Memoirs
OF THE
Queensland Museum
BRISBANE
MARCH, 1984
VOLUME 21
PART 2
VOLUME 21
PART 2
Memoirs
OF THE
Queensland Museum
PUBLISHED BY ORDER OF THE BOARD
NOTE
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THE
QUEENSLAND MUSEUM
Gregory Terrace
Fortitude Valley
Brisbane, Qld., 4006
Telephone: 52 2716-8
TRUSTEES
J.C.H. Gill, B.A., LL.B. (Chairman)
I. G. Morris, C.M.G. (Vice Chairman)
R.I. Harrison, E.D., B.Comm.
J. T. Maher, F.R.Hist.S.Q.
Professor D.J. Nicklin, B.Sc.App., B.Sc., B.Econ., Ph.D.,
FT.E.Aust., F.I.Chem.E.
Professor J.M. Thomson, D.Sc.
D.M. Traves, O.B.E., B.Sc.
A. Bartholomai, M.Sc., Ph.D., {ex officio . Secretary)
SCIENTIFIC STAFF
Director: A. Bartholomai, M.Sc., Ph.D.
Deputy Director; B.M. Campbell, M.Sc.
Senior Curators: L.R.G. Cannon, B.Sc., Ph.D., Curator of Lower Invertebrates
Jeanette Covacevich, B.A., M.Sc., Curator or Reptiles
E.C. D.AHMS, B.Agr.Sc., B.Sc,, M.Sc., Curator of Higher Insects
Valerie Davies, M.Sc., D.Phil., Curator of Arachnids
Patricia Mather, Ph.D., D.Sc., Curator of Higher Invertebrates
R.J, McKay, B.A., Curator of Fishes
R.E. Molnar, Ph.D., Curator of Mammals (Editor)
G.B. Monteith, B.Sc., Ph.D., Curator of Lower Insects
M.C. Quinnell, M. a. (H ons.), Curator of Anthropology and Archaelogy
D. Robinson, B.Sc. (Hons.), Ph.D., Curator of History and Technology
LG. Sanker, B.Sc., Dip. Ed., Curator of Industrial Technology
Mary Wade, B.Sc. (Hons.), Ph.D., Curator of Geology
Curators: R.A. CoLEMAN, B.F.A., Curator of Maritime Archaeology
P.J. Davie, B.Sc., Curator of Crustacea (Assist. Editor)
R.G. Hardley, B. a, (H ons.), Curator of Australian Ethnography
G.J. Ingram, B.Sc., Curator of Amphibians and Birds
J. Stanisic, M.Sc., Curator of Molluscs
Conservator: N.H. Agnew, M.Sc., Ph.D.
Mem. QdMus. 21(2): 257—59. [1984]
NEW RECORDS OF BOPYRIDAE (CRUSTACEA : ISOPODA : EPICARIDEA)
FROM QUEENSLAND WATERS
S.P. Nearhos and RJ.G. Lester
Department of Parasitology University of Queensland
ABSTRACT
Two species of bopyrids were recovered from commercially caught prawns in Queensland.
Epipenaeon ingens Nobili 1906, previously recorded from Darwin, was found in four places in
Queensland on its type host Penaeus semisulcatus, or on P. merguiensis. On the basis of a
high degree of variability in specimens, E. ingens latifrons Bourdon 1979 is considered a junior
subjective synonym of E. ingens. Other members of the genus Epipenaeon are listed for
comparison.
Parapenaeon expansus was found on Penaeus plebejus in Moreton Bay, and on Penaeus
sp. from Karumba.
These constitute new site and host records, for both species.
INTRODUCTION
Parasites of prawns have recently attracted
interest as they may be useful as biological
markers for prawn sub-populations (Owens 1981,
1983). Bopyrid isopods are among the most
obvious parasites carried by Australian prawns.
Although several bopyrid species occur in
Australian waters only two have been found on
commercially valuable prawns. These are
Epipenaeon ingens Nobili 1906, and
Parapenaeon prox. expansus Bourdon 1979a,
both from ‘tiger prawns’ taken in the vicinity of
Darwin, N.T. (Bourdon 1979b). Recently these
bopyrid species were recovered from collections
of penaeids made in Queensland waters. This note
records where and on what species of hosts they
have been found.
Prawns trawled from the Gulf of Carpentaria,
Rosslyn Bay, and Maryborough were obtained
frozen from fish marketing boards. In Moreton
Bay prawns were obtained fresh from trawlers.
Parasites were removed and preserved in 70%
alcohol. Specimens have been lodged at the
Queensland Museum (QM).
EPIPENAEON Nobili 1906
Epipenaeon ingens Nobili 1906
Epipenaeon ingens Nobili 1906, p. 1099-101 Fig.
1, 1 a-e. Bourdon 1968, p. 327-33, Figs.
145-50.
Epipenaeon nobili Nierstrasz and Brender a
Brandis 1929, p. 299-302, Figs. 5-9.
Epipenaeon grande Nierstrasz and Brender a
Brandis 1929, p. 157-58, Fig. 18; 1932, p. 91,
Fig. 1.
Epipenaeon ingens latifrons Bourdon 1979, p.
429-30, Fig. 4 a-c.
MATERIAL EXAMINED
QM W10438, $ + 2, ex Penaeus
semisulcatus, Karumba, Gulf of Carpentaria,
NW.Q., S.P. Nearhos, I4.vii.79; QM W10445,
15 ^ -E 19 V ex, P. semisulcatus, Karumba, S.P.
Nearhos, 4.vii.78; QM W10436, 3' + e, ex P,
merguiensis, Karumba, S.P. Nearhos, I4,vii.79;
QM W 1043 7, S + 9, ex P. merguiensis,
Karumba, S.P. Nearhos, 14.vii.79; QM W10439,
■S -E 9, ex P. merguiensis, Karumba, S.P.
Nearhos, 14.vii.79; QM W10446, 1 1 + 22 9, ex
P. merguiensis, Karumba, S.P. Nearhos,
17.vii.79; QM W10448, 8 2, 2 rj, ex P.
semisulcatus, Maryborough, SE.Q., S.P.
Nearhos, 10.iv.78; 10 (7 -e 13 2, ex P.
merguiensis, Rosslyn Bay ME.Q., S.P. Nearhos
17.vi.78.
OTHER MATERIAL
E. nobili i + 5, ex P. semisulcatus. Red Sea,
Nierstrasz and Brender a Brandis.
E. grande 9, ex P. semisulcatus, Hong Kong,
23.V.1890, Nierstrasz and Brender a Brandis.
E. ingens latifrons, 4 <7 -E 4 2 , ex P,
semisulcatus, Darwin, Bourdon.
DISTRIBUTION
Mediterranean Sea and the Indo-Pacific region
— from Hong Kong in the north to southeast
Queensland.
258
MEMOIRS OF THE QUEENSLAND MUSEUM
REMARKS
E. ingens was originally described from
Penaeus semisulcatus from the Red Sea (Nobili
1906). It was redescribed from the same host in
the Mediterranean by Bourdon (1968). Bourdon
(1979b) synonomized E. nobili and E. grande
with E. ingens and described a new subspecies E.
ingens latifrons from a Penaeus sp. for its well
developed frontal plate and unusually wide lateral
plates.
Examination of 24 specimens from P.
semisulcatus from the Gulf of Carpentaria and
from Maryborough showed that development of
frontal plates varied between individuals. The
mean ratio of frontal plate length to total length
was 0.074 (range 0,03 - 0,10). The types of E.
ingens latifrons fall within the range. Lateral
plates showed similar variation. Specimens from
P. merguiensis from both eastern and northern
Queensland have less developed frontal and
lateral plates, and in this respect more closely
resemble E. ingens from the Red and
Mediterranean Seas than they do E, ingens
latifrons. The males from both hosts are very
similar and correspond closely to the male
described from the Mediterranean, the only E.
ingens male described.
It appears then, that the development of the
frontal and lateral plates vary from individual to
individual and from host to host. On this basis we
propose that E. ingens latifrons be considered a
junior subjective synonym of E. ingens.
Two other species of Epipenaeon also occur on
P. semisulcatus — E. elegans Chopra, 1923 and
E. pestae Nierstrasz and Brender a Brandis, 1932.
The published descriptions of these species are
inadequate to clearly separate them from E.
ingens, considering the highly variable nature of
E. ingens individuals.
E. ovalis Pillai 1954, was recorded from
Parapenaeopsis stylifera and characterized by
having a broader frontal plate and greater
development of the pleonal lamellae than E.
ingens. Both of these features were found to have
been variable for specimens of E. ingens
examined in this study. The type material of E.
ovalis has been lost and is unavailable for
comparative study (Pillai, pers. comm.).
E. oviformis Nierstrasz and Brender a Brandis,
1931, described from a Penaews" sp. is obviously a
juvenile specimen which makes comparison with
mature adults impossible. Its status is therefore
doubtful.
Of the species described in the genus, at this
only E. fissurae Kensley 1974, seems
definitely separable from E. ingens. It can be
distinguished by the shape of the antennae, the
blunt digitations of the posterior margin of the
cephalon and first oostegite, and the knobbed
nature of the pleopods.
PARAPENAEON Richardson 1904
Parapenaeon expansus Bourdon 1979
Parapenaeon expansus Bourdon 1979, 495-8,
Figs. 15-8.
MATERIAL EXAMINED
QM W 10440, ?, QM W1044I, + 9, QM
W10442, 2 ^ QM W10443, 9 ex Penaeus
plebejus, Moreton Bay, SE.Q. S.P. Nearhos,
9.ii.79, 9.ii.79, 12.ii.79 and l.v.78; QM W10444,
^ + , ex P. plebejus, Moreton Bay, R.J.G.
Lester, 23.i.78; QM W10447 ^ ex P. plebejus,
Moreton Bay, S.P. Nearhos, 24.iv.78; QM
W10451, 6 2, ex Penaeus sp. Karumbu, Gulf of
Carpentaria, NW.Q.S.P. Nearhos, 9.ii.79.
DISTRIBUTION
Indo-Pacific Oceans from Madagascar to
northern Australia and south to Moreton Bay in
Queensland.
REMARKS
Present material of Parapenaeon expansus
(both males and females) correspond closely to
those described by Bourdon (1979a) from the type
host Penaeus teraoi in Madagascar. This
constitutes a new host and distribution record for
the species.
ACKNOWLEDGMENTS
We thank Dr R. Bourdon for his help, advice
and the loan of the type material of E. ingens
latifrons and P. expansus. Dr H.E. Gruner,
Zoology Museum, East Berlin, kindly loaned
specimens of E. nobili and Dr T. Wolff, Zoology
Museum, Copenhagen loaned the 2 type
specimen of E. grande. P.J.F. Davie,
Queensland Museum is thanked for his assistance
in the preparation of this manuscript.
The study formed part of the Masters
Qualifying Thesis for the senior author.
LITERATURE CITED
Bourdon, R., 1968. Les Bopyridae des mers
Europeenes. Mem. Mas. Nat. d’Hist. Paris,
L2; 327-33.
1979a. Epicarides de Madagascar 11. Bull. Mus.
Nat. d'Hist. Nat. Paris, 43 Ser. 1 Sect. A:
471-506.
NEARHOS AND LESTER: NEW RECORDS OF BOPYRIDAE
259
1979b. Sur le taxonomic et Pethologie de
quelques Orbionines. (Isopoda : Epicaridea).
Int. Rev. Ges. Hydrobiol 64: 425-35.
Chopra, B., 1923. Bopyrid Isopods parasitic on
Indian Decapod Madura. Rec. Ind. Mus. 25:
411-542.
Kensley, B., 1974. Bopyrid Isopoda from
Southern Africa. Crustaceana 26: 259-66.
Nierstrasz, H.F. and G.A. Brender a Brandis,
1929. Neue Epicaridea. Zool. Anz. 85:
295-302.
1931. Papers from Dr Th Mortenson’s Pacific
Expedition. LVII Epicaridea II. Vidensk.
Meddr. dansk. naturh. Foren. 91: 147-225.
1932. Alte und Neue Epicaridea. Zool. Anz.
101: 90-100.
Bobili, Y., 1906. Nuovi Bopiridi. Atti Acad Sci
Torino 41: 1098-113.
Owens, L., 1981. Relationships between some
environmental parameters and trypanorynch
cestode loads in banana prawns (Penaeus
merguiensis de Man). Aust. J. Mar. Freshw.
Res. 32: 469-74.
1983. Bopyrid parasite Epipenaeon ingens
Nobili as a biological marker for the banana
prawn Penaeus merguiensis de Man. Aust. J.
Mar. Freshw. Res. 34: 477-81.
PiLLAi, N.K., 1954. A preliminary note on the
Tanaidaceae and Isopoda of Travancore.
Bull. Cent. Res. Inst. Univ. Travancore. C3:
23-4.
Mem. QdMus. 21(2); 261—69. [1984]
PITONGA GEN. NOV., A SPIDER (AMAUROBIIDAE : DESINAE) FROM
NORTHERN AUSTRALIA.
Valerie Todd Davies
Queensland Museum
ABSTRACT
A 3-clawed spider from the mangroves of the Northern Territory, Australia is described and
provisionally placed in the Desinae. It has untoothed tarsal claws and a long inferior claw, a
copulatory spur on tibia II and no serrula.
INTRODUCTION
These spiders were collected by members of the
Australian Littoral Society during a survey of the
mangrove areas of Northern Australia. The
female was found in a crab-hole on the mud, the
male walking on the mud and two of the three
juveniles collected were in silk cells on the
mangrove leaves along the river, the third on
mud.
Pitonga gen. nov.
Medium sized, 3-clawed spider. Eyes small, in 2
almost straight rows occupying median third of
head. Clypeus narrow. Promargin of chelicera
with teeth, retromargin without teeth but with a
secretory protuberance opposite last promarginal
tooth. Labium and maxillae elongate, no serrula.
Spinnerets sub-terminal; no colulus. Legs 1423,
anterior trochanters un-notched. One long distal
trichobothrium on metatarsi, none on tarsi. Male
with thick spines on Tibia I and ventral spur-like
spine on Tibia 11.
‘Pitonga’ is an aboriginal word meaning
mangrove.
Pitonga woolowa sp. nov.
Holotype; In crab hole on mud bank, Flying
Fox Is., East Alligator River, Northern Territory,
W. Houston, 15.vi.81, I ?, QM S1300.
Paratypes: On mud, East Alligator River,
N.T., W. Houston, P. Davie, 23.vi.82, 1 c?, QM
S1301; in silk cells on mangrove (Avicennia sp.)
leaves, 2 juvs QM S1302, On mud. Point
Farewell, East Alligator River, N.T., W.
Houston, P. Davie, 18.vi.82, 1 juv., QM S1303.
Description of female
CL 3.13, CW 2.13, AL 2.87, AW 2.00
(abdomen shrunken).
Colour; The spider is pale and straw-coloured
resembling Clubiona. Chelicerae, metatarsi and
tarsi light brown. Closely adpressed hair on
cephaloihorax and abdomen, longer hairs on
abdomen. Legs hairy, especially laterally on distal
segments. Viewed from above anterior row of
eyes slightly recurved, posterior row straight;
from the front, anterior row straight, the
posterior row procurved (Figs. 1, 2). Ratio of eyes
AME;ALE;PME;PLE is 9;9.*7:I0. Clypeus
narrower than diameter of AME. Chelicerae large
and long (2.0 mm) with boss. Retromargin
without teeth, promargin 10 teeth. Small medium
tooth opposite 4-5 and retromarginal
protuberance opposite the last promarginal tooth
between which the tip of the fang rests (Figs. 3,
14). Magnification shows that this boss has
regular pores opening on it (Fig. 15). Labium
longer than wide 1;0.81. Sternum longer than
wide 1;0.71. Spinnerets; small, short and sub-
terminal (Fig. 4).
TABLE I: ? Leg Measurements.
Femur
Patella
Tibia
Metatarsus
Tarsus
Total
palp
1.50
0.57
0.69
-
0.94
3.70
I
2.72
1.00
2.50
2.41
0.72
9.35
II
2.13
0.84
1.56
1.81
0.63
6.97
III
1.84
0.84
1.09
1.50
0.53
5.80
IV
2.75
1.00
2.31
2.19
0.63
8.88
262
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGS. 1-6: 9 Pitonga woolowa (holotype). 1, cephalothroax, dorsal; 2, cephalothorax, lateral; 3, cephalothorax
and mouthparts, ventral; 4, spinnerets; 5, epigynum, external; 6, epigynum, internal.
■^-9; ^ Pitonga woolowa (paratype). 7, tibia I, ventral; 8, tibia II, retrolateral; 9, 1. palp, dorsal.
DAVIES: PITONGA GEN. NOV.
263
TABLE II S Leg Measurements
Femur
Patella
Tibia
Metatarsus
Tarsus
Total
I
3.09
1.03
2.97
3.09
0.94
11.12
II
2.41
0.97
1.97
2.25
0.66
8.26
III
2.03
0.94
1.25
1.75
0.53
6.50
IV
3.38
1.06
2.50
2.47
0.75
10.16
Legs 1423 (Table I) tarsi short, about 1/3
metatarsi; hairs plumose (sensu Lehtinen).
Posterior trochanters shallowly notched. Tarsus
with 3 smooth claws; inferior claw long (Figs. 10,
11). Two stouter hairs lateral to it and cluster of
short hairs ventral to it (Fig. 12). Between 2-4
Irichobothria in irregular rows on tibiae, one long
distal trichobothrium on metatarsi (Fig. 16), none
on tarsi. Capsulate tarsal organ (Fig. 17) with
pyriform opening.
Notation of Spines: First leg: Femur, p 1 distal,
d 1.1.1, Tibia, p 1.1. 1.1. 1(0), v 2.2.2. Metatarsus,
p 1 . 1 , 1 , r 0. 1 . 1 , V 2 proximal. Second leg: Femur,
p 1 distal, d 1.1.1. Tibia, p 0.1. 1(0), v 2.1.1.
Metatarsus, p 1.0.1., d 0.1.1, r 1.0.1, v 1.2. 1(0).
Third leg: Femur, d 1.1.2. Tibia, p 0.1,1, r 0.1.1,
V 2(1). 0.0(1). Metatarsus, scattered proximal 5-6,
distal whorl 6. Fourth leg: Femur, d
I.l.l.l.l.I.O.O. Tibia, r O.l.O.l, v 2.O.I.O.
Metatarsus, scattered proximal 5, distal whorl
5-6.
Epigynum: The fossa has a sclerotized rim
which makes it difficult to trace the course of the
ducts to the spermathecae (Figs. 5,6).
There are 4 unbranched abdominal tracheal
tubes.
Description of male
CL 3.18, CW 2.35, AL 3.13, AW 1.73.
Similar to female in colour, eye ratios,
cheliceral teeth and trichobothrial pattern. Legs
1423 (Table II). The prolateral and ventral spines
on tibia I (Fig. 7) are enlarged and there is a large
ventral spur-like spine on tibia II (Fig. 8) which is
presumably used during mating.
Notation of Spines. First leg: Femur, p 1 distal,
d 1.1. 1.1. Tibia, p 2(1).1.1.0(1).L0, d 1.0.1, r
1(0). 1. 0(1). 1, V 2. 2. 2. 0.0. Metatarsus, p 1.0.1., v
2.2(1). 1. Second leg: Femur, p 0.0(l).l, d 1.1. 1.1.
Tibia, p O.l.O.l(spur), d 0.1.0, r 0.1.0, v 2.1.1.
Metatarsus, p 1.1.1, v 2.1.1. Third leg: Femur, p
0.1.0, d 1.1. 1.0.1, r 0.1.0. Tibia, p 0.1.1, d 0.0.1,
r 0.1.1, V 2.0.0. Metatarsus, scattered proximal 6,
distal whorl 6. Fourth leg: Femur, d
1.1.1.1.0(l).l. Tibia, p O.l.O.l, v 2.0. 1.0.
Metatarsus, scattered proximal 6, distal whorl 6.
Palp. Embolus thick and spear-shaped,
membraneous median apophysis (Fig. 13). Small
prolateral posterior extension of cymbium.
Elaborate tibial apophyses (Fig. 9).
The species name ‘woolowa’ is an Aboriginal
word meaning mud flat.
DISCUSSION
It is difficult to decide whether characters in
Pitonga indicate relationship to a recognised
family or are specialized adaptations to the
mangrove area in which the spider lives. It seems
likely that the long inferior tarsal claw and lack of
teeth on any of the claws are adaptations for
running on mud.
Lehtinen (1980 : 493) regards the trichobothrial
pattern of 2 rows on tibiae, 1 on metatarsi and
none on tarsi as the plesiomorphic state; it is
found in hypochiloids, haplogynes araneoids and
some others. In Pitonga the tibial trichobothria
are few and irregularly placed. The structure of
the trichobothrial base and the ridged (rather than
scale-like) cuticle around it, as well as the
pyriform aperture of the tarsal organ, suggest
amaurobioid rather than araneoid affinities
(Lehtinen 1978 : 267, Forster 1980 : 273).
Spination and copulatory spurs on the anterior
tibiae of males are common in mygaiomorphs and
in a few araneids — several other familes have
metatarsal spurs. A retrolateral tibial apophysis is
found in the palp of most $ spiders, the main
exceptions being most of the araneoid spiders and
the lycosids. In Pitonga there are 2 apophyses, the
second one arises prodorsally and turns
retrolaterally. Many amaurobioid spiders have
complex tibial apophyses. The absence of a
serrula in Pitonga is regarded as an apomorphy.
Although Pitonga resembles araneoid spiders
in trichobothrial pattern and possession of tibial
copulatory spurs in the male these characters are
regarded as plesiomorphic and thus may not
264
MEMOIRS OF THE QUEENSLAND MUSEUM
indicate relationship. The structure of the
trichobothrial base and cuticle, the pyriform
opening of the tarsal organ and the plumose hairs
suggest amaurobioid affinities. Many dictynoids
(among the amaurobioids) show a like reduction
in tarsal trichobothria. The complex tibial
apophysis further supports this view and although
there is no colulus the anterior spinnerets are well
separated suggesting recent cribellate ancestors.
The hairiness of the legs, the long chelicerae
and maxillae, and the coastal locality have
influenced the provisional placing of Pitonga in
the Desinae.
LITERATURE CITED
Lehtinen, P.T., 1978. Definition and Limitation
of Supraspecific Taxa in Spiders. Symp. zool.
Soc. Lond. 42: 255-71.
1980. Trichobothrial patterns in high level
taxonomy of spiders. Verh. 8. Internationaler
Arachnologen-Kongress Wien ; 493-8.
Forster, R.R., 1980. Evolution of the tarsal
organ, the respiratory system and the female
genitalia in spiders. Verh. 8. Internationaler
Arachnologen-Kongress Wien : 269-84.
266
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
FIGS, 10-13: Pitonga woolowa. 10, tarsus IV (QM S1302); 11, tarsus
III claws and hairs (holotype), scale line = 71 um; 12, same,
ventral, scale line = 83.3 um; 13, S palp, prolateroventral, scale
line = 50 um.
DAVIES: PITONGA GEN. NOV.
267
268
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
FIGS. 14-17 Pitonga woolowa. 14, cheliceral teeth and process (QM
S1032), scale line = 50 urn; 15, process with pore openings, short
scale line = 5 urn; 16, trichobothrial base metatarsus III
(holotype), short scale line = 12.5 um; 17, tarsal organ tarsus III
(holotype), short scale line + 10 um.
DAVIES: PITONGA GEN. NOV.
269
« is#
Mem. QldMus. 21(2): 271 —336. [1984]
REVISION OF THE GENUS MELITTOBIA (CHALCIDOIDEA : EULOPHIDAE)
WITH THE DESCRIPTION OF SEVEN NEW SPECIES.
Edward C. Dahms
Queensland Museum
ABSTRACT
This taxonomic revision of the genus Melittobia contains 2 new combinations, Tachinobia
diopsisephila (Risbec), Cirrospilus {Atoposomoidea ) cosmopterygi (Risbec); a generic
diagnosis; the redescription of 7 species, davicornis (Cameron), acasta (Walker), chalybii
Ashmead, megachi/is (Packard), hawaiiensis Perkins, australica Girault and bekiliensis
Risbec; description of 7 new species, evansi, scapata, femoraia, assemi, sosui and one from
Argentina not named because of lack of suitable specimens for type selection; synonymies,
strandi Wolff and Krausse and Anthophorabia fasdata Newport become acasta (Walker),
Japonica Masi becomes davicornis (Cameron), sceliphronidis (Brethes) becomes hawaiiensis
Perkins, Tachinobia gradwdli Bou6ek becomes T. diopsisephila (Risbec); osmiae Thompson
and hawaiiensis peles Perkins remain unidentifiable in the absence of types and definitive
descriptions; figures and keys are provided to
MATERIALS AND METHODS
Specimen Mounting
The methods for mounting Melittobia are
standards used for other Chalcidoidea involving
air dried specimens glued to a card rectangle or
cleared in 10*^0 NaOH for mounting on a
microscope slide.
Air drying specimens after mounting on a card
results in totally collapsed specimens and
alteration of subtle colour differences. Added to
this is the problem of leaching in ethyl alcohol,
e.g. after 12 months in 75% ethyl alcohol some
specimens become off-white. Gordh and Hall
(1979) reported excellent results using a critical
point dryer for specimen preparation of
Chalcidoidea before mounting. The procedure
involves collecting specimens into 75% ethyl
alcohol, slowly dehydrating with alcohol,
substitution of liquid CO^ for absolute alcohol
under pressure, raising the temperature until the
liquid CO; dissolves to gas and finally bleeding
off the CO; gas. The result with Melittobia was
beautifully inflated specimens with natural
colours which then were mounted on card
rectangles. Unfortunately. I have only had access
to a critical point dryer for 12 months, therefore
most of the specimens at my disposal for this
revision were air dried and others were slightly
leached because of storage in alcohol. Fortunately
some are in very good condition. Notes on
specimen preservation are given with the colour
notes on each species.
Microscope slide preparations were made for
each species depending upon availability of
aid in identification of species.
specimens, e.g. in the case of M. diopsisephia
only 2 2 2 , 1 $ exist and no slides were prepared.
Wings were first removed from the specimens and
placed in Euparal on a microscope slide. The head
and body were soaked in 10% NaOH until clear
then taken in 15 minute steps through 15% acetic
acid, distilled water after which they were
dehydrated in ethyl alcohol. When dehydrated
they were transferred to a 1:1 mixture of absolute
alcohol and Terpineol and placed under an
incandescent bulb until the ethyl alcohol had
evaporated. Antennae, heads and bodies were
separated and mounted in Euparal. This
procedure is discussed more fully by Prinsloo
(1980).
Figures
All figures except 1-3 were drawn from
cleared, microscope slide-mounted speiemens and
each has the scale indicated. They were drawn
with a camera lucida fitted to a Wild M20
compound microscope and constant
magnifications were used for the same part or
appendage of all species. Figures 1-3 were drawn
from freshly killed, dry-mounted specimens with
a camera lucida fitted to a Leitz TS
stereomicroscope .
TERMINOLOGY
The terminology used follows that of de V.
Graham (1969) except that the body is divided
into head, mesosoma (thorax and propodeum)
and metasoma (remainder of the abdomen).
Figures 1-13 serve to illustrate the general
Ill
MEMOIRS OF THE QUEENSLAND MUSEUM
morphology of Melittobia species and the
terminology used.
HISTORICAL RESUME
The genus Melittobia belongs to the chalcidoid
family Eulophidae, sub-family Tetrastichinae.
Ferriere (1960) grouped it with five other genera
of tetrastichine eulophids on the basis of their
common possession of a dorsoventrally flattened
thorax and large pronotum (Crataeopoides
Zinna, 1955; Crataepus Forster, 1878;
Aceratoneuromyia Girault, 1917; Pronotalia
Gradwell, 1957; Crataepiella T>oxncmc\nn\, 1956).
Peck, Boutek and Hoffer (1964) sank two of
these — considering Crataepoides a junior
synonym of Elachertus Spinola, 1811 and
Pronotalia a junior synonym of Crataepiella.
Domenichini (1966) divided the Tetrastichinae
into two tribes; the Tetrastichini with a genal
sulcus and Meiittobiini without a genal sulcus.
The latter comprised Melittobia,
Aceratoneuromyia, Crataepus and Crataepiella
— the four genera from Ferriere’s grouping
above. In the same year, BouCek described
Kocourekia and placed it with genera lacking a
genal sulcus. Boufcek (1977) described the genus
Tachinobia and although he made no reference
concerning its tribal placement it clearly fits with
these genera as the description mentions the lack
of a genal sulcus.
None of these treatments take into account
many of Girault’s genera except
Aceratoneuromyia. Of the large number of
genera described by Girault from Australia, it is
not known if any more could be placed in the
group lacking a genal sulcus. A revision of
Girault’s eulophid genera is presently being
undertaken by Dr. Boufcek. Clarification of this
point therefore rests with him.
In summary then, we have six tetrastichine
genera, Aceratoneuromyia, Crataepus,
Crataepiella, Kocourekia, Melittobia and
Tachinobia, grouped on the absence of a genal
sulcus. Crataepus separates out easily because of
the possession of a longitudinal median groove on
the mesoscutum and two fore tibial spurs. The six
genera show variation in the presence or absence
of facial and ocellar lines, and delimitation of the
vertex (a groove between the posterior ocelli and
the eyes). Facial lines converging from the vertex
or ocellar area to the scrobes occur in all except
Kocourekia. The ocellar area is delimited into an
ocellar plate by a groove in Melittobia and
Crataepiella. All genera except
Aceratoneuromyia have a delimited vertex.
Boutek (1977) in his description of Tachinobia
does not mention whether it has a delimited
vertex, but his figure of the dorsal aspect of the
female’s head shows it to be not delimited. In
ethyl alcohol preserved specimens of T. repanda,
the type-species, the vertex is clearly delimited. In
facial, ocellar and vertex grooves therefore
Melittobia most closely resembles Crataepiella.
The genera also vary in the number of grooves on
the scutellum. In Tachinobia there are no
grooves; Kocourekia and Aceratoneuromyia have
2 sub-lateral grooves; Melittobia, Crataepus and
Crataepiella have 2 sub-lateral and 2 sub-median
grooves.
Except for Kocourekia all of the genera are
known from both sexes. Of these Melittobia and
Tachinobia show pronounced sexual dimorphism
with the males greatly modified. Male Melittobia
are brachypterous with eyes reduced to a single
spot whereas male Tachinobia are apterous with
eyes reduced to several facets. In both, the
antennal scapes of the male are greatly enlarged.
Tachinobia males have a strongly inflated scape
with a large clear area ventrally (Fig. 15). The
scape of male Melittobia is also swollen but with a
ventral groove, a cup-shaped depression or hardly
grooved at all with a large ventral clear area (Figs
16-19). In the remaining genera, where males are
known, they closely resemble the females, are
macropterous and have scapes only slightly
modified, if at all.
There are other features which separate the
genera, e.g., the number of teeth on the
mandibles, the degree of flattening of the
prothorax, setation and so on, Boutek (1977)
gave a tentative key to the Tetrastichinae which
separates all six genera very well.
This taxonomic revision arose out of the
necessity to establish the identity of a species of
Melittobia whose biology and behaviour was
under study Dahms (1983a). Comparison of my
specimens with the type of the single Australian
species, M. australica Girault, 1912, showed that
they were conspecific. However, establishing the
validity of Girault’s species proved much more
difficult as the following account from the
literature reveals.
The generic name Melittobia Westwood (1847)
arose in an atmosphere of confusion because of
an argument between Mr G. Newport and Mr.
J.O. Westwood over the authorship of this new
genus. In 1849 the argument surfaced in the form
of a series of letters from the two antagonists to
the editors of the Annals and Magazine of
Natural History published in that journal. The
first letter was written by Newport in which he
claimed to have been studying the insect since
DAHMS: REVISION OF MELITTOBIA
273
FIGURES 1-3, Melittohia australica. 1 — Dorsal view female and male; 2 — side view male head; 3 — side view
female head and mesosoma.
274
MEMOIRS OF THE QUEENSLAND MUSEUM
1832 and implied plagiarism by Westwood.
However, Newport did not publish a description
of the species until 1849 when he called the genus
Anthophorabia and the species retusa after the
host Anthophora retusa Le Peletier and Serville.
His description was preceded by Westwood (1847)
in his ‘Introduciton to the Classification of
Insects’ where he mentioned the same species
from specimens forwarded to him by Mr Audouin
from the nests of Odynerus, Anthophora and
Osmia. Westwood also exhibited these specimens
in 1847 to the Entomological Society of London
and a brief descriptive note appeared with the
name Melittobia audouinii in that Society’s
Proceedings for 1847. He later published a more
formal description in the Proceedings of the
Unman Society in 1849.
Of the two generic names Melittobia stands,
but neither specific name stands because both had
been preceded by Walker who described the
species as acasta in 1839 and incorrectly assigned
it to the genus Cirrospilus basing his description
on a male that was in fact a female. From all of
this, the genus is Melittobia Westwood 1847 and
the type-species Cirrospilus acasta Walker 1939
by synonymy.
Although this confusion was removed fairly
early (Smith (1853); Dalla Torre (1898)) further
confusion has arisen at the species level. This
appears to be related to the relative uniformity of
the females. Ferriere (1933) thought it probable
that several of the described species were
synonyms of M. acasta or M. hawaiiensis.
Examination of females of the various species
shows they are difficult to separate and leads one
to agree with Ferriere. However, if the greatly
modified males are examined, it is clear there are
more than two species. Males do not emerge from
the host cell or puparium, therefore females are
more commonly encountered and their apparent
uniformity has led to many misidentifications,
not only in collections, but also in the literature,
e.g., the name M. chalybii Ashmead has been
applied to at least 2 species of Melittobia from
North America, neither of which is the true M.
chalybii.
Perusal of the literature revealed 13 described
species at the start of this revision. In addition
there are M. peloepi Ashmead, 1892 (published
ACASTA GROUP
acasta
evansi sp. nov.
scapata sp. nov.
digit at a sp. nov.
femorata sp. nov.
megachilis
chalybii
without a description which means it is a nomen
nudum) and M. hawaiiensis peles Perkins, 1907.
Of the 13 species, 9 are known from both sexes
and the remainder from females only. The initial
goal was to build up a collection of species based
on associated sexes and to use the males to
separate species. Decisions were checked against
notes on courtship behaviour generously provided
by Dr van den Assem from his own work on this
aspect of the genus. When the males were suitably
sorted, females were checked for reliable
morphological differences.
As a result of this study I am recognising 7 of
the previously described species M. clavicornis
(Cameron), M. acasta (Walker), M. chalybii
Ashmead, M. wegac/zZ/iy (Packard), M. australica
Girault, M. hawaiiensis Perkins and M.
bekiliensis Risbec. Two of the three species
described by Risbec were incorrectly placed by
him in the genus Melittobia. M. cosmopterygi 1
have transferred to Cirrospilus Westwood and M.
diopsisephila to Tachinobia BouCek. Four new
synonymies occur; M, japonica Masi becomes M.
clavicornis (Cameron), M. sceliphronidis
(Brethes) becomes M. hawaiiensis, M, strandi
Wolff and Krausse and M. fasciata (Newport)
become M. acasta (Walker). This leaves M.
osmiae Thompson and M. hawaiiensis peles
Perkins neither of which can be placed in the
absence of diagnostic descriptions, and I have not
been able to locate type-material.
Seven new species are described: M. scapata,
M. evansi, M. femorata, M. digitata (from North
America); M. assemi (Seychelles); M. sosui
(Japan); and a new species from Argentina for
which no types can be selected because the
specimens are fragmentary.
As a result of ethological work, van den Assem
(pers. comms., 1974-81) and van den Assem and
Maeta (1978, 1980) divide the genus into species
groups: acasta group, hawaiiensis group and
Mahe (assemi) group. They regard M. clavicornis
as the most primitive and keep it separated from
these groups. Using morphological grounds the
species separate easily on males into the same
groups and this is discussed more fully later. For
the purposes of the following discussions the
species groupings are as follows:
ASSEMI GROUP
assemi sp. nov.
sosui sp. nov.
bekiliensis
SP. NOV. Argentina
HA WAIIENSIS GROUP
hawaiiensis
australica
Kauai
DAHMS: REVISION OF MELITTOBIA
275
INTRODUCTORY MORPHOLOGY
Sexual dimorphism in the genus is so extreme
that the sexes cannot be associated by
morphology alone (Fig. 1). Males are greatly
modified and the modifications, involving
appendages and body regions, can be related to
the restriction of male activity to reproduction,
i.e. fighting, courting and mating.
Females are a fairly typical tetrastichine
eulophid form with fully developed wings and
eyes except in the case of second-form females
(see under Polymorphism Dahms, 1983a). They
show no gross modifications for courtship and
their most marked features are probably related
to host seeking in confined spaces and excavation
into or out of host enveloping membranes, e.g.,
enlarged prothorax and dorsoventral flattening of
the body. In addition, the head is
anteroposteriorally flattened and it articulates
with the prothorax close to the vertex which
allows the head to fold back almost in the same
plane as the body (Fig. 3). The antennae are
inserted low on the head and can be pushed
forwards when the head is in the flattened
position.
Males show a greater range of easily
discernible, reliable, morphological features for
separation of most species. They are
brachypterous, lack compound eyes and are less
pigmented than females (Fig. 5). These reductions
are related to their not emerging from the host
cocoon or puparium. Modifications have taken
place to heads, antennae and legs, and these are
related to the function of these parts in courtship
and fighting. Variations in these modifications
are related to specific variations in courtship
repertoires.
At the generic level, the male scape is
expanded, with a ventral groove or cup and is
open distally (Figs 16-19). The groove or cup is
used to house the female’s antenna during
courtship. Covering the distal opening of the
scape is a flap-like pedicel which is used in
manipulations of the female’s antenna. The
funicle is four-segmented with specific variation
in the relative proportions of the segments (Figs
12, 187, 190, 193, 196, 199, 202, 205, 208, 213,
216, 219). Females have a funicle composed of
three relatively uniform segments (Fig. 9).
In frontal aspect (Figs 5, 151-160) the male
head is shorter than that of the female (Figs
38-48); the eyes are reduced to single spots, and
the ocelli, although present, may be faint. In
lateral aspect (Fig. 2) the head is greatly inflated
in comparision to that of the female. This can be
related to the need for large muscles to carry out
the complicated courtship movements of the
enlarged scape and for effective use in combat of
the relatively larger mandibles. The prothorax
remains large but the remainder of the mesosoma
is reduced in correlation with the reduced wings
and inability to fly.
Legs play an important part in courtship
behaviour. The fore tarsi are used to hold the
female by the neck. Fusion of tarsal segments 3
and 4 on the fore leg has occurred in most species,
but there is fusion of 2, 3 and 4 in one group and
fusion of all segments in another (Figs 20-22).
There is a curving of the fused parts of segments 3
and 4 to accommodate the neck of the female.
Mid legs are used by all species during all or part
of the proceedings and the posterior, ventral
surface of the mid femur in all species bears a very
long fringe of setae. In some species a long fringe
of setae also occurs on the mid trochanter
ventrally. These setae appear to be used to brush
the female’s body during courtship. Their pattern
and distribution varies between species (Figs
220-230).
The genitalia of males show very little variation
except in size and are of little use taxonomically.
REVISION
Introductory Remarks
One of the major difficulties in dealing
taxonomically with the genus was the apparent
uniformity of females. This is mostly due to the
gross distortion of the body of females on drying
so that differences in the relative proportions of
head and mesosoma could not be appreciated.
Added to this has been the use by authors of
unreliable characters e.g. many aspects of
setation. This is again complicated by the
description of a few species from the female sex
only e.g. M. megachilis and M. japonica.
As discussed under Introductory Morphology,
males show a great range of morphological
features for separation of species and by
obtaining sexes bred out together it has been
possible to separate females of the different
species. Distortion of females on drying was
overcome by using a critical point dryer (Gordh
and Hall, 1979), and several useful proportional
features were revealed. In addition colours were
preserved without fading, provided the specimens
have not been kept for too long in ethyl alcohol.
Preparation of microscope slides has allowed
appreciation of differences in clypeal margins,
antennae, palps, mandibles and wing venation in
both sexes.
In spite of these techniques, difficulties still
exist. Colours vary with the method of
276
MEMOIRS OF THE QUEENSLAND MUSEUM
facial grooves
scrobes
toruli
clypeal margin
prothorax
mesoscutum mid lobe
mesoscutum lateral lobe
axilla
scutellum sublateral groove
cutellum submedian groove
dorsellum
FIGURES 4-8, Melittobia australica. 4 — Dorsal female mesosoma; 5 — Frontal aspect, male head; 6 — Female
mandible; 7 — Male mandible; 8 — Frontal aspect, female head; POL = posterior ocellar line, OOL = ocellar —
ocular line.
DAHMS: REVISION OF MELITTOBIA
111
FIGURES 9-14, Melittobia australica. 9 — Female antenna; 10 — Club segment 3, female antenna; 11 — Male
scape; 12 — male pedicel plus flagellum; 13 — Male wing; 14 — Female fore wing.
278
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 15-19, Male antennal scapes. 15 — Tachinobia repanda; 16 — Melittobia clavicornis; 17 — Melittobia
australica; 18 — Melittobia assemi (sp. nov.); 19 — Melittobia acasta.
FIGURES 20-22, Male fore legs, Melittobia spp. 20 — assemi (sp. nov.); 21 — sp. nov. Argentina; 22 — australica.
DAHMS: REVISION OF MELITTOBIA
279
preservation, for example, in older air-dried
specimens males tend to become a fairly uniform
brown whereas in fresh material subtle
infuscations and colour differences occur.
Prolonged preservation in alcohol causes leaching
of specimens converting dark browns to pale
yellows. Hence in this part of species descriptions
to follow mention is made of the method of
preservation as a reference point for comparison
and future descriptions. Although critical point
drying has removed the problem of shrinkage
there is still a tendency for the heads of females to
fold transversely along the line in front of the
ocellar plate. This occurs also in some males,
which gives a false impression of head shape and
proportions. For this reason all L:W proportions
were taken from slide mounted specimens. Care
was exercised in the preparation of slide material
not to over-inflate the specimens during clearing
and critical point dried specimens were used as a
check that this did not happen. The only L;W
proportion not taken from slides is that of the
prothorax.
In males, most of the characters that proved
useful in separating species have functional
significance in courtship. The following features
were of use: head shape (frontal and lateral),
clypeal margins, mandibles, palps, antennal
scapes (plus shape and position of pheromone
gland, Dahms (1983b)), relative proportions of
funicle segments, distribution of muitiporous
plate sensilla on the flagellum (MPS formula),
presence or absence of a short setal tuft ventrally
on the fore trochanters, presence or absence of a
very long setal tuft ventrally on the mid
trochanters, distribution and differentiation of
the long setal fringe ventrally on the mid femora,
shape of fore wings, shape of stigmal vein if
present. In general, the male mesosoma in dorsal
aspect lacks many of the sutures present in the
female except for M. clavicornis where all are
present.
In females, the differences useful for separating
species appear to have no functional differences
related to courtship except perhaps setation of the
eyes and the breadth of separation of the facial
grooves (Courtship Dahms, 1983a). The
following features were of use: head shape and
proportions in frontal aspect, eye setation, degree
of convergence of the frontal lines and the
distance apart of the upper arms, scrobe length,
clypeal margins, mandibles, palps, antennal scape
proportions, scape and pedicel colour, number of
muitiporous plate sensilla per segment of
flagellum (MPS formula), position of
subterminal seta on terminal nipple of club
segment 3, proportions of nipple, mesosomal
proportions and sculpture patterns in dorsal
aspect, number of setae on scutellum, wing shape,
ratio of submarginal to marginal vein length, and
shape of stigmal vein.
Stigmal veins of females show slight variations
in each species and the figures 127-139 are of the
most common shape. Apart from the usefulness
of the nipple on club segment 3 and the setae on
this nipple in females, the proportions of the
segment itself proved useful. Since the margin
between club segment 2 and 3 is undulating, the
proportion is worked out on the shortest length
and the ratio becomes shortest length to width.
The MPS formula is slightly variable ( ± I) and
the figures given in the descriptions represent the
maximum counted for 4 antennae in each species.
Care should be exercised when counting them as
those on the margins of a segment can be easily
overlooked and those which wrap around
segments (as in female funicular segments) may
be counted twice. In all cases descriptions are
based upon the type-form only (Polymorphism
Dahms, 1983a). Holotype and lectotype selection
where possible has been made on the male since,
of the sexes, this is the more distinctive.
References given at the start of each description
are taxonomic only. Those dealing with biology
occur in Dahms (1983a) as do references to
recorded hosts. The following abbreviations are
used for institutions.
ANIC: Australian National Insect Collection,
C.S.I.R.O., Canberra; Australia.
BM(NH): British Museum of Natural History,
London, England.
CU, NY: Cornell University, New York, USA.
DPIQ; Queensland Department of Primary
Industries, Brisbane.
DSIR: Department of Scientific and
Industrial Research, Christchurch,
New Zealand.
HDA: Hawaiian Department of Agriculture,
Honolulu, Hawaii.
KU: Kyushu University, Kyushu, Japan.
MCZ: Museum of Comparative Zoology,
Harvard, USA.
MDA: Museum de La Plata, La Plata,
Argentina
MNHP: Musee National d’Histoire Naturelle,
Paris, France.
NMC: National Museum (Nat. Hist.),
Prague, Czechoslavakia.
QM: Queensland Museum, Brisbane,
Australia.
280
MEMOIRS OF THE QUEENSLAND MUSEUM
UCR: University of California, Division of
Biological control, Riverside,
California, USA.
UG: University of Georgia, Athens,
Georgia, USA.
USNM: United States National Museum,
Washington D.C., USA.
NEW COMBINATIONS
Two species previously referred to the genus
Melittobia were found to be misplaced. These are
redescribed in their correct genera.
GENUS TACHINOBIA BOUCEK 1977
Type-species: Tachinobia repanda Boubek,
1977 : 27.
Tachinobia diopsisephila (Risbec, 1956)
(Figs 23, 24, 26-31)
Melittobia diopsisephila Risbec, 1956 : 118.
Tachinobia diopsisephila (Risbec). COMB NOV
Tachinobia gradwelli Boutek, 1977 : 28. SYN
NOV
This species, only known from females, can be
readily separated from the genus Melittobia by
the following features: head slightly wider than
long, vertex almost fiat, non-delimited ocellar
area, upper facial lines wider than posterior
ocelli, shape of clypeus and mandible, pronotum
shorter than wide, undifferentiated setae on the
mid lobe of mesoscutum, scutellum lacking
submedian and sublateral grooves, wings with
relatively longer setation and postmarginal vein
barely developed or absent.
There is considerable agreement between
Risbec’s specimens and the description of T
gradwelli by Boubek (1977); lower frontal lines
separated by twice the diameter of the median
ocellus and meeting distinctly below the middle of
the eyes, malar space almost equal to mouth
breadth, antennal toruli below lower eye margin;
setation on dorsal mesosoma more sparse and
shorter than T. repanda, the type species, fore
wings about 2.8 times longer than wide, stigmal
vein not much longer than the longest marginal
fringe. 1 have subsequently examined the
holotype of T. gradwelli and confirmed this
synonymy.
Dry and slide mounted specimens of T.
diopsisephila clearly show the vertex delimited by
a groove passing from the eyes to the ocelli. In his
description of the genus Tachinobia, Boubek
(1977) makes no mention of a groove across the
vertex and his figure of the dorsal head of T.
repanda does not show a groove. Slide mounted
material I have made of T. repanda do not show
this groove either. However, examination of ethyl
alcohol specimens of T. repanda clearly show this
groove (Fig. 25) and dry specimens of an
undescribed species (USNM Oollection) also have
this groove. This feature should therefore be
added to the generic description of Tachinobia.
MATERIAL EXAMINED
Holotype 5, 'Tachinobia gradwelli Boubek
(1977)’, ‘Venezuela, Estado Aragua Turmero 500
mts, xii.1948, H.E. Box’, ‘Hyper of Paratheresia
claripolopus (Wulp) ex Diatraea lineolata
(Walker) in Zea mays ’.
Four microscope slides containing a total of 24
specimens dry-mounted under coverslips sealed
with wax. I have remounted 6 specimens from 2
slides either on cards or in Euparal on the original
slides. Details of the labels and remounts are as
follows:
1) Melittobia diopsisephila Risbec ex dipt. par.
de Diopsis thoracica, Garoua 12.54,
Descamps 213’. (2 5 9 cleared and remounted
in Euparal on the original slide) (Garoua is in
the Cameroon, W. Africa).
2) Melittobia diopsisephila Risbec, epip. de
Diopsis thoracica ex dip. par., Descamps 200,
Garoua’. (15 5 5 under one coverslip and a
fragmentary head under a separate coverslip).
3) Melittobia diopsisephila Risbec ex pupa
Pachylopus sp.. Descamps 243, Garoua,
3.54’. (4 5$ remounted; 2 separately on
cards, one of which is selected as lectotype,*
the remaining 2 cleared and mounted in
Euparal on the original slide).
4) as (3). (7 9 5).
There is some disparity between Risbec’s
published notes and the information on his labels;
notably the lack of the name Steleocerus
lepidopus Becker on his labels and absence of the
label name Pachylopus in his published account.
Neave (1939) lists Pachylopus as a genus of
Coleoptera which seems doubtful as a host. A few
names before Pachylopus in Neave is Pachylopus
a genus of Chloropidae which is more likely to be
the true host. Steleocerus lepidopus is not
mentioned on the labels, but is a chloropid fly.
My examination of the series shows 24 specimens
plus some cylindrical debris which Risbec may
have counted to bring his total to 25. I feel that
these 4 slides represent his syntypical series of T.
diosisephila with some error with respect to the
chloropid host. From this series a lectotype has
been selected and card mounted on a pin. The
remaining specimens are on slides and selected as
DAHMS: REVISION OF MELITTOBIA
281
paraleclotypes. Labels have been attached
indicating these selections and the specimens
reside in the collections of the Musee National
d’Histoire Naturelle, Paris.
GENUS CIRROSPILUS WESTWOOD, 1832
Type-species: Cirrospilus elegantissimus
Westwood, 1832
Cirrospilus (Atoposomoidea) cosmopterygi
(Risbec)
(Figs 32-37)
Melittobia cosmopterygi Risbec, 1951 : 90
Cirrospilus (Atoposomoidea) cosmopterygi
(Risbec, 1951) COMB. NOV.
Females of this species are readily separated
from the genus Melittobia by their
colouration, lack of facial grooves, non-
delimited ocellar area, non-emarginate
clypeus, antennal insertions above lower
margins of eyes, 2 segmented funicle,
terminal style of club without a terminal seta
and with more than 3 subterminal setae,
scutellum squarish with only 2 longitudinal
grooves, wings with admarginal setae, long
marginal fringe and relatively longer discal
ciliation on the fore wing.
The black and yellow' colouration is typical of
Cirrospilus. Using Peck, BouCek and Hoffer
(1964) the species keys readily to Cirrospilus
(Atoposomoidea). De V. Graham’s key (1959)
also easily places the species in Cirrospilus. Risbec
(1951) in his collecting data states that C
cosmopterygi was bred from leaf mining
lepidopterous larvae; a typical host of
Cirrospilus. This type of host is not recorded for
Melittobia,
There has been some difference of opinion as to
the status of Atoposomoidea, e.g. Delucchi
(1958) treats Atoposomoidea as a genus. Other
workers, e.g. De V. Graham (1959), Boufcek
(1959) and Kerrich (1969) place it as a sub-genus
of Cirrospilus. This question is beyond the scope
of the present work and it has been decided to
follow the direction of BouCek (1959) and leave it
as a sub-genus of Cirrospilus.
The following redescriplion has been based
upon the syntypical material of Risbec; no other
specimens being available. The two females are
not cleared and are badly flattened on a
microscope slide.
Female: 1.4 mm long; head, antennae brown;
mesosoma yellow with black markings (Fig.
33); legs yellow; wings hyaline; metasoma
yellow with 2 transverse black bands dividing
metasoma equally into thirds.
Head in frontal aspect (Fig. 32); 0.3 mm
wide, length to width about 1:1; vertex
collapsed, clearly elevated above eyes, not
immargined, not vaulted as in
Zagrammosoma Schulz; POL approximately
equals OOL. Eyes dark, probably red in life,
oval, sparsely pilose, closer to vertex than
clypeus. Clypeus concave, not emarginate.
Antennae (Figs. 34, 36); inserted above level
of lower eye margins; scape cylindrical, 3.25
times longer than wide, slightly arched in
lateral view; pedicel expanded distally, equal
to funicle 1; ring joint compound, of several
lamellae; funicle 2-segmented, I and 2
cylindrical, equal in length; club
3-segmented, 1 and 2 cylindrical, equal in
length, 3 shorter, conical bearing a terminal
style with 5 subterminal setae but without a
terminal seta. Head lateral and dorsal aspects
not visible.
Mesosoma (Fig. 33) in dorsal aspect;
pronotum large, slightly wider than long,
campanulate; posterior border with four
large setae, inner pair closer to one another
than to pair at posterior lateral angles.
Mesoscutum with clearly defined, sigmoidal
parapsidial sutures; mid lobe with 2 pairs of
long setae, anterior pair finer and closer
together than posterior pair; axillae
advanced; scutellum large, wider than long,
more angular than in Melittobia, posterior
margin convex; one pair of sublatera! grooves
anteriorly in line with posterior parapsidial
sutures; 2 pairs of long setae situated on the
lateral lobes; 1 pair about mid-way and 1 pair
on posterior margin, Dorsellum triangular,
apex directed posteriorly, base anteriorly
convex. Propodeum appears shallowly
inclined, without carinae, length at mid line
approximately 1/6 width at posterior lateral
angles. Wings hyaline (Fig. 35); fore wings
about 2.6 times longer than wide, costal
margin almost straight, slightly curved at
junction of parastigmal and marginal veins;
submarginal 0.27 mm long, postmarginal
0.04 mm long, sligmal 0.06 mm long;
si\bmarginal veins with 6 long, erect setae in
both specimens; costal cell with 9-10 setae
ventrally plus two anterolateral setae
dorsally; discal ciliation even, moderately
long; 8 long, admarginal setae
posteroventrally of marginal vein; marginal
fringe long. Hind wing narrow, 0.75 mm long
X 0.09 mm wide; apex acute; marginal fringe
longer than fore wing marginal fringe;
282
MEMOIRS OF THE QUEENSLAND MUSEUM
stigmal vein absent. Lateral aspect not
visible.
Metasoma in dorsal aspect, ovoid, 0.69 mm
long X 0.35 mm wide, more acute
posteriorly; ovipositor over half length of
metasoma, not extruded. Lateral aspect not
visible.
Male: Unknown
MATERIAL EXAMINED:
One slide bearing 2 5 $ , 2 pupal cuticles plus
one pupa 'Melittobia cosmopterygi Risbec,
Syntypes; Eulophinae, G.B., ex Cosmopteryx
attennatella. III .77’.
In his paper, Risbec (1951) gives the locality
as M’ Bambey, Senegal, and mentions further
specimens from Cosmopteryx in Niebe.
However, he goes on to state that the Niebe
material was not in his possession at the time
of description. I have been unable to locate
the Niebe material and have taken the slide
above to include the entire syntypical series.
The two specimens on this slide fit Risbec’s
description of C. cosmopterygi. The
specimen closest to the Risbec label is selected
as the lectotype and the other as the
paralectotype. The slide has been labelled
accordingly and it is to be found in the
collections of the Musee National d’Histoire
Naturelle, Paris.
GENERIC DIAGNOSIS
GENUS MELITTOBIA WESTWOOD, 1847
Type-species :C/r/‘05/7//«.s’ acasta Walker, 1839
by synonymy with Melittobia audouinii
Westwood, 1847
Melittobia Westwood, 1847. Type-species,
Melittobia acasta (Walker, 1839) by
synonymy.
Anthophorabia Newport, 1849. Type-species,
Anthophorabia retusa Newport by
monotypy.
Philopison Cameron, 1908. Type-species,
Philopison clavicornis Cameron, 1908 by
monotypy.
Sphecophagus Brethes, 1910. Type-species,
Sphecophagus sceliphronidis Brethes, 1910
by monotypy.
Sphecophilus Brethes, 1910. 311, new name
proposed for Sphecophagus, Brethes.
Generic Description:
Female: Black to dark brown, shining in most
species; moderately setose; 1-1.6 mm long.
Head in frontal aspect (Figs 38-48); variable
in shape, about as long as wide with
alutaceous sculpturing in most species; vertex
elevated above eyes, rounded; facial grooves
present, converging ventrally to antennal
scrobes, distance between upper arms and
degree of convergence variable; antennal
scrobes shallow, area between slightly raised;
clypeus mostly bilobed, rarely truncate
emarginate (Figs 59-69); mandibles (Figs
49-58) tridentate, anterior tooth the largest,
acute; palps I -segmented (Figs 70-81). In
lateral aspect (Fig. 3); very narrow,
converging ventrally; eyes oval, longer than
wide; genae well developed, genal sulci
absent. In dorsal aspect ocelli arranged in a
shallow triangle positioned on an ocellar plate
delimited by grooves; vertex with transverse
grooves connecting the ocellar plate to the
eyes. Antennae (Figs 82-101); 8 segmented
plus 1 compound ring joint, inserted below
eyes; toruli closer to one another than to eyes;
scape nearly reaching top of eyes, elongate,
slighly expanded distally, dorsoventrally
flattened, ventrally slightly concave; pedicel
pyriform, sub-equal to funicle 1; ring joint
thin, compound, of 4 lamellae; flagellum
dorsoventrally flattened; funicle
3 -segmented, segments sub-equal, about as
long as wide, all bear MPS; club
3-segmented, all bearing MPS, segment 2
longest, segment 3 shortest (Figs 102-114),
conical with terminal nipple, in some species
nearly as long as segment 3; terminal nipple
with 2-3 setae shorter than nipple, 1 terminal,
the others of variable position from 1/2 way
down nipple to base.
Mesosoma in lateral aspect (Fig. 3) relatively
flat, prothorax triangular. In dorsal aspect
(Figs 1 , 4) prothorax large, slightly wider than
long, campanulate, posteriorly fringed with
relatively long, recurved setae; mesoscutum
without a longitudinal median groove,
parapsidial sutures well defined; mid lobe
contracting posteriorly, posterior margin
truncate with 1 large recurved seta at each
posterior lateral angle; axillae advanced,
acute anteriorly, well defined on mesoscutal
and scutellar margins; scutellum transverse,
submedian and sublateral grooves present,
evenly spaced, inner lobe bearing at least 1
pair (rarely more) of large, recurved setae
along outer margin of each submedian
groove; dorsellum well defined in non-
collapsed specimens, ovoid; propodeum
DAHMS: REVISION OF MELITTOBIA
283
normally developed, wider than long, not
steeply inclined, without a median,
longitudinal carina; spiracles freely exposed,
round; legs normal, fore and hind coxae the
longest, hind coxae the broadest; fore wings
(Figs 115-126); norma! tetrastichine type,
hyaline, discal ciliation evenly scattered, setal
lines occur along basal, cubital and
subcubital vein positions; submarginal vein
shorter than marginal, with 4-6 long setae;
postmarginal vein poorly developed;
marginal vein fringed with long setae which
extend to end of postmarginal vein; remaider
of wing around apex to distal subcubital setal
line fringed with short setae, the fringe
longest on posteroapical margin; stigmal vein
short (Figs 127-139), just longer than
postmarginal; uncus well developed; hind
wings narrow, apex acute; poslmarginal and
marginal vein not developed; fringed with
very long setae from end of marginal vein
around apex and along posterior margin.
Metasoma elongate, sides sub-parallel,
segments of fairly even size; ovipositor not
exserted.
Male: Brachypterous, 1.0- 1.5 mm long, dark
brown to honey yellow in colour.
Head in frontal aspect (Figs 151-160) variable
in shape, inflated, wider than long in most
species, facial lines absent; clypeus mostly
bilobed, rarely truncate emarginate;
mandibles tridentate, anterior tooth as in
female but much larger (Figs 161-173); palps
1 segmented (Figs 174-184). In lateral aspect
(Fig. 2) greatly inflated; eyes reduced to scar-
like spots; genal sulcus absent. In dorsal
aspect (Fig. 1) ocelli variously reduced;
delimiting lines around ocelli and across
vertex absent in most species. Antennae (Figs
185-219) greatly modified, 9 segmented, in
some a 10th appearing at ring joint (Fig. 202);
scape enormously developed, longer than
pedicel plus funicle, pyriform to
subpyriform, ventrally with either a groove
running full length or a distal cup-shaped
depression; pedicel produced laterally to
form a cap over the distal end of the scape
groove or cup; ring segment, compound, thin
or in some cases expanded; funicle 4
segmented, with or without plate organs, size
and shape of segments variable; club
3-segmented, segments of variable size,
terminal segment with a small terminal nipple
hardly differentiated in some species;
terminal nipple with 2 setae.
Mesosoma of different proportions from
female (Fig 1); in lateral aspect flattened,
prothorax triangular. In dorsal aspect
prothorax large, wider than long,
campanulate, posteriorly fringed with
relatively long recurved setae, mesoscutum
reduced, much wider than long, parapsidial
sutures in most species indefinite anteriorly; 1
pair of strong recurved setae on posterior
margin; axillae poorly delimited in most
species; scutelium in most species without
submedian or sublateral grooves, with 4
(rarely more) stiff recurved setae; dorsellum
well defined in non-collapsed specimens,
more rounded than in female; propodeum
well developed, not steeply inclined, smooth,
median longitudinal carina absent; spiracles
freely exposed, rounded; legs sturdier than in
female, fore coxae broad, fore tarsi with at
least segments 3 and 4 fused; mid tibiae and
in some species mid trochanters ventrally
fringed with long setae (Figs 220-230); fore
wings reduced (Figs 231-241), of variable
width, never longer than mesosoma; marginal
vein longer than submarginal; postmarginal
and stigmal veins poorly developed; long stiff
setae along submarginal and marginal veins,
hind wings very reduced, elongate;
postmarginal and stigmal veins absent,
setation reduced.
Metasoma (Fig. 1) in lateral aspect arched; in
dorsal aspect ovoid in life becoming flattened
on drying; segments evenly proportioned.
Keys for Identification
Females
1) Facial grooves running separately to scrobes
(Figs. 38-45, 48) 2
Facial grooves meeting just above middle of
eyes then passing as one line to scrobes, upper
arm equal to or wider than POL (Figs. 46, 47)
10
2) Scape and pedicel dark, concolorous with
flagellum (Figs. 83, 84), fore wing with costal
margin almost straight (Fig. 116)
acasta (Walker)
Scape and pedicel paler than flagellum, fore
wing coastal margin noticeably bent at
junction with parastigmal vein 3
3) Head broad, length to genal width about
1.1:1 (Figs. 38,41) 4
Head relatively narrow, length to genal
width greater than 1.1:1 5
284
MEMOIRS OF THE QUEENSLAND MUSEUM
4) Upper arms of facial grooves widely
separated, approximately equal to POL,
lower arms of facial grooves separated by a
distance equal to the diameter of the median
ocelleus (Fig. 38), clypeal lobes with small
lateral undulations (Fig. 59), subterminal seta
on antennal nipple situated basally (Fig. 102)
clavicornis (Cameron)
Upper and lower arms of facial grooves much
closer than above (Fig. 41), clypeal lobes
without lateral undulations (Fig. 62),
subterminal seta on antennal nipple not basal
(Fig. 105) scapataSP.^OV .
5) Eyes densely clothed with long setae (Figs. 44,
48) 6
Eyes relatively bare, with a few short
scattered setae 7
6) Head and mesosoma densely setose (Fig. 44);
clypeal margin bilobed, lobes narrow each
with a small, lateral, lobe-Hke undulation
(Fig. 65); nipple on club segment 3 with 1 sub-
terminal seta (Fig. 108).... chdlybii
Head and mesosoma not as densely setose
(Fig. 48); clypeal margin bilobed, lobes
broad, without a small, lateral, lobe-like
undulation (Fig. 69); nipple on club segment
3 with 2 subterminal setae (Fig. 114)
sp. nov. Argentina
7) Terminal seta on postmarginal vein
noticeably longer than those on marginal vein
(Figs. 129, 131), head sculpture normal,
surface shining 8
Terminal seta on postmarginal vein not
noticeably longer than those on marginal vein
(Figs. 132, 134), head sculpture fine, surface
dull, shagreened 9
8) Clypeal margin bilobed each lobe without a
lateral, lobe-like undulation (Fig. 61);
sculpture pattern on mesoscutum and
scutellum very open (Fig. 142); mid lobe of
scutellum broad, L:W 1.4:1; MPS formula
on flagellum 355:553 evansi SP. NOV.
Clypeal margin bilobed, each lobe with a
lateral, lobe-like undulation (Fig. 63),
sculpture pattern on mesoscutum and
scutellum mid lobes less open particularly on
scutellum (Fig. 144);- mid lobe of scutellum
narrow, L:W 1.9:1 MPS formula on
flagellum 567-653 digitata SP. NOV.
9) Scape and pedicel yellow-brown, dorsally
dark, reddish-brown (Fig. 90); nipple on club
segment 3 long, L:W 4:1, subterminal seta on
antennal nipple not basal (Fig. 107)
femorataSP. NOV.
Scape and pedicel yellow-brown, not strongly
darkened dorsally (Fig. 93); nipple on club
segment 3 short, L:W 2.5:1, subterminal seta
almost basal (Fig. 109)
Megachilis (Packard)
10) Clypeus bilobed (Fig. 67); upper arms of
facial grooves wider than POL (Fig. 47) 11
Clypeus truncate emarginate (Fig. 68); upper
facial grooves as wide as POL (Fig. 46)
. . . australica Girault and hawaiiensis Perkins
11) Submarginal vein with 5 setae; proximal pair
distinctly shorter than rest . assemi SP. NOV.
Submarginal vein with 5 long setae of equal
length sosuiSP. NOV.
Males
1) Scape ventrally with a distal cup-shaped
depression (Fig. 19) 8
Scape ventrally with a longitudinal groove
(Figs. 16-17) 2
2) Head large (Fig. 151) dark brown; scape
yellow, distinctly club-shaped (Figs. 185,
186), ventral groove shallow; glandular area
large, circular clavicornis (Cameron)
Head and scape concolorous, yellow-brown;
not distinctly club-shaped as (Fig. 151), scape
groove deep; glandular area not circular .... 3
3) Head transversely elliptical, lateral margins
broadly rounded (Fig. 158); funicular
segmental proportions (Fig. 208), 1 the
smallest, narrow, 2-1-3 the largest, 4 cup-
shaped, closely applied to club segment 1;
clypeus without lobes 4
Head more or less rectangular (Figs. 159,
160); funicular segmental proportions not as
above 5
4) Flange overhanging scape groove on the side
of pedicel attachment with up to 5 setae, only
1 or 2 of which are on the inner margin, 1-2
setae on the proximal floor of groove (Fig.
206) australica Girault
Flange overhanging scape groove with more
than 5 setae of which most are on the inner
margin, flange longer than australica or
Kauai, 4 setae on the proximal floor of
groove (Fig. 209) hawaiiensis Perkins
Flange overhanging scape groove with more
than 5 setae spread evenly along flange, most
of which are on the inner margin, 2 setae on
the proximal floor of groove, (Fig. 210)
Kauai
DAHMS: REVISION OF MELITTOBIA
285
5) Fore wing apex acute (Fig. 239) 6
Fore wing apex rounded (Fig. 241) 7
6) Mid femoral fringe sparse (Fig. 228)
assemi SF . NOV ,
Mid femoral fringe denser (Fig. 230)
505W/SP. NOV.
7) Mandibles very broad (Fig. 173), projecting
well below clypeus when closed; face below
toruli with tufts of long, stiff setae (Fig. 160);
Maxillary palp broad, distally excavated (Fig.
184); clypeus with 2 broad lobes
sp. nov. Argentina
Mandibles not projecting well below clypeus
when closed; face below toruli without stiff
setae, maxillary palp tapered distally as
assemi (Fig. 182); clypeal margin with narrow
lobes as assemi (Fig. 159) . . bekiliensis Risbec
8) Distal scape margin deeply excavated to
produce a thumb-like appendage (Figs. 197,
198) digitataSF. NOV.
Distal scape margin not as deeply excavated
9
9) Distal scape strongly oblique with a broad
excavation overhung by a long setal fringe
(Figs. 188, 189) (Walker)
Distal scape not strongly oblique 10
10) First funicular segment only slightly wider
than segments 2-3 (Figs. 193, 196) 11
First funicular segment much wider than
segments 2-3; the distal ring-joint expanded
slightly to give a small segment before funicle
1 (Figs. 202, 205) 12
11) Fore wing relatively narrow, L:W 2.6:1;
costal cell long, L;W 8:1 (Fig. 233); marginal
to submarginal vein length 1.4:1; mid femoral
fringe relatively even, longest setae about as
wide as femur (Fig. 223); head narrowed
above eye spots (Fig. 153) L:W 1:1 evansi
SP. NOV.
Fore wing relatively broad, L:W 2.4:1;
coastal cell shorter, L:W 7.5:1; (Fig. 234)
marginal to submarginal vein length 1:1; mid
femoral fringe of uneven length, proximal
half shorter, just shorter than width of
femur, distal half longer, about 1.5 times
width of femur (Fig. 222); Head not
contracted above eye spots, wider than long,
L:W 1:1.3 (Fig. 154) SP. NOV.
12) Fore wing relatively narrow (Fig. 237), L:W
2.9:1, posterior margin almost straight;
coastal cell long, L;W 11.7:1; mid femoral
fringe uneven, proximal 1/3 shorter than
width of femur, distal 2/3 about as wide as
femur (Fig. 226); head densely setose, genae
below^ eye spots contracting to clypeal margin
(Fig. 157) chalybii Ashmead
Fore wing relatively broad, L:W 2.6:1; costal
cell short, L:W 7.6:1 (Fig. 236); mid femoral
fringe uneven, proximal 1/2 of fringe about
as long as width of femur, distal 1/2 of fringe
dense, extremely long nearly 2 x width of
femur (Fig. 225); head not as densely setose,
genal margins below eye spots straight,
parallel, not contracting towards clypeus
(Fig. 156) femorataSF. NOV.
SPECIES DESCRIPTIONS
Melittobia clavicornis (Cameron)
(Figs 38, 49, 59, 70, 82, 102, 115, 127, 140, 151,
161, 174, 185, 186, 187, 220 231)
Philopison clavicornis Cameron, 1908 : 559.
Melittobia clavicornis : Ferriere, 1933 : 103.
Melittobia japonica Masi, 1966 : 38, SYN NOV.
Melittobia japonica : Iwata and Tachikawa, 1966
: 6 .
Melittobia Japonica : Domenichini, 1966 : 57.
TYPE SPECIMENS;
I have examined the syntypical series of
Cameron which is in the collections of the British
Museum (Natural History) London. Details are in
the MATERIAL EXAMINED section. I have
examined also the syntypical series of M. japonica
Masi which are in the collections of the Kyushu
University, Kyushu Japan. From both these
syntypical series I have selected lectotypes. Details
occur in the MATERIAL EXAMINED section.
DISTRIBUTION:
Borneo, Ceylon, Japan (= species 3, van den
Assem and Maeta (1978) = Species 1 van den
Assem, Bosch and Prooy (1982)).
DESCRIPTION:
Female: Critical point dried specimens 1.3-1. 4
mm long. Head, antennal flagellum,
mesosoma, coxae dark brown; trochanters,
proximal 2/3 femora, metasoma paler
brown; scape, pedicel, remainder of legs
yellow-brown.
Head in frontal aspect (Fig. 38) relatively
broad, length to genal width 1.1:1; genal-
clypeal margin broadly rounded; clypeal
margin (Fig. 59) bilobed, each lobe with a
small lateral lobe-like undulation; eyes
286
MEMOIRS OF THE QUEENSLAND MUSEUM
relatively bare, with a few short scattered
setae. Facial grooves remaining separate to
meet scrobes well below middle of eyes, upper
arms widely separated, maximum distance
between arms 3.5 times diameter of median
ocellus, greater than POL, converging
gradually to scrobes remaining broadly
separated but contracting suddenly just
before scrobes. Scrobes relatively short,
scrobe to eye length 1:3. Mandibles (Fig. 49);
anterior tooth long, narrow, 2 and 3 well
defined, 2 the more definite. Maxillary palps
(Fig. 70) elongate, cylindrical, L:W 5:1.
Antennae (Fig. 82); scape L:W 3.7:1; MPS
formula on flagellum 354:773; club segment 3
(Fig. 102) shortest length to width 1:2.5;
nipple short, broad, L:W 3:1, barely reaching
above the MPS; subterminal seta basal.
Mesosoma in dorsal aspect. Setation
relatively long. Prothorax wider than long,
L:W 1:2. Posterior margin of mesoscutum
mid lobe 1 .4 times wider than anterior margin
of scutellum. Scutellum mid lobe L:W 1.7:1;
1 pair of setae on each submedian lobe,
posterior seta almost on hind margin.
Sculpture pattern on mid lobes of
mesoscutum and scutellum (Fig. 140).
Propodeum in dry specimens rectangular,
wider than long; posterior margin truncate
emarginale; posterolateral angles 90°. Fore
wings (Fig. 115); costal margin noticably
angled at junction with parastigmal vein;
L:W 2.2:1; marginal to submarginal vein
length 1.3:1; marginal to stigmal vein length
3.4:1; submarginal to stigmal vein length
2.7:1; stigmal vein (Fig. 127); terminal seta on
postmarginal vein as long as those on
marginal vein.
Male: Critical point dried specimens 1.4 mm
long. Head, body and legs dark brown, head
darkest; scape and legs pale yellow-brown.
Head in frontal aspect (Fig. 151) large,
slightly wider than long, L;W 1:1.1; vertex
depressed medially; lateral margins more or
less broadly rounded, slightly contracted
below eye spots; clypeal margin bilobed,
lobes broad. Mandibles (Fig. 161); anterior
tooth of moderate length, relatively close to
second, third tooth poorly defined. Maxillary
palps (Fig. 174) elongate, cylindrical, slightly
curved, L:W 4.8:1. Antennae (Figs.
185-187); scape strongly club-shaped, scape
to head length 1:1.8, L:W 1.6:1; ventral
surface with a shallow groove, distally
truncate not excavated, glandular area large,
circular; funicular segment proportions (Fig.
187) 1 very large, L;W 1:1.8, 2-4 sub-equal,
relatively small, about as wide as length of
segment 1 ; MPS formula on flagellum
0122:342.
Mesosoma in dorsal aspect. Prothorax L:W
1:2. Mesoscutum with clearly defined
parapsidial sutures; axillae well defined.
Scutellum without submedian grooves;
sublateral grooves present; 2 pair of large
setae present, situated as in female.
Fore trochanters without a ventral tuft of
short, stiff setae; tarsal segments 3 + 4 fused.
Mid legs (Fig. 220); trochanters without a
dense tuft of long, fine setae; femora with a
cluster of long, fine setae distally, a few short
setae proximally; mid femur L:W 2.7:1; mid
tibia relatively short and wide, shape quite
distinctive, L:W 2.3:1; mid tarsal joints
unfused. Fore wings (Fig. 231) broad, L:W
2.5:1; marginal to submarginal vein length
I. 2:1; stigmal vein well developed; costal cell
L:W 9:1, costal margin slightly arched.
MATERIAL EXAMINED:
BM(NH) I on a card, minus wings; left
antennal flagellum separated; dark blue BM
‘LECTOTYPE’ label, ‘Kuching Nov. 07
J. H.’, ‘This also from Pison sarawakensis
cocoons’, "Philopison ciavicornis Cameron
Type Borneo’, ‘B.M. TYPE HYM 5.1354’.
LECTOTYPE.
1 ^ glued ventral surface down on a card
(only 1 leg attached, and 1 leg glued
separately on the card; only 1 fore wing
present — torn); blue BM ‘PARA-
LECTOTYPE’ label, ‘Kuching, Nov. 07,
J.H. P. Cameron Coll., 19I4-I00’,
'Philopison ciavicornis Cam., Type, Borneo’.
PARALECTOTYPE.
1 V glued on a card, intact; blue BM
‘PARALECTOTYPE’ label, ^Philopison
ciavicornis Cam.’, ‘Kuching J.H.’, ‘Borneo,
J. Hewitt, 1910-380’. PARALECTOTYPE.
2 ? V glued ventral surface down on a card,
both incomplete (outer minus metasoma, all
wings except I hind wing separated in the
glue; 1 antenna complete, the other
separated, fragmentary ; inner complete
except for metasoma); blue BM
‘PARALECTOTYPES’ label, ‘Bred from
cocoons of Pison sarawakensis Kuching,
Nov. 07, J.H,’ ‘This seems to be same as my
J.19 is it not?’, ‘A.B. 2’, *Philopison
ciavicornis Cam., Type, Borneo’, ‘P.
DAHMS: REVISION OF MELITTOBIA
287
Cameron collection, 1914-110’. PARA-
LECTOTYPES.
1 9 on a card buried in glue; blue BM
TARALECTOTYPE’ label, ^Philopison
clavicornis. Cam. Type Borneo’, ‘P.
Cameron Coll. 1914-110’. PARALECTO-
TYPE.
I 2 on a card, intact, blue BM
TARALECTOTYPE’ label, ‘Kuching J.H.
[this label has July crossed out]’, ‘P.
Cameron Coll. 1914-110’.
PARALECTOTYPE.
II microscope slides with various parts of
both sexes as follows:
Slide 1 — 4 coverslips containing a
disembered S . 'Melittobia (Philopison)
clavicornis. Cam., 'Pelopaeus madra-
spatarum Borneo, Kuching. (Edinburgh
Mus.) J. Hewitt Coll.’ PARALECTOTYPE.
Slide 2 — 1 coverslip containing a $ head
minus mouthparts and antennae. ‘Head,
Meliitobia (Philopison) clavicornis. Cam.’,
‘Kuching, Borneo Nov. 1907 J. Hewitt
1910-380’. PARALECTOTYPE.
Slide 3 — 3 coverslips containing S antennae
and mouthparts. ‘Mandibles, Trophi, &
Antennae, i . Meliitobia (Philopison)
clavicornis. Cam.’, ‘Kuching, Borneo, Nov.
1907 J. Hewitt 1910-380’. PARA-
LECTOTYPE.
Slide 4 — 1 coverslip containing a $
mesosoma + metasoma; minus prothorax,
legs, wings and genitalia. ‘Meso-Metathorax
& Abdomen, $ Meliitobia (Philopison)
clavicornis. Cam.’, ‘Kuching, Borneo Nov.
1907 J. Hewitt 1910-380.’ PARA-
LECTOTYPE.
Slide 5 — 1 coverslip containing S wings (2
pairs). ‘Wings, $. Meliitobia (Philopison)
clavicornis. Cam.’, ‘Kuching, Borneo Nov.
1907 J. Hewitt 1910-380’. PARA-
LECTOTYPE.
Slide 6 — 1 coverslip containing 1 set of $
legs. ‘Legs, ct. Meliitobia (Philopison)
clavicornis Cam.’, ‘Kuching, Borneo, Nov.
1907 J. Hewitt 1910-380’. PARA-
LECTOTYPE.
Slide 7 — 2 coverslips containing $ prothorax
and genitalia. ‘Prothorax & Genitalia, S.
Meliitobia (Philopison) clavicornis Cam.’,
‘Kuching, Borneo Nov. 1907 J. Hewitt
1910-380’. PARALECTOTYPE.
Slide 8 — 1 large coverslip containing an
intact 2 . Meliitobia (Philopison)
clavicornis. Cam. CO-TYPE’, ‘Kuching,
Borneo J. Hewitt, 1909-182’. PARA-
LECTOTYPE.
Slide 9 — 4 coverslips containing 1 2 head
with separated antennae and mouthparts.
‘Head, Antennae, Mandibles & Trophi 2
Meliitobia (Philopison) clavicornis Cam.’,
‘Kuching, Borneo J. Hewitt 1909-182’.
PARALECTOTYPE.
Slide 10 — 4 coverslips containing 1 2
mesosoma (minus legs and wings), metasoma
and ovipositor. ‘Prothorax, Mesothroax,
Metathorax, Propodeum, Abdomen &
Ovipositor 2 Meliitobia (Philopison)
clavicornis. Cam.’, ‘Kuching, Borneo J.
Hewitt 1909-182L PARALECTOTYPE.
Slide 11 — 2 coverslips containing 2 wings (2
pairs) and legs (1 set). ‘Legs & Wings, 2
Meliitobia (Philopison) clavicornis Cam.’,
‘Kuching, Borneo J. Hewitt 1909-182’.
PARALECTOTYPE.
The parts on slides 2-7 all make up a single $
and were probably from a single specimen,
similarly the parts on slides 9-11 may all be
from a single 2.
USNM 2 29 intact glued on separate cards on
separate pins in Dr. K. Krombein’s voucher
specimen collection and labelled — ‘SRI
LANKA : Colombo, 3.x. 1977, K.V.
Krombein, 10377 A, ex Paraleptomenes
mephitis'; a pink label, ‘10377A’.
1 2 1 ^ on separate pins with above data; $
also labelled "Meliitobia clavicornis (Cam.),
det. Z. Bou(:ek, 1978’.
KU 6 points on one pin, 5 each bearing 1 2
(upper minus head) and 1 bearing plant
material. These are labelled, "Melittobia
japonica MS., Cotypi!, det. L. Masi’. From
this syntypical series I have selected the
female on the bottom point as the lectotype
and marked its point with red. The remaining
4 specimens have been selected as
paralectotypes. Labels were applied
indicating these selections.
QM 2 microscope slides, I with 2 2 2 the other
with 1 ^ i and both labelled, "Meliitobia
clavicornis (Cam.) E. Dahms det. 1981,
Chiisageta-gun, Nanango Pref., Japan, Jan.
1977, T. Kitamura ex Trypoxylon malaiseV .
In addition to these there is a series of card-
mounted specimens each with 1 S 4 2 2 ;
‘Chiisagata-gun, Nanango Pref., Japan, Jan.
1977, T. Kitamura ex Trypoxylon malaiseV,
" Meliitobia clavicornis (Cdxnexon), E. Dahms
det. 1981’. These are all critical point dried
specimens and one card has been deposited in
288
MEMOIRS OF THE QUEENSLAND MUSEUM
each of the following institutions: ANIC,
BM(NH), DSIR, KU, MDA, MNHP, NMC,
QM, UCR and USNM.
NOTES: The figures of this species were taken
from the QM slides of specimens from Japan.
The syntypical slides came to hand after
figures were assembled.
Melittobia acasta (Walker)
(Figs 39, 50, 60, 71, 83. 84, 103, 116, 128, 141,
152, 162, 175, 188, 189, 190, 221, 232)
Cirrospilus acasta Walker, 1839 : 328.
Melittobia audouinii Westwood. 1840 : 160.
Melittobia audouinii : Westwood, 1847 : 18.
Anthophorabia retusa'Hew^X^ovi, 1849a : 183.
Melittobia audouinii : Westwood, 1849a : 295.
Melittobia audouinii : Westwood, 1849b : 37.
Melittobia audouinii : Westwood, 1849c : 65.
Anthophorobia retusa : Westwood, 1849c : 65.
Anthophorabia retusa : Newport, 1849b : 513.
Melittobia audouinii : Newport, 1849b : 514.
Melittobia audouinii : Westwood, 1849d : 39.
Anthophorabia retusa : Newport, 1849c : 122.
Melittobia audouinii : Newport, 1849c : 122.
Anthophorabia retusa : Newport, 1852a : 63.
Melittobia audouinii : Newport, 1852a : 65.
Anthophorabia fasciata Newport, 1852b : 81
SYN NOV.
Anthophorabia fasciata : Newport, 1853 : 165.
Melittobia acasta ; Smith, 1853 : 248.
Melittobia acasta : Giraud, 1869 : 151, 155.
Melittobia audouinii : Ashmead, 1892 ; 228.
Anthophorabia retusa : Ashmead, 1892 : 228.
Melittobia acasta : Dalla Torre, 1898 : 84.
Melittobia acasta : Ashmead, 1904 : 348, 380.
Melittobia acasta : Schmiedeknecht, 1909 ; 466.
Melittobia acasta : Morely, 1910 ; 57.
Melittobia acasta : Waterston, 1917 : 190.
Melittobia strandi Wolff and Krausse, 1922 : 16
SYN NOV.
Melittobia acasta : Peck, 1963 : 161.
Melittobia acasta : Peck, Boutek and Hoffer,
1964 : 98.
Melittobia acasta : Domenichini, 1966 ; 56.
Melittobia acasta : De Santis, 1957 : 109.
Melittobia acasta : De Santis, 1973 : 16.
Melittobia audouinii : Boubek, 1977 : 24.
Melittobia acasta ; Boufcek, 1977 : 24.
Melittobia audouinii ; Gordh, 1979 : 1005.
TYPE SPECIMENS:
I have not seen Walker’s type-specimen of M.
acasta which is in the British Museum (Natural
History), London nor have 1 seen any specimens
of Westwood (M audouinii) and Newport
(Anthophorabia retusa, A. fasciata). The
specimens of the last two authors, if they exist,
are probably also in the British Museum. My
recognition of this species is based upon material
identified by Dr Z. Boutek and the exhaustive re-
description by Waterston (1917). It is a distinctive
species and there can be little doubt of its identity.
Newport (1852b) described the species A, fasciata
provisionally, in the absence of any of the A.
retusa specimens for comparison, because new
specimens to hand showed characters he could
not remember in A. retusa. However, from the
figures in this paper and his 1853 paper, A.
fasciata is clearly a junior synonym of M. acasta.
The type of M. strandi Wolff and Krrausse
(1922) could not be located. From their
description and figures this species is clearly a
junior synonym of M. acasta.
DISTRIBUTION:
England, Europe, Japan (= M. japonica of
Maeta and Yamane (1974)) (= species 1 of van
den Assem and Maeta (1978)), South America,
Canada (= chalybii of Hobbs and Krunic (1971))
and New Zealand, (= species 2 of van den
Assem, Bosh and Prooy (1982)).
DESCRIPTION:
Female: Critical point dried specimens
1.4-1. 5 mm long; air dried specimens
1.3-1. 5mm long. Colour air dried specimens:
head, scape, pedicel, flagellum, mesosoma,
coxae, trochanters, proximal Vz femora,
metasoma dark brown, remainder of legs
yellow brown.
Head in frontal aspect (Fig. 39) of moderate
width, length to genal width about 1.2:1;
genal-clypeal margin not as broadly rounded
as M. clavicornis clypeal margin (Fig. 60)
bilobed, lobes broad; eyes relatively bare,
with a few short scattered setae. Facial
grooves remaining separate to scrobes,
maximum distance between arms not as wide
as POL, 2.6 times diameter of median
ocellus, converging gradually to meet scrobes
well below middle of eyes, lower arms
separated by a distance 0.5 times the width of
median ocellus. Scrobe to eye length 1:2.
Mandibles (Fig. 50); anterior tooth relatively
short, narrow, middle tooth more prominent
than 3, acute, narrow, third tooth broad not
as clearly defined. Maxillary palps (Fig. 71)
elongate, cylindrical, slightly widest at
middle, of moderate length, L:W 4:1.
DAHMS: REVISION O? MELITTOBIA
289
Antennae (Fig. 84); scape (Fig. 83) narrow,
L:W 3.4:1; MPS formula on flagellum
345:563; club segment 3 (Fig. 103) shortest
length to width 1:2; nipple elongate, standing
well above MPS, L:W 4.7:1; subterminal seta
about mid-way down nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.8. Posterior margin of mesoscutum mid
lobe 1.2 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.7:1; 1 pair of setae on each submedian lobe,
posterior setae situated forward of posterior
margin by a distance of 1/lOth length of
submedian lobe. Sculpture pattern on
mesoscutum and scutellum mid lobes (Fig.
141). Propodeum wider than long,
rectangular, posterior margin transverse,
emarginate, posterolateral angles 90°. Fore
wings (Fig. 116) relatively long, L:W 2.5:1
costal margin almost straight; marginal to
submarginal vein length 1.4:1; stigmal vein
(Fig. 128), marginal to stigmal vein length
4.8:1, submarginal to stigmal vein length
3.2:1; terminal seta on post marginal vein as
long as those on marginal vein.
Male: Critical point dried specimens 1.4-1. 7
mm long. Head, antennae, mesosoma, legs
and metasoma medium brown except
flagellum infuscated, mesoscutum and axillae
very pale; vertex, prothorax, scutellum,
coxae, trochanters, proximal 2/3 femora,
metasoma lightly infuscated. In air dried
specimens the colour becomes a fairly
uniform medium to dark brown.
Head in frontal aspect (Fig. 152) vertex
broad, almost straight especially in air dried
specimens, contracting strongly below eye
spots to clypeal margin, L:W 1:1.07; clypeal
margin bilobed, lobes broad. Mandibles (Fig.
162) elongate, anterior tooth long, narrow,
widely separated from second, second and
third teeth broad, 2 the largest. Maxillary
palps (Fig. 175) elongate, cylindrical, of
medium length, L:W 4:1. Antennae (Figs.
188-190); scape club-shaped, less so than M.
clavicornis, scape to head length 1.1:1, L:W
2:1; ventral surface distaily with a deep, cup-
shaped depression, glandular area
transversely elongate (Fig. 189); distal scape
margin strongly oblique, broadly excavated,
excavation overhung by long setae (Fig. 188).
Funicular segmental proportions (Fig. 190), 1
the largest, L:W 1:1.3; segments 2-4 small,
sub-equal, 2 the smallest, width of 2-4
slightly larger than length of 1; MPS formula
0001:342.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.3. Parapsidial sutures and axillae not as
well defined as in M. clavicornis. Scutellum
without submedian grooves, sublateral
grooves faintly defined; 2 pairs of setae
positioned as in female. Fore trochanters
without a dense tuft of short, stiff setae
ventrally; tarsal segments 3 + 4 fused. Mid
legs (Fig. 221) trochanters with a dense tuft of
long, fine setae ventrally; femoral fringe
strongly differentiated, proximal half of
femora with fringe of setae shorter than
width of femur, distal fringe with long setae,
slightly longer than width of femur; mid
femur L:W 3.8:1; mid tibia shape distinctive,
L:W 3.8:1; mid tarsi without fused segments.
Fore wings (Fig. 232) broad, L:W 2.6:1;
marginal to submarginal vein length 1.4:1;
stigmal vein well developed, broad; costal cell
relatively broad, L:W 6.7:1, costal margin
strongly arched.
MATERIAL EXAMINED:
BM(NH) 12 2 2 ‘L.S. Kensington, iii.1948, R.B.
Benson, B.M. 1948. 145’; 3 2 2 ‘England,
Hants. Basingstoke, Long Sutton, 4.vi.l948,
F.D. Goodliffe’, ‘Com. Inst. Ent. Coll. No.
11069’, ‘Pres, by Com. Inst. Ent. B.M. 1948
— 550’, ‘ex leaf cutter bee’; I S ‘Basingstoke,
Hants., 29.vi.1904, A.H. Hamm’,
"Melittobia acasta Wlk. Melittobia
acasta (Walk.) Det. Z. BouCek 1974’, ‘Bred
from Odynerus sinuatus'; 4 SS ‘Cambridge,
1915, Dr G.S. Graham Smith’, ‘J. Waterston
Det. Melittobia acasta Walk (?’, ‘Bred from
Calliphora'; 2 5$ ‘Haywards Heath, Sussex,
v.iii.l919, K.G.B. 1919 — 222’, ‘Bred from
Tachinid puparia ex A. pyramidea\
USNM 1 2, 4 / ‘ex Cnidocampa flavescens'
‘Melrose Hills Mass. 9-29’, ‘Gip. Moth Lab.
13043’; 3 ‘ex Tachinid puparium’, ‘Phila
Pa 9-20-24’, ‘Gip. Moth Lab 11741 J13’.
QM 2 2 2 1 S on a microscope slide ‘Jose C.
Paz. Prov. de Bs.As., S/ Trypoxylon sp. col.
Ibarra Grasso 10/III/1956 2 and
"Melittobia acasta Det. De Santis’.
NMC 20 2 $ 4 SS ‘Boh. Majdalena, Treboh,
V.1963 K. Denes’, ‘ex pupa Coroebus undatus
F. (Buprestidae)’; 22 2 2 1 i ‘Majdalena
Bohemia, v.63, lat. K. Denes’, ‘ex pupa
Coroebus undatus F. Buprestidae’; 1122 4
$ $ ‘Bohemia : Sobeslav, ex Vespa crabro,
290
MEMOIRS OF THE QUEENSLAND MUSEUM
X.1950, K. Pfleger ed^; 5 22 ‘Italia : Sasso,
Furbara, 1948, Sceliphron destill.*
In addition critical point dried material as
follows:
1) ‘Losser, Overijssel Prov., Netherlands,
18. vi, 1974, G.A. Bekke, ex pompilid/ 4 22 1
in each of the following institutions:
ANIC, BM(NH), DSIR, USNM, MNHP,
QM, THAES, UCR, USNM.
2) ‘1 km W, Taitapu New Zealand, 20 March,
1980 R.P. Macfarlane\ ‘ex nest Bombus
hortorum nest in box No. 031” 4 22 1 S in
each of the following institutions: ANIC,
BM(NH), DSIR, QM.
3) ‘Morioka Exp. Stn., Iwate Pref., Japan,
1974, Y. Maeta*, ‘ex Osmia imaii* 4 2 2 1 ^ in
each of the following institutions: ANIC,
BM(NH), DSIR, KU, QM, THAES, UCR,
USNM.
4) ‘Lethbridge, Alta, Canada, xi.l974, G.A.
Hobbs’, ‘ex Megachile relativa* 4 22 1 c? in
each of the following institutions: ANIC,
BM(NH), DSIR, QM, UCR, USNM.
Melittobia evansi SP NOV
(Figs 40, 51, 61, 72, 85, 86, 104, 117, 129, 142,
153, 163, 176, 191, 192, 193, 223, 233)
TYPE SPECIMENS:
The types were selected from a laboratory
culture maintained at the University of Georgia,
Athens, Georgia, U.S.A. which was established
from specimens collected at Athens, Georgia,
U.S.A., by D.A. Evans, out of nests of
Trypoxylon striatum. The holotype -S (marked H
on card-mount) is air dried from alcohol and card
mounted with 2 Si 2 2 2 paratypes (USNM).
Additional paratypes consist of 15 cards each
with 4 2 2 and 5 cards each with 1 (J 4 2 2 critical
point dried from alcohol and are deposited in the
following institutions USNM BM(NH), QM,
UCR, UG. The paratypes are rather pale as a
result of leaching in alcohol.
DISTRIBUTION:
Athens, Georgia, U.S.A. ; Goshen, N.Y.,
U.S.A. ( = species 3 of van den Assem, Bosch and
Prooy (1982)).
DESCRIPTION:
Female: Critical point dried specimens 1.1-1. 4
mm long. Colour from air dried material ex
alcohol. Head, antennal flagellum.
mesosoma, coxae, proximal 2/3 femora dark
brown; metasoma medium brown; scape,
pedicel and remainder of legs yellow-brown.
Head in frontal aspect (Fig. 40) relatively
narrow, length to width of gena about 1.3:1;
genae almost parallel, genal-clypeal margin
angled rather than broadly rounded; clypeal
margin (Fig. 61) bilobed, lobes broad; eyes
relatively bare, with a few short, scattered
setae. Facial grooves remaining separate to
scrObes, maximum distance between arms
about 2.2 times diameter of median ocellus;
grooves converge gradually to meet scrobes
below middle of eyes, minimum distance
between grooves 0.5 times the diameter of
median ocellus. Scrobe to eye length about
1:2.5. Mandibles (Fig. 51); anterior tooth
long, narrow; second and third well defined,
second long, narrow. Maxillary palps (Fig.
72) elongate, cylindrical, relatively long, L:W
5:1. Antennae (Figs 85, 86); scape narrow,
L:W 4.2:1; MPS formula on flagellum
355:553; club segment 3 (Fig. 104) length to
shortest length 1:1.4, nipple broad, L:W
2.5:1, barely projecting above MPS;
subterminal seta just below middle of nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.4. Posterior margin of mesoscutum mid
lobe 1.2 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.4:1; 1 pair of setae on each submedian lobe
of scutellum, posterior setae on posterior
margin of lobe. Sculpture pattern on
mesoscutum and scutellum mid lobes (Fig.
142), pattern very open. Propodeum wider
than long, posterior margin an open V-shape,
posterolateral angles obtuse. Fore wings (Fig.
117) L:W 2.4:1; costal margin bent at
junction with parastigmal vein; marginal to
submarginal vein lengths 1.5:1; 4-5 setae on
submarginal vein; stigmal vein (Fig. 129)
marginal to stigmal vein length 4.1:1;
submarginal to stigmal vein length 2.8:1;
terminal seta on postmarginal vein much
longer than those on marginal vein.
Male: Air dried specimens ex alcohol 1.4 mm
long. Head, antennae, mesosoma, legs pale
brown, head darkest, mesoscutum and axillae
palest; metasoma dark brown.
Head in frontal aspect (Fig. 153), L:W 1:1,
vertex rounded, narrower than width below
eye spots, genae broadly rounded, not
contracted, meeting clypeus in an obtuse
angle; clypeal margin bilobed, lobes broad.
Mandibles (Fig. 163) long, narrow; anterior
DAHMS: REVISION OF MELITTOBIA
291
tooth very long; second and third well
defined, second the longest. Maxillary palps
(Fig. 176) elongate, cylindrical, L:W 4.8:1.
Antennae (Figs. 191-193) scape club-shaped,
less pronounced than M. acasta, scape to
head length 1:1.6, L:W 1.7:1; ventrally (Fig.
192) with a distal, deep, cup-shaped
depression; glandular area transverse
expanding on side opposite pedicel
attachment; distal scape broadly excavated,
very slightly oblique; funicular segmental
proportions (Fig. 193) 1 large, L;W 1:1.6; 2-4
sub-equal, width to length of first 1:1.2; MPS
formula 0022:232.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.8. Parapsidal sutures and axillae not as
clearly defined as in M. clavicornis.
Submedian grooves of scutellum absent,
sublateral grooves faintly marked; 2 pair of
setae positioned as in female. Fore
trochanters without a ventral tuft of short,
stiff, setae; fore tarsi with segments 3 h- 4
fused. Mid legs (Fig. 223); mid trochanters
with a dense tuft of long, fine setae; femoral
fringe evenly distributed along femur, setae
of even length, about as wide as femur, mid
femur L:W 3.9:1; mid tibia L:W 3.2:1; mid
tarsi without fused segments. Fore wings
(Fig. 233) broad, L:W 2.6:1, costal and
posterior margin almost parallel; marginal to
submarginal vein length 1.4:1; stigmal vein
well developed, broad; costal cell narrow,
L:W 8:1, costal margin slightly arched.
MATERIAL EXAMINED:
Holotype $; and paratype $9 from
Georgia deposited as in TYPE SPECIMENS
section.
USNM 5 99 ‘Goshen, N.Y., 3.22.40, vial 1,
R.G. Schmieder*, ‘Ex Trypoxylon politum ’;
3 99 data as above vial 2; 1 99 ‘Goshen,
N.Y. 7-27-36, vial 3, R.G. Schmieder’, ‘Ex
Trypoxylon politum ’ . The last specimen is a
second form with crumpled wings.
This species is named evansi after Dr. D.A.
Evans, Kalamazoo College, U.S.A. who
brought this species to my attention and has
been of help in many other ways.
Melittobia scapata SP NOV
(Figs 41, 52, 62, 73, 87, 105, 118, 130, 143, 154,
164, 177, 194, 195, 196, 222, 234)
TYPE MATERIAL:
Holotype $ plus 4 paratype 2 2 on a single card
with the original alcohol labels ‘Tompkins Co.,
N.Y. late Apr., 1974, Suellen Vernoff, ‘ex nest
Trypoxylon \ ‘CU Lot No. 1040, site C, Nest 88’;
4 29 1 paratypes on a single card with holotype
data (USNM); 22 2 9 2 paratypes with
holotype data (CU, NY); 2 c? (7 , 3 9 9 paratypes on
slides (QM) ‘New York, U.S.A., 8.May.l971, G.
Eichardt, reared nest of mason wasp. Euparal,
E.C. Dahms 1980’.
DISTRIBUTION:
Tompkins County, New York, U.S.A.
DESCRIPTION:
Female: Critical point dried specimens 1.6-1. 7
mm long. Head, flagellum, mesosoma,
coxae, femora dark brown; scape, pedicel,
metasoma, remainder of legs medium brown.
These specimens have been in alcohol since
1974 and the antennae have leached. The
scape and pedicel were probably fairly dark
when fresh but not as dark as the flagellum.
Head in frontal aspect (Fig. 41) relatively
broad, length to genal width 1.1:1; genae
broadly rounded, genal-clypeal margin more
or less sharply angled; clypeal margin (Fig.
62) bilobed, lobes broad, relatively long; eyes
relatively bare, with a few short setae. Facial
grooves remaining separate to scrobes,
maximum distance between arms 2.1 times
diameter of median ocellus; grooves
converging sharply to just above middle of
eyes then running close and parallel to meet
scrobes well below middle of eyes; minimum
distance between arms 0.3 times diameter of
median ocellus. Scrobe to eye length 1:2.4.
Mandibles (Fig. 52) broad, anterior tooth
short, narrow, very widely separated from
second; second and third broad, equal.
Maxillary palps (Fig. 73) elongate,
cylindrical, relatively long L:W 4.6:1,
Antennae (Fig. 87); scape narrow, L:W 3.4:1;
MPS on flagellum 356:673; club segment 3
(Fig. 105) shortest length to width 1:1.8;
nipple relatively broad, L:W 2.6:1;
subterminal seta just below middle of nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.7. Posterior margin of mesoscutum mid
lobe 1 .6 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
2:1; 1 pair of setae on each submedian lobe,
posterior setae on posterior margin of lobe.
Sculpture pattern on mesocutum and
292
MEMOIRS OF THE QUEENSLAND MUSEUM
scutellum mid lobes (Fig. 143). Propodeum
wider than long, posterior margin an open V-
shape, posterolateral angles obtuse. Fore
wings (Fig. 118) L;W 2.2:1; costal margin
bent at junction with parastigmal vein; 5-6
long setae on submarginal vein; marginal to
submarginal vein length 1.4:1; stigmal vein
(Fig. 130) marginal to stigmal vein length
4.7:1; submarginal to stigmal vein length
3.3:1; terminal seta on postmarginal vein
slightly longer than those on marginal vein.
Male: Critical point dried material, 1.4 mm
long. Head, body and legs a fairly uniform
pale brown; mesosoma lightly infuscated.
Head in frontal aspect (Fig. 154) rounded in
shape, genae slightly flattened, L:W 1:1.3;
clypeal margin bilobed, lobes broad.
Mandibles (Fig. 164); anterior tooth long,
narrow, widely separated from second;
second and third well defined, relatively
narrow. Maxillary palps (Fig. 177) elongate,
widest basally, L:W 4.5:1. Antennae (Figs
194-196) more or less evenly expanded
distally, length to head length 1:1.6, L:W
1.9:1; ventral surface (Fig. 195) with a distal,
deep, cup-shaped depression; glandular area
transverse, narrow, relatively small; distal
end of scape broadly excavated, only slightly
oblique; funicular segmental proportions
(Fig. 196) segment 1 the largest, L:W 1:1.6;
segments 2-4 smaller, sub equal, not much
narrower than segment 1; MPS formula of
flagellum 0011:122.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.4. Parapsidial sutures and axillae not as
well defined as in M. clavicornis. Submedian
grooves of scutellum absent, sublateral
grooves poorly defined; 2 pair of large setae
positioned as in female. Fore trochanters
without a ventral tuft of short, stiff setae;
tarsal segments 3 + 4 fused. Mid legs (Fig.
222); trochanters with a dense tuft of long,
fine setae; femoral fringe of uneven length,
proximal half of fringe short, setae a little
shorter than width of femur, distal fringe
with longer setae about 1.5 times width of
femur; L:W femur 4:1; tibia L:W 3.8:1; mid
tarsi without fused segments. Fore wings
(Fig. 233) broad, L:W 2.4:1; marginal to
submarginal vein length 1:1; stigmal vein well
developed; costal cell relatively broad, L:W
7.5:1, costal margin arched.
MATERIAL EXAMINED:
Holotype and paratypes as listed in TYPE
SPECIMENS section.
This species is named scapata to draw attention
to the relatively short, narrow scapes in the male.
Melittobia digitata SP NOV
(Figs 42, 53, 63, 74, 88, 89, 106, 119, 131, 144,
155, 165, 178, 197, 198, 199, 224, 235)
TYPE MATERIAL:
Holotype $ (indicated by arrow) mounted on a
card with 4 ? W paratypes ‘5 mis W. Tallahasse,
Florida, U.S.A., J. Trexler, 26. xi. 1980, ex
Trypoxylon politum ’ (USNM). 15 2 ^
paratypes ‘Leon Co. Fla., U.S.A., from culture
begun 11.26.1980’ in the following institutions:
BM(NH), QM, UCR, USNM.
DISTRIBUTION:
U.S.A. — Florida, Connecticut, Michigan,
Te.xas, Virginia, Mississippi. (= chalybii of
Buckell (1928), (= species 4 of van den Assem
Bosch and Prooy (1982)).
DESCRIPTION:
Female: Critical point dried material. 1.6 mm
long. Specimens unleached. Head, antennal
flagellum, mesosoma, coxae dark brown;
trochanters, proximal 2/3 femora, metasoma
lighter brown; scape, pedicel, remainder of
legs yellow-brown.
Head in frontal aspect (Fig. 42) relatively
narrow, length to genal width 1.4:1; genae
long, almost parallel; genal-clypeal margin
angled; clypeal margin (Fig. 63) bilobed,
lobes broad, each with a small, lateral, lobe-
like undulation; eyes relatively bare, with a
few, short, scattered setae. Facial grooves
separate to scrobes; maximum distance
between arms 1.8 times diameter of median
ocellus; contracting evenly to meet scrobes
below middle of eyes; minimum distance
between arms 0.5 times diameter of median
ocellus. Scrobe length to eye length 1:1.8,
Mandibles (Fig. 53); anterior tooth relatively
short, narrow; second and third well defined,
second the longest, both relatively acute.
Maxillary palps (Fig. 74) elongate,
cylindrical, of medium length, L:W 3.8:1.
Antennae (Figs 88, 89). Scape narrow, L:W
3.9:1; MPS formula on flagellum 567:653;
club segment 3 (Fig. 106) shortest length to
width 1:2; nipple relatively broad, L:W 3:1;
subterminal seta situated well below half way
down nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.2. Posterior margin of mesoscutum mid
DAHMS: REVISION OF MELITTOBIA
293
lobe 1.3 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.9:1; 1 pair of setae on each submedian lobe,
posterior setae situated on posterior margin
of lobe. Sculpture pattern on mesoscutum
and scutellum mid lobes (Fig. 144).
Propodeum wider than long, posterior
margin an open V-shape, posterolateral
angles obtuse. Fore wings (Fig. 119) L:W
2.5:1, costal margin bent at junction with
parastigmal vein; 4 setae on submarginal
vein; marginal to submarginal vein length
1,3:1; stigmal vein (Fig. 131) marginal to
stigmal vein length 4.3:1; submarginal to
stigmal vein length 3.3:1 ; terminal seta on
postmarginal vein longer than those on
marginal vein.
Male: Critical point dried specimens 1.5 mm
long. Specimens unleached. Head,
mesoscutum, axillae, scutellum yellow, paler
than rest of body; antennae, remainder of
mesosoma, legs pale brown; metasomal
segments each with a posterior broad,
transverse, infuscated band. In air dried
specimens the whole insect becomes a
medium brown, metasoma black.
Head in frontal aspect (Fig. 155) vertex
broadly rounded, almost straight in air dried
specimens; genal margins relatively straight,
contracting slightly to an angular genal-
clypeal junction; clypeal margin bilobed,
lobes broad, well defined. Mandibles (Fig.
165); anterior tooth of median length, broad,
well separated from second; second and third
teeth well defined, broad, 3 the broadest.
Maxillary palps (Figs 178) elongate,
cylindrical, of medium length, L:W 3:1.
Antennae (Figs 197-199). Scape club-shaped,
length to head length 1:1.5, L:W 1.9:1;
ventral surface (Fig. 198) with a distal, cup-
shaped depression; glandular area elongate,
transverse, narrow; distal club margin very
deeply excavated producing a thumb-like
projection on the side opposite pedicel
attachment. Funicular segmental proportions
(Fig. 199) I largest, L:W 1:1.7; segments 2-4
sub-equal, 2 the smallest; width of 2-4
approximately equal to length of segment 1;
MPS formula on flagellum 021:142.
Mesosoma in dorsal aspect. Prothorax L;W
1:1.6. Parapsidial sutures and axillae not as
well defined as in M. clavicornis. Submedian
grooves of scutellum absent, sublateral
grooves poorly defined. Fore trochanters
without a ventral tuft of short, stiff setae;
fore tarsi with segments 3+4 fused. Mid legs
(Fig. 224); trochanters with a dense tuft of
long, fine setae; femoral fringe of uneven
length, proximal 1/3 much shorter than width
of femur, medial 1/3 about as wide as femur
and distal 1/3 about twice width of femur;
L:W femur 4.1:1; tibia L:W 3.4:1; mid tarsi
without fused segments; setae on tarsal
segments relatively long. Fore wings (Fig.
235) broad, L:W 2.7:1; marginal to
submarginal vein length 1.2:1; costal cell
narrow, L:W 7.6:1, costal margin slightly
arched; stigmal vein well developed.
MATERIAL EXAMINED:
Holotype and paratypes as listed in TYPE
SPECIMENS section.
USNM 3 3 ‘Connecticut, U.S.A.,
6.i.l978, T.M. O’delF, ‘ex Lab. culture
Tachinid Blepharipa sp., U.S. Dep. Agr. &
Forest Service’; 3 2? 1 ^ ‘Gainesville, Fa.,
2/17/26’, ‘ex Tromotobia rufopectus on
Argiope sp.\ ‘W.A. Murrill coll.’; 2 ?£ I t?
‘Norfolk, Va., 5.20.31, L.D. Anderson’,
‘Reared from larvae and pupae, Sphecoid
wasp’; 1 2 4 ‘College Station, Brazos
Co., Texas, viii.8.1976, Ha! Reed’; 40 99 9
S S ‘Hillsdale, Michigan, U.S.A., Sept. 1977,
B.K. Nubel’, ‘ex Megachile sp. nesting in old
Trypoxylon nests’; 1 $ ‘State College, Miss.,
Ex sphecidae, R.E. Hutchings’.
QM 3 99 2 on microscope slides with the
Hillsdale data above.
This species has been named digitata to draw
attention to the deeply excised distal margin of
the scape which results in a thumb-like projection
on the side opposite the pedicel attachment.
Melittobia femorata SP NOV
(Figs 43, 54, 64, 75, 90, 91, 107, 120, 132, 145,
156, 166, 179, 200, 201, 202. 225, 236)
TYPE MATERIAL:
Holotype S 1 paratype £ mounted together
‘Jackson and Franklin Counties, North Carolina,
U.S. A., C.E. Hinton, 21. vi — ll.viii.l979, ex
Trypoxylon politum ’. (USNM); 44 paratype 9 5
bearing the same data as holotype in the following
institutions: BM(NH), QM, UCR, USNM; 4 99 2
paratype ^ ^ on slides bearing the same data as
holotype (QM).
294
MEMOIRS OF THE QUEENSLAND MUSEUM
DISTRIBUTION:
North Carolina, U.S.A.
DESCRIPTION:
Female: Critical point dried specimens 1. 5-1. 6
mm long. Head, antennal flagellum,
mesosoma, coxae, trochanters dark brown;
proximal 2/3 femora and metasoma paler
brown; remainder of legs, dorsal scape and
pedicel rufous brown; ventral scape and
pedicel yellow-brown. Sculpture pattern on
head relatively fine giving the surface a dull
shagreened appearance as in M. megachilis.
Head in frontal aspect (Fig, 43) relatively
narrow, length to genal width 1.3:1; genal-
clypeal margin angled; clypeal margin (Fig.
64) bilobed, lobes broad; eyes relatively bare,
with only a few short scattered setae. Facial
grooves remaining separate to scrobes,
maximum width between arms 1.2 times
diameter of median ocellus; contracting
gradually to meet scrobes below middle of
eyes; minimum distance between arms 0.2
times diameter of median ocellus. Scrobe to
eye length 1:2.7. Mandibles (Fig. 54); anterior
tooth short, very broad, second and third well
defined, broad, equal. Maxillary palps (Fig.
75) elongate, cylindrical, relatively long 4.2: 1 .
Antennae (Figs 90, 91); scape narrow, L:W
3.7:1; MPS formula on flagellum 577:763;
club segment 3 (Fig. 107) shortest length to
width 1:1.8; nipple elongate, L:W 4:1;
subterminal seta situated just below middle of
nipple,
Mesosoma in dorsal aspect. Prothorax L:W
1:1,5. Posterior margin of mesoscutum mid
lobe 1.3 limes wider than anterior margin of
scutellum. Scutellum mid lobe L:W 1.9:1; 1
pair of setae on each submedian lobe,
posterior seta situated on posterior margin of
lobe. Mesoscutum and scutellum mid lobe
sculpture pattern (Fig. 145). Propodeum with
posterior margin truncate emarginate,
posterolateral angles 90°. Fore wing (Fig.
120) L:W 2.3:1; costal margin bent at
junction with parastigmal vein; 3-5 setae on
submarginal vein; marginal to submarginal
vein length 1.3:1; stigmal vein (Fig. 132) in
some specimens quite distinctive, in others it
is not unlike that of M. scapata (Fig. 130);
marginal to stigmal vein length 5,3:1;
submarginal to stigmal vein length 4.0:1 ;
terminal seta on end of postmarginal vein not
longer than those on marginal vein.
Male: Critical point dried specimen 1.5 mm
long. Head, body and legs rufous brown
except for infuscations on distal scape,
pedicel, vertex, mesosoma and metasoma.
Head in frontal aspect (Fig. 156) vertex
broadly rounded, lateral margins flat,
parallel, L:W 1:1; clypeal margin bilobed.
Mandibles (Fig. 166); anterior tooth long,
broad, widely separated from others; second
and third teeth well defined, broad, equal.
Maxillary palps (Fig. 179) elongate,
cylindrical, L:W 4.8:1. Antennae (Figs
200-202); scape gradually expanded from
proximal end, distal 1/3 showing a slightly
greater expansion; length to head length
1:1.5, L:W 1.8:1; ventral surface with distal,
deep, cup-shaped depression; glandular area
transverse, broad, expanded at side opposite
pedicel attachment; distal end of scape
transverse, with a deep, relatively narrow
excavation; 5 funicular segments, the first is
one of the ring joints expanded and segment 2
is equivalent to segment 1 of other species;
funicular segmental proportions 1 small, 2
largest, L:W 1:1.4, segments 3-5 sub-equal
with 3 the smallest; width of segments 4-5 to
length of segment 2 is 1:1,5; MPS formula on
Oagellum 00142:432.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.4. Parapsidial sutures and axillae not as
well defined as M. clavicornis. Scutellum
without submedian grooves; sublateral
grooves clearly marked; 2 pair of long setae
situated as in female. Fore trochanters
without a ventral tuft of short, stiff setae;
fore tarsi with segments 3 + 4 fused. Mid legs
(Fig. 225); trochanters with a ventral tuft of
long, fine setae; femoral fringe uneven,
proximal fringe about equal to width of
femur and extends for approximately half the
femur; distal fringe is dense with extremely
long setae about 1.75 times width of femur;
L:W femur 3.9:1; tibia L;W 3:1; mid tarsi
without fused segments. Fore wings (Fig. 236)
broad, L:W 2.6:1; marginal to submarginal
vein length 1.3:1; costal cell narrow, L:W
7.6:1, costal margin above, straight; stigmal
vein well developed.
MATERIAL EXAMINED:
The specimens listed in TYPE SPECIMENS
section.
This species is named femorata to draw
attention to the extremely long mid femoral setae
of the male.
DAHMS: REVISION OF MELITTOBIA
295
Melittobia chalybii Ashmead.
(Figs 44, 55, 65, 76, 92, 108, 121, 133, 146, 157,
167, 180, 203, 204 205, 226, 237)
Melittobia chalybii Ashmead, 1892 : 231.
Melittobia chalybii : Dalla Torre, 1898 : 85.
Melittobia chalybii : Schmiedeknecht, 1909 : 466.
Melittobia chalybii : Peck, 1951 : 452.
Melittobia chalybii : Burks, 1958 ; 67.
Melittobia chalybii ; Peck, 1963 : 162.
Melittobia chalybii : Gordh, 1979 : 1005.
TYPE MATERIAL:
Ashmead (1892) did not select a primary type
and merely said, *... from many specimens of
both sexes, reared September 14 from cells of
Chalybion caerubum Linn, collected in Virginia’,
I have selected a lectotype and paralectolypes
from his point-mounted, syntypical series in the
USNM as follows:
1 cJ minus right antennae flagellum and left
wings ‘Bladensb. Va. Sept. 14.91’, ‘Paratype
No. 2135 U.S.N.M.’. LECTOTYPE.
1 c? minus left antenna, right antennal
flagellum and right wings ‘Va.’,
‘Allotype No. 2135 U.S.N.M.’, ^Melittobia
chalybii Kshm: . PARALECTOTYPE.
1 $ minus both antennal flagella, all wings
and part of left mesosoma ‘Va.’, ‘c?’,
‘Paratype No. 2135 U.S.N.M.’. PARA-
LECTOTYPE.
1 2 intact ‘Bladensb. Va. Sept. 14.91’,
‘Paratype No. 2135 U.S.N.M.’, ^Melittobia
chalybii Ashm: . PARALECTOTYPE.
1 2 intact ‘Va.’, ‘ 2 ’, ‘Paratype No. 2135’.
PARALECTOTYPE.
1 2 minus right antennal flagellum and all
wings ‘Va.’, ‘ 2 ’, ‘Paratype No. 2135
U.S.N.M.’. PARALECTOTYPE.
In addition to the above there is 1 point
bearing only some legs from a $ syntype
labelled ‘Va.’, ‘Paratype No. 2135
U.S.N.M.’. The USNM labels on these
specimens incorrectly indicate that the series
contains an allotype and several paratypes.
Presumably the specimen with a USNM
holotype label no longer exists.
DISTRIBUTION:
U.S.A. — Virginia and New Jersey.
DESCRIPTION:
Female: Air dried specimens 1.3- 1.5 mm long.
Head, antennal flagellum, mesosoma, coxae,
proximal 2/3 femora dark brown; mesosoma
paler; scape, pedicel, remainder of legs yellow
brown.
Head in frontal aspect (Fig. 44) extremely
setose, relatively broad, length to genal width
1.2:1; genal -clypeal margin broadly rounded;
clypeal margin (Fig. 65) bilobed, lobes
relatively small, each with a small, lateral,
lobe-like undulation; eyes densely clothed in
longish setae. Facial grooves remaining
separate to scrobes; maximum distance
between arms 1.2 times diameter of median
ocellus; arms converge gradually to meet
scrobes just below middle of eye; minimum
distance between arms 0.2 times diameter of
median ocellus. Scrobe to eye length 1:1.9.
Mandibles (Fig. 55); anterior tooth relatively
short, narrow; second and third well defined,
2 the narrowest. Maxillary palps (Fig. 76)
elongate, cylindrical, of medium width, L;W
3.4:1. Antennae (Fig. 92); scape narrow, L:W
3.3:1; MPS formula on flagellum 346:663,
club segment 3 (Fig. 108) shortest length to
width 1:2.6; nipple elongate, L:W 3:1;
subterminal seta just below middle of nipple.
Mesosoma in dorsal aspect. Prothorax L;W
1:1.4. Posterior margin of mesoscutum mid
lobe 1.5 limes wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.9:1; submedian lobes each with 3-5 setae,
sometimes varying between left and right on
the same specimen. Mesoscutum and
scutellum mid lobes sculpture pattern (Fig.
146). Propodeum posterior margin an open
V-shape, posterolateral angles obtuse. Fore
wing (Fig. 121) L:W 2.4:1; costal margin not
as sharply bent at junction with parastigmal
vein as other species (except M. acasta );
marginal to submarginal vein length 1.5:1;
5-6 long setae on submarginal vein; stigmal
vein (Fig. 133); marginal to stigmal vein
length 4.6:1; submarginal to stigmal vein
length 3.2:1; terminal seta on postmarginal
vein as long as setae on marginal vein.
Male: Air dried specimens 1.1 mm long. Head,
body, legs uniform golden brown. There are
indications that the head may be paler than
the rest and that the mesosoma is lightly
infuscated, but confirmation requires fresh
material.
Head in frontal aspect (Fig. 157) rather
globose in slide-mounted specimens, L:W
1:1. In air dried specimens head folds
transversely just below ocelli which gives the
head a shape more like M acasta (Fig. 152).
296
MEMOIRS OF THE QUEENSLAND MUSEUM
Genae contracted well below eye spots;
clypeal margin bilobed. Mandibles (Fig. 167)
elongate, anterior tooth long, narrow, not
widely separated from second; second and
third unequal, acute, 2 the longer. Maxillary
palps (Fig. 180) elongate cylindrical, of
medium length, L:W 3.3:1. Antennae (Figs.
203-205). Scape relatively broad, evenly
expanded from proximal end, length to head
length 1:1.5; L:W 1.6:1; ventral surface (Fig.
204) with a distal, deep, cup-shaped
depression; glandular area geniculate; distal
end of scape with a relatively shallow
excavation, not as deep as in M. digitata;
funicular segmental proportions 1 the largest,
L:W 1:1.3; segments 2-4 sub-equal, 2 the
smallest, width of 2-4 to length of 1 1:1.7;
MPS formula on flagellum 021:242.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.3. Parapsidial sutures and axillae not as
well defined as in M. clavicornis. Scutellum
without submedian grooves; sublateral
grooves weakly defined; 2-3 pairs of large
setae positioned as in female. Fore
trochanters with a tuft of short, stiff setae,
less dense, but longer than M. australica; fore
tarsi with segments 3 + 4 fused. Mid legs
(Fig. 226); trochanters with a ventral tuft of
long, fine setae; femoral fringe uneven,
proximal 1/3 shorter than width of femur,
distal 2/3 approximately as long as width of
femur; L:W femur 3.7:1; tibia L:W 3.4:1;
mid tarsi without fused segments. Fore wing
(Fig. 237) broad though relatively elongate,
posterior margin relatively straight and
parallel to costal margin, L:W 2.9:1;
marginal to submarginal vein length 1.6:1;
costal cell narrow, L:W 11.7:1, costal margin
above not arched; stigmal vein well
developed.
MATERIAL EXAMINED:
USNM 3 9 3 syntypes as in TYPE
SPECIMENS section. 2 2^ 3 SS card-
mounted ‘Marlton, N.J., 3.17.40, vial 6,
R.G. Schmieder’; 2 SS same data but from
vial 5.
QM 3992c?^ona microscope slide, same data
as above, vial 5.
Over the years the name M. chalybii has been
applied to more than one species of Melittobia
from North America. Buckell (1928) applied it to
M. digitata. Although I have not examined his
specimens his figures are quite clear and the scape
of M. digitata males is very distinctive. However,
the most common species confused with M.
chalybii is M. australica from which it is easily
distinguished using the following characters:
Female:
Scape L:W
Scape & pedicel
Clypeal margin
Facial grooves
Setae on submarginal
vein
Setae on each sub-
median lobe of
scutellum
Male:
Scape
Segment 1 of flagellum
Head shape
Clypeal margin
Mid-leg trochanters
Fore-wings L:W
Stigmal vein
chalybii
3.1:1
yellow brown
bilobed
converge separately to scrobes
5-6
3-4
ventral cup
the largest
Fig. 174
bilobed
with a dense tuft of long fine
setae
2.9:1
well developed
australica
2.5:1
infuscated
truncate
converge to meet then pass to
scrobes as a single line
4
2
ventral groove
the smallest
Fig. 175
without lobes
without this tuft
3.7:1
reduced to a swelling on
marginal vein
DAHMS: REVISION OF MELITTOBIA
297
Although M. chalybii is the commonest name
used in the literature for North American
Melittobia it was the least common species
encountered amongst collections borrowed for
this revision or forwarded for identification. In
fact the only specimens available were the types
and some dried specimens from USNM.
Melittobia megachilis (Packard)
(Figs 45, 66, 77, 93, 109, 122, 134, 147)
Anthophorabia megachilis Packard, 1864 : 134.
Anthophorabia megachilis : Packard, 1868 : 204.
Anthophorabia megachilis : Packard, 1869 : 206.
Pteromalus gerardi 1875 : 131.
Anthophorabia megachilis : Howard, 1885 : 46.
Melittobia megachilis : Cresson, 1887 : 244.
Melittobia megachilis : Ashmead, 1892 : 229.
Melittobia megachilis : Ashmead, 1894 : 26.
Melittobia megachilis : Dalle Torre, 1898 ; 85.
Chrysocharis aeneus Brues, (1909) : 161.
Melittobia megachilis : Schmiedieknecht, 1909 :
466.
Miotropis megachilis : Viereck, 1916 : 465.
Chrysocharis aeneus : Girault, 1925 : 3.
Melittobia megachilis : Girault, 1925 ; 3.
Melittobia megachilis : Peck, 1951 : 452,
Melittobia gerardi : Burks, 1958 : 67.
Melittobia megachilis : Burks, 1958 : 67.
Melittobia megachilis : Peck, 1963 : 162.
Melittobia megachilis : Gordh, 1979 : 1005.
TYPE SPECIMENS:
Packard’s species is represented by a syntypical
series of 5 females and a tube of dried larvae.
These specimens reside in the collections of MCZ,
Harvard. Two of the five females 1 recovered
from amongst the dried larvae and mounted on
one card with a paralectotype label. The three
remaining females were mounted separately on
points as follows:
1) A reasonably complete female minus right
flagellum and labelled ‘‘Anthophorabia
megachilis Pack. F.W.P.’, Type 529’. I have
removed one fore wing from this specimen
and mounted it on a microscope slide.
2) A damaged female of which only wings and
mesosoma remain mounted upside down in
glue and labelled ^megachilis ’, ‘Type 529”.
3) A female without wings labelled ‘megachilis\
‘Type 529’. 1 have cleared and mounted this
specimen on the slide with the wing of 1 .
From these three I have selected (1) as the
lectotype and (2-3) as paralectotypes. All
specimens have been labelled accordingly.
I have not examined the types of Pteromalus
gerardi Hickok and Chrysocharis aeneus Brues to
confirm these synonymies.
DISTRIBUTION:
The type-locality is Brigport, Vermont, U.S.A.
DESCRIPTION:
Female: Air dried specimens 1.3 mm long; 2 $ ?
ex larvae 1.5 mm long. Head, antennal
flagellum, mesosoma, metasoma dark
brown; coxae, femora lighter brown; scape,
pedicel, remainder of legs yellow-brown.
Sculpture pattern on head relatively fine
giving the surface a dull shagreened
appearance as in M. femorata.
Head in frontal aspect (Fig. 45) relatively
narrow, length to genal width 1.4:1; genal
margin relatively straight, more or less
parallel; clypeal margin (Fig, 66) bilobed;
eyes relatively bare, with a few, short,
scattered setae. Facial grooves remaining
separate to scrobes; greatest width between
arms 1.8 times diameter of median ocellus;
arms converging gradually to meet scrobes
well below middle of eyes; minimum distance
between arms 0.2 diameter of median ocellus.
Scrobe to eye length 1:2.2 but may be larger
because the eyes are folded transversely.
Mandibles not visible. Maxillary palps (Fig.
77) elongate, cylindrical, long, L:W 5:1.
Antennae (Fig. 93); scape narrow, L:W 3.7:1;
MPS formula on flagellum 565:663; club
segment 3 (Fig. 109) shortest length to width
1:1.4; nipple relatively short, L:W 2.5:1;
subterminal seta almost basal.
Mesosoma in dorsal aspect. Prothorax L:W
1:1,4. Posterior margin of mesoscutum mid
lobe 1.5 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.9:1; 1 pair of setae on each submedian lobe,
posterior seta on posterior margin of lobe,
Propodeum posterior margin an open V-
shape, posterolateral angles obtuse.
Mesoscutum and scutellum mid lobe
sculpture pattern (Fig. 147). Fore wing (Fig.
122) L:W 2.5:1; marginal to submarginal vein
1.3:1; 4 setae on submarginal vein; marginal
to stigmal vein length 5.2:1 ; submarginal to
stigmal vein 4.0:1; terminal seta on
postmarginal vein not longer than those on
marginal vein.
Male: Unknown.
298
MEMOIRS OF THE QUEENSLAND MUSEUM
MATERIAL EXAMINED;
Only the types of this species were available
as in the section TYPE SPECIMENS.
Meiitfobia australica Girault
(Figs 46, 56, 68, 78, 96, 97, 112, 124, 135, 148,
158, 169, 181, 206, 207, 208 227, 238)
Melittobia australica : Girault, 1912 : 203.
Melittobia australica : Girault, 1913 : 205, 250.
Melittobia australica : Girault, 1914 ; 8.
Melittobia australica : Girault, 1915 : 216, 259.
Melittobia australica : Dahms, 1973 : 411.
TYPE SPECIMENS:
The syntypical series of Girault consists of 7 £ 2
2 c? c? on a microscope slide ''Melittobia australica
2 s 2 <?’, TYPE Hy/997, A. A. Girault’ (QM).
There are in DPIQ 5 slides containing numerous
specimens of both sexes and all labelled, ‘Dep.
Ag. & Stk., Qld. CHALCIDIDAE Melittobia
australica Ex Pison spinolae (Hym.) Tambourine
11/12/11 H.T. No. Hy.58’, ‘Mt. Tambourine S.
Queensland Dep. Ag. & S. 11.12.11’. These are
all part of the original series bred by Tryon, but
have no Girault labels. One of these slides
contains 3 females and 1 male therefore fitting
Girault’s published information for his ‘Cotypes’
and it would be safe to assume that he saw the
remainder of the slides. I am therefore labelling
the 5 slides as containing paralectotypes. The QM
slide has been relabelled by someone other than
Girault and the error in the number of females (2
as opposed to 7) is no doubt one of
transliteration. From this series 1 have selected the
intact male as the lectotype and the remaining
specimens (7 9 2 1 as paralectotypes. No
locality data occurs on this slide, but the
published data read, ‘Host, Pison spinolae
(Hym.) Mt. Tambourine, S. Queensland, Dept.
Ag. & S., 11; 12; IT.
DISTRIBUTION:
South Africa, Australia, Japan {= species 2 of
van den Assem and Maeta (1978)), North
America (= M. chalybii of Hermann(1971),
Evans and Matthews (1976)), Jamaica (= M.
chalybii of Freemann and Parnell (1973),
Freemann (1977)) (= hawaiiensis complex of
Freeman and Ittyeipe (1976), Jayasingh and
Freeman (1980)), (= species 8 of van den Assem,
Bosch and Prooy (1982)).
DESCRIPTION;
Female: Critical point dried specimens 1.1-1. 2
mm long. Head, antennal flagellum,
mesosoma, coxae, proximal 2/3 femora dark
brown; scape and pedicel barely paler;
metasoma paler; remainder of legs pale
brown.
Head in frontal aspect (Fig. 46) length to
genal width 1.4:1; genal-clypeal margin
broadly rounded; clypeal margin (Fig. 68)
truncate emarginate; eyes densely covered
with long setae. Facial grooves converging to
meet just above middle of eyes then passing
as a single line to scrobes; maximum width
between arms 2.9 times diameter of median
ocellus (approximately equals POL). Scrobe
to eye length 1:2.8. Mandibles (Fig. 56);
anterior tooth long, narrow relatively close to
second; second and third tooth well defined,
2 longer and narrower. Maxillary palps (Fig.
78) cylindrical, short, L:W 2.8:1. Antennae
(Figs 96, 97); scape broad, L:W 2.5:1; MPS
formula on flagellum 445:573; club segment 3
(Fig. 112) shortest length to width 1:1.2;
nipple elongate, L:W 4:1; subterminal seta in
distal half of nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.5. Posterior margin of mesoscutum mid
lobe 1.3 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.9:1. Mesoscutum and scutellum mid lobe
sculpture pattern (Fig. 148). Submedian lobes
of scutellum each with I pair of setae,
posterior seta well forward of posterior
margin of lobe. Propodeum posterior margin
an open V-shape, posterolateral angles
obtuse. Fore wing (Fig. 124) L:W 2.3:1;
marginal to submarginal vein length 1.6:1; 4
setae on submarginal vein; sligmal vein (Fig.
135); marginal to stigmal vein length 4.2:1;
submarginal to stigmal vein length 2.6:1;
terminal seta on postmarginal vein slightly
longer than those on marginal vein.
Male: Critical point dried specimens 1. 2-1. 3
mm long. Entirely honey brown in colour
except upper face in an area equivalent to that
between the facial grooves in female pale,
mesoscutum lightly infuscated, funicle
segment 4 plus club strongly infuscated.
Head in frontal aspect (Fig. 158) wider than
long L;W 1:1.4, transversely elliptical, lateral
margins broadly rounded; clypeus deeply
impressed, area above impressed clypeus and
below toruli with a dense tuft of fine setae
(similar to head setation); clypeal margin
without lobes, as Fig. 158. Mandibles (Fig.
169); anterior tooth relatively broad and
widely separated from second; second and
DAHMS: REVISION OF MELITTOBIA
299
third well defined, sub-equal, 2 slightly
longer. Maxillary palps (Fig. 181) as M.
hawaiiensis, cylindrical, short and broad,
L:W 2.5:1. Antennae (Figs 206-208); scape
broad, expanded evenly from proximal end
except for a pronounced constriction about
mid-way along scape; length to head length
1:1; scape L:W 1.8:1; ventral surface (Fig.
207) with a deep longitudinal groove,
proximal end of groove with only one seta;
glandular area geniculate, one arm more or
less transversely across distal scape, the other
along the side of groove opposite pedicel
attachment; flange overhanging scape groove
on same side as pedicel attachment with up to
5 setae, most of which are not on the margin;
distal scape margin more or less transverse,
not oblique; funicular segmental proportions
(Fig. 208) 1 short the smallest segment; 2 and
3 large, wider than long; 4 short, transverse,
cup-shaped, closely applied to club segment
1; MPS formula on flagellum 0000:141.
Mesosoma in dorsal aspect. Prothorax 1:1.5.
Parapsidial sutures and axillae reasonably
well defined though not as well as in M
clavicornis. Submedian grooves of scutellum
absent, sublateral grooves faint; 2 pair of
setae present, positioned as in female. Fore
legs with a dense tuft of short, stiff setae
(Dahms 1983b: Plate 4b); tarsal segments 3 +
4 fused. Mid leg (Fig. 227) trochanters
without a ventral, dense tuft of long, fine
setae; femoral fringe divided into 2 sections, a
short proximal tuft on basal 1/4 of femora, a
space without setae about equal to 1/4
femoral length followed by a fringe
occupying distal 1/2 of femur; setal fringe
slightly longer than 1.5 times width of femur;
L:W femur 4.3:1; tibia L:W 4.1:1; tarsi
without fused segments. Fore wings (Fig. 238)
elongate, apex rounded, L:W 3.7:1; marginal
to submarginal vein length 1.9:1; stigmal vein
reduced to a swelling on end of marginal vein;
costal cell narrow L:W 13.5:1; costal margin
slightly arched above.
MATERIAL EXAMINED:
QM Lectotype and paralectotypes as in TYPE
SPECIMENS section; 6 5? 4 SS card-
mounted ‘Brisbane SEQ, E.C. Dahms, Dec.
1979, ex Sce/iphron laetum 2 ? ? 2 ^ slide-
mounted 'Melittobia australica Grit. E.
Dahms det. 1980, Brisbane, SEQ, E.C.
Dahms, Dec. 1979 ex Sceliphron laetum,
Euparal E.C.D.'; 27 slide-mounted
'Melittobia australica Girault, Bred from
cells of Sceliphron laetum, 20.1.1918, H.
Hacker'; 16 4 second-form individuals
‘Brisbane, January 1981, E.C. Dahms ex
laboratory colony on Sceliphron formosum ’;
17 9? 4 card-mounted ‘Acacia Ridge,
Brisbane, SEQ, E. Dahms, 21.1.1979’, ‘ex
Stenarella victoriae 2 2 2 2 SS card-
mounted, 4 22 5 slide-mounted ‘Arita-
gun, Wakayama Pref., Japan, on Megachile
subalbuta, M, Matsuura, iii.1976’; 8 22 2
card-mounted, 4 2 2 2 slide-mounted
‘Kingston, Jamaica, West Indies, on
Sceliphron assimili, K, Ittyeipe, v.1975’; 8 22
2 S S card-mounted, 3 2 2 3 S S slide-
mounted ‘ex culture maintained at University
of Georgia, D.A. Evans, on Trypoxylon
striatum, v.l975’;2222c?<3 on a microscope
slide ‘Brits., Transvaal, Rep. South Africa on
Sceliphron sp., R.T. Simon Thomas,
xi.l974’.
In addition 4 2 2 and I S from the following
localities are deposited in BM(NH) and
USNM.
1) ‘Brisbane, SEQ, E.C. Dahms, Dec. 1980,
ex Sceliphron laetum ’.
2) ‘Brisbane, SEQ, E.C. Dahms, January
1981’, ‘ex laboratory colony on
Sceliphron formosum, second-forms’.
Melittobia hawaiiensis Perkins
(Figs 94, 95, 110, 111, 123, 136, 168, 170, 209,
210)
Melittobia hawaiiensis : Swezey, 1907 : 125.
Melittobia hawaiiensis Perkins, 1907 : 126.
Melittobia hawaiiensis peles Perkins, 1907 : 127.
Sphecophagus sceliphronidis Brtihes, 1911 : 209.
Sphecophilus sceliphronidis : Brethes, 1911 : 311
SYN NOV
Melittobia hawaiiensis : Masi, 1917 : 226.
Melittobia hawaiiensis : Ferriere, 1933 : 103.
Melittobia hawaiiensis : Gradwell, 1958 : 277.
Melittobia peles : Yoshimoto, 1965 : 683.
Melittobia hawaiiensis : Yoshimoto, 1965 ; 683.
Melittobia hawaiiensis : De Santis, 1973 : 18.
TYPE SPECIMENS:
Perkins (1907) did not indicate the location of
the type of this species and did not consider it
necessary ... ‘because the specimens could not be
preserved in satisfactory condition for subsequent
comparison’. Gradwell (1958) selected a neotype
300
MEMOIRS OF THE QUEENSLAND MUSEUM
of this species from a slide containing 112 2 1^
in the collections of the British Museum (Nat.
Hist.). I have examined this slide and confirmed
the separation of M. hawaiiensis and M.
australica. The separation is not easy, but it was
aided by fresher material of M. haw^aUensis and
M. australica , in conjunction with the fact that
the two will not interbreed (van den Assem pers.
comms. 1974-1980). Brethes (1911) described the
species Sphecophagus sceliphronidis and in 1911
changed the generic name to Sphechophilns
believing Sphecophagus to be preoccupied. De
Santis (1949) transferred this species to the genus
Melittobia. In 1957 De Santis made S.
sceliphronidis a junior synonym of M. acasta. I
have been unable to locate the type of S.
sceliphronidis, but from Brethes’ figures and
description, the species is definitely not M. acasta
— the scape has a ventral longitudinal groove.
The only species it could possibly be, given the
present state of knowledge of the world
Melittobia fauna, are M. hawaiiensis, M.
australica or sp, nov. Argentina. From Brethes’
figures it is clearly not the latter, De Santis (1973)
records M. hawaiiensis from Argentina but this
species is difficult to separate from M. australica
without slide preparations of the male scape. In
the absence of any material of these species from
South America and the type of S. sceliphronidis I
have provisionally placed M sceliphronidis as a
junior synonym of M. hawaiiensis subject to
confirmation.
DISTRIBUTION:
South America, Hawaii, New Zealand (= M
clavicornis Donovan (1953) and Cowley (1961))
( = species 7 of van den Assem, Bosch and Prooy
(1982)), Seychelles, Guam. In the literature there
are a great many localities given for this species
especially around the Pacific region, e.g.
Yoshimoto (1965). Because of the ease of
confusion of this species with M, australica I have
listed only the distribution of specimens I have
examined.
DESCRIPTION:
Female: In all aspects examined the females of
M. hawaiiensis and M. australica tend to
grade into one another e.g. variations in
shape of the stigmal vein overlap, the degree
of infuscation of the scape and pedicel is
variable within each species and overlaps
between species and so on. Given the present
state of knowledge of the two species I cannot
separate them on females.
Male: Again M. hawaiiensis and M. australica
males are very similar in most respects and
their variations overlap between species. Only
one consistent feature serves to distinguish
the species and that is setation on the scape.
Compare figures 206, 209 and 210. The
flange overhanging the groove of the scape,
on the same side as the pedicel is attached, is
relatively longer, and has more than 5 setae,
most of which are arranged on the edge of the
flange; the proximal floor of the groove has
more than 2 setae (generally 5 or more) as
opposed to 1-2 in M. australica. The last
mentioned setal arrangement appears very
reliable in all specimens of the species that I
have examined.
In addition to these species I have specimens of
an hawaiiensis -type species from Kauai, Hawaii
(= species 7/8 of van den Assem et alia (1982)).
Again the females appear very similar to M.
hawaiiensis and M, australica except that the
nipple on club segment 3 is longer and narrower
(Figs 110-112). Males also are very similar except
in the setation patterns on the scapes. The flange
overhanging the scape groove is relatively short as
in M. australica, it has more setae than in M,
hawaiiensis and these setae are distributed along
the entire length of the flange, whereas they are
more restricted in M. australica and M.
hawaiiensis. The mandibles of males also show
some differences (Figs 168-170). However 1 am
rather hesitant to describe the Kauai material as a
new species. This whole complex is in need of a
detailed morphometric study which could be tied
in with ethological work of van den Assem et alia
(1982). In the summary of this paper aspects of
crossing experiments by van den Assem et alia
(1982) with hawaiiensis group species and Kauai
are discussed.
MATERIAL EXAMINED:
BM(NH) Neotype slide with 1 1 2 ? 1 c? as
figured by Grad well (1958).
1 9 card-mounted ‘Mahe ’08-9, Seychelles
Exp.’, ‘Percy Sladen Trust Exped., BM
1913-170’, 'Melittobia hawaiiensis Perkins,
L. Masi det.’
QM 7 2 2 1 c? on a microscope slide 'Melittobia
hawaiiensis Perk., Gahan, ex Pison
argentatum, Piti, Guam 9-27-36, O.H.S.’; 3
2 9 3 <?<? on microscope slides, 4 22 2
card-mounted 'Melittobia hawaiiensis
Perkins, ex Lab culture Est. from Te Pirita,
Cantebury, New Zealand, Sept. 1974, B.J.
Donovan ex Pison spinolae, E.C. Dahms
DAHMS: REVISION OF MELITTOBIA
301
EuparaF; 6 2? 3 on microscope slides, 1
2 1 S card mounted ‘ex lab culture est. from
Kilauea, Kauai, Hawaii, 21, xi. 1976, S.L.
Montgomery and J. Maciolek 100’ ex mud
nest, E.C. Dahms Euparal*.
Melittobia assemi SP NOV
(Figs 67, 79, 125, 137, 149, 159, 171, 182, 211,
212, 213, 228)
TYPE SPECIMENS:
Holotype ct 5 2 paratypes on the one card
‘Anse Bazarca, Mahe Island, Seychelles, ex
eumenid species, R.T. Simon Thomas, ii.l976’
BM(NH); 2 2 paratypes card-mounted, 4 2 2
paratypes on a slide with data as holotype (QM).
DISTRIBUTION:
Mahe Island, Seychelles ( M, hawaiensis (in
part) of Masi (1917)), Kerala Forest Reserve,
India (= species 5 of Assem, Bosh and Prooy
(1982))
DESCRIPTION:
Female: Critical point dried specimens 1.3 mm
long. Head, flagellum, mesosoma, coxae
dark brown; proximal 2/3 femora, metasoma
slightly paler; scape, pedicel, remainder of
legs pale yellow-brown; upper scape and
pedicel lightly infuscated.
Head in frontal aspect as M sosui (Fig, 47)
length to genal width 1.3:1; genal-clypeal
margin broadly rounded; clypeal margin (Fig.
67) bilobed, lobes small, each sharply
separated from a small, lateral, lobe-like
undulation. Eyes densely clothed with long
setae. Facial grooves converging to meet just
above middle of eyes then passing as a single
line to scrobes; maximum distance apart of
arms greater than POL. Scrobe to eye length
2.6:1. Mandibles as M. sosui (Fig. 57)
anterior tooth very small, broad; second and
third well defined, broad, 3 the broadest.
Maxillary palp (Fig. 79) short, broad, L:W
2.5:1. Antennae as M. sosui (Figs 100, 101)
scape broad, relatively strongly curved, L:W
2.8:1; MPS formula on flagellum 335:263;
segment 3 of club as M. sosui (Fig. 113)
shortest length to width 1:2; nipple relatively
long, L:W 3:1; subterminal seta about mid-
way down nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.5. Posterior margin of mesoscutum mid
lobe 1 . 1 times wider than anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1,9:1; 3-4 setae on each sublateral lobe.
Propodeum posterior margin an open V-
shape, posterolateral angles obtuse. Fore
wing (Fig. 125) L:W 2.4:1; marginal to
submarginal vein length 1.5:1; 5 setae on
submarginal vein, the proximal 2 about 1/2
length of others; stigmai vein (Fig. 137);
marginal to stigmai vein length 3.4:1;
submarginal to stigmai vein 3,7:1.
Male: Air dried ex alcohol about 1.3 mm long.
Head, body and legs pale golden brown;
antennae also, except funicle 4 and club
which are infuscated.
Head in frontal aspect (Fig. 159) broad,
vertex broadly rounded, hardly raised, lateral
margins broadly rounded contracting slowly
below eye spots; genal-clypeal margin
broadly rounded; clypeal margin bilobed,
lobes long, narrow, clypeus deeply excised
between; clypeus deeply impressed, area
above impression and below toruli with a
dense cluster of long, fine setae. Mandibles
(Fig. 171) broad, anterior tooth of medium
length, broad, well separated from second;
second and third tooth shallowly defined,
broad. Maxillary palps (Fig. 182) tapered
distally, short, broad, L:W 2.2:1. Antennae
(Figs 211-213); scape more or less gradually
expanded from proximal end, whole scape
curved, concave on outer margin; ventral
surface with a deep, longitudinal groove,
more open than M. australica (Fig. 206),
flange over-hanging groove on side of pedicel
attachment narrow, without long setae;
glandular area geniculate, but distal arm not
as transverse as in M. australica (Fig. 207);
distal scape transverse, not excavated (Figs
211-212); scape length to head length 1.3:1;
scape L:W 1.6:1; pedicel tends to be concave
on inner margin as in M. sosui (Fig. 216);
funicular segmental proportions (Fig. 213)
1-3 sub-equal, 4 transverse, cup-shaped,
closely applied to segment 1 of club; MPS
formula on flagellum 0000:162.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.3. Parapsidial sutures absent, axillae
poorly defined. Scutellum without submedian
or sublateral grooves; 3-4 pairs of setae on
scutellum positioned as in females. Fore
trochanters without a ventral, dense tuft of
stiff, short setae; tarsi (Fig. 20) with 2
segments, 2-t-3-f-4 fused. Mid legs (Fig. 228);
trochanters with 6-12 long, curved, stiff
setae; femoral fringe of uneven length,
302
MEMOIRS OF THE QUEENSLAND MUSEUM
sparse, proximally absent, followed by a few
short setae, then a median fringe of setae not
quite long as width of femur, the distal c. 1/4
of fringe consists of short setae, width of
femur to length of distal fringe is about 2.5:1;
L:W femur 3.6:1; tibia L:W 3.7:1; mid tarsi
with 3 segments, segments 1 and 2 with a
posterior comb of long thick setae. Fore wing
as M. sosui (Fig. 239) long, narrow, L:W
4.1:1, apex acute; marginal to submarginal
vein length 1 .7:1; costal cell narrow, L:W 6:1;
stigmal vein reduced to a large swelling at the
end of marginal vein.
MATERIAL EXAMINED:
Type specimens as in TYPE SPECIMENS
section.
QM 1 5 W H- W head on microscope slides
‘Nilambur Kerala State, South India, from
H. van den Assem, Feb. 1980, E.C. Dahms
EuparaP.
BM(NH) 4 2$ card-mounted ‘Percy Sladen
Trust expedition, BM 1913-170*, ‘Mahe
*08-9, Seychelles Exp.’, '‘Melittohia
hawaiiensis Perkins, L. Masi det.’.
This species has been named in recognition of
Dr van den Assem, Leiden who has been very
generous with notes from his ethological studies
and with specimens.
Melittobia sosui SP NOV
(Figs 47, 57, 80, 100, 101, 113, 138, 172, 183,214,
215, 216, 230, 239, 240)
TYPE MATERIAL:
Holotype 5 2 paratypes (one 2 minus head)
card-mounted ‘Sosu, Okinawa IsL, Ryuku Arch.,
Japan, Y. Maeta, 23.xii.1978 ex Eumenid’ (KU);
3 2 2 paratypes on slides with data of holotype
(QM).
DISTRIBUTION:
Okinawa IsL, Japan (= species 4 of van den
Assem and Maeta (1980)) ( = species 6 of van den
Assem et alia (1982)).
DESCRIPTION:
This species is very close to M. assemi and the
description to follow merely consists of diagnostic
differences from M. assemi.
Female: Critical point dried specimens 1.3-1. 4
mm long. Coloration as in M. assemi.
There are some proportional differences
between M. assemi and M. sosui females, but
these are not very significant given the natural
variation in Melittobia and the polymorphism
in the type-forms of M. sosui recorded by van
den Assem and Maeta (1980). Therefore it
would be unwise to rely upon these
proportional differences for species
separation. As with the M. hawaiiensis
complex this species group is in need of
detailed morphometric analysis.
In females that I have for examination there
appear to be three consistent features which
are of use:
1) Of the 5 setae on the submarginal vein in M.
assemi the proximal 2 are about 1/2 the
length of the remaining 3 whereas in M. sosui
the 5 are of equal length.
2) On each submedian lobe of the scutellum, M.
assemi has 3-4 setae, occasionally with
variation between left and right on the one
specimen, whereas in M. sosui I observed a
consistent 2 on the left lobe and 3 on the
right.
3) Stigmal veins are different (Figs 137, 138).
Males of the two species are also very similar
but there are a few consistently different features:
1) The mandibles of M. sosui (Fig. 172) are
shorter and broader than M. assemi (Fig.
171).
2) The maxillary palps in M. sosui (Fig. 183) are
shorter than in M. assemi (Fig. 182); L:W
1.4:1 as opposed to 2.2:1 for M. assemi.
3) The scape of M. sosui (Figs 214, 215) is
narrower than M. assemi (Figs 211, 212);
L:W 2:1 as opposed to 1.6:1 for M. assemi.
In M. sosui the flange overhanging the scape
groove is broader than in M. assemi and bears
several long setae (up to 5).
4) The mid femoral fringe has more setae and
covers a greater length of the femur in M.
sosui than in M. assemi (Figs 228, 230). The
shapes of the femora are different; L:W M.
sosui 3.8:1, M. assemi 3.6:1.
MATERIAL EXAMINED:
As in TYPE SPECIMENS section.
This species is named M. sosui after its type-
locality.
Melittobia bekiliensis Risbec.
Melittobia bekiliensis Risbec, 1952 : 253.
DAHMS: REVISION OF MELITTOBIA
303
TYPE MATERIAL:
2 2 $ I ^ on minutien pins ‘MADAGASCAR,
Bekily, REG SUD DE LTLE’, ‘MUSEUM
PARIS, vi.36, A. SEYRIC’, ^Melittobia
bekiliensis Risb\ ‘Syntype’. From this syntypical
series I have selected the male as the lectotype and
the females as paralectotypes. They reside in the
collections of the Musee National d’Histoire
Naturelle, Paris.
DISTRIBUTION;
Madagascar.
DESCRIPTION:
Female: Air dried specimens 1. 0-1.1 long.
Head, pedicel, flagellum, mesosoma dark
brown; metasoma slightly paler; scape, legs
pale yellow-brown.
These specimens appear to be very similar to
females of M. assemi. It is difficult to
separate them without making slides. This
difficulty is increased due to disruption of the
thorax by the minutien pin and reliable
diagnosis of the female must await fresh
material.
Male: Air dried specimen. About 1.2 mm long,
specimen curled.
Head in frontal aspect resembling that of M.
assemi (Fig. 159); not contracting ventrally as
strongly as M. assemi and the clypeal region
is barely impressed; setation below toruli not
dense as in M, assemi; clypeal margin
bilobed, lobes not as long as M. assemi
Mandibles not projecting below clypeal
margin when closed. Maxillary palps
contracting distally as M. assemi (Fig. 182).
Antennal scapes quite distinctive, expanding
evenly from proximal end (no constrictions as
in M, australica (Fig. 207)), pyriform, not
curved as in assemi (Fig. 212), dorsal surface
smoothly rounded; ventral surface with a
deep longitudinal groove, relatively open as
in M. assemi; flagellum difficult to see;
segment 1 very small; 2-1-3 slightly larger; 4
very wide, asymmetrically cup-shaped,
closely applied to club, longest side of 4
nearly covering club segment 1.
Mesosoma in dorsal view. Prothorax wide,
triangular, L:W 1:3. Fore legs as in M. assemi
(Fig. 20). Fore trochanters without a ventral
tuft of short, stiff setae; fore tarsi 2
segmented. Mid legs as M. assemi except
distal setae on femoral fringe are not shorter
than those of medial fringe; tarsal segments
appear unfused. Fore wings resemble those of
SP NOV Argentina (Fig. 241), apex broadly
rounded; stigmal vein well developed.
This is quite a distinctive species in the male. Its
scape, maxillary palp and femoral fringe place it
with M, assemi and M. sosui.
Argentina SP NOV
(Figs 48, 58, 69, 81, 98, 99, 114, 126, 139, 150,
160, 173, 184, 217, 218, 219, 229, 241)
TYPE SPECIMENS:
Material too poor for type selection.
DISTRIBUTION:
Argentina.
DESCRIPTION;
Female: Air dried specimens from alcohol 1.1
mm long. Head, mesosoma, antennal
flagellum, coxae, proximal 1/3 of femora
dark brown; scape, pedicel and remainder of
legs yellow-brown; metasoma slightly paler
than mesosoma.
Head in frontal aspect (Fig. 48) broad, length
to genal width 1.2:1; genal-clypeal margin
broadly rounded; clypeal margin (Fig. 69)
bilobed, lobes broad, weakly developed; eyes
densely covered with long setae. Facial
grooves converging separately to scrobes;
maximum width between arms 2 times
diameter of median ocellus; arms converging
gradually to meet scrobes just below middle
of eyes; minimum distance between arms 0.25
times diameter of median ocellus. Scrobe to
eye length 1:1.9. Mandibles (Fig. 58); anterior
and median tooth long, narrow, third shorter
and broader. Maxillary palp (Fig. 81)
elongate, of medium length, L:W 4:1.
Antennae (Figs 98, 99); scape broad, L:W
2.5:1; MPS formula on flagellum 445:463;
club segment 3 (Fig. 114) shortest length to
width 1:2; nipple relatively long, L:W 3:1; 2
subterminal setae situated in proximal 1/3 of
nipple.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.8. Posterior margin of mesoscutum mid
lobe equal to width of anterior margin of
scutellum mid lobe. Scutellum mid lobe L:W
1.8:1; 1 pair of setae on each submedian lobe,
posterior situated on posterior margin of this
lobe. Mesoscutum and scutellum mid lobes
sculpture pattern (Fig. 150). Fore wing (Fig.
126) L;W 2.2:1; marginal to submarginal vein
304
MEMOIRS OF THE QUEENSLAND MUSEUM
length 1.6:1; 4-5 setae on submarginal vein;
stigmal vein (Fig. 139); marginal to stigma!
vein length 4.8:1; submarginal to stigmal vein
length 3.1:1; terminal seta on postmarginal
vein no longer than those on the marginal
vein.
Male: Air dried specimen from alcohol,
metasoma absent, length of head mesosoma
0.5 mm. Head, scape, funicular segments
2-3, legs pale brown; funicular segment 4
club strongly infuscated.
Head in frontal aspect (Fig. 160) broad, more
or less rectangular, lateral margins not
contracting strongly to clypeus, genae slightly
indented below eye spots, L:W almost 1:1;
clypeus impressed, area above clypeus and
below toruli with a dense tuft of long, thick
setae; clypeal margin bilobed, lobes broad,
well defined. Mandibles (Fig. 173) very
broad, projecting well below clypeus when
closed. Maxillary palps (Fig. 184) very
distinctive, broad, distally excavated, L:W
1.4:1. Antennae (Figs 217-219); scape
relatively evenly expanded from proximal
end, distal 1/2 expanding rather suddenly;
ventral surface (Fig. 218) with a deep
longitudinal groove, more open than M.
australica; glandular area rather amorphous,
extending along groove; scape to head length
1:1.2; scape L:W 1.9:1; distal scape more or
less truncate, without an excavation;
funicular segments (Fig. 219) transverse, 1 the
narrowest; 2 + 3 sub-equal; 4 as wide as 2 and
3, cup-shaped, closely applied to club
segment 1; MPS formula on flagellum
0000:131.
Mesosoma in dorsal aspect. Prothorax L:W
1:1.6. Parapsidial sutures and axillae poorly
defined. Scutellum without submedian and
sublateral grooves. Fore legs (Fig. 21);
trochanters without a dense tuft of short,
stout setae; tarsal segments fused into 1. Mid
legs (Fig. 229); trochanters with a few,
curved, short, thick setae; femoral fringe not
completely lining femora, basal 1/4 with very
short, normal setation, distal 3/4 of even
length, slightly longer than width of femur;
ventral surface of femur appears grooved to
receive tibia; L:W femur 7.2:1; tibia L:W
5.7:1; mid tarsi of 2 segments, 2 + 3 + 4 fused.
Fore wing (Fig. 241) relatively broad, L:W
3:1; marginal to submarginal vein length
1.8:1; costal cell narrow L:W 6:1, costal
margin above slightly arched; stigmal vein
well developed.
MATERIAL EXAMINED;
CU,NY 5 5fragments card mounted as follows
— 3 heads with antennae; 2 heads plus
mesosoma and 1 fore wing; 1 complete
specimen except for antennae. 1 S head plus
mesosoma without wings; several
fragmentary females in alcohol ‘Ascasubi
Argentina, Dec. 1976, R.H. Gonzalez’, ‘ex
Megachile rotundata 3 25 1 c? on a
microscope slide with above data.
I have not named this species because the
specimens are too poor for type-selection.
INCERTAE SEDIS
Melittobia osmiae Thompson
Melittobia osmiae Thompson, 1878 : 204.
Melittobia acasta (?) : Domenichini, 1966 : 56.
TYPE SPECIMENS:
Not located.
DISTRIBUTION:
Europe.
Domenichini (1966) provisionally placed this
species as a junior synonym of M. acasta. It may
well fit here, but the description is not diagnostic
and I could not locate the type. For these reasons
it was decided to leave it as a separate species
awaiting confirmation.
Melittobia hawaiiensis peles Perkins
Melittobia hawaiiensis peles 1907 ; 125.
Melittobia peles : Yoshimoto, 1965 : 683.
TYPE SPECIMENS:
Not located.
DISTRIBUTION:
Oahu, Hawaii.
I have been unable to locate the type of this
variety and the brief description by Perkins is
insufficient to allow recognition of this taxon.
SUMMARY:
Van den Assem and Maeta (1978, 1980) using
ethological criteria have divided the genus into
acasta group, hawaiiensis group and Mahe group
(= assemi group) and have kept M. clavicornis
separate as the most primitive species. When one
looks at the comparative morphology of the
males a similar grouping applies using the
following characteristics:
DAHMS: REVISION OF MEUTTOBIA
305
acasta
hawaiiensis
assemi
1) scape:
cup
grooved
grooved
2) gland in scape:
transverse
geniculate
geniculate
3) proportions of funicular
segments:
1 largest
1 smallest
all equal
4) funicular seg. 4 cup shaped and
closely applied to club 1:
-
+
+
5) presence of plate organs on
funicle:
+
-
-
6) mid trochanters with a dense tuft
of long fine setae:
+
-
-
7) wings broad:
+
~
-
hawaiiensis group
hawaiiensis
australica
Kauai
assemi group
assent i
sosui
bekiliensis
Argentina
The grouping on this basis is as follows:
acasta group
acasta
evansi
scapata
femorata
digitata
chalybii
megachilis (?)
Of these groups, hawaiiensis and assemi appear
closest. I regard the unifying characters as derived
relative to M. clavicornis and considering the
shared possession of these derived characters,
these two groups could confidently be regarded as
monophyletic sister groups. Species within these
groups can be sorted on a mixture of derived and
relatively primitive features, and this again points
to the monophyiy of each group. The division is
as follows:
1) head
2) mandibles
3) funicle segments
4) mid femoral fringe
5) fore trochanters with a dense tuft of short setae
6) mid trochanters with 6-12 long, stiff curved seta
7) ventral fore trochanters with a dense setae tuft
of stiff setae
hawaiiensis group
elliptical *
narrow
1 small 2 + 3 large *
long proximally *
+ *
no long setae *
+ *
assemi group
rectangular *
broad *
1 — 3 sub-equal *
short proximally
+ *
(* = characters I regard as derived)
The hawaiiensis group has relatively few species
which may be a reflection of our state of
knowledge of the world fauna. The species M.
hawaiiensis and M. australica is very close
morphologically and ethologically, but van den
Assem et alia (1982) found that they are
reproductively isolated when they tried crossing
them. However, they discovered that females of
both M. hawaiiensis and M. australica when
crossed with males of Kauai produced fertile
female offspring. Reciprocal crosses gave similar
results. It appears therefore that male courtship in
M. hawaiiensis and M. australica is not the
same, but there are elements of Kauai male
courtship which make them reproductively
compatable with either M, australica and M.
hawaiiensis and vice versa. This is a rather
interesting situation since both Af. hawaiiensis
and Kauai occur on the islands of Hawaii and
given the capability of Melittobia to be wind
dispersed it is hard to imagine that geographical
barriers operate. Given the highly polyphagous
nature of Melittobia one can argue against
ecological isolation. It appears therefore that the
306
MEMOIRS OF THE QUEENSLAND MUSEUM
hawaiiensis group is a relatively young group in
the process of speciation. In contrast the assemi
group contains more species showing greater
morphological diversity. The comparative
ethology of M. assemi and M. sosui only, is
known for this group. These two species are very
close morphologically and ethologically, (van den
Assem and Maeta 1980), which indicates a fairly
recent separation. M. bekiliensis is close to M.
assemi on the basis of head and palp shape but its
scape and funicle shape I regard as more derived.
Sp. nov. Argentina is the most derived species on
the basis of mandible and palp shape. The greater
number of species in the assemi group and their
greater morphological diversity suggests that this
group is relatively older than the hawaiiensis
group.
The acasta group, however, is less easy to
divide phylogenetically. The relatively larger
number of species in this group and their
morphological and ethological diversity suggests
an earlier origin for this group. The characters
used for grouping the species are relatively
primitive ones (many are shared with M.
clavicornis ) and therefore the group may be
paraphylelic rather than monophyletic. Male
morphological differences are closely related to
the use of appendages and body parts during
courtship. Without this knowledge it is difficult
to place characters which separate species into a
phylogenetic order with any confidence.
Two species, M acasta and M. digitata can be
grouped on their oblique distal scape, the
excavation of this margin and the transverse,
narrow shape of the scape gland. I regard M.
digitata as the most derived and van den Assem
(pers. comms. 1974-1980) regards it to be derived
on ethological data also. The reduction in relative
size of funicle segment I and the size of the scape
I regard as derived and unify M. evansi and M.
scapata. Two species, M. femorata and M.
chalybii both possess an extra funicle segment as a
result of expansion of one of the ring segment
lamellae. Their scape shapes are more similar to
one another than to any other species. Figs 203,
204 of M. chalybii were from a slide in which the
scapes are slightly rolled. In dry specimens the
excavation of the scape in M chalybii is of the M.
femorata type but not so deep. I regard M,
chalybii as the more derived since the scape gland
is geniculate and the mid femoral fringe is more
even (the primitive condition appears to be distal
fringe much longer than proximal fringe).
The situation however, may not be this simple.
Two species, M. chalybii and sp. nov. Argentina,
do not entirely fit the species groupings on
morphological data. The courtship patterns of
these two species are not known, but correlating
morphology with known courtship patterns
allows some interpretive discussion.
Males of M. chalybii have setae on the ventral
fore trochanters not unlike those of the
hawaiiensis group although not as dense, short or
stiff and the male scape gland is geniculate
although not as well developed as in the assemi
and hawaiiensis groups. However, in scape shape
and proportions of the funicular segments (even
to the extra, expanded ring segment) M. chalybii
is extremely similar to M, femorata which
morphologically is very definitely an acasta group
species. The long setae on the male mid femora in
M. chalybii are also of the acasta group pattern.
If we look at the females of M. chalybii we find
that they have the acasta group narrowly spaced
facial grooves, but the eyes are densely setose as
in the hawaiiensis and assemi groups. Thus we
find a mixture of morphological features in M.
chalybii which can be found in all three groups.
Sp. nov. Argentina males are easily placed in the
assemi group on all features except for relatively
broad wings. The females differ in that their
facia! grooves are narrowly spaced as in the acasta
group.
Turning now to courtship, we find that in M.
australica {hawaiiensis group) the male courtship
position involves close application of his
mouthparts onto the relatively broad area
between the facial grooves of the female, in fact I
observed that this area of the female is pushed
inwards by the male’s mandibles. I have not
observed courtship of the assemi group species
but in M. assemi and M. sosui the male position
as reported by van den Assem and Maeta (1980)
and van den Assem et alia (1982) resembles that
of the hawaiiensis group. From their discussions
it is not clear whether the male’s mandibles
impinge on the area between the facial arms of the
female in these two species, but since the area
between the facial arms in females of these species
is broad there may be some correlation between
breadth of this area and male position. If this is so
then the courtship position of sp. nov. Argentina
it not as in the hawaiiensis group but may be more
like the acasta group where this area in females is
relatively narrow. Another factor in both sp. nov.
Argentina and M. bekiliensis (both assemi group)
is the relatively broad male wings more like acasta
group males than males of the hawaiiensis group
or M. assemi and M. sosui. Broad male wings and
wing vibration by males during courtship are a
DAHMS: REVISION OF MELITTOBIA
307
correlation in the acasta group as are narrow male
wings and no wing vibration during courship by
hawaiiensis group species as well as the species M.
assemi and M. sosui. Perhaps male wing vibration
also occurs during courtship in sp. nov. Argentina
and M. bekiliensis.
Species where the male scape gland is geniculate
involve permanent antennal contact during
courtship (hawaiiertsis group, M. assemi, M.
sosui ) which contrasts with permanent contact
either through only part of the courtship (M.
acasta, M. evansi ) or not at all (M digitata ). In
M. chalybii the geniculate nature of the male
scape gland may indicate that permanent antennal
contact is more important during courtship in this
species than in other acasta group species. The
presence of the setal tuft on the male ventral fore
trochanters in M. chalybii indicates that there
may be some similarities between male courtship
position in M. chalybii and the hawaiiensis group
where these setae rest on the female’s mesosoma.
However, it may not be entirely so as M. chalybii
females have narrowly separated facial grooves.
The presence of densely setose eyes in females of
the hawaiiensis and assemi groups correlates with
a predominance of mid leg action during
courtship. M. chalybii females have densely
setose eyes which perhaps indicates that mid leg
action during courtship in this species assumes a
more important role than it does in the coursthip
of other acasta group species.
From this speculative evidence there is some
suggestion of convergent evolution in courtship
behaviour and associated morphology in
Melittobia. We may therefore be dealing with a
polyphyletic group rather than a monophyletic
one. Of key importance in understanding this are
the coursthip patterns of M. chalybii and sp. nov.
Argentina coupled with a more thorough
knowledge of the world fauna. Africa and South
America will no doubt yield many more species
than are presently known.
ACKNOWLEDGEMENTS
This paper was taken from my M.Sc. thesis
submitted to the University of Queensland in
1982. My superviser, Dr Elizabeth Exley,
University of Queensland, was extremely helpful
in providing constructive comments and editorial
remarks. Dr T. Woodward. University of
Queensland and Dr G. Gordh, University of
California as examiners, provided corrections and
advice towards publication of the thesis. Dr
Gordh was also of great assistance, imparting to
me many of his illustration techniques.
My Technician Miss Gudrun Sarnes has been
very helpful in manuscript corrections, German
translations and numbering figures. The typists
whose patience I tried severely were Miss P.
Tinniswood and Miss E. Proberts. My wife
Judith assisted with manuscript checking and
figure assembly.
Many people were of assistance with specimens
for examination: Dr L. Hoberlandt, National
Museum (Nat. Hist.) Prague; Dr D.A. Evans,
Kalamazoo College, U.S.A.: Mr K. Ittyeipe,
University of West Indies, Jamaica; Dr G.
Funisaki, State Department of Agriculture,
Hawaii; Dr J. Noyes, British Museum (Nat.
Hist.), London; Dr L. De Santis, Museum de La
Plata, Argentina; Dr P. Hurd, United States
National Museum, Washington D.C., U.S.A.;
Mr J.C. Hall, University of California, Riverside,
California; Dr Y. Maeta, Tohoku National
Agricultural Experiment Station, Japan; Dr T.
Tachikawa, Ehime University, Japan; Dr Y.
Hirashima, Kyushu University, Japan; Dr J.
Beardsley, University of Hawaii, U.S.A.; Prof.
H. Morge, Deutsche Akademie der
Landwertschaftswissenschaften, German Demo-
cratic Republic; Dr R. Macfarlane and Dr B.
Donovan, D.S.I.R., New Zealand; Dr R.
Quentin, Musee National d’Histoire Naturelle,
Paris, France; Miss M. Pearce, Museum of
Comparative Zoology, Harvard U.S.A. and Dr
G. Eickwort, Cornell University, New York,
U.S.A.
Special thanks are due to Dr J. van den Assem,
University of Leiden, Holland. We have
corresponded freely since 1974 and he has been of
the greatest assistance with notes from his
ethological studies on Melittobia and provision of
specimens for study.
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Smithsonian Press, Washington D.C., 1967 :
430-3.
Meata, Y, 1978. A preliminary study of the
physical control of Melittobia acasta
(Walker) by cold treatment (Hymenoptera :
Eulophidae). Bull. Tohoku natn. agric. exp,
stn. 58:211-29.
and S. Yamane, 1974. Host records and
bionomics of Melittobia japonica Masi
(Hymenoptera, Eulophidae). Bull, Tohoku
natn. agric. exp. stn. 47: 115-31.
Masi, L., 1917. Chalcididae of the Seychelles
Islands. Novit. zool. 24: 121-230.
1936. On a supposed new species of Melittobia
from Japan. Mushi 9; 38-9.
Michener, C.D., 1956. Hymenoptera. In
Taxonomists glossary of genitalia in insects.
Ed. Tuxen, S.L., 1956, Ejnar Munksgaard,
Copenhagen: 131-40.
Morely, C., 1910. Catalogue of British
Hymenoptera of the family Chalcididae.
British Museum (Nat. Hist.) Lond. 1910, 28
pp.
Neave, S.A., 1939. Nomenclator Zoologicus.
Zool. Soc. Lond., 1939 Vols 1-4.
Newport, G. 1849a. Anthophorabia retusa.
Gdnrs* Chron., 1849 : 183.
1849b. On the identification of a new genus of
parasitic insects, Anthophorabia. Ann. Mag.
nat. Hist. (2) 3 : 513-7.
1849c. On the identification of the parastiic
genus of insects, Anthophorabia. Ann. Mag.
nat. Hist. (2) 4: 122-4.
1852a. The anatomy and development of
certain Chalcididae and Ichneumonidae,
compared with their special oeconomy and
instincts; with descriptions of a new genus
and species of bee parasites. Trans, Linn.
Soc. Lond. 21: 61-77.
1852b. Further observations on the genus
Anthophorabia. Trans. Linn. Soc. Lond. 21:
79-83.
1853. On the ocelli in the genus
Anthophorabia. Trans. Linn. Soc. Lond. 21:
161-5.
Packard, A.S., 1864. Humble bees etc. of New
England. Proc. Essex Inst. 4: 134.
1868. The parasties of the Honey-bee. Am.
Nat. 2: 195-205.
310
MEMOIRS OF THE QUEENSLAND MUSEUM
1869. Guide to the study of insects. Henry Holt
& Co. 1st ed. 1869, 715 pp.
Peck, O., 1951. Hymenoptera of America North
of Mexico. Synoptic catalogue. Ed.
Muesebeck, C.F.W. and Krombein, K.V.,
1951. U.S, Dept, Agric. Monograph (2);
410-594.
1963. A catalogue of the Nearctic Chalcidoidea
(Insecta : Hymenoptera). Mem. ent, Soc.
Can. 30: 1-1092.
Z. BoufeK, and A. Hoffer, 1964. Keys to the
Chalcidoidea of Czechoslovakia (Insecta :
Hymenoptera). Mem. ent. Soc. Can. 34:
1 - 120 .
Perkins, R.C.L., 1907. Melittobia hawaiiensis
sp. nov. (Hymen). Proc. Hawaii, ent. Soc. 1:
124-5.
Prinsloo, G.L., 1980. An illustrated guide to the
families of African Chalcidoidea (Insecta ;
Hymenoptera). Sci. Bull. Dep. Agric. Fish.
Repub. S. Afr. (395): 1-47.
Risbec, J., 1951. I. Les chalcidoides d’ A.O.F.,
Mem. Inst. fr. Afr. noire (13): 7-409.
1952. Contribution a Fetude des chalcidoides
de Madagascar. Mem. Inst. Scient.
Madagascar 2: 1-499.
1956. Hymenopteres parasties du Cameroun
(2nd and 3rd contributions). Mem. Inst. fr.
Afr. noire (18): 97-164.
Santis, L., De, 1949. Dos notas sobre
chalcidodieos Argentines (Hymenoptera :
Chalcidoidea), Notas Mus. La Plata (ZooL)
14: 275-81.
1957. Anotaciones sobre chalcidoideos
Argentinos (Hymenoptera). Notas Mus. La
Plata 19: 107-19.
1973. Himenopteros parasitos de Megaquilos
en la republica Argentina. Revta Cienc.
Abejas 2: 15-9.
Schmiedeknecht, O., 1909. Hymenoptera Fam.
Chalcididae. Genera Insectorum 97: 1-548.
Smith, F., 1853. Notes on the habits of a bee
parasite Melittobia acasta. Trans, ent Soc.
Lond. (N.S.) 2: 248-53.
Swezey, O.H., 1907. Odynerus parasites. Proc.
Hawaii, ent. Soc. 1: 121-3.
Thompson, C.G., 1878. Melittobia osmiae.
Hym. Scandin. 5: 204.
ViGGiANi, G., 1971. Ricerche sugli Hymenoptera
Chalcidoidea XXVIII. Studio morfologico
comparativo delF amatura genitale esterna
maschile dei Trichogrammatidae. Boll Lab.
Ent. agr. Filippo Silvestri 29: 181-222.
1971. The significance of the male genitalia in
the Trichogrammatidae (Hymenoptera :
Chalcidoidea). Proc. 13th. Int. Congr. Ent.,
Moscow 1; 313-5.
Viereck, H.L., 1916. Miotropis megachilis. Bull.
Conn. Si. geol. nat. Hist. Surv. 22: 465.
Walker, F., 1893. Monographia chalciditum 1:
1-328.
Waterson, J.W., 1917. Notes on the
morphology of Chalcidoidea bred from
Calliphora. Parasitology 9: 190-8.
Westwood, J.O., 1840. An introduction to the
modern classification of insects; founded on
the natural habits and corresponding
organisation of the different families, 2: 587
pp.
1847. Melittobia audouinii. Proc. ent. Soc.
Lond. (1847) : XVII.
1849a. Melittobia audouinii. Gdrns'; Chron.
(19), 1849 : 295.
1849b. Description of Melittobia audouinii, a
bee parasite, Proc. Linn. Soc. (1849) : 37.
1849c. Melittobia audouinii. Proc. ent. Soc.
Lond. (1849) : LXV.
1849d. On the identification of a genus of
parasitic Hymenoptera. Ann. Mag. nat. Hist.
(2) 4: 39-41.
Wolff, M. and A. Krausse, 1922. Nachtrag zu
unserem Aufsatz "uber Melittobia strandi
n.n.sp. Arch, Naturgesch. 86, Abt. A, H.6:
16-21.
Yoshimoto, C., 1965. Synopsis of Hawaiian
Eulophidae including Aphelinidae (Hym. :
Chalcidoidea). Pacif. Insects 1: 665-99.
DAHMS: REVISION OF MELITTOBIA
311
FIGURES 23, 24, 30-31, Tachinobia diopsisephila Female; 23 — Head, frontal aspect; 24 — Mesosoma, dorsal
aspect; 26 — Scape and pedicel; 27 — Flagellum, dorsal aspect; 28 — Club, lateral aspect; 29 — Club segment 3;
30 — Fore wing; 31 — Stigmal vein. Fig. 25 — Tachinobia repanda female head, dorsal aspect.
312
MEMOIRS OF THE QUEENSLAND MUSEUM
36
0.05
FIGURES 32-37, Cirrospilus (Atoposomoidea) cosmopterygi female; 32 Head, frontal aspect, 33 Mesosoma,
dorsal aspect; 34 - antenna; 35 - wings; 36 - Club segment 3; 37 - Stigmal vein.
DAHMS: REVISION OF MELITTOBIA
313
FIGURES 38-43, Female heads, Melittobia spp. 38 — clavicornis; 39 — acasta; 40 — evansi; 41 — scapata; 42 —
digitata; 43 — femorata.
314
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 44-48, Female heads, Melittobia spp. 44 — chalybii; 45 — megachilis; 46 — australica; 47 — sosui;
sp. nov. Argentina.
DAHMS: REVISION OF MELITTOBIA
315
FIGURES 49-58, Female mandibles, Melittobia spp. 49 — clavicornis; 50 — acasta; 51 — evansi; 52 — scapata; 53
— digitata; 54 — femorata; 55 — chalybii; 56 — australica; 57 — sosui; 58 — sp. nov. Argentina.
316
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 59-69, Female clypeal margins, Melittobia spp. 59 — clavicornis; 60 acasta; 61
scapata; 63 — digitata; 64— femorata; 65 — chalybii; 66 — megachilis; 67 — assemi; 68 — austrahca; 69
nov. Argentina.
DAHMS: REVISION OF MELITTOBIA
317
73 74 75
FIGURES 70-81, Female maxillary palps, Melittobia spp. 70 — davicornis; 11 — acasta; 12 — evansi; 73 —
scapata; 74 digitata; 75 — femorata; 76 — chalybii; 11 — megachilis; 78 — australica; 79 — assemi; 80 —
sosui; 81 — sp. nov. Argentina.
318
MEMOIRS OF THE QUEENSLAND MUSEUM
92
FIGURES 82-92- Female scapes and antennae, Melittobia spp. 82 — clavicornis; 83 scape, 84 acasta,
85 - evansi; 86 - evansi scape; 87 - scapata; 88 - digitata; 89 - digitata, scape; 90 - femorata; 91 -
femorata, scape; 92 — chalybii.
DAHMS: REVISION OF MELITTOBIA
319
FIGURES 93-101 , Female scapes and antennae, Melittobia spp. 93 — megachilis; 94 — Kauai, scape; 95 — Kauai;
96 — australica, scape; 97 — australica; 98 — sp. nov. Argentina, scape; 99 — sp. nov. Argentina; 100 — sosui,
scape; 101 — sosui.
320
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 102-114, Female antennae, terminal segment, Melittobia spp. 102 clavicornis;^
evansi; 105 — scapata; 106 — digitata; 10? - femorata; 108 — chalybii; 109 — megachilis; 110
111 — Kauai; 112 — australica; 113 — sosui; 114 — sp. nov. Argentina.
acasta; 104 —
— hawaiiensis;
DAHMS: REVISION OF MELITTOBJA
321
FIGURES 115-119, Female fore wings, Melittobia spp. 115 — clavicornis; 116 — acasta; 117 — evansi; 118
scapata; 119 — digitata.
322
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 120-126, Female fore wings, Melittobia spp. 120 —femorata; 121 — chalybii; 122 — megachilis; 123 —
hawaiiensis; 124 — australica; 125 — assemi; 126 — sp. nov. Argentina.
DAHMS: REVISION OF MELITTOBIA
323
FIGURES 127-139, Female stigmal veins, Melittobia spp. 127 — clavicornis; 128 — acasta; 129 — evansi; 130 —
scapata; 131 — digitata; 132 — femorata; 133 — chalybii; 134 — megachilis; 135 — australica (paralectotype);
136 — hawaiiensis; 137 — assemi; 138 — sosui; 139 — sp. nov. Argentina.
324
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 140-150, Female sculpture, mid lobe of mesoscutum, mid lobe of scutellum, Melittobia spp. 140
clavicornis; 141 — acasta; 142 — evansi; 143 — scapata; 144 — digitata; 145 —femorata; 146 — chalybii; 147
megachilis; 148 — australica; 149 — assemi; 150 — sp. nov. Argentina.
DAHMS: REVISION OF MELITTOBIA
325
FIGURES 151-155, Male heads, Melittobia spp. 151 — clavicornis; 152 — acasta; 153 — evansi; 154 ~ scapata;
155 — digit ata.
326
MEMOIRS OF THE QUEENSLAND MUSEUM
160
FIGURES 156-160, Male heads, Melittobia spp. 156 —femorata; 157 — chalbyii; 158 — australica; 159 — assemi;
160 — sp. nov. Argentina.
DAHMS; REVISION OF MELITTOBIA
327
FIGURES 161-170, Male mandibles, Melittobia spp. 161 — clavicornis; 162 — acasta; 163 — evansi; 164 —
scapata; 165 — digitata; 166 — femorata; 167 — chalybii; 168 — hawaiiensis; 169 — australica; 170 — Kauai.
328
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 171-173, Male mandibles, Melittobia spp. 171 — assemi; \12 — sosui; 173 _
FIGURES 174-184, Left male palps, Melittobia spp. 174 — clavicornis; 175 — acasta; lib
scapata; 178 - digitata; 179 -femorata; 180 - chalybii; 181 - australica; 182 - a55e/w/ (reverse); 183 -
184 — sp. nov. Argentina.
DAHMS: REVISION OF MELITTOBIA
329
FIGURES 185-193, Male antennae, Melittobia spp. 185 — clavicornis ventral scape; 186 — clavicornis dorsal
scape; 187 — clavicornis pedicel and flagellum; 188 — acasta ventral scape; 189 — acasta dorsal scape; 190 —
acasta pedicel and flagellum; 191 — evansi ventral scape; 192 — evansi dorsal scape; 193 — evansi pedicel and
flagellum.
330
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 194-202, Male antennae, Melittobia spp. 194 — scapata ventral scape; 195 — •scapata ^
— scapata pedicel and nagellum; 197 — digitata ventral scape; 198 — digitata dorsal scape; 199 digitata ped
and nagellum; 200 — femorata ventral scape; 201 — femorata dorsal scape; 202 — femorata pedicel and
flagellum.
DAHMS: REVISION OF MELITTOBIA
331
FIGURES 203-210, Male antennae, Melittobia spp. 203 — chalybii ventral scape; 204 — chalybii dorsal scape; 205
— chalybii scape and flagellum; 206 — australica ventral scape; 207 — australica dorsal scape; 208 — australica
pedicel and flagellum; 209 — hawaiiensis ventral scape; 210 — Kauai ventral scape.
203
204
205
332
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 211-216, Male antennae, Melittobia spp. 211 — assemi
assemi pedicel and flagellum; 214 — sosui ventral scape; 215
flagellum.
sosui dorsal scape; 216 — sosui pedicel
DAHMS: REVISION OF MELITTOBIA
333
FIGURES 217-219, Male antennae, Melittobia spp. 217 — sp. nov. Argentina ventral scape; 218 — sp. nov.
Argentina dorsal scape; 219 — sp. nov. Argentina pedicel and flagellum.
FIGURES 220-224, Male mid legs Melittobia spp. 220 — clavicornis; 221 — acasta; 222 — scapata; 223 — evansi;
224 — digitata.
334
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 225-230, Male mid legs, Melittobia spp. 225 — femorata; 226 — chalybii; 227 — australica; 228
assemi; 229 — sp. nov. Argentina; 230 — sosui.
DAHMS: REVISION OF MELITTOBIA
335
FIGURES 231-235, Male fore wings, Melittobia spp. 231 — clavicornis; 232 — acasta; 233 — evansi; 234 —
scapata; 235 — digitata.
336
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 236-241, Male fore wings, Melittobia spp. 236 — femorata; 237 chalybii; 238 australica, 239
sosui large form; 240 — sosui small form; 241 sp. nov. Argentina.
Mem. QdMus. 21(2): 337—60. [1984]
A REVIEW OF THE BIOLOGY OF SPECIES IN THE THE GENUS MELITTOBIA
(HYMENOPTERA : EULOPHIDAE) WITH INTERPRETATIONS AND ADDITIONS
USING OBSERVATIONS ON MELITTOBIA AUSTRALICA.
Edward C. Dahms
Queensland Museum
ABSTRACT
This paper reviews published accounts of Melittobia biology and contains observations on
the single Australian species M. australica. The species was found to be a highly polyphagous
primary ectoparasite attacking the immature stages of nesting Hymenoptera, both solitary and
social. It is also hyperparasitic on the immature primary parasites within the primary host’s
nests, and are reported parasitising the immature stages of hosts belonging to a variety of
orders other than Hymenoptera. Females gain access to the host by entering the host cell before
it is sealed (delaying oviposition until the host has reached a suitable developmental stage),
excavation through the cell wall and enveloping membranes or by oviposition directly through
enveloping membranes. Any one species can show this range of behaviour and the female’s
nutritional condition is important as a factor in deciding which occurs. Females puncture the
host with their ovipositors to feed and to subdue an active host but not for oviposition. Males
do not feed and are highly aggressive although male aggression seems to vary both within and
between species. Courtship is extremely complex and the three basic patterns reported in the
literature are summarised and some morphological features are correlated with them.
Reproduction is also complex and the importance of parthenogenesis, sib-mating (including
mother-son matings), multiple settling by females and sex ratios shifts is discussed. A
nutritionally induced polymorphism occurs, as well as sexual dimorphism, and varies with
species. The result is type-form and second-form individuals of both sexes which differ
morphologically and physiologically. Second-form specimens of both sexes of an acasta group
species are described and compared with second-form specimens of both sexes of M.
australica. Dispersal is by flight and evidence suggests that it is wind assisted. The capability of
Melittobia to use man’s transport for dispersal is also discussed. A brief account of the life
cycle of M. australica is included and compared with published accounts of other species.
INTRODUCTION
Species in the genus Melittobia are very
efficient organisms. In all stages of their
development they show remarkable plasticity of
behaviour and adaptability to prevailing
conditions. Theoretically, uninseminated females
can survive and eventually produce progeny of
both sexes even in the absence of preferred hosts.
They make very good laboratory animals and
their plasticity coupled with arrhenotokous
parthenogenesis make them ideal subjects for
laboratory investigations into the genetics of
speciation and evolution.
Some of the reports on the biology of
Melittobia species in the literature proved either
confusing or inconsistent. Detailed study of the
biology of Melittobia australica allowed many of
the confusing and inconsistent aspects to be
clarified. The following account is therefore a
blend of previously published accounts on several
species and my recent observations on the single
Australian species. The outcome of this review
has been of great assistance in understanding
phylogeny in the genus and therefore of great
assistance in making taxonomic decisions for my
revision of the genus (Dahms 1983a)
MATERIALS AND METHODS
Cultures of Melittobia australica were
maintained in excavated blocks with glass covers
in the laboratory without controlled temperature
or humidity. Behavioural observations were made
with a Leitz TS stereomicroscope with fibre-
optics, cold-light illumination. Hosts used for
culture of M. australica were Pison spp., and
Sceliphron spp. Larvae of the ant genus
338
MEMOIRS OF THE QUEENSLAND MUSEUM
Camponotus were tried as hosts but proved
unsuitable. Although larvae of Apis meiiifera
proved suitable hosts for cultures they suffered
high mortality due to mechanical damage during
extraction and from not being in a controlled
environment.
Investigations into the life history of M.
australica were carried out using Sceliphron
formosum prepupae, a supply of which was
maintained in the refrigerator without
deterioration. In this case the trials were carried
out in plastic-stoppered glass vials. A constant
temperature room was not available and the
colonies were kept in a room with an air
conditioner. Under these circumstances the
temperatures recorded were 25 °C (± 5°) and
humidity (± 5^o).
All figures were drawn from cleared
microscope slide-mounted specimens and each
has the scale indicated. They were drawn with a
camera lucida fitted to a Wild M20 compound
microscope.
BIOLOGY
Hosts
To say that species of Melittobia are not host
specific is a gross understatement. Waterston’s
1917 view of M. acasta that it is remarkably
polyphagous attacking everything within its
limited range of action is more realistic.
In the main, Melittobia are primary parasites
within the nests of wasps and bees, both solitary
and social. Amongst the social species are:
Vespula acadica (Salden) (H.C. Reed, USA, pers.
comm. 1978); Vespula germanica (JEdbucms) ^.n(^
Bombus sp. (R. Macfarlane, New Zealand, pers.
comm. 1980); Polistes exclamans Viereck (H.C.
Reed, USA, pers. comm. 1977); Bombus
pennsylvanicus (DeGeer) (A.C. Haman, USA,
pers. comm. 1977) and Apis meiiifera Linnaeus
(E.H. Erickson, USA, pers. comm. 1978). The
last mentioned of course has serious economic
implications although van den Assem (pers.
comm. 1981) considers that sperm inside the
spermathecae of female Melittobia do not survive
at the relatively high temperatures found inside
the hive oi A. meiiifera. Melittobia species have
reached economic pest status wherever the Alfalfa
Leaf-cutter Bee (Megachile rotundata (Fabricius)
is cultured (Prof. Thorp, University of California
Davis pers. comm. 1981). Four species, M. acasta
(Walker 1839), M. chalybii Ashmead, 1892, M.
japonica Masi 1966 (= M. clavicornis (Cameron
1908)) and M. megachilis (Packard 1864) have
been recorded in the literature as being
hyperparasitic within nests of Hymenoptera and
in the present study M. australica Girault, 1912
was found to be hyperparasitic also.
There are published records of Melittobia
naturally parasitising hosts belonging to orders
other than Hymenoptera. Rau (1940) reports
breeding M. chalybii from the ootheca of the
cockroach Periplaneta americana (Linnaeus).
Howard and Fiske (1911), and Graham-Smith
(1916, 1919) have bred M. acasta from dipteran
puparia. Swezey (1909) discovered M. hawaiiensis
Perkins, 1907 breeding on the larvae of the bud-
moth Ereunetis flavistriata Wilson. Howard
(1892) reported a species from dipteran puparia
within the cells of a mud-dauber’s nest and M.
japonica (= M. clavicornis ) is noted as utilising
similar dipteran hosts by Iwata and Tachikawa
(1966). In the present study M. australica was
bred from dipteran puparia within Sceliphron
spp. nests.
Laboratory trials by various workers have
shown a remarkable range of hosts that
Melittobia will utilise under these conditions.
Balfour-Browne (1922) and Thompson and
Parker (1927) found M. acasta to be highly
polyphagous in the laboratory, even attacking
spiders and lepidopteran larvae taken from mud
nests. However, the progeny failed to mature and
Balfour-Browne felt this may have been due to
desiccation of the hosts. These papers contain a
very large number of species successfully
parasitised from the insect orders Coleoptera,
Hymenoptera, Lepidoptera and also from the
arachnid order Araneae. Peck (1963) and Burks
(1979) provide comprehensive host lists for North
American species; Domenichini (1966) has host
lists for M. acasta and M. Japonica; (= M.
clavicornis ) and Thompson (1955) listed hosts of
M. acasta, M. hawaiiensis and an unidentified
species using Commonwealth Agricultural
Bureau records. These lists are extremely long and
it is not practical to duplicate them here.
Not all species of Hymenoptera, however, are
successful hosts for Melittobia. Balfour-Browne
(1922) found that Osmia rufa (Linnaeus) was
rarely attacked in the wild. In the laboratory
naked larvae and pupae of O. rufa were readily
accepted by M. acasta females which fed and laid
eggs. The eggs often failed to hatch and, if they
did hatch, the resulting larvae failed to reach
maturity. If he placed M. acasta females in a cell
with larval O. rufa just before the cocoon was
spun the M. acasta females often became
entangled in the outer layers of the cocoon. This
did not happen under similar circumstances with
DAHMS: BIOLOGY OF MELITTOBIA
339
other hosts. Where A/, acasta females were
presented with O. rufa pupae inside cocoons they
were not attacked and Balfour-Browne suggested
that this was due to the toughness of the cocoon.
Malyshev (1911) had earlier suggested that some
species may escape attack by Melittobia as the
result of a mechanical barrier related to the type
of nesting material used, e.g. those species which
use resin for nest construction.
Jayasingh and Freeman (1980) also draw
attention to the importance of nest material in
host succeptibility to attack by Melittobia. They
found the resinous nests of Chalicodoma
rufipennis (Fabricius) to be a total barrier to
Melittobia. They also found that another factor
was direct attack by the mother on Melittobia e.g.
females of Pachodynerus nasidens (Latreille)
were observed to crush Melittobia females in their
mandibles. I have observed parasitic mites which
can normally be found on Secliphron spp. larvae
acting in competition with M. australica larvae
and in one case the mite larvae were feeding upon
the M. australica larvae. From these observations
it is clear that Melittobia do not have it all their
own way.
There are a few indications in the literature that
Melittobia spp. may be endoparasitic. Girault
(1912), at the end of his descripton of M.
australica quotes the collector, ‘Mr. Tryon
informs me that the parasites emerged from the
host in its cocoon but not until after it had
transformed into the adult, the latter died. A
number of parasitic larvae make their way out of
each Pison and pupate nakedly’. This record I
regard as an error based upon an assumption. In
all cases I have observed, M. australica is very
definitely ectoparasitic. Perhaps Mr. Tryon on
opening the host cocoon saw prepupal M.
australica larvae with meconium and assumed
that the larvae had emerged from within the host.
Malyshev (1911) states that under certain
circumstances M. acasta is endoparasitic e.g.
when the female oviposits through the cocoon of
hymenopteran hosts or through the puparial wall
of a dipteran host. Thompson and Parker (1927)
found that M. acasta would not oviposit in fresh
puparia of Sarcophaga sp. Oviposition occurs
only after the body of the fly has separated from
the wall of the puparium creating air spaces. Eggs
are placed directly onto the surface of the pupa
within. They found the same with some living but
slightly desiccated pupae of the ant genus
Camponotus. Air spaces had developed beneath
the cuticle resembling the situation with dipteran
puparia. Maeta and Yamane (1974) reported that
one method of oviposition used by M. japonica
(identification corrected to M. acasta by Maeta
(1978)) was to oviposit through the wall of
cocoons of species belonging to the
hymenopteran genera Osmia, Monodontomerus,
Nematopoideus, Trypoxylon and Chalicodoma.
In all of these situations insertion of the
ovipositor through enveloping membranes
implies endoparisitism, but the eggs are placed on
the surface of the body of the host which means
they are in fact ectoparasitic.
In the present investigation M. australica was
bred from the following hosts:-
1) Pison aureosericeum Rohwer
2) Pison spp.
3) Sceliphron laetum Smith
4) Sceliphron formosum Smith
5) Megachile sp.
6) Stenarella victoriae Cameron
7) Dipteran puparia in Sceliphron spp. nests
8) Camponotus sp.
9) Apis meilifera Linnaeus
10) Anthrax angularis Thomson
The host given by Girault for the type
specimens of M. australica was Pison spinolae
Schuckard. In the above list, 1-6 were naturally
infested. Stenarella victoriae is an ichneumonid
parasite on Sceliphron spp. The dipteran puparia
in Sceliphron nests are thought to be parasites on
the provisioned spiders since they are always
found in cells fully stocked with dry spiders and
without a Sceliphron larva. Hosts 8-10 were
presented in the laboratory. Larvae of the ant
genus Camponotus were tried as substitute hosts
for laboratory work. Although the M. australica
progeny developed through to maturity the
resulting adults were small and lacked vigor.
Honey bee {Apis meilifera ) larvae were also tried
as alternative hosts. They were readily accepted
and produced vigorous parasite adults, but
proved difficult to extract from the comb without
a high percentage of deaths. Anthrax angularis
was found as a parasite in Sceliphron nests. Two
larvae were presented to fertilised M. australica
females and were readily accepted. The resulting
progeny were of normal size and vigour. Since
Anthrax angularis is a natural parasite of
Sceliphron spp. it is fairly safe to assume that it
would be naturally attacked by M. australica.
Access to the Host
In the literature, workers have put forward
several behavioural patterns associated with
gaining access to the host as follows:-
340
MEMOIRS OF THE QUEENSLAND MUSEUM
1) excavation into the host cell and cocoon
2) entrance into a host cell before closure
3) oviposition through enveloping membranes
1) Excavation
Melittobia females have well developed,
tridentate mandibles and there are several records
in the literature which indicate that they have well
developed excavatory powers.
Howard and Fiske (191 1) stated that female M.
acasta in search of a host (sarcophagid puparia in
this case) entered damp soil for a distance of
several inches. Graham-Smith (1916) however,
suggested that the fly puparia buried in the soil
were possibly connected to the surface by minute
passages sufficiently large to admit Melittobia,
There may in fact be minute passages left as the
sarcophagid larvae dig into the soil and this may
not be a case of true excavation by Melittobia.
More direct evidence was provided earlier by
Howard (1892) quoting observations by Giraud.
The latter noted that a M. acasta female after
walking around on the intact cell of the bee
Chalicodoma sp., stopped and gnawed the
membrane until a perforation was made through
which she entered the cell. Malyshev (1966)
observed that a M. acasta female in a host nest
moved from one cell through the cell wall into the
next cell and through the cocoon to gain access to
another host. Graham-Smith (1916) stated that
females of M. acasta emerged from intact fly
puparia through a small hole which one of them
excavated. He also noticed that females of M.
acasta confined in glass lubes with cork stoppers
immediately began to excavate a tunnel in the
cork stopper. Similarly, Buckell (1928) found
females of M. chalybii (= digitata Dahms 1983a)
excavated their way out of glass vials through a 25
mm cork stopper. Torchio (1963) recorded
excavation holes in the cell partitions of
Megachile rotundata (Fabricius) made by M.
chalybii (which I suspect was M. acasta ). Cowley
(1961) mentioned that M. clavicornis (= M.
hawaiiensis Perkins) will excavate a hole in
cocoon walls to gain access to Pison spinolae
pupae within. Iwata and Tachikawa (1966)
observed 1-5 excavation holes each of 0.5 mm
made by M. japonica ( = A/, clavicornis ) females
in a series of mud cells of Auplopus sp. They also
found several incomplete holes whose bottoms
were obstructed by sand grains and from this
postulated that the excavations were from the
outside in. Observations by Maeta and Yamane
(1974) on M. japonica (= A/, acasta see Maeta
(1978)) led them to conclude that female
Melittobia have the capacity to excavate holes in
plugs or partitions either of leaf fragments or
mud, even if they were fairly thickly constructed.
In this investigation, inseminated female M.
australica were presented with sealed nests of
Pison sp. and Sceliphron spp. After presentation
of the Pison nests, M. australica females were
noted excavating the mud walls. Only one hole
was constructed in each cel! and several females
were observed working at each site. Only one
female worked at any one time at the one site with
the others taking turns. Graham-Smith (1916)
mentioned that M. acasta females produce one
exit hole in each puparium, rarely two. Also when
confined in glass vials he found that only one
excavation tunnel was made in each cork stopper
and that females worked singly at the excavation.
For practical reasons one would expect that the
economy in number of holes excavated per cell
would be fairly general in the genus, although
Iwata and Tachikawa (1966) observed 1-5 per
host cell for M. japonica ( = Af, clavicornis ) as
mentioned above.
As soon as the hole in a Pison cell was large
enough, the female M, australica passed through.
The next day when the cells were broken open the
host cocoon was seen to have a single excavated
hole and the parasite females were inside on the
body of the host. In the case of the Sceliphron
spp. nests, excavation was not directly observed,
but 24 hours after exposure to inseminated M.
australica females there were no parasites to be
seen. Examination of the host cell walls showed a
single excavation in each and on breaking open
the cells, I found that the parasite females had
penetrated the cocoons to reach the host within by
a further single excavation. Under these
conditions more than one female had entered
each cell. Similarly inseminated M. australica
females gained access to Megachile sp. larvae
within a sealed leaf nest lying uncovered on a
bench about 2 metres from the release site.
In one instance where plastic stoppered tubes
were used for cultures of M. australica, I found
that adult females were capable of escaping by
excavating their way through three sealing flanges
on the inserted part of the cap and the rim of the
cap where it fitted against the top of the glass
tube. They did this in each of the 10 tubes being
used.
If presented with Sceliphron cocoons outside
their mud cells, inseminated M. australica females
gnawed a hole in the cocoons and oviposition
followed feeding. If naked Sceliphron prepupae
and pupae were presented, oviposition followed
feeding without delay. Therefore, as Thompson
DAHMS: BIOLOGY OF MELITTOBIA
341
and Parker (1927) found with M. acasta, the
presence and penetration of enveloping
membranes are not necessary prerequisites for
oviposition in M. australica.
2) Entrance before the host cell is closed
Several workers have shown that Melittobia
enter unsealed nests of their hosts and are able to
delay oviposition until the host is at a suitable
developmental stage.
Schmieder (1933), working with M. chalybii
reported that female parasites gained access to the
larvae of bees and wasps by entering host cells
before they were completed. The only evidence to
support this in his paper is the fact that
examination of a Trypoxylon sp. cocoon did not
reveal whether or not it contained Melittobia. He
took this to indicate that the parasite gained
access to the host before the cocoon was spun and
became enclosed with the host. However, it does
not necessarily mean that M. chalybii females
entered before the nest was closed since they
could just as easily have excavated their way into
a sealed host cell before the cocoon was spun and
then become enclosed with the host. Therefore,
this is not conclusive evidence.
Balfour-Browne (1922) noticed M. acasta
females becoming sealed up in cells being
constructed by bees and wasps in elder stems and
glass tubes he had provided in his garden. From
observation of those in the glass tubes he
discovered that their being sealed in the host nests
was not accidental. He found that females can
delay oviposition for up to 60 days when placed in
a cell with an unhatched host egg. The parasite
commenced oviposition only when the host
reached full-grown larval condition. Feeding by
the parasite on the developing host appeared not
to affect the latter’s development and he had
many examples of eggs being pierced by the
female’s ovipositor for food without affecting
development of the host. During his trials he
placed up to 15 M. acasta females in a cell with a
newly hatched Osmia sp. larva and allowed them
to feed freely on the host for 14 days without
apparently affecting the host which completed its
development. When he placed M. acasta females
with older larvae there were no ill effects on the
host as long as the parasites were only feeding. He
felt quite satisfied that feeding by M. acasta
females was not necessarily injurious to the host.
Malyshev (1966) also mentioned M. acasta
females entering host cells before they were
closed.
Maeta and Yamane (1974) found that, in most
Trypoxylon sp. cells infested with M. japonica ( =
M. acasta see Maeta (1978)), the closing plugs did
not show entrance holes. They concluded that the
parasite had gained access to the host cell before it
was sealed. When discussing oviposition, they
mentioned the capacity of M. japonica (= M.
acasta see Maeta (1978)) females to delay feeding
until the host reached a suitable stage for
parasitism; in fact they kept females of this
species alive for more than 2 months without
food.
In this investigation, M. australica females
were not directly observed entering host cells
before they were closed, but there is indirect
evidence that this may occur. On numerous
occasions M. australica females were kept for
periods up to 3 weeks without food. At the end of
this period, when a suitable host was provided,
they fed and subsequently laid fertile eggs. Thus
they can survive long periods without ovigenesis
being adversely affected. Females accidentally
released in the laboratory were later found
residing in empty host cells of old Sceliphron
formosum nests lying on the laboratory bench.
When these cells were broken open the parasites
showed the usual negative reaction to light which
is displayed in the presence of a host. Feeding by
M. australica females does not affect
development of the host e.g. when inseminated
females were allowed to feed on prepupal
Sceliphron formosum larvae for a few days then
removed the host successfully passed to pupa! and
adult stages. Several Sceliphron formosum early
pupae were supplied each to 10 inseminated M.
australica females and all hosts continued to
develop to full adult colouration in spite of
feeding by the parasites and their progeny. Death
of the host pupae resulted ultimately due to
feeding pressure of the parasites. Therefore, M.
australica females will enter empty host cells, can
delay feeding and oviposition for long periods,
and are able to feed on the host larva or pupa
without affecting its development.
This capability with its attendant behaviour
patterns probably occurs in all species.
3) Oviposition directly through enveloping
membranes
As mentioned before, Thompson and Parker
(1927) found that M. acasta oviposits directly
through the puparial wall of Diptera and that this
takes place only after the fly pupa has separated
from the puparial wall. Malyshev (1966) also
mentions this. In all cases where I have reared M.
australica from fly puparia there were no
excavation holes in the puparial walls until
emergence of the parasite, these being the exit
342
MEMOIRS OF THE QUEENSLAND MUSEUM
holes of the progeny. Maeta and Yamane (1974)
stated that M. japonica (= acasta see Maeta
(1978)) oviposited directly through the cocoon of
species of the hymenopteran genera Osmia,
MonodontomeruSy Trypoxylon and
Chalicodoma. However, there appears to be some
versatility of behaviour here since they also found
that in some cases the parasites entered the host
cocoons of Trypoxylon and Chalicodoma before
oviposition.
Malyshev (1966) provided an explanation.
When a M. acasta female’s work was finished in
one cell of a host’s nest she made her way into the
next cell with her jaws. If the host cocoon was in
close contact with the cell partition the parasite
gnawed through both the partition and the
cocoon. However, if the cocoon was not in
contact with the ceil partition and the body of the
host was some distance from the cocoon wall the
parasite gnawed through the cocoon. Where the
cocoon was close fitting he found that the parasite
oviposited directly through the cocoon wall. Thus
closeness of fit of the cocoon to the host appears
to be important, i.e. it is necessary for the tip of
the ovipositor to reach the host within and this is
substantiated by my observations on M.
australica outlined below.
For hosts with spacious cocoons this behaviour
would not be possible, e.g., it is difficult to
imagine A/, australica ovipositing through the
cocoon walls of Sceliphron spp. In all cases,
whether the M. australica females had fed or not,
when Sceliphron spp. cocoons were provided they
were always entered before oviposition.
Although the hosts mentioned above by Maeta
and Yamane (1974) are all small with close fitting
cocoons, some were found to be entered also and
I feel the nutritional condition of the female is
important in these cases as well as the closeness of
the cocoon to the cell partition. When I presented
inseminated unfed females of M. australica with
Pison sp. cocoons, which are close fitting, each
was entered by the parasite. On one occasion on
breaking open a Pison sp. cell collected from the
wild I found two M. australica females with
distended metasomas on the cocoon surface.
They were observed to insert their ovipositors
through the cocoon wall. The point of insertion
was always on the side of the cocoon about 1/2 to
2/3 the way down the wall. The ovipositor was
fully inserted followed by a pause of about 3-5
seconds, half withdrawn, reinserted followed by a
pause of about 3-5 seconds then fully withdrawn.
On one occasion, a female inserted her ovipositor
at the upper, anterior end of the cocoon and was
noticed to indulge in partial withdrawals and re-
angling the direction of the ovipositor. No
pausing occurred and the ovipositor was
eventually withdrawn. This end of the cocoon
housed the narrow, anterior end of the prepupal
larva which from the upper surface of the cocoon
was not accessible to the ovipositor of the female.
No attempt was made by the females to enter the
cocoon. On breaking open the cocoon about 15
eggs were visible on the lateral portions of the
prepupal Pison larva — none on the anterior
portion. It appears therefore that contact of the
ovipositor with the host within is necessary before
oviposition occurs and that oviposition through
enveloping membranes occurs with close fitting
cocoons where the parasite female has previously
fed. No published records are available on
penetration of fly puparia by the female parasite.
1 have tried M. australica on blow fly puparia but
without success. Tachinid or sarcophagid puparia
were not available. In the case of puparia, the
parasite female may feed on the early pupa before
it separates from the puparial wall or the pupa
within may be close enough to the puparial wall in
some areas to allow some body fluids to well out
of a puncture site, e.g., Graham-Smith (1919)
mentioned that fertilised or unfertilised females
of M. acasta confined with fly puparia lived for
long periods (up to 36 days) and seemed to derive
nourishment from fluid exuding from the puparia
at ovipositor puncture sites. Van den Assem
(pers, comm. 1981) has confirmed this behaviour
in all Melittobia species in his cultures. However,
in some cases, the parasites gnawed their way into
fly puparia. He found that in crossing
experiments involving the assemi group, females
gnawed holes in fly puparia and walked on the
surface of the pupa within. Van den Assem (1976)
found that virgin M. acasta females gnawed their
way into fly puparia containing males of this
species and mated with them.
Migration from one cell to another appears to
be nutritionally governed as well. The relatively
large eggs (0.3 mm long; females 1. 1-1.5 mm
long) mean that a female cannot produce her
entire egg batch in 1 or 2 days, Oviposition and
feeding were observed to be progressive
throughout the life of female M. australica. It is
reasonable to assume, therefore, that competition
for food with her progeny may be an important
factor in governing the number of eggs per host.
On a relatively large host e.g. Sceliphron spp.
there is probably enough food to support the
larvae and the mother for the length of her life.
On smaller hosts e.g. TrypoxyloUy Osmia, Pison
DAHMS: BIOLOGY OF MELITTOBIA
343
etc. competition for food with her progeny would
necessitate her migration from one cell to
another. In this situation, if she has sufficient
food for maturation of eggs she may oviposit
directly through the enveloping membrane of the
host in the next cell. What determines the number
of progeny in this case is not known. Perhaps as
she approaches the time for nutritional
replenishment she might again migrate then
penetrate the next cocoon. The whole process of
oviposition and nutritional requirements is one
deserving close study.
In summary, oviposition behaviour of
inseminated Melittobia females is very flexible
and is dependent upon a number of conditions. If
the female parasite encounters a host cell before it
is closed, she enters and feeds upon the
developing host without affecting its
development. She can delay oviposition until the
host is at a suitable stage, i.e., the prepupal larva
or pupa. If the host is large with a spacious
cocoon she can become incorporated within
during construction or gnaw in afterwards. She
stays with this one host all her life and is assisted
in its utilisation by specialised second-form
progeny (discussed later). Where the host is small
with a close fitting cocoon she can either become
incorporated or oviposit through the cocoon wall.
Because of the limited food supply on a small host
she must seek another to attain her full egg laying
potential and moves to another cell. If the cocoon
is touching the cell partition and/or she requires
additional food she gnaws through the cocoon
wall. However, if the cocoon is not in contact
with the cell partition and she does not require
more food she can continue ovipositing through
the enveloping membrane. Where the host cell is
sealed she gnaws through the cell wall. If the host
has not spun a cocoon she can follow the
behaviour patterns above depending upon the size
of the host and the closeness of fit of the cocoon.
Should she enter a cell and encounter a cocoon,
no matter how close fitting she gnaws through it
to feed upon the host.
FUNCTIONS OF THE OVIPOSITOR
Female Melittobia use their ovipositors for
feeding, to paralyse the host and for egg laying.
1) Feeding
When inseminated M. australica females were
presented with quiescent larvae or pupae, I
noticed the ovipositor was fully inserted and
within a few seconds, withdrawn. The females
moved back and fed on the drop of body fluid
which issued from the host. Old wounds, visible
as dark brown spots, were frequently revisited by
the females who fed on the congealed body fluids
of the host. There appeared to be no favoured
spot for puncture of the host’s body and on one
occasion a female punctured the head capsule of a
host larva. Torchio (1963) observed M. chalybii
(which I feel was probably M. acasta ) feeding on
congealed host body fluids at old puncture sites.
Balfour-Browne (1922) observed this behaviour
in M. acasta and even the eggs of the hosts were
used as a food source. Malyshev (1966) also
mentioned the habit of M. acasta females feeding
on the body fluids of the host oozing from
ovipositor penetration points. Schmieder (1933)
mentioned this feeding behaviour in M. chalybii.
Maeta and Yamane (1974) noted dark brown
spots on the body of the host and assumed these
to be the feeding spots of M. japonica (= M.
acasta see Maeta 1978) females although they did
not directly observe this feeding. It was recorded
also for M. japonica (— M. clavicornis ) by Iwata
and Tachikawa (1966).
This behaviour is no doubt a general one for all
species of Melittobia and feeding upon the host
by the female is recorded amongst other parasitic
Hymenoptera. In the case of Melittobia it can
occur without death of the host and this, together
with the female’s ability to delay oviposition for
long periods is a decided advantage when a host in
an early stage of development is encountered.
Doutt (1959) in his review of the biology of
parasitic Hymenoptera mentioned this feeding
behaviour and that it is well established that
feeding on the host body fluids is necessary to
obtain protein for ovigenesis. In support he
mentioned the work of Flanders (1942, 1953) on
Metaphycus helvolus (Compere). Over a 3 week
period at 80° F and away from its host, ovigenesis
ceased in this species. When presented with a host
at the end of this period the parasite fed without
delay and oviposition began a few days later.
In this investigation, newly emerged,
inseminated M. australica females when deprived
of a host remained as they emerged, i.e., without
distended metasomas. When presented with a
host pupa after 7 days all females immediately
inserted their ovipositors and fed at the puncture
sites. Within 24 hours their metasomas were
distended and well developed eggs were clearly
visible through the intersegmental membranes of
the metasoma. They began laying eggs 2-3 days
after feeding. It would appear therefore that
feeding upon the host is essential for egg
maturation in Melittobia.
344
MEMOIRS OF THE QUEENSLAND MUSEUM
2) Preparation of the host
Buckell (1928) presented M. chalybii (= M.
digitata ) with active host larvae which became
quiescent after 24 hours. He postulated that some
paralysing fluid was injected. Balfour-Browne
(1922) found that once an A/, acasta female had
oviposited on a host the latter was doomed even
though the eggs were removed before hatching
and the adult females removed as well. We have
seen before that he found feeding by the adult
females did not affect host development. He
described a fluid oscillating in the ovipositor as it
was being inserted. I have noticed movement in
the ovipositor of M. australica but consider it
more likely to be rotation of the valves of the
ovipositor as the female works at insertion. A
similar movement was seen during insertion of the
ovipositor of M. australica for feeding. Balfour-
Browne also noticed that the ovipositor was held
fully inserted for a period before withdrawal and
that the females did not feed at these sites.
When active last instar larvae of Anthrax
angutaris and Sceliphron spp. were presented to
inseminated M australica females they became
very agitated and continually performed twisting
and rolling movements. The parasites inserted
their ovipositors in spite of the activity and within
24 hours the host larvae were quiescent. In these
cases the ovipositor was inserted and held in
position for some time before extraction. After
withdrawal of the ovipositor the females moved
away without attempting to feed at these sites.
Once the host larvae were quiescent, the M.
australica females were noted to insert their
ovipositors and feed at the puncture sites after
withdrawal.
The minute size of Melittobia relative to its
hosts makes suppression of an active host seem an
impossible task. Beard (1952) working with
Habrobracon hebetor (Day) found that one part
of the venom of this species to 200,000,000 parts
of the host’s blood was sufficient to cause
permanent paralysis. If the levels of potency are
similar in Melittobia, the task of subduing an
active host would not be impossible.
Given the capacity of Melittobia to delay
oviposition and its ability to feed on the host
without affecting the latter’s development one
wonders whether paralysis of the host is necessary
under natural conditions. In all cases where host
cells have been broken open and M. australica
found, the host has been able to produce a cocoon
and in some cases development had reached the
pupal stage and attained adult colouration before
death. Malyshev (1966) suggested that the
stinging by M. acasta was for preservation of the
host and it may be that under certain
circumstances the injection of venom prevents
further development of the host. This is an aspect
that requires further investigation.
4) Oviposition
Last but not least, the ovipositor is used for egg
deposition. The ectoparasitic status of the genus
has already been discussed. Oviposition
therefore, does not involve insertion of the
ovipositor into the host. In M. australica the tip
of the ovipositor was braced against the surface
of the host and the metasoma raised releasing the
inner ovipositor valves which therefore became
arranged at right angles to the metasoma. The
relatively large egg appeared to flow down the
ovipositor valves onto the host. No particular site
on the host appeared to be favoured, but the eggs
tended to be deposited in clusters. The surfaces of
the eggs were moist, and this coating kept them
attached to the host and to each other. This
procedure for M, australica appears to be fairly
standard for the genus.
HABITS OF THE MALE
In all species for which the male is known he
has reduced wings, modified antennae and
reduced eyes. His sole function appears to be
reproduction. Important aspects of his behaviour
are feeding, aggression and courtship.
1) Feeding.
Waterston (1917) wrote ... The male is at first
of a transparent yellowish brown colour, the head
sometimes darker but after feeding, the abdomen
may be opaque Other workers (Balfour-
Browne (1922) with M. acasta, Buckell (1928)
with M. chalybii (= M. digitata ), Schmieder
(1933) with M. chalybii and Dahms (1973) with
M. australica ) have not observed males to feed.
In most cases when males emerge the host is fully
utilised leaving only brothers and sisters as
potential food. Male aggression has been
mentioned by different workers and Matthews
(1975) suggested this aggression may be important
for male nutrition as the opponent’s body fluids
could serve as an additional energy source. As
more direct evidence in support he drew attention
to the occasional killing by males of virgin female
M. chalybii {— M. australica ) presented to them
in mating chambers. The male usually tore a hole
in the female’s metasoma and chewed vigorously
on her for several minutes. Graham-Smith (1919)
found that in some battles between male M.
acasta the victor buried his mandibles in the
DAHMS: BIOLOGY OF MELITTOBIA
345
dorsal part of his adversary’s head and continued
to bite for several minutes.
I agree with van den Assem, Gijswijt and
Niibel (1980) who felt that this suggestion is
questionable. Although male aggression has been
reported for several species there appears to be
some variation in whether an opponent is
mutilated or not, i.e., it is apparently not
consistent in the genus. Balfour-Browne (1922)
observed female mutilation by males in M. acasta
but felt this was due to experimental conditions.
He also noted that male to male aggression was
less prominent where the cell was full of emerging
females. In all the years I have been culturing M.
australica {M. chalybii of Matthews (1975)) on
only 2 occasions have I noted male aggression
causing mutilation of other males and on only one
occasion did I observe female mutilation. On
these occasions I did not notice males pausing to
gnaw on a victim.
The suspicion that males do not feed is
substantiated by indirect evidence from my
observations with M. australica. When males of
this species emerge their metasomas are
distended, but become increasingly deflated until
finally they are very flat. Deflation of the
metasoma would result from utilisation of food
reserves for spermatogenesis and courtship. That
this deflation would be dramatic can be seen from
the extremely biased sex ratios recorded in the
literature for several Melittobia species — 1-13%
males. To quantify this — van den Assem,
Gijswijt and Niibel (1980) found that the progeny
from 29 host puparia each with a single M.
japonica, {= M. clavicornis ) female was 1843
individuals of which only 72 were males. The sex
ratio of M. australica I found to be 3-4% males.
If feeding were occurring without being observed
then deflation of the metasoma would not have
occurred or been so marked, Schmieder (1933)
found the males of M. chalybii to be short lived.
He attributed this to rapid depletion of food
reserves resulting from abstinence from food
during constant activity, which agrees with my
assumption. Male aggression and feeding are
topics deserving more detailed investigation.
2) Male aggression.
In the genus, males have not only undergone
radical modification, e.g. head capsule and
antennae, but also have undergone major
reductions in non-required organs, e.g., eyes and
wings. If males do not feed one would expect a
reduction of the mouth parts. However, in all
species, the mandibles of males are larger than
those of females and each has a well
differentiated, sickle-shaped, anterior tooth. That
these mandibles function as weapons in male
aggression is reported in many species. Graham-
Smith (1919) found that M. acasta males were
very aggressive and encounters between males
resulted in the death of one of the opponents.
Only rarely did he find more than one live male in
each host puparium. Balfour-Browne (1922) also
found M. acasta males very aggressive and bouts
between males often resulted in death. However,
he also noted that in a cell full of emerging
females, the males were very busy and paid little
attention to each other. Malyshev (1966) found
Af. acasta males to be very aggressive. Hobbs and
Krunic (1971) found that some male M. chalybii
M. acasta ) fought and died before the first
females emerged. Often all were dead before the
last female emerged. This in addition to the
biased sex ratio often meant that late-developing
females had no males with which to mate. Buckell
(1928) recorded aggression in M. chalybii M.
digitata ) and he found the males to be extremely
pugnacious. They fight until only one is left and,
as Graham-Smith (1919) found with M. acasta,
dead pupae or parts of males were readily
attacked. Schmieder (1933) did not observe such
fierce fighting between males of M. chalybii when
confined with or without females. The males,
when they met, engaged in a brief excited tussle
and then separated. Hermann (1971) did not
observe duels between males of M chalybii ( =
M australica ) confined together in gelatin
capsules. However, she did find that the first male
to emerge touched other male pupae frequently
and that these failed to emerge. This same species
in Kalamazoo (the M. chalybii of Evans and
Matthews (1976)) is very aggressive. When I
visited Dr. Evans in 1974 I observed battles
between these M. australica males which
frequently resulted in mutilation. The other
species kept in culture by Dr. Evans, M. evansi
(Dahms 1983a), according to him was not as
aggressive. Matthews (1975) confirmed that adult
male M. chalybii ( = M. australica ) in his cultures
are highly aggressive and more so than M. evansi.
In the latter case the first male to emerge
systematically decapitates others just prior to
emergence from the pupa or immediately after.
However, when adult male M. evansi met, one
adopted an inert or passive posture and the
aggressor abandoned it without inflicting injury.
In my cultures of M. australica over several
years, encounters between males resulted in a
brief excited tussle with the males rolling about.
After a few seconds the males disengage and go
346
MEMOIRS OF THE QUEENSLAND MUSEUM
their separate ways a little faster than usual. I
have not observed males paying any attention to
male pupae. Males which emerged first walked
over the pupal mass palpating it with their
antennae and paused only at close-to-emergence
females. On two occasions I have noticed male
aggression resulting in mutilation of other males
and occasionally males confined without females
indulged in fatal encounters.
It appears that male aggression is a standard
behaviour pattern in the genus and that some
species are more aggressive than others. It
appears also that male aggression can vary in
intensity within a species. Matthews (1975)
remarks on M. evansi indicate that there may be
some variation in the stage at which other males
are attacked and his description of males
adopting passive postures when encountered by
another male is the first record of this type of
behaviour in the genus. This aspect of male
behaviour would make a very nice study. The
implications of male aggression are discussed
later under ‘Reproduction’.
A peculiar aspect of male aggression is reported
for M. acasta and M. chalybii ( = M, australica ).
Balfour-Browne (1922) found that the killing of
females by males was not uncommon, but he
thought that this was related to experimental
conditions. Hermann (1971) found that males of
M. chalybii ( = M. australica ) eight days or older
when placed with a receptive female would grasp
her and feed on her. After feeding upon her for a
few minutes the males began copulatory
behaviour. Such females generally died during
courtship or before oviposition. Matthews’ (1975)
observation on the same species where males chew
on a females’s metasoma for several minutes has
been mentioned under ‘Feeding’ above. In my
colonies of this species male aggression resulting
in female mutilation was noticed on only one
occasion and several females were affected. I did
not observe males pausing to chew or feed upon
females which they mutilated. Perhaps Balfour-
Browne is correct in assuming male aggression
towards females was due to experimental
conditions. In the wild, fertilised females disperse
fairly soon after mating, but in the laboratory
they are kept crowded and confined for several
days. With increasing numbers of mated females,
presumably with remnants of male odour (see
Dahms 1983b), there is an increase in aggression
some of which may be directed towards females.
Whatever the cause, it appears to be a rare
occurrence and is certainly not what one would
expect.
3) Courtship.
In Melittobia, courtship is a lengthy and
involved process. Detailed accounts of a few
species can be found in Parker and Thompson
(1928), Hermann (1971), Hobbs and Krunic
(1971), Dahms (1973), van den Assem (1975),
Evans and Matthews 0976), van den Assem and
Maeta (1978, 1980), and van den Assem, Gijswijt
and Nubel (1980), van den Assem, et alia (1982).
Van den Assem has been investigating this aspect
of behaviour in several species of Melittobia. His
published work and personal communications
over the years have been of immense value in
guiding taxonomic decisions in the genus.
Van den Assem’s work proves that courtship
patterns in the genus show specific characteristics.
Within the genus there appear to be three basic
patterns (plus another demonstrated only by M.
clavicornis Cameron 1908). The three basic
patterns, acasta group, hawaiiensis group and
assemi group, together with that of M. clavicornis
have been discussed by van den Assem and Maeta
(1978, 1980) and van den Assem et alia (1982), but
I will briefly outline the situation for the sake of
completeness. The reader is referred to Dahms
(1983a) for an explanation of the species groups.
In M. australica {hawaiiensis group) the male
stands well forward on the female with his
mouthparts depressing her facial triangle just
below the ocelli. His scapes, placed over the
flagella of the female, lie close to her face.
Antennal contact is permanent during courtship
and antennation has only one pattern i.e.
alternating up and down movements of the flap-
like pedicel. The female is held around the neck
by the fore tarsi of the male, his mid legs are held
forwards with their tarsi alongside the eyes of the
female and his hind legs are braced against the
wings or hind legs of the female. In M. acasta
{acasta group), males stand with their heads a
little further down the face of the female without
the close contact of M. australica. The flagella of
the female fit into cup-shaped depressions of the
male scapes which are not pressed against the face
of the female. Antennal contact is broken during
the antennation sequence which has two
consecutive phases: knocking, jerky movements
involving the pedicel and at the end of this phase a
strong pinch involving the pedicel plus the first
funicle segment. Antennal contact is broken after
the pinch when the male raises his antennae
sideways. The female is held around the neck by
the fore tarsi of the male, the mid legs are braced
against the thorax of the female and the hind legs
are held forwards alongside the thorax of the
female.
DAHMS: BIOLOGY OF MELITTOBIA
347
In these two groups there is an alternation of
antennation and leg movements. On the basis of
these leg movements, the groups can be called mid
leg courters (ha^^'a^iensis and assemi groups) and
hind leg courters {acasta group). During
antennation, the mid legs of M. australica are
held laterally and forward with their trembling
tarsi alongside the female’s eyes. Van den Assem
and Maeta (1978) observed that at the start of the
display in their ‘species 2’ (= M. australica ) the
male’s middle legs arc braced against the female’s
thorax, but after the first antennation sequence
they are brought forward towards the female’s
head for the mid leg sequence. They do not fully
return to the original position after this but are
held out trembling and gradually move to the
frontal position at the start of the mid leg
sequence. From my observations the mid leg
sequence involves an upward swing of the mid
legs and a return to half way down the female’s
eyes slightly brushing them. They pause here for a
few seconds then are suddenly swung down and
backwards and this is accompanied by a strong
jerk of the male’s body. As with antennation
there is no change in the pattern of movements in
the mid leg phase until the finale when the male’s
body undergoes a series of convulsive movements
accompanied by up and downward swings of the
mid legs beside the female’s eyes.
After M. acasta males break antennal contact
they stretch their fore legs increasing the distance
between the heads of the courting couples. At this
point the hind legs move forward making swaying
movements beside the mesosoma of the female.
This sequence is ended by a push against the
female’s mid legs. The alternation of antennation
and leg movements continues for a period, but
they begin to overlap at which time there is a
change in behaviour pattern. Antennal contact
becomes permanent and co-ordination of the hind
leg movements change. The hind legs begin to rub
up and down on the side of the female’s
mesosoma. In the finale, the male places his hind
legs on the female’s wing or metasoma and brings
his mid legs forward to stroke the female’s eyes
with a downward movement. This is done with his
antennae stretched downward over the female’s
face. He then breaks antennal contact, raises his
wings at which point the female signals
receptivity.
The assemi group comprises a new species
complex from the Seychelles, India and Japan
(van den Assem and Maeta (1980)). Here the
courtship pattern resembles that of the
hawaiiensis group. The male’s scape is ventrally
grooved and he is a mid leg courter. The male
stands further forward over the female’s head so
that the distal part of his scapes touch her mouth
parts. Antennation involves a quivering motion as
in M. australica alternating with a pinch using the
pedicel. Alternating with antennation van den
Assem and Maeta describe the mid leg movements
as a very rapid kick involving the synchronous
movement of both legs as far forward as his own
head. During this movement parts of the female’s
body are brushed by long bristles on the ventral
surface of the femur of the male’s mid legs and
his tarsi brush the female’s pilose eyes. The mid
legs return to their initial position except that they
are held out laterally from the female’s mesosoma.
As the sequence proceeds the alternation of
antennation and mid leg movements accelerates
up to the last quiver which ends in a prolonged
pinch. Hereafter the mid leg movements become
an asynchronous to and fro rubbing motion
which lasts for a few seconds. In the finale, the
mid legs are moved synchronously back and forth
at which point the female may signal receptivity.
The species which stands alone is M. japonica
Masi, 1936 (= M. clavicornis ) and its courtship is
reported by van den Assem and Maeta (1978) and
van den Assem et alia (1982). Unlike that of the
other species, the male scape lacks an obvious
groove or cup-shaped depression but has a large
clear area distally opposite the attachment of the
pedicel. Male courtship position is the same as in
M. acasta but his scape presses the female’s
flagellum against her face. Antennation involves
a series of knocking movements as in M, acasta
and alternates with leg movements, but in this
species both mid and hind leg movements are
prominent. The mid leg movements are rigidly
stereotyped involving a rapid flick-like motion
towards the female’s eyes followed by a pause. At
this point the male may raise his antennae
sideways and break antennal contact, but this is
not always done. The hind leg movements are less
stereotyped and involve a walking motion
alongside the female’s metasoma or folded wings.
Leg movements are carried out during antennal
raising. There is no finale by the male and the
female signals receptivity during the sequence,
but always after a mid leg flick.
The courtship pattern in the genus is very
complicated and in some species can last up to 30
minutes. I have timed M. australica up to 15
minutes.
It is possible to draw some tentative
correlations between morphology and courtship
patterns in the genus. The following discussion is
restricted to those species for which courtship is
known and where a species group is mentioned it
348
MEMOIRS OF THE QUEENSLAND MUSEUM
includes only M. australica and A/, hawaiiensis
{hawaiiensis group), M. assemi and M. sosui
{assemi group) and M. acasta, M. evansi and M.
digitata {acasta group). The reader is referred to
Dahms (1963a) for figures illustrating
morphology.
Broadly spaced facial grooves and densely
setose eyes in females correlate with male position
(mouth parts impinging on upper face of female)
and mid leg courting in the hawaiiensis and
assemi groups. The presence of a dense tuft of
stiff setae on the ventral fore trochanters of males
of the hawaiiensis group seems to indicate some
difference in courtship position between males of
this group and the assemi group where this tuft is
absent. Dahms (1983b) discusses the application
of this setal tuft by M. australica males. Narrowly
spaced facial grooves and the sparcity of setae on
the eyes of females of the acasta group and M.
clavicornis correlate with the male head not
closely applied to the head of the female and the
predominance of hind leg action during
courtship.
Narrow male fore wings correlates with the
absence of male wing vibration during courtship
in the hawaiiensis and assemi groups in contrast
to broad male wings and male wing vibration
during courtship in the acasta group. A grooved
ventral scape and a geniculate scape gland in the
male correlates with permanent antennal contact
during courtship {hawaiiensis and assemi groups).
A cup-shaped depression in the ventral scape and
a non-geniculate scape gland in males correlates
with antennal contact through only part of
courtship (acasta group).
Amongst acasta group males there is some
variation in the size of the scape gland relative to
that in M. acasta; it is expanded in M. evansi, M.
femorata and M. chalybii; similar in M. digitata;
or reduced in M. scapata. Dahms (1983a, b)
discusses the possible implications. Also in the
acasta group there is variation in the size of the
first funicle segment in males; large in A/, acasta,
M. digitata. A/, femorata and Af . chalybii (the last
2 also have an extra expanded ring segment) and
relatively small in M. evansi and M. scapata. At
first it was thought that a large first funicular
segment in males might correlate with a pinch by
the male at the end of each antennal vibration
phase, but this does not appear to hold for A/.
digitata where, according to van den Assem et alia
(1982), there is no pinch at the end of a series of
antennal vibrations.
The mid femoral fringe in males varies between
and within species groups. It would be interesting
to see if these correlate with variations in male
mid leg movements and/or parts of the female
stroked during mid leg action in courtship.
There are a number of puzzling combinations
of these correlatable features, e.g. in M. chalybii
(acasta group) the male scape gland is geniculate,
his ventral fore trochanters have a setal tuft
resembling that of M, australica and the female
has densely setose eyes (hawaiiensis group); the
male scape has a ventral cup-shaped depression,
his antennal flagellum has a large first funicle
segment, his mid legs have an acasta group setal
fringe, females have narrowly spaced facial
grooves and a relatively thin scape in dorsal view'
(acasta group). It appears therefore that we are a
long way from understanding species
relationships within the genus and further study is
required to confirm or rearrange correlations
between morphology and courtship. Dahms
(1983a) in his summary discusses the matter in
greater detail.
REPRODUCTION
In the parasitic Hymenoptera, several aspects
of reproduction are important in understanding
evolution: parthenogenesis, sib-mating, biased
sex ratios, and sex ratio shifts.
Parthenogenesis
It is widely accepted that all species of
Hymenoptera reproduce parthenogenetically.
Gordh (1979) lists three types of parthenogenesis:
thelytoky, deuterotoky and arrhenotoky. A few'
species are thelytokous and the population
consists of only females or females plus a few.
non-functional males. Deulerotokous species are
also relatively few in number and unfertilised eggs
develop into both sexes. Most species are
arrhenotokous, i.e. the population consists of
diploid females and haploid males. The latter
develop from unfertilised eggs and are therefore
impaternate. In this case uninseminated females
can and do produce eggs from which only males
emerge.
The Melittobia species acasta, chalybii and
digitata have been shown to be arrhenotokous —
Howard and Fiske (1911), Malyshev (1911),
Graham-Smith (1919), Balfour-Browne (1922),
Buckell (1928) and Schmieder (1933). In the
present study, eggs from uninseminated M.
australica females produced males only, and
those from inseminated females resulted in both
sexes all of which indicates arrhenotokous
parthenogenesis.
Sib-mating
In the parasitic Hymenoptera, particularly the
Chalcidoidea to which Melittobia belongs, sib-
DAHMS: BIOLOGY OF MELITTOBIA
349
mating or close inbreeding appears to be the rule.
Hamilton (1967), Askew (1968a) and Crozier
(1977) list several biological features which
indicate that a species practices close inbreeding:
a) males are apterous or brachypterous and
therefore confined to the immediate area of their
emergence. Male Melittobia are brachypterous
and do not leave the host cell or puparium in
which they emerge.
b) close inbreeders are gregarious with eggs laid
in batches isolated from one another ensuring
that males and females from the one mother
emerge in spatial and temporal proximity to one
another. Melittobia are gregarious ectoparasites
and their host enveloping membranes (cell walls,
cocoons or puparia) ensure isolation of the egg
batches.
c) there is a tendency for mating to take place
on emergence before dispersal. Female M.
australica in my colonies would not disperse until
after insemination. Dahms (1973) mentioned that
uninseminated, freshly emerged females of M.
australica were observed to solicit the attention of
males and that it was not uncommon to observe
groups of females standing around a male
engaged in courtship, palpating him with their
antennae. They made no attempt to disperse.
Therefore Melittobia fit the biofacies for close
inbreeding.
Askew (1968), discussing speciation in the
Chalcidoidea, pointed out that the effectiveness
of sib-mating as an isolating mechanism is
increased by monandry in females, i.e.
unreceptivity after an insemination. Gordh and
De Bach (1978) found that male polygony and
female monandry are common in the
Hymenoptera. Female monandry requires
extreme economy of sperm utilisation and this has
been demonstrated in the arrhenotokous eulophid
Dahlbominus fuscipennis (Zetterstedt) by Wilkes
(1965). He found that from a single mating
involving 150 sperm, the female can produce as
many off-spring, over 90*7o of which are females.
Out of another batch of 254 eggs which he
stained, only 4 contained more than one sperm.
Such economy involved the synchronous release
of ova and sperm from the storage organs.
In laboratory cultures of M. australica I
noticed males frequently courting previously
inseminated females. Dahms (1973) felt this was
due to laboratory conditions where inseminated
females could not disperse. In all cases where I
have observed male M. australica courting
previously inseminated females attempts at
copulation by the male failed. The normal
situation is that females disperse after
insemination which precludes the attempted
second mating by a male. This is general for the
genus and therefore the species show male
polygony and female monandry. However, I have
found that M. australica females can and do mate
a second time apparently when their sperm supply
is depleted. Balfour-Browne (1922) considered
that M. acasta females also are able to mate a
second time when their sperm supply is depleted.
In both cases the females mate with a son. Under
laboratory conditions Melittobia exhibit another
facet of sib-mating behaviour which appears to be
widespread amongst arrhenotokous organisms
i.e. virgin females lay only a few eggs which
develop into males with which they mate
(Hamilton 1967). Howard and Fiske (191 1) found
that virgin females of M. acasta laid 4-5 eggs only
and these developed into males. The number of
unfertilised eggs laid was equivalent to the
number of unfertilised eggs laid if the female had
mated. They found also that virgin females lived
longer than fertilised females and survived to
mate with their sons after which normal egg
laying began. Balfour-Browne (1922) observed
the same behaviour with virgin M. acasta. By
removing unfertilised eggs from the hosts as they
were laid by uninseminated females he was able to
more than double the life of the female (up to 202
days) and increase the number of unfertilised eggs
laid.
The habit of uninseminated females laying only
a few eggs has been recorded for M. chalybii by
Schmieder (1933) and M. chalybii ( = M. digitata)
by Buckell (1928). They do not, however,
mention whether they mate with their sons. In the
present investigation 5 uninseminated M.
australica females were confined singly with a
host and produced only one egg each after 5 days.
When their sons emerged they mated and normal
egg production began. The ability of
uninseminated females to lay only a few eggs and
mate with a son is probably general throughout
the genus.
The economy of male production by
uninseminated females is easy to understand as an
adaptation for conservation of food supply that
would be depleted by production of superfluous
males (Schmieder and Whiting (1946)). In
Melittobia, mother — son mating has at least tw'o
advantages. Where species have a high level of
aggression between males, combat may result in
total annihilation of males or the surviving males
may die before all the females are fertilised.
Hobbs and Krunick (1971) found that in M.
350
MEMOIRS OF THE QUEENSLAND MUSEUM
chalybti(= M. acasta ) all males were often dead
before the last of the females became adult, which
meant that the last females to emerge had no
males with which to mate.
Balfour-Browne (1922) felt that mother-son
mating is part of the normal life cycle when a
female exhausts her sperm supply. He placed 5
freshly emerged, inseminated M. acasta females
in separate cells with a host. At the end of 6 to 7
weeks the females had ceased to lay eggs and he
noted that the last eggs to be laid produced males
only, indicating a depletion of sperm. After
providing each female with a male, a second
normal egg laying period began. Maeta (1978) has
confirmed this with M acasta. In this
investigation I noticed egg laying had ceased in a
stock colony containing 5 inseminated M.
australica females on a S. formosum host. All of
the progeny were in the larval stage. The host was
not completely utilised, indicating that egg laying
had ceased. The five females were separated and
each supplied with a fresh S. formosum pre-pupa.
Three of the females continued oviposition and
produced progeny of both sexes indicating that
the original host was probably unsuitable for
further oviposition. The two remaining females
produced 1 egg each which developed into males
with which they mated and normal egg laying
followed. It seems, then, that a second mating can
occur after sperm depletion. If the host is
nutritionally unsuitable for further oviposition,
migration within the host nest may occur.
Balfour-Browne (1922) with M. acasta felt that a
female migrates from a cell only when her
spermatheca is full. Once a female has completed
her first egg laying, she waits for a second mating
before migrating. As evidence he noted that in his
glass cells the female was often to be seen on the
cotton-wool plug after her second mating and
that this occurred generally when the host was
fully stocked with progeny or fully utilised.
Schmieder’s work in 1933 on the polymorphic
forms of M, chalybii presents a different
procedure in host utilisation. The normal or type-
form female produces from its first 12-20 eggs
rapidly developing second-form females and
males which are morphologically and
physiologically different from the type-form.
Second-form females begin laying eggs
immediately after fertilisation and assist the
mother in full host utilisation. The procedure
adopted in host utilisation may be related to host
size. In the case of larger hosts such as Sceliphron
spp. Schmieder’s system operates, and with
relatively smaller hosts, e.g. Pison spp.,
migration of females occurs due to competition
for food with her progeny. If sperm depletion
occurs in the latter case a female may mate with a
son. It is clear that such a close sib-mating
situation would ensure maximum host utilisation
and maximisation of a female’s reproductive
capacity. It also means that it is theoretically
possible for a virgin female to colonise an area by
mating with her son.
From the discussion above and from direct
observations on M australica^ it is clear that sib-
mating is an important part of the normal pattern
of reproduction in Melittobia. Askew (1968),
discussing evolution in the Chalcidoidea,
concedes that a small amount of outcrossing
occurs which mitigates against any tendency
towards inbreeding depression. Crozier (1977)
also considered that some outcrossing occurs. He
argued that the continued production of males is
puzzling if indeed there is no outcrossing.
Hamilton (1967) regards male aggression as
evidence that some outcrossing occurs in species
which exhibit the biofacies of extreme inbreeding
and arrhenotoky. He felt that outbreeding was
brought about by male migration or multiple
settling by females. In Melittobia, as males are
brachyplerous and non-dispersive, multiple
settling of females must be the method by which
outbreeding occurs. The host to parasite size ratio
in Melittobia, in some cases, would certainly
allow multiple settling and during the years I have
been culturing M. australica there has been no
reluctance by a female to oviposit on a previously
parasitised host even in the presence of more than
20 other females. That multiple settling occurs in
Melittobia can also be inferred from the
occurrence of male aggression within the genus.
That multiple settling of females is a fairly
common event in Melittobia can be seen from the
high degree of male aggression reported for some
species and the enlargement of male mandibles —
the weapons used in aggression.
Sex ratios
Amongst insects which exhibit extreme
inbreeding and arrhenotoky, female biased sex
ratios appear to be the norm, i.e., there is extreme
economy in the production of males. In
Melittobia spp. various workers have recorded
depressed ratios of 1-13*70 males and these are
made more biased by male aggression. In M.
australica I have found ratios of l-4*7o males.
Multiple parasitism has been shown by Wilkes
(1966) to result in a shift of sex ratio in the
pteromalid wasp Nasonia vitripennis Walker, a
parasite of house-fly pupae. Increasing the
number of females per host resulted in a reduced
percentage of female progeny. He postulated
three causes for this shift:
DAHMS: BIOLOGY OF MELITTOBIA
351
1) Superparasitism resulted in a greater number
of eggs per host and thus the number of eggs
per host was in excess of the number of larvae
the host could support. He assumed that
supernumaries were eliminated by starvation
and that reduced female progeny resulted
from stronger male competition. This
mechanism has been recorded for a number
of hymenopterous species and Wilkes (1966)
lists papers covering this subject.
2) Detection of previous parasitism.
3) Interference from other females on the host.
The last two mentioned result in a higher
percentage of unfertilised eggs being laid. Wylie
0965) on reviewing the literature found that
females of many hymenopterous species can
distinguish between parasitised and unparasitised
hosts. There are also cases in the literature where
females mark a host that they have parasitised.
In M. Qustralica where there is superparasitism,
there appears to be a greater production of males
but I have not quantified this. If there is a shift in
sex ratio then differential larval mortality could
be part of the shift since I have observed larval
cannibalism on numerous occasions where the
host was very crowded. Balfour-Browne (1922)
observed similar larval cannibalism in A/, acasta.
Dr, van den Assem is currently working upon
various aspects of sex ratio shifts in parasitic
Hymenoptera e.g. Charnow, Hartogh, Los-den,
Jones, van den Assem (1981). For this reason I
have not pursued this aspect of Melittobia biology
any further.
Therefore Melittobia exhibit the biofacies of
extreme inbreeding and arrhenotokous
reproduction. Outcrossing due to multiple settling
by females appears to be part of the normal
pattern of reproduction, and this is clearly
indicated by male aggression perhaps coupled
with sex ratio shifts in favour of males. The
normal sex ratio is strongly female biased and this
bias may be increased by male aggression. Males
are polygonous and females monandrous, the
latter dispersing after insemination. Mother-son
mating occurs when the female is uninseminated
or if she depletes her sperm supply.
POLYMORPHISM
Schmieder (1933) found two forms of each sex
in M. chalybii which he called the type - ( =
typical) form and the second-form. The two
forms showed marked morphological differences
which he described and figured. To. summarise
male
^ TABLE
Type Form
1)
pale
2)
3 ocelli
3)
eye spot pigmented
4)
wing normal for male, uncrumpled
female
1)
normal dark colour
2)
wings normal, uncrumpled
3)
cuticle normal, no fusion of sclerites
He found that in addition to these
morphological differences there were L.. equally
striking differences in their physiological
characteristics and in their behaviour’. Courtship
behaviour of the male second-form was less
regular than in the type-form and he found that
when he tried mating males of one form with
females of another, the lack of synchrony proved
troublesome. Van den Assem (pers. comm. 1981)
does not agree with Schmieder’s observations on
second form male courtship. He has had no
difficulty in mating one form maie with the other
form female. As there seems some doubt about
this aspect and since Dr. van den Assem is
working on the courtship behaviour of Melittobia
I have not pursued the matter further.
Second Form
dark reddish-brown
ocelli may be absent
eye spot unpigmented
wing smaller, uncrumpled
paler than type form
wings small, crumpled as they emerged
from pupa
cuticle thinner, some fusion of sclerites e.g.
on abdomen and antennae
The physiological differences between females
of the two forms in M. chalybii are quite
pronounced. Females of the second form have
larger metasomas in the pupal stage and
Schmieder (1933) suggested this was due to eggs
developing within the pupa. Egg laying began on
the day of emergence after mating in second-form
females. He found the life span of second-form
females to be shorter than that of the type-form
and that they make no effort to disperse, whereas
type-form females, after mating make their way
out of the host cell and disperse.
Schmieder’s investigations led him to conclude
that the causal factor was nutritional. The first
eggs laid develop rapidly to emerge as second
forms and the ‘... interpolation of an additional
352
MEMOIRS OF THE QUEENSLAND MUSEUM
generation of adults in the life history is thus seen
to constitute a remarkable biological adaptation
which effects a more complete utilisation of the
host and, as a corollary, secures the production of
the maximum number of offspring from each
host The type-form he saw as being
morphologically and physiologically the
dispersive phase.
Van Lith (1955) found a polymorphism in M.
acasta in which he mentioned only females which
had distended metasomas full of eggs and short,
often crumpled, wings. He did not feel there was
any connection between the production of these
females and nutrition.
Van den Assem and Maeta (1980) recorded
male dimorphism in M. sosui Dahms 1983a but
made no mention of dimorphic females. They did
not find any overlap between the two forms of
males and felt the causal factor was not
nutritional since males of both types emerged
from the same host at the same time. Van den
Assem (pers. comm. 1981) has informed me that
these imorphic males are actually distinct
morphs of the type-form. They are separate from
type and second-form males which also occur in
this species. This is a rather unusual phenomenon
in the genus and is one under study by Dr. van
den Assem. Detailed examination of the two
morphs of the type-form male shows few
differences except in size. In the larger morph the
forewings are larger and slightly crumpled (cf.
Figs 1 and 2) and the scape is about 1.2 times
larger than that of the smaller morph.
I have received for identification some slide-
mounted specimens from the U.S. National
Museum which are obviously second-form males
and females. There is little to use for
identification since the dimorphism has affected
most of the diagnostic morphological features.
Those that appear to be unchanged cause
uncertainty, e.g., the mid leg bristle pattern (Fig.
9) and the proportions and shape of the mid tibia
of males resemble those of M. acasta males
whereas the most common scape morphology is
that of M. evansi (Fig. 1 1). For the present I have
decided to label these specimens as acasta group
and positive identification must await breeding of
second-forms of all species in the acasta group. In
the following discussion therefore, the features
described are compared to those of the acasta
group rather than to any particular species.
Female: Larger than type-forms, 1. 7-2.1 mm
long. Colour brown except flagellum which is
infuscated. Head in frontal aspect quite
broad and more rounded than in type-forms.
Eyes relatively smaller. Ocelli variously
reduced as follows: 2 normal posterior ocelli
with either a very small or absent median
ocellus; normal right, posterior ocellus and
with median ocellus; or small right ocellus
only. Scrobes shorter than type-forms.
Mandibles (Fig. 4) more like those of the
male. Antennae variable (Figs 13-15) even
between right and left on the same specimen;
scapes of variable shape with some showing
expansion similar in form but not size to
those of some males; flagellum showing
fusion of segments in some specimens e.g.
fusion of funicle 2 and funicle 3 is the
commonest, but fusion of funicle 3 and club 1
also occurred and in some, the delimitation of
club segments is imperfect; plate organs in
some specimens are modified to peg-like
structures (Fig. 16). In lateral aspect the head
appears to be more inflated than in type-
forms.
Mesosoma in dorsal aspect (Fig. 10) appears
broader and shorter than in type-forms; setal
fringe on posterior margin of prothorax
shorter; sutures on mesonotum less distinct
than type-form particularly those delimiting
the axillae; position of setae on scutellum
variable even from right to left on the one
specimen e.g. normal position as in type
forms or with anterior setae moved close to
posterior setae; propodeum much broader
and shorter than type-form, more angular in
shape resembling the propodeum of the male.
Legs similar to those of type-forms. Wings
reduced (Fig. 6), crumpled, remaining as they
emerge from the pupa; postmarginal and
stigmal veins poorly developed, the stigmal in
some specimens closely resembling that in
type-form male wings. Lateral aspect not
visible.
Metasoma in dorsal aspect much larger than
that of pre-feeding type-form females.
Male: 1.6-1. 8 mm long. Colour light brown.
Head in frontal aspect (Fig. 8) rounded, not
contracted ventrally as in some type-forms.
Mandibles (Fig. 5) not unlike those of the
female second-form. Eyes are much larger
than those of the type-forms. Ocelli variously
developed as follows: median ocellus reduced
or absent, posterior ocelli normal; only the
right, posterior ocellus developed, the others
absent; or all ocelli absent. Antennae (Figs
11, 12, 17, 18) variable, the predominant
scape morphology is as in Fig. 1 1, but there is
variation even between left and right on the
same specimen (Figs 17,18), scape glands vary
DAHMS: BIOLOGY OF MELITTOBIA
353
FIGURES 1,2 — Melittobia assemi (sp. nov.) male fore wings.
FIGURES 3-9, Melittobia acasta group second-form male and female; 3 — Male fore wing; 4 — Female mandible;
5 — Male mandible; 6 — Female fore wing; 7 — Frontal aspect, female head; 8 — Frontal aspect, male head; 9 —
Male mid leg.
354
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURES 10-18, Melittobia acasta group second-form male and female. 10 — Dorsal female thorax; 11 — Male
scape; 12 — Male flagellum; 13,14 — Female scapes; 15,16 — Female flagella; 17,18 — Male scapes from the
same specimen.
DAHMS: BIOLOGY OF MELITTOBIA
355
in shape and development even between right
and left on the same specimen funicular
segments all fairly uniform in size unlike
those of type-form M. acasta group where
segment 1 is enlarged; flagellum showing
different degrees of segmental fusion even
between right and left on the same specimen
as follows: funicle segments I and 2; 1, 2 and
3; 2 and 3; 4 plus club segment I ; and in some
the delimitation of club segments is
imperfect; plate organs appear to be reduced.
Lateral aspect not visible.
Mesosoma in dorsal aspect similar to the
type-forms. Legs not greatly modified; mid
leg (Fig. 9) similar to that of M. acasta, some
specimens showing fusion of tarsal segments
3 and 4; in some specimens tarsal segments 3
and 4 of hind legs are also fused. Wings (Fig.
3) not much reduced in size, stigmal vein
absent in most specimens, poorly developed
in others.
Metasoma of normal proportions.
Material Examined:
14 $ 9, 2 ^ , on microscope slides labelled 20
mi. South Washington D.C. December 1974
Trypoxylon sp. nest Col Gordh; 8 2 9, 8 >3 .5
on microscope slides, data as before but
collected 10 January, 1975; 10 9?, 6 3^^ on
microscope slides labelled, Augusta West
Virginia February 1975 ex Trypoxylon nest
Col. A. Menke. These are in the collections of
the U.S. National Museum, Washington,
D.C.
A polymorphism without the marked
morphological differences of M. chalybii and the
M. acasta group discussed above occurs in M.
australica. In my trials where up to 20 M.
australica larvae per host were bred on Sceliphron
formosum prepupae, second-form progeny
resulted. Trials were not carried out to determine
the upper limit of parasite to host larvae for
second-form production. Males of M. australica
second-form were larger than the type-form but
otherwise appeared morphologically similar to
the latter. Second-form females were larger than
the type-form with reduced eyes, shortened wings
and enlarged metasomas. In neither sex was there
any evidence of fusion of tarsal or antennal
segments and the scapes of both forms were
normal. The shortened wings of the second-form
females were not crumpled, but fully expanded
without any alteration of venation.
Second-form M. australica females differ
behaviourally and physiologically from type-form
females. They are ready to lay eggs immediately
after insemination, i.e. on the day of emergence.
Type-form, inseminated females begin laying 4-5
days after being placed on a host. Second-form
females make no attempt to disperse from the
breeding chamber, but remain on or under the
host and show a negative reaction to light typical
of laying, type-form females with a host. Type-
form females, after insemination, make their way
to the top of breeding jars and show a positive
reaction to light, e.g. if released they move in a
direct line towards windows.
Some of the features of second-form M. acasta
group females are similar to those in type-form
males. In comparison to type-form females, the
head is shorter, broader and more inflated and
the eyes are relatively smaller. The mandibles
more closely resemble those of a male; the scapes
are expanded and some resemble the sorts of
expansions found in male scapes (Fig. 13); the
mesosoma, particularly the propodeum, is shorter
and broader as in males; and in some cases the
stigmal vein in the crumpled wings resembles that
in males.
It is interesting that the M. acasta group
females on hand show modifications resembling
those from which the present male morphology
appears to have arisen and that these
modifications are nutritionally induced, at least in
part. Males tend to emerge first e.g. Buckell
(1928) found that M. chalybii (= M. digitata )
males emerge after 21 days and females after 37
days. Schmieder (1933) found that the first
progeny to emerge in M. chalybii were second-
form individuals. Differential development of the
sexes is no doubt related to their dispersive and
non-dispersive roles, and in the male subsequent
modifications plus embellishments would be
related to the restriction of their role to combat,
courtship and copulation.
In addition, the second-form males of the
acasta group on hand show quite an amount of
variation in both head and scape morphology,
e.g. Figs 17, 18 are right and left scapes of the
same specimen. This is in contrast to males of the
hawaiiensis group. I have not seen polymorphic
forms of the assemi group. From our knowledge
of species at the moment the acasta group
contains the greatest number of species (7), the
hawaiiensis group contains 2 and the assemi
group contains 4. Perhaps this apparently greater
diversity of species in the acasta group is related
in part to the variability found amongst second-
form acasta group males. However, this is
speculative since the world fauna is not properly
known and the full implications of polymorphic
forms in Melittobia require much more study.
356
MEMOIRS OF THE QUEENSLAND MUSEUM
LIFE HISTORY OF M. AUSTRALICA.
No particular part of the host was favoured for
oviposition and eggs were laid in clusters
generally in the intersegmental grooves of
prepupae. Eggs are large relative to the female’s
metasoma; 0.38 mm long by 0.1 mm wide. The
length of the metasoma of inseminated, unfed
females is 0.6 mm. Eggs are elongate, slightly
curved with broadly rounded ends, one end being
slightly wider than the other. They fit the
hymenopteriform type of Clausen (1962). They
are white and translucent with a thin, smooth
chorion. The surface appears moist and coated
with a substance that makes them loosely adhere
to each other and to the host.
Larvae hatched in 3-4 days and the newly
hatched larvae fitted the hymenopteriform type
of Clausen (1962), i.e., white, translucent, visibly
segmented grub unadorned by obvious spines,
setae etc. The head and mouth parts are relatively
small. The eggs, larvae and larval head of M.
australica resembles the figures of M. acasta
(Balfour-Browne 1922). As feeding proceeds
waste material can be seen accumulating within
the larva. No attempt was made to determine the
number of larval moults, but Balfour-Browne
(1922) recorded 2 larval moults plus the larva to
pupal moult in M. acasta. After 7-9 days, larvae
were fully grown and measured 1.6 mm long by
0,5 mm wide. They were distended and smooth
without obvious segmentation. When feeding
finished, the larvae rolled from the host remains
and voided waste material as faecal pellets
resembling strings of beads. One day later they
pupated. At this stage it was easy to distinguish
the sexes because of the enlarged scape and
absence of eyes in the male. The pupal stage
lasted 3-4 days and the total life cycle was
therefore 14-18 days. In the case of second-form
progeny, the life cycle duration was 12-13 days.
The above figures were obtained from rearings at
25-30^C.
The average total production from 10 type-
form females, each on a separate S. fortnosum
prepupa was 370 females and 7 males. The
percentage males varied between 1 and 4*^0 in
newly emerged adults.
The literature reports a wide range for life cycle
duration, Balfour-Browne (1922) obtained a time
of 17-23 days for M, acasta at an unspecified
temperature which compares with 25-29 days
(second-form) and 37-47 days (type-form)
recorded by van Lith (1955) for the same species
at 18-19°C. Buckell (1928) found that male M.
chalybii (= M. digitata ) took a total of 21 days
compared to 37 days for females but did not
specify any rearing temperatures. Schmieder
(1933) bred M. chalybii at 19-25°C and recorded
a life cycle length of 90 days for type-form
individuals and 14 days for second-form
individuals. It appears therefore that some
standardisation of rearing temperatures is
required before results can be compared. Even so,
the result of 90 days for M. chalybii type-form
individuals obtained by Schmieder (1933) seems
excessively long in comparison with other figures.
My results with M. australica show very little
difference in life cycle time between type and
second-form individuals.
DISPERSAL
There are two aspects to dispersal, natural and
man assisted. The latter is important since the
plastic behaviour exhibited by Melittobia has
allowed it to avail itself of man’s travelling
facilities.
Males do not disperse, but die in the host cell or
puparium in which they emerge. Inseminated
type-form females escape from the host cell or
puparium either by excavation or through
entrance holes made by the mother. From this
point on workers provide a variable story.
Graham-Smith (1916, 1919) observed that
female M. acasta can fly for a considerable
distance. Malyshev (1911) in contrast, found that
M. acasta females could fly only a few
millimetres. Balfour-Browne (1922) observed that
female M. acasta fly only 25 mm or so at a time
and for the most part do not use their wings. He
suggested they might disperse by phoresy, but
there is no evidence to support this. Buckell
(1928) did not observe M. chalybii ( = M. digitata)
flying, but noted they hop like fleas when
disturbed and concluded that although they were
winged they were flightless, relying on their legs
for dispersal. Krombein (1967) found that M.
chalybii females do not fly frequently but rely on
walking. Van den Assem (pers. comm. 1981)
considers that flying in Melittobia spp. is partially
a matter of temperature. At higher temperatures
or in direct sunlight Melittobia females will fly
away, but at temperatures less than 20°C they will
not.
Evidence in the literature seems to suggest that
the dispersal power of female Melittobia is
limited, but my observations and some recent
work by Freeman (1977) and Freeman and
Parnell (1973) indicate that this is not so. When I
released inseminated M. australica females in the
laboratory they dispersed initially by hopping and
DAHMS: BIOLOGY OF MELITTOBIA
357
running. Later they flew. They were noted to be
capable of flying 3 metres towards a closed
window where they accumulated. Within 5
minutes there were no females left in a ! metre
radius of the release area. This area had been
cleared prior to release to avoid females hiding or
being unobserved. When the window was opened
the females flew outside. I suggest that all
emerging fertilised female Melittobia have
functional wings used for dispersal. This dispersal
is no doubt assisted by air currents.
Freeman and Parnell (1973), investigating the
mortality of Sceliphron assimile Dahibom caused
by M. chalybii (= australica ) in Jamaica, found
that the parasite accounts for 16% of
developmental mortality in the host. Moreover,
where Sceliphron forms large breeding
populations the parasite kills a higher proportion
of them. Freeman (1977) found some variations
in the percentage mortality expected on the basis
of a linear density-dependent relationship and
that these were often partly due to the effects of
the prevailing easterly or south-easterly winds
carrying Melittobia across Jamaica. Each host
cell can yield up to 300 alate Melittobia which
means that large numbers of females can be
released into the air from host nests. Freeman
(1977) argued that the further inland or westward
a host cell might be the greater its chances of
being found by a flying Melittobia since there
would be an increasing number up-wind of host
nests producing Melittobia. Conversely, nests
near the sea shore or towards the east would have
less chance of being found. He concluded there is
circumstantial evidence that the higher
percentages of Melittobia parasitism observed
away from the shore and to the west were caused
by dispersal of the parasite by the wind.
Further circumstantial evidence exists to
support long range dispersal. Using figures
provided by Freeman (1977) it is seen that each
host cell can produce up to 300 alate females. At
10 high-density sites the host had 3499 cells with
3458 eggs laid of which 1430 were killed by
Melittobia. The maximum yield from these cells is
nearly 500,000 alate female Melittobia. Dispersal
would be necessary just to find enough hosts and
if it did not occur one could probably expect a
higher percentage developmental mortality by
Melittobia than the 41. 4% recorded by Freeman
and Parnell (1973) in areas of host density. Since
Melittobia are delicate insects one would expect
passive wind dispersal to result in high mortality.
The production of large numbers of alate females
could offset the risk factors in wind dispersal.
Man’s ability to travel on a global scale has
provided Melittobia with an added means of
dispersal. Several features of its biology allow it
to take advantage of man’s travelling facilities.
1) Melittobia are highly polyphagous. One
could reasonably expect to find mud nesting
Hymenoptera and cockroach oothecae
associated with ships and packing crates. In
the past, hygiene on sailing ships was
probably not of a high standard and some fly
puparia were no doubt present. On long
journeys more hosts could be taken on board
at port stops. All of the above hosts are
recorded for Melittobia.
2) Females are able to delay feeding and
oviposition for several weeks until a host is at
a suitable stage for oviposition or until a
suitable host is located. Modern, rapid
transport reduces the risk for Melittobia and
packing crates provide the necessary hosts
rather than air craft e.g. the North American
Sceliphron caemenlarium (Drury) is
spreading rapidly through the Pacific region
and in July 1979 was intercepted at Alice
Springs, Australia in packing crates from
North America (Naumann 1980 unpublished
report). In December 1980 this species was
collected from Eight Mile Plains near
Brisbane from nests in a dwelling.
3) Melittobia females have the capacity to be
very efficient founder organisms. It is
theoretically possible for an uninseminated
female to begin a new population by laying a
few unfertilised eggs. These develop into
males with whom she then mates. This aspect
has been discussed more fully under
‘Reproduction’.
ACKNOWLEDGMENTS
This paper was taken from my M.Sc. thesis
submitted to the University of Queensland in
1982. My superviser, Dr Elizabeth Exley,
University of Queensland, was extremely helpful
in providing constructive comments and editorial
remarks. Dr T. Woodward, University of
Queensland and Dr G. Gordh, University of
California as examiners provided corrections and
advice towards publication of the thesis. Dr
Gordh was also of great assistance, imparting to
me many of his illustration techniques. Dr H.
Townes, American Entomological Institute,
U.S.A. kindly provided identifications of
ichneumonid hosts.
358
MEMOIRS OF THE QUEENSLAND MUSEUM
My Technician, Miss Gudrun Sarnes, was of
great assistance checking manuscripts and
numbering figures. The typists whose patience I
tried severly were Miss P. Tinniswood and Miss
E. Proberts. My wife Judith assisted with
manuscript checking and figure assembly.
Special thanks are due to Dr J. van den Assem,
University of Leiden, Holland. We have
corresponded freely since 1974 and he has been of
the greatest assistance with notes from his
ethological studies.
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Georgia, U.S.A., 1971, 52 pp.
Hobbs, G.A. and M.D. Krunick, 1971.
Comparative behaviour of three chalcidoid
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cutter bee, Megachile rotundata, in the
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Howard, L.O., 1892. The habits of Melittobia.
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the United States of the parasites of the gipsy
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report on progress, with some consideration
of previous and concurrent efforts of this
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312 pp.
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superfamilies Chalcidoidea, and Procto-
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1-29.
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of Melittobia chalybii Ashmead and the
determining factors involved in their
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Woods Hole 65: 338-52.
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economy in the chalcidoid wasps Melittobia.
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and H.L. Parker, 1927. The problem of host
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360
MEMOIRS OF THE QUEENSLAND MUSEUM
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1966. Some mechanisms that affect the sex
ratio of Nasonia vitripennis (Walk.)
(Hymenoptera : Pteromalidae) reared from
superparasitised housefly pupae. Can. Ent.
98: 645-53.
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97: 279-86.
Mem. QdMus. 21(2): 361—85. [1984]
AN INTERPRETATION OF THE STRUCTURE AND FUNCTION OF THE
ANTENNAL SENSE ORGANS OF MELITTOBIA AUSTRALICA (HYMENOPTERA :
EULOPHIDAE) WITH THE DISCOVERY OF A LARGE DERMAL GLAND IN THE
MALE SCAPE.
Edward C. Dahms
Queensland Museum
ABSTRACT
The importance of the antennae during courtship behaviour of Melittobia species prompted
an investigation into the histology of the enlarged male scape using the single Australian
species Melittobia australica. The application of the male scape during courtship suggests a
possible chemical communication between the antennae of the two sexes. Histological and
SEM work reveal the presence of a large dermal gland in the male scape. SEM work and
chemical applications reveal the presence of long thin unfluted setae, tapering fluted setae,
multiporous plate sensilla and short basiconic capitate pegs on the antennae of both sexes (the
short basiconic capitate pegs are absent in most males). Together with behavioural
observations these are used to suggest the possible structure and function of the antennal sense
organs and the most likely receptor for the male scape pheromone.
MALE SCAPE
Amongst the parasitic Hymenoptera, male
antennation is a common component of
precopulatory behaviour and it reaches a high
expression in Melittobia (Gordh and DeBach
(1978)). The enlargement of the male scape and its
application discussed by Dahms (1983b) suggest
that it has a function in stimulating the female’s
antennae. In males of all species cleared in 10%
NaOH the scapes show a clear delimited zone (PI.
la, M. australica). In M. australica the surface
appears to have a cellular pattern at higher
magnifications. Since all internal tissue is
removed in this process, the clear delimited zones
must be cuticular which suggests mechanical
stimulation of the female’s flagellum. However,
freshly killed M. australica males used for AgNOj
staining to test for touch chemoreceptors when
examined after 30 minutes in Toluene were found
to be not completely cleared. In the scapes of
these males could be seen a cellular-like zone
occupying the same area as the clear delimited
zone in NaOH cleared specimens. (PI. la) is a
lateral view of the side opposite pedicel
attachment and (PI. lb) is an end view of the same
side. This cellular-like zone was absent in
specimens fully cleared in Toluene indicating that
it was internal tissue. Serial sectioning of male M.
australica scapes clearly shows large dermal
glands which follow the clear delimited zone in
NaOH cleared specimens (PI. 3b, c; 4a). The
sections also show that the inner cuticular lining
on the scape groove is much thinner than the
outer cuticle and that there are cuticular
infoldings along the length of the gland. These
cuticular infoldings form the limits of the clear
zone in NaOH cleared specimens.
To prepare them for SEM examination, males
were treated with 10% NaOH until cleared to
remove any glandular secretions which might
obscure the cuticular surface of the gland. They
were then dehydrated in alcohol and finally
treated with Toluene to remove any wax which
might obscure cuticular pores. Males were air
dried and mounted upside down on stubs in
preparation for gold coating. It was noticed that
after removal from Toluene and air drying the
flexible intersegmental cuticular areas became
white while the thicker cuticular sclerites
remained yellow-brown. The lining of the scape
groove became white indicating that it was
flexible thin cuticle, which would explain the
cuticular infoldings around the gland for support.
SEM photomicrographs (PI. 2b, c; 3a) show the
cuticular surface over the gland to be well
differentiated and perforated by numerous pores.
At higher magnifications the cuticular surface
over the gland shows a somewhat reticulate
362
MEMOIRS OF THE QUEENSLAND MUSEUM
appearance. Noirot and Queenedey (1974)
mention cuticular specialisations in Heteroptera,
Blattodea and butterflies associated with dermal
glands and that these serve as evaporative areas
ensuring rapid evaporation of secretions. The
cuticular area over the scape gland in M.
australica fits this pattern.
The shape, size and position of the scape gland
varies with species and provides a very useful
taxonomic tool. I refer the reader to my
taxonomic revision of the genus (Dahms 1983a)
for a fulle- discussion together with figures and a
discussion of species groups.
Van den Assem et alia (1982: 458) raise a rather
interesting point. They found that males with
their antennae removed do court females and
induce receptivity. From this they concluded that
stimuli which might arise from them are by no
means necessary. The data they present are from
mutilation experiments with M. acasta (Walker)
and it is not clear from their account if they
carried out similar trials with all species at their
disposal. If we look at the male scape gland in M.
acasta it is not as extensive as in the hawaiiensis
and assemi groups. The cuticle over the scape
gland in M. acasta as shown by Van den Assem et
alia (1982, PI. I) does not appear to have an
evaporative function as it does in M australica
(PL 2c, 3a). Perhaps the scape gland does not
have the importance in the acasta group that it
has in the hawaiiensis and assemi groups. It is
interesting to note also that antennal contact is
permanent throughout courtship in the
hawaiiensis and assemi groups but only through
part of the cycle in M. acasta and M. evansi
Dahms (1983a) but not in M. digitata Dahms
(1983a). There may be some variation in the
importance of permanent antennal contact in
some members of the acasta group (not studied by
Van den Assem et alia (1982)) which seem to have
relatively expanded scape glands eg. M. femorata
Dahms (1983a) and M. chabylii Ashmead. In the
latter the scape gland in genicualte as in the
hawaiiensis and assemi groups although that of
M. chabylii is not as extensive.
Goodpasture (1975) observed pores in the
modified scapes of the torymid chalcidoid wasps
Monodontomerus montivagus Ashmead and M
dementi Grissell which were applied to the tip of
the female flagellum during the climax phase of
courtship. He concluded that the male scapes
might be the source of chemical communication
as a behavioural cue and further suggested that
the pores indicated either a chemical sensory
function or pheromone elaboration sites. From
my studies on M. australica I suggest that they are
probably the latter. Antennation during courtship
is a common phenomenon in Chalcidoidea and is
often accompanied by antennal modifications.
Pheromone glands in male antennae may also be
a common occurrence. Houston (1975) has found
antennal modifications containing derma! glands
in several Australian species of the bee genus
Hylaeus. Antennal modifications containing
glands may be more widespread in the
Hymenoplera than current knowledge indicates.
ANTENNAL SENSE ORGANS
During the course of a biological study on M.
australica Dahms (1983b) it was decided to
investigate the sense organs on the antennae since
the latter play an important part in precopulatory
behaviour. The following discussion is based on
SEM work, behavioural observations and a few
chemical applications. It does not have the benefit
of histological or electro-physiological data,
therefore the structures and functions of the sense
organs are suggested rather than conclusively
proven.
Males of all species of Melittobia have their
compound eyes reduced to a single ocellus-like
spot, Picard (1922) examined M. acasta males
histologically and found that the reduced eyes
lacked the normal structural elements of even an
ocellus. The optic ganglia were also reduced
relative to the female. He related this reduction,
together with shortened wings and relatively
reduced pigmentation, to the male’s restriction to
the host cell or puparium in which they emerge.
This reduction in functional elements in the eyes
of males is no doubt general in the genus.
The sole function of the male is related to
reproduction. Van den Assem and Putters (1980)
found that sound production is not involved in
the courtship of Melittobia. Presumably males
rely on chemical and tactile stimuli for locating
females and for precopulatory behaviour.
Chemical information appears to play an
important role in the behaviour of both sexes.
The size of the complex gland in the male scape
and the role of this segment during precopulatory
behaviour suggests that the female receives a
considerable chemical input during antennation.
Behavioural observations (mentioned under
‘Long thin unfluted setae’ below) indicate that the
sexes are chemically different and there are easily
discernible behaviour patterns depending on the
sex encountered by individuals of both sexes.
Females have additional occasions in which
olfactory reception could be important e.g. host
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
363
location and feeding. The antennae of both sexes
have setae whose structure suggests tactile
receptivity.
SEM examination of the antennae of males and
females of M. australica revealed the presence of
the following sensory structures:-
1) long thin unfluted setae
2) tapering fluted setae
3) multiporous plate sensilla
4) short basiconic capitate pegs
1) Long thin unfluted setae (PL 5b,)
These are readily distinguished by the absence
of both a basal socket and fluting. They are only
present on the club of both sexes especially at the
tip of the terminal segment.
Lack of a socketed base suggests they are not
tactile in function. A possible contact
chemoreceptive function is suggested by their
concentration at the tips of the antennae,
particularly noticeable in males, and by
behavioural observations.
A male can instantly distinguish between the
sexes by tapping another individual with the tips
of his antennae. His behaviour varies
dramatically according to the sex encountered;
another male induces aggression, a female is
mounted.
Females can also distinguish between the sexes.
Virgin female M. australica when confined
without males stand around with their mandibles
open. When provided with a dead male pupa they
immediately become active and begin searching
behaviour. When another female is encountered
they stop, palpate the encountered female with
the tips of the antennae then resume searching
behaviour. When the dead male pupa was
encountered and palpated, searching behaviour
ceased. The females stood around the pupa
continually palpating it with the tips of their
antennae. Some of the females opened their
mandibles. In cultures, similar reactions occurred
and it was not uncommon to see groups of
females standing around a male engaged in
courtship, palpating him with the tips of their
antennae. Mandibular opening was also observed
in these groups of virgin females and it suggests
that mandibular glands could be the source of a
female odour. Gordh and DeBach (1978)
mentioned that mandibular involvement in
courtship appears to be an adaptation in some
Chalcidoidea; also studies showed that olfaction
could be used for mate attraction. They suggested
that the gland-like ducts in the mandibles of
chalcidoids may indicate exocrine glands. The
mandibles of both sexes in Melittobia have two
such gland-like ducts. During courtship in M.
australica, mandibular opening by females occurs
towards the end of the sequence which brings
them into close proximity with the flagellum of
the male.
Female behaviour on a host also suggests a
contact chemoreceptive function for thse sensilla.
A fertilised female uses her ovipositor to puncture
the host then feeds on the droplet of host body
fluid that wells forth. After withdrawal of the
ovipositor the female moves backwards palpating
the surface of the host with the tips of her
antennae until the droplet is encountered. At this
point feeding begins. Female M. australica revisit
old puncture sites to feed upon congealed host
body fluids which they relocate by palpation with
the tips of their antennae.
SEM examination of the tip of these setae does
not reveal the typical pores of contact
chemoreceptor sensillae. Following the
procedures of Slifer, Prestage and Beams (1957)
very good results were obtained using AgNo,.
After 60 minutes, the AgNo, had penetrated the
tips of these setae (PI. 5a). The penetration was
more rapid in males than in females which is
probably related to the setae being of larger
diameter in the males. These tests, behavioural
observations and location of these setae suggest
therefore that they are touch chemoreceptors.
In the female, touch chemoreceptors on the
terminal antennal segments would also be of use
in host identification. Female M. australica walk
over a host nest palpating it with their antennae.
Once it has been identified and entered these
sensillae may be of use in detecting the presence
of enveloping membranes, e.g. cocoons. They
may also help differentiate hosts e.g. oviposition
behaviour differs between hymenopteran and
some dipteran hosts. Should a Melittobia female
enter a host cell when the host is immature these
sensillae would allow her to distinguish between
the host and its provisioned food. The relative
amount of food provision and its state of
preservation may be of use in distinguishing if the
host had failed or its stage of development. On
the other hand these sensillae may be used directly
to ascertain the age of the host. No information is
available on whether there are chemical
differences between the different stages of a host,
but since the hymenopteran hosts I have observed
accumulate waste internally and pass it out just
before pupation as meconium, the relative
amount of internally accumulated waste or the
presence of meconium may be an important
sensory signal for oviposition in Melittobia.
364
MEMOIRS OF THE QUEENSLAND MUSEUM
Finally these sensilla may be useful in detecting
the suitability of a host i,e. whether the host is
diseased.
2) Tapering fluted setae (PI. 5b, ii)
These arise from a socketed base and show a
slightly whorled fluting on the surface (PI. 6d).
They do not take up AgNO, stain. The fluting
provides rigidity allowing the setae to resist
bending thus transferring maximum movement to
the socketed bases. In the male they are present
on all antennal segments with marked
differentiation. On the proximal segments of the
flagellum they are long and numerous, but are
reduced in length and number towards the
terminal club segment (PI. 6a). On the other
segments of the antenna they are also relatively
shorter and are fairly evenly distributed except for
their absence on the lining of the scape groove
and for a relatively denser arrangement of
shortish setae on the upper surface of the lateral
expansion of the pedicel. In the female they are
present on all segments of the antenna and have a
fairly even distribution (slightly fewer on the club)
without any size differentiation. In general, they
are shorter and finer than in the male and they are
shorter and finer than the long thick non-fluted
setae on the clubs of both sexes.
Because of the fluting and socketed base I
assume they are touch receptors. Their general
uniformity in size and shape in the female
indicates they have no specific function, just
providing generalised tactile information, e.g.
they would be of assistance in estimating the size
of the hole the female excavates in the host cell. In
this way the female would be able to estimate if
the excavation is wide enough and when
penetration has been effected. Female M.
australica when excavating in a Sceliphron
formosum nest were noted to insert their
antennae periodically and touch the walls of the
excavation with the sides of the flagellum.
Differentiation of these sensilla on the male
flagellum suggests a specific function. When a
male encounters a female he mounts her first then
searches for her head with his antennae. When he
mounts a female he orientates longitudinally on
her and taps his flagella either side of the female’s
extremity like a pair of cupped hands thus
engaging the long setae on the proximal flagellar
segments. When the female’s posterior metasoma
is touched the male turns 180° and repeats the
procedure at the head then scoops her antennae
into his scape groove. Where the female's head is
touched no turning occurs. This may not be the
only sensory input e.g. if the female produces a
female scent from mandibular glands then
orientation on the female could also involve
olfactory information via his multiporous plate
sensilla.
Before passing on to the other sensilla, mention
should be made of two clusters of differentiated,
socketed, fluted setae occuring on leg segments in
the male. In M. australica and M, hay^aiiensis
males the ventral surfaces of the fore-trochanters
bear a dense tuft of thick, short, socketed setae
with whorled fluting (PI. 4b). In males of all other
known species except M. chalybii where they are
not as w'ell developed the fore-trochanters have a
few fine, undifferentiated, scattered setae
ventrally. During courtship of M. australica and
M. hawaiiensis these setae press firmly down on
the pronotum of the female. In all other species
for which courtship is known, the position of the
male is such that the fore trochanters are not in
contact with the female. It is difficult to suggest
the use that these serve, but since modifications in
male Melittobia morphology are closely linked to
some aspect of courtship there must be some
important sensory input via these setae. Perhaps
they are useful in positioning the male for
courtship.
Another group of differentiated, fluted setae
with socketed bases occur on the posterior ventral
surfaces of the mid femora of males of all species
and ventrally on the mid trochanters of all species
except M. australica and M. hawaiiensis. Those
of M. australica are definitely socketed with
shallow, unwhorled fluting (PI. 4c). These setae in
all species are much longer than the general body
setation and there is some differentiation amongst
them. The mid legs are used by the males of all
species during courtship and in M. australica
these setae were noted to brush the ‘shoulder’
junction of the pro- and mesonotum of the
female. The pattern of distribution and the degree
of differentiation amongst these setae varies with
the species Dahms (1983a). It would be interesting
to see if these variations are related to specific
differences in male mid leg movements and/or the
parts of the females brushed. Their function in
the male may be to signal contact with that pan of
the female to be stroked and their input to the
female would most likely be tactile also via her
undifferentiated general body setation. Females
do show differentiated long setae on the posterior
pronotum, the mid-lobe of the mesoscutum and
the scutellum, but these are not in a position to be
stimulated by the differentiated mid leg setae of
the male.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
365
3) Multiporous plate sensilla (plate organs) (PI.
5b, iii).
These are raised sensilla which appear as pale
areas on dry and slide-mounted antennae of most
Hymenoptera. In Melittobia they are elongate
(0.009 X 0.003 mm) on female club segments of
M. australica) and are present on all flagellar
segments in both sexes except for males of the M
hawaiiensis and M. assemi groups where they are
restricted to the club segments only. On the
funicle segments they tend to be orientated
transversely and in general they are fewer in
number than on the middle club segment. On the
club segments they tend to be longitudinal in
arrangement and the transverse arrangement on
the funicle segments may be related to the smaller
number per segment. Distribution on each
segment is uniform, i.e. there is no accumulation
on any one side. The pattern of distribution along
the antenna seems to vary with species and there is
a small amount of intraspecific variation.
Multiporous plate sensillae have been assigned
various functions, e.g. mechanoreceptors (Merlin
1941), auditory receptors (Ruland, 1888), air
pressure receptors (Mclndoo 1914, 1922),
photoreceptors (Booth, 1963) and so on. More
recent work indicates an olfactory function. The
reaction of M. australica multiporous plate
sensilla to ethyl acetate (discussed later) certainly
indicates a reaction to chemicals.
Studies by Slifer, Prestage and Beams (1959)
showed that some of the peg sensilla on the
flagellum of grasshoppers had numerous fine
pores in their cuticular walls and these could be
demonstrated by soaking the antenna in 0.5<^^o
methylene blue. Sections revealed fine nerve
fibres running to each pore. Further work has
shown these to be olfactory receptors. Electro-
physiological work by Lacher and Schneider
(1963) and Lacher (1964) has shown that the
multiporous plate sensilla on honey bee {Apis
mellifera) antennae are olfactory. Slifer and
Sekhon (1961) demonstrated fine pores in the
cuticle of these, but not the fine fibrils. Slifer
(1969) felt that further examination will probably
reveal fine fibrils passing to these pores.
Multiporous plate sensilla in aphids examined by
Slifer, Sekhon and Lees (1964) differ in structure
from those of the honey bee partly in the
possession of an inner and outer cuticular layer.
Dedrites pass singly or in groups through pores in
the inner cuticular layer and enter a fluid filled
chamber between the two cuticular layers in
which they branch repeatedly. The fluid filled
chamber has some importance in interpreting
results obtained when I treated M. australica
females with ethyl acetate (discussed later). The
outer cuticular layer of aphid multiporous plate
sensilla are penetrated by numerous fine pores
each supplied with fine fibrils. It could not be
determined if these fine fibrils were connected to
the dendrites, but Slifer, Sekhon and Lees (1964)
felt this was probably the case. Freshly moulted
specimens admitted crystal violet dye through
these pores. The presence of fine filaments
terminating in the pores plus the staining via these
pores were taken to indicate an olfactory function
for aphid multiporous plate sensilla,
Slifer (1969) examined the sense organs on the
antennae of the pteromalid wasp Nasonia
vitripennis (Walker). When treated with 0,5%
crystal violet dye the stain rapidly entered the
multiporous plate sensillum and examination
showed the presence of numerous fine pores in
the surface, in cross section these multiporous
plate sensilla were seen to have an inner
membrane similar to the multiporous plate
sensilla of aphids. The inner cuticle lies just above
two shelf-like invaginations of the cuticle. She
noticed a large group of dendrites just below the
proximal end of the inner membrane and
presumed that these passed through the inner
membrane, as in the aphid, and sent filaments
into the pores in the outer surface.
The multiporous plate sensilla of M. australica
resemble those of Nasonia vitripennis in overall
shape and appearance. In cleared specimens, two
longitudinal cuticular invaginations can be seen
on either side of the multiporous plate sensilla as
in N. vitripennis, I have not carried out
histological investigation of the M. australica
multiporous plate sensilla to confirm the presence
of an inner membrane but am fairly certain there
is one. Attempts to demonstrate pores in the outer
wall using 0.5% crystal violet solution as used by
Slifer (i960, 1969) were not successful even in
freshly moulted specimens and SEM
investigations did not reveal pores either. Slifer,
Sekhon and Lees (1964) found that the plate
organs of aphids showed crystal violet
penetration when fixed four hours after the final
moult but no penetration at 24 hours and 48
hours after final moult. They suggested that the
minute pores in the older specimens might be
rimmed with a waxy or hydrophobic material
which prevents entry of the dye. Locke (1964)
when discussing the formation of the insect
cuticle states that the secretion of the endocuticle
and the secretion of wax can occur concurrently
and extend through the intermoult period.
Therefore, it is possible that wax secretions
constrict the pores after moulting. PI. 8a-c show
366
MEMOIRS OF THE QUEENSLAND MUSEUM
the thin walled basiconic pegs of the grasshoppers
Atractomorpha similis (Bolivar) freshly moulted
and Valanga irregularis (Walker) several hours
after moulting. The difference between the two is
thought to be a result of wax encroachment. Gold
coating would further fill the fine pores and
obscure them in older specimens.
The normal method used for preparing M.
australica for SEM examination involved killing
by immersion in 75*^0 ethyl alcohol before gold
coating. When freshly moulted specimens were
killed using ethyl acetate vapour before gold
coating the result showed multiporous plate
sensilla with a crumb-like surface which at higher
magnifications look like exudations from pores in
the surface (PI. 6b-d). It can be seen that the
surface of the antenna lacks the usual crazed
appearance of older antennae and this is thought
to be because the wax layer was absent or very
thin. Older specimens when killed with ethyl
acetate showed multiporous plate sensilla with a
blistered appearance and distribution of the
blisters matched distribution of the exudations in
freshly moulted specimens (PI. 6d). It is thought
that the exudations arise from the fluid filled
space between the outer and inner membranes of
the multiporous plate sensilla and that this is in
response to fairly high ethyl acetate concentration
being an attempt to protect the sensitive nerve
ending from a pungent vapour. The analogy is to
the mammalian nasal mucosa which produces
copious mucus in response to pungent vapours.
Barlin and Vinson (1981) investigated the
multiporous plate sensilla on the antennae of
several species of Chalcidoidea. In some cases
they also found that the presence of pores in the
outer plate was shown by exudations. Their
investigations revealed two types of multiporous
plate sensilla; Type 1 - present in both sexes and
possessing a thin outer cuticle with numerous
pores; Type 2 - present in females only and
possessing a thick outer cuticle with fewer pores.
Both types were found in all species studied
except three i.e. not all species were found to have
Type 2 multiporous plate sensilla. In M.
australica, the females appear to have only Type
1. However, my work on M. australica antennal
sense organs was carried out some time before the
appearance of the paper by Barlin and Vinson
and time does not permit a more thorough
investigation of this aspect. I have not
investigated the multiporous plate sensilla on the
male antennae.
The similarity of the multiporous plate sensillae
of M. australica to those of Nasonia vitripennis,
the presence of pores on the outer surface in
conjunction with the results of Slifer et alia on
other receptors having porous surfaces and the
results of Barlin and Vinson with various
chalcidoids suggest strongly that the multiporous
plate sensilla of M. australica are olfactory in
function. From their reaction to ethyl acetate they
are probably very sensitive.
Before discussing their function we should look
at the short basiconic capitate pegs as these
appear also to be olfactory in function.
4) Short basiconic capitate pegs (PI. 5b, iv).
These organs are absent from the antennae of
males of most species but are present on the
funicle and club of females where they
predominate on the dorsal or outer surface. They
arise from a circular, shallow, relatively broad
depression in the cuticle. Thereafter the peg
tapers into stalk which terminates in a spherical
knob, the whole resembling a champagne cork.
They occur sub-marginally on the distal portions
of the segments and are directed towards the
distal tip of the antenna.
These were not recorded on the antennae of
Nasonia vitripennis by Slifer (1969) but were
found subsequently in this species and another
pteromalid wasp Peridesmia discus (Walker) by
Miller (1972). Weseloh (1972) found them on the
antennae of the encyrtid wasp Cheiloneurus
noxius Compere. Neither gave any details of their
ultrastruciure but Miller (1972) assumed they
were not touch receptors because of their
sheltered location. Slifer, Prestage and Beams
(1957,1959) suggest that basiconic capitate pegs
may function in olfaction if they are thin walled
or in the perception of irritant substances if they
are thick walled.
When females of M. australica were killed by
immersion in 15% ethyl alcohol, SEM
examination showed no details other than a
crazed surface thought to be wax (PL 7a), Freshly
moulted females killed with ethyl acetate vapours
showed weeping from slits or rows of pores
arranged along the longitudinal axis of its capitate
tip or caput. In PL 7b-d these slits or rows of
pores can be seen quite clearly. The distal tip of
the caput was devoid of exudations and resembled
a tonsure. Around the base of the peg were
exudations resembling those of the multiporous
plate sensilla.
If one is to accept the reasoning put forward
earlier to explain the exudations from
multiporous plate sensillae then the internal
structure and the function of the short basiconic
capitate pegs may be the same as the multiporous
plate sensilla. They are undoubtedly olfactory in
function. Therefore there are two
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
367
morphologically different olfactory sensilla on
the antennae of all females and some males.
Schneider and Steinbrecht (1968) when
discussing insect olfactory sensilla indicated there
are two physiological types of olfactory cells —
odour specialists and odour generalists. The
former respond to biologically important odours,
e.g. sex attractants, warning or specific food
odours. Both types may be found in the one
sensillum and this has been demonstrated in the
multiporous sensilla of the honey bee Apis
mellifera.
In Melittobia, evidence suggests specific and
sexually different chemical signals. The sources
are the male scape gland and circumstantial
evidence indicates mandibular glands in both
sexes. These would suggest the presence of odour
specialist, olfactory cells. Electro-physiological
work would be required to identify the presence
and location of these cells but some speculation is
possible.
In the males of most species the antennae lack
short basiconic capitate pegs. In females these are
located mostly on the upper surface of the
antennae which is the surface applied to the inner
lining of the male scape groove or cup containing
the dermal gland. The short basiconic capitate
pegs therefore might contain odour specialist
olfactory cells for perception of a male
pheromone. Females are able to detect males
from a distance, e.g. when a male is placed in with
a group of inactive virgin females the latter
immediately become active and move fairly
directly towards the male. Hermann (1971)
mentions male calling in M. chalybii (= M.
australica). It was noticed in my colonies of M,
australica that males walk about with their scapes
raised laterally and flagellar segments extended so
that the tip of the scape groove was open. They
also stand around in this pose. It could be that
they are exposing their scape gland to attract
females. The rearing jars are much larger than the
host cell or puparium and it is difficult to imagine
the need for such a system in the confines of a
host cocoon or puparium.
The distribution of multiporous plate sensilla
varies in males. In the ha'waiiensis and assemi
groups they occur on the club segments only, but
in the acasta group they occur on all flagellar
segments. During courtship M. australica females
were noticed to open their mandibles. Initially I
thought this to be the signal for the female’s
readiness to copulate thus inducing the male’s
finale. However, van den Assem does not agree
(Pers. comm. 1980). Mention has been made
previously of virgin females without males
standing with open mandibles. When provided
with a dead male pupa females located it and
again stood with open mandibles. It was argued
that mandibular glands may be the source of
female scent and if this is so then opening
mandibles during courtship probably means some
chemical input by the female. If virgin females
call by mandibular glands then the only olfactory
receptors in most males are the plate organs and
these would contain the odour specialist olfactory
cells. During courtship of species in the
hawaiiensis and assemi groups the male position
is such that his clubs are in close proximity to the
female’s mandibles when open. In these groups
only male club segments bear plate organs.
During courtship of the acasta group the male
does not stand so far forward and his funicular
segments would be in contact with her open
mandibles. This is thought to have some bearing
on retention of plate organs on the funicle as well
as the club segments in the acasta group males.
ACKNOWLEDGMENTS
This paper is taken from my M.Sc. thesis
submitted to the University of Queensland in
1982. My superviser, Dr Elizabeth Exiey,
University of Queensland, was extremely helpful
in providing constructive comments and editorial
remarks. Dr T. Woodward, University of
Queensland and Dr G. Gordh, University of
California, Riverside, as examiners provided
corrections and advice towards publication of the
thesis.
Plates 5b-d, 6a-d, 8a-c were provided by Mr J.
Hardy, Electron Microscope Department,
University of Queensland and Plates 2a-b, 3a,
4b-c were provided by Dr R. Raven, Queensland
Museum. Sections of the male scape of Melittobia
australica (Plates 3b-c) were cut by Mr N. Hall,
Queensland Museum and photographed by Dr L.
Cannon, Queensland Museum. My technician,
Miss Gudrun Sarnes, was of great help checking
manuscripts and numbering plates. The typists
whose patience I have tried severely were Miss P.
Tinniswood and Miss E. Proberts of the
Queensland Museum. My wife Judith provided
assistance with manuscript checking and plate
assembly.
Special thanks are due to Dr J. van den Assem,
University of Leiden, Holland. We have
corresponded freely since 1974 and he has been of
the greatest assistance with notes from his
ethological studies on Melittobia.
368
MEMOIRS OF THE QUEENSLAND MUSEUM
LITERATURE CITED
Assem, J. van den and F.A. Putters, 1980.
Patterns of sound produced by courting
chalcidoid males and its biological
significance. Entomologia exp. appl. 27:
293-302.
H.A.J. IN DEN Bosch and E. Prooy, 1982.
Melittobia courtship behaviour: a
comparative study of the evolution of a
display. Neth. J. Zool. 32: 427-71.
Barlin, M.R. and S.B. Vinson, 1981.
Multiporous plate sensilla in the antennae of
the Chalcidoidea (Hymenoptera). J. Insect
Physiol, and Embryol. 10: 29-42.
Booth, C.O., 1963. Photokinetic function of
aphid antennae. Nature, Lond. 197: 265-6.
Dahms, E.C., 1983a. Revision of the genus
Melittobia (Hymenoptera : Eulophidae) with
the description of seven new species. Mem.
Qd Mus. 21: 241-306.
1983b. A review of the biology of species in the
genus Melittobia (Hymenoptera :
Eulophidae) with interpretations and
additions using observations on Melittobia
australica. Mem. Qd Mus. 21: 307-30.
Goodpasture, C., 1975. Comparative courtship
behaviour and karyology in
Monodontomerus (Hymenoptera : Tory-
midae). Ann. ent. Soc. Am. 68: 391-7.
Gordh, G. and P. de Bach, 1978. Courtship
behaviour in the Aphytis lignanensis group,
its potential usefulness in taxonomy, and a
review of sexual ibehaviour in the parasitic
Hymenoptera (Chalcidoidea : Aphelinidae).
Hilgardia 46: 37-75.
Hermann, L.D., 1971. The mating behaviour of
Melittobia chalybii (Hymenoptera :
Eulophidae). Unpublished Thesis, Univ.
Georgia, U.S.A., 1971, 52 pp.
Houston, T.F., 1975. A revision of the
Australian Hylaine bees (Hymenoptera :
Colletidae) 1. Introductory material and the
genus Heterapoides Sandhouse,
Gephyrohylaeus Michener, Hyleoides Smith,
Pharohylaeus Michener , Hem irh iza
Michener, Amphylaeus Michener and
Meroglossa Smith. Aust. J. Zool., Suppl.
Ser., (36): 1-135.
Lacker, V., 1964. Elektrophysiolgische
Untersuchungen an einzelnen Rezeptoren f ur
Geruch, Kohlendioxyd, Luftfeuchligkeit und
Temperatur auf den Antennen der
Arbeitsbiene und der Drohne (Apis mellifica
L.) Z. vergl. Physiol. 48: 587-623.
and D. Schneider, 1963. Elektrophysio-
logischer Nachweis der Riechfunktion von
Porenplatten (sensilla placodea) auf den
Antennen der Drohne und Arbeitsbiene (Apis
mellifica L.) Z. vergl. Physiol. 47: 274-8.
Mcindoo, N.E., 1914. The olfactory sense of the
honey bee. J. exp. Zool. 16: 265-346.
1922. The auditory sense of the honey bee. J.
Comp. Neurol. Psych. 34: 173-99.
Melin, D., 1941. The function of pore-plates in
Hymenoptera. Zool. Bidr. Upps. 20: 304-44.
Miller, M.C., 1972. Scanning electron
microscope studies of the flagellar sense
receptors of Peridesmia discus and Nasonia
vitripennis (Hymenoptera : Pteromalidae)
Ann. ent. Soc. Am. 65: 1119-23.
Picard, F., 1923. Recherches biologiques et
anatomiques sur ^"Melittobia acasta'^ Walk.
(Hymenoptere chalcidien). Bull. biol. Fr.
Belg. 57: 469-508.
Ruland, R., 1888. Beitrage zur Kenntnis der
antennalen Sinnesorgane der Insekten. Zeit.
wiss. Zool. 46: 602-28.
Schneider, D., 1969. Insect olfaction:
deciphering system for chemical messages.
Science, NY. 163: 1031-7.
and R.A. Steinbrecht, 1968. Checklist of
insect olfactory sensilla. Sym. zool. Soc.
Lond. 23: 279-97.
Slifer, E.H., 1960. A rapid and sensitive method
for identifying permeable areas in the body
wall of insects. Ent. News 71: 179-82.
1969. Sense organs on the antenna of a
parasitic wasp, Nasonia vitripennis
(Hymenoptera, Pteromalidae). Biol. Bull,
mar. biol. Lab., Woods Hole 136: 253-63.
J.J. Prestage, and H.W. Beams, 1957. The
fine structure of the long basiconic sensory
pegs of the grasshopper (Orthoptera :
Acrididae) with special reference to those on
the antenna. J. Morph. 101: 359-81.
1959. The chemoreceptors and other sense
organs on the antennal flagellum of the
grasshopper (Orthoptera : Acrididae). J.
Morph. 105: 145-66.
and S.S. Sekhon, 1961. Fine structure of the
sense organs on the antennal flagellum of the
honey bee, Apis mellifera Linnaeus. J.
Morph. 109: 351-81.
S.S. Sekhon, and A.D. Lees, 1964. The sense
organs on the antennal flagellum of aphids
(Homoptera), with special reference to the
plate organs. Q. Jl micros. Sci. 105: 21-9.
Weseloh, R.M., 1972. Sense organs of the
hyperparasite Cheiloneurus noxius
(Hymenoptera : Encyrtidae) important in
host selection processes. Ann. ent. Soc. Am.
65: 41-6.
370
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
Melittobia australica male scape, AgN 03 stain, Euparal slide
mount, X 900
a) Lateral view.
b) Ventral view.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
371
b
372
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
Melittobia ausiralica male scape.
a) Dorsal view, NaOH cleared, Euparal slide mount,
X 340.
b) Ventral view, showing transverse arm of gland,
SEM, X 340.
c) Cuticular surface over gland, SEM, x 2,000.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
373
Plate 2
374
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 3
Melittobia australica male scape.
a) Cuticular surface over gland showing pores, SEM,
X 5,200.
b) TS male scape just proximal of scape attachment —
section through transverse arm of gland, Euparal
slide mount, x 650.
c) TS more proximal region of scape-section through
longitudinal area of gland, Euparal slide mount, x
700.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
375
Plate 3
a
376
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 4
Melittobia australica male scape and setae on legs.
a) TS longitudinal area of scape showing cuticular
invaginations to support gland, Euparal slide
mount, X 2,000.
b) Dense setal tuft on ventral fore-trochanters, SEM,
X 1,000.
c) Seta in mid-femoral fringe, SEM, X 4,000.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
378
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 5
Melittobia australica male, female antennae.
a) Male club, AgNO, Stain, Euparal slide mount.
b) Female club, SEM, x 900.
i) long thin unfluted setae.
ii) tapering fluted seta.
iii) multiporous plate sensillum.
iv) short basiconic capitate peg.
c) Male club, segments 2 and 3, SEM, x 2,000.
d) Male tapering fluted setae, SEM, x 2,800.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
379
Plate 5
a b
c
d
380
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 6
Melittobia australica male flagellum and female sensillae.
a) Male flagellum, SEM, x 600.
b) Distal female club after exposure to ethyl acetate.
SEM, X 1,300.
c) Multiporous plate sensillum of freshly moulted
female after exposure to ethyl acetate, SEM, x
10 , 000 .
d) Multiporous plate sensillum of older female after
exposure to ethyl acetate, SEM, x 5,500.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
381
Plate 6
c
d
382
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 7
Melitiobia australica female short basiconic capitate pegs,
a) Older female killed by immersion in ethyl alchol,
SEM, X 13,000.
b-d)Freshly moulted female after exposure to ethyl
acetate, SEM, (b x 10,000; c x 18.000; d x
13,000).
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
383
Plate 7
a b
c
d
384
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 8
Grasshoper basiconic pegs.
a) Freshly moulted Atractamorpha simitis, SEM, x
6 , 000 .
b) Freshly moulted Atractamorpha similis, SEM, x
21 , 000 .
c) Several hours after moulting Valanga irregularis,
SEM, X 7,500.
DAHMS: ANTENNAL SENSE ORGANS OF MELITTOBIA
385
C
Mem. QdMus. 21(2): 387—80. [1984]
A NEW SPECIES OF APTENOCANTHON MATTHEWS FROM NORTH
QUEENSLAND. (COLEOPTERA : SCARABAEIDAE : SCARABAEINAE)
R.L Storey
Department of Primary Industries,
MP.O. Box 1054, Mareeba, Queensland
ABSTRACT
Aptenocanthon monteithi sp. nov. is described from the Atherton Tableland area in
northern Queensland. The nearest relatives are from mountains in eastern New South Wales.
INTRODUCTION
The dung beetle genus Aptenocanthon was
erected by Matthews (1974) for the species A.
hopsoni Carter (formerly Panelus and a second
new species, A, rossi. Both species were known
only from wet, high altitude localities in central
New South Wales, Barrington Tops and Mt
Wilson respectively. Both species were poorly
known, but A. hopsoni has since been taken in
numbers at Barrington Tops, from altitudes of
500 m to 1400 m, in both dung baited pitfall traps
and sieved litter (T.A. Weir, pers. comm.), and
A. rossi has been recorded at Mt Irvine at 750 m
at dung baited pitfall traps (Williams and
Williams 1982).
It is thus with some interest that a new species
was taken recently on top of the Bellenden-Ker
Range in north Queensland, 1900 km north of the
previous records for the genus. The specimens
were taken by an expedition, organized jointly by
the Queensland Museum and the Earthwatch
Organization, to study the change in insect fauna
with altitude on Mt Bellenden-Ker, a locality
noted for relict and other interesting species in
both the botanical and insect world.
Subsequently, further specimens of the same
species were taken on mountain areas on the
Atherton Tableland proper, namely the Mt Fisher
and Mt Haig areas, 45 km S.W. and 40 km N.W.
from the original Bellenden-Ker locality
respectively.
Genus Aptenocanthon Matthews
Aptenocanthon Matthews 1974, Aust. J. ZooL,
Supp. Ser. No 24, pp. 93-97. Type species
Panelus hopsoni Carter, 1936.
Aptenocanthon monteithi sp. nov. (Figs. 1 and 2)
Holotype: Queensland Museum T8592
Bellenden-Ker Range, N. Qld., Summit TV Stn.,
1560 m, 25-31. X. 1981, Earthwatch/Qld.
Museum, QM Berleseate No, 372, 17.16S,
145. 51E, Rainforest. Sieved litter.
Allotype: Queensland Museum T8593 $,
Bellenden-Ker Range, N. Qld., Summit TV Stn.,
1560 m, 25-3 l.x. 1981, Earthwatch/Qld.
Museum, Pitfall trap in rainforest.
Paratypes (34): Bellenden-Ker Range, Summit
TV Stn., 1560 m, 17.x-5.xi.l981,
Earthwatch/Qld. Museum (5 ^ 6 , 4 ■??), same
data 1-7. xi. 1981, (1 S), same data 25-31. x. 1981
(3 =3). Mt Bartle Frere, 0.5 km north of South
Peak, 1500 m, 6-8. xi. 1981, Earthwatch/Qld.
Museum, (4 Mt Bartle Frere, NW/Centre
Peak ridge, 1400-1500 m, 7-8. xi. 1981,
Earthwatch/Qld, Museum, (3 S S). Mt Fisher, 7
km SW Millaa Millaa, N. Qld., 1050-1100 m,
27-29. IV. 1982, Monteith, Yeates and Cook, (1
3, 1 ^), 26 km up Tinaroo Ck Rd., via Mareeba,
N. Qld. Il.xi.l982, Morgan, Brown and Storey,
(1 2 ), same data 10-30.xi.i982, (2 9$), same data
1-23. xii. 1982, (1 2 ), same data 23.xii,1982-
12.i.l983, (2 ?9), same data I2-28.i. 1983, (1 s),
same data 28.l-16.ii.1983, (1 9). (Paratypes
placed in Queensland Museum, ANIC Canberra,
South Australian Museum, Queensland D.P.I.,
Coll. R.I. Storey, Mareeba, H.F. Howden,
Ottawa, A. Walford-Huggins, Julatten, P.
Allsopp, Toowoomba, G. Williams, Taree).
Total length 3.3-4. 1 mm. colour black, legs
reddish brown antennae yellow brown.
Male
Head: Clypeal teeth small, close together, U-
shaped between, rest of margin feebly sinuate to
genal angles which are angulate, margin very
feebly beaded. Surface nitid, densely punctuate
with moderate simple punctures, effaced along
edge of clypeus, glabrous. Dorsal part of eyes
small, about 4 facet rows wide, separated by
388
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 1: Aptenocanthon monteithi sp. nov., male paratype.
about 8-10 eye widths, canthus incomplete.
Pronotum: Anterior angles quadrate, feebly
projecting, lateral angles very obtuse, rounded,
posterior angles obtuse. Pronotal surface smooth,
nitid, punctate with moderate simple punctures
which are effaced along lateral margin and
posterior angles, reduced in centre of disk,
glabrous. Lateral edge rounded to deflexed
portion, finely margined. Elytra: Striae on disk
effaced, almost undetectable, impunctate,
intervals feebly convex, very finely punctate,
surface feebly shagreened, glabrous. Surface
deflexed outside 7th stria, forming
pseudepipleura about 2/3 length of elytra, the
edge beaded with a slightly raised nitid area
between it and the 7th stria, with two fine striae
between edge of pseudepipleura and the feebly
sinate epipleura. Hind Wing: Entirely absent.
Sterna: Mesosternum with a few large superficial
umbilical punctures on edges. Median lobe of
metasternum nitid, impunctate, lateral lobes with
numberous large, shallow umbilical punctures.
Legs: Fore tibia with inner edge broadly concave,
outer edge with three short teeth on apical 1/3,
distal edge straight, transverse, with inner apical
angle produced into a broad inwardly projecting
truncate lobe, tibal spur short, triangular. Middle
and hind legs unmodified. Abdomen: Pygidium
STOREY: NEW SPECIES OF APTENOCANTHON
389
Strongly convex, smooth nitid, scattered very fine
punctures, glabrous. Each slernite with several
large punctures near ends, in a shallow depression
which is deepest on 6th. Aedeagus with parameres
asymmetrical, as in Fig. 2.
Female
Fore tibia with inner apical angle not produced
into a truncate lobe, tibial spur and teeth on outer
edge slightly larger. Abdominal sternites
expanded. Otherwise similar to male.
COMMENTS
The specimens from the Bellenden-Ker Range
were all taken above 1400 m in a habitat described
by Tracey and Webb (1975) as simple Microphyll
Vine-Fern Forest grading into thicket at the
absolute summit. It was not taken at lower
altitudes at this locality even though sites at 5,
100, 500 and 1054 m were collected intensively.
However, the two additonal localities are at lower
attitudes, in the vicinity of 1000 m. Individuals
were taken in pitfall traps, dung baited traps and
in sieved litter.
The species is named for Dr G.B. Monteith
who was involved with the organizing of the
Earthwatch expedition and has contributed much
to the knowledge of Australian Scarabaeinae.
The following adaptations of Matthews (1974)
key should aid in separating the three species:
1) Pygidium with a strong transverse basal
groove; elytral striae distinct, with prominent
punctures on intervals; central N.S.W 2
Pygidium without a basal groove; elytral
striae almost obsolete, intervals very finely
punctate; north Queensland
monteithi sp. nov.
2) Elytral intervals flat, glabrous; lateral elytral
Carina sharply defined basally; outer edge of
epipleura strongly sinuate hopsoni Carter
Elytra! intervals convex, densely setose;
lateral elytral carina feeble, broadly
rounded; epipleura normal.... rossi Matthews
DISCUSSION
Some problems were encountered in placing
monteithi within the genus Aptenocanthon.
Specimens of both A. Aops-o/?/ and A. rossi were
available for comparison. It agrees with these
species in size and general appearance, secondary
sexual characters, shape of mentum and labial
palps, size of eyes and flightless condition.
Differences include shape of the lateral edge of
the pronotum, shape of pseudepipleura, nine as
opposed to eight elytral striae, and lack of
pygidial sculpturing. It is this author’s opinion
that grouping the three species in one genus is
more useful than creating a new genus as they are
closer to each other than to related genera. The
lack of basal pygidial groove in monteithi is
interesting, as it is a feature of Aptenocanthon
and related genera — the Australian Tesserodon
Hope and the New Caledonian Onthobium
Reiche, though in the Australasian Ignambia
Heller and New Zealand Saphobius Sharp it is
lacking. The Australian genus Lepanus Balthasar
is similar in having variable pygidial sculpturing
and is more heterogeneous than Aptenocanthon
as envisaged here.
Australian dung beetles of the tribe Scarabaeini
all fall within the subtribe Canthonina, which
though found almost worldwide has a basically
southern distribution, being best represented in
the neotropical region. Australian species make
up 14^0 of the world species diversity (Matthews,
1974). Australian Canthonina can be further
divided into two groups — those with simple
claws and pseudepipleura and those with dentate
or subdendale claws and without pseudepipleura.
Aptenocanthon falls within the first group,
usually referred to as the mentophilines.
Matthews (1974) considered this the most
primitive of the two groups, both structurally and
in behaviour, with ball-rolling being unknown in
the group.
The mentophiline group shows a basic
Gondwanan distribution, with genera in
Australia, New Zealand and New Caledonia. The
genus Ignambia also occurs in New Guinea, as
well as Australia and New Caledonia (Howden,
1981). Scarabaeinae as a whole occur in warm
temperate to tropical climates. Some genera in
Australia are southern {Aulacopris White and
Cephalodesmius Westwood) and others such as
Tesserodon and Ignambia are essentially
tropical. Amphistomus Landsberge has a
complex of species along the eastern and northern
coasts. The situation in Aptenocanthon, with
such widely separated flightless mountain top
species is unique in Australian Scarabaeini,
though a flightless species of Aulacopris is now
known to occur in north Queensland and will be
described in a subsequent paper. This distribution
in flightless species of insects with southern
genera being found on north Queensland
mountain tops has been recorded in other
families: Hacker iella (Peloridiidae) and
Kumaressa (Aradidae) in Hemiptera and
Lissapterus (Lucanidae) in Coleoptera. Recent
390
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 2: Aedeagus, A. monteithi sp. nov.
publications by Monteith (1980) and Kikkawa et
al. (1981) have given summaries of the
phenomenon and its importance to studies in
Australian biogeography.
The discovery of A. monteithi gives further
support to the view that high mountain tops in
north Queensland, and the Bellenden-Ker range
in particular, are major refugia for wet area
plants and insects in Australia.
ACKNOWLEDGMENTS
The author wishes to thank Mr Geoff
Thompson of the Queensland Museum for the
drawing in Figure 1 , Dr Geoff Monteith,
Queensland Museum and Mr Tom Weir of
CSIRO, Canberra for helpful comments on the
manuscript and Mr G. Williams of Taree for the
loan of specimens. The Earthwatch Organization
rates thanks for their assistance in funding of the
field work involved.
LITERATURE CITED
Howden, H.F. 1981. Zoogeography of some
Australian Coleoptera as exemplified by the
Scarabaeoidea. in A. Keast (ed.). ‘Ecological
Biogeography of Australia’. (The Hague: W.
Junk publishers) pp. 1009-35.
Kikkawa, J., G.B. Monteith and G. Ingram,
1981. Cape York Peninsula: Major region of
faunal intercharge, in A. Keast (ed.).
‘Ecological Biogeography of Australia’. (The
Hague: W. Junk publishers) pp. 1697-742.
Matthews, E.G. 1974. A revision of the
scarabaeine dung beetle of Australia IT Tribe
Scarabaeini. Aust. J. Zool. SuppL Ser. 24:
1 - 211 .
Monteith, G.B. 1980. Relationships of the
genera of Chinamyersiinae, with description
of a relict species from mountains of north
Queensland (Hemiptera : Heteroptera :
Aradidae). Pac. Insects 21: 275-85.
Tracey, J.G. & L.J. Webb, 1975. ‘Key to the
vegetation of the humid tropical region of
North Queensland, plus 15 maps at 1:100,000
scale.’ (Canberra; CSIRO Division of Plant
Industry).
Williams, G.A. and T. Williams, 1982. A
survey of the Aphodiinae, Hybosorinae and
Scarabaeinae (Coleoptera : Scarabaeidae)
from small wet forests of coastal New South
Wales. Part 1: Nowra to New Castle. Aust.
ent. Mag. 9 (4): 42-8.
Mem. QdMus. 21(2): 391—99. [1984]
SPAWNING OF THE AUSTRALIAN LUNGFISH, NEOCERATODUS FORSTERI
(KREFFT) IN THE BRISBANE RIVER AND IN ENOGGERA RESERVOIR,
QUEENSLAND.
A. Kemp
Queensland Museum
ABSTRACT
The Australian lungfish, Neoceratodus forsteri (Krefft) breeds annually between mid-
August and December. The onset of oviposition is not related to rainfall, temperature, pH or
dissolved content of the water in which they live. Lungfish begin to spawn when daylength
has been increasing for 6-1 1 weeks, provided that suitable weeds are growing, and the rate of
flow of the water may also have an effect on the site chosen for spawning.
INTRODUCTION
Caldwell (1884) was the first to report finding
eggs of the Australian lungfish, Neoceratodus
forsteri (Krefft), in the Burnett River, 14 years
after Krefft (1870) had described the adult fish
from the Mary River. Semon, who studied the
development of N. forsteri (1893), found eggs in
shallow water amongst weeds in the Boyne River,
a tributary of the Burnett (1899:96). Eggs are
deposited among growing water weeds or on the
sides and bottom of submerged logs (Illidge
1892). Plants important for egg laying in the
Burnett River are Hydrilla verticillata, Vallisneria
spiralis and Nitella sp. (Bancroft 1911). Spawn
has also been found on the roots of the water
hyacinth, Eichornia crassipes, in Enoggera
Reservoir, Brisbane (Bleakly, per. comm., 1969),
from “weeds” in a tributary of the Burnett River
(Grigg 1965) and on the submerged roots of
Callistemon saligna growing beside the Brisbane
River (Kelly, per. comm., 1977). Spencer (1925)
states that eggs are laid separately amongst
vegetation, but not attached to it, and that they
finally lie on the mud.
There is only one published report on the
breeding behaviour of N. forsteri in the wild
(Grigg 1965). The breeding season has been
recorded as August-October in Enoggera
Reservoir (Bleakly, cited by Grigg 1965),
September in the Boyne River (Semon 1899),
August-November in Enoggera Reservoir (Kemp
1977), September-October (Spencer 1926),
November-December (Illidge 1893), August-
October in the Burnett River (Bancroft 1911 and
1928) and August in a tributary of the Burnett
River (Grigg 1965). Factors which influence
spawning are not known.
This work was undertaken to determine the
times and places of spawning of N. forsteri in a
river and a lake environment in southeast
Queensland, and to discover any possible
environmental regulation of breeding. In this
respect the Australian lungfish was compared
with the African and South American species,
Protopterus annectens (Owen), Protopterus
aethiopicus Heckel and Lepidosiren paradoxa
Fitzinger.
MATERIALS AND METHODS
Information on breeding, stages of eggs
collected and weeds used for oviposition were
recorded from Enoggera Reservoir from
1971-1973 and from the Brisbane River from
1978-1981. Rainfall data for these years was
obtained from the Bureau of Meteorology in
Brisbane, Queensland.
Information on hours of daylight was obtained
from the records of the Department of Mapping
and Surveying, Brisbane, Queensland.
Observations were made during the period July
1979 to November 1979 at two breeding sites in
the Brisbane River: one was a bed of Vallisneria
spiralis and the other, submerged roots of
Callistemon saligna 1 km downstream.
Temperature and dissolved oxygen were
measured in the field between 11 am and 1 pm.
Water samples were taken to measure pH in the
laboratory. Eggs were collected from weeds or
roots in the area from which the samples were
taken. Plants were identified from Aston’s (1973)
392
MEMOIRS OF THE QUEENSLAND MUSEUM
description. Further observations on temperature
and dissolved oxygen in relation to spawning were
made during the period May 1980 to January
1981.
Eggs were kept in insulated containers and
assessed for their stage of development in the
laboratory, 1-2 hours after collection. The onset
of oviposition was determined by subtracting the
age of the oldest eggs found from the date of
collections, e.g. minus 1 day if the oldest eggs
were cleavage stages or minus 6 days if the oldest
were neurulae of stage 18. Ages were based on
times of development of eggs maintained in the
laboratory at 18-22°C, temperatures comparable
to those of the river at the time of collection
(Kemp 1981).
RESULTS
Breeding Behaviour
i) Plants in the breeding areas and their
potential as oviposition sites. A list of plants in
the lake and river breeding areas is given in Table
1 , with observations on the use made of the weeds
for spawning. The weeds were present at all times
except for Eichornia crassipes, Hydrilla
verficillata and Potamogeton Javaniciis, all of
which die off in winter.
Weeds used consistently for spawning are E.
crassipes (in river and lake), Vallisneha spiralis
and submerged roots of Callistemon saligna (river
only). Fewer eggs were found on H. verticillata,
Potamogeton perfoliatus or Nitella sp. Use of
Ceratophyllum sp. or filamentous algae was
incidental and occurred only when these weeds
grew amongst C. saligna and V. spiralis
{Ceratophyllum ) or coated the C. saligna roots
(algae).
TABLE 1: Utilisation of Weeds for Spawning in
Enoggera Reservoir and in the Brisbane
River.
Species of Weed
Eichornia crassipes
Hydrilla verticillata
Vallisneria spiralis
Callistemon saligna
(submerged roots)
Nitella sp.
Potamogeton crispus
Potamogeton perfoliatus
Potamogeton javanicus**
Ceratophyllum sp.***
Nymphoides indica
Nymphaea capensis
Nymphoea flava
Ludwigia peploides
Brochiaria mutica
Rumex bidens
Sedge - c
Filamentous algae*** - *
* used for spawning
c not used
weed has not been found
? weed loo deep to sample
** breeding behaviour observed but no eggs found
*** growing with or on C. saligna roots or K spiralis
plants
ii) Oviposition in the lake. Eggs were always
found attached to the roots or submerged floats
of E. crassipes. Eggs have not been found on or
among any other weeds in this locality, or lying
free at the bottom of the lake, as measured by
dredging samples or by diving. H. verticillata,
used by fish in the Burnett River for spawning
(Bancroft 191 1 and 1928), also occurs in the lake
but it was too deep to be easily sampled.
During the breeding season of N. forsteri, air
breathing is frequent, and is accompanied by a
distinct loud burp, made in the air with the lips
clear of the water. During two periods of
observation groups of fish appeared to be
responding to each other. On the first occasion,
one fish in the centre of a group ‘sounded’, then a
fish to one side, then a fish to the other side, then
the central fish, and finally one ahead of the first
fish. On the second occasion one fish after
another breathed air, in no particular order,
along a weed bank about 20 m in length. The fish
w-ere spaced at 2-4 m intervals, and several
minutes elapsed between each breath. During the
period of observation (2 hours during the day),
individual fish breathed at regular intervals of
about 20 minutes. Pairs of fish also perform
circling movements at the surface of the water
close to weed banks.
Fish manoeuvre into the root mass of E.
crassipes to spawn, and may lay eggs deep within
the mass of roots as well as on the edges.
Oviposition was observed once at 11 am in the
lake. The female turns on her side when laying
eggs and the male, entwined around her, fertilises
the eggs as they emerge. Eggs less than 3 hours old
(uncleaved, or in stage 1 or 2) were often found
during daytime collecting trips.
Eggs laid at one time are found in a
circumscribed area of the roots, for example all at
the top. It is possible to distinguish individual
clutches by their different ages, i.e. a set of late
neurulae compared with a set of early cleavage
stages. Eggs are laid over areas of 1-5 sq.m, on
the roots and occasionally on partly submerged
floats of the hyacinth, in varying positions from
near the surface to a depth of one metre. They are
Lake River
* *
9 *
_ *
_ *
_ *
c
c *
c
c
c c
c
c
c
KEMP: SPAWNING OF NEOCERATODUS FORSTERI
393
attached quite firmly by the outer jelly layer
which is sticky when first laid. Eggs were usually
placed singly, or occasionally in pairs.
Fish performing frequent air breathing and
circling movements have been observed in beds of
Potamogeton javanicus in the lake, but eggs have
not been found on this weed despite extensive
searches. Adult lungfish have also been seen at
night in para grass, Brachiaria mutica, which
grows thickly in shallow water beside the shore,
but again no eggs have been collected from this
plant.
iii) Oviposition in the Brisbane River. Eggs are
laid on the submerged roots of Callistemon
saligna, with or without a covering of filamentous
algae, in beds of Vallisneria spiralis, mixed stands
of V. spiralis and Hydrilla verticillata, or on the
upper parts of the alga Nitella sp. which also
grows with V. spiralis. Eggs have also been found
on fronds of Ceratophyllum sp. growing amongst
C. saligna roots, and on E, crassipes.
Eggs were not found on sedge or on Ludwigia
peploides which occur in shallow water near
consistently used V. spiralis beds, nor on the H,
verticillata growing in deep water nearby. Also
fish did not appear to lay on V. spiralis growing in
a substrate of fine black mud, but used weeds
rooted in fine sand or gravel. In 1980 and 1981,
logging operations upstream resulted in a deposit
of silt throughout the V. spiralis bed, and
although this was fine, it was not as fine as the
mud and the fish continued lo lay eggs.
Nyrnphoides indica, Nymphaea flava and
isolated fronds of Ceratophyllum are also ignored
by the fish, as is Potamogeton crispus which
occurs in deeper or faster flowing water.
Eggs may be found close to the surface to a
depth of 1.5 m. They occur on leaves or on the
partly exposed upper roots of V. spiralis,
sometimes partly buried in the substrate, and on
any part of the C. saligna root mass, on or under
the mat of filamentous algae if this is present.
Sometimes the eggs are laid so high on the root
mass that a drop in the water level after laying
leaves the eggs exposed.
During 1980 E. crassipes was plentiful in the
river and the fish used it for spawning. In one
area they seemed to prefer it to C. saligna nearby
laying eggs exclusively on the water hyacinth
when it grew sufficiently dense (Fig. 1), and in
another locality they used both. River fish often
laid eggs in clusters of 4-14 eggs (i.e. close
together but not touching) on E. crassipes.
In localities containing V. spiralis or C.
saligna, different clutches of eggs could not
usually be distinguished. Eggs of various ages
appeared to be randomly distributed.
On one occasion only, in 1978, eggs were found
lying free on the bottom of the river, in shallow,
weed free areas amongst the V. spiralis plants.
Many of these eggs were dead and all were
exposed to sunlight and to higher temperatures
than the eggs which were hidden in the V. spiralis
leaves. The weed free areas did not look like nests
and no adults were in attendance.
Other Observations
Both the lake and the river have a permanent
inflow of water. Enoggera Reservoir is situated
on a creek fed by springs in the D’Aiguilar Ranges
to the west of Brisbane. Although the creek is
reduced in times of drought, it is permanent.
Water levels in the Brisbane River fluctuate
because it supplies water for Brisbane, but the
flow is continuous.
In both lake and in river, fish lay eggs in areas
with a slow or moderate current of water, free of
floating debris. Eggs are not found in still water,
even if suitable weeds are available.
Fish do not guard the eggs in any of the
spawning localities.
Suitable weeds for spawning are always present
in the river. This was not always the case in the
lake as E. crassipes dies off in winter. In 1973, as
there was a mild winter and the weeds did not die
off, so suitable weeds were present throughtout
the year.
The timing of oviposition
Data on the duration of breeding by fish in the
lake and in the river (calculated from the age of
the oldest eggs found and the last date on which
new laid eggs w^ere collected) are shown in Figure
1. Changes in hours of daylight and the daily
rainfall are included.
In the lake, breeding began in mid-September
in 1971 and in early September in 1972. In 1973
oviposition began early in August, lasted for
about one w eek and began again in early October
(Fig. I). In 1971 and 1972 breeding had been in
progress for some time when the first eggs were
collected, as a proportion of the eggs collected
were in late stages of development (over 10 days
old) in contrast to the first collection of 1973
when almost all the eggs found were at stage 1 1
(less than 2.5 days old) or younger. At the first
part of the spawning season of 1973 progressed
the high proportion of young eggs gave way to
peaks of older embryos and finally to collections
consisting entirely of larvae close to hatching.
This progression is not always so obvious. Most
collections from the lake show a proportion of
older eggs, unlike collections from the river.
394
MEMOIRS OF THE QUEENSLAND MUSEUM
In the river, the breeding season started earlier
and lasted longer, normally from mid to late
August until November (Fig. 1). V. spiralis beds
were used for spawning a little earlier than C.
saligna roots in 1979 and at much the same time in
1980. The V. spiralis beds were discovered in the
middle of the 1978 season, some time after
breeding had begun. Oviposition is continuous in
the V. spiralis beds throughout the breeding
season, but not on the C. saligna roots where
breaks may occur. In 1978 a small number of eggs
appeared very late, in December, on K. spiralis
plants (Fig. I). In every collection from the river,
even late in the season, a proportion of young
eggs was present.
Leading Stimulus for Oviposition
The start of oviposition did not appear to be
related to rainfall. In the lake during the 1971 and
1972, oviposition began after a dry winter, but
before heavy rains. In 1973, spawning followed a
heavy rainfall in the previous month, stopped
after 2 dry months and started again during a
month of moderate rainfall (Fig. 1).
On the river, rainfall was generally lower than
on the lake, and oviposition began in mid or late
August or early September in each year in both
breeding areas (Fig. 1). In 1979, in the V. spiralis
beds, there was a lag of two months between a
peak of heavy rainfall and production of eggs. In
1980 heavy rain fell in May, very little in June,
July or August and eggs were not found until the
end of August (Fig. 1). In 1981, the start of
spawning followed a dry month and came to an
end before much rain fell.
In the C. saligna area, a peak of rain followed
the start of oviposition in 1978, and preceded
spawning by 3 and 4 months respectively in 1979
and 1980 (Fig. 1). This area was destroyed by
logging operations before the start of the 1981
season.
Eggs less than 3 days old were found following
rain within the preceding 3 days in 6 out of 15
collections that yielded new laid eggs in the lake.
In the river 6 out of 24 and 3 out of 21 collections
contained newly laid eggs in C. saligna and V.
spiralis beds respectively. The rainfalls which did
precede the finding of newly laid eggs were
usually light, less than 25 mm. In both areas the
heaviest rains occurred in January and February,
when the fish do not spawn.
In every year, in both areas, N. forsteri begins
to breed at a time of increasing daylength, in nine
out of ten cases within 10 weeks after the shortest
day and once in the lake after 1 1 weeks (Fig. 1),
Other conditions in the Brisbane River at the time
of spawning.
The numbers of eggs found and the physical
conditions at the time of collection on successive
dates in two different localities in the Brisbane
River in 1979 and 1980 are given in Fig. 2.
Temperatures are moderate initially and fairly
steady but influenced by cold water from
Somerset Dam higher up the river, released at
irregular intervals in response to requirements in
Brisbane. There was no marked rise until mid-
October.
pH remains steady and slightly alkaline.
Dissolved oxygen levels fluctuated, low at first
and then higher, and were normally reasonably
high during the day in the weed beds, probably
because of photosynthesis by water plants. Levels
of dissolved oxygen fell in November when water
temperatures in the weed beds where the fish
spawned were consistently over 24®C.
A large number of eggs were laid in the V.
spiralis area in August 1979, at the same time as
the level of dissolved oxygen rose, and fewer eggs
were found in late September when the level of
dissolved oxygen fell. There was a second rise in
the number of eggs collected and in the level of
dissolved oxygen before egg laying stopped in
mid-November when levels of oxygen were low. A
similar but less exact correspondence between
number of eggs collected and levels of dissolved
oxygen was found in the C. saligna area in 1979.
Conversely, in 1980, oviposition began when
levels of dissolved oxygen were falling in both
areas, and reached a peak as the level of dissolved
oxygen continued to fall.
DISCUSSION
Lungfish in the lake have been observed to
spawn by day, and recently laid eggs are
frequently found late in the morning in the river
and the lake. This suggests that lungfish spawn
during the day in these localities. This conflicts
with the observation of Grigg (1965) who
observed courtship behaviour in the evening and
found eggs the following morning. Differences in
timing are probably not important and more
information may show' that the time of
oviposition is variable in both areas.
The significance of increased air breathing is
also hard to assess. Normally this occurs rarely
(Bancroft 1918 and Longman 1926). Possibly
oxygen requirements are higher in the breeding
season, or perhaps the sound made represents a
“mating call” as Kesteven states (1944: 221).
Johnels and Svensson (1955: 158) mention that “a
KEMP: SPAWNING OF NEOCERATODUS FORSTERI
395
rainfall in mm.
o o
CNJ o
o
00
o
hours of daylight
FIGURE 1. Hours of daylight, daily rainfall and duration of oviposition in Enoggera Reservoir (1971-73) and the Brisbane River (1978-1981). The
first day of each third month is marked; rainfalls of less than 3 mm are not included; the start of oviposition is calculated by subtracting the age
of the oldest eggs found from the date of collection, and the end from the last day on which new laid eggs were found.
396
MEMOIRS OF THE QUEENSLAND MUSEUM
substantial shrieking sound, which is very audible
in the swamps” can occasionally be heard when
Protopterus annectens breathes air, but they do
not associate this sound with breeding behaviour.
Lepidosiren paradoxa is supposed to be able to
make a cry like a cat (Natterer, cited by Kerr
1900). Circling movements at the surface are
presumably part of courtship, as eggs are actually
laid when the fish are entwined. This has also
been observed when lungfish have spawned in
captivity (Hegedus 1970 and Moreno 1968).
N. forsteri does not build a nest, unlike the
South American lungfish, L. paradoxa (Kerr
1900) or the African lungfish, P. annectens
(Budgett 1901 and Johnels and Svensson 1955)
and P, aethiopicus (Greenwood 1958). Also,
unlike the males of other species of lungfish (Kerr
1900, Budgett 1901 and Johnels and Svensson
1955), N. forsteri does not appear to care for its
young. There is little similarity in the breeding
behaviour of the Australian lungfish and that of
African or South American species except
perhaps the laying of separate clutches of eggs in
one place (Johnels and Svensson 1955). Whether
this suggests that male N, forsteri have territories
for oviposition W'hich are visited by successive
females for spawning, as appears to be the case in
P. annectens, or whether it is fortuitous, remains
to be seen.
Suitable weeds are available all the year round
in the Brisbane River, but they occur seasonally in
the lake, and this may affect the time of spawning
in the latter area. In 1973 in the lake weed was
available during the winter and early spring and in
this year spawning first occurred in August and
again in early October. Also, availability of
suitable weed for oviposition for a long time may
determine the length of the breeding season in the
river. Presence of suitable weed is known to affect
the timing of spawning in goldfish, Carassius
auratus Linnaeus (Stacey, Cook and Peter
1979a).
Lungfish are specific in their choice of weeds
for oviposition. In the lake, eggs are laid on
Eichornia crassipes roots, and perhaps also on
Potamogeton javanicus and para grass. In the
river, eggs were found attached to submerged
roots of Callistemon saligna or Eichornia
crassipes, to Vallisneria spiralis, Potamogeton
perfoliatus, Nitella sp. and Hydrilla verticillata
plants, occasionally to Ceratophyllum associated
with C. saligna or V. spiralis, and also on
filamentous algae covering the C. saligna roots.
Some weeds, like Potamogeton crispus, are not
used for spawning, perhaps because they only
occur in fast flowing water. Weeds which do not
form dense banks, e.g. Nymphaea capensis and
TV. flava, Nymphoides indica, Ludwigia peploides
and Rumex bidens, which all have submerged
stems, are likewise not used for egg laying. Some
of the results reported here are in agreement with
those of Semon (1893 and 1899), Bancroft (1911,
1918 and 1928) and Illidge (1893).
P. aethiopicus and P. annectens lay large
numbers of eggs, several thousand in one season,
in their nests (Budgett 1901, Greenwood 1958 and
Johnels and Svensson 1955). This does not appear
to be the case with TV. forsteri, which produces
hundreds of eggs at the most in the wild. In
captivity, numbers of 200 and 500-600 eggs laid
at one time have been reported (Hegedus 1970
and Moreno 1968).
Substrate and current may also be important in
the choice of a spawning area. Eggs are found on
weeds growing in areas with a slow or moderate
current of fresh water, where there is a substrate
of fine sand or gravel. Such areas provide a
suitable micro-environment for the larva when it
hatches i.e. a place to hide in dense cover, with
readily available food. There is also an adequate
level of dissolved oxygen maintained by plants
which are carrying out photosynthesis. Eggs of
other species of lungfish are laid in stagnant water
with low oxygen tension (Greenwood 1958) and it
appears to be necessary for the adult to agitate the
water (Budgett 1901 and Greenwood 1958), or
otherwise oxygenate it (Kerr 1900).
Eggs laid loose on the river bed were found
once only, at the height of the spawning season.
This is regarded as an abnormal feature, the result
perhaps of crowding in the weed bed. The eggs
may even have been washed into the exposed
pools where they were found, after original
deposition in the weed bed. Observations
reported by Macieay (1884) at second hand that
lungfish pair, scoop out an identation in the mud,
spawn there and remain together nearby have not
been confirmed. Also, contrary to Spencer’s
(1925) results, eggs were found to be firmly
attached to weeds in most cases and sometimes
quite difficult to remove.
Successive collections of young eggs in the
breeding areas followed by periods without newly
laid eggs or without any eggs at all must reflect
either individual fish becoming ready to breed or
the same female spawning several times. The
number of old eggs in lake collections is probably
a result of delayed hatching in eggs from this
source (Kemp 1981).
200 r * temperature ★ number of eggs collected
I ■ % dissolved oxygen • pH
KEMP: SPAWNING OV NEOCERATODUS FORSTERI
397
pH / temperature
Number of eggs/7o dissolved oxygen
SONDJFMAMJJASON January
1979 1981
FIGURE 2. Physical conditions and number of eggs collected in 1979 and 1980 in two areas of the Brisbane River.
398
MEMOIRS OF THE QUEENSLAND MUSEUM
Differences in spawning times between the lake
and the river are probably related to the
availability of suitable weeds. As the season of
1973 showed, if weed is present spawning may
occur in the lake as early as August, as it does in
the river. Differences in breeding times between
the lake, the river and more northerly river
systems may arise for similar reasons.
Attempts to relate dissolved oxygen content of
the water, temperature, pH and water level were
inconclusive, but none of these factors appeared
to act as a trigger for spawning. In one season, in
the river, there was an apparent correlation
between oviposition and a raised oxygen content
in the water. However, in the following year,
spawning occurred while oxygen content in the
water was falling. Water levels fluctuated but did
not appear to be related to spawning.
Temperatures remain the same level in the months
preceding spawning and in the early part of the
breeding season, and pH stays the same
throughout. There are no sudden changes of pH
or temperature to correspond with the beginning
of spawning, unlike the situation with certain
other fish, e.g. Carassius auratus where
temperature is involved in the stimulus for
spawning (Stacey, Cook and Peter 1979a) and
Carassius klungingeri (Lake 1967).
Breeding in the Australian lungfish appears to
be associated with a rising photoperiod and with
the presence of suitable aquatic weeds.
Oviposition in the river and in the lake begins
when the daylength has been increasing for up to
11 weeks, if suitable weeds are present. A similar
situation has been reported in other fish (Stacey,
Cook and Peter 1979a and b, Urasaki 1973 and
Pike 1973).
Rainfall sufficient to flood the environment of
the lungfish does not usually occur in the months
before they spawn, and the fish lay eggs
irrespective of rain. This differs from the
behaviour of other fish, e.g. Cyprinus carpio
(Pike 1973), or Protopterus annectens and
Lepidosiren paradoxa which spawn when the
dried out swamps where they aestivate are
flooded (Kerr 1900 and Johnels and Svensson
1955) or Protopterus aethiopicus which breeds
after rain (Greenwood 1958). Spawning in
response to flood and a minimum temperature is
an adaptive characteristic of some Australian
freshwater fish e.g. the golden perch Plectroplites
ambiguous^ the silver perch Bidyanus bidyanus
and the Murray cod Maccullochella
macquariensis^ all of which live in the Murray-
Darling River system which often dries out to
become chains of water holes (Lake 1967).
However, a response to flood is not to be
expected in a species living in Enoggera Reservoir
or the Brisbane River both of which have a
permanent inflow of water. Most features of the
oviposition of TV. forsteri appear to be related to
the particular environment.
ACKNOWLEDGMENTS
I would like to thank the large number of
friends who acted as field assistants during this
project, and Dr D.H. Kemp who reviewed the
text.
LITERATURE CITED
Aston, H. 1973. Aquatic plants of Australia.
Melbourne University Press: Melbourne.
Bancroft, T.L. 191 1 . On a weak point in the life
history of Neoceratodus forsteri Krefft. Proc.
R. Soc. Qd,, 23; 251-6.
1918. Some further notes on the life history of
Ceratodus (Neoceratodus) forsteri. Proc. R.
Soc. Qd. 30: 91-4.
1928. On the life history of Ceratodus. Proc.
Linn. Soc. N.S.W. 53:315-7.
Budgett, J.S. 1901. On the breeding habits of
some West African fishes with an account of
the external features in the development of
Protopterus annectens and a description of
the larva of Polypterus lapradei. Trans. Zool.
Soc. Lond. 16: 115-36.
Caldwell, W.H. 1884. On the development of
the monotremes and Ceratodus. JL Proc, R.
Soc. NS. W. 18:117-22,
Greenwood, P.H. 1958. Reproduction in the
East African Lungfish, Protopterus
aethiopicus Heckel. Proc. Zool. Soc. Lond.
130: 547-67.
Grigg, G. 1965. Spawning behaviour in the
Queensland Lungfish, Neoceratodus forsteri.
Aust. Nat, Hist. 15: 75.
Hegedus, A.M. 1970. Australian lungfish
spawns. Anchor 4 (7): 207-9.
Illidge, T. 1893. On Ceratodus forsteri. Proc.
R. Soc. Qd. 10: 40-4.
Johnels, A.G. and G.S.O. Svensson 1955. On
the biology of Protopterus annectens (Owen).
Ark. Zool. 7(7): 131-64.
Kemp, A. 1977. The development of Ceratodus
forsteri Krefft, with particular reference to
the dentition. Ph.D. thesis. University of
Queensland.
1981. Rearing of embryos and larvae of the
Australian Lungfish, Neoceratodus forsteri.
KEMP: SPAWNING OF NEOCERATODUS FORSTERI
399
under laboratory conditions. Copeia 1981:
776-84.
Kerr, J.G. 1900. The external features in the
development of Lepidosiren paradoxa Fitz.
Phil. Trans R. Soc. B. 172: 299-330.
Kesteven, H.L. 1944. The evolution of the skull
and the cephalic muscles. Part II. The
Amphibia. Memoirs of the Australian
Museum, 8 (3): 133-236.
Lake, J.S. 1967. Rearing experiments with 5
species of Australian freshwater fishes. 1.
Inducement of spawning. Aust. J. Mar.
Freshw. Res. 18: 155-73.
Longman, A.H. 1928. Discovery of juvenile
lungfishes with notes on Epiceratodus. Mem.
QdMus. 9:161-73.
Macleay, W. 1884. Notes on a collection of
fishes from the Burdekin and Mary Rivers,
Queensland. Proc. Linn. Soc. N.S.W. 8:
199-213.
Moreno, D.H. 1968. Letter to Queensland
Fisheries Service.
Pike, T. 1973. Delayed spawning of carp.
Lammergeyer 1973: 32.
Semon, R. 1893. Die aussere Entwickelung des
Ceratodus forsteri. Denkschr. med.-naturw.
Ges. Jena 4: 29-50.
1899. In the Australian Bush. Macmillan and
Co. London.
Spencer, B.W. 1925. Ceratodus. The Australian
Encyclopaedia, Angus and Roberston,
Sydney. Vol. i: 248-50.
Stacey, N.E., A.F. Cook and R.E. Peter
1979a. Spontaneous and gonadotrophin
ovulation induced in the goldfish Carassius
auratus L.: effects of external factors. J. Fish
Biol. 15:349-61.
1979b. Ovulatory surge of gonadotrophin in
the goldfish, Carassius auratus. Gen. Comp.
Endocrinol, 37: 246-9.
Urasaki, H. 1973. Effect of pinealectomy and
photoperiod on oviposition and gonadal
development in the fish Oryzias latipes.
J.E.Z. 185 (2): 241-46.
Mem. QdMus. 21(2): 401—11. [1984]
A BIOGEOGRAPHICALLY SIGNIFICANT NEW SPECIES OF LEIOLOPISMA
(SCINCIDAE) FROM NORTH EASTERN QUEENSLAND
Jeanette Covacevich
Queensland Museum
ABSTRACT
The rock-dwelling skink Leiolopisma jigurru sp. nov. is described from the summit of Mt
Bartle Frere, on southern Cape York Peninsula, northeastern Queensland. The combinations
of 30 mid-body scales and paired fronto-parietals distinguishes this species from all but one of
its Australian congeners. Colour, pattern, and 4th toe lamellae count distinguish it from L.
entrecasteauxii.
Zoogeographic studies of the vertebrates of Cape York Peninsula have focussed on the New
Guinea influence and on the high proportion of endemic species in the area. The discovery of a
species of Leiolopisma at 1620 m in tropical Queensland and separated from its congeners by a
gap of 1500 km highlights a new aspect of the zoogeography of vertebrates of the Cape —
‘temperate’ taxa occurring as relicts in the tropics. The distribution of Leiolopisma species is
paralleled in several other, mainly invertebrate, taxa.
L. jigurru sp. nov. shares many morphological features with other skinks confined to rock
‘islands’.
The species is ‘rare’ but its habitat is well protected.
INTRODUCTION
In October — November 1981, curators from
the Queensland Museum and ‘Earthwatch’
volunteers undertook an altitudinal survey of
invertebrates of the Belienden Ker Range, on
southern Cape York Peninsula, northeastern
Australia. (See definitions of Cape York
Peninsula by Covacevich and Ingram 1980, and
Covacevich et al. 1982). This range, the second
highest in Australia, supports dense rainforest
and has not been methodically surveyed before.
During this survey, it was possible to collect frogs
and reptiles on Mt Bartle Frere, the highest and
most southern peak of the range. Amongst the
material collected and now located in the
Queensland Museum are two new species of
skinks belonging to the genera Lampropholis and
Leiolopisma. The Lampropholis sp. nov. occurs
widely in the rainforests of the Belienden Ker
Range and is, along with other members of this
genus, the subject of revision of Mr Mark
Schuster of the University of New England.
Greer (1974, 1979) has discussed the
relationships of the skinks, including those in the
genera Leiolopisma and Lampropholis. He has
shown that Leiolopisma spp. have alpha palates
(with the inner edges of the palatal rami diverging
posteriorly along the smooth curve) while
Lampropholis spp. have beta palates (with the
rami having a large recurving process anteriorly).
Such a major difference; Greer’s (1974) diagnoses
of the genera; Cogger’s (1979) additional
diagnostic feature of narrowly separated nasals
for Leiolopisma V5 widely separated nasals for
Lampropholis; and the fact that Leiolopisma spp.
are generally viviparous (Rawlinson 1976) while
Lampropholis spp. lay eggs communally (Ingram,
pers. comm.) suggest that assigning skinks to
these two genera would be a simple task. Such is
not the case however.
The palate in small skinks cannot be examined
easily and there is considerable overlap in the
characters used by Greer and Cogger to
distinguish these genera. Further, Ingram and
Ehmann (1981) have recently described an egg-
laying species of Leiolopisma, L. zia, from
southeastern Queensland and northeastern New
South Wales. Examination of the karyotypes of
most currently recognised species of Leiolopisma
(including L. Jigurru ), Lampropholis, and most
other genera in the Eugongylus group sheds no
further light on the problem of separating
Leiolopisma from Lampropholis. All species of
Leiolopisma examined have 30 chromosomes and
are very similar karyotypically. The only
variation is in pairs 6-9 which is also a
402
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 1: Occurrence of Leiolopisma species in Australia.
characteristic of Lampropholis spp. (S. Donellan,
pers. comm.).
The Bartle Frere skink described here has an
alpha palate, a characteristic of Leiolopisma. It
also has the widely separated nasals of
Lampropholis. The degree of separation of the
nasals in Leiolopisma as presently defined
apparently varies considerably (e.g. L. zia,
narrow vs L. trilineata, wide, but not as wide as in
the species described here). No data on breeding
biology for this new species are available because
only a handful of specimens are known.
In the light of this information it is reasonable
to assign the Bartle Frere species to Leiolopisma.
Leiolopisma jigurru sp. nov.
(Pis la, b; 2a, b; 3)
Material Examined
Holotype: QM J40040, 2 , near summit of
South Peak of Mt Bartle Frere, NE.Q., 1620 m,
on granite boulders; J. Covacevich, R. McKay,
D. Marshall; 7-8 Nov., 1981.
Paratypes: AM R95553, Mt Bartle Frere,
1524 m, 23 Jan., 1977; J39494-99, Northwest
Peak of Mt Bartle Frere, 1440 m, under
exfoliated granite, 7-8 Nov., 1981; J39492-3 as
for holotype.
COVACEVICH: SIGNIFICANT NEW LEIOLOPISMA
403
DIAGNOSIS
A mid-body scale count of 30 and paired
frontoparietal scales distinguishes Leiolopisma
jigurru from other Australian species of
Leiolopisma except L. entrecasteauxii (Dumeril
and Bibron). L. entrecasteauxii has a lower
lamellae under the 4th toe count (16-22) than L.
jigurru (26-29) and lacks the distinctive dark
brown to black, and white to cream colour
pattern of L. jigurru.
Ten New Zealand species of Leiolopisma have
the combination of 30 mid-body scale rows and
paired frontoparietals, but only 3 species also
have a 4th toe subdigital lamellae count which
overlaps with that of L. jigurru. These are L.
infrapunctatum (Boulenger), L. nigriplantare
(Peters), and L. lineoocellatum (Dumeril and
Dumeril). Colour and pattern quickly distinguish
L. jigurru from these species. See PI. la-b, 2a-b
and Hardy (1977, figs 27, 30-32, 33).
Description of Holotype
Snout-vent length 68.9 mm, tail 126.3 mm;
T/SVL'!^ 183.3; tip of snout - forelimb/axilla -
groin = 22.2/37.0 (.59); head width 7.5 mm.
Head slender. Rostral broad, in contact with
the nasals and frontonasal. Frontonasal broader
than long, bordered by two large prefrontals
which do not meet. Frontal twice as long as
broad, narrow posteriorly, and equal in length to
the frontoparietals and parietal together. Frontal
in contact with first and second supraocular.
Supraoculars 4, the second largest and the fourth
smallest. Seven supraciliaries. Two
frontoparietals, which are distinct from and
larger than interparietal. Seven supralabials, 5th
largest and, with sixth, contacting eye. Lower
eyelid scaly with a large oval palpebral disc. Ear
opening large, nearly round, with a deeply set
tympanum, and without auricular lobules.
Mid-body scales 30. Mid-dorsal scales slightly
larger than ventral and lateral scales, and lightly
striated. Limbs and digits long. Twenty-six
lamellae under 4th toe.
Colour (in life): Basically brown and black
dorsally and cream ventrally, with a metallic
sheen. See PI. la, b. and 2a, b for distinctive
pattern.
Variation in the Paratypes
SVL 34.5 - 67.2, Tail 63.5 - 115.5 (part of the
tail of J39495 has been lost), tip of snout -
forelimb/axilla - groin .54 - .88, head width 4.5 -
8.2, T/SVL 135 - 189%. There is little variation.
Lamellae under the fourth toe, 27-29. In six
paratypes there are 8 supraciliaries. One specimen
(J39492) has an extruded columnar hemipenis.
DISTRIBUTION AND HABITAT
Leiolopisma jigurru is known from only one
locality — Mt. Bartle Frere, on southern Cape
York Peninsula, NE.Q. It is found amongst
granite boulders which occur as large ‘fields'
surrounded by dense rainforest near the mountain
summit. Specimens were collected at 1440 m,
1524 m, and 1620 m. The type locality is cool to
cold throughout the year and is frequently
covered in mist. Climatic data are not recorded on
Mt Bartle Frere but average annual rainfall
(1974-1980) on the adjoining peak, Mt Beilenden
Ker (1560 m), is 7736 mm. In early November,
1981 when all but one of the skinks in the type
series were collected, daily temperatures ranged
from 7° - 20'^C.
ETYMOLOGY
‘Jigurru’ is both the Mamu and the Ngajan
name for this lizard, acccording to Molly
Ramond and George Watson, the last people to
speak these languages well. Their people lived in
the rainforest country at the headwaters of the
Mulgrave and Russell Rivers on the slopes of the
Beilenden Ker Range and their territories
overlapped in the high mountains such as Bartle
Frere. ‘For more than ten thousand years they
lived in harmony ... with their environment. One
hundred years ago many of them were shot and
poisoned (Dixon, 1972).
DISCUSSION
Biogeographic studies of the herpetofauna of
Cape York Peninsula have focussed on
Pleistocene New Guinea migrations and on the
high proportion of taxa endemic to the area (e.g.
Keast 1959; Storr 1964; Tyler 1972; Covacevich
and Ingram 1980; Kikkawa et al. 1981;
Covacevich et al. 1982). The discovery of
Leiolopisma jigurru on the ‘temperate’ summit of
Mt. Bartle Frere on southern Cape York
Peninsula in tropical Queensland highlights
another aspect of its biogeography.
Several taxa whose present distribution is
concentrated in the southern, temperate zones of
Australia, are known to have relict
representatives in cool, montane habitats in the
tropics. This pattern has been observed for
certain landsnails (Ohdner 1917) and plants and
insects (Monteith 1980, Storey 1983) but has not
404
MEMOIRS OF THE QUEENSLAND MUSEUM
been previously recorded for vertebrates. Spiders
(Gradungulidae, Migidae) and the marsupial
Antechinus stuartii also have similar
distributions. (V. Davies, S. Van Dyck, pers.
comm.). The occurrence of Leiolopisma spp. is of
special interest because, with the discovery of L.
jigurru, it is a parallel of the southeastern
Australia — montane northeastern Queensland
— New Caledonia — New Zealand occurrence
noted for some insects and plants (Monteith,
1980).
Forty-two species of Leiolopisma are now
recognised. They occur in Tasmania. Lord Howe
Island, mainland southeastern and southwestern
Australia; New Zealand and the Chatham
Islands; New Caledonia and the Loyalty Islands;
and Mauritius (Greer 1979). The present
‘stronghold’ for the genus is temperate
southeastern Australia — New Zealand (Hardy
1977). The Australian distribution of members of
this genus is shown in Fig. 1. There is a gap of
some 1500 km between the Mt Bartle Frere
population of L. jigurru and the other two species
occurring in Queensland, L. platynota (Peters)
and L. zia Ingram and Ehmann, both of which
occur in Queensland only at high altitudes in the
e.xtreme southeast of the state. L. platynota has a
fairly broad coastal distribution from
southeastern Queensland to northeastern Victoria
(Cogger 1979). L. zia, on the other hand, is
restricted to high altitude (above KXK) m)
rainforests and Antarctic Beech (Nothofagus )
forests of southeastern Queensland and
northeastern New South Wales (Ingram and
Ehmann 1981).
Greer (1974) has suggested a southern.
Tasmanian/southeastern Australian, centre of
diversity for Leiolopisma. Hardy (1977) revised
New Zealand species of the genus and suggested a
northern. New Guinean, centre of diversity. The
discovery of the temperate relict species,
Leiolopisma jigurru, in tropical Queensland may
be used to support either hypothesis.
L. jigurru, is an agile and fast-moving
posturing heliotherm like other lygosomine skinks
endemic to isolated rocky areas. It is a typical
rock dweller in being flattened dorsoventrally,
large in relation to its congeners, and in having
long digits and limbs and a highly achromatic
pattern (Covacevich and Ingram 1980).
Frog and reptile species have been described as
‘rare’ for conservation purposes if they are
known from 20 or less museum specimens or
from five or less localities (Covacevich, et al.
1982). L. jigurru, qualifies as a ‘rare’ species on
both counts, but is well protected. The type
locality and other potential habitats on the
Bellenden Ker Range lie in State Forest and
National Park.
ACKNOWLEDGMENTS
‘Earthwatch’ is a private, American-based,
non-profit organization which makes available
volunteer workers to assist research programmes
involving field expeditions. Without the support
of ‘Earthwatch’, L. jigurru would not have been
collected. Dr G. Monteith led the expedition and
made the trip to Bartle Frere possible.
V. Davies. S. Donnellan, S. Van Dyck, G.
Hardy, G.J. Ingram, G. Monteith, K. McDonald
and M. Schuster has assisted in preparing this
paper, either by providing information or
constructive criticism.
Photographs were taken by A.J. Easton.
LITERATURE CITED
Bull, P.C. and A.H. Whitaker, 1975. The
amphibians, reptiles and mammals, in
Kuschel, G. (ed.) ‘Biogeography and Ecology
in New Zealand’. (Junk: The Hague).
Covacevich, J. and G.J. Ingram, 1980. The
endemic frogs and reptiles of Cape York
Peninsula, in Stevens, N.C. and Bailey, A.
(eds.) ‘Contemporary Cape York Peninsula’
(Royal Society of Queensland : Brisbane).
G.J. Ingram and G.V. Czechura, 1982. The
‘rare’ frogs and reptiles of Cape York
Peninsula, Australia. Biol. Conservation 22
(4): 283-94.
Cogger, H.G. 1979. ‘Reptiles and frogs of
Australia’. (A.H. and A.W. Reed: Sydney).
Dixon, R.M.W. 1972. ‘The Dyirbal language of
North Queensland’. (Cambridge University
Press: Cambridge).
Greer, A.E. 1974. The generic relationships of
the scincid lizard genus Leiolopisma and its
relatives. Aust. J. Zool. SuppL Ser. no. 31:
1-67.
1979. A phylogenetic subdivision of Australian
skinks. Rec. Aust. Mus. 32 (8): 339-71.
Hardy, G.S. 1977. The New Zealand Scincidae
(Reptilia : Lacertilia); a taxonomic and
zoogeographic study. N.Z. J, Zool. 4 (3):
221-325.
Ingram, G. and H. Ehmann, 1981. A new
species of scincid lizard of the genus
Leiolopisma (Scincidae : Lygosominae) from
southeastern Queensland and northeastern
New South Wales. Mem. Qd Mus. 20 (2):
307-10.
COVACEVICH: SIGNIFICANT NEW LEIOLOPISMA
405
Keast, a. 1959. The reptiles of Australia in
Keast, A., R.L. Crocker and O.S. Christian
(eds.) ‘Biogeography and Ecology in
Australia’ (Junk: The Hague).
Kikkawa, J., G.B. Monteith and G. Ingram,
1981. Cape York Peninsula: major region of
faunal interchange. Ch. 60, in Keast, A. (ed.)
‘Ecological Biogeography of Australia’.
(Junk: The Hague).
Monteith, G.B. 1980. Relationships of the
genera of Chinamyersiinae, with description
of a relict species from mountains of north
Queensland (Hemiptera : Heteroptera :
Aradidae). Pacific Insects 21 (4): 275-85.
Ohdner, N.H.J. 1917. Results of Dr E.
Mjoberg’s Swedish Scientific Expedition to
Australia, 1910-1913, XVII Mollusca. KungL
Svenska. Vetenskapsakademiens. Handlingar
52 (16): 1-115, pis. 1-3.
Rawlinson, P.A. 1974. Biogeography and
ecology of the reptiles of Tasmania and the
Bass Strait area, in Williams, W.D. (ed.)
‘Biogeography and Ecology in Tasmania’.
(Junk: The Hague).
1976. The endemic Australian lizard genus
Morethia (Scincidae : Lygosominae) in
southern Australia. Mem, Nat. Mus. Vic. 37:
27-42.
Storey, R.L 1983. A new species of
Aptenocanthon Matthews from north
Queensland (Coleoptera : Scarabaeidae :
Scarabaeinae). Mem. Qd Mus. 21:
357-60.
Storr, G.M. 1964. Some aspects of the
geography of Australian reptiles.
Senckenbergiana bioL 45: 577-89.
Tyler, M.J. 1972. An analysis of the lower
vertebrate faunal relationships of Australia
and New Guinea, pp. 231-56 in Walker, D.
(ed.) ‘Bridge and Barrier: the natural and
cultural history of the Torres Strait’. (Aust.
Nat. Univ.: Canberra),
406
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
a,b Leiolopisma jigurru sp. nov. from the summit of
Mt Bartle Frere, NE.Q., showing highly
achromatic pattern, dorsoventral flattening, and
long digits.
COVACEVICH: SIGNIFICANT NEW LEIOLOPISMA
407
b
408
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
a Close-up lateral view of L. Jigurru sp. nov. showing
ear opening, eye detail and colour pattern,
b Dorsal view of head scales of L. jigurru sp. nov.
COVACEVICH: SIGNIFICANT NEW LEIOLOPISMA
409
410
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 3
Mist-covered granite boulders near the summit of Mt. Bartle Frere,
type locality of the temperate relict, L. jigurru sp. nov.
COVACEVICH: SIGNIFICANT NEW LEIOLOPISMA
4II
Mem. QdMus. 21(2): 413—517. [1984]
DINOSAUR TRACKWAYS IN THE WINTON FORMATION (MID-CRETACEOUS)
OF QUEENSLAND
Richard A. Thulborn,
Department of Zoology, University of Queensland
and
Mary Wade
Queensland Museum
ABSTRACT
Dinosaur trackways have been discovered at several closely-grouped sites in mid-Cretaceous
sediments of the Winton Formation, central W. Queensland. At one of these sites about 209 m^
of bedding plane was exposed to reveal trackways of more than 150 bipedal dinosaurs. One of
these trackways is very much larger than any of the others; it is attributed to a large theropod
dinosaur (carnosaur) and is identified as cf. Tyrannosauropus. The remaining trackways are
referred to two new ichnotaxa — Wintonopus latomorum and Skartopus australis — which are
attributed to ornithopods and coelurosaurs (respectively). The sizes of the track-makers are
estimated by means of allometric equations derived from osteological data; the speeds of the
track-makers are estimated by using the mathematical relationships of size, speed and gait that
have been determined for living tetrapods. The carnosaur is estimated to have been about 2.6
m high at the hip, and to have been walking at a speed of about 7 km/h. The ornithopod track-
makers ranged from 14 to 158 cm in height at the hip; these animals were using a fast running
gait equivalent to cantering or galloping in mammals, and their mean speed is estimated at 16
km/h. The coelurosaur track-makers ranged from 13 to 22 cm in height at the hip; these too
were using a fast running gait, and their mean speed is estimated to have been 12 km/h. The
trackways of the ornithopods and coelurosaurs are interpreted as those of animals caught up in
a stampede — which was presumably triggered by the approach of the carnosaur. It is
suggested that relative stride length (i.e. stride length relative to height at the hip) is the best
available criterion for appraising the locomotor performances of dinosaurian track-makers. By
this criterion the performances of the Winton ornithopods and coelurosaurs are outstandingly
good. There is an indication that these animals were running at or near their maximum speeds
— with relative stride lengths in the range 3.9 to 5.0. If the most highly adapted of cursorial
dinosaurs (the ornithomimids or ‘ostrich dinosaurs’) attained such figures for relative stride
length their speeds would have been up to about 60 km/h.
INTRODUCTION
In June 1976 Mr Ron McKenzie showed us
some well-preserved dinosaur footprints that he
had collected from a site about 120 km SW of
Winton, central west Queensland. The footprint
site, which was later named Seymour Quarry, was
in sediments of the Winton Formation (mid-
Cretaceous) and its existence was known to many
residents in the Winton area. In 1971 a small field
party including Dr R.H. Tedford (American
Museum of Natural History) and Dr A.
Bartholomai (Queensland Museum) paid a brief
visit to the site; this party established that the
footprint horizon extended to a second site some
100 metres away (Knowles 1980). This second
site, which was later named Lark Quarry,
subsequently proved to be of very great interest.
The footprints at these localities attracted our
attention because of their abundance, their
excellent preservation and their remarkably small
size (by dinosaurian standards). In 1976 we
carried out preliminary excavations at both sites,
and in the following year a large labour-force of
volunteers co-operated in a major excavation at
Lark Quarry. This excavation revealed several
thousands of footprints representing the
trackways of well over 100 bipedal dinosaurs --
many of them apparently no bigger than
chickens. In preliminary accounts the Lark
Quarry trackways have been interpreted as
414
MEMOIRS OF THE QUEENSLAND MUSEUM
evidence of a dinosaur stampede (Thulborn and
Wade 1979, Wade 1979). If this interpretation is
correct it may carry important implications for
current understanding of dinosaur biology. The
present work has three aims: 1) to provide a
systematic account of the trackways at Lark
Quarry and its surroundings; 2) to offer some
interpretations regarding the sizes, speeds and
behaviour of the track-makers; 3) to justify those
interpretations and to consider their implications
for the understanding of dinosaur biology.
LOCALITIES
The footprints described in this paper were
found at three localities on Mt Cameron
property, SW of the town of Winton in central
west Queensland. About 95 km SW of Winton the
road to Jundah and Stonehenge runs along the
crest of a west-facing scarp (the Tully Range)
which is formed by sediments of the Winton
Formation capped by duricrust. The localities are
close to the foot of the scarp, alongside a track
leading NW to Cork Station. The maximum
distance between any two of the localities is about
200 m (see map. Fig. 1).
Seymour Quarry. A deep hillside cutting
alongside the track to Cork Station. The site is
identified as an opal mine on the Brighton Downs
sheet of the BMR 1:250,000 series of geological
maps (sheet SF/54-15; map reference 23°0rS,
142^24’W). Footprints occur as natural casts
below' a thin bed of red arkosic sandstone that
outcrops at the foot of the hill. This friable
sandstone overlies a weathered mudstone, and its
lower surface is infiltrated by dark brown
ironstone which prevents the footprints from
crumbling on exposure. Traces of plant rootlets
are preserved along with the footprints, while
these themselves are very well preserved and may
even show indications of skin texture (see p.
422, PL 1). The footprints are attributed to
small bipedal dinosaurs of two types
(coelurosaurs and ornithopods), and they appear
to represent continuations of trackways at
another site to the SW (Lark Quarry). This first
site is named for Mr Glen Seymour, its discoverer
and former manager of Cork Station.
Lark Quarry (PL 3). A large excavation
revealing more than 200 m’ of a single bedding
plane. This site is located to the SW of Seymour
Quarry, and has been the subject of preliminary
descriptions (Thulborn and Wade 1979, Wade
1979). The Lark Quarry bedding plane carries
well over 3000 footprints, representing the
trackways of at least 150 bipedal dinosaurs. The
trackways are almost entirely unidirectional: one
track-maker was headed to the SW, whereas all
the others were headed to the NE (in the direction
of the present Seymour Quarry). The footprints
occur as natural moulds in a bed of laminated
claystone which varies between 6 and 12 cm in
thickness. The footprint itself will be referred to
as a mould, and the filling of the footprint as a
cast, in conformity with standard ichnological
usage. The claystone is generally bright pink in
colour (though individual laminae range from
pink through red to purple), and its upper surface
appears to be stained dark red-brown by
ironstone infiltration. This ‘surface stain’ is in
fact an extremely thin adhesion from the base of
the overlying sandstone. Below the claystone is a
thick bed of arkosic sandstone; this is buff in
colour and finely cross-bedded. Similar
sandstone/claystone couplets occur above and
below' the trackway horizon. The next claystone
bed below' the trackway horizon also bears
footprints in the form of natural moulds, though
these are uncommon and seem to have no
preferred orientation. The footprint horizon at
Seymour Quarry seems to be an extension of the
main trackway surface at Lark Quarry; it is
possible to trace the footprint horizon through
intermediate outcrops, though there is a complete
break of about 30 metres caused by a creek bed.
Moreover there is a uniform dip to the NW of
about 4^, and by taking direct line of sight along
the Lark Quarry bedding plane this will be found
to coincide with the footprint horizon at Seymour
Quarry. It may be mentioned that the reverse
procedure (extrapolating the dip of the beds at
Seymour Quarry) was used to locate the Lark
Quarry trackways in the first instance (Knowles
1980). At both sites the footprints are similar in
diversity, in abundance, in morphology and in
their singular orientation. Lark Quarry has been
designated an Environmental Park by the
National Parks and Wildlife Service, Queensland,
and is now' roofed for its protection. The site is
named for Mr Malcolm Lark, of Miles, who
played a leading role in its excavation.
New' Quarry. One of a series of small hillside
exposures scattered from 100 to about 120 m due
S of Lark Quarry. At the New Quarry site the
trackway of a single bipedal dinosaur was
measured in situ. In its preservation this trackway
is identical to those at Lark Quarry. There is a
major erosional gap between New Quarry and
Lark Quarry, but at both sites the footprints
occur at equivalent levels in similar sequences of
sandstone/claystone couplets. Moreover at both
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
415
FIGURE 1. Map showing location of footprint sites. Contours are at 5 m intervals, and dashed lines indicate dry
creek-beds. Inset map shows location of quarry in Queensland. B, Brisbane; C, Cairns; M, Mt. Isa; R,
Rockhampton; T, Townsville.
416
MEMOIRS OF THE QUEENSLAND MUSEUM
sites there is a marked change in sediment type
about 5 or 6 m above the footprints — the
appearance of a yellowish arkose containing
scattered plant fragments. The evidence is not
conclusive, but it suggests that the New Quarry
trackway is at about the same stratigraphic level
as the Lark Quarry trackways. Footprints also
occur in the next claystone layer below the New
Quarry trackway; at this lower level the claystone
is thoroughly trampled and churned up by deep
footprints without preferred orientation.
METHODS
Excavation and collection of footprints
At Seymour Quarry the footprint horizon was
reached by digging through the overburden of soil
and weathered rock. The thin sandstone layer
bearing the footprints (natural casts) proved
rather fragile; most footprints collected from this
site are on small slabs or are in the form of
detached casts (see PL 1). One large slab (QM
FI 2266) is approximately 95 by 40 cm and carries
casts of at least 28 footprints — representing
about 19 trackways.
At Lark Quarry the footprints were exposed by
breaking up and removing a thick overburden of
sandstone. Fortunately the sandstone was well
jointed (as is the footprint surface — see PL 4),
and it could be removed in blocks once these had
been levered out with crowbars and jack-hammer.
About 60 tons of overburden was removed,
exposing an area of more than 209 mL It was then
necessary to clean the footprints (natural moulds)
be removing the sandstone that filled them. This
sandstone filling was soft enough to be broken up
with an awl. At the bottom of each footprint
mould the colour of the sandstone filling changed
from orange-red to bright yellow-green — a
useful guide to ensuring that the footprints were
fully excavated. More than 3300 footprints were
exposed and cleaned in this way. A portion of the
footprint bed was removed from the eroded NE
margin of the site and was transferred to the
Queensland Museum (QM F10321). In addition
several individual footprints were collected —
including holotypes of the new ichnotaxa
described below.
Fibreglass replicas
After the Lark Quarry bedding plane had been
exposed, and its footprints were thoroughly
excavated, it was swept free of dust and rock
debris; large parts of the bedding plane were then
coated with liquid latex, which was reinforced
with a cloth backing. Once it had set, the latex
was stripped off in the form of large ‘peels’
(Wade 1979). These latex ‘peels’ were later used
as a basis for moulding a fibreglass replica of the
bedding plane and its footprints. The. entire area
shown in Fig. 3 was included in this replica.
Individual footprints were also replicated (see, for
example, PL 5, Fig. A). These high-fidelity
replicas are much lighter and more durable than
plaster casts; they enabled us to undertake a long-
term study of the Lark Quarry footprints, even
though our total expenditure of time at the site
was no more than a few weeks.
Illustrations
The Lark Quarry bedding plane is almost
horizontal, and its footprints are under natural
low-angle lighting for only a few minutes after
dawn and before dusk. Even at these times of day
it may be difficult to obtain worthwhile
photographs because the direction and intensity
of lighting cannot be adjusted. We obtained few
good quality photographs of individual footprints
in situ. Most of our illustrations (PL 5 to 16) show
fibreglass replicas of the footprints — though
some do show original material (including type
specimens; see plate captions).
Most of the area exposed at Lark Quarry was
marked out with a grid of chalk lines, and each
quadrat was photographed from a height of 1
metre (with the camera mounted on a rigid iron
frame). The resulting photographs were then
assembled into an accurate and detailed
photomosaic — a representative portion of which
is reproduced as PL 4. The same photographs
were later used in drawing up a chart to show the
distribution of footprints at Lark Quarry (PL 17).
Descriptions
There are no universally accepted methods for
describing footprints and trackways (Sarjeant
1975, Leonard! 1979a), and it is necessary to
define the measurements and statistics we
employ. All linear measurements are expressed in
centimetres.
Footprint length (abbreviation FL) — the
maximum footprint dimension measured along,
or parallel to, the axis of the longest digit (see
Figs. 2A, B).
Footprint width (FW) — the maximum
footprint dimension measured at a right angle to
footprint length (Figs. 2A, B).
The ratio footprint width / footprint length
(ratio FW/FL) is used to express footprint
proportions.
We discovered that FL and FW were quite
variable within each trackway, so that neither of
these measurements could be regarded as a
completely reliable indicator of the track-maker’s
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
417
size. Consequently we calculated an index of
footprint size (SI) for each footprint:
SI = (FL X Fwy’^
This index was found to be remarkably consistent
within each trackway, and it seems to be a useful
guide to the relative sizes of two or more track-
makers. The index is expressed in centimetres.
Interdigital angles (expressing divarication of
digits) are commonly cited in descriptions of
vertebrate footprints, but they are difficult to
measure consistently (see Sarjeant 1975) and are
often so variable that they are of questionable
value (see comments of Welles 1971). We have
not attempted to compile detailed measurements
of interdigital angles, and they will be mentioned
only as approximate averages.
Pace length (PL) — the distance between
corresponding points in two successive footprints
(left and right, or right and left; see Fig, 2C).
Stride length (SL) — the distance between
corresponding points in two successive prints of a
single foot (see Fig. 2C).
With measurements of two successive paces
(PL' and PL"), and of the stride (SL) that they
encompass, it is possible to calculate pace
angulation (ANG) as follows:
cos ANG = _(PL-)^+ (PLT- (SL)^
2 X (PL^) X (PL^)
The more nearly pace angulation approaches to
180° the narrower is the trackway and the less
obvious is the zig-zag arrangement of its
footprints (see Fig. 2C).
The ratios pace length / footprint length
(PL/FL) and stride length / footprint length
(SL/FL) are often used in definitions of
ichnotaxa (see, for example, Lull 1953, Haubold
1971) and are also provided here. These ratios
tend to increase as a track-maker accelerates, and
they can therefore give a useful indication of a
track-maker’s gait. In calculating these ratios FL
was taken to be the mean for the two footprints
defining each pace or stride.
To calculate means and other statistics it was
usually necessary to reduce sample sizes (N) by
excluding data from damaged or badly distorted
footprints.
DESCRIPTIONS
At first glance the dinosaur trackways at Lark
Quarry present a rather confusing picture (PI. 4).
However, it soon becomes apparent that the
trackways can be sorted into several natural
groups on the basis of size, orientation,
preservation and footprint shape. Five such
groupings may be recognized.
A. Remnants of a few trackways made by fairly
large bipedal dinosaurs. These remnants comprise
scattered footprints which are very poorly
preserved and have no preferred orientation. The
footprints appear to have been tridactyl, with
rather short, thick and bluntly-rounded toes, and
they are tentatively attributed to ornithopod
dinosaurs. They seem to have been formed, then
eroded and filled with water-laid sediment, well
before the substrate was exposed to the air and
the other footprints were formed at Lark Quarry.
It was not possible to obtain any accurate
measurements, and these remnants of old
trackways will not be considered further.
B. A single trackway of a medium-size bipedal
dinosaur (B in Fig. 3). This trackway extends
across the southern part of Lark Quarry from
WSW to ENE, and is attributed to an ornithopod
dinosaur. The tridactyl footprints have relatively
short, broad and well-rounded digits (see example
a in Fig. 4) and are referred to the same
ichnotaxon as the many small footprints in group
D (below). However, this trackway is much larger
than any of those in group D, and it was certainly
formed at an earlier date: some of its footprints
were deeply impressed in soft waterlogged mud
and others (in lower-lying areas) were partly
destroyed by scouring. This trackway seems to
have been formed at about the time the substrate
was draining free of surface water and was
becoming exposed to the air.
C. A single trackway of an exceptionally large
bipedal dinosaur (C in Fig. 3). This trackway
extends across the northern part of Lark Quarry
from NE to SW, and is attributed to a carnosaur
— a large representative of the Theropoda. The
footprints are very obvious basin-like structures
(PI. 5, Fig. B), and some of them show clear
traces of three tapering or V-shaped digits (Fig.
4).
D. Numerous trackways of small to medium-
sized bipedal dinosaurs; extending from SW to
NE (Fig. 3, PI. 4). The footprints are well
preserved and each of them comprises three fairly
short, thick and bluntly rounded digits. These
trackways are attributed to ornithopod dinosaurs,
and their footprints may be found superimposed
upon those of the carnosaur (C, above) and upon
those of coelurosaurs (E, below).
E. Numerous trackways of small (and
sometimes very small) bipedal dinosaurs;
extending from SW to NE (Fig. 3, PI. 4). Each of
these trackways comprises footprints with three
fairly long, narrow and sharply-pointed digits.
The trackways are attributed to coelurosaurs
(small dinosaurs of the suborder Theropoda), and
418
MEMOIRS OF THE QUEENSLAND MUSEUM
their footprints may be found superimposed upon
those of the carnosaur (C ) and upon those of the
ornithopods (group D ).
Footprints in the latter three groups are equally
well preserved, and all of them seem to have been
formed at about the same time. These footprints
were formed after the muddy substrate had been
exposed long enough to have attained a firm
plastic consistency. From the evidence of
superimposed footprints it is clear that the
carnosaur traversed the Lark Quarry area before
some, at least, of the ornithopods and
coelurosaurs did so.
The trackways attributed to ornithopods
(group D ) and coelurosaurs (group E ) extend in a
single direction, and among them it is common to
find trackways coinciding, intersecting at low
angles, or weaving together inextricably (PI. 4 and
14). Moreover in some places the trackways of
small, and even medium-size, individuals quite
simply disappear: apparently these smaller
dinosaurs were so light that their feet failed to
break through the firmer patches of surface
sediment. These discontinuities are especially
noticeable among the coelurosaur trackways
(group E ); the coelurosaur track-makers seem to
have had relatively large feet (by comparison with
the ornithopod track-makers), and their widely-
spread and probably rather springy toes seem to
have functioned as analogues of snow-shoes.
Many of the track-makers, both ornithopods and
coelurosaurs, seem to have been roughly similar
in size (the majority having hip height estimated
at less than 50 cm), and the footprints in any one
trackway are not always consistent in their shape
or spacing. This combination of factors makes it
difficult TO trace any single trackway, with
confidence, for more than a few strides.
Consequently our descriptions and analyses are
based, in the main, on relatively short sections of
trackways. For the ornithopod dinosaurs (group
D ) we examined 57 sections of trackways; on
average each of these comprises 3 strides (a
sequence of 5 footprints). The longest section of
ornithopod trackway studied here comprises 17
strides (a sequence of 19 footprints). For the
coelurosaurs (group E ) we examined 34 sections
of trackways; here the average number of strides
per section of trackway is between 3 and 4
(between 5 and 6 footprints). The longest section
of coelurosaur trackway studied here comprises
22 strides (a sequence of 24 footprints). The
difference in sample size (57 ornithopod
trackways as opposed to 34 coelurosaur
trackways) does not indicate that ornithopods
were more abundant than coelurosaurs. On the
contrary, the ornithopod track-makers were
probably outnumbered by the coelurosaur track-
makers (see p. 443). The sample sizes differ for
two reasons. First, the coelurosaur trackways are
affected by so many discontinuities that it is
difficult to find sequences of more than a few
paces. Second, the coelurosaur footprints show
much less variation in size and shape than do the
ornithopod footprints: coincident or intersecting
trackways of ornithopods could usually be
separated through differences in footprint size or
footprint shape, but coincident or intersecting
trackways of coelurosaurs were usually
inextricable.
The trackways of the carnosaur, the
ornithopods and the coelurosaurs are described in
turn. But before proceeding to these descriptions
it will be useful to consider the circumstances
under which the trackways were formed. It is
important to determine these circumstances
because some of them (e.g. the consistency of the
substrate) have a direct bearing on footprint
morphology, while others (e.g. the physical
geography) are pertinent to the behaviour of the
track-makers.
The Winton Formation is a series of
continental sediments that reaches a thickness of
more than 10(X) feet in the area around Winton
(Casey 1966). The sediments are mainly
sandstones, siltstones and mudstones, though
there are some intraformational conglomerates
and coal seams. Fragments of fossil wood are
common, but other well preserved fossils are rare;
these include angiosperm leaves, conifers,
freshwater bivalves, lungfish toothplates and
fragmentary remains of sauropod dinosaurs
(Senior, Mond and Harrison 1978, Coombs and
Molnar 1981). The Winton Formation is mid-
Cretaceous in age (probably uppermost Albian to
Cenomanian), and the conditions under which its
sediments accumulated were well summarized by
Senior et al. (1978, p. 15): ‘Terrestrial-fluviatile,
paludal, lacustrine. Low relief, wide river flats,
local development of short lived lakes and
swamps.’
The sediments at and around Lark Quarry are
of lacustrine and fluviatile origin. At the time the
dinosaur trackways were formed the Lark Quarry
site seems to have repre.sented part of a major
drainage channel; it was most probably part of a
platform lying on point bar deposits, a sand-spit,
that had built out into a lake which deepened to
the SW' (see Fig. 25A). The Lark Quarry bedding
plane now dips NW at 4°, but it originally had a
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
419
run-off to the SW — as is indicated by the
orientation of drag marks and prod-marks
produced by floating vegetation (PI. 4). At times
of flood the lake would have spread over a wide
area (including all three footprint localities) to
deposit sand followed by muddy sediments. In the
intervening periods the lake would have receded
to become little more than a remnant water-hole
surrounded by newly-exposed mud. It was during
such a period that the trackways seem to have
been formed.
The mud began to compact under water, and
before it was fully exposed a single dinosaur
traversed the southern part of the future Lark
Quarry site from WSW to ENE. In this trackway
(B ) some footprints were formed as the animal
crossed slightly elevated and newly-exposed
patches of very soft sediment; the other footprints
were formed in lower-lying areas of mud which
were still covered by water. These lower-lying
footprints were subsequently scoured and eroded
as the remaining surface water drained off to the
SW. At the future site of Lark Quarry the
recently-laid sand and overlying mud were
penetrated by narrow, vertical and unbranched
tubes that probably mark the escape of buried
arthropods (PL 2, Fig. B). Traces of similar
escape burrows may be found at the Seymour
Quarry site, along with horizontal and oblique
tubular structures that seem to represent plant
rootlets of various sizes (PL 1, Figs. A, B). The
presence of plant rootlets might indicate that the
mud was exposed sufficiently long for terrestrial
vegetation to take hold. After the mud had been
exposed for some time it was traversed by a single
carnosaur and by numerous ornithopods and
coelurosaurs (trackway groups C, D and E
above). The mud was exposed long enough to
achieve a firm plastic consistency, but not long
enough for desiccation cracks to appear. The
period of exposure would certainly have been a
matter of hours, if not of days or weeks.
Evidently the mud was not waterlogged at the
time it was traversed by the dinosaurs: none of the
thousands of footprints collapsed or slumped
after withdrawal of the track-maker’s foot. Nor
does the mud seem to have been very tenacious,
for there are very few instances in which it
adhered to a track-maker’s foot. In one of these
(footpriat No. 8 in Fig. 4) the mud adhering to the
underside of a single toe was drawn up into a
longitudinal crest; in another (PL 8, Fig. C) mud
adhered to one toe in the form of a ‘cusp’ or
‘bubble-like’ structure. However, we suspect that
in many cases the imprints of one or more digits
have been narrowed by suction created during
withdrawal of the track-maker’s foot. Overall it
seems that the mud may have had the consistency
of potter’s clay at the time it was traversed by the
dinosaurs.
CARNOSAUR TRACKWAY
Ichnogenus cf. Tyrannosauropus Haubold 1971
Eleven footprints at Lark Quarry are far bigger
than any others, and form a single trackway
extending from NE to SW (Figs 3 and 4, Pis 4 to
6). This trackway is attributed to a carnosaur — a
large bipedal predator of the dinosaur suborder
Theropoda (order Saurischia).
It was not feasible to collect any of the
carnosaur footprints, for to do so it would have
been necessary to destroy many other trackways.
In any case, the footprints do not show sufficient
detail to warrant their assignment to any new or
existing ichnospecies. Measurements of the
carnosaur trackway were taken directly from the
bedding plane at Lark Quarry and were checked
on fibreglass replicas (QM F10322) at the
Queensland Museum.
Description.
The trackway comprises deep, basin-like and
rather ‘messy’ footprints, often with poorly
defined margins. Evidently the track-maker’s feet
plunged right through the muddy surface layer
and churned up the underlying sandy sediment. In
most cases the impact of the foot caused sediment
to bulge up between the toes and behind the foot
to leave a prominent raised rim to the footprint
(PL 5, Fig. B). The sandy sediment in the floor of
the footprint is usually raised into a series of
irregular ripples. Two of the carnosaur footprints
have been illustrated elsewhere (Thulborn and
Wade 1979, fig. 2), and two more examples are
shown here (Pis 5 and 6). The following
description is a generalized one, based on
information from all better-preserved footprints
in the trackway.
Each footprint is tridactyl, with clear imprints
of digits 2, 3 and 4, but with no trace of the hallux
(digit I). The three digits are relatively short,
emerging from a large basin-like depression
representing a ‘sole’ or ‘pad’ to the foot; in
footprint No. 3, for example, the length of digit
3, as a free entity, represents only about A\% of
total footprint length (PL 5, Fig. A). The digits
are usually quite sharply defined, but often there
is no clear outline to the back of the foot, making
it difficult to obtain a measurement of total
footprint length (see, for example, PL 5). All the
better-preserved footprints are slightly narrower
420
MEMOIRS OF THE QUEENSLAND MUSEUM
than long, with footprint width equivalent to
some 85-9OV0 of footprint length. The digits are
broad and straight, and taper sharply to V-shaped
tips. In none of the footprints are there traces of
digital swellings or nodes that might indicate the
phalangeal formula. Digits 2 and 4 are distinctly
shorter than digit 3, and are roughly mirror-
images — being almost complementary in shape,
size and angle of divergence from digit 3. In one
of the best-preserved footprints (No. 3; PL 5, Fig.
A) the interdigital angles are about 33° (2-3) and
30° (3-4).
Mean measurements of footprint size, pace
length and stride length are as follows (each with
standard deviation and coefficient of variation):
Mean FL: 51.4 ± 6.5 cm (CV 13^o; N 7)
Mean FW: 46.1 ± 4.0 cm (CV 9»7o; N 10)
Mean PL: 166.6 ± 26.5 cm (CV 16%; N 10)
Mean SL: 330.6 ± 37.4 cm (CV 11%; N 9)
Mean ratio FW/FL: 0.88 ± 0.05 (CV 6%; N 7).
The near-symmetrical footprints are arranged
with slight positive rotation (i.e. they point not
only forwards, but slightly inwards). They form a
narrow trackway, with mean pace angulation
calculated at 170°47’ (SD 9°26’; CV 5.5% N 9).
The trackway has a slightly sinuous course from
NE to SW. From the spacing and orientation of
the first few footprints we suspect that the animal
actually approached the present Lark Quarry site
from the NNE; the orientation of the last (llth)
footprint indicates that the animal made an
abrupt right-hand turn and moved off to the NW
(see Fig. 3). There is no trace of a tail drag.
Some of the footprints show additional details
of interest. Footprint No. 7, for example, consists
of little more than shallow imprints of the three
digits (PL 6), and seems to have been formed on a
relatively resistant patch of sediment. In this
footprint there is evidence that the tips of the
digits extend forwards, beneath the surface of the
sediment, as conical tunnels about 4 cm in length.
These tunnels appear to be marks of long, robust
and sharply-pointed claws. Traces of similar
claws occur in several other footprints of the
carnosaur. In footprint No. 8 the central digit (3)
is unusually broad and contains a longitudinal
crest of mudstone in the midline (Fig. 4; see also
Thulborn and Wade 1979, fig. IB). This crest was
presumably formed by mud adhering to the
underside of the middle toe as the animal’s foot
was lifted from the substrate; other prints from
the same foot do not show this feature. Fig. 4
illustrates variation in shape of the carnosaur’s
footprints.
Status and Affinities.
The occurrence of a carnosaur trackway at
Lark Quarry is not unduly surprising. Carnosaurs
seem to have had an almost world-wide
distribution during the Cretaceous period, with
their footprints having been reported as far afield
as Spilzbergen (Edwards et al. 1978) and Western
Australia (Colbert and Merrilees 1968). No
skeletal remains of carnosaurs are recorded from
Queensland, though footprints of large theropod
dinosaurs are well known in the Jurassic rocks of
the state (Ball 1933, 1934a, 1934b, 1946;
Anonymous 1951, 1952a, 1952b; Staines 1954;
Bartholomai 1966). For the sake of convenience
we may distinguish two major groupings of
carnosaur footprints in general: those with
relatively long and slender toes, and those with
comparatively short and thick toes. Examples of
these tw'o groupings are, respectively,
Megalosauropus and Tyrannosauropus (see
Haubold 1971 and references cited therein). The
former are probably footprints of smaller and
more gracile carnosaurs, such as AUosaurus and
MegalosauruSy while the latter probably represent
bigger and more robust forms like
Tyrannosaurus. The footprints of the Lark
Quarry animal have rather short thick toes, and
they appear to be closer in appearance to
Tyrannosauropus than to any other form of
carnosaur footprint so far described. The Lark
Quarry footprints resemble Tyrannosauropus in
general shape and proportions (mean ratio
FW/FL of 0.88 as opposed to approximately 0.86
in Tyrannosauropus ), but they differ in the
following respects: in size (FL up to 80 cm in
Tyrannosauropus ), in pace angulation (170° as
opposed to approximately 150°), and in the ratio
SL/FL (6.4 as opposed to 5.0). On the basis of
these similarities and differences we recommend
that the Lark Quarry footprints should be
referred to as cf Tyrannosauropus. This
identification does not imply that the theropod
dinosaur Tyrannosaurus was responsible for the
Lark Quarry trackway; the track-maker can be
identified no more precisely than ‘carnosaur’.
It must be mentioned here that footprints of
carnosaurs have often been confused with those
of ornithopod dinosaurs (bipedal herbivores of
the suborder Ornithopoda, order Ornithischia).
The source of this confusion is partly historical:
the first footprints to be attributed to a particular
genus of dinosaur — ihe ornithopod Iguanodon
— happened to be large tridaclyl examples from
the Lower Cretaceous of Europe. Subsequently,
there grew a common tendency for any large
tridactyl footprints to be ascribed to Iguanodon
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
421
or to some similar ornithopod dinosaur (see
comments by Heckles 1862, Charig and Newman
1962, Sarjeant 1974). Such confusion has, in fact,
occurred over footprints from the Australian
Cretaceous: large tridactyl prints from the
Broome Sandstone (Lower Cretaceous) of
Western Australia were attributed to iguanodonts
by McWhae et al. (1958), but were later identified
as those of theropod dinosaurs (Colbert and
Merrilees 1968). Iguanodontids and some
theropods both produced large iridactyl
footprints which may, in some circumstances, be
difficult to distinguish — particularly if
preservation is poor. It has been suggested to us
(by Dr D.B. Norman, pers. comm.) that some
doubt may attach to the footprints of the Lark
Quarry animal, and that these might actually be
footprints of an iguanodontid ornithopod.
However, several distinctive features of the
footprints lead us to conclude that they are almost
certainly those of a carnosaur. First, the
footprints are slightly longer than broad, whereas
those of ornithopods are commonly broader than
long (see, for example, Langston 1960, Currie
and Sarjeant 1979, and the many ornithopod
footprints described below). Next, the three digits
have an almost symmetrical arrangement; in
many ornithopod footprints digit 2 is more widely
spaced from digit 3 than is digit 4 (see same
examples). Third, the Lark Quarry footprints
have traces of a large pointed claw on each toe;
the ungual phalanges of the larger ornithopods
were blunter, and sometimes rather hoof-like,
structures. Finally, the central digit (3) is V-
shaped in outline; in ornithopod footprints digit 3
tends to have roughly parallel margins that curve
round to form a U-shaped extremity. These basic
distinctions seem to confirm that the large
trackway at Lark Quarry is that of a carnosaur
(see Fig. 4).
ORNITHOPOD TRACKWAYS
Ichnogenus Wintonopus ichnogen. nov.
Type and only ichnospecies W. latomorum
ichnosp. nov.
HOLOTYPE: single right footprint, preserved as
natural mould; QM F103I9 (PI. 7, Fig. A).
REFERRED MATERIAL: QM FI0320 (single
left footprint, as natural mould; PI. 11, Fig. A);
QM F10321 (rock slab with footprints and
trackways as natural moulds); QM FI 2264 (single
right footprint, as natural cast; PI. 1, Figs. A, B);
QM F10322 (fibreglass replicas of footprints and
trackways preserved as natural moulds; PI. 8 to
10; PI. II, Figs. B, C, D; PI. 13, Fig. C; PL 14,
Fig. A; PL 16, Figs. B, C).
LOCALITIES: Lark Quarry (QM F10319, QM
F10320, QM F10321. QM F10322); Seymour
Quarry (QM F12264). See Fig. 1 for location of
quarries.
HORIZON: interbedded sandstones and
mudstones about the middle of the Winton
Formation; early Upper Cretaceous
(Cenomanian).
ETYMOLOGY: Ichnogenus name derived from
the name Winton and the Greek pous (foot);
ichnospecies name (from Latin lalomus, stone-
mason) as tribute to the many volunteers who
worked at the Lark Quarry excavation.
DIAGNOSIS (ichnogenus and ichnospecies):
narrow trackway of small to medium-size
digitigrade biped, with pace angulation about
160°. Footprint size index (SI) usually between
3.2 and 11.1 cm, but occasionally as high as 26.6
cm. No imprints of hand or tail. Footprints
tridactyl (digits 2, 3 and 4), slightly broader than
long (ratio FW/FL about 1.15), showing distinct
positive rotation. Digits broad, with rounded or
bluntly angular tips, without indications of
phalangeal pads. Digit 3 longest, with sub-parallel
sides. Digit 4 shorter and slightly narrower than
digit 3, extended as blunt posterior salient. Digits
3 and 4 close together, parallel or only slightly
divergent. Digit 2 shortest, and widely separated
from digit 3 (with interdigital angle often about
60°). Imprints of digits 2 and 3 sometimes
completely separated. Posterior margin of foot
convex forwards. Ratio PL/FL usually between
8.0 and 13.5, rarely as low as 4.0 or as high as
15.0; ratio SL/FL usually between 16.0 and 24.0,
rarely as low as 8.0 or as high as 27.0.
DESCRIPTION. The holotype is a sharply
defined footprint, probably formed by the foot
penetrating and leaving the substrate with
minimal disturbance (PL 7, Fig. A). Few of the
footprints referred to Wintonopus latomorum are
identical to the holotype: the majority have less
complete imprints of the digits, and many appear
to have been disfigured by withdrawal of the
track-maker’s foot from the sediment.
Nevertheless all these disfigured and less complete
examples can be interpreted as variants of the
footprint pattern exemplified by the holotype (see
Fig. 5). Variation in shape of the footprints is
described first; thereafter we describe variation in
size, proportions and spacing of the footprints.
All the footprints are tridactyl. They are often
several centimetres deep, yet none of them shows
any trace of digit 1. If this digit was present in the
422
MEMOIRS OF THE QUEENSLAND MUSEUM
track -maker’s foot it must have been quite short
and non-supportive, so that it failed to touch
down even when the three weight-bearing digits
sank quite deeply into the mud. Digit 1 was
probably no longer than it is in oniithopods such
as the Lower Cretaceous Hypsilophodon (Fig.
6B). The imprint of digit 4 extends farther back
than the imprints of digits 2 and 3, giving the
impression of a distinct salient or ‘spur’ at the
posterolateral corner of the footprint. This ‘spur’
is unlikely to indicate the presence of a functional
digit 5 (which is vestigial in even the earliest
ornithopod dinosaurs) and probably reflects that
the distal end of metatarsal 4 was located well to
the rear of the foot, in standard ornithopod
fashion (see, for example, the foot of
Fabrosaurus — Thulborn 1972, fig. 12B).
Nearly all the footprints are asymmetrical, with
a strongly divergent digit 2, so that isolated
examples are readily identified as left or right.
This method of identification has been verified in
at least 60 Wintonopus trackways. Only a few
footprints (in otherwise typical trackways) show
any close approach to a symmetrical arrangement
of the three digits. These near-symmetrical prints
could have been produced in any of several ways:
by some degree of spreading and/or closure of
digits in the foot; by partial flexion of the
divergent digit 2; by rotation of the foot around
the long axis of digit 3 (so that digit 2 was carried
laterally and slightly underneath digit 3); by the
foot meeting the substrate at an unusual oblique
angle (with the long axis of digit 3 directed
forwards, downwards and slightly outwards).
The imprints of the digits are usually quite
short and relatively broad. In most footprints the
three digits are about equally broad (as, for
example, in the holotype), but in a few cases digit
3 is much broader than digits 2 and 4 (e.g. PI. 9,
Fig. D; PI. 10, Fig. D). In these examples digit 3
seems to have borne most of the track -maker’s
weight, while the flanking digits splayed out to
form smaller and shallower imprints. A very
similar effect was described by Sternberg (1932) in
an ornithopod footprint {Gypsichnites pacensis )
from the Lower Cretaceous of British Columbia.
Digit 3 is longest, and in the least -disfigured
footprints digits 2 and 4 extend about equally far
forwards. The imprint of digit 3 is often straight,
but sometimes shows very slight curvature
(convex laterally; e.g. PI. 11, Fig. A; PL 15, Fig.
B). The hindmost margins of digits 2 and 3 lie on
a line approximately normal to the long axis of
digit 3, whereas digit 4 extends well behind this
line to form the posterior salient or ‘spur’. In
consequence the posterior margin of the footprint
(or the line connecting the hindmost points of the
three digital imprints) is arched forwards. In some
footprints, such as the holotype, the three digital
imprints are joined together posteriorly, and the
arched rear margin is continuous. Evidently these
footprints were formed by the foot sinking into
the mud up to, or beyond, the distal end of the
metatarsus. In many other examples the foot did
not sink so deeply, so that the three digital
imprints are partly or completely separated and
there is no continuous rear margin to the
footprint (e.g. Figs 5D, E, F; PL 8, fig, B). The
imprint of digit 2 is often completely separated
from that of digit 3, whereas the imprints of digits
3 and 4 are usually joined together (e.g. PL 8, Figs
A, B, D). This difference probably indicates that
digit 2 diverged from digit 3 higher up the
metatarsus than did digit 4 (see fool skeletons of
ornithopods, Fig. 6). There are no certain
indications of phalangeal pads in any of the
footprints. The tips of the digital imprints are
generally well-rounded in outline — except where
they have been extended forwards as scrape-
marks (see below) — but are sometimes a little
sharper in the smallest footprints (PL 11).
Interdigilal webbing is limited in extent; the
holotype shows traces of a small web between
digits 3 and 4, and less certain traces of another
between digits 2 and 3.
Wintonopus material from Seymour Quarry
comprises natural casts reinforced by superficial
infiltration of ironstone. The surfaces of the casts
are wrinkled and finely tuberculate, and
somewhat reminiscent of reptilian skin texture
(PI. 1). However, it is not certain that these
specimens do show preservation of skin texture:
an apparently identical texture is found on
footprint casts attributed to the coelurosaurs and
on some areas of seemingly undisturbed
sediment, and it may be no more than a by-
product of ironstone formation.
Practically every Wintonopus footprint at Lark
Quarry seems to have been disfigured to some
extent as the track-maker’s foot was withdrawn
from the mud. Basically, each foot sank quite
deeply into the mud, and as it was lifted clear at
the end of the stride the tips of one or more digits
tended to drag and scrape through the rim of the
newly-formed footprint. So, in many cases, there
are scrape-marks extending forwards from one or
more of the digital imprints. Digit 3 was longest in
the fool and, for that reason, tended to produce a
scrape-mark most frequently (e.g. PL 8, Fig. A).
Digit 4 was intermediate in length between digits 2
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
423
and 3, and was next most likely to produce a
scrape-mark, whereas digit 2 was shortest
(parallel to the long axis of digit 3) and rarely did
so (e.g. PL 10, Fig. D). In most cases the scrape-
marks are fairly short, but in a few examples they
are longer than the digital imprints (PL 8, Fig. A,
and PL 10, Fig. D). The scrape-marks are not
straightforward extensions of the digital imprints,
but veer away from them at a distinct angle.
Sections of trackways show that each foot was
planted into the mud with slight positive rotation
(i.e. with the toes pointing forwards and
inwards). But as the foot was lifted from the mud
it evidently turned to face directly ahead, so that
the tips of the toes swept forwards and slightly
outwards. In other words the foot was placed in
the mud at one angle and w^as withdrawn at a
different angle, and it is for this reason that the
digital imprints have a different orientation from
their scrape-marks. In some footprints this effect
is so marked that the tip of digit 3 appears to be
forked or Y-shaped (e.g. PL 8, Fig. D, and, to a
lesser extent, in the holotype). In such examples
the medial branch of the fork was formed when
the digit was placed into the mud, and the lateral
branch is a scrape-mark produced when the digit
was withdrawn in a different direction. An
exactly comparable type of scrape-mark was
illustrated by Sarjeant (1970, Fig. 5e) in an
ornithopod footprint (? Satap/iasaurus cf. S.
dsocenidzei ) from the Middle Jurassic of
Yorkshire, England.
Clear examples of backwardly-directed scrape-
marks are less common. Again, the longest digit
(3) seems to have produced a scrape-mark most
frequently whereas the shortest digit (2) rarely
produced one. These scrape-marks are also
aligned at a slight angle to the long axes of the
digital imprints (e.g. PL 8, Fig. B), confirming
that the tips of the digits swung laterally as the
foot was lifted from the substrate.
The development of these scrape-marks
(whether forwards or backwards) is best
understood in relation to the sequence of events
during the track-maker’s stride cycle (Fig. 7). At
the start of this cycle the forwardly-extended foot
would have been planted into the sediment with
slight positive rotation (Stage 1 in Fig. 7). The
initial footprint would have been quite shallow.
At mid-stride the track-maker’s centre of gravity
would have passed forwards above the foot,
which would then have sunk deeper into the
substrate (Stage 2 in Fig. 7). In many instances the
foot also slipped backwards a little, so that the
front margins of the footprint are distinctly
‘stepped’ or ‘terraced’ (see, for example, PL 8,
Figs. A, B; PL 9, Fig. D). Shortly thereafter the
foot began to rotate (so that the long axis of digit
3 was directed straight ahead), and the rear part
of the fool started to lift clear of the substrate.
Sometimes the toes continued to slip backwards
as they were lifted from the footprint: in these
cases the toe-tips incised deep slots in the floor of
the footprint (Stage 3A in Fig. 7) or even
breached the rear wall of the footprint to leave
backwardly-directed scrape-marks (Stage 3B).
More commonly there was limited back-slip of
the toes (Stages 1 to 2) and the toes-tips dragged
through the front wall of the footprint to produce
forwardly-directed scrape marks (Stage 3C).
Wintonopus footprints are typically broader
than long, even though many examples have their
total length exaggerated by scrape-marks. In
some cases the track-maker’s foot was planted
into the mud at a steep angle, to leave relatively
short and stubby imprints of the toes (e.g. PL 8,
Fig. B). In other cases the foot seems to have lost
its purchase in the muddy substrate, and the toes
slithered back to form deep scratches that
exaggerate the total length of the footprint (e.g.
PL 9, Fig. B). In still other cases only the distal
parts of the toes entered the mud, and then
skidded backwards to produce a footprint
consisting of little more than three long scratch-
marks (e.g. PL 10, Fig. A). A few footprints
consist of three puncture-marks apparently
formed by the toes entering and leaving the mud
almost vertically (e.g. PL 15, Fig. C).
Measurements of footprints, paces and strides
were taken from parts of 57 different trackways
of Wintonopus (see ‘Methods’ for descriptions of
measurements). Fifty-six of these trackways are
on the Lark Quarry bedding plane; the other
section of trackway is at a different site — New
Quarry. The 56 trackway sections at Lark Quarry
comprise 284 footprints, representing 228 paces
and 172 strides. This sample provides the
following mean figures for dimensions of
footprints, paces and strides:
Overall means
Mean FL: 6.71 ± 3.39 cm (CV 51*^0; N 200)
Mean FW: 7.58 ± 4.51 cm (CV 60'^^o; N 214)
Mean PL: 68.3 ± 32.2 cm (CV 47%; N 215)
Mean SL: 131.7 ± 63.4 cm (CV48%;N162)
The high coefficients of variation reflect
considerable ranges in size. Nearly all the
trackways are those of small animals, with
footprint lengths between 2 cm and 16 cm, and
stride lengths in the range 49-271 cm; but the size
424
MEMOIRS OF THE QUEENSLAND MUSEUM
distribution, as a whole, is attenuated by the
presence of a single very large trackway with
footprints up to 33 cm long and stride lengths
reaching 345 cm (trackway ‘B’ in Fig. 3; see Figs 8
and 10).
The index of footprint size, the pace angulation
and various ratios were calculated from the basic
measurements listed above (see ‘Methods’); they
have the following means:
Overall means
Mean SI: 7.20 ± 3.93 (CV 55*70; N 173)
Mean ANG: 161 °24’ ± 11°20’ (CV 1% N
145)
Mean ratio FW/FL: 1.15 ± 0.25 (CV 22*7o; N
173)
*Mean ratio PL/FL: 10.18 ± 2,00 (CV 20%; N
120 )
’'Mean ratio SL/FL: 19.84 ± 3.69 (CV 19%; N
88 )
(* FL is mean for two footprints defining each
pace or stride)
Footprint proportions (FW/FL) and pace
angulation appear to show relatively little
variation, and are probably good diagnostic
characters; that is, Wintonopus trackways are
characterized by being narrow, and fairly
straight, and by having footprints that are usually
broader than long.
There is a strong positive correlation between
any two measurements of size in the Wintonopus
trackways and footprints; for example:
product moment
correlation coefficients
untransformed log transformed
variables
data
data
FL
: FW (N 174)
0.87
0.89
*FL
: PL (N 112)
0.86
0.87
*FL
: SL (N 79)
0.87
0.89
*FW
; SL (N 89)
0.89
0.93
*SI :
SL (N 59)
0.90
0.92
(*mean for two footprints defining each pace or
stride)
The correlations between stride length and
footprint dimensions (FL, FW or SI) are worthy
of note. It is only to be expected that bigger
animals would take bigger strides, but stride
length varies according to the gait and speed of an
animal — and not simply to its size alone. The
impressive correlations between foot size and
stride length imply that the Wintonopus track-
makers at Lark Quarry were all using a similar
gait; in a random sample of dinosaur trackways
one might expect to find a somewhat looser
correlation between stride length and footprint
dimensions.
The correlation between footprint size (SI) and
footprint proportions (ratio FW/FL) is much
poorer: 0.22, with untransformed data, N 172.
So, too, is that between footprint length and pace
angulation (0.23, with untransformed data, N
73). These poor correlations seem to confirm our
observation that footprint proportions and pace
angulation tend to remain fairly constant
throughout the entire size range of Wintonopus
trackways (see further discussion below).
All the preceding estimates and statistics are
based on pooled data from the Wintonopus
trackways (i.e. on every available example of the
284 footprints and their paces and strides). If the
data are grouped, and mean figures are taken for
each of the 56 trackways studied, there emerges a
somewhat similar pattern of size distribution and
correlations (see Figs 9 and 10). Means based on
the grouped data may be summarized as follows:
Means per trackway
Mean FL: 6.64 ± 3.07 cm (CV 46%; N 56)
Mean FW: 7.55 ± 4.36 cm (CV 58%; N 56)
Mean PL: 67.1 ± 30.6 cm (CV 46%; N 55)
Mean SL: 128.0 ± 59.6 cm (CV 47%; N 52)
Mean ST: 7.05 ± 3.58 cm (CV 51%; N 56)
Mean ANG: 162^^47’ ± 8°46’ (CV 5%; N 51)
Mean ratio FW/FL: 1.14 ± 0.19 (CV 18%; N 56)
Mean ratio PL/FL: 10.18 ± 1.97 (CV 19%; N
49)
Mean ratio SL/FL: 19.75 ± 3.50 (CV 18%; N
44)
Most coefficients of variation remain very high.
An analysis of variance (see Sokal and Rohlf
1969, p. 204 et seq. ) will reveal how much of this
variation lies within the Wintonopus trackways:
variation
variation
within
among
trackways trackways
variable
(%)
(%)
FL
15.1
84.9
FW
6.0
94.0
PL
7.5
92.5
SL
2.4
97.6
SI
0.8
99.2
ratio FW/FL
52.1
47.9
ANG
77.9
22.1
Evidently footprint dimensions, pace
length and
stride length remain
fairly constant
within the
Wintonopus trackways. Footprint
length is
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
425
somewhat more variable than footprint width
within the trackways, but the index of footprint
size is virtually constant. (Footprint length is
more variable than footprint width because it is
more strongly affected by the angle at which the
foot enters and leaves the substrate, and by the
development of scrape-marks; see Fig. 5). Stride
length appears to be remarkably consistent within
trackways, though pace length is more variable.
Footprint proportions (ratio FW/FL) and pace
angulation appear to vary more within trackways
than they do between trackways; but these two
features show little overall variation in the first
place — so they may still be regarded as good
diagnostic characters. In general, most of the
variation in footprint dimensions, paces and
strides can be attributed to the difference in size
between one trackway and another.
STATUS AND AFFINITIES. The footprints
referred to Wintonopus latomorum vary a good
deal in size and in appearance, yet they all show
some at least of the following diagnostic
characters: a widely spaced or divergent digit 2, a
backwardly-projecting ‘spur^ behind digit 4, a
forwardly-arched rear margin, and a width that
equals or exceeds footprint length. Where the
footprints can be connected into trackways they
are arranged with distinct positive rotation, and
the trackways are always narrow and rather
straight (with pace angulation about 160°).
Moreover, footprint dimensions are strongly
correlated one with another, and with stride
length. In short, all the footprints may be
regarded as those of animals sharing one
distinctive pattern of foot structure (see Fig. 6C)
and using the same gait. For these reasons it
seems justifiable to assemble all these footprints
in a single ichnospecies. Differences in shape
between one footprint and another appear to be
no more than circumstantial (see preceding
descriptions and Fig. 5). According to recent
recommendations for the nomenclature of trace
fossils (see Article 40 in Basan 1979) it might be
legitimate to define several ichnospecies of
Wintonopus on the basis of footprint shape alone
— e.g. a ‘scratchy-toed’ species, a ‘stubby-toed’
species, and so on. In the present circumstances,
where footprint shape varies within a single
trackway, such a measure would be rather
confusing. Moreover there is no clear evidence
that Wintonopus footprints of different
morphology were ‘produced in different phases
of behavior’ on the part of the track-maker (as
Basan’s Article 40 seems to require). In addition it
may be noted that the range of variation in
Wintonopus is no greater than that in some
existing ichnotaxa — e.g. the ichnogenus
Anomoepus (as defined by Lull 1953), and the
ichnospecies Grallator variabilis and G. olonensis
(as defined by de Lapparent and Montenat 1967).
The makers of the Wintonopus trackways were
almost certainly dinosaurs of the suborder
Ornithopoda (bipedal herbivores of the order
Ornithischia). Ornithopods had a world-wide
distribution during the Mesozoic era: their
skeletal remains and their footprints have been
reported from every continent except Antarctica.
The following features of Wintonopus
latomorum seem to be characteristic of very many
ornithopod footprints: the footprint is tridactyl,
and its width rivals or exceeds its length; the
digital imprints are relatively short, thick and
blunt (indicating the presence of ‘hoof-like’
unguals rather than sharp claws); the space
between digits 2 and 3 is distinctly greater than
that betw^een digits 3 and 4; the outer margins of
digits 2 and 4 diverge only slightly from the
longitudinal axis of digit 3, so that the footprint
has sub-parallel sides; there are sometimes traces
of small interdigital webs. In all these features W.
latomorum resembles other footprints attributed
to ornithopod dinosaurs — e.g. Amblydactylus
ichnospp. from the Lower Cretaceous of Canada
(Sternberg 1932, Currie and Sarjeant 1979);
unnamed types from the Late Jurassic/Early
Cretaceous of Mexico (Ferrusquia-Villafranca et
al. 1979); footprints of Iguanodon, from the
Lower Cretaceous of Europe (Beckles 1862, Dollo
1906). However, the footprints described here are
distinctly smaller than many others attributed to
ornithopod dinosaurs (see Table 1). Of the 57
Wintonopus trackways examined in this study
only two have mean SI greater than 12 cm
(actually 12.7 cm and 26.6 cm); among other
footprints attributed to ornithopods only those of
Anomoepus ichnospp. are commonly found to be
so small. Aside from this Wintonopus differs
from most other ornithopod footprints in one
other respect — in the absence (or weak
development) of an imprint representing a ‘sole’
or ‘heel’ to the foot. The rear margin of the
footprint is concave (arched forwards) rather
than convex (arched backwards) and presumably
corresponds to the natural arch formed by the
distal ends of the metatarsals. This distinctive
footprint shape seems to indicate that the
Wintonopus track-makers were thoroughly
digitigrade, whether they were walking or running
(see later discussion of speeds and gaits). These
differences in size and shape are sufficient to
426
MEMOIRS OF THE QUEENSLAND MUSEUM
distinguish Wintonopus from most other tracks
attributed to ornithopod dinosaurs. Anomoepus
ichnospp. are comparable in size to Wintonopus,
but are distinguished by narrower and more
acutely pointed digits with obvious phalangeal
nodes (see Lull 1953). In addition most examples
of Anomoepus have the ratio PL/FL much lower
than it is in Wintonopus (Fig. 11).
However, two types of footprint described by
de Lapparent and Montenat (1967) from the
Rhaeto-Liassic of Vendee (W France) bear some
definite resemblances to Wintonopus. One of
these types, Anatopus palmatus, was also
attributed to an ornithopod dinosaur but is,
unfortunately, represented by only three isolated
footprints. In all three cases footprint size index is
about 9.0 cm — well within the range described
for Wintonopus; the ratio FW/FL is about 1.14
— practically identical to the mean for
Wintonopus (1.15). Anatopus appears to
resemble Wintonopus not only in size, shape and
arrangement of the three digits, but also in
possessing what seem to be anterolaterally
directed scrape-marks at the tips of digits 3 and 4
(see de Lapparent and Montenat 1967, fig. 16, but
note the different identification of digits).
Nevertheless Anatopus certainly differs from
Wintonopus in that the digits are relatively
narrow and show distinct outlines of phalangeal
pads. Moreover, de Lapparent and Montenat
identified traces of very extensive interdigital
webbing in the type specimen of Anatopus. A
second type of footprint, Saltopoides iga/ensis,
was attributed to a Iheropod dinosaur but is, once
again, rather similar to Wintonopus (see de
Lapparent and Montenat 1967, Fig. 15).
TABLE 1: A Comparison of Size Among
Footprints Attributed to Ornithopod
Dinosaurs.
mean index of
footprint size
(cm)
68.5 ‘Ornithopoda’, Jurassic of Brazil
(Leonard! 1980)
61.4 Amblydactylus gethingi, L Cretaceous of
Canada (Sternberg (1932)
51.3 Iguanodon, L Cretaceous of Portugal
(Antunes 1976)
46.3 Irenesauripus acutus, L Cretaceous of
Canada (Sternberg 1932)
28.6 Gypsichnites pacensis, L Cretaceous of
Canada (Sternberg 1932)
24.7 ‘Ornithopod morphotypes’,
Jurassic/Cretaceous of Mexico
(Ferrusquia-Villafranca et al. 1978)
23.5 Satapliasaurus cf 5. dsocenidzei, M
Jurassic of England (Sarjeant 1970)
18.5 Satapliasaurus dsocenidzei, L Cretaceous
of Georgia, USSR (Gabouniya 1951)
18.3 Amblydactylus kortmeyeri, L Cretaceous
of Canada (Currie and Sarjeant 1979)
17.6 Iguanodon, U Jurassic of England (Delair
and Brown 1974)
17.5 ?cf Satapliasaurus, M Jurassic of England
(Sarjeant 1970)
15.7 Irenichnites gracilis, L Cretaceous of
Canada (Sternberg 1932)
15.6 Sauropus barrottii \
15.0 Anomoepus crassus f Triassic of
10.7 Anomoepus isodactylus ) Connecticut
I (Lull 1953)
9.2 Anomoepus intermedius ]
9.0 Anatopus palmatus, Rhaeto-Liassic of
France (de Lapparent and Montenat 1967)
8.7 Anomoepus curvatus, Triassic of
Connecticut (Lull 1953)
7.7 Anomoepus scambus, Triassic of
Connecticut (Lull 1953)
7.2 Wintonopus latomorum
6.2 Anomoepus gracillimus, Triassic of
Connecticut (Lull 1953)
5.4 Anomoepus minimus, Triassic of
Connecticut (Lull 1953)
Saltopoides has a footprint size index about 13.4
cm, but the footprints differ from those of
Wintonopus in being distinctly longer than wide
(FW/FL ratio about 0.75). In addition the lateral
and media! margins of the footprints are
divergent (rather than parallel as in Wintonopus),
and there is no very marked positive rotation of
the footprints. Saltopoides also differs in showing
faint indications of phalangeal pads, but in two
other respects it is very like Wintonopus — in
having high values for pace angulation (almost
180°) and for the ratio PL/FL (11.1). In
summary, Wintonopus is similar to both
Anatopus and Saltopoides in some features, but
in neither case is there an exact correspondence in
footprint morphology.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
427
COELUROSAUR TRACKWAYS
Ichnogenus Skartopus ichnogen. nov.
Type and only ichnospecies S. australis ichnosp.
nov.
HOLOTYPE: single left footprint, preserved as
natural mould; QM F10330 (PI. 7, Figs. B, C).
REFERRED MATERIAL: QM F10321 (rock
slab with footprints and trackways as natural
moulds); QM FI 2265 (single right footprint, as
natural cast; PI. 1, Figs. C, D); QM F10322
(fibreglass replicas of footprints and trackways
preserved as natural moulds; PI. 10, Figs B, D;
PI. 12; PI. 13, Figs A, B; PI. 14; PI. 15, Fig. A; PI.
16).
LOCALITIES: Lark Quarry (QM F10330, QM
FI 0321, QM FI 0322); Seymour Quarry (QM
FI2265). See Fig. 1 for location of quarries.
HORIZON: interbedded sandstones and
mudstones about the middle of the Winton
Formation; early Upper Cretaceous
(Cenomanian).
ETYMOLOGY: Ichnogenus name derived from
Greek skartes (nimble) and pous (foot);
ichnospecies name refers to southern (Australian)
provenance.
DIAGNOSIS (ichnogenus and ichnospecies):
trackway of small digitigrade biped, with pace
angulation about 150®. Footprint size index (SI)
between 2.9 and 5.7 cm. No imprints of hand or
tail. Footprints tridactyl (digits 2, 3 and 4),
slightly longer than broad (ratio FW/FL about
0.95) showing distinct positive rotation. Digit
imprints narrow, straight and sharply pointed,
without indications of phalangeal pads. Digit 3
longest; digits 2 and 4 about equal in length, and
almost equally divergent from digit 3 (both
interdigital angles betw^een 25® and 30®). Imprint
of digit 4 extends slightly farther back than
imprint of digit 2, but does not form a posterior
salient or ‘spur’. Traces of small interdigital webs
sometimes present. Posterior margin of footprint
is an oblique line (posterolateral to anteromedial),
either straight or arched forwards, in some
examples there is an imprint of the metapodium:
this is sub-rectangular in outline and roughly
equivalent in length to digit 3. Ratio PL/FL
usually between 5.5 and 8.5, rarely as low as 5.2
or as high as 9.1; ratio SL/FL usually between
11.0 and 16.0, rarely as low as 10.6 or as high as
17.3.
DESCRIPTION: The holotype is a well-defined
footprint, impressed in the substrate with
minimal disturbance. In general the footprints
identified as Skartopus show much less variation
in shape than do those referred to Wintonopus.
Once again, however, all the variations that do
exist may be interpreted as circumstantial
modifications of the footprint pattern shown by
the holotype. Variation in footprint shape is
described first; thereafter we describe variation in
size, proportions and spacing of the footprints.
All footprints referred to Skartopus are
tridactyl, with clear traces of digits 2, 3 and 4. The
footprints sometimes reach a depth of 2 cm, or
more, but none of them shows any certain trace
of the hallux (digit 1). If the hallux was present in
the track-maker’s foot it must have been
relatively short and without a major supportive
role; presumably it extended no farther than the
line of the metatarso-phalangeal contacts in digits
2, 3 and 4 (see Figs 6D-F). Where the three digits
are deeply impressed, as in the holotype, it may be
seen that their rear ends do not fall on a straight
line. Digits 2 and 4 extend slightly farther back
than digit 3, to form a curve (convex forwards)
that presumably reflects the natural arch formed
by the distal ends of the metatarsals.
Skartopus footprints are almost bilaterally
symmetrical. It is sometimes difficult to identify
isolated examples as left or right, but the
following features are often useful guides: the
space between digits 2 and 3 is slightly greater
than that between 3 and 4; digit 2 is slightly more
divergent than digit 4; scratch-marks, often
present at the tips of the digit imprints, extends
forwards and laterally. All these features are well
shown in the holotype (PL 7, Figs B, C). Where
the footprints can be connected into sections of
trackway they are easily identified as left and
right on account of their positive rotation (see PL
14, Fig. B).
The imprints of the digits are straight, and
relatively long and narrow (by comparison with
those of Wintonopus ). In all cases the three digit
imprints are about equally broad. Digits 2 and 4
are roughly equal in length, and digit 3 is longer.
In nearly all examples the tips of the digit imprints
are quite sharply pointed. The interdigital angles
are small, and in some footprints digits 3 and 4
are sub-parallel (e.g. the holotype). There are no
definite indications of phalangeal pads in any of
the footprints. The holotype shows traces of small
interdigilal webs, as do several other footprints in
the referred material (e.g. PI. 12, Fig. D).
Skartopus material from Seymour Quarry
comprises natural casts with a finely wrinkled
ironstone surface; it is not certain if this wrinkling
is a representation of original skin texture (see PL
1 and p. 422).
428
MEMOIRS OF THE QUEENSLAND MUSEUM
Variation in footprint shape is less marked in
Skartopus than in Wintonopus. Divarication of
the digits is slightly more pronounced in some
footprints than in others, but in all cases the digits
form a near-symmetrical pattern. In most
Skartopus footprints the digits terminate in sharp
scratch-marks; apparently similar marks have
been illustrated (though not described as such) in
coelurosaur footprints from the mid-Cretaceous
of Israel (Avnimelech 1966, pi. 7, fig. 2).
Variation in the shape of Skartopus footprints is
most easily explained by reference to events
during the track-maker’s stride cycle (Fig. 12). At
the start of this cycle the forwardly-extended foot
would have been planted on the sediment, and
there would have been a very shallow initial
footprint, or no footprint ar all (Stage I in Fig.
12). At mid-stride the track-maker’s centre of
gravity passed forwards above the foot, which in
some cases sank into the substrate (Stage 2A in
Fig. 12). Later, as the rear part of the foot started
to lift clear of the sediment, the claws pressed
down and slightly backwards to produce sharp
outlines to the tips of the digital imprints (Stage
3A in Fig. 12). At this point the toes sometimes
started to slip backwards, their claws incising
grooves in the floor of the footprint (Stage 4A in
Fig. 12). This sequence of events produced sharp-
toed tridactyl footprints such as the holotype (PL
7, Figs. B, C), some of which are secondarily
deepened by backwardly-directed scratch-marks
(e.g. PL 12, Figs. C, D; PL 13, Fig. A). However,
in many other cases the foot did not sink into the
substrate at mid-stride (Stage 2B in Fig. 12). To
judge from the number of discontinuities (or
‘missing’ footprints) in Skartopus trackways this
seems to have been a very frequent occurrence.
There are at least two obvious reasons why the
feet of Skartopus track-makers did not always
leave recognizable footprints. First, the track-
makers seem to have been remarkably small, and
presumably light, dinosaurs (see Fig. 15). Second,
it appears that coelurosaurs have bigger feet
(relative to hip height) than many other bipedal
dinosaurs (see later discussion concerning sizes of
track-makers). It seems, then, as if the Skartopus
track-makers may have been lightweight
dinosaurs with large spreading feet that acted as
analogues of snow-shoes. Even if the entire foot
did not sink into the substrate the tips of the toes
sometimes left imprints as the track-maker
‘kicked off’ at the end of its stride (Stage 3B in
Fig. 12). The toes then slithered back through the
mud to leave a series of curved parallel scratches
(Stages 4B and 5B in Fig. 12). In some cases only
one or two of the toes left such traces (e.g. PL 16,
Fig. A).
A few Skartopus footprints are noteworthy in
that they appear to include an imprint of the
metapodium (e.g. PL 12, Figs A, B; PL 13, Fig.
B). In these examples the imprint of the
metapodium is a large sub-rectangular depression
behind the three digit imprints. The metapodium
imprint is no wider than the maximum spread of
the digits, and it is roughly as long as the imprint
of digit 3; it is widest at the rear, where it is
broadly rounded in outline (convex backwards).
Footprints with such traces of the metapodium
are uncommon, and most of them occur singly
and at random in the Skartopus trackways.
However, one short section of trackway (a
sequence of 3 paces) is composed entirely of such
footprints (PL 14, Fig. B).
Measurements of footprints, paces and strides
were taken from parts of 34 trackways of
Skartopus australis (see ‘Methods’ for description
of the measurements). All 34 trackways are on the
Lark Quarry bedding plane, and they comprise a
total of 191 footprints (representing 157 paces
and 123 strides). This sample provides the
following mean figures for measurements of
footprints, paces and strides:
Overall means
Mean FL: 4.46 ± 0.70 cm (CV 16*70; N 131)
Mean FW: 4.10 ± 0.56 cm (CV 14*7o; N 158)
Mean PL: 32.1 ± 4.0 cm (CV 12<7o; N 151)
Mean SL: 61.7 ± 7.8 cm (CV 13*70; N 122)
The coefficients of variation are considerably
lower than those for equivalent measurements in
Wintonopus — in consequence of the much
smaller size range in Skartopus. The index of
footprint size, the pace angulation, and standard
ratios were calculated from the basic
measurements listed above (see ‘Methods’); they
have the following means:
Overall means
Mean SI: 4.29 ± 0.52 cm (CV 12*70; N 126)
Mean ANG: 152°38’ ± ir44’ (CV 8*7o; N
112 )
Mean ratio FW/FL: 0.94 ± 0.16 (CV 17*70; N
126)
*Mean ratio PL/FL: 7.27 ± 1.33 (CV 18*7o; N
97)
*Mean ratio SL/FL: 14.03 ± 2.26 (CV 16*70; N
85)
(* FL is mean for two footprints defining each
pace or stride)
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
429
Pace angulation appears to show relatively little
variation, and is probably a good diagnostic
character. In general terms pace angulation is
about 10° greater in Wintonopus than in
Skartopus — so that trackways of the latter tend
to be slightly broader and to have a more obvious
zig-zag arrangement of the footprints. In addition
the footprints of Skartopus are commonly longer
than broad whereas the reverse is true in
Wintonopus.
There is generally a poor correlation between
any two measurements of size in the Skartopus
material; for example:
product moment
correlation coefficients
untransformed log transformed
variables
data
data
FL :
: FW (N 131)
0.36
0.34
*FL :
: PL (N 70)
0.04
0.16
*FL
: SL (N 64)
0.07
0.07
*FW
: SL (N 88)
0.23
0.20
*SI :
SL (N 57)
0.15
0.10
(*mean for two footprints defining each pace or
stride)
The correlations are not improved by
transformation of the data. These poor
correlations may be attributed, once again, to the
limited size range of the footprints and trackways
(see further discussion below). The correlation
between footprint size index (SI) and footprint
proportions (ratio FW/FL) is also poor (-0.19,
with untransformed data; N 126), as is that
between footprint length and pace angulation
(-0.18, with untransformed data; N 61). The
significance of these poor correlations will be
examined later (p. 430).
A somewhat similar pattern of size distribution
and correlations emerges if the data are grouped
and mean figures are taken for each of the 34
Skartopus trackways (see Figs 14 and 15). Means
derived from the grouped data may be
summarized as follows:
Means per trackway
Mean FL: 4.46 ± 0.50 cm (CV ll^^o; N 34)
Mean FW: 4.14 ± 0.49 cm (CV 12%; N 34)
Mean PL: 32.0 ± 2.6 cm (CV 8%; N 34)
Mean SL: 61.8 ± 4.8 cm (CV 8%; N 34)
Mean SI; 4.28 ± 0.43 cm (CV 10%;, N 34).
Mean ANG; 153°37’ ± 9°35^ (CV 6%; N 34)
Mean ratio FW/FL: 0.94 ± 0.12 (CV 13%; N
34)
Mean ratio PL/FL: 7.19 ± 1.08 (CV 15%; N
29)
Mean ratio SL/FL: 13.73 ± 1.73 (CV 13%; N
32)
Analysis of variance (below) reveals that there is
as much, or more, variation within trackways as
there is among trackways;
variation
variation
within
among
trackways
trackways
variable
(%)
(%)
FL
57.9
42.1
FW
46.2
53.8
PL
79.9
20.1
SL
50.6
49.4
SI
52.1
47.9
ratio FW/FL
74.5
25.5
ANG
49.9
50.1
However, all these characters show little overall
variation in the first place (see coefficients of
variation) — so that most, if not all, of them may
still be regarded as of diagnostic value.
STATUS AND AFFINITIES. The footprints
designated Skartopus australis do not vary a great
deal in size or in shape. They consistently show
the following distinctive features: a near-
symmetrical arrangement of three long, relatively
narrow and sharply pointed digits, a forwardly
arched rear margin (except where there is an
imprint of the metapodium), and a length that
equals or exceeds footprint width. Where the
footprints can be connected into trackways they
are found to be disposed with distinct positive
rotation; the trackways are moderately broad,
with pace angulation about 150°. By comparison
the trackways of Wintonopus appear to be
narrower, with pace angulation about 160°.
Variation in shape of the Skartopus footprints
appears to be circumstantial — the occasional
appearance, in otherwise normal trackways, of
footprints represented only by scratches or of
footprints including a trace of the metapodium.
The scratch-like footprints were probably formed
when the track-maker’s foot slipped backwards
across the surface of the muddy substrate (Stages
2B to 5B in Fig. 12); footprints with a trace of the
metapodium were presumably formed when the
track-maker inadvertently came down ‘flat-
footed’, or perhaps when the foot sank deeply in
the mud. (Note, however, that one short section
of trackway consists entirely of footprints with
traces of the metapodium (PI. 14, Fig. B). This
sequence of footprints could be fortuitous, or it
could derive from any of several factors — e.g. a
pathological condition of the track-maker, or an
accumulation of mud on the animal’s feet.)
There is limited variation in size of the
Skartopus footprints, paces and strides. The
biggest footprint is less than twice the size of the
430
MEMOIRS OF THE QUEENSLAND MUSEUM
smallest (in terms of SI); by contrast the biggest
example of Wintonopus is nearly 12 times the size
of the smallest. The coefficients of variation
indicate that dimensions of footprints, paces and
strides vary much less in Skartopus than they do
in Wintonopus — yet the correlation between any
two of these dimensions is in most cases very
much poorer in Skartopus (compare Figs 10 and
15). These poor correlations do not necessarily
indicate that the Skartopus material is a
heterogeneous assortment of footprints and
trackways: rather, they reflect the very limited
range in size. For comparative purposes the entire
sample of Skartopus tracks might be regarded as
equivalent to a small size class selected from the
Wintonopus sample. A very similar relationship
between two ichnotaxa has been well illustrated
by de Lapparent zmd Montenat (1967, fig. 8). In
other words the Skartopus footprints and
trackways are all roughly similar in their
dimensions, so that there is as much (or more)
variation within a trackway as there is between
one trackway and the others (see analysis of
variance). In addition it must be noted that the
Skartopus footprints and trackways are, on the
whole, much smaller than the Wintonopus
footprints and trackways — yet both have been
measured within the same limits of error. Small
measurement errors would be of negligible
importance in the large Wintonopus tracks, but
they would certainly tend to blur correlations in
the absolutely smaller Skartopus tracks. Overall
the Skartopus footprints are quite consistent in
size, shape, and their spacing within trackways;
for these reasons it seems justifiable to assemble
them in a single ichnospecies. The range of
variation seen in this assemblage is no greater
than that in several other ichnospecies attributed
to coelurosaurs (e.g. Grallator olonensis and G.
variabilis, de Lapparent and Montenat 1967),
The Skartopus footprints were very probably
made by coelurosaurs — small representatives of
the dinosaur suborder Theropoda. Skeletal
remains and footprints of theropods have been
recorded from every continent except Antarctica;
theropod body fossils have not yet been reported
from Queensland, though their footprints are well
known in the state (for references see p. 420). In
their size and general appearance the Skartopus
footprints are comparable with those attributed
elsewhere to coelurosaurs (see review by
Haubold, 1971). The examples listed in Table 2
will illustrate the basic agreement in size. All these
examples (including Skartopus) share the
following similarities: the imprints of digits 2, 3
and 4 are rather narrow and quite sharply
pointed; the digits diverge (usually) at low angles,
and often have a near-symmetrical arrangement.
However, Skartopus differs from nearly all other
trackways attributed to coelurosaurs in having
exceptionally high values for the ratios PL/FL
and SL/FL (Fig. 16). Skartopusls also distinctive
in its footprint morphology. It differs from
Columbosauripus ichnospp. in the lesser
divarication of the digits; in addition the digital
TABLE 2: A Comparison of Size Among Footprints
Attributed to Small Theropod
Dinosaurs.
mean index of
footprint size
(cm)
24.3 Anchisauripus minusculus, Triassic of
Connecticut (Lull 1953)
14.1 Grallator formosus, Triassic of
Connecticut (Lull 1953)
12.2 Columbosauripus (2 ichnospp.),
Cretaceous of Canada and Algeria (both
ichnospp. illustrated by Haubold 1971,
a.v.)
10.2 CoelurosQurichnus (5 ichnospp.), Triassic
of Europe (all ichnospp. illustrated by
Haubold 1971, q.v.)
8.1 Grallator ci G. variahiliSy Triassic of
Algeria (Bassoullet 1971)
7.5 Anchisauripus hitchcocki, Triassic of
Connecticut (Lull 1953)
5.8 Otouphepus magnificus, Triassic of
Connecticut (Lull 1953)
4.3 Skartopus australis
3.8 Plesiornis pilulatuSy Triassic of Connecticut
(Lull 1953)
3.7 Wildeichnus navesi, Jurassic of Argentina
(Casamiquela 1964)
3.4 Grallator graciliSy Triassic of Connecticut
(Lull 1953)
2.5 Stenonyx lateralis , Triassic of Connecticut
(Lull 1953)
0.4 Coelurosaurichnus ichnosp., Triassic of
England (Wills and Sarjeant 1970)
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
431
imprints of Columbosauripus tend to be broader
than those of Skartopus, and digit 4 is often
considerably longer than digit 2 (see Haubold
1971, fig. 47). Coelurosaurichnus ichnospp. are
distinguished from Skartopus by very distinct
imprints of claws and phalangeal pads, by the
common preponderance of digit 4 over digit 2,
and (in many cases) by having digit 3 noticeably
broader than the flanking digits. In a few
instances, however, Coelurosaurichnus does
resemble Skartopus in having the rear margin of
the footprint arched to the front. Similarly
Grallator ichnospp. differ from Skartopus in
possessing well developed phalangeal pads, traces
of acuminate claws and, very often, a pronounced
‘spur^ formed by the backwards extension of digit
4 (see Lull 1953). Stenonyx may be distinguished
from Skartopus by the same features and, in some
instances, by the presence of a hallux imprint.
Wildeichnus appears to differ in possessing a
prominent ‘spur’ behind digit 3 and in showing a
clear imprint of the hallux. To summarize,
Skartopus is similar in basic morphology to other
coelurosaur footprints, but it may be
distinguished from these through differences in
divarication and relative lengths of the digits,
through lacking imprints of phalangeal pads or of
acuminate claws, through the absence of a
posterior ‘spur’, through the absence of a hallux
imprint, through the relative straightness of the
digits, and through its high values for the ratios
PL/FL and SL/FL.
DISCUSSION
Sizes and speeds of track-makers
From studies of locomotion in living terrestrial
vertebrates Alexander (1976) determined the
following relationship between stride length (A, in
metres), height at the hip {h, also in metres) and
speed (w, in metres per second):
(/) X/h = 2.3 {u/gh )
In this equation A represents our measurement for
stride length (SL), and g is a constant — the
acceleration of free fall; the ratio X/h is termed
‘relative stride length’ (Alexander 1976).
Alexander indicated that this relationship seems
to hold true, at least in general terms, for large
and small animals, both bipeds and quadrupeds,
at gaits from slow walk to fast run. In addition he
observed that the relationship does not seem to be
seriously affected by variation in the consistency
of the substrate. With additional data from fast-
moving African ungulates Alexander, Langman
and Jayes (1977) refined expression (7 ) to give:
(2) X/h =
These authors concluded that expression (2 ) is
best applied to animals that are cantering or
galloping, whereas expression (7 ) is appropriate
for animals using slower gaits. In mammals the
change from a walking gait to a trotting gaiF
occurs when X/h is approximately 2.0 (Alexander
1976); the change from trotting to galloping
follows when A / h has increased to about 2.9. This
latter figure is derived from two generalizations
presented by Alexander (1977). The first of these
is that mammals tend to shift from a trotting or
racking gait to a galloping or cantering gait when
the quantity u reaches a value of about 1.5. The
quantity ii, or ‘dimensionless speed’, was defined
by Alexander as follows:
(3) ii= u(ghr
where h is expressed in metres and u is in m/s.
The second generalization, which seems to apply
to a wide variety of animals through a wide range
of speeds, is that
(4) X/h =
where X/h represents mean relative stride length.
From these generalizations it may be assumed
that the shift from trotting to galloping occurs
when
(5) X/h = 2.3 (1.5)'’-^
\/h = 2.9
Consequently it is possible to identify the gaits of
dinosaurian track-makers on the basis of relative
stride length, as follows;
walk: X/h < 2.0; locomotor performance
equivalent to walking in mammals.
trot: X/h = 2.0 to 2.9; locomotor performance
equivalent to trotting or racking in mammals.
run: K/h > 2.9; locomotor performance
equivalent to cantering, galloping or sprinting in
mammals.
To estimate the speeds of certain dinosaurs
Alexander (1976) transformed expression (7 ) to
give:
(6) u = 0.25g'’W'-^/2
This equation was then applied to data from
dinosaur trackways, where A could be measured
directly (Alexander’s A = SL of our descriptions)
and where h might be estimated from the size of
the footprints. This method has since been
applied to more than 50 dinosaur trackways,
including a sample of those at Lark Quarry
(Russell and Beland 1976; Tucker and Burchette
1977; Coombs 1978; Thulborn and Wade 1979;
432
MEMOIRS OF THE QUEENSLAND MUSEUM
Farlow 1981; Kool 1981; Thulborn 1981, 1982).
Table 3 presents a summary of the speeds so far
estimated in this way.
Figures listed in Table 3 for the Lark Quarry
dinosaurs are preliminary estimates, and they will
be revised in the present work. Our discussion will
focus on the problem of estimating h on the basis
of footprint dimensions. It is desirable that h
should be estimated with reasonable care, because
an underestimate will generate an overestimate of
the track-maker’s speed; conversely an
overestimate of h will generate an underestimate
of speed. In some instances the hindlimb length of
a dinosaurian track-maker has been estimated
from the evidence of pace length or stride length
(e.g. Avnimelech 1966); an estimate of this type is
of questionable value, simply because pace length
and stride length vary according to the gait and
speed of the track-maker (Lull 1953, p. 146). In
other cases hindlimb length has been estimated on
the basis of footprint dimensions; for example,
Avnimelech (1966, p. 5) suggested that in the
footprints of bipedal dinosaurs the length of digit
3 represented about 18% of hindlimb length.
Elsewhere Alexander suggested (1976) that h
could be calculated as approximately four times
footprint length for a variety of dinosaurian
track-makers, both bipeds and quadrupeds, and
this suggestion has been rather widely accepted
(see all sources cited in Table 3). However,
Coombs (1978) expressed some reservations
about this generalization, and Alexander (1976)
did mention that footprint length could represent
anything between 0.23A and 0.28/i in the bipedal
dinosaurs that he examined. In discussing the
sizes and speeds of the Lark Quarry track-makers
we will investigate some other methods for
estimating h.
In comparing the sizes, weights and speeds of
various dinosaurs Coombs (1978, Table 2) drew a
TABLE 3: Summary of Sizes and Speeds Previously Estimated for Dinosaurian Track-Makers.
ichnoiaxa or track-
makers
(N)
h
(m)
u
(m/s)
Bipedal dinosaurs
(6)
0.6-2.1
1.2-3. 6
Sauropods
(2)
1. 5-3.0
l.O-l.l
Ornithomimid
(1)
1.2
1.8
^Giant ornithopod
(1)
3.4
7.5
^Giant ornithopod
(1)
3.4
2.4
lAnchisauripus
(2)
0.4-0.6''
x
1.3-2. 2
Carnosaur
(1)
2.6
2,3
Ornithopods
(10)
<L0
4.3""
Coelurosaurs
(10)
<1.0
3.6'"
Gypsichnites pacensis
(1)
1.2
2.0
Irenesauripus spp.
(3)
1. 5-2.1
L4-2.7
Irenichnites gracilis
(1)
0.6
2.8
Amblydactylus kortmeyeri
(1)
0.5
1.1
Tetrapodosaurus borealis
(I)
1.4
0.9
'^Theropods
(3)
1.2-1. 5^
8.3-11.9
^Theropods
(3)
1.5-1. 8^
1.8-2. 5
Theropods
(15)
1.5""^
4.2'"
u
(km/h)
X/h
gait
source
4.3-13.0
1.2-2. 5
walk (4)
trot (2)
Alexander 1976
3. 6-4.0
0. 8-1.1
walk
6.4
1.5"
walk
Russell and Beland 1976
27.1
2.7"
trot
8.5
1.3
walk
Thulborn 1981
4.5-7. 9
1. 3-1.4"
walk
Tucker and Burchette
1977
8.2
1.4
walk
Thulborn and Wade 1979
15.5'"
13.0*"
>2.0
>2.0
trot/run
(Lark Quarry)
7.2^
1.8
walk
5.0-9.7’‘
L2-L6
walk
10.1’'
2.3
trot
Kool 1981
4,0
1.5
walk
3.2’'
0.9
walk
29.9-42.8
3.7-4.9
run
6.4-8.9
1.5-1. 8
walk
Farlow 1981
15.2'"
2.3'"
trot
a: two interpretations of single trackway,
f: three fastest of Farlow’s 15 track-makers,
s: three slowest of Farlow’s 15 track-makers,
e: figures estimated from published data,
m: mean.
x: Coombs (1978) provides different speed estimates,
apparently through computational error (Farlow 1981).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
433
distinction between ‘height at hips* (as had been
estimated by Colbert 1962) and ‘standard [or
skeletal] hindlimb length’ (the sum of the lengths
of femur, tibia and metatarsal 3). He indicated
that there was sometimes a considerable
difference between these two dimensions —
particularly among large dinosaurs. In addition a
rather similar distinction has been made between
skeletal hindlimb length {h ) and ‘height of the
hindlimb’ (H) — the latter being defined as ‘the
combined lengths of femur, tibia and longest
metatarsal, plus an increment of 9% to account
for ankle bones and for soft tissues at knee, ankle
and sole* (Thulborn, 1982, p. 228). For present
purposes these various dimensions are assumed to
be roughly equivalent, on the grounds that they
are likely to be of great significance only in large
dinosaurs. Our estimates of h for the Lark Quarry
track-makers (Tables 4 and 5) are based on
osteometric data and may be regarded as
TABLE 4: Estimates of Size. Speed and Relative Stride Length for 57 Wintonopus Track-Makers.
estimated h
estimated speed
(cm)
(m/s)
(km/h)
estimated X/h
1.
13.7
2. 8-3. 5 (3.1)
10.0-12.5 (11.2)
3. 6-4.2 (3.9)
2.
16.8
3. 3-4.0 (3.6)
11.9-14.3 (12.9)
3. 8-4.3 (4.0)
3.
16.3
(3.5)
(12.6)
(4.0)
4.
16.8
3.0-3.3 (3.1)
10.6-11.8 (11.3)
3. 5-3. 8 (3.6)
5.
18.9
2.9-3.3 (3.1)
10.5-11.9 (11.2)
3. 3-3.6 (3.4)
6.
18.6
(3.2)
(11.6)
(3.5)
7.
18.4
3.4-3.9 (3.7)
12.1-14.0 (13.5)
3.7-4. 1 (4.0)
8.
19.7
3.0-3. 1 (3.0)
10.6-11.3 (11.0)
3.3-3.4 (3.3)
9.
20.1
3. 8-4.4 (4.1)
13.7-15.8 (14.7)
3.9-4.4 (4.2)
10.
18.4
3. 1-3.8 (3.4)
11.2-13.8 (12.4)
3. 5-4.1 (3.8)
11.
20.4
3. 8-4.2 (4.0)
13.6-15.2 (14.4)
3.9-4.2 (4.1)
12.
23.7
2.6-3.2 (2.9)
9.4-11.5 (10.5)
2. 7-3. 2 (3.0)
13.
27.1
4.0-4.5 (4.3)
14.5-16.1 (15.5)
3. 7-4,0 (3.8)
14.
25.5
4.4-4.9 (4,6)
15.8-17.6 (16.7)
4.0-4.4 (4.2)
15.
29.1
4.5-5.0 (4.7)
16.1-18.0 (17.0)
3.9-4.2 (4.0)
16.
22.0
3. 7-4.3 (3.9)
13.2-15.3 (14.0)
3. 7-4.1 (3.9)
17.
29.6
(4.1)
(14.6)
(3.5)
18.
32.6
4. 8-5.2 (5.0)
17.3-18.5 (17.9)
3.9-4. 1 (4.0)
19.
27.4
4.3-5.0 (4.7)
15.3-17.8 (16.8)
3.8-4.3 (4.1)
20.
29.7
4.1-5.4 (4.6)
14.7-19.3 (16.5)
3.6-4.4 (3.9)
21.
29.2
(6.3)
(22.7)
(5.0)
22.
26.4
4.3-4.4 (4.3)
15.6-15.8 (15.6)
(3.9)
23.
30.7
3. 8-4.3 (4.1)
13.7-15.6 (14.6)
3.3-3. 7 (3.5)
24.
33.2
(3.5)
12.5-12.7 (12.6)
(3.0)
25.
34.7
4.1-4.9 (4.6)
14.6-17.8 (16.4)
3.3-3.9 (3.7)
26.
30.1
3.5-4.7 (4.2)
12.7-16.9 (15.2)
3.2-4.0 (3.7)
27.
33.6
(4.1)
(14.8)
(3.4)
28.
31.0
3.7-3. 9 (3.8)
13.3-14.1 (13.7)
3. 3-3.4 (3.3)
29.
34.2
3. 8-5.0 (4.3)
13.7-18.0(15.5)
3.2-4.0 (3.5)
30.
29.5
(4.3)
(15.5)
(3.7)
31.
32.4
4.3-4.5 (4.4)
15.6-16.2 (15.8)
3.6-3. 7 (3.6)
32.
38.8
3. 9-5.0 (4.5)
14.0-18.0 (16.2)
3. 1-3.8 (3.5)
33.
29.2
3.2-3.4 (3.3)
11.6-12.3 (11.9)
3.0-3. 1 (3.1)
34.
33.4
3.7-4.6 (4.1)
13.3-16.5 (14.9)
3.2-3. 7 (3.4)
35.
33.7
(3.9)
(14.0)
(3.3)
36.
28.1
3. 8-5. 5 (4.9)
13.8-19.8 (17.7)
3. 5-4.6 (4.2)
434
MEMOIRS OF THE QUEENSLAND MUSEUM
37.
45.6
5. 8-6. 3 (6.1)
20.9-22.7 (21.9)
4.0-4.2 (4.1)
38.
44.6
(5.7)
(20.4)
(3-9)
39.
37.1
(4.2)
(15.2)
(3.4)
40.
38.1
4.2-4.8 (4.5)
15.0-17.3 (16.2)
3.3-3.7 (3.5)
41.
35.8
3.2-4.9 (4.0)
11.4-17.7 (14.4)
2.7-3. 8 (3.3)
42.
39.6
3.4-4.5 (3.9)
12.3-16.0 (14.0)
2.8-3.4 (3.1)
43.
49.3
5.6-6. 5 (6.0)
20.0-23.2 (21.6)
3.7-4.2 (3.9)
44.
41.8
2.5-3. 3 (3.1)
9.2-12.0 (11.0)
2.2-2.7 (2.5)
*45.
47.4
(1.1)
(4.0)
(1.5)
46.
49.4
(7.8)
(28.2)
(4.9)
47.
58.7
(4.0)
(14.4)
(2.7)
48.
61.5
6.0-7.6 (6.8)
21.8-27.3 (24.4)
3.6-4. 3 (4.0)
49.
53.6
6. 1-6.6 (6.4)
22.1-23.7 (22.9)
3.9-4. 1 (4.0)
50.
54.9
5.0-7 .0 (5.7)
18.0-25.0 (20.4)
3.3-4.2 (3.6)
51.
52.0
5.4-5.7 (5.5)
19.4-20.5 (19.9)
3.6-3. 7 (3.6)
52.
60.3
6. 1-6.5 (6.3)
22.1-23.4 (22.8)
3.7-3.9 (3.8)
53.
53.9
8.2-8.3 (8.2)
29.4-29.9 (29.7)
(4.9)
54.
61.9
(5.1)
(18.5)
(3.2)
55.
54.7
5.0-6.3 (5.7)
18.2-22.6 (20.7)
3. 3-3.9 (3.7)
56.
70.0
5.4-7.0 (6.3)
19.6-25.1 (22.6)
3.2-3. 9 (3.6)
57.
158.4
4.6-5.0 (4.8)
16.7-18.1 (17.2)
2. 1-2.2 (2.1)
Means
4.3-4.8 (4.6)
15.4-17.4 (16.4)
3. 5-3.9 (3.7)
’•'New Quarry trackway.
For each track-maker we show the range and the mean (in parentheses) of speed and relative stride length. A single
figure (in parentheses) indicates that only one stride could be measured, or that there was little or no variation in
stride length. Figures for the New Quarry track-maker (No. 45) were excluded from calculations of overall means.
equivalent to skeletal hindlimb length. If these
estimates were increased by 9% (to provide
estimates of H, or ‘height of the hindlimb*) the
mean increment for the omithopod track -makers
would be 3.1 cm; for the coelurosaur track-
makers the mean increment would be 1.5 cm.
These small increases in estimated body size
would not affect the general conclusions that we
draw regarding the speeds and gaits of the track-
makers.
Carnosaur trackway
In the trackway of the Lark Quarry carnosaur
footprint length ranges from 41 cm (estimated) to
64 cm; mean footprint length is 51.4 cm. With the
assumptions used by Alexander (1976) h could be
estimated to lie in the range 1.^ to 2.56 cm —
with the mean estimate at 2.06 m. However, it
seems legitimate to base our estimate of h on the
best-preserved and most complete footprint; this
particular footprint (number 3 in the trackway) is
64 cm long, providing estimated h of 2.56 m. The
footprint has a well-defined rear margin, and its
length is not exaggerated by scrape-marks; other
footprints in the carnosaur trackway appear to
have less complete impressions of the rear part of
the foot, and they would probably generate
underestimates of h (and, in consequence,
overestimates of the carnosaur’s speed).
There are at least two other ways to estimate h
for the Lark Quarry carnosaur. First it is possible
to compare the sizes of carnosaur footprints to
the sizes of carnosaur skeletons. Footprints
attributed to tyrannosaurs are reported to reach a
maximum length of about 80 cm (Haubold 1971),
while the largest well-known tyrannosaur,
Tyrannosaurus rex, has a skeletal hip height
about 3.17 m (representing the sum of the lengths
of femur, tibia and longest metatarsal). With the
admittedly untestable assumption that the largest
known footprints were made by animals about
the size of Tyrannosaurus, skeletal hip height
could be predicted as 3.96 times footprint length.
In the case of the Lark Quarry carnosaur this
method would indicate a skeletal hip height of
about 2.54 m.
Next it is possible to make use of the fact that
metatarsus length (MT) is strongly correlated with
skeletal hip height in carnosaurs — see Fig. 17,
where the least squares regression line represents
the following allometric equation:
(7) h = 4.15MT + 28.52 cm
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
435
(In this equation all measurements are expressed
in cm; Bartlett’s three-group method (see Sokal
and Rohlf 1969, p. 483) yields a virtually identical
equation). To apply equation (7 ) in the case of
the Lark Quarry carnosaur it is first necessary to
estimate MT on the basis of footprint size. It
should be possible to make such an estimate with
a fair degree of accuracy because in many bipedal
dinosaurs MT is roughly equivalent to the
summed lengths of phalanges in digit 3 (EP; see
Figs 6 and 18, and various illustrations given by
Coombs 1978); moreover, in a digitigrade animal
such as a dinosaur total footprint length (FL)
should be slightly greater than EP (Fig. 18). In
carnosaurs MT seems to be a little greater than SP
(see Russel! 1970, Table 1, for data on Canadian
carnosaurs), but would probably have been less
than FL (which comprised SP, claw sheath, joint
capsules and, possibly, a ‘heel’ region supported
by the distal part of the metatarsus). Evidently
MT would have been somewhat less than total
footprint length. For practical purposes we will
assume that MT is roughly equivalent to footprint
size index (SI) — which is also less than FL
TABLE 5: Estimates of Size, Speed and Relative Stride Length for 34 Skartopus Track-Makers.
estimated h
estimated speed
(cm)
(m/s)
(km/h)
estimated X/h
1.
13.3
3.0-4.2 (3.7)
10.8-15.1 (13.3)
3. 8-5.0 (4.5)
2.
13.3
3. 8-4.4 (4.2)
13.6-15.9 (15.0)
4.6-5.2 (4.9)
3.
14.4
2. 8-3. 3 (3.1)
10.2-12.0 (11.2)
3.6-4.0 (3.8)
4.
14.5
3.2-3.9 (3.5)
11.4-14.0 (12.6)
3.9-4.5 (4.2)
5.
14.8
3. 5-4.1 (3.8)
12.5-14.7 (13.6)
4. 1-4.7 (4.4)
6.
14.9
2. 8-3.4 (3.0)
10.2-12.2 (10.8)
3. 5-4.0 (3.7)
7.
15.0
2.9-3. 3 (3.1)
10.6-11.9 (11.2)
3.6-3. 9 (3.8)
8.
15.1
3.4-3. 6 (3.5)
12.1-13.0 (12.5)
4.0-4.2 (4.1)
9.
15.2
3. 1-3.8 (3.4)
11.3-13.8 (12.3)
3. 8-4.4 (4.0)
10.
15.3
2. 8-3. 5 (3.0)
10.1-12.5 (10.9)
3.4-4. 1 (3.7)
11.
15.5
3. 3-3.4 (3.3)
11.8-12.3 (12.1)
3.9-4.0 (3.9)
*12.
15.5
3.4-3.6 (3.4)
12.1-12.8 (12.4)
3.9-4.! (4.0)
13.
15.5
3.6-4.1 (3.9)
12.8-14.9 (14.0)
4.1-4.6 (4.4)
14.
15.6
3.4-3. 8 (3.6)
12.2-13.5 (13.0)
4.0-4.3 (4.2)
15.
16.2
3.6-3. 9 (3.7)
12.9-13.9 (13.3)
4.1-4.3 (4.2)
16.
16.3
2. 8-3. 2 (3.0)
10.2-11.4 (10.6)
3.4-3.7 (3.5)
17.
16.4
3. 1-3. 3 (3.2)
11.3-11.8 (11.6)
3. 7-3. 8 (3.7)
18.
16.6
3.3-3.5 (3.4)
11.9-12.4 (12.1)
3. 8-3.9 (3.8)
19.
16.6
(3.2)
(11.4)
(3.7)
20.
17.2
2.5-3.7 (3.1)
8.8-13.3 (11.3)
3.0-4. 1 (3.6)
21.
17.7
2. 8-3.9 (3.3)
9.9-13.9 (12.0)
3.2-3. 8 (3.7)
22.
17.9
3. 1-3.9 (3.3)
11.0-14.1 (12.0)
3. 5-4.2 (3.7)
23.
17.9
2.5-3. 1 (2.8)
9.0-11.3 (10.1)
3.0-3.5 (3.2)
24.
18.2
2.3-3.2 (2.6)
8.3-11.6 (9.5)
2.8-3.6 (3.1)
25.
18.2
2.2-2. 8 (2.5)
8.0-10.2 (9.1)
2,7-3.2 (3.0)
26.
18.3
(3.2)
(11.7)
(3.6)
27.
18.5
2.6-2. 8 (2.7)
9.4-10.1 (9.7)
3.0-3.2 (3.1)
28.
19.3
2.9-3. 5 (3.2)
10.4-12.6 (11.4)
3.2-3.7 (3.5)
29.
19.3
2. 5-2. 6 (2.5)
8.9-9.3 (9.1)
2.9-3. 0(2.9)
30.
19.6
2.7-2. 9 (2.8)
9.8-10.5 (lO.l)
3. 1-3.2 (3.1)
31.
19.9
3.2-3. 7 (3.5)
11.7-13.2 (12.6)
3. 5-3. 8 (3.7)
32.
20.1
2.6-3.0 (2.8)
9.2-10.7 (10.1)
2.9-3.2 (3.1)
33.
21.6
2.5-3. 1 (2.9)
9.1-11.3 (10.3)
2.8-3.3 (3.1)
34.
21.9
2.7-3.4 (3.0)
9.6-12.2 (10.6)
2.9-3.5 (3.1)
Means
3.0-3. 5 (3.2)
10.7-12.5 (11.6)
3.5-3.9 (3.7)
For each track-maker we show the range and the mean (in parentheses) of speed and relative stride length. A single
figure (in parentheses) indicates that only one stride could be measured, or that there was little or no variation in
stride length.
* Trackway No. 12 was made by an animal with consistent ‘flat-footed’ gait (see Plate 14, fig. B). The length of each
footprint is exaggerated by an imprint of the metatarsus, and to estimate the animal’s size and speed our
measurements of total footprint length were reduced by 50®7o.
436
MEMOIRS OF THE QUEENSLAND MUSEUM
because the footprint is longer than wide. The
best-preserved carnosaur footprint at Lark
Quarry has SI of 57.69 cm; by substituting this
figure for MT in equation (7 ) we can estimate
skeletal hip height to have been about 2.68 m.
These various estimates are in close agreement,
and they indicate that the Lark Quarry carnosaur
was between 2.54 and 2.68 metres in height at the
hip. The mean figure, which we will use for
estimating the animaLs speed, is 2.59 m.
Apparently the animal was about the same size as
one specimen of the Canadian carnosaur
Albertosaurus libratus (National Museum of
Canada, No. 2120, with h about 2.63 m; Lambe
1917); it would have been intermediate in size
between specimens of Daspletosaurus torosus {h
2.40 m, estimated from data of Russell 1970) and
Tyrannosaurus rex (h about 3.17 m; Osborn
1917).
The strides of the Lark Quarry carnosaur range
in length from 2.82 m to 3.74 m (mean 3.31 m).
Consequently relative stride length (X/h) is
estimated to range from 1.09 to 1.44 (mean 1.28).
In every stride X/h is well below 2.0, which
indicates that the animal was using a walking gait
(Alexander 1976) and that its speed is most
appropriately estimated with equation (6 ). The
carnosaur’ s progress may be plotted in some
detail, as follows;
stride
(m/s)
speed
(km/h)
X/h
1
2.34
8.43
1.44
2
2.13
7.65
1.36
3
2.35
8.47
1.44
4
2.34
8.43
1.44
5
1.76
6.32
1.21
6
1.67
6.03
1.18
7
1.74
6.26
1.20
8
1.54
5.54
1.12
9
1.47
5.28
1.09
means
1.93
6.93
1.28
The animal took a slightly weaving course (Fig,
3), during which it showed a defmile tendency to
decelerate (i.e. to shorten its strides). Its first four
strides are relatively long and are defined by very
deep footprints; the animal then switched quite
abruptly to a series of shorter strides (Nos. 5-9,
Fig. 22A) and its footprints became noticeably
shallower. The last two strides are the shortest
and were taken as the animal made a sharp turn to
its right.
Ornithopod trackways
In our preliminary account of Lark Quarry
(1979) the makers of the ornithopod trackways
{Wintonopus ) were estimated to range from the
size of bantams to the size of ostriches; their mean
speed was calculated to be about 4.31 m/s (15.52
km/h). These preliminary estimates of size and
speed were derived from a small sample (parts of
10 trackways) with the methods described by
Alexander (1976).
In the 57 trackways studied here mean
footprint length ranges from 2.40 cm to 22.75 cm
(mean 6.68 cm); by excluding the earlier-formed
trackway of the exceptionally large animal (No.
57 in Table 4) the range of means for footprint
length is reduced to 2.40 — 10.86 cm (with overall
mean 6.39 cm). Alexander assumed (1976) that
height at the hip {h) could be estimated as
approximately four times footprint length for a
variety of dinosaurian track-makers; if we apply
this assumption to the Wintonopus data the
track-makers are estimated to have had h ranging
between 9.60 and 43.44 cm (mean 25.58 cm), with
the single large individual at 91.0 cm. However,
Alexander indicated that footprint length (FL)
could represent anything between 0.23/i and 0.28/i
in the bipedal dinosaurs that he studied (1976),
and it seems worthwhile to investigate an
alternative method for estimating h.
Analysis of variance (p. 424) revealed
considerable variation in footprint length within
the Wintonopus trackways. Within a single
trackway there may occur foreshortened (‘stubby-
toed’) prints, normal prints and attenuated toe
scratches from a single foot (Figs 5L and 7, PI.
lOD). Consequently mean footprint length for a
trackway may not be a satisfactory indicator of
the track-maker’s size — because the mean will be
affected by the relative frequencies of
foreshortened and attenuated footprints.
Fortunately footprint size index (SI) seems to be a
reliable guide to the relative sizes of two or more
track-makers: the index differs from one
trackway to the next, but is virtually constant
within any one trackway. Footprint size index is
usually a little greater than footprint length in
Wintonopus (because the footprints are usually
broader than long), and it has the following
range: 2.89 to 12.69 cm (mean 6.74 cm), with the
single large individual at 26.59 cm. Two
observations make it possible to estimate h on the
basis of SI: first, in the foot skeletons of many
ornithopods the summed lengths of phalanges in
digit 3 (SP) is roughly equal to the length of
metatarsal 3 (MT; see Figs 6A, C); and, second,
there exists a strong correlation between MT and
skeletal hip height in ornithopods (see Fig. 19).
So, to calculate h for the Wintonopus track-
makers there seems to be only one prerequisite —
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
437
an estimate of MT (or of SP) derived from SI. By
dinosaurian standards the Wintonopus track-
makers seem to have been fairly small animals, so
that absolute differences between FL and 2P (and
hence MT) are likely to have been small. For the
sake of convenience we will assume that SI is
roughly equal to MT. It may be noted that SI is
usually a little greater than FL, because the
Wintonopus footprints are normally a little
broader than long. Consequently our estimates of
h (based on SI) will tend to be slightly greater than
similar estimates based on FL. This difference
will, on the whole, have a conservative effect — in
the sense that the sizes of track-makers may
overestimated, and their speeds may be
underestimated.
Figure 19A shows the relationship between MT
and skeletal hip height in a sample of 32
ornithopod skeletons (with MT ranging from 6.3
to 38.1 cm); the least squares regression line
represents the following allometric equation;
(8) h = 3.76MT“^
where both h and MT are expressed in
centimetres. By substituting SI for MT in this
equation we estimate that the Lark Quarry
ornithopods ranged in skeletal hip height from
12,86 to 71.57 cm (mean 34.41 cm), with the
solitary large individual at 168.81 cm. However,
the regression equation for cursorial ornithopods
(with femur shorter than tibia) is slightly different
from that for graviportal ornithopods (with
femur longer than tibia); for cursorial
ornithopods the equation is:
(9) h = 3.97MT'“
while for graviportal ornithopods it is:
(70) h = 5.06MT'^
The two regression lines, which are shown in Fig.
19B, have similar slopes but different intercepts.
In general terms equation (70 ) provides estimates
of h about 20 to 25 greater than those derived
with equation (9 ). The Lark Quarry ornithopods
seem to have been small animals, judging from
the size of their footprints, and this might indicate
that equation (P ), based on data from cursorial
ornithopods, should be used to estimate h. It
must be admitted, however, that the exact identity
of the Wintonopus track-makers is unknown —
beyond that fact that they seem to have been
ornithopods of some sort. For this reason we have
obtained three estimates of h for each track-
maker (by substituting SI for MT in each of the
preceding equations), and we will use the mean
figure in calculating the speed of each animal.
These mean values for h range from 13.70 to
69.98 cm (overall mean 34.82 cm), excluding the
single large animal with h estimated to be 158.59
cm.
Of the 57 Wintonopus trackways examined
here only one appears to have been made by an
animal more than 1 metre high at the hip. Forty-
six of the track-makers (Sl^^^o) are estimated to
have h less than 50 cm, and 24 of them (42%) h
less than 30 cm. In all cases Alexander’s
assumptions (1976) would provide smaller
estimates of h.
In terms of general body size the Lark Quarry
animals may be compared with cursorial
ornithopods of the families Fabrosauridae,
Hypsilophodontidae and Heterodontosauridae
(see skeletal reconstructions or flesh restorations
given by Galton 1974, Thulborn 1972, Santa Luca
et al. 1974); they might also be compared with
juvenile specimens of some bigger graviportal
ornithopods (e.g. the juvenile hadrosaurs
described by Horner and Makela 1979) and with
the juvenile psittacosaurs recently described by
Coombs (1980a, 1982). From the evidence of
trackways it does not seem possible to decide with
certainty whether the Lark Quarry ornithopods
were small cursorial forms ranging up to adult
status, or whether they were juveniles of some
bigger graviportal ornithopod. Neither of these
possibilities can be dismissed entirely: two types
of small ornithopod, resembling
hypsilophodontids, are reported from the
Cretaceous of Victoria (Flannery and Rich 1981),
and a large ornithopod with h about 2.44 m was
recently described from the Lower Cretaceous of
Queensland (Muttaburrasaurus, Bartholomai and
Molnar 1981). In either case it would still seem
reasonable to regard the smallest (at least) of the
Lark Quarry track-makers as juvenile animals. It
has sometimes been supposed that juvenile
dinosaurs were rare (Richmond 1965, Leonard!
1981), but Horner and Makela (1979) found that
more than 80% of dinosaur specimens collected
from the Two Medicine Formation (Upper
Cretaceous) of Montana could be identified as
juveniles or subadults. It seems that juvenile
dinosaurs may have been quite common, at least
in some localities.
The curve illustrating size frequency
distribution for the Wintonopus track-makers is
distinctly skewed and is similar to the type of
curve derived by Boucot (1953) from ‘life
assemblages’ of fossils (see Fig. 21A). The curve
might be interpreted in any of several ways. First
it could simply be regarded as a survivorship
curve for a population of small cursorial
ornithopods. According to this interpretation the
438
MEMOIRS OF THE QUEENSLAND MUSEUM
animals would normally have grown to achieve h
of about 30 cm (peak of curve); thereafter the
mortality rate would have reached its maximum
(steepest part of curve), with fewer and fewer
animals surviving to reach greater and greater size
(by virtue of the indeterminate growth prevailing
among reptiles). This interpretation assumes, of
course, that the sample of 57 track-makers is truly
representative of the dinosaur population from
which it was drawn; it also assumes the existence
and constancy of some strong correlation between
size and age. Secondly it is possible to interpret
Fig. 21A as a survivorship curve based on a
sample drawn from a population of big
graviportal ornithopods (with assumptions as
above). In this case it would appear that there was
a high rate of mortality among juveniles or
subadults once these had attained h of about 40
cm. The adults, presumably with /j of 1.5 metres
or more, would then have been comparatively
rare and relatively long-lived. Next there is the
possibility that our sample of 57 track-makers is
not a representative one: it could comprise
animals from two or more species, or there could
be serious under-representation of bigger animals
in a single species. (It seems scarcely possible that
smaller animals could be under-represented).
There is no obvious reason to assume that the
Wintonopus trackways were produced by two or
more different types of dinosaur: none of the
frequency distributions is strongly bimodal (Figs
8 and 9), there are impressive correlations
between any two dimensions of the footprints and
trackways (Fig. 10), and all the footprints can be
interpreted as those of animals sharing one
distinctive pattern of foot structure (Fig. 6C) and
using the same gait (Fig. 11). We cannot discount
entirely the possibility that our sample of 57
Wintonopus trackways might be heterogeneous
(in which case we could deduce nothing about the
population structure and possible affinities of the
track-makers), but this possibility does seem
rather unlikely: it supposes the co-existence of
two or more dinosaurian species, each of which is
represented by juveniles or is characteristically of
small size, and each of which produced
Wintonopus -like trackways. The final possibility
is that bigger animals did exist, but that they are
under-represented in our sample. Under-
representation of bigger animals could not be
regarded as an effect of sampling: trackways of
large animals are not common at Lark Quarry,
and we ensured that our sample contained data
from the largest and second-largest examples of
Wintonopus, Consequently this final possibility
must imply that large and small animals were
segregated, either fortuitously or through some
deliberate strategy on the part of the potential
track-makers. Of all these possible interpretations
the simplest would certainly seem to be the first —
that the sample of 57 trackways is representative
of a dinosaur population in which animals rarely
grew to hip heights estimated at greater than 70
cm. Finally there is some evidence of three size
classes in the Wintonopus sample (note the three
clumps of data points in Fig. 10). In terms of
estimated hip height these three size classes have
approximate limits of 13 to 20 cm, 25 to 40 cm,
and 50 to 60 cm. There is no way to investigate the
possibility that these size groupings might be
equivalent to age classes.
From our estimates of h it is calculated that the
mean value of \/h per trackway ranges from 2.69
to 5.03 (overall mean 3.69); these figures exclude
values for the solitary large animal {X/h 2.10),
and for the New Quarry track-maker. It seems
that all the Wintonopus track-makers at Lark
Quarry were using a gait faster than a walk (\/h
greater than 2.0). Two individuals were
apparently moving at a trot or a slow run (with
mean X/h at 2.1 and 2.7), whereas all the others
(96%) were using a fast running gait equivalent to
a mammalian gallop (with X/h at 3.0 or greater).
The 56 ornithopod track-makers at Lark
Quarry all have X/h estimated to be greater than
2.0, so that their speeds are most appropriately
estimated by means of equation (2 ) re-written as:
(11) u = [gh(X/\.ShY^^]'^
In this equation A and h are expressed in metres
and u is in metres per second. Mean speed per
trackway is estimated to range from 2.92 m/s
(10.52 km/h) to 8.24 m/s (29.66 km/h), with the
overall mean for 55 animals (excluding solitary
large individual) at 4.48 m/s (16.12 km/h). The
one exceptionally large animal has an estimated
mean speed of 4.78 m/s (17.22 km/h). The
minimum speed estimated for any of these track-
makers, on the basis of its single shortest stride, is
2.22 m/s (8.00 km/h); the maximum speed
estimated for any of the track-makers, on the
basis of its single longest stride, is 8.30 m/s (29.88
km/h).
The single Wintonopus trackway at New
Quarry has mean SI of 8.91 cm; for this track-
maker h is estimated to have been 47.4 cm,
indicating X/h of about 1.54. The New Quarry
animal w'as not associated with those at Lark
Quarry, and it seems to have been using a
different gait (walking rather than running). Its
speed, which is best calculated with equation (6),
is estimated to have been 1.11 m/s (3.98 km/h).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
439
Even if its speed is estimated, inappropriately,
with equation (11 ) the New Quarry animal would
still seem to have been moving more slowly than
any of the Lark Quarry track-makers (i.e. at 1.76
m/s, or 6.34 km/h).
The estimated sizes, speeds and relative stride
lengths of all 57 Wintonopus track-makers are
summarized in Table 4. The animals seem to have
maintained fairly constant speeds, and in most
cases mean stride length for the second half of the
trackway differs only slightly from mean stride
length for the first half (see Fig. 23A). On average
the difference between these two means in an
increase of 1.3%. An analysis of stride lengths in
the longest section of ornithopod trackway (No.
41 in Table 4, comprising 17 strides) reveals no
consistent trend towards acceleration
(lengthening of strides) or deceleration
(shortening of strides). Mean stride length for the
second half of this trackway is 6.8% less than
mean stride length for the first half. Short
sections of this trackway (Fig. 22B) might give the
impression of a strong tendency to lengthen or
shorten the strides, yet the track-maker seems,
overall, to have maintained a reasonably
consistent stride length (116.82 ± 11.13 cm; CV
9.5%).
To summarize, the Wintonopus trackways at
Lark Quarry seem to have been made by small
ornithopod dinosaurs, mostly less than 60 cm in
height at the hip. All these dinosaurs seem to have
been using a fast running gait, with \/h in most
cases between 3.2 and 4.1. For the majority
speeds are estimated between 3.4 m/s (12.2 km/h)
and 5.6 m/s (20.2 km/h), and there is no clear
evidence that the animals were accelerating or
decelerating.
COELUROSAUR TRACKWAYS
In our preliminary study of Lark Quarry (1979)
we estimated that the makers of the coelurosaur
trackways (Skartopus) had a size range equivalent
to that between bantams and half-grown emus;
their mean speed was estimated to have been
about 3.62 m/s (13.04 km/h). These preliminary
estimates of size and speed were obtained by
applying Alexander’s method (1976) to parts of
10 trackways.
With Alexander’s working assumption (1976)
that footprint length (FL) represents about 0.25/?
it may be estimated that the 34 trackways studied
here were made by animals ranging from 14.5 to
22.5 cm in height at the hip (overall mean 17.8
cm). But again, as with the ornithopod track-
makers, it may be worthwhile to investigate
another method for estimating h.
Analysis of variance (p. 429) reveals that FL
is highly variable within and among the Skartopus
trackways; so, too, are footprint width (FW) and
footprint size index (SI). Consequently there
seems to be little advantage in selecting SI, rather
than FL or FW, as an indicator to the relative
sizes of the track-makers. None of these variables
appears to be a very reliable guide to the relative
sizes of the track-makers, but this fact is not
particularly important because all these animals
seem to have been much the same size anyway.
Our estimates of size and speed for the track-
makers will be based on mean FL per trackway. It
might not be appropriate to base these estimates
on mean SI (as was done for the ornithopod
trackways) because this is usually less than actual
footprint length — on account of the footprints
usually being longer than broad. This preference
for FL, rather than SI, will have a conservative
effect — in that estimates of h will tend to be
increased and estimates of speed will tend to be
decreased. In the 34 trackways studied here mean
FL ranges from 3.62 to 5.62 cm, with the overall
mean at 4.46 ± 0.50 cm.
In a variety of coelurosaurs MT (the length of
metatarsal 3) is strongly correlated with skeletal
hip height; this relationship illustrated in Fig. 20,
where the least squares regression line represents
the following allometric equation:
(12) /? = 3.06MT"^
Both MT and h are expressed in centimetres. To
estimate h for the Lark Quarry coelurosaurs we
will substitute FL for MT in this equation, on the
assumption that these two dimensions were
roughly equal. This assumption is reasonable
because both dimensions were probably a little
greater than SP (summed lengths of phalanges in
digit 3). The slight preponderance of FL over SP
is apparent from Fig. 18; from various
illustrations of coelurosaur foot skeletons it
appears that MT is also a little greater than IP —
in a ratio about 11:10 (see Figs 6D, E, and
references given in caption to Fig. 20).
By substituting FL for MT in equation (12 ) we
estimate that the Lark Quarry coelurosaurs
ranged from 13.27 cm to 21.93 cm in height at the
hip (with mean for the 34 animals at 16.92 ± 2.23
cm). These estimates are slightly smaller than
those obtained with Alexander’s assumption that
h is approximately four times FL; for the smallest
track-maker the two estimates differ by 12 mm,
and for the largest they differ by 6 mm. These
differences are unlikely to be of great
significance, especially since Alexander’s work
(1976) indicates that h could represent anything
440
MEMOIRS OF THE QUEENSLAND MUSEUM
from about 3.6FL to about 4.3FL in the various
bipedal dinosaurs that he studied. Moreover these
differences will not affect the general conclusions
that we draw regarding the gaits and speeds of the
track-makers (see Table 6, where our estimates
for h are increased by about 40 to SO^’/o).
From these estimates is seems that the
Skartopus track-makers were somewhat smaller
than the familiar coelurosaurs Coelophysis (h
from 33.7 to 55.9 cm in 13 specimens listed by
Colbert 1964), Ornitholestes (h about 48.3 cm,
Osborn 1917) and Podokesaurus (h 25.5 cm,
Talbot 191 1). The only well known coeiurosaur of
comparable size would seem to be
Compsognathus, with h as little as 21.1 cm (for
holotype, estimated from measurements given by
Ostrom 1978). However, the Skartopus footprints
do agree in size with many other footprints
attributed to coelurosaurs (see Table 2).
The size frequency distribution for the
Skartopus track-makers is distinctly skewed (Fig.
2 IB) and is open to the several interpretations
that were considered earlier for the Wintonopus
track-makers. The possible interpretations may
be summarized as follows:
1. The 34 Skartopus track-makers represent a
coeiurosaur population in which individual
animals grew to a maximum hip height estimated
at 22 cm. This interpretation assumes that the
sample of 34 track-makers is truly representative
of the population from which it was drawn (i.e.
that it includes both juveniles and adults), and
that there existed a strong unvarying correlation
between size and age.
2. The 34 track-makers are juveniles of some
type of theropod dinosaur that grew to greater
size — though the bigger individuals are not
represented at Lark Quarry. This implies that
large and small animals were segregated, either by
chance or through their behaviour.
3. The sample of 34 trackways is
heterogeneous. This assumes that two or more
types of small (or juvenile) dinosaurs made
identical trackways at a single site, and that they
did so at approximately the same time.
All these possible interpretations involve
untestable assumptions, but the first of them
would seem to be the simplest. In any case it is
noteworthy that the Skartopus track-makers must
have been remarkably small animals by
dinosaurian standards. Even if it is assumed that
the track-makers were exceptionally long-legged
animals resembling ornithomimids (or ‘ostrich
dinosaurs’) it may be estimated that the largest of
them was less than 32 cm in height at the hip (see
Table 6 and further discussion below).
With our estimates of h the mean value of \/h
per Skartopus trackway is found to range from
2.90 to 4.94, with the overall mean at 3.71. In
only three trackways is minimum X/h (based on
the shortest stride) estimated to be less than 2.9,
and in no case does it fall below 2.7. Evidently all
34 track-makers w'ere using a fast running gait.
These figures for \/h indicate that the speeds of
the track-makers are most appropriately
estimated by means of equation (I I ). Mean speed
per trackway is estimated to range from 2.53 m/s
(9.09 km/h) to 4.16 m/s (14.98 km/h), with the
overall mean at 3.22 m/s (11.58 km/h). The
minimum speed estimated for any of the track-
makers, on the basis of its single shortest stride, is
2.23 m/s (8.03 km/h); the maximum speed
estimated for any of the track-makers, on the
basis of the single longest stride, is 4.42 m/s
(15.91 km/h).
Table 5 presents a summary of estimated size,
speed and relative stride length for each of the 34
track-makers. There is no clear indication that the
animals were either accelerating or decelerating.
In most cases mean stride length for the second
half of a trackway differs only slightly from mean
stride length for the first half (Fig. 23B). On
average this difference between the two means is a
decrease of about 2.1 %. From Fig. 23B it appears
that the majority of track-makers showed a slight
reduction in stride length during their progress.
However, this tendency to shorten the strides is
neither consistent nor well-marked, and it is
probably of little significance. By way of
illustration Fig. 22C shows an analysis of the
longest section of coeiurosaur trackway (No. 20
in Table 5, comprising 22 strides): in the latter
half of this trackway mean stride length is 2.5'^^o
less than mean stride length in the first half, but
there is no consistent trend towards progressive
shortening of the strides. Short sections of the
trackway could give the impression of a strong
trend to shortening or lengthening the strides, yet
the track-maker seems, overall, to have
maintained a fairly consistent stride length (61.77
± 5.40 cm; CV 8.7<^^o).
In summary, the Skartopus trackways seem to
have been produced by small coelurosaurs, all of
which are estimated to have been less than 22 cm
high at the hip. All of these animals seem to have
been using a fast running gait; estimates of X/h
are in most instances greater than 2.9, though a
few trackways include short strides indicating
occasional lapses of X/h as low as 2.7. Mean
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
441
TABLE 6: Estimates of Size, Speed and Relative Stride Length for 34 Skartopvs Track-Makers — With
THE Assumption That These Were Animals Resembling Ornithomimids. Ranges and Means of
Estimates Shown as in Table 5.
h (cm)
speed (m/s)
speed (km/h)
X/h
1.
19.5
2.2-3. 1 (2.8)
8.0-11.1 (9.9)
2.6-3.4 (3.1)
2.
19.6
2. 8-3. 3 (3.1)
10.1-11.8 (11.1)
3. 1-3.5 (3.4)
3.
20.9
2.1-2. 5 (2.3)
7.6-8.9 (8.4)
2.4-2. 8 (2.6)
4.
21.1
2.4-2. 9 (2.6)
8.5-10.4 (9.4)
2.7-3. 1 (2.9)
5.
21.5
2.6-3.0 (2.8)
9.4-10.9 (10.2)
2. 8-3.2 (3.0)
6.
21.6
2. 1-2.5 (2.3)
7.6-9.1 (8.1)
2.4-2.8 (2.5)
7.
21.7
2. 2-2. 5 (2.3)
7.9-8.9 (8.3)
2. 5-2.7 (2.6)
8.
21.9
2. 5-2.7 (2.6)
9.0-9.7 (9.4)
2.7-2.9 (2.8)
9.
22.0
2.3-2.9 (2.6)
8.4-10.3 (9.2)
2.6-3.0 (2.8)
10.
22.1
2. 1-2.6 (2.3)
7.5-9.3 (8.2)
2.4-2. 8 (2.5)
11.
22.4
2. 5-2.6 (2.5)
8.9-9.2 (9.1)
2. 7-2.8 (2.7)
*12.
22.4
2. 5-2.8 (2.6)
9.0-9.6 (9.3)
2. 7-2.9 (2.8)
13.
22.4
2.7-3. 1 (2.9)
9.6-11.2 (10.5)
2.9-3.2 (3.1)
14.
22.6
2.6-2. 8 (2.7)
9.2-10.2 (9.8)
2. 8-3.0 (2.9)
15.
23.2
2.7-2. 9 (2.8)
9.7-10.5 (10.0)
2. 8-3.0 (2.9)
16.
23.4
2.1-2.4 (2.2)
7.7-8. 5 (8.0)
2.4-2.6 (2.4)
17.
23.5
2.4-2. 5 (2.4)
8.5-8.9 (8.7)
(2.6)
18.
23.8
2.5-2.6 (2.5)
9.0-9.4 (9.1)
(2.7)
19.
23.8
(2.4)
(8.6)
(2.6)
20.
24.6
1. 9-2.8 (2.4)
6.7-10.0 (8.5)
2. 1-2.8 (2.5)
21.
25.3
2. 1-2.9 (2.5)
7.5-10.6 (9.1)
2. 3-2.9 (2.6)
22.
25.4
2. 3-3.0 (2.5)
8.4-10.7 (9.1)
2.4-2.9 (2.6)
23.
25.4
1.9-2.4 (2.1)
6.8-8.5 (7.7)
2. 1-2.5 (2.3)
24.
25.8
1. 7-2.4 (2.0)
6.3-8. 8 (7.2)
1. 9-2.5 (2.2)
25.
25.9
1. 7-2.1 (1.9)
6.1-7.7 (6.9)
1. 9-2.3 (2.1)
26.
26.0
(2.5)
(8.9)
(2.5)
27.
26.2
2.0-2. 1 (2.0)
7.2-7.7 (7.4)
2.1-2.3 (2.2)
28.
27.2
2.2-2. 7 (2.4)
7.9-9.6 (8.7)
2. 3-2.6 (2.5)
29.
27.3
1.9-2.0 (1.9)
6.8-7. 1 (6.9)
2.0-2. 1 (2.0)
30.
27.6
2. 1-2.2 (2.2)
7. 5-8.0 (7.8)
2.2-2.3 (2.2)
31.
27.9
2. 5-2.8 (2.7)
8.9-10.1 (9.7)
2.5-2. 7 (2.6)
32.
28.2
2.0-2.3 (2.1)
7. 1-8. 2 (7.7)
2.1-2.3 (2.2)
33.
30.1
1. 9-2.4 (2.2)
7.0-8.7 (8.0)
2.0-2.4 (2.2)
34.
30.5
2. 1-2.6 (2.3)
7.4-9.4 (8.2)
2.1-2. 5 (2.2)
Means
2.3-2.6 (2.4)
8. 1-9.5 (8.7)
2.4-2.7 (2.5)
estimated speeds of the track-makers are
generally in the range 2.82 m/s to 3.61 m/s (10.16
to 13.00 km/h), and there is no clear indication
that the animals were either accelerating or
decelerating.
The preceding estimates of size and speed
depend on the assumption that the Lark Quarry
track-makers had hindlimb proportions similar to
those in ‘typical’ coelurosaurs such as
Coelophysis, Compsognathus and Ornitholestes.
However, one group of small to medium-sized
theropod dinosaurs — the ornithomimids or
‘ostrich dinosaurs’ — is characterized by unusual
hindlimb proportions: these animals have
exceptionally long hindlimbs terminating in
relatively short toes. For the sake of completeness
we will examine the possibility that Skartopus
trackways might have been made by animals
resembling ornithomimids.
From published measurements of orni-
thomimid skeletons (Russell 1972 — Orni-
thomimus, Struthiomimus, Dromiceiomimus;
Osmolska et al. 1972 — Gallimimus ) it appears
that MT represents something between 1.45 and
1.72 times SP (the mean figure derived from 9
specimens being 1.57). Since FL was probably a
little greater than SP (see Fig. 18) we may assume,
for the sake of convenience, that MT was
equivalent to about 1.5FL. From the same
osteometric data it seems that there is a strong
positive correlation between MT and skeletal hip
height (r = 0.997, N = 9); the latter may be
442
MEMOIRS OF THE QUEENSLAND MUSEUM
predicted by means of the following allometric
equation:
(13) h = 3.49MT'"^
where h and MT are expressed in centimetres. If
the Lark Quarry coelurosaurs resembled
ornithomimids in limb proportions their skeletal
hip heights might, then, be estimated as follows:
(14) h = 3.49(1. 5FL)'“
This method provides estimates of h ranging from
20.44 to 31.96 cm (with overall mean of 25.27
cm). Consequently estimates of mean X/h per
trackway would range from 1.96 to 2.88 (with
overall mean 2.49). In only two of the 34 cases
would the estimate for mean X/h be less than 2.0
(actually 1.99 and 1.96). These figures would
indicate that all the Skartopus track-makers were
trotting or running (or. in two cases, were at least
on the point of breaking into a trot). Estimates of
mean speed per trackway, obtained with equation
(11 ), range from 1.85 m/s to 2.97 m/s (6.67
km/h to 10.69 km/h), with the overall mean at
2.33 m/s (8.40 km/h). These particular estimates
of size, speed and relative stride length are
summarized in Table 6.
It seems difficult to escape the conclusion that
Skartopus trackways are those of running
dinosaurs — even with the assumption that these
might have been exceptionally long-legged
dinosaurs resembling ornithomimids. There are at
least two good reasons for believing that the
track-makers were not ornithomimids. First there
is simply the matter of size: the Skartopus
footprints are much smaller than the foot
skeleton in any specimen of ornithomimid
dinosaur so far described. In the smallest of the
complete foot skeletons listed by Osm61ska et al.
(1972, p. 131) SP is 12.8 cm. One smaller example
is listed by these authors, but it lacks the
penultimate phalanx in digit 3; for this specimen
we estimate SP to have been about 9.0 cm. In the
ornithomimids described by Russell (1972) SP
ranges from 21.5 cm to more than 25.5 cm.
Footprints attributed to ornithomimids, or to
unknown but presumably similar dinosaurs, have
FL from about 10 cm (Hopiichnus shingi, Welles
1971) to about 28 cm (Ornithomimipus angustus,
Sternberg 1926). By comparison the largest figure
for mean FL in any of the Skartopus trackways is
5.62 cm. In the second place there seem to be no
certainly-identified skeletal remains of
ornithomimids from the Gondwana continents, A
few bones (dorsal vertebrae and a phalanx) from
the Lameta Formation of India were described by
von Huene and Matley (1933) as
Ornithomimoides mobilis and O. barasimlensis.
but Osmolska et al. regarded the material as
‘systematically insufficient’ (1972, p. 104).
Russell (1972) considered the two species to be
nomina vana. Molnar (1980), examining these
and other records, concluded that there was no
evidence suggesting that ornithomimids existed
on the southern continents. In summary, the
Skartopus footprints seem rather too small to be
those of ornithomimids, and there is little
evidence to suggest that such dinosaurs existed in
the southern continents. Consequently we may
assume the Lark Quarry trackways to have been
made by some other type of theropod dinosaur —
which presumably had ‘typical’ limb proportions.
A DINOSAUR STAMPEDE?
The most distinctive features of the Lark
Quarry trackway site are: (1) that the dinosaurian
track-makers were very numerous; (2) that nearly
all these track-makers seem to have been small by
dinosaurian standards; (3) that trackways of the
two main types ( Wintonopus and Skartopus ) are
in some places coincident, superimposed or
interwoven; (4) that all the trackways (except that
of the carnosaur) head in a single direction; and
(5) that all the track-makers (except the
carnosaur) seem to have been running. This
combination of features appears to be unique,
and it previously led us to interpret the Lark
Quarry trackways as the result of a dinosaurian
stampede (Thulborn and Wade 1979). The
evidence underlying this interpretation may now
be examined in more detail.
The Lark Quarry bedding plane carries one of
the densest accumulations of dinosaur footprints
yet reported (see Table 7). In our preliminary
description of the site we estimated that these
footprints represented the trackways of at least
130 dinosaurs (excluding the carnosaur), with
coelurosaurs (= Skartopus trackways)
outnumbering ornithopods (= Wintonopus
trackways) in a ratio about 55:45. It proved
impossible to assign every footprint at Lark
Quarry to a particular trackway, and so our
estimate for the number of track-makers must be
checked in some other way. In a 1 metre wide
transect al a right angle to the direction of the
trackways (between points X-X in Fig. 3) we
counted a total of 350 footprints. In practically all
cases mean pace length for a track-maker
(whether ornithopod or coelurosaur) is less than 1
metre — which means that nearly every animal
crossing the line of the transect should have left at
least one footprint in the metre-wide strip.
Consequently the maximum number of animals
that crossed the transect could be estimated at
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
443
350. However, mean pace length for Wintonopus
is 68 cm (i.e. about 1 Vi footprints per metre), and
for Skartopus it is 32 cm (i.e. about 3 footprints
per metre). By assuming that ornithopods and
coelurosaurs were present in equal numbers we
can estimate the number of animals to have
crossed the transect as:
350
= 156 animals
(3 X 0.5) + (1.5 X 0.5)
This very rough estimate must be regarded as an
absolute minimum. We suspect that coelurosaurs
outnumbered ornithopods, but it is impossible to
make allowance for this without identifying every
footprint and assigning to it a particular
trackway. This proved an impossible task (for
reasons described earlier; see p. 418). Moreover
it is certain that many smaller animals (both
coelurosaurs and ornithopods) crossed our
transect without leaving recognizable footprints:
these animals would have been so light that their
broad-spreading and rather springy feet simply
failed to break through the surface of the
sediment. This seems to have happened very
commonly, to judge from the number of
discontinuities or ‘gaps’ in the trackways at Lark
Quarry. The area of bedding plane exposed at
Lark Quarry is delimited partly by erosion and
partly by undisturbed overburden (mainly to S
and SW, see PI. 3). Within this area the footprints
are fairly evenly distributed (PL 4), and it seems
almost certain that more of them could be
revealed by extending the quarry to the SSW. Our
estimate of 156 animals may well represent a
fraction of the number of dinosaurs that
traversed Lark Quarry and its environs.
Trackways are certainly abundant, but this fact
alone does not support the hypothesis of a
dinosaur stampede. However, this fact does
assume some significance when one considers that
all the trackways may have been formed
simultaneously by animals running in a single
direction.
There are several reasons for believing that all
the Wintonopus and Skartopus trackways at Lark
Quarry were formed at, or about, the same time
(excepting the single unusually large example of
Wintonopus ). First, all these trackways are very
similar in preservation; there is no evidence (such
as scouring or erosion) to indicate that some
tracks are much older than others. Second, all the
trackways are impressed to about the same depth
— evidently in sediment of uniform consistency.
If the trackways had accumulated over a lengthy
period one might expect to find evidence of a
change in the consistency of the substrate (i.e.
some tracks more deeply impressed than others).
Finally there is the evidence of superimposed
footprints; these are quite common, and in all
cases the later-formed print is similar in its depth
and state of preservation to the earlier-formed
one. These similarities are apparent even where
three or more animals have trodden the same spot
(see PI. 13, Fig. C; PL 16, Figs B, C). Ornithopod
footprints {Wintonopus) may be found
superimposed on footprints of other ornithopods
or of coelurosaurs {Skartopus ), and the same is
true for the coelurosaur footprints. Footprints of
both types may be found superimposed upon
those of the carnosaur. From the evidence of
superimposed footprints it may be deduced (a)
that the carnosaur traversed the area before some
(at least) of the ornithopods and coelurosaurs did
so; (b) that some ornithopods preceded some
coelurosaurs: and (c) that some coelurosaurs
preceded some ornithopods. But from the
evidence as a whole it is possible to reach a more
general conclusion: that both ornithopods and
coelurosaurs traversed the Lark Quarry site at (or
about) the same time, and that they did so after
the passage of the solitary carnosaur. This is
exactly the chronological sequence to be expected
if the approach of the carnosaur had triggered a
stampede of the ornithopods and coelurosaurs.
Of course there is no absolute proof that the Lark
Quarry trackways were formed in exactly this
sequence, but it is difficult to imagine any other
when one considers that all the ornithopod and
coelurosaur track-makers were running in a single
direction.
Perhaps the most striking feature of the Lark
Quarry site is that all the animals responsible for
Wintonopus and Skartopus trackways were
headed in a single direction — about 55^" E of true
N, and in almost direct opposition to the course
taken by the solitary carnosaur (see PL 4). None
of these abundant ornithopod and coelurosaur
trackways deviates more than a few degrees from
a single compass bearing, and in this respect the
Lark Quarry site appears to be unique. Dinosaur
tracks uncovered at other prolific localities are
randomly oriented (e.g. see de Lapparenl and
Montenal 1967, Tucker and Burchette 1977) or
show some less obvious tendency to sub-parallel
alignment (e.g. see Avnimelech 1966, Ostrom
1972). And at sites where sub-parallel trackways
do predominate these often represent a ‘two-way
traffic’ — with some trackways diametrically
opposed to others (e.g. Avnimelech 1966, pi.
Vlll; Ostrom 1972, fig. 4). By contrast the Lark
444
MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 7: Comparison of Statistics for Various Trackway Sites.
number of number of
site
age
area(m^)
Ain-Sefra, Algeria:
Triassic
1
St Laurent de Treves,
France;
L Jurassic
25
Bendrick Rock, S Wales:
U Triassic
25
Rocky Hill, Connecticut:
Rhaetic
930
Swanage, S England:
U Jurassic
47^
Kerman area, Iran:
Jurassic
5
Tocantins River, Brazil:
Jurassic —
Cretaceous
6-8000
Beth Zayit, Israel:
Cenomanian
400
F6 Ranch, Texas:
Aptian —
Albian
7
Serrote do Letreiro, Brazil:
Triassic
‘spacious’
Mt Tom, Massachusetts:
Rhaetic
800
Lark Quarry, Queensland:
Cenomanian
209
( ^ — estimated figures.)
Quarry trackways are more nearly parallel and
(excepting the carnosaur trackway) entirely
unidirectional. The coincidence of so many
trackways certainly implies that some external
factor controlled the behaviour of the track-
makers (cf. Ostrom 1972). At the time the
trackways were formed the Lark Quarry site
appears to have been part of a broad drainage
channel, and it is conceivable that the track-
makers might have been funnelled along a
common route by physical barriers such as levees
or steep banks. However, there is no direct
evidence of such barriers, and it may be recalled
that remnants of a few randomly oriented
trackways do occur at the site. These scattered
and eroded remnants of trackways testify that
some medium-sized bipedal dinosaurs (probably
ornithopods) traversed the area before the
carnosaur made its appearance, and that they did
so randomly — that is, seemingly without the
control of physical barriers. Apparently the
behaviour of the Wintonopus and Skartopus
track-makers was influenced by some factor that
had not previously affected the movements of
dinosaurs across the same area. Next, it is obvious
that the Lark Quarry site cannot have been part
of some established route along which dinosaurs
were accustomed to move in either direction. Nor
is it possible to believe that the dinosaurian track-
makers could have adhered with absolute fidelity
to a system of ‘one-way’ routes. One general
conclusion seems inescapable: that the singular
footprints
trackways
source
12
7
Bassoullet 1971
24
7
Thaler 1962
400
7
Tucker and Burchette 1977
1000 +
7
Ostrom 1972
46
3
Charig and Newman 1962
8
?6"
de Lapparent and
Davoudzadeh 1972
47 +
6
Leonardi 1980
200 +
?10
Avnimelech 1966
76
15
Farlow 1981
9
17
Leonardi 1979
137
28 +
Ostrom 1972
3300 +
130 +
Thulborn and Wade 1979
orientation of trackways at Lark Quarry must
reflect some unusual behaviour on the part of the
track-makers.
There can be little doubt that all the
Wintonopus and Skartopus trackways at Lark
Quarry were made by running animals. In most
cases relative stride length is estimated to have
been well over 2.9 — apparently indicative of a
fast running gait equivalent to a mammalian
gallop or sprint. In the few remaining cases
relative stride length is estimated to have been
between 2.0 and 2.9 — indicative of a gait
equivalent to mammalian trotting. Our estimates
of relative stride length (and hence of speed)
depend in turn upon estimates of hindlimb height.
The piling of estimate upon estimate may, indeed,
have introduced and multiplied some errors, but
these are unlikely to be of very great significance
for our general conclusions. For example, in the
case of the Wintonopus track-makers our
estimates of hindlimb height are consistently
greater than those that would be obtained by
straightforward application of Alexander’s
method (1976). Consequently relative stride
length (and speed) may, if anything, be under-
estimated for these animals. Our estimates of
hindlimb height for the Skartopus track-makers
are slightly smaller than those that would be
obtained with Alexander’s method; but even if
our estimates are increased by as much as 40 or
50% it still appears that these dinosaurs would
have been running (compare Tables 5 and 6). In
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
445
any event, it is not necessary to estimate hindlimb
height in order to demonstrate that the
Wintonopus and Skartopus track-makers had an
exceptionally long-striding gait: this is fully
apparent from simple ratios of SL/FL and
PL/FL (see Figs 1 1 and 16). Once again it appears
that the Lark Quarry dinosaurs were indulging in
unusual behaviour, for reports of trackways
attributed to running dinosaurs are otherwise very
few (see Farlow 1981, Thulborn 1982). Here it is
worth recalling that the single Wintonopus
trackway at another site (New Quarry) is that of a
walking animal.
In summary, the Lark Quarry site has revealed
a most unusual assemblage of dinosaur
trackways, and to account for their origin it seems
legitimate to postulate some exceptional pattern
of behaviour on the part of the track-makers. The
findings of this study seem to confirm, or even
strengthen, our earlier interpretation of the
trackways as evidence of a dinosaur stampede.
Indeed, it is not easy to propose (let alone justify)
an alternative interpretation. This difficulty arises
for two reasons: (1) because all the Wintonopus
and Skartopus trackways are unidirectional, and
(2) because all these trackways seem to have been
made by running dinosaurs. It seems very unlikely
that this assemblage of trackways could represent
a series of events — i.e. the passage, at intervals,
of individual animals or of small groups. Any
such series of events would have involved the
most remarkable coincidences: at different times
various dinosaurs would have traversed the same
area, but always in exactly the same direction,
and always at a run. Consequently one is forced
to conclude that the Lark Quarry trackways
represent a single event — a conclusion supported
by the uniform preservation of the trackways. We
submit that a group of more than 150 animals
running in one direction must constitute a
stampede or some similar event. Several
experienced stockmen have examined the Lark
Quarry trackways; all of them agreed that the
trackways could well have resulted from a
stampede, though on two occasions we were
offered an alternative explanation (as a joke) —
that the track-makers were being ‘herded’ or
‘driven’.
There arise some intriguing questions. First,
what caused the stampede ? Only one piece of
fossil evidence seems to hint at a plausible answer
— the trackway of the single carnosaur. It is quite
conceivable that a gathering of ornithopods and
coelurosaurs, drinking or foraging round a water-
hole, might have been startled by the approach of
a large predatory dinosaur. We have reservations
about reading too much significance into the
evidence of a single dinosaur trackway, but the
only alternative is to admit that the unusual
behaviour of the Wintonopus and Skartopus
track-makers is inexplicable. A second question
arises: why did the ornithopods and coelurosaurs
run to the NE if this was the very direction from
which the carnosaur had approached them? Here
we can only offer speculations. The ornithopods
and coelurosaurs had reached the water-hole, to
the SW of the present Lark Quarry site, by some
unknown and presumably preferred or ‘normal’
route. One might have expected these animals to
have made their escape by such a route. The fact
that some, at least, did not do so seems to imply
that their preferred route had been blocked —
perhaps by the manoeuvres of the carnosaur,
which certainly made a sharp right turn. In
making this turn to its right the carnosaur would
simultaneously have opened up a new escape
route — to the NE, and along the broad drainage
channel that extended over the present Lark
Quarry site towards Seymour Quarry. This
reconstruction of events (Fig. 25) accords with all
available evidence and seems to be fairly
parsimonious. The motives of the carnosaur
remain unknown; it may simply have been
approaching the water-hole to drink, or it may
have been hunting — perhaps attempting to
corral its prey on the point extending SW into the
water-hole. If the animal were hunting it is
possible to speculate a little further. First, it seems
likely that this large predator would have selected
its prey from among the ornithopods; the
coelurosaurs, insofar as they are known from
their trackways, would seem to have been rather
small game. Next it is conceivable that the
carnosaur may not have been hunting alone; it
might possibly have been assisted by another
carnosaur (or perhaps even more than one)
strategically placed to forestall the escaping
ornithopods and coelurosaurs. This is no more
than a speculation, but it might help to account
for the remarkably close grouping of the
ornithopods and coelurosaurs — especially since
their progress over the Lark Quarry site seems not
to have been constrained by physical barriers.
These suggestions are not inconsistent with
previous speculations on the hunting behaviour of
large theropod dinosaurs (Farlow 1976).
Whatever the carnosaur’s maneouvres or
intentions may have been, a number of
Wintonopus and Skartopus track-makers did run
to the NE, across the present Lark Quarry and
446
MEMOIRS OF THE QUEENSLAND MUSEUM
Seymour Quarry sites. In doing so these animals
seem to have traversed an area that they had not
trodden before (or, at least, not in the immediate
past). Their trackways at Lark Quarry are strictly
unidirectional, and none of them seems to have
been made by a walking animal. Evidently these
ornithopods and coelurosaurs were not following
some well-trodden path, and they may in fact
have been moving across an area that was
formerly unattractive to them. It is not difficult to
imagine in what sense the Lark Quarry area might
have been unattractive to small bipedal dinosaurs:
it was covered by a layer of soft mud, into which
the animals might (and certainly did) sink to a
depth of several centimetres. This would have
been of little consequence to a very large animal
such as the carnosaur; as this animal traversed the
area its feet plunged right through the mud to rest
on the firmer sandy sediments below. But it is
conceivable that the small coelurosaurs (mean hip
height about 17 cm) and ornithopods (mean hip
height about 35 cm) ran some risk of becoming
bogged. If these small dinosaurs were crossing an
unattractive or even dangerous area, it is
reasonable to suppose that they were doing so
under some quite unusual and compelling
circumstances.
Our reconstruction of the events at and around
Lark Quarry is illustrated in Fig. 25. This
reconstruction takes into account all the
peculiarities of the Lark Quarry trackways, and
we have found no evidence that conflicts with it.
A central assumption of this reconstruction is that
the behaviour of the carnosaur was responsible,
at least in some measure, for the unusual
behaviour of the ornithopods and coelurosaurs.
This assumption naturally implies that very little
time would have elapsed between the formation
of the carnosaur’s trackway and the formation of
the ornithopod and coelurosaur trackways. The
uniform preservation of all these trackways seems
to indicate that they were formed at about the
same time ... but there is no certain way to
discover if the carnosaur preceded the
ornithopods and coelurosaurs by a matter of
minutes or by a matter of hours. However there is
one suggestive clue, provided by ornithopod and
coelurosaur footprints superimposed on those of
the carnosaur. In their preservation these
superimposed prints are identical to those
elsewhere, yet they were formed in thin streaks
and pockets of mud remaining in the floor of the
carnosaur’s prints. The fact that these remnant
patches of mud had not dried out, despite their
thinness, suggests that little time elapsed (or that
drying was very slow). If the intervening period
were to be estimated in terms of hours, rather
than minutes, we could no longer maintain our
suggestion that the carnosaur’s behaviour
prompted the stampede of ornithopods and
coelurosaurs. Even so, it would remain clear that
a stampede (or some similar event) did occur —
though its cause would be unknown.
The minimum distance travelled by the
stampeding ornithopods and coelurosaurs is more
than 95 metres — from the SW end of Lark
Quarry to the NE end of Seymour Quarry. From
the .speed estimates presented earlier it may be
calculated that most of the ornithopods and
coelurosaurs covered this distance in less than 30
seconds; at their minimum speeds the slowest
ornithopod and the slowest coelurosaur would
have done so in 38 seconds and 45 seconds
respectively. There is no indication of either the
start or the end of the stampede.
Finally, it might be objected that our use of the
term ‘stampede’ is inappropriate because the
animals involved seem to have been moving at
rather low speeds (mean 16 km/h for
ornithopods, and mean 12 km/h for
coelurosaurs). In a present-day stampede,
comprising ungulate mammals, one might expect
to find animals moving several times faster than
the dinosaurs at Lark Quarry. However, such
comparisons of absolute speeds are of limited
significance — simply because the dinosaurs that
traversed Lark Quarry were, on the whole, much
smaller than living ungulates. To measure and
compare the locomotor performances of
different-sized animals it is necessary to select
criteria that will reduce or eliminate the effects of
size-differences. Such criteria will be examined
below.
IMPLICATIONS FOR THE
UNDERSTANDING OF DINOSAUR
BIOLOGY
It has sometimes been supposed that juvenile
dinosaurs were rare (Richmond 1965, Leonard!
1981), and that small (but adult) dinosaurs were
equally rare or ‘unknown* (Bakker 1972).
However, there have recently been reports of
juvenile dinosaurs, some no bigger than rats or
pigeons, and even embryonic dinosaurs (see
Bonaparte and Vince 1979, Kitching 1979,
Coombs 1980a, 1982, Carpenter 1982, among
others). The Lark Quarry trackways provide
striking confirmation that small dinosaurs —
whether adults or juveniles, or both — may have
been abundant in some localities. This fact should
certainly have some bearing on the debate over
thermoregulation in dinosaurs, though at this
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
447
point we cannot pursue so large and complicated
a subject (see Thomas and Olson 1981 for a
comprehensive review).
It seems impossible to distinguish with certainty
between trackways made by small (but adult)
dinosaurs and those made by juveniles.
Consequently it is fruitless to use ichnological
data in speculating about the reproductive
strategies and population dynamics of dinosaurs.
These aspects of dinosaur biology should more
properly be investigated on the basis of body
fossils — where it might prove possible to
determine relationships between age, sexual
maturity and body size. Nevertheless we have
outlined some possibilities regarding population
structure of the Wintonopus and Skartopus track-
makers (Fig. 21). It must be emphasized that these
are only possibilities, for we cannot agree entirely
with the assumption that footprints of different
sizes were made by animals of different ages
(Leonard! 1981). Such an assumption is valid only
if growth proceeded at a constant rate through the
lives of the track-makers. Moreover the
assumption would seem to be particularly suspect
where there is a limited range in size of footprints
(e.g. Skartopus, where the largest footprint is less
than twice the size of the smallest). Even so, it
may be legitimate to identify very small footprints
as those of juveniles in cases where there is a very
great range in footprint size (e.g. Wintonopus,
where the largest footprint is nearly 12 times the
size of the smallest).
Ostrom (1972) compiled data on dinosaur
trackways in order to examine the possibility that
some dinosaurs might have been gregarious. At
various sites he found trackways grouped in near-
parallel arrangement, and he concluded that
several different kinds of dinosaurs may well have
been gregarious in habit. Ostrom’s conclusion is
supported, incidentally, by the identification of
dinosaurian adaptations for intra-specific
combat, display and vocalization (Galton 1970,
Hopson 1975, Molnar 1977, Weishampel 1981).
The trackways at Lark Quarry provide strong
evidence of gregarious behaviour, for they seem
to have resulted (with the exception of the
carnosaur trackway) from the movement of a
single large group of dinosaurs. This group
seems, however, to have been heterogeneous and,
perhaps, rather disorganized. The trackways of
coelurosaurs (Skartopus) and ornithopods
(Wintonopus ) are thoroughly intermingled, and
there is no clear indication that the two types of
dinosaur were segregated. This could indicate that
these coelurosaurs and ornithopods were in the
habit of moving together as a ‘mixed herd’, as
was suggested by Krassilov (1980). In this case
one might evisage the coelurosaurs as
opportunists — ready to seize insects and other
small animals as they were flushed from
vegetation by an ornithopod herd moving
through its feeding grounds. Alternatively the
trackways at Lark Quarry could have resulted
from accidental mingling of an ornithopod herd
with one or more foraging parlies of
coelurosaurs. Even so it would be reasonable to
regard both the ornithopods and the coelurosaurs
as gregarious, for it seems unlikely that so many
individuals could have gathered independently
and at random in the vicinity of the Lark Quarry
site. Nevertheless it is just conceivable that all the
Lark Quarry track-makers were normally solitary
animals: they might have been attracted to the
Lark Quarry water-hole during a period of
drought. However, we have found no desiccation
cracks or other evidence of drought, and it may
be recalled that all the trackways seem to have
been formed in moist sediment. Moreover, the
much-trampled claystone layer at New Quarry
seems to confirm that the Lark Quarry area was
quite commonly frequented by large numbers of
dinosaurs.
Several studies of dinosaur trackways have
employed Alexander’s method (1976) to estimate
the speeds of the track-makers (Russell and
Beland 1976, Tucker and Burchette 1977,
Coombs 1978, Thulborn and Wade 1979, Kool
1981, Farlow 1981, Thulborn 1981, 1982), These
studies have made it possible to compare the
speeds of various dinosaurs under various
circumstances (e.g. see Russell and Beland 1976),
or to compare the speeds of dinosaurs to those
recorded for mammals and ground-dwelling birds
(e.g. see Farlow 1981). Such comparisons of
absolute speeds are certainly interesting, but they
often give a poor indication of relative locomotor
performances. A mammalian analogy will make
this clear: if a horse, a dog and a mouse all move
at 6 km/h, the horse will be walking, the dog will
be trotting, and the mouse will literally be
galloping (Heglund et al, 1974). Absolute speed is
identical for all three animals, but their
locomotor performances are obviously very
different. Exactly similar relationships between
size, speed and gait would have prevailed in
dinosaurs, despite KooTs generalization (1981)
that different-sized animals, including dinosaurs,
‘all walk at roughly the same speed’
To evaluate and compare the locomotor
performances of dinosaurs it is desirable to adopt
some criterion that will reduce or eliminate the
effect of differences in body size. In comparing
448
MEMOIRS OF THE QUEENSLAND MUSEUM
the locomotor performances of fishes it is
common practice to express burst speeds in terms
of body lengths per second, and by analogy the
speeds of dinosaurian track-makers might
conveniently be expressed in terms of h /s (a ‘size-
related' speed, where h is height at the hip). Table
8 presents examples of such ‘size-related' speeds,
compared to absolute speeds, for a variety of
dinosaurian track-makers. Evidently dinosaurs
with similar absolute speeds may have very
different ‘size-related' speeds, and vice versa. In
terms of such ‘size-related' speed the locomotor
performances of the Lark Quarry ornithopods
and coelurosaurs are outstanding, although in
terms of absolute speed these animals seem to
have been moving rather slowly. However, such
‘size-related’ speed is no more useful for
comparing locomotor performances than is
absolute speed. The following (hypothetical)
example, where three different-sized animals are
moving at the same ‘sized-related' speed, will
make this clear:
‘size-related’
relative
h
speed
absolute speed stride length
gait
(m)
{h/s)
(m/s)
iX/h)
0.5
3.0
1.5
1.8
walk
1.5
3.0
4.5
2.5
trot
3.0
3.0
9.0
3.1
run
Evidently the effects of size-differences are
undiminished, because large animals need to
attain faster gaits and higher absolute speeds in
order to match the ‘size-related’ speeds of small
animals.
Heglund, Taylor and McMahon (1974)
proposed that ‘speed at the trot-gallop transition
point is a “physiologically similar speed” for
animals of different size’. We may extend this
proposition to identify two points at which
animals of different sizes would attain
‘physiologically similar' speeds: the walk-trot
transition, and the trot-run transition. At such
points different-size animals will have different
absolute speeds (and different ‘size-related’
speeds), but their locomotor performances may
be regarded as equivalent. These two points may
be defined in terms of relative stride length (X/h
about 2.0 and 2.9 respectively). They may also be
defined in terms of Froude number (u/gh; see
Alexander 1976), or in terms of ‘dimensionless
speed’ u(gh)^^\ see Alexander 1977). It is probably
most convenient to compare locomotor
performances in terms of X/h, because estimates
of this ratio have been cited in previous studies of
speed in dinosaurian track-makers. From the
estimates of X/h presented here (Tables 4, 5 and
8) it is clear that the locomotor performances of
the Lark Quarry ornithopods and coelurosaurs
were superior (and often far superior) to those of
most other dinosaurian track-makers. Estimates
of Froude number and of ‘dimensionless speed’
(equivalent to the square root of Froude number)
point to the same conclusion.
Relative stride length would seem to be a useful
and fairly realistic basis on which to evaluate and
compare the locomotor performances of
different-size animals. But even on this basis an
element of bias will emerge because there
probably exists a negative correlation between
body size and maximum X/h; this seems to be the
case among some living mammals (see data
presented by Alexander el al. 1977), and it is
reasonable to suppose that a similar relationship
between size and gait prevailed among dinosaurs.
Consequently straightforward comparisons of
X/h might in some cases be a little misleading; for
example, two dinosaurian track-makers with X/h
estimated at 2.0 could scarcely be regarded as
maintaining equivalent performances if one of
them were a large dinosaur moving at maximum
speed and the other were a small dinosaur capable
of accelerating to greater speeds. Unfortunately
there is no way to eliminate this bias, because
there is insufficient evidence (either from living
animals or from dinosaur trackways) to
determine the regression of maximum X/h on
body size (represented by h or by body mass). All
that may be said, in general terms, is that small
dinosaurs probably attained higher values for
maximum X/h than did large dinosaurs. Indeed, it
has been maintained (Thulbom 1982) that giant
dinosaurs were unable to extend X/h beyond 2.0
and were physically incapable of running.
Nevertheless comparisons of X/h would seem, for
the present at least, to give the best indication of
relative locomotor performances among
dinosaurs. And on this basis the performances of
the Lark Quarry ornithopods and coelurosaurs
appear to be exceptional.
If the ornithopods and coelurosaurs at Lark
Quarry were caught up in a stampede, or some
similar event, one might expect these animals to
have been running at or near their maximum
speeds. And if this were the case it might be
possible to determine a relationship between body
size {h ) and maximum running speed. Such a
relationship might then be used to gain some idea
of the maximum speeds of dinosaurs in general.
Fig. 24A is a plot of estimated mean speed against
estimated hip height for ornithopods and
coelurosaurs at Lark Quarry. In this diagram it is
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
449
TABLE 8: A Comparison of the Locomotor Performances of Various Dinosaurian Track-Makers.
A. LARGE THEROPODS
relative
size-
absolute
stride
related
Froude
dimension-
ichnotaxon or h
speed
length
speed
number
less speed
track-maker (m)
(m/s) (km/h)
(X/h)
(h/s)
(uVgh)
{u(gh)-0-5)
source
*Megalosaurus insignis: 1 . 5
2.4
8.6
1.7
1.6
0.38
0.62
de Lapparent and
Zbyszewski 1957
Irenesauripus mcleani: 1.7
0.7
2.4
0.8
0.4
0.03
0.17
Sternberg 1932
*Megaiosaurus: 1.3
2.2
8.0
1.6
1.3
0.29
0.54
de Lapparent and
Zbyszewski 1957
*Irenesaunpus acutus: 2.2
2.5
8.9
1.6
l.I
0.28
0.53
Sternberg 1932
Vrenesauripus
occidentalis: 2.3
0.9
3.3
0.9
0.4
0.04
0.19
Sternberg 1932
cf Tyrannosauropus: 2.6
1.9
6.9
1.3
0.7
0.15
0.38
this paper
* Tyrannosauropus
petersoni: 3.6
2.7
9.6
1.4
0.7
0.20
0.45
Haubold 1971
B. SMALL AND MEDIUM-SIZED
THEROPODS
*Gratlator gracilis: 0.17
0.8
2.9
1.7
4.8
0.41
0.64
Lull 1953
*PIesiornis pilulatus: 0.20
0.9
3-3
1.8
4.7
0.43
0.66
Lull 1953
*Grallator cursorius: 0.32
2.4
8.7
2.3
7.5
1.86
1.36
Lull 1953
*Hopiichnus shingi: ^0.55
13.1
47.2
7.0
23.8
31.99
5.66
Welles 1971
*Hopiichnus shingi: ^1 *00
8.2
29.5
3.8
8.2
6.86
2.62
*Anchisauripus
sillimani: 0.69
0.5
1.9
0.9
0.8
0.04
0.20
Lull 1953
*Grallator formosus: 0.82
1.6
5.8
1.6
2.0
0.33
0.57
Lull 1953
*Saltopoides igalensis: 0.83
8.4
30.1
4.2
10.1
8.70
2.95
de Lapparent and
Montenat 1967
*Anchisauripus
exsertus: 1-07
2.1
7.7
1.8
2.0
0.43
0.66
Lull 1953
*Dilophosauripus
williamsi: 1.37
1.3
4.8
1.3
1.0
0.13
0.37
Welles 1971
*Ornithomimipus
angustus: L56
1.3
4.8
1.2
0.9
0.12
0.34
Sternberg 1926
^Theropod Q94/98:
12.1
10.1
43.6
36.4
4.9
3.9
10.4
6.9
12.94
7.16
3.60
2.68
Farlow 1981
1 48
"Theropod BLV/A3:
9.4
7.9
33.8
28.3
3.7
3.0
6.4
4.3
6.12
3.42
2.47
1.85
Farlow 1981
‘^Theropod 86/0-82:
11.9
10.0
42.8
35.9
4.3
3.5
7.8
5.4
9.55
5.34
3.09
2.31
Farlow 1981
Skartopus Nr 25: 0.18
2.2
8.0
2.7
12.1
2.73
1.65
Skartopus (vcitdin): 0.17
3.2
11.6
3.7
18.8
6.18
2.49
this paper^
Skartopus lAr 1: 0.13
4.4
15.9
4.9
33.1
14.92
3.86
C. ORNITHOPODS
*Anomoepus minimus: 0.28
0.3
1.2
0.9
1.2
0.04
0.20
Lull 1953
Amblydactylus
kortmeyeri: 0.64
0.8
2.8
1.1
1.2
0.10
0.31
Currie and Sarjeant
1979
450
MEMOIRS OF THE QUEENSLAND MUSEUM
*Anomoepus crassus: 0.84
0.6
2.3
0.9
0.7
0.05
0.22
Lull 1953
*Sauropus barrattii: 0.88
1.9
6.7
1.7
2.1
0.40
0.63
Lull 1953
Irenichnites gracilis: 0.89
1.6
5.9
1.6
1.9
0.31
0.56
Sternberg 1932
*Satapliasaurus
0.30
Gabouniya 1951
dsocenidzei: 1 .08
1.0
3.5
1,1
0.9
0.08
‘Gypsichnites pacensis: 1 .74
1.3
4.8
1.2
0.8
0.10
0.32
Sternberg 1932
3 44
^Ornithopod:
7.6
2.4
27.2
8.6
2.7
1.3
2.2
0.7
1.70
0.17
1.30
0.41
Brown 1938
Wintonopus Nr 44: 0.42
2.5
9.2
2.2
6,0
1.53
1.24
Wintonopus (mean): 0.35
4.6
16.5
3.7
13.3
6.26
2.50
this paper
Wintonopus Nr 21: 0.29
6.3
22.7
5.0
21.7
14.03
3.75
Wintonopus (New
this paper
Quarry): 0.47
1.1
4.0
1.5
2.3
0.26
0.51
In most cases h is estimated by methods introduced in this paper.
*stride length estimated from published data (e.g. pace length or ratio of pace length: footprint length).
^with h estimated as for an ornithomimid; equation (14).
^with h estimated by Welles (1971).
‘^firsl estimate of h represents four limes footprint length, after Farlow (1981); second estimate is the mean of
two, calculated as for a carnosaur and as for a coelurosaur (equations (7) and (12) respectively).
'^all track-makers at Lark Quarry. First estimate represents the worst-performing track-maker (in terms of \/h),
and the third estimate represents the best-performing track-maker. Track-makers are listed by number in
Tables 4 (IVintonopus) and 5 (Skarlopus).
^two interpretations of single trackway (Russell and Beland 1976; Thulborn 1981), both with h estimated as
four times footprint length.
immediately obvious that the trend (or first
principal axis) of the distribution is roughly
parallel to the regression lines defining size/speed
relationships at the walk-trot transition {\/h 2.0)
and the trot-run transition (X/h 2.9). This
parallelism is not an artefact generated by our
methods for estimating size and speed (note
size/speed relationship for the New Quarry
ornithopod); nor does it appear to be fortuitous.
Instead it demonstrates very clearly that most
animals at Lark Quarry were running and,
moreover, that animals of different sizes were
maintaining equivalent locomotor performances
(in terms of X/h ). Mean X/h for the ornithopod
track-makers is estimated to be 3.69; for the
coelurosaur track-makers it is estimated to be
3.71. Fig. 24B is similar to Fig. 24A, except that it
illustrates the relationship between estimated
maximum speed and estimated hip height. In this
diagram the line drawn through the distribution is
not derived from our data: it is a line defin.ng the
theoretical regression of speed on size (/?) when
X/h is at a value of 3.93. (The figure of 3.93 was
selected on the basis of our findings: among the
ornithopod track-makers the mean figure for
maximum X/h per trackway is 3.94, and among
the coelurosaurs it is 3.92). Evidently the actual
relationship between size and speed conforms
quite closely to the theoretical relationship at this
particular value for X/h, In other words most of
the Lark Quarry track-makers seem to have been
running at a ‘physiologically similar [or standard]
speed’ — even though the track-makers were of
various sizes and had different absolute speeds. It
seems quite probable that the ‘physiologically
similar’ speed shared by the Lark Quarry
dinosaurs did represent maximum or near-
maximum speed. If this were not so one might
reasonably expect that small animals would have
matched the absolute speeds of larger ones.
Further, it is difficult to conceive of any
circumstances that might have led different-sized
dinosaurs to run at a ‘physiologically similar’
speed less than maximum speed. From the
evidence presented in Fig. 24 we may deduce that
small bipedal dinosaurs, with /; up to about 60
cm, could attain maximum X/h of at least 3.93. In
our preliminary account of the Lark Quarry site
(Thulborn and Wade 1979) we attempted to
account for the ‘rather low’ absolute speeds of the
track-makers by suggesting that the animals
might have been fatigued, or that they might have
been retarded by sinking deeply into the muddy
substrate. However, we have found no evidence
that the track-makers were decelerating to any
marked degree, and Alexander pointed out (1976)
that relationships between body size, speed and
stride length did not seem to be seriously affected
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
451
by the consistency of the substrate. Even so, it
may be more accurate to re-phrase our general
conclusion as follows: that the Lark Quarry
animals were running at maximum or near-
maximum speed under the conditions that then
prevailed. We cannot determine to what extent
those prevailing conditions might have affected
the locomotor performances of the Lark Quarry
dinosaurs.
If small bipedal dinosaurs were capable of
attaining maximum X/h about 3.93 we may
estimate the maximum speeds of these animals by
substituting 3.93/? for A in equation (2 ). This
equation may then be re-written as follows:
(75) u = [gh0.9^h
and simplied to give:
(76) u = (72.22/1 r
where h is in metres and u is solved in metres per
second. This equation may be applied equally well
to trackway data (with h estimated from footprint
dimensions) or to osteometric data (with h
measured directly). The maximum estimate of
X/h for any of the Lark Quarry track-makers is
5.03 (for a Wintonopus track-maker. No. 21 in
Table 4). If this figure (rather than 3.93)
represents maximum X/h for small bipedal
dinosaurs we may estimate their maximum speeds
by substituting it for A in equation (2 ). The
equation may then be re-written and simplified to
give:
(77) u = (U6hr
However, the assumption behind this equation is
that nearly all track-makers at Lark Quarry
would have been running at somewhat less than
their maximum possible speeds. Consequently it
seems reasonable to qualify our conclusions as
follows: the Lark Quarry track-makers seem to
have attained maximum X/h of at least 3.93 and,
in some instances, as high as 5.03. If these
conclusions do have more general application it
should be possible to predict maximum running
speed for any small bipedal dinosaur {h < 70 cm)
that is known from a skeleton or a trackway: its
maximum speed would probably lie between the
two estimates to be obtained with equations (76 )
and (77 ). It might also be legitimate to make use
of these equations in estimating maximum speeds
for some medium-size bipedal dinosaurs (with h
up to 1.5 or 2 m), but it is certainly not
appropriate to do so for very large bipeds or
quadrupeds. This is because a dinosaur moving
with \/h as high as 3.93 must incorporate an
unsupported interval in each stride, and the
ability to use unsupported intervals is generally
restricted to animals with body mass less than
500-800 kg (see discussions by Coombs 1978,
Thulborn 1982). Many large bipedal dinosaurs
were certainly above this critical weight limit, as
were nearly all of the quadrupedal forms (see, for
example, the body weights estimated for
dinosaurs by Colbert 1962). Coombs indicated
(1978) that the best mammalian runners had
optimum body mass of about 50 kg — but not
over 500 kg or below 5 kg ~ and it seems likely
that dinosaurs would have been under similar
physical constraints. In addition it is possible that
the maximum speeds of quadrupedal dinosaurs
were restricted by structural peculiarities of the
limbs and their girdles (see Thulborn 1982). Even
so, it may be legitimate to apply equations (76 )
and (77 ) in the case of some quite large bipedal
dinosaurs that seem to have been very lightly
constructed. Notable among these are the
ornithomimids or ‘ostrich dinosaurs'; these
animals possess striking cursorial adaptations and
are commonly supposed to have been the swiftest
of all dinosaurs (Russell 1972, Coombs 1978).
One example of the ornilhomimid
Dromiceiomimus has skeletal hip height of 1.22
metres and is estimated to have had a live body
weight of about 154 kg (Russell and Beland 1976);
with equation (76 ) the maximum speed of this
animal may be estimated at 9.31 m/s (33.5 km/h).
Among the ornithomimids described by
Osmolska et al. (1972) the largest example
{GaUimimus) has skeletal hip height of 1.94
metres; this dinosaur's maximum speed may be
estimated at 1 1 .82 m/s (42.6 km/h). These speeds
(c. 35-45 km/h) could conceivably be the highest
attained by any of the dinosaurs. However, it is
certainly possible that the ornithomimids were
able to extend relative stride length beyond 3.93
— particularly in view of their cursorial
adaptations. Russell and Bdand (1976) used
Alexander’s method (1976) to consider the
hypothetical example of an ornithomimid {h
about 1.22 metres) running at 80 km/h; at this
speed the animal’s stride length would have been
about 8.6 metres, indicating X/h about 7.05. It is
difficult to imagine that any dinosaur could have
extended stride length to such a degree. Among
living mammals such a high figure for X/h is
achieved only by the most highly adapted of
quadrupeds — which are able to employ stride-
lengthening techniques unavailable to bipeds (e.g.
scapular rotation and flexion/extension of the
vertebral column). An example may help to make
this clear. Al a speed of 10 m/s (36 km/h) a
452
MEMOIRS OF THE QUEENSLAND MUSEUM
human sprinter with hip height about 95 cm will
have relative stride length in the region of 4.6 —
as estimated with equation (2 ). To attain X/h of
7.0 a human athlete .must perform a leap. We
suspect that similar constraints apply to ratites,
though we have been unable to find suitable data
on these animals. By comparison it is unlikely
that a bipedal dinosaur could have maintained a
running gait with X/h as high as 7.05, even though
the ornithomimids may have reached maximum
values of X/h somewhat higher than 3.93. If the
largest ornithomimid mentioned above
(Gallimimus, with h of 1.94 metres) had been
capable of achieving maxmium X/h of 5.0 its
maximum speed would have been about 16 m/s
(58 km/h) — as estimated by means of equation
(17).
The general conclusion to be drawn from the
Lark Quarry trackways is that small bipedal
dinosaurs (h < 70 cm) attained maximum X/h of
at least 3.93, and possibly as high as 5.03. These
same figures for maximum X/h might also apply
to somewhat larger bipedal dinosaurs, providing
that these had live body weights less than 500-800
kg. Larger and heavier dinosaurs, both bipeds
and quadrupeds, probably attained lower figures
for maximum X/h — simply because they would
have been too heavy to have made use of
unsupported intervals. If the most highly adapted
of dinosaurian cursors — the ornithomimids —
did have maximum X/h of about 3.93 their
maximum speeds might have been about 35-45
km/h. Even if ornithomimids were capable of
attaining X/h as high as 5.03 their meiximum
speeds might still have been no greater than about
60 km/h. These estimates fall rather short of the
maximum possible speeds attributed to
ornithomimids on the basis of anatomical
comparisons (70-80 km/h, or even more; see
Russell 1972).
Are these general conclusions supported or
contradicted by evidence from other dinosaur
trackways? Trackways attributed to running
dinosaurs appear to be uncommon, but we will
examine those few examples that have come to
our attention. In describing a short section of
ornithopod trackway from the Cretaceous of
Colorado, Brown (1938) mentioned that each
footprint measured 34 inches (c. 86 cm) in width
and length, and that the track-maker had
‘stepped’ a distance of 15 feet (c. 4.6 metres).
assumed status of
h
X/h
track-maker
(m)
‘typical’ coelurosaur
0.70
4.9
carnosaur-like
0.93
3.7
ornithomimid-like
0.85
4.0
Browm did not suggest that this trackway had
been made by a running dinosaur: instead he
accounted for the remarkably long stride by
suggesting that the track-maker was a gigantic
creature nearly twice the height of Tyrannosaurus
in its classic standing pose (i.e. 35 feet as opposed
to 18 feet). By using Alexander’s methods (1976)
to determine speed and hip height Russell and
Beland (1976) estimated that this trackway had
been made by a very large animal (h about 3.44
metres) running at a speed of 7.54 m/s (27.1
km/h). Russell and Beland estimated that the
Colorado ornithopod had weighed about 1 1
tonnes, and from their figures it may be
calculated that X/h was in the region of 2.7. These
estimates have, at best, an indirect bearing on our
general conclusions: if a giant dinosaur was
capable of running at 27 km/h one might
reasonably expect the much smaller dinosaurs at
Lark Quarry to have matched, or even surpassed,
such a speed. However, it may be recalled that the
best available measure for comparing locomotor
performances seems to be relative stride length
(X/h), rather than absolute speed. Moreover there
is a suspicion that Brown may have
misinterpreted the Colorado trackway, and that
the dinosaur responsible for it was actually
walking (X/h 1 .34) at a speed no greater than 8.5
km/h (see Thulborn 1981).
In Saltopoides igalensis, the trackway of a
bipedal dinosaur from the Rhaeto-Liassic of
France, de Lapparent and Montenat (1967) found
the ratio PL/FL to be as high as 11/1 (see Fig.
16). These authors considered that the track-
maker was most probably a long-legged
coelurosaur; they gave a figure of 15.5 cm for
footprint length, and from their diagram of the
trackway (1967, fig. 15B) we estimate stride
length to have been about 344 cm. The
Saltopoides footprints are rather large by
coelurosaurian standards (see Table 2), and they
could quite possibly have been made by a
medium-size iheropod closer in appearance to
carnosaurs. Alternatively the track-maker might
have been a rather large coelurosaur with ‘typical’
hindlimb proportions, or a coelurosaur with
hindlimb proportions resembling those of
ornithomimids. By considering all these
possibilities we may obtain several estimates of
size and speed for the track-maker:
estimated speed equations used
(m/s)
(km/h
(h)
(speed)
9.45
34.0
(12)
(11)
7.57
27.3
(7)
(11)
8.10
29.2
(14)
(11)
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
453
In each case X/h is found to be greater than 2.0,
so that speed is most appropriately estimated with
equation {11 ). These estimates of speed and
relative stride length are in fair agreement with
the conclusions we have drawn from the Lark
Quarry trackways, and they might be taken to
indicate that the Saltopoides track-maker was
running at or near its maximum speed. It is
noteworthy that all three estimates of X/h are
below 5.0.
From the Kayenta Formation of Arizona (Early
Jurassic or Late Triassic) came a sequence of
three dinosaur footprints described by Welles
(1971) as Hopiichnus shingi. The maker of this
assumed status of
h
x/h
track-maker
(m)
‘typical’ coelurosaur
0.42
9.0
ornithomimid-like
0.55
7.0
ornithomimid-like
1.00
3.8
ornithomimid-like
1.00
3.8
In the last two cases h is the estimate by Welles
(1971); and in one of these speed is estimated by
means of equation (6 ), which is appropriate for
walking animals, even though X/h is greater than
2.0. It seems impossible to draw any firm
conclusions from these estimates of size, speed
and relative stride length. For the animal to have
been walking (with X/h less than 2.0) it would
need to have been at least 1.9 metres high at the
hip; this improbably high figure is equivalent to
19 times footprint length. It may be recalled that
Alexander (1976) found h to be approximately 4
times footprint length among bipedal dinosaurs;
and even on the assumption that the track-maker
could have been an ornithomimid-like dinosaur
we estimate h to have been less than 6 times
footprint length (see the second of the cases listed
above). If we adopt Welles’s estimate for h
(equivalent to 10 times footprint length) the
Hopiichnus track-maker is found to be similar to
the Lark Quarry track-makers in terms of
size/speed relationship {X/h 3.82 as opposed to
mean of 3.93). This close correspondence in
relative stride length might indicate that the
Hopiichnus track-maker was running at or near
its maximum speed — if it were indeed an
extremely long-limbed dinosaur. Estimates of h
obtained with equations {12) and {14) are
equivalent to 4.2 and 5.5 times footprint length
(see the first two cases listed above), but these
indicate that X/h was as high as 7.0 or 9.0. It is
difficult to imagine that any bipedal dinosaur
could have attained such values for relative stride
trackway was evidently a long-striding bipedal
dinosaur: footprint length was about 10 cm
whereas pace length was found to be 191 cm.
Welles commented that pace length was
‘tremendous* in relation to the size of the
footprints (compare data in Fig. 16), but he did
not suggest that the track-maker had been
running. He considered, instead, that the track-
maker had been an exceptionally long-limbed
animal (perhaps an ornithomimid) with h about 1
metre. Dr Welles also informs us (pers. comm.)
that the morphology of the footprints seems to
indicate a walking gait rather than a running gait.
We may consider several estimates of size and
speed for the Hopiichnus track-maker:
estimated speed equations used
(m/s)
(km/h)
{h)
(speed)
16.03
51.1
(12)
(11)
13.10
47.2
(14)
(11)
8.18
29.5
-
(11)
7.32
26.4
-
(6)
length. In summary, we are unable to offer any
satisfactory interpretation of the Hopiichnus
trackway. If the track-maker had hindlimb
proportions resembling those in any known
dinosaur it must have been progressing in a series
of phenomenal leaps. If the track-maker had been
using a running gait {X/h from 2.9 up to about
5.0) it must have had hindlimbs about twice as
long as those of an ornithomimid with
comparable foot length. And if the track-maker
had been walking {X/h less than 2.0) it must have
had hindlimbs about 3 times as long as those of an
ornithomimid with comparable foot length.
There is no indication that the animal might have
been swimming, and only touching down
occasionally with its feet (cf. theropod trackways
described by Coombs 1980b).
Alexander’s methods (1976) have recently been
applied by Farlow (1981) to a series of 15
dinosaur trackways in the Cretaceous of Texas.
Three of these trackways seem to have been made
by fast-running animals, with X/h in the range 3.7
to 4.9 and speeds estimated from 30 to 43 km/h.
Once again it is noteworthy that estimates of X/h
are less than 5.0. Farlow identified the Texas
track-makers as theropod dinosaurs, and from
the size of their footprints they might be
envisaged either as exceptionally large
coelurosaurs or as small to medium-size
carnosaurs. Consequently it is possible to
compare several estimates of size and speed for
these track-makers:
454
MEMOIRS OF THE QUEENSLAND MUSEUM
assumed status of
h
X/h
estimated speed
equations used
track-maker
(m)
(m/s)
(km/h)
(h)
(speed)
*Q94/98:
theropod
1.16
*4.9
*11.9
*42.8
-
(6)
theropod
1.16
*4.9
*12.1
43.6
-
(11)
coelurosaur-Iike
1.42
4.0
10.3
37.0
(12)
(11)
carnosaur-like
1.49
3.8
9.9
35.7
(7)
(11)
*BLV/A3:
theropod
1.48
*3.7
*8.3
*29.9
-
(6)
theropod
1.48
*3.7
*9.4
33.8
-
(11)
coelurosaur-like
1.87
2.9
7.7
28.0
(12)
(11)
carnosaur-like
1.82
3.0
8.0
28.6
(7)
(11)
*86/0-82:
theropod
1.52
♦4.3
*11.1
♦39.9
-
(6)
theropod
1.52
*4.3
*11.9
42.8
-
(11)
coelurosaur-like
1.93
3.4
9.8
35.4
(12)
(11)
carnosaur-like
1,86
3.5
10.1
36.4
(7)
(11)
(*trackway identification numbers and estimates taken from Farlow, 1981)
None of these estimates seems to be in serious
conflict with our general conclusions. If h is
estimated as 4 times footprint length only one of
the track-makers (Q94/98) is found to have
attained X/h very much greater than 3.93. If h is
estimated with the methods introduced in this
paper it appears that this same track-maker would
have rivalled the Lark Quarry dinosaurs in its
locomotor performance [X/h 3.8 to 4.0).
To summarize, we have found no certain
evidence that any bipedal dinosaur greatly
surpassed the locomotor performances of the
Lark Quarry dinosaurs. The Colorado
ornithopod (Brown 1938) may have been walking
with X/h about 1.34 (Thulborn 1981); even if the
track-maker had been trotting or running (Russell
and Beland 1976) X/h would have been no greater
than 2.7. The Saltopoides track-maker (de
Lapparent and Montenat 1967) seems certainly to
have been running, with mean X/h of 4.2 (based
on three estimates for h). If this track-maker had
resembled carnosaurs or ornithomimids in body
build it would appear to have matched the
locomotor performances of the Lark Quarry
animals, having X/h in the range 3.7 to 4,0. But if
the Saltopoides track-maker is envisaged as an
exceptionally large coelurosaur with ‘typical’
hindlimb proportions its locomotor performance
[X/h 4.9) is matched by only a few of the Lark
Quarry dinosaurs. The Hopiichnus trackway
(Welles 1971) presents intractable problems of
interpretation. If this track-maker resembled any
known dinosaur in hindlimb proportions its
locomotor performance must have been
phenomenal: X/h would have been at least 7.0,
and possibly 9.0 or higher. It is difficult to believe
that any bipedal animal could sustain a running
gait with such figures for X/h. But if the
Hopiichnus track-maker had achieved this feat it
would be necessary to abandon, or at least
modify, the conclusions we have drawn from the
Lark Quarry trackways. In this case further
problems would arise. If the Hopiichnus track-
maker and the Lark Quarry track-makers were
running at or near maximum speed we might be
forced to question Alexander’s findings (1976,
1977) on the relationships of size, speed and gait
in living tetrapods. Alternatively we must suppose
that the Lark Quarry dinosaurs were very severely
retarded by sinking into the muddy substrate
(with the effect of reducing X/h from at least 7.0
to 4.0 or less). If the Lark Quarry track-makers
had been running well below their maximum
possible speeds (but were not seriously retarded
by the muddy substrate) another question will
emerge: what circumstances caused these animals
to run at a ‘physiologically similar speed’ [X/h =
3.93) a good deal less than maximum speed? We
cannot find a satisfactory answer. Next it might
be surmised that the Hopiichnus track -maker had
been travelling with X/h no greater than 5.0 (the
maximum estimate from any trackway considered
here); in this case the track-maker must be
envisaged as an animal with hindlimbs very much
longer (relative to foot length) than those in any
known dinosaur. Evidently all these
interpretations of the Hopiichnus trackway
present difficulties; for the present we must
regard the significance of this trackway as
uncertain or equivocal. Finally we estimate that
the three fastest of the Texas theropods (Farlow
1981) were travelling with X/h between 2.9 and
4.0. These theropods, whether coelurosaurs or
carnosaurs, seem to have maintained locomotor
performances equivalent or inferior to those of
the Lark Quarry dinosaurs.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
455
CONCLUSIONS
The most reliable guide to the size of a
dinosaurian track-maker is probably footprint
size index (SI) — rather than footprint length
(FL) or any similar dimension. This conclusion is
based on analysis of variance in a sample of 57
WintonopiiS trackways, and it remains to be
tested elsewhere. The Skartopus trackways
cannot be used to lest this conclusion because
they do not show sufficient variation in size.
In bipedal dinosaurs the anatomy and posture
of the foot were such that metatarsus length (MT)
can be estimated on the basis of footprint
dimensions (Fig. 18). Such an estimate of MT can
then be used to predict skeletal hip height {h)
because these two dimensions are strongly
correlated in each major group of bipedal
dinosaurs. We provide allometric equations to
predict h in the following groups of dinosaurs;
coelurosaurs (with ‘typicaT hindlimb
proportions), ornithomimids, carnosaurs,
ornithopods in general, cursorial ornithopods,
and graviportal ornithopods. These equations
were used to obtain the following estimates of h
for dinosaurian track-makers at the Lark Quarry
site: about 2.6 m for the single carnosaur
(trackway identified as cf. Tyrannosauropus)\
from 14 cm to 70 cm for the numerous
ornithopods (trackways identified as Wintonopus
latomorum ichnogen. et ichnosp. nov.), but with
one large individual at about 1.6 m; from 13 to 22
cm for the numerous coelurosaurs (trackways
identified as Skartopus australis ichnogen, et
ichnosp. nov.).
The carnosaur traversed the Lark Quarry area
from NE to SW, and a mixed group of
ornithopods and coelurosaurs subsequently
crossed the same area in the opposite direction.
This mixed group comprised at least 150 animals.
The gaits of these (and other) dinosaurian track-
makers must be defined arbitrarily: this is because
bipedal dinosaurs had the same sequence of limb
movements at all speeds (and because the
sequence of limb movements is unknown in
quadrupedal dinosaurs). We define three
dinosaurian gaits on the basis of relative stride
length (A//?): a walking gait {X/h< 2.0), a trotting
gait (A/A between 2.0 and 2.9), and a running gait
(A/A> 2.9). These may be regarded as
‘physiologically similar’ to the walking, trotting
and running gaits of mammals (see Heglund et al.
1974, Alexander 1977). On this basis it is
determined that the carnosaurian track-maker
was walking (A/A 1.3) whereas the ornithopods
and coelurosaurs were using a fast running gait
equivalent to cantering or galloping in mammals
(mean A/A about 3.7). An ornithopod track-
maker at a second site (New Quarry) was found to
have been walking (A/A 1,5).
The relationships between size (A), speed and
gait in living tetrapods (see Alexander 1976, 1977;
Alexander et al. 1977) were used to estimate the
speeds of the track-makers. It is estimated that
the carnosaur was walking at a speed of about 7
km/h; for a sample of 56 ornithopods mean speed
is estimated to have been about 16 km/h, and for
a sample of 34 coelurosaurs it is estimated to have
been about 12 km/h. For the single ornithopod at
New Quarry estimated speed is 4 km/h.
The findings of this study support our
preliminary interpretation of the Lark Quarry
trackways — that the ornithopods and
coelurosaurs were caught up in a stampede, which
may have been generated by the approach of the
carnosaur (Thulborn and Wade 1979, Wade
1979). We can find no evidence that conllicts with
this interpretation. Indeed, it is difficult to
imagine any other circumstances that might
account for a mixed group of 150 dinosaurs
running in a single direction. Moreover there is
some indication that the ornithopods and
coelurosaurs were running at or near their
maximum speeds (under the conditions that
prevailed): different-sized individuals were
moving at different absolute speeds, but they
seem to have maintained a ‘physiologically
similar speed’ measured in terms of relative stride
length (A/A= 3.7). It is difficult to believe that so
many animals could have maintained and shared
a ‘physiologically similar speed’ other than
maximum speed.
To measure and compare the locomotor
performances of dinosaurian track-makers it is
desirable to adopt some criterion that will
eliminate (or at least reduce) the effects of
differences in body size. Direct comparisons of
absolute speed are of little value because they are
biased in favour of larger animals; comparisons
of a ‘size-related’ speed (A/s, analogous to body
lengths per second in studies of fish locomotion)
are equally biased in favour of small animals. Of
the criteria that are available for appraising
locomotor performance the most suitable would
seem to be relative stride length (A/A), Froude
number (see Alexander 1976), and ‘dimensionless
speed’ (see Alexander 1977). In terms of these
criteria the locomotor performances of the Lark
Quarry ornithopods and coelurosaurs are
outstandingly good; their performances are better
(and usually far better) than those of most other
dinosaurian track-makers.
456
MEMOIRS OF THE QUEENSLAND MUSEUM
The ornithopods and coelurosaurs at Lark
Quarry attained maximum \/h of at least 3.9, and
possibly as high as 5.0. This latter figure might
represent the maximum limit of relative stride
length for any bipedal dinosaur: it is difficult to
imagine that any bipedal animal could extend A / h
much beyond 5.0, and we have found no certain
evidence of any dinosaur having done so. If the
most highly adapted of dinosaurian cursors — the
ornithomimids — attained X/h of 5.0 their
maximum speeds might have been about 60
km/h.
Finally it is clear that small dinosaurs, whether
juveniles or adults (or both), may have been
abundant in some localities. It may not be
legitimate to identify small footprints as those of
juvenile dinosaurs, because dinosaurian rates of
growth are unknown and may not have been
constant. For this reason it is probably fruitless to
investigate dinosaurian demography on the basis
of ichnological data.
ACKNOWLEDGMENTS
Many residents in and around Winton took an
active interest in the work at Lark Quarry and
made generous offers of materia! assistance: in
particular we thank Peter Knowles (‘Namarva’),
Eric and Marjorie Bryce (‘Colston’), Roslyn and
Bob Blackett (‘Amelia Downs’), Arthur and
Roslyn Wallace (‘Cork’), and Ron McKenzie
(Winton). Among the many persons who played
roles in the discovery and excavation of the
trackways we extend particular thanks to
Malcolm Lark, Barbara Molnar, Duncan
MePhee, students from the University of New
South Wales and the University of Queensland,
members of the 6th Battalion, Royal Australian
Regiment, and local friends too numerous to list.
Our work relied heavily on support from
members of the Queensland Museum staff,
especially Alan Bartholomai, Errol Beutel, Don
Dale, Yvonne Evans, Ralph Molnar, Howard
Plowman, Andrew Rozefelds and Terry Tebble.
Most photographs are the work of Alan Easton,
and Fig. 1 is based on an aerial survey
commissioned by the Queensland National Parks
and Wildlife Service. Dave Norman (London),
Philip Currie (Alberta), Jim Farlow (Michigan)
and Sam Welles (California) offered useful
comments and information about dinosaur tracks
elsewhere in the world. One of the authors
(R.A.T.) received continuing financial support
from the Australian Research Grants Committee.
In June 1982 Lark Quarry and a surrounding
area (about 374 hectares) were designated an
Environmental Park under the joint trusteeship
of the Winton Shire Council and the Queensland
Museum, This end was achieved through the
goodwill and co-operation of Roslyn and Bob
Blackett (‘Amelia Downs’), members of the
Winton Shire Council, and officers of the
Queensland National Parks and Wildlife Service
(especially Alan Chenoweth and Warren Oxnam).
The Winton Shire Council has constructed an
access road to the trackway site, which is now
furnished with a permanent walkway and a
protective roof (designed by Duncan MePhee,
who also worked on site during excavations),
Neville Agnew (Conservator, Queensland
Museum) is undertaking a long-term study to
monitor and retard any deterioration of the
trackway surface.
To all these individuals and organizations we
express our sincere thanks.
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Buenos Aires 1978, vol. 1:215-22.
1981. Ichnological data on the rarity of young
in North East Brazil dinosaurian populations.
An. Acad, brasii Cienc. 53: 345-6.
Lull, R.S., 1953. Triassic life of the Connecticut
Valley (revised edn). Bull. Connecticut State
Geol. nat. Hist. Surv. 81: 1-331.
and N.E. Wright, 1942. Hadrosaurian
dinosaurs of North America. Geol. Soc.
Amer. Spec. Pap. 40: 1-242.
McWhae, J.R.H., P.E. Playford, A.W.
Lindner, B.F. Glenister, and B.E. Balme,
1958. The stratigraphy of Western Australia.
J. geol. Soc. Aust. 4: 1-161.
Molnar, R.E., 1977. Analogies in the evolution
of combat and display structures in
ornithopods and ungulates. Evol. Theory 3:
165-90.
1980. Australian late Mesozoic terrestrial
tetrapods: some implications. Mem. Soc.
geol. France 139; 131-43.
Osborn, H.F., 1917. Skeletal adaptations of
Ornitholestes, Struthiomimus and
Tyrannosaurus. Bull. Amer. Mus. nat. Hist.
35: 733-71.
1924. Psittacosaurus and Protiguanodon: two
Lower Cretaceous iguanodonts from
Mongolia. Amer. Mus. Novit. 127: 1-16.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
459
OSMOLSKA, H., E. Roniewicz, and R.
Barsbold, 1972. Results of the Polish-
Mongolian palaeontological expeditions, part
IV: A new dinosaur, Gallimimus bullatus n.
gen., n. sp, (Ornithomimidae), from the
Upper Cretaceous of Mongolia. Palaeont,
Polonica 27: 103-43.
OsTROM, J.H., 1972. Were some dinosaurs
gregarious? Palaeogeogr., Palaeoclimatol.,
Palaeoecol. 11: 287-301.
1978. The osteology of Compsognathus
/ong/pes" Wagner. Zitteliana 4: 73-118.
Parks, W.A., 1920. The osteology of the
trachodont dinosaur Kritosaurus
incurvimanus. Univ. Toronto Stud. Geol.
Ser. 11: 1-76.
Richmond, N.D., 1965. Perhaps juvenile
dinosaurs were always scarce. J. Paleont. 39:
503-5.
Russell, D.A., 1970. Tyrannosaurs from the
Late Cretaceous of western Canada. Nat.
Mus. Canada Palaeont. Putins 1: 1-34.
1972. Ostrich dinosaurs from the Late
Cretaceous of western Canada. Can. J. Earth
Sci. 9: 375-402.
and P. Beland, 1976. Running dinosaurs.
Nature, Lond. 264: 486.
Santa Luca, A.P., A.W. Crompton, and A.J.
Charig, 1976. A complete skeleton of the
Late Triassic ornithischian Heterodonto-
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Sarjeant, W.A.S., 1970. Fossil footprints from
the Middle Triassic of Nottinghamshire and
the Middle Jurassic of Yorkshire. Mercian
Geologist 3: 269-82.
1974. A history and bibliography of the study
of fossil vertebrate footprints in the
British Isles. Palaeogeogr., Palaeoclimatol.,
Palaeoecol. 16: 265-378.
1975. Fossil tracks and impressions of
vertebrates. Pp. 283-324 in ‘The study of
trace fossils’ (Ed. R.W. Frey). (Springer:
Berlin).
Senior, B.R., A. Mono, and P.L. Harrison,
1978. Geology of the Eromanga Basin.
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Bull. 167: 1-102.
SOKAL, R.S. and F.J. Rohlf, 1969. ‘Biometry’.
776 pp. (Freeman: San Francisco).
Staines, H.R.E., 1954. Dinosaur footprints at
Mount Morgan. Qd Govt Mining J. 55:
483-5.
Sternberg, C.M., 1926. Dinosaur tracks from
the Edmonton Formation of Alberta.
Canada, Mus. Bull., Geol. Ser. 44: 85-7.
1932. Dinosaur tracks from Peace River,
British Columbia. Ann. Kept Nat. Mus.
Canada 1930: 59-85.
Talbot, M., 1911. Podokesaurus holyokensis, a
new dinosaur from the Triassic of the
Connecticut Valley. Amer. J, Sci. (ser. 4) 31:
469-79.
Thaler, L., 1962. Empreintes de pas de
dinosaures dans les dolomies du Lias
inferieur des Causses (note prdliminaire).
C.R. somm. seanc. Soc. geol. France 1962:
190-2.
Thomas, R.D.K. and E.C. Olson, (Eds), 1980.
‘A cold look at the warm blooded dinosaurs’.
Amer. Assn Adv. Sci., Selected Symposium
Nr 28.
Thulborn, R.A., 1972. The post-cranial
skeleton of the Triassic ornithischian
dinosaur Fabrosaurus australis. Palaeon-
tology 15: 29-60.
1981. Estimated speed of a giant bipedal
dinosaur. Nature, Lond. 292: 273-4.
1982. Speeds and gaits of dinosaurs.
Palaeogeogr., Palaeoclimatol., Palaeoecol.
38: 227-56.
and M. Wade, 1979. Dinosaur stampede in the
Cretaceous of Queensland. Lethaia 12:
275-9.
Tucker, M.E. and T.P. Burchette, 1977.
Triassic dinosaur footprints from South
Wales: their context and preservation.
Palaeogeogr., Palaeoclimatol., Palaeoecol.
22: 195-208.
Wade, M., 1979. Tracking dinosaurs; the
Winton excavation. Aust. nat. Hist. 19:
286-91.
Weishampel, D.B., 1981. Acoustic analyses of
potential vocalization in lambeosaurine
dinosaurs (Reptilia : Ornithischia). Paleo-
biology 7:252-61.
Welles, S.P., 1954. New Jurassic dinosaur from
the Kayenta Formation of Arizona. Bull.
Geol. Soc. Amer. 65: 591-8.
1971. Dinosaur footprints from the Kayenta
Formation of northern Arizona. Plateau 44:
27-38.
Wills, L.J. and W.A.S. Sarjeant, 1970. Fossil
vertebrate and invertebrate tracks from
boreholes through the Bunter Series (Triassic)
of Worcestershire. Mercian Geologist 3;
399-414.
460
MEMOIRS OF THE QUEENSLAND MUSEUM
X
h-
o
z
IJJ
cc
Q.
}-
o
o
FIGURE 2. Diagrams to illustrate measurements of footprints and trackways. A, outline of ornithopod footprint
showing footprint length measured along or parallel to the axis of digit 3 (dotted line); footprint width is measured
at a right angle to footprint length. B, outline of coelurosaur footprint showing corresponding measurements ot
length and width. C, short section of ornithopod trackway showing measurements of two successive paces and a
single stride. Pace angulation (ANG) is calculated from the lengths of the paces and the stride (see ‘Methods ).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
461
FIGURE 3. Outline chart of the study area at Lark Quarry. Scale bar indicates 2 m. Trackway of the solitary
carnosaur is shown at left (C), and partly eroded trackway of an exceptionally large ornithopod is shown at right
(B). A representative portion of the bedding plane is enlarged (bottom centre) to illustrate the abundance and
orientation of small footprints attributed to ornithopods (D) and coelurosaurs (E). Trackways are identified by
corresponding letters in descriptions (p. 417). The number of track-makers was estimated by counting footprints
in a metre-wide transect between the points marked X. The area shown in outline was photographed (see PI. 4),
replicated in fibreglass and studied in detail. Adjoining areas of bedding plane (mainly to S and SW) were exposed
during excavations but were not studied in detail. An outlier or ‘island’ of overburden was left undisturbed in the
area indicated.
462
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 4, Variation in size and shape of the footprints at Lark Quarry. Scale bar indicates 20 cm. The four large
footprints are from the trackway of the carnosaur (cf. Tyrannosauropus ) and are identified by their number in
the sequence 1-11. Footprint 7 shows traces of large pointed claws (see also PI. 6); footprint 8 shows a
longitudinal crest formed by mud adhering to the underside of the track-maker’s middle toe. a, the largest
ornithopod footprint {Wintonopus) found at Lark Quarry, b, an ‘average’ ornithopod footprint {Wintonopus )
at Lark Quarry, based on mean dimensions in 284 examples, c, an ‘average’ coelurosaur footprint {Skartopus ) at
Lark Quarry, based on mean dimensions in 191 examples.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
463
FIGURE 5. Diagrams to illustrate variation in shape of ornilhopod footprints {Wintonopus ) at Lark Quarry. A,
complete and undislorted imprint of a right foot; all other diagrams illustrate variation on this basic footprint
shape. Examples B, C and D are fore-shortened or ‘stubby-toed* footprints formed by the toes entering and
leaving the sediment at a steep angle. In example D the toes entered the sediment vertically but did not sink to the
level of the interdigital web between digits 2 and 3. In examples E and F the foot has not sunk deeply enough to
leave traces of one or both of the inlerdigital webs. In examples G and H footprint width is reduced because the
foot entered and left the sediment obliquely — with the track-maker’s weight carried mainly on the outer two
digits. Examples I and J show backwardly directed scrape-marks; examples K and L show scrape-marks directed
anterolaterally.
464
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 6. Foot structure in small bipedal dinosaurs. In all cases the right fool is shown in anterior view and the
scale bar indicates 2 cm. A, foot skeleton in the Upper Jurassic ornithopod Nanosaurus; B, foot skeleton in the
Lower Cretaceous ornithopod Hypsilophodon; C, attempted restoration of foot structure in the Wintonopus
track-maker (based on mean dimensions in 284 footprints); D, foot skeleton in the Triassic coelurosaur
Coelophysis; E, foot skeleton in the Upper Jurassic coelurosaur Compsognathus; F, attempted restoration of foot
structure in the Skartopus track-maker (based on mean dimensions in 191 footprints). A and B after Gallon and
Jensen (1973); D and E after Ostrom (1978).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
465
/
•
\ C
3C
FIGURE 7. Morphological features of ornithopod footprints (Wintonopus ) related to events during the track-
maker’s stride cycle. Each diagram shows position of foot (at top, with distal end of metatarsus indicated by a
spot), longitudinal section of corresponding footprint (at middle), and corresponding plan view of right footprint
(at bottom). Stage 1: start of stride, with forwardly extended foot; initial footprint (shaded) is shallow and shows
positive rotation. Stage 2: as the track-maker moves forwards the foot sinks deeper, rotates to face directly ahead,
and slips backwards a little (unshaded footprint). Stage 3A: as foot starts to lift from the substrate the toes
continue to slip backwards, incising slots in the floor of the footprint. Stage 3B (following Stage 2, or via Stage
3A): toes slip back far enough to breach rear wall of footprint, producing backwardly-directed scrape-marks.
Stage 3C (following Stage 2): toes do not slip backwards but drag through front wall of footprint to produce
forwardly-directed scrape-marks.
466
MEMOIRS OF THE QUEENSLAND MUSEUM
FOOTPRINT LENGTH FOOTPRINT WIDTH:LENGTH STRIDE LENGTH
p,, RATIO CM
FOOTPRINT WIDTH FOOTPRINT SIZE INDEX
CM CM
FIGURE 8. Frequency distributions based on pooled data from Wintonopus (ornithopod) trackways at Lark
Quarry. All modal classes drawn to uniform height, and vertical scales are absolute frequencies. Diagrams for
footprint length, footprint width and footprint size index exclude data from single exceptionally large trackway
(No. 57 in Table 4; see also Fig. 9).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
467
MEAN FOOTPRINT LENGTH MEAN FOOTPRINT MEAN STRIDE LENGTH
CM WIDTH:LENGTH RATIO cm
CM CM CM
CM CM
FIGURE 9. Frequency distributions based on grouped data from Wintonopus (ornithopod) trackways at Lark
Quarry. All modal classes drawn to uniform height, and vertical scales are absolute frequencies.
MEAN STRIDE LENGTH cm MEAN FOOTPRINT WIDTH cm
468
MEMOIRS OF THE QUEENSLAND MUSEUM
%•
I •
L...
I
300
I
K-
o
200
Q
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I —
CD
100
• •••{^
•• -y. •
5 10 20
MEAN FOOTPRINT LENGTH cm
L....
5 10 20
MEAN FOOTPRINT LENGTH cm
400
300
200
100
50
L...-
. ^
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•
400
300
X
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100
50
V*
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5 10 20
MEAN FOOTPRINT SIZE INDEX cm
L...-
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5 10 20 30
MEAN FOOTPRINT WIDTH cm
FIGURE 10. Scatter diagrams based on grouped data from Wintonopus (ornithopod) trackways. In all cases both
axes have logarithmic scales. Note that the New Quarry track-maker (open circle) resembles Lark Quarry track-
makers (solid circles) in footprint proportions, but is distinguished by its relatively short stride. The same is true
(but less obviously so) for the single exceptionally large track-maker at Lark Quarry (star).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
469
10 20 30 40 50
MEAN FOOTPRINT LENGTH
FIGURE 11. Scatter diagram to illustrate relationship between pace length and footprint length in trackways
attributed to ornithopod dinosaurs. Solid circles — Wintonopus (at Lark Quarry; No. 57 is the single large
trackway formed at slightly earlier date); open circle — Wintonopus (at New Quarry); stars — Anomoepus;
triangles — Irenesauripus; open squares — various trackways including Amblydactylus, Gypsichnites and
Sauropus. Incorporating data from Sternberg 1932, Lull 1953, Currie and Sarjeant 1979.
470
MEMOIRS OF THE QUEENSLAND MUSEUM
1 ► 2A ► 3A ► 4A
I
FIGURE 12. Morphological features of coelurosaur footprints {Skartopus ) related to events during the track-
maker’s stride cycle. Each diagram shows position of foot (at top, with distal end of metatarsus indicated by a
spot), longitudinal section of corresponding footprint (at middle), and corresponding plan view of footprint.
Stage 1: start of stride, with forwardly extended foot; initially there is no footprint, or a very shallow one. Stage
2A; as the track-maker moves forwards the foot sinks deeper. Stage 3A: the foot lifts from the substrate, leaving
sharp imprints of the claws. Stage 4A (frequently follows Stage 3A): the toes slip backwards, incising slots in the
floor of the footprint. The sequence of Stages 2B to 5B is equivalent to the sequence 2A to 4A, but the foot does
not sink into the substrate; the only traces are scratches produced by the toes slipping backwards.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
471
FOOTPRINT LENGTH
CM
FOOTPRINT WIDTH:LENGTH
RATIO
CM
CM CM
FIGURE 13. Frequency distributions based on pooled data from Skartopus (coelurosaur) trackways at Lark
Quarry. All modal classes drawn to uniform height, and vertical scales are absolute frequencies.
472
MEMOIRS OF THE QUEENSLAND MUSEUM
MEAN FOOTPRINT WIDTH MEAN PACE LENGTH
CM CM
CM CM
50 60 70
MEAN STRIDE LENGTH
CM
to
5
50 60 70 80
MAXIMUM STRIDE LENGTH
CM
to
5
teo"^ 180 “
MEAN PACE ANGULATION
FIGURE 14. Frequency distributions based on grouped data from Skartopus (coelurosaur) trackways at Lark
Quarry. All modal classes drawn to uniform height, and vertical scales are absolute frequencies.
MEAN STRIDE LENGTH cm MEAN FOOTPRINT WIDTH cm
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
473
L....
MEAN FOOTPRINT LENGTH cm
MEAN FOOTPRINT SIZE INDEX cm
MEAN FOOTPRINT WIDTH cm
FIGURE 15. Scatter diagrams based on grouped data from Skartopus (coelurosaur) trackways at Lark Quarry. In
all cases both axes have logarithmic scales. Polygons define distributions for Wintonopus (ornilhopod) trackways
at the same site.
474
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 16. Scatter diagram to illustrate relationship between pace length and footprint length in trackways
attributed to small and medium-sized theropod dinosaurs. Solid circles — Skartopus (at Lark Quarry); sters —
Grallator; triangles — Anchisauripus; solid squares — Plesiornis; Sa — Saltopoides; St — Stenonyx; 1
unidentified theropods. Incorporating data from Lull 1953, de Lapparent and Montenat 1967, Bassoullet 1971,
Tucker and Burchette 1977, Farlow 1981 (two smallest of 15 unidentified theropods).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
475
METATARSUS LENGTH CM
FIGURE 17. Relationship between hindlimb height and metatarsus length in large theropod dinosaurs. Product-
moment correlation coefficient (r 0.979) is not improved by transformation of data. Least squares regression line
represents equation (7 ) in text. Based on data from Lambe 1917 (Gorgosaurus ), Osborn 1917 {Tyrannosaurus ),
Gilmore 1920 {AUosaurus, Ceratosaurus ), von Huene 1932 {Megalosaurus ), Welles 1954 {Dilophosaurus ),
Russell 1970 {Daspletosaurus ).
476
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 18. Diagrammatic comparison of dimensions in the foot of a bipedal dinosaur. The diagram represents a
vertical section along digit 3, with bones stippled and other tissues in outline. SP represents the sum of the lengths
of phalanges in digit 3. FL (footprint length) comprises 2P together with claw sheath, joint capsules, base of the
metatarsus and (perhaps) a fleshy ‘heel’ at point X. MT (length of metatarsus) is often about the same length as
HINDLIMB HEIGHT cm
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
477
METATARSUS LENGTH cm
METATARSUS LENGTH cm
FIGURE 19. Relationship between hindlimb height and metatarsus length in ornithopod dinosaurs. Logarithmic
scale on both axes. A, heterogeneous sample of 32 ornithopod dinosaurs; r 0.988; least squares regression line
represents equation {8 ) in text. B, same data* but with graviportal ornithopods (23 specimens) separated from
cursorial ornithopods (9 specimens); for graviportal ornithopods r = 0.997 and least squares regression line
represents equation {10 ) in text; for cursorial ornithopods r = 0.997 and least squares regression line represents
equation (9 ) in text. Based on data from Gilmore 1915 {Thescelosaurus ) and 1924 {Stegoceras ), Parks 1920
{Kritosaurus ), Osborn 1924 {Protiguanodon, Psitlacosaurus ), Hooley 1925 {Iguanodon ), Lull and Wright 1942
{Anatosaurus, Corythosaurus ). Thulborn 1972 {Fabrosaurus ), Gallon 1974 {HypsUophodon, Dryosaurus,
Parksosaurus ), Gallon and Jensen 1973 {Nanosaurus ), Santa Luca el al. 1974 {Heterodontosaurus ), Dodson
1980 {Camptosaurus, Tenontosaurus ).
478
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 20 Relationship between hindlimb height and metatarsus length in coelurosaurs Logarithmic scale on
both axes- r 0 989 and least squares regression line represents equation (12 ) in text. Based on data from Talbot
1911 (Podokesaurus),Oshom \9\l(Ormtholestes), CoVotxX \9(A(Coelophysis),OiUom I 9 n (Compsognathus).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
479
ESTIMATED HINDLIMB HEIGHT (CM)
FIGURE 21. Comparison of size-frequency curves for (A) Wintonopus track-makers and (B) Skartopus track-
makers at Lark Quarry. Vertical scale is percentage frequency. Insets show examples of similar curves for
populations with different mortality rates and different initial population sizes (adapted from Boucot 1953).
7o DEVIATION FROM MEAN
480
MEMOIRS OF THE QUEENSLAND MUSEUM
STRIDE NUMBER
FIGURE 22. Variation in
stride length for three track-makers at Lark Quarry. A, carnosaur trackway (^.
Tyrannosauropus ) comprising 9 strides. B, ornithopod trackway (Wintonopus ) comprising 17
coelurosaur trackway (Skartopus ) comprising 22 strides. In each case the scale at left indicates percentage
deviation from mean stride length (horizontal line, M).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
481
%
30
20
10
7o
30
20
10
PERCENTAGE DEVIATION OF 2^^^ MEAN SL FROM 1®^ MEAN SL
FIGURE 23. Consistency of stride length for (A) 56 Wintonopus track-makers and (B) 34 Skartopus track-makers
at Lark Quarry. Horizontal scale indicates percentage deviation ( + /-) of mean stride length for second half of
trackway (2nd mean SL) from mean stride length for first half of trackway (1st mean SL). Trackways showing no
deviation are shared equally between +(0-5%) and -(0.5%) classes. Vertical scale is percentage frequency.
482
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 24. A, relationship between estimated mean speed and estimated hindlimb height for Wintonopus track-
makers (solid circles) and Skarlopus track-makers (triangles) at Lark Quarry. For the single Wintonopus track-
maker at New Quarry (star) speed is estimated with equation (6 ) in text. Logarithmic scale on both axes. Lines
defining gaits correspond to size/speed relationships when X/h is 2.0 (walk-trot transition) and 2.9 (trot-run
transition). B, relationship between estimated hindlimb height and estimated maximum speed (based on single
longest stride per trackway). Small symbols indicate that maximum speed is also mean speed (i.e. all strides in
trackway are equal in length, or only a single stride could be measured). Speed for the New Quarry track-maker
(star) is maximum possible estimate, derived (perhaps inappropriately) with equation {II ) in text. Line drawn
through distribution indicates the theoretical regression of speed on hindlimb height when X/h is 3.93.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
483
FIGURE 25. Reconstruction of geographic features and events leading to formation of (he Lark Quarry trackways.
A, outline reconstruction of geographic features at the time the trackways were formed. Sites of Lark Quarry and
Seymour Quarry are superimposed. B, ornithopods and coelurosaurs congregate to drink or to forage in the area
marked by stars (possibly also further to SW). C, carnosaur traverses future site of Lark Quarry from NE to SW;
it turns sharp right to approach the ornithopods and coelurosaurs, which begin to disperse. D, ornithopods and
coelurosaurs take fright and stampede, presumably on account of carnosaur’s subsequent behaviour (unknown);
some may escape via their entry route (?), but at least 150 are driven round the point to the SW and can only
escape by running to the NE — across the future sites of Lark Quarry and Seymour Quarry.
484
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
Wintonopus latomorum ichnogen. et ichnosp. nov.,
and Skartopus australis ichnogen. et ichnosp. nov.
Referred specimens, preserved as natural casts at Seymour Quarry.
All X 1.0.
FIGURES A-B; Wintonopus latomorum; right footprint in posterior
(A) and inferior (B) views. The specimen is a natural cast
detached from overlying sandstone. Distal parts of all three digits
are broken away; so too is the inferior part of digit 3 (which in
Fig. B reveals sandstone filling and ironstone cortex). Adherent
tubular structures are plant rootlets and/ or burrows of
invertebrates. Fine tubercles and wrinkles may represent skin
texture. In Fig. B note concave posterior margin (uppermost).
(QM FI 2264).
FIGURES C-D: Skartopus australis; right footprint in inferior (C)
and anterior (D) views. A natural cast still attached to a small
portion of overlying sandstone. (QM F12265).
486
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
Sediments at Lark Quarry.
FIGURE A; Freshly-broken hand-specimen showing finely laminated
claystone in which dinosaur footprints occur as natural moulds.
Dark-coloured sediment below the level of the scale-bar (marked
in cm) is part of the underlying sandstone.
FIGURE B: Natural section (joint face) through laminated claystone
and the underlying sandstone bed. Scale bar marked in cm.
Colour contrast between claystone and sandstone is slightly
masked by iron-staining (especially at upper left). The three slot-
like cavities (across centre) are pick-marks.
Abbreviations; c, thin ferruginous adhesion from overlying
sandstone: f, dinosaur footprint, still filled with overlying
sandstone; t, tubular structures (possibly escape burrows of
arthropods).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
487
488
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 3
Lark Quarry, viewed from the NW. Carnosaur footprints (cf.
Tyrannosauropus ) are visible in front of kneeling figure at centre.
The site is now roofed for its protection.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
489
490
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 4
Portion of Lark Quarry bedding plane to show abundance, relative
sizes and orientation of dinosaur footprints. All footprints are natural
moulds, and lighting is from the lower left. Area shown is near the W
corner of the quarry (see Fig. 3 in text) and is approximately 2.8 by 3.8
m. The sequence of three large footprints (numbered 6 to 8) is from
the trackway of a carnosaur that travelled to the SW (towards bottom
of page). The numerous small footprints (350 + ) are attributed to
coelurosaurs and small ornithopods, all of which travelled in the
opposite direction.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
491
492
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 5
Carnosaur footprints, cf. Tyrannosauropus.
FIGURE A: Single left footprint preserved as natural mould, x 0.22.
Photographed from fibreglass replica (QM F10322/I), lighting
from N. This is footprint number 3 in the carnosaur trackway (1 1
prints) at Lark Quarry. Note ripples of sandy sediment in the
floor of the print, and surrounding footprints of small dinosaurs.
FIGURE B: Portion of Lark Quarry bedding plane (NW margin)
showing first four footprints in carnosaur trackway.
Photographed obliquely under natural low-angle illumination.
Scale indicated by stride of the carnosaur (3,31 m or
approximately 1 1 feet). The animal moved from NE (top right) to
SW (lower left); note the upwelHng of sediment around each
footprint, and the numerous footprints of small dinosaurs that
moved in the opposite direction.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
493
494
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 6
Carnosaur footprint* cf, Tyrannosauropus.
Single left footprint preserved as natural mould, x 0.25.
Photographed from fibreglass replica (QM F10322/II), with lighting
from NW. This is footprint number 7 in the trackway at Lark Quarry.
In diagrammatic key (below): ART, artefact (a pick-mark); ORN 1,
ornithopod footprint still filled with sandstone; ORN 2, ornithopod
footprint with scrape-marks extending forwards from digits 3 and 4.
All other footprints appear to be those of coelurosaurs.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
495
496
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 7
Wintonopus latomorum ichnogen. et ichnosp. nov.
and Skartopus australis ichnogen. et ichnosp. nov.
FIGURE A: Wintonopus latomorum, holotype (QM F10319). A right
footprint preserved as natural mould, x 1. Lighting from NE.
Attributed to an ornilhopod dinosaur.
FIGURES B and C: Skartopus australis, holotype (QM FI0330). A
right footprint preserved as natural mould, x 1. In Fig. B
lighting is diffuse, from above; in Fig. C lighting is from E.
Attributed to a small theropod dinosaur (coelurosaur).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
497
498
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 8
Wintonopus latomorum ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry,
and all photographed from fibreglass replicas.
FIGURE A: Right footprint, x 0.5. Lighting from E. Showing
pronounced anterolateral scrape-mark from digit 3, and a shorter
scrape-mark from digit 4. No trace of interdigital web between
digits 2 and 3. (QM F10322/II).
FIGURE B: Left footprint, X 0.5. Lighting from NE. Footprint fore-
shortened by toes entering and leaving sediment at steep angle.
Interdigital web is clearly imprinted between digits 3 and 4,
faintly imprinted between digits 2 and 3. Note backwardly-
directed scrape-marks from digits 3 and 4. (QM FI 0322/11).
FIGURE C: Left footprint, x 0.5. Lighting from NE. Digit 2
contains raised ‘cusp’ formed by sediment adhering to underside
of track-maker’s toe. Footprint has poorly defined outline
because it was badly damaged during excavation. (QM
F10322/B).
FIGURE D: Left footprint, X 0,5. Lighting from SE. Anterolateral
scrape-mark produces forked or Y-shaped outline to digit 3.
Interdigital web is clearly imprinted between digits 3 and 4,
absent between digits 2 and 3. (QM F10322/II).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
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500
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 9
Wintonopus latomorum ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry,
and all photographed from fibreglass replicas.
FIGURE A: Left footprint, x 1. Lighting from E. Shallow imprint
with digit 2 exceptionally broad, and digit 4 represented by a
furrow. Probably formed with the track-maker’s body weight
carried mainly on the two inner toes. (QM F10322/B).
FIGURE B: ?Right footprint, x 0.75. Lighting from E. Showing
backwardly-directed scrape-marks from all three digits. (QM
F10322/A).
FIGURE C; Right footprint, x 1. Lighting from NW. From the
smallest trackway referred to W. latomorum. Note the distinct
‘spur’ behind digit 4, and the faint indication of an anterolateral
scrape-mark from digit 3. (QM F10322/B).
FIGURE D: Right footprint, x 0,75. Lighting from N. From the
second-largest trackway referred to fP. latomorum. (QM
F10322/1).
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502
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 10
Wintonopus latomorum ichnogen. et ichnosp. nov.,
and Skartopus australis ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry.
and all photographed from fibreglass replicas.
FIGURE A: Wintonopus latomorum; X^Hiooi^unU x 0.75. Lighting
from NW. With all three digits represented by furrows, and with
distinct trace of interdigital web between digits 3 and 4. There
may also be a very faint imprint of the metapodium. (QM
F10322/A).
FIGURE B: Skartopus australis; ?right footprint, x 0.66. Lighting
from W. Characteristically divergent digits, but one of them (?2)
unusually exaggerated in width. Presumably formed with track-
maker’s body weight carried mainly on the two inner toes. (QM
F10322/B).
FIGURE C: Wintonopus latomorum; two right footprints, x 1.
Lighting from NE. The slightly larger print (below) is fore-
shortened by toes entering the sediment at a very steep angle. As
toes were withdrawn the central one scraped the sediment
forwards — so that it folded over to conceal digit 4 of a smaller
and earlier-formed print. (QM F10322/n).
FIGURED: Wintonopus latomorum; Ml iooipxmU X 0.33. Lighting
from E, Showing exceptionally broad imprint of digit 3, and
forwardly directed scrape-marks from all three digits. The scrape-
mark from digit 3 is extremely long and is deflected slightly as it
runs through the earlier-formed footprint of a coelurosaur
{Skartopus australis). (QM F10322/II).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
503
504
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 11
Wintonopus latomorum ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry.
FIGURE A: Left footprint, x 1. Lighting from NE. Note deeply
imprinted interdigital webs, and distinct curvature of digit 3.
Photographed from rock slab (QM F10320).
FIGURE B: ?Left footprint, x 1. Lighting from N. Extremely fore-
shortened on account of toes entering and leaving sediment at a
very steep angle. Trace of posterior ‘spur’ at left (behind digit ?4)
seems to confirm identification as left footprint. Fibreglass
replica (QM F10322/II).
FIGURE C: ?Left footprint, x 1. Lighting from SE. Three small pits
seem to indicate brief touch-down of toe-tips after they had been
withdrawn from the footprint. Location of these pits indicates
that the foot was shifted forwards and slightly to the right, and
that it was rotated around the axis of digit 3. Fibreglass replica
(QM FI0322/I1).
FIGURE D: Three left footprints (representing three track-makers),
X !. Lighting from W. All three examples show characteristic
‘spur’ behind digit 4. Uppermost example is very fore-shortened,
with shallow imprint of digit 2; imprints of digits 3 and 4 are
amalgamated. Lowermost example (bisected by joint ) shows Y-
shaped tip to digit 3. Fibreglass replica (QM F10322).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
505
506
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 12
Skartopus australis ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry,
and all photographed from fibreglass replicas.
FIGURES A and B: Single right footprint, X 1. In A lighting is from
the NE, in B lighting is from the E. Showing full imprint of the
metapodium. Note .-harply pointed tips of digits (QM F10322/1).
FIGURE C: ?Right footprint, x 1. Lighting from NE. Somewhat
fore-shortened, and with deeply incised scratches formed by
backwards sweep of the track-maker’s foot. (QM F10322/B).
FIGURE D: Right footprint, X 1. Lighting from E. Characteristic
symmetrical arrangement of narrow and sharply pointed digits.
Showing faint imprint of metapodium, traces of interdigital
webs, and scratch-marks behind digits. (QM F10322/II).
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508
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 13
Wintonopus latomorum ichnogen. et ichnosp. nov.,
and Skartopus australis ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry,
and all photographed from fibreglass replicas.
FIGURE A: Skartopus australis; ?left footprint, x 0.66. Lighting
from SW. Showing deeply incised scratch in the floor of each
digit imprint. (QM F10322/B).
FIGURE B: Skartopus australis; ?Ieft footprint, X 0.5. Lighting from
N. Much fore-shortened example with imprint of metapodium.
Fore-shortening accounts for apparently unusual thickness and
bluntness of digit imprints (QM FI0322/B).
FIGURE C; An amalgam of at least four footprints, x 0.66. Lighting
from NW. Two outer digits of a left ornithopod footprint
{Wintonopus latomorum ) are visible at left, with a forwardly
directed scrape-mark from digit 3. Note that all footprints are
similar in depth, in state of preservation, and in orientation. (QM
F10322).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
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510
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 14
Wintonopus latomorum ichnogen. et ichnosp. nov.,
and Skartopus australis ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry,
and all photographed from fibreglass replicas.
FIGURE A: Group of three footprints (representing three animals),
X 0.66. Lighting from SW. At top is a right footprint referred to
Wintonopus latomorum; this shows exaggerated imprint of digit
2, and was presumably formed with track-maker’s body weight
carried mainly on the inner side of the foot. At centre is a
footprint comprising three long scratches — probably from the
left foot of a Wintonopus track-maker. Spacing of the three
scratches would correspond to spacing of three digits in a left
footprint of Wintonopus type. At bottom left is a ?right footprint
referred to Skartopus australis. (QM FI0322/B).
FIGURE B: Portion of Lark Quarry bedding plane, X 0.175.
Lighting from S. Three footprints connected by lines are referred
io. Skartopus australis and form part of a single trackway. AH
three footprints (and two others not shown) have traces of the
metapodium. Despite its ‘flat-footed’ gait this animal seems to
have kept pace with other track-makers at Lark Quarry (see
estimates of speed in Table 5). (QM F10322/B).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
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512
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 15
Wintonopus latomorum ichnogen. et ichnosp. nov.,
and Skartopus australis ichnogen. el ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry.
FIGURE A: Skartopus australis; ?right footprint, x 1. Lighting from
W. Note sharply pointed digits. Fibreglass replica (QM
F10322/II).
FIGURE B: Wintonopus latomorum; left footprint, x 0.5. Lighting
from NE. Showing characteristic proportions and spacing of the
digits; note very slight curvature of digit 3. The outer side of the
track-maker*s foot (digit 4) was much more deeply impressed
than the inner side (digit 2). Photographed from rock slab (QM
F10320).
FIGURE C: A mixture of at least three footprints (representing at
least three animals), X 1. Lighting from N. Apparently one
example of Wintonopus latomorum is superimposed obliquely on
another. The line of three puncture-like marks (top left) is a
smaller and extremely fore-shortened footprint — formed by the
track-maker’s toes entering the sediment vertically.
Photographed from rock slab (QM F10320).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
513
514
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 16
Wintonopus latomorum ichnogen. et ichnosp. nov.,
and Skartopus australis ichnogen. et ichnosp. nov.
Referred specimens, all preserved as natural moulds at Lark Quarry,
and all photographed from fibreglass replicas.
FIGURE A: Skartopus australis; left footprint, X 0.66. Lighting
from SE. All three digits represented by scratches; note
characteristic divergence of the digits (QM F10322/B).
FIGURE B: A group of between 6 and 8 footprints, X 0.5. Lighting
from W. At lower left is an example of Wintonopus latomorum
(somewhat damaged during excavation); at top right is a second
example showing characteristic Y-shaped tip to digit 3. This
second example has largely obliterated an earlier-formed
footprint (to right), and is in turn partly distorted by a later-
formed print (to left). At centre is a little-distorted example of
Skartopus australis. Towards bottom right is a complete
amalgam of at least two unidentifiable footprints (QM
F10322/B).
FIGURE C: Three footprints (representing three animals), x 1.
Lighting from W. A footprint at lower right {?Skartopus ) is
extended into anterior scrape-marks which partly disrupt two
earlier-formed footprints. At lower left is a very characteristic
example of Wintonopus latomorum (left footprint); at upper
right is a smaller example (?right footprint). (QM F10322/A).
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
515
m
516
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 17
Plan of footprints at Lark Quarry. Arrow indicates north.
THULBORN AND WADE: WINTON DINOSAUR TRACKWAYS
517
Mem. QdMus. 21(2): 519—39. [1984]
THE THORNTON PEAK MELOMYS, MELOMYS HADROURUS (RODENTIA :
MURIDAE): A NEW RAINFOREST SPECIES FROM NORTHEASTERN
QUEENSLAND, AUSTRALIA.
J.W. Winter
Queensland National Parks and Wildlife Service
Townsville
ABSTRACT
Melomys hadrourus is described from six specimens collected from Thornton Peak and the
McDowall Range, northeastern Queensland. It is a large species within the size range of
described Melomys such as M. levipes and M. leucogaster . The well developed white-tipped
tail and the robust skull, as demonstrated by the thickness of l‘, suggest affinities with Uromys,
but in general body size it is distinctly smaller than Uromys porculus, the smallest Uromys. M.
hadrourus is considered an upland relict species with no vicariants. All four collection
localities are in rainforest in the upland (> 300 m) zone of the Thornton Peak massif, isolated
from other upland areas of the Daintree and Bloomfield valleys.
INTRODUCTION
Rainforest in Australia occurs as a series of
discontinuous blocks throughout coastal eastern
Australia (Webb and Tracey 1981). Within
tropical Australia the most extensive area of
rainforest is in the Townsville to Cooktown
region. This region is of particular interest to the
zoogeographer because of the relatively high
number of endemic species and of a faunal
affinity with New Guinea (vide Kikkawa,
Monteith and Ingram 1981 for a recent review).
The region is geomorphologically diverse and,
although extensive collections of mammals have
been made from the region (vide Lumholtz 1889,
Cairn and Grant 1890, Tate 1952, Taylor and
Horner 1973), some areas remained unworked by
mammalogists. The upland area of the Thornton
Peak massif, isolated by the Daintree and
Bloomfield valleys (Fig. 1) was one such area.
Thus the opportunity of helicopter transport to
the summit of Thornton Peak, offered by the
Commonwealth Forestry and Timber Bureau
(now CSIRO, Division of Forest Research),
Atherton, was accepted. The circumstances of the
visit and the subsequent capture of a new species
of rodent have been given elsewhere (Winter
1978). This newly discovered species is described
here.
SYSTEMATICS
Melomys hadrourus sp. nov.
Holotype: Queensland Museum JM504 adult
female, skin and torso in spirit, skull extracted,
collected 16 November 1973, by J.W. Winter.
Type Locality; Thornton Peak summit area at
altitude 1220 m, latitude 16°09’30”S, longitude
145°2r45”E (Mossman sheet, 1:100,000 series
R631, grid reference CC250126), northeastern
Queensland, Australia (Fig. 1).
Paratypes: From Thornton Peak type locality
Queensland Museum JM3837 adult female (Watts
and Aslin 1981, PI. 2), skin and torso in spirit,
skull extracted, collected 5 June 1974, by J.W.
Winter; from southern face of Thornton Peak at
altitude 640 m, latitude 16°I0*30”S, longitude
145°2r45”E (Mossman CC250108) Australian
Museum M12520 adult female, skin in spirit,
skull extracted, collected between 3 and 13
November 1975, by H. Posamentier; from
southern face of Thornton Peak at altitude 1020
m: latitude 16°10M5”S, longitude 145°22’00”E
(Mossman CC2521 15), Australian Museum
M12521 subadult male, skin and torso in spirit,
skull extracted, collected 13 November 1975, by
H. Posamentier; from McDowall Range crest,
northeastern Queensland, at altitude 520 m,
latitude 16°06’20’’S, longitude 145°20’00’T
(Mossman CC218190), Queensland Museum
JM2173 subadult male, puppet skin and extracted
skull, with torso in spirit, colour photograph
(Anon. 1977), and subadult male (escaped), both
collected 20 October 1976, by J.W. Winter and
R.G. Atherton.
Diagnosis:
A large Melomys with head-body length to 180
mm, condylobasal length to 42.7 mm
520
MEMOIRS OF THE QUEENSLAND MUSEUM
O ^
145 10
o f
145 20
145°30 ,
15 50
16 ° 10 '
16 20
FIGURE 1 . Map of the Thornton Peak and adjacent uplands above 300 m (light stippling) in altitude, showing type
locality (circled star) and other capture localities (solid stars) of Melomys hadrourus, cleared land (heavy
stippling), and national park (cross hatching).
WINTER: THORNTON PEAK MELOMYS
521
(occipitonasal length to 45.3 mm); tail longer than
the head-body length, relatively thick (> 4.7 mm
diameter at the base), apical quarter white, scales
non-abutting and subtending one hair per scale.
Distinguished from Uromys by smaller size
(condylobasal length < 43 mm, occipitonasal
length < 46 mm) and from juvenile U.
caudimaculatus by shorter pes length (< 39 mm).
Distinguished from other large Melomys by the
thicker tail; from M. leucogaster group by the
longer rostrum (nasal length: condylobasal length
< 1: 2.45); from M. levipes group by greater
thickness of V (> 2.3 mm) and tail longer than
the head-body length. Distinguished from other
Australian Melomys by greater size (head-body
length > 165 mm, pes s.u. > 35 mm,
occipitonasal length > 44 mm) and by the thicker
tail with only one hair per scale.
Description:
Pelage generally soft and guard hairs not
prominent, colour light fawn becoming lighter
ventrally, no prominent markings other than a
white patch on the throat and sternum. A detailed
description of the holotype follows (colour names
after Ridgway 1912): dorsally median fur is 15-18
mm long in lumbar region to c. 14 mm between
ears and 4-5 mm on nasals, basal two thirds of
fur Pale Mouse Gray with apical third Cinnamon-
buff on back, Ochraceous Orange between ears
and Tawny to Light Buff on nasals giving it a
slightly grizzled appearance; terete guard hairs
20-24 mm long on back and c. 15 mm on nasals
with basal one third Pale Mouse Gray and apical
two thirds Tawny tending to a colourless tip of up
to 2 mm; laterally mid-body fur grades to
Ochraceous Buff on apical third and Pale Mouse
Gray on basal two thirds, on side of face below
eye it is Warm Buff for apical third and Pale
Mouse Gray for basal two thirds, and in region of
mysticeal vibrissae fur Light Buff apically with
basal two thirds White; guard hairs greatly
reduced in number, but otherwise as dorsally,
ventrally fur 8-9 mm long on abdomen with basal
half Pallid Mouse Gray median quarter Fawn
Colour and apical quarter Light Buff, from mid-
thoracic region to chin fur is White throughout its
length, width of white patch 10-15 mm with
constriction at level of fore-legs.
Ears: Prominent and rounded, skin Fuscous,
sparsely covered with Tawny to colourless hairs c.
1 mm long.
Vibrissae: Approximately 30 mysticeal vibrissae
on each side and up to 73 mm long, from Mars
Brown basally through Tawny medially to
colourless apically in varying proportions; three
supraorbital vibrissae on right side (none on left)
up to 15 mm long, colour as for mysticeal
vibrissae; one postorbital on each side 21-23 mm
long, colour as for mysticeal vibrissae;
submentals numerous and up to 6 mm long,
colourless; ulnar carpals 4 each side and up to 12
mm long, colourless.
Manus: Skin Cream-Buff in preserved specimen
and sparsely covered dorsally with colourless
hairs c. 2 mm long, except for mid-dorsal line in
which median section of hair is partially Tawny;
for foot pads see PI. 1.
Pes; Skin Cream-Buff in preserved specimen and
sparsely covered dorsally with colourless hairs c. 2
mm long, except for line of hairs outward of mid-
dorsal line in which median section of hair
partially Tawny; for foot pads see PI. 1.
Tail: Longer than head-body (Table 1), diameter
at base c. 5 mm, scales round and reduced (not
abutting) with slight sculpturing and subtending
one hair each (PL 1), length of scale hair 1.5
scales on basal portion and less than 0.5 on apical
portion; colour of scales Tawny dorsally for basal
three quarters, slightly paler ventrally especially
at base, apical quarter pure white dorsally and
ventrally (PI. 1), hairs Tawny to colourless. The
general impression is of a large thick tail more
reminiscent of that of a juvenile Uromys
caudimaculatus caudimaculatus than of a typical
Melomys. {vide PI. 3).
Skull: Characteristic of genus but generally larger
and more robust, particularly incisors (Table 2,
PI. 2); crown pattern of check teeth characteristic
of genus (PL 2), alveolar pattern M', M\ M^:
4,4,3 (= pattern D of Knox 1976), anterior face
of incisors orange.
Mammary formula 0-2 = 4, vagina perforate,
teats small and not lactating.
Variation of Paratypes:
Pelage of paratypes similar to that of holotype
including ventrally a patch of fur, white to base,
from chin to the mid-thoracic region; tail of
JM2173 and M12521 with white tip of c. 28 mm
(25 + c. 3 mm which withered away — vide
photograph of this specimen in Anon. 1977) and
52 mm respectively; JM3837’s tail (complete on
capture) and that of individual in PI. 3 also had
extensive white tips; apical third of M12520’s tail
missing on capture; measurements of external
features and skulls given in Tables 1 and 2;
mammary formula of JM3837 0-2 = 4, not known
for M 12520.
A subadult male captured at the McDowall
Range locality, and which subsequently escaped,
is illustrated in PI. 3.
522
MEMOIRS OF THE QUEENSLAND MUSEUM
Etymology:
The specific name is derived from the Greek,
hadros (well-developed, bulky, stout, large,
strong and great) and oura (tail), and refers to the
well-developed tail of M. hadrourus, which is its
most characteristic external feature.
TAXONOMIC POSITION
Comparative Material Examined:
Melomys levipes shawmayeri Riimmler 1935,
type, British Museum (Natural History), London
(BM) No. 35.12.20.2 (field No. S.M. 368 in Tate
1951), specimen and photographs; Melomys
levipes lanosus Thomas 1922, type, BM 22.2.2.26
(not 22.2.22.26 as given in Tate 1951), specimen
and photographs; Melomys levipes levipes
(Thomas 1897), cotype, BM 97.8.7.72, specimen
and photographs; Melomys levipes rattoides
Thomas 1922, type BM 22.2.2.25, specimen and
photographs; Melomys levipes naso (Thomas
1911), type, BM 11.11.11.54, specimen and
photographs; Uromys sapientis Thomas 1902,
type BM 2. 5. 1.4, specimen and photographs;
Uromys porculus Thomas 1904, type BM
89.4.3.8. Uromys caudimaculatus aruensis Gray
1873, Museo Civico di Storia Naturale, Genoa
(MSNG) No. 3605a and type. No. 3248 (skull
missing), specimens only; Melomys levipes levipes
(Thomas 1897), cotype, MSNG 3600a, specimen
only; Melomys leucogaster leucogaster (Jentink
1909), American Museum of Natural History
(AMNH) No. 105723, specimen and
photographs; Melomys leucogaster latipes Tate
and Archbold 1935, type AMNH 104273,
photograph only. Melomys levipes lorentzii
(Jentink 1909), type, Rijksmuseum van
Natuurlijke Historic, Leiden (RNHL) No. 25494
(Field No. 132), photograph only; Melomys
leucogaster leucogaster (Jentink 1909), type,
RNHL 25493 (Field No. 119), photographs only.
Specimens of Uromys caudimaculatus (Krefft
1867), Melomys cervinipes (Gould 1852),
Melomys capensis Tate 1952, and Melomys
burtoni (Ramsay 1887) were on hand from field
collections in northeastern Queensland by the
author, and Melomys rubicola Thomas 1924
from field collections on Bramble Cay by C.J.
Limpus (pers. comm,).
Measurements used for comparative purposes
in Figs. 2 and 4 were those given in Tate (1951)
unless otherwise stated.
Generic Position:
Melomys hadrourus belongs to the mosaic-
tailed rats lacking a distinct prehensile tail, within
the Uromys group of genera of Tate (1951). This
group consists of relatively small rats within the
genus Melomys Thomas 1922 and much larger
rats of the genera Uromys Peters 1867 and
Solomys Thomas 1922. Tate (1951) included
Solomys as one of several subgenera of Uromys,
but Laurie and Hill (1954) accorded it full generic
rank (type species Uromys sapientis Thomas
1922). M. hadrourus lies at the top of the size
range of the described Melomys as indicated by
its skull size (Fig. 2). The skull is generally more
heavily built than in other Melomys, although M.
levipes naso is similar. The depth of I', is
generally greater than in other Melomys the only
overlap being with M. 1. leucogaster (AMNH No.
105723) (Fig. 3). The tail is distinctly thicker than
in all other Melomys examined and is similar to
that of juvenile Uromys caudimaculatus
caudimaculatus and the juvenile male Uromys
caudimaculatus aruensis Gray 1873 (type, MSNG
No. 3248). Nevertheless M. hadrourus is
distinctly smaller than Uromys porculus Thomas
1904 (the smallest Uromys ) as is shown by skull
size (Fig. 2) and I' depth (Fig. 3). {Uromys
porculus Thomas 1904, is still retained in the
genus Melomys by Laurie and Hill (1954), but 1
agree with Tate’s (1951) decision to include it
within Uromys, My agreement with Tate is based
on skull size of porculus which fits into the
Uromys group rather than the Melomys group
(Fig. 2)). Although Solomys has relatively
inflated bullae as in Melomys, its larger body size
and V-shaped rear margin to the palate (Laurie
and Hill 1954) distinguish it from Melomys.
Therefore, on the basis of general body size
which is within the range described for Melomys,
I have placed M. hadrourus in that genus. Other
features such as the w'ell-developed tail and thick
r, indicate affinities with Uromys; the
configuration of the tail in particular is aberrant
for Melomys. These features may well become
significant in determining the generic status of M.
hadrourus should the two genera at some future
time be distinguished on anything other than size.
Specific Position;
Melomys hadrourus is similar in size to the
larger New Guinean forms within the M. levipes
and M. leucogaster groups (Fig. 2). It differs
from the members of the M. leucogaster group
by having a distinctly longer rostrum, as shown
by the ratio of the nasal length to condylobasal
length of the skull (Fig. 4), and by the skull longer
relative to its breath (Fig. 2).
Five of the subspecies of M. levipes recognised
by Tate (1951) are close to M. hadrourus in size.
WINTER: THORNTON PEAK MELOMYS
523
viz M. 1. rattoides, M. I. naso, M. 1. lanosus, M. 1.
shawmayeri, and M. /. lorentzH (M. 1. levipes
with a condylobasal length of 37.0 mm and
zygomatic width of 19.4 mm, cotype, BM
97.8.7.72 is not one of the larger members of the
M. levipes group (Fig. 2)). All five subspecies
have slender tails typical of the genus Melomys,
and all have tails shorter than the head-body
length, in contrast to the thick tail of M.
hadrourus which is longer than the head-body
length. The depth of V in M. hadrourus is
significantly greater than in these five large M.
levipes subspecies (Fig. 3). In the photograph of
the skull of M. /. lorentzH (RNHL 25494) the
posterior margin of the palate is obscured by
remnants of the soft palate, but in the other four
subspecies the margin lies forward of the
posterior end of the molar row, whereas in M.
hadrourus it lies well behind (PL 2).
Melomys hadrourus has one hair per tail scale
as do Melomys leucogaster M. levipes naso, and
M. levipes levipes, but differs from these
respectively by having a longer rostrum, a longer
thicker tail, and a larger body size. Melomys
levipes rattoides, M. L shawmayeri, M. 1. lanosus
all have three hairs per tail scale, as does M. L
lorentzH except for one specimen (Tate 1951).
Melomys levipes lorentzH has a mammary
formula of 0- 1 = 2 (Zeigler 1 972) in contrast to the
0-2 = 4 which is typical for the genus and M.
hadrourus.
Melomys hadrourus differs from other
Australian Melomys, which are placed into the
M. cervinipes (including capensis and rubicola )
and M. lutillus (= burtoni vide Knox 1978)
groups by Tate (1951), by being distinctly larger
(Fig. 2) and by having only one hair per tail scale
in contrast to the three in the other two groups.
Baverstock, Watts and Hogarth (1977)
examined the chromosomes of the paratype
(JM3837) (their specimen no. IMVS 181F) of
Melomys hadrourus. The karyotype had a
diploid number of 48, which is the standard
number for the Australian species they examined
in the M. cervinipes and M. burtoni complexes.
The karyotype differed from M. ?littoraiis ( =
burtoni vide Knox 1978) by two fixed
rearrangements (pairs 2 and 4) and from M.
cervinipes by three fixed rearrangements (pairs 1,
2 and 4) (Baverstock et aL 1977). From their
chromosomal work on the Australian Melomys,
Baverstock, Watts, Adams and Gelder (1980)
concluded that three karyotypic forms occurred
in Australia; M. burtoni, M. cervinipes
(including capensis ), and the Thornton Peak
melomys (M. hadrourus ).
The alveolar pattern (type D) of Melomys
hadrourus is the same as that of M. cervinipes
(including rubicola ) and M. rufescens (Alston
1877) but differs from that of the M. burtoni
group (Knox 1976).
It is concluded, therefore, that Melomys
hadrourus is a valid species, quite distinct from
other described Melomys.
HABITAT AND DISTRIBUTION
All specimens of Melomys hadrourus were
caught in rainforest on the Mareeba Granite of
the Thornton Peak massif. The type locality,
where JM504 and JM3837 were caught within 100
m of each other, was within 200 m of the head of
a gully, with numerous boulders, on the western
face of Thornton Peak, and which was one of the
northernmost tributaries of Hilda Creek. The
gully originated at a fern-covered saddle
(Glyeichenia sp.) at the northwestern end of the
summit valley. The vegetation (PL 4) was simple
microphyll vine-fern thicket (Tracey 1982) with a
canopy height of 6-12 m. Thin wiry lianes and
tree ferns were common, with Lacospadix palms
abundant in the understorey. Ground cover was
sparse leaf litter with scattered ferns and tree
seedings between boulders and fallen logs. Mosses
and lichens were abundant from the ground layer
through to the canopy. The summit area of
Thornton Peak is wet with a rainfall likely to be
similar to that of the summit of Bellenden Ker,
130 km to the southeast, which has an annual
average of 8529 mm recorded over a six year
period (Tracey 1982). Even in the dry season
months of April to October the summit is
enveloped in cloud much of the time.
JM2173 and a subadult male, which
subsequently escaped, were caught on the crest of
the McDow'all Range within 100 m of the road in
simple notophyll vine forest (Tracey 1982).
Canopy height was 20-25 m and tree diameters
mainly in the 40-50 cm range with a few up to 70
cm. There was a straddled understorey and a
scattered shrub layer which consisted mainly of
Calamus clumps without the climbing tendrils.
Lacospadix palms and tree ferns were scarce,
woody lianes moderately common, but wiry
lianes absent. Ground cover was sparse and leaf
litter and bare soil with sparse tree seedlings and
very sparse ferns. This site is below the cloud line
that generally envelopes the summit of Thornton
Peak, and mosses and lichens were relatively
sparse. Rainfall is less than for the summit region,
and the locality comes between the 2500 and 3750
mm isohyets (Tracey 1982). Ml 2520 was caught
in mesophyll vine forest and Ml 2521 in simple
524
MEMOIRS OF THE QUEENSLAND MUSEUM
mesophyll vine forest — simple notophyll vine
forest at sites 39 and 40 respectively (Broadbent
and Clark 1976) on a steep southerly ridge
immediately to the west of the main branch of
Hilda Creek.
JM504 and JM3837 were captured in
aluminium Elliott traps (33 x 9 X 9 cm) (Pi. 4)
set on the ground and baited with sweet-potato in
linseed oil. Both McDowall Range animals were
caught in cage traps (one aluminium and one wire
with slightly larger dimensions than for the
above) set on the ground and baited with rolled
oats plus aniseed oil and sweet-potato in linseed
oil respectively. Both the Australian Museum
animals were caught in snap traps (one a
Conibear) set on the ground; the bait was either
peanut butter compound or aniseed oil
(Broadbent and Clark 1976). Table 5 in
Broadbent and Clark (1976) lists 5 'Melomys
**levipes'* group' as being captured at site 39.
There was some confusion over the identity of M
hadrourus with M. cervinipes at the time. In fact
only one specimen of M. hadrourus (Ml 2520)
was kept from this site and one adult male,
probably attributable to M. hadrourus, was
discarded because of damage received on capture.
All four localities at which M. hadrourus has
been captured are in the upland (> 300 m) zone
of' the Thornton Peak massif. The area of this
upland zone (measured as a flat surface from the
1:100 000 vegetation map, Tracey and Webb
1975) is approximately 24,780 ha and 96 per cent
is rainforest. The Thornton Peak upland zone is
isolated from other upland zones to the southwest
and northwest by the Daintree and Bloomfield
valleys respectively, and the latter, with open
forest vegetation, also acts as an ecological
barrier (Fig. 1).
Thornton Peak itself (altitude 1374 m) and the
eastern fall of the massif are national park (Fig.
1). Approximately 5620 ha (22.7<7o) of the upland
zone is within the national park, the remainder is
within timber reserve. The relative isolation and
rugged terrain of the area have, to date, protected
the upland zone from major forestry, mining and
agricultural developments. The clearing of
rainforest that has taken place in the area has
been restricted to the lowlands (Fig. 1).
HABITS
All specimens of M. hadrourus were caught in
traps set on the ground, but like M. cervinipes
and Uromys caudimaculatus it is probably
scansorial. The stomach of Ml 2521 was filled
with a creamy coloured endospern of a nut. Adult
female JM3837 was captured 5 June 1974 and
kept alive in captivity until 2 February 1975. She
did not give birth to young. Adult female JM504,
captured 16 November 1973, had a perforate
vagina and small teats that were not lactating
(contents of uteri unknown). The breeding
condition of adult female Ml 2520 was not
recorded. The three males (JM2173, M12521 and
the escaped individual) were all subadults with
testes abdominal and captured in October and
November. Subadult male, JM2173, when
handled, gave squeaks similar to that of M.
cervinipes but deeper in pitch, and not a Uromys
-like growl.
At the type locality M. hadrourus was recorded
as living sympatrically with three other rodents;
M. cervinipes (Gould 1852), Rattus fuscipes
(Waterhouse 1839) and Rattus leucopus (Gray
1867). At the McDowall Range locality, in
addition to these three species, Uromys
caudimaculatus was also recorded.
DISCUSSION
Melomys hadrourus represents a third
phylogenetic line of Melomys in Australia with
clear differences from the M. burtoni and M.
cervinipes /capensis species groups both in
karyotype and in morphology (larger size, well-
developed tail, on hair/tail scale). On size alone
its closest affinities would seem to be the New
Guinean species groups, M. leucogaster and M.
levipes. At one stage it was thought that M.
hadrourus was closely allied to M. levipes
(Baverstock et al. 1977) and therefore a vicariant
with New Guinean affinity (Kikkawa et al. 1981).
Melomys hadrourus, however, has certain
affinities with Uromys, namely the general
configuration of the tail and the more robust
skull, particularly in the thickness of T. It also
has other clear differences from M. leucogaster
(longer rostrum) and M. levipes (tail longer than
head-body length, one hair per tail scale in
contrast to the three in four of the large
subspecies, palate margin posterior to tooth row).
It is suggested here that M. hadrourus,
although included with the genus Melomys on
size, is an aberrant form for the genus. Whether
these differences warrant removing M. hadrourus
from Melomys as a third genus, or alternatively
using it to synonymise Melomys and Uromys,
requires an extensive review of the mosaic-tailed
rats, beyond the scope of this paper. For the
present, it is judged to be sufficiently different
from other described species for it to be treated as
a species without vicariants.
WINTER: THORNTON PEAK MELOMYS
525
This being the case, M. hadrourus belongs to
the group of species, endemic to the Townsville to
Cooktown region, and considered to be relicts of
a wet- and cool-adapted fauna which may have
originated in Australia from a common pre-
Pleistocene stock of Australia and. New Guinea;
mammalian representatives of this group are
Antechinus godmani (Thomas 1923),
Pseudocheirus herbertensis (Collett 1884),
Pseudocheinis lemuroides (Collett 1884) and
Hypsiprymnodon moschatus Ramsay 1876
(Kikkawa et al. 1981). Their survival has been
dependent on the continuous existence of
rainforest refugia, of which the uplands of the
Thornton Peak massif is one example (Webb and
Tracey 1981).
ACKNOWLEDGMENTS
I wish to thank the Commonwealth Forestry
Timber Bureau (now CSIRO Division of Forest
Research), Atherton, for making available the
helicopter transport which enabled me to make
the first visit to the summit of Thornton Peak,
and to those people who subsequently helped in
the search for further specimens of the Thornton
Peak melomys — Rob Atherton. Louise
Atherton, Cherie Daniel, Curly Matthew, Dan
Norris, Ian Shield, Mark Weaver, David Winter
and Margaret Winter. I also wish to thank Mr H.
Posamentier, the Australian Museum, for making
available to me the specimens of M. hadrourus
that he captured, and Dr P.D. Dwyer, University
of Queensland, for the skull of Uromys anak. Ms
E. J. Knox and Mr J.A. Mahoney very kindly read
early drafts of the manuscript and made many
useful suggestions for its improvement.
Photographs in PI. 1 and 2 were supplied by the
Department of Primary Industries, Photographic
Department, and Hans and Judy Beste took the
photographs used in PL 3. Finally I wish to thank
the staff of the British Museum (Natural
History), London, and Museo Civico di Sloria
Naturale, Genoa, the Rijksmuseum van
Natuurlijke Historic, Leiden, the American
Museum of Natural History, the Australian
Museum, and the Queensland Museum for their
help in examining their collections and in
supplying photographs.
LITERATURE CITED
Anon., 1977. Eye on the world. Wildlife, Lond.
19: 391.
Baverstock, P.R., C.H.S. Watts, M. Adams
and M. Gelder, 1980. Chromosomal and
electrophoretic studies of Australian
Melomys (Rodentia : Muridae). Aust, J.
ZooL 28: 553-74.
C.H.S. Watts and J.T. Hogarth, 1977.
Chromosome evolution in Australian
rodents. I. The Pseudomyinae, the
Hydromyinae and the Uromys /Melomys
group. Chromosoma, Berl. 61: 95-125.
Broadbent, j. and S. Clark, 1976. A faunal
survey of East Australian rainforests. Interim
Report. (Australian Museum: Sydney).
Cairn, E.J. and R. Grant, 1890. Report of a
collecting trip to north-eastern Queensland
during April to September, 1889. Rec. Aust.
Mus. 1: 27-31.
Corbet, G., 1964. Preservation and
measurements of mammals, pp. 116-28 in
‘The handbook of British mammals’. H.N.
Southern (ed.) (Blackwell Scientific
Publications: Oxford).
Kikkawa, J., G.B. Monteith and G. Ingram,
1981. Cape York Peninsula: Major region of
faunal interchange, pp. 1695-1742, in A.
Keast, (ed.), ‘Ecological biogeography of
Australia’. Vol. 3 (Dr W. Junk bv Publishers:
The Hague).
Knox, E., 1976. Upper molar alveolar patterns of
some Muridae in Queensland and Papua New
Guinea. Mem. Qd Mus. 17: 457-9.
1978. A note on the identification of Melomys
species (Rodentia : Muridae) in Australia. J.
ZooL Lond. 185: 276-7.
Laurie, E.M.O. and J.E. Hill, 1954. ‘List of
land mammals of New Guinea, Celebes and
adjacent islands 1758-1952’. (British
Museum (Natural History): London).
Lumholtz, C., 1889. ‘Among cannibals, an
account of four years’ travels in Australia
and of camp life with the aborigines of
Queensland’. (John Murray: London).
Mahoney, J.A., 1972. The identity of Hapalotis
personata Krefft, 1867 (Muridae, Rattus )
from Cape York, Queensland. Aust.
Mammal. 1: 14-9.
Ridgway, R., 1912. ‘Color standards and color
nomenclature’. (Publ. by author:
Washington).
Tate, G.H.H., 1951. Results of the Archbold
Expeditions. No. 65. The rodents of
Australia and New Guinea. Bull. Amer. Mus.
Nat. Hist. 97: 183-430.
1952. Results of the Archbold Expedition. No.
66. Mammals of Cape York Peninsula, with
notes of the occurrence of rainforest in
Queensland. Bull. Amer. Mus. Nat. Hist. 98:
563-616.
526
MEMOIRS OF THE QUEENSLAND MUSEUM
Taylor, J.M. and B.E. Horner, 1973. Results
of the Archbold Expeditions. No, 98.
Systematics of native Australian Rattus
(Rodentia, Muridae). Bull. Amer. Mus. Nat.
Hist 150 : 1-130.
Tracey, J.G., 1982. ‘The vegetation of the
humid tropical region of North Queensland’.
(CSIRO: Melbourne).
and L.J. Webb, 1975. ‘Vegetation of the humid
tropical region of North Queensland’. (15
maps at 1:100 000 scale and key). (CSIRO:
Indooroopilly).
Watts, C.H.S., and H.J. Aslin, 1981. ‘The
rodents of Australia’. (Angus and Robertson:
Sydney).
Webb, L.J. and J.G. Tracey, 1981. Australian
rainforests: patterns and change, pp. 605-94,
in A. Keast (ed.), ‘Ecological biogeography
of Australia’. Volume 1. (Dr W. Junk bv
Publishers: The Hague).
Winter, J.W., 1978. The rainforest, pp. 113-46,
in H.J. Lavery (ed.), ‘Exploration North’.
(Richmond Hill Press: Richmond).
Zeigler, A.C., 1972. ‘Guide to native land
mammals of northeast New Guinea.
(Preliminary version for use of Wau Ecology
Institute)’. Mimeographed, 28 pp.
Department of Entomology, Bernice P.
Bishop Museum, Honolulu, Hawaii.
TABLE 1 : External Body Measurements. Weights and Hairs per Tail Scale of Melomys hadeourus.
♦Measurements taken by H. Posamentier.
Holotype
Museum no.
JM504
JM3837
Status
Adult 2
Adult 2
Weight (g)
149
164
Head-body length (mm)
180
177
Tail-vent length (mm)
196
-
Pes s.u. (mm)
38
37
Ear (to notch) (mm)
24
23
Tail diameter at base (mm)
5.1
5.2
Tail scale rows/cm
12.5
13
Hairs/tail scale
1
1
Paratypes
Escaped
JM2173 M12520*
S-Adult S Adult 2
M12521*
S-Adult (3 S-Adult S
125
162
149
152
174
171
173
184
184
-
188
197
38
38
38
37
25
24.5
23.5
23
4.7
5.6
5.1
-
11
12
13.5
-
1
1
1
-
WINTER: THORNTON PEAK MELOMYS
527
TABLE 2: Skull Measurements (in mm) of Melomys hadrourus. Definition of Measurements as given by,
— Taylor and Horner (1973),^ — Mahoney (I 972 ). or^- Corbet ( 1964 ). 1' Thickness Measured
Parallel to Horizontal. Plane from Posterior Point of Emergence from Premaxilla.
Holotype Paraiypes
JM504
JM3837
M12520
M12521
JM2173
'Occipilonasal length
45.3
44.2
44.6
44.2
44.7
'Condylobasal length
42.2
42.7
41.2
41.2
42.0
'Basal length
39.5
40.1
38.8
38.4
39.0
'Zygomatic width
22.6
22.5
21.5
21.4
22.2
'Interorbital width
7.1
6.8
6.9
7.0
7.2
^Width of rostrum
7.1
6.7
6.7
6.8
7.0
^Nasal length
17.8
17.4
17.4
17.2
17.7
^Maximum width across paired
nasals
5.0
5.4
4.9
5.1
5.0
^Maximum width across paired
parietals
16.6
15.8
16.5
16.9
16.0
'Mastoid width
16.0
16.1
15.7
15.8
16.2
interparietal length
6.1
5.7
6.0
6.6
5.1
interparietal width
11.6
11.2
11.6
11.2
11.8
Zygomatic plate minimum width
5.2
5.8
5.4
5.2
5.6
‘Palatal length
25.2
25.2
24.9
24.7
24.9
^Diastema length
13.2
13.6
13.9
13.6
13.2
'Anterior palatal foramen length
6.6
6.5
6.0
5.8
6.4
'Anterior palatal foramina width
2.5
2.8
2.5
2.6
2.7
^Palate width between
anterointernal roots of M'
4.9
4.8
4.8
4.2
4.7
^Palate width between anterior
roots of M^
5.3
5.2
5.4
5.2
5.7
‘Bulla length
4.9
4.9
5.0
5.1
5.0
'M'"^ length (crowns)
7.3
7.6
7.3
7.2
7.1
'M'"^ length (alveoli)
8.1
8.2
8.3
7.8
8.0
M' length x width (crowns)
3.4x2. 3
3.8x2. 1
3. 6x2.2
3.4x2.3
3.3x2.3
M^ length x width (crowns)
2.7x2.1
2.5x2.0
2 . 8 x 2.2
2 . 8 x 2.1
2.6x2.3
M^ length x width (crowns)
1 . 6 xl .6
1 . 6 x 1 . 5
1 . 7x1.6
1.7xl.6
1.5X1.6
1 ' thickness
2.5
2.5
2.3
2.3
2.4
^Length of mandibular ramus from
tip of incisor
31.1
32.3
30.5
30.0
30.8
^Height of condyle above ventral
surface of mandibular ramus
11.5
11.6
12.0
11.5
12.0
M |.3 length (crowns)
7.5
7.4
7.6
7.8
7.1
M ,.3 length (alveoli)
7.8
8.3
7.7
8.1
7.9
Mj length x width (crowns)
3.2x2. 2
3.3xl.9
3.1x2.!
3. 1x2.0
3.1x2.!
M 2 length X width (crowns)
2.5x2. 1
2.2xl.7
2 . 6 x 2. 1
2.7x2. 1
2.5x2. 1
M 3 length X width (crowns)
1.9xl.7
1.9x1. 4
1 . 8 x 1.6
1 . 8 x 1.6
2 . 0 xl .6
528
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 2. The relationship between condylobasal length and zygomatic width in Vromys, Melomys hadrourus
and other Melomys species groups.
Condylobasal length mm
Incisor thickness mm
WINTER: THORNTON PEAK MELOMYS
529
FIGURE 3. The relationship between condylobasal length and incisor' thickness in Melomys hadrourus {•), M.
leucogaster (A), M. levipes (□), and Uromys spp. (o). T measurements: M. leucogasier latipes AMNH 104273
(1.7’), M. leucogaster RNHL 25493 (2,1 ) M. /. leucogaster AMNH 105723 (2.3), M. levipes lanosus BM
22.2.2.26 (1.7*), M. levipes shawmayeri BM 35.12.20.2 (1*7*), M. /. levipes BM 97.8.7.72 (1.75*), A/. /. levipes
rattoides BM 22.2.2.25 (1.8*), M. levipes naso BM 1 1 .11.11.54 (1.85*), A/, levipes lorentzii RNHL 25494 (1.9 ‘),
Uromys caudimaculatus caudimaculatus juv. QM JM3839 (2.6), C/.c. caudimaculatus adult QM No. JM3840
(4.6), U. porculus BM 89.4.3.8 (2.65*), U. sapientis BM 2.5.1.4 (3.0*), U. anak QM JM3838 (4.3).
Measurements supplied by P.D. Jenkins, BM (*); C. Smeenk, RNHL ( + ); C. Smeenk, RHNL ( + ); M.A.
Lawrence, AMNH (’).
530
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 4. The relationship between nasal length and condylobasal length expressed as a ratio, in Melomys
hadrourus (o), M. levipes (□), and M. leucogaster (A). The upper and lower limits of the ratio are shown.
Condylobasal length mm
532
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
External features of Melomys hadrourus (QM JM504, holotype)
from Thornton Peak, N.E. Queensland. A; pes, B; manus, C: tail, D:
tail detail about one third from the base.
WINTER: THORNTON PEAK MELOMYS
533
534
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
Skull features of Melomys hadrourus (QM JM504, holotype) from
Thornton Peak, N.E. Queensland. A-E; cranium and mandible, F:
upper left molars, G: lower left molars.
WINTER: THORNTON PEAK MELOMYS
535
536
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 3
Subadult male of Melomys hadrourus from the McDowall Range,
N.E. Queensland. The animal subsequently escaped.
WINTER: THORNTON PEAK MELOMYS
537
538
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 4
Type locality of Melomys hadrourus. Simple microphyll vine-fern
thicket, Thornton Peak, N.E. Queensland.
WINTER: THORNTON PEAK MELOMYS
539
Mem. QdMus. 21(2): 541—59. [1984]
THE MT. INGLIS CACHE: A NEW PERSPECTIVE ON ABORIGINAL MATERIAL
CULTURE IN THE CENTRAL HIGHLANDS OF QUEENSLAND.
M.J. Morwood
University of New England
Armidale, N.S.W.
ABSTRACT
This paper describes a cache of Aboriginal material recovered from ‘Mt. Inglis’ Station in
the Central Highlands of Queensland. The cache includes bones, skins and lithic materials.
Rockshelter caches, particularly of organic items provide evidence for a range of Aboriginal
material culture, activities and practices which were never ethnographically described. The Mt.
Inglis cache of ceremonial and decorative items was probably hidden for later re-use during the
final phase of traditional Aboriginal culture in the region.
INTRODUCTION
In 1901 Archibald Meston, then Protector of
Aborigines for southern Queensland, visited the
upper Maranoa River in the Central Queensland
Highlands ‘to obtain some ethnological
specimens said to exist in sandstone caves’
(Meston 1901). Meston well appreciated the
importance of these finds, and a mere 40 years
after the first European occupation of the area,
he states —
In the caves and rockshelters of our
mountain ranges there are still hundreds of
specimens specially valuable to ethnology,
and the value is incalculable when we regard
them as among the last available memorials
of a primitive race rapidly vanishing from the
face of the earth (Meston 1901).
This was the first published account of a cache
of Aboriginal material culture from shelters in the
Queensland Central Highlands, and remains one
of the few.
In 1975, such an Aboriginal cache was
discovered on “Mt. Inglis” Station, northeast of
Carnarvon Gorge (latitude 24°46”S, longitude
148®18”E). The find became known to local
residents, but fortunately much of the material
was left in situ. The Archaeology Branch,
D.A.I.A. was notified of the find, and in
accordance with the Aboriginal Relics Protection
Act of 1967-76, it was decided to recover the
cache to prevent its unauthorised removal. In
January 1976, I removed the cache, assisted by
Jeff Pratt (then Aboriginal Relics Ranger for
Central Queensland).
DESCRIPTION
The material was located in a small shelter on
the southern slope of a rocky hillside, overlooking
a black-soil flat (PI. 1). The surrounding
vegetation comprised an open woodland of
ironbark ( Eucalyptus melanophloia ), round-
leafed box (£'. populnea ), bloodwood (E.
terminalis ), and yellowjack, with an Acacia
understory. Water was available at a permanent
spring some 300 metres to the north.
The shelter faced north and measured 6 by 4.5
metres with a maximum dripline height of 1.5
metres. The cache was positioned on a small ledge
in the roof, 50 cm back from the entrance and
facing east. The ledge measured 70 by 30 cm and
was 92 cm above the shelter floor. The cave was
an obvious one, but to a casual observer it was
‘obviously empty* as the shelf was not visible
from the entrance.
Originally the cache had been concealed with
three sandstone blocks (25 cm maximum
dimension) placed at the front of the ledge. These
had been shifted and at the time of the removal
sections of marsupial skin and twine were hanging
down from the ledge. The material had already
been removed several limes for inspection after its
initial discovery, and had suffered some damage
because of this. On the shelter floor beneath the
ledge, lay a deposit of red ochre, necklace reels,
lengths of twine, skin and feathers. As part of the
recovery procedure, this area of the floor deposit
was sifted through to a depth of 3 cm. The poor
condition of the organic items recovered from the
shelter floor would indicate that deterioration
542
MEMOIRS OF THE QUEENSLAND MUSEUM
here was rapid. All the material found on the
floor had probably fallen from the ledge recently.
A photographic record was kept as each item
was removed, described, numbered, then double-
sealed in polythene bags. Foam chips were placed
between the bags and the exterior bag partially
inflated. No noticeable damage occurred to the
material during transit to Brisbane, and it is now
held in the collections of the Queensland Museum
(S181/1-88).
The cache comprised 29 items as well as 26
incomplete lengths of twine (5181/6,10,14,27,28,
31,35,26,37,41,42,46,47,49) and 8 fragments of
skin (8181/7,10,13,14, 17,26). The more complete
items are described below in the order of
registration by the Queensland Museum.
(S181/1): A juan knife comprising a large,
silcrete blade of trigonal cross-section and backed
by steep, bi-directional retouch along the thick
back. The distal end tapers to a point, whereas the
proximal end has a haft of Whiptail wallaby
{Macropus parryi ) skin which extends for 3.5 cm.
The skin is attached by a black resin which has red
ochre embedded in it. This classic juan knife
(Mulvaney and Joyce 1965, p. 190) is 15.5 cm in
length, 4.4. cm wide and 2.5 cm thick.
(SJ8I/2): A small, sandstone grindstone
measuring 10.3 by 7.4 by 2.0 cm. Both major
facets exhibit grinding and smoothing, the
obverse side being slightly convex and the reverse
slightly concave. A thick coating of red ochre on
both ground surfaces shows that the grindstone
was used as a palette.
(SI81/3): A water-rolled, quartzite pebble
measuring 7.8 by 7.2 by 3.1 cm. It has red ochre
and kaolinite adhering to the flat, reverse side and
probably functioned as an upper grinding stone.
(S181/4): A lump of kaolinite measuring 7.0 by
5.7 by 6.0 cm. It appears to have been used as a
source of white pigment, and has abraded areas
and striations. Six holes 8 mm in diameter and 10
mm deep, and 10 smaller holes 2 mm in diameter
have also been drilled into the material.
(S181/5): A cylindrical mass of compressed, emu
feathers which has been consolidated with red
ochre grease and unidentified adhesives (PI. 2a).
Tests undertaken by the Pathology Division (Qld.
Health Dept.), have shown that human blood was
not the adhesive used. The remnants of a skin
wrapping have the fur side out. The original
feather mass has now disintegrated into two
sections, the largest of which measures 25 by 11
by 7 cm and the smaller 13 by 7 by 4 cm. Three
smaller wads of emu feathers and red ochre wash
(8181/8,9,30), are almost certainly fragments of
the same original item.
A section of long-bone can be seen embedded
in the largest feather mass, while on one side three
cylindrical objects tightly wrapped in possum fur
twine, with a dense red ochre coating are partially
revealed. On the upper side of this mass there are
the negative impressions of two similar objects.
Another impression can be seen on the second
largest feather mass. It is 13 cm in length and 1 cm
in diameter.
(S181/12): A length of necklace comprising 13
small reels of cut reed sections which have an
outside diameter of 4.5 mm and vary in length
from 4 to 6 mm. Eleven reels are threaded on
2-ply bark fibre 2 mm in diameter. The twine and
reels have been coloured with red ochre (PI. 2b).
A further 31 reels of the same necklace (8181/25)
were recovered from the shelter floor.
(S18I/15): A lump of prepared red ochre which
has fragmented into 2 sections. Each measures 3
by 3 by 1.5 cm.
(8181/ 16): A small section of string bag
measuring 3 by 2 cm. The fabric is of 2-ply, bark
fibre twine of variable diameter, which is woven
in the knotted netting technique. The bag has a
mesh size of 1.5 by 1.1 cm and is probably a
fragment of 8181/34.
(8181/18): A bag made from Brush-tailed
possum {Trichosurus vulpecula ) skin which has
the fur facing inwards (PL 2c). The bag was
originally cylindrical in shape with a circular base
sewn across the centre, with sinew threaded
through pierced holes, and measures 22 cm in
length and 9 cm in diameter. The top of the bag is
made from the rear portion of the animal and is
open with 3 small pierced holes around the
periphery for a draw string. Two holes where the
limbs have been detached, the cloaca and the
remains of the pouch, can still be seen.
When removed the bag contained 34 artefacts
of opaline chert (8181/55-88; Fig. 2; Table 1) and
a lump of yellow ochre measuring 3,8 by 3.5 by
2.3 cm. The ochre has an ovate depression in one
face.
(8181/19): A pouch of Whiptail wallaby {M.
parryi ) skin. The fur faces inwards but most of
the fur has been removed (PL 2d), Traces of red
ochre occur on the interior and exterior. The
pouch measures 21 by 14 by 5 cm and contains an
emu feather head-dress, a bone point and a small
long-bone shaft.
The head-dress has been manufactured by
attaching the emu feathers to a stem of 7 lengths
of bark fibre, which have been tightly bound
transversely with a strand of bark fibre. A small
tassle of bark fibre with a large knot at one end
originates from the top of the fibre stem. The
MORWOOD: MT. INGLIS CACHE
543
FIGURE 1. S181/1 hafted juan knife; 3 water-rolled quartzite pebble; 54 large silcrete scraper. 52 silcrete flake;
53 opaline chert flake; 51 amorphous quartz pebble.
544
MEMOIRS OF THE QUEENSLAND MUSEUM
stem is 0.7 mm in diameter. Its length was not
determined as the item was not removed from the
pouch.
The bone point (SI 81/20) is made from the
proximal end of a juvenile macropod fibula. The
point is polished as well as ground. It is 17 cm in
length.
The long-bone shaft (S181/2!) is from a bird of
pelican or brolga size. It has been snapped at both
ends and has traces of red ochre adhering. The
shaft is 12 cm in length and 0.6 cm in diameter.
(SI8I/22): A section of Brush-tailed possum (7.
vulpecula ) skin. It has the fur facing out and the
anterior has been coloured with red ochre wash.
The base of the tail has been reversed, plugged
with skin, fur and sinew, then tightly bound. This
‘plug’ measures 1.4 cm in length and 0.4 cm in
diameter. The inverted testes are evident on the
inner surface near the tail plug. The skin measures
34 by 6 by 5 cm.
(S181/23): An elongate parcel wrapped in Brush-
tailed possum skin (7. vulpecula ) with the fur
facing outwards (PI. 2e). The parcel contains a
red-ochred mass of emu feathers and at least 8
strands of 2-ply, fur twine (3 mm diameter), A
section of the skin wrapping has been bound with
sinew. The parcel measures 29 by 10 by 5 cm.
(S181/24): A dried cloaca of a large macropod
which has been compressed to form a circular
pendant (PI. 20- Cloacal hairs form a fringe
around the periphery of the disc. A hole has been
pierced near the edge and threaded with a length
of sinew which has been knotted at one end. The
disc has a maximum diameter of 51 mm and is 4
mm in thickness.
(SI81/25): Thirty-one necklace reels of cut reed
sections recovered from the shelter floor. These
are part of the same necklace as S181/12.
(S18J /28d): A wad of Brush-tailed possum fur
(7. vulpecula ) which measures 5 by 2 by 0.5 cm.
(S181/29): A fragment of string bag
manufactured from bark fibre and coloured with
red ochre. It measures 10 by 7 cm. The fabric is
compacted so that the original mesh size is no
longer visible but it is of the loop and single twist
weave (See Davidson 1933, p. 262; Roth 1901, pi.
17). The siring is 1.5 mm in diameter. The
remains of the original opening are still visible.
This is 2.5 cm in diameter and has a draw string
0.9 mm in diameter.
(8181/32): A close-meshed armband which is
woven in the loop and single twist technique. The
fabric is of 2-ply, fur twine 1.5 mm in diameter
and coated with red ochre (PI. 2g). Remnants of
the finishing off weave are still evident at the top
and bottom of the band which is 14 cm in width,
II cm in expanded diameter and has a
circumference of about 30 cm.
(SI81/33): A head-net which is woven in the
knotted netting technique. The fabric is of 2-ply,
bark fibre twine which is 1 mm in diameter and
coloured with red ochre (PI. 2h).
The net is bell-shaped and is 15 cm in diameter
at the open end and 9 cm in depth. The body of
the head-net is woven from a circular ring at the
top which is 2 cm in diameter. The fabric has a
mesh size of 5 mm.
(8181/34): The remains of a string bag which
measures 42 by 29 cm (PL 3a). It is manufactured
in the knotted netting technique with a mesh size
of 2 cm. The fabric is of 2-pIy, bark fibre twine
which is 1.5 mm in diameter. A small portion of
the neck remains.
A piece of macropod skin adheres to the bag
(probably due to insect activity). This measures 16
by 9.5 cm.
(8181/38): A cylindrical roll of budgeroo bark
{Lysicarpus angustifolius ) which measures 28 cm
in length and 8 cm in diameter (PL 3b). The roil
contains four bundles of ‘pins’ bound up with
bark fibre, and a packing of dry grass. One of the
bundles was removed for examination and was
found to contain 20 slicks, each about 9 cm in
length, 3-4 mm in diameter and tapering to a
point at one end. At the other, blunt end, some of
the sticks are split and down and feathers of the
sulphur-crested cockatoo have been inserted.
These splits have then been bound up with bark
fibre strands or fur twine. Other sticks remain
undecorated.
The bark cylinder has been loosely bound with
2-pIy, bark-fibre twine (2.5 mm diameter) which
encircles the cylinder four times.
(8181/39): A quid comprising a continuous
strand of coarse bark fibre. It measures 4.3 by 2,5
by 1.5 cm and appears to have been chewed.
(8181/40): A quid of bark fibre measuring 5.5 by
2.0 by 0.5 cm. It appears to have been chewed and
small indented tooth marks are visible.
(8181/43): A section of tibia shaft from a small
macropod of pademelon size. The shaft has been
snapped at both ends and measures 6.4 cm in
length and 0.9 cm in diameter. This item was
recovered from the shelter floor beneath the cache
shelf. It may therefore, have been deposited by a
natural predator.
(8181/44): A belt made from a single strand of
2-ply, bark fibre twine (1. 5-2.0 mm diameter)
arranged in three parallel loops, then transversely
bound with 2-ply, fur twine which is 1-2 mm
diameter (PL 3c). Each end of the bark fibre
MORWOOD: MT. INGLIS CACHE
545
strand terminates in a small loop around the
composite inner core of the belt.
The belt is 6 mm in diameter and 82 cm in
length.
(S181/45): Four parallel lengths of ochred, 2-ply,
bark fibre twine which have been knotted
together at each end (PI. 3d). The ‘tassle’ is 18 cm
in length and each length of twine is 4 mm in
diameter. The knots measure 2.7 by 1.7 cm.
(S181/48): Four parallel lengths of ochred, 2-ply,
bark fibre twine of which only one remains
complete. The lengths are knotted together at
both ends. The ‘tassle’ is 18 cm in length and each
length of twine is 3 mm in diameter.
(S181/50): A parcel wrapped in Rock wallaby
(Petrogale penicillata ) skin with the fur facing
inwards (PI. 3e). At one end the skin has been
folded in on itself, while the other has been sewn
up with 2-ply, bark fibre strand.
The parcel measures 35 cm in length, 13 cm in
width and 8 cm in depth. It contains 1045 gram of
finely powdered, red ochre and has been bound
with 2-pIy, bark fibre twine around the mid-
section. This twine is of variable diameter and has
been knotted once.
(S181/51): A water-rolled pebble of pink,
amorphous quartz measuring 4.5 by 4.0 by 1 .9 cm
(Fig. 1).
(8181 /52): A flake of fine-grained, white silcrete
measuring 4.2 by 3.5 by 1.0 cm (Fig. 1).
(S181/53): A flake of opaline chert measuring 2.9
by 2 by 0.5 cm (Fig. 1).
(8181/54): A unifacially retouched scraper of
fine-grained, white silcrete. It measures 8.5 by 4,0
by 1.5 cm and has two worked edges of 63° and
72° respectively (Fig. 1).
(8181/55-88): Thirty-four stone artefacts of
opaline chert found in the possum skin bag
(S181/18). Details of these are given in Table 1
and Fig. 2.
TABLE 1. Attributes of thirty-four opaline chert artefacts found in the possum skin bag. (Queensland reference
numbers S181/55-88).
S181
Description
Dimensions (cm)
Edge Damage
Edge Angle
length,
width,
thickness
55
Utilised flake
2.5
2.1
.5
Bifacial utilisation
52°
Unifacial utilisation
69°
56
Utilised flake
2.1
1.5
.4
Bifacial utilisation
34°
57
Utilised flake
2.0
1.2
.3
Bifacial utilisation
30°
58
Tula adze slug with
adhering resin
1.7
.6
.4
Step fractured retouch
48°
59
Utilised flake
3.0
1.7
.5
Unifacial utilisation
46°
60
Utilised flake
2.7
1.7
.5
Bifacial utilisation
24°
61
Utilised flake
1.7
0.6
.3
Unifacial utilisation
o
o
62
Tula adze slug with
adhering resin
2.1
.6
.3
Step fractured retouch
56°
63
Utilised flake
2.5
1.9
.4
Bifacial utilisation
52°
Bifacial utilisation
41°
64
Utilised flake
2.1
1.1
.2
Bifacial utilisation
32°
65
Utilised flake
2.0
1.7
.2
Bifacial utilisation
35°
66
Tula adze slug with
adhering resin
1.5
.6
.3
Step fractured retouch
56°
67
Utilised flake
2.5
2.1
.4
Bifacial utilisation
45°
Bifacial utilisation
52°
Unifacial utilisation
69°
68
Unused fragment
1.9
1.3
.5
69
Utilised flake
1.9
1.6
.4
Unifacial utilisation
42°
70
Utilised flake
3.0
2.1
.6
Bifacial utilisation
36°
Unifacial utilisation
62°
71
Utilised flake
2.6
1.2
.4
Unifacial utilisation
54°
72
Utilised flake
2.7
1.5
.3
Bifacial utilisation
30°
Bifacial utilisation
28°
546
MEMOIRS OF THE QUEENSLAND MUSEUM
73
Utilised flake with resin
adhering
3.1
1.4
74
Utilised flake
2.1
2.1
75
Utilised flake
1.8
1.1
76
Utilised flake
1.7
1.0
77
Utilised flake
2.9
2.8
78
Utilised flake
2.5
1.9
79
Utilised flake
2.5
2.1
80
Utilised flake
2.0
1.7
81
Utilised blade
4.3
1.5
82
Utilised flake
2.3
2.1
83
Utilised flake
2.7
1.9
84
Utilised flake
1.6
1.6
85
Utilised flake
2.9
1.9
86
Utilised flake
3.4
2.3
87
Utilised flake
2.5
1.6
88
Unused fragment
2.1
1.4
DISCUSSION
Ethnographic context
Mt. Inglis is located within the boundary of the
Kanaloo linguistic group which occupied the
headwaters of the Comet River, from below
Rolleston to the Carnarvon Range (Oates and
Oates 1970; Quinnell 1976, p. 14). The
demographic history of this population is briefly
described by Josephson (1887), who states that
around 1860, when Europeans first settled the
area, ‘the tribe’ numbered 500 persons. By 1869
this was reduced to 300 and by April 1879 the
numbers had fallen to 200.
More detailed information on Aboriginal
groups within the area is provided by Priddle
(n.d.), who lived in Rolleston. People camped
near Rolleston still went on ‘walk-about’ and
carried out ‘tribal’ ceremonies until about 1920.
During walk-about the Rolleston group went to a
series of swamps 32 km to the south. They did not
go to Lake Nuga Nuga owned by the
Moolayamber group — the Bemburraburra
(Goddard 1940/41, p. 368), nor did they go to
Fifteen Mile Swamp on ‘Consuelo’ Station owned
by the ‘Carnarvon tribe’ (Priddle n.d., p. 6-7).
On this evidence, it is possible to sketch in the
territorial boundaries of local, land-owning
Aboriginal groups, or patricians. Mt. Inglis
.5
Bifacial utilisation
45°
.5
Unifacial utilisation
50°
.3
Unifacial utilisation
35°
.3
Bifacial utilisation
32°
.9
Bifacial utilisation
50°
Bifacial utilisation
50°
.7
Bifacial utilisation
15°
.6
Bifacial utilisation
32°
.4
Bifacial utilisation
39°
Unifacial utilisation
45°
.8
Unifacial utilisation
61°
Bifacial utilisation
42°
.8
Bifacial utilisation
65°
Bifacial utilisation
25°
.2
Bifacial utilisation
30°
Bifacial utilisation
28°
.2
Bifacial utilisation
36°
Bifacial utilisation
42°
.7
Bifacial utilisation
32°
1.3
Unifacial utilisation
32°
.6
Unifacial utilisation
o
0
1
.2
occurs within, or immediately adjacent to the
territory of the group which utilised Carnarvon
Gorge — possibly the Goon-garee (Winterbotham
1958, p. 219).
Ethnographic details on the material culture,
social organisation and ceremonies of Aborigines
in the upper Comet region are sparse (see
Josephson 1887, p. 96-7; Winterbotham 1958).
However, the range of material culture in general
use throughout the Central Queensland
Highlands can be partially reconstructed by
synthesizing several sources of information (see
Morwood 1979, p.49-80). These sources include
the reports and collections of early observers (e.g.
Ahern 1887; Landsborough 1862; Mitchell 1848)
and the work of salvage ethnographers (e.g. Curr
1887; Donovan 1976; Howitt 1904; Kelly 1935).
The recorded material culture included hand-
thrown spears, clubs, boomerangs, softwood
shields, axes, bags, baskets, containers, hunting
and fishing nets, bone and stone tools, bark
paintings, bullroarers, message sticks, bone
points, pubic aprons, possum skin rugs,
necklaces, pendants, head-dresses, bracelets,
burial cylinders, huts, burial platforms and wells.
In common with other areas of Australia, the
majority of observations and collections are
biased towards the hunting and fighting
MORWOOD: MT. INGLIS CACHE
547
FIGURE 2. Thirty-four opaline chert artefacts found in the possum skin bag (S181/18) and described in Table 1.
548
MEMOIRS OF THE QUEENSLAND MUSEUM
implements of men (see McBryde 1978, p. 185).
The list can be extended by including surviving
field evidence (e.g. dams, stone arrangements,
scarred trees, stone tool scatters), but this field
evidence is usually biased towards the larger and
less perishable elements of the culture, and often
comprises the by-products rather than the end-
products of manufacturing activities. Stenciled
objects in the numerous rock art sites of the
Central Highlands provide further evidence of
Aboriginal material (e.g. Beaton and Walsh
1977). However, stencil art also tends to be biased
towards men’s equipment (Morwood 1979, p.
347). The range of goods recovered from Mt.
Inglis and other rockshelter caches provides a
complementary perspective to the more
traditional sources of evidence on the material
culture of this region.
Stone tools
There are no ethnographic observations of
stone tool use in the region, but the Mt. Inglis
assemblage compares well with the most recent
assemblages recovered from archaeological
excavations (e.g. Beaton 1977; Morwood 1979;
Mulvaney and Joyce 1965). Most notably, blade
technology is evident although the majority of
tools are amorphous flakes. The last 2000 years of
Central Highland stone tool use are characterised
by an increase in the frequency of tula adze slugs
(Morwood 1979, p. 227), and these are also well
represented in the Mt. Inglis collection
(S181/58, 62, 66). Similarly, juan knives are
distinctive implements occurring only in the most
recent industries of the area — the oldest
specimen recovered is less than 600 years in age
(Mulvaney and Joyce 1965, p. 192).
The principal difference between Mt. Inglis
stone artefacts and excavated assemblages in the
area, are the high proportion with retouch or use-
wear, the small size of the specimens in the
possum skin bag (5181/ 18), and the preservation
of hafting medium on the tula adze slugs and the
juan knife. Other hafted juan knives are known
(e.g. Tindale 1957, p. 28; Mulvaney and Joyce
1965, p. 190), but the Mt. Inglis specimen
(S181/1) appears to be the largest specimen yet
recorded, as well as the only hafted example
remaining in an Australian collection.
The function of the amorphous quartz pebble
(5181/51) is uncertain, although similar pebbles
have been found in other caches. For instance, the
Keegan collection includes several quartz and
quartzite pebbles. All have clear indications of
percussive use, and several have vegetable mastic
adhering. The cultural context of the Mt. Inglis
example and three recorded from a shelter in
Moolayamber Gorge (Queensland Museum Reg.
QE 3171), would suggest that some specimens
were also of ceremonial use. 5pencer (1922, p.
105) described one such ceremonial stone
collected from the adjacent Springsure area. This
was carried about wrapped in possum skin and
was not allowed to be seen by women or
uninitiated men. Ethnographic observations
throughout Queensland also state that quartz
pebbles were often used as ‘magic stones’ for a
variety of purposes, including the healing of the
sick (e.g. Hamlyn-Harris 1915, p. 6).
Both the small grindstone (5181/2) and the
quartzite pebble (5181/3) have adhering red
ochre, testifying to their use in the preparation of
pigment for art or decoration. This conclusion is
supported by their association in the cache with
red, yellow and white pigment.
Containers
Bags, baskets, etc. are poorly represented in
Australian museum collections, and the Mt. Inglis
examples add considerably to the range
previously described for the region. All of the
woven material is of 2-ply string. Occasionally,
3-ply is found in Central Highland caches but is
rare (Peter Keegan, pers. comm.). The dilly-bags
(5181/16,29,34) are all manufactured from plant
fibre, most probably from the kurrajong
{Brachychiton populneum ) or from reeds, as
recorded ethnographically (Donovan 1976, p.
112; Josephson 1887, p. 96; MacGlashan 1887, p.
19; 5heridan and Bay 1887, p. 252). Both the
knotted netting and the loop and single twist (see
Davidson 1933, p. 258) were used in the
manufacture of the bags. The same techniques
were employed for a range of woven items in this
region including the Mt. Inglis head-net (knotted
netting), the armband (loop and single twist) and
a hunting net (knotted netting) recovered from a
cave near Springsure and now in the Queensland
Museum collections (QE 3167).
The only previous reference to the use of skin
containers is by Donovan (1976, p. 112) who
states that bags for carrying infants on the upper
Nogoa River were made from kangaroo skin
rubbed with wood ash. Each end was tied with
sinew or fibre and a handle attached. The skin
wallets and wrappers from Mt. Inglis
(5181/5,18,19,23,50) are not unique however, as
similar items are known from other caches. For
instance, the Keegan collection includes an
inverted, possum skin bag very similar to the Mt.
Inglis example (5181/18). It has the leg openings
closed by sinews. The mouth of this bag has a
MORWOOD: MT. INGLIS CACHE
549
‘stopper’ comprising a tassle of emu feathers, and
it contains about a dozen stone flakes of varying
materials, some of which have adhering resin.
Containers made from the belly section of a
goanna skin are also known, one end being finely
stitched up and the other fitted with a string
handle (Peter Keegan, pers. comm.).
The use of bark for manufacturing shallow
dishes, buckets and cylindrical burial cylinders is
ethnographically described (e.g. Josephson 1887,
p. 96; Lethbridge 1885), so the use of this material
for ‘wallets’ (S181/38) is not surprising. Skin and
bark containers are not specifically mentioned by
early observers, but the Mt. Inglis examples are
very similar in type and content to those reported
elsewhere. In Central Australia, for instance,
Spencer and Gillen (1938, p. 611) described
examples made from skin, or from small slabs of
bark tied round with string. These contained emu
feathers, tendon, stone tools, lumps of ochre,
pendants, nose-bones, armlets, necklets and
charms.
The function of the emu feather mass
(8181/5,8,9,30) is uncertain. It is clearly not a
kadaitcha shoe as described by Porter (1961, p.
50) in the Aramac region, but may have
functioned to protect and conceal the length of
bone and other objects contained within it (cf.
Spencer 1922, p. 107, 120). The latter are very
similar in appearance to several objects found in
the Moolayamber cache (QE3i71), as well as to
the Mandu-kuya amulets described by Roth
(1903, p. 37) for N.W. Queensland.
Adornments
Several of the cached adornments have specific
ethnographic references. Necklaces of stong grass
or reed stems cut into lengths (cf. 5181/12,25),
were said to have been common in the region (e.g.
Sheridan and Bay 1887, p. 252). Decorative loops
or reed beads also occur on one of the string bags
held in the Keegan collection.
Ochre and kaolinite (5181/4,5,18,50) were used
for decoration of the body, implements and
rockshelter walls (e.g. Josephon 1887, p. 96-7).
In fact, the practice of coating implements with
ochre may be one factor in the excellent
preservation of cached organic items in this
region. Under normal conditions such material
would be subject to attack by insects, fungi,
bacteria and other micro-organisms and would
deteriorate rapidly. The fact that the Mt. Inglis
cache was saturated with red ochre suggested that
the ochre could have played a role in
preservation. X-ray fluorescence spectroscopy
was undertaken on a sample of the cached ochre
by Dr John Kleeman (Geology Dept., U.N.E.).
This demonstrated the presence of significant
concentrations of copper, zinc and lead, as well as
traces of mercury. The results are detailed in
Appendix 1. Such heavy metal ions are bio-toxins
and are active constituents in many insecticides
and fungicides (Mr Peter Gregg, Microbiology
Dept., U.N.E. pers. comm.; A/Prof John
Brown, Botany Dept., U.N.E. pers. comm.). In
the concentrations present they could have
inhibited, if not prevented, biological damage to
organic material.
The feathers of the emu, white cockatoo and
other birds were used for personal adornment. In
1847, for instance, Mitchell (1848, p. 160) saw
Aborigines coloured with ochre, and with white
cockatoo feathers in their hair and beards. The
use of feathered ‘pins’ as found in the roll of
budgeroo bark (S181/58) was not recorded for
the Central Highlands, although similar examples
also occur in the Keegan collection. Roth (1897,
p. 108) describes their use in North West
Queensland thus —
Feather-tufts or “aigrettes” are formed with
various birds feathers tied on a small sprig,
which is stuck indiscriminately here and there
into the hair: among birds so utilised are the
emu, eagle-hawk, pelican, turkey, crow, etc.
These feather-tufts are very generally used in
times of rejoicing, at corroboree: they may
sometimes be stuck into the waist-belt either
at its side or back, or may be fixed under the
armlets.
Given this documented association between the
use of feather-tufts, armlets and belts, it is
significant that all of these items also occurred in
the Mt. Inglis cache.
The hair net (S181/32), armlet (S181/32), and
bell (S181/44), are identified on the basis of their
similarity to those described by Roth (1897, p.
109) in northwest Queensland, as well as on
general characteristics and size: their use was not
recorded in the Central Highlands. Roth states
that the hair-net was a sort of netted cap used to
prevent the hair dangling in the eyes. It had a
circular ring at the top from which the body of the
net was woven from flax fibre string, then coated
thickly with red ochre grease. When
manufactured by men, the body was woven using
the simple loop weave. Another type was made by
the women using the knotted netting technique, as
used for the Mt. Inglis example.
Many different items were used as pendants in
the Central Highlands, including shells, eagle-
claws, and even ‘a copy of last year’s Nautical
550
MEMOIRS OF THE QUEENSLAND MUSEUM
Almanac’ (Middleton and Noble 1887, p. 90;
Mitchell 1848, p. 358). The use of a dried,
compressed macropod cloaca as a pendant
(S181/24), however, was never noted in this
region or elsewhere.
The bone point {S181/20) and bird-bone tube
(SI 8 1/21) contained in the skin pouch, can be
matched both in the ethnographic and
archaeological records. Beaton (1977, p. 122)
found bone points during excavations at
Cathedral Cave in Carnarvon Gorge, and
suggested that they were utilitarian items used for
piercing skins. Porter (1961, p. 50) described the
ceremonial use of bone points near Aramac in
Central Queensland for inducing sickness and
death. He states that the ceremonial user wore
appropriate make-up as well as kadaitcha shoes
fashioned from emu feathers and held together
with gum and dried blood (cf. Roth 1897, p. 152).
A decorative function is also possible, as it is
known that Central Highland initiation
ceremonies included piercing of the nasal septum
of the novices (e.g. Josephson 1887, p. 97;
Looker, et al. 1887, p. 273). The Mt. Inglis bone
artefacts are indeed very similar to the nose-pins
described by Roth (1897, p. 1 10) in northwest and
Petrie (19()4, p. 20) in southeast Queensland.
These could be a sharp pointed bone of a turkey,
pelican, kangaroo or emu. Other objects such as
grass or reeds could also be used.
The cultural context of many cached examples
suggests a decorative or ceremonial role. For
instance, one bone point was illegally removed
from the Goat Rock site on the upper Warrego
River where it was associated with a bark, burial
cylinder (Fred Cameron, pers. comm.). Another
was found with 3 human skeletons and
ceremonial items in Moolayamber Gorge
(QE3171). The fact that the Mt. Inglis bone point
and tube occurred in a pouch, with a presumed
emu feather head-dress, strongly suggests that
they were of decorative function.
The function of Central Highlands caches
Central Highland caches add significantly to
the range of Aboriginal material culture known
from the region, but just as important is the
evidence that they provide for economic and
social/ceremonial practices. Two types of caching
behaviour were ethnographically described in the
region —
1) The temporary storage of useful, valued
items.
2) The permanent disposal of burial
cylinders.
Examples of temporary storage include the
hanging of large, hunting nets in trees or on
platforms (Donovan 1976, p. 121; Landsborough
1862, p. 101; Mitchell 1848, p. 303, 367). While
exploring the upper Maranoa River, Mitchell also
found a club and a shield stored on a platform
(British Museum of Mankind Reg. 48/2-2/1 and
48/2-2/2). Two hardwood clubs and a hunting
net (QE 3617-9), found in a shelter on the
Staircase Range (Springsure area), may have been
deposited in this way. Elsewhere, the practice of
leaving grindstones as ‘appliances’ at campsites
where they were re-used, has been described (e.g,
Gould 1977, p. 173; Peterson 1968, p. 568).
Grindstones have been found on the floors of
many Cential Highland rockshelters where they
appear to have been deliberately left as
‘appliances’ (e.g. the Art Gallery, Cathedral
Cave). The dispositon of functional ground
implements (axes, mullers) recovered in
archaeological excavations, suggests that many
were originally placed against shelter walls for
later retrieval. In fact, caching of implements
appears to have been a major depositional
mechanism for ground stone artefacts in shelters
(Morwood 1979, p. 219-20). Other items found
cached near occupation sites include stone knives,
cores and wooden implements (pers. obs.; G.
Walsh, pers. comm.). There are also widespread
reports from other areas, of the caching of sacred
items which were periodically removed for
ceremonial use (e.g. Spencer and Gillen 1912, p.
208). Archaeological excavations have shown that
the (presumed) temporary caching of implements
has a long history — one huge silcrete core
positioned against the rear wall of Native Well 1 is
approximately 6000 years old (Morwood 1979, p.
203).
The permanent caching of burial cylinders was
also described by early European observers.
Depending on the status of the deceased several
different means of disposing of the dead were
used in the Central Highlands including
cremation (Looker, et al. 1887, p. 273) and burial
(Lethbridge 1885). Sometimes final disposal of
the remains was delayed for considerable periods
(two or three years), during which time they were
carried tightly bound up in a sheet of bark (see
Robins and Walsh 1979). The common method
for finally disposing of such burial cylinders was
to drop them into a pipe of a hollow tree
(Muirhead and Lowe 1887, p. 27; Looker, et al.
1887, p. 273). However, the fact that many burial
cylinders have been found cached in Highland
shelters and crevices, suggests that this was an
MORWOOD: MT. INGLIS CACHE
551
alternate means (e.g. Gaukrodger 1924; Goddard
1940/41).
It is significant that many caches of material
culture have been found in direct association with
human remains. The earliest report of this
association was by A.S. MacLellan (1901), who
wrote of Aboriginal art and burials at ‘the
Tombs’ rocksheller on the upper Maranoa River
Many a skeleton I saw in the caves there,
and hand and foot imprints and other
impressions on the walls and roofs of the
caves; and fishing nets made out of fibre or
bark. These caves served as a vault for this
wild race.
More recent finds include a sewn marsupial skin
blanket, a bone point, and “a witch-doctor’s
skin-bag” associated with a painted burial
cylinder at Goat Rock, on the upper Warrego
River (QE 6422; Morwood 1979). Another cache
found in Moolayamber Gorge comprised a bone
point, three amorphous quartz pebbles (with
percussion marks and adhering ochre), nine
amulets tightly wrapped in ochred possum-fur
string, and a small steel blade. This material was
found in association with 3 human skeletons and
it is now in the collections of the Queensland
Museum (QE 3171). The mortuary context of
these caches suggest that they were unlikely to be
merely temporary storage of valued items, but
were intended as ‘grave goods’. Looker, et al.
(1887, p. 273) reports that when deceased persons
were cremated, their belongings were burnt also,
so similar principles of disposal may have applied
for other burial practices. However, the ‘burials’
associated with mortuary caches are often of
children, who were too young to have used the
material in life (Peter Keegan, pers. comm.).
Obviously mortuary practices in the Central
Highlands were far more complex and variable
than those ethnographically observed. The
cultural context (rock art, occupation deposits),
content of caches, and age structure of associated
human remains could provide valuable evidence
for these undocumented activities if properly
researched.
Unfortunately, the potential of this source of
cultural data has never been realised as most
burials and caches of material culture were, and
continue to be, desecrated and dispersed without
proper study. Research and management
priorities for this region must include detailed
recording of in situ cached material, plus
documentation and description of finds already in
private and public collections. Some of the ethical
problems in dealing with mortuary evidence have
already been discussed by Robins and Walsh
(1979).
CONCLUSIONS
There is no evidence that the Mt. Inglis cache
was ever associated with human remains. This
collection of ceremonial/decorative items is,
therefore, unlikely to have been a permanent,
mortuary cache, but was probably hidden for
later re-use.
The age of the material is unknown but such
‘de facto’ refuse (see Schiffer 1973, p. 60) most
probably relates to the terminal phase of
Aboriginal occupation. On the evidence of
Priddle (n.d.. p. 34), elements of traditional life
in this region continued until 1920, and this
provides the most recent possible dale for the
material. A similar cache from Moolayamber
Gorge contained a metal knife with a resin haft,
indicating a post-European contact date — i.e.
later than 1840 for this region. The Mt, Inglis
material can, therefore, be compared and
contrasted with ethnographic observations of the
contact period. Clearly this material provides
evidence for a range of material culture and
activities, many of which were never documented.
Such cached material also provides a timely
reminder to researchers. Most of the evidence for
‘recent* Aboriginal culture in the Central
Highlands is based on superficial and biased
ethnographic accounts and collections. There is
therefore, a tendency to equate the simplicity of
surviving evidence with a simplicity of life-style
and material culture (cf. White 1977). It is a
sobering thought that for a minimum of 19,000
years (see Mulvaney and Joyce 1965), successful
Aboriginal occupation of the Highlands
depended on a finely-honed economic and
ideological adaptation. This was based on non-
material, esoteric knowledge about a wide range
of resources, yet both the ideological and organic
components of Aboriginal culture are beyond the
usual scope of archaeological investigation.
ACKNOWLEDGMENTS
The Mt. Inglis material was removed and
researched under the auspices of the Archaeology
Branch, D.A.LA. I wish to thank Kate Sutcliffe
and Jeff Pratt for organising logistic support.
Richard Robins and Michael Quinnell
(Queensland Museum) and Peter Keegan (Roma
resident) gave considerable assistance while the
following people also provided expertise on
various aspects of the work — Dr John Kleeman
552
MEMOIRS OF THE QUEENSLAND MUSEUM
(X-Ray fluorescence spectroscopy), A/Professor
John Brown (biocides), Dr Hans Brunner (hair
identification), Dr Michael Archer and Steve Van
Dyke (faunal identification), Dr Neville Stevens
(geological advice), Kathy Morwood
(draughting), Alan Easton (photography) and
Wendy Chappell (typing).
Appendix 1 — Results of X-ray fluorescence
spectroscopy of red ochre.
Dr John Kleeman, Geology Dept., U.N.E.
Item S181/50 of the Mt. Inglis cache contained
1045 grams of finely powdered, red ochre. Three
grams of this was removed and prepared as a
pressed sample mount. The basis of the technique
used to test for heavy metal ions is fully described
in Norrish and Chappel (1967). The following
results were obtained —
Cu 45 ± 4 ppm
Zn 79 ± 6 ppm
Pb 66 ± 5 ppm
Hg 1-3 ppm (semi-quantitative)
As not detected at or location, say less
than 20 ppm
Cd not detected at location, say less than
10 ppm
Note that the lower limits of detection for As and
Cd are not well known as we do not analyse them
routinely. Subject to a (perhaps) imprecise “less
than” figure, they are not present in the sample.
LITERATURE CITED
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west of Blackall. In E.M. Curr, ‘The
Australian Race’, Vol. Ill; 72-5.
Beaton, J.M. 1977. Dangerous harvest.
Unpublished Ph.D. thesis, A.N.U.
and G.L. Walsh, 1977. Che-ka-ra. Mankind
11 (1): 46-8,
Curr, E.M. 1887. ‘The Australian Race’, Vol.
Ill, (Melbourne).
Davidson, D.S. 1933. Australian netting and
basketry techniques. J, Polynesian Soc. 42:
257-99.
Donovan, H.L. 1976. The Aborigines of the
Nogoa Basin. An ethnohistorical/
archaeological approach. Unpublished B.A.
Hons, thesis, University of Queensland,
Gaukrodger, D.W. 1924. Queensland
Aboriginal tombs in the Great Dividing
Range. Sydney Mail 27 February: 19-20.
Goddard, R.H. 1940/41. Aboriginal rock
sculpture and stencilling in the Carnarvon
Ranges. Oceania 11: 368-72.
Gould, R.A. 1977. Puntutjarpa Rockshelter and
the Australian Desert Culture. Anthrop.
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Hamlyn-Harris, R. 1915. On certain
implements of superstition and magic:
illustrated by specimens in the Queensland
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Howitt, A.W. 1904. ‘The Native Tribes of
south-east Australia’, (London: MacMillan).
JOSEPHSON, T, 1887. Head of the Comet River.
In E.M. Curr, ‘The Australian Race’, Vol.
Ill: 96-9.
Kelly, C.T. 1935. Tribes on Cherburg
Settlement, Queensland. Oceania 5 (4):
461-73,
Landsborough, W. 1862. ‘Journal of
Landsborough’s Expediton from Carpentaria
in Search of Burke and Wills.’ (Melbourne:
F.F. Bailliarra).
Lethbridge, R.C. 1885. Letter to A.W. Howitt,
20 July 1885, Courtesy of D.J. Mulvaney.
Looker, W.H., et al. 1887. Paroo and Warrego
Rivers north of Lat. 27 30 and Mungallella
Creek, In E.M. Curr, ‘The Australian Race’
Vol. Ill: 27-29.
McBRYDE, I. 1978. Museum collections from the
Richmond River District. In I. McBryde (ed.)
‘Records of Times Past’, (Canberra:
A.I.A.S.).
MacGlashan, j. 1887. Main range between the
Belyando and Cape Rivers waters. In E.M.
Curr, ‘The Australian Race’, Vol. Ill: 18-25.
MacLellan, A.S. 1901. The Queenslander, 9
February 1901.
Meston, a. 1901. Among the Myalls. Maranoa
Aborigines. The Queenslander, 12 January
1901.
Middleton, T. and E.I, Noble, 1887. Nogoa
River. In E.M. Curr, ‘The Australian Race’,
Vol. Ill: 90-5.
Mitchell, T.L. 1848. ‘Journal of an expedition
into the Interior of Tropical Australia’.
(London: Longman),
Morwwd, M.J. 1979. Art and stone: towards a
prehistory of central western Queensland.
Unpublished Ph.D. thesis, ANU.
Muirhead, j. and C. Lowe, 1887. Belyando
River. In E.M. Curr, ‘The Australian Race’,
Vol. Ill: 26-35.
Mulvaney, D.J. and E.B. Joyce, 1965.
Archaeological and geomorphological
investigations on Mt. Moffat Station,
Queensland, Australia. Proc. Prehistoric
Society, 31: 147-212.
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NoRRiSH, K. and B. Chappell, 1967, X-ray
fluorescence spectrography. In J. Zussman,
‘Physical methods in Determinative
Mineralogy’, (London; Academic Press),
161-214.
Oates, W.J. and L.F. Oates, 1970. *A revised
linguistic survey of Australia’. (Canberra:
A.I.A.S.).
Peterson, N. 1968. The pestle and mortar: an
ethnographic analogy for archaeology in
Arnhem Land. Mankind 6 (11): 567-70.
Petrie, C.C. 1904. ‘Tom Petrie’s reminiscences
of early Queensland’. (Brisbane: Watson,
Ferguson and Co.).
Porter, J.A. 1961. ‘Roll the Summers Back’.
(Brisbane; Jacaranda Press).
Priddle, V., (n.d.) ‘Dung on his boots’.
(Brisbane).
Quinnell, M.C. 1976. Aboriginal rock art in
Carnarvon Gorge, South Central
Queensland, Unpublished M.A. thesis,
University of New England.
Robins, R.P. and G.L. Walsh, 1979. Burial
cylinders. The essence of a dilemma in public
archaeology. Austral. Archaeol. 9: 62-76.
Roth, W. 1897. ‘Ethnographical studies among
the North-west-central Queensland
Aborigines’. (Brisbane: Government
Printer).
1901. String, and other forms of strand,
basketry, woven bag and net-work. ‘North
Queensland Ethnography’, Bulletin No. 1.
(Brisbane: Government Printer).
1903. Superstition, magic and medicine. ‘North
Queensland Ethnography’, Bulletin No. 5.
(Brisbane: Government Printer).
Schiffer, M., 1973. Cultural formation
processes of the archaeological record:
applications at the Joint Site, East-Central
Arizona. Ph.D. thesis, University of Arizona.
Sheridan, R. and F.B. Bay, 1887. Part of the
Maranoa River and country around Roma. In
E.M. Curr, ‘The Australian Race’, Vol. Ill:
251-7.
Spencer, W.B. 1922. ‘Guide to the ethnological
collection exhibited in the National Museum
of Victoria’, (3rd ed.) (Melbourne:
Government Printer).
and F.J. Gillen, 1912. ‘Across Australia’,
(London: MacMillan).
and F.J. Gillen, 1938. ‘The native tribes of
central Australia’, (London: MacMillan).
Tindale, N.B. 1957. Cultural succession in
south-eastern Australia from Late
Pleistocene to the present. Rec. S.A. Mus.
13: 1-49.
Winterbotham, L.P. 1958. The initiation of
Mirianbuddy. Mankind 5 (5): 219-20.
554
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
Context of the Mt. Inglis cache.
Top — general view of Mt. Inglis shelter (central background).
Middle — general view of site during cache removal.
Bottom — cache after removal of three concealing sandstone blocks.
MORWOOD: MT. INGLIS CACHE
555
556
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
Items from the Mt. Inglis cache, a consolidated emu feather mass and
skin wrapping (S181/5): b length of necklace (S181/I2); c possum
skin bag and ball of yellow ochre (SI8I/I8); d pouch of Whiptail
wallaby skin containing bone tube, bone point and emu feather head-
dress (SI 8 1/19); e possum skin parcel containing emu feathers and
fur twine (S181/23); f pendant made from dried cloaca of a large
macropod (S181/24); g fur twine armband (S18I/32); h bark-fibre
head-net (S181/33). Photos courtesy Queensland Museum.
MORWOOD: MT. INGLIS CACHE
557
558
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 3
Items from the Mt. Inglis cache, a remains of string bag (S181/34); b
roll of budgeroo bark containing feathered ‘pins’ (S181/38); c waist-
belt of fur and bark-fibre twine (S181/44); d tassle of bark-fibre twine
(S181/45); e parcel of Rock wallaby skin containing powered red
ochre (S181/50). Photos courtesy Queensland Museum.
MORWOOD: MT. INGLIS CACHH
559
Mem. QdMus. 21(2): 561—65. [1984]
INCISED STONES FROM GLENORMISTON STATION, S.W. QUEENSLAND
M.J. Morwood
Univeristy of New England
and
M. Gibson
Archaeology Branch, D.A.I.A.
ABSTRACT
This paper describes an Aboriginal cache of incised stones and an associated myth. The
stones, incised designs, and story are then discussed in the light of other evidence for
Aboriginal culture in S.W. Queensland.
INTRODUCTION
Many years ago Charlie Trottman, an elderly
Aborigine from Glenormiston Station showed
Jim Newman (widely known as ‘Old
Kookaburra’) an Aboriginal cache in a small
rockhole near Lake Wonditti, northeast of
Glenormiston homestead, S.W. Queensland (Lat.
22°55’S, Long 138°48’E). The cache comprised
natural stones which were said by the local
Arraringa people to be a stone boomerang, knife,
healing stone, and kadaitcha shoe. A solitary
mulga tree adjacent to the rockhole was said to
have been a spear.
The cache site is close to a tailing yard and over
the years the material was removed several times
for examination by musterers camped nearby. As
a result the stone knife had disappeared and Mr
Newman, who is of Kalkadoon descent, feared
for the safety of the remaining items. In 1977, he
contacted one of the authors (M.G.) and asked
for the items to be removed to a place of safe-
keeping. They are now in the collection of the
Queensland Museum (Reg. No. S362/1-3). This
paper is written with the consent and co-operation
of Mr Newman.
The associated mythology of the material
concerns a Kalkadoon man who had been
promised a wife by the local people, and who
travelled down to Lake Wonditti for this purpose.
Finding nobody there he threw his equipment into
the rockhole and vanished. The story is best told
as explained by Jim Newman to Kate Sutcliffe
(Archaeology Branch, D.A.I.A.) during a taped
interview.
‘A man had been around here cooking and
got friendly with this dark old fella and he tell
me a story then. The black fella call it religion.
You see it is a religion to them. He showed me
these things — stone boomerangs, stone knife,
what they cut the kidneys out with, and a
healing stone, what they heal the wound up
with. You got to put it in the fire like a
.soldering iron. You put the stone in the fire
then you put it on the wound and heal it up and
its as good as new again.
So I want to gel to the bottom, to get the full
story how he got there to leave no feathers
there. Well he said he come across from here
between Cloncurry and the Georgina.
According to this old fella telling the story,
telling it to me, he had to go and pick up his
little wife what the black fella gave him in black
fella law. He had to go and get his wife. When
he go there, there was no one there at this one
little lake-waterhole. It’s a lovely big lake fresh
water, but it was milky, the colour w'as milky.
So, I’m baking bread one afternoon and this
old fella said to me, “I’ll show you devil
directly”. I wasn’t interested about the devil at
all, I went about cooking my bread. So at last I
gave up.
Not very far away from where our camp was,
he had a little shallow cave with all these stones
in it — the boomerang, the kadaitcha shoe, like
these T It was all red stone.
Well I said, “How he get there then?” He
couldn’t find this girl, nobody there, tribe’s
gone. So, well he pulls his boots off. Threw
them into this cave — stone knife, and the
boomerang and this healing stone. In they go.
He stuck his spear in the ground alongside the
cave. And he told me that from the spear this
little mulga tree grow. And there’s no mulga
around the place within 40 mile around in the
area. All the rest of the trees are whitewood,
bloodwood, coolibah, gum.
And 1 said “What happened to him then?”, I
said. Well, he said he just went like this
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MEMOIRS OF THE QUEENSLAND MUSEUM
“choo!”. He said, he went straight up into the
cloud. Well I said “He must be still up there
then. How could he disappear into the cloud, a
Kalkadoon black fella and leave all his gear
behind.” He said “choo!” like somebody give
him a bump and away he went.
Well, that’s the end of that old story. It was
handed down to one another. They handed it
down to me and now I’m giving it to you on
tape’.
DESCRIPTION
The Glenormiston cache comprised three
limestone fragments of unusual shape. Each has a
natural surface staining/paiina of red
colouration. Almost certainly the material is
derived from an exposure of Georgina limestone
which outcrops immediately south of
Glenormiston Station (Dr. Neville Stevens; pers.
comm.).
The 'boomerang' is an elongate, arc-shaped
fragment measuring 48.0 cm in length, 1 1.O cm in
width and 4.2 cm in thickness. At one end sinuous
lines, a circle, and a bird track have been incised
to a depth of approximately 0.2 mm (Fig. la).
The incised motifs are of the same colour as the
unmodified limestone surface.
The 'kadaitcha shoe* is a weathered fragment
23.5 cm in length, 11.0 cm in width and 4.9 cm in
thickness. Fine bedding lines 2 mm apart occur at
right angles to the long axis, while the upper
surface has an irregular topography of ridges and
grooves formed by solution grooving. A pattern
of shallowly incised concentric circles and
connecting lines occurs on the upper surface (Fig.
lb) . The lower surface is slightly convex in cross-
section and bears a finely incised figure of an
Aboriginal warrior with shield, spear, spear-
thrower, head-dress, and body decoration (Fig.
l c) . All of the incised designs on the fragment are
shallow and approximately 0.1 mm in depth and
thickness. The incised designs are similar in
colouration to the remainder of the fragment, and
accurate discernment requires oblique lighting. It
is apparent that the incised designs are of some
antiquity as more recent scratches of similar
depth are white in colour.
The 'healing stone ’ is water-rolled and roughly
spherical in shape. It has a maximum diameter of
6.9 cm and exhibits bruising and patina loss in
one area. However, the white colouration of this
suggests that it is probably modern damage.
DISCUSSION
McCarthy (1976, p. 66) notes that Aboriginal
incised stones have been found in New South
Wales and Queensland ‘but they are very rare’.
The Glenormiston examples are significant not
only because such artefacts are uncommon, but
also because the associated mythology is known.
‘Old Kookaburra’ (pers. comm.) states that he
knows of a similar cache located between
Glenormiston and Tobbomoree (N.T.) Stations,
so the find is not unique. For example, Stubbs
(1974, p. 86) illustrates a fine-grained, igneous
rock with an incised, linear pattern which was
found on Glenormiston Station. An incised
pebble was also recovered from a rock hollow at a
rock engraving site near Mt. Isa (pers.
knowledge). Designs on this pebble include a grid,
a dot series, a concentric circle, and a 5-tiered
chevron (see Armstrong n.d., p. 58).
Unfortunately, the ceremonial context and
function of such items is unknown.
The incised motifs are of particular interest as
they probably relate to the associated mythology.
Those on the stone ‘boomerang’ and the upper
surface of the ‘kadaitcha shoe’ can be closely
matched in motif emphasis and composition with
designs found on other items of material culture
from the area — e.g. boomerangs, tjurunga.
Kelly (1968, p. 565) for instance, describes
concentric circles and a sinuous line incised on a
tjurunga of an elder of the Mulligan river woma
snake totem. Circles, concentric circles, spirals,
sinuous lines, and bird tracks are also widely used
in the numerous rock art assemblages of western
Queensland (Elkin 1949/50; Morwood 1979).
Typically, there is a high proportion of non-
figurative designs and many of these are shared
with assemblages of central Australian art (cf.
Edwards 1965, 1966; Mountford 1960;
Mountford and Edwards 1963).
The incised figure on the ‘kadaitcha shoe’ (Fig.
Ic) is very different from other figurative motifs
known from the area. Normally the figurative
component of western Queensland Aboriginal art
would fall into Maynard’s ‘Simple Figurative’
category. (Maynard 1976). It is crudely
naturalistic, rigid, standardised, and comprises
simplified silhouettes of humans and animals.
Humans are usually male and depicted from the
front with splayed legs and exaggerated penises.
Anatomical detail is minimal and the figures lack
facial features and body contours.
By contrast the Glenormiston incised figure is
depicted in twisted perspective and conveys a
MORWOOD AND GIBSON: INCISED STONES FROM GLENORMISTON
563
a
cm.
FIGURE 1. Designs incised on items from the Glenormiston cache
a) Track-line-circle composition on the ‘boomerang’.
b) Line-circle composition on the upper surface of the ‘kadaitcha shoe’.
c) Figure of an Aboriginal warrior on the lower surface of the ‘kadaitcha shoe’.
564
MEMOIRS OF THE QUEENSLAND MUSEUM
sense of movement. The amount of detail shown
and the fact that the principal outlines of the body
were carefully incised several times suggests that
the task was not undertaken casually but was of
some importance. Facial features and body
contours are shown, while body decoration and
associated weapons are accurately depicted — the
figure advances clutching a decorated shield with
the right arm poised to propel a spear from a
spear-thrower. Although the associated story
concerns a Kalkadoon man, it is interesting that
the spear-thrower illustrated is of the flattened,
leaf-shaped type used on the upper Mulligan and
upper Georgina Rivers and along the Toko Range
(i.e. local). It is quite different from the linear
lath type used by the Kalkadoon in the Boulia,
Leichhardt-Selwyn and Cloncurry districts (Roth
1897, pp. 148-9 and Fig. 372). Body decoration
includes a head-dress (possibly feathers or
ceremonial items), a series of vertical lines down
the chest (probably body paintings) a belt (?), and
horizontal lines across the thighs. The figure
appears to be wearing kadaitcha (?) boots as
described in the associated myth. In the attention
to detail, style and depiction of movement this
incised human figure differs markedly from the
basic naturalism characteristic of western
Queensland figurative art.
The associated mythology also has several
features of interest. It includes a covert
explanation for the unexpected absence of the
wife. In the story the hero arrives to collect his
wife but finds her absent. The colour of Lake
Wonditti, is then described as ‘milky’. This had
seminal connotations and suggests that the wife
may have absconded with another man.
The Glenormiston myth also differs in an
important detail from other myths recorded in
this region. The mythology of the Nappamerrie
engraving site on Cooper Creek, provides an
interesting contrast, as this appears to be the only
rock art site in southwest Queensland for which
details of the associated mythology have been
recorded. These engravings comprise concentric
arcs, upright lines, and circles, and refer to a
‘murra murra’ myth of the dog cult-totem of the
Yanruwanto tribe (Elkin 1949/50).
‘The petroglyphs are said to have been made by
two “dog women”, Widjini and Kilki mura
(heroines), who camped at the spot and used to
sit under the two big ti-trees nearby. A third
slab was similarly and distinctly marked. My
informant said it represented poa, a grass seed.
The concentric arcs represented the falling of
the grass seed on a heap under the grinding
stones.’ (Elkin 1949/50, p. 141).
The Nappamerrie myth ends with the two
heroines travelling up-stream to a spot called
Malgera where they can still be seen as white
stone. This type of transformation and residence
at a specific locality is characteristic of the murra,
or ‘western’ myths found throughout the
Australian arid zone (See Allen 1972, p. 112). In
northeast South Australia and southwest
Queensland, murra myths were associated with
patrilineal cult-totems, together with a
philosophy of localised totem centres and clans
(Elkin 1933, p. 138). Although not stated in
Elkin’s account, Malgera would certainly have
served as a ceremonial centre of the local dog
totem (cf. Spencer and Gillen 1912, pp. 96-7).
The out-come of the myth associated with the
Glenormiston incised stones differs in an
important detail: the hero did not remain at a
specific locality but left the earth and disappeared
into a cloud. This trait is far more characteristic
of Aboriginal mythology in SE. Australia with
matrilineal, ‘social’ totemism and non-localised
clans (Elkin 1933, p. 138).
To conclude, the Glenormiston material adds
significantly to the little information available on
the cultural and mythological context of
Aboriginal art in southwest Queensland. Aspects
of the cache also extend the range of material
culture, art and mythology of the region beyond
that previously recorded.
ACKNOWLEDGMENTS
This paper could not have been written without
the considerable assistance of Mr Jim Newman of
Mt. Isa. We also wish to thank Dr Neville Stevens
(Mineralogy Dept., Queensland University) for
his geological identifications, Dr Nicolas Peterson
(A.N.U.) for his anthropological advice, and
Wendy Chappel for typing.
LITERATURE CITED
Allen, H., 1972. Where the crow flies
backwards. Unpublished Ph.D. thesis.
A.N.U.
Armstrong, E.M., n.d. ‘The Kaladoons: a study
of an Aboriginal tribe on the Queensland
frontier.’ William Brooks and Co., Brisbane.
Co., Brisbane.
Edwards, R., 1965. Rock engravings and incised
stones: Tiverton Station, northeast South
Australia. Mankind 6(5): 223-31.
MORWOOD AND GIBSON: INCISED STONES FROM GLENORMISTON
565
1966. Comparative study of rock engravings in
South and Central Australia. Trans. Roy.
Soc. South Australia. 90: 33-6.
Elkin, A.P., 1933. Studies in Australian
totemism. The Oceanic Monographs, No. 2.
Australian National Research Council.
1949/50. The origin and interpretation of
petroglyphs in southeast Australia. Oceania
20: 119-57.
Kelly, J.D., 1968. Hut sites, rock engravings,
stone arrangements and tjurunga. Mulligan
River, Queensland. Mankind 6{\\)\ 563-6.
Maynard, L., 1976. An archaeological approach
to the study of Australian rock art.
Unpublished M.A. thesis, University of
Sydney.
Morwood, M.J., 1979. Art and stone: towards a
prehistory of central western Queensland.
Unpublished Ph.D. thesis, A.N.U.
MOUNTFORD, C.P., 1960. Simple rock engravings
in Central Australia. Man 66: 145-7.
and R. Edwards, 1963. Rock engravings of
Panaramitee Station, northeastern South
Australia. Trans. Roy. Soc. South Australia
86: 131-46.
Roth, W.E., 1897. ‘Ethnological studies among
the North-west-central Queensland
Aborigines.’ Govt. Printer, Brisbane.
Spencer, B. and F.J. Gillen, 1912. ‘Across
Australia.’ Macmillan and Co., London.
Stubbs, D., 1974. ‘Prehistoric art of Australia.’
Macmillan, Melbourne.
Mem, QdMus. 21(2): 567—75. [1984]
TWO ABORIGINAL SHELTERS IN SOUTHWESTERN QUEENSLAND
E. Durbidge
Box 8, Pt. Lookout, Queensland;
and
R. Robins
Queensland Museum
ABSTRACT
Two dome-shaped shelters made of Gidgee {Acacia cambagef) by Aborigines, post-
European contact, have been found in southwestern Queensland. Shelter 1 is in good
condition. From this the general sequence of construction can be established.
Shelter 2 is similar, but in poor condition. Such shelters are important and should be
preserved.
INTRODUCTION
Few aboriginal shelters can be seen in areas of
Queensland where Aborigines no longer make or
use such shelters. Despite their ephemeral
appearance and the short term usage envisaged by
their builders, some shelters can still be found in
remote parts of southwestern Queensland where
the dry climate and isolation have retarded
processes of decay and destruction. Age, frail
construction, and general lack of protective
measures afforded these structures makes detailed
description of them necessary if data about site
location, site use, dwelling construction and
Aboriginal history is to be recorded and
preserved.
Two previously unrecorded shelters were
examined by one of us (E.D.) in mid 1982 in the
Birdsville area. These are reported here. Further
information is held in the Queensland Museum,
and the Archaeology Branch Department of
Aboriginal and Islanders Advancement.
THE SITE
The shelters are situated approximately 85 m
apart, 100 m up a gentle slope from a seasonal
drainage line which, during floods, forms part of
the Diamantina River Channels. The general area
is treeless, lightly grassed, ‘undulating stony
downs* (Dawson 1974). Stands of Gidgee {Acacia
cambagei) grow along the drainage line. The
nearest permanent water to the site is at
Nerathella Waterhole, 3.5 km to the north. No
artefacts were observed near either shelter.
SHELTER 1 (PI. 1, Fig. 1)
The shelter lies on a small mound. It was based
on four forked interlocking branches (Fig. 1, a, b.
c, d), presumably from the nearby Acacia
cambagei stand.
The frame of the dome-shaped shelter was
formed by burying the stout ends of four forked
branches (Fig. I, a, b, c, d) in the ground. Three
of these branches are long, and curved. These
(Fig. 1 a, c, d) interlocked. A ‘ridge pole* (Fig, le)
was placed from the remaining structural branch
which is short, straight and stout, (Fig. Ib) to rest
near the top of the ‘dome* formed by the
interlocking branches. (Fig. I a, c, d). The
stoutest of the main supports has a maximum
diameter of 16 cm. Diameter of the most slender
structural branch is 7 cm. The basic frame was
overlaid with curved branches (of diameters
varying between 3 cm and 15 cm). The butt ends
of these non-struclural branches were also buried
in the earth. The entrance (Fig. 1) faces east. All
branches used were cut by metal axes (PI, 3c).
This shelter is in good condition generally,
although some of the small, lighter, lateral ‘wall
liners’ have collapsed and now lie at random
round the base. Charcoal fragments indicating
the former fire place, lie 4.5 m from the shelter
entrance.
SHELTER 2 (PI. 2b)
This is very similar to shelter 1 , but is less well
preserved. Only one main structural support
branch and the ‘ridge pole’ are still upright. All
the curved ‘wall’ timbers have collapsed and lie
within 4 m of the standing support branch. A
fireplace consisting of charcoal and baked clay
was located 8.0 metres from the former entrance.
DISCUSSION
There are no known station records, published
accounts, or archival material on these shelters.
568
MEMOIRS OF THE QUEENSLAND MUSEUM
FIGURE 1. Plan of shelter 1 showing position and size of structural and other branches. Main structural branches
are drawn in full. Position of embedded ends of other branches are indicated by open circles. Entrance (X).
Metal axe cuts (PI. 3c) on all timbers indicate that
the shelters were constructed after European
settlement (i.e. at least since the 1870’s). Local
opinion suggests the shelters were probably
constructed at a temporary camp site by
shepherds when floods restricted movement (J.
Evans, pers. comm.).
Shelters such as these are important for two
reasons: Firstly, they may be seen as historical
documents of recent Aboriginal occupation for
which few written records exist (e.g. Duncan-
Kemp 1933, 1964; Robins 1981). As tangible
evocative evidence of Aboriginal occupation they
may become a focal point for folk sentiment.
Secondly, they offer archaeological evidence for
such factors as technology, site location, campsite
size and seasonality. Such evidence may assist in
the formulation of models of Aboriginal
settlement and subsistance. Although this
evidence may not be able to be applied in the form
of direct analogy to prehistoric archaeological
evidence it may help explain anomalies observed
in the archaeological record in other areas.
ACKNOWLEDGMENTS
Jim Evans of Durrie Station and Genevieve and
Ashley Daley of Mt. Leonard provided
information on the shelters and on early history
of the area. One plant specimen was identified by
the Government Botanist. Jeanette Covacevich
(Queensland Museum) assisted in preparing the
manuscript. Plates were prepared with the
assistance of Maureen Kelly and David Bligh
(Queensland Museum).
LITER.ATURE CITED
Dawson, N.M., 1974. Land Systems in Western
Arid Region Land Use Study — Part 1 . Tech,
Bull 12, Div. Land Utilisation.
Duncan-Kemp, A.M., 1933. ‘Our Sandhill
Country’. (Angus and Robertson: Sydney).
1964. ‘Where Strange Paths Go Down*. (W.R.
Smith and Paterson Pty. Ltd.: Brisbane).
Robins, R., 1981. Four Aboriginal dwelling sites
in southwest Queensland. Australian
Archaeology 12: 79-90.
570
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 1
Aboriginal shelters near Birdsville, SE.Q.
a) Shelter 1 and Shelter 2, 85 m apart in undulating,
stony downs.
b) Shelter 1 showing size and general appearance.
DURBIDGE AND ROBINS: ABORIGINAL SHELTERS
571
572
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 2
General appearance of the shelters.
a) Shelter 1
b) Shelter 2
DURBIDGE AND ROBINS: ABORIGINAL SHELTERS
573
574
MEMOIRS OF THE QUEENSLAND MUSEUM
PLATE 3
Detail of Shelter 1
a) Entrance
b) Structural detail near the entrance. One of the main
structural branches (Fig, lb) and the ‘ridge pole’
(Fig. le) can be seen slightly to the right of centre.
c) Metal axe cut Gidgee {Acacia cambagei) ends.
DURBIDGE AND ROBINS: ABORIGINAL SHELTERS
575
576
WILLIAM ALEXANDER MCDOUGALL
1905-1982
The death occurred on 7th January 1982 of the
retired Government Entomologist for Queensland
and noted authority on native Muridae, Dr
William Alexander McDougall. Mac had served
as an Entomologist and Zoologist in the
Queensland Department of Primary Industries
for 45 years and after his retirement in 1971 was
made an Associate of the Queensland Museum in
which capacity he continued to work in
Australian native rats especially Melomys.
Mac’s zoological career commenced in 1926
when he was appointed to the Bureau of Sugar
Stations at that time a branch of the Department
of Agriculture and Stock (now Primary
Industries). He trained as an entomologist under
Professor Goddard in the Zoology Department of
the University of Queensland and subsequently
served at Meringa 1928-32 and Mackay 1932-49.
Whilst at Mackay he undertook research on the
range of insect problems in sugar cane including
major work on wireworms which earned him the
degree of MSc in 1934. However, following a
period of study on rats at the University of
Sydney his main interest became an investigation
of the biology, ecology and control of the
canefield rat {Rattus conatus Gould). For this
work he was awarded the degree of DSc in 1949
and acknowledged as the Australian authority on
rodent ecology and control. Some of his early rat
studies had also involved collaborative work with
medical researchers on the control of Weil’s
Disease in canefield workers.
In 1949 he transferred from the Sugar Bureau
to take charge of the Entomology Section within
the Science Branch of the Department. During the
next 22 years he was to make totally different but
equally important contributions in entomology
and zoology. He recruited and guided the work of
a group of entomologists and zoologists who were
to make agricultural entomology and native
fauna research into the vigorous disciplines they
are today in Queensland. He excercised a very
personal style of leadership and all of his staff
were actively encouraged to seek further training
with the result that their record for higher degrees
and scientific publications was outstanding. It
was due in part to the high standards of
experimentation he demanded but also to his
superb skills as a scientific editor.
Parts of his Departmental duties involved
service on interstate committees. He was often
controversial but the minutes of these committees
carry frequent evidence of the salutary effect of
his comment. Most important would have been
his contributions Australia-wide as a foundation
member of the Committee of Commonwealth and
State Entomologists. Within Queensland he was a
member of the Agricultural Requirements Board
from its inception in 1952. This Board carried the
responsibility of regulating the use of pesticides
within Queensland and he instituted and followed
a rigid ethical code in the approval of insecticides.
Towards the end of his career Mac was able to
expand his interest in vertebrate zoology by the
formation of a zoology group within Entomology
to work on native fauna. This group became the
Fauna Branch of the Department of Primary
Industries at his retirement and subsequently
merged with National Parks to form the
independent National Parks and Wildlife Service
of Queensland. This provided him personally
with the opportunity to develop his original
interest in the Muridae, and after retirement he
undertook a reorganisation of the Queensland
Museum reference collection based on newly
available chromosome and skeletal studies by
members of his previous staff.
Mac was very much a Queenslander. He was
born in Ipswich but raised in Goondiwindi
although he returned to Ipswich for his secondary
education as a boarder at the Ipswich Grammar
577
School. He excelled at sport and in his final year
was a member of every school sporting team and
captain of all but one. Later at the University of
Queensland as a student of King’s College his
sporting prowess was again apparent with the
award of the blues for cricket and football. In
North Queensland he was similarly successful at
inter-city cricket and tennis.
His personal interests were mainly his family,
the staff who worked for him and their families,
and sport, latterly as a spectator, but he
participated in a wide range of community
activities. To Mrs McDougall, their daughters
and grandchildren we express our appreciation of
Mac as a friend and mentor and of his
contributions to his chosen discipline.
N.W. HEATHER.
CONTENTS
Nearhos, S.P. and R.J.G. Lester
New records of Bopyridae (Crustacea: Isopoda: Epicaridea) from Queensland Waters 257
Davies, Valerie Todd
Pitonga gen. nov., a spider (Amaurobiidae: Desinae) from northern Australia 261
Dahms, Edward C.
Revision of the genus Melittobia (Chalcidoidea: Eulophidae) with the description of seven new species 271
Dahms, Edward C.
A review of the biology of species in the genus Melittobia (Hymenoptera: Eulophidae) with interpretation
and additions using observations on Melittobia australica 337
Dahms, Edward C.
An interpretation of the structure and function of the antennal sense organs of Melittobia australica
(Hymenoptera: Eulophidae) with the discovery of a large dermal gland in the male scape 361
Storey, R.I.
A new species of Aptenocanthon Matthews from north Queensland (Coleoptera: Scarabaeidae;
Scarabaeinae) 3g7
Kemp, A.
Spawning of the Australian lungfish Neoceratodus forsteri (Krefft) in the Brisbane River and in Enoggera
Reservoir, Queensland 39I
CovACEvicH, Jeanette
A biogeographically significant new species of Leiolopisma (Scincidae) from north eastern Queensland ....401
Thulborn, R.A. and Mary Wade
Dinosaur trackways in the Winton Formation (Mid-Cretaceous) of Queensland 413
Winter, J.W.
The Thornton Peak Melomys, Melomys hadrourus (Rodentia; Muridae): a new rainforest species from
northeastern Queensland, Australia 519
Morwood, M.J.
The Mt. Inglis cache; a new perspective on aboriginal material culture in the central highlands of
Queensland 54 j
Morwood, M.J. and M. Gibson
Incised stones from Glenormiston Station, S.W. Queensland 561
Durbidge, E. and R. Robbins
Two aboriginal shelters in southwestern Queensland 567
Obituary — William Alexander McDougall 1905-1982 576
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