from a point midway
between the hook and the fulcrum and appears to be an integral and functional part of the hook.
Armillifer australis Riley & Self 1981 is described from infections in four pythons (Morelia
amethistina and Morelia spilota) ; the latter is a new host record. A single nymph recorded from the
body cavity of Rattus leucopus is identified as A. australis on the basis of abdominal annulus
counts.
FURTHER OBSERVATIONS ON PENTASTOMIDS (ARTHROPODA)
PARASITIC IN AUSTRALIAN REPTILES AND MAMMALS
J. RILEY & D, M, SPRATT
RILEY, J. & SPRATT. D, M, 1987. Further observations on Penjastomids (Arthropoda) parasitic
in Australian reptiles and mammals, Ree, 8, Aust, Mus. 21(2): 139-147.
A collection of adult and tiymphal pentastomids representative of at least four genera is
described. Two species of Raillietiella and one species OF Parasambonia from snakes (Pseudechis
australis, Pseudonaja textilis and Cryptophis nigrescens) are probably new, but more specimens
are required before their status can be confirmed. Mature and immature Waddyecephalus
fongicauda, W; superbus and W. puncrulatus (all Riley & Self 1981) are identified trom snake
hosts Dut specific determination of five lots of specimens was not possible, Evidence endorses
an earlier suggestion that there may be two species of Waddycephalus in liger snakes; Wi scufata
from island populations and an unnamed species from mainland populations of the Notechis
sculalius/ater complex, Nympha) specimens of Waddycephalus from marsupials (Parantechinus
apicalis and Dasykaluta rosamondae), a snake (Cryptaphis nigrescens), a gecko (fleteronotia
binoei), a skink (Hemiergis decresiensis) and. frogs (Ranidella remota and Palmatorappia
solomonis) all bear characteristic double hooks, The accessory spine above the hook arises from
a point midway between the hook and the fulcrum and appears to be an integral and functional
part of the hook. Armmillifer ausiralis Riley & Self, 1981 is described from infections in four
pythons (Morelia amethistina and Morelia spitota); the latter is a new host record. A single nymph
recorded from the body cavity of Rattus /eucopus is identified as A, gusiralis on the basis of
abdominal annulus counts,
J. Riley, Department of Biological Sciences, University of Dundee, Dundee DDL 4HN, Scoiland,
UK. & D. M. Spratt, Division of Wildlife and Rangelands Research, CSIRO, P.O, Box 84,
Lyneham, A.C.) 2602. Manuscript received 21 October 1986.
In an historical review of Australian
Pentastamida, Riley, Spratt & Presidente (L985)
recorded seven genera comprising 17 species occur-
ring in Australian reptiles and mammals, and identi-
fied nymphal Weddycephalus spp. and Armillifer
spp. from marsupials, This paper reports primarily
on a pentastomid collection in the South Australian
Museum (SAM) and describes further adult and
nymphal material, attributed to the genera
Waddycephalus and Armillifer, from reptiles,
amphibians and mammals. The double nature of
the hook of nymphal Waddycephalus (Riley et al.
1985) is. confirmed, as are earlier observations (Riley
& Self 198ib) of significant anatomical differences
beiween mainland and island forms of Waddy-
cephalus infecting the same species of snake, Two
Jarge raillietiellids and two parasambonids from
snakes are described, however more material. is
required before their specific status can be con-
firmed as new.
MATERIALS AND METHODS
‘The material exantined in this study was collected
primarily in eastern mainland Australia, Tasmania
and neighbouring offshore islands, It is
supplemented by nymphal Waddycephalus (or
Elenia?) spp, from ibe Solomon Islands and New
Guinea.
The methods are those outlined in Riley e¢ af-
(1985) and the hooks of railhietiellids were measured
accotding to the convention of Ali ef al, (1982) i.
barb length AB (notation AD in error, p. 42, Riley
et al, 1985), shank length BC, Overall hook length
of the double hooks of nymphal Waddycephalus
spp. was measured according to the convention
illustrated in Figure 2. All measurements are in
micrometers with the exception of body length,
which is in millimetres. Most specimens are
deposited in the South Australia Museum (SAM),
Adelaide; two lots are deposited In the Queensland
Museurn (QM), Brishane Reptile nomenclature
follows Cogger, Cameron & Cogger 1983; dasyund
marsupial nomenclature follows Archer 1982,
Order CEPHALOBAENIDA
Raillietiella spp. trom snakes
Railiietielta sp. a
Material Examined
From lung of Pseudechis qustralis (Gray), locality
unknown (died in Melbourne Zoo), in SAM No,
NI9BO183,
SPRATT
J. RILEY & D. M.
140
AUSTRALIAN PENTASTOMIDS 14]
Description
Female (n = 4). Length 33-52 (K = 42), with 45
or 6, 41 or 2, 36 (2) and 39 (?) annuli respectively.
Posterior hook of 52 mm specimen, AB 240, BC
370.
Male (n = 1). Length approx. 9, annuli uncount-
able and therefore slide-mounted. Posterior hook,
AB 135; BC 220. Base of copulatory spicule
massive, maximum diameter 520 and covered with
a reticulum of tubular elevations (Fig. 1A).
Discussion
The heavily ornamented male spicule, 520 um
across at the base (Fig. 1A), is virtually identical
to those of R. orientalis and R. agcoi (Ali, Riley
& Self 1982). R. orientalis infects colubrid, viperid
and elapid snakes in south-east Asia and Taiwan
whereas R. agcoi is found only in cobras in the
Philippines. The females of these two species are
distinguished in a number of ways: the former has
bigger hooks (see Fig. 3, in Ali et a/. 1982), more
annuli (33-47 contra 30-35) and is generally longer
and stouter than R. agcoi. The overall shape of the
present species, and its hooks, are reminiscent of
R. agcoi but its annulus count, though variable, is
within the range of R. orientalis. The host, the king
brown snake, Pseudechis australis, is an endemic
species and its raillietiellid parasite may be unique
by virtue of geographic isolation. This is probably
a new species, but, because it combines important
characteristics of these two closely related species
we have left it unnamed, pending more specimens
and more refined diagnostic techniques.
Raillietiella sp. b
Material Examined
From lung of Pseudonaja textilis (Dumeéril,
Bibron & Duméril), Townsville, Queensland, in
SAM No. N1985149.
Description
Female (n = 2). One headless abdomen; length
other specimen 50, annuli uncountable, specimen
therefore slide-mounted. At least 30-40% of eggs
in uterus contain fully-developed primary larvae;
the specimen is therefore mature. One posterior
hook measured, AB 400; BC 510.
FIGURE 2. Diagrammatic representation of hook of
nymphal Waddycephalus sp. The distance between the
larger arrows indicates our measurement of the overall
length (a = apodeme; ah = accessory hook; f = fulcrum;
hb = hook barb) (scale bar = 120 ym).
Discussion
Apart from the species recorded above, only two
raillietiellids are known from Australia; R.
amphiboluri from the bearded dragon,
Amphibolurus barbatus (Cuvier), (Mahon 1954,
Riley et a/. 1985) and R. scincoides from the eastern
blue-tongued lizard, Tiliqgua scincoides (White),
(Ali, Riley & Self 1984): the latter has blunt-tipped
posterior hooks and R. amphiboluri is smaller than
the present species with much smaller hooks (AB
200-220; BC 370) (Ali et al. 1985) (notation AD
in error).
The intact specimen clearly belongs to the Group
VI taxon of raillietiellids (Ali et a/. 1985) which
includes all of the species from snakes. The two
species, R, orientalis from south-east Asia and
Indonesia and R. agcoi from the Philippines are,
zoogeographically, most proximate to the present
specimen, but it has much longer hook barbs
(dimension AB) than either of these species
(compare with Fig. 3 in Ali, Riley & Self 1982). This
is almost certainly a new species but the poor state
jn ee
FIGURE 1. A, Copulatory spicules of male Raillietiella sp. from Pseudechis australis showing
massive base covered with tubular elevations (scale bar = 500 um). B. Cephalothorax of male
nymph of Waddycephalus sp. from mesentry of Satanellus hallucatus [described by Riley ef
al. (1985)]. The double nature of the outer hooks is obvious but the inner hooks are not in the
plane of focus (m = mouth) (scale bar = 500 pm). C. Outer hook of nymph of Waddycephalus
sp. from Hemiergis decresiensis showing spinous extension. The back of the extension forms
a long apodeme (a) (f = fulcrum) (scale bar = 100 zm). D. Hook of adult Armillifer sp. from
python (S.A.) illustrating typical unornamented porocephalid hook (a = apodeme; f = fulcrum)
(scale bar = 200 pm).
142 J, RILEY & D. M. SPRATT
of preservation and lack of males precludes specific
identification,
Order POROCEPHALIDA
Parasambonia spp. from snakes
Parasambonia bridgesi Riley & Self
Material Examined
From lung of Pseudechis porphyriacus (Sha\w),
Healesville Sanctuary, Victoria, in SAM No.
N1980173.
Description
Female (n = 2), Length 26, with 50 and 54 pre-
vaginal and post-vaginal annuli.
Male (n = 1). Length 7, with 53 annul
Discussion
There are no uncertainties regarding the status
of these specimens, All of the characters fall well
within the ranges described for P bridgesi by Riley
& Self (1982).
Parasambonia sp. a
Material Examined
From lung of Austrelaps superbus (Gimther),
5 kim south of Bowral, New South Wales, in SAM
No, N1986192,
Description
Male (n = 1 plus 2 anterior ends), Length 8, with
51 annuli. Heads slide-mounted, outer hook with
finger-like extension, AD 175, 190; BC 95, 100.
Discussion
The outer hooks possess the projecting spine
characteristic of the genus Parasambonia, The low
annulus count is more characteristic of P minor
than P bridgesi, however the hook dimensions are
much smaller than those of A minor, the
dimensions of which do not overlap with the much
larger hooks of P. bridgesi (Riley & Self 1982). The
absence of fully gravid females precludes confident
specific identification. AR minor, but not RB bridgesi,
has been recorded from the copperhead (Riley &
Self 1982).
Parasambonia sp, b
Material Examined
From lung of Cryptephis nigrescens (Ginther),
Mogo 5.K, New South Wales, in SAM Noa
N1986191,
Description
Female (n 1). Length 29, annuli uncountable.
Outer hook with projecting spine, AD 465; BC 245,
Discussion
Hook dimensions in this gravid specimen are
larger than those reported in species of
Parasambonia from Australian snakes (Riley & Self
1982), suggesting that it represents a new species.
Additional specimens in good condition are
required to resolve this matter. The stomach of this
small-eyed snake contained a partly-digested eastern
water skink, Sphenomorphus quoyii (Duméril &
Bibron)- -
Waddycephalus spp. from shakes
Waddycephalus longicauda Riley & Self
Material Examined
From lung of Demansia psammophis (Schlegel),
Queensland National Parks and Wildlife Service,
Moggill, Queensland, in QM No. W12193.
Description
Female (n = 1). Length 27, with 49 pre-vaginal
and 7 post-vaginal annuli, preadult,
Male (n = 6). Length 8-11, with 56-59 annuli
(X = 57).
Discussion
The features of this material, particularly the long
and finely tapered post-vaginal tail, are
characteristic of HW longicauda (Riley & Self 1981b),
Waddycephalus sp, a
Material Examined
From lung of Pseudonaja lextilis (Duméril,
Bibron & Duméril) taken at Halls Gap, Grampians,
Victoria, in SAM No, N1980204.
Description
Female (n = 1). Length 37, with 63 pre-vaginal
and 4 post-vaginal annuli, Hooks removed from one
side, BC 530, 540; AD 860, 890.
Discussion
The only pentastomid described from PF textilis
is a female which was tentatively identified as
porphyriacus purely on the basis of similarities in
annulus counts: hooks were not measured (Riley &
Self 1981b). The present fully mature specimen has
far fewer pre-vaginal annuli (63 contra 75) than the
type series of H4 porphyriacus. Hook dimensions
are very much smaller than those of H¢
porphyriacus and similar to W. superbus from a
AUSTRALIAN PENTASTOMIDS i]
copperhead, Austrelaps superbus (Giimher), and
to an unnamed species from a mainland tiger snake,
Notevhis scutatus (Peters) (Riley & Self 1981b), Two
other characters, the number of annuli and the
attenuated caudal extremity, also place it close to
these species. However, the type series of HW.
superbus was denved from copperheads taken in
Tasmania and nearby islands (Riley & Self 1981b),
as was a tnore recently described infection (Riley
eral. 1985). To date, WH. superbus is recognised only
as an island species, although copperheads are
known on the mainland where their range overlaps
that ot F fextilis. Different dietary preferences have
been reported in these snakes (Cogger 1983)
however, the food habits of different populations
of a snake species over ils geographic range are not
well known. Recent evidence suggests strongly that
dietary preference is different in regions where food
resources are skewed or Limiting (Schwaner 1985),
Until more specimens become available the status
of the Haddycephalus from Pseydonaja textilis
remains uncertain,
Waddycephalus sp, bh
Material Examined
Froes lung of Notechiy ater atger Kinghorn,
Reevesby |, South Australis, in SAM No. NIS8SISI,
Description
Female {(n = 2). Both specimens in very poor
condition, length about 36-38, na detail of
annulation could be discerned. Hooks fram one
Specimen, BC 400, 410; AD 705, 690.
Discussion
The hooks are much smaller than ihase of all
recognised species of Waddycephalus except VW
Scusata, algo taken from a tiger snake, on St Francis
Island, South Australia, Riley & Self (1981b)
recorded the host of the type as Notechis scutata
(= scufaius), based on the collector’s label in the
vial. Cogger (1983) recognised N. sculatus as a
purely mainland species being replaced by N. afer
on islands off the coast of South Australia.
However, recent studies of morphological variation
in tiger snakes on Kangaroo Island have revealed
banded, unbanded, red-bellied and melanistic forms
believed fo belong Lo the same species complex
(Schwanec 1984), Thus W{ scufata js currently
recopnised as a parasite only of island populations
of the N. seu/atns/ater complex, its principal
distinguishing characteristic being its particularly
small hooks. Hook dimensions in the present
specimens form a cluster distinat from. Wt scwtata
(compare with Fig. 6 In Riley & Self 1981b and
below) and sugyestive of a new species of
Waddyeephalus.
Wiaddycephalus sp. c
Moterial Examined
From lung of Nolechis scutatus (Peters),
Grampians. Victoria, in SAM No, NISROI71.
Description
Female (n = 7), Length 32-43 (k = 38), with
60-63 pre-vaginal (x = 61.5) arid 2-3 (X — 3.2) past-
vaginal annuli.. Hooks taken from one side of 32
mim and 40 mm females, BC 495, 535; AD 885, 910
respectively.
Male (n = 4), Length 14-20 (R = 16), 62-68
annuli (X — 64),
Discussion
Jn their review of the genus Waddycephalus Riley
& Self (J98)b) separated the species from tiger
snakes (Wolechis spp.) into two groups distinguished
principally by marked differences in hook size:
small hooks are churacteristic of WH. seu/ata from
island populations of the Notechis scutatus/ater
complex whereas larget hooks are found in
specimens from mainland tiger snakes (see Fig. 6
in Riley & Self 1981b), ‘They concluded that
geographical isolation was responsible for the
observed differences and that two hosts may be
involved. The present findings substantiate these
differences and combine to suggest indeed that there
may be iwo species of Waddycephalus infecting
tiger snakes. More sophisticated diagnostic
techniques, preferably utilizing live matenal, are
required 10 confirm this postulate.
Waddycephalus superbus Riley & Self
Material Examined
From hing of Ausirelaps superbus (Giinther), (a)
Launceston, (b) Longford, Tasmania, in SAM Nos.
(a) N1980175, (by NI9BO205.
Description
Female fa) (n = 3), Length 37-41, with $9-63 pre-
vaginal and 3-4 post-vagiral annuli, Hooks from
one side of 40 mm specimen, BC $30, 530; AD 900,
970,
Female (6) (n = 2). Most of abdomens missing,
annuli uncountable; both apparently mature. Hooks
dissected from one side of both females, BC 525,
$25; AD 840, 840 respectively.
Male (b) (n = 1). Length 15, possibly 62 annus.
Discussion
These specimens are Very similar to the type serles
of HW! superbus (from the same host species also
taken in Tasmania) except that one specimen From
Launcesion has rwo more abdominal annuli and the
hooks are shightly larger than those described by
144 J. RILEY & D. M, SPRATT?
Riley & Self (J981b) and Riley er wl. (L985),
Nevertheless, all of the honk dimensions measured
to date combine to form a discrete cluster group
and this species at least, is now well characterized.
All specimens recovered thus far come [ram
Tasmania, endorsing the suggestion that
Waddycephalus teretiusculus Baird, 1862, the type
species of the (axon and also occurring in the
copperhead, |s prabably a maintand species (Riley
& Self 1981b, Riley ef a/. 1985). Lungs of specimens
of A. superbus held in the Australian National
Wildlife Collection were examined for
teretiusculus from the following mainland localities
(numbers of Specimens in parentheses) but
pentastomids were not recovered: Mi Gingera, ACT
(1), Ginini Plats, ACT (1); Captain’s Flat, NSW (3);
Pepper Creek on Big Badja Mountain via
Numeralla, NSW (1); Kosciusko National Park near
Kiandra entrance (1) and near Peak River, NSW (1);
Tumbarumba, NSW (1) Portland, Vic. (2); Flinders
1. (1).
Waddycephalus sp. d
Material Examined
From lung of Drysdalla coronoides (Giinther),
Fenelon !., South Australia, in SAM No, N1985152,
Description
Female (a = 1). Immature, slide-mounted.
Length 10, with 56 pre-vaginal and 5 post-vaginal
annuli. Hook measurements BC 280, 300; AD 430,
420.
Male (n = 1). Immature, slide-mounted, Length
9, with 64 annuli.
Discussion
The host snake was originally recorded as
Denisania coronoides but species of Drysdalia were
formerly Included in the genus Denisonia. The
immature state of the present specimens precludes
specific identification,
Waddycephalus punctulatus Riley & Self
Materiul Examined
From lung of Dendrelaphis puncrlata (Gray),
Northern Territory, in SAM Noa. N1984153.
Deseription
Female (n = 1). Length 33, with 52 prevaginal
and 11 posl-vaginal annuli,
Male (n 1), Length 14, possibly 4! annuli.
Discussion
There is no confusion concerning the status of
these specimens from the common tree snake; their
size and annulus number agree well with the original
description of WY punctulatus (Riley & Self 1981b),
Waddycephalus sp. e
Material Examined
Fram lung of Morelia spilota (Lacépéde}, Sa
Francis §., South Australia, in SAM No, N1985154,
Descriptian
Female (n = 1). Immature, length 9.5, with 52
pre-vaginal and 4 or 5 post-vaginal annuli,
Discussian
The anterior part of the cephalothorax, inchiding
the pair of inner hooks and the mouth are missing.
The outer hooks lack the projecting spine
characterisuic of Parusambonia spp. (Riley & Sell’
1982). The position of the vagina places the
specimen in the family Sambonidae and the annulus
count indicates that it is a species of Waddycephalus
hut, it has far fewer annuli than the immature
female described previously from the same host
species and tentatively identified as HW. porphyriacus
(Riley & Sell 1981b),
Nymphal Waddycephalus spp.
Material Examined
{i) from Parantechinus apicalis (Gray), locality
unknown, in SAM Ne, NI9B0210.
(li) Encysted in a skink, Hentierpis decresiensis
(Cuvier), South Australia, in SAM Na, (985155,
(ui) Eneysted in abdomen of Daspkalura
resatnondae Ride (a) Woodstock Station, (b)
Abydos Station, near Marble Bar, Western
Australia, in SAM Nos, (a) 1985156, (b) NI980182.
(iv) One nymph, from below posi-orbital skin of
a frog, Ranidella remota Tyler & Parker, Papua New
Guinea, in SAM No. N1985157.
(Vv) 14 nymphs encysted in intestinal connective
tissue af Cryplophis nigrescens (Giinther), Mogo
S.F, New South Wales, in SAM No, N1986190.
(vi) 3 nymphs eneysted in gecko, Heteronotia
binvei (Gray), Girraween National Park, Wyberba,
Queensland, in QM Na. W12194,
Description
(i) Three nymphs dissected from cysts and slide-
mounted. Length about 6, | specimen (sex
unknown) with 74 annuli, 1 male with 70 annuli.
All hooks double, overlain by accessory spine
(Fiz, 2) the base of which arises from a point
between fulcrum and hook, Spine an integral part
af hook and attached to fulcrum only by thin,
flexible sheet of cuticle, Overall hook length
(measurement as |llustrated in Fig, 2) 300-340,
(ii) Three nymphs dissected from cysts and slide
mounted, Length about 4-5, with 62-63 annuli, sex
indeterminable. Hooks double, overall length
210-250 (Fig. 1C).
AUSTRALIAN PENTASTOM(DS 145
(iia) All dissected from cysts and slide-miounted,
Length about 6, with 56-62 annuli (% = 60), Hooks
double, overall length 250-280,
(iiib) Six small cysts opened and nymphs slide-
mounted, large composite cyst containing many
larvae left intact, Length 4-5, wilh 56-59 annuli
(3 counted). Hooks double, overall length 230-260,
(iv) Male nymph, length approximately 4, with
56 annuli, Hooks double, not dissected and
measured.
(v) Three nyoiphs.slide-mounted. Length about
§, with 56-58 annuli (% = 57), Hooks dauble,
overall length 170-180.
(vi) Length about 5, with 56-61 annuli (% = 58).
Hooks double, overall length 195-215,
Discussion
Riley ef a/. (1985) ascribed a double-hooked male
nymph from the northern quoll, Satare/lus.
hallucatus (Gould), to the genus Waddycephalus
exclusively on the number of annuli, which vary
from 5$-78 in adult males of the genus (Riley &
Sell 1981b). This is considerably more than occurs
in the two other genera which may have double-
hooked larvae, Elenia and Parasambonia (Reymons
(929, Riley & Self 1982, Riley ef a/. 1985). All of
the present specimens are placed in the genus
Haddveephalus for the same reason, although
specific identification is nol possible.
The specimens from D. rosamondae probably
belong (0 the same species, those from Abydos
Station being at a slightly earlier stage of
development.
The nymph trom &. remota may be
punctulatus, This species was first described from
the common tree snake Dendrelaphis punctiulata
(Gray) in Australia (Riley & Self 1981) bul this host
also occurs in New Guinea (Cogger 1983), The prey
of tree snakes consist of frogs and birds, although
reptiles and small mammals are occasionally eaten
(Cogger 1983). Clearly frogs are probable
intermediate hosts of Ho punciulatus,
Either Waddycephalus or Elenia sp
Material Examined
One nymph, from submandibular lveophatic sac
of a frog, Palmatoreppia solamenis (Sternfeld)
Solomon. Islands, in SAM No, NL98SIS58..
Deseripiion
Male. Length 5, with 48 annuli, Hooks double,
overall length 220-230.
Discussion
‘The generic status of this nymph 1s uncertain.
Waddycephalus komodoensis and Wo radiata are
known from Indonesia (Riley & Self 198th) and
Elenia vitiensis is known from the tslands of Fiji
(Heymons 1932). The low annulus count of the
specimen may just preclude it being a species of
haddyeephalus, the lowest annulus number known
in mature males is 52, occurring in HW Kormodaensis
(Riley & Self 1981b),
Armillifer sp. fram snakes and rodents
Armillifer australis Riley & Self
Material Examined
(i) From yiscera (the specimens probably in-
habited the membranous lung which fs often
mistaken for the abdominal cavity) of a python
(species unknown), South Australia, in SAM Na.
N1980207,
(i) From lung of Morelia amethistina
(Schneider), Melbourne Zoo, in SAM No
N1980206,
Gili) From lung of Morelia spilota (Lacépéde),
Queensland, in SAM No, Ni980208.
(ivy) From lung of Morelia spiloia (Lacépéde),
Melbourne Zoo, in SAM No. N1I980172.
(v) Encysted nymph from body cavity of Ratius
lercopus cooktownensis Tate, Queensland, in SAM
No, N1980209,
Descriplion
(i) Female (n = 2) Length 63 and 67, both with
3L annuli and 2 incomplete annul on terminal
segment. Hooks removed from one side of one
female, AC 410; AD 624 (Fig. 1D), Male (n = 2),
Length 21 and 22, with 40 snnull; first 12 annuli
with pair of projections pointing backward from
posterior lateral angles.
(ii) Female (n = 15), Length 34-53 (one punctured
female not included) (X = 42.5), with 29-32 annuli
(% = 30,6) and 2 (or in two cases, 3) incomplete
armuli on terminal segment, Hooks from a 49 mm
specimen, AC 440; AD 635.
(iii) Female — mature (n = 1). Length 47, with
32 annuli (plus two incomplete segments
terminally). Female — immature fn = 4). Lensth
16-27 (X = 19), with 29-32 annuli (plus 2 incom-
plete). Male (n = 3). Length 16-17, with 36-37
annuli, anterior 1-12 bearing backward-pointing
projections,
(iv) Female (n = 1), Length 46, with 32 annuli
(plus 3 incomplete).
(v) Female?) (n » 1). Length 6, with 31 or 32
annuli. Hooks simple but could not be measured.
Discussian
Adult specimens from the four snakes are
vomistakably 4, australis and all characters accord
perfectly with those of the type series (Riley & Self
198la). Morelia spilota is a new host record,
146 J, RILEY & BM. SPRATT
Hooks of the nymph from R, /evcoprs are simple
(i.e, without an accessory spine) and (he annulus
count is within the range (29-35) of mature female
A, atistralis (Riley & Sell 19814), The present
specimen almost certainly belongs to this taxon, as
other species of Armillifer described from
Australian hosts have more annuli (Riley & Self
198la, Riley ef wil. 1985),
DiscUSSION
In our earlier review of pentastomid parasites in
Australian reptiles and mammals (Riley e/ a/, 1985)
we noted that the state of our knowledge of
taxonomy ts embryonic, and this is particularly true
of the genus Waddycephalus, Our original finding
of a double-hooked larva, which we attributed to
the genus Haddycephalus rather than Blenia solely
on the basis of the number of abdominal annuli,
was the first implication of mammals as
intermediate hosts in this genus.
This single male larva, from the mesentery of
Satanellus hallucatus was cleared and mounted (Fig.
1B) and we observed that the sharp spinous
extension overlying the hook appeared to be an
integral part of it and separate from the fulcrum.
This is unlike the situation in the related genus
Sambonia where the accessory spine is clearly an
extension of the fulcrum (Fain & Mortelmans 1960),
The relative abundance of nymphal Waddycephalus
material in the present study has permitted more
detailed observations of hook morphology and
these have confirmed our earller interpretation. The
spine is a functional part of the hook, 1b possesses
an apodeme, onto which muscles attach and extend
from it down into the fulcrum (contrast Figs IC and
D), The relative positions of the hook, its spinous
extension and the fulcrum sré presented diagram-
matically (Fig, 2).
The seven species of Maddycephalus currently
recognised in Australia infect boid, colubrid ane
elapld snakes (Riley & Self 19816) which prey upon
a variely of vertebraies (mostly frogs, lizards or
mammals — see Cogger 1983) and the present
report of Waddycephalus nyniphs from these three
classes of vertebrates is to be expected, particularly
since vertebrate intertriediate hosts are usital in
porocephalid life-cycles (Nicoli & Nicoli 1966), Also,
there is growing evidence from experimental
infections (Esslinger 1962, Vargas 1970, Winch &
Riley 1986), and from recoveries of nymphs in
intermediate hosts (Sachs, Rack & Woodtord 1973},
that the definitive number of annuli is present in
porocephalids by the infective stage. Our tentative
diagnoses are based on the assumption that this
occurs in the genus Waddycephalus. From the
viewpoint of host dietary regimen, itis equally likely
thal the related genera Parasambonia and Elenia
also utilize vertebrate intermediate hosts but in all
cases, experimental evidence of these life-cycles is
required,
ACKNOWLERGMENTS
We gratefully acknowledge che co-operation of Dr David
Lee who arranged loan of specimens from the South
Australian Museum, Mrs F, Walter who examined lungs
of mainland copperheads for pentastomids, Br Terry
Schwaner who read an earlier draft of the manuscript and
offered! valuable comment on the biology of snake species,
and Dr (an Beveridge who collected many of the speciinens
and ¢riticized an earlier draft of this work.
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the taxonomy of Ruillictiella boulengeri (Vaney &
Sambon, 1910) Sambon, (410, 8. erientalis{Hett, 1915)
Sambon, 1922 and R. weca/ Tubangul & Masilungan,
1956 (Pentastomida: Cephalobaenida). Sys/. Parasitol.
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ALL J. H., RILEY, J, & SELB, J.T) 4984. Further
observations Of bilint-hooked raijlietielNds
(Pentasiomida: Cephatobaenidal from lizards, with
descriptions of (hree new species. Syst. Puresifol. tr
(47-160.
ALI, J. H., RILEY, J, & SELF, fT, 1985. A review of
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ARCHER, M_ 1982. Review of the dasyurid (Marsupialia)
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Archer (Bu.). Carnivorous Marsupials’, Volume 2. R.
Zool. Sac N.SW., Sydney. Pp. 297-343.
COGGER, H. G. 1983 Reptiles and Amphibians af
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caperimental intermediate fiosts, 4 Parasitol, 48;
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le cycle evolulil de Sambonio fohrimanni chez le -varan.
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46: 518-541.
AUSTRALIAN PENTASTOMIDS 147
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Z. wiss. Biol. 4: 409-430,
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barbatus (Cuv). Proc. Zool. Soc. Lond. 124: 509-516,
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pentastomides. Annis Parasit, hum. comp. 41: 255-277.
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Australian snakes. Syst. Parasitol. 2: 171-179.
RILEY, J. & SELF, J. T. 198ib. A redescription of
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LOOKING FOR DITJI-MINGKA
BY L. A. HERCUS
Summary
Ditji-mingka was an important Aboriginal site in the north-east of South Australia. The present
paper contains a Wangkangurru text describing what has happened to this site.
LOOKING FOR DITJI-MINGKA
L. A. HERCUS
L. A. HERCUS 1987. Looking for Ditji-mingka. Res. S. Aust. Mus. 21(2): 149-156,
Ditji-mingka was an important Aboriginal site in the north-east of South Australia. The present
paper contains a Wangkangurru text describing what has happened to this site.
L. A. Hercus, Faculty of Asian Studies, Australian National University, P.O. Box 4, Canberra,
Australian Capital Territory 2600. Manuscript received 13 May 1985.
Ditji-mingka, ‘Sun-Cave’ (lit. ‘Sun-Hole’), south-
west of Etadunna on the Birdsville Track in north-
eastern South Australia, was one of the most
important places in Diyari mythology. Evidence of
this comes from J. G. Reuther who worked for
eighteen years (1888-1906) as a missionary amongst
Diyari and neighbouring people at Killalpaninna
on the lower Cooper. There was nobody who could
rival the achievements of the Reverend Reuther in
documenting the languages and traditions of the
Lake Eyre Basin. Without his great work of
fourteen volumes, recently translated by P, Scherer
(Reuther 1981), much information would be totally
lost. He gives the following account of the Ditji-
mingka site in his list of place-names.
Ditji = ‘sun’; minka = ‘cave’.
‘There is a cave at this spot, where the female sun
muramura is said to have first risen. But since she made
conditions too hot for her followers (‘people’) she
wandered inside the earth towards the east and has been
rising there ever since’ (Reuther 1981; VII: 29).
There are two expanded accounts of the Ditji-
mingka story given by J. G. Reuther (1981, VIII:
20 and X: 20). In the latter he describes how Ditji,
the Sun, was one of the two wives of the Wild
Onion Ancestor ‘Jelkabalubaluna’ (i.e. Yalka parlu-
parlunha, ‘a wild Onion peeled’); Ditji crawled into
the earth at Ditji-mingka. He makes other
references to the site; the most telling of these is
in XI: 163 where he describes ‘tthe enchantment of
the sun’, ie. a form of magic for creating hot
weather. This magic was carried out with the use
of ‘ditjipupara’ (earth from Ditji-mingka). Reuther
adds: ‘Everyone who is a devotee of the female
muramura, Ditji, has some of it in his wurley’ (1981,
XI: 163).
There can thus be no question about the
importance of the site. The Reverend Reuther —
as is clear from Wangkangurru evidence — usually
gives a ‘cleaned up’ version of mythological
traditions. We cannot be certain whether he himself
censored the myths, or whether Aboriginal people,
who respected him, were reluctant to tell him things
they knew would offend him. His accounts are dull
and humourless: all the spice has been taken out.
There was certainly more to the story of Ditji-
mingka than he implies, and there were associated
rituals. The police trooper Samuel Gason was the
first to publish details of Diyari traditions. He was
not a missionary, but after all, he lived in the
Victorian era. He was clearly shocked by the Sun
rituals and gives the following version:
‘Their traditions suppose that man and all other beings
were created by the moon, at the bidding of the
Mooramoora. (This term simply means “Ancestor”).
Finding the emu pleasant to the sight, and judging it
to be eatable (but unable, owing to its swiftness to catch
it during the cold that then prevailed), the Mooramoora
was appealed to to cast some heat on the earth so as
to enable them to run down the desired bird. The
Mooramoora, complying with their request, bade them
perform certain ceremonies (yet observed, but too
obscene to be described), and then created the sun’
(Gason 1879: 260).
Other versions of the sun-legend, all involving
Ditji-mingka, the cave of the sun, are given by
Siebert (1910: 44-45) and by Howitt (1904: 427),
Aiston gives yet another, different account of the
Sun History, in which the Sun Ancestor is regarded
as male. The cave however still figures prominently:
‘The moora, however, escaped, but so annoyed was he,
that he sank and sank, until at last, at the place now
called in memory of the event Ditchaminka, he plunged
into the ground. To this day is shown the hole in the
stony plain where the sun disappeared’ (Horn & Aiston
1924:131).
Diyari rituals gradually fell into oblivion because
of the disintegration of the group and because of
pressure against ‘paganism’ from the missionaries.
Nevertheless the myth and the knowledge of the
Ditji-mingka site did not immediately fade from the
minds of Diyari people. H. K. Fry (1937: 189-194)
published a brief Diyari text, with English
translation, of a myth called ‘Ditji-mingka Mura
and Pinja, Sun Cave Mura Revenge’. Apart from
the name in the title, this text does not actually
mention that the female Sun Ancestor is involved.
There can however be no doubt that it is an
150 1. A. HERCUS
uncensored version of the same myth as in Reuther
(1981, X: 20): the Old Woman in Fry’s story, like
Reuther’s ‘Ditji’, is one of the two wives of the Wild
Onion Man ‘Jelkapalupaluna’. This text was given
to Fry and Vogelsang by an old Diyari man ‘Sam’
Dintibana Kinjmilina, who owned the myth at that
lime, in the mid-1930s, Sam also told Fry that he
had a ‘sun-bag’ filled with light blue earth from
Ditji-mingka which he claimed could be used to
‘sing the sun and make it very hot’ (Fry 1937: 193).
[t is of the nature of cult-heroes that they may
be larger and more extreme than the ordinary things
of life. The female Sun Ancestor and her older sister
in Sam Dintibana’s story certainly were: they are
depicted as grotesque and fearsome old women who
got their husband ‘Jelkapalupaluna’ killed, cut him
into small pieces and mutilated the body, and then
mutilated themselves. The Diyari text only gives us
an outline, but we can begin to imagine why the
full story does not appear in Reuther’s work and
why Gason was shocked. The important matter is
that despite the missionary disapproval of earlier
years, the myth was still alive in the 1930s in Sam
Dintibana’s mind. Even after Sam’s death the
knowledge that Ditji-mingka was a special place,
a ritual centre, lingered on and the older people with
Diyari associations had a deep affection for it. This
became clear to me over the years that I was
travelling on the Birdsville track for language work
at both Marree and Birdsville.
Ditji-mingka was often mentioned by Mick
McLean Jrinjili, by Jimmy Russell Wanga-mirri
(‘Many mornings’), and also by Ben Murray Palku-
nguyu (‘One mass of clouds’). Whenever we headed
north on the Birdsville track we would pass Blaze’s
Well, a well which is now totally silted up. It was
in a small depression which they called Thidna-
kurduni [‘Making a (deep) Footprint’) because it
was there that Ditji, the Sun Ancestor, stepped into
muddy ground while looking for wild onions. As
we continued north towards Cannuwaukaninna
Bore those older men would point nostalgically
towards the west, saying; ‘Over there is Ditji-
mingka’, Mick and Jimmy were Wangkangurru,
Ben's mother was Arabana, but they had all been
associated with Diyari people (Hercus 1980, 1986;
Austin 1981), They had learnt from them to admire
the cave of the sun and were anxious to see it again.
They often described it: it was in one of the more
inhospitable areas on Etadunna station, in rough
country. [t was high up on the western slope of the
ridge that faces the bed of the Ditji-mingka creek
as it nears Lake Palankarinna. There was a soakage
in the creek nearby and the old people used to camp
there for their ceremonies (see Fig. 1).
The walls and the ceiling of the cave were of
glistening gypsum which reflected the last rays of
the setting sun: it was the home of the sun. There
were rock-carvings inside, circles which symbolised
the sun; Jimmy Russell had seen them. The two
PIGURE 1. The edge of the bed of the Ditji-mingka Creek, where the soakage used Lo be. (Photo B. Jeffery.)
LOOKING FOR DITJI-MINGKA 151
cave-openings were slightly upward; this meant that
occasionally animals, particularly feral goats, fell
in and died, not being able to get out again. It was
a haven for snakes and this irritated the owners of
nearby Etadunna station. Mick McLean and Jimmy
Russell spoke of the cave so often that I wrote letters
to have it protected; Ditji-mingka was truly a site
of significance. Moreover it was only a few miles
from Lake Palankarinna which was a declared
geological reserve because of important fossil finds
(Rich, van Tets & Knight 1985: 46 ff).
We were determined to see Ditji-mingka and tried
a number of times in the early 1970s. Mick McLean
was then well over eighty years old, he could not
scramble up over the ridge, and on our own we
failed to find it despite his instructions. Next time
in August 1974, Jimmy Russell came to help, he was
only in his seventies and his sense of direction was
uncanny, like Mick’s. He was deeply ashamed,
because he too could not find it. Later of course
it became clear why! Finally in June 1976 we had
quite a group of people, which included Jimmy
Russell and Ben Murray, as well as members of the
South Australian Government’s Aboriginal Heritage
Unit, along with linguists Tamsin Donaldson and
Peter Austin who was then writing his grammar of
Diyari (Austin 1981). We again went looking for
Ditji-mingka (see Fig. 2). Later in camp Jimmy
Russell spoke in Wangkangurru about what
happened:!
Text
1. thika-rna arniri wadna-ya-rna Ttatinha yadla-ku
Come-IMP we run -SP-IMP Etadunna close-DAT
kari-rnda uta arniri yani-ngura mudlu-nga-thu arniri
see-PRES now we say-CONT sandhill-LOC-EMPH we
marrili tharni-thika-Ihuku Ditji-mingka-ruku mingka
this side eat-return-PURP Ditji-mingka-ALL cave
nhanhi-lhiku.
see -PURP.
FIGURE 2. Ben Murray (left) and Jimmy Russell (right) setting out to look for Ditji-mingka. (Photo B. Jeffery.)
152
2. wantali
Separately
yurrakati-nga
west side-LOC
intja Ditji-mingka?
Where Ditji-mingka?
L. A. HERCUS
thika-lhuku arniri
come-HIST we
yuka-lhuku.
go -HIST.
3. arniri yuka-ka partjarna wadlhu yurrakati-nga
We go -PAST all place west side -LOC
thika-lhuku arniri, wadlhu pidla Thita-pulumanha.
return-HIST we, place name Thita-pulumanha,
4. uta arniri partjarna karla-nga pathara midla-nga,
Now we all creek-LOC box tree nose-LOC,
thidna-ra -ki yuka-rna mingka-thu wapa-rna thika-rna
foot-CAUS-EMPH go-IMP cave-EMPH seek-IMP come-IMP
thadna-rna, wadni-rnda watungunta nguthi thika-lhuku.
leave-IMP, follow-PRES rest reverse come-HIST.
5. nhararda waru katha-liparna, ipali katha-rna,
Here long ago walk-ANC, before walk-IMP,
malka-thu mingka nhanhi, Ditji-mingka.
not-EMP cave see, Ditji-mingka.
kutha irtjirtja thuntiripa-rna kutha kathiwiRi-ri
Water soakage cover over-IMP water big -ERG
tjiRi-ri.
flood-ERG,
padni-li punga nhanhi-ra, thanpi-liparna thiRi-ri
Not-ADV humpy see-PUNC, knock down-ANC flood-ERG
wanpa-rna.
carry -IMP.
6. kayi kadnha awukinta kadnha arniri wadna-yi-rnda.
Here rock this-ALL tock we run-TR-PRES.
yadla witji-yangu, ayi! tjarlpa tharka-tharka-rnda
close become-PLUP, hey! tree stand-stand-PRES
nhararda yada...
here close..
7. thanpi-thanpi-la-rda
Destroy-destroy-ALT-PRES
Ditji-mingka!
Ditji-mingka!
LOOKING FOR DITJI-MINGKA
& L, What was it like before?
J. ngurka arla, parluru, antha iparli katha-nangka-rda
Good true, smooth. I before travel-CONT S-PRES
marna parkulu nhanhi-ka neurku-nhaku! kanhangarda
mouth two see-PAST good-EMPH! There
mingka tharka-tharka-rnda mingka-rda.
cave stand-stand-PRES gape-PRES.
9. antha nguyu katha-liparna nhantu-ra, marna-nga
I alone travel-ANC horse-CAUS, mouth-LOC
tharka-rnda. mintjt-mintji-rnda, muyu katinari yantakara
stand-PRESS. Shine-shine-PRES, sun beyond west
muyu mintjiyva-ra.
sun shine-TR-PUNC.
10. malka antha wintaku-ra, antha nguyu-nguyu.
Not I go in-PRES, I alone-alone.
kudju-ru pirda-lira nguyu-nguyu mingka-nga.
kurdaitcha-ERG kill-LEST alone-alone cave-LOC,
11. antha thadla-ra waya-rna, thadla nhinka-rna-li.
1 fear-CAUS want-IMP, frightened squint-IMP-EMPH.
mintji-mintji yalkiri-ri. muy round one.
shine-shine kopi-INST. Sun.
12. thanpi-thanpi-rda, partjarna thanpi-la-rda
Knock down-PRES, all desiroy-ALT-PRES
parluru-ku
level-DAT.
Translation
1. We came back (from looking at the lower Cooper)
and drove in close to Etadunna. We had a look around
and then started saying ‘we'll have our lunch this side
of the sandhill and then go to Ditji-mingka to have a
look at the cave.”
2. (We split up) and walked separately coming and going
over the west side of the sandhill. Where is Ditji-
mingka?
3. We went all over the country on the west side and
back again, that area is called Thita-pulumanha.
4. Then we all walked around in the creek bed, where
the box-trees run out to a point. We were going about
on foot, looking for the cave, we went and turned back
again, we left, we followed the others but they came
back as if they had got to a dead end.
5. ] used to travel around here long ago, but I can’t see
that cave, Ditji-mingka, The soakage had been washed
out by big rains, by flood-waters. [ couldn't see the
humpies (that had been there), they must have been
knocked down and carried away by the floods a long
time ago.
6. ‘Ah, here is that rock!!’ (I said). So we all ran straight
over towards the rock. We got near: ‘Ah, that tree is
standing over there, we’re really close!”
7, They have blown up Ditji-mingka!
8. L. What was it like before’?
J. It was lovely, all smooth.
When I was travelling about a long time ago I saw the
two openings of the cave. It was beautiful. I stood there
for a while just looking at the cave.
9. |] was travellng about alone on my horse, and I stood
in the mouth of the cave. It was gleaming and glistening,
the sun was shining in from the west.
10. I didn’t go in, | was on my own, (I thought) a
kurdaitcha [revenge killer] might come and kill me if
I was all alone in that cave.
154 L, A, HERCUS
11. | was terrified. I peered in, very frightened. It was
glistening with kopi. There was (engravings of) the sun
there, round ones.
12. It has been destroyed, completely blown apart (with
dynamite), razed to the ground! (see Fig, 3).
CONCLUSION
In 1879 Gason, as was typical of the period, had
a very restricted view of Aboriginal religion, when,
as quoted above he states with obvious lack of
enthusiasm: ‘Their traditions suppose that man and
all other beings were created by the moon, at the
bidding of the Mooramoora’ (Gason op, cil.), We
tend to take an equally simplistic view nowadays
when we identify sites on a one to one basis with
traditions: we are often told for instance in popular
reporting that the destruction of a particular tree
might result in the Aboriginal myth associated with
it being forgotten. There is more to traditional
mythology than that. This has often been said, and
it was expressed with special clarity in 1965 by
W.E.H. Stanner in his well-known article
‘Religion, Totemism and Symbolism’, where he
shows the subtlety and intricacy of Aboriginal
religious thought. Speaking of the totemic symbol-
function he states that it has: ‘four elements (i) living
men (totemists) serving as interpreters of (ii) signs
(totems and totem-places), by using (iii) vehicles that
form and express affective conceptions of (iy) sign-
objects, which are the significance of the Dream
Time marvels’ (Stanner 1965: 228).
If we look at these elements with regard to the
Diyari Sun Myth we can see that the first to be
impaired was (iii), in that the ritual, the dances, the
songs and the detail of the stories fell into oblivion,
Therefore (iv), the deeper symbolism, the marvel
of the Dream Time Sun Myth has disappeared for
ever, All we are left with is the bare outline of the
story. Until recently we still had (ii) the totem-place.
The situation was no worse than for much of the
mythology of eastern Australia, where we usually
just have minimal stories and the totem place.
Although we lacked the vehicle of interpretation
provided by the chants and dances, we might still
have had a glimpse of one aspect of the marvels
of the Dream Time through the engraving and
through the appearance of the cave, but this has
now gone through the destruction caused by
European activity. This leaves us only with (i), the
living men. For many years, since the death of Sam’
Dintibana, there has not been anyone who
‘belonged’ to the Sun Myth and identified with it.
There are only two people living to whom the
mythology of Ditji-mingka and the surrounding
areas still means something: they are Jimmy Russell
who has suffered a major stroke and Ben Murray
who is now in his nineties. Therefore it is sadly true
that soon there will be no one who can stop on the
Birdsville Track at the right spot south of Etadunna,
look across to the west, and visualise that not far
away is Lake Palankarinna- an expanse of white
fading away into the distance into an endless plain
(see Fig. 4). No one will remember the humour of
the story of the wicked Old Man Markanjangkurla
who chased the Seven Sisters across the lake and
across the plain beyond and recall what he did there,
FIGURE 3. The heap of rubble that used to be Ditjimingka. (Photo B. Jeffery.)
LOOKING FOR DITJI-MINGKA 155
FIGURE 4. Looking towards Lake Palankarinna from near Ditji-mingka. (Photo B. Jeffery.)
(Lake Palankarinna, which Europeans know mainly
as a fossil reserve, is really parla-ng-kari-nha, a
crude name, since parla means ‘semen’ and kari
means ‘to chase’). Very soon there will be no one
that can even visualise the cave south of Lake
Palankarinna and point and say: ‘Over there was
Ditji-mingka’. We have lost the Sun Myth with all
its symbolism.
ENDNOTES
1. The text transcribed in this paper was recorded in June
1976 as Hercus field-tape 739, a copy of which has been
deposited with the Australian Institute of Aboriginal
Studies, Canberra. For ease of reference the text has been
split into numbered sections. The divisions are on the
whole in accordance with intervals in speech, In this paper
a practical orthography has been used for Wangkangurru:
Plosive consonants other than the retroflex plosive have
been written as unvoiced (k, p, th, ¢), but prestopped
consonants have been written with voiced plosives as this
corresponds most closely to the pronunciation, hence bm,
dn, dnh, dnij, dl, dth.
Retroflexes have been written as r + consonant, i.e.
rl is retroflex /
rn is retroflex n
rd is retroflex f
Interdentals have been written as consonant + A, hence
nh, th, lh,
Palatals have been written as consonant +/, hence ¢/, nj, //.
ng has been used for velar 7.
The three r-sounds have been transcribed as follows:
r = the alveolar flap
rr = the trilled r
R = retroflex r.
The following abbreviations have been used for linguistic
terms in the interlinear gloss:
ABL Ablative case
ACC accusative case
ACT active stem-forming suffix
ADV adverbial suffix
ALL allative case
ALT altruistic stem-forming suffix
ANC ancient past
CAUS causative case
CONT continuous participle
CONT $ continuous stem-forming suffix
DIST aspect showing distance
EMPH emphatic clitic
ERG ergative case
EXCL exclusive pronoun
HAB habitual aspect
HIST historical past
IMP imperfective
LOC locative case
NAR narrative past
PAST past tense
PERF perfect
PLUP pluperfect
POS possessive suffix
PRES present tense
PUNC punctiliar present
PURP purposive
SP speed form, indicating action
undertaken before departing
TR transitory aspect
156 L, A. HERCUS
REFERENCES
AUSTIN, P. 1981. ‘A Grammar of Diyari, South Australia.’
Cambridge University Press, Cambridge.
FRY, H. K. 1937 Dieri legends. Folklore 48: 187-206;
269-287.
GASON, S. 1879. The Manners and Customs of the
Dieyerie Tribe of Australian Aborigines, Jn J. D. Woods
(Ed.). ‘The Native Tribes of South Australia’. Pp, 258-306.
E. S. Wigg, Adelaide.
HERCUS, L. A. 1980. How we danced the Mudlunga.
Aboriginal History 4: 5-31.
HERCUS, L. A. 1985, Leaving the Simpson Desert.
Aboriginal History 9: 22-43.
HORNE, G. A. & AISTON, G. 1924, Savage Life in
Central Australia’. Macmillan, London,
HOWITT, A. W. 1904. ‘The Native Tribes of Southeast
Australia’. Macmillan, London.
HOWITT, A. W. & SIEBERT, O. 1904. Legends of the
Dieri and kindred tribes of Central Australia. J. R. Anth.
Inst. 34: 100-129.
REUTHER, J. G, 1981. The Diari’, translated by P.
Scherer. Microfiche edition, Australian Institute of
Aboriginal Studies, Canberra.
RICH, P. V., VAN TETS, G. F. & KNIGHT, F. 1985.
‘Kadimakara’, Pioneer Design Studio, Melbourne.
SIEBERT, O. 1910. Sagen and Sitten der Dieri und
Nachbarstaemme in Zentral Australien. Globus 97: 44-50;
53-9.
STANNER, W. E. H. 1965. Religion, totemism and
symbolism. Jn R. M. & C. H. Berndt (Eds). ‘Aboriginal
Man in Australia’. Pp. 207-237. Angus & Robertson,
Sydney.
THE SCALED-SQUID LEPIDOTEUTHIS GRIMALDII JOUBIN FROM
SOUTH AUSTRALIAN WATERS
BY W. ZEIDLER
Summary
The first substantiated record of Lepidoteuthis grimaldii from Australian waters (i.e. within the 200
nautical mile fishing zone) was by Lu & Phillips (1985) but they gave no details of specimens. The
purpose of this paper is to provide more details of these specimens, in particular of one in the South
Australian Museum (SAM) which was not seen by Lu & Phillips, and thus highlight this interesting
record from Australian waters.
THE SCALED-SQUID LEPIDOTEUTHIS GRIMALDIT JOURIN
FROM SOUTHERN AUSTRALIAN WATERS
The first substantiated record of Lepidereuris
arimalaii from Australlaiy waters (1G within the 20)
nautical mile fishing zone) was by Lu & Phillips
(1985) bur they gave no details of specimens, The
purpose of this paper is to provide more details of
these specimens, in particular of one in the South
Australian Museum (SAM) Which was not seen by
Lu & Phillips, and thus highlight this interesting
record from Australian waters,
Lepidoteuthis grimaldil was first deseribed by
Joubin (1895) from two mantles froin a sperm
whale’s stomach and from a fragment of @ Risso’s
dolphin, both caught off the Azores, However, up
until 1960 there had been only four records of this
species and only two included the head (Clarke
1960) and a complete description was not available
until 1962 (Clarke & Maul 1962), Since then the
species has been recorded from most of the world's
oceans, North and South Atlantic (Clarke 1966),
Indian Ocean (Clarke 1980), Pacifie Ocean between
the New Hebrides and New Caledonia (Rancurel
1970), Pacific Ocean olf Japan (Okutam ef al.
1976), Tasman Sea (Clarke & Macleod 1982) and
Southern QOvean (Lu & Phillips 1985)-
Complete specimens are still rare and apart fram
juveniles caught in nets (Clarke 1964 & 1980, Lu
& Clarke 1975, Roper & Young 1975) and the reoord
of Lu & Phillips (1985) all specimens have been
obtained {rom the stomachs of predators, ovainly
the sperm Whale, Plivsefer catoden, but also fran
Rissa’s dolphin, Granyus griseus (Joubin 1895); the
taneet fish, Alepisaurus ferox (Rancurel 1970}; the
black-scabbard lish, 4phanopus curbo (Clarke &
Maul 1962) and the tuna, Gera ahesus (Clarke
& Maul 1962),
The first published evidence that 1, grrmvaldu
might oecur in Australian waters was provided by
Clarke (1980) who recorded the buccal mass of two
specimens from the stomachs of sperm whales
caught by whaling, Ships Operating out of Albany,
Wesiern Australia, Clarke & MacLeod (1982) also
recorded the remains of specimens from the
stomachs of sperm whales killed ii the Tasman Sea
between 3305, 172°R and 40S, 155°R but this is
at least 550 km south-east of Cane Huwe, eastern
Australia, The Australian specimens referred to by
Lu & Phillips (1985) and the one iy SAM are
noteworthy in (hat they are the first records of
adulls from other than predators’ stomachs und
indicate that the species oceurs relatively close in-
shore along the south coast well within teach of
commervial crawlers.
Derails ol speciinens are as follows:
1. Male, 122 mm dorsal nant leneth, 97 km cast
olf Broken Bay, New South Wales (33° 28'S, [52°
43’ E), depth 0-1000 m, Engel mid-water trawl,
FRY ‘Kapala’, J. Paxton, 14 December 1977
(Australian Museum, Sydney AM, C111782).
2. Sex undetermined (viscera missing), 753 mm
mantle Jength, approx. 30 km south-west off
Beachport, South Australia, trawled in 550 m by
‘Margaret Phillipa’, 6-10 September 1982
(Museum of Victoria MY, F53159),
3. Sex undetermined {viscera decayed), 790 mm
dorsal mantle length, approx, 40 km south-west
off Beachport, South Australia, trawled in 220 m,
obtained fresh from fish processor in
Portland, Victoria by W. Zeidler, 22 October 198t
(SAM, DI7589),
Specimen I, a juvenile, is in relatively good
condition and only the tips of the arms are missing.
Generally it agrees with the description of young
stages given by Clarke (1964) and some body
measurements are given if Table |.
TABLE |. Lemdoteuthis erimaldii body measurements,
Character Measurement (mm)
Specimen | Specimen 3
Manele length (darsal) 122 ‘790
Mantle length (Ventral) 115 740)
Mantle width (max) 26 210
Fie length 60 410
Fin widih tmax) sy 240
Gladius length - 790
Gladius width (max) - 3
Rachis length - ang
Rachis width (max) - 28
Max. width of scales 14 Ww
Specimen 2
measurement®
Specimen 3 (Pig. 1) when collected was in good
condition with only the tips al the arms missing.
However, it was inadvertently left out of the freezer
and deteriorated considerably before being
measured and preserved, The head js too damaged
for accurate measirement, olher bady
measurements (Table 1) are according ro Roper &
Voss (1983) and beak dimensions (Table 2) are
according to Wolff (1984), Some measurements arc
inaccurate due ta the damaged nature of the
specinien a. the fins are contracted, dorsal mantle
length probably is longer as Ihe up oF the tail fs
damaped and abou! 70 mm is missing and the
ntantle is probably not as wide when all internal
is too damaged for accurate
158 W. ZEIDLER
FIGURE 1. Ventral view of Lepidoreuthis grimaldi, SAM,
D. 17589.
organs are intact. The beaks (Fig. 2a-c), radula (Fig.
2d) and gladius have been adequately described for
this species by Clarke & Maul (1962) and the SAM
specimen does not differ from that description.
TABLE 2, Lepidoteuthis erimaldii (SAM, D17589) beak
dimensions,
Character Measurement (mm)
Upper
Hood length 39.5
Rostral length 17.6
Wing width 9.5
Rostral tip to inner margin of wing 26.0
Wing to crest length 43.2
Crest length 59.6
Jaw angle width 10.0
Lower
Rostral tip to inner posterior corner of
lateral wall 41,2
Rostral length 17.3
Rostral tip to inner margin of wing 35.2
Wing length 18.3
Jaw angle width 93
ACKNOWLEDGMENTS
| wish to thank Tan Loch, The Australian Museum, for
the loan of Specimen 1, Dr C, C, Lu, Museum of Victoria,
for information on the MV specimen and for his
constructive criticism of the manuscript and John Glover,
SAM, for providing common names of fish predators. The
photographic expertise of Roman Ruchle (Fig. 1) and Jan
Forrest (Figs 2 and 3), both of SAM, is also gratefully
acknowledged.
REFERENCES
CLARKE, M. R. 1960. Lepidoteuthis grimaldii — a squid
with scales. Nature, Land. 188: 955-956.
CLARKE, M. R. 1964. Young stages of Lepidoteuthis
grimaldii (Cephalopoda, Decapoda). Proc. Malac. Soc.
Lond. 36: 69-78.
CLARKE, M. R. 1966, A review of the systematics and
ecology of oceanic squids. Jn FE. S, Russell (Ed,),
‘Advances in Marine Biology’, Vol. 4. Academic Press,
London and New York. Pp. 91-300.
CLARKE, M. R. 1980, Cephalopods in the diet of sperm
whales of the southern hemisphere and their bearing
on sperm whale biology, Discovery Rep, 37: 1-324,
CLARKE, M, R, & MACLEOD, N. 1982. Cephalopod
remains from the stomachs of sperm whales caught in
the Tasman Sea. Mem. Nain. Mus. Viet. 43: 25-42.
CLARKE, M. R. & MAUL, G, E. 1962. A description
of the ‘scaled’ squid Lepidoteuthis grimaldii Joubin
1895, Proc. Zool. Soe. Lond. 139: 97-118.
JOUBIN, L, 1895, Céphalopodes recucillis dans l’estomac
d’un Cachelot capture aux iles Acores. CR. Acad. Sci.
Paris 121: 1172.
THE SCALED-SQUID LEPIDOTEUTHIS GRIMALDI JOUBIN 159
LU, C. C. & CLARKE, M. R. 1975. Vertical distribution
of cephalopods at 11°N, 20°W in the North Atlantic.
J. mar. biol. Ass. U.K. 55: 369-389.
LU, C. C. & PHILLIPS, J. U. 1985. An annotated checklist
of the Cephalopoda from Australian waters. Occ. Pap.
Mus. Vict, 2: 21-36.
OKUTANI, T., SATAKI, Y., OHSUMI, S. &
KAWAKAMI, T. 1976. Squids eaten by sperm whales
caught off Joban District, Japan, during
January-February. Bull. Tokai, reg. Fish. Res. Lab. 87:
67-113.
RANCUREL, P., 1970. Les contenus stomacaux
d’Alepisaurus ferox dans le sud-ouest Pacifique
(Cephalopodes). Cah. ORSTOM sér Oceanogr. 8: 4-87.
ROPER, C. F. E. & YOUNG, R. E. 1975. Vertical
distribution of pelagic cephalopods. Smithson. Contrib.
Zool. No. 209: 1-51.
ROPER, C. F. E. & VOSS, G. L. 1983. Guidelines for
taxonomic descriptions of cephalopod species. Mem.
Natn. Mus. Vict. 44: 49-63.
WOLEFE, G. A. 1984. Identification and estimation of size
from the beaks of 18 species of cephalopods from the
Pacific Ocean. NOAA Tech. Rep. NMFS 17: 1-50.
W. ZEIDLER, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Rec.
S. Aust. Mus. 21(2): 157-159.
FIGURE 2. Lepidoteuthis grimaldii (SAM, D17589); a. Upper beak; b, c. Lower beak; d. Radula.
REVIEW : FROM HORIZONTAL TO PERPENDICULAR : TWO RECENT
BOOKS ON CENTRAL AUSTRALIAN ABORIGINAL PAINTING
BY P. SUTTON
Summary
Albert Namatjira, the life and work of an Australian painter by Nadine Amadio, Anne Blackwell,
Jonah Jones & Daniel Thomas. Macmillan, Melbourne, 1986. ix + 102 pp. illus. Cloth only.
$A29.95.
Dot and circle : a retrospective of the Aboriginal acrylic paintings of Central Australia edited by
Janet Maughan & Jenny Zimmer. Royal Melbourne Institute of Technology Communications
Services Unit, Melbourne, 1986. 207 pp. illus. Paper only. $A20.00
REVIEW
FROM WORIZONTAL TO PERPENDICULAR:
TWO RECENT BOOKS ON CENTRAL AUSTRALIAN ABORIGINAL PAINTING
Albert Namatjira, the Life and Work of an
Australian Painter by Nadine Amadio, Anne
Blackwell, Jonah Jones & Damel Thomas,
Macmillan, Melbourne, 1986, ix + 102 pp. Illus.
Cloth only, $429.98.
Dot and Circle: a Retrospective of (he Aboriginal
Acrylic Paintings of Central Australia edited by
Janet Maughan & Jenny Zimmer. Royal Melbourne
Institute of Technology Communications Services
Unit, Melbourne, (986. 207 pp- illus. Paper only.
$A 20,00,
“The interaction between the European and
Aboriginal artistic tradulons can produce a
renaissance potentially as significant (or Australian
life as that which was laifiched Upon Europe by the
spread of the “new knowledge” from Constan-
tinople in the sixteenth century’. With these
dramatic words Dr H.C. Coombs launches his
introduction to the most recent book on the wark
of Albert Namaliira, compiled and edited by
Nadine Amadio, arts editor of Sydney's Sunduy
Telegraph. De Coombs is certainly right ta point
out that Aboriginal graphic.and sculptural imagery
conslilules a distinctive body of sources for the arts
in Australia generally, allhough their role thug far
might more aptly be likened to that of African and
Oceanie works in the rise of twentieth century
primitivism in Western Eupopean art, than to that
of the manuscripts of Constantinople cirea 1453.
The Renaissance was, among olher things, a
scholarly revolution. By contrast, Picasso was-able
to remark, in effect: “Everything | need to know
about Africa is in those objects’ (Rubin 1984: 74).
However, Albert Namatjira, on the face of it, is
a reverse case: an Aboriginal person who painted
in the Buropean-Australian watercolour landscape
style. The handsome new book of around 30 colour
plates and nearly 40 pages of text which is part of
the subject of this article begins with a ‘celebration’
of the painter’s life by Nadine Amadio, Here we are
fhelplully told of the important retrospective exhi-
bition of Namatjira's work held at the Araluen Arts
Centre in Alice Springs in 1984, and curated by
Mona Byrnes. This signalled a revival of interest
ing painter who had become an Australian house-
hold name by the 1950s bur whose reputation had
never been very secure among curarors and scholars
of art and whose reputation declined afier his death
in Iragie circumstances in 1959.
Amadio's chapter carries the obligatory
post-l960s enlightenment regarding the falsity of
applying the wrdinary sense of the term ‘primitive’
to Aboriginal art, While bending a knee briefly to
Namatyira’s belatedly recognised (and poorly
documented) concern with cvuniry and sites of
mythological significance, this chapler succeeds
nevertheless in rescuing and promoting Namatiira
as a haive artist (my term, not hers) in another
sense: instead of High Chocolate Box we now have
Grandma Moses,
Amadio is especially concerned to frame
Namatjira’s work within the traditional key symbols
of western European art mythology. He was a ‘true
painter’ (p. 5) with his ‘own vision! (p. 6), ho ‘Knew
the joy of a full creative life, He must have possessed
an extraordinary drive and passion’ (p. 2). His ‘drive’
or ‘creative drive’ is referred to repeatedly, Strangely
posed next to this supposed ‘drive'is Namatjira as
Nature’s Gentleman, a man whose ‘dignity’ and
‘natural dignity’ (p. 10) are to be remarked upon,
and whose pictures are noted for their pellucid, airy
calm and balanced repose, They are also pictures
of beautiful (and largely unpeopled) landscapes,
excluding even a hint of the nasty side of tile
available in and areund the Alice Springs of his
times
Jonah Jones’ short picce on the 1984 exhibition,
where a remarkable 56 Namatjiras were assembled,
is of particular historical value, althoagh one could
have done without che rmystificatian of the
comparlson with Piero della Francesea on page 18
(‘It is all there’, What Is? And what is it?),
Daniel Thomas's chapter re-evaluales Namatjira’s
relationship ro the art. world. tt is the best chapter
by far and say$ same new and substantial things
abou( Namatjira’s wark, He points out some snajor
reasons Why Namatjira’s work was anly slowly
recognised by art professionals; it came our of rhe
tourist industry; it Was exhibited in unprestigious
locales for much of his hte; yice-regal patronage
inade art scholars suspicious, watercolours were
strongly associated (as they still are) with
amateurisin; and Namatjira painted in a style that
was already considered conservative and backneyed
by ‘professional, vocal, modernist’ arrists. Yet he
painted with the best of those vharavteristically
Aboriginal artistic qualities of “extrenie delicacy,
refinement, and gentleness’ (p. 26), Finally, Thomas
makes Lhe important observation that Namatjira’s
ant. which was seen in his own lime a$ “Buropean’,
Io2
is now ‘re-Ayboriginalised’ because of three maitt
factors: iis reference to signifeant country; is
symbolic repetitiveness; and 18 fineness of touch,
‘Our altered understanding pertnits us now (o
admire it more fully, and lo be moved by i (p26),
The other exeellent chapter in the book ts that
by Anne Blackwell, @ young archaeologist and
historian who sadly was killed ina war accident in
Central Australia in 1986. While Thomas’ chapter
excels in ideas, Blackwell's excels in historical fact.
The history of Hermanusburg, che roles of Carl
Strehlow and FE W. Albrecht there, the creation of
the Aboriginal uraft industry, the biography of
Namatjira, and the emergence of 'a Hermannsburg,
watercolouris! school’, are presented with scholarly
atrention to detail and handsomely dlustrated. Iv
is here that we find, amazingly, the only remark on
Namatjira’s work by an Aboriginal person: ‘That
old man — he painted’ This is reminiscent of
Amadio’s comment about the Papunya acrylics
painters on p. 33: ‘Every painting they point to, they
say simply, “this is my country” .. .', Typically, s
lengthy explanatory discourse or even a brief
gushing eulogy or eritival sideswipe is an unlikely
Central Australian Aboriginal response to a
question about a painting. This does not mean that
exlensive records cannot be made of what people
know, think and teel about a picture, Nh does mean
Thac much patient and laborious Investigation has
16 be invested in the process by the outside enquirer,
Forrning the last section of the book, the colour
pistes are perhaps the best guide to the aesthetic
upparatus of lovers of Namatiira’s work, because
Iliey are accompanied by capuons which use phrases
such as; Tichly-toned —, . warmth and sensuality
.., simple beauty of form —., an earthy power’
{p. 46); delicacy and economy’ (p. 48): a gentle and
lyrical painting, teslrained in volour' fp, 89),
Throughout evalirations like these both here and
earlier in the bawk the recurring thenves ate:
1. Innocence, naivete, gentleness, restraint
2. Fineness, delicacy, tonal subtlety, economy,
accuracy
3. Lightness, enerey, vitality, drama, vibrancy,
urgency
4_ Optimism, celebration, fanctfulness, warmth,
sensual respose.
5. Vasiness, distance, openhess of Space
6 Repetinion,
1 think Chis lends some weight to my suggestion
that Namatjira has emerged as 2 naive but
technically proficient and respectable painter, in the
assessment of fnany of his admirers. [in the past,
il was his Falsely suppased lack of sophisheation
Which played such a commanding rate in his lion-
isttion and demotie fame among suburban Ans-
Traliatis. His lotellectual sophistication is now seen
10 lie in the relanouship between his pictures and
the geographical sites and geographical religion they
represent, rather than in some happy fusion of his
High Culture with his manner of representation,
Had he done the same works in the 1980s, though,
this may not Have been the case al all,
Namatyira's is the gentle innocence and ‘warural
dignity’ of someone ‘close to the earth’ and
Unpolluted (or just about ta be polluted) by the
industrial civic culture. This is not the gentleness
(gentility, geoteelness, gentrification, gentleman-
liness} of restrained passion so characteristic of
‘civilisation’, where some sign of the wildness that
has been domesticated must remain i art is to avoid
being too wishy-washy. This is the clarity of Eden.
Ard | suggest, also, that this analysis might to
a lesser degree apply to the generally positive
response by art lovers to the Aboriginal acrylics
which have so rapidly displaced Hermannsburg
watercolourism from centre stage in the Centre,
Technically variable but usually Far from technically
incompetent, the acrylics are — especially when
documented — genuifely appealing depictions of
the concerns of a tradition which seems [ree of the
tired baggage of self-comment.
While Albert Namatjina painted (lor sale, at least)
the typically horizontal view of landscape of the
Enropean art tradition, the acrylics painters presenc
us with images of place viewed [ram a perpen-
dicular, aerial photographer's perspective.
Papunya paintings, Papunya Tula Ari, Prntup!
paintings, Western Desert paintings, Aboriginal
acrylics, the dot paintings — the tera) has yet to
settle down, perhaps reflecting two persistent
uneertaindes. Que of Ufese wicertainiies resis on
the taet chat most of us who talk about the works
haye at best a partial grasp of their social and
cultural location, or indeed that of any art or
artefactual style in Aboriginal Australia, Amather
ineertainry arises from the fact that, fifteen years
after the birth of che Aboriginal acrylics ‘mave-
went’, i is sill manly Known through exhihirion
epheniera, TV programmes, bric! mayurine arpctes,
and spall sections of survey works such as the
excellent introductions to Aboriginal art by Berndt,
Berndt & Stanton (1982) and tsaacs 41984).
In other wards, there is no mayor, susiaimed,
scholarly study of these paintings available, and
which is on a scale comparable to Nancy Munn's
Halbiri fcunography (1973) or Howard Morphy's
Unpublished thesis on northeast Arohem Land
bark paintings (1977), Such a study would form an
authoritative basis for characterising the works. An
historically ortented study of this kind ix now
urgently needed, (ur the acrylics are not only a
‘srvle’, uw ‘genre’. & ‘Thovemen!” (spreading phena-
oienon, at east) and an “indusiry’ but alse a rapidly
developing ‘phase’ in a very old graphie tradition
— in feet, the very one Munn was writing about
iwerily Years ago
The catalogue of book Bar arid Circle, edited by
Janel Maughan & Jenny Zimmer, lies Somewhere
between ihe handy but hghiweight ephemera, which
usually Throws up unintegrated fragments of face
(and fiction), and the fullecale scholarly study
Which turns an array of facts into a body of
knowledge and interpretation,
In one-of its facets, Dot anc Circle isa listina and
reproduction of the 102 pictures shown in the
exhibition of [he same name lent by the Flinders
Universtiy Acl Mrseum to the Royal Melbourne
Insotute af Teclunology Gallery in April-May 1985_
They later returned to be shown at Flinders
University in Adelaide, The catalogue is valuable
nol only because it lists the works and their atten
considerable documentation, but also because it
reports the exhibviion itself in the form of
photographs of the installation and 1s opening, lists
of case exhibit¢ and exhibition photographs, and
4 reproduction of the exhibition text labels,
Inventories of the firse year's consignments of
aervlics from Papunya te the Stuarr Art Centre
1971-2, provided here by Pat Hogan, and the
appendices detailing works painted by David Corby
Kjapaltjarrt and Tiirkey Tolsen Tyupurrita while
Aboriginal Artists in Residence at Flinders in 1979,
may alse be mere lists but they will be of much
greaicr historieal interest as Gme goes by.
In its ather, more discursive role the book does
provide a collection of brief essays on the ongins
and sigoificance of the acrylics movement by a
variety of authors, There is tile imtelleciual
coherence berween the essayists here, however. Geoff
Rardon’s account of the sudden adoption of aeryhe
painting at Papunya durjig his time there it 1971,
here reprinted from his 1979 work (Bardon 1979},
is uw reminiscence of importance to Australian art
history and also a work of greac honesty, even if
at times it shows an unnervingly slight grasp of the
culuire of Aboriginal people. Dick Kimber's much
mare savvy recollections of the same period follow
those of Bardon and very usefully cite the names
of the key European supporters of the ‘movement’
(As Kimber says, rather paradoxieally, iris ‘only a
movement in Buropean eyes accustomed to crowds
ralher than individuals’ (p. 19).) lis clear that the
aerylics ure, and always have been, very much a
dveet prodact of particular Aboriginal-European
interactions, nol stmply ‘Aboriginal products’
sought and boughl by Europeans.
Another reminiscence in this colleetion, Radney
Morice’s ‘The Kdnekayunt experience’, is the
exceptional essay in the lilerary sense if as
beuurifally written, it is alive and conveys the quality
of daily experence in a remote Aboriginal camp,
wd ib rings true, ay Ure work of someone Who was
Wis
there long enough and had sufficient sensibility to
get it nght, As a means of orienting the reader to
an unfamiliar World it is excellent, even though its
reiation to the business at hand ls rather gerieral
(and like the Bardon piece itis a reprint (in Uns case,
from Over/and |978)), The heavy dominance of the
overview in the presentation of the Aboriginal arts
is, 1] hope, soon coming to an end,
Andrew Crovker’s essay, ‘Potential and pitfalls',
promotes |he view Lhat Aboriginal works should be
allowed to take ‘their place in the contemporary
arnstic Jorum’, aiid supports the view that the
evolution of Aboriginal art “enables those
Aboriginal artists with an exceptional artistic flair
to (loursh'(p. 47}, opposing the restrictive attitude
of ‘traditional is better’. The possibility that the
Papunya paintings emerged from an assertion of
traditionalism by older men sits uneasily here —
it does not invalidate Crocker'’s view, bul it does cry
out for consideration,
The sections of the book which will perhaps. be
ol greatest iaterest fo scholars of art history and
material culture are those by Janet Maughan and
Vinvent Megaw. (Ry ‘material culture’ here | meat
io include aesthetic anthropology, as well as the less
theoretically-oriented branches of that field.)
Megaw, an archacologist by training, has
published a number of papers on the Aboriginal
acrylics phenomenon. This one, ‘Dreamtime
diseipline or alien adulteration?’, stresses a theme
similar to that of Crocker’s piece which precedes
it} Chal the “fine art” status achieved by Aborginal
works is legitimate, and that commercial and insti-
tutional recognition (or should it be vonstitition”?}
of thal fact is inevitable, People who complain of
capitalist penetration of Aboriginal society via art
are idealising, romanticising and alrempring to
fossilise its culture, Megaw suggests that change in
Aboriginal art is not adulteration of a pristine
Table culture ty a greedy, foreign one bul 4 sign
of Aboriginal ealtural vitality tn the modern world,
Like his other papers on similar themes, Megaw's
essay in this case questions the rationality and the
moraliry of distinctions such as fine art/tourist art,
art artefact, art /crall and so on, meanwhile raising
the problem of applying aesthetic judgements
across cultures. Several pasitions on these subjects
are adumbrated, none of them is particularly
rigorously pursued, and few of them are plumped
for ottier (han gingerly, The profession of arehac
ology is Mentioned jnan act of dissociation of the
author from the philosophy of art or art criticism,
Jenny Zimmer's preface (oddly, one of two, rhe
other being by Maughan & Megaw) provides much
usetul information on jhe creation of the exhibition
itself, As a performance it must go down as one
of the most archestrally complex in Abonginal art
history, She alse offers some revealing thoughts an
164
why such an exhibition was considered useful as
part of the education of some Melbourne art
students. There is not room here to argue all the
points she raises, although most of them are
arguable.
Her most interesting point is summarised in her
last paragraph: ‘Cultural Convergence can only serve
to make our culture richer and more authentic. This
exhibition and booklet are dedicated to the concept
of Cultural Convergence’ (p. 13, bold print original).
Perhaps this is an implicit stand against the recent
avalanche of Left criticism in this field, which
resounds with phrases like ‘cultural genocide’,
‘astheticisation’ and ‘neo-colonialist encapsulation’.
It may also be a recognition of the fact that a new
Australian sub-culture has been emerging for fifteen
years or so among people of no Aboriginal ancestry,
many of them resident in Central Australia. They
are creating a partly new way of life, one affected
by a perception of Aboriginal culture and by
interaction with Aboriginal people, yet distinct. This
theme is reflected in the final section of the
exhibition and catalogue, which is called variously
‘White artists using Aboriginal designs’ (p. 18), ‘The
Western Desert image and the European art context’
(p. 25), ‘Across the cultural divide’ (p. 191) and
‘Across the cross-cultural divide’ (p. 205).
Tim Johnson is a non-Aboriginal artist who has
used graphic devices drawn from the Aboriginal
acrylics and who has painted jointly with an
Aboriginal artist (see cat. nos. 97, 98). Whether this
is best described as ‘convergence’ or ‘incorporation’
(etc.) has been a matter of debate. Whether it has
yet resulted in ‘good works of fine art’ has also been
a matter of debate. The latest wave of European
artists influenced by Aboriginal graphs can
probably claim a moral, if not an aesthetic, advance
over predecessors like Preston, whose contact with
Aboriginal people was generally much more
marginal and whose borrowing was done from
photographs and museum collections rather than
‘with the people’.
Not only artists but collectors and curators
interested in the Aboriginal acrylics are beginning
to talk of them as the first truly Australian art, and
as a case of cultural convergence between
Aborigines and European Australians. This
perception, problematic as it may be, is gathering
force.
I have left Janet Maughan’s contribution to the
end, both because it is the longest and because it
is comparatively rich, She outlines the history of
the genre and the referential meaning system of its
images, and attempts a style analysis which, as she
herself points out, fails to get off the ground (p. 17).
I do not agree that such an analysis is not possible,
however. The effect of variety perceivable among
the pictures is great, certainly, but it is one which
rests on the recombination of a limited set of motifs
and of ways of placing them in a symmetrical field.
The list of motifs or recurrent symbols in Figure
2.(p. 16) seems to me seriously incomplete, and also
conflates sets of symbols which ought to be kept
distinct. No structural typology of the images is
offered, although the works are eminently suitable
for this kind of first-sort. It is difficult to see how
a style analysis could procede without some kind
of sorting into the various kinds of axial symmetry,
for example.
Maughan then goes on to discuss the major
subgroups of works in the exhibition and the
rationale for their subgrouping, beginning at the
beginning (the early paintings), and moving on to
look at two artists specially represented in the
exhibition, Tim Leura Tjapaltjarri and Toby Brown
Tjampitjinpa. This concentration on two indivi-
duals was important to establishing an idea of the
style and output of particular artists over time.
Institutional roles in the acceptance and
promotion of Aboriginal acrylics are highlighted
here. The particular efforts of Visual Arts staff and
students at Flinders University are given their due,
though for better balance there could have been
more emphasis, for example, on the historic role of
Robert Edwards and the Aboriginal Arts Board in
promoting both the production and the sale or
public collecting of these works in the 1970s and
1980s.
‘Children’s paintings’ (for, not by) come next,
followed by ‘Women’s paintings’ (by, not for), where
it is made clear that women are under-recognised
as painters in the genre and tend to produce for the
‘craft’ market. The question of how much this may
be a reflection of gender roles in matters of religious
authority is not opened up.
Maughan ends by relating Papunya paintings to
conceptual art, but, in the absence of a discussion
of what that is, many readers may find the connec-
tion obscure. It is a pity there was not room enough
in her essay for a substantial discussion of the
relation of the acrylic designs to those largely very
similar ones of the same region which have been
recorded since the earliest European times, but in
other media: the rock intaglios, rock paintings,
body decorations, shield designs, and the sand
drawings of daily conversation and play.
Among Maughan’s more interesting observations
are the view that the complexity of figure-ground
relations in the acrylics has increased over time, and
her important statement that there is a gaping lack
of published information on Aboriginal criteria for
merit in paintings. Great credit is due also for her
long and detailed work on the catalogue annota-
tions.
The production quality of the book leaves much
to be desired, but most of my reservations there
would have been pre-empted by more generous
publication funding of typesetting and layout,
printing and binding. The page size (A4) is
unattractive; it needs to be more squat. The 13
colour plates are fine but the monochrome plates
are rather muddy. In one case a signature has been
chopped in half (p. 104) and in another the
alignment marks have intruded (p. 111). Lines
wobble.
Perhaps more serious are editorial lapses such as
uncaptioned plates (pp. 1, 4, 8, 12, 41, 48, 93),
typographic errors, and the entirely wrong
information under the plate on p. 20 which says:
‘Western Desert: Demonstration of circular designs
which have in part replaced the more complex
traditional ground paintings. Photo: Penny
Tweedie.’ This is actually a photograph by C.P.
165
Mountford taken at Haasts Bluff in 1942. This
mistake is perhaps partly compensated for by the
humour value of the caption to the upper photo
on p. 46: ‘Old Mick Tjakamarra with Daphne
Williams recording the story of the painting, 1982.
Photo: Vincent Megaw’. Ms Williams, apparently
short of paper, is on close inspection writing down
the details in her Bank of New South Wales
cheque-book. . .
ACKNOWLEDGMENTS
I am grateful to Christopher Anderson, Philip Jones
and Andrew Pekarik for their helpful comments on an
earlier draft of this paper.
REFERENCES
BARDON, G. 1979. ‘Aboriginal Art of the Western
Desert’. Rigby, Adelaide.
BERNDT, R.M., BERNDT, C.H. & STANTON, J.E. 1982.
‘Aboriginal Australian Art: A Visual Perspective’.
Methuen Australia, Sydney.
ISAACS, J. 1984. ‘Arts of the Dreaming: Australia’s Living
Heritage’. Landsdowne, Sydney.
MORPHY, H. 1977. ‘ “Too many meanings”: an Analysis
of the Artistic System of the Yolngu of Northeast
Arnhem Land’. Ph.D. thesis, Australian National
University, Canberra.
MUNN, N.D. 1973. ‘Walbiri Iconography’. Cornell
University Press, Ithaca.
RUBIN, W. 1984. ‘Modernist Primitivism: An
Introduction’. Jn W. Rubin (Ed.). ‘ “Primitivism” in 20th
Century Art: Affinity of the Tribal and the Modern’.
Museum of Modern Art, New York.
P. SUTTON, Head, Division of Anthropology, South Australian Museum, North Terrace, Adelaide 5000. Rec. S.
Aust. Mus. 21(2): 160-164.
RECORDS
NY
THE
SOUTH
AUSTRALIAN
MUSEUM
VOLUME 21 PART 2
NOVEMBER 1987
ISSN 0081-2676
CONTENTS:
ARTICLES
69 D.S. TRIGGER
85
119
139
149
157
161
Inland, coast and islands: traditional Aboriginal society and material culture
in a region of the southern Gulf of Carpentaria
C. PATTERSON & P. V. RICH
The fossil history of the emus, Dromaius (Aves: Dromaiinae)
S. J. EDMONDS
Echiurans from Australia (Echiura)
J. RILEY & D. M. SPRATT
Further observations on pentastomids (Arthropoda) parasitic in Australian
reptiles and mammals
L. HERCUS
Looking for Ditji-mingka
NOTES
W. ZEIDLER
The scaled-squid, Lepidoteuthis grimaldii Joubin, from southern Australian
waters
P. SUTTON
From horizontal to perpendicular: two recent books on central Australian
Aboriginal painting
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